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Temporal Trends in Juvenille Alosa Spp. Abundance and Relation to Predator Diets at the St. Johns River, Florida

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Temporal Trends in Juvenille Alosa Spp. Abundance and Relation to Predator Diets at the St. Johns River, Florida
Copyright Date:
2008

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Subjects / Keywords:
Anchovies ( jstor )
Catfish ( jstor )
Fish ( jstor )
Fisheries ( jstor )
Freshwater bass ( jstor )
Human geography ( jstor )
Juveniles ( jstor )
Menhaden ( jstor )
Predators ( jstor )
Shad ( jstor )

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University of Florida
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University of Florida
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7/24/2006

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TEMPORAL TRENDS IN JUVENILE ALOSA SPP. ABUNDANCE AND RELATION
TO PREDATOR DIETS AT THE ST. JOHNS RIVER, FLORIDA













By

NICHOLAS A. TRIPPEL


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

UNIVERSITY OF FLORIDA


2006



























Copyright 2006

by

Nicholas A. Trippel
































This document is dedicated to my family: Mom, Dad, and Aimee.















ACKNOWLEDGMENTS

This thesis could not have been completed without the hard work and dedication of

many people. I thank Drew Dutterer, Mo Bennett, Christian Barrientos, Galen Kaufman,

Mark "Captain No Fun" Rogers, Jason Dotson, Kristin Henry, Kristin Maki, Ginni

Chandler, Steve Larsen, Travis Tuten, Porter Hall, Matt Catalano, Greg Binion, Vince

Politano, Patrick Cooney, Kevin Johnson, Julie Harris, Adam Richardson, Vaughn

Maciena, Eric Nagid, and Jay Holder for their help with field work, lab processing,

finding literature, and coming along on the late-night trawling trips. I thank my

supervisory committee members (M. Allen, D. Murie, and R. McBride) for the assistance

and instruction they have given me throughout this study. I would like to thank my

supervisory committee chair (M. Allen) for all the valuable knowledge he has shared with

me, for encouraging me, for teaching me to work hard, for being such a knowledgeable

mentor, and for being the only other tiger fan in the lab.

I want to especially thank my parents (Don and Deb Trippel) for everything they

have done for me throughout the years, believing in me, and motivating me to be

successful.
















TABLE OF CONTENTS

page

A C K N O W L E D G M E N T S ................................................................................................. iv

L IST O F T A B L E S .. .... ........................................................................ ..... vi

LIST OF FIGURES ..................................... ................ .......... viii

A B ST R A C T ................. ...................................................................................... ......

INTRODUCTION .............. ................................... ..............

H history of A m erican Shad Fisheries........................................................ .................1
Am erican Shad Distribution and Biology.................................... ....................... 3

M E T H O D S ............................................................................ . 9

S ite S e le c tio n ................................................................................................................ 9
D ata C collection ............................................................................................. ....... 10
A n aly se s.................................................... 13

R E S U L T S ................................................................................19

Traw l Collections ............................................. 19
Electrofishing Samples ...... ......... ......... ......... .........21

D IS C U S S IO N .............. ..... ............ ................. ...................................................4 6

Juvenile Alosa spp. Abundance ............... ............ .......................46
Relative Prey Abundance and Occurrence in Predator Diets ...................................49
M anagem ent Im plications .............................................................. 53

LIST OF REFERENCES .......................... ..................58

B IO G R A PH ICA L SK ETCH ....................................................................... ...................64









v















LIST OF TABLES


Table page


1. Prey species classified as resident (collected in sample area year around) and
seasonal (marine species collected seasonally in sample area) in the Palatka area
of the St. Johns River, Florida from April 2004 through March 2005....................33

2. Summary of monthly mean trawl and electrofishing catch rates (fish/hour) for
juvenile Alosa spp. and total number of hours sampled using each gear by month
in the Palatka area of the St. Johns River from April 2004 through March 2005.... 35

3. Mean total lengths of juvenile American shad collected with trawl gear by month
in the Palatka area of the St. Johns River, Florida, from September through
December of 1969, 1970, and 2004. Number collected (N) and Standard Error
(S.E.) are shown by month for 2004 data. Months with no fish collected and no
data available are labeled as Na. Data from 1969 and 1970 were from Williams
and Bruger (1972). ............................................................... ............... 37

4. Number of predators sampled with electrofishing and how many contained
stomach contents (number and percent) in the Palatka area of the St. Johns
River, Florida from April 2004 through March 2005. .........................................38

5. Presence (x) or absence of prey items in diets collected from largemouth bass
Micropterus salmoides of the small size class (<365 mm) in the Palatka area of
the St. Johns River, Florida from April 2004 through March 2005......................39

6. Total numbers (N) of largemouth bass Micropterus salmoides from each of three
size classes (small < 365 mm, medium 366-432 mm, and large > 432 mm)
sampled each month containing stomach contents and the total number of prey
items found in these stomachs in the Palatka area of the St. Johns River, Florida
from April 2004 through M arch 2005 ....................................... .............. 42

7. Presence (x) or absence of prey items collected from predator species (in order
of number of diet samples collected from most on the left to least on the right)
other than largemouth bass Micropterus salmoides in the Palatka area of the St.
Johns River, Florida from April 2004 through March 2005 ...............................43









8. Total number, mean and median legnth (mm), and mean and median weight (g)
of each of the four most common prey species: Atlantic croaker (ATCR),
Atlantic menhaden (ATME), bay anchovy (BAAN), and threadfin shad (THSH),
found in each of three size classes (small < 365 mm, medium 366-432 mm, and
large > 432 mm) of largemouth bass, Micropterus salmoides, diets from April
2004 through March 2005 in the Palatka area of the St. Johns River, Florida........45















LIST OF FIGURES


Figure page

1. Map of Florida showing the St. Johns River in its entirety, along the Atlantic
coast of Florida .............. ......... ......... ........... ..................... 17

2. Map showing the Palatka area of the St. Johns River, Florida, including the
location of the state road 100 bridge, which was the mid-point of the study
sampling area. The sampling area was 5 km north and south of this bridge. This
figure shows that the Palatka area is the last narrow stretch of river before
widening and it remains this wide or wider until reaching the Atlantic Ocean....... 18

3 Comparing mean monthly trawl catch rates (+ standard error) of juvenile
American shad between 1970 and 2004 with 2004 mean monthly water
temperatures for the Palatka area of the St. Johns River, Florida.........................25

4. Mean number of Atlantic croaker, Atlantic menhaden, bay anchovy, threadfin
shad, and other fish species collected per five minute trawl with standard error
bars throughout 12 months of sampling on the Palatka area of the St. Johns
River Florida. ................................... ............................... .......... 26

5. Percent composition of each prey species from total trawl catches from January
through December at the Palatka area of the St. Johns River, Florida. The figure
shows only species that accounted for at least 5% of monthly trawl catches..........27

6. Percent by number of Atlantic croaker, Atlantic Menhaden, bay anchovy, and
threadfin shad found by month in diet samples (solid line) of all species of
predators collected by electrofishing and in trawl samples (dashed line) in the
Palatka area of the St. Johns River, Florida from April 2004 through March
2005 ....................... ................................. 28

7. Percent by number of prey items that were not Atlantic croaker, Atlantic
menhaden, bay anchovy, or threadfin shad found by month in diet samples of
small (<356 mm), medium (356 mm 432 mm), and large (>432 mm)
largemouth bass collected by electrofishing and in trawl samples in the Palatka
area of the St. Johns River, Florida from April 2004 through March 2005. ...........29

8. Percent frequency by number for crayfish, armored catfish (plated and sailfin
catfish), and blue crab found by month in all diet samples of largemouth bass
collected by electrofishing in the Palatka area of the St. Johns River, Florida
from April 2004 through M arch 2005 ........................................ .............. 30









9. Mean LOGIT values for prey items (>0 means more seasonal diet items, <0
means more resident diet items) by month found in stomach contents of three
size classes of largemouth bass (small <356 mm, medium 356 mm-432 mm and
large >432 mm TL) and for trawl catches in the Palatka area of the St. Johns
River, Florida from April 2004 through March 2005. ......................................... 31

10. Total percent caloric content (left panels, caloric values estimated only for four
species of prey fish therefore does not always represent total caloric intake) and
percent by number (right panels) for Atlantic croaker, Atlantic menhaden, bay
anchovy, and threadfin shad found in three size classes of largemouth bass
diets,(small < 356 mm, medium 365 mm-432 mm, and large > 432 mm)
collected by electrofishing in the Palatka area of the St. Johns River, Florida
from April 2004 through March 2005. Note : Only parts of croaker were found
in large size class largemouth bass. These were counted in total prey counts,
however parts were too small to estimate total weight and size and therefore
were left out of caloric estimates. .......... .. .... ............................... .............. 32















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

TEMPORAL TRENDS IN JUVENILE ALOSA SPP. ABUNDANCE AND RELATION
TO PREDATOR DIETS AT THE ST. JOHNS RIVER, FLORIDA


By

Nicholas A. Trippel

May 2006

Chair: Micheal S. Allen
Major Department: Fisheries and Aquatic Sciences

The St. Johns River, Florida, was once the largest recreational American shad

Alosa sapidissima fishery on the Atlantic coast. This fishery has drastically declined due

to decreased abundance of American shad. I assessed the temporal trends in juvenile

American shad relative abundance leaving the river, compared catch rates to those of a

similar study completed 35 years ago, and evaluated diets of piscivorous fish in the

sample area before, during, and after the juvenile American Shad had moved through the

area. I also compared predator diets to prey availability over 2004-2005 and estimated

caloric values of more-common prey to determine the seasonal variation in the relative

importance of these species to predator diets within the Palatka area of the St. Johns

River.

Trawl catch rates of juvenile American shad and other juvenile Alosa spp. were

extremely low. Only 23 American shad were collected during 12 months of sampling.

Highest catch rates occurred in October, which was similar to historic catch rates 35









years ago using similar trawling gear. Only one American shad and one hickory shad A.

mediocris were found in predator diets in 12 months of sampling and 1,747 total predator

diets measured.

The four most common species collected in trawl and diet samples were threadfin

shad Dorosoma petenense, bay anchovy Anchoa mitchilli, Atlantic croaker Micropogon

undulatus, and Atlantic menhaden. The number of these species found in largemouth

bass Micropterus salmoides (the most common predator) diets varied significantly by

month and size class of largemouth bass. Atlantic menhaden were found to be the most

energetically beneficial to predators, and I found that during months when they were

present all size classes of largemouth bass used them as prey. Correlation analysis

revealed that trawl catches and occurrence of individuals found in diet samples were

positively correlated for several species (a = 0.01), including Atlantic menhaden, (P

<0.01), sailfin catfish Pterygoplichthys multiradiatus (P < 0.01), and white catfish

Ameiurus catus (P = 0.01).

Management implications of this study include helping to successfully manage

the American shad fishery in this river, and to better relate the life history of common

prey items in this coastal river system to seasonal and ontogenetic diet shifts for the

common predators. I identified low abundance of juvenile Alosa spp. These juveniles

must deal with the predatory gauntlet, but were not seemingly preyed upon

disproportionately to their own abundance. Researchers need to look into flow and

habitat issues related to spawning success of adult American shad, water quality, and

pollution issues to see if these may be reasons we saw such low juvenile abundance.















INTRODUCTION

Five species of anadromous shad belonging to the genus Alosa are native to

Florida. Three of these species (American shad Alosa sapidissima, hickory shad A.

mediocris, and blueback herring A. aestivalis) are found within the St. Johns River

system. The other two (Alabama shad A. alabamae, and the skipjack herring A.

chrysochloris) are found on the Gulf of Mexico coast of northwest Florida (McBride

2000).

Clupeids are among the most economically important fishes, and worldwide no

other family of fishes is consumed or harvested in larger quantities (Scharpf 2003).

Florida's St. Johns River was once estimated to support the largest recreational fishery

for anadromous shad species on the Atlantic coast (Walburg and Nichols 1967; McBride

2005). Thus, research investigating the ecology of anadromous shads in Florida is

important for their conservation and management.

History of American Shad Fisheries

Until recent years, American shad supported some of the most important

commercial fishing industries in the United States (Williams and Bruger 1972; McBride

2000; McPhee 2002; Limburg et al. 2003; Scharpf 2003; McBride 2005). Although

Atlantic salmon Salmo salmar were first targeted by early Americans, American shad

soon replaced them and was favored for its flavorful roe and meat (McPhee 2002).

In the United States during the 1800s, many fisheries (along with the shad fishery)

grew rapidly (Walburg and Nichols 1967; McBride 2000). American shad were









harvested using multiple methods including fish dams, shad floats, fyke nets, seines,

pound nets, and gill nets (Scharpf2003). More-successful gear types overexploited

American shad populations for over a century now (Limburg et al. 2003; Scharpf 2003;

McBride 2005). Some American shad stocks in the New England states were reportedly

over-exploited as far back as 1830 (Gerstell 1998). In both 1889 and 1890, the shad

harvest in Florida alone was over 2 million pounds (McBride 2000; McBride 2005). In

1896, over 45 million pounds of American shad were harvested in the United States

(Stevenson 1899). Between 1904 and 1930, U.S. shad fisheries along the Northeastern

Atlantic coastline collapsed (Walburg and Nichols 1967) because of overfishing, water

pollution, and construction of dams (Leggett and Whitney 1972).

As catch rates declined, commercial fishers looked for new ways to harvest shad,

eventually developing the ocean-intercept fishery in the last 20 years. Even though shad

populations in rivers had declined, most commercial fishers now harvested shad in

offshore areas where all shad populations were congregated together, leading to a short-

term increase in landings (ASMFC 1999). Effectively, this meant that fishers were

simultaneously harvesting American shad populations from all of the rivers along the

Atlantic coast (McBride 2005). Commercial harvest of American shad was closed in

Maryland in 1982, and Virginia in 1994. As stocks became severely depleted, harvest

was also prohibited in Maine, New Hampshire, Massachusetts, and Rhode Island

(Scharpf2003). Inshore netting regulations imposed in the 1990s led to virtual closure of

Florida's commercial shad fishery (McBride 2005). To regulate the ocean-intercept

fishery, the Atlantic States Marine Fisheries Commission (ASMFC) mandated a 40%

reduction in effort for this fishery by the end of 2002 (ASMFC 1999). Froml995 to









2003, harvests declined, average annual U.S. domestic landings dropped to 2.7 million

pounds (NOAA 2005); compared to 50 million pounds in the 1890s, and 9 million

pounds the 1950s and 1960s (NOAA 2005). Because of these severe catch declines, and

concerns about the shad stock, all U.S. Atlantic shad ocean-intercept fisheries were

closed at the end of 2004 (ASMFC 1999; McBride 2005) thereby directing any remaining

targeted effort to within river systems.

American Shad Distribution and Biology

American shad are native to the Atlantic coast of North America, and are the

largest species of the family Clupeidae, reaching a maximum size of 5.5 kg, length of 76

cm, and age of 11 years (Froese and Pauly 2005). The native range of American shad is

the U.S. Atlantic coast from the St. Lawrence River (on the border of the United States

and Canada) southward to the St. Johns River in Florida (Brown et al. 1999; Limburg et

al. 2003; McBride 2005).

American shad are river-specific, with each river having a distinct spawning stock

(ASMFC 1999; McBride 2005). American shad on the southern end of their range are

semelparous, spawning once and dying; whereas fish at higher latitudes are iteroparous,

spawning repeatedly (Limburg 1996). All populations south of the Neuse River, North

Carolina, including the St. Johns River, are semelparous (Limburg et al. 2003; McBride

2005). Life history characteristics of these separate stocks vary from north to south,

depending on the environment of their home river (Limburg 1996). Characteristics that

vary include fecundity, size, and age at maturity (Leggett and Carscadden 1978; Limburg

et al. 2003). Fecundity is inversely related to latitude and semelparity. In general,

American shad at lower latitudes produce 3-5 times more eggs per kilogram of body

weight then their northern counterparts (Limburg et al. 2003). American shad at lower









latitudes are also smaller, and mature at younger ages than their northern counterparts

(Limburg et al. 2003).

Talbot and Sykes (1958) used tagging studies to learn that American shad from

Florida to Maine congregate together during summer and fall seasons in the Gulf of

Maine, whereas populations from Canada spend their summers in the St. Lawrence

estuary (Limburg et al. 2003). As water temperatures drop during winter, American shad

move south, with northern populations congregating on the Scotian Shelf; and southern

populations (those from Florida to Maine) congregating in the Middle Atlantic Bight, off

the coast of Florida (Limburg et al. 2003). American shad remain offshore in these

congregations until reaching maturity, then return to their natal streams to reproduce and

start the cycle again (Limburg et al. 2003).

Differences in spawning traits are partly due to the energy reserve needed for

spawning migration. Southern stocks migrate thousands of kilometers farther than

northern stocks, making migration energetically expensive. Nearly 50% of the Florida

American shad's total somatic energy reserves are used just to make it to their spawning

grounds (Leggett and Carscadden 1978; Glebe and Leggett 1981a; Glebe and Leggett

1981b). Another 20 to 30% of their somatic reserve is then expended during the

spawning process. Thus, southern American shad do not have the energy reserve needed

to return to sea as the northern populations do, resulting in semelparity (Leggett and

Carscadden 1978). Although southern stocks of American shad die during the spawning

season they return to the river, they may spawn multiple times during that spawning

season (Olney and McBride 2003; McBride 2005).









Shad migrations to spawning grounds correlate with water temperature (Leggett

and Whitney 1972). Timing of shad arrival at their spawning grounds along the Atlantic

coast varies with latitude but is similar in temperature (Leggett and Whitney 1972;

McBride 2000), which accounts for the earlier spawning in the south than in the north.

American shad begin to reach their spawning grounds as early as November in Florida,

and this run may continue through May (Leggett and Whitney 1972; Davis 1980;

McBride 2005). American shad near Canada begin spawning in May and continue

through July (Limburg et al. 2003).

American shad are broadcast spawners, with communal spawning lasting up to 10

weeks (Scharpf2003). Release of the demersal, free-drifting eggs (Williams and Bruger

1972), occurs at depths up to 10 meters, usually over sand or gravel substrate (Walburg

and Nichols 1967). Limburg et al. (2003) found the following conditions conducive to

larvae survival: water temperature above 200C, pH above 7, salinity level at least 10 ppt,

and minimum zooplankton levels of 50 organisms/L. Larvae drift downstream, maturing

into juveniles (Scharpf 2003).

During their first summer, juvenile American shad remain in a nursery area of the

river where they were spawned (Leggett and Whitney 1972). This nursery area is in the

natal stream; downstream from the spawning grounds where the river is tidally influenced

but contains relatively low salinity. Movement of juveniles from the spawning area to

the nursery area is triggered by increasing water temperatures and current (Williams and

Bruger 1972; McBride 2005). Movement from the nursery area to the ocean is triggered

by decreasing water temperatures. After juvenile American shad emigrate to sea, they

spend anywhere from three to six years migrating up and down the Atlantic coast until









eventually returning to their natal river (Limburg et al. 2003; Scharpf 2003). American

shad emigrate from rivers earlier in the north and may occur as late as December in

Florida (Williams and Bruger 1972).

