FWS/OBS-82/05

September 1984

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THE ECOLOGY OF THE

APALACHICOLA

BAY SYSTEM:

AN ESTUARINE PROFILE

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Fish and Wildlife Service

U.S. Department of the Interior

FWS/OBS-82/05 '^ ' ''' September 1984

THE ECOLOGY OF THE APALACHICOLA BAY SYSTEM:

AN ESTUARINE PROFILE

by

Robert J. .J.iv inQston

Department of Biological Science

Florida State University

Tallahassee, PL 32306

> ) ~^^= Project Officer

Wi ley M. Kitchens

National Coastal Ecosystems Team

U.S. Fish and Wildlife Service

1010 Gause Blvd.

Slidell, LA 70458

Prepared for

National Coastal Ecosystems Team

Division of Biological Services

Research and Development

Fish and Wildlife Service

U.S. Department of the Interior

Washington, DC 20240
Library of Congress CarH Mo. R4-'S01077

This report should be cited as:

Livingston, R.J. 1984, The ecology of the Apalachicola Bay system: an estuarine

profile. U.S. Fish Wildl. Serv. FWS/OBS 82/05. 148 pp.
PREFACE

This paoer represents a synthesis of

knowledqe concerninq the Apalachicola

drainage system, which is located in

Florida, Georgia, and Alabama. The

Apalachicola Bay complex is only one part

of a ma.ior drainage area that includes the

Apalachicola, Chattahoochee, and Flint

River systems on one side and the

northeastern Gulf of Mexico on the other.

The boundaries that separate various

components (i.e., the river and its

associated wetlands, the bay system, and

the open gulf) are artificial in an ecological sense. Likewise, the

traditional boundaries that have separated

various scientific discipl ines--such as physics, chemistry, meteorology, and

biology--are somewhat arbitrary when a systems approach is used to determine the

functional interactions among interacting

subsystems. Thus various boundaries must

be crossed when the investigator attempts

to understand an entire aquatic ecosvstem.

Over the past 12 years, researchers

in the Apalachicola system have carried

out a series of multidisciplinary and

interdisciplinary studies to determine the

response of the Aoalachicola estuary to a series of environmental variables. Such

an effort can be likened to the growth of

concentric layers of a snowball as it

rolls down a hill. The solution of each

problem forms the foundation for a new guestion, which, in turn, serves as the

template for new hypotheses and tests.

The combination of background field

analyses and experiments in the laboratory

and the field have been used as the basis

of this effort. Eventually, we can view

the overall picture by cutting through the

snowball of ideas, hypotheses, and

resolutions to form models of how the

ecosystem works. As of this writing, 12

years of continuous field and experimental

data have been transformed into

computerized files, which are now being

used to develop models of how the

Aoalachicola Ray system works in

comparison with other such systems in the

southeastern United States.

The scientific work on the

Apalachicola estuary is only the first

step in our understanding of system

functions. Increasingly, humans are having an important influence on natural

aquatic systems. Urbanization,

industrialization, and agricultural

activities can lead to habitat

destruction, pollution, and severe restrictions on productivity, which, in

turn, can be translated into very real

socioeconomic problems. The Apalachicola

area is a multiple-use system.

Accordingly, sound land planning and

progressive resource management are best

carried out with a comprehensive base of

objective scientific and economic

information. With the recent

establishment of the Apalachicola River

and Bay National Estuarine Sanctuary--the

largest such sanctuary in the nation--the

Apalachicola drainage system has been

designated by law as a special area, a place of refuge and shelter for important

aquatic species as well as humans as integral parts of the ecosystem. As one

of the last relatively natural biq river

areas in the United States, the highly

productive Apalachicola system is small

enough to analyze in a comprehensive

scientific fashion while being extensive

enough to be used as a natural model for

other such areas. The Apalachicola valley

is currently part of a major experiment to determine whether scientific data can be

translated into a comprehensive resource management program that will accommodate

economic development while perpetuating

the natural resources of the region.

m
SUMMARY

The results of 1? years oi" continuous

field studies and exoeriments in the

Apalachicola Bay system are reviewed and

summarized in this paoer. Included are data concerninq the geography, hydrology,

chemistrv, geology, and biology of the

Apalachicola drainage system with particular emphasis on the estuary and associated

waters.

The Apalachicola Bay system is part of

a major drainage area that includes four

rivers and their associated wetlands in

Georgia, Alabama, and Florida. The Bay is

a shallow coastal lagoon fringed by

barrier islands and dominated by wind

effects and tidal currents. River bottomlands that include the channels, sloughs,

swamps and backwaters, and periodically

flooded lowlands are important components

of the system. Principal influences on the biological processes in the estuary

are the physiography of the basin, river

flow, nutrient input, and salinity distribution in space and time. Water

quality is affected by periodic wind and

tidal influences and freshwater inflows.

Compared to most of the estuaries in

the United States, the Aoalachicola Ray

system is in a relatively natural state,

although hardly pristine. However,

economic development and pooulation growth

are beginning to put pressure upon the

region, threatening it with destructive

changes. The economic and ecological

importance of the area as a producer of

food and as shelter for diverse species is

such that it has inspired a movement to protect its natural resources. Broadening

the economic base of the region while

maintaining its biological productivity

will require the development of a comprehensive management plan based on the

deepest possible understanding of the

basis for that productivity, supported by

ongoing study, close monitoring, and

continued cooperation from local

interests.

Research efforts to acquire the

necessary understanding are not yet com- plete, but have nonetheless given rise to one of the most extensive computerized

data bases so far assembled on an estuarine system. Powerful programs for

wo»-king with these data have also beei^ developed; because of the extreme com- plexity of their interplay, computer

analysis has been and will continue to be

a primary tool in understanding how

physical and biological processes work in

the estuary.

Rased upon the data obtained thus

far, some efforts have been initiated to preserve and protect important freshwater

and estuarine wetlands. Included in these

efforts are the followina:

• State and federal land-purchase

programs

• Integration of local (county) land- use regulations into a comprehensive

plan for new and existing

development

t Creation of the Apalachicola River

and Bay National Estuarine Sanctuary,

the largest such sanctuary in the

country.

The effort to manage the Apalachicola

Bay system is an ambitious one; only time

will tell whether it will be successful in

its effort to protect important wildlife

values as the region undergoes economic

development.

TV
CONTENTS

Page

PREFACE 1 i 1 SUMMARY i V FIGURES VI i TABLES X CONVERSION TABLE xi i ACKNOWLEDGMENTS xi i i

1. INTRODUCTION (HISTORICAL PERSPECTIVE AND OVERVIEW) 1 1.1. Geographic Setting and Classification 1 1.2. Driving Forces and Human Influence 2

2. ENVIRONMENTAL SETTING 6 2.1. Origin and Evolution of the Estuary 6 2.1.1. Geological Time Frame 6 2.1.2. Geomorphology and Regional Geology 7 2.1.3. Watershed Characterization 9 2.1.4. Barrier Islands 10

2.2. Climate 11 2.2.1. Temperature 11 2.2.2. Precipitation 11 2.2.3. Wind 12

2.3. Hydrology 13

2.3.1. Freshwater Input 13

2.3.2. Tides and Currents 13

2.4 Physical/Chemical Habitat 14

2.4.1. Temperature and Salinity 14 2.4.2. Dissolved Oxygen 18

2.4.3. pH 18

2.4.4. Water Color and Turbidity 18

2.5 Biological Habitats 19

2.5.1. Wetlands 19

2.5.2. Seagrass Beds 24

2.5.3. Soft-Bottom Substrates 25

2.5.4. Oyster Bars 25

2.5.5. Nearshore Gulf Environment 25

2.6 Natural Resources of the Apalachicola Drainage System 26

3. PRIMARY PRODUCTIVITY AND NUTRIENT CYCLING 28

3.1. Primary Producers 28

3.1.1. Allochthonous Sources 28

3.1.2. Autochthonous Sources 31

3.2. Detritus Flux and Nutrient Dynamics 36

3.3. Microbial Ecology 41

4. SECONDARY PRODUCERS 43

4.1. Zooplankton 43

4.2. Larval Fishes 46
4.3. Benthos 49

4.4. Oysters 60

4.5. Nekton 60

5. NICHE DIVERSITY, TROPHIC INTERACTIONS, AND COMMUNITY STRUCTURE 76

5.1 Habitat-Specific Associations 76

5.1.1. Marshes 76

5.1.2. Seagrass Beds 76

5.1.3. Litter Associations 77

5.1.4. Oyster Bars 79

5.1.5. Subtidal (Soft-Sediment) Communities 79

5.2 Physical Control of Biological Processes 80

5.3. Trophic Relationships and Food-Web Structure 83

5.4. Predator-Prey Interactions and Community Response 88

6. LONG-TERM ECOLOGICAL RELATIONSHIPS 90

7. THE ESTUARY AS A RESOURCE 99

7.1. Fisheries 99

7.2. Socioeconomic Factors 101

7.3. Existing and Projected Impact by Man 103

7.3.1. Physical Alterations 103

7.3.2. Toxic Substances 104

7.3.3. Municipal Development 105

7.4. Land Planning and Resource Management 107

7.4.1. Public Land Investment 108

7.4.2. The Apalachicola Estuarine Sanctuary 109

7.4.3. Local Planning Efforts and Integrated Management 110

7.4.4. Integration of Management Efforts 110

8. COMPARISON WITH OTHER ESTUARIES 112

LITERATURE CITED 118

APPENDICES 131

A. Overview of Sampling Program in North Florida Coastal Areas 131

1. Apalachicola Bay System 131

2. Apalachee Bay System 132

B. Computer Programs for Analyzing Field and Laboratory Data 134

1. Special Program for Ecological Science (SPECS):

System Overview 134

2. "MATRIX" Program System: Summary of Capabilities 137

C. Review of Ongoing Research Programs of the Center for Aquatic

Research and Resource Management (Florida State University) 144

1. Overall Scope of the Program 144

2. Center for Aquatic Research and Resource Management:

Personnel (1984) 146

VI
FIGURES

Number Page

1 The tri -river drainage area 1

2 Location of the tri-river drainage system in the southeastern

United States 2

3 Important features of the Apalachicola Bay system, the major

contributing drainages, and the barrier island complex 3

4 Impoundments along the tri-river system 4

5 The Apalachicola estuary 6

6 Geological features of the Apalachicola drainage system 7

7 Natural areas of the Apalachicola basin 9

8 Aerial view of St. Vincent Island 11

9 Seasonal averages of Apalachicola River flow and rainfall from

Columbus, GA, and Apalachicola, FL 11

10 Six-month and thirty-six month moving averages of Apalachicola River

flow and Apalachicola rainfall 12

11 Net water current patterns in the Apalachicola estuary as indicated

by flow model s 14

12 Apalachicola River flow and monthly average minimum air temperature 15

13 SYNMAP projections of average levels of salinity, dissolved oxygen,

turbidity, and color at permanent stations in the Apalachicola

estuary 16

14 Surface salinity at stations 1 and 5 in the Apalachicola estuary from

1972 through 1982 17

15 Surface dissolved oxygen at stations 1 and 5 in the Apalachicola

estuary from 1972 through 1982 18

16 Water color at stations 1 and 5 in the Apalachicola estuary from

1972 through 1982 19

17 Turbidity at stations 1 and 5 in the Apalachicola estuary from

1972 through 1982 19

18 Frequently flooded areas and soil associations in the Apalachicola

River Basin 23

v11
Number Page

19 Distribution of the marshes and submergent vegetation in the

Apalachicola estuary 24

20 Distribution of oyster bars and sediments in the Apalachicola

estuary 26

21 Nutrient/detritus transport mechanisms and long-term fluctuations in

detrital yield to Apalachicola River flow 28

22 Regression analysis of microdetri tus and Apalachicola River flow by season 31

23 Average seasonal variation in phytoplankton productivity for the

Apalachicola estuary 36

24 Monthly averages of daily litterfall on intensive transect plots across

the Apalachicola wetlands 38

25 Tentative model of microbial interactions with various physical and

biological processes in the Apalachicola River estuary 42

26 Seasonal distribution of total zooplankton biomass in the Apalachicola

estuary and associated coastal areas during 1974 45

27 Summed numerical abundance and number of species of benthic

infauna and epibenthic fishes and invertebrates taken in the

Apalachicola estuary 59

28 Life cycle of the blue crab along the gulf coast of Florida 65

29 Average monthly distribution of anchovies in the Apalachicola estuary

from 1972 through 1979 69

30 Average monthly distribution of croaker in the Apalachicola estuary

from 1972 through 1979 70

31 Average monthly distribution of sand seatrout in the Apalachicola

estuary from 1972 through 1979 71

32 Average monthly distribution of spot in the Apalachicola estuary

from 1972 through 1979 72

33 Average monthly distribution of penaeid shrimp in the Apalachicola

estuary from 1972 through 1979 73

34 Average monthly distribution of blue crabs in the Apalachicola

estuary from 1972 through 1979 74

35 Numerical abundance and species richness of invertebrates taken

in leaf-litter baskets at various permanent sampling sites in

the Apalachicola estuary 78

36 Regression of numbers of species of litter-associated macroinvertebrates

on salinity at three stations in the Apalachicola estuary 78

viii
Number Page

37 Simplified feeding associations of four dominant fishes (bay anchovy,

sand seatrout, Atlantic croaker, spot) and blue crabs in the

Apalachicola estuary 85

3S Generalised simplified model of seasonal relationships of the dominant

macroinvertebraes and fishes in the Apalachicola Bay system 86

39 Long-term fluctuations of squid abundance, salinity and temperature

taken in the Apalachicola estuary from June 1972 through March 1979 91

40 Monthly frequencies of blue crabs and variations in key physico-chemical

parameters at the 10 day-time stations in the Apalachicola estuary

frcif March 1972 thi . :h March 1978 92

41 Long-term abundance patterns in the dominant trawlable fish populations

in the Apalachicola estuary from March 1972 through February 4, 1982 95

42 Relative importance of four dominant species of invertebrates and

fishes taken in the Apalachicola Bay System from March 1972 through

February 1975 96

43 Temporal arsociations of fishes taken in Apalachicola estuary from

March 1972 to February 1976 98

44 Dredge spoil bank along the Apalachicola River 100

45 Ditching and diking associated with agricultural activities in the

lower Apalachicola floodplain 105

46 The extent of :. Mng by agricultural interests along the western

bank of the lower Apalachicola River 105

47 Portions of St. George Island showing housing development on the

Gulf side and dredging on the bay side 106

48 Major public investiments and specially designated areas in the

Apalachicola basin 108

49 Boundaries of the Apalachicola River and Bay Estuarine Sanctuary

with inclusion of real and proposed purchases according to the

Environmentally Endangered Land (EEL) Program (state) and

current federal holdings 110

IX
TABLES

Number Page

1 Distribution and area of major bodies of water along the coast of

Franklin County (north Florida) with relative area of oysters,

grassbeds, and contiguous marshes 15

2 Bottom salinities at stations in the Apalachicola estuary 17

3 Terrestrial habitats and land-use patterns in the immediate watershed

of the Apalachicola Bay system 20

4A Tree species found within the Apalachicola floodplain 21

4B Areas of each mapping category for five reaches of the

Apalachicola River 22

5 Linear regression of total microdetri tus and river flow by month/year by season (August 1975-April 1980) 30

6 Net above-ground primary production of marsh plants in various

salt marshes 32

7 Presence/absence information for net phytoplankton taken from the

Apalachicola estuary by month from October 1972 through September 1973 .... 33

8 Physical, chemical, and productivity data taken from locations along the

northwest gulf coast of Florida 37

9 Total annual net productivity and net input to the Apalachicola estuary

and the Apalachicola Bay system 38

10 Nutrient yields for various drainage areas in the ApalachicolaChattahoochee-Fl int River system 39

11 Nutrient values for stations in the Apalachicola estuary and River 40

12 Distribution of the major zooplankton groups in the Apalachicola estuary

and associated coastal areas 44

13 Pearson correlation coefficients for significant zooplankton relationships

in East Bay, Apalachicola Bay, and coastal areas 46

14 Distribution of ichthyoplankton in the Apalachicola estuary as indicated

by the presence of eggs and larvae 47

X
Number Page

15 Numbers of ichthyoplankton taken at various stations within the

Apalachicola estuary 48

16 Invertebrates taken in cores, leaf-baskets, dredge nets, and otter

trawls in the Apalachicola Bay system (1975-1983) 50

17 General abundance information and natural history notes for the

dominant organisms in the Apalachicola estuary 56

18 Fishes and invertebrates commonly taken with seines in oligohaline and

mesohaline marshes of the Apalachicola estuary 61

19 Epibenthic fishes and invertebrates in the Apalachicola estuary from

1972 through 1982 63

20 Epibenthic fishes and invertebrates in the Apalachicola estuary from

June 1972 to May 1977 66

21 Factor analysis of physico-chemical variables in the Apalachicola system

taken monthly from March 1972 to February 1976 81

22 Correlation coefficients of linear regressions of nitrate,

orthophosphate, silicate, and ammonia on salinity 82

23 Results of a stepwise regression analysis of various independent

parameters and species (population) occurrence in the Apalachicola

estuary from March 1972 to February 1975 84

24 Parametric and nonparametric correlations of seasonal variations of

blue crab frequencies and abiotic variables 93

25 Multiple stepwise regression of seasonal variations of frequencies of

blue crabs of three size groups and selected abiotic variables 93

26 Land use inventory of the Apalachicola River basin 102

27 Approximate dimensions of selected estuarine systems 113

28 Estimates of particulate primary production in various estuaries in

the Uni ted States 113

29 Approximate land use distribution and population density surrounding the

estuarine study areas 114

30A Approximate annual input from land drainage and point source discharge

of dissolved inorganic nitrogen per unit area and per unit volume in

various estuaries 115

308 Approximate annual input from land drainage and point source discharges

of dissolved inorganic phosphate per unit area and per unit volume in

the study areas 116

31 Total numbers of fishes per trawl sample taken at permanent stations in

the Apalachicola estuary, the Econfina estuary, and the Fenholloway

estuary 117

xi
CONVERSION TABLE
ACKNOWLEDGMENTS

The research on which this paper is

based heqan as a modest monitoring proiect

in Apalachicola Bay in March 197?. Since

that time, more than 1000 peoole--

scientists, research aides, graduate and

undergraduate students, and professional

staff peoole--have participated in a series of projects carried out within a broad spectrum of disciplines. The

research effort has included chemistry,

hydrological engineering, physical

oceanography, biology, geology, geography,

fisheries, computer programming,

statistics, resource planning and

management, and economics. Many of the

data have been retained and organized into

a series of computer files, which T am currently holding at the Florida State

Uniyersity Computer Center. A complete

list of this information is given in the

appendices to this paper.

Although funding for this program has come from various sources, the maior

contributions have been made by the

Florida Sea Grant College (National

Oceanic and Atmospheric Administration)

and the Franklin County Board o^" Commissioners. Supplementary funds have

been provided by private industry and

state and federal agencies. The list

includes local developers, forestry

interests, the Florida Department of

Environmental Regulation, the Florida

Department of Community Affairs, the

Coastal Plains Regional Commission, the

U.S. Environmental Protection Agency, the

National Science Foundation, the Florida

Department of Natural Resources, the

Northwest Florida Water Management

District, the U.S. Geological Survey, the

Florida Game and Fresh Water Fish

Commission, the U.S. Fish and Wildlife

Service, and the Man in the Biosphere

Program of the U.S. Department of State.

Special credit should be given to the

Department of Biological Science (Florida

State University) for its long-running

support of the research. It is somehow

consistent that the main impetus for the

research effort has come from local concerns (the fishermen of Franklin

County, i^lorUa) and a federal agency (the

Florida Sea Grant College, NCAA) that has

always sought to apply basic scientific

knowledge to practical problems. The

people of Franklin County, depending on the sea for their livelihood, recognized

early that, as land development

accelerates in Florida, a forward-looking

management program will be necessary to protect the resource that has been at the

center of their way of life for

generations. The combination of basic and

applied science, local, state, and federal

involvement, and a multidisciplinary,

long-term research program has led to a series of resource management/planning

actions that are unprecedented in the

nation.

xm

CHAPTER 1 INTRODUCTION (HISTORICAL PERSPECTIVE AND OVERVIEW)

1.1. GEOGRAPHIC SETTING AND

CLASSIFICATION

The ADalachicola estuary (Fioures

1-3) is part of a tri-river system that

includes the Apalachicola River in Florida

and the Chattahoochee and Flint Rivers in

Georgia and Alabama. The Chattahoochee

River originates at the base of the

Appalachian Mountains in the Piedmont

upland, and traverses three geologic

provinces: the Piedmont, the Appalachian,

and the Coastal Plain. The Flint River

begins in the lower Piedmont Plateau just

north of the fall line and flows through

the Coastal Plain.

The Apalachicola-Chattahoochee-Flint

(ACF) drain_age basin includes an estimated

48,484 km"

Georgia,

northern Florida (Figure 1). The

Chattahoochee River drains approximately

-^ (19,^00 mi2) in western

southeastern Alabama, and

?1,840 km^ (8,650 mi^) and the Flint River

drains an estimated ?1,444 km^ (8,4P4

mi^). The Jim Woodruff dam, which forms

Lake Seminole at the confluence of the

Flint and Chattahoochee rivers,

constitutes the headwaters of the

Apalachicola River. The Apalachicola

River is approximately 171 km (108 mi)

long, with a fairly uniform slope of 0.15

m/km (0.5 ft/mi); it falls approximately

12 m in its course from Lake Seminole to the Gulf of Mexico. The Apalachicola

River drains an area of about 2,500 km^

(1,030 mi2). The Chipola River, which

ioins the Apalachicola River near its

southern terminus (Figure 1), has a watershed equal to that of the

Apalachicola. About 3% of the ACF basin

is in the Blue Ridge mountains, 38% in the

Piedmont Plateau, and 59% in the coastal

plain below the fall line (Figure 2). The

lower coastal plain is nearly flat, with

extensive wetlands development.

en.'!""""

TRI-RIVER

SYSTEM

jkj y-
ChoMohoochee River COAST

Figure 1. Location of the tri-river

drainage system in the southeastern United

States showing the relative positions of

upland features and the Apalachicola

estuary.

A detailed review of the dimensions

of the Apalachicola Bay system (^QOis'n to ?9055'N; 84O20'W to 85O?0'W) (Figure ?) is

given by Livingston (1980a). This system

is composed of six major subdivisions:

3,Q81 ha (9,837 acres)

?0,959 ha

East Bay

Apalachicola Bay

('51,792 acres)

St. Vincent Sound 5, "^40 ha

(13,689 acres)

West St. George Sound (to Dog Island)

14,747 ha (36,440 acres)

East St. George Sound

16,016 ha (39,576 acres)

Alligator Harbor 1,637 ha

(4,045 acres)

The entire area totals 6?, 879 ha (155,374

acres). A natural shoal forms a submerged

boundary between Apalachicola Bay and

St. George Sound. The bay is bounded on

its extreme southern end by three barrier

islands: St. Vincent, St. George, and Dog

Island. There are four natural openings

to the gulf: Indian Pass, West Pass, East

Pass, and a pass between Dog Island and

Alligator Harbor. A man-made opening

(Sike's Cut) was established in the

western portion of St. George Island. The

3.6-m- (12-ft-) deep Intracoastal Waterway

extends northwestward from St. George

Sound through Apalachicola Bay, up the

Apalachicola River to Lake Wimico and then

along an artificial channel to St. Andrews

Bay to the west.

The Apalachicola estuary is a lagoon

and barrier island complex. It has been

classified as a shallow coastal plain

estuary oriented in an east-west direction

(Dawson 1^55). Because of the placement

of the barrier island complex, it coul'^ be

called a coastal lagoon. The average

depth is between 7 and 3 m at mean low

tide (Gorsline 1^163).

In terms of Pritchard's (1^67)

estuarine classification scheme, the

Apalachicola Bay system is a widthdominated estuary controlled by lunar

tides and wind currents. As such, it is a type D estuary (Conner et al. 1981) in

that it is dominated by physical forces

(i.e., tidal currents, wind) as a function

of its shallow depths. As a result, the

bay system is relatively well mixed,

although various portions of the estuary

are periodically (seasonally) stratified

(Livingston 1984a).

1.^. DRIVING FORCES AND HUMAN INFLUENCE

The principal driving forces that

determine the habitat structure and

biological processes of the estuary are river flow, physiography of the basin,

seasonal changes of nutrients, and

salinity as modified by wind, tidal

influences, and freshwater inflows. Tidal

influence extends approximately 40 km (?5

miles) up the river. As a biological

entity (Odum et al. 1'374),

(which includes East Bay,

Bay, St. Vincent Sound,

portions of St. George

characterized by upland marshes that grade

into soft-sediment areas, vegetated

shallow bottoms, and oyster reefs. The

oligohaline East Bay merges with

mesohaline and oolyhaline portions of

Apalachicola Bay, St. Vincent Sound, and

St. George Sound.

The Apalachicola River, the largest

in Florida in terms of flow, is the

principal source of fresh water to the

estuary. The average flow rate is about

the estuary

Apalachicola

and western

Sound), is
P;65 m3 sec-l (23, '^00 ft^ sec'l) measured

at Blountstown, Florida. Maximum and

minimum discharqes over the pa<;t 1? years

are 4,600 m^ sec"! (162,^^00 cfs) and 178

m^ sec"l (6,280 cfs), respectively. The

river and, secondarily, local rainfall

determine the distribution o^" salinity in

the estuary. The placement of the barrier

islands also has a maior influence on the

salinity reqime of the estuary (Livinqston

1^79, iq84a). The islands limit the

outflow of the low-sal initv water to the

outer Gul-P of Mexico.

The Aoalachicola basin occupies the

last sparsely inhabited and undeveloped

drainaqe system and coastal reqion in

Florida (Livingston lQ83a, b, c).

Franklin County, with a population of only

8,403 in 197Q, encompasses the lower river

and bay system. Forested uplands,

wetlands, and aquatic habitats comprise

most of the land area in Franklin County.

The local economy is based larqely on the

sport and commercial fisheries of the

Apalacbicola River and Ray system.

According to recent estimates (Florida

Department of Administration l'^77),

commercial fishinq, recreation, forestry

and timber processing, aqriculture, and

light manuf acturinq characterize the

regional economv of the entire

Apalacbicola basin. The human population

of the six counties along the river has

grown slowly since li^GO, increasing only

7% (from 101,782 to 10Q,?S4) from 1%^ to

1974. State government is a major

employer in the region, while industrial

or commercial land use is confined to only

0.2% of the basin area.

The Apalacbicola drainage system is one of the least polluted in the country

(Livingston 1974a, b, iq77a-d, 1Q78, 1979,

1980a-c; Livingston and Thompson 1975;

Livingston and Duncan 1979; Livinqston et

al. 1974, 1976a, b, 1Q77, 1Q78_).

Some

problems, however, have emerged in recent

years (Livinqston l'583d).

CAPE SAN BLAS

Figure 3. Detailed features of the Apalachicola Bay system including the major contributing drainages, the barrier island complex, and the major passes in the bay complex.
1, A 13,352-ha (33,000-acre) cattle

ranch was established in the Apalachicola

River floodplain about '^-10 km (6 mi)

Much of the area was

and drained, while waste

over the dikes into the

The potential impact of

above the bay.

cleared, ditched,

water was pumped

river system.

this operation is under study and review,

although farming has continued, and water

quality has deteriorated in some of the

upland creeks.

?. Portions of the drainage system

have historically been subjected to forestry operations, which include

ditching, draining, clearcutting, and

reforestation. These activities have been

associated with local changes in water

quality and short-term adverse effects on aquatic biological associations

(Livingston 1^78). A long-term

mul tidiscipl inary study has iust been

completed by the Florida Sea Grant College

(Livingston lQ83c) along with proposed

management practices which are designed to

mitigate adverse impacts.

3. Recent population increases along

the north Florida coast have stressed

regional coastal counties in terms of

municipal development, sewage disposal,

and storm water runoff (Livingston lP83d).

The recognition of such potential impact

has led to the development of relatively

advanced local land use plans such as that

adopted by Franklin County in 1P81

(Livingston lP80a, b, iq83c).

Implementation of the comprehensive plan

has not been carried out, however. During

1^8^, sewage spills closed down the

Apalachicola oyster industry for prolonged

periods. Meanwhile, proposals to bring

high-density construction projects to coastal areas of Franklin Count v have

proliferated.

4. A continuing problem in the

region involves proposals to either

channelize or dam the Apalachicola River

to make a corridor for barge traffic and

industrial development. These

developments would serve as a north-south

link between upriver ports on the

Chattahoochee and Flint Rivers in Alabama

and Georgia and the Gulf of Mexico.

Authorization for a maintained channel

(30. B m or 100 ft wide, ?.7 m or 8.8 ft

deep) by the U.S. Army Corps of Engineers

(USAGE) was part of the amended Rivers and

Harbor Act of 1^46. A system of 13 dams

is already in place on the Chattahoochee

River and three dams are currently in use

on the Flint River (Figure 4). Associated

with these activities are a series of

barge terminal facilities and offloading

systems. Rock outcrops in the

Apalachicola River have been removed as part of ongoing, extensive dredging and

channelization of the river. Superimposed

over these activities is the increasing

municipal water use in areas such as Atlanta, Georgia, where sustained

population growth could reduce water flow

in the tri -river system in the near future.

Federal a Other

Figure 4. Distribution

along the tri-river

information provided by

Corps of Engineers).

of impoundments

system (after

the U.S. Army
5. Past studies on pesticide

distribution in the estuary (Livingston

and Thompson 1^175; Livingston et al. 1<578)

have indicated relatively low levels of

organochlorine contamination in the

Apalachicola Bay system by the mid lQ70's.

Winger et al. (1984) found that biota from

the Apalachicola River had moderately high

levels of total DDT, total PCB's, and

toxaphene in 1978. Animals from the upper

river had higher organic residues than

those taken in the lower river. Such

levels exceeded recommended permissible

levels for the protection of aquatic life. A recent review of the heavy-metal

distribution (Livingston 1^83d; Livingston

et al. 1982) indicates local increases of

metals in the sediments and biota of Lake

Seminole, parts of the Chipola drainage,

and areas in the bay system that receive

municipal runoff. These increases are due

to local point sources such as battery

recycling operations (upper Chipola),

industrial sources in Georgia, marinas,

and municipal outfalls. Winger et al.

(1984) found metal residues in riverine

organisms generally below 1 pg g-^. A recent analysis of data on long-term

monitoring of the metal concentrations in

oysters (Crassostrea virginica ) in the

estuary (Florida D¥partment of Natural

Resources, personal communication)

indicates no undue increases of such

metals in shellfish over the past decade.

^. Dredging and spoil placement take

place in the Apalachicola River and Rav

system (Livingston 1984a). These

operations are being reviewed bv the

Florida Department of Environmental

Regulation (S. Leitman et al. 1982). The

immediate impact of long-term dredge and

spoil activities on the estuary is given

by Livingston (1984a).

In summary, the Apalachicola drainage

basin is currently lightly populated with

an economic system dominated by renewable

natural resources. However, over the next

few decades, the essential Iv rural economy

will probably give way to more energydependent industrial and urban

development, which might lead to increased

stress on the natural system due to growing population pressure, residential

development, agricultural activities,

toxic waste disposal, erosion and

sedimentation, and alteration of the

physical structure of the drainage basin.
CHAPTER 2

ENVIRONMENTAL SETTING

?.l. ORIRIN AND EVOLUTION OF THE ESTUARY

?,1.1. Geological Time Frame

The Dhysiograohic structures of most

estuaries are ephemeral in terms of

geological time. Climatological forces

are continuously at work shaping and

reshaping the basin features.

Characteristics of the Apalachicola

estuary are dependent on the interaction

of an upland drainage system wi^h offshore

marine conditions. The estuary is, in

effect, an extension of the upland river

or drainage area, and its origin and

evolution are inextricably linked to the

dynamic geological history of the land/sea

interaction.

approximately '^?.^ km (14 mi) northeast of

Apalachicola, Florida. These islands were

located in what is now the Tate's Hell

Swamp (Figure 1). The general dimensions

of the Apalachicola vallev as we see them

today were established in the Pleistocene.

The maior drainages of the Florida

panhandle (which includes the Apalachicola

drainage system) are alluvial in that they

carry sediment loads that eventually end

UD in the coastal estuaries (Figures 1,

^). The geological structure of the

Apalachicola River estuary is of Recent

and Pleistocene origin. Marine sediments

comprise a maior physical feature of the

region. The Apalachicola estuary is

bounded by well-developed beach-ridge

The Aoalachicola River is the only

drainage area in Florida that has its

origin in the Piedmont, which, as will be

explained later, is of biological

importance to the region. The geological

history of this area is well known in

general terms. Rv the Cretaceous period

(about 135 million years ago), most of the

tri -river valley was submerged under

ancient seas (Tanner 19fi2). The origin of

the Apalachicola River or its antecedents

occurred some time in the Miocene epoch

about 25 million years ago (W. F. Tanner,

Florida State University, pers. comm.).

There has been a gradual decline in sea

level through Cenozoic time (70 million

years ago to present); sea level has

dropped an estimated 70-100 m from the

middle of the Miocene (Tanner 1968).

Olsen (1^68) gives evidence that the uoper

Apalachicola River basin (the area around

Blountstown, Florida; Figure 1) was a deltaic or coastal environment during the

Miocene. By the Pleistocene epoch (1

million years ago), there was evidence of

an arcuate chain o^ barrier islands

Figure 5. The Apalachicola estuary with

details of upland drainage areas and the

placement of permanent sampling sites for

the long-term field studies of the Florida

State University research team (after

Livingston et al. 1974).
plains of late Holocene oriqin (Fernald

1Q81). The linear, qentlv curving beach

ridges of the area attest to the changes

in orientation of the estuary through

geological time in resoonse to wide

fluctuations of sea level. The

Apalachicola estuary is part of a broad,

sandv shore plain, which is constantly

being changed by a combination of

climatological elements such as wind,

rainfall and sea level alterations. The

present structure of the bay is around

10,000 years old (Tanner 1^83). Sea level

reached its modern position about SOOO

years ago when the construction of the

present barrier island chain was underway.

Exceot for the southward migration of the

delta front, the general outline of the

bay system was established at this time

(Tanner 1^83).

?.!.?. Oeomorphology and Regional Geology

a. Upland areas . The maior

formations in the upper Chattahoochee

River system are underlain by igneous

rocks and crystalline schists. The area

is characterized by Tertiarv limestone

outcroppinqs, which add to the habitat

diversity of the region (Figure 6). The

lower division of Piedmont upland, defined

as the Opelika Plateau, is underlain by

Archean (i.e., Precambrian) rocks.

Tributaries of the Ohattahoochee River

have subsequently eroded these formations

with some valleys cut approximately 6? m

(?00 ft) below the general surface. The

rocks of the Appalachian province pass

under the Coastal Plain formations. Along

the border between the Appalachian

province and the Coastal Plain,

Appalachian rocks are overlain by

Cretaceous formations, more deeply buried

Quaternary sediments further north

Coastal Plain is covered with a

These rocks

by Tertiary

are

and

The

thick

layer of clastic (erosion produced)

sediments as well as limestone

(nonclastic) sediments, some of which mav

be crystalline.

