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Full text of "The ecology of the Apalachicola Bay system : an estuarine profile"

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 particu- 
lar 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 bottom- 
lands 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 dis- 
tribution 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 estu- 
arine 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 Apalachicola- 
Chattahoochee-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 





Metric to U.S. Customary 




Multiply 


BiL 


To Obtain 


millimeters (mm) 
centimeters (cm) 
meters (m) 
kilometers (km) 


0.03937 
0.3937 
3.281 
0.6214 


inches 
inches 
feet 
miles 


2 

square meters (m ) 
square kilometers (km ) 
hectares (ha) 


10.76 
0.3861 
2.471 


square feet 
square miles 
acres 


liters (1) 

cubic meters (m^) 

cubic meters 


0.2642 
35.31 
0.0008110 


gallons 
cubic feet 
acre- feet 


milligrams (mg) 
grams (g) 
kilograms (kg) 
metric tons (t) 
metric tons 
kilocalories (kcal ) 


0.00003527 
0.03527 
2.205 
2205.0 
1.102 
3.968 


ounces 

ounces 

pounds 

pounds 

short tons 

British thermal units 



Celsius degrees 



l.SC'C) + 32 



Fahrenheit degrees 



U.S. Customary to Metric 



inches 


25.40 




mill imeters 


inches 


2.54 




centimeters 


feet (ft) 


0.3048 




meters 


fathoms 


1.829 




meters 


miles (mi) 


1.609 




kilometers 


nautical miles (nmi) 


1.852 




kilometers 


square feet (ft^) 


0.0929 




square meters 


acres ^ 
square miles (mi ) 


0.4047 




hectares 


2.590 




square kilometers 


gallons (gal) 


3.785 




liters 


cubic feet (ft^) 


0.02831 




cubic meters 


acre- feet 


1233.0 




cubic meters 


ounces (oz) 


28.35 




grams 


pounds (lb) 


0.4536 




kilograms 


short tons (ton) 


0.9072 




metric tons 


British thermal units (Btu) 


0.2520 




kil ocal cries 


Fahrenheit degrees 


0.5556(°F • 
xii 


- 32) 


Celsius degrees 



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- 




) y 


CHATX4ilOOCHE< 




^jS.j j^ 


Y DA 
CHIPOLA JK , 


*t— ^-'^ SEMINOLl/ y 


V! 


Uu >• /'^^ •-( 


DEAD \. ) J 
LAKE vjf /f" 


/apalaci 


HrfCOLA H 
\oCrt\.0CRONE£ (4 


^ 


m 


' .>*-DOG 1 


^-M-^^J^^S, GEORGE , 

SI /7*>"'"^^APALACHICOLA 
VINCENT 1 / BAY 
APALACHICOLA 



Figure 1. The tri-river (Apalachicola, Chattahoochee, Flint) drainage area showing 
the distribution of the important habitats and the position of key cities and 
municipalities within the Apalachicola-Chipola drainage system. 



1 



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 width- 
dominated 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 contri- 
buting 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 energy- 
dependent 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 rela- 
tively 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 princi- 
pal 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 first- 
order 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 small- 
order 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 rain- 
derived 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 time- 
series (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 















(1920-77) 








Columbus (rain) 




1200- 




(1920-77) 






s.d 








i 


A r% o t Af^ k I ^^ 


Ola (rain) 




<D 


L Mpaiacnic 




« 




\ (1937-77) 




"11000- 




\ 




^ 


J 


\ 






O 


/ 


\ 




■30 


— 1 800- 




\ 






u. 




\ 






<r 




\ 


s.d. 


-25 


ill 




\ 




O 


> 600- 




\^,/ 


\ 1 


•20 Z 


E 


s.d. 




< 




./ 


/ \ 


\ / 


■15 ^ 


400- 


/ 


\ '' /-• 








./ 




\ 






''-•■•■■■ 


'■<-J-^ \ 


\ 




-ID 


200- 


.--■• 


\/ 


s.d. 


/ 


•5 




J F A 


A A M J J A « 


; o N 


D 





MONTHS 

Figure 9. Seasonal averages of 
Apalachicola River flow (Blountstown, 
Fla.) and rainfall from Columbus, Georgia, 
and Apalachicola, Florida. Standard 
deviations (S.D.) are given for selected 
months (after Meeter et al. 1979). 



11 



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 reoccur- 
rence, 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 



UJ 

o 

< 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 



—I 

1925 



-| 1 \ 

1930 1935 1940 



-I \ 

1945 1950 



1955 



-1 1 

i960 1965 



"1 1 

1970 1975 



YEARS 



UJ 
C3 
< 

cr 

UJ 
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 (clock- 
wise 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 bottom- 
land 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 rain- 
fall 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 flood- 
plain 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 long- 
term 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. 





Area 


Oysters 


Grassbeds 


Marshes 


Water body 


(ha) 


(ha) 


(ha) 


(ha) 


St. Vincent Sound 


5.539.6 


1.096.5 


— 


1,806.9 


Bay 


20,959.8 


1,658.5 


1.124.7 


703.4 


East Bay 


3,980.6 


66.6 


1,433.5 


4,606.1 


St. George Sound (West) 


14,746.8 


1.488.8 


624.3 


751.9 


St. George Sound (East) 


16,015.5 


2.6 


2,767.3 


810.8 


Alligator Harbor 


1.637.0 


36.7 


261.3 


144.3 


Total 


62,879.3 


4,349.7 


6,211.1 


8,850.4 


Percent of total water area 


100 


7 


10 


14 



temperature minima occur from December to 
February; monthly variance is highest 
during winter. Whereas peak summer 
temperatures are comparable from year to 
year, winter minima vary annually (Figure 
1?). During years of extreme cold, 
temperature ranged from 5^ C to a maximum 
of 33° C over a P-month period. In 
addition to strong seasonal components of 
changes in water temoerature, periodic 
winter lows occurred at relatively regular 
(8-11 yr) intervals. In recent times, the 
winter of 1Q76-77 was particularly cold. 
The seasonal temoerature cycles are 
evidently superimposed over long-term 
temperature trends. 

The distribution of salinity in the 
bay at any given time is affected 
primarily by river flow, local rainfall, 
basin configuration, wind speed and 
direction, and water currents. The 
principal source o^ fresh water for the 
estuary is the Apalachicola River, 
although there is evidence that local 
runoff and ground water flows affect the 
habitat characteristics of the bay system 
in local areas (Livingston unpublished 
data). In terms of salinity, the bay 
system may be divided into two main 
provinces: the open Gulf waters of 
eastern St. George Sound and the brackish 
(river-diluted) oortions of western St. 
George Sound, Apalachicola Bay, East Bay, 
and St. Vincent Sound. 



o 

X 200 



o 100 



UJ 

> 

cc 




50 



|iiiiiiiiiii| run nil i|Miiiiiiiii|iiiriiiiiii|iniriiiMi|liillililll|niilliilll|iillllillljl 

MjSOMJS;iM.i:, ;. MJSDMJSDMJSOMJSOMJSOMJSO 
1972 1973 197-4 197^ i976 i977 1976 1979 1980 





LU 


?4 




LC 






D 






1- 




III 


< 




(0 


tr 


18 


< 


LlJ 




rr 


G. 