Anadromous fishes such as American shad may play an important role as seasonal

prey item for predatory fish in coastal river systems. Not only do the spawning fish

stimulate primary production (Durbin et al. 1979), but migrating juveniles may provide

cyclic influxes of prey. These influxes may promote higher growth and survival of

predators (Yako et al. 2000). Juvenile anadromous fishes are faced with the survival

challenge of having to pass through a gauntlet of relatively stationary predators (Petersen

and DeAngelis 2000). Therefore, predators may have a large impact on the number of

emigrating juvenile shad in coastal river systems.

Few studies have assessed effects of piscivores on juvenile prey fish in coastal

river systems (Buckel et al. 1999). In many systems, predation may be the main factor

determining community structure (Raborn et al. 2003). Studies have been conducted in

salt ponds or enclosures (Wright et al. 1993), but few have measured the impacts in

natural systems. Diets of juvenile bluefish Pomatomus saltatrix consisted largely of

anadromous fishes (Buckel et al. 1999). Buckel et al. (1999) demonstrated that mortality

of juvenile striped bass Morone saxatilis was most strongly influenced by bluefish

predation.

Many studies have evaluated effects of predator mortality on the anadromous

salmonids of the United States' Pacific coast. Predation plays an important role in

juvenile mortality rates for anadromous salmonids (Ricker 1941). Parker (1968) found

that up to 85% of a year class of juvenile salmonids may be consumed by predators on









their way to the Pacific. Beamesderfer et al. (1990) determined that at John Day

Reservoir, Columbia River, British Columbia, northern squawfish Ptychocheilus

oregonensis consumed 1.7 million salmonids annually. At the same reservoir, Rieman et

al. (1991) found that the number of deaths from all predators totaled 2.7 million juvenile

salmonids annually. The high predation rates seen for juvenile salmonids in this

reservoir may be high due a large population of predators and the amount of time

juveniles remain in the reservoir before moving down river (Beamesderfer et al. 1990).

Predation during the seaward migration may greatly reduce each year's cohort, but the

seasonal influx of prey is likely to be highly beneficial to predators (Durbin et al. 1979).

Anadromous fishes may encourage higher survival and growth rates for predators

(Yako et al. 1999). Yako et al. (1999) found that largemouth bass Micropterus salmoides

in headwater lakes with seasonally available juvenile anadromous river herring, including

alewife Alosa pseudoharengus and blueback herring, may grow faster and attain larger

sizes than in lakes where these species were absent. Seasonally available herrings were

the most consumed fish species for both lakes examined (Yako et al. 1999). Largemouth

bass generally did not begin feeding on river herrings until August, probably due to

ontogenetic changes in the foraging behavior of young bass. Using bioenergetics models,

Yako et al. (1999) found that a diet of herring increased growth rates, and the presence of

trophy-size largemouth bass was positively correlated with the seasonal availability of

anadromous herrings in coastal Massachusetts. However, no studies have examined the

migration of juvenile American shad in southern populations and the impacts predators

have on their abundance and survival.









I evaluated the summer migration of juvenile American shad during their

downstream migration to sea, at the St. Johns River, Florida. Objective 1: estimate the

relative abundance and timing of juvenile shad migrating back to the Atlantic Ocean,

Objective 2: compare historical trawl catch rates of juvenile American shad from 1969-

1970 (Williams and Bruger 1972) to my results, Objective 3: examine and compare

differences between predator diets and prey availability over one year, and Objective 4:

examine predator diets before, during, and after American shad migration to assess

potential effects of predation on migrating juveniles. Effects of predation were measured

in terms of frequency of predator events. Estimated caloric values of more common prey

were also used to determine seasonal variation in the relative importance of these species

to predator diets within the Palatka area of the St. Johns River.















METHODS

Site Selection

The St. Johns River flows from south to north along Florida's Atlantic coast,

eventually entering the Atlantic Ocean near Jacksonville (Figure 1). It is the longest

river in Florida at about 500 km and has a watershed area of over 20,000 km2 (McBride

2005). This tannic-stained river is very slow flowing, having a slope averaging only 2

cm/km with a total gradient of less than 10 m (McBride 2005). It starts off as a small

meandering channel near Melbourne, then forms a series of lakes, and eventually reaches

a width of over three kilometers and spills into the Atlantic near Jacksonville (Figure 1).

I selected the Palatka area of the St. Johns River (Figure 2) as my study site

because it was described by Williams and Bruger (1972) as a summer nursery area used

by juvenile Alosa spp. before they emigrate to sea in the fall. The Palatka area of the river

is approximately 127 river km from the Atlantic Ocean (McBride 2005). It is tidally

influenced; however the salt wedge does not typically extend as far upriver as Palatka.

The State Road 100 bridge crosses the river in this area and was the midpoint of my

sampling area, with the sampling stretch extending 5 km north and 5 km south of this

bridge (Figure 2). This sample area was selected because of the overlap with Williams

and Bruger (1972) and because the area has a natural constriction of the river channel

(Figure 2), which could concentrate anadromous fishes and improve catches compared to

wider sections of the river.









Data Collection

I sampled the Palatka stretch of river using trawl gear to sample juvenile Alosa

spp. and prey abundance from April of 2004 through March of 2005. Prey abundance and

juvenile Alosa spp. populations were sampled using trawling techniques modified from

Williams and Bruger (1972). A surface trawl (3.7 m wide mouth, 4.6 m long body, 25.4

mm mesh body, 19.1 mm mesh bag, and 12.7 mm mesh liner) of the same dimensions as

used by Williams and Bruger (1972), was pulled between two jon boats (Williams and

Bruger used one boat) using 30.5 m warp lines and otter boards modified with hydrofoils

to keep the net on the surface (Massman et al. 1952, Trent 1967; Loesch et al. 1982).

Using a surface trawl has been proven to be an effective technique for sampling clupeids

offshore (i.e., open water) (Massman et al. 1952).

Sampling effort was partitioned by season and on either side of the bridge.

During months when Alosa spp. were expected, July through December, trawl samples

were conducted biweekly. Throughout the rest of the year, January through June, trawl

samples were conducted monthly. Williams and Bruger (1972) pulled trawls for 15

minute intervals after dark. For this study, trawls were pulled for 5 minutes after dark as

per Massman et al. (1952), as Loesch et al. (1982) reported increased capture of

American shad with surface trawls at night. Twelve trawls were pulled each sampling

trip. The areas north and south of the state road 100 bridge were each divided into six

sections from east to west across the river. Each section was sampled once in random

order on each sampling trip for a total of twelve trawls per trip. Trawls were pulled at an

average speed of 1.3 m/sec with trawl direction varied randomly and alternating against

and with the current. Trawling with the current was possible as current was often

minimal to non existent.









All fish from each trawl were identified to species and counted. I brought all

Alosa spp. back to the lab where they were weighed, measured, and otoliths were pulled

and mounted on glass slides using Thermo Shandon synthetic mountant cement.

Abundances of shad were indexed using the trawl catch rates (fish/minute), which was

the same index used by Williams and Bruger (1972). I kept several specimens of all prey

species in the area for reference and to construct an otolith key for identification of

stomach content analysis of predators (Whitfield and Blaber 1978).

I recorded total lengths to the nearest millimeter from all American shad collected

both trawling and electrofishing (described below). For the months of September

through December I compared these to mean lengths of American shad collected by

Williams and Bruger (1972) from 1969 and 1970. Because Williams and Bruger

recorded fork lengths, I measured fork length and total length on fish from 2005 and used

linear regression to estimate total lengths for fish from Williams and Bruger's (1972)

study.

Water temperature was monitored in the sample area throughout the study period

using Onset Computer Corporation Hobo model temperature loggers. The temperature

logger was located at the state road 100 bridge approximately 1 meter below the surface.

Water temperature was recorded every six hours throughout the year to the closest 1 C.

I used a 5.5 m aluminum electrofishing boat, model SR-18 with a GPP 9.0

electrofisher and 16 horsepower, 9,000 watt generator to collect only fish which were

potential predators. Juvenile Alosa spp. were also targeted for collection during

electrofishing samples, however no other prey species were collected. Predators were

sampled in the same weeks that trawls were pulled. I attempted to collect at least 50









predators of all species combined with stomach contents each sampling trip.

Electrofishing was conducted evenly throughout the shorelines of the sampled river

stretch. Pilings at bridge and powerline crossings were also sampled while electrofishing

to collect pelagic predators. I recorded total length (mm) and weight (g) of all predators

collected. I used transparent acrylic tubes to obtain stomach contents of live predatory

fish as described by Van Den Avyle and Roussel (1980). Diets were removed and

immediately placed on ice and returned to the lab for examination. Predators were

immediately released alive after diet samples were collected. Predators that could not

have contents removed via the tube were placed on ice and returned to the lab for diet

analyses.

Before analysis, diets were thawed and blotted dry. Organisms in the stomach

were identified using a microscope, dichotomous key, and otolith key, and then measured

and counted. I recorded total diet weight, individual diet item lengths and weights, prey

species, and body part if only partial prey items were found following the guide of Storck

(1986). Prey species were also classified as either resident (freshwater and estuarine

species found in the sample area of the river year round) or seasonal (usually marine

species and found in sample area only seasonally) (Table 1).

I retained 50 individuals of common prey species from the trawl sampling for

energetic information and diet identification keys, including Atlantic croaker

Micropogonias undulatus, Atlantic menhaden Brevoortia tyrannus, bay anchovy,

Anchoa mitchilli, and threadfin shad Dorosoma petenense. For these fish I measured

total length, standard length, vertebral column length, body depth, and length of the head.

Predictive regressions were determined based on fish size to then estimate total lengths









and weights for partial fish of these species found in diet samples. Total caloric content

rates for each of these four species were based on previous studies. I estimated caloric

content of prey found in diet samples using dry weight caloric values of: 4,638 cal/g and

76.3% moisture for Atlantic croaker (Thayer et al. 1973), 5,115 cal/g and 65% moisture

for Atlantic menhaden (Steimle Jr. and Terranova 1985), 5,067 cal/g and 72% moisture

for bay anchovy (Steimle Jr. and Terranova 1985), and 4,835 cal/g and 79.2%for

threadfin shad (Strange and Pelton 1987). These caloric values were then multiplied by

the total measured weight of prey or total estimated weight for partial prey items to

determine total calories predators consumed of each prey type.

These caloric values were calculated at various locations and various times of the

year; however, I applied them as general estimates. Atlantic croaker caloric rates were

determined from samples of adult fish collected in estuaries near Beafort, North Carolina

(Thayer et al. 1973). Atlantic menhaden and bay anchovy caloric rates were determined

from the most commonly collected sizes of these species along the Atlantic coast from

Nova Scotia to North Carolina (Steimle Jr. and Terranova 1985). Threadfin shad caloric

rate estimates were from fish collected throughout the year in two Tennessee reservoirs

(Strange and Pelton 1987). These values were determined for bay anchovy and threadfin

shad of similar size to those I collected, while most Atlantic croaker and Atlantic

menhaden I collected were juveniles, thus smaller than those used to determine caloric

values.

Analyses

Raw data from Williams and Bruger's trawl catch rates were archived and made

available to me by Julie Harris of Florida Fish And Wildlife Research Institute. Trawl

catch rates for American shad for this project and those of Williams and Bruger (1972)









were converted to fish caught per hour to allow a qualitative comparison of catch rates

between time periods. Total trawl catch rates were also summarized. Percent

composition of fish species in the trawl and diet samples were estimated for each month.

A cumulative list of all species found in diets and trawls was compiled from which

presence or absence was recorded for each trawl and diet. I determined the percent each

species made up by month of the total trawl catch and total diet samples.

Statistical Analysis Software (SAS 2000) was used to run statistical analyses. I

used correlation analysis to assess relationships between the percent each prey species

made up of the monthly trawl catches and the percent that those same species made up of

the total monthly diet samples. Correlations were done for each prey type across

months.

Largemouth bass was by far the most common predator in this study, and thus

allowed much more detailed diet analyses. I evaluated whether diet contents varied with

largemouth bass size and season for the most common prey species. Four prey species

represented 95% of all prey collected in trawls and they were used to examine

largemouth bass diets in greater detail: Atlantic croaker, Atlantic menhaden, bay

anchovy, and threadfin shad. I compared the frequency of occurrence in the diets

monthly and also the total estimated caloric values that each species comprised for that

month's diet samples. Largemouth bass were divided into three size classes: small

(<356mm), medium (356mm-432mm) and large (>432mm), and energetic comparisons

of total kilocalories consumed from each of the four prey species were made among the

three largemouth bass size classes throughout the year I used the four common prey

species and collectively grouped all other prey items into a category defined as "other". I









used a log transformation plus 0.5 to improve normality of the variance around each

mean. Two-way analyses of variance (ANOVA, Procedure GLM, SAS 2000) ANOVA's

assessed how percent frequency by number and caloric value of each species found in

diets (dependent variables) varied by month (12 months sampled), largemouth bass size

class (3 size classes) and the interaction between these factors. Separate ANOVA's were

used for each prey species and the "other" category. When a two-way ANOVA was

significant, Least Squared Means test (LS Means, SAS 2000) was performed to

determine where significant differences occurred. Differences were declared significant

at P < 0.05 for all analyses, and this value was selected to reduce the chance of Type II

error.

To analyze the importance of seasonal versus resident prey for predators, prey

fishes were categorized as resident or seasonal. Resident fishes were prey species

collected throughout the entire year in the sample area, and prey species classified as

seasonal were the marine species found only seasonally in the sample area. I estimated a

ratio defined as the LOGIT for each individual predator diet collected as:


Log (number of seasonal prey items + 0.5/number of resident prey items + 0.5).









Thus, if this value was positive there were more seasonal items in that particular diet,

whereas if it were negative there were more resident items in that diet. A two-way

ANOVA (Procedure GLM SAS 2000) was used to assess differences in LOGIT values

(dependent variable) by month (12 months sampled), largemouth bass size class (3 size

classes) and the interaction between these two factors. A least squares means test (LS

Means) was used to separate the means if the ANOVA was significant (SAS 2000). This

analysis was used to assess whether largemouth bass diet contents shifted when seasonal

prey were available.

I also assessed how trawl catches of seasonal versus resident prey fish varied through the

year. A repeated measures ANOVA (Procedure Mixed SAS 2000) was used to test if the

total number of individuals (log n) collected in trawl samples (dependent variable) varied

by month, prey species class (resident or seasonal), and the interaction between these two

independent variables. I used the autoregressive order 1 covariance matrix structure,

which models correlations decreasing with distance in time (Littell et. al. 1996). Each

trawl repetition was nested randomly within site (six transverse river sampling as sections

described above). If significant, the LS Means test was used to test for differences

among means.





















0 50 100 200 Kilometers
I I I I I I I I I


I I 1 1i 1


1" I I 1.1 I IIn


created by NickTrippel, October of2005


Figure 1. Map of Florida showing the St. Johns River in its entirety, along the Atlantic
coast of Florida.




















































Figure 2. Map showing the Palatka area of the St. Johns River, Florida, including the
location of the state road 100 bridge, which was the mid-point of the study
sampling area. The sampling area was 5 km north and south of this bridge.
This figure shows that the Palatka area is the last narrow stretch of river
before widening and it remains this wide or wider until reaching the Atlantic
Ocean.















RESULTS

Trawl Collections

I collected a total of 27 fish species in the trawls representing 13 families (Table

1). Most species were classified as resident because they were collected in the sample

area throughout all 12 months of the year, but six species were classified as seasonal

based on the fact that they were only collected seasonally in this stretch of the St. Johns

River, Florida. Table 1 also includes prey items found in diets and not collected in

trawls.

I collected all three Alosa spp. while trawling. Blueback herring occurred earliest

in the year and accounted for a majority (N = 37) of the Alosa spp. collected (Table 2).

American shad ranked second in total catch with 23. I collected a total of six hickory

shad between the months of May and October (Table 2). Trawl catch rates were highest

in May for blueback herring whereas catch rates for both American shad and hickory

shad were highest in October (Table 2).

Throughout the 12-month sampling period a total of 23 juvenile American shad

were collected using the trawl (Figure 3). American shad were collected beginning in

June with the last catches occurring in December. Most juvenile American shad were

collected from September through November with catch rates being the highest in

October, when nine American shad were collected. Trawl catch rates and temporal

trends of juvenile American shad occurrence were similar to those of Williams and

Bruger in 1970 (Figure 3). It appeared that decreasing water temperatures triggered









juvenile American shad to emigrate (Figure 3). However, I never detected large pulses of

juvenile shad moving through the area in the trawl catches. Mean total length of

American shad was similar between this study and Williams and Bruger (1972) for all

sample months (Table 3).

Juvenile seasonal species were commonly collected in the sample area during

spring and fall of the sample period. Mixed model analyses determined that mean

catches of resident and seasonal prey species in the trawl samples varied throughout the

year, with significant interaction between month and species classification (resident or

seasonal) (P<0.01). This meant that both the number of total prey fish caught and prey

species (resident or seasonal) caught varied by month. For example, seasonal species had

lower occurrence in summer than in spring or fall, whereas resident species catch was

highest in summer and winter.

Monthly patterns in abundance of the most common resident and seasonal species

were therefore further examined using ANOVA and LS MEANS tests. The four most

abundant species collected with the trawl were threadfin shad, bay anchovy, Atlantic

menhaden, and Atlantic croaker. I collected numerous other species infrequently (Figure

4). Threadfin shad and bay anchovy appeared in trawl samples consistently throughout

the year. Atlantic croaker appeared in all 12 months but catch rates were much higher

during February June (Figure 4). I sampled only a few mature Atlantic croaker and

most were small juveniles (<50 mm TL). Atlantic menhaden appeared in trawl samples

only during the months of September, October, and November. During January and

November, white catfish Ameiurus catus accounted for 25% and 9% respectively of total

trawl catch (Figure 5).









Electrofishing Samples

Some species of prey targeted by trawling were also captured with electrofishing

gear. While electrofishing for predators I collected 55 juvenile American shad, 194

juvenile blueback herring, and 4 juvenile hickory shad. More juvenile Alosa spp. were

collected while electrofishing for predators than while targeting them using the trawl, but

catch rate via electrofishing was highest later in the fall than the trawl catches of Alosa

spp. (Table 2).

Predator species sampled by electrofishing included: largemouth bass, striped

bass Morone saxatilis, black crappie, Florida gar Lepisosteus platyrhincus, longnose gar

Lepisosteus osseus, mangrove snapper Lutjanus griseus, red drum Sciaenops ocellatus,

ladyfish Elops saurus, channel catfish Ictalurus punctatus, white catfish, brown bullhead

A. nebulosus, bowfin Amia calva, southern flounder Paralichthys lethostigma, and chain

pickerel Esox niger. I collected a total of 1,747 predators of which 714 contained

stomach contents, and largemouth bass was by far the most common predator collected

(Table 4).

Prey items collected from potential predators by tubing were often highly digested

and difficult to identify. These items were given the classification unidentifiable (UID).