Adams et al. (1'526) have presented a detailed account of the Paleozoic,

Mesozoic, and Cenozoic formations in

Alabama, which is generally applicable to the Apalachicola valley. The Cenozoic

formations are confined to the Coastal

Plain and represent deposits at the bottom

of an ancient sea, which consist of sand,

clay, mud, or calcareous ooze. Fossil

marine mollusks and echinoderms are interspersed with remnants of fossil

plants from flood plains, marshes, and

swamps. Pleistocene marine sands and

clays overlie older formations along the

coast, and estuarine and fluvial deposits

extend up the main river valley. Swamps

immediately upland of the Apalachicola

estuary are underlain by quartz sand

(Brenneman and Tanner 1958).

Figure 6. Geological features of the

Apalachicola drainage system showing (A) a line north and west of which there are thin patches of Tertiary limestone near

the land surface and (B) a line beyond

which the limestone thickens and is more deeply buried. The top of the Tertiary

limestone is shown in feet below sea level, while Tertiary limestone that occurs in or near the land surface is also

outlined (modified from Means 1977).
The coastal geomorpholoqy of the

Apalachicola reqion is extremeiv complex;

major features are developed from wind and

current modified beach ridqes (Clewell

1977). These formations are complicated

by considerable Pleistocene sea-level

fluctuations. The northern qulf coastal

lowlands are dominated by Pliocene epoch

marine sands. The flood plain of Holocene

(recent) sediment reaches depths

approximating 24.3 m (80 ft) near the

river mouth and 13.7 m (45 ft) near Blountstown, Florida (Fiqure 1). These

sediments lie directly on Miocene strata

because much of the Pliocene and

Pleistocene sediments were eroded during

periods when sea level was lower and river

flow was greater. The sea level

approximately 20,000 years ago was over

125 m (410 ft) lower than that found

today, and the coastline was considerably

seaward of its current position.

The Florida panhandle is an uneven platform of carbonate bedrock (limestone

with dolomite) overlain by one or more

layers of less consolidated elastics

(Fiqure 6, Puri and Vernon 1«564; Clewell

1^78). Superficial strata are of Eocene,

Oligocene or early Miocene origin.

Considerable solution activity has led to the formation of sinks, caves and other

karst features (Means 1977). The elastics

consist of Fuller's earth (primarily the

clays montmoril linite and attapulqite),

phosphatic matrix, sand, silt, clay, shell

marl, qravel, rock fragments, and fossil

remains. The elastics with shell marl are sediments of ancient shallow seas and

estuaries. Various clastic strata were deposited during the early Miocene, while

others were fluvial and aeolian deoosits

or sediments in lake bottoms. These

elastics form terraces sloping toward the

Gulf. Such terraces are altered by

erosion and dissection by streams and

rivers. In spite of various

post-Pleistocene sea-level fluctuations,

elevations in this area have changed less

than 10 m as a result o^ erosion,

deposition, and sedimentation. Dunes,

spits, bars, and beach ridqes became

stranded inland as the sea receded.

Soils and sediments. The

Apalachicola River floodplain lies wholly

within the Florida Coastal Plain and is in

contact with Tamoa Limestone (early

Miocene). The river just below the Jim

Woodruff Dam flows through the Citronelle

formation (Pliocene) that borders the

western edge of the Pleistocene bed from

16 to 20 km below the dam to Blountstown.

The eastern portion of the river is

influenced by the Hawthorn formation

(Fuller's earth and phosphatic limestone)

and Duplin marl (sandy marine and clayey,

micaceous shell marl). The clays in

particular and fine sands cause considerable turbidity. The river bed is

composed primarily of remnants of

Pleistocene deposits (sand to coarse

gravel) that are covered by fine clay

sediments. The lower river valley is

composed largely of Plio-Pleistocene

marine sands, which lie over the Aucilla

Karst Plain, the Jackson Bluff formation,

and the lower part of the Citronelle

formation.

Upland soil composition reflects the

geological history of the Apalachicola

valley. Soils in the titi swamps and

savannahs of the Apalachicola National

Forest are strongly acidic and low in

extractable cations (Mooney and Patrick

1915; Coultas 1976, 1C)77, IQRO). Total

phosphorus is low in all soils of the

basin. Cypress and gum swamps are also

highly acidic and low in extractable

bases, while more alluvial soils are less

acidic. Estuarine marsh soils are relatively hiqh in organic matter, especially

at the river mouth. These soils are derived largely from the erosion of the

northern Piedmont-Appalachian soils, which

have been deposited on the sea floor and,

at times, have been uplifted above sea

level. Floodplain soils are composed of a broad range of textures and colors. They

are predominantly clay with some silty

clay and minor clay loams (Leitman, 1978).

Point bars in the river bed are composed

largely of fine and very fine sands.

Soils in wetlands directly associated

with the Apalachicola River have been

analyzed. Swamp soils are wet, moderately

acidic, high in clay content, and low in

salinity (Coultas in press). The principal clay-sized minerals include kaolinite,

vermiculite, quartz, and mica. These

areas are poorly drained and contain

considerable amounts of clay and organic

matter. The soils are formed from recent

accumulations of sediments deposited in
stream channels and estuarine meanders.

The pH values ranqe from 4.^ to f^.6.

Studies of the marshes above East Bay

(Coultas 1^80; Coultas and Gross 197^^)

indicate that the deltaic soils are slightly acidic and become alkaline with

depth. The dense mats of roots and

rhizomes from the predominant sawgrass

( Cladium .iamai cense ) and needlerush

( Juncus roemerianus ) along the eastern

portions of the estuary tend to hold the

soils in place. The soils are composed of

thin organic deposits mixed with clay and

overlie loamy sands of fine-textured

materials. Considerable amounts of silt occur in some soils, and most have poor

load-bearing capacity because of the high

organic content and high field moisture

levels. Vegetation differences are attributed to soil salt content. Sawgrass

is dominant in areas most affected by

river flow (i.e., with low salinity), and

needlerush is predominant in tidal areas

(i.e., those with higher salinity)

(Coultas 1P80).

Sediments in the estuary are characterized bv mixtures of sand, silt,

and shell components (Livingston 1^78).

Present sediments are accumulating over tertiary limestones and marls that outcrop

in the scoured central channels of West

Pass and Indian Pass. St. Vincent Sound

and northern portions of Apalachicola Bay

are silty areas that grade into sand/silt

and shell gravel toward St. George Island.

The thickness of these sediments (10-^0 m)

(Gorsline 1^63) may be the result of

erosion of older deltaic deposits during

periods of higher sea level. East Ray is

composed of silty sand and sandy shell.

Areas near the river mouth have varying

quantities of woody debris and leaf

matter, especially during winter and

spring months of heavy river flooding

(Livingston et al. 1976a). The floor of

the bay is thus formed largely of quartz

sand with a thin (but varying) cover of

silt, clay, and debris depending on the

proximity to land runoff.

The estuarine sediments originated in

the southern Appalachians and have

undergone a complex history of deposition

and reworking in the coastal plain

deposits, coastal marshes, beaches, and

dunes. Fine sediments flow out of the bay

into the Gulf of Mexico while sand is

moved by tidal currents within the bay and

at the mouths of the western inlets. The

cusp of the Aoalachicola Bay coastline has

been built by river sediments deposited

during Tertiary and Pleistocene times with

modification bv waves and long-shore

drift. Puri and Vernon (19fi4) and Clewell

(1978) have made a detailed review of the

geological formations and soil

distribution in the region.

? . 1 . 3 . Watershed Characterization

Numerous physiographic, geological,

and bioqeographic features contribute to the biotic richness of the Apalachicola

drainage system (Clewell 1^77; Means

1977). While the Apalachicola basin

(Figure 7) lies entirely within the

Coastal Plain, it is subdivided into upper

and lower regions; the Marianna lowlands.

New Hope Ridge, Tallahassee Hills and

MARIANNA

LOWLANDS

WESTERN

RED HIL

GRAND

RIDGE

|><^PALACHICOLA ' ^-RAVINES

RIVER

BOTTOM

LANDS

/ APALACHICOLA

LOWLANDS

\ COASTAL

^ ^ MARSHES

OFFSHORE

AREAS

Figure 7. Natural areas Apalachicola basin based

physiography, vegetation types,

geography, and distribution of

(after Means 1977).

of the

on the

regional

organisms
Beacon Slope are part of the Gulf-Atlantic

rollinq plain, while the lower coastal

lowlands are part of the Gulf-Atlantic

Coastal Flats (H, M. Leitman et al. l^^S?).

The drainaqe system contains streams of

various types, which range from firstorder ravine streams (Means 1*^77) to the

hiqher order low-qradient, meanderinq

types. The latter contain hiqh orqanic

acid levels in the flatwoods or are calcareous and clear in the Marianna

Lowlands karst plain. Extensive lake

systems are lacking in the valley;

Ocheesee Pond is located in an abandoned

bed of the Apalachicola River, and two

other natural lakes (Lake Wimico, Dean

Lake) occur in the basin. The upper river

region, cutting through Miocene sediments,

has a flood plain 1.^-3 km (O.P-l.q mi)

wide. This floodplain widens to 3-5 km

(1.9-3.1 mi) along middle portions of the

river, with the lower river having the

widest floodplain (7 km; 4.4 mi). The

upstream tidal influence in the floodplain

does not extend above km 40 (mi PS). The

Chipola River joins the Apalachicola at km

45 (river mi ?8). The delta is about 16

km (in mi) wide and is surrounded by a broad marsh.

The previously described geological

processes have led to hiqh physical

diversity of the land forms in the

Apalachicola basin. "Steepheads" or amphitheatre-shaped valley heads with very

steep walls (Means 1^77) occur in small

drainages that dissect the eastern

escarpment between Bristol and Torreya

State Park within a narrow east-to-west

alignment through the Florida panhandle.

These constant environments are important

habitats for various species. The

Apalachicola Ravines (Figure 7) (Hubbell

et al. 1956) are drainages that form

another unique habitat associated with the

river basin. These ravines include smallorder stream bottoms and steep valley

slopes; the vegetation qrades upward from

hydric plant communities near the bottom

to xeric vegetation at the top of small

divides between ravines. The Marianna

lowlands form a karst plain containing

more vadose (i.e., above water table) cave ecosystems than any other part of the

coastal plain (Means 1977). The

Aoalachicola lowlands, a flatwoods region

with little relief, is a low, sliahtlv

inclined plain with extensive swamplands.

The eastern portion of the Aoalachicola

lowlands contains parts of the Tate's Hell

Swamp, which is undergoing extensive

changes due to forestry operations. The

western lowlands are part of a cattle

ranch and farming operation. The Western

Red Hills are separated from the other

natural areas by the Chipola River valley.

This area is high in elevation but not as deeply dissected as the Apalachicola

Ravines. Grand Ridge (Figure 7) is a wedge-shaped area bounded by the Chipola

and Apalachicola Rivers. While originally

part of the same upland mass that extended

from the Apalachicola Ravines westward.

Grand Ridge has been eroded. This area is

associated with springs, caves, and

troglodyte (i.e., subterranean) fauna.

The river bottomlands represent a floodplain habitat characterized by the

river channel, sloughs, swamps and

backwaters, and the periodically flooded

lowlands. Many springs and aquatic cave systems empty directly into the river

bottomlands.

?.1.4, Barrier Islands

At the mouth of the Apalachicola

River is a well developed barrier-island

system composed of three islands (St.

Vincent, St. George, Dog) (Figure 3).

These islands roughly parallel the

coastline and are characterized bv sets of

sand dunes of differing geological ages.

While the shore system is based on dunes

that date back some 3000 to 6000 years,

the barrier islands are no older than 3000

years. They consist of quartz sand that

has been transported from the southern

Appalachian Piedmont by the river system

and that currently rests on an eroded

Pleistocene surface (Zeh 1980). On St.

Vincent Island, for example, qently

curvinq lines of beach ridges (Figure 8)

up to 1 m (3 ft) hiqh serve as the base

for small dunes; such ridqes represent the

geological history of sand deposition in

the region, with the oldest (northernmost)

ridqes indicatinq where sea level achieved

its earliest position.

St. Georqe Island is about 48 km (30

mi) lonq and averaqes less than 0.5 km

(1/3 mi) in width. It consists of ?,Q73

ha (7,340 acres) of land and 486 ha (1,?00

acres) of marshes. The medium to fine

grain sands provide for relatively poor

10
Figure

Island.

Aerial view of St. Vincent

aquifer conditions; all fresh water is

derived from rainfall. Silty clav

sediments at depths between 7.6 and P.? m

(25-30 ft) below the sandy surface create

an impermeable barrier to separate rainderived fresh water from the surrounding

salt water. There is a shallow lens of

fresh water beneath the island. Some of

this fresh water, modified by

transpiration and evaporation, is

eventually discharged into the Gulf and

lagoonal marine systems.

?.2. CLIMATE

2.2.1. Temperature

The climate in the Apalachicola basin

is mild, with a mean annual temperature of

20° C (68° F). Temperature varies with

elevation and proximity to the coast. The mean annual number of days with

temperatures at or below freezing is 20 at Lake Seminole and 5 along the Gulf Coast

(National Oceanic and" Atmospheric

Administration, unpublished data; Clewell

1°77). Livingston (unpublished

manuscript), working with long-term

(40-year) climatological data, found that

temperatures usually peak in August with

lows from December to February, at which

time monthly variance is maximal. While

peak summer temperatures are similar from

year to year, winter minima vary. A timeseries (spectral) analysis indicates that

there is a long-term period of recurring

low winter temperatures of 118 months (P. 8

Precipitation

Mean annual rainfall in the

Aoalachicola River basin is approximately

150 cm (59 inches). There are, however,

considerable local differences in monthly

precipitation totals. In the Apalachicola

delta, areas west of the river receive

almost one-third less rainfall than those

east of the river (i.e., Tate's Hell

Swamp). Rainfall in the Georgia portion

of the watershed is 130 cm/yr (51

inches/yr).

The rainfall patterns of Florida and

Georgia (Figure 9; Meeter et al. 1979) are basically similar exceot for the timing of

rainfall peaks. Georgia rainfall has two
peaks: one in March and another of equal

tnaqnitude in July. The Florida rainfall

peak in March is not as great as that of

Georgia, but the primary difference is the

much larger, sustained rainfall peak in summer and early fall in Florida. In both

areas, there are drought periods during

mid to late fall. Spectral analysis of

long-term trends (Figure 10) indicate

that, while rainfall is highly variable,

there are certain long-term trends.

Florida (Apal achicola) rainfall has

RO-month ('i."'-yr) cycles in peak reoccurrence, while Georgia rainfall has a slightly different spectrum.

2.^.3. Wind

Wind direction is predominantly from

the southeast during the spring

(March-May) and southwest to west during

the summer (June-August). Winds come from

the north or northeast during the rest of

the year. However, analysis of long-term

wind data indicates that there is wide

variability of wind velocity and direction

over the Apalachicola watershed at any

given time. Tn the shallow estuary, winds

can cause rapid changes in the normal

tidal current patterns. Southerly winds

tend to augment astronomical tides and

UJo

< 560

a:u

>

<

CO (5 400-

"I

o

2 240-

z

o

«>

80-

RIVER FLOW

1920

I 1925 1930

I I 1935 1940

-400

LJ<

ir

LlI >

<

300 CP

200

-100

T T T

1945 1950 1955 I960 1965 —1 1970 —1 1975

O)

> o

o

liJ O

<

UJ

o

o

CD

24 -

16-

6 MONTH - 36 MONTH

RAINFALL

1920 —I1925 -| 1 \ 1930 1935 1940 -I \ 1945 1950 1955 -1 1 i960 1965 "1 1 1970 1975 YEARS

UJ

C3<

crUJ

iS) z

>

o

o

CO

fO

Figure 10. Six-month and 36-month moving averages of Apalachicola River flow

(cfs; 1920-1977) and Apalachicola rainfall (1937-1977). Data are taken from Meeter

et al. (1979).

12
cause abnormally hiqh water without the

usual ebb.

The air circulation over the Gulf of

Mexico is primarily anticvclonic (clockwise around an atmospheric high-pressure

region) during much of the year. However,

strong air masses of continental origin

often move through the northern Florida

area, especially during the winter. From

November to March, an average of 30 to ^0

polar air masses penetrate the Gulf each

year. Storms are usually formed along

slow-moving cold fronts in winter.

Tropical storms or hurricanes may occur in summer and early fall. Lesser storms

often occur as extratropical cyclones,

which tend to move across the Gulf from

west to northeast during winter oeriods

(Jordan 1973). Winter storms tend to be more pervasive in a geographic sense,

while summer storms are often intensive,

short-lived, localized events. The

likelihood of the occurrence of a hurricane in the northeast Gulf is about once every 17 years with fringe effects

about once every S years (Clewell 1^78).

The last hurricane to hit Apalachicola,

Hurricane Agnes, occurred in June 197?.

Overland (1975) showed that basin

orientation (relative to wind direction,

headlands, and marsh areas) can produce

variations in surge heights, which are responsible for much damage. Livingston

(unpublished data) found that Hurricane

Agnes had no sustained effect on water

quality or the biota o^ the Apalachicola

estuary.

''.3. HYDROLOGY

?.3.1. Freshwater Input

The Apalachicola River has the

highest flow rate (690 m^ sec"l at Chattahoochee, Florida; 1958-1^80) and

broadest flood olain (450 km? of bottomland hardwood and tupelo-cypress forests)

of any river in Florida (H. M. Leitman et

al. 1982). Apalachicola River discharge

accounts for 35^ of the total <'reshwater

runoff on the west coast of Florida

(McNulty et al. 1'572). Seasonal variation

(Figure Q) is high, with peak flows from

January through April and low flows from

September through November. The absence

of a summer river-flow peak (despite rainfall peaks in the basin at this time) may

be related to higher evapotranspiration

rates in the vegetation of the watershed

(Livingston and Loucks l'^78). A spectral

analysis using data from 1920 to 1977

(Figure 10) indicated river-flow cycles on the order of 6-7 years (Meeter et al.

1<^79). Indications of longer-term cycles were shown along with the abnormally low

river flow during the mid-1950's.

In a cross-spectral analysis of

Georgia rainfall with river flow, the two

patterns were in phase (Meeter et al . 1979; Figure °) . The analysis indicated

that the Apalachicola River flow patterns more closelv resembled cycles of Georgia

rainfall than they did those of Florida

rainfall. This pattern should be expectd

since only 11.6% of the drainage basin is

in Florida, and the remainder is in

Georgia. Stage fluctuations vary greatly

from upper to lower river with the

narrowest ranges (from peak to low) at downstream stations (H. M. Leitman et al.

1982). Such flooding patterns are essential to elements of the hydrology of

the estuary.

Floodplain inundation varies with

location on the river and reflects the

influence of natural riverbank levees

(H. M. Leitman et al. 1982). Natural

levees within the flood plain are inundated only at high stages of river

flow. The level of the water table also

depends on river stage. Fluctuations are damped by water movement through floodplain soils. The levees of the upper

river, where there is a greater range of

water fluctuation, are higher than those

in the lower river where the flood plain

is guite flat. Flood depths tend to decrease from the upper to the lower river

and rates of flow in the upper river

floodplain are generally less than those

along the middle and lower reaches of the

river. The height of the natural levees

and the size and distribution of breaks in

the levees all control the hydrological

conditions of the river flood plain. Such

hydrological conditions, in turn, control

the form and distribution of floodplain

vegetation (H. M. Leitman et al . 1982).

2,3.2. Tides and Currents

Franklin County straddles a region of

transition between the diurnal tides of

13
west Florida and the semidiurnal tides on the Gulf peninsula. Tides at Aoalachicola

are diurnal to semidiurnal, with

"uncertainties" concerninq the selection

of a "typical" tide pattern for each month

(Conner et al. I'^Sl). Tides in the

Apalachicola estuary are influenced by the

main entrances and smaller passes. Tidal

ranqes vary from 0.13 m (0.^3 ft) at Doq

Island near the eastern end of the estuary

to 0.?3 m (0.75 ft) at East Pass.

Gorsline (1963) classified this estuary as "unsymmetrical and semidiurnal except

durinq periods of stronq wind effect."

While currents in the Apalachicola estuary

are tide-dominated, they are also

dependent on local physioqraphic

conditions and wind soeed and direction

(Livinqston 1<578). River discharqe has

little influence on the hydrodynamics of

the partially stratified estuary (Conner

et al. 1°81). Shallow estuaries such as the Apalachicola are wind dominated in

terms of flushing and current movement.

The wind can be up to three times more

important than the tidal input in the

determination of current strength and

direction (Conner et al. 1981).

Net flows tend to move to the west

from St. George Sound; East Ray water

merqes with the westward flow (Fiqure 11).

West Pass appears to be a maior outlet for

the discharge of estuarine water to the

Gulf, especially when influenced by longterm or high velocity winds from the east.

Water movement through Indian Pass also occurs in a net westward direction,

although the Picoline Bar may retard

passage (Dawson 1955). Estuarine currents

Figure 11. Net water current patterns in

the Apalachicola estuary as indicated by

flow models developed by B. A. Christensen

and colleagues. (A detailed analysis of

such currents can be found in Conner et

al. (1981).)

may be affected by excessive land runoff

or high velocity winds from the east or west. Strong north to northeast winds

deflect water downwind and to the west.

Gorsline (1963) estimated a tidal

prism equal to about ?0^ of the bay water

volume, and he suggested that the

residence time of river water in the

estuary ranges from a few days to a month.

The two western passes account for over

669$ of the total bay discharqe, even though they account for only 10°4 of the

inlet area (Gorsline 1'563). The bulk of

river flow exits through these passes, and

the effects of river flow on salinity can

be felt ?65 km (165 miles) offshore in the

gulf. Tidal deltas extend seaward from

Indian Pass, West Pass, and East Pass,

indicating appreciable sediment transport

through these areas. Current velocities

in the bay rarely exceed 0.5 m sec"^,

while velocities in the passes may reach

2-3 m sec"l.

?.4. PHYSICAL/CHEMICAL HARITAT

Important habitat features of the

Apalachicola Bay system include physio-

• graphic, climatic, and river-flow

conditions. While marshes (emergent

vegetation), oyster beds, and qrassbeds

(submerqent veqetation) represent

important biological habitats of the

estuary. the primary physical habitat in

terms of areal extent is the shallow,

unvegetated soft sediment bottom (Table

1). Within the myriad of rapidly changing

gradients of physical and chemical

features of the estuary, there are certain

recurrent patterns and general trends that

remain more or less constant in soace and over time. Such water-quality features

and nutrient distributions are important

determinants of the habitat conditions in

the Apalachicola Bay system.

?.^.\. Temperature and Salinity

Because of the shallowness of the bay

system and wind-mixing of the water

column, there is little thermal

stratification in the estuary. Water

temperature is hiqhlv correlated (r = O.QO, p < 0.00001) with air temperature

(Livinqston 1983c), which indicates rapid

mixing. Summer temperature peaks are similar from year to year, with seasonal

highs usually in August. Water

14
Table 1. Distribution and area of major bodies of water along the coast of Franklin

County (north Florida) with areas of oysters, grassbeds, and contiguous marshes.
Mean salinity values are lowest at the mouth of the river and in East Bay

(Table ?, Figure 13). According to the

Venice system of brackish water

classification, the lower reaches of the

Apalachicola River constitute the limnetic

zone, with salinities reaching O.S parts

per thousand (ppt). During periods of

high river flow, the zone expands to include East Bay and considerable portions

of Apalachicola Bay. Because of extreme

seasonal and annual variability, there are

no clear-cut zones that remain stable in

the bay system. Rather, the salinity

gradients move through the bay area according to upland runoff conditions.

East Bay, lying northeast of the river

head, is oliqohaline (O.'i-'i.O ppt) durinq

most of the year (Figure 13).

Apalachicola Bay, St, Vincent Sound, and

western portions of St. George Sound vary

between mesohaline (5-18' ppt) and

polyhaline (18-30 ppt) conditions,

depending on river flow and upland runoff

(Livingston 1983d). Areas near the passes

and in the eastern sections of St. George

SALINITY (ppt)

FWOuEW oiSTRiauriON or o»ia <K»n values im each uexei TURBIDITY (Jackson turbity units)

FREOUENCY aSTSIBUTION Of OA'A OOINT VAIUES IN EACH LEVEL

LEVEL L id^iiH

Mm %. 000 300 600 iO£K> IIOQ TOOL: Mo. TU 3X 60O tOOO 1500 2000 3500

I I HiHiIB 1^1

000 12 00

.2 00 1500

15.00 700

1900 2100 2100 2400

DISSOLVED OXYGEN (ppm) COLOR (Pt-Co units)

fHiO^itNCt r>iSt»<iBU*iON Oi L-iTi i.-:.!NT VA^UtS IN EACH Ltvtl

-r---] QABil

fRfOufWCY OtSTHISUTlON OF '.^ATa oOINt VALUES IN EACH LtVSl

LEVCL I 2_

Vr pe*r OOC 5 00 '5C 3 00 y ^C> 9 00

»o. p(n^ 500 75C eOO 850 900 iOOO

"li fSEQ 4:]diiiM ? •>

M.r ptco 000 2000 4QCX) 550C moO 9000

Moi III CO 2000 4000 5500 7000 9000 16000

Figure 13. SYNMAP projections of average levels of salinity, dissolved oxygen,

turbidity, and color at permanent stations in the Apalachicola estuary, based on data

taken monthly from 1972-1980.

16
Sound vary from polyhaline to euhaline (>

10 ppt) conditions. Gorsline (1'363)

alluded to the vertical isohaline

conditions of the estuary except for areas

that are deep or near the inlets.

Livinqston (1*^78, 1984a), however, has

documented seasonal vertical salinity

stratification in various parts of the

estuary, especially in areas affected most

directly by the river. Differences of

surface and bottom salinities of as much

as 5-10 ppt during periods of

stratification further complicate the

exact dimensions of the salinity regime in

a given area of the bay system through

time. However, by most statistical

measures, river flow is the chief

determinant of the salinity structure of

the estuary (Meeter and Livingston 1P78).

periods of high local precipitation

(Figure 14). Salinitv qenerally peaks

during the fall drought

(October-November), Long-term salinity

trends follow river flow fluctuations; low

salinity was noted for a prolonged period

throughout the estuary during the heavy

river flow conditions of the winter of

1972-73, although various factors combine

to shape the long-term (multiyear)

salinity trends in the estuary. Various

statistical analyses (Meeter and

Livingston 1978; Meeter et al . 1°79) have

made a strong association of Apalachicola

River flow with the spatial/temporal

distribution of salinitv throughout the

bay system.

There are persistent seasonal

patterns of salinity in the Apalachicola

estuary, although such patterns are modified by annual variation of river flow

and fluctuations of local rainfall. Low

bay salinities coincide with high river

flows during winter and spring periods;

secondary salinity reductions occur in the

bay system during late summer-early fall

Table 2. Bottom salinities in parts per

thousand at stations in the Apalachicola

estuary. All data represent 5-year means

(1972-77) with maxima and minima for this

period. A cluster analysis was made to group the stations according to salinity

type.

Apalachicola

estuary areas

Bottom salinities (ppt)

Sta- Mini- Maxi- 5-yr

tion mum mum mean

Outer BayRiver dominated-

-1

-lA

-IE

-IC

-IX

[—2

-3

0.0

3.0

6.9

1.4

0.0

33.7

35.6

31.6

33.7

32.0

I—4

0.0 28.1

0.0 22.0

0.0 31.8

Upper (East) bay—

Sike's Cut4A

5 h— 5A

5B

5C

6

0.0

0.0

0.0

0.0

0.0

0.0

26.2

28.0

27.3

25.7

27.8

23.0

15.7

22.1

15.7

20.4

17.8

10.4

4.8

9.6

3.6

7.4

5.1

3.8

4.3

3.6

Stotion 1

Q.

a.

<

(A

i "r iiiii' " iii "" i| I

' "" I "

I

' I "" I |i " iii 'M ii| 1972 1973 1974 1975 1976 1977 I97e 1979 1980 1981 1982

Station 2

a.

a.

<

CO

IB 10.6 35.5 28.6

"

I ""

I I """ I' "" I I I I 'I I I 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 YEAR and MONTH

Figure 14. Surface salinity (5-month

moving averages) at stations 1 and 5 (Apalachicola Bay, East Bay) taken monthly

from 1972 through 1982.

17
?.4.'. Dissolverl Oxygen

Diurnal and seasonal variations of

dissolved oxygen (Figure 1"^) reflect

biological and physical processes in the

svstem. Maximum levels usually occur during winter and spring months because of

low water temperature and, to a lesser

degree, low salinity. During summer and

fall periods, vertical stratification of

dissolved oxygen is evident in various

parts of the estuarv. Spatial

distribution of mean dissolved oxvgen

values (Figure 13) is not uniform; the

highest values occur in the upper reaches

of East Bav (i.e.. Round Bay), iust off

St. George Island (i.e., Nick's Hole), and

alona the eastern side of St. Vincent

Island. Concentrations of dissolved

oxygen in most of the estuary during the

10-vr period of observation are sufficient

to support most forms of estuarine biota

(Figure 1'^) . No sign of cultural

eutrophication is evident. The long-term

pattern of dissolved oxyqen maxima

followed the long-term temperature trends,

with dissolved oxygen peaking during the

cold winters from \^''6 to 1'378. Such

changes represent an indirect effect of

temperature on long-term habitat variation

in the estuary.

?.4.3. _pH

From 1^7? to I'^S?, the pH throughout

most of the bay system ranged between fi and P (Livingston lQ?^3c, unpublisheddata). However, relatively low pH levels

(4-5) were observed in upper portions of

East Bay during periods of heavy local

rainfall and runoff from newly cleared

lands in Tate's Hell Swamp (Livingston

19 78). Such changes were temporary and,

overall, the pH of the Apalachicola Bay

system remains within a range that is not

limiting to most life forms.

"^

. ^ . ^ . Water Color and Turbidit y

Light transmission, as determined by

color (measured in platinum-cobalt units)

and turbidity (in .lackson Turbidity

Units), is a key variable in the timinq

and distribution of orimary and secondary

productivity in the estuarv. The spatial

and temporal distributions of water color

and turbiiHity (Figures 13, 16, 17) are related to patterns of fresh-water flow

into the bav system. The highest levels

of both factors are found at the mouth of

the river and throughout upper East Bay

with clear-cut gulfward gradients. Both

color and turbidity reach seasonal hiqh

levels during winter and early spring

periods of hiqh river flow and overland

runoff. During the major flooding in the

winter of 197?-73, turbidity and color in

the estuary reached a 10-yr high point at most stations. While the general pattern

of color in the estuary follows river flow

fluctuations, the highest levels occurred

in eastern East Bay. The color was directly associated with forestry

activities and runoff from the Tate's Hell

Swamp (Livingston 1"578). Various

compounds such as tannins, lignins, and

„
fulvic acid complexes, which occur naturally in the upland swamps, are washed

into the estuary durinq periods of high

local precipitation. Such water-qual itv

changes, associated with river flow and

local rainfall, affect the biological

organization of the bay system in terms of

primary productivity and food web

structure (Livingston ic)83b-d).

'.S. BIOLOGICAL HABITATS

The Aoalachicola drainage system as a whole is an almost unbroken series of

natural habitats, which include upland

vegetation, swamps, marshes, and flood

plain wetlands. Much of the basin

vegetation has the appearance of a mature

forest because of rapid regrowth. Slash

and longleaf pine are abundant in upland

areas. Although several municipalities

are located near or within the

Apalachicola and Chipola flood plains. none is a major urban center; there is

little industrialization in the basin.

The dimensions of the biological habitats

within the bay system and its associated

watershed (i.e., Franklin County) are given in Tables 1 and ?. Aquatic areas,

together with forested and nonforested

wetlands, comprise about 42% of the total area of Franklin County. As noted

previously, aquatic areas are dominated by

unvegetated soft-bottom substrates.

2.5.1. Wetlands

a. Bottomland hardwoods . The

Apalachicola flood plain (Figure 18) of

the upper river is relatively narrow

a.

o

o

o

I

z>

>- ^ I- "2 5 ^ 5 ^

1972 1973 1974 1975 1976 1977 1978 B79 1980

1972 1973 1974 1975 1976 1977 1979 1979 1980 1981 1982

Oo

cmo

_io

o

<

1974 1975 1976 1977 1978 1979 1980 YEAR and MONTH

« 140

1972 1973 1974 1975 1976 1977 ra78 1979 1980 1981 1982 YEAR and MONTH

Figure 16. Color (5-month moving

averages) at stations 1 and 5 in the

Apalachicola estuary taken monthly from

1972 through 1982.

Figure 17. Turbidity (5-month moving

averages) at stations 1 and 5 in the

Apalachicola estuary taken monthly from

1972 through 1982.

19
Table 3. Terrestrial habitats and land-use patterns in the immediate watershed of the

Apalachicola Bay system (Florida Bureau of Land and Water Manaqement 1977).

Category

Residential

Commercial, services

Transportation, utilities

Mixed urban or built-up areas Other urban or built-up areas

All urban or built-up areas

Total area (ha)

2,461

178

218

27

39

2,922

% of total

1.3

0.1

0.1

0.0

0.0

1.5

Cropland and pasture

Other agriculture

All agricultural land

78

4 82

0.0

0.0

0.0

Herbaceous rangeland

Rangeland

13

13

0.0

0.0

Evergreen forest land

Mixed forest land

All forest land

68,598

36,396

104,994

35.7

18.9

54.6

Streams and canals

Lakes

Reservoirs

Bays and estuaries

All water

1,469

452

10

62,879

64,810

0.8

0.2

0.0

24.3

25.4

Forested wetland

Nonforested wetland

All wetlands

25,562

8,465

34,027

13.3

4.4

17.7

Beaches

Quarries and pits

Transitional areas

All barren land

Total area of Franklin County:

1,441

25

110

1,575

198,398

0.7

0.0

0.1

0.8

(l.'^-S.O km or O.^'-l.Q mi wide). The

forested flood plain broadens along the

lower river (up to 7 km or 11.3 mi wide),

with most of the flood-plain wetlands

located in the lower delta (H. M. Leitman

et al. 1982). The forested flood plain of

the Apalachicola basin is the largest in

Florida (450 km?, 173 mi?; Wharton et al . 1977), and 60 of the 211 tree species in

north Florida are found there (Table 4).

The predominant soecies in terms of cover include water tupelo, ogeechee tupelo,

baldcypress, Carolina ash, swamp tupelo,

sweetgum, and overcuo oak. These soecies

are typical of southeastern alluvial flood

plains and occur in such areas partially

because of their adaptive response to restricted availability of oxygen in

saturated and inundated soils. Despite

continuous logging for over a century, the

20
Table 4. A. Tree species found in the Apalachicola floodplain (from Leitman 1983 and

H. M. Leitman et al. 1982). Included is the relative basal area (in percent) of the

top 25 species. B. Area, in acres, of each mapping category for five reaches of the

Apalachicola River (from Leitman 1983).

Common name Scientific name

Ash, Carolina

Green

Pumpkin

Baldcypress

Birch, river

Box elder

Bumelia, buckthorn

Buttonbush

Chinaberry

Cottonwood, swamp

Cypress

Dogwood, stiff Cornell

(swamp dogwood'^)

Elm, American

Slippery

Winged

Grape

Hawthord, green

Parsley

Hickory, water

Hornbeam, American

Locust, water

Maple, red

Mulberry, red

Oak, cherrybark

diamond- leaf

laurel

overcup

swamp chestnut

water

Palmetto, cabbage

Persimmon, common

Pine, loblolly

spruce

Planertree (water-elm^)

Possumhaw holly

Silverbell, little

Sugarberry (hackberry)

Swamp- privet

Sweetbay

Sweet gum

Sycamore, American

Titi

Fraxinus caroliniana Mill. (5.4)

Frax in us pennsyl vanica Marsh. (2.9

Fraxinus profunda (Bush.) Bush. (1.9)

Taxodium distichum (L.) Rich. (10.6)

Betula nigra L, TO,

Acer nequndo L. (0,

Bumelia lycioides (L.) Pers.