UJ 


5 




-> 


LlJ 


12 


< 


(- 






TIME 

Figure 12. Apalachicola River flow and 
average minimum air temperature data 
provided by U.S. Army Corps of Engineers 
and the NOAA Environmental Data Service, 
Apalachicola, Fla. 



15 



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 Hi Hi IB 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 




:]diiiM 



■"li fSEQ 4 ? •> 

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 Bay- 



River 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 Cut- 



4A 
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, unpublished- 
data). 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 



„ 


is- 




















Station 1 


E 
























o. 
























a. 


les- 






















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(1 








u 












^ ^ 












o 


10 - 




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. 


\ f 






, / 






> 






\ 


J 


\ 




\ 


/ 


1 


\ / 


\ 


X 






\ 


rs 1 


\ 


\ 


\ 


/ 


\ 




\ ^ 


o 






^ 


N 


\ 


\ 


\ 






I 


\ A 


Q 


75- 




V 




\/ 


V 


\J 


I 


\J 


\/ 


V U 


LJ 




^-_ 






v 






V 


V 


V 


1/ 


> 
























_l 
























O 
























V) 


5- 






















CO 
























Q 


25- 
























1972 


'! 1" 

1973 


1971 


1975 


1976 


1977 


197B 




1979 


I960 


1 1 

1981 1982 



E 
a. 

a. 125 



> 

o 

Q 
Ul 

o 

</} 

a 




1972 1973 1974 1975 1976 1977 1978 1979 1960 1981 1962 

YEAR and MONTH 

Figure 15. Surface dissolved oxygen 
(5-month moving averages) at stations 1 
and 5 in the Apalachicola estuary taken 
monthly from 1972 through 1982. 



18 



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 



O 

o 



cm 
o 

_i 
o 
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, 

(0, 



Acer nequndo L. 

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.^) 
Carpinu s 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) 
Dalmetto (Walt.) 



Sabal 

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) 

Pla tanus 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. 






Lower 
river from 


Lower 
river from 


Lower 
river from 




Mapping 


Upper 


Middle 


Wewahitchka 


Sumatra 


mile 10 




category 


river 


river 


to Sumatra 


to mile 10 


to mouth 


Total 


Pine 


136 


672 





204 





1,010 


Sweetgum- 














Sugarberry- 














Water oak- 














Loblolly Pine 


642 


1,440 


154 


474 





2,710 


Water hickory- 














Sweetgum- 














Overcup oak- 














Green ash- 














Sugarberry 


12,500 


32,200 


15,800 


1,770 


48.0 


62,300 


Tupelo-cypress 














with mixed 














hardwoods 


1,170 


1,860 


8,310 


15,800 


6,920 


34,100 


Tupelo-cypress 


2,420 


2,270 


6,240 


10,300 


456 


21,700 


Pioneer 





150 


19.2 








169 


Marsh 














9,030 


9,030 


Open water 


2,730 


3,110 


1,540 


2,010 


1,260 


10,700 


Unidentified 


1,020 


748 


81.3 


76.8 


19.2 


1,950 


Total 


20,600 


42,500 


32,100 


30,600 


17,700 


144,000 



22 



Apalachicola flood plain remains relative- 
ly 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 


J <""' 


f 


Ml BASIN 


^^^-p LIMITS 


\h 


1 r 




/ 



-=c^^ 




ALLUVIAL 
SOILS 



Figure 18. Frequently flooded areas and soil associations in the Apalachicola River 
Basin (taken from the Florida Department of Administration 1977). 



23 



are dependent to a larqe degree on water- 
shed 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 needle- 
rush, 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 light- 
limited 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, soft- 
bottom 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, bottom- 
land hardwood wetlands of the Apalachicola 
River have the highest litter-fall 
production rates of the worldwide warm- 



26 



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 approxi- 
mately 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., autoch- 
thonous: phytoplankton, benthic plants). 
Tnorqanic nutrients (phosphorus, nitrogen) 
and orqanic matter (dissolved, particu- 
late) 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 intri- 
cately 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 ^ 

SOILS- 



FLOODPLAIN 
f INUNDATION 



> TREE 

DISTRIBUTION 




POSITION 

O^ V LEAF 

PRODUCTION 



ATMOSPHERE 



^NUTRIENTS 
DETRITUS 



RIVER- 
BAY 



TRANSPORT 



fJ 



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 egua- 
torial 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 













A 




o 




« 


2.5- 






* 






o 


• 


o> 


















~-^ 








A 


O 








(A 












* 






3 




























o 




• 




w 


2.0- 




m 


C 





• 


• 




a> 


















T3 














° 




O 






A 












O 








o° 










E 


1.5- 


 


m 

AA 


 










a> 




* 


m 


 


m 








o 


1.0- 


A 


 

A  


• 

• * 
* 

• A 


• 


• 


• 

-I 





2.5 

Ash free dry weight 



3.0 



log river flow ( cfs) 



Figure 22. Regression analysis of the 
relationship of microdetri tus to 
Apalachicola River flow by season (totals 
taken from station 7, surface) (after 
Livingston 1981a). 



systems (Mattraw and Elder 
include headwater inflow, 
ground-water inflow, upland 
atmospheric fallout, and 
within the aguatic system 
hvdrological characteristics 



1Q8?). These 

tributary and 

productivity, 

productivity 

itself. The 

of the river 



system influence both the type of detritus 
produced and the guantity transported, 
since the wetland distribution is 
determined by patterns of flooding, and 
the same flooding provides an energv input 
as a transport medium. The Jim Woodruff 
Dam removes practically all the 
particulate matter from the Flint and 
Chattahoochee drainages (Mattraw and Elder 
1982), so the Chioola-Apalachicola wetland 
area is the primary contributor of organic 
detritus to the bay system.^ 

b. Coastal marshes . The primarv 
nonforested area in the bay system 
consists of freshwater and brackish 
marshes in the Apalachicola delta just 
above East Bay (Figure 19). In parts of 
East Bay, marshes are dominated by 
bullrushes ( Scirpus spp. ), cattails ( Typha 
domingensis ) , and other freshwater species 
such as sawgrass ( Cladium iamai cense ) . 
Brackish-water species such as cordgrass 
and needle rush are also found. The 



northeast section of St. Vincent Island 
has a well-developed brackish-water marsh. 