The percent of UID stomach items varied by month and comprised anywhere from 11-

45% of total monthly stomach contents by number. Throughout the entire year only one

juvenile American shad and one juvenile Hickory shad were found in predator diets, both

from largemouth bass diets. Prey items that comprised more than five percent by number

of all diets for each month's total diet samples included: threadfin shad, bay anchovy,

Atlantic croaker, Atlantic menhaden, bluegill Lepomis macrochirus, blue crabs

Callinectes sapidus, clown goby Microgobius gulosus, white catfish, plated catfish









Hoplosternum littorale, sailfin catfish Pterygoplichthys multiradiatus, and unidentified

crayfish species. The most common species collected in trawls compared to those found

in diet samples by month are shown in Figure 6. This showed that as prey availability

increased, occurrence in diets also increased. Figure 7 shows the percent frequency by

number of diet items from anything other than the four most commonly collected species

in trawl samples, collectively grouped as "other". Large size class largemouth bass

commonly preyed upon Lepomis spp. which were grouped as "other" (Table 5). The

number of "other" species peaked in fall when we experienced extremely high water.

During this time, the most common prey items included crayfish species, plated catfish,

and sailfin catfish (Figure 8).

Trawl catches and occurrence of individuals found in predator diet samples were

significantly correlated for several species. These species included Atlantic menhaden

(P<0.0001), sailfin catfish (P < 0.0001), and white catfish (P= 0.0009) (Figures 4, 6, and

7). For all other species (n = 38) the Proc CORR procedure revealed no significant

correlations.

Diet contents for largemouth bass varied with fish size and across months. There

were 114 largemouth bass in the large size class, 205 in the medium size class, and 267 in

the small size class (Table 6). The GLM procedure revealed that LOGIT values for diets

varied significantly for the interaction between month and size class (P= 0.0003).

Throughout the year different size classes of largemouth bass were feeding on different

prey species, and that also throughout the year these prey varied from seasonal to

resident. During the winter all size classes of largemouth bass were preying upon bay

anchovy when other prey species were less prevalent. Mean LOGIT values by month









revealed that trawl and diet samples collected contained higher proportions of resident

species than seasonal species, except for large and medium size class bass diets in April

and large size class bass diets in June (Figure 9). This was due to high numbers of

threadfin shad and bay anchovy collected in trawls and largemouth bass diets.

Exceptions included the month of April when medium and large size class fish

commonly preyed upon blue crabs and June when Atlantic croakers were found in diets

of large size class largemouth bass. After July all LOGIT values were negative as

threadfin shad became the most common prey item through summer and fall.

Throughout fall and winter LOGIT values remained negative as all size classes of fish

preyed upon armored catfish, crayfish, and threadfin shad. Trawl catches had negative

LOGIT values throughout the entire year of sampling compared to diets, indicating that

diet contents of all predators contained higher seasonal prey composition than the trawl

samples.

Next to largemouth bass, black crappie was the most abundant predator collected

with 56 containing stomach contents (Table 7). The most common prey items found in

black crappie diets were bay anchovy (20%), also threadfin shad and insects each

accounted for 9% of the total prey items by number. Bay anchovy (35%) and threadfin

shad (18%) were the most common items found in the diets of 16 striped bass. Thirteen

longnose gar sampled contained prey items with threadfin and gizzard shad were the

most common prey items accounting for 56% of total prey items by number. Atlantic

croaker made up for another 19% of prey items found in the longnose gar diets. Channel

catfish consumed various Lepomis spp., crayfish, white catfish, and even pork chops.






24


Using published caloric values for the four most common prey species, I

estimated from diets the monthly caloric intake that each species comprised for each

largemouth bass size class by month. Numbers and sizes of prey consumed by each size

class of largemouth bass are shown in Table 8. I also compared this to the number of

individuals of each species found in diets by month. Bay anchovy and threadfin shad

were often the most common prey item by number while threadfin shad and menhaden

often made up for a majority of consumed energy (Figure 10).























9 35


30


25 -
o 6 -

5- 1970 Trawls I 20
.c 2004 Trawls
c, -*-Water Temperature E
o 4 -15 .

E3
10
2



0 0-

Apr May Jun Jul Aug Sep Oct Nov Dec




Figure 3 Comparing mean monthly trawl catch rates (+ standard error) of juvenile
American shad between 1970 and 2004 with 2004 mean monthly water
temperatures for the Palatka area of the St. Johns River, Florida.















Atlantic croaker


Atlantic menhaden











other species








< A- :zf5


bay anchovy





t A ,




threadfin shad








L S C ) -
.- .* I ^ = < -


Month


Figure 4. Mean number of Atlantic croaker, Atlantic menhaden, bay anchovy, threadfin
shad, and other fish species collected per five minute trawl with standard error
bars throughout 12 months of sampling on the Palatka area of the St. Johns
River Florida.











100
90
80

70 1 white catfish
60 threadfin shad
S50o bay anchovy
40 0 Atlantic menhaden

30 Atlantic Croaker
20
10
0
a- ^ o a o Mu 0
< z a -, L





Figure 5. Percent composition of each prey species from total trawl catches from
January through December at the Palatka area of the St. Johns River, Florida.
The figure shows only species that accounted for at least 5% of monthly trawl
catches.




























Atlantic croaker


100

80

60

40

20


0


W 0 0 . "-



It
0 l

LL-


w 1 bay anchovy
a- I


60

40

20


ai ^ a C n > 0 = -0 E
<2-, .QCOZ )LJI


Atlantic menhaden


- >' = C3 aM C C =
CLO 0 W) M W3
1 0 ;z4O )ZO -W


threadfin shad


U I I I
S>. C .3 0)*
a S -,


Diet
......... Trawl
















Figure 6. Percent by number of Atlantic croaker, Atlantic Menhaden, bay anchovy, and

threadfin shad found by month in diet samples (solid line) of all species of

predators collected by electrofishing and in trawl samples (dashed line) in the

Palatka area of the St. Johns River, Florida from April 2004 through March

2005.


a > Ca C n
0) 0 C) Cu 03 r
d 0 Q -W M
a) z~~ L


Month





























I


/ D#I 7\ -


r : :)5


0 z


II
I'
I'
I'
I

I'


- Small
- Medium
- Large
- -Trawl


U_ a
Gi CU I


Figure 7. [o]Percent by number of prey items that were not Atlantic croaker, Atlantic
menhaden, bay anchovy, or threadfin shad found by month in diet samples of
small (<356 mm), medium (356 mm 432 mm), and large (>432 mm)
largemouth bass collected by electrofishing and in trawl samples in the
Palatka area of the St. Johns River, Florida from April 2004 through March
2005.










20
18
16
14
12 Crayfish
10- Armored Catfish
8- \ Blue Crab
6- \ '
4
2 \
0 A\

I< M < ) Z a -' u_ a




Figure 8. Percent frequency by number for crayfish, armored catfish (plated and sailfin
catfish), and blue crab found by month in all diet samples of largemouth bass
collected by electrofishing in the Palatka area of the St. Johns River, Florida
from April 2004 through March 2005.



























S- Trawl
'Small
-. Medium
O Large
-4-


-5


-6


-7 ..
Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar





Figure 9. Mean LOGIT values for prey items (>0 means more seasonal diet items, <0
means more resident diet items) by month found in stomach contents of three
size classes of largemouth bass (small <356 mm, medium 356 mm-432 mm
and large >432 mm TL) and for trawl catches in the Palatka area of the St.
Johns River, Florida from April 2004 through March 2005.















100 100
80 80
60 60
40 40
20 20
0
o -o


U Medium Medium
100 1 100
I--
S80 s80
60 0 60
L 40 4E 40
20 20







80 80


40 40
20 20
IL O_ I _
o m o



100 100 0















threadfin shad






Figure 10. Total percent caloric content (left panels, caloric values estimated only for
four species of prey fish therefore does not always represent total caloric
intake) and percent by number (right panels) for Atlantic croaker, Atlantic
menhaden, bay anchovy, and threadfin shad found in three size classes of
largemouth bass diets,(small < 356 mm, medium 365 mm-432 mm, and large

> 432 mm) collected by electrofishing in the Palatka area of the St. Johns
River, Florida from April 2004 through March 2005. Note: Only parts of

croaker were found in large size class largemouth bass. These were counted
in total prey counts, however parts were too small to estimate total weight and

size and therefore were left out of caloric estimates.










Table 1. Prey species classified as resident (collected in sample area year around) and
seasonal (marine species collected seasonally in sample area) in the Palatka
area of the St. Johns River, Florida from April 2004 through March 2005.


Family
Fishes
Atherinidae



Belonidae


Callichthyidae

Centrachidae


Clupeidae






Cyprinidae

Cyprinodontidae

Eleotridae


Elopidae


Engraulidae


Gerreidae

Gobiidae


Resident


Brook silverside (Labidesthes sicculus)
Atlantic silverside (Menidia menidia)


Atlantic needlefish (Stronvglura marina)


Plated catfish (Hoplosternum littorale)

Black crappie (Pomoxis nigromaculatus)
Bluegill (Lepomis macrochirus)
Largemouth bass (Micropterus salmoides)
Redbreast sunfish (Lepomis auritus)
Redear sunfish (Lepomis microlophus)
Spotted sunfish (Lepomis punctatus)
Warmouth (Lepomis gulosus)

Gizzard shad (Dorosoma cepedianum)
Threadfin shad (Dorosoma petenense)


Seasonal


American Shad (Alosa sapidissima)
Atlantic menhaden (Brevoortia
tyrannus)
Blueback herring (Alosa aestivalis)
Hickory shad (Alosa mediocris)


Golden shiner (Notemigonus crvsoleucas)

Seminole killifish (Fundulus seminolis)

Fat sleeper (Dormitator maculatus)


Ladyfish (Elops saurus)


Bay anchovy (Anchoa mitchilli)


Irish Pompano (Diapterus auratus)


Clown goby (Microgobius gulosus)
Naked goby (Gobiosoma bosc)












Table 1. Continued.

Family Resident Seasonal
Ictaluridae Channel catfish (Ictalurus punctatus)
White catfish (Ameirus catus)

Loricariidae Sailfin catfish (Pterygoplichthvs
multiradiatus)

Mugilidae Striped mullet (Mugil cephalus)


Poeciliidae Sailfin molly (Poecilia latipinna)


Sciaenidae Atlantic croaker (Micropogonias
undulates)
Silver perch (Bairdiella chrysoura)
Crustaceans
Portunidae Blue crab (Callinectes sapidus)











Table 2. Summary of monthly mean trawl and electrofishing catch rates (fish/hour) for
juvenile Alosa spp. and total number of hours sampled using each gear by
month in the Palatka area of the St. Johns River from April 2004 through
March 2005.



American shad


Hours Trawled

1
1
1
1
3
2
2
2
1
1
1
1


Electrofishing
Catch Rate
0
0
0
0
0.6
2.4
1.98
6.96
3.24
0
0
0


Hours of Pedal
Time
1.7
1.0
1.7
2.3
1.7
3.8
4.1
4.6
1.6
4.2
3.4
3.0


Blueback Herring


Trawl Catch Rate

1
12
1
1
0.3
4
4
1.5
0
2
0
0


Hours Trawled

1
1
1
1
3
2
2
2
1
1
1
1


Electrofishing
Catch Rate
0
0
0
0
0
6.1
3.9
13.5
36.1
4.9
0.3
0


Hours of Pedal
Time
1.7
1.0
1.7
2.3
1.7
3.8
4.1
4.6
1.6
4.2
3.4
3


Month

April
May
June
July
August
September
October
November
December
January
February
March


Trawl Catch
Rate
0
0
2
0
0.33
2.5
4.5
2
2
0
0
0


Month

April
May
June
July
August
September
October
November
December
January
February
March











Table 2. Continued



Hickory Shad


Hours Trawled

1
1
1
1
3
2
2
2
1
1
1
1


Electrofishing
Catch Rate
0
0
0
0
0
0.54
0.24
0.24
0
0
0
0


Hours of Pedal
Time
1.7
1.0
1.7
2.3
1.7
3.8
4.1
4.6
1.6
4.2
3.4
3.0


Month

April
May
June
July
August
September
October
November
December
January
February
March


Trawl Catch
Rate
0
2
0
0
0.33
0
1.5
0
0
0
0
0







37


Table 3. Mean total lengths of juvenile American shad collected with trawl gear by
month in the Palatka area of the St. Johns River, Florida, from September
through December of 1969, 1970, and 2004. Number collected (N) and
Standard Error (S.E.) are shown by month for 2004 data. Months with no fish
collected and no data available are labeled as Na. Data from 1969 and 1970
were from Williams and Bruger (1972).

Month 1969 1970 2004 2004 (N) 2004 (S.E.)
June Na Na 66 2 1
July Na Na Na 0 Na
August Na Na 80 1 Na
September 69 58 70 5 1.93
October 75 82 85 9 2.47
November 89 95 79 4 2.27
December 94 106 93 2 8













Table 4. Number of predators sampled with electrofishing and how many contained
stomach contents (number and percent) in the Palatka area of the St. Johns
River, Florida from April 2004 through March 2005.


Predator Species

Largemouth Bass
Black Crappie
Longnose Gar
Florida Gar
Striped Bass
White Catfish
Channel Catfish
Chain Pickerel
Ladyfish
Southern Flounder
Brown Bullhead
Red drum
Mangrove Snapper
Totals


Number Examined
for Prey
1,443
120
36
35
29
25
18
14
10
8
5
3
1
1,747


Number With Prey


Percent With Prey











Table 5. Presence (x) or absence of prey items in diets collected from largemouth bass
Micropterus salmoides of the small size class (<365 mm) in the Palatka area
of the St. Johns River, Florida from April 2004 through March 2005.


Size Class
Small
American shad
Atlantic croaker
Atlantic menhaden
Atlantic needlefish
Atlantic silverside
bay anchovy
blue crab
brook silverside
channel catfish
clown goby
crayfish
fat sleeper
gizzard shad
golden shiner
grass shrimp
hickory shad
Irish pompano
ladyfish
largemouth bass
Lepomis spp.
mud crab
naked goby
pink shrimp
plated catfish
sailfin molly
Seminole killifish
silver perch
striped mullet
sailfin catfish
threadfin shad
white catfish


Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec


x x x x
x x


x x
x x x
x
x


x x x
x


X X X


x x


x x x


x x


x x
x


x x


x x


x x x x x x
X X X X
X


x
x x x x







40



Table 5, continued. Presence (x) or absence of prey items in diets collected from
largemouth bass Micropterus salmoides of the medium size class (366 mm -
432 mm) in the Palatka area of the St. Johns River, Florida from April 2004
through March 2005.


Size Class
Medium
American shad
Atlantic croaker
Atlantic menhaden
Atlantic needlefish
Atlantic silverside
bay anchovy
blue crab
brook silverside
channel catfish
clown goby
crayfish
fat sleeper
gizzard shad
golden shiner
grass shrimp
hickory shad
Irish pompano
ladyfish
largemouth bass
Lepomis spp.
mud crab
naked goby
pink shrimp
plated catfish
sailfin molly
Seminole killifish
silver perch
striped mullet
sailfin catfish
threadfin shad
white catfish


Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec


x x


x x


x
x x


x x
x
x


x x x


x x


x x x


x x


x x x x x x x x x







41



Table 5, continued. Presence (x) or absence of prey items in diets collected from
largemouth bass Micropterus salmoides of the large size class (>432 mm) in
the Palatka area of the St. Johns River, Florida from April 2004 through
March 2005.


Size Class
Large
American shad
Atlantic croaker
Atlantic menhaden
Atlantic needlefish
Atlantic silverside
bay anchovy
blue crab
brook silverside
channel catfish
clown goby
crayfish
fat sleeper
gizzard shad
golden shiner
grass shrimp
hickory shad
Irish pompano
ladyfish
largemouth bass
Lepomis spp.
mud crab
naked goby
pink shrimp
plated catfish
sailfin molly
Seminole killifish
silver perch
striped mullet
sailfin catfish
threadfin shad
white catfish


Jan Feb Mar Apr May


x x


Jun Jul Aug Sep Oct Nov Dec
x


x
x x
x x x


x x


x x x x x


x x


x x x x


x x


x
X X X X X X


x X X


x x x x


x
x x
x












Table 6. Total numbers (N) of largemouth bass Micropterus salmoides from each of
three size classes (small < 365 mm, medium 366-432 mm, and large > 432
mm) sampled each month containing stomach contents and the total number
of prey items found in these stomachs in the Palatka area of the St. Johns
River, Florida from April 2004 through March 2005.


Total N Total N of
Prey items
Small Small
21 24
16 37
28 111
23 44


Total N Total N of
Prey items
Medium Medium
5 7
9 20
14 23
10 14


Total N Total N of
Prey items
Large Large
5 6
5 11
5 6
8 11
7 12
17 36
14 25
25 48
17 35
4 14
2 2
5 13
114 219


Month


April
May
June
July
August
September
October
November
December
January
February
March
Total










Table 7. Presence (x) or absence of prey items collected from predator species (in order
of number of diet samples collected from most on the left to least on the right)
other than largemouth bass Micropterus salmoides in the Palatka area of the
St. Johns River, Florida from April 2004 through March 2005.



Predator Species black crappie striped bass longnose channel Florida gar
(n=56) (n=16) gar catfish (n=10) (n=9)
Prey Species (n=13)
Atlantic croaker x x
Atlantic menhaden x x
Atlantic needlefish x
bay anchovy x x
blue crab x
brook silverside x
channel catfish
crayfish x
grass shrimp x
gizzard shad x
insects x
Irish pompano x
ladyfish x
largemouth bass
Lepomis spp. x x
plated catfish x
pork chops x
threadfin shad x x x x
sailfin catfish
white catfish x










Table 7. Continued


Predator Species bowfin chain white ladyfish redfish
(n=8) pickerel catfish (n=3) (n=2)
(n=7) (n=6)
Atlantic croaker x
Atlantic menhaden
Atlantic needlefish x
bay anchovy x
blue crab x x
brook silverside
channel catfish x
crayfish x
grass shrimp x x
gizzard shad
insects
Irish pompano
ladyfish
largemouth bass x
Lepomis spp. x
plated catfish
pork chops
threadfin shad x x
sailfin catfish x
white catfish











Table 8. Total number, mean and median length (mm), and mean and median weight (g)
of each of the four most common prey species: Atlantic croaker (ATCR),
Atlantic menhaden (ATME), bay anchovy (BAAN), and threadfin shad
(THSH), found in each of three size classes (small < 365 mm, medium 366-
432 mm, and large > 432 mm) of largemouth bass, Micropterus salmoides,
diets from April 2004 through March 2005 in the Palatka area of the St. Johns
River, Florida.