Cephalanthus occidentalis L. Melia azedarac h L.^

Populus heterophyl la L . (0.4) see baldcypress

Cornus foemina Mill . rCornus stricta Lam.b)

Ulmus americana L. (2 . 4 ) Ulmus rubra Muhl.

Ulmus alata Michx.

V i t i s spp.c

Crataegus viridis L. Crataegus marshal lii Egqle.

Carya aquatica (Michx. f.) Nutt. (2.^)

Carpinus carolinian a Walt. (2.0)

Gleditsia aquatica Marsh.

Acer rubrum L. (1.5)

Morus rubra L. Quercus falcata Michx., var. paqodaefolia Ell

Quercus laurifolia Michx. (2.5)

Quercus hemisphaerica Bartr. (£. laurifol ia

Michx. '^) Quercus lyrata Walt. (3.2)

Quercu s prinus L. (Q_. michauxii Nutt.^) (0.3)

Quercus nigra L. (1.8)

Sabal Dalmetto (Walt.)

Diospyros virqiniana L Pinus taeda L. Pinus glabra Walt.

Planera aquatica Gmel . Ilex decidua Walt. (0.

Lodd.

(2.9)

Halesia tetraptera Ellis. (_H. parvif lora Michx.'')

Celtis laevigata Willd. (2.8)

Forestiera acuminata (Michx.) Poir,

Magnolia virqiniana L. (1.0)

Liquidambar styraciflua L. (4.8)

Platanus occidental is L . (0.6)

Cyril la racemif lora L.

(continued)

21
Table 4. (Concluded.

Common name Scientific name

Tupelo, Ogeechee

water

swamp (blackgum)

black (sourgum)

Viburnum, witherod

Walnut, black

Willow, black

Nyssa ogeche Bartr. (11.0)

Nyssa aquatica L. (29.9)

Nyssa biflora Walt. (N. sylvatica var. bif lora

(Walt.) Sarg.b) X5.0)

Nyssa sulvatica Marsh. (_N. sylvatica Marsh.

var. sylvatica^ ) Viburnum cassmoides L. Juglans nigra L. Salix nigra Marsh. (0.4)

^Introduced exotic species.

"^According to Little (1979).

CRadford and others (1968).

<^Little (1979) does not recognize Quercus hemisphaerica as a separate

species.

Acres

B.
Apalachicola flood plain remains relatively intact as a functional bottomland

hardwood system.

Tupelo, gum, and cypress species are dominant in the upper flood plain (Table

4). The lower flood plain is

characterized by coastal plain pine

flatlands, coastal dunes (shortleaf pine,

titi, and bayhead) and freshwater and

brackish marshes. Various forest

associations occur in different regions of

the basin (Table 4) (Leitman 1^83, H. M. Leitman et al. 1982): (1) The

sweetgum/sugarberry/ water oak/loblolly

pine association is found in dry to damp

soils or wetland-toupland/transition

areas. These forest types decrease in the area within the basin as the river

approaches the coast. (?) The water

hickory/sweetgum/overcup oak/green

ash/sugarberry association covers about

78% of the floodplain mainly in the uoper

and middle reaches of the river basin.

This association is not common in the

lower reaches of the valley. (^) The

water tupelo/ogeechee tupelo/baldcypress

association is found in dry to saturated

soils and is concentrated along waterways

and relict waterways in the lower reaches

of the river basin. (4) The water

tupeln/baldcypress association is located

in damp to saturated soils along the

entire length of the river. Pioneer

associations are dominated by a narrow

zone of black willow in areas inundated more than ?^% of the time. Marsh areas

are located along the lower river. Water

depth, duration of inundation and

saturation, and fluctuations in water

levels all contribute to the composition

of the wetland forests. These conditions

100-YR

FLOOD

HURRICANE

FLOOD

^^~~l 1
are dependent to a larqe degree on watershed runoff, flood plain toooqraphic

relief, and drainage characteristics.

b. Marshes . Most of the intertidal

areas around the estuarv are surrounded by

freshwater, brackish, and saltwater

marshes (Figure 19). The freshwater and

brackish-water marshes are characterized

by bull rushes ( Scripus spp.), cattails

( Tvpha spp.), saw grasses ( Cladium spo.),

cordgrass ( Spartina spp.), and needlerush

( Juncus roemerianus ) . Salt marshes of the

region are represented by black needlerush, cordgrass, Distichl is spicata , and

Sal i corn i a spp. Maior marsh develooment

is found along the lower flood plain and areas adjacent to East Bay. These marshes

are dominated by mixed freshwater soecies.

Similar marsh associations are found in

the New River and Ochlochonee River

drainages to the east. Narrow stands of

brackish water marshes occur intermittently along the lagoonal

interface of the Alligator Point peninsula

(at the extreme east end of the system;

Figure 3) and along the bayside portion of

the barrier islands. Limited marshes are located along the mainland east and west

of the Aoalachicola River mouth. The East

Bay marshes dominate the system by area

(Table 1) with lesser marsh development

along St. Vincent Sound and along the

lagoonal oortions of St. George Island and

Dog Island. The marshes in the entire bay

system comorise approximately l^% of the

total water surface.

The Apalachicola marshes are significant feeding and reproductive zones for various aquatic and terrestrial

species (Livingston lQR3c). Vertical and

lateral stratification of this habitat has

provided conditions that house and feed

some of the most important species

(ecologically and commercially) in the

river- bay system.

?.S.?. Seagrass Beds

Grassbeds in the Apalachicola estuary

(Figure 19) account for about 10% of the

"=#^ :

:^»^Mt>'*

Figure 19. Distribution of the marshes and submer'^ent vegetation in the Apalachicola

estuary (data compiled from aerial photographs and ground-truth observations by divers)

(see Livingston 1980a).

24
total water area (Table 1). Except for

certain areas alonq the eastern portions

of St. Georqe Sound, submerqed vegetation

in the Apalachicola estuary is lightlimited by hiqh turbidity and water color.

High sedimentation and resuspension of

sediments in the estuary may also affect

the seagrass bed distribution. Seagrasses

and algal associations are largely

confined to frinqes of the estuary at depths of less than 1 m. The largest

concentration of these submerged grassbeds

is in eastern St. George Sound; such

seagrass beds also occur in upper East

Bay, inside St. George Island in

Apalachicola Bay, and in western St,

George Sound. In East Bay, freshwater and

brackish-water species ( Val 1 isneri a americana , Ruppi a maritima , and

Potamogeton sp. ) are oredominant. Grass

beds along the mainland east of the river

are dominated by Halodule wrightii , Syringodium fil i forme , and Thalassia

testudinum . The shallow lagoonal flats of

Alligator Point, Dog Island, and St.

George Island are populated by Halodule

wrighti i, Gracilaria spp., and Syringodium

fil i forme . Few if any grassbeds are found

in St. Vincent Sound.

As a habitat, seagrass beds provide

organic matter and shelter for various

infaunal and epibenthic invertebrates and

fishes.

?.S.3. Soft-Bottom Substrates

Muddy, soft bottom substrates

comprise about 78% of the ooen water zone

of the Apalachicola Bay system and are thus the dominant habitat form in the

area. The relative composition of the

sand, silt, clay and shell fractions of

the sediments depends on proximity to land, runoff conditions, water currents,

and trends of biological productivity.

Sediment type and associated water-gual ity

conditions in the benthic habitat

determine the composition of infaunal and

epifaunal biological components.

Recruitment and community composition of

the benthic invertebrates (meio^auna and

macrofauna) may depend on the distribution

of flocculent resuspended sediments and

bedload transport. The unvegetated, softbottom habitat in the Apalachicola Bay

system represents the basis for imoortant

food web relationships in the estuary.

?.5.4. Oyster Bars

The Apalachicola estuary is an ideal

environment for the growth and culture of

the oyster ( Crassostrea virqinica )

. The

oyster bars that cover about 1% of the

aquatic area of the bay system (Table 1) are an important habitat for various

assemblages of estuarine organisms. Malor

oyster beds are located in St. Vincent

Sound, west St. Georqe Sound, and the East

Bay-Apalachicola Bay complex (Figure ?0).

New (constructed or artificial) oyster

reefs stq located in eastern portions of

St. Vincent Sound. The highly productive

natural oyster bars of St. Vincent Sound

and western St. George Sound represent the

primary concentrations of commercial

oysters in the estuary. The waters of

both areas are well circulated by the

prevailing currents and are characterized

by salinity conditions optimal for oyster

propaqatinn and qrowth (l.ivinqston 1983c,

d). The reefs near the seaward edqe of

the bay thrive when the river is high

while those near the river mouth do well

during conditions of low water.

Whitfield and Beaumariage (1977)

estimate that about 40''^ of Apalachicola

Bay is suitable for growing oysters but

that substrate type is a major limiting

factor. Rapid oyster growth due to

favorable environmental conditions

accounts for the fact that over 90% of

Florida's oysters (8%-10% nationally) come from the Apalachicola estuary.

?.5.5. Nearshore Gulf Environment

The shallow nearshore gulf is a drowned alluvial plain grading into a limestone plateau to the east and south

(McNulty et al. 1972). The eastern Gulf

of Mexico is characterized by moderately

high-enerqy sand beaches. The north qulf

coast sedimentary province contains relict

sand west of the Apalachicola delta. The

Miocene relict sands and clays off the

Apalachicola embayment qrade into quartz

sand and mud over limestone characteristic

of the extreme eastern gulf region. Much

of the water motion along the shallow West

Florida Shelf is due to tides, although

wind effects are evident, especially in

winter when cold fronts move through the

area. The high-salinity coastal waters

are well mixed except durinq warmer months

25
Figure 20. Distribution of oyster bars and sediments in the Apalachicola estuary (data

from historic records, personal information from oyster dealers in Apalachicola, field

observations by F.S.U. field personnel, and records from the Florida Department of

Natural Resources) (Livingston 1980a). (This chart is currently being updated.)

when a thermocline separates the cooler

bottom waters from the surface-waters.

Organisms in near-shore areas are part of a temperate sand community (Jones

et al. 1<573; Smith 1P74). The shallow

(10-20 m) shelf benthos reflects the

intrusion of tropical species in both

sandy areas and rocky outcrop substrates.

The northeastern gulf lies in the Carolina

Zoogeographic Region with a warm-temperate

fish fauna. Fish assemblages are characterized by high endemism and hiqh

species diversity due, in part, to a number of eurythermic tropical species.

The northeastern Florida gulf coast has a relatively high fishery potential for

crustaceans and finfishes (Jones et al.

1973; Smith 1974).

2.6. NATURAL RESOURCES OF THE

APALACHICOLA DRAINAGE SYSTEM

There are several natural attributes

of the Apalachicola drainage system that

make it unique among Florida and North

American river estuaries (Livingston and

Joyce 1977). The strategic placement of

the drainage, together with the relatively

unspoiled natural comDonents--streams,

rivers, wetlands, estuary, offshore gul^--

have combined to create the conditions for

speciose and unique assemblages of

terrestrial and aguatic organisms. In many ways, the Apalachicola system is an important dispersal route for temperate

species of plants and animals from the

high elevations of the southeastern United

States to the Gulf of Mexico.

The following is an summary of such attributes:abbreviated

1. The Aoalachicola ranks as one of

the great rivers of

is the largest river

Florida. It is the

to stretch from the

of Mexico.

the United States and

(in terms of flow) in

only river in Florida

Piedmont to the Gulf

2. The area of forested floodplain

is the greatest of all river systems in

Florida. The densely forested, bottomland hardwood wetlands of the Apalachicola

River have the highest litter-fall

production rates of the worldwide warm26
temperate systems that have been studied

(Mattraw and Elder 1<380).

3. Nutrient levels are higher in the

Aoalachicola wetlands than in most

comparable systems throuqhout the northern

hemisphere. The Aoalachicola wetlands

contribute significant quantities of

nutrients and organic matter to river and

bay areas. Regular seasonal flooding by

the currently free-flowing river is

necessary for mobilization of particulate

organic matter (POM) and nutrients out of

the floodplain (Mattraw and Elder 1980).

4. The Apalachicola drainage system

includes a grouo of ecological regions

that contribute to soeciose and unique

plant associations. The flora comorises

117 plant species, of which 17 are endangered, ?8 are threatened, and 30 are

rare. Nine species are narrowly endemic

while 71 are endemic to the general

Apalachicola area (Means 1977).

5. The Apalachicola wetlands provide

habitat for rich faunal assemblages. The

basin receives biotic exchanges and input

from the Piedmont, the Atlantic Coastal

Plain, the Gulf Coastal "lain, and

peninsular Florida. The floodplain

forest, with over ^50 species of

vertebrates, is one of the most important

animal habitats o-f the Southeast (Means

1977).

6. Of the drainages of the

Apalachicolan and West Floridian molluscan

province (from the Escambia River to the

Suwannee River), the Apalachicola River

contains the largest total number of

species of freshwater gastropod and

bivalve mollusks. The river contains the

greatest proportion of endemics to the

total fauna in the province, with at least

six VZV9, and endangered soecies (two

Amblemids, four Unionids) (Heard 1977).

7. The tri -river valley is

characterized by a rich fish fauna (ll^i

species) (Yerger 1977). The Apalachicola

basin contains more fish species (85) than

any other Florida river. Three species

( Notropis cal litaenia , N_. zonistius , Moxostoma sp. ) are restricted to the

Apalachicola River and its ma.ior

tributaries, while a fourth species (the

"handoaint" bluegill, Lepomis macrochirus)

originated in the system. Existing

freshwater sport and commercial fisheries

are diverse and rich. The Apalachicola

River is the only river on the Florida

gulf coast that supports a striped bass

( Morone saxatil is ) fishery (Livingston and

Joyce 1977). TlTis fishery is based on a population that is endemic to the river

and considered a separate race from the

Atlantic coast striped bass.

8. Excluding fishes, the

Apalachicola River system contains over

?50 species of vertebrates. The highest

species density of amphibians and reptiles

in North America (north of Mexico) occurs

in the upper Aoalachicola basin (Means

1977). The abundant and diverse bird

fauna is concentrated in the floodplain

forests. Two species considered extinct,

the ivory-billed woodoecker ( Campephilus

principal is ) and Bachman's sparrow

( Aimophi la aestivalis ), were last siqhted

in the Apalachicola system. These species

are part of a growing list of approximately fifty soecies of amphibians,

reptiles, birds, and mammals that are considered endangered, threatened, rare,

of special concern, or of undetermined

status.

^. The Apalachicola estuary, with

its barrier islands, represents a ma.ior

flyway for gulf migratory bird species.

The estuary has the highest density of

nesting osoreys ( Pandion haliaetus ) along

the northeast Florida gulf coast (Eichholz

10. The Apalachicola Ray system is one of the richest and least polluted such areas in the United States. The estuary

now provides over ^0% of Florida's oysters

and is part of a major spawning ground for

blue crabs along the Florida gulf coast

(Livingston and Joyce 1977). The bay

serves as an important nursery for penaeid

shrimp and finfishes and is characterized

by some of the highest densities of

infaunal invertebrates of any comparable

area in the United States.

11. The hiqhly profitable

Apalachicola oyster industry and various

sport and commercial fisheries directly

and indirectly provide the economic and

cultural basis for a high oroportion of

the peoole in the region (Livinqston

iq83c).

27
CHAPTER 3

PRIMARY PRODUCTIVITY AND NUTRIENT CYCLING

Most aquatic systems such as rivers

and estuaries depend on sources of organic

matter outside the system (i.e.,

allochthonous: dissolved and particulate

orqanic matter from associated wetlands)

and within the system (i.e., autochthonous: phytoplankton, benthic plants).

Tnorqanic nutrients (phosphorus, nitrogen)

and orqanic matter (dissolved, particulate) are swept into aquatic systems by

rainfall, overland runoff, and river

flooding. The extremely complex chemical

processes involved in the transformation

of nutrients into plant and animal biomass

are not well understood and are intricately related to microbioloqical

activity. One important qeneral ization

based on the long-term field studies is

that the Apalachicola estuary is

inextricably linked to the river in terms

of freshwater input and the movement of

dissolved and particulate orqanic material

into the estuary. River input is sea- sonally and annually pulsed, and such

influx of materials has an important

influence on allochthonous and

autochthonous sources of orqanic matter

throuqhout the Apalachicola estuary.

et al. 1P77; Elder and Cairns 1982; Elder

and Mattraw 1^82; Mattraw and Elder 1980,

1982). Over time, the Apalachicola River

has meandered in broad curves through the

flood plain, Erosional and depositional

processes have led to the development of

shoals, backswamps, channels, sloughs,

levees, and oxbow lakes. The dynamics of

the Apalachicola River affect the

transport of dissolved and particulate

substances into receiving aquatic areas. However, such transport of allochthonous

substances depends on complex interactions

of river flooding with factors such as wetland productivity, decomposition

processes, the timing and relative heights

of the flood staqe, the heiqhts of

surroundinq lands, soil types, and

drainaqe characteristics of the flood

plain. The unifyinq characteristics of

the wetland inputs are the distribution

and environmental functions of the

bottomland hardwood forests of the

Apalachicola floodplain (Figure 21).

Nutrient fluxes and primary

productivity of the river-estuary system

have been studied for over a decade; the

followinq is a review of the available

information concerning the Apalachicola

system.

3.1. PRIMARY PRODUCERS

3.1.1. Allochthor-ous Sources

a. Freshwater wetlands . The

production and decomposition of orqanic

matter in the floodplain wetlands

represents one facet of estuarine

productivity (Livinqston l<581a; Livinqston

EVAPO-TRANSPIRATION

RflIN - DISCHARGE

STAGE^___^ ELEVATION ^

SOILSFLOODPLAIN

f INUNDATION

> TREE

DISTRIBUTION

POSITION O^ V LEAF PRODUCTION

ATMOSPHERE

^NUTRIENTS

DETRITUS

RIVERBAY

TRANSPORTfJ

Fiqure 21. Nutrient/detritus transport

mechanisms and lonq-term fluctuations in

detrital yield to the Apalachicola River

flow (modified from Mattraw and Elder 1980

and Livingston unpubl.).

28
General plant distribution in the

riverine wetlands is associated with

topographic features of the flood plain

and surrounding forested lowlands (Clewell

1978). H. M. Leitman et al. (1^8?) showed

that the height of natural riverbank

levees and the size and distribution of

levee breaks control floodplain hvdrologic

conditions. Vegetative composition is

highly correlated with depth of water,

duration of inundation and saturation, and

water level. Leitman (1978, 1983) and

Leitman and Sohm (1981) described in

detail the distribution of floodplain

trees in the Apalachicola drainage.

According to these studies, pine flatwoods

and loblolly pine-sweetgum associations

are often found on elevated slopes while more mesic hardwoods inhabit the levees.

River banks are occupied by willows and

birches. Terraces or basin depressions

are inhabited by hardwood swamp species.

Cypress-tupelo associations are often

located in sloughs. Backswamps are characterized by blackgum and sweetbay

associations.

The bottomland hardwood community of

the Apalachicola floodplain produces larae

amounts of potentially exportable material

(Elder and Cairns 1982). The weighted

mean of litterfall was 800 grams m'^ with

overall annual deposition within the 454

km-^ bottomland hardwood flood plain of

360,000 metric tons (mt) (396, 7?0 tons) of

organic matter. These production levels

are similar to those observed in eguatorial forests but are higher than those

noted in cool temoerate forests and most

warm-temperate forests. Levee vegetation

produced more litterfall per ground

surface area than did the swamp

vegetation. The seasonal distribution of

litterfall was characterized by a sharp

late autumn peak. The three most abundant

flood plain tree species (tupelo, cypress

and ash) accounted for over 50% of the

total leaf-fall, even though these species

were the least productive of those

analyzed on the basis of mass-per-stem

biomass.

Annual flooding is a major factor for

mobilization of substances out of the

flood plain. Flooding leads to immersion

of litter material, enhanced decomoosi tion

rates, and transfer of the breakdown:

products (nutrients and detritus) to

associated aquatic systems (Cairns I'^Bl,

Elder -and Cairns 1^82). The river is thus

closely associated with the rich

productivity of the Apalachicola wetlands

and is the primary agent for movement of

organic matter out of the floodplain. In this way, the forested Apalachicola River

flood plain is an important source of

organic carbon for the estuary. Spring

floods during March and April of 1980

deposited 35,000 mt (38,570 tons) of

detritus derived from litterfall into the

Apalachicola estuary (Mattraw and Elder

1982). During one year of observation,

total organic carbon deposits in the bay

amounted

' to 214,000 mt (235,830 tons).

Total nitrogen and total phosphorus inputs

to the river during the same period were

21,400 (23,593) and l,fi50 mt (1,818 tons),

respectively (Mattraw and Elder 1<582).

The annual detrital organic carbon input was 30,000 metric tons (Mattraw and Elder

1982). Mattraw and Elder (1^82) estimated

that an 86-day period of winter and spring

flooding accounted for 53, 60, 48, and 56

percent of the annual total organic

carbon, particulate organic carbon, total

nitrogen, and total phosphorus transport,

respectively. Flood characteristics are important determinants of the amounts and

forms of transported materials. While

there was an annual net export of

nutrients to the estuary, it is likely

that the wetland system acted as a nutrient sink during certain periods of

the year. Although nutrients are released

to the river by flood-plain vegetation,

such compounds are subiect to active

recycling within the receiving aquatic

systems.

The considerable exoort of

particulate matter from the flood plain is

consistent with previous findings.

Livingston (1981a) and Livingston et al . (1976a) found a direct relationship

between river flooding and the appearance

of micro- and macroparticulate matter in

the estuary. Results of long-term studies

of the significance of river-derived

particulate organic matter to the estuary

(Livingston 1981a, b) indicate that the

exact timing of the peak river flows and

the seasonal changes in the oroductivity

of wetlands vegetation are key

determinants of short-term fluctuations

and long-term trends of the input of

allochthonous organic matter into the

29
Apalachicola estuary (Figure 21). A linear regression of microdetri tus and

river flow by season (Table 5; Figure 22)

showed seasonal differences in the

relationship of detrital concentration and

river flow (Livingston 1981a). During

summer periods, there was no direct

correlation of river flow and detritus in

the estuary. By the fall, there was still

no significant relationship although there were occasional influxes of detritus with

minor peaks in the river flow. By winter,

however, a strong direct relationship was apparent between microdetrital loading and

rivpr-flow peaks,

differed from that

loading,

associated

required

comoarable

detritus,

the degree

a seasonal

The winter rearession

of the spring detrital

though significantly

river-flow levels,

river levels for

concentrations and loading of

This analysis indicates that

and timing of river flooding on basis affects the level of

which,

with

higher

detrital loading to the estuary.

There are various additional sources

of al lochthonous nutrients and detritus

for the Apalachicola River and estuary

Table 5. Linear regression (log/log) of total microdetritus (ash-free dry weight) and

river flow (m^ sec"l) by month/year by season (August 1975-April 1980), at station 7,

located at the mouth of the Apalachicola River. Data are taken from Livingston

(1981a). r = Pearson correlation coefficient.

Station/month (Significance of r)

Station 7 (Surface)

June-August

September-November

December- February

March-May

0.08

0.48

0.70

0.77

0.?3

0.23

0.49

0.60

0.39863

0.03469

0.00188

0.00057

Station 7 (Mid-depth)

June-August

September-November

December-February

March-May

0.35

0.19

0.64

0.68

0.12

0.04

0.40

0.46

0.11809

0.25542

0.00570

0.00397

Station 7 (Bottom)

June-August

September-November

December- February

March-May

0.08

0.21

0.77

0.55

0.01

0.04

0.60

0.30

0.40243

0.22867

0.00037

0.02253

30
JUNE -AUG O DEC -FEB

A SEPT -NOV •MAR -MAY
the Apalachicola estuary throughout the oredominant in the spring, while

year (Table 7) (Estabrook 1P73). In East Skeletonema costatum , Rhizosolenia alata

Bay, Melosira granulata is the dominant and Coscinodiscus radiatus prevail during

species; Chaetoceros lorenzianus is fall and winter months. Although the

dominant in Apalachicola Bay. Species phytoplankton standinq crop is quite low

such as Chaetoceros lorenzianus , at any given time, phytoplankton

Bacteriastrum delicatulum , and productivity is often quite high in areas Thalassiothrix frauenfeldii are such as the Apalachicola Bay system.

Table 6. Net above-ground primary production of marsh plants in various salt marshes

(Kruczynski et al. 1978b).

Marsh plant and Net primary productivity g/m^/yr

location LM UM HM Authors

Spartina alternif lora

FL
Table 7. Presence/absence information for net phytoplankton taken from the

Apalachicola estuary by month from October 1972 through September 1973 (Estabrook

1973). X

= presence.

1 = 10/14/72

2

= 12/02/72

3

= 01/06/73

4

= 03/19/73

5

= 04/22/73

6

= 05/19/73

7 = 06/11/73

8 = 07/12/73

9

= 08/22/73

10 = 09/10/73

Phytoplankter 10

PHYLUM CHRYSOPHYTA

Melosira sulcata

Melosira granulata

Melosir a nummuloides

Melosira dubia

Melosira varians

Skeletonema costatum

Coscinodiscus radiatus

Coscinodiscus spp.

Coscinodiscus apiculatus

Coscinodiscus wailessi

Coscinodiscus excentricus

Coscinodiscus marqinatus

Coscinodiscus central is

Coscinodiscus oculus iTidis

Coscinodiscus nitidus

Coscinodiscus concinnus

Actinocyclus chrenberqii

Actinocyclus undulatus

Biddulphia sinensis

Biddulphia rhombus

Biddulphia aurita

Biddulphia alternans

Biddulphia longicruris

Eupodiscus radiatus

Bellarochia malleus

Triceratium f avus Tricerati urn reticulum

Hemiaulus hauckii

Chaetoceros spp.

Chaetoceros lorenzianum

Chaetoceros decipiens

Chaetoceros didymus

Chaetoceros curvisetus

Chaetoceros coarctatus

Chaetoceros bravis

Chaetoceros aff inis

Chaetoceros compressus

Chaetoceros peruvianum

Chaetoceros glandazii

Chaetoceros pelagicus

Chaetoceros dan i cum Chaetoceros constrictum

Bacteriastrum delicatulum

X

X

XX

X

X

XX

X

X

XX

XX

X X X X

(continued)

33

X X

X X

X X

X

X

XX

X

X

X

X

X

X

X

X

X

X

X

XX
Table 7. (Continued. )

Phytoplankter
Table 7, (Concluded. )

Phytoplankter 9 10

Dinophysis caudata

Dinophysis diagenesis (?)

Dinophysis tripos

PHYLUM CHLOROPHYTA

Pediastrum simplex

Pediastrum duplex

Pediastrum tetras var. tetraodon

Scenedeslrius quadricauda

Studies by R. L. Iverson and his

students indicate that phytoplankton

productivity is an important source of

organic matter in the Apalachicola

estuary. In general, phytoplankton growth

depends on temperature, light, and

avail able nutrients (nitrogen, ohosphorus)

(Figure 23). Temperature is the primary

limiting factor for phytoplankton

productivity in the estuary during the

winter months. Nutrient concentrations

and possibly predation pressure control

production from late soring

The usually low levels of

productivity during the

way to peaks in April,

are noted during summer

phytoplankton

to the fall,

phytoplankton

winter give

Secondary peaks

and fall months.

The average cA^ phytoplankton

productivity (Figure 23) ranged from fi3 to

1,694 mg C m"? dayl (Estabrook 1973;

Livingston et al. 1974). The relationship

of phytoplankton productivity and

predation pressure from zooplankton has

not been determined. However, since river

discharge is strongly associated with

nutrient concentrations in the estuary

(Livingston et al. 1<^74), such factors as river flow and nutrients, together with

the general ecological conditions in the

estuary, combine to control the phvtoolankton productivity of the bay system.

Despite considerable spatial and

temporal variability of phytoplankton

productivity, Eastabrook (1973) estimated

an annual productivity value of 371 g C

m-^ for the Apalachicola estuary. This

figure was taken from averaged data (five

bay stations) sampled monthly over a 12-month period. Based on these figures,

the phytoplankton productivity from the

bay system approximates 233, ?84 t C vr"l

(257,070 tons C yr-1); for the immediate

estuary (East Bay, Apalachicola Bay), this

figure is 103,080 t C yr-1 (113, S94 tons C yr"l). When compared to production values

in other estuaries of the region (Table

8), the phytoplankton productivity and

chlorophyll _a levels in the Apalachicola

estuary are relatively high.

b. Submerged vegetation . The

relatively high levels of color,

turbidity, and sedimentation tend to limit

submerged macrophytes to the shallowest

portions of the Apalachicola estuary

(Livinqston 1980c, ia83c). Species

composition and distribution of seagrass

beds are given by Livingston (l^SOc,

1983c). A major concentration of

seagrasses occurs in eastern St. George

Sound, which remains outside of the

influence of river drainage (Table 1,

Figure 19). Such areas are dominated bv

turtle grass ( Thalassia testudinum ), shoal

grass ( Halodule wrighti i ), and manatee

grass ( Syrinqodium f il iforme )

. Seagrass

beds are also located in upper portions of

East Bay. Such assemblages are dominated

by tape weed ( Vail isneria americana ),

widgeon grass ( Ruppia maritimaTJ and sago

pondweed ( Potamogeton sp. )

. Since the

early 1980's Eurasian watermil^oil

( Myriophyllum spicatum ) has taken over

35
3

O

o '=

CL

_>» O

o

2000
Table 8. Physical, chemical, and productivity data taken from locations along the

northwest gulf coast of Florida (from R. L. Iverson and his students, unpublished data,

Myers 1977). Standard deviations () are also given.

Station
Table 9. Total annual net productivity and net input to the Apalachicola estuary (East

Bay, Apalachicola Bay, St. Vincent Sound) and the Apalachicola Bay system (Aoalachicola

estuary, St. George Sound, Alligator Harbor). Productivity includes (metric tons)

organic carbon produced by the Apalachicola River wetlands, coastal marshes, phytoplankton, and seagrass beds.

Vegetation

Apalachicola estuary Apalachicola Bay system

Net in situ Net input Net in situ Net input

productivity mt C yr~l productivity mt C yr"l

mt C yr~l mt C yr~^

Season of

maximum input

Freshwater

wetlands

Coastal

marshes

Phytoplankton

Seagrass

beds

360,000

37,714

103,080

8,953

30,000 360,000 30,000

37,714(?) 46,905

103,080

8,953 27,213 27,213

winter/spring

46,905(?) late summer,

fall(?)

233,284(?) 233,284(?) spring and

fall

summer-fal 1

A review of the phytoplankton ecologv

of the Apalachicola estuary (Estabrook

1973; Livingston et al. 1974, lQ76a; Myers

and Iverson 1977) indicates that ohytoplankton productivity is relatively

restricted to conditions of optimum

temperature and ample (available)

nutrients. Such conditions occur princioally in the spring, summer, and

fall. Multiple regression analysis (Myers

and Iverson 1977) indicated that river

discharge explained 209S-50^ of the

variability of chlorophyll _a and phvtoplankton productivity. Nutrients were positively correlated with river

discharge. Temperature accounted for 26^

to 49% of the variability in phytoplankton

productivity. Water temperature was also

positively correlated with phytoplankton

productivity. Wind speed was positively

correlated with suspended sediments and

phosphate concentrations, increases in

which were followed by increases in ohytoplankton productivity. Nutrient

enrichment experiments indicated that

nutrients are limiting only during summer

and fall (Estabrook 1^73) and that

phosphate is the primary nutrient that

limits phytoplankton productivity in East

Bay and Apalachicola Bay (Myers and

Iverson 1*^77), although both nitrates and

phosphates may be limiting in summer (Livingston et al. l^^a).

2

<

N D J F M A MONTH

Figure 24. Monthly averages of daily

litterfall on intensive transect plots across the Apalachicola wetlands (after

Elder and Cairns 1982).

38
Recently, certain revisions have been

proposed of early concepts of detritus

outwellinq from coastal marshes (Haines

197Q). There is evidence of no net exportof particulate organic matter (POM) from

salt marshes under certain conditions

(Woodwell et al. 1Q77). Odum et al.

(1^179) have hypothesized that net fluxes

of POM from coastal marshes depend on the

qeomoroholoqy of the wetland basin, the

maqnitude of the tidal ranqe, and upland

freshwater inout. In the Apalachicola

estuary, the tidal ranqe is relatively

small. Marsh distribution is limited

larqely to the delta area (East Bay) and

laqoonal portions of the barrier islands.

The considerable river runoff and the

associated export of orqanic matter due to floodinq would amplify the importance of

the East Ray marshes accordinq to the Odum

model (Odum et al . l^^yg).

The salt marshes of the bay svstem

contribute only a small fraction of the

particulate orqanic loadinq to the bay

system (Livinqston et al . 1974), althouqh

such areas are important nurseries for

estuarine fishes and invertebrates

(Livinqston 1^80c). However, the marshes

may olay a role in the export of orqanic

material to the bay system. Ribelin and

Collier (1P79) showed that local marshes

export detrital agqreqates or films that

averaqe ?5-50 m in thickness and are produced by benthic alqae rather than by

microbial decomposition of the marsh

plants. Tidal action lifts these films of

alqae out of the marshes, especially

durinq late summer ebb flows. Thus, while

the vascular tissue of the marsh qrasses

is decomposed beneath a layer of benthic

alqae, it is essentially retained within

the marsh proper. Amorphous aqqreqates of

"nanodetritus" composed of microalqae may

play a more important role in the nutrient

budget of the bay svstem than previously

thouqht, esoecially during late summer and

early fall periods.

The seasonal abundance and spatial

distribution of nutrients and detritus in

the Apalachicola Bay system result from a combination of forces, some of which are quite localized and specific in nature.

For example, the timing and magnitude of

localized hydrologic events such as

Table 10. Nutrient yields for various drainage areas in

Chattahoochee-Flint River system. Data are presented on an areal

Mattraw and Elder 1982).

the Apalachicolabasis (adapted from
passinq thunderstorms, wind effects, and

tidal actions are superimposed over basin

characteristics such as depth and bottom

morphology. These, in turn, may

significantly influence larqer-scale

conditions such as temperature, salinity,

and light penetration. The large-scale

seasonal fluctuations of important

climatic features, in combination with the

influence of local habitat distribution

and basin configuration, produce an array

of processes whereby organic matter is

incorporated into the estuarine food webs.

The seasonal cycle of nutrientdetritus flux in the Apalachicola estuary

has been well established (Livingston et

al . 1976a; Livingston and Loucks 1978).

Huring winter and spring periods of hi ah

river flow, dissolved nutrients and

particulate organic matter are washed into

the estuary. The influx is concurrent

with salinity reductions. Peak levels of

leaf matter are present during these

periods. One to two months later, wood

debris and other forms of particulates

appear in the bay system. In the spring,

as river flow diminishes, temperature

increases, and the water becomes clearer,

phytoplankton blooms occur. As

principally phosphorus, become

during summer/fall months,

productivity becomes

wind-mixed transfers of

the spring

nutrients,

1 imi ti ng

phytoplankton

dependent on nutrients from the sediments into the

water column. During the summer and early

fall, local rainfall enhances nutrient

enrichment. At this time, benthic

macrophytes begin to die off. The peak

levels of macrophyte organic debris and

Table 11. Nutrient values (winter and summer) for stations in the Apalachicola estuary

(means ± one standard deviation of five stations) and River (Station 2) (Livingston et

al. 1974).