Kruczynski (1978) and Kruczynski et 
al. (l'578a, b) have analyzed the primary 
production of tidal marshes dominated by 
Juncus roemerianus in the St. Marks 
National Wildlife Refuge .iust east of the 
Apalachicola estuary. The authors 
considered such marshes representative of 
undeveloped wetlands in northwest Florida. 
Aboveground production was measured in 
each of three zones based on soil 
characteristics, elevation, and species 
assemblages. The high marsh areas were 
located approximately 600 m (l,Qfi9 ft) 
inland; middle marsh areas were located 
approximately 240-360 m (787-1,181 ft) 
from the bay; and low marsh areas were 
placed 0-120 m (0-394 ft) from the bay. 
Based on carbon-14 methods, the authors 
found that total aboveground production of 
a north Florida Juncus marsh is S.S t C 
ha"l yr-1 (3.8 tons/acre/yr) (low marsh), 
5.7 t C ha-1 yr-1 (2.5 tons/acre/yr) 
(upper marsh), and 1.8 t C ha-1 yr-1 (0.8 
tons/acre/yr) (high marsh). Using average 
figures weighted by area for an 
extrapolated estimate of marsh 
productivity in the Apalachicola marshes 
(Table 1), there is an estimated net 
production of 37,714 t yr-1 (41,561 
t/yr-1) in the Apalachicola estuary (East 
Bay, Apalachicola Bay, St. Vincent Sound) 
and 46,905 t yr-1 (51, 68^ tons/year) in 
the entire bay system. 

A comparison of these figures with 
those from other areas (Table 6) indicates 
that production of Juncus and Spartina 
systems along the northeast Gulf coast is 
comparable to that in other marsh areas. 
According to Kruczynski et al. (1978b), 
Soartina decomposes faster than Juncus , so 
nutrients from the former may be more 
readily available to associated estuarine 
systems. 

3.1.2. Autochthonous Sources 



a. Phytoplankton . Phytoplankton are 
ubiguitous in rivers, estuaries, and 
coastal systems. The phytoplankton 
community represents an important part of 
aguatic ecosystems both from the 
stamtpoint of primary production and as a 
key element in food webs. Diatoms are 
dominant in the net phytoplankton taken in 



31 



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 




700 


335 


130 


NJ 




— 


.-- 


300 


DE 




— 




445 


NY 




827 


508 


— 


GA 




985 


— 


— 


New 


England 


800-1300 


200-300 


— 


GA 




1158 


— 


— 


MD 




1207 


— 


— 


NC 




1296 


329 


— 


NC 




1300 


610 


— 


LA 




1410 


1005 


— 


GA 




2000 


— 


— 


LA 




2960 


1484 


— 


VA 




— 


500 


— 


DE 




— 


— 


445 


NC 




— 


— 


650 


VA 




— 


— 


1332 


GA 




— 


— 


2883 


GA 




— 


— 


3000 


Juncus roemerianus 









Kruczynski et al. 1978a 

Good 1965 

Morgan 1961 

Udell et al. 1969 

Smalley 1959 

Shea et al . 1975 

Teal 1962 

Johnson 1970 

Stroud & Cooper 1%8 

Marshall 1970 

Day et al. 1973 

Schelske & Odum 1961 

Kirby 1971 

Keefe & Boynton 1973 

Morgan 1961 

Williams & Murdoch 1972 

Wass & Wright 1969 

Odum & Fanning 1973 

Odum 1971 



Fit 949 595 243 Kruczynski et al. 1978a 

MS — — 390 Gabriel & de la Cruz 197' 

NC — — 560 Foster 1968 

NC — — 754 Williams & Murdoch 1972 

NC — — 796 Stroud & Cooper 1968 

NC — — 849 Heald 1969 

NC — - — 895 Waits 1967 

NC — - — 870-1900 Kuenzler & Marshall 1973 

MS — — 2106 Willingham et al. 1975 



LM = low marsh. 

UM = upper marsh. 

HM = high marsh. 

+ = estimate by change in biomass method. 



32 



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 
X 
X 

X 
X 
X 
X 
X 
X 
X 
X 
X 
X 



X X X X 

(continued) 
33 



X X 



X X 



X X 



X 
X 
X 
X 
X 
X 
X 

X 
X 
X 
X 
X 

X 
X 

X 
X 
X 



Table 7. (Continued. ) 



Phytoplankter 


1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


Bacteriastrum elonqatum 


X 

X 
X 

X 

X 
X 

X 

X 

X 

X 
X 

X 


X 
X 
X 

X 

X 
X 

X 

X 

X 
X 

X 

X 

X 
X 
X 


X 
X 
X 
X 

X 

X 
X 

X 

X 

X 

X 

X 
X 
X 
X 
X 

X 

X 
X 
X 
X 

X 


X 
X 

X 

X 

X 

X 
X 

X 

X 
X 
X 

X 

X 
X 

X 
X 
X 
X 
X 


X 
X 
X 

X 
X 
X 

X 

X 
X 

X 

X 
X 

X 

X 
X 


X 
X 
X 

X 

X 

X 
X 

X 

X 
X 
X 

X 

X 

X 

X 
X 

X 
X 


X 
X 

X 

X 
X 
X 
X 
X 

X 
X 

X 
X 
X 
X 

X 
X 

X 
X 
X 
X 


X 
X 

X 

X 
X 

X 

X 
X 
X 

X 
X 

X 
X 
X 
X 

X 


X 
X 
X 

X 

X 
X 

X 
X 

X 
X 

X 
X 

X 
X 
X 
X 

X 




Rhizosolem 


a alata 




Rhizosoleni 


a imbricata 


X 


Rhizosolem 


a setiqera 


X 


Rhizosoleni 


a berqonii 




Rhizosoleni 


a spp. 




Rhizosoleni 


a robusta 


X 


Rhizosoleni 


a stotterfothii 




Rhizosolen 


a calcar-avis 


X 


Rhizosolenia hebetata 




Guirardia flaccida 




Asterionella formosa 




Thalassiothrix frauenfeldii 




Thalassiothrix mediterranea 


X 


Thalassiothrix lonqissima 




Thalassiothrix nitzschioides 




Licmophora abbreviata 




Rhabdonema adriaticum 




Pleurosigma spp. 
Gyros igma spp. 
Amphiprora paludosa 
Navicula lyra 


X 
X 


Navicula spp. 
Lithodesmium undulatum 
Fraqilaria spp. 
Diatoma spp. 
Nitzschia pungens 


X 

X 
X 

X 
X 


Nitzschia spp. 
Nitzschia sigmoidea 


Nitzschia closterium 


Nitzschia paradoxa 


Grammatophora marina 




Cymbella tumida 




Cymatosira belgica 




Pinnularia spp. 
Synedra spp. 
Surirella fastuosa 




Cocooneis disculoides 




Schroederella delicatula 


X 


Eucampia cornuta 
PHYLUM PYRROPHYTA 


Ceratium furca 


X 


Ceratium tripos 




Ceratium massiliense 




Ceratium fuses 




Ceratium concilians 




Ceratium trichoceros 




Peridimium spp. 
Peri dimi urn grande(?) 