Size Class Species Number Mean Median Mean Median
Length Length Weight (g) Weight (g)
(mm) (mm)
Small ATCR 5 63 58 3.8 2.0
Medium ATCR 2 80 80 10.3 10.3
Large ATCR 3 N/A N/A N/A N/A
Small ATME 7 109 110 13.1 12.3
Medium ATME 17 117 115 28.2 15.6
Large ATME 7 112 108 12.8 12.9
Small BAAN 152 34 31 0.5 0.2
Medium BAAN 33 54 56 1.2 0.9
Large BAAN 17 45 46 0.7 0.7
Small THSH 100 63 63 2.7 2.0
Medium THSH 104 70 70 3.3 2.7
Large THSH 29 72 72 3.8 2.9















DISCUSSION

Juvenile Alosa spp. Abundance

Night-time trawl samples collected low numbers of juvenile American shad,

blueback herring, and hickory shad. However, catch rates were similar to those of

Williams and Bruger (1972) using the same trawling methods in 1969 and 1970. Both

studies found highest catch rates occurring in October. Most American shad were

collected from September through December near Palatka, which is described as the

northern extent of the nursery area (Williams and Bruger 1972). A few juvenile

American shad were collected in June. Williams and Bruger (1972) sampled the entire

river and found some juveniles entered brackish areas of the river during summer but did

not find shad leaving the mouth of the river until November.

From September through December mean monthly water temperatures ranged

from 27 C down to 19 OC. Mean water temperature for the month of October when most

fish were collected was 24 C similar to mean temperatures in 1970. Emigration this

time of year begins much later than fish at higher latitudes (October through November in

North Carolina: Davis and Cheek 1966; November through March in Florida: Williams

and Bruger 1972; June through November in New York: Limburg et al. 2003). My

occurrence of American shad corresponded with declining water temperatures similar to

Williams and Bruger (1972), although it is not clear that declining water temperatures

caused movement of American shad in this system relative to other potential mechanisms

(e.g., photoperiod, light availability).









While electrofishing I collected juvenile American shad from August through

December with the highest number being collected in November. On 18 November

2004, I collected 26 juvenile American shad while electrofishing for predators. This

suggests that this may have been a time when large numbers of shad were moving

through the area. However, trawl sampling in November did not reveal a similar trend.

As I collected many more fish electrofishing than trawling, the surface trawl might not be

an effective sampling gear for collecting juvenile American shad in this region.

Williams and Bruger (1972) and McBride (2000) reported that hickory shad were

the first of the Alosa spp. to make their spawning runs up the St. Johns River, November

through February, followed by the American shad and blueback herring in December

through April. Loesch et al. (1982) found that in Virginia Rivers American shad

spawning precedes that of blueback herring. O'Leary and Kynard (1986) stated that

juvenile blueback herring will emigrate slightly earlier than American shad. My data

revealed this same trend as I found juvenile blueback herring as early as April with 12

being collected in May, the highest number ofbluebacks collected during any month.

Juvenile hickory shad appeared in May before the first American shad arrived in June

following the suggested trend.

Although catch rates were low for juvenile Alosa spp., trawl gear did effectively

sample various other prey species. The sample area was impacted by two major

hurricanes during the fall of my sample period. Twice and only weeks apart, the river

flooded and experienced extremely high flow rates. I did not collect high numbers of

juvenile Alosa spp. during these time periods, evidence that the high flows did not trigger









juveniles to emigrate earlier than normal. However, the high flow conditions could have

reduced my sampling efficiency and contributed to low catch.

Nevertheless, low catches of juvenile Alosa spp. may be indicative of low

population size. Reduced populations of American shad have occurred along the Atlantic

coast and can be attributed to factors such as decreasing water quality, changes in flow,

increased sediment load, overharvest by recreational anglers and previous commercial

fishers, and dams (Walburg and Nichols 1967; Williams and Bruger 1972; Hightower et

al. 1996; Ross et al. 1997; McBride 2000; Olney and Hoenig 2001; Limburg et al. 2003;

Scharpf2003). My catch of Alosa spp. was similar to values from the early 1970's,

indicating a lack of large changes in juvenile abundance between time periods. Thus, my

results do not indicate a large reduction in abundance, albeit with both time periods

having a relatively low sample size.

Atlantic estuaries are important nursery grounds for various anadromous species

(Juanes et al. 1993), and seasonal influxes of anadromous prey species such as juvenile

American shad and blueback herring may play an important role in the diets of relatively

stationary predators such as largemouth bass (Yako et al. 2000). Yako et al. (2000) found

that largemouth bass in coastal Massachusetts rivers with such calorie rich prey available

seasonally may obtain larger sizes and have faster growth rates than bass in similar river

systems without anadromous prey available. Pine et al. (2005) determined that flathead

catfish Pylodictus olivaris in coastal North Carolina Rivers selected juvenile American

shad and hickory shad over resident freshwater species. Other studies on the Atlantic

coast have shown that juvenile striped bass play an important role as prey for young

bluefish (Juanes et al. 1993).









Throughout an entire year of sampling I found only one juvenile American shad

in a predator diet, which was a largemouth bass collected in December. One hickory

shad was found in a largemouth bass diet in August. The effect of predation on a

particular species during a year is likely to reflect the relative abundances of predator and

prey and the food preferences of the predator (Juanes et al. 1993). I did not observe a

large pulse of Alosa spp. moving through during the fall, and I did not observe predators

switching over to this energetically beneficial prey. Thus, it appears that Alosa spp. were

relatively uncommon in the river and thus were uncommon as prey for predators.

Relative Prey Abundance and Occurrence in Predator Diets

There is a wide variety of prey species found in this area of the St. Johns River.

Not only are all of the common freshwater species found here but also many marine

species spend all or part of the year in this area. The presence of salt springs that drain

into the river cause salinity to rise upriver from Palatka to Lake George (Odum 1953),

making the area suitable for seasonal species. Some seasonal species in this study were

abundant all year and therefore classified as resident species (e.g., bay anchovy), whereas

others occurred only seasonally and were classified as seasonal species (e.g., Atlantic

croaker and Atlantic menhaden). This was also true for abundance of freshwater prey

species found in this study, as some were present in about the same abundance all year

whereas abundance of others spiked seasonally (e.g., threadfin shad and white catfish).

Juvenile Atlantic croaker spend the first months of life in low salinity to

freshwater areas (Murdy et al. 1997). In the fall both juvenile and mature Atlantic

croakers leave rivers and migrate to bays (Murdy et al. 1997). These life history traits

explain why abundance was highest February through May, but they were collected from

January through August and were composed primarily of small juveniles.









Threadfin shad spawn in the spring and often again in the fall (Jenkins and

Burkhead 1993). Threadfin shad were not collected in trawls from December through

February, partly because mesh was too large to collect age-0 fish and partly because

mature shad may have moved deeper and become unavailable to the surface trawl or were

avoiding the trawl. During July and August threadfin shad accounted for a vast majority

of fish caught in trawls and the second most abundant species year round.

Bay anchovy reproduce throughout the year when water temperatures exceed

about 120 C, which allows nearly year round spawning in Florida (Murdy et al. 1997).

Bay anchovy were collected in trawls in all months of this study. Only during April did

the percentage of total catch that anchovy comprised drop below 20%. Eight months out

of the year anchovy made up for more than 50% of each month's total trawl catch, and

they were by far the most collected species using the surface trawl gear.

Larval Atlantic menhaden move into freshwater areas where they metamorphose

into juveniles. These juveniles will remain in freshwater areas using them as nursery

areas until fall when they move out to the bays and ocean (Murdy et. al 1997). Juvenile

menhaden will congregate into dense schools as they leave these nursery areas from

August through November (Rogers and Van Den Avyle 1983). Menhaden were only

collected in trawls from September through November, presumably when they were

emigrating from the low salinity river to the sea.

Prey abundance and size strongly influence predator diets and growth. If the

abundance of a preferred prey increases, importance in the diets of predators increases

(Adams et al. 1982; Storck 1986; Hartman and Margraf 1992; Michaletz 1997). I found a

similar relationship between largemouth bass (predator) and Atlantic menhaden (prey).









Bay anchovy were the most abundant and available prey in the sample area

throughout the entire sampling period. At various times of the year high numbers of

anchovy were consumed with extremely high numbers consumed by small fish in spring

and summer. Bay anchovy were eaten by all size classes of largemouth bass in the winter

when other prey species were less prevalent and during the summer by small size

largemouth bass which may be because juvenile threadfin shad had exceeded their gape

limits although this was not measured.

Threadfin shad were found in diets during all 12 months of the year, although

highest numbers were found in diets from July through December (Figure 6). This

follows the same trend as was seen in trawl collections. Although no threadfin shad were

collected during December-February with trawls this does not necessarily mean their

abundance had dropped but is more than likely because they were no longer on the

surface and available to capture by a surface trawl, as they were still found in predator

diets. Threadfin shad were the most common occurrence in medium and large size bass

diets and second to bay anchovy in small size class fish. Although bay anchovy were

consumed in higher numbers by small size class largemouth bass, threadfin shad

accounted for most of the caloric intake. Medium and large size largemouth bass diets

revealed threadfin shad occurring most by number throughout the year and making up for

most of caloric intake. Numbers of threadfin shad found in diets peaked in late summer

and early fall.

Although common in trawl catches from February through August, Atlantic

croaker never appeared frequently in largemouth bass diets except for May when they

were the most common prey species by number and also accounted for most of the









caloric intake found in large size class fish diets. There was a small spike in numbers of

Atlantic croaker found in small and medium size class largemouth bass diets during the

summer. I did not detect that predators were switching to croaker as prey even during

times of year when they were relatively abundant in trawl samples.

Atlantic menhaden were collected in diet and trawl samples only during the months

of September through November. During these months I did find menhaden in the diets

of all size classes of bass. Also, menhaden were found in diets of striped bass and

longnose gar during this same time period. For large size class largemouth bass in

September, bay anchovy were the most common prey consumed by number, followed

closely by Atlantic menhaden, which made up for the highest caloric intake of any

month. Atlantic menhaden then accounted for most of the caloric intake for large size

class fish in November. This suggests that predators did switch to menhaden and took

advantage of their presence and high caloric content during the fall.

Caloric values used were measured from only adult samples of Atlantic croaker

(Thayer et al. 1973) which I rarely collected, so caloric estimates may vary. Atlantic

menhaden (Steimle Jr. and Terranova 1985), bay anchovy (Steimle Jr. and Terranova

1985), and threadfin shad (Strange and Pelton 1987) caloric estimates were made from

most commonly collected size fish. Bay anchovy and threadfin shad of all sizes were

sampled during this project so these estimates should be accurate while only juvenile

Atlantic menhaden were sampled meaning their caloric estimates are likely less accurate.

Throughout certain times of the year several prey items occurred regularly in diets

that were not collected in trawl samples. During April and May blue crabs Callinectes

sapidus made up more than 10 % of items found in all predator diets. Also from October









- December, crayfish Procambarus spp. and armored catfish (sailfin catfish and brown

hoplo) occurred frequently in diets. This switch to crayfish and armored catfish was

likely due to extremely high water after hurricanes. Predators at this time were often

collected in areas of newly flooded timber where these prey items were likely abundant.

Small fish often consumed non-fish prey, such as insects and various invertebrates.



Management Implications

Estimates of stock abundance are crucial to the assessment and effective

management of freshwater, anadromous, catadromous, and marine fish populations.

Massman et al. (1952) found surface trawls to be an effective technique to sample young

fishes in tidal rivers. Loesch et al. (1982) proved this technique even more effective on

juvenile Alosa spp. when used after dark as juveniles will be feeding on the surface at

this time.

I was fortunate enough to have data collected by Willams and Bruger (1972)

using the same size surface trawl pulled at night and in the same sampling area. My

catch rates for juvenile American shad in the trawl samples were similar to what they

found over 30 years ago. This could mean that recruit abundance is similar to what it was

back then.

However, in a river as big as the St. Johns, over 2 km wide in many areas, it may

be hard to effectively sample with a 3 m wide trawl net. If the net was pulled too fast a

pressure wave would build in front of it causing fish to avoid the net, while if pulled slow

enough to minimize the pressure wave, fish were seen jumping out of the trawl. Thus,

the trawl appeared to not be a highly effective gear for sampling the open water

planktivorous fishes, especially considering that electrofishing collected more than twice









as many juveniles in the daytime while looking for predators. Juveniles were in the area,

but my collections were not as successful as planned. If abundance was relatively high, I

would have seen higher numbers trawling, shocking, and in diets. Although the trawl

may not have effectively sampled all fishes in the water column, I was able to detect

trends between trawl catch and occurrence in predator diets for several species.

After Florida's net ban in 1995, Florida's American shad fishery was reduced to

recreational anglers fishing on the spawning grounds (McBride 2005). McBride (2000,

2005) found angler catch per unit effort (CPUE's) of American shad increased in 1995-

1996 and 1997-1998 but then decreased to the lowest levels of the creel study in 2004-

2005. Although this decrease in CPUE was observed, angler catch rates have remained

stable throughout the last 15 years, at about one fish per hour. This however, comes

along with a decrease in angler effort (McBride 2005). Current harvests of American

shad are well below historical levels, and although unlikely, even these extremely low

current exploitation rates could be excessive because population sizes of American shad

are thought to be low and historically depressed (Hightower et al. 1996; McBride 2005).

Methods for rebuilding of shad stocks include reduction of harvest of adults as

well as improvement and protection of spawning and nursery habitats (Walsh et al.

2005). Commercial landings in the St. Johns River have been at nearly zero since the

mid-1990s (McBride 2000). All offshore commercial fishing directed at American shad

was shut down in 2004. Even with this reduction it does not appear that stocks will

regain historical levels seen at the turn of the century 20th century (Hightower et al.

1996).









Because current fishing mortality is not likely to be causing the extremely low

population levels, other factors need to be considered. Walsh et al. (2005) determined

that in the Roanoke River flooded forest land would be beneficial to blueback herring and

alewife eggs and larvae. During my research I did not see extremely high river levels

until after the hurricanes in the fall of 2004. I was unable to detect how factors from

these storms affected survival and abundance of juvenile American shad during 2004.

Crecco et al. (1986) concluded that major climatic events can overshadow the

compensatory mechanism of American shad populations. They also noted that turbulent

June flow rates promote unfavorable feeding conditions for larvae, eventually reducing

survival. In the St. John's River, high flow rates cause the river to have high color from

tannic acids, potentially limiting primary and secondary production. These extremely

high flow rates seen in the fall of my sampling period may have caused similar problems

for juveniles and contributed to low catch rates/abundance.

Usually, increasing fall river flow would trigger juveniles to emigrate (O'Leary

and Kynard 1986). However, I saw no such trends, suggesting that temperature or

something such as photoperiod may be more important factors to trigger emigration than

flow. O'Leary and Kynard (1986) also suggested that when juveniles reach a certain size

they will leave the river regardless of environmental conditions, which I did not see.

Increased industry, residential growth, and organic pollution have all caused

decreases in the quality of habitat available to American shad (Limburg et al. 2003;

McBride 2005). Williams and Bruger (1972) found some of the major problems the St.

Johns River American shad population faced was changes in flow due to an extensive

series of water control structures, degradation of spawning areas, and increased industrial









and domestic effluents. Spawning grounds have remained the same for over 50 years, so

it is crucial that areas be protected from future degradation (McBride 2005). Although

only one dam was constructed on the main river channel, these problems still affect the

St. Johns River today. (McBride 2005).

One natural problem facing this shad stock is that they are at the southernmost

extent of their native range which may cause temporal and ecological problems.

Commercial fishing is non-existent now, however its effects are still present. McBride

(2005) found female shad considerably less abundant than males throughout the

spawning season. This may be due to the commercial fishers who targeted the female

shad for roe. Although the commercial American shad fishery has been closed, non-

target mortality from other fisheries may be significant (McBride 2005) The recreational

fishery has also drastically declined so only time will tell if the American shad population

in this river rebounds (McBride 2000; McBride 2005). Crecco et al. (1986) showed that

American shad can produce large year classes from few adults, but I did not find

evidence of abundant juvenile American shad in this system.

In the future we need to attempt to get better population estimates for spawning

fish and the resulting juvenile recruits. I believe we need to look more closely at

available spawning habitat and also into the larval life stage to examine if water quality is

having an effect on survivability or if there is lack of some critical food available to

juveniles in the upper St. John's River. Stocking could be a future option to help bring

the population back up to historical levels. All of these issues need to be looked into and

are crucial to successfully manage and maintain the St. Johns River American shad

population.









Prey availability and amount of energy consumed govern the proportion of

consumed energy allocated to the principal physiological functions (Adams et al. 1982).

In many freshwater systems threadfin and gizzard shad are the primary prey species.

These species undergo large temporal fluctuations in abundance due to temporal changes

(Adams et al. 1982; Storck 1986; and Michaletz 1997). This was not a problem in this

area as threadfin shad were found in diets throughout the entire year. Also, there was

usually more than one abundant prey species each month.

Prey availability drives the success of a fish population (Krohn et al. 1997; Yako

et al. 2000) and prey selectivity of predators is an important mechanism structuring

aquatic communities (Juanes et al. 2001). In coastal systems anadromous fishes may

provide a seasonal influx of high energy prey (Durbin et al. 1979). I did not find that

anadromous shads were important in predator diets due to their apparent low abundances,

however seasonally Atlantic croaker and Atlantic menhaden were important prey items.

High growth rates can increase the size of largemouth bass, in turn increasing the

range of prey they can ingest and their probability of survival during the winter through

spring period (Adams et al. 1982). The influx of nutrient-rich Atlantic menhaden in the

fall may help this cause, especially as all size classes of largemouth bass were actively

feeding on them when present.

Managers should focus on managing the river as a whole. This includes looking

into flow issues, water quality and pollution concerns, and habitat quality. This research

will help not only predator (e.g., largemouth bass) and American shad populations of this

area but also all fish and wildlife along this unique 500-km long river.















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BIOGRAPHICAL SKETCH

Nicholas Aaron Trippel was born January 5, 1981 in Goshen, Indiana, the son of

Donald and Debra Trippel. He graduated from Northwest Guilford High School, North

Carolina in 1999. He received his Bachelor of Science degree in Fisheries Management

from Auburn University. He became interested in the field of fisheries management

while completing a year long mentorship project with a district fisheries biologist during

his senior year of high school. While completing his Bachelor of Science degree he

gained fisheries management experience working on research projects while employed

with Alabama Fish and Wildlife Cooperative Research Unit. After completing his

undergraduate degree he received a job working as a full time lab technician at the

University of Florida for Dr. Mike Allen. After working as a lab technician for eight

months Dr. Mike Allen took him on as graduate research assistant to pursue a Master of

Science degree in fisheries management. After graduation in May 2006, he hopes to

pursue a successful career in fisheries management working as a state freshwater fisheries

biologist.




Full Text

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TEMPORAL TRENDS IN JUVENILE ALOSA SPP ABUNDANCE AND RELATION TO PREDATOR DIETS AT THE ST. JOHNS RIVER, FLORIDA By NICHOLAS A. TRIPPEL A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2006

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Copyright 2006 by Nicholas A. Trippel

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This document is dedicated to my family: Mom, Dad, and Aimee.

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iv ACKNOWLEDGMENTS This thesis could not have been completed without the hard work and dedication of many people. I thank Drew Dutterer, Mo Bennett, Christian Barrientos, Galen Kaufman, Mark Captain No Fun Rogers, Jason Dotson, Kristin Henry, Kristin Maki, Ginni Chandler, Steve Larsen, Travis Tuten, Porter Hall, Matt Catalano, Greg Binion, Vince Politano, Patrick Cooney, Kevin Johnson, Julie Harris, Adam Richardson, Vaughn Maciena, Eric Nagid, and Jay Holder for their help with field work, lab processing, finding literature, and coming along on the late-night trawling trips. I thank my supervisory committee members (M. Allen, D. Murie, and R. McBride) for the assistance and instruction they have given me throughout this study. I would like to thank my supervisory committee chair (M. Allen) for all the valuable knowledge he has shared with me, for encouraging me, for teaching me to work hard, for being such a knowledgeable mentor, and for being the only other tiger fan in the lab. I want to especially thank my parents (Don and Deb Trippel) for everything they have done for me throughout the years, believing in me, and motivating me to be successful.