Nutrient

Nutrient values ( g/1)

Site
microaggreqates from the marshes occur during the fall as river flow and rainfall

are minimal. By late fall (November),

temperature drops and salinity

coincidental ly increases to an annual

maximum throughout the estuary. Bv

winter, temperature is low as river flow

once again rises.

Even though the input from various

sources is variable in terms of magnitude

over time, the input of particulate

organic matter to the estuarv from all sources is fairlv constant. Thus, there

is a generally continuous influx of

dissolved and particulate organic and

inorganic matter to the estuary throughout

the year; this matter is then subiect to various processes, physical and

biological, which are dependent on specific spatial-temporal habitat

conditions.

3.3. MICROBIAL ECOLOGY

In the Apalachicola estuarv,

approximately 0.005'^ of the sediment dry

weight is composed of bacterial biomass

(organic carbon) and COQ"^ is composed of

extracellular carbohydrates (D. C. White,

Florida State University; pers. comm.).

Usually, these microbes are concentrated

on particulate surfaces as morphologically

diverse prokaryotic and microeukarvotic

assemblages (White 1*^83). The ecological

importance of microbes to the estuary is

defined by microbial biomass (which forms

the basis of food webs) and microbial

metabolic activity (which contributes to various bioqeochemical and recycling

processes). White and his coworkers have

quantified the biochemical "siqnature"

components of specific microbial community

associations. These components include

phospholipids, adenine- containing

components, muramic acid, and hydroxy

fatty acids, which orovide biomass

estimates. Community composition has been

evaluated by analysis of phosoholipid

alkyl fatty acids (prokaryotes

microeukaryotes) and "signature" lioids

(anaerobic-aerobic bacteria). Fatty acids

are an excellent measure of algae, and

other groups of microeukaryotes can be

characterized by the polyenoic fatty acid

composition (Federle et al. 1°83).

Nutritional status was analyzed by

measurement of poly-beta-hydroxy alkonates

(PHA), extracellular qlycolalyn, and other

microbial byproducts (White 1<583). These

methods were used to analyze microbial

activity in the Apalachicola estuarv.

A series of experiments have been

carried out to learn the fate of

particulate organic matter deposited in

the estuary as a result of river flooding.

Morrison et al. (I'^^y) demonstrated a succession of microbiota that colonized

oak leaves deposited in the estuarv.

Initially, colonization is by bacteria

with a hiqh ratio of muramic acid to ATP.

These bacteria are succeeded by diatoms

and fungal mycelia that do not contain

muramic acid. Thus, initial bacterial

colonization is succeeded by a community

of fungi and microeukaryotes. Bobbie et

al. (1078) found that microbial

communities on biodeqradable substrates

such as leaf matter are biochemically and

morpholoqicall y more diverse than those on bioloqically inert substrates. A 10-fold

increase in biomass on the bioloqical substrates was also noted. Grazinq amphipods

removed microbiota without affectinq the

morphology of oak leaves (Morrison and

White 1980). The colonization of mixed

hardwood leaves from the Apalachicola

flood plain in the estuary varied more as

a function of leaf surface than of

location (White et al . 1977, 1979a, b).

However, macroorganisms were attracted to the litter baskets as a function of

location rather than microbial biomass

(Livingston unpublished data).

The activities of microbes are inextricably linked with orqanisms at hiqher levels of the estuarine food web

(Figure ?5). Amphipod distribution was significantly correlated with concentrations of certain bacterial fatty acids

(White et al. 1979a, b). Amphioods

grazing at natural densities induced

increases in microbial biomass, oxygen

utilization, PHB synthesis, lioid synthesis, and I'^CO^ release from simple substances by microbes (Morrison and White

1980). These chanqes caused qrazinq

shifts in community structure from diatomfunqal-bacterial associations to bacterially dominated ones. Within

limits, grazing thus stimulates microbial

growth and alters the microbial communitv.

Indications are that organisms graze on detrital and sedimentary microbiota and

41
substantially affect the microbial

associations. Studies of microbes in the

absence of their predators are not

sufficient if comparisons with natural

^unctions are intended (White 1Q83).

Recent studies indicate that

estuarine microbial associations in

polvhaline areas of the bay are actually

controlled by epibenthic predators

(Federle et al. 1'383). Replicate areas (^

m'') of mud-flat sediment were caqed in the

field to confine and exclude predators.

Uncaqed areas were used as controls. The

microbiota of the sediments was characterized at weeks 0, ?, and fi by

measurement of the concentrations o^" phospholipid and analysis of the fatty

acids of the microbial lipids extracted

from the sediments. The data were analyzed using an analysis of variance and

step-wise discriminant analysis. After ? weeks, the microbiota of the predatorexclusion area was significantly different

from that in the control and predator

inclusion areas. After R weeks, these

differences became more pronounced. There

were no demonstrable caqing effects that

could account for the treatment

differences. The results indicated that

removal of predators had a profound effect

on the microbial communities in estuarine

sediments. Thus, we see that the

intermediate trophic levels (epibenthic

predators) of the estuarine food webs are part of the control mechanism that defines

the structure and level of productivity of

the microbial communities.

Sediments and particulate matter

deposited in the estuary form a substrate

for microbial productivity, which is

stimulated by dissolved nutrients in

various forms (Figure ?'i). The

transformation of dissolved substances

into living particulate matter produces

the food of important grouos of grazing

organisms, which, in turn, represent the

base of the detrital food webs in the

estuary. Grazing and other physical

disturbances enhance microbial

productivity and alter the qualitative

composition and succession of the

microbial community. The oeriodic input

of particulate organic matter and

RIVER FLOW

RUNOFF LOW SALINITY

DISSOLVED NUTRIENTS

INCREASED MICROBIAL

BIOMASS

TIDAL SUBSIDY

WIND

DISTURBANCE

II

PHB/LIPIDS

'"COj RELEASE

Oj UTILIZATION

BASE, DETRITAL yFOOD WEBS

Figure 25. Tentative model of microbial

interactions with various physical and

biological processes in the Apalachicola

River estuary (Livingston 1983c).

dissolved nutrients into a shallow bay

ecosystem characterized by gradients of

salinity is seen to provide the appropriate components for a highly productive

system. Tidal and wind-induced currents,

periodic flooding, and predation all

provide a series of disturbances that,

together with the periodic enrichment of

the system from upland runoff, increase

microbial productivity. River flow and

fresh water runoFf from associated

wetlands, together with the shallowness of

the system and tidal/wind energy

subsidies, all contribute to the observed

high productivity of the estuary.

Considering their immense biomass and

their role as processors of nutrients into

biologically active material, the microbes

are an important component in the energy

transformations within the system.

42
CHAPTER 4

SECONDARY PRODUCERS

4.1. ZOOPLANKTON

The diverse zooplankton represent an important link between the phytoplankton

and higher levels of the estuarine food

webs. Almost every ma.ior qroup of

organisms is represented in the

zooplankton, either as larvae or as adults; great variety is also evident in

the relatively extensive size range of

individuals. Zooplankton have marked

differences in swimming ability and are often dispersed in patch v, somewhat

irregular spatial distributions.

Zooplankton repackage organic matter

produced by phytoplankton into larger

particles, thereby concentrating energy

into forms more useful to higher

predators. At the same time, they excrete

nutrients that may again contribute to phytoplankton productivity.

of the estuary (Figure 26). Overall

seasonal peaks of copepods in Apalachicola

Bay are noted from March to August with

minimum densities in January and February.

Optimal salinities for the dominant

species, Acartia tons a , range from Ifi to

22 ppt. East Bay, characterized by low

but variable salinity, has the highest

variability in zooplankton numbers over

time. Coastal waters have been most

stable in terms of seasonal changes in

zooplankton abundance. Apalachicola Bay

also has the highest species richness of

the three areas studied. Cladocerans and

chaetoqnaths are located primarily in

coastal waters. Decapod larvae throughout

the estuarv are primarily crab zoeae; other zooplankton include polychaete

larvae, ostracods, amphipods, isopods,

mysids, echinoderms, ctenophores, and

coelenterates.

Zooplankton (Table 12) are among the

least known assemblages in the

Apalachicola estuary. While the

dimensions and interrelationships of the

zooplankton community are relatively

poorly understood in the Apalachicola

estuary, certain factors such as temperature, salinity, wind, nutrients,

primary (phytoplankton) productivity, and

predator-prey relationships are known to contribute to processes involving this

group of organisms. Net zooplankton are composed largely of holoplankton (plankton

for entire life cycle; about '^0%) , while

meroplankton (temporary plankton)

constitute less than lO?;! of the total

(Table 12; Edmisten 1QZ9). The

holoplankton are composed mainly of

copepods, cladocerans, larvaceans, and

chaetognaths. Copeoods, notably Acartia

tonsa , are dominant throughout the

estuary. Apalachicola Bay supports higher

numbers of copepods than any other portion

The zooplankton mean standing crop

(dry weight) in East Bay approximates 4.0

3 mg m"-^ annually; in Apalachicola Bay, 32.1

mg m m~3 yr~l

yr

-1. in coastal areas, 16.7 mg

Peak dry-weight biomass occurs

in May throughout most of the study area with secondary increases during .luly and

August (Figure 26). Zooplankton

distribution is influenced by changes of

temperature and salinity through time

(Table 13). Edmisten (1979), using

analysis of covariance with temperature

and salinity as covariates for factors

such as Acartia numbers, percent abundance

(of Acarti a ) , total zooplankton numbers,

zooplankton biomass, and Shannon

diversity, found significant station and

month differences in all cases (p - 0.02).

Temperature significantly influenced

numbers of Acartia , total zooplankton

numbers (p < 0.01), and biomass. Salinity

significantly affected zooplankton

numbers, biomass, and diversity (p < 0.01)

43
Table 12. Distribution of the major zooplankton qrouDs in the Apalachicola estuary and

associated coastal areas (after Edmisten, 1979). Average values are given from 1973

through 1974. The symbol (+) means l/m^ or less than 0.1%.

Average 1973-1974 values

Apalachicola Bay

^. , (6 stations)

Zooplankton groups No./m^ % No./m^ %

Copepods

Acartia tonsa

Paracalanus

crossirostris

Paracal
Table 12. (Concluded.)
Table 13. Pearson correlation coefficients (r) for significant (p < 0.05) zooplankton

relationships in East Bay, Apalachicola Bay, and coastal areas (Edmisten 1979).

Variable East Bay Apalachicola Bay Coastal areas

Temperature vs. Acarti a tonsa

Total zooplankton

Zooplankton biomass

Salinity vs. Acarti a tonsa

% Acarti a tonsa

Total zooplankton

Zooplankton biomass

Zooplankton diversity

0.45

0.50a

0.45

0.58

0.58

-0.30

0.31

0.40

0.51

0.463

^Significant at p

' 0.10.

less than ?0% of the overall abundance.

The evenness factor is hiqher in the more

stable marine environment with increased

representation by cladocerans, decapod

larvae, and other cooepods (i.e., Temora

turhinata , Paracl inus parvus , P_. crassirostris , Oithona nanlT) (Tdmisten

1<379) . Zooplankton biomass in coastal

waters is correlated with temperature (r = 0.46).

Zooplankton in Apalachicola Bav has

characteristics of both the inshore and

offshore components (Edmisten 1979).

Overall numerical abundance was highest in

Apalachicola Bay (Figure ?fi). Numbers of

and total zooplankton

biomass follow general

of water temperature,

the spatial distribution

any

Acarti a tonsa

abundance and

seasonal trends

Salinity affects

of zooplankton in Apalachicola Bay at given time. Salinity increases appear to

be associated with decreased relative

abundance of Acarti a tonsa . At low

salinities, lower numbers of Acarti a are taken although this species still comorise

a hiqher percentage of the overall zoo- Dlankton assemblage at such times. Thus,

while temperature influences overall

trends of abundance through time, salinity

is associated with the spatial

distribution and relative abundances of

zooplankton in Apalachicola Bay at any

given time.

4.?. LARVAL FISHES

Planktonic fish larvae, derived from

either demersal or planktonic eggs, are common among various marine teleost

species. While it is well known that

estuaries have relatively high levels of

phytoplankton productivity and that such

levels are necessary for feeding

aggregations of zooplankton (Mann 1982),

the relationship o-f such high productivity

to developing stages of marine fishes is

not quite as well known. Lasker (1975)

has shown that larvae of the northern

anchovy ( Engraul is mordax ) feed on phytoolankton and that there is a direct

association between feeding activity and

phytoplankton concentration. Thus, there

may be close relationships between the

highly oroductive inshore waters of the

Gulf and develooing stages of various

teleost fishes.

The relatively high numbers of

ichthyoplankton in the Apalachicola

estuary indicate the importance of this

system as a nursery for fishes. The most

abundant planktonic form is the bay

anchovy ( Anchoa mitchil li ), which accounts

for 92% of the eggs and 7S% of the larvae

taken during a year-long survey (Tables

14, 15; Blanchet 1978). Other relatively

abundant larvae include silversides

46
Table 14, Distribution of ichthyoplankton in the Apalachicold estuary as indicated by the presence of eggs and larvae. Dotted lines indicate

sparse breeding activity. Solid lines indicate widespread and/or intensive breeding as indicated by large numbers of eggs or larvae. Data are taken from Blanchet (1978),

Month

Species N D M A M N D

Brevoorita sp. Harenqula jaguana

Anchoa mitchili

Anchoa hepsetus

Gobiesox strumosus . Atherinidae

Syngnathus s co ve 1 1 i .. . . Syngnathus louisianae

Chloroscombrus chrysura

Lagodon rhomboides

Bairdiella chrysura

Cynoscion arenarius

Cynoscion nebulosu?

Leiostomus xanthurus

MenticirTWus sp. Micropogonias undulatus

Poqonias chromis

Sciaenop's ocellata .. Hypleuroc"hilus geminatus

Hypsoblennius hentzi

Gobiosoma sp. Prionotus sp. .... Trinectes maculatus

(Atherinidae), skilletfish ( Gobiesox

strumosus ), gobies ( Gobiosoma spp. ), aruT various warm-season spawners. Winter to early spring types are dominated by

Atlantic croaker ( Micropogonias

undulatus ), spot ( Leiostomus xanthurus ),

and Gulf menhaden ( BrevoorTTa patronus ) . Various other sciaenid larvae are taken,

including red drum ( Sciaenops ocellatus ),

southern kinqfis"h CMenticirrhus

americanus ), and the sand seatrout

( Cynosci"on" arenarius ). The abundance of

total larvae is highest in western

portions of Apalachicola Bay, largely

because of the high numbers of Anchoa

mitchil li .

Eggs of most species (except

anchovies) are generally found offshore,

indicating that few species actually spawn

within the estuary. The developinq stages

of fishes usually appear within the bay

system at different times of the year.

Areas in the estuary away from the passes

are characterized by the presence of

species that soawn within the hay

(anchovies, atherinids, blennies and

gobies). Relatively large numbers of goby

larvae are found at West Pass.

With the exception of the gulf

oipefish ( Syngnathus scovell i ), which

appears to breed throughout the year, most

species have specific breeding seasons extending from one to several months.

Anchovies have an extended breeding season

although they are considered warm-season

spawners. Two peaks in total larval

abundance (April-May and July-September)

occur (Table IB). Larval abundance and

species richness are higher during spring

months, however. Peak numbers of

47
Table 15. Numbers of ichthyoplankton with larvae and without anchovy larvae (in

parenthesis) taken at various stations within the Apalachicola estuary (after

Blanchet 1978).
Table 15. (Concluded.)

Date

Station

IC Offshore IB

Inshore

Little

St. George lA

10/17
Table 16. Invertebrates taken in cores, leaf-baskets, dredge nets, and otter trawls in

the Apalachicola Bay system (1975-1983). Data are derived from Livinqston et al.

(1976c, 1977), McLane (1980), Purcell (1977), Mahoney (198?), and Sheridan (1^78,

1979). Recent taxonomic updates are noted in Livingston et al. (1983).

Phylum - Mollusca

Class - Gastropoda

Subclass - Prosobranchia

Order - Archaeogastropoda

Family - Neritidae

Neritina reel i vat a Order - Mesogastropo^a

Family - Calyptraeidae

Crepidula fornicata

Crepidula plana

Family - Naticidae

Polinices duplicatus

Family - Epitoniidae

Epitonium rupicola

Family - Hydrobiidae

Texadina

sphinctostoma

Family - Cerithiidae

Bittium varium

Order - Neogastropoda

Family - Fasciolari idae

Fasciolaria tulipa

Family - Melongenidae

Busycon contrarium

Busycon spiratum

Melongena corona Family - Muricidae

Urosalpinx perrugata

Family - Col umbel 1 idae

Anachis avara Mitrella lunata

Family - Olividae

1 i ve 1 1 a sp. Family - Thaididae

Thais haemastoma

Family - Marginellidae

Prunum apicinum

Subclass - Opisthobranchia

Order - Cephalaspidea

Family - Bull idae

Bulla striata

Family - Retusidae

Retusa canal iculata

Family - Pyrami dell idae

Odostomia laevigata

Order - Anaspidea

Family - Aplysiidae

Aplysia wiUcoxi

Order - Nudibranchia

Nudibranch sp.

Class - Bivalvia

Bivalve sp. ? Bivalve sp. x Order - Mytiloida

Family - Mytil idae

Amygdalum papyri a Brachidontes exustus

Brachidontes sp.

Order - Arcacea

Family - Arc idae

Anadara

Anadara

Anadara

brasil iana

sp. transversa

Order - Ostreoida

Family - Ostrei idae

Crassostrea virginica

Order - Veneroida

Family - Cyrenoididae

Pseudocyrena f loridana

Family - Mactridae

Mactra fragil is

Mul inia lateral is

Rangia cuneata

Family - Sol en idae

Ensis mino r Family - Tellinidae

Macoma bal thica

Macoma mitchel li Tell ina texana

Famil y

- Semelidae

Abra aequalis

Family - Solecurtidae

Tagelus plebeiu s Family - Dreissenidae

Mytil ops is leucophaeta

Family - Corbiculidae

Polymesoda carol ini ana Family - Card i idae

Dinocardium robustum

Class - Cephalopoda

Order - Teuthoidea (= Decapoda)

Family - Loliginidae

Loll iguncula brevis

Class - Polyplacophora

Family - Chitonidae

Chiton tuberculatus

Phylum - Annelida

Class - Polychaeta

Polychaete (unident.)

(continued)

50

Table 16. (Continued. )

Phylum - Arthropoda

Subphylum - Crustacea

Class - Malacostraca

Superorder - Peracarida

Order - Mysidacea

Mys i do ps i s almyra

Mysidopsis bahia

Mysi dopsis b 1 qe 1 ow Taphromysis bowmani

Taphromysis

louisianae

Order - Tan a i dace a Harqeria rapax

Order - Cumacea

Cumacea sp. Order - Isopoda

Family - Anthuridae

Cyathura polita

Xenanthura

brevitelson

Family - Sphaeromatidae

Cassidinidea ovalis

Sphaeroma

quadridentatum

Sphaeroma terebrans

Family - Idoteidae

Edotea montosa

Edotea sp. (cf . montosa ) Erichsonella sp. (cf. filTf"ormis ) Family - Munnidae

Munna reynoldsi

Order - Amphipoda

Family - Ampeliscidae

Ampelisca abdita

Ampelisca vadorum

Ampelisca

verril 1 i Family - Melitidae

Melita

appendiculata

Melita elonqata

Melita fresnelii

Family

Family

Suborder

Family

Suborder

Fami ly

Family -

Family -

Caprell idea

Caprellidae

Paracaprella

tenuis

Gammari dea

Haustoridae

Lepi dactyl us sp. Haustoridae sp. Gammari dae

Gammarus

macromucronatus

Gammarus

mucronatus

Gammarus so. Bate i dae

Batea

catharinensis

Carinobatea sp.

Melita

intermedius

Me"TTta lonqisetosa

Melita nitida

Mel ita sp. Ischyroceridae

Cerapus sp. (cf. tubularis ) Erichthonius

brasil ie"ris"is Erichthonius sp. 2 Aoridae

Grandidierella

bonnieroides

Grandidierella

sp. Lembos sp. Microdeutopus sp. Corophiidae

Corophium

louisianum

Corophium sp. Family - Cranqonyctidae

Cranqonyx

richmondensis

Family - Amphil ochidae

Gitanopsis sp. Family - Ampithoidae

Cymadusa compta

Cymadusa sp. Family - Tal itridae

Orchestia qril lus

Orchestia uhleri

Family

(continued)

52

Table Ifi. (Concluded.)

Family - Porcel lanidae

Petrol isthes

arm at us Clibanarius

vittatus
evident. Tnfaunal numerical abundance and

dry weiqht biomass (Figure ?7) in East

Bay, Apalachicola Bay, and St. George

Sound usually peak during winter and early

spring months (Mahonev and Livingston

1982; Livingston l'583b, c; Livingston et

al. 1983). Numbers of infaunal species

reach the highest levels during winter and

spring months (Figure ?J) . Monthly

variance follows the trends of numerical

abundance and soecies richness. Sheridan

and Livingston (1983), working in shoal

grass ( Halodule wrightii ) meadows on the

north shore of St. George Island, found

infaunal densities exceeding 104,000

individuals m"'' in April 1^15.

Spatial gradients of salinity,

oroductivity, and sediment types influence

the infaunal community composition

(Livingston et al. 1983). While physical

•"actors appear to predominate in the

infaunal community relationships in the

upper estuary near the river mouth, other

factors such as predation pressure and

competition mav be important determinants

of such interspecific interactions in

polyhaline portions of the bay system

(Livingston et al . 1983).

Overall, infaunal soecies fall into

four general categories: crustaceans,

polychaetes, mollusks, and a miscellaneous

qroup that includes insect larvae and

oligochaete worms. Predominant species in

East Bay include Mediomastus ambiseta , Steblospio benedicti , Heteromastus

fil iformis , Ampel isca vadorum , Hobsonia

florida , Hargeria rapax , and

Grandidierel la bonnieroides . The tanaid

Hargeria raoax is most abundant in or near grass beds in Apalachicola Bav from

February to April. Other dominant grassbed soecies include Heteromastus

fil iformis and Hobsoni a florida . The

bonnieroides amphipod

ranges

Grandidierel 1 a throughout the East

Bay-Apal achicola Bay complex, with peak

abundances during early spring and late summer. Soft-sediment polyhaline

assemblages are dominated by Mediomastus

ambiseta , Paraorionospio pinnata , and

immature tubificid worms (Livingston et

al. 1^83). The sedentary polychaete

Heteromastus fil iformis is largely

restricted to grass beds and is most

abundant during April. The amphipod

Ampel i sca vadorum occurs primarily in the

Apalachicola Bay seagrass meadows during

winter and early fall months. The polychaete Mediomastus ambiseta is found in

fine mud bottoms throughout the bay, with

peaks of abundance in March. The

ubiquitous polychaete Streblospio

benedicti utilizes a variety of habitats

throughout the estuary, with peak

abundance during winter months. The

polychaete Hobsonia florida is found

throuqhout the bay from qrass beds to soft

sediment (unveqetated areas). Peak

abundance is noted during early fall

months. Tn general, the polychaete

species are eurythermal and euryhaline and

include selective and nonselective deposit

feeders. Sheridan and Livingston (1983)

noted that the dominant tanaids and

amphipods are detritivores and deposit

feeders.

Because considerable amounts of

detrital matter are usually swept into the

estuary by the Apalachicola River during

winter-spring periods, the organic litter

forms an important habitat for various

macroinvertebrates. Organisms associated

with leaf litter and detritus have been

described by Livingston (1978) and

Livingston et al. (1976b, 1977). Litter

fauna is dominated by isopods, amphipods,

and decapods, which utilize particulate

matter and litter-associated microbes for

food and/or shelter. Dominant species in

East Bay and Apalachicola Bay include

Neritina reclivata , Palaemonetes spp.,

Corophium louisianum , Gammarus spo.,

Grandidierella bonnieroides , Mel ita spo.,

and Munna reynoldsi . Salinity appears to be an important orqanizinq feature of

litter associations (Livinqston unpubl.).

Life-history strategies of dominant

infaunal and litter-associated

macroinvertebrate populations are dictated

by substrate type, temperature, salinity,

and biological factors (Table 17). Most

dominant infaunal populations reach peaks

of numerical abundance during late winter

and spring periods of low salinity and

increasing temperature. Most such soecies

are euryhaline and eurythermal.

Reproduction of some infaunal populations

occurs throughout the year while others

reproduce only between spring and fall.

Individual species have different patterns

of distribution within the estuary depending on recruitment patterns and response

55
• I 10 s- c <u 3 +-> 10 O "O

4- <0 OJ C J- •!-> •r- 1^ VI ^ x: -f- u I— E vo a> CO q; Si. •f— .,- (O c u

«o OJ ''^ en Q. 0)

J. <rt ••-' o o ^ °^

5 8

4'

>.

O) O) O .rI— -c "t: TO <-> * 3 (0

••-

-o a; "Z

<0 *^ 3

O •- 0)

(O V> I—

E 01 3

S- <-> cn o <o M- S- S-

•r— OJ XI

O) 1. o

c > c

(O c "" ?" -

•— *

'^

n3 vT ?

5n <^ i

c trt -, <U .r- -g

r-^ r '5

<U <U O

I— .Q •!- ^ -r- -!-> fO Q. W

C t/l

*J .— -i^ ^

u •» (u o r^

_ r-l Q. -t-J CTi u r-. O 1/1 •—

^ .— TO 4-j cj^ u a

"O c •— -^ q; C — ^ 3

>-U-, . ^ "C

J QJ l/l

: t: oi

3 C O

• CD

4_> f-- IT?

O^ -C •—< CD

J O -— CT ^ x: a c

3 a' fc x: 3

,
Oj .^ .

?500

6000-

1500

3000

1500- ilM

Ki ,9:

CNJ
to stress. However, there is relatively

little in the way of detailed life-history

information concerninq these invertebrate

species.

4. a. OYSTERS

Oysters ( Crassostre a virqinica ) represent an important part of the biota

of the Apalachicola estuary (Fiqure 20).

Such factors as temperature, rainfall/

river flow (and hence salinity),

productivity (al lochthonous and

autochthonous), bottom type, and predation

define the life history of oysters in the

Apalachicola estuary. Ingle and Dawson

(1951, 1952) noted that temperature is

rarely limiting and that the spawning

season is one of the longest in the United

States (April through November). The

f ree-swimminq larval stage persists for

two weeks. Ingle and Dawson (1952) found

that oyster growth in Apalachicola Bay is

the fastest in the United States and is

continuous throughout the year because of

the relatively high year-round

temperatures. Successful oyster

development depends on an appropriate

substrate such as oyster shells, which can

be d1 anted throughout the estuary as cultch to enhance growth. Whitfield and

Beaumariage [l^^ll) estimate that nearly

40^ of Apalachicola Bay is suitable for

growing oysters. The ample nutrients and

primary production of the bay also enhance

oyster growth.

Oyster-bar associations also include

various organisms that prey on oysters

(Menzel et al. 1^58, 1966). These include

boring sponges, polychaete worms,

gastropod mollusks (such as Thais

haemastoma and Melongena corona ), and

crustaceans ( Menippe" mercenari a )

. Salinity is the most important limiting

factor for oyster populations, but it has

been hypothesized that such influence is

indirect in that low salinity limits

predation by excluding important species

such as Thais and Menippe . During periods

of high salinity, oyster predation is

enhanced and can be considerable.

Experiments have shown that oysters over

50 mm in length are rare in unprotected

areas of high salinity relative to areas where oysters are shielded from predation

by baskets at similar salinities (Menzel

et al. 1%6).

4.5. NEKTON

Nekton are those organisms that are strong enough swimmers that they can move

through the water column, even aqainst

water currents. In the Apalachicola Bay

system, the nekton comprise the bulk of

the sport and commercial fisheries and are among the more consoicuous biological

components of the estuarv. Eoibenthic

fishes and invertebrates in the

Aoalachicola marshes (Table IS) and ooen water areas (Table 1°) are characterized

by high numbers of oredominant soecies,

with the top three species of each group

accounting for 70'?;-RO'3^ of the total

numbers taken throughout the year. The

relatively low number of fish and

invertebrate species in the bay system at any given time, together with the high

dominance of a relatively few extremelv

successful species, contribute to the low

species diversity throughout the estuary

(Livingston l'376b).

In a given year, peak numbers of

fishes tend to occur from February through

Aoril (Figure 27). This situation is due

largely to the presence of juvenile soot

and Atlantic croaker. Species numbers, on the other hand, tend to oeak during

October. Epibenthic invertebrates reach

abundance peaks from August through

October, largely because of high numbers

of penaeid shrimp and, secondarily, blue

crabs (Figure 27). Seasonal patterns of

invertebrate species richness tend to follow those of the fishes. The highest

numbers of invertebrate soecies usually

occur in October. The peaks of abundance

and species richness of fishes and

invertebrates are characterized by monthly

high variances.

Various organisms appearing in the

estuary may not be estuarine dependent

throughout their life histories. Many

such organisms are migratory. The

anadromous soecies in the Apalachicola

drainage system include the Atlantic

sturgeon ( Acipenser oxvrhynchus ), Alabama

shad ( Alosa alabamaej^ and striped bass

( Morone saxatil iT) (Yerqer 1<^77). The

skipiack herrinq ( Alosa chrysochloris ) is

another possible anadromous species.

Other species, such as the Atlantic

needlefish ( Stronqylura marina ) may be

diadromous. Catadromous species include

60
Table 18. Fishes and invertebrates commonly taken

with seines in oligohaline (East Bay) and mesohaline

(Apalachicola Bay) marshes of the Apalachicola estuary

(from Livingston and Thompson 1975).

Species

Scientific name Common name

East Bay

Fishes

Ictalurus natalis

Micropterus salmoides

Lepomis microlophus

Lepomis punctatus

Poecil ia 1 at i pinna

Adinia xenica

Cyprinodon variegatus

Fundulus qrandis

Fundulus confluentus

Fundulus si mil is

Notemigonus crysoleucas

Lucama parva

Lucania goodei

Notropis sp. Lepisosteus osseus Cyprinus carpi o Anguil la rostrata

Pomoxis nigromaculatus

Menidia beryl 1 ina

Anchoa mitchil li Brevoorti a patronus

Mugil curema

Mugil cephalus

Micropogomas undulatus

Bairdiella chrysoura

Stellifer lanceolatus

Cynoscion arenarius

Paralichthys lethostigma

Trinectes maculatus

Eucinostomus gula

Lutjanus griseus

Gobiosoma bosci

Microgobiu"s~gu1osus

Archosargus probatocephalus

Invertebrates

yellow bullhead

largemouth bass

redear sunfish

spotted sunfish

sailfin molly

diamond killifish

sheepshead minnow

gulf killifish

marsh killifish

longnose killifish

golden shiner

rainwater killifish

bluefin killifish

shiners

longnose gar common carp

American eel

black crappie

inland silverside

bay anchovy

gulf menhaden

white mullet

striped mul let

Atlantic croaker

silver perch

star drum

sand seatrout

southern flounder

hog choker

silver jenny

gray snapper

naked goby

clown goby

sheepshead

Callinectes sapidus

Palaemonetes pugio

Penaeus setif erus Penaeus aztecus

blue crab

grass shrimp

white shrimp

brown shrimp

(continued)

61
Table 18. (Concluded.)

Species

Scientific name Common name

Aoalachicola

Fishes

Bay

Anchoa mitchil 1 i Anchoa hepsetus

NIemdia beryll ina

Eucinostomus quia

Synodus foetens

Stronqyiura marina

Lucania parva

Fundulus similis

Synqnathus floridae

Lagodon rhomboides

Leiostomus xanthurus

BairdieTTa chrysoura

Cynoscion ne^bulosus

Mugil cephalus

Orthopristis chrysoptera

Opsanus beta

Invertebrates

Callinectes sapidus

Palaemonetes pugio

Palaemonetes vulqaris

Palaemonetes intermedium

Penaeus setiferus

Penaeus duorarum

Penaeus aztecus

Neopanope texana

bay anchovy

striped anchovy

inland silverside

silver jenny

inshore lizardfish

Atlantic needlefish

rainwater kil lif ish

longnose kil lif ish

dusky pipefish

pinf ish

spot

silver perch

spotted seatrout

striped mullet

pig fish

gulf toad fish

blue crab

grass shrimp

grass shrimp

grass shrimp

white shrimp

pink shrimp

brown shrimp

mud crab

the American eel ( Anquil la rostrata ) , hoqchoker ( Trinectes maculatus ), and

mountain mulTet ( Agonostomus monitcola )

. Various other freshwater species and some marine forms, such as strioed mullet

( Muqil cephalus ) and the southern flounder

( Paral ichthys lethostiqma ), occur in the

lower river and estuary although thev do

not make true migrations.

The estuarine dominants such as sciaenid fishes, penaeid shrimp, and blue

crabs have annual migrations during which

the adults spawn offshore, the larval and

juvenile stages move into the estuarine

nursery, and finally the subadults return

to the open gulf to spawn as adults. Most

such species are either marine-estuarine

or estuarine. Oesterling and Evink (1°77)

studied migratory habits o^ blue crabs

along the Gulf coast of Florida (Figure

?R). Adult blue crabs spawn offshore and

the larvae, after going through a series

of zoeal (planktonic) stages, metamorphose

into a single megalops staqe that has both

Dlanktonic and benthic features (Figure

28). The meqalops eventually molts into

the first crab stage, which develops

mainly within the estuarine nursery

grounds. The authors found that female

crabs move northward alonq the oulf coast

of Florida, some as far as SDO km. Few

males move more than 40 or 50 km. Such

migrations appear to be linked to spawning

within the Apalachicola offshore area (from the Ochlockonee River drainage to the Apalachicola River drainaqe). Larqe

numbers of egg-bearing females are

62
Table 19. Epibenthic fishes and invertebrates taken in otter trawls and

trammel nets at various stations in the Apalachicola estuary from 1972

through 1982 (Livingston unpublished data). Species are listed in order of

numerical abundance.

Species

Fishes

1, Anchoa mi tchil 1 i 2. Micropogonias undulatus

3 . Cynoscion arenarius

4 . Leiostomus xanthurus

5. Polydactylus octonemus

6. Arius feli s 7. Chloroscombrus chrysurus

8. Menticirrhus amencanus

9. Symphurus plagiusa

10. BairdielTa chrysura

11. Etropus crossotus

12. Trinectes maculatus

13. Prionotus tribulus

14. StellifeF lanceo Tatus

15. Anchoa hepsetus

1 6 . Porichthys porosissimus

17. Prionotus scitulus

18. Eucinostomus aula

19. Paralichthys letiTostigma

20. Synodus foetens

21 . Eucinostomus arqenteus

22. Dasyatis sabina

23. Cynoscion nebulosus

24. Microgobius thalassinus

25. Urophycis floridanus

26 . Lagodon rhomboides

27. Gobiosoma bosci

28. Chaetodipter us f aber

29. Orfhopristis chrysoptera

30. Brevoortia pat ronus

31 . Dorosoma petenense

32. Peprilu s burti

33. Peprilus paru

34 . Stephanolepis hispi dus

3 5 . Sphaeroides nephe 1 us_ 36 . Ophichthu s gomes f

~

37. Synqnathus louisTanae

38 . Syngnathu s scovelli

39. Gobionellus boleosoma

40. Harengul a pensacolae

41.