[continued) 
34 



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 phvto- 
olankton 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 maritima TJ 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 




















.+ 




1000 




: 


:: 


> : 


:: = 


; 




n 




+ 



ONDJ FMAMJ JAS 

Month 

Figure 23. Average seasonal variation in 
daily phytoplankton productivity for the 
Apalachicola estuary (taken from Estabrook 
1973; Livingston et al. 1974). 



various bayous along the northeastern 
margin of the bay (Livingston, unoublished 
data). There is little or no submerqed 
vegetation in St. Vincent Sound. Seagrass 
beds in Apalachicola Bay and western St. 
George Sound are restricted to shallow 
lagoonal oortions of Hog Island and St. 
George Island and are dominated by 
Halodule wrighti i , Gracil aria spp., and 
Syringodium f il iforme . Thus the 
distribution of submerged vegetation 
generally reflects previously described 
depth characteristics, water-quality 
features, drainage and current patterns, 
and salinity distribution. 

Seagrass beds undergo regular sea- 
sonal cycles of productivity and standing 
croD. The ecology of the East Bav 
Vallisneria beds has been well studied 
(Livingston and Duncan 1'57Q; Purcell I'^ll; 



Sheri dan 

Livingston 

production 

q C m-^ 

species 

standing 

months. 

growth. 



1Q78, 1^70; 

1079, 1P83). 
of Vallisneria 
yr-1 to 3^0 g C 
undergoes sharp 
crop biomass 
After a period 
maximum leaf 
maintained from May 
August, considerable 
plant standing crop occurs 
by new growth during 



Sheridan and 

Net annual 

varies from 3?0 

m-2 yr-1. This 

reductions of 

durina winter 

of rapid sprinq 

development is 

through July. By 

degeneration of the 

and is followed 

September and 



October. Similar cycles of growth occur 
in the Thai ass i a -domina ted qrassbeds in 
areas of higher salinity (Bittaker 107S; 
Livingston 10R?a; Zimmerman and Livingston 
1976a, b, 1Q79). Net annual production 
has been estimated to be 'iDO g C 
fn-2 vr-1 (Iverson unpublished data). 
Rapid growth occurs during soring and 
early summer. Standing crop biomass 
usually peaks during summer months with 
rapid degeneration as water temperature 
falls (November, December). Durina winter 
months, productivity and standing croD are 
relatively low in the various types of 
seagrass beds in shallow coastal areas of 
the northeast Gulf coast of Florida. 

Based on the productivity figures and 
the seagrass distribution (Table 1), the 
grassbeds in the East Bay-Apalachicola Bay 
area produce 8,053 t C yr-1 (P,866 tons C 



-ll 



yr-J-). Grassbed production in the 

remaining portions of the Apalachicola Bay 
system approximates 18,?60 t C yr-1 
(?0,1?? tons r yr-1). Total production 
for the entire system is ?7,?1'' t r y-1 
(?Q,'^89 r. y-1). 



3.?. DETRITUS FLUX AND NUTRIENT DYNAMICS 

Availability of organic matter does 
not explain the processes involved in 
transformation of energy as it moves 
through the complex food webs of the 
ri ver-estuarv system. Since relatively 
few organisms feed directly on living 
macrophytes, the degradation processes, 
which include mechanical fragmentation, 
chemical leaching, autolysis, hydrolysis, 
oxidation, and microbial activity, are 
important in the dynamic transfer of 
estuarine nutrients from available organic 
matter. Input to the immediate estuarv 
and the bay system as a whole is 
seasonally timed to specific 
meteoroloqical factors (Table o). Most of 
the river input occurs during winter and 
sprinq periods, while major phytoplankton 
blooms take place in the spring and fall. 
Input of organic matter from the seagrass 
beds occurs during the summer and fall. 
The transfer of organic materials from the 
coastal marshes is not as well understood 
as that of the other sources. In general, 
the contribution of plant detritus to the 
nutrient dynamics of the estuary is ex- 
tremely complex in terms of timing and 



36 



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 




Temp, 
or 


Salin. 

0/00 


Turb. 
,1TIJ 


Light 
Iv hr-1 


NO^ 


NO? 


°f^4 


Pri. prorl. 
•nn r rn-3 hr-1 


Chl-a 






q at:Tl 1-1 




mq m-3 


Econf ina 
estuary 




?a.4 

(1.01) 


26.? 
('.48) 


3.15 
(0.35) 


?6.5 
(5.60) 


0.3? 
(0.14) 


0.01 
(0.0^) 


0.04 
(0.01) 


6.00 
(1.75) 


0.61 
(0.17) 


F.S.U. Marine 
Laboratory 




(1.7R) 


?q.7 

(3.'S3) 


3.15 
(0.4q) 


37.8 
(3.7?) 


0.55 
(0.10) 


0.0? 
(0.07) 


0.10 
(0.04) 


0.7O 

(o!58) 


0.5? 

(o.?l) 


Ochlockonee 
River estuary 


(1) 


(O.PO) 


4.70 

(l.ofi) 


4.Q7 

(0.78) 


37.0 
(7.") 


1.83 

(0.?7) 


0.05 
(0.01) 


0.37 
(0.07) 


30.8 
('.^^7) 


7.14 
(0.41) 


Ochlockonee 
River estuary 


(?) 


(0.80) 


10.^ 
(0.70) 


4.0? 
(0.51) 


37.0 
(7.7?) 


7 74 

(0.8^) 


0.17 

(0.05) 


0.36 
(0.00) 


76.4 
(1.74) 


3.00 
(0.51) 


Apalachicola 
estuary (5) 




?7.S 
(1.1<?) 


3.74 
(7.S8) 


16.5 
(8. 06) 


33. Q 

(^.17) 


? 08 
('.63) 


0.15 
(0.16) 


0.34 
(0.08) 


40.7 
(10.7) 


5.13 
(1.1') 


Apalachicola 
estuary (?) 




?7.S 
(1.^4) 


11.7 


11.7 
(5.88) 


36.0 
(3.50) 


3.55 
(3.60) 


0.71 
(0.16) 


0.40 
(O.OQ) 


36.7 

(5.^1) 


4.11 
(0.84) 



Drocessinq 
al. 1979). 