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v TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................iv LIST OF TABLES.............................................................................................................vi LIST OF FIGURES.........................................................................................................viii ABSTRACT....................................................................................................................... ..x INTRODUCTION...............................................................................................................1 History of American Shad Fisheries.............................................................................1 American Shad Dist ribution and Biology.....................................................................3 METHODS........................................................................................................................ ..9 Site Selection................................................................................................................9 Data Collection...........................................................................................................10 Analyses......................................................................................................................1 3 RESULTS........................................................................................................................ ..19 Trawl Collections........................................................................................................19 Electrofishing Samples...............................................................................................21 DISCUSSION....................................................................................................................4 6 Juvenile Alosa spp. Abundance..................................................................................46 Relative Prey Abundance and Occu rrence in Predator Diets.....................................49 Management Implications..........................................................................................53 LIST OF REFERENCES...................................................................................................58 BIOGRAPHICAL SKETCH ............................................................................................64

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vi LIST OF TABLES Table page 1. Prey species classified as resident (collected in sample area year around) and seasonal (marine species collected seasonally in sample area) in the Palatka area of the St. Johns River, Florida from April 2004 thro ugh Marc h 2005. ...................33 2. Summary of monthly mean trawl and electrofishing catch rates (fish/hour) for juvenile Alosa spp. and total number of hours sampled using each gear by month in the Palatka area of the St. Johns Ri ver from April 2004 th rough Marc h 2005 ....35 3. Mean total lengths of juvenile American shad collected with trawl gear by month in the Palatka area of the St. Johns River, Florida, from September through December of 1969, 1970, and 2004. Number collected (N) and Standard Error (S.E.) are shown by month for 2004 data. Months with no fish collected and no data available are labeled as Na. Data from 1969 and 1970 were from Williams and Bruger (1972). ...............................................................................................37 4. Number of predators sampled with electrofishing and how many contained stomach contents (number and percent) in the Palatka area of the St. Johns River, Florida from Apr il 2004 through March 2005. ...........................................38 5. Presence (x) or absence of prey items in diets collected from largemouth bass Micropterus salmoides of the small size class (<365 mm) in the Palatka area of the St. Johns River, Florida fr om April 2004 thro ugh Marc h 2005. .......................39 6. Total numbers (N) of largemouth bass Micropterus salmoides from each of three size classes (small < 365 mm, medium 366-432 mm, and large > 432 mm) sampled each month containing stomach contents and the total number of prey items found in these stomachs in the Palatka area of the St. Johns River, Florida from April 2004 thro ugh Marc h 2005. ..................................................................42 7. Presence (x) or absence of prey items collected from predator species ( in order of number of diet samples collected from most on the left to least on the right) other than largemouth bass Micropterus salmoides in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005. .................................. 43

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vii 8. Total number, mean and median legnth (mm), and mean and median weight (g) of each of the four most common prey species: Atlantic croaker (ATCR), Atlantic menhaden (ATME), bay anchovy (BAAN), and threadfin shad (THSH), found in each of three size classes (small < 365 mm, medium 366-432 mm, and large > 432 mm) of largemouth bass, Micropterus salmoides, diets from April 2004 through March 2005 in the Palatka area of the St. Johns Ri ver, Flor ida........45

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viii LIST OF FIGURES Figure page 1. Map of Florida showing the St. Johns River in its entirety, along the Atlantic coast of Florida.....................................................................................................17 2. Map showing the Palatka area of the St. Johns River, Florida, including the location of the state road 100 bridge, which was the mid-point of the study sampling area. The sampling area was 5 km north and south of this bridge. This figure shows that the Palatka area is the last narrow stretch of river before widening and it remains this wide or wider until reaching the Atlantic Ocean.......18 3 Comparing mean monthly trawl catch rates (+ standard error) of juvenile American shad between 1970 and 2004 with 2004 mean monthly water temperatures for the Palatka area of the St. Johns River, Florida...........................25 4. Mean number of Atlantic croaker, Atlantic menhaden, bay anchovy, threadfin shad, and other fish species collected per five minute trawl with standard error bars throughout 12 months of sampling on the Palatka area of the St. Johns River Florida........................................................................................................26 5. Percent composition of each prey species from total trawl catches from January through December at the Palatka area of the St. Johns River, Florida. The figure shows only species that accounted for at least 5% of monthly trawl catches. .........27 6. Percent by number of Atlantic croaker, Atlantic Menhaden, bay anchovy, and threadfin shad found by month in diet samples (solid line) of all species of predators collected by electrofishing and in trawl samples (dashed line) in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005. .................................................................................................................... 28 7. Percent by number of prey items that were not Atlantic croaker, Atlantic menhaden, bay anchovy, or threadfin shad found by month in diet samples of small (<356 mm), medium (356 mm – 432 mm), and large (>432 mm) largemouth bass collected by electrofishing and in trawl samples in the Palatka area of the St. Johns River, Florida from April 2004 th rough Marc h 2005. ...........29 8. Percent frequency by number for crayfish, armored catfish (plated and sailfin catfish), and blue crab found by month in all diet samples of largemouth bass collected by electrofishing in the Palatka area of the St. Johns River, Florida from April 2004 thro ugh Marc h 2005. ..................................................................30

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ix 9. Mean LOGIT values for prey items (>0 means more seasonal diet items, <0 means more resident diet items) by month found in stomach contents of three size classes of largemouth bass (small <356 mm, medium 356 mm-432 mm and large >432 mm TL) and for trawl catches in the Palatka area of the St. Johns River, Florida from Apr il 2004 through March 2005. ...........................................31 10. Total percent caloric content (left panels, caloric values estimated only for four species of prey fish therefore does not always represent total caloric intake) and percent by number (right panels) for Atlantic croaker, Atlantic menhaden, bay anchovy, and threadfin shad found in three size classes of largemouth bass diets,(small < 356 mm, medium 365 mm-432 mm, and large > 432 mm) collected by electrofishing in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005. Note : Only parts of croaker were found in large size class largemouth bass. These were counted in total prey counts, however parts were too small to estimate total weight and size and therefore were left out of caloric estimates..........................................................................32

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x Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science TEMPORAL TRENDS IN JUVENILE ALOSA SPP ABUNDANCE AND RELATION TO PREDATOR DIETS AT THE ST. JOHNS RIVER, FLORIDA By Nicholas A. Trippel May 2006 Chair: Micheal S. Allen Major Department: Fisheries and Aquatic Sciences The St. Johns River, Florida, was once the largest recreational American shad Alosa sapidissima fishery on the Atlantic coast. This fishery has drastically declined due to decreased abundance of American shad. I assessed the temporal trends in juvenile American shad relative abundance leaving the river, compared catch rates to those of a similar study completed 35 years ago, and evaluated diets of piscivorous fish in the sample area before, during, and after the juvenile American Shad had moved through the area. I also compared predator diets to prey availability over 2004-2005 and estimated caloric values of more-common prey to determine the seasonal variation in the relative importance of these species to predator diets within the Palatka area of the St. Johns River. Trawl catch rates of juvenile American shad and other juvenile Alosa spp. were extremely low. Only 23 American shad were collected during 12 months of sampling. Highest catch rates occurred in October, which was similar to historic catch rates 35

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xi years ago using similar trawling gear. Only one American shad and one hickory shad A. mediocris were found in predator diets in 12 months of sampling and 1,747 total predator diets measured. The four most common species collected in trawl and diet samples were threadfin shad Dorosoma petenense bay anchovy Anchoa mitchilli Atlantic croaker Micropogon undulatus and Atlantic menhaden. The number of these species found in largemouth bass Micropterus salmoides (the most common predator) diets varied significantly by month and size class of largemouth bass. Atlantic menhaden were found to be the most energetically beneficial to predators, and I found that during months when they were present all size classes of largemouth bass used them as prey. Correlation analysis revealed that trawl catches and occurrence of individuals found in diet samples were positively correlated for several species ( = 0.01), including Atlantic menhaden, ( P <0.01), sailfin catfish Pterygoplichthys multiradiatus (P < 0.01), and white catfish Ameiurus catus ( P = 0.01). Management implications of this study include helping to successfully manage the American shad fishery in this river, and to better relate the life history of common prey items in this coastal river system to seasonal and ontogenetic diet shifts for the common predators. I identified low abundance of juvenile Alosa spp. These juveniles must deal with the predatory gauntlet, but were not seemingly preyed upon disproportionately to their own abundance. Researchers need to look into flow and habitat issues related to spawning success of adult American shad, water quality, and pollution issues to see if these may be reasons we saw such low juvenile abundance.

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1 INTRODUCTION Five species of anadromous shad belonging to the genus Alosa are native to Florida. Three of these species (American shad Alosa sapidissima hickory shad A mediocris and blueback herring A aestivalis) are found within the St. Johns River system. The other two (Alabama shad A alabamae and the skipjack herring A chrysochloris) are found on the Gulf of Mexico coast of northwest Florida (McBride 2000). Clupeids are among the most economically important fishes, and worldwide no other family of fishes is consumed or harvested in larger quantities (Scharpf 2003). FloridaÂ’s St. Johns River was once estimated to support the largest recreational fishery for anadromous shad species on the Atlantic coast (Walburg and Nichols 1967; McBride 2005). Thus, research investigating the ecology of anadromous shads in Florida is important for their conservation and management. History of American Shad Fisheries Until recent years, American shad supported some of the most important commercial fishing industries in the United States (Williams and Bruger 1972; McBride 2000; McPhee 2002; Limburg et al. 2003; Scharpf 2003; McBride 2005). Although Atlantic salmon Salmo salmar were first targeted by early Americans, American shad soon replaced them and was favored for its flavorful roe and meat (McPhee 2002). In the United States during the 1800s, many fisheries (along with the shad fishery) grew rapidly (Walburg and Nichols 1967; McBride 2000). American shad were

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2 harvested using multiple methods including fish dams, shad floats, fyke nets, seines, pound nets, and gill nets (Scharpf 2003). More-successful gear types overexploited American shad populations for over a century now (Limburg et al. 2003; Scharpf 2003; McBride 2005). Some American shad stocks in the New England states were reportedly over-exploited as far back as 1830 (Gerstell 1998). In both 1889 and 1890, the shad harvest in Florida alone was over 2 million pounds (McBride 2000; McBride 2005). In 1896, over 45 million pounds of American shad were harvested in the United States (Stevenson 1899). Between 1904 and 1930, U.S. shad fisheries along the Northeastern Atlantic coastline collapsed (Walburg and Nichols 1967) because of overfishing, water pollution, and construction of dams (Leggett and Whitney 1972). As catch rates declined, commercial fishers looked for new ways to harvest shad, eventually developing the ocean-intercept fishery in the last 20 years. Even though shad populations in rivers had declined, most commercial fishers now harvested shad in offshore areas where all shad populations were congregated together, leading to a shortterm increase in landings (ASMFC 1999). Effectively, this meant that fishers were simultaneously harvesting American shad populations from all of the rivers along the Atlantic coast (McBride 2005). Commercial harvest of American shad was closed in Maryland in 1982, and Virginia in 1994. As stocks became severely depleted, harvest was also prohibited in Maine, New Hampshire, Massachusetts, and Rhode Island (Scharpf 2003). Inshore netting regulations imposed in the 1990s led to virtual closure of FloridaÂ’s commercial shad fishery (McBride 2005). To regulate the ocean-intercept fishery, the Atlantic States Marine Fisheries Commission (ASMFC) mandated a 40% reduction in effort for this fishery by the end of 2002 (ASMFC 1999). From1995 to

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3 2003, harvests declined, average annual U.S. domestic landings dropped to 2.7 million pounds (NOAA 2005); compared to 50 million pounds in the 1890s, and 9 million pounds the 1950s and 1960s (NOAA 2005). Because of these severe catch declines, and concerns about the shad stock, all U.S. Atlantic shad ocean-intercept fisheries were closed at the end of 2004 (ASMFC 1999; McBride 2005) thereby directing any remaining targeted effort to within river systems. American Shad Distribution and Biology American shad are native to the Atlantic coast of North America, and are the largest species of the family Clupeidae, reaching a maximum size of 5.5 kg, length of 76 cm, and age of 11 years (Froese and Pauly 2005). The native range of American shad is the U.S. Atlantic coast from the St. Lawrence River (on the border of the United States and Canada) southward to the St. Johns River in Florida (Brown et al. 1999; Limburg et al. 2003; McBride 2005). American shad are river-specific, with each river having a distinct spawning stock (ASMFC 1999; McBride 2005). American shad on the southern end of their range are semelparous, spawning once and dying; whereas fish at higher latitudes are iteroparous, spawning repeatedly (Limburg 1996). All populations south of the Neuse River, North Carolina, including the St. Johns River, are semelparous (Limburg et al. 2003; McBride 2005). Life history characteristics of these separate stocks vary from north to south, depending on the environment of their home river (Limburg 1996). Characteristics that vary include fecundity, size, and age at maturity (Leggett and Carscadden 1978; Limburg et al. 2003). Fecundity is inversely related to latitude and semelparity. In general, American shad at lower latitudes produce 3-5 times more eggs per kilogram of body weight then their northern counterparts (Limburg et al. 2003). American shad at lower

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4 latitudes are also smaller, and mature at younger ages than their northern counterparts (Limburg et al. 2003). Talbot and Sykes (1958) used tagging studies to learn that American shad from Florida to Maine congregate together during summer and fall seasons in the Gulf of Maine, whereas populations from Canada spend their summers in the St. Lawrence estuary (Limburg et al. 2003). As water temperatures drop during winter, American shad move south, with northern populations congregating on the Scotian Shelf; and southern populations (those from Florida to Maine) congregating in the Middle Atlantic Bight, off the coast of Florida (Limburg et al. 2003). American shad remain offshore in these congregations until reaching maturity, then return to their natal streams to reproduce and start the cycle again (Limburg et al. 2003). Differences in spawning traits are partly due to the energy reserve needed for spawning migration. Southern stocks migrate thousands of kilometers farther than northern stocks, making migration energetically expensive. Nearly 50% of the Florida American shadÂ’s total somatic energy reserves are used just to make it to their spawning grounds (Leggett and Carscadden 1978; Glebe and Leggett 1981a; Glebe and Leggett 1981b). Another 20 to 30% of their somatic reserve is then expended during the spawning process. Thus, southern American shad do not have the energy reserve needed to return to sea as the northern populations do, resulting in semelparity (Leggett and Carscadden 1978). Although southern stocks of American shad die during the spawning season they return to the river, they may spawn multiple times during that spawning season (Olney and McBride 2003; McBride 2005).

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5 Shad migrations to spawning grounds correlate with water temperature (Leggett and Whitney 1972). Timing of shad arrival at their spawning grounds along the Atlantic coast varies with latitude but is similar in temperature (Leggett and Whitney 1972; McBride 2000), which accounts for the earlier spawning in the south than in the north. American shad begin to reach their spawning grounds as early as November in Florida, and this run may continue through May (Leggett and Whitney 1972; Davis 1980; McBride 2005). American shad near Canada begin spawning in May and continue through July (Limburg et al. 2003). American shad are broadcast spawners, with communal spawning lasting up to 10 weeks (Scharpf 2003). Release of the demersal, free-drifting eggs (Williams and Bruger 1972), occurs at depths up to 10 meters, usually over sand or gravel substrate (Walburg and Nichols 1967). Limburg et al. (2003) found the following conditions conducive to larvae survival: water temperature above 20C, pH above 7, salinity level at least 10 ppt, and minimum zooplankton levels of 50 organisms/L. Larvae drift downstream, maturing into juveniles (Scharpf 2003). During their first summer, juvenile American shad remain in a nursery area of the river where they were spawned (Leggett and Whitney 1972). This nursery area is in the natal stream; downstream from the spawning grounds where the river is tidally influenced but contains relatively low salinity. Movement of juveniles from the spawning area to the nursery area is triggered by increasing water temperatures and current (Williams and Bruger 1972; McBride 2005). Movement from the nursery area to the ocean is triggered by decreasing water temperatures. After juvenile American shad emigrate to sea, they spend anywhere from three to six years migrating up and down the Atlantic coast until

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6 eventually returning to their natal river (Limburg et al. 2003; Scharpf 2003). American shad emigrate from rivers earlier in the north and may occur as late as December in Florida (Williams and Bruger 1972). Anadromous fishes such as American shad may play an important role as seasonal prey item for predatory fish in coastal river systems. Not only do the spawning fish stimulate primary production (Durbin et al. 1979), but migrating juveniles may provide cyclic influxes of prey. These influxes may promote higher growth and survival of predators (Yako et al. 2000). Juvenile anadromous fishes are faced with the survival challenge of having to pass through a gauntlet of relatively stationary predators (Petersen and DeAngelis 2000). Therefore, predators may have a large impact on the number of emigrating juvenile shad in coastal river systems. Few studies have assessed effects of piscivores on juvenile prey fish in coastal river systems (Buckel et al. 1999). In many systems, predation may be the main factor determining community structure (Raborn et al. 2003). Studies have been conducted in salt ponds or enclosures (Wright et al. 1993), but few have measured the impacts in natural systems. Diets of juvenile bluefish Pomatomus saltatrix consisted largely of anadromous fishes (Buckel et al. 1999). Buckel et al. (1999) demonstrated that mortality of juvenile striped bass Morone saxatilis was most strongly influenced by bluefish predation. Many studies have evaluated effects of predator mortality on the anadromous salmonids of the United StatesÂ’ Pacific coast. Predation plays an important role in juvenile mortality rates for anadromous salmonids (Ricker 1941). Parker (1968) found that up to 85% of a year class of juvenile salmonids may be consumed by predators on

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7 their way to the Pacific. Beamesderfer et al. (1990) determined that at John Day Reservoir, Columbia River, British Columbia, northern squawfish Ptychocheilus oregonensis consumed 1.7 million salmonids annually. At the same reservoir, Rieman et al. (1991) found that the number of deaths from all predators totaled 2.7 million juvenile salmonids annually. The high predation rates seen for juvenile salmonids in this reservoir may be high due a large population of predators and the amount of time juveniles remain in the reservoir before moving down river (Beamesderfer et al. 1990). Predation during the seaward migration may greatly reduce each yearÂ’s cohort, but the seasonal influx of prey is likely to be highly beneficial to predators (Durbin et al. 1979). Anadromous fishes may encourage higher survival and growth rates for predators (Yako et al. 1999). Yako et al. (1999) found that largemouth bass Micropterus salmoides in headwater lakes with seasonally available juvenile anadromous river herring, including alewife Alosa pseudoharengus and blueback herring, may grow faster and attain larger sizes than in lakes where these species were absent. Seasonally available herrings were the most consumed fish species for both lakes examined (Yako et al. 1999). Largemouth bass generally did not begin feeding on river herrings until August, probably due to ontogenetic changes in the foraging behavior of young bass. Using bioenergetics models, Yako et al. (1999) found that a diet of herring increased growth rates, and the presence of trophy-size largemouth bass was positively correlated with the seasonal availability of anadromous herrings in coastal Massachusetts. However, no studies have examined the migration of juvenile American shad in southern populations and the impacts predators have on their abundance and survival.