42.

43.

44.

45.

46.

47.

48.

4q.

50.

51.

52.

53.

54.

55.

56.

57.

58.

60.

60.

61.

62.

63.

64.

65.

66.

67.

68.

69.

70.

71.

72.

73.

74.

75.

76.

77.

78.

79.

1 . Penaeus setif eru s 2. Call inectes sapidus

3. Palaemonetes puqio

Archosarqus probatocephalus

Microgobius gulosus

Bagre marinus

Menidia beryl 1 ina

Monacanthus cil iatus

Caranx hippos

Centropristis melana

Syngnath us flondae

Ancyclopsetta quadrocellata

Chilomycterus schoepf i Pi plectrum formosum

Tctalurus catus

Sciaenops ocellata

Astroscopus y-qraecum

Hippocampus erectus

Leoisosteus osseus Lucanis parva

Lutjanus qriseus

Opsanus beta

Paralichthys albigutta

Ophidion beani

Aluterus schoepf i Diplodus" holbrook'i

Gobionellus hastatus

Hypsoblennius hentzi

Menticirr hus saxatil is

Myrophis punctatus

Oqilbia cayorum

Oligopl it es sauru s Pomatomus saltatrix

Rhinoptera bonasus

Scomberomorus maculatus

Selene vomer Sphyraena boreal i s Sphyrna tiburo

Sardinella anchovia

Caranx bartholomaei

B. Invertebrates

4.

5.6.

(continued)

63

Mugil sp. Gymnura mi crura

Penaeus duorarum

Trachypenaeus constrictus

Chrysaora quinquecirrha
Table IP. (Concluded.)

Species

B. Invertebrates (continued]

7. Lolliquncula brevis

8. Penaeus aztecus

^. Palaemonetes vulgaris

10. Portunus gibbesi i 11. Stomolop'hys me1e¥gris

12. Neritina reclivata

13. Squil la empusa

14. Callinectes similis

15. Rhithropanopeus harrisi i 16. Neopanope texana

1 7 . Polinices duplicatus

18. Neopanope packardii

19. Mulinia Taterali s 20 . Acetes americanus

21. Pagurus pollicarTs

22. Rangia cuneata

23. Menippe mercenaria

24. Xiphopeneus kroyeri

25. Alpheus heterochaelis

26. Latreutes parvuTTTs

27. Palaemonetes intermedius

28. Metoporhaphis calcarata

29. Crassostrea virginica

30. Palaemon floridanus

31. Periclimenes longicaudatus

32. Ogyrides li mi col a 33. Trachypenaeus similis

34. Busycon contrarium

35. Branchiosychis americana

36.

37.

38.

39.

40.

41.

42.

43.

44.

45.

46.

47.

48.

49.

50.

51.

52.

53.

54.

55.

56.

57.

58.

59.

60.

61.

62.

63.

64.

Brachiodontes exustus

Hexapanopeus angustifrons

L u i d i a clathrata

Persephona mediterranea

ClibanarTus vittatus

Libinia dubia

Periclimenes americanus

Ambidexter symmetricus

Busycon spiratum

Procabarus paeninsulanus

Eupleura~u1cidentata

Hemipholus elonqata

Alpheus normanni

Eurypanopeus dep^ressus

Lysmata wurdemanni

Pentacta sp. Petrolisthes armatus

Podochela rii se i Tozeuma carolinense

Nudi branch sp. Alpheus armil latus

Sesarma cinereum

Sicyonia dorsalTs

Anadara brasil iana

Dinocardium robustum

Cantharus cancel 1 aria

Urosalpinx perrugata

Ova li pes guadulpensis

Pagurus lonqicarpus

concentrated in this area in winter. The

authors hypothesized that larval dispersal

from the Apalachicola area takes place

along clockwise (Lood) currents that

eventually wash onto the Florida Shelf

(Figure 28). '^oea larvae then disperse

along the coast, with the megalops stage

settlinq into the coastal estuaries.

Livingston et al. (1°77) used daytime

trawling to estimate winter populations of

juvenile blue crabs in the Apalachicola

estuarv of approximately 30,000,000

individuals. Miqration of spawning

females aopears to coincide with flooding

of the north Florida drainaae system,

which makes particulate organic matter

available as food to the young crabs

(Lauqhlin 107Q). Thus, the migration of

blue crabs along the gulf coast could be

tied to both the reproductive

characteristics of the species and the

trophic organization of the Apalachicola

estuary.

Life-history features of the dominant

epibenthic soecies in the Apalachicola

estuary have the same patterns as elsewhere in the northern Gulf of Mexico

(Table 17). Spawning and recruitment

generally vary from species to species

according to different combinations of

seasonal physical factors. The bay

anchovy is the most abundant fish and is one of the few fish species that does not

show regular seasonal recruitment

progressions. In contrast, the Atlantic

64
megalops

Figure 28. Life cycle of the blue crab

along the gulf coast of Florida.

Ovigerous females move toward the

Apalachicola estuary. It is hypothesized

that developing stages move back down the

gulf coast of Florida with offshore

currents (after Oesterling and Evink

1P77).

croaker spawn near passes during fall and

early winter; the iuveniles occupy the

estuary in peak numbers during late winter

and early spring when salinities are usually less than 10-15 opt. Spot also

spawn near passes, and peaks of abundance

in the estuary generally coincide with

those of the Atlantic croaker. Sand

seatrout are usually most abundant during

summer months after spawning offshore

during the spring. This species is taken

at various salinities, but temperature

appears to be limiting; high catches are generally taken in ?0O-350-C water.

White shrimp are dominant from August

to November, with spring spawning and

recruitment. Other penaeids usually reach

peak numbers during late spring (brown

shrimp: Penaeus aztecus ) or late summer

(pink shrimp: P_. duorarum )

. The blue

crab shows a bimodal annual peak of

recruitment; numbers peak during winter

and summer periods. Oeoth and specific

microhabitat conditions are the principal

determinants of blue crab distribution at

any given time (Laughlin 1<570; Livingston

unpubl.). The brief sguid (Lol liguncula

brevis ), is limited in spatial/temporal

distribution by salinitv ('O-'^O ppt) and

other habitat characteristics and complex

trophic relationshios (Laughlin and

Livingston 1P8?). In summary, these

species-specific responses to multifactor

complexes demonstrate the difficulty of

trying to design linear models to explain

and oredict spatial/temporal patterns of occurrence.

The spatial distributions of nektonic

fishes and invertebrates in the

Apalachicola estuary (Table 20) tend to be

associated with freshwater runoff into the

system. Relative dominance at a given

station varied according to salinity

gradients and habitat type. Regular

seasonal changes in distributions are evident for most of the dominant nektonic

species. For example, anchovies are relatively uniformly distributed within

the estuarv during January and February

(Figure 7°). By the spring, anchovies ^re

concentrated in upper portions of East

Ray. Huring the early summer, there are minor population peaks with orimary

concentrations in eastern portions of East

Bay. Bv the fall, the anchovies

concentrate around the mouth of the

Apalachicola River as well as in portions

of East Bay, and during early winter, the

anchovies become uniformly distributed

throughout East Bay and Apalachicola Bay.

In January, Atlantic croaker tend to conqregate at the mouth of the

Apalachicola River and upper oortions o*" East Bay (Figure ?0). Bv February, this

distribution is more uniform throughout

East Bav and northern Apalachicola Bay, a situation that appears to hold during

ensuing winter and spring months until,

by May or June, the croakers move out of

the bay.

The spatial distribution of sand

seatrout through a given seasonal cycle is

guite regular (Figure 31). As the young

seatrout move into the bay svstem in May,

thev concentrate in upper portions o^^ East

Bay and just off the mouth of the

Apalachicola River. Secondary concentrations are found throuqhout East Bay and

northern oortions of Apalachicola Bav. The distribution changes little in June,

65
Table ?0. Epibenthic fishes and invertebrates taken in otter trawls at permanent

stations in the Apalachicola estuarv from June 1^7? to Mav 1'377. Stations have been

ordered by cluster analysis according to relative abundance of fishes and

invertebrates. Data are given concerning numbers/sample, dry weight biomass/sampl e, percent dominance (by numbers), and Margalef richness. Dominant species are also

enumerated by station.

Number Biomass per % Dominper sample (a, ance (by

Station samp le dry wei qh t^) numbers) Dominant species

Margalef

richness

OUTER BAY1— IX 73.?

A. FISHES

- 1
Table 20. (Continued.

Number Biomass per % Dominper sample (q, ance (by

Station sample dry weight) numbers) Dominant species

Margalef

richness

A. FISHES (continued)

SIKE'S CUT IB 20.6 129.3 36 ANCHOA MITCHILLI

CYNOSCION ARENARIUS

ETROPUS CROSSOTUS

4.92

B. INVERTEBRATES

OUTER BAYRIVER

DOMINATED

- 1 7.0 7.2

I— lA 5.5 5.3

I— 4 14.7 16.8

47 CALLINECTES SAPIDUS 2.58

PENAEUS SETIFERUS

38 PENAEUS SETIFERUS 1.86

CALLINECTES SAPIDUS

LOLLIGUNCULA BREVIS

TRACHYPENAEUS CONSTRICTUS

IE
Table 20. (Concluded.)

Number Biomass per % Domi ri- per sample (q, ance (by

Station sample dry weight) numbers) Dominant species

Marqalef

richness

I— 4A 13.0 — 5

UPPER -

(EAST) BAY

5A — 58

12.2

13.7

6.8 — 5C 12.5

I— 6 45.8

SIKE'S CUT IB 10.0

B. INVERTEBRATES (continued)

16.0






be attracted to the upper freshwater

portions in the estuary.

Although the major peaks in numbers

of juvenile blue crabs occur during the

winter, secondary increases are often

noted during the summer and fall (Figure

34). As the young blue crabs enter the

Aoalachicola estuary during the winter

months, they concentrate in East ^ay and

off the Nick's Hole drainage (St. George

Island). During May and June, oeaks in

the number of blue crabs occur in these

areas. By the summer and fall months, the

blue crabs are concentrated in East Bay.

Blue crabs appear to be attracted to areas

that receive overland runoff although they

are not attracted by direct river flow.

While there is a qeneral pattern of

concentration of the dominant epibenthic

fishes and invertebrates in areas that

receive direct input of freshwater runoff

from upland areas, it is simplistic to assume that runoff per se is the primary

factor that influences the temporal and

soatial asoects of the distribution of

such organisms in the estuary. There are,

in fact, a complex o^" species-specific

limiting factors that are associated with

the trophic organization of the bay

system.

75
CHAPTER 5

NICHE DIVERSITY, TROPHIC INTERACTIONS, AND COMMUNITY STRUCTURE

5.1. HABITAT-SPECIFIC ASSOCIATIONS

The Apalachicola estuary, as an ecosystem, can be defined as a series of

habitats with associated assemblages of

organisms. Such assemblages (or communities) live in the same general habitat,

compete for space and food, and are part

of the highly complex trophic structure of

the river-bay system. The dimensions of a given community are difficult to define

precisely because the component

populations vary considerably in their

distribution and community function in

space and time. However, selected factors

can be used to characterize the various

estuarine assemblages. Sources of primary

productivity, habitat features, the

physical and chemical environment

(including pollutants), modes of

reproduction and recruitment, feeding

interactions, predator-prey relations, and

competition are some of the features that

shape the estuarine communities.

The distribution of most of the

estuarine assemblages may be partitioned

into the following habitats: marshes,

seagrass beds, litter associations, oyster

bars, and subtidal unvegetated (softsediment) areas. Many of the long-term

biological studies in the Apalachicola

estuary have concentrated on the macro- invertebrates (benthic, epibenthic) and

fishes that are found in these areas.

S.1.1. Marshes

The marshes, which include complex

patterns of tidal channels and small

creeks, provide food and habitat for a number of organisms in the Apalachicola

estuary (Table 18). Marsh complexes

include insects, mollusks, crustaceans,

fishes, birds, and mammals. Topminnows of

various species are dominant in such areas. Many species that are important to the sports and commercial fisheries of the

region spend at least part of their life

histories in the estuarine marshes. Such

species include blue crabs, penaeid

shrimp, large-mouth bass, lepomids,

striped mullet, spotted and sand seatrout,

and anchovies. Few species spend their

entire lives within the marshes, however,

and the marsh habitat is best

characterized as a nursery for migratory

species during summer and fall months.

5.1.2. Seagrass Beds

The distribution of grassbeds in the

Apalachicola estuary (Figure 19) is the

result of a number of environmental

controlling factors. Even though it is

limited to only about 10% of the aquatic

area by the high turbidity and

sedimentation associated with the river,

this habitat's productivity is high.

Grassbed productivity is also limited by

water temperature, salinity, and the

activity of certain invertebrates.

However, grassbeds also have an effect on certain water quality indices. Various

studies in East Bay (Livingston 1978;

Purcell 1977) indicate that water quality

factors such as dissolved oxygen and pH are higher in the grassbeds than in

associated mudflats.

The oligohaline grassbeds of East Bay

are dominated by tapeweed ( Val isneria

americana ), a freshwater species. Other

species found in conjunction with tapeweed

are Potamogeton pusil lus , Ruppia maritima

(locally dominant in western bayous of

East Bay), Cladophora sp., and Halophila

engelmanni . In recent years, some parts

of East Bay are being taken over by the

Eurasian watermilfoil ( Myriophyllum

76
spicatum ). During the period 1980-1981,

this introduced species became dominant in

Round Bay, one of the eastern bayous. By

1982-1983, the Myriophyllum had become

rooted throughout the upper East Bay area (Livingston unpubl.). It is unclear how

spread of Eurasian watermilfoil will

affect the distribution of plants and

animals in the East Bay seagrass beds.

Currently, the oligohaline seagrass

beds serve as a nursery for benthic

species such as the snail Neritina

reclivata (a major dominant! and

epibenthic species ( Udostomia sp.,

Gammarus macromucronatus and Taphromysis

bowman i )". Infaunal assemblages are dominated by polychaetes ( Loandalia

americana , Mediomastus ambiseta ),

amphipods ( Grandidierella bonnieroides ) and chironomid larvae ( Dicrontendipes

sp.). Fish populations are dominated by

rainwater killifish ( Lucania parva ),

pipefish ( Syngnathus scovell i ),

silvers ides ( Menidia beryl lina")^ gobies

(Microgobius gulosus ), and centrarchids.

Many species utilize these areas (Duncan

1977; Livingston and Duncan 1979; Purcell

1977). Of the 28 dominant benthic species

of fishes that comprised over 98% of the

abundance in the area, most consumed

detritus, small mollusks, crustaceans,

epiphytes, and insect larvae. Most of the

penaeid shrimp, insect larvae, and fishes

that are found here are seasonally

abundant at early stages of their

reproductive cycles, which indicates the use of these areas as primary nursery

grounds. Peaks of abundance are staggered

throughout the year.

The predominant macrophyte species in

mesohaline or higher-salinity areas off

St. George Island in Apalachicola Bay is

Halodule wrighti i (Sheridan and Livingston

1983) . Infaunal macroinvertebrates,

dominated by Hargaria rapax , Heteromastus

f i 1 if ormis , Ampel isca vadorum and various

oligochaetes, reach peaks of abundance

during early spring. Predominant fishes

include silver perch ( Bairdiel la

chrysoura ), pigfish ( Orthopristis

chrysoptera ), pinfish ( Lagodon rhomboides ) and spotted seatrout ( CynoscTon

nebulosus ). These species are abundant

from May through September. Blue crabs

( Cal linectes sapidus ) , pink shrimp

( Penaeus duorarum ) ^and grass shrimp

( Palaemonetes vulgaris ) are the dominant

invertebrates. Their densities are bimodal, peaking in the winter and summer

months. These areas are also

characterized by the year-round presence

of larval and juvenile nekton.

5.1.3. Litter Associations

Leaf litter associations are dominated by omnivores and detriti vores. The fraction of particulate organic matter

(POM) large enough to be identified as litter is populated with gastropod

mollusks ( Neritina reclivata ), amphipods

( Gammarus mucronatus . Me 1 i t a spp.,

Grandidierella bonnieroides , Corophium

louisianum , Gitanopsis sp.), isopods

( Munna reynoldsi ) , and decapods

( Palaemonetes pugio , P^. vulgaris , Penaeus

setiferus , Cal linectes sapidus ).

Species richness of the litterassociated fauna in upper East Bay

(station 5A), the river mouth (station 3),

and the shoal qrassbeds off St. George

Island (station IX) peaks during August

and September (Figure 35). Such peaks are strongly associated with salinity levels

at the respective study sites (Figure 36).

Dominant species vary from location to location. The level and timing of peaks

of abundance also vary spatially (Figure

35). Upper East Bay, which is outside of

the direct influence of the Apalachicola

River, appears to be the least productive

part of the estuary in terms of litterassociated macroinvertebrates. Areas rich

in detritus, such as station 3, are most

highly populated during March and

September, periods when the river is

flooding or macrophytes are dying off.

The highest numbers of litter-associated

macroinvertebrates occur in the Halodule

beds off St. George Island from April to June, a period of high macrophyte

productivity. These data indicate that

while species richness may be strongly

influenced by salinity, the numerical

abundance of the litter associations is more strongly aligned with the

availability of detritus.

While physical factors such as salinity and temperature are important

determinants of the distribution of

litter-associated organisms in the

estuary, recent experiments by Florida

77
LU

<

LU

CO

L.O

a:

UJm

3

<Q

Z

m

<

_i<o

tr

iij

25

15-

20,000-

15,000-

10,000-

5000-

Stotions

I I I I I I I I I I I I JFMAMJJASOND

TIME

Figure 35. Numerical abundance and

SHPcies richness of invertebrates taken in

leaf-litter baskets at various permanent

sampling sites in the Apalachicola

estuary, monthly from January, 1976,

through Pecember, 1976. After Livingston

(1978) and Livingston et al. (1977).

State University researchers indicate that

biological associations are also

important. Macroinvertebrates appear to utilize the detritus as shelter and a source of food (White in press). In a series of experiments with the leaf litter

community. White et al. (1979a) found

that, whereas the biomass (as measured

bv lipid phosphate and

poly-beta-hydroxybutyrate), nutritional

history, and respiratory activity of

microbes are correlated with substrate

type, the macrofaunal populations are more

often associated with specific water

quality features such as salinity.

Numbers, biomass, and species richness of

detritus-associated microfauna are associated with the mass and community

structure of the macrofaunal food web.

These macroinvertebrates apparently seek

out microbial populations rich in
5.1.4. Oyster Bars

Oyster bars represent a relatively

significant habitat in the estuary (Table

1). The main concentrations of oysters

( Crassostrea virqinica ) (Fiqure 20) lie in

St. Vincent Sound and western portions of

St. George Sound, Oyster distribution is

dependent upon substrate, temperature,

salinity, and available food. Oyster

bars, themselves, provide habitat and food

for a variety of organisms. The oyster

associated community includes sponges

( Cliona vastifica), bryozoans

( Membranipora sp.), flatworms ( Stylochus

frontalis ), annelids ( Neanthes succinea . Polydora websteri ), various arthropod

crustaceans ( Callinectes sapidus , Menippe

mercenaria , Neopanope spp., Petrolisthes

armatus ),~ gastropods ( Crepidula plana , Melongena corona , Thais haemastroma and

pelecypods ( Brachidontes exusta, Chi one cancel lata ) (Menzel et al . 1966). Fishes

include blennies ( Hypsoblennius spp.) and

toadfish ( Opsanus beta ). These organisms

use the reef for shelter and/or feeding.

Salinity controls oyster-bar

community organization. When salinities

are high, various stenohaline gulf species

are able to move Into the oyster-rich

areas and feed on the oysters. Low

salinity limits such predation by acting

as a barrier to those organisms. Species

richness and diversity of the oysterassociated populations vary directly with

seasonal increases in salinity. During warmer months, extensive oyster mortality

in the Apalachicola estuary has been

attributed to infestation by the pathogen

Perkinsus marinus (formerly called

Dermocyctidium marinum ) (Menzel 1983).

Young oysters are unaffected by this

disease, although up to 505^ of adult

oysters may be killed annually. The

relatively long period of high water

temperature in the gulf estuaries

contributes to such mortality. A longterm study is currently under way to determine the response of the Apalachicola

oyster associations to various stimuli

including habitat features (water quality,

substrate), predation, competition,

disease, and possible over-fishing

(Livingston et al., unpubl.).

5.1.5. Subtidal (Soft-Sediment)

Communities

Almost 70% of the Apalachicola Bay

system can be characterized as a subtidal,

unvegetated, soft-sediment area (Table 1).

The muddy bottom substrate is inhabited

primarily by polychaetes ( Mediomastus

ambiseta , Streblospio benedict i ) and" amphipods ( Grandidierella bonnleroides ). The polychaetes are deposit and suspension

feeders with a high reproductive capacity

and considerable tolerance for low

salinity and variable environmental

conditions. Productivity trends, habitat

type, and the ecological characteristics

of the various populations contribute to

what is a temporally variable but highly

persistent assemblage of organisms in

terms of species richness, relative

abundance, and recruitment. In oligohaline areas of the estuary, the

benthic macroinvertebrate assemblages are characterized by high dominance, low

species richness, low diversity, and

varying standing-crop biomass and

numerical abundance (Livingston 1983c, d).

Areas around the mouth of the river have

much higher numbers of infaunal

macroinvertebrates than areas outside of

the region of general flow. Such

differences have been attributed

(Livingston 1983c, d) to the deposition of

nutrients and detritus by the river during

periods of flooding (Figure 9) and

Increased activity and abundance of the

benthic macroinvertebrates (Figure 27).

The general community characteristics

of the soft-bottom assemblages change as salinities increase temporally and

spatially. In mesohaline and polyhaline

portions of the system, overall numerical

abundance Is lower than in oligohaline

areas, but species richness and diversity

increase significantly (Livingston et al.

1983). Such trends are evident in the

associations of epibenthic fishes and

invertebrates, which are an Important part

of the soft-sediment communities.

Dominant populations such as Atlantic

croaker, spot, penaeid shrimp, and blue

crabs feed extensively on organisms within

the muddy bottom of the estuary.

The soft-sediment community

(invertebrates and fishes) of the

79
Apalachicola estuary reflects the response

of hundreds of species to a complex

combination of physical, chemical, and

biological factors. Physical control,

together with productivity features,

recruitment patterns, predator-prey

interactions, and competition for various resources determine to a considerable

degree the form and functions of the softsediment communities in the Apalachicola

Bay system. Because the majority of the

research in the Apalachicola Bay system

has been carried out with the fishes and

macroinvertebrates of the soft-sediment

estuarine habitat, the interrelationships

of the dominant features of these

biological systems will be treated in a more detailed fashion below.

5.2. PHYSICAL CONTROL OF BIOLOGICAL

PROCESSES

For some time, ecologists have argued

about the relative importance of physical

and biological control of aquatic

populations and communities. Clearly, the

problem is extremely complex, based on the

fact that each species is a product of a given habitat while also having an input,

through predation and competition, to the

community. It is generally agreed that

temperate estuaries such as the

Apalachicola system are highly productive

and physically unstable in space and time.

Temperature and salinity have a major

influence on the form and processes of the

estuarine biota in such a system. At the same time, various populations interact

with each other and their environment with

almost continuous feed-back to the system

as a whole.

evaluation of hypotheses derived from

observational data can then be used to determine the processes that define and

ultimately control the observed structural

components of the system.

Various attempts have been made to delineate the relationships of physical

and biological variables in the

Apalachicola estuary (Livingston 1975,

1976b, 1979, 1982b; Livingston and Loucks

1978; Livingston et al. 1974, 1976b, c, 1978; Mahoney and Livingston 1982; Meeter

and Livingston 1978; Meeter et al. 1979).

Most analyses indicate that Apalachicola

River flow has a major influence on the

physical and biological relationships in

the estuary. For example, statistical

analysis of the principal physico-chemical

variables (Table 21) indicates that the

main factor or component could be called

"river flow." This river flow is

associated with low salinity, increased

color and turbidity (and reduced Secchi

readings), and reduced chlorophyll _a. River flow alone explained 32% of the

total variance and about half of the

variance explained by the four factors.

Average bay values of major nutrients vary

seasonally; high nutrient concentrations

are found during high (winter) river

discharge and low salinity conditions

(Table 22). The Apalachicola River

controls to a considerable degree various

factors such as nutrient and detritus

concentrations, salinity, color and

turbidity, and other water quality

factors. Tn turn, these conditions

control the level and pattern of

productivity fluctuations in the bay

system.

The timed interactions of multiple

physical and biological components of an estuarine system are difficult to differentiate for a variety of reasons. Individual physical events follow

different temporal patterns. Often such

phenomena are essentially cyclic although

"cycle" does not necessarily imply that

there is a complete return to a previous

condition. Biological responses are not

that simple and often follow nonlinear or curvilinear patterns of response to varying controlling factors. Analysis of

biological responses requires the initial

delineation of key dependent and

independent variables. Experimental

Studies of temperate estuaries

indicate that the combination of high

primary productivity and extremely

variable environmental conditions is often

associated with relatively low species

richness and diversity and high secondary

productivity of a few dominant species.

No matter which group of organisms is

considered, from phytoplankton to fishes,

salinity appears to be the primary

regulator of species numbers at a given

location in the estuary. Dominants are able to adapt to low or highly variable

salinity conditions. Salinity is a major

determinant of species richness (S) of

80
Table 21. Factor analysis of physico-chemical variables in the Apalachicola system

taken monthly from March 1972 to February 1976. Color (Pt-Co units), turbidity

(J.T.U.), Secchi readings (m), salinity (ppt), temperature C^C), and chlorophyll _a (mg

1"1) were noted at Station 1. Tidal data included stages of the tide on the day of

collection while the wind variable was represented by two vector components (speed,

direction) (from Meeter and Livingston 1978).

Variable
Table 22. Correlation coefficients of linear regressions of nitrate, orthophosphate,

silicate, and ammonia on salinity (from Livingston et a1. 1974).

Date
different times of the year. Estabrook

(1973) noted that grazing zoopiankton also

may control phytoplankton productivity

since experiments removing zooplankton and

net plankton enhanced nannoplankton

productivity greatly. The possibility

exists that competition for nutrients

among various species also is an important

determinant of relative phytoplankton

dominance.

Among the zooplankton, copepods are dominant. The copepod Acartia tonsa

constitutes 95.5% of total zooplankton in

East Bay, 68.2% in Apalachicola Bay and

19.8% in coastal waters (Edmisten 1979).

Salinity and temperature control the

composition of zooplankton communities in

the estuary. Populations of Acartia vary

inversely with distance from the mouth of

the Apalachicola River and are concentrated in Apalachicola Bay.

Temperature is associated with significant

(p < 0.01) differences in Acartia numbers.

Salinity significantly (p < 0.01) affects

the overall relative abundance of the

dominant populations. Edmisten (1979)

showed that temperature, salinity, station

and month had a multiple r value of 0.775.

In East Bay, Acartia numbers (as well as zooplankton numbers and biomass) peak

during periods of high salinity. Thus,

temperature usually determines overall

numbers in the bay system, while salinity

determines their spatial distribution at any given time. The response to midrange

salinities explains the nonlinear

(parabolic) relationship of Acartia with

salinity,. It appears that other

organisms can successfully complete with

Acartia at higher and lower salinities.

Life history strategies of various

nektonic estuarine species depend to some degree on spatial/temporal gradients of

substrate type, salinity, food

availability, and energy flow. The

spatial distribution and abundance of

brief sguid ( Lolliguncula brevis ) is

determined to a considerable degree by

salinity and temperature (Laughlin and

Livingston 1982). Optimal salinities

range between 25 and 30 ppt. Squid tend

to congregate near the passes during

summer and fall periods of high salinity.

Distribution within the estuary is

associated with the distribution of

zooplankton in the bay. Population trends

of squid followed long-term (9-year)

salinity trends that, in turn, were associated with climatic features. There

were sharp decines in squid abundance

during periods of low salinity.

Overall, attempts to correlate

patterns of species abundance with

individual physical, chemical, and

productivity variables have not been

entirely successful. A multiple

regression analysis of individual

population densities with combinations of

independent variables indicates that such

components accounted for less than 50% of

the population variability (Table 23). No

single set of physical conditions

explained population variation through

time. While factors such as temperature,

salinity, productivity, and water quality

characteristics are important determinants

of general habitat availability, it is

clear that other factors, presumably

biological in nature, may be important to our understanding of the processes that

determine the community structure of the

Apalachicola Bay system.

5.3. TROPHIC RELATIONSHIPS AND FOOD-WEB

STRUCTURE

Community structure is determined in

part by predator-prey interactions,

especially among dominant estuarine

populations. Comprehensive studies of the

feeding habits of dominant fishes

(Sheridan 1978; Sheridan and Livingston

1979) and invertebrates (Laughlin 1979)

have been carried out (Figure 37).

Pelagic anchovies feed preimarily on calanoid copepods throughout their lives.

Seventy percent of the diet of young

anchovies (standard length (SL), 10-39 mm)

is composed of these copeoods. Larger

fish (SL 40-69 mm) eat mysids, insect

larvae and juvenile fishes. A seasonal

progression of food item consumption

follows trends of available prey species.

The Atlantic croakers progress through a series of distinct ontogenetic trophic

stages. Young fish (SL 10-30 mm) eat

insect larvae, calanoid copepods, and

harpacticoid copepods. "Midrange fish (SL

40-99 mm) consume detritus, mysids, and

isopods; larger fish (SL 100-159 mm) eat a high proportion of juvenile fishes, crabs,

and infaunal shrimp. Croaker at all

stages eat polychaete worms. Spot, which

83
Table 73. Results of a stepwise regression analysis of various independent parameters

and species (numerical abundance) in the Apalachicola estuary from March 1972 to

February 1^75. Independent variables are listed bv order of importance with R- expressed as a cumulative function of the given parameters (from Livingston et al.

1976b). Independent variables were run with and without lag periods of 1-:^ months.

Species Independent variables R?

Anchoa mitchil li Micropogonias undulatus

Cynoscion arenanus

Polydactylu

Arius felis

s octonemus

Leiostomus xanthurus

Chloroscombrus chrysurus

Menticirrhus americanus

Symphurus plagiusa

Bairdiella chrysura

Penaeus setif erus Palaemonetes pugio

Callinectes sapidus

Penaeus duorarum

Lolliguncula br?yis

Portunus gibbesii

Palaemonetes vuTgaris

Rhithropanopeus harrisi i Callinectes similis

ChloroDhyll a, Secchi

River flow (Tag), Secchi (lag)

Chlorophyll _a, wind, Secchi (lag) temp.

Chlorophyll _a (lag), salinity, Secchi

Temp., wind

Turbidity (lag), Secchi, salinity, temp.

Temp, (lag), temp., salinity

Temp, (lag)

Color (laa), color, Secchi

Wind, temp., color

Wind, chlorophyll _a, incoming tide, color

Turbidity

Secchi, incoming tide

Chlorophyl 1 a_, Secchi

a (lag), temp.

a (lag), Secchi

Chlorophyl 1 Chlorophyll

Turbidity

Wind

Chlorophyl 1 a, temp.

0.38

0.46

0.83

0.=^8 0.30

0.85

0.44

0.19

0.63

0.40

0.48

0.40

0.43

0.41

0.43

0.39

0.32

0.18

0.34

are also benthic omnivores, consume polychaetes, harpacticoid copepods, bivalves,

and nematodes. Spot have a more diverse

diet than croaker and do not concentrate

on single prey types. Trends across size

classes are not as clearcut, although

there is decreased specialization with

growth. The sand seatrout is a watercolumn predator of fishes and mysid shrimp

( Mysidopsis bahia ). Small trout (SL 10-29

mm) tend to eat mysids and calanoid

copepods, while larger fish (SL 30-8° mm) consume more juvenile fishes. Anchovies

( Anchoa mitchil li ) comprise 70^ of all

fishes taken.

Fishes regularly undergo ontoqenetic

dietary shifts encompassing planktivory,

carnivory, omnivory, and herbivory within

the same species (Sheridan 1978; Sheridan

and Livingston 1979; Livingston 1'379,

1982). Sheridan and Livingston (1979)

indicated that temporal differences in

feeding progressions were a major factor

in the lack of overlap in food types among

species. Laughlin (1979) found that blue

crabs also undergo trophic progressions.

Juveniles, abundant during winter months,

feed largely on plant matter, detritus,

and bivalve mollusks such as Rangia

cuneata , Brachidontes exustus , and

Crassost r ea virginica . As the crab grows,

bivalves and fishes become progressively

more important in the diet. Larger blue

crabs feed primarily on bivalves, fishes,

and crabs (i.e., blue crabs, mud crabs

such as rhithropanopeus harrisi , and

xanthid crabs of the genus Neopanope )

. Cannibalism is a significant mode of

foraqing in the older blue crabs. Diet

generally reflects seasonal shifts of prey

abundance.

Although the distinctive nutrient

sources for the estuary have been

identified, the rate functions of energy

movement through the system are little

understood. The periodic inputs of

nutrients and detritus into the estuary

are transformed into biological matter.

Such integrative processes continuously

smooth out the episodic nature of energy

84
Fiqure 37. Simplified feeding

associations of four dominant fishes--bay

anchovy, sand seatrout, Atlantic croaker,

spot--and blue crabs in the Apalachicola

estuary. Four food compartments are shown: phytopl ankton (P), holoplankton

(H), meroplankton and benthos (MB), and

sediments (S). Major food items in the

compartments are: DE=detritus,

8I=bivalves, HC=harpacticoid copepods,

NE=nematodes, IN=insects, PO=polychaetes,

SH=shrimp, MY=mysids, CR=crabs, FS=fishes,

CC=calanoid copepods, OI=diatoms. Numbers

indicate dry-weight contribution of

particular food items (within boxes) and

food contributions of major food

compartments (after Laughlin 1979 and

Sheridan 1978).

transfer from upland systems. The

planktonic and detrital pathways come together at the sediment level through

repackaging of fecal material and the

activity of the microorganisms. The

microbes transform dissolved nutrients

into available particulate matter. Over

2% of the dry-weight mass of the sediments

is composed of organic carbon, bacterial

biomass, and extracellular polysaccharides

(D. C. White personal communication). The

sediment organic matrix and POM form the

basis of the benthic (detrital) food webs.

The grazing of detritus and its microbial

populations enhances nutrient quality for

subsequent microbial development by

stimulating further microbial productivity

and enhancing the nitrogen and phosphorus

content of the POM. Physical disturbance,

through wind and tidal action and active

predation and biological activity, is one

of the reasons why the Apalachicola

estuary is such a productive system.

Seasonal relationships among the

various physical and biological factors in

the bay system have been developed (Figure

38). Although the biological response to

a given event usually follows a nonlinear

or curvilinear pattern, certain relationships have become evident after many years

of observation. Seasonal variations of

temperature and the pulsed river flow are usually out of phase. Local rainfall

(Florida) peaks during summer months.

Salinity in the estuary is highest during

summer and fall months. The timing of the

river flow, and the resultant loading of

nutrients and POM, is critical to the

seasonal biological successions in the

estuary, especially during winter and

early spring. During such periods of low

winter temperature and salinity and high

river flow and detrital movement into the

estuary, benthic infaunal abundance is

high. Epibenthic organisms (especially

fishes) reach peak levels during late

winter as temperature starts to increase

and macroinvertebrates available for food

are abundant. Benthic omnivores such as spot and the Atlantic croaker are favored

by such conditions. Although these

sciaenids overlap in their temporal distribution, food size partitioning by these

two bottom-feeding fishes results in

distinctive differences in prey type and

size (Sheridan 1978). A larger apparatus

allows croaker to penetrate deeper into

the substrate and consume larger polychaetes, shrimp, and crabs. Spot tend to exploit smaller organisms, such as nematodes, harpacticoid copepods, juvenile

bivalves, and smaller forms of polychaetes. There is enough dietary overlap,

however, to allow the potential for

competition between these two species.