(Odum and Heald 1972; Odum et 



Amonq the maior litter producers of 
the Apalachicola flood plain. Cairns 
(1Q81) and Elder and Cairns (1^82) found 
decomposition rates of floodplain leaf 
matter to he soecies-specif ic. Tupelo 
( N vs s a spp.) and sweetgum ( Liquidambar 
styracif lua ) leaves decomposed completely 
in R months. Leaves of baldcypress 
(Taxodium distichum ) and diamond-leaf oak 
( Quercus'lauri^'ol ia ) were more resistant. 
Water hickory fCarya aquatica ) had 
intermediate decomposition rates. Rates 
of carbon and biomass loss were linear 
over a 6-month period, but phosphorus and 
nitrogen leachinq was nearly complete 
within a month. Periods of river floodinq 
were particularly important for 
mobilization of the litterfall into the 
aquatic system. Floodinq immerses litter 
material, increases decomposition rates, 
and provides a transport medium. Because 
of the high diversity of floodplain tree 
species, the autumn peak of leaf fall is 
relatively prolonged (September-December) 
(Figure 24). Compared to the ACF system 
as a whole, the Apalachicola flood plain 
is extremely high in nutrient yield per 
unit "area, especially for carbon and 
phosphorus (Table 10). Mattraw and Elder 



(1982) postulated that the upper 
ChattahoocheeFlint watersheds yielded 
fewer nutrients because the 16 reservoirs 
act as nutrient retention ponds. Although 
headwater inflow provides substantial 
loads of dissolved nutrients to the 
estuary, particulate matter delivered from 
the river is derived almost exclusively 
from the Apalachicola/Chipola wetlands. 
Approximately \^% of the organic carbon 
delivered to the estuary is derived from 
less than 1^ of the ACF basin (Mattraw and 
Elder 1082). 

Particulate organic matter is 
transferred from the river to the estuary 
primarily during winter/spring floods, 
athouqh there is no direct correlation 
between microdetritus in the estuary and 
river flow by season (Table 5). 
Microdetritus flow is generally low during 
summer and fall periods and highest during 
the first river floods of winter (Figure 
22). In the estuary, surface dissolved 
nitrogen and phosphorus concentrations 
peak during periods of high river flow 
(Estabrook 1^73; Livingston et al . 1074, 
1976a; Table 11). Thus, the degree and 
timing of river flooding on a seasonal 
basis determines the form and level of 
nutrient fluxes into the estuary from the 
river wetlands. 



37 



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, phyto- 
plankton, 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 

Phyto- 
plankton 

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 ohyto- 
plankton 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 phvto- 
plankton 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 ohyto- 
plankton 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 export- 
of 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 Apalachicola- 
basis (adapted from 





Area 

(km?) 


Annual output minus 
(metric tons) 


input 




Are 
(q 


al yie 
m~'^ yr' 


Id 


Drainage basin 


Carbon 


Nitrogen 


Ph 
ph 


os- 
orus 


Carbo 


m N 


Phos- 
itroqen phorus 


Apalachicola- 

Chattahoochee- 

Flint 


50,800 


213,800 


21,480 


1 


,652 


4 




0.4 


0.03 


Chattahoochee- 
Flint 


44,600 


142,700 


17,860 




1.340 


3 




0.4 


0.03 


Apalachicola- 
Chipola 


6,?00 


71,100 


3,620 




312 


12 




0.6 


0.05 


Apalachicola 


3,100 


41,500 


1,060 




237 


13 




0.3 


0.08 


Chipola 


3,100 


29,600 


2,560 




75 


10 




0.8 


0.02 


Apalachicola 
flood plain 


393 


34,300 


674 




206 


87 




1.7 


0.52 



39 



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 nutrient- 
detritus 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 


17 February 1973 




12 July 


1973 


Bay T 


179.53 ± 13.11 




2.25 ± 


2.84 


B 


186.79 ± 19.48 




4.24 ± 


2.25 


River 


232.90 




219.54 




Bay T 


26.13 ± 18.53 




8.05 ± 


3.30 


B 


38.15 ± 30.61 




14.26 ± 


4.40 


River 


7.81 




7.57 




Bay T 


6.92 1 1.17 




4.03 ± 


.76 


B 


6.93 ± 1.29 




5.78 ± 


1.69 


R i ver 


12.63 




9.53 




Bay T 


2,531.80 ± 57.59 


1, 


,939.66 ± 


413.15 


B 


2,534.08 ± 62.88 


L 


,216.67 ± 


802.98 


River 


2,632.55 


3, 


,109.12 





N03 



NH4 



P04 



Silicate (Si04) 



40 



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 sub- 
strates 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 concentra- 
tions 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 syn- 
thesis, and I'^CO^ release from simple sub- 
stances by microbes (Morrison and White 
1980). These chanqes caused qrazinq 
shifts in community structure from diatom- 
funqal-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 predator- 
exclusion 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 



y 



BASE, DETRITAL FOOD 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 appro- 
priate 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 


anus parvus 


Temora 


turbinata 


Oithona 
Oirhona 


nana 
colcarva 



Pseudodiaptomus 
coronatus 

Centropagestus 

Centropagestus hamatus 

Euterpina actifrons 

Corycaeus americanus 

Carycaeus amazonicus 

Labidocera aesti va 

Other copepods 

Cirripedia larvae 

Decapod larvae 

Cladocerans 

Molluscan larvae 

Larvaceans 



East Bay 
(1 station) 
No./m3 % 


1696 


94.1 


1666 


92.5 


2 


+ 








+ 


+ 


1 


+ 


9 


+ 


9 


+ 














4 


0.2 




















3 


0.2 


49 


2.7 


50 


2.8 


2 


0.1 


+ 


+ 


+ 


+ 



6522 


80.2 


5546 


68.2 


352 


4.3 


48 


0.6 


101 


1.2 


35 


0.4 


60 


0.7 


217 


2.7 


25 


0.3 


15 


0.2 


25 


0.3 


9 


0.1 


14 


0.2 


60 


0.7 


21 


0.3 


94 P 


11.7 


79 


1.0 


168 


2.1 


166 


2.1 


74 


0.9 



Coastal 

(1 station) 

No./m3 % 


2286 


71.4 


635 


IP. 8 


244 


7.6 


342 


10.7 


567 


17.7 


194 


6.0 


11 


0.4 


17 


0.5 


36 


1.1 


64 


2.0 


44 


1.4 


28 


0.8 


17 


0.5 


25 


0.8 


61 


1.9 


180 


5.6 


26 


0.8 


460 


14.4 


58 


1.8 


95 


3.0 



'continued) 
44 



Table 12. (Concluded.) 









Average 1973-74 values 






Zooplankton groups 


East 
(1 Stat 
No./m^ 


Bay 
ion) 

t 


Apalachicola Bay 
(6 stations) 
No./m3 % 


Coast 
(1 Stat 
No./m-' 


al 
ion) 


Chaetognaths 





0.0 


27 0.3 


52 


1.6 


Polychaete larvae 


1 


+ 


92 1.1 


10 


0.3 


Fish eggs & larvae 


1 


+ 


92 1.1 


10 


0.3 


Other zooplankton 


2 


0.1 


35 0.4 


16 


0.5 



(Table 13). Although direct correlations 
were lacking, there was a strong positive 
relationship between salinity and 
diversity. Temperature and salinitv had 
no significant effect (at the 0.05 level) 
on the various dependent variables in East 
Bay or coastal areas. 