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8 I evaluated the summer migration of juvenile American shad during their downstream migration to sea, at the St. Johns River, Florida. Objective 1: estimate the relative abundance and timing of juvenile shad migrating back to the Atlantic Ocean, Objective 2: compare historical trawl catch rates of juvenile American shad from 19691970 (Williams and Bruger 1972) to my results, Objective 3: examine and compare differences between predator diets and prey availability over one year, and Objective 4: examine predator diets before, during, and after American shad migration to assess potential effects of predation on migrating juveniles. Effects of predation were measured in terms of frequency of predator events. Estimated caloric values of more common prey were also used to determine seasonal variation in the relative importance of these species to predator diets within the Palatka area of the St. Johns River.

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9 METHODS Site Selection The St. Johns River flows from south to north along FloridaÂ’s Atlantic coast, eventually entering the Atlantic Ocean near Jacksonville (Figure 1). It is the longest river in Florida at about 500 km and has a watershed area of over 20,000 km2 (McBride 2005). This tannic-stained river is very slow flowing, having a slope averaging only 2 cm/km with a total gradient of less than 10 m (McBride 2005). It starts off as a small meandering channel near Melbourne, then forms a series of lakes, and eventually reaches a width of over three kilometers and spills into the Atlantic near Jacksonville (Figure 1). I selected the Palatka area of the St. Johns River (Figure 2) as my study site because it was described by Williams and Bruger (1972) as a summer nursery area used by juvenile Alosa spp. before they emigrate to sea in the fall. The Palatka area of the river is approximately 127 river km from the Atlantic Ocean (McBride 2005). It is tidally influenced; however the salt wedge does not typically extend as far upriver as Palatka. The State Road 100 bridge crosses the river in this area and was the midpoint of my sampling area, with the sampling stretch extending 5 km north and 5 km south of this bridge (Figure 2). This sample area was selected because of the overlap with Williams and Bruger (1972) and because the area has a natural constriction of the river channel (Figure 2), which could concentrate anadromous fishes and improve catches compared to wider sections of the river.

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10 Data Collection I sampled the Palatka stretch of river using trawl gear to sample juvenile Alosa spp. and prey abundance from April of 2004 through March of 2005. Prey abundance and juvenile Alosa spp. populations were sampled using trawling techniques modified from Williams and Bruger (1972). A surface trawl (3.7 m wide mouth, 4.6 m long body, 25.4 mm mesh body, 19.1 mm mesh bag, and 12.7 mm mesh liner) of the same dimensions as used by Williams and Bruger (1972), was pulled between two jon boats (Williams and Bruger used one boat) using 30.5 m warp lines and otter boards modified with hydrofoils to keep the net on the surface (Massman et al. 1952, Trent 1967; Loesch et al. 1982). Using a surface trawl has been proven to be an effective technique for sampling clupeids offshore (i.e., open water) (Massman et al. 1952). Sampling effort was partitioned by season and on either side of the bridge. During months when Alosa spp. were expected, July through December, trawl samples were conducted biweekly. Throughout the rest of the year, January through June, trawl samples were conducted monthly. Williams and Bruger (1972) pulled trawls for 15 minute intervals after dark. For this study, trawls were pulled for 5 minutes after dark as per Massman et al. (1952), as Loesch et al. (1982) reported increased capture of American shad with surface trawls at night. Twelve trawls were pulled each sampling trip. The areas north and south of the state road 100 bridge were each divided into six sections from east to west across the river. Each section was sampled once in random order on each sampling trip for a total of twelve trawls per trip. Trawls were pulled at an average speed of 1.3 m/sec with trawl direction varied randomly and alternating against and with the current. Trawling with the current was possible as current was often minimal to non existent.

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11 All fish from each trawl were identified to species and counted. I brought all Alosa spp. back to the lab where they were weighed, measured, and otoliths were pulled and mounted on glass slides using Thermo Shandon synthetic mountant cement. Abundances of shad were indexed using the trawl catch rates (fish/minute), which was the same index used by Williams and Bruger (1972). I kept several specimens of all prey species in the area for reference and to construct an otolith key for identification of stomach content analysis of predators (Whitfield and Blaber 1978). I recorded total lengths to the nearest millimeter from all American shad collected both trawling and electrofishing (described below). For the months of September through December I compared these to mean lengths of American shad collected by Williams and Bruger (1972) from 1969 and 1970. Because Williams and Bruger recorded fork lengths, I measured fork length and total length on fish from 2005 and used linear regression to estimate total lengths for fish from Williams and BrugerÂ’s (1972) study. Water temperature was monitored in the sample area throughout the study period using Onset Computer Corporation Hobo model temperature loggers. The temperature logger was located at the state road 100 bridge approximately 1 meter below the surface. Water temperature was recorded every six hours throughout the year to the closest 1o C. I used a 5.5 m aluminum electrofishing boat, model SR-18 with a GPP 9.0 electrofisher and 16 horsepower, 9,000 watt generator to collect only fish which were potential predators. Juvenile Alosa spp. were also targeted for collection during electrofishing samples, however no other prey species were collected. Predators were sampled in the same weeks that trawls were pulled. I attempted to collect at least 50

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12 predators of all species combined with stomach contents each sampling trip. Electrofishing was conducted evenly throughout the shorelines of the sampled river stretch. Pilings at bridge and powerline crossings were also sampled while electrofishing to collect pelagic predators. I recorded total length (mm) and weight (g) of all predators collected. I used transparent acrylic tubes to obtain stomach contents of live predatory fish as described by Van Den Avyle and Roussel (1980). Diets were removed and immediately placed on ice and returned to the lab for examination. Predators were immediately released alive after diet samples were collected. Predators that could not have contents removed via the tube were placed on ice and returned to the lab for diet analyses. Before analysis, diets were thawed and blotted dry. Organisms in the stomach were identified using a microscope, dichotomous key, and otolith key, and then measured and counted. I recorded total diet weight, individual diet item lengths and weights, prey species, and body part if only partial prey items were found following the guide of Storck (1986). Prey species were also classified as either resident (freshwater and estuarine species found in the sample area of the river year round) or seasonal (usually marine species and found in sample area only seasonally) (Table 1). I retained 50 individuals of common prey species from the trawl sampling for energetic information and diet identification keys, including Atlantic croaker Micropogonias undulatus, Atlantic menhaden Brevoortia tyrannus, bay anchovy, Anchoa mitchilli and threadfin shad Dorosoma petenense For these fish I measured total length, standard length, vertebral column length, body depth, and length of the head. Predictive regressions were determined based on fish size to then estimate total lengths

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13 and weights for partial fish of these species found in diet samples. Total caloric content rates for each of these four species were based on previous studies. I estimated caloric content of prey found in diet samples using dry weight caloric values of: 4,638 cal/g and 76.3% moisture for Atlantic croaker (Thayer et al. 1973), 5,115 cal/g and 65% moisture for Atlantic menhaden (Steimle Jr. and Terranova 1985), 5,067 cal/g and 72% moisture for bay anchovy (Steimle Jr. and Terranova 1985), and 4,835 cal/g and 79.2%for threadfin shad (Strange and Pelton 1987). These caloric values were then multiplied by the total measured weight of prey or total estimated weight for partial prey items to determine total calories predators consumed of each prey type. These caloric values were calculated at various locations and various times of the year; however, I applied them as general estimates. Atlantic croaker caloric rates were determined from samples of adult fish collected in estuaries near Beafort, North Carolina (Thayer et al. 1973). Atlantic menhaden and bay anchovy caloric rates were determined from the most commonly collected sizes of these species along the Atlantic coast from Nova Scotia to North Carolina (Steimle Jr. and Terranova 1985). Threadfin shad caloric rate estimates were from fish collected throughout the year in two Tennessee reservoirs (Strange and Pelton 1987). These values were determined for bay anchovy and threadfin shad of similar size to those I collected, while most Atlantic croaker and Atlantic menhaden I collected were juveniles, thus smaller than those used to determine caloric values. Analyses Raw data from Williams and BrugerÂ’s trawl catch rates were archived and made available to me by Julie Harris of Florida Fish And Wildlife Research Institute. Trawl catch rates for American shad for this project and those of Williams and Bruger (1972)

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14 were converted to fish caught per hour to allow a qualitative comparison of catch rates between time periods. Total trawl catch rates were also summarized. Percent composition of fish species in the trawl and diet samples were estimated for each month. A cumulative list of all species found in diets and trawls was compiled from which presence or absence was recorded for each trawl and diet. I determined the percent each species made up by month of the total trawl catch and total diet samples. Statistical Analysis Software (SAS 2000) was used to run statistical analyses. I used correlation analysis to assess relationships between the percent each prey species made up of the monthly trawl catches and the percent that those same species made up of the total monthly diet samples. Correlations were done for each prey type across months. Largemouth bass was by far the most common predator in this study, and thus allowed much more detailed diet analyses. I evaluated whether diet contents varied with largemouth bass size and season for the most common prey species. Four prey species represented 95% of all prey collected in trawls and they were used to examine largemouth bass diets in greater detail: Atlantic croaker, Atlantic menhaden, bay anchovy, and threadfin shad. I compared the frequency of occurrence in the diets monthly and also the total estimated caloric values that each species comprised for that month’s diet samples. Largemouth bass were divided into three size classes: small (<356mm), medium (356mm-432mm) and large (>432mm), and energetic comparisons of total kilocalories consumed from each of the four prey species were made among the three largemouth bass size classes throughout the year I used the four common prey species and collectively grouped all other prey items into a category defined as “other”. I

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15 used a log transformation plus 0.5 to improve normality of the variance around each mean. Two-way analyses of variance (ANOVA, Procedure GLM, SAS 2000) ANOVA’s assessed how percent frequency by number and caloric value of each species found in diets (dependent variables) varied by month (12 months sampled), largemouth bass size class (3 size classes) and the interaction between these factors. Separate ANOVA’s were used for each prey species and the “other” category. When a two-way ANOVA was significant, Least Squared Means test (LS Means, SAS 2000) was performed to determine where significant differences occurred. Differences were declared significant at P 0.05 for all analyses, and this value was selected to reduce the chance of Type II error. To analyze the importance of seasonal versus resident prey for predators, prey fishes were categorized as resident or seasonal. Resident fishes were prey species collected throughout the entire year in the sample area, and prey species classified as seasonal were the marine species found only seasonally in the sample area. I estimated a ratio defined as the LOGIT for each individual predator diet collected as: Log (number of seasonal prey items + 0.5/number of resident prey items + 0.5).

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16 Thus, if this value was positive there were more seasonal items in that particular diet, whereas if it were negative there were more resident items in that diet. A two-way ANOVA (Procedure GLM SAS 2000) was used to assess differences in LOGIT values (dependent variable) by month (12 months sampled), largemouth bass size class (3 size classes) and the interaction between these two factors. A least squares means test (LS Means) was used to separate the means if the ANOVA was significant (SAS 2000). This analysis was used to assess whether largemouth bass diet contents shifted when seasonal prey were available. I also assessed how trawl catches of seasonal versus resident prey fish varied through the year. A repeated measures ANOVA (Procedure Mixed SAS 2000) was used to test if the total number of individuals (log n) collected in trawl samples (dependent variable) varied by month, prey species class (resident or seasonal), and the interaction between these two independent variables. I used the autoregressive order 1 covariance matrix structure, which models correlations decreasing with distance in time (Littell et. al. 1996). Each trawl repetition was nested randomly within site (six transverse river sampling as sections described above). If significant, the LS Means test was used to test for differences among means.

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17 Figure 1. Map of Florida showing the St. Johns River in its entirety, along the Atlantic coast of Florida.

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18 Figure 2. Map showing the Palatka area of the St. Johns River, Florida, including the location of the state road 100 bridge, which was the mid-point of the study sampling area. The sampling area was 5 km north and south of this bridge. This figure shows that the Palatka area is the last narrow stretch of river before widening and it remains this wide or wider until reaching the Atlantic Ocean.

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19 RESULTS Trawl Collections I collected a total of 27 fish species in the trawls representing 13 families (Table 1). Most species were classified as resident because they were collected in the sample area throughout all 12 months of the year, but six species were classified as seasonal based on the fact that they were only collected seasonally in this stretch of the St. Johns River, Florida. Table 1 also includes prey items found in diets and not collected in trawls. I collected all three Alosa spp while trawling. Blueback herring occurred earliest in the year and accounted for a majority (N = 37) of the Alosa spp collected (Table 2). American shad ranked second in total catch with 23. I collected a total of six hickory shad between the months of May and October (Table 2). Trawl catch rates were highest in May for blueback herring whereas catch rates for both American shad and hickory shad were highest in October (Table 2). Throughout the 12-month sampling period a total of 23 juvenile American shad were collected using the trawl (Figure 3). American shad were collected beginning in June with the last catches occurring in December. Most juvenile American shad were collected from September through November with catch rates being the highest in October, when nine American shad were collected. Trawl catch rates and temporal trends of juvenile American shad occurrence were similar to those of Williams and Bruger in 1970 (Figure 3). It appeared that decreasing water temperatures triggered

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20 juvenile American shad to emigrate (Figure 3). However, I never detected large pulses of juvenile shad moving through the area in the trawl catches. Mean total length of American shad was similar between this study and Williams and Bruger (1972) for all sample months (Table 3). Juvenile seasonal species were commonly collected in the sample area during spring and fall of the sample period. Mixed model analyses determined that mean catches of resident and seasonal prey species in the trawl samples varied throughout the year, with significant interaction between month and species classification (resident or seasonal) ( P <0.01). This meant that both the number of total prey fish caught and prey species (resident or seasonal) caught varied by month. For example, seasonal species had lower occurrence in summer than in spring or fall, whereas resident species catch was highest in summer and winter. Monthly patterns in abundance of the most common resident and seasonal species were therefore further examined using ANOVA and LS MEANS tests. The four most abundant species collected with the trawl were threadfin shad, bay anchovy, Atlantic menhaden, and Atlantic croaker. I collected numerous other species infrequently (Figure 4). Threadfin shad and bay anchovy appeared in trawl samples consistently throughout the year. Atlantic croaker appeared in all 12 months but catch rates were much higher during February – June (Figure 4). I sampled only a few mature Atlantic croaker and most were small juveniles (<50 mm TL). Atlantic menhaden appeared in trawl samples only during the months of September, October, and November. During January and November, white catfish Ameiurus catus accounted for 25% and 9% respectively of total trawl catch (Figure 5).

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21 Electrofishing Samples Some species of prey targeted by trawling were also captured with electrofishing gear. While electrofishing for predators I collected 55 juvenile American shad, 194 juvenile blueback herring, and 4 juvenile hickory shad. More juvenile Alosa spp. were collected while electrofishing for predators than while targeting them using the trawl, but catch rate via electrofishing was highest later in the fall than the trawl catches of Alosa spp. (Table 2). Predator species sampled by electrofishing included: largemouth bass, striped bass Morone saxatilis black crappie, Florida gar Lepisosteus platyrhincus longnose gar Lepisosteus osseus mangrove snapper Lutjanus griseus red drum Sciaenops ocellatus ladyfish Elops saurus channel catfish Ictalurus punctatus white catfish, brown bullhead A. nebulosus bowfin Amia calva southern flounder Paralichthys lethostigma and chain pickerel Esox niger I collected a total of 1,747 predators of which 714 contained stomach contents, and largemouth bass was by far the most common predator collected (Table 4). Prey items collected from potential predators by tubing were often highly digested and difficult to identify. These items were given the classification unidentifiable (UID). The percent of UID stomach items varied by month and comprised anywhere from 1145% of total monthly stomach contents by number. Throughout the entire year only one juvenile American shad and one juvenile Hickory shad were found in predator diets, both from largemouth bass diets. Prey items that comprised more than five percent by number of all diets for each monthÂ’s total diet samples included: threadfin shad, bay anchovy, Atlantic croaker, Atlantic menhaden, bluegill Lepomis macrochirus blue crabs Callinectes sapidus clown goby Microgobius gulosus white catfish, plated catfish

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22 Hoplosternum littorale sailfin catfish Pterygoplichthys multiradiatus and unidentified crayfish species. The most common species collected in trawls compared to those found in diet samples by month are shown in Figure 6. This showed that as prey availability increased, occurrence in diets also increased. Figure 7 shows the percent frequency by number of diet items from anything other than the four most commonly collected species in trawl samples, collectively grouped as “other”. Large size class largemouth bass commonly preyed upon Lepomis spp. which were grouped as “other” (Table 5). The number of “other” species peaked in fall when we experienced extremely high water. During this time, the most common prey items included crayfish species, plated catfish, and sailfin catfish (Figure 8). Trawl catches and occurrence of individuals found in predator diet samples were significantly correlated for several species. These species included Atlantic menhaden ( P <0.0001), sailfin catfish ( P < 0.0001), and white catfish ( P = 0.0009) (Figures 4, 6, and 7). For all other species (n = 38) the Proc CORR procedure revealed no significant correlations. Diet contents for largemouth bass varied with fish size and across months. There were 114 largemouth bass in the large size class, 205 in the medium size class, and 267 in the small size class (Table 6). The GLM procedure revealed that LOGIT values for diets varied significantly for the interaction between month and size class ( P = 0.0003). Throughout the year different size classes of largemouth bass were feeding on different prey species, and that also throughout the year these prey varied from seasonal to resident. During the winter all size classes of largemouth bass were preying upon bay anchovy when other prey species were less prevalent. Mean LOGIT values by month

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23 revealed that trawl and diet samples collected contained higher proportions of resident species than seasonal species, except for large and medium size class bass diets in April and large size class bass diets in June (Figure 9). This was due to high numbers of threadfin shad and bay anchovy collected in trawls and largemouth bass diets. Exceptions included the month of April when medium and large size class fish commonly preyed upon blue crabs and June when Atlantic croakers were found in diets of large size class largemouth bass. After July all LOGIT values were negative as threadfin shad became the most common prey item through summer and fall. Throughout fall and winter LOGIT values remained negative as all size classes of fish preyed upon armored catfish, crayfish, and threadfin shad. Trawl catches had negative LOGIT values throughout the entire year of sampling compared to diets, indicating that diet contents of all predators contained higher seasonal prey composition than the trawl samples. Next to largemouth bass, black crappie was the most abundant predator collected with 56 containing stomach contents (Table 7). The most common prey items found in black crappie diets were bay anchovy (20%), also threadfin shad and insects each accounted for 9% of the total prey items by number. Bay anchovy (35%) and threadfin shad (18%) were the most common items found in the diets of 16 striped bass. Thirteen longnose gar sampled contained prey items with threadfin and gizzard shad were the most common prey items accounting for 56% of total prey items by number. Atlantic croaker made up for another 19% of prey items found in the longnose gar diets. Channel catfish consumed various Lepomis spp., crayfish, white catfish, and even pork chops.