Benthic macroinvertebrates occupy an important trophic link between the primary

producers (and microbes) and the upper

85
REGIONAL I RAINFALLj

EVAPOTRANSPIRATION

Soil Composition

Floodploin Inundotion /

VEGETATION DISTRIBUTION

MICROBIAL

SUCCESSION

LOCAL RAINFALL

E mer genlVeoetation

Leaf Pfoduction Decomposition

Ffogmenlofion Nutrient/ Detritus Flo' S

y Ttdal Subsidy W.nd^V

o.s.rba'r'y ISUBH/IERGED VEGETATION PRODUCTIVITY

Conditioning

VER PEAKS I w.rdY

>

/ M N

^ ^ B.oiccoi^ ISUBH/IERGED VEGETATION nenl/OelcMus Flo* Oisluibance \ | ^'•'"

Salinity Gradients ,,_\, / 3

'"" — ^'\

/ \ ^-^ >

| PHYTOPLANKTON PRODUCTIVITY | ^

Potticulate Organic MatterV Wind Subsidy ; j- -; .-y' ^--;

I MICROBES -^DE

ganic Matter^ Wind Subsidy ; .- --

3ETR|TUS ' W T WT

~^LITTEr1- ASSOCIATED f

INVERTEaRATES i I

Detritus Product ion 1—

f

I 1

OETRITLIS-FEEDING INVERTEBRATES

CALANOID COPEPQDS -^ OTHER _iO0PL^NK.T0N_

, INVERTBBRATES

ANCHpyiES \ ^ ANCHOVIES

PENAElp SHRIMP

SEATROUT

TOP PREDATORS ( Incluqing mon )

DEC JAN. FEB MAR APR MAY i JUN

MONTH

JUL. AUG. SEPT I OCT NOV

Figure 38. Generdlized, simplified model of seasonal relationships of the dominant

macroinvertebrates and fishes in the Apalachicola Bay system. The model associates

population distribution with seasonal changes in key physical variables, productivity

features, and the predator-prey relationships of the estuary.

trophic levels of the estuary. Of the 10

numerically dominant infaunal species

(representing over 83% of the total

number), five are detrital feeders, four

are deposit feeders (surface and subsurface), and one is a filter feeder. Of the

entire infaunal assemblage, there are fifteen omnivore/carni vore types, seven subsurface deposit feeders, eleven surface

deposit feeders, twelve (generalized)

deposit feeders, and seven filter feeders.

There are high numbers of the various

filter-feeding mollusks such as Rangia

cuneata and Crassostrea virginica .

The important role of detritus and

its associated microbial components is

indicated by the predominance of the

detriti vore/omni vore feeders in the

macroinvertebrate assemblages. Of the

dominant litter-associated organisms, the

polychaetes are generally omnivorous,

consuming fine detritus, microalgae,

copepods, and amphipods. The gastropods

in the litter include omnivores, filter

feeders, scavengers, suspension feeders,

and carnivores. The herbivorous snail

Neritina reel i vata is a ma.ior species in

the grassbeds of East Bay. The amphipods

found among the litter assemblages include

omnivores, detritus feeders (or leaf

scavengers) and, in the case of some gammarids, filter feeders. A few species

such as Hyalel la azteca , Gammarus

86
lacustris , and Mel ita spp. are known to be

leaf ThreddeFs [i.e., herbivores),

although other amphipods are predaceous,

feeding on hydroids, bryozoans, and

(possibly) zooplankton. Crustaceans such

as the tanaid Hargeria rapax are generally

omnivores, but some are shredders or parasites. Mysid shrimp generally feed on fine detritus and diatoms. Decapod

crustaceans found in the litter

associations are largely omnivores and

detritus feeders, although certain

dominants, such as penaeid shrimp and blue

crabs, are predominately carnivorous

during certain life stages.

During the spring months, river flow

discharge decreases, salinity increases,

and the water clears. These conditions

trigger the late spring phvtoplankton

blooms and associated zooplankton

increases. The spring plankton peaks are concurrent with increased relative

abundances of plankti vorous fishes such as anchovies and menhaden. As the

temperature increases and river flow

falls, the high numbers of infaunal

macroinvertebrates fall precipitously. As

a result, by the end of spring there are few spot and Atlantic croaker in the bay,

and the sand seatrout, feeding on anchovies, becomes the dominant scianid.

Sheridan (1978) postulated that the summer anchovy peaks are truncated bv sand

seatrout. There is little trophic

Interaction of the sand seatrout with

other dominant fish predators; likewise,

there is little dietary overlap of these

species during their concurrent periods in

the estuary (May-August). During such

periods, predation pressure on penaeid

shrimp and crabs is low. Ry fall, most of

the sand seatrout have moved out of the

estuary and anchovies become dominant.

As temperature peaks during the

summer, the numbers of invertebrates

(penaeid shrimp, blue crabs) increase

(Figure 27). During this time, local

rainfall reaches seasonally hiqh levels.

Benthic macrophytes attain peak

productivity and standing crop. Ry the

end of summer, macrophytes start to die

off, and estuarine detritus levels

increase as the temperature begins to decline and salinity increases throughout

the estuary. By early fall, the numbers

of species of fishes and invertebrate

species reach high levels. One possible

explanation for this situation is that

those species limited by low salinity

during most of the year are able to enter

the shallow portions of the estuary at this time. Other factors that could

enhance the observed high numbers of

species during the fall could be falling

temperatures (to optimal levels) and the

availability of detritus and/or

detriti vorous invertebrates as food.

An overwhelming majority of the

estuarine nekton is omnivorous at some life-history stage, and detritus forms an important component of stomach contents at any given time (Sheridan 1978; Sheridan

and Livingston 1979; Livingston 1982b).

Of the seven dominant macroinvertebrates,

representing over 90% of the trawlsusceptible catch, five ( Peaneus

setiferus , Palaemonetes pugio , Callinect es sapid us, Penaeus aztecus and Loll iguncula

brevisT are omnivore/carni vore types;

Ner i ti na reel i vata is an herbivore, and

Lolliguncula brevis is a zooplankti vore. While the nutritional importance of the

detritus remains in doubt, omnivory

appears to be an important characteristic

of the predominant feeding patterns at intermediate levels of the estuarine food

webs.

Top predators, feeding largely on decapod crustaceans and fishes during the

fall, include spotted seatrout ( Cynoscion

nebulosu s), flatfishes ( Paral ichthys

spp. ), adult silver perch ~

T"Bairdiel1a"

chrysoura ), searobins ( PrionotTTs spp. ), and various shark types.

During November, as the temperature

drops rapidly, epibenthic organisms

decrease and various migratory species

leave the estuary for nearshore gulf

waters as part of their annual migration.

Penaeid shrimp are an example of this type

of population behavior. River flow starts

to increase during the early winter, and

salinity goes down. Benthic infaunal

species richness and abundance increase as winter progresses (Figure 27).

The seasonal succession of habitat

change, energy distribution, soeciesspecific recruitment patterns, predatorprey relationships, and the resulting food

web configurations contribute to the

87
biological organization of the estuary.

Infaunal macroinvertebrates reach maximum

abundance from November through March,

although species richness is highest in

May. As indicated previously,

phytoplankton and zooplankton are abundant

during spring months and summer periods.

Fish abundance peaks during winter and

early spring although fish and

invertebrate species richness indices

reach their highest level in October.

Epibenthic invertebrate abundance, on the

other hand, is high during August when

penaeid shrimp and blue crabs are prevalent. In general, the dominant fish

species, while overlapping in abundance to some degree, tend to predominate during

different times of the year; high croaker

and spot abundance occurs in winter and

early spring, sand seatrout in summer, and

anchovies in the fall and early winter.

Water column feeders such as anchovies are linked to plankton outbursts and predation

pressure from species such as sand

seatrout. Benthic feeders occur primarily

during periods of detritus/

macroinvertebrate abundance. Croakers and

spot feed largely on polychaetes, while

blue crabs concentrate on bivalves.

Directly or indirectly, most such species

take advantage of the detritus that is

brought into the estuary by the river.

The combination of low salinity, high POM,

and low predation pressure contributes to the observed high relative abundance of

these species.

5.4. PREDATOR-PREY INTERACTIONS AND

COMMUNITY RESPONSE

Although productivity trends and

habitat characteristics are important

factors in the development and control of

food web and community structure,

biological features such as predator-prey

relationships and competition for resources can be extremely important in

affecting the biological organization of

the estuary. Predation within aguatic

associations can lead to changes in

relative abundance, species diversity, and

other important community indices.

Peterson (1979) reviewed factors that

relate the impact of predation and

competitive exclusion to the response of

benthic macroinvertebrates in unvegetated,

soft-sediment estuarine habitats.

Previous work with various marine assemblages (largely rocky intertidal

communities) has indicated that isolation

from predation (through manipulative

processes such as caging) should lead to increased total density, increased species

richness, and restriction of competitive

exclusion by particular dominant species

(Peterson 1979). According to this model,

manipulative predator exclusion should cause simplification of the prey community

as a result of enhanced competition due to

increased population densities. Various

authors have found that soft-bottom

associations of benthic macroinvertebrates

do not always follow such a paradigm

(Peterson 1979). A series of tests of

this basic hypothesis has been carried out

in the Apalachicola Bay system over the

past 3 years.

Inverse correlations between predator

and prey population do exist in the

Apalachicola estuary (Sheridan and

Livingston 1983). Macroinf aunal abundance

often declines precipitously during

periods of peak abundance of the chief

sciaenid predators (Mahoney and Livingston

1982). Such correlative results suggest

that fishes may have a direct influence on the infaunal assemblages through

predation. In grassbed areas, however,

infaunal biomass is not affected because

larger species (burrowing deeper in the

sediments) are not influenced by such

predation. Also, recent experiments

indicate that macroinvertebrate

assemblages in East Bay remain largely

unaffected by predation pressure from

fishes in the late winter and spring and

by motile invertebrates (penaeid shrimp,

blue crabs) in the summer/fall (Mahoney

and Livingston 1982; Livingston unpubl.).

Thus, predation does not appear to play a decisive role in the regulation of prey

density or macroinvertebrate community

structure in oligohaline portions of the

estuary during periods of peak predation

pressure.

One possible explanation of the

apparent contradiction of the predation

paradigm could lie in the recruitment

potential of the dominant infaunal

species. In a series of experiments with

azoic sediments (i.e., devoid of

macroinvertebrates), Mahoney (1982) found

that infaunal larval recruitment was a deciding factor in the population dynamics
of various macroinvertebrate species such

as Streblospio benedicti and Capi tella

capitata . Such organisms are characterized by extremely short life

cycles. Rapid reproduction and larval

settlement could mask the impact of

and biological disturbances,

often important features of

estuaries. Heavy larval

not always followed by

a given species, however,

habitat suitability

also be implicated

physical

which are temperate

recruitment is

predominance of

Other factors such as and competition could

in the

structure.

determination of community

At various levels of biological

organization in the estuary, the dominant

macroinvertebrate populations are opportunistic and are influenced to varying degrees by the high productivity

and physical instability of the system.

Such populations have adapted well to habitat instability and variability.

Response time to disturbance remains

little understood, however. Recent

experiments in polyhaline portions of the

bay system (Livingston et al. 1983)

indicate that salinity could be a factor

in the influence of predation on benthic

infaunal associations. Infaunal

macroinvertebrates in the field were manipulated using a series of treatments

that involved exclusion cages (i.e.,

predators were kept out), inclusion cages

(i.e., predators were returned to exclusion cages), and field controls.

These treatments were compared to laboratory microcosms taken from the

field. Preliminary results indicate that, over a 6-week period of observation, there were increased numbers of

macroinvertebrates in the laboratory

microcosms and exclusion cages. Species

diversity was reduced in such treatments

relative to field controls and inclusion

cages. Thus predation in polyhaline areas

of high macroinvertebrate diversity and

low dominance may affect infaunal

macroinvertebrate community structure.

The influence of salinity on species

diversity and relative dominance could

thus be a factor in the relative influence

of predation pressure on dominant

populations in various portions of the

estuary. In areas of low dominance, the

influence of predation may be enhanced

relative to oligohaline areas where

dominance is naturally high. In any case,

few generalizations of predation effects

can be made without due consideration to local habitat conditions.

89
CHAPTER 6

LONG-TERM ECOLOGICAL RELATIONSHIPS

Although diurnal and seasonal changes

in population and community structure in

the estuary are relatively well documented

(Livingston IQ/Sb, 1977a, lQ77d, 1Q78;

Livingston et al. 1974, 1977), the longterm biological relationships, measured in

decades, are still under consideration

(Livingston unpublished data; Appendix A).

Seasonal changes in important physical and

chemical factors are relatively stable in

terms of timing (Figures 9, 1?); however,

there is considerable annual or year-toyear variation of such factors (Figures

10, 14, 15, 16, 17). The coupling between

climatological features such as river flow

and long-term changes in the commercial

catches of oysters, shrimp, and crabs

(Meeter et al. 11^79) is often complicated

by socioeconomic influences on such data

(Whitfield and Beaumariage 1977).

The specific short-term distribution

of a given species is often associated

with complex habitat variables and the

avail ablility of food. At the same time,

long-term changes in a given population in

the estuary may be influenced by

climatological cycles. Thus, the monthly

distribution of brief squid ( Lol 1 ingun cula

brevis ) depends to a considerable degree

on fluctuations of zooplankton abundance,

but the timing and annual abundance of

this species is also associated with

recurrent cycles of salinity and

temperature (Fiaure 39; Laugh! in and

Livingston 1<582). Spring migration into

the estuary has been correlated with

specific changes in both temperature and

salinity, while the fall emigration

largely depends on temperature changes.

Timing of the succession of climatological

changes is important since a specific

temperature has entirely different

meanings to a given species in the spring

and in the fall.

Long-term patterns of blue crab

( Callinectes sapidus ) recruitment cannot

be determined solely by the physical and

chemical environment (Figure 40; Laughlin

and Livingston, unpubl.). For any given

year, the winter recruitment was inversely

related to blue crab population abundance

and to summer recruitment levels. The

variable size 1 (monthly mean frequencies

of crabs of 1-30 mm; Table 24) was inversely correlated with temperature (p

0.01) and with variable size 3 (monthly

mean frequencies of crabs 61 mm) (p

O.OS). No significant correlations were

found with river flow or local rainfall,

which were associated with peak

recruitments at different times of the

year. In a multiple regression with

variable size 1 as the dependent variable

(Table 25A, N

= 12 months), temperature,

rainfall, and variable size 2 explained

about 89°^ of the variability of relative

abundance. The variable size 2 was weakly

correlated with all other variables (Table

26R). In a multiple regression with

variable size 3 as the dependent variable,

temperature, river flow, size 1 and size 2 explained about 70^ of the variability of

relative abundance (Table 25C).

Winter recruitment was below the

6-year average (59 crabs/month) in

1972-73, 1974-75 and 1975-75. A single

high peak, however, occurred in 1<373 and was correlated with the highest peak of

river flow of the 6-year period (Figure

40). During the winter months of these

years, river flow (which largely

determines salinity values in the estuary)

reached high (1^73), intermediate (1975),

and low (1°76) values, whereas water

temperatures deviated little {+_ 1° C) from

the 6-year temperature mean (14.9° C). By

contrast, summer recruitment for each of

these years was well above the 6-year

90
YEAR and MONTH

Figure 39. Long-term fluctuations of squid abundance, salinity, and

temperature at stations lA, IB, and IC in the Apalachicola estuary from

June 1972 through March 1979 (Laughlin and Livingston 1982).

average (51 crabs/month) and was not

directly correlated with abiotic or physico-chemical factors; summer rainfall

varied from minimal (1976) to maximal

(1975) values and temperature varied

little. The total population abundance

(all sizes) following the winter

recruitments of 1972-73, 1974-75, and

1975-76 was above the 6-year average (59

crabs/ month). Summer recruitment values were not included in these calculations.

Alternatively, winter recruitment was above the annual mean in 1973-74, l'376-77.

and 1977-78, and was correlated with

relatively high (1974, 1977) and low

(1978) winter river flow. Water

temperatures were just above the average

in 1974 and markedly low in 1977 and 1978,

Summer recruitment levels and total

population abundance following the winters

of these years were all below the 6-year

average. Tn fact, dramatic decreases in

total numbers of crabs occurred in 1974

and 1978. Again, none of these values was significantly correlated with any single

abiotic factor. With the exception of

91
~]iiiiiiiii i |iiii n i iiii p

ii ffiiiiii|iiiiiiii iii|iiiiiiiiiii[iiiiiiiiiii[ i i i iiiiii'~ m s J 1972

m s 1973

ms)msjmsjms)

1974 1975 1976 1977

m s 1978

( ) l-50mm

( ) All Sizes YEAR and MONTH

Figure 40. Monthly frequencies of blue crabs and variations in important

physicochemical parameters at the 10 day-time stations in the Apalachicola

estuary from March 1972 through March 1978 (Laughlin and Livingston,

unpubl . ) .

92
Table 24. Parametric (r) and nonparametric (T) correlations of seasonal variations of

blue crab frequencies and abiotic variables. Variables represent monthly averages of

monthly data for 7 years. Salinity and temperature means are baywide over 14 stations

in the Apalachicola estuary (from Laughlin and Livingston unpubl.). Correlation matrix

- Seasonal variations (N = 12).

Variables
1978, years with high levels of winter

recruitment were preceded by years of hiqh

population abundances; however, the

opposite was not true for winters of low

recruitments.

Unlike the brief squid, there was no significant linear relationships between

blue crab population parameters and

abiotic factors. Including 1-, ?-, and

3-month time lags of the abiotic variables

did not improve such linear relationships.

However, for a given year, there was a significant inverse correlation between

winter recruitment and the following

summer recruitment (p < 0.1). In other

words, in any given year, above-average

winter recruitment was usually followed by

a sharp decrease in total population and

by low summer recruitment levels.

Conversely, relatively hiqh population

abundances and high levels of summer recruitments followed winters of low

recruitment levels. Thus, long-term

population features of these dominant

invertebrate species (brief squid and blue

crabs) are dependent on different factors.

Temporal variability is extremely

complex since, at any given instant, a natural system represents a composite ot different sequences of varyinq periods

superimposed over one another as the

result of an almost infinite number of

cause-and-effect reactions. Determining

causality is difficult because these

overlapping cycles may differ along

habitat gradients and at different levels

of biological organization. Consequently,

the term "backqround noise" has become a euphemism for our inability to determine

the temporal or sequential cause and

effect relationships. Modeling efforts

often assume that systems are in a state

of equilibrium, without defininq the

actual extent of temporal variability.

Terms such as stability, resilience, and

diversity are used to give a theoretical

^'ramework to what is essentially a lack of

consistent observations of organisms under

field conditions.

Annual variability among dominant

fish populations in the Apalachicola

estuary was considerable (Figure 41).

Each species followed a distinct, longterm pattern of abundance; no single

aspect of the physical environment was apparent as the controlling factor of the

long-term changes. Bay anchovies were

most dominant during periods of high

salinity. The sand seatrout population

tended to follow the anchovy pattern with

particularly low numbers during the vear

of peak flooding when anchovies were also

low (1973). The Atlantic croaker followed

no obvious pattern relative to temperature

or salinity. Spot showed the highest

year-to-year variability with relatively

high numbers taken during the winterspring months of 1981. The cold winters

of 1976-77 and 1977-78 did not appear to affect any of the dominant fish

populations in the Apalachicola estuary.

It is clear that factors other than

temperature and salinity are important in

the control of long-term fluctuations of

these populations.

Although generalized temperature and

salinity preferences are well established

for various estuarine species (Table 17),

most such organisms have a relatively wide

tolerance for these factors. Tolerance of

this kind could explain the lack of

importance of these factors in the

determination of long-term population

variability (Table 23). When viewed from

the aspect of relative (percentage)

abundance, a certain temporal regularity

of the appearance of the dominant fishes

and invertebrates becomes apparent

(Livingston et al. 1976b; Figure 42). For

example, relative occurrence of

Palaemonetes pugio is high during spring

while Penaeus setif erus was dominant

during late summer and fall. The blue

crab is abundant during winter periods.

Among the fishes, sand seatrout are dominant during the spring and summer while bay anchovies (after the first year

of sampling) predominate in the fall and

Atlantic croaker prevail during the late

winter and soring. When a comparison is

made among the dominant fishes for peaks

of abundance, such increases tend to be

evenly distributed over a 12-month period.

However, of the top invertebrates, most

abundance peaks occur during fall periods

(September-November) with secondary

concentrations of peaks during early

summer (May-June). The major dominants

for both fishes and invertebrates thus

94
Bay Anchovy { Anchoa mitchilli )

Atlontic Croaker ( Micropogonias undulolus )

aoo

Z2

24

16 0-

Spot ( Leiostomus xanthurus )

979 1980 1981 1982

Bottom Water Temperoture

—1—m—I—\ I I

I97J 1974 075 in7p 1977 ,9?e 1979 I98C 1981

Surface Water Salinity

Figure 41. Long-term abundance patterns in the dominant

trawlable fish populations in the Apalachicola estuary

from March 1972 through February 4, 1982, with reference

to temperature and salinity variations (Livingston 1983

unpubl . ) .

95
X

o

h-

<

o

ILi_ o

hUJ

o

I I I r A

I I I I J 1—1—I I I I t I I I I I I I I I I I I I I I I I I

T—1—I—r M A M J T—I I I I I—r I I I I—I I I I—r—n—i i"-i i i i—r i i i i i JASONDJFMAMJJASONDJFMAMJJASONDJF

1972 I97J 1974 1975 YEAR and MONTH

Figure 42. Relative importance of four dominant species of invertebrates

and fishes taken in the Apalachicola Bay system from March 1972 through

February 1975. These species represent 32.4% and 86.0% of the respective

3-yr totals (Livingston et al . 1976).

show distinct patterns of relative

abundance throuqh a given seasonal period.

Various independent ecological

factors operate to determine the temporal

distribution of the dominant estuarine

organisms. Biological functions, such as adaptive response to the physical and

trophic environment, determine

distributional patterns, thereby allowing

a somewhat orderly temooral succession of

dominant forms within certain broad

trophic spectra. Patterns of reproduction

of various dominant estuarine species have

evolved in such a way as to permit longterm partitioning of the estuarine

environment. Superimposed on these

patterns of response are varying levels of resource division based on vertical and

horizontal distribution of the component

96
species. Various microhabitat phenomena

such as salinity, bottom type, currents,

and the availability of detritus and food

are important. Thus, no single parameter

Drevails in the determination of the

community structure of the estuary, which

itself undergoes predictable seasonal

changes as part of a physically forced

system.

Although there are appreciable shortterm fluctuations in the numbers of

individuals of different populations, the

system maintains a temporal constancy

which, according to a traditional view of

such phenomena, could be termed stability.

This does not mean that the system is not

in a constantly transient state. On the

contrary, through natural and unnatural

mechanisms such as habitat alteration and

destruction, hurricanes, cold winters, and

periodic flooding, the various population

equilibria continuously shift. Each

population fluctuates around a specific

point of equilibrium, and the fluctuations

reflect the adaptive response to the

specific aspects of the estuarine

environment.

The

physically

Apalachicola

unstable in

estuary

time but

IS

is

characterized by epibenthic populations

which maintain relatively stable temporal

interspecific relationships. The dominant

fishes and invertebrates are temporarily

partitioned in time. Particular groups of

fishes tend to co-occur (Figure 43).

Generally, three main clusters were arranged around the top dominants, Anchoa

mitchilli (I), Micropogonias undulatus

(II), and Cynoscion arenarius ( IV) . The

anchovy group is abundant during the fall.

The Micropogonias group predominates

during winter and early spring periods,

and the Cynoscion group prevails during

the summer and early fall.

Studies are currently being

undertaken to model the response of the

major groups of fish with respect to physical stress, abundance of prey

(Mahoney and Livingston 1982; Livingston

et al. 1983), long-term changes of

concurrent populations, and experimental

manipulations of a variety of associations

within the estuary (Livingston et al.

1983; Livingston, unpubl.; Appendices A,

B, C). These studies will be based on occurrence patterns over a 1?- to 13-year

period.

97
0.9

Similarity

0.5 01 -0.3 —-0.5

r~

m

m

m

YR

'PTTT

Anchoo mitchilli Etropus crossofus Symphurus pfogiuso

Dasyatis sabina Pnonotus fnbulus

Archosargus probotocephafus

Lepisosfeus osseus

DorosofDc pefeoense

Mtcropogonios undulatus Lewstomus xanfhurus

Trinectes maculofus

Parohchthys lethostigmo Oob/osomo bosci Microgobius gulosus Microgobius fhafassinus GobiontHus boleosoma

Syngnafhus scovetii -

Lucartia parva

L agodon rhomboides

Brevoof fio pafronus

Mbnidia berylfina

Urophycis f/oridonus

Ictalurus cofus

Cynosdon arenarius Anus felis M»nticirrhus amenconus

Synodus foetens

Chloroscombrus chrysurus Choefodipferus faber

Peprilus paru

Syngnotbus flondae Eucmostomus argenfeus Syngnatbus louisianae CynosC'on nebulosus Pnonotus scitulus

Boirdiello chrysura

Sphoeroides nap^elus Or thoprisfis Cbrysopfero

Porichthys porosissimus

Eucmostomus gula

Opistbonerria oglmum

Anchoa hepse lus Monocanfhus hispidus

Poiydactyius octonemus Bagre mannus

Figure 43. Temporal associations of fishes taken in Apalachicola

estuary from March 1972 to February 1976. Only top 45 species in

terms of total numbers of individuals are shown. Clusters represent

species that occur together from one year to the next (Livingston et

al. 1978).

98
CHAPTER 7

THE ESTUARY AS A RESOURCE

7.1. FISHERIES

There ^re relatively few studies of

fisheries in the Apalachicola River

system. Early surveys (Cox and Auth,

1^70-1973) of the upoer Apalachicola River

noted increasing stress to various species

of fishes as a result of physical

alterations such as damminq, dredqing, and

eutrophication. Studies of striped bass

(Barkuloo 1967, 1970; Crateau et al. 1981)

indicated that, be^'ore the construction of

the Jim Woodruff Dam (1955) at the

confluence of the Flint and Chattahoochee

Rivers, there was a viable sport fishery

for striped bass in the Apalachicola

River. Since that time, the striped bass

fishery has declined "drastically." The

dams in Georgia (Figure 4), together with

dredging and spoil deposition along the

upoer Apalachicola River, have eliminated

spawning grounds in the Flint and

Chattahoochee Rivers. Pesticides from

agricultural runoff and industrial

effluents (Livingston I'^SAb) are also

suspected of reducing these populations.

The native Gulf of Mexico race of striped

bass, once widespread throughout the

rivers of the northern gul''', is now limited to a small population in the

Apalachicola River. Recent stocking of

Atlantic coast striped bass has further

diluted the gulf strain and has resulted

in only limited success (Crateau et al . 1981). Wooley and Crateau (1<383) conclude

that the native Apalachicola striped bass

represent the only existing remnant of a population that historically was present

in numerous Gulf of Mexico drainages. For

this reason, the authors recommend

conservation of the existing stock as a "gene bank."

A commercial catfish fishery still

exists along the Apalachicola River.

However, Miller et al. (1977) cite studies

that related snagging (i.e., stump removal

from the river bed for navigation) to the

decline of the commercial catfish harvest

from the river. This activity, together

with the massive excavation and

maintenance activities associated with

nagivation projects (Figure 44), has

reduced or modified the riverine habitat

substantially (Miller et al . 1977).

Recent studies of the Apalachicola River

(Ager et al . 1984) indicate that sand bars

and spoil disposal sites are now common throughout the river; in the upper river,

the gently sloping natural bank habitat

has become "scarce" because of dredging

activities over the past 30 years (Ager et

al. 1984). It has been projected that,

because of such habitat alterations, the

fish species composition will continue to

shift from game species (characteristic of

natural habitats) to rough and forage

species (characteristic of sand-bar

habitats). This loss of habitat has also

been associated with the recent decline of

the sturqeon fishery. Accordinq to recent

studies (Wooley and Crateau 1982), Florida

sturgeon landings in the Apalachicola

River have declined raoidly (U.S.

Department of Commerce 1976 landing

statistics) relative to neiqhboring gulfcoast rivers. The fishery effectively

ended in 1^70 when only five fish were

taken. The Apalachicola sturgeon

copulation appears to be in trouble,

although it is believed that at least a relict sturqeon population still remains

in the Apalachicola River. Recently,

Wooley et al. (li^B?) reported the first

recorded capture of a larval gulf sturgeon

about 3.3 km below the Jim Woodruff Dam in

May, 1977. Wooley and Crateau (1982)

reported that relatively few sturgeon

(35-40) were harvested by angling during

1981. An important spawning area has been

99
Figure 44. Dredge spoil bank along the

Apalachicola River--a result of channelmaintenance efforts of the U.S. Army Corps

of Engineers. Note dead trees in what was once the river flood plain.

located in the upper Apalachicola at the

end of the usual spring flooding. Recent

studies (Wooley and Crateau 1984 in

review) indicate seasonal migrations of

sturgeon between freshwater and estuarine

portions of the Apalachicola system. A strong homestream tendency is apparent.

The tailwaters of the Jim Woodruff

Dam still support some sport fishing in

the spring, especially for the white bass

( Morone chrysops ) and the hybrid or sunshine bass (M^. saxati 1 is x

M_. chrysops). Largemouth bass and various

forms of bream and shellcrackers are also

important sport fishes. The yellow perch

( Perca f lavescens ) is taken occasionally

by freshwater fishermen. The Alabama shad

( Alosa alabamae ) is the most abundant

anadromous fish along the river. As pointed out by Miller et al. (1977), the

general decline of the freshwater

fisheries is inevitable if habitat

destruction along the river continues.

Habitats are destroyed by dredging and

channelization, damming, urban and

agricultural runoff, toxic substances, and

other forms of river modification. There

is a need for a comprehensive assessment

of the current status of the Apalachicola

River fisheries and the current and future

effects of river modifications and habitat

loss on such productivity. However, as of

this writing, the channelization of the

upper Apalachicola River by dredging and

rock removal for navigation purposes

continues, and there is little hope of a return to former levels of productivity of

the once-viable freshwater fisheries.

The commercial fisheries of the

Apalachicola Bay system are diverse and

substantial. According to the summaries

of commercial marine landings in Franklin

County (Florida Department of Natural

Resources, 1952-1976) and

projections of commercial

there is considerable annual

such landings over the

observation (1952-1977)

Prochaska 1977). Shrimp,

analyses of

populations,

variation of

period of

(Cato and

together with

oysters and blue crabs, provide over 80^

of the annual catch by weight. Black

mullet and grouper contribute almost m

of the remaining catch. Whiting,

menhaden, flounder, red snapper, and

spotted seatrout all contribute to the

overall landings. In terms of total

value, shrimp (53.9°/$), oysters (33%), and

blue crabs (5.1%) constitute the backbone

of the commercial fishery value in

Franklin County, which itself accounts for over 90% of Florida's oyster landings and

the third highest catch of shrimp

statewide.

The oyster fishery in the

Apalachicola estuarv has historical

significance (Swift 1896; Ruge 1897;

Danglade 1917). Many of the historic

observations were similar to today's in

that floods and droughts have an important

impact on the viability of individual

oyster bars. The present distribution of

oyster bars does not differ substantially

from that depicted on maps produced during

the early part of this century (Whitfield

and Beaumariage 1977). However, the

current maps (Figure 20) need to be

updated, as they are based largely on obsolete surveys. Commercially valuable

oyster bars currently cover only half the area estimated to be available at the turn

of the century. Shell planting with

"cultch" or shucked shells has proven to

be a successful management technique for

encouraging oyster bar development

(Whitfield 1973). Approximately 40% of

the Apalachicola Bay area is suitable for

growing oysters if cultched in an appropriate manner (Whitfield and

Beaumariage 1977). The actual and

potential productivity has been attributed

to the unique geographical and physical

100
attributes of the largely unpolluted

Apalachicola drainage system. More

sanitary (safe) harvesting waters for

oysters exist in the Apalachicola estuary

than in any other Florida estuary.

Considerable support exists for this

industry as a regional and statewide

natural resource. This fact, added to recent information that the Apalachicola

Bay system appears to be a major spawning

or source area for the entire Florida Gulf

blue crab fishery (Oesterling and Evink

1977), has stimulated various research

investigations concerning future fishery

potential.

The overall Apalachicola fishery

resource has grown substantially over the

past decade. During the period from 1Q77

to 1981, all previous oyster production

records were broken on an annual basis

(Joyce 1983). The record landings were

due largely to an increase in the fishing

effort (Prochaska and Mulkey 1983),

although newly instituted programs of summer oyster ing (1977) and an oyster

relay program (Futch 1983) have added to the annual crop. Although oyster

production has increased to Al% of the

total Franklin County landings, the

relative value of the oyster crop has

declined to 365;!, partly as a result of

increased county shrimp landings and

considerable increases in shrimp prices

(Prochaska and Mulkey 1983). Blue crabs

constitute about 5% of the total value of

the commercial fishery in Franklin County.

Of the commercial finfish catch, striped

mullet ( Mugil cephalus ) is the most

important. Grouper, menhaden, and whiting

are also taken, although the commercial

''infish industry has declined in recent

years (Livingston 1983c).

Sport fishing in the Apalachicola Bay

system remains largely undeveloped,

although the potential exists for a highly

productive industry. Sport fisheries

associated with the estuary include

spotted seatrout ( Cynoscion nebulosus ),

red drum ( Sciaenops ocellatus ), tarpon

( Megalops atlanticus ), sheepshead

( Archosargus probatocephlus ), black drum

( Pogonias cromis ) and flounder

( Paral ichthys spp.). Fishes taken ott the

barrier islands and Alligator Point

include various species of sharks, cobia

( Rachycentron canadum ), bluefish

( Pomatomus sal tatrix ), red snapper

( Lutjanus campechanus ), and different

species of grouper. The development of

artificial offshore reefs in the region

could add considerably to the continued

development of sport fisheries in the area.

7.2. SOCIOECONOMIC FACTORS

The Apalachicola valley depends to a considerable degree on a rather narrow economic base. A land-use inventory

(Table ?6) is indicative of the regional

socioeconomic conditions. Forestry and

agriculture account for nearly 80% of the

land use in the basin. Forestry,

agriculture, sport and commercial

fisheries, recreation, and light

manufacturing are the chief industries of

the region. In Franklin and Gulf

Counties, commercial and industrial land use are only 0.9% and 0.^% of the total

area, respectively. In the entire river

basin, the population was 109,254 in 1974,

with only modest projected increases for

the next 10-20 years. Per capita income

is low, averaging only 65% of the state

level in 1974. Despite a historic trend

of emigration of workers, the natural

features of the river and bay system

continue to attract new residents,

especially in the coastal areas. The

Apalachicola system contributes an important part of the regional economy and

culture, with unique sociological

conditions characterized by the close

relationship between the natural attributes of the drainage system and the local

inhabitants. The slight investment needed

to maintain the rich renewable resources

of the area is an important factor in anv review of the value (economic and

cultural) of the natural productivity of

the valley.