Coastal areas are physically stable 
when compared to the estuary; salinity 
varies little throughout the year in the 
offshore systems. In such areas, 
zooplankton standing crop is generally 
higher than that in East Bay. Hiversitv 
tends to increase because Acartia averages 



The general lack of definitive 
statistical relationships between 
individual zooplankton indicators or 
indices and dominant physical variables 
such as temperature and salinity reflects 
the considerable diel, seasonal, and 
annual variability in the distribution of 
zooDlankton in the estuary. Other factors 
^iTQ. almost certainly important to such 
distribution during various periods o^^ the 
year. Peaks o^ zooplankton biomass tend 
to be associated in some way with 
ohytODl ankton peaks, especially in 
Apalachicola Bay and coastal areas (Figure 
2fi). Predator-orev relationshios may play 
an important role in zooplankton 
distribution and abundance throughout the 
year. Such trends are obviously affected 
by habitat differences, however. The 
relatively small East Bav is characterized 
by low salinity and high sedimentation and 
turbidity. Salinitv changes, derived 
largely from river flow and storm-water 
runoff, are raoid. Most of the oeaks of 
zooplankton abundance correspond to 
salinity increases in this area. The 
copepod Acarti a tonsa has a maior 
influence on abundance curves and 
diversity indices in East Bay; it averages 
92''^ of the zooplankton taken throughout 
the year. 



E 80+ 

CO 

CO 
< 

2 60+ 

CD 

z 
o 



40- • 



Q. 
O 
O 
M 



O 



20 



■»- PHYTOPL AN KTON 
PEAKS 

* — * EAST BAY 

• — • COAST 

 _■ APALACHICOLA 
BAY 





\ 



NDJ FMAMJJ ASOND 

MONTH 

Figure 26. Seasonal distribution of 
zooplankton biomass in the Apalachicola 
estuary and associated coastal areas 
during 1974 (after Edmisten 1979). 



45 



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 inten- 
sive 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). 











Station 










Date 


3 


IC 


2 


offshore 


IB 


Inshore 
Little 
St. Georqe lA 


1 


11/21/73 


0.8 
(0.8) 


8.4 
(8.4) 


?.7 
(2.7) 


0.8 
(0.8) 


4.1 
(4.1) 


1.5 
(1.5) 


6.2 


1.7 
(1.7) 


12/9 




0.7 
(0.7) 




1.4 
(1.4) 


1.9 
(1.9) 


3.4 
(3.4) 


4.3 
(4.3) 


0.7 
(0.7) 


12/27 


0.3 
(0.3) 


1.3 
(1.3) 


1.0 
(1.0) 


11.3 
(11.3) 


12.0 
(12.0) 


0.4 
(0.4) 


0.7 
(0.7) 


— 


1/5/74 


3.0 
(3.0) 








— 








1/12 


— 


0.3 
(0.3) 


-- 


-- 






— 


12.3 
(12.3) 


2/26 


6.8 
(0.4) 


1.2 

(0.7) 


4.7 
(4.2) 






0.4 
(0.0) 


3.1 
(1.2) 


(2.0) 


2/27 


0.5 
(0.5) 


0.8 
(0.8) 


0.2 
(0.2) 




2.5 
(2.2) 


7.1 
(7.1) 


1.4 
(1.4) 


0.5 
(0.3) 


3/28 


14.3 
(1.8) 


61.3 
(40.3) 


115.1 
(0.9) 


10.1 
(6.1) 


47.7 
(7.3) 


265.^ 
(3.0) 


222.6 
(33.?) 


2^8. 4 
(10.?) 


4/20 


~ 


-- 


90.4 
(1^.8) 


— 


-- 


-- 


-- 


■'41.5 
(24.1) 


4/26 


13.4 
(8.4) 


163.0 
(7.8) 


171.0 
(25.3) 


2.4 
(1.7) 


84.0 
(7.7) 


2580.8 
(11. M 


1010.6 
(25.4) 


108.0 
(8.4) 


5/17 


98.9 
(52.8) 


70.5 
(51.0) 


8.3 
(0.0) 


62.8 
(52.7) 


241.5 
(50.6) 


1325.2 
(31.2) 


1234.5 
(283.8) 


54.0 
(12.?) 


6/18 


34.7 
(1.6) 


3.5 
(0.4) 


32.4 
(4.0) 


55.5 
(50.6) 


16.1 
(0.7) 


136.7 
(16.1) 


2.3 
(1.7) 


5.^ 
(1.3) 


7/18 


0.5 
(0.0) 


-- 


-- 


3.5 
(3.5) 


Q.5 
(2.4) 


20.3 
(5.1) 


1119.4 
(38.7) 


61.0 
(0.0) 


8/22 


16.4 
(9.9) 


150.7 
(4.1) 


72.8 
(23.3) 


-- 


16.2 
(1.6) 


141.1 
(9.7) 


75.5 
(10.3) 


18.1 
(0.7) 


9/12 


5.5 
(3.7) 


194.9 
(92.0) 


99.2 
(2.1) 


746.6 
(738.2) 

(continue 
48 


217.8 
(75.1) 

d) 


51.1 
(6.9) 


1032.6 
(20.6) 


46.6 
(0.0) 



Table 15. (Concluded.) 



Date 



Station 



IC 



Offshore 



IB 



Inshore 

Little 

St. George 



lA 



10/17 


5.1 
(4.1) 


4.1 
(4.1) 


2.5 
(1.4) 


7.8 
(7.8) 


2.4 
(2.4) 


4.2 
(4.2) 


3.5 
(3.2) 


3.8 
(0.8) 


11/7 


0.6 
(0.6) 








0.5 
(0.5) 


0.2 
(0.2) 


0.2 
(0.2) 


— 


12/3 


2.8 
(2.8) 




0.5 
(0.5) 


2.5 
(2.5) 


0.7 
(0.7) 


1.6 
(1.6) 


7.0 
(7.0) 


10.1 
(9.8) 



ichthyoplankton (25.8 m"'') 
beyond Sike's Cut in April. 



are found iust 



Fishes that live in a qiven estuary 
can be organized into various categories 
according to their life history (McHugh 
1^67). Estuarine-deoendent forms include 
truly estuarine species, anadromous and 
catadromous soecies, marine soecies that 
live and often spawn offshore but use the 
estuarv as a nursery, and marine species 
that enter the estuary seasonally as 
adults but remain offshore as juveniles. 
Tn the Apalachicola estuarv, the estuarine 
eggs and larvae are dominated by one 
estuarine species, the bay anchovy. At 
stations that are not near the passes (3, 
2, 1; Table 15) numbers of larvae of 
species other than anchovies are usually 
low. Such areas tend to be dominated by 
species that spawn within the estuary 
(i.e., atherinids, blennies, skilletf ish). 
Blanchet (1^78) attributed the low number 
of eggs in the estuary to the flushing of 
the bay system. It is also possible that 
the generally low salinities within the 
estuary prevented spawning by most 
species. Overall, the pattern and 
distribution of the fish larvae within the 
bay system would indicate that, while 
specific causative factors remain unknown, 
the primary function of the bay is its use 
as a nursery by true estuarine soecies and 
marine species that soawn offshore. 