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24 Using published caloric values for the four most common prey species, I estimated from diets the monthly caloric intake that each species comprised for each largemouth bass size class by month. Numbers and sizes of prey consumed by each size class of largemouth bass are shown in Table 8. I also compared this to the number of individuals of each species found in diets by month. Bay anchovy and threadfin shad were often the most common prey item by number while threadfin shad and menhaden often made up for a majority of consumed energy (Figure 10).

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25 0 1 2 3 4 5 6 7 8 9 AprMayJunJulAugSepOctNovDecAmerican Shad / Hour0 5 10 15 20 25 30 35Water Temperature (C) 1970 Trawls 2004 Trawls Water Temperature Figure 3 Comparing mean monthly trawl catch rates (+ standard error) of juvenile American shad between 1970 and 2004 with 2004 mean monthly water temperatures for the Palatka area of the St. Johns River, Florida.

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26 Figure 4. Mean number of Atlantic croaker, Atlantic menhaden, bay anchovy, threadfin shad, and other fish species collected per five minute trawl with standard error bars throughout 12 months of sampling on the Palatka area of the St. Johns River Florida.

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27 0 10 20 30 40 50 60 70 80 90 100Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Percent white catfish threadfin shad bay anchovy Atlantic menhaden Atlantic Croaker Figure 5. Percent composition of each prey species from total trawl catches from January through December at the Palatka area of the St. Johns River, Florida. The figure shows only species that accounted for at least 5% of monthly trawl catches.

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28 Figure 6. Percent by number of Atlantic croaker, Atlantic Menhaden, bay anchovy, and threadfin shad found by month in diet samples (solid line) of all species of predators collected by electrofishing and in trawl samples (dashed line) in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005.

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29 0 10 20 30 40 50 60 70 80 90 100Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Percent Frequency Small Medium Large Trawl Figure 7. [0]Percent by number of prey items that were not Atlantic croaker, Atlantic menhaden, bay anchovy, or threadfin shad found by month in diet samples of small (<356 mm), medium (356 mm – 432 mm), and large (>432 mm) largemouth bass collected by electrofishing and in trawl samples in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005.

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30 0 2 4 6 8 10 12 14 16 18 20Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb MarPercent Crayfish Armored Catfish Blue Crab Figure 8. Percent frequency by number for crayfish, armored catfish (plated and sailfin catfish), and blue crab found by month in all diet samples of largemouth bass collected by electrofishing in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005.

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31 -7 -6 -5 -4 -3 -2 -1 0 1 AprMayJunJulAugSepOctNovDecJanFebMarLOGIT Values Trawl Small Medium Large Figure 9. Mean LOGIT values for prey items (>0 means more seasonal diet items, <0 means more resident diet items) by month found in stomach contents of three size classes of largemouth bass (small <356 mm, medium 356 mm-432 mm and large >432 mm TL) and for trawl catches in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005.

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32 Figure 10. Total percent caloric content (left panels, caloric values estimated only for four species of prey fish therefore does not always represent total caloric intake) and percent by number (right panels) for Atlantic croaker, Atlantic menhaden, bay anchovy, and threadfin shad found in three size classes of largemouth bass diets,(small < 356 mm, medium 365 mm-432 mm, and large > 432 mm) collected by electrofishing in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005. Note : Only parts of croaker were found in large size class largemouth bass. These were counted in total prey counts, however parts were too small to estimate total weight and size and therefore were left out of caloric estimates.

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33 Table 1. Prey species classified as resident (collected in sample area year around) and seasonal (marine species collected seasonally in sample area) in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005. Family Resident Seasonal Fishes Atherinidae Brook silverside (Labidesthes sicculus ) Atlantic silverside (Menidia menidia ) Belonidae Atlantic needlefish (Strongylura marina ) Callichthyidae Plated catfish (Hoplosternum littorale ) Centrachidae Black crappie (Pomoxis nigromaculatus ) Bluegill (Lepomis macrochirus ) Largemouth bass (Micropterus salmoides ) Redbreast sunfish (Lepomis auritus ) Redear sunfish (Lepomis microlophus ) Spotted sunfish (Lepomis punctatus ) Warmouth (Lepomis gulosus ) Clupeidae Gizzard shad (Dorosoma cepedianum ) American Shad (Alosa sapidissima ) Threadfin shad (Dorosoma petenense ) Atlantic menhaden (Brevoortia tyrannus ) Blueback herring (Alosa aestivalis ) Hickory shad (Alosa mediocris ) Cyprinidae Golden shiner (Notemigonus crysoleucas ) Cyprinodontidae Seminole killifish (Fundulus seminolis ) Eleotridae Fat sleeper (Dormitator maculatus ) Elopidae Ladyfish (Elops saurus ) Engraulidae Bay anchovy (Anchoa mitchilli ) Gerreidae Irish Pompano (Diapterus auratus ) Gobiidae Clown goby (Microgobius gulosus ) Naked goby (Gobiosoma bosc )

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34 Table 1. Continued. Family Resident Seasonal Ictaluridae Loricariidae Channel catfish (Ictalurus punctatus ) White catfish (Ameirus catus ) Sailfin catfish (Pterygoplichthys multiradiatus ) Mugilidae Striped mullet (Mugil cephalus ) Poeciliidae Sailfin molly (Poecilia latipinna ) Sciaenidae Atlantic croaker (Micropogonias undulates ) Silver perch (Bairdiella chrysoura ) Crustaceans Portunidae Blue crab (Callinectes sapidus )

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35 Table 2. Summary of monthly mean trawl and electrofishing catch rates (fish/hour) for juvenile Alosa spp. and total number of hours sampled using each gear by month in the Palatka area of the St. Johns River from April 2004 through March 2005. American shad Month Trawl Catch Rate Hours Trawled Electrofishing Catch Rate Hours of Pedal Time April 0 1 0 1.7 May 0 1 0 1.0 June 2 1 0 1.7 July 0 1 0 2.3 August 0.33 3 0.6 1.7 September 2.5 2 2.4 3.8 October 4.5 2 1.98 4.1 November 2 2 6.96 4.6 December 2 1 3.24 1.6 January 0 1 0 4.2 February 0 1 0 3.4 March 0 1 0 3.0 Blueback Herring Month Trawl Catch Rate Hours Trawled Electrofishing Catch Rate Hours of Pedal Time April 1 1 0 1.7 May 12 1 0 1.0 June 1 1 0 1.7 July 1 1 0 2.3 August 0.3 3 0 1.7 September 4 2 6.1 3.8 October 4 2 3.9 4.1 November 1.5 2 13.5 4.6 December 0 1 36.1 1.6 January 2 1 4.9 4.2 February 0 1 0.3 3.4 March 0 1 0 3

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36 Table 2. Continued Hickory Shad Month Trawl Catch Rate Hours Trawled Electrofishing Catch Rate Hours of Pedal Time April 0 1 0 1.7 May 2 1 0 1.0 June 0 1 0 1.7 July 0 1 0 2.3 August 0.33 3 0 1.7 September 0 2 0.54 3.8 October 1.5 2 0.24 4.1 November 0 2 0.24 4.6 December 0 1 0 1.6 January 0 1 0 4.2 February 0 1 0 3.4 March 0 1 0 3.0

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37 Table 3. Mean total lengths of juvenile American shad collected with trawl gear by month in the Palatka area of the St. Johns River, Florida, from September through December of 1969, 1970, and 2004. Number collected (N) and Standard Error (S.E.) are shown by month for 2004 data. Months with no fish collected and no data available are labeled as Na. Data from 1969 and 1970 were from Williams and Bruger (1972). Month 1969 1970 2004 2004 (N) 2004 (S.E.) June Na Na 66 2 1 July Na Na Na 0 Na August Na Na 80 1 Na September 69 58 70 5 1.93 October 75 82 85 9 2.47 November 89 95 79 4 2.27 December 94 106 93 2 8

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38 Table 4. Number of predators sampled with electrofishing and how many contained stomach contents (number and percent) in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005. Predator Species Number Examined for Prey Number With Prey Percent With Prey Largemouth Bass 1,443 586 41 Black Crappie 120 56 47 Longnose Gar 36 13 36 Florida Gar 35 10 29 Striped Bass 29 16 55 White Catfish 25 6 24 Channel Catfish 18 10 55 Chain Pickerel 14 7 50 Ladyfish 10 3 30 Southern Flounder 8 3 38 Brown Bullhead 5 2 40 Red drum 3 2 67 Mangrove Snapper 1 0 0 Totals 1,747 714

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39 Table 5. Presence (x) or absence of prey items in diets collected from largemouth bass Micropterus salmoides of the small size class (<365 mm) in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005. Size Class Small Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec American shad Atlantic croaker x x x x Atlantic menhaden x x Atlantic needlefish Atlantic silverside x x bay anchovy x x x x x x x x x x x blue crab x x x x brook silverside x x x channel catfish clown goby x x x crayfish x x x x x fat sleeper gizzard shad x x golden shiner x x grass shrimp x x x x x hickory shad x Irish pompano ladyfish largemouth bass Lepomis spp. x x x x x x x mud crab x x x naked goby x x pink shrimp plated catfish x sailfin molly Seminole killifish x x silver perch striped mullet sailfin catfish x x threadfin shad x x x x x x x x x x x x white catfish x

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40 Table 5, continued. Presence (x) or absence of prey items in diets collected from largemouth bass Micropterus salmoides of the medium size class (366 mm – 432 mm) in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005. Size Class Medium Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec American shad Atlantic croaker x x x x Atlantic menhaden x x x Atlantic needlefish x x Atlantic silverside x x bay anchovy x x x x x x x x blue crab x x x x x x brook silverside channel catfish clown goby x x crayfish x x x x x x fat sleeper x gizzard shad x x x golden shiner x x grass shrimp x x hickory shad Irish pompano ladyfish largemouth bass x Lepomis spp. x x x x x x x mud crab x naked goby pink shrimp plated catfish x x x sailfin molly x Seminole killifish silver perch striped mullet sailfin catfish x threadfin shad x x x x x x x x x x x white catfish x

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41 Table 5, continued. Presence (x) or absence of prey items in diets collected from largemouth bass Micropterus salmoides of the large size class (>432 mm) in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005. Size Class Large Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec American shad x Atlantic croaker x x x x Atlantic menhaden x x x Atlantic needlefish x x x x x Atlantic silverside bay anchovy x x x x blue crab x x x x x x x brook silverside channel catfish x x clown goby crayfish x x x x fat sleeper gizzard shad x x x golden shiner x x grass shrimp x hickory shad Irish pompano x ladyfish x largemouth bass x Lepomis spp. x x x x x x x x x x x mud crab x naked goby pink shrimp x plated catfish x x x sailfin molly Seminole killifish x silver perch striped mullet x sailfin catfish x x threadfin shad x x x x x x x x white catfish x x x

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Table 6. Total numbers (N) of largemouth bass Micropterus salmoides from each of three size classes (small < 365 mm, medium 366-432 mm, and large > 432 mm) sampled each month containing stomach contents and the total number of prey items found in these stomachs in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005. Month Total N Total N of Prey items Total N Total N of Prey items Total N Total N of Prey items Small Small Medium Medium Large Large April 21 24 5 7 5 6 May 16 37 9 20 5 11 June 28 111 14 23 5 6 July 23 44 10 14 8 11 August 7 12 6 11 7 12 September 25 48 35 59 17 36 October 37 60 46 74 14 25 November 27 82 34 93 25 48 December 14 58 20 49 17 35 January 7 37 8 17 4 14 February 32 75 11 12 2 2 March 30 56 7 13 5 13 Total 267 644 205 392 114 219

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43 Table 7. Presence (x) or absence of prey items collected from predator species ( in order of number of diet samples collected from most on the left to least on the right) other than largemouth bass Micropterus salmoides in the Palatka area of the St. Johns River, Florida from April 2004 through March 2005. Predator Species Prey Species black crappie (n=56) striped bass (n=16) longnose gar (n=13) channel catfish (n=10) Florida gar (n=9) Atlantic croaker x x Atlantic menhaden x x Atlantic needlefish x bay anchovy x x blue crab x brook silverside x channel catfish crayfish x grass shrimp x gizzard shad x insects x Irish pompano x ladyfish x largemouth bass Lepomis spp. x x plated catfish x pork chops x threadfin shad x x x x sailfin catfish white catfish x

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44 Table 7. Continued Predator Species bowfin (n=8) chain pickerel (n=7) white catfish (n=6) ladyfish (n=3) redfish (n=2) Atlantic croaker x Atlantic menhaden Atlantic needlefish x bay anchovy x blue crab x x brook silverside channel catfish x crayfish x grass shrimp x x gizzard shad insects Irish pompano ladyfish largemouth bass x Lepomis spp. x plated catfish pork chops threadfin shad x x sailfin catfish x white catfish

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45 Table 8. Total number, mean and median length (mm), and mean and median weight (g) of each of the four most common prey species: Atlantic croaker (ATCR), Atlantic menhaden (ATME), bay anchovy (BAAN), and threadfin shad (THSH), found in each of three size classes (small < 365 mm, medium 366432 mm, and large > 432 mm) of largemouth bass, Micropterus salmoides, diets from April 2004 through March 2005 in the Palatka area of the St. Johns River, Florida. Size Class Species Number Mean Length (mm) Median Length (mm) Mean Weight (g) Median Weight (g) Small ATCR 5 63 58 3.8 2.0 Medium ATCR 2 80 80 10.3 10.3 Large ATCR 3 N/A N/A N/A N/A Small ATME 7 109 110 13.1 12.3 Medium ATME 17 117 115 28.2 15.6 Large ATME 7 112 108 12.8 12.9 Small BAAN 152 34 31 0.5 0.2 Medium BAAN 33 54 56 1.2 0.9 Large BAAN 17 45 46 0.7 0.7 Small THSH 100 63 63 2.7 2.0 Medium THSH 104 70 70 3.3 2.7 Large THSH 29 72 72 3.8 2.9

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46 DISCUSSION Juvenile Alosa spp. Abundance Night-time trawl samples collected low numbers of juvenile American shad, blueback herring, and hickory shad. However, catch rates were similar to those of Williams and Bruger (1972) using the same trawling methods in 1969 and 1970. Both studies found highest catch rates occurring in October. Most American shad were collected from September through December near Palatka, which is described as the northern extent of the nursery area (Williams and Bruger 1972). A few juvenile American shad were collected in June. Williams and Bruger (1972) sampled the entire river and found some juveniles entered brackish areas of the river during summer but did not find shad leaving the mouth of the river until November. From September through December mean monthly water temperatures ranged from 27 C down to 19 C. Mean water temperature for the month of October when most fish were collected was 24 C similar to mean temperatures in 1970. Emigration this time of year begins much later than fish at higher latitudes (October through November in North Carolina: Davis and Cheek 1966; November through March in Florida: Williams and Bruger 1972; June through November in New York: Limburg et al. 2003). My occurrence of American shad corresponded with declining water temperatures similar to Williams and Bruger (1972), although it is not clear that declining water temperatures caused movement of American shad in this system relative to other potential mechanisms (e.g., photoperiod, light availability).

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47 While electrofishing I collected juvenile American shad from August through December with the highest number being collected in November. On 18 November 2004, I collected 26 juvenile American shad while electrofishing for predators. This suggests that this may have been a time when large numbers of shad were moving through the area. However, trawl sampling in November did not reveal a similar trend. As I collected many more fish electrofishing than trawling, the surface trawl might not be an effective sampling gear for collecting juvenile American shad in this region. Williams and Bruger (1972) and McBride (2000) reported that hickory shad were the first of the Alosa spp. to make their spawning runs up the St. Johns River, November through February, followed by the American shad and blueback herring in December through April. Loesch et al. (1982) found that in Virginia Rivers American shad spawning precedes that of blueback herring. OÂ’Leary and Kynard (1986) stated that juvenile blueback herring will emigrate slightly earlier than American shad. My data revealed this same trend as I found juvenile blueback herring as early as April with 12 being collected in May, the highest number of bluebacks collected during any month. Juvenile hickory shad appeared in May before the first American shad arrived in June following the suggested trend. Although catch rates were low for juvenile Alosa spp trawl gear did effectively sample various other prey species. The sample area was impacted by two major hurricanes during the fall of my sample period. Twice and only weeks apart, the river flooded and experienced extremely high flow rates. I did not collect high numbers of juvenile Alosa spp. during these time periods, evidence that the high flows did not trigger

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48 juveniles to emigrate earlier than normal. However, the high flow conditions could have reduced my sampling efficiency and contributed to low catch. Nevertheless, low catches of juvenile Alosa spp. may be indicative of low population size. Reduced populations of American shad have occurred along the Atlantic coast and can be attributed to factors such as decreasing water quality, changes in flow, increased sediment load, overharvest by recreational anglers and previous commercial fishers, and dams (Walburg and Nichols 1967; Williams and Bruger 1972; Hightower et al. 1996; Ross et al. 1997; McBride 2000; Olney and Hoenig 2001; Limburg et al. 2003; Scharpf 2003). My catch of Alosa spp. was similar to values from the early 1970Â’s, indicating a lack of large changes in juvenile abundance between time periods. Thus, my results do not indicate a large reduction in abundance, albeit with both time periods having a relatively low sample size. Atlantic estuaries are important nursery grounds for various anadromous species (Juanes et al. 1993), and seasonal influxes of anadromous prey species such as juvenile American shad and blueback herring may play an important role in the diets of relatively stationary predators such as largemouth bass (Yako et al. 2000). Yako et al. (2000) found that largemouth bass in coastal Massachusetts rivers with such calorie rich prey available seasonally may obtain larger sizes and have faster growth rates than bass in similar river systems without anadromous prey available. Pine et al. (2005) determined that flathead catfish Pylodictus olivaris in coastal North Carolina Rivers selected juvenile American shad and hickory shad over resident freshwater species. Other studies on the Atlantic coast have shown that juvenile striped bass play an important role as prey for young bluefish (Juanes et al. 1993).

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49 Throughout an entire year of sampling I found only one juvenile American shad in a predator diet, which was a largemouth bass collected in December. One hickory shad was found in a largemouth bass diet in August. The effect of predation on a particular species during a year is likely to reflect the relative abundances of predator and prey and the food preferences of the predator (Juanes et al. 1993). I did not observe a large pulse of Alosa spp. moving through during the fall, and I did not observe predators switching over to this energetically beneficial prey. Thus, it appears that Alosa spp. were relatively uncommon in the river and thus were uncommon as prey for predators. Relative Prey Abundance and Occurrence in Predator Diets There is a wide variety of prey species found in this area of the St. Johns River. Not only are all of the common freshwater species found here but also many marine species spend all or part of the year in this area. The presence of salt springs that drain into the river cause salinity to rise upriver from Palatka to Lake George (Odum 1953), making the area suitable for seasonal species. Some seasonal species in this study were abundant all year and therefore classified as resident species (e.g., bay anchovy), whereas others occurred only seasonally and were classified as seasonal species (e.g., Atlantic croaker and Atlantic menhaden). This was also true for abundance of freshwater prey species found in this study, as some were present in about the same abundance all year whereas abundance of others spiked seasonally (e.g., threadfin shad and white catfish). Juvenile Atlantic croaker spend the first months of life in low salinity to freshwater areas (Murdy et al. 1997). In the fall both juvenile and mature Atlantic croakers leave rivers and migrate to bays (Murdy et al. 1997). These life history traits explain why abundance was highest February through May, but they were collected from January through August and were composed primarily of small juveniles.