Franklin County, which surrounds the

Apalachicola Bay System , has a relatively

limited scope of employment with primary

dependence on products from the aquatic resource base and tourist expenditures

(Colberg et al. 1968). Commercial

fisheries alone provide jobs for over 65%

of the Franklin County work force.

Fishing is an "export" industry for

Franklin County because practically all

sales are outside the region (Prochaska

and Mulkey 1983). Export sales trigger a

101
Table 26. Land use inventory of the Apalachicola River basin (from Florida Department

of Administration 1<^77).
contributions of the entire family

(husband and older boys as tonqers, wife

and older daughters as shuckers, joint

management of the business). Such a family-oriented business structure has

strengthened the traditional bond between

the community and the industry to an extent that is not common elsewhere in

today's society. Thus, family and kinship

bonds underlie and strengthen the

dependence of the area on the natural

industries.

Some of the more important prospects

for regional growth are based on residential develooment of areas such as St. George Island and industrialization

of the river watershed. This situation

has resulted in a direct confrontation

between local and outside developmental

interests. Future planning initiatives

will have to be based on a reasonable

evaluation of the natural renewable- resource base if the local industry is to be protected. The potential for

destruction of these resources through

environmental alterations and pollution is

high. At the same time, the potential for

expanding the highly profitable oyster

industry with updated management of the resource is excellent.

7.3. EXISTING AND PROJECTED IMPACT BY MAN

A number of publications have

addressed the problem of environmental

alteration and pollution in the

Apalachicola drainage system (Livingston

1^74, 1P75, 1976a, b, lQ77a-d, 1978,

1980a-c, 1983d; Livingston and Duncan

1979; Livingston et al. 1974, 1976a,

1978). The Apalachicola estuary depends

on three basic elements for its

productivity: (1) the Apalachicola River

system, (2) the Tate's Hell Swamp and

surrounding freshwater/brackish wetlands,

and (3) the barrier islands. Physical

alterations of these areas or changes in

water quality or quantity due to human

activities could affect the natural

processes that define and control the

productivity of the river-bay system.

7.3.1. Physical Alterations

Darnell (1976) reviewed the effects

of structural changes on a range of

aquatic systems. Impoundment,

channelization, dredge and spoil

operations, diking, and other physical

modifications have the capacity to alter

natural aquatic systems. Since the early

1970's, there has been considerable

controversy concerning efforts to dam

and/or channelize the currently freeflowing Apalachicola River. Georgia and

Alabama industrial interests want to maintain an authorized 9-ft channel so that barge traffic can move from the Gulf

of Mexico to upriver cities along the

Flint and Chattahoochee Rivers.

Currently, this system is deep enough for

barge traffic only 83^ of the time (U.S.

Army Corps of Engineers, 1975), which is

not enough for the upriver interests.

There are 13 hydroelectric dams on the

Chattahoochee River and 3 dams on the

Flint River, some of which are privately

owned (Figure 4). Publicly owned dams and

dredging and maintenance activities have

cost in excess of $700 million.

According to a 1975 environmental

impact statement by the U.S. Army Corps of

Engineers, dredging has had adverse

effects on the Apalachicola River.

Livingston and Joyce (1977) point out that

impoundments such as the Jim Woodruff Dam cause aquatic weed problems, water guality

degradation due to the accumulation of

herbicides and insecticides, continued

need for dredging due to sedimentation,

reduction of habitat due to spoil

disposal, and restriction of the movement

of nutrients and particulate matter to downriver areas. Dredging and snagging

(removal of submerged stumps) operations

along the Apalachicola River are blamed

for habitat loss (Stevenson 1977),

destruction of benthic organisms (Miller

et al. 1977), loss of flood-plain

vegetation (Clewell and McAninch 1977),

reduction of bank overflow, blocked

migrations of migratory fishes,

restriction of striped bass from thermal

refuges and sturgeon from former ranges,

and increased pollution due to oil and

chemical spills (Figure 44).

Stabilization of a river usually

leads to industrialization and municipal

development in the former flood plain with

associated effects on water availability

and quality. The development of the

Apalachicola floodplain is uneconomical in

terms of the cost-benefit analysis

103
(Rockwood and Leitman, 1977). A 1<382 comparison of federal subsidies prepared

by the Congressional Budget Office shows

that waterways in general receive the

highest level of public transportation

support of all industries. On the basis

of cost-per-ton mile, the

Apalachicola-Chattahoochee-Flint (ACF)

system is the most expensive maintenance

operation in the country (45.5 mills per

ton mile), being almost twice as expensive

as the second highest and 41.36 times the

national average. The cost to the public

of moving a barge through the Jim Woodruff

Dam is around $2,040. The 1981 cost for

maintenance of the Jim Woodruff Dam and

dredging of the Apalachicola River

exceeded $6,735,000, and recent cost

increases have not been offset by revenue

from increased barge traffic. Despite all

this information, the Corps of Engineers

has recently been authorized to blast tons

of rock from the river (a form of

channelization) at a cost exceeding

$1,000,000.

There are few available data for

evaluating the environmental impact of

physical alteration of the tri-river

system. Cox (1970) and Cox and Auth

(1971-1973) indicate that dredging (Figure

44) has contributed to local habitat

destruction on the Apalachicola River

along with associated simplification of

the fauna and reduced productivity. As indicated above (Ager et al .

, 1984), the

long-term dredging of the river is a significant ecological occurrence. These impacts include altered habitat,

shortening of the river, and redirected

natural river flow. Operations associated

with these activities include construction

of training dikes, maintenance dredging,

spoil deposition, bendway elimination, and

snag removal. The river has already been

shortened by past activities, and

channelization continues.

In the Apalachicola estuary, dredging

of Sike's Cut has been related to increased salinity in the bay and reduced

productivity due to a loss of nursery

habitat (Livingston 1979). A review by

state and federal agencies (Florida

Department of Environmental Regulation,

pers. comm. ) is currently in progress

(Livingston 1984a) to determine the

potential impact of dredging along the

Intracoastal Waterway on the salinity

regime and oyster productivity in the

estuary. Proven dredging effects include

deterioration of water-sediment quality in

dredged channels near areas of urban

runoff and effects on the natural salinity

regime of the estuary (Livingston 1984a).

In the lower Apalachicola valley, a 33,000-acre cattle ranch was established

along the west bank of the river in the

early 1970's (Figures 45, 46). This

operation was accompanied by extensive

clearing, ditching, and diking. Land was drained by periodic pumping of turbid,

sediment-laden water over the dikes.

Extensive forestry operations have been

carried out in the Tate's Hell Swamo above

East 8ay. After clearcuttina of large

tracts of trees, the land was ditched,

drained, olowed and replanted with pine

trees. Livingston et al. (1978) found

that during periods of heavy local

rainfall, cleared areas caused increased

levels of runoff leading to increases in

color and turbidity and reductions in pH

and dissolved oxygen. Analyses of the

problem indicate short-term adverse impact

on certain biological associations in upper East Bay. The long-term

implications of forestry activities for

water resources are currently being

evaluated (Livingston unpubl.).

Overall, the primary wetlands of the

Apalachicola valley remain intact,

although dredging and associated

construction activities, especially in the

upper reaches of the river, are continuing. These activities include the

construction of bridges across the river

and development of a barge terminal

facility and offloading system.

Currently, state and federal agencies are attempting to purchase portions of the

remaining wetlands for preservation.

7.3.?. Toxic Substances

The limited industrial and

agricultural activity in the region has

contributed to the relatively low levels

of pollutants found in the Apalachicola

drainage system. However, the water

quality of the Flint and Chattahoochee

Rivers has been adversely affected by

waterway maintenance activity,

urbanization, and the discharge of

104
Figure 45. Ditching and diking associated

with agricultural activities in the lower

Aoalachicola floodplain.

Figure 46. The extent of diking by

agricultural interests along the western

bank of the lower Apalachicola River.

industrial and agricultural wastes

(Georgia Department of Natural Resources

1Q78, 1^82). A thorough scientific

analysis of the biological resoonse to eutrophication and the influx of toxic

substances to these rivers is lacking,

however. Recent studies by the L). S. Geological Survey (H. Mattraw pers. comm. ) concerning the levels of toxic

substances in the Apalachicola River

indicate relatively low levels of heavy

metals and negligible concentrations of

herbicides. In the Apalachicola estuary,

from l'^72-1976, there was a orecipitous

decrease of organochlorine residues in

sediments and associated estuarine

organisms. This decrease was attributed

to the banning of DDT in 1Q72, the

flushing action of the river, and the

heavy sedimentation associated with the

estuary (Livingston et al. l'^78).

Recent studies (Winger et al. 1^582)

indicate that residue concentrations of

organochlorine insecticides (DDT, toxaphene), ool ychlorinated biphenyls, and

heavy metals in aguatic biota are higher

in the upper Apalachicola River than in

the lower river. Total organic

contaminant residues, particularly from

the upper river, exceeded permissible

levels for the protection of wildlife.

The authors considered that such

moderately high residues indicated that

the Apalachicola River "may be in the

early stages of contamination." The

highest levels of cadmium and lead in

sediments and biota of the

Apalachicola-Chipola drainage system are found in tributaries leading to the

Chipola River below an industrial plant

that discharged battery wastes into the

svstem (Livingston et al . 1°82). The oH levels of runoff water approximated 1.2 to

1.4. Concentrations of lead and cadmium

in sediments of the Little Dry Creek-Dry

Creek tributary to the Chipola River were particularly high. Studies are currently

under way to evaluate the biological resoonse to this contamination (R. J. Livingston unoubl.). Recent analyses

indicate that this contamination has not

reached the Apalachicola Bay system

(Florida Department of Natural Resources,

pers. comm.).

7.3.3. Muni cipal Development

Municipal development in Florida is

concentrated along the coast. The Big

Bend region, which includes the

Apalachicola Bay system, remains one of

the last undeveloped coastal areas in

Florida. In Franklin County, urbanization

is restricted to the cities of

Apalachicola (approximately 3,000 people)

and Carrabelle (aoproximatel v 1,000

people). A municipal waste system is

currently under construction in

Apalachicola to eliminate point sources of

waste discharge (Scipio Creek) into

surrounding areas. Nutrient,

phytoplankton, and dissolved oxygen data

indicate no discernible tendency for

105
cultural eutrophication in the estuary

(Livingston unpubl.). l^ost of the

construction activity in the Apalachicola

Bay system has occurred in Apalachicola

and East Point and on St. George Island

(Figure 47). While there is considerable

pressure for construction on the island,

population density is still relatively

low. The outlook for future growth,

however, remains uncertain, as portions of

the estuarv have already been contaminated

with municipal and agricultural runoff and

waste (Livingston l'383d).

Coastal development is often

accompanied by the loss of natural

vegetation, increased levels of solid

waste, and enhanced effluent discharge.

These activities often lead to increased

runoff, erosion, physical alterations,

changes in water circulation, increased

deposition of sediments, and the

introduction of various pollutants into

the river-bay system. Such changes can have an adverse effect on the natural resources of the area. According to Bell

and Canterbery (1'574, 1^75), "The ma.ior cause of closing of commercial shellfish

areas is bacterial pollution at sublethal

contamination levels." Closings of

Louisiana's shellfish beds went from 5, POO

acres in 1%5 to lciR,812 acres in 1^71, a

S^OO"-^ increase. In Florida, considerable

areas of shellfish grounds are closed each

year because of pollution. Of over ? million acres of available shellfish areas

in Florida, only 27% are approved for

harvesting; 13% are prohibited, ^t are conditionally approved, and about 60% are unclassified. The national figures show over 3 million acres of clam and oyster

beds closed, at a loss of over $3R.4

million (Bell and Canterbery, 1975).

Septic tank effluents, sewage waters, and

municipal and industrial runoff account

for most of these problems. Since

commercial fisheries account for 65% of

the Franklin County income, there is cause

for concern (Florida Department of

Administration 1P77).

St. George Island (Figure 47) forms

the gulfward perimeter of Apalachicola Bay

and is of critical importance to bay

productivity because its orientation

determines the distribution of salinity

and other water-quality features of the

estuary. In 1965, a bridge was completed

Figure 47. Portions of St. George Island

showing housing development and other

human activities.

from the mainland to St. George Island at public expense. The bridge caused the

island's value as real estate to escalate

tremendously. Today, portions of St. George Island are currently under considerable pressure for municipal development

(Livingston 1976a). Based on past

experience in Florida and other coastal

states, the outlook for St. George Island

is to be the center of the growth for

Franklin County. On St. George Island, as elsewhere in the drainage area, there is a real need for planned development if the

natural resources of the estuary are to be

maintained.

Recently, there have been a number of

incidents in which oyster ing in the bay

has been closed down because of high

coliform bacteria counts (Livingston et

al. 1978). This situation has caused

local economic problems and represents a continuing threat to the oyster industry

in the Apalachicola estuary. The

combination of dredging and municipal

development has led to localized pollution

of portions of the estuary (Livingston

1983d). Dredged channels south of

Apalachicola and East Point have acted as sinks for nutrients (nitrogen and

phosphorus compounds), oils and greases,

and heavy metals (Livington 1983b). Such

substances have been associated with the

silt (i.e., fine) fractions of the

sediments and have led to conditions of

high biochemical oxygen demand (BOD). The

degree of urban development, the heavy

106
boat traffic, and the dredging activities

have been directly associated with local

destruction of near-shore grassbeds,

deterioration of water and sediment

guality, and the loss of biological

productivity (Livingston 1983b, d).

Municipal drainages contribute

significantly to the pollution burden of

the Apalachicola River and Bay area (Livingston 1983d). Scipio Creek

(Apalachicola), Eagle (or Indian) Creek

(East Point), and runoff from East Point

into near-shore areas of St. George Sound

have been affected by a combination of

high biochemical oxygen demand (BOD) and

chemical oxygen demand (COD), low

dissolved oxygen, and heavy-metal

contamination of sediments. Areas of

northern Apalachicola Bay that receive

runoff from the city of Apalachicola also

show signs of low water quality. The

dredged canals of St. George Island are polluted. The boat basins at St. George

Island and Apalachicola have been

contaminated with organic input and heavy

metals in the sediments. The lowest

dissolved oxygen in the entire system

occurs at the St. George boat basin (just

west of the causeway as it enters the

island; Figure 47) during periods of high summer rainfall and overland runoff.

There are signs of organic runoff in the

vicinity of St. George Sound receiving

input from construction sites, although more analysis is necessary to qualify this

observation. At all of the above sites,

the biological indices (benthic infaunal

macroinvertebrates) indicated moderate to high biological stress.

Other major sources of pollutants are located in areas receiving drainage from

agricultural operations (Murphy Creek and

Clark's Creek off the Jackson River; West

Bayou in East Bay from the Tate's Hell

Swamp). Aerial reconnaisance of the study

area indicates that forestry interests

have drained extensive areas of the Tate's

Hell Swamp into East Bayou and West Bayou

in eastern portions of East Bay. High

organic input and heavy-metal

contamination of the sediments have been

noted in areas of the drainage system

receiving agricultural runoff. Biological

indices have indicated severe stress.

Various stations along the lower

Apalachicola River, while having relatively low levels of pollution in the

water and sediments, also appear to be

biologically stressed (Livingston 1983d).

These sandy areas could be naturally

stressed by the heavy currents and the

shifting qualities of the sandy substrate.

Dredging activities along the Apalachicola

River could contribute to the observed

paucity of benthic macroinvertebrates

noted in these areas, although the exact cause of the observed biological

conditions remains unknown.

Overall, the Apalachicola River and

Bay system remains relatively pollution

free at this time. Some areas, such as eastern portions of St. Vincent Sound,

have been characterized by relatively high

levels of heavy metals in the sediments,

the source of which is not immediately

apparent. These areas could be points of

sedimentation (such as the dredged

channels in Apalachicola Bay), which

naturally concentrate contaminants such as heavy metals as part of the fallout of

silt/clay fractions from river input and

urban runoff. Such small particles are known to adsorb chemicals such as heavy

metals. The dredged channels serve as silt traps within the system. The

cumulative effect of municipal and

agricultural activities in the region

could be especially significant to the

rather sensitive oyster industry in

Franklin County. It will take imaginative

and progressive planning and resource management action if the fisheries

potential of the Apalachicola estuary is to be preserved and enhanced.

7.4. LAND PLANNING AND RESOURCE

MANAGEMENT

Resource management, based on comprehensive scientific data, depends on complex socioeconomic factors and cultural

trends. The mere identification of a given natural resource does not

necessarily ensure enlightened planning

for its perpetuation. There have been a series of reviews of the resource problems

in the Apalachicola basin. The history of resource planning and management in the

Apalachicola basin has been well

documented over the past decade

(Livingston 1974b, 1975, 1976a, 1977a-d,

1978, i980a-c; 1982a; Livingston and Joyce

1977). Overall, there has been a

107
relatively qood relationship between

researchers, manaqers, and local user groups. The wel 1-inteqrated (local,

state, federal) planning initiatives have

been based largely on preservation (land

purchases) and conservation approaches.

Whether such efforts will maintain the resource remains to be seen.

7.4.1. Publi c Land Investment

Public and private parks, designed to conserve or preserve areas in the

Apalachicola Valley, are scattered

'. I

' iSlT

Figure 48. Major public investments and

specially designated areas in the

Apalachicola basin.

throughout the area (Figure 4R). The

Torreya State Park includes unique plant

species such as the Florida Torreya cedar

and Florida yew. The Apalachicola

Forest and

One of the major land-acquisition

projects, the bottomland hardwoods in the

lower basin, was the result of research

funded by the Florida Sea Grant College

and the Franklin County Commission

(Livingston et al. 1976a). In 1^76,

portions of the Apalachicola River

floodplain were considered for purchase

Legend

1. Three Rivers State Park

2. Jim Woodruff Lock and Dam

3. Jackson County Port Authority

4. Torreya State Park

5. Gaskin Wildlife Refuge (private)

6. G. IJ. Parker Wildlife Management

Area (private)

7. Apalachicola National Forest

R. Environmentally Endangered Land

Purchase

9. Ed Ball Wildlife Management Area

(private)

10. Apalachicola Bay Aquatic

Preserve

11. St. Vincent Island National

Wi Idlife Refuge

12. Little St. George Island EEL

Purchase

13. Dr. Julian Bruce State Park

14. Dead Lake Recreational Area

15. Proposed purchase (estuarine

sanctuary)

16. Unit 4, EEL purchase

17. Dog Island, Nature Conservancy

18. Proposed bottomland hardwood

purchase: Nature Conservancy

and "Save Our Rivers" program

(state).

108
through the Environmentally Endangered

Lands Program (EEL) of the State of

Florida, The environmental background and

justification for purchase was based on data concerning the movement of nutrients

and POM from floodplain areas (Livingston

et al. 1^77; Pearce 1977). Ecological

associations were made between the

hardwood forests of the lower floodplain

and the productivity of the Apalachicola

River-Bay system. Based on the data and

the need to protect this ecologically

sensitive portion of the system, the

Florida Cabinet approved the purchase of

?8,044 acres of the lower Apalachicola

floodplain for $7,615,250 in December,

1976. While this purchase represented

only a small percentage of the total

floodplain and could not hope to achieve a total approach to management of the system

as a whole, it provided a much needed

state presence in the area.

Considerable effort has been expended

in the preservation of barrier islands

bordering the Apalachicola estuary. Based

on information concerning the importance

of the islands to the bay productivity

(Livingston et al. 1976a), portions of the

eastern end of St. George Island were

added to the existing state nark. In March 1977, the State of Florida

authorized the purchase of Little St. George Island for S8, 838, 000.

Approximately 1,300 acres of undeveloped

land on Dog Island were purchased by the

Nature Conservancy in 1982 for the

implementation of an island conservation

program. In addition, the Trust for

Public Land purchased that portion of St. George Island known as Unit 4 which

borders the highly productive oyster beds

of East Hole. This land was recently

repurchased by state agencies as part of

the EEL program. The balance of St.

George Island is still in private

ownership, Ma.ior portions of the holdings

on western portions of this island are already restricted by planning regulations

to 1 unit/acre. Thus, much of the barrier

island system is currently under public

ownership or within the jurisdiction of

the comprehensive plan of Franklin County

(see below).

In summary, there has been a continuous and guite successful effort

over the past decade to purchase and place

in public stewardship those oortions of

the Apalachicola drainage system which

have been identified as important for

maintaining the high productivity of the area.

7,4.2, The Apalachicola Estuarine

Sanctuary

After years of effort by local, state

and federal agencies, the Apalachicola

River and Bay Estuarine Sanctuary was established in September 1979. The

sanctuary is the largest in the country

and includes 192,750 acres of submerged

waters and associated wetlands (Figure

49).

The approval of the Estuarine

Sanctuary was the legal equivalent

(Section 315, Coastal Zone Management Act;

P. L. Q2-583) of setting this area aside

as a natural field laboratory "for longterm scientific and educational purposes,"

With the establishment of the Sanctuary

came a federal grant of SI. 8 million, to

be matched by $1,95 million of Florida's

EEL funds (the previous wetlands purchase

on the Lower Apalachicola River) for the

acquisition of the additional wetlands

surrounding the East Bay system (the

nursery portion of the Apalachicola

estuary) (Figure 49), After the

acquisition of the final 12,467 acres around East Bay and portions of the M. K. Ranch along the lower Apalachicola River

by the state of Florida, the public land

perimeter of the estuarine sanctuary will

be nearly complete. Recently, state

agencies have entered into negotiations

for another tract of wetlands along the

Apalachicola River. If successful, this

land will become part of the "Save Our

Rivers" program administered by the

northwest Florida Water Management

District.

Currently, in a close cooperative

effort between local interests and state

environmental agencies, the Apalachicola

Sanctuary program is involved in the

development of a resource atlas

(Livingston 1983c) and management plan,

several ongoing research projects, public

educational programs, and continuous input

into local planning problems and public

interest issues. Mot the least of this

effort is the potential development of

109
training programs and curricula in the

Franklin County secondary school system.

A group of educational films on the

Apalachicola drainage system has been

developed for showing throughout the

valley. The close interaction of aquatic

research with local and regional elements

has been one of the keys to the successful

a management program for

effort will be carried

the auspices of

Apalachicola Estuarine Sanctuary if

effective mode of administration can establ ished.

development of

the area. This

largely under

out

the

anbe

7.4.3. Local Planning Efforts and

Integrated Management

A series

Commissions have

establishment of

local development

These

zoning

plans have

restrictions

of Florida County

been responsible for the

comprehensive plans for

and resource management.

the legal

which have

status of

been upheld

ST GEORGE

ISLAND

SANCTUARY

PROPOSED

iiji EEL

FEDERAL

Figure 49. Boundaries of the Apalachicola

River and Bay Estuarine Sanctuary, including actual and proposed purchases according to the Environmentally Endangered

Lands (EEL) Program (state) and current

federal holdings.

in recent court decisions. For some years, agencies such as the Apalachee

Regional Planning Council, the Washington,

D.C. -based Conservation Foundation,

Florida State University, and the Florida

Sea Grant College have aided local

officials in the development of a comprehensive management plan for Franklin

County. During the summer of 1981, the

Franklin County Commission passed a plan

which installed various restrictions on the level and tyoe of construction

activities in the area and established low

density requirements in environmentally

sensitive areas. These areas include

wetlands, barrier islands, and portions of

the county that drain into oyster bars and

grass beds (Livingston 19R3c). This plan,

in conjunction with the estuarine

sanctuary program and state and federal

management, could eventually provide for

the orderly development of the area while

managing the natural resources of the

region. Passage of the plan is only the

first step in the planning process.

Successful implementation of the Franklin

County Comprehensive Plan has not yet been

achieved, and the status of local resource management in the estuarine sanctuary

remains in doubt.

7.4.4. Integration of Management Efforts

A diverse series of management

approaches coordinated through local user associations and the estuarine sanctuary

could provide the key for broadening the

economic base of the region while

conserving the unique natural assets of

the Apalachicola drainage system. This resource use will have to be subiect to specific internal controls as the

population grows to prevent overfishing

and other problems related to the fishing

industry.

Long-term scientific data have been

used to address local problems such as pesticide use, aquatic weed control,

shoreline development, and other aspects

of human activity around the bay. Such

problems have often been solved through

close cooperation between researchers and

local elected officials. The initial

studies, funded through a series of grants

administered by the Florida Sea Grant

College, provided needed information

concerning the ecologically sensitive

points in the drainage system. These

110
areas include the Apalachicola River, the

upland wetlands (including the Tate's Hell

Swamp), and the barrier islands--all

features that control the hydroloqical

regime, nutrient structure, and physicochemical environment (salinity, water

quality), which, together with other

specific habitat conditions, provide the

appropriate environment for the seasonal

and annual progressions of prominent

estuarine populations. Through contact

with public officials, state and federal

administrators, and leaders of private

industry, researchers have been able to

channel scientific information into public

use. Through close cooperation with local user groups, the Apalachicola research

effort is gradually being applied to

regional problems.

The real test for this management

effort, however, remains in the future.

As of this writing, the issue is

unresolved.

Ill
CHAPTER 8

COMPARISON WITH OTHER ESTUARIES

The Apalachicola estuary has been

included in a comparison of 14 estuaries

on the Atlantic, Gulf of Mexico, and

Pacific coasts of the United States (Nixon

1983). This study indicated that

Apalachicola Bay is a relatively small and

shallow estuary, rapidly flushed, with a considerable watershed area (Table 21)

when compared to other estuaries in the

United States. The cross-sectional area

of the Apalachicola estuary (18.1 x 10^

m^) is relatively small comoared to most

of the other estuaries. Because of the

dimensions of the bay and the volume of

freshwater input, Nixon (1983) estimates

that dissolved and suspended materials are likely to remain in Apalachicola Bay for a shorter time than in many of the other

estuaries in the survey. The relatively

high level and strong seasonality of the

rainfall in the Apalachicola drainage

basin would contribute to the high river

discharge rates to the estuary.

Approximately f^2% of the surface area of

the estuary has salinities that average

less than 15 ppt. Apalachicola Bay stands

out, along with Mobile Bay and Northern

San Fransisco Ray, as a system that

resDonds to river discharge in "a major

way" (Nixon 1983).

Because of the physical

characteristics and the relatively high

annual level of solar radiation,

Apalachicola Bay and Kancohe Ray (Hawaii)

are the only estuaries of those surveyed

in which the bay bottoms fall within the

euphotic zone (Nixon 1983). This fact,

together with the major impact of the

river on the estuary, could help to explain the apparently high productivity

of the Apalachicola system. The

phytoplankton productivity in the

Apalachicola estuary is moderately high

(Table ?8). Estabrook (1973) found that

such production is similar to that found

in Tampa Bay. The importance of

phosphorus as the limiting nutrient for

phytoplankton productivity for various

estuaries, including the Apalachicola

system, is evident (Nixon 1983).

Relatively little of the Apalachicola

primary productivity is due to cultural

eutrophication from input of nutrients

from human wastes. The Apalachicola is

the least developed of the estuaries

surveyed, with an extremely low population

density (Table ?9). The contribution of

nutrients from point source discharges to the Apalachicola estuary is extremely low

(Table 30). These data indicate that the

Apalachicola estuary remains in a relatively natural state compared to other

such systems around the country.

A comparison of zooplankton abundance

in different estuaries is difficult

because distribution and abundance depend

to some degree on mesh size of the nets

used to take the samples. A wide variety

of mesh sizes has been used in such

studies. When compared with other

estuaries in the gulf, Apalachicola Bay

has a similar or larger zooplankton

assemblage in terms of numbers and biomass

(Edmisten 1^79). Such numbers are comparable to those taken in various

estuaries in the United States (Nixon

1983). A comparison of ichthyplankton in

the other estuaries indicated that the bay

anchovy ( Anchoa mitchil 1 i) as a dominant

species is a common characteristic in half

the estuaries surveyed (Nixon 1983). The

low numbers of fish eggs in the

Apalachicola system, relative to other

areas such as Tampa Bay, has been

attributed to the relatively low

salinities in the Apalachicola estuary

(Blanchet 1978). Attempts to make

comparisons between the level of primary

production and abundance of organisms at higher trophic levels indicate no direct

or simple correlation (Nixon 1983).

112
Table 27. Approximate dimensions of

selected estuarine systems (Nixon 1983).

Estuarine system

Watershed

area (km^)

Surface Mean

area depth

(kni^) (m) (m)

Mean Flushing

tide factor

(days)

Narragansett Bay

Long Island Sound

New York Bay. Delaware Bay Chesapeake Bay Patuxent Estuary

Potomac Estuary

Pamlico Estuary Apalachicola Bay Mobile Bay Barataria Bay

San Francisco Bay*- Suisun Bay plus San Pablo Bay South Bay Kaneohe Bay

4.8 X
Table 29. Approximate land use distribution and population density surrounding the

estuarine study areas (Nixon 1983).
Table 30. A. Aoproximate annual input from land drainaqe and point source discharge

of dissolved inorganic nitrogen (NH4''", NO2", NO3") per unit area and per unit volume in

various estuaries. '^ The top number of each entry is in mmol m~'^- y^, the bottom number

is in mmol m"3 y-1 (Nixon 1983).

Estuary

Land

drainaqe
Table 30. (Concluded.)

B: Approximate annual input from land drainage and point source discharges of

dissolved

areas. inorganic phosphate (PO43-) per unit area and per unit volume in the studyThe upper entry for each estuary is area (mmol m"'^ y"l) and the lower is

volume (mmol m-3 yl) (Nixon 1P83).

Estuary

Land

drainage Sewage Total

Percent

sewage

Narragansett Bay

Long Island Sound

New York Bay

Delaware Bay

Chesapeake Bay

Patuxent Estuary

Potomac Estuary

Paml ico Estuary

Apalachicola Bay

Mobile Bay

Barataria Bay

Northern San Francisco Bay

South San Francisco Bay

Kaneohe Bay

28

3

55

8

18

2

40

1

67

12

114

34

14

5

240

74

104

22

mi nor

38

4

1500

210

62

6

9

1

170

32

55

7

minor

minor

66

7

1555

218

80

6q

2

237

44

55

> 7

114

34

14

5

250

77

58

96

78

13

72

mi nor

minor

3.6

216
Table 31. Total numbers of fishes per trawl sample taken at permanent stations in the

Apalachicola estuary (3/72-7/80), the Econfina estuary (6/71-5/79), and the Fenhollowav

estuary (6/71-5/79). Numbers per trawl are averaged over the entire period of study

with percentages of the total number of fishes taken indicated by brackets. The 25

numerically dominant species in each estuary were used for the analysis. Sciaenids are marked with asterisks (from Livingston I'^sib).

Species

Total numbers

per sample

{% of total) Species

Total numbers

per sample

{% of total)

Apalachicola estuary Econfina estuary (continued)

Anchoa mitchil 1 i Leiostomus xanthurus

Micropogoni as undul atus

Brevoorti a patronus

Cynoscion arenarius

'^

Harengula pensaco lae

Bairdiella chrysura

*

Trinectes maculatus

Arius felis

Lagodon rhomboides

Symphurus plaqiusa

Chloroscombrus chrvsurus

Etropus crossotus

Microgobius gulosus

Lucania parva

Polydactylus octonemus

Paral icht hys lethostigma

Menti cirrhus americanus

Syngnathus scovel 1 i Stellifer lanceolatus*

Anchoa hepsetus

Eucinostromus argenteus

Pri not us trihulus

Menidi a beryl 1 i na Gobiosoma bosci

2511

1766

1513

1214

498

54

50

4137

37

35

35

33

32

30

27

24

24

19

16

14

14

14

13

13

30.8

21.6

18.5

14. Q

5.1

0.7

0.6

0.5

0.5

0.5

0.4

0.4

0.4

0.4

0.4

0.3

0.3

0.3

0.2

0.2

0.2

0.2

0.^

0.2

Econfina estuary

Lagodon rhomboides

Leiostomus xanthurus

Bairdiel 1 a chrvsura

Monacanthus cil i atus

Gobiosoma robustum

Dipl odus holb ro oki

Ort ho prist is chrysoptera

Eucinostomus gul a Micrognathus cri nigerus

Syngnathus floridae

Opsanus beta

Eucinostomus argenteus

Stephanolepis hispidus

1418 (
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Shea, M. L., R. W. Warren, and W. A. Niering. 1975. Biochemical and

transplantation studies of the growth

form of Spartina alternif lora in

Connecticut salt marshes. Ecology

56:461-466.

Sheridan, P. F. 1978. Trophic relationships of dominant fishes in the

Apalachicola Bay system (Florida).

Ph.D. Dissertation. Florida State

University, Tallahassee.

Sheridan, P. F. 1979. Three new species

of Melita (Crustacea: Amphipoda), with

notes on the amphipod fauna of the

Apalachicola estaury of northwest

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Sheridan, P., and R. J. Livingston. 1979.

Cyclic trophic relationships of fishes

in an unpolluted, river-dominated

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in R. J. Livingston, ed.. Ecological

processes in coastal and marine systems.

Plenum Press, New York.

Sheridan, P. F., and R. J. Livingston.

1983. Abundance and seasonality of

infauna and epifauna inhabiting a Halodule wrightii meadow in Apalachicola

Bay, Florida. Estuaries 6:407-419.

Smalley, A. E. 1959. The growth cycle of

Spartina and its relation to the insect

populations in the marsh. Pages ci6-l00

in Proceedings of the Salt Marsh

Conference, Sapelo Island, Ga.

Smith, R. E. 1974. Marine environment

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eastern Gulf of Mexico. Institute of

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Springer, V. G., and K. D. Woodburn.

1960. An ecological study of the fishes

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in R. J. Livingston and E. A. Joyce,

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26.

Stokes, G. M. 1974. The distribution and

abundance of penaeid shrimp in the lower

Laguna Madre of Texas with a description

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Parks Wildl. Dep. Tech. Series 15. 32

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Stroud, L. M., and A. W. Cooper. 1968.

Color-infrared aerial photographic

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Apalachicola Bay, and St. George Sound,

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Swingle, H, A., and D. G. Bland. 1974. A study of the fishes of the coastal

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Sykes, J. E., and J. H. Finucane. 1%6.

Occurrence in Tampa Bay, Florida, of

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Serv. Fish. Bull. fi5:369-37Q.

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67:17-33.

Tanner, W. F. 1962. Upper Cretaceous

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S. Andree, ed., Apalachicola oyster

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Florida Sea Grant College.

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Ecology 43:614-624.

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Productivity and nutrient values of

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on the composition of the detrital

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in R. J. Livingston, ed.. Ecological

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sturgeon ( Acipenser oxyrhynchus

Desotoi). Northeast Gulf Sci. 5:57-58.

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Apalachicola River. Pages 22-23 in R.

J. Livingston and E. A. Joyce, Jr.,

eds. Proceedings of the Conference on the Apalachicola Drainage System. Fla.

Dep. Nat. Resour. Mar. Res. Publ. 26.

Zeh, T. A. 1980. Sikes Cut

inlets report No. 7. Sea

(Rep. No. 35). 39 pp.

; glossary of

Grant College

Zimmerman, M. S., and R. J. Livingston.

1976a. The effects of kraft mill

effluents on benthic macrophyte

assemblages in a shallow bay system

(Apalachee Bay, North Florida,

U. S. A.). Mar. Biol. 34, 297-312.

Zimmerman, M. S., and R. J. Livingston.

1976b. Seasonality and physico-chemical

ranges of benthic macrophytes from a north Florida estuary (Apalachee Bay).