4.3. BENTHOS 

Considerable information is available 
concerning benthic macroinvertebrates in 



estuarine and coastal systems (Mann 1P82). 
Benthic infauna, which live within the 
sediments, are usually separated according 
to size into macrobenthos, meiobenthos, 
and microbenthos. Although there are 
differing opinions as to the exact 
dimensions of each size category, most 
workers agree that the macrobenthos 
includes those organisms taken in 250-500 
micrometer ( m) sieves. Meiobenthic 
organisms are those taken between 62 m 
and 250 m, and organisms 
m are classified as 
Macroinvertebrates living 
sediments or at the 
interface are called 
eoi benthic invertebrates, 
will be treated as nekton 



smaller than 62 

microbenthos. 

just above the 

sediment-water 

epifauna or 

These organisms 

in this review. 



The relative composition of any given 
benthic macroinvertebrate collection 
depends to a considerable degree on the 
form of sampling gear. In the 
Aoalachicola Bay system, benthic 
macroinvertebrates have been taken by 
cores and ponars (McLane 1980; Mahoney and 
Livingston li^S?), leaf packs (Livingston 
et al. 1977), otter trawls (Livingston 
1976a, b; Livingston et al . lQ76b), and 
dredge-nets and seines (Purcell 1P77), 
The benthic macroinvertebrates in the 
Apalachicola Bay system represent a 
diverse fauna (Table 16) with distinct 
oatterns of temporal and spatial 
distribution (Livingston et al. 1^77). 
Although considerable seasonal and year- 
to-year variation in species composition 
and relative abundance is found at any 
given sampling area, certain trends are 



49 



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. ) 




Order - Or 


bini ida 


Family 


- Pilarqiidae 


Family - 


Orbiniidae 




Ancistrosyllis 




Haploscoloplos 




hartmanae 




foliosus 




Ancistrosyllis sp. 




Haploscoloplos 




Parandalia americana 




frazil is 




Siqambra bassi 




Scoloplos rubra 


Family 


- Syllidae 


Family - 


Paraonidae 




Pionosyllis sp. 




Paraonis sp. 




Syllidae sp. 


Order - Sp 


ion ida 


Famil y 


- Nereididae 


Family - 


Spionidae 




Laeonereis culveri 




Carazziella hobsonae 




Nereid sp. A 




Paraprionospio 




Nereis succinea 




pinnata 




Stenoni nereis martini 




Spiophanes bombyx 


Family 


- Glyceridae 




Streblospio benedicti 




Glycera americana 




Scololepis texana 


Family 


- Goniadidae 


Family - 


Magelonidae 




Glycinde sol it aria 




Magelona polydentata 


Order - Amphinomida 




Magelona sp. 


Family 


- Amphinomidae 


Family - 


Cirratulidae 




Amphinome rostrata 




Chaetozone sp. 


Order - Terebellida 


Order - Capitellida 


F am i 1 y 


- Amphictenidae 


Family - 


Capitellidae 




Cistena gouldi 




Capitella capitata 


Family 


- Ampharetidae 




Capitella sp. 




Hobsonia florida 




Capi tell ides jonesi 




Melinna maculata 




Heteromastus 


Order - E 


!unicida 




f il if ormis 


Family 


- Onuphidae 




Mediomastus ambiseta 




Diopatra cu^rea 




Notomastijs hemipodus 


Fami ly 


- Eunicidae 




Polydora liqni 




Marphysa sanguine a 




Polydora socialis 


Family 


- Lumbrineridae 




Polydora websteri 




Lumbrineris sp. 


Family - 


Arenicolidae 




Lumbrineris tenuis 




Arenicola cristata 


Order - S 


;abellida 


Family - 


Maldanidae 


Family 


- Sabellidae 




Branchioasychis 




Fabricia sp. 




amencana 


Class - Oliaochaeta 




Clymenella sp. 




Oliqochaeta spp. 


Order - Phyllodocida 


Order - F 


laplotaxida 


Family - 


Phyllodocidae 


Family 


- Tubificidae 




Eteone heteropoda 




Limnodriloides sp. 




Paranaitis soeciosa 




Peloscolex benedeni 




Phyllodoce fra£|lis 




Phallodrilus sp. 


Family - 


Hesionidae 




Tubif icoides 




Gvptis brevipalpa 




heterochaetus 




Oohiodromus abscura 




Tubificoides sp. 




Podarke sp. 


Family 


- Naididae 








Paranais litoralis 



(continued) 
51 



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 16. (Continued.) 




Superorder - 


Eucarida 


Family - 


 Processidae 


Order - De' 


capoda 




Ambidexter 


Family - 


Penaeidae 




symmetricus 




Penaeus aztecus 




Processa 




Penaeus duorarum 




fimbriata 




Penaeus setiferus 




Processa 




Trachypenaeus 




hemph i 1 1 i 




constrictus 




Processa sp. 




Trachypenaeus 


Fami ly - 


• Cambaridae 




similis 




Procambarus 




Xiphopenaeus 




penaensalanus 




kroyeri 


Family - 


• Callianassidae 




Sicyonia 




Call ianassa 




brevirostris 




atlantica 




Sicyonia dorsal is 




Call ianassa 


Family - 


Sergestidae 




jamaicense 




Acetes americanus 


Family - 


 Paquridae 


Family - 


Palaemonidae 




Paqurus 




Leander 




bonairensis 




tenuicornis 




Pagurus 

"lonqicarpus 




Macrobrachium 






oh i one 




Pa^^urus 




Palaemonetes 




poll i car is 




intermedius 


Family  


- Ma.iidae 




Palaemonetes 




Libinia dubia 




pugio 
Palaemonetes 




L i b i n i a 






emarginata 




vulgaris 
Periclimenes 




Metaporhaphis 






calcarata 




americanus 




Podochela riisei 




Periclimenes 


Family • 


- Portunidae 




lonqicaudatus 
Alpheidae 




Callinectes 


Family - 




sapidus 




Alpheus 




Callinectes 




armillatus 




simil is 




Alpheus formosus 




Ova li pes 




Alpheus 




guadulpensis 




heterochaelis 




Portunus gibbesii 




Alpheus normanni 


Family • 


- Xanthidae 


Family - 


Oqyrididae 




Eurypano^eus 




Ogyrides li mi col a 




depressus 


Family - 


Hippolytidae 




Hexapanopeus 




Hippolyte 




angustifrons 




zostericola 




Menippe 




Latreutes 




mercenana 




parvulus 




Neopanope 




Lysmata 




packardii 




wurdemanni 




Neopanope texana 




Thor dobkini 




Panopeus herbstii 




Tozeuma 




Rhithropanopeus 




carolinense 




harrisi i 






Family 


- Grapsidae 
Sesarma cinereum 






Family 


- Ocypodidae 
Uca minax 



53 



Table Ifi. (Concluded.) 