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50 Threadfin shad spawn in the spring and often again in the fall (Jenkins and Burkhead 1993). Threadfin shad were not collected in trawls from December through February, partly because mesh was too large to collect age-0 fish and partly because mature shad may have moved deeper and become unavailable to the surface trawl or were avoiding the trawl. During July and August threadfin shad accounted for a vast majority of fish caught in trawls and the second most abundant species year round. Bay anchovy reproduce throughout the year when water temperatures exceed about 12 C, which allows nearly year round spawning in Florida (Murdy et al. 1997). Bay anchovy were collected in trawls in all months of this study. Only during April did the percentage of total catch that anchovy comprised drop below 20%. Eight months out of the year anchovy made up for more than 50% of each monthÂ’s total trawl catch, and they were by far the most collected species using the surface trawl gear. Larval Atlantic menhaden move into freshwater areas where they metamorphose into juveniles. These juveniles will remain in freshwater areas using them as nursery areas until fall when they move out to the bays and ocean (Murdy et. al 1997). Juvenile menhaden will congregate into dense schools as they leave these nursery areas from August through November (Rogers and Van Den Avyle 1983). Menhaden were only collected in trawls from September through November, presumably when they were emigrating from the low salinity river to the sea. Prey abundance and size strongly influence predator diets and growth. If the abundance of a preferred prey increases, importance in the diets of predators increases (Adams et al. 1982; Storck 1986; Hartman and Margraf 1992; Michaletz 1997). I found a similar relationship between largemouth bass (predator) and Atlantic menhaden (prey).

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51 Bay anchovy were the most abundant and available prey in the sample area throughout the entire sampling period. At various times of the year high numbers of anchovy were consumed with extremely high numbers consumed by small fish in spring and summer. Bay anchovy were eaten by all size classes of largemouth bass in the winter when other prey species were less prevalent and during the summer by small size largemouth bass which may be because juvenile threadfin shad had exceeded their gape limits although this was not measured. Threadfin shad were found in diets during all 12 months of the year, although highest numbers were found in diets from July through December (Figure 6). This follows the same trend as was seen in trawl collections. Although no threadfin shad were collected during December-February with trawls this does not necessarily mean their abundance had dropped but is more than likely because they were no longer on the surface and available to capture by a surface trawl, as they were still found in predator diets. Threadfin shad were the most common occurrence in medium and large size bass diets and second to bay anchovy in small size class fish. Although bay anchovy were consumed in higher numbers by small size class largemouth bass, threadfin shad accounted for most of the caloric intake. Medium and large size largemouth bass diets revealed threadfin shad occurring most by number throughout the year and making up for most of caloric intake. Numbers of threadfin shad found in diets peaked in late summer and early fall. Although common in trawl catches from February through August, Atlantic croaker never appeared frequently in largemouth bass diets except for May when they were the most common prey species by number and also accounted for most of the

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52 caloric intake found in large size class fish diets. There was a small spike in numbers of Atlantic croaker found in small and medium size class largemouth bass diets during the summer. I did not detect that predators were switching to croaker as prey even during times of year when they were relatively abundant in trawl samples. Atlantic menhaden were collected in diet and trawl samples only during the months of September through November. During these months I did find menhaden in the diets of all size classes of bass. Also, menhaden were found in diets of striped bass and longnose gar during this same time period. For large size class largemouth bass in September, bay anchovy were the most common prey consumed by number, followed closely by Atlantic menhaden, which made up for the highest caloric intake of any month. Atlantic menhaden then accounted for most of the caloric intake for large size class fish in November. This suggests that predators did switch to menhaden and took advantage of their presence and high caloric content during the fall. Caloric values used were measured from only adult samples of Atlantic croaker (Thayer et al. 1973) which I rarely collected, so caloric estimates may vary. Atlantic menhaden (Steimle Jr. and Terranova 1985), bay anchovy (Steimle Jr. and Terranova 1985), and threadfin shad (Strange and Pelton 1987) caloric estimates were made from most commonly collected size fish. Bay anchovy and threadfin shad of all sizes were sampled during this project so these estimates should be accurate while only juvenile Atlantic menhaden were sampled meaning their caloric estimates are likely less accurate. Throughout certain times of the year several prey items occurred regularly in diets that were not collected in trawl samples. During April and May blue crabs Callinectes sapidus made up more than 10 % of items found in all predator diets. Also from October

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53 – December, crayfish Procambarus spp. and armored catfish (sailfin catfish and brown hoplo) occurred frequently in diets. This switch to crayfish and armored catfish was likely due to extremely high water after hurricanes. Predators at this time were often collected in areas of newly flooded timber where these prey items were likely abundant. Small fish often consumed non-fish prey, such as insects and various invertebrates. Management Implications Estimates of stock abundance are crucial to the assessment and effective management of freshwater, anadromous, catadromous, and marine fish populations. Massman et al. (1952) found surface trawls to be an effective technique to sample young fishes in tidal rivers. Loesch et al. (1982) proved this technique even more effective on juvenile Alosa spp when used after dark as juveniles will be feeding on the surface at this time. I was fortunate enough to have data collected by Willams and Bruger (1972) using the same size surface trawl pulled at night and in the same sampling area. My catch rates for juvenile American shad in the trawl samples were similar to what they found over 30 years ago. This could mean that recruit abundance is similar to what it was back then. However, in a river as big as the St. Johns, over 2 km wide in many areas, it may be hard to effectively sample with a 3 m wide trawl net. If the net was pulled too fast a pressure wave would build in front of it causing fish to avoid the net, while if pulled slow enough to minimize the pressure wave, fish were seen jumping out of the trawl. Thus, the trawl appeared to not be a highly effective gear for sampling the open water planktivorous fishes, especially considering that electrofishing collected more than twice

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54 as many juveniles in the daytime while looking for predators. Juveniles were in the area, but my collections were not as successful as planned. If abundance was relatively high, I would have seen higher numbers trawling, shocking, and in diets. Although the trawl may not have effectively sampled all fishes in the water column, I was able to detect trends between trawl catch and occurrence in predator diets for several species. After FloridaÂ’s net ban in 1995, FloridaÂ’s American shad fishery was reduced to recreational anglers fishing on the spawning grounds (McBride 2005). McBride (2000, 2005) found angler catch per unit effort (CPUEÂ’s) of American shad increased in 19951996 and 1997-1998 but then decreased to the lowest levels of the creel study in 20042005. Although this decrease in CPUE was observed, angler catch rates have remained stable throughout the last 15 years, at about one fish per hour. This however, comes along with a decrease in angler effort (McBride 2005). Current harvests of American shad are well below historical levels, and although unlikely, even these extremely low current exploitation rates could be excessive because population sizes of American shad are thought to be low and historically depressed (Hightower et al. 1996; McBride 2005). Methods for rebuilding of shad stocks include reduction of harvest of adults as well as improvement and protection of spawning and nursery habitats (Walsh et al. 2005). Commercial landings in the St. Johns River have been at nearly zero since the mid-1990s (McBride 2000). All offshore commercial fishing directed at American shad was shut down in 2004. Even with this reduction it does not appear that stocks w ill regain historical levels seen at the turn of the century 20th century (Hightower et al. 1996).

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55 Because current fishing mortality is not likely to be causing the extremely low population levels, other factors need to be considered. Walsh et al. (2005) determined that in the Roanoke River flooded forest land would be beneficial to blueback herring and alewife eggs and larvae. During my research I did not see extremely high river levels until after the hurricanes in the fall of 2004. I was unable to detect how factors from these storms affected survival and abundance of juvenile American shad during 2004. Crecco et al. (1986) concluded that major climatic events can overshadow the compensatory mechanism of American shad populations. They also noted that turbulent June flow rates promote unfavorable feeding conditions for larvae, eventually reducing survival. In the St. JohnÂ’s River, high flow rates cause the river to have high color from tannic acids, potentially limiting primary and secondary production. These extremely high flow rates seen in the fall of my sampling period may have caused similar problems for juveniles and contributed to low catch rates/abundance. Usually, increasing fall river flow would trigger juveniles to emigrate (OÂ’Leary and Kynard 1986). However, I saw no such trends, suggesting that temperature or something such as photoperiod may be more important factors to trigger emigration than flow. OÂ’Leary and Kynard (1986) also suggested that when juveniles reach a certain size they will leave the river regardless of environmental conditions, which I did not see. Increased industry, residential growth, and organic pollution have all caused decreases in the quality of habitat available to American shad (Limburg et al. 2003; McBride 2005). Williams and Bruger (1972) found some of the major problems the St. Johns River American shad population faced was changes in flow due to an extensive series of water control structures, degradation of spawning areas, and increased industrial

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56 and domestic effluents. Spawning grounds have remained the same for over 50 years, so it is crucial that areas be protected from future degradation (McBride 2005). Although only one dam was constructed on the main river channel, these problems still affect the St. Johns River today. (McBride 2005). One natural problem facing this shad stock is that they are at the southernmost extent of their native range which may cause temporal and ecological problems. Commercial fishing is non-existent now, however its effects are still present. McBride (2005) found female shad considerably less abundant than males throughout the spawning season. This may be due to the commercial fishers who targeted the female shad for roe. Although the commercial American shad fishery has been closed, nontarget mortality from other fisheries may be significant (McBride 2005) The recreational fishery has also drastically declined so only time will tell if the American shad population in this river rebounds (McBride 2000; McBride 2005). Crecco et al. (1986) showed that American shad can produce large year classes from few adults, but I did not find evidence of abundant juvenile American shad in this system. In the future we need to attempt to get better population estimates for spawning fish and the resulting juvenile recruits. I believe we need to look more closely at available spawning habitat and also into the larval life stage to examine if water quality is having an effect on survivability or if there is lack of some critical food available to juveniles in the upper St. JohnÂ’s River. Stocking could be a future option to help bring the population back up to historical levels. All of these issues need to be looked into and are crucial to successfully manage and maintain the St. Johns River American shad population.

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57 Prey availability and amount of energy consumed govern the proportion of consumed energy allocated to the principal physiological functions (Adams et al. 1982). In many freshwater systems threadfin and gizzard shad are the primary prey species. These species undergo large temporal fluctuations in abundance due to temporal changes (Adams et al. 1982; Storck 1986; and Michaletz 1997). This was not a problem in this area as threadfin shad were found in diets throughout the entire year. Also, there was usually more than one abundant prey species each month. Prey availability drives the success of a fish population (Krohn et al. 1997; Yako et al. 2000) and prey selectivity of predators is an important mechanism structuring aquatic communities (Juanes et al. 2001). In coastal systems anadromous fishes may provide a seasonal influx of high energy prey (Durbin et al. 1979). I did not find that anadromous shads were important in predator diets due to their apparent low abundances, however seasonally Atlantic croaker and Atlantic menhaden were important prey items. High growth rates can increase the size of largemouth bass, in turn increasing the range of prey they can ingest and their probability of survival during the winter through spring period (Adams et al. 1982). The influx of nutrient-rich Atlantic menhaden in the fall may help this cause, especially as all size classes of largemouth bass were actively feeding on them when present. Managers should focus on managing the river as a whole. This includes looking into flow issues, water quality and pollution concerns, and habitat quality. This research will help not only predator (e.g., largemouth bass) and American shad populations of this area but also all fish and wildlife along this unique 500-km long river.

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58 LIST OF REFERENCES Adams, S. M., R. B. McLean, and M. M. Huffman. 1982. Structuring of a predator population through temperature-mediated effects on prey availability. Canadian Journal of Fisheries and Aquatic Sciences 38:387-393. ASMFC (Atlantic States Marine Fisheries Commission). 1999. Amendment 1 to the interstate fishery management plan for shad & river herring. Fishery Management Report No. 35. Washington D.C. 77pp. Beamesderfer, R. C., B. E. Rieman, L. J. Bledsoe, and S. Vigg. 1990. Management implication of a model of predation by resident fish on juvenile salmonids migrating through a Columbia River reservoir. North American Journal of Fisheries Management 10:290-304. Brown, B. L., P. E. Smouse, J. M. Epifanio, and C. J. Kobak. 1999. Mitochondrial DNA mixed-stock analysis of American shad: coastal harvests are dynamic and variable. Transactions of the American Fisheries Society 128:977-994. Buckel, J. A., D. O. Conover, N. D. Steinburg, and K. A. McKown. 1999. Impact of age-0 bluefish ( Pomatomus saltatrix ) predation on age-0 fishes in the Hudson River estuary: evidence for density-dependent loss of juvenile striped bass ( Morone saxatilis ). Canadian Journal of Fisheries and Aquatic Sciences 56:275-287. Crecco, V., T. Savoy, W. Whitworth. 1986. Effects of density-dependent and climatic factors on American shad, Alosa sapidissima recruitment: a predictive approach. Canadian Journal of Fisheries and Aquatic Sciences 43:457-463. Davis, J. R., and R. P. Cheek. 1966. Distribution, food habits, and growth of young clupeids, Cape Fear River System, North Carolina. Proceedings of the 20th Annual Conference of Southeastern Association of Game and Fish Commissions 250-260. Davis, S. M. 1980. American shad movement, weight loss and length frequencies before and after spawning in the St. Johns River, Florida. Copeia (4)889892. Durbin, A. G., S. W. Nixon, and C. A. Oviatt. 1979. Effects of the spawning migration of the alewife, Alosa pseudoharengus on freshwater ecosystems. Ecology 60:817.

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59 Froese, R., and D. Pauly. Editors. 2000. Fishbase 2000: concepts, design and data sources. ICLARM, Los Banos, Laguna, Phillipines. World wide web electronic publication. www.fishbase.org version (06/2005). Gerstell, R. 1998. American shad in the Susquehanna River Basin: a three hundred year history. Pennsylvania State University Press, University Park, PA. Glebe, B. D., and W. C. Leggett. 1981a. Temporal, intra-population differences in energy allocation and use by American Shad Alosa sapidissima during the spawning migration. Canadian Journal of Fisheries and Aquatic Sciences 38:795-805. Glebe, B. D., and W. C. Leggett. 1981b. Latitudinal differences in energy allocation and use during the freshwater migrations of American shad Alosa sapidissima and their life history consequences. Canadian Journal of Fisheries and Aquatic Sciences 38:806-820. Hartman, K. J., and F. J. Margraff. 1992. Effects of prey and predator abundances on prey consumption and growth of walleyes in western Lake Erie. Transactions of the American Fisheries Society 121:245-260. Hightower, J. E., A. M. Wicker, and K. M. Endres. 1996. Historical trends in abundance of American shad and river herring in Albermarle Sound, North Carolina. North American Journal of Fisheries Management 16:257-271. Jenkins, R. E., and N. M. Burkhead. 1993. Freshwater fishes of Virginia. American Fisheries Society, Bethesda, Maryland. Juanes, F., J. A. Buckel, and F. S. Scharf. 2001. Predatory behaviour and selectivity of a primary piscivore: comparison of fish and non-fish prey. Marine Ecology Progress Series 217:157-165. Juanes, F., R. E. Marks, K. A. McKown, and D. O. Conover. 1993. Predation by age-0 bluefish on age-0 anadromous fishes in the Hudson River estuary. Transactions of the American Fisheries Society 122:348-356. Krohn, M., S. Reidy, and S. Kerr. 1997. Bioenergetic analysis of the effects of temperature and prey availability on growth and condition of northern Cod ( Gadus morhua ). Canadian Journal of Fisheries and Aquatic Sciences 54 (S1):113-121. Leggett, W. C., and J. E. Carscadden. 1978. Latitudinal variation in reproductive characteristics of American shad ( Alosa sapidissima ): evidence for population specific life history strategies in fish. Journal of the Fisheries Research Board of Canada 35:1469-1478.

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60 Leggett, W. C., and R. R. Whitney. 1972. Water temperature and the migrations of American shad. Fishery Bulletin 70 (3):659-670. Limburg, K. E., 1996. Modelling the ecological constraints on growth and movement of juvenile American shad ( Alosa sapidissima ) in the Hudson River estuary. Estuaries 19 (4):794-813. Limburg, K. E., K. A. Hattala, and A. Kahnle. 2003. American shad in its native range. Pages 125-140 in K. E. Limburg and J. R. Waldman, editors. Biodiversity, Status, and Conservation of the WorldÂ’s Shads. American Fisheries Society, Symposium 35, Bethesda, Maryland. Littell, R. C., G. A. Milliken, W. W. Stroup, and R. D. Wolfinger. 1996. SAS system for mixed models. SAS Institute. Cary, North Carolina. Loesch, J. G., W. H. Kriete, Jr., and E. J. Foell. 1982. Effects of light intensity on the catchability of juvenile anadromous Alosa species. Transactions of the American Fisheries Society 111:41-44. Massman, W. H., E. C. Ladd, and H. N. McCutcheon. 1952. A surface trawl for sampling young fishes in tidal rivers. Transactions of the Seventeenth North American Wildlife Conference. March 17-19. Wildlife Management Institute, Washington D.C. McBride, R. S. 2000. FloridaÂ’s shad and river herrings ( Alosa species): a review of population and fishery characteristics. Florida Fish and Wildlife Conservation Commission. FMRI Technical Report TR-5, Tallahassee, Florida. McBride, R. S. 2005 Develop and evaluate a decision-making tool for rebuilding American and Hickory shad. Project F-106. Three-year Final Performance Report for Federal Aid in Sportfish Restoration Act. November 30. 60pp. FWC/FWRI File Code: 2459-02-05-F. McPhee, J. 2002. The Founding Fish. Farrar, Straus and Giroux. New York, New York. Michaletz, P. H. 1997. Influence of abundance and size of age-0 gizzard shad on predator diets, diet overlap, and growth. Transactions of the American Fisheries Society 126:101-111. Murdy, E. O., R. S. Birdsong, and J. A. Musick. 1997. Fishes of the Chesapeake Bay. Smithsonian Institution Press, Washington and London. National Oceanic and Atmospheric Administration (NOAA). 2005. Annual Commercial Landing Statistics. U. S. Department of Commerce. Washington D. C. USA.

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64 BIOGRAPHICAL SKETCH Nicholas Aaron Trippel was born January 5, 1981 in Goshen, Indiana, the son of Donald and Debra Trippel. He graduated from Northwest Guilford High School, North Carolina in 1999. He received his Bachelor of Science degree in Fisheries Management from Auburn University. He became interested in the field of fisheries management while completing a year long mentorship project with a district fisheries biologist during his senior year of high school. While completing his Bachelor of Science degree he gained fisheries management experience working on research projects while employed with Alabama Fish and Wildlife Cooperative Research Unit. After completing his undergraduate degree he received a job working as a full time lab technician at the University of Florida for Dr. Mike Allen. After working as a lab technician for eight months Dr. Mike Allen took him on as graduate research assistant to pursue a Master of Science degree in fisheries management. After graduation in May 2006, he hopes to pursue a successful career in fisheries management working as a state freshwater fisheries biologist.