Contr. Mar. Sci. Univ. Tex. 20, 34-45.

Zimmerman, M, S., and R. J. Livingston.

1979. Dominance and distribution of

benthic macrophyte assemblages in a north Florida estuary (Apalachee Bay,

Florida). Bull. Mar. Sci. 29, 27-40.

130
APPENDIX A

OVERVIEW OF SAMPLING PROGRAM IN NORTH FLORIDA COASTAL AREAS

1. Apalachicola Bay System

a. Physico-chemical measurements . (All stations, surface and bottom; March,

1972-present; minimum at monthly intervals. Temperature (air), river flow and

rainfall data from Atlanta, Georgia, to Apalachicola, Florida, (monthly, 1920--

present) are also on files in the data base)

temperature (^C)

salinity (ppt)

dissolved oxygen (ppm)

turbidity (J.T.U.)

color (Pt-Co units)

depth (m)

pH (since 1974)

Seech i readings (m)

chlorophyll _a (discontinued 9/76) (pg e,"l)

orthophosphate (discontinued 9/76) (yg P «."!)

nitrite (discontinued 9/76) (ng N «."!)

nitrate (discontinued 9/76) (^g N s,"l)

silicate (discontinued 9/76) (yg Si e,"^)

ammonia (discontinued 9/76) (yg NH3 n-l)

organochlorine compounds (pesticides, PCB's, etc.) (monthly, 1972-74)

heavy metals (1983)

B.O.D., C.O.D. (1983

b. Sediments , (representative stations, monthly intervals, 3/75-2/76)

grain size (phi units)

organic content {% dry weight)

c. Detritus , (macroparticulates: all stations, monthly from 1/75 to present),

microparticulates: mouth of Apalachicola and Little St. Marks Rivers, monthly

from 8/75 to present)

macroparticulates (by species or type, g dry weight)

microparticulates (sieve intervals; 45 y , 88 p , 125 y , 250 m, '^00 p, 1 mm, 2 mm;

g ash-free dry weight)

d. Phytoplankton analysis (Iverson et al.). (selected stations, monthly intervals;

7/72-9/76)

qualitative (species) analysis

productivity (ng C m"^ hr"^)

limiting factor analysis

131
e. Grassbed ( Vallisneria americana ) analysis , (macrophyte samoles, m^, monthly

from 11/75 to 7/77)

By species biomass (g dry weight)

f. Litter-associated assemblages , (stations 5A, 3, and IX; quarterly and/or

monthly from 4/74 to 1/77)

By species (numbers and biomass, q ash-free dry weight)

g. Benthic infauna . (stations 1, IX, 3, 4, 4A, BA, SB, 6); 10 repetitive

cores/station; monthly, 3/75 to present); weekly (station 3, 5A, Marine

Laboratory: 10/82-present )

"y species (numbers and biomass g ash-free dry weight)

h. Grassbed assemblages , (stations 4A and 4B; monthly from 11/75 to 7/77)

By species (numbers and biomass in g dry weight)

i. Epibenthic fishes and invertebrates , (otter trawls; all stations, 3/72 to present. Trammel nets and seines, various stations)

By species (numbers and biomass in g dry weight)

j. Stomach contents, fishes (dominant species) and blue crabs , (all stations,

monthly from 3/75 to 12/78)

By group or species according to month, size class, and station biomass (g ashfree dry weight) (Peter F. Sheridan, Roger A. Laughlin)

k. Zooplankton . (202 m mesh nylon net; monthly from 11/73 to 12/74)

By species (numbers, biomass, g dry weight) (H. Lee Edmisten)

1. Larval fishes . (505 p plankton net; monthly from 11/73 to 12/^4)

By species (numbers) (Harry Blanchet)

m. Meroplankton . (303 m plankton net; weekly, 10/82 to present; stations 3, 5A,

Marine Laboratory)

n. Fisheries data , (key commercial species; Florida Department of Natural

Resources)

(monthly from 1Q55 to present)

2. Apalachee Bay System

a. Physico-chemical measurements , (all stations, surface and bottom; June 1971-May

1979; at (minimum) monthly intervals)

temperature (OC)

salinity (ppt)

dissolved oxygen (ppm)

turbidity (J.T.U.)

color (Pt-Co units)

depth (m)

132
pH (discontinued in 1974)

Secchi readings (m)

chlorophyll a_ (discontinued in 1975) (pg i-l)

orthophosphate (discontinued in 1975) Lg P n'^)

nitrite (discontinued in 1975) (^g N n"^)

nitrate (discontinued in 1975) (pg N j,-!)

Sediments , (representative stations, October 1972; November 1972; February

1973; monthly, November 1976 - December 1978)

Phytoplankton analysis (Iverson and Bittaker). (selected stations, monthly

intervals, ElO-Fll, E11-F14, T21; January 1972-1975)

qualitative (species) analysis

productivity (ng C m^ hr"l)

d. Benthic macrophytes: long-term changes , (monthly from March 1972 - May 1979,

at certain permanent stations)

by species, m^, g dry weight

e. B enthic infauna: seasonal variability , (same stations as sediments; 10 repetitive cores/station; monthly, 11/1976 to 12/1978)

by species (numbers and ash-free dry weight/m^)

f. Short- and long-term variablity of epibenthic fishes and invertebrates (numbers

and biomass)

Seine: marsh stations, 1972-1975

Trammel nets: Offshore stations, 1974-1975, 1976-1978

Multiple otter trawl tows (7.2 min. /station) monthly, E7, E8, ElO, E12; F9, FIO,

Fll, F12; 6/72-5/79; quarterly, all stations, 6/72-5/79)

g. Trophic relationships (stomach contents) of fish assemblages in Apalachee Bay , (top 28 species, by numbers, all stations, monthly from 6/1972 to 12/1978)

biomass by group or species, according to month, size class, and station (g ashfree dry weight)

h. Trophic interactions of the pinfish (Lagodon rhomboides) with key biological

variables such as macrophytes and benthic invertebrates in Apalachee Bay

(Allan W. Stoner)

i. Nocturnal feeding habits of fish assemblages in Apalachee Bay (Joseph D. Ryan)

j. Day/night and seasonal varibility of epibenthic invertebrate distribution (Holly

S. Greening)

'

k. Seasonal variability of larval fishes in Apalachee Bay (Kathleen Brady)

1. Trophic relationships of decapod crustaceans (K. Leber)

133
APPENDIX B

COMPUTER PROGRAMS FOR ANALYZING FIELD AND LABORATORY DATA

1. Special Program for Ecological Science (SPECS): System Overview

a. Introduction

Long-term field studies in which diverse habitats are regularly sampled

for a variety of organisms and physical-chemical factors amass large amounts of

data. Organization and presentation of such data in a useful form has been

aided significantly by modern high-speed computers.

At Florida State, we have designed and developed a computer software

system specifically for use with long-term biological data. Primary design

criteria have been storage of a large data base, retrieval of virtually any

subset of the data, and rapid access to a diverse group of biological,

statistical, and graphical data.

The SPECS system has been written mostly in the FORTRAN programming

language. A few subroutines are written in the Control Data Corporation (CDC)

COMPASS assembly language. SPECS operates on a CDC 6500 or CYBER 74 computer

under the KRONOS operating system.

b. Organization of the System

Data storage

Field and laboratory data on physical-chemical parameters and fish, invertebrate, and plant populations are assembled and punched on standard 80-column

cards or entered directly via a computer terminal. Upon completion of a preliminary edit a program is executed to add the raw data to a data-base tape.

Two data base tapes are maintained, each with four files (one each for the

four types of data). One tape is always the "current" data base, the other serves as a backup. Upon each addition of new information the taoes reverse

roles.

Raw data information is also copied to a raw-data tape. This tape serves

as an additional backup copy of information (although it is not in data-base

format).

User Programs

All user programs, procedure files (predefined sets of often-used

operating system commands), program libraries, and active data files reside on computer-center disk packs (for rapid access). Most of the SPECS system is stored as a single file on one of- these disks.

134
This file contains one large program which has been structured in an overlay format having one main overlay and nine secondary overlays. Secondary

overlays perform the majority of system functions, such as loading data,

sorting, calculating biological indices, preparing for graphics and statistics,

etc. The main overlay simply fields a SPECS system command and calls for the

loading of a secondary overlay.

Library Programs

The F.S.U. Computer Center program library contains many routines accessed

by the SPECS system. Among these are the Statistical Package for the Social

Sciences (SPSS), the FSU plotting package, a mapping package (SYMAP), and a SORT/MERGE routine. The function of some SPECS secondary overlays is therefore

to prepare data base information for input to these higher level routines.

Operation of the System

All programs in the system are designed to be operated from a remote teletype or CRT terminal. System operation is interactive in that there is two-way

communication between the user and the program. The user guides the program

through each steo of analysis by entering commands or other information in

response to questions displayed by the program.

Terminal Session

A terminal session with the SPECS system begins with a user call of the

INIT (initiate) procedure file. This procedure first asks the user for the

location of the data to be used in this run (possibly a data base tape or an active data file). It then gets the SPECS program and initiates its execution.

The main overlay of SPECS writes a "COMMAND?" message to the terminal

screen. In response the user enters a SPECS system command. The LOAD

(retrieve) and SORT commands are used to create an active data file from a database tape. If the user began this run with an active data file (created in a previous run), the LOAD and SORT commands are not needed. Once an active data

file is available for use, the user selects from among a group of commands that

initiate execution of secondary overlays which perform analyses of active data.

Upon completion of an analysis, the user may wish to load more data

(create an additional active data file), request another type of analysis on the same data file, or terminate SPECS system operation. When system operation

is ended file disposition is under user control. Active data files or other

intermediate files may be saved if they will be used again. This ootion is

especially valuable if an important file has taken a long time to generate

(that time need not be invested again).

Summary

The SPECS system consists of a collection of programs written expressly

for the storage, retrieval, and analysis of long-term ecological data. Some

programs perform direct calculations or data manipulations while others serve

as interface programs that prepare data for higher level (and widely available)

program packages.

Interactive design affords a person with limited computer background

immediate access to a broad-based data file. It also facilitates a raoid,

relatively inexpensive yet comprehensive analysis with great flexibility of access to data and forms of analysis. All operations are carried out at the

135
terminal; new options can be added easily; and routine periodic updates of the

data base are easily made. This gives the biologist the use of a sophisticated

computerized software system as a research tool.

e. Capabil i ties

(1) Data Storage

(a) Physical-chemical data (by area, station, date, time of day, and depth)

-dissolved oxygen, color, turibidty, Secchi disk depth, temperature, pH,

river flow, rainfall, bottom type

-nitrate, phosphate, ammonia, water-column productivity

(b) Fish and invertebrate data (by area, station, date, and time of day):

-genus and species, number of individuals, mean size (with standard

deviation), biomass (ash-free dry wt.), sex (invertebrates only)

(c) Plant data (by area, station, date, and time of day):

-genus and species, total wet and dry weight stems and roots (wet and

dry weight), tops (wet and dry weight)

(2) Data Processing

(a) Retrieval

-for any area, station or group of stations, date or range of dates

(b) Sorting

-by area, date, station, time of day, or any combinaton of these

-biological data sorted by species

(c) Calculation of biological indices (based on numbers of individuals or biomass per species for any area, station or group of stations, date or range of dates, or time of day):

-Species Richness (number of species, Margalef Index)

-Species Diversity (Simpson index, Brillouin Index, Shannon Index,

Mclntosh/indices, Hurlbert's E(Sn))

-Species equitablity (Brillouin J,' Shannon J')

(3) Graphics

-for any area, station or group of stations, range of dates, or time of

day): plotted as a function of time or any other variable

-all physical chemical variables

-fish and invertebrates

a) number of individuals (single species or collective total)

b) average size

c) dry weight biomass (single species or collective total)

d) number of species

-plants

a) dry weight biomass (single species or collective total)

b) number of species

-Versatec high-resolution electrostatic plotter

(4) Statistics

-for virtually any set(s) of numbers that can be generated by any other

routine in the system

136
-linear regression. Student's t-tests, non-oarametric correlations,

discriminant analysis, factor analysis, scattergrams, analysis of

variance (one, two, and three-way), multivariate ANOVA, canonical

correlations, etc.

(a) Cluster analysis

-cluster by species, station, or time

-total flexibility in how species, stations, and dates are grouped prior

to analysis

-selection of similarity index from among Orloci's standard distance,

product moment correlation, Fager, Jaccard, Sorenson's, Webb, Kendall,

Czekanowski, Canberra metric, C-lambda, rho, and tau

-selection of clustering strategy from among unweighted pair group (grp

avg), weighted pair (centroid) grouping, nearest neighbor grouping,

furthest neighbor grouping, median grouping, and flexible grouping (with

beta)

(b) Dendrogram

-for any output from cluster analysis

-three scales available

(c) Data reduction by summary (for any area, station or group of stations,

range of dates, and times of day)

-number of individuals or dry weight biomass by species, month, and year

(fish, invertebrates, and plants)

-mean, standard deviation, and range of values over any specified time

period (for each of 12 physical-chemical parameters)

-trophic analysis - diet summary of food items (user-defined classes)

-C-lambda (for any area, station or group of stations, date or range of

dates, and times of day)

(d) Data smoothing

-moving average (number of time units optional)

-seasonal adjustment

-data tapering and trend adjustment

(e) Time-series analysis

-autoregressive moving average approach (Box-Jenkins methodology)

-spectral analysis

(f) Mapping

-physical-chemical data, macrophyte data, fish or invertebrate species

population totals mapped for all stations in study areas (by month)

(g) Data base update

-modification of any field in a data base record or records

-deletion of data records

2. "MATRIX" Program System: Summary of Capabilities

a. Introduction

The term "matrix" as used here refers to a form for holding numbers . It does

not have any algebraic connotations. A two-dimensional array (or table) is one very useful and frequently encountered form for the presentation of numbers. In

a table (see below), basic units ( cells ) that contain numbers are arranged in rows and columns , where the cells of any single row or column (vector) are generally related in some way. A table of numbers can be considered a two137
dimensional matrix. A three-dimensional matrix (see below) comprises a series

(or set) of tables, where each table ( plane ) contains the same number of rows

and columns. All the numbers in a single matrix plane are usually related in some way.

COLUMNS > COLUMNS >

ROWS
system was developed as a utility (i.e., a tool) to aid an investigator in

pulling together all the data required for a desired analysis and preparing the

data for direct use by other analytic systems.

c. MATRIX System Design Considerations

The principal design consideration for MATRIX was flexibility in input

data formats, retrieval and grouping of raw-data file values, and in manipulation and presentation of matrix file contents. Flexibility was achieved mostly

through generalization of program code; MATRIX was written without any fixed

input file formats so that the system could be used on a variety of input data

types. Furthermore, when a matrix is produced from raw data, the user is

offered a high degree of flexibility regarding which file values are retrieved,

where they are positioned along a matrix dimension, and how they are "pooled"

in the matrix cells. Once a matrix has been created, any of several manipulative operations can be performed on the data. Since these operations simply

act on matrix rows, columns, and planes, they are effectively available for use with any MATRIX-compatible input file, regardless of the original format.

Other design considerations were adaptability and allowance for user creativity. The MATRIX system has been coded in such a way that as new higherlevel package programs become available or new functions are desired of MATRIX,

the changes necessary to incorporate the new features will require a minimum of

programming time. There is considerable room for creativity in the use of the

MATRIX system; manipulative functions currently available under MATRIX can take

matrices apart, "twist" them around, change the contents, and piece them

together. It is left entirely up to the user to become familiar with the power

of these operations and to envision their application to specific problems.

d. Summary of MATRIX Functions

Listed below are brief descriptions of the functions performed by MATRIX

system operations.

GENERATE — Produces a numeric data matrix file of 2 or 3 dimensions from an input file containing alphanumeric storage keys and numeric data variables.

The program provides for complete user definition of row, column, and plane

contents, automatic insertion of missing values, and pooling of qualified

retrieval values by summation or averaging. Storage keys are written along

with data to serve as row, column and plane labels.

READ — Loads the data and label information from a previously generated matrix

file.

VIEW — Displays (to the terminal) a subsection of the data points contained in

the currently active matrix file. User defines the extent of row, column,

and plane dimensions for a desired submatrix (which may be the entire

matrix if it is 2-dimensional )

.

DESCRIBE — Lists the labels assigned to rows, columns, or planes. This function is helpful in determining the contents of a matrix.

EDIT — Allows the user to modify contents of a matrix. A user may change

labels, cell values, contents of a vector ( single row, column, or plane),

or the missing value code assigned to a matrix. He may also add a vector

to an existing matrix.

139
REPORT -- Similar to VIEW, but the display is written to a separate file that

is suitable for printing. The display is also more informative than that

of VIEW because:

(1) labels are written along with data points;

(2) an optional title is provided;

(3) the program performs report paging; and

(4) marginal totals can be reported (at user option).

SUBMATRIX -- Extracts a user-specified subsection of a larger matrix. A new matrix file (complete with labels) is created containing only the selected

portion.

MERGE -- Combines two existing matrices into one, with the following

restrictions:

(1) Both matrices must have the same missing-value code;

(2) At least two dimensions of the matrices must be equal (e.g.,

each matrix has 25 rows and 3 planes).

A new matrix file (complete with labels) is created.

TRANSPOSE -- Reorients the dimensions of a matrix in one of 2 ways:

(1) interchanging the rows, columns, or planes;

(2) making a three-dimensional matrix into two dimensions.

A new matrix file (complete with labels) is created.

STATISTICS -- Computes and (optionally) displays matrix marginal statistics

including total, mean, standard deviation, number of missing points, and

number of nonzero values. Statistics can be computed for either rows or columns over all planes or a selected plane. A matrix file (suitable for a MERGE operation) can also be produced if row statistics (all planes) have

been selected.

TRANSFORM -- Allows a user to perform data transformation (e.g., log, square

root, unit conversion) and/or standardization (i.e., to mean = 0, st. dev.

= 1). Also permits computation of linear combinations of variables.

PREPARE FOR PACKAGE -- Strips a matrix file of label and header information.

This function leaves a file containing data points only, which is the most

convenient form of input to the BMDP, SPSS, and MINITAB statistical

packages.

GRAPHICS — Prepares matrix row or column data for the EZGRAF graphics system.

A series of EZGRAF "EN"ter data commands are generated and written to a file (which is saved) suitable for EZGRAF entry with the "RUN" command.

MAPPING -- Prepares matrix data for spatial mapping with the SYMAP system.

Matrix columns must correspond to predefined spatial locations (i.e.,

stations). The user selects which matrix rows are to be mapped.

SUBSAMPLE SPECIES — A very specialized function, which performs "soecies

accumulation" according to the method described by Livingston et al. (1^76)

and "rarefaction" according to the method of Simberloff (1^78).

MENU — Displays a "full" menu of available system operations (descriptions of

options are more complete).

END -- Terminates the MATRIX program system and returns the user to interactive

communication with the operating system (NOS).

140
SPECS Interfaces

The SPECS computer program system (Special Program for Ecological Science)

was developed for use with the experimental and long-term biological data of

Dr. Robert J, Livingston at Florida State University. While SPECS provides the

capability to retrieve and sort data-base information and to calculate values

of biological indices, it has only a limited ability to make these results

available in a form compatible with higher-level packages such as BMDP, SPSS,

EZGRAF, and MINITAB. MATRIX can act as a powerful interface between SPECS and

these programs. The SPECS data base comprises the following types of data:

fish, invertebrate, plant, trophic, and physical-chemical. Using the SPECS

LOAD and SORT commands, these data can be retrieved for any area(s), station

(or group of stations), and date (or range of dates). The resulting file is

called a load/sort file and may be input to MATRIX GENERATE using one of the

predefined formats described in Table A. Notice that, for each data type,

there are several date options. Prudent selection of one of these can greatly

reduce the user effort required for the collapse procedure soecif ication. For

example, suppose a load/sort invertebrate file is input to GENERATE and the rows of the matrix file are to be individual months from January 1978 through

December 1982 (60 months). If the full date format is used, the date key

values will be listed as individual days (YYMMDD). It could be tedious here to specify a monthly collapse procedure, because all the numerical assignments for

the days in 01/78 would have to be entered, then all the assignments for 02/78,

and so forth for possibly all of the 60 months. If the data are read with the

year/month format, the day field would be skipped and the listed values would

be YYMM (i.e., the monthly collapse is accomplished by the format instead of a laborious user response). The user could then simply enter 999*1 and a new row would be generated for each month. If each row were to represent one of the 12

months of the calendar year (i.e., row 1 would represent all January's, row 2 all February's, etc.), the "month only" format would be appropriate. This format causes the day and year parts of the date to be ignored, leaving only 12

possible values for the date key.

The SPECS CALC command computes ten separate diversity, richness, and evenness indices along with the total number of individuals and number of species. These variables may be calculated for any area(s), station(s), date(s),

or time(s) of day or any combination thereof (see SPECS manual for details).

CALC outputs two files. One (keyword OUTPUT) is suitable for printing; the

other (keyword PLOTDAT) is suitable as input to MATRIX GENERATE. The use of

the MATRIX program on a SPECS CALC output file is the simplest way to make

these computed variables available for plottinq and/or statistical analysis.

The SPECS and MATRIX systems can be run with maximum efficiency if the user gives forethought to exactly what information is needed for his analysis.

A combination of LOAD, SORT, and SLECT orocedures in SPECS can be used to get

an input file for MATRIX with little or no extraneous data. If, for example,

the fish data for all dates and stations were retrieved to a load/sort file and

input to GENERATE when only the data for stations 3 and 5A from February 1978

through June 1980 were needed, two things would happen. First, MATRIX would

have to read a great deal of nonrelevant data, which would result in wasted

computer time and money. Second, there would be a very large number of key

values listed in the collapse procedure, so more user time and effort would be

required to specify the collapse correctly. The LOAD command causes an entire

data base to be read. The records that match the load parameters are written

to an output file. The SLECT command reads a load/sort file and writes the

records that match its parameters to a smaller load/sort file. If many subanalyses are to be run on a group of data, a LOAD command should be used to retrieve all the data that will be required for all the analyses; therefore the

141
Table A. Predefined file formats (including lists of key and variable names) to accomplish a number of SPECS-MATRIX interfaces.

File format Key names Variable names

SPECS Load/Sort File

-- Inverts (Full Date) — Fish (Full Date)

SPECS Load/Sort File

-- Inverts (Date is Year/

Month only)

-- Fish (Date is Year/Month

Only)

SPECS Load/Sort File — Inverts (Date is Month

Only)

SPECS Load/Sort File

-- Plants (Full Datel

SPECS Load/Sort File — Plants (Date is Year/

Month Only)

SPECS Load/Sort File

-- Phys/Chem Data (Full

Date)

AREA
Table A. (Concluded.

File format Key names Variable names

SPECS Load/Sort File

-- Phys/Chem (Date is Year/

Month Only)

SPECS CALC Output File

AREA

YRMON

STATION

TOD

DEPTHCODE

AREA

DATE (YYMMDD)

STATION

TOD

DEPTH
APPENDIX C

REVIEW OF ONGOING RESEARCH PROGRAMS OF THE CENTER FOR AQUATIC

RESEARCH AND RESOURCE MANAGEMENT (FLORIDA STATE UNIVERSITY)

1. Overall Scope of Program

Since 1971, together with undergraduate and graduate students, a multidisciplinary array of scientists, and a permanent staff of post-doctoral fellows

and full-time personnel, R. J. Livingston has put together a series of multidisciplinary and interdisciplinary studies concerning various aguatic systems in

the southeastern United States. Simultaneous laboratory and field studies

(descriptive, trophic, experimental) have been carried out, and the resulting data

have been entered into a series of computerized files. Simultaneously, computer

programs have been developed over the past 10 years that are designed to handle

short- and long-term multidisciplinary data from various aguatic systems.

Currently, the data from the 13-year research effort are being compiled and

organized for publication. These data are also being utilized to design and carry

out an ongoing field experimental program in a series of freshwater, estuarine,

and marine habitats.

Laboratory and Field Bioassays

A. Single-species tests (seagrasses, macroinvertebrates, fishes; fresh-water and

marine animals).

B. Multiple-species tests (macroinvertebrates; freshwater and marine)

C. Seagrass microcosms

Field Surveys

A. Habitat analyses (including pollutants) and biological components

(productivity, epibenthic fishes and macroinvertebrates, infaunal

macroi nvertebrates )

1. Spatial comparisons among rivers and associated estuaries

a. Flint River (Georgia), Chipola River (Florida), Econfina River

(Florida), Fenholloway River (Florida), Mobile River (Alabama),

Escatawpa--East Pascagula Rivers (Mississippi), Pee Dee--Sampit

Rivers, Winyah Bay (South Carolina)

2. Temporal comparisons (daily, weekly, and monthly intervals; 10-12 years of

continuous data)

a. Apalachicola River-estuary

144
b. Econfina River-estuary

c. Fenholloway River-estuary

B. Food-web structure of infaunal macroinvertebrates and epibenthic macroinvertebrates and fishes (freshwater and marine systems)

1. Transformation of species-specific abundance and biomass data into trophic

units by feeding mode and trophic position in food web

2. Comparative analysis among systems by feeding mode and trophic position in

food web (trophic unit)

3. Analysis of long-term (10-12) changes of food web structure in different

systems (with and without effects of pollution and habitat alteration)

4. Interaction of habitat features, primary production, and food web features

C. Impact Analysis (freshwater, estuarine, marine)

1. Pulp mill effluents (6 riverine and 5 estuarine systems)

2. Storm-water runoff (Apalachicola River and Bay systems)

3. Toxic substances (pesticides, heavy metals) (Flint River, Chipola River,

Hogtown Creek, Apalachicola River and Bay systems)

4. Dredging and spoiling (Apalachicola River and Bay system)

5. Forestry management (Apalachicola River and Bay system)

Experimental Ecology (Laboratory and Field)

A. Validation of freshwater bioassays with field data at toxic waste sites along

two rivers (Chipola River, Hogtown Creek): infaunal macroinvertebrates, epibenthic fishes and macroinvertebrates (ongoing)

B. Validation of bioassays using multi-species microcosms of soft-sediment,

marine infaunal macroinvertebrates (Apalachicola Bay system and the Yorktown

estuary, Virginia) (ongoing)

C. Predator-prey interactions (soft-sediment areas and seagrass beds) (ongoing)

1. Behavioral ecology

2. Field effects of predation on prey assemblages

3. Influence of predator-prey relationships on community structure under

varying environmental conditions (intra- and intersystem comparisons with

and without pollution variables)

4. Relation of predator-prey relationships to community structure and food

web patterns

Models : time-series changes of physical, chemical, and biological variables in

various aquatic systems (ongoing)

Application of research findings to resource management and public education

145
Development of the Apalachicola River and Bay National Estuarine Sanctuary

A. Apalachicola Atlas.

B. Continuing integration of regional research projects and a broad spectrum

of educational activities (secondary, undergraduate, graduate),

C. Input of research data to local, state, and regional planning/management

authorities.

2. Center for Aquatic Research and Resource Managment : Personnel (1984)

Robert J. Livingston (Director)

Glenn C. Woodsum (Associate Director)

DATA PROCESSING/ANALYSIS

Duane A. Meeter (Associate Investigator: Statistical Analysis)

Loretta E. Wolfe (Computer programming, statistical analysis)

Shelley J. Roberts (Project coordination, data transmission, formation of computer

files)

FIELD OPERATIONS

Robert L. Howell IV (Field collections, epibenthic fishes/invertebrates)

BIOLOGICAL ANALYSIS

Christopher C. Koenig (Bioassay, experimental protocols, biology of fishes)

Kenneth R. Smith (Oligochaete worms, benthic invertebrates)

Gary L. Ray (Polychaete worms, benthic invertebrates)

Bruce M. Mahoney (Benthic invertebrates, experimental ecology)

William H. Clements (Benthic invertebrates, feeding habits of fishes, experimental

ecology)

William R. Karsteter (Aquatic insects, benthic invertebrates, water/sediment

chemi stry)

John Epler (Aquatic insects)

Akshintala Prasad (Aquatic plants)

GRADUATE STUDENTS

Joseph Luczkovich (Ph.D.) (Predator-prey interactions, fish foraging, experimental

ecologyl

Jon Schmid t (Ph.D.) (Benthic invertebrates, experimental ecology)

David Bone (Ph.D.) (Experimental ecology, food web interactions)

Felicia Coleman (Ph.D.) (Physiological and behavioral ecology)

146
Kelly Custer (M.S.) (Feeding habits of decapod crustaceans, food processing by benthic invertebrates)

David Mayer (M.S.) (Ecology of penaeid shrimp, benthic invertebrates)

GRADUATE STUDENTS (continued)

Susan Mattson (M.S.) (Benthic invertebrates, experimental ecology)

Carrie Phillips (M.S.) (Benthic invertebrates, experimental ecology)

_J. Michael Kuperberg (M.S.) (Interactions of benthic macrophytes and animals)

LABORATORY ANALYSIS

Kim Burton (Rough sorting, sample preparation)

Howard L. Jelks (Rough sorting, sample preparation)

Mike Hollingsworth (Sediment analysis, algal studies)

Stephen B. Holm (Rough sorting, sample preparation)

John B. Montgomery (Sample preparation)

Brenda C. Litchfield (Sample preparation)

Mike Goldman (Sample preparation)

Frank Jordan (Fish identification)

Sam Cole (Sample preparation)

Hampton Hendry (Sample preparation)

Kline Miller (Sample preparation)

Melanie Saunders (Data punching)

Joanna Greening (Sample preparation, oligochaete mounting)

Carl Felton (Sample preparation)

David Ringelberq (Sample preparation)

Sharon Solomon (Sample preparation)

Sandy Vardaman (Sample preparation)

Erica Meeter (Sample preparation)

Carol Meeter (Sample preparation)

Julia Beth Livingsto n (Sample preparation)

Sara Van Beck (Sample preparation)

Cathy Wallace (Data preparation)

147
POST-DOCTORAL ADVISORS

Kenneth Leber (Feeding habits of decapod crustaceans, experimental ecology)

Kevan Main (Predator-prey interactions, behavioral ecology)

RESEARCH PROGRAM OF R. J. LIVINGSTON ET AL.

Pol lut ion

FEEDING STUDIES

(Stom^ich contents)

(ModfS of tcoding . function.il morphology, bt'h.ivior)

TRORHIC ORGANIZATION AND

FOOD WEB INTERACTIONS

( Bv t roph i c units)

VALIDATION EXPERIMENTS

{Single species, multipl- *"

species, microcosms' MODELS OF ENERGY CYCLING HYPOTHESES

FOR EXPERIMENTAL ECOLOGY

LABORATORY

EXPERIMENTS

(Bioassay, predator-prey

interactions; feeding patterns)

SPATIAL/TEMPORAL EFFECTS

( Wi t h i n-sy St cm variation;

be tween-syscem variation)

PREDATOR-PREY RELATIONSHIPS

(Effects of predators on prev assemblages)

Compe t i t ion

Figure A. An overview of the ongoing research program of the Florida State University

Aquatic Research Group concerning- long-term studies in nine river systems and six

estuaries in the southeastern United States.

148
50?77-101 REPORT DOCUMENTATION

PAGE

1. REPORT NO. FWS/ORS-R?/OS

3. Recipient's Accession No.

4. rrti. .nd subtiti. 7he Ecology of the Apalachicola Bay System: An Estuarine ProfileS. Report Date September 1984

7. Aothor(») Robert J. Livingston

9. Author's Afti Nation:

8. Performrng Organization Rept, No.

10. Project/Task/Work Unit No. Department of Biological Science

Florida State University

Tallahassee, FL 32306

11. Contract(C) or Grant(G) No.

(C)

(G)

12. Spontortng Organization Name and Address National Coastal Ecosystems Team

Research and Development

Fish and Wildlife Service

Washington, DC 20240

13. Type of Report & Period Covered

15. Supplementary Notes

16. Abstract (Limit: 200 words) Twelve years of studies in the Apalachicola Bay

data on geography, hydrology, chemistry, geology, and

The system is part of a major drainage area incl

wetlands in Georgia, Alabama, and Florida. It is a s barrier islands and dominated by wind effects and tid

(channels, sloughs, swamps, and backwater) and period

tant components. Principal influences on biological

river flow, nutrient input, and salinity. Water qual

tidal influences and freshwater inflows.

The system is in a relatively natural state, tho

development and population growth are beginning to th ecological importance as a food producer and shelter

movement to protect its natural resources, including

programs, integration of county land-use regulations

and creation of the Apalachicola River and Bay Nation

Research has produced an extensive computerized

working with these data have been developed.

system are reviewed. Included are biology,

uding four rivers and associated

hallow coastal lagoon fringed by

al currents. River bottomlands

ically flooded lowlands are imporprocesses are basin physiography,

ity is affected by periodic wind and

ugh hardly pristine. But economic

reaten it. The area's economic and

for diverse species has inspired a State and Federal land-purchase

into a comprehensive development,

al Estuarine Sanctuary,

data base. Computer programs for

17. Document Analysis a. Descriptors Geology „•

,,

, ^^ River

Hydrology r j. d1^i„^, Estuaries

Bioloay ,- , '^ Ecology .. Fisheries

b. Identifiers/Open-Ended Terms Bottomlands Apalachicola Bay

Flooded lowlands Florida

Nutrient input

c. COSATI Field/Group

16. Availability Statement 19. Security Class (This Report) unclassified

unl i mi ted

20. Security Class (This Page) unclassified

No. of Pages 150

(See ANSI-Z39.18) OPTIONAL FORM 272 (t-77) (Formerly NTIS-35) Department of Commerce

»^ r'">€^

^ Headquarters Division o' Biological Services, Washington, DC X Eastern Energy and Land Use Team

Leetown. WV

National Coastal Ecosystems Team

Slidell LA

Western Energy and Land Use Team

Ft Collins, CO

Locations ot Regional Offices

REGION 1

Regiimul Direckir US. Fish and Wildlite Service Lloyd Five Hundred Building. Suite Ib'^'^ 500 N.E. Multnomah Street Portland, Oregon 47232

REGION 2

Regional Director

U.S. Fish and Wildlife Service P.G.Bo.x 130(1 Albuquerque, New Mexico K 7 103

REGION 3

Regional Director

U.S. Fish and Wildlife Service Federal Building, Fort Snelling

Twin Cities, Minnesota S.'i 1 II

REGION 4

Regional Director

U.S. Fish and Wildlife Service Richard B. Russell Building 75 Spring Street, S.W.

Atlanta, Georgia 30303

REGION 5

Regional Director

U.S. Fish and Wildlife Service One Gateway Center Newton Corner, Massachusetts 021 5X

REGION 6

Regional Director

U.S. Fish and Wildlife Sei^ice PC, Box 25486

Denver Federal Center

Denver, Colorado 80225

REGION 7

Regional Directoi

U,S. Fish and Wildlife Service

101 1 t, Tudor Road

Anchorage, Alaska 4i)503
"""sSS^^"

DEPARTMENT OF THE INTERIOR

U.S. FISH AND WULIFE SERVICE

As the Nation's principal conservation agency, the Department of the Interior has responsibility for most of our nationally owned public lands and natural resources. This includes

fostering the wisest use of our land and water resources, protecting our fish and wildlife, preserving thAenvironmentai and cultural values of our national parks and historical places,

and providing for the enjoyment of life through outdoor recreation. The Department as- sesses our energy and mineral resources and works to assure that their development is in the best interests of all our people. The Department also has a major responsibility for American Indian reservation communities and for people who live in island territories under

U.S. administration.