Family - Porcel lanidae 
Petrol isthes 

arm at us 

Cliba narius 

vittatus 







Family - 


Leucosi idae 
Persephona 
mediterranea 




Superorder - 


Hoplocarida 






Order - Stomatopoda 






Family - 


Squil lidae 
Squil la empusa 


CI 


ass 


- Ostracod 


a 
Ostracoda sp. 


CI 


ass 


- Branch iura 








Argulus sp. 



Subphylum - Hexapoda 
Class - Insecta 

Insect larvae 
(several unident.) 
Order - Diptera 

Family - Chironomidae 
Chironomidae 
Ablabesmia sp. 
Chironomus sp. 
Cladotanytarsus so. 
Clinotan'ypus sp. 
Coelotanypu s sp. 
Cryptochironomus 

ful vus 
Cryptochironomus 

sp. 
Dicrotendipes sp. 
Glyptot endipes sp. 
Har nis chia sp. 
Microtendipes sp. 
Nanocladi us sp. 
Orthoclad ius sp. 
Parachironomus sp. 
Polypedilum sp. 
Procladiu s sp. 
ProcladiTis sp. 
Tanypus sp. 
Tanytarsus sp. 
Family - Heleidae 
Rezzi a sp. 
Order - Odonata 

Suborder - Anisoptera 

2 unident. spp. 
Suborder - Zygoptera 

1 unident. sp. 
Order - Ephemeroptera 
Family - Caenidae 
Caenis sp. 



Phylum 



Family - Heptaqeni idae 
1 unident. sp. 
Family - Baetidae 

Cal libaeti s sp. 
Order - Plecoptera 

1 unident. sp. 
Order - Hemiptera 
Family - Corixidae 

1 unident. sp. 
Order - Lepidoptera 
Family - Pvr al idae 

Nymphula sp. 

Echinodermata 

Echinarachinus 



parma 



Echinaster sp. 
Hemipholus 

elongata 
Luidia clathrata 
Ophiothr ix 

anqulata 



54 



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 grass- 
bed soecies include Heteromastus 
fil iformis and Hob soni 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 poly- 
chaete 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 depen- 
ding on recruitment patterns and response 



55 



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Bosed on 48 2 -mm otter trowl tows token monthly In Apolochicolo Estoory 1972 - 1982 

Bosed on 48 2 - mm otter trowl tows token monthly m Apolochicolo Esluory 1972 - 1982 

Figure 27. Summed numerical abundance and number 
of species of benthic infauna and epibenthic fishes 
and invertebrates in East Bay and Apalachicola Bay 
from 1972 to 1982 (from Livingston unpubl.). Data 
are presented as monthly means _+l standard devia- 
tion of the mean. 



59 



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 alabamae j^ and striped bass 
( Morone saxatil i T) (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 



Anchoa mi tchil 1 i 



1, 

2. Micropogonias undulatus 

3 . Cynoscion are narius 

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. Paralichthy s letiTostigma 

20. Synodus foetens 

21 . Eucinostomus arqenteus 

22. Dasyatis sabina 

23. Cynoscion nebulosus 

24. Microgobius thalassinus 

25. Urophy cis floridanus 

26 . Lagodon rhomboides 

27. Gobiosoma bosci 

28. Chaetodipter us f aber 

29. Orfhop ristis 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 pensacola e 



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 
Olig opl 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. Busyco n 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 concentra- 
tions 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 % Domin- 

per sample (a, ance (by 

Station samp le d ry wei qh t^) num bers) 



Dominant species 



Margalef 
richness 



OUTER BAY- 



1— IX 73.? 



A. FISHES 



- 1 


43.4 


46.? 


39 


MICROPOnONIAS UNDULATUS 
ANCHOA MITCHILLT 


3.77 


- lA 


18.0 


47.5 


41 


ANCHOA MITCHILLT 
MICROPOnONIAS UNDULATUS 
LEIOSTOMUS XANTHURUS 


3.43 


- IE 


55.9 


53.9 


77 


LEIOSTOMUS XAMTHURUS 


3.54 


- IC 


51.5 


75.1 


43 


MICROPOGONIAS UNDULATUS 


3.48 



171.8 



ANCHOA MITCHILLI 

34 LAGODON RHOMROIOES 
RAIRDIELLA CHRYSURA 
ORTHOPRISTIS CHRYSOPTERA 



3.55 



RIVER 
DOMINATED 



1— 2 



-- 3 



96.4 



44.5 



100. Q 



'^5.6 



31.3 



46.0 



46 ANCHOA MITCHILLT ?.88 

MICROPOGONIAS UNDULATUS 

44 ANCHOA MITCHILLT 3.8? 

LEIOSTOMUS XANTHURUS 

49 ANCHOA MITCHILLT 3.14 

MICROPOGONIAS UNDULATUS 
BREVOORTTA PATRONUS 



UPPER 
(EAST) BAY 



4A 64.6 48.0 
5 74.3 76.6 



5A 101.4 



— 5B 



74.1 



60.9 



28.? 



— 5C Q0.8 27.0 

— 6 109.9 53.5 



47 LEIOSTOMUS XANTHURUS 3.30 

44 ANCHOA MITCHILLT 3.Q0 

MICROPOGONIAS UNDULATUS 
LEIOSTOMUS XANTHURUS 3.01 

47 ANCHOA MITCHILLT 

LEIOSTOMUS XANTHURUS 
MICROPOGONIAS UNDULATUS 

47 ANCHOA MITCHILLT 2.99 

LEIOSTOMUS XANTHURUS 

47 LEIOSTOMUS XANTHURUS 3.0^ 

33 ANCHOA MITCHILLT 3.98 

LEIOSTOMUS XANTHURUS 
MICROPOGONIAS UNDULATUS 
BREVOORTTA PATRONUS 



66 



Table 20. (Continued. 



Number Biomass per % Domin- 
per 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 BAY- 



RIVER 
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 


10.1 


11.9 


48 


CALLINECTES SAPIDUS 
PENAEUS AZTECUS 


1.81 


IC 


6.4 


9.5 


27 


PENAEUS DUORARUM 
LOLLIGUNCULA BREVIS 
CALLINECTES SAPIDUS 


2.82 


IX 


16.3 


8.8 


57 


ACETES AMERICANUS 
CALLINECTES SAPIDUS 
PENAEUS DUORARUM 


1.86 


2 


38.5 


28.0 


70 


PENAEUS SETIFERUS 


1.68 


3 


12.2 


6.2 


49 


CALLINECTES SAPIDUS 


1.43 



52 



PENAEUS SETIFERUS 

PENAEUS SETIFERUS 
CALLINECTES SAPIDUS 



1.38 



(continued) 
67 



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 


67 


PENAEUS SETIFERUS 
PALAEMONETES PUGIO 


1.24 


9.0 


57 


PENAEUS SETIFERUS 
CALLINECTES SAPIDUS 


1.45 


3.9 


65 


PENAEUS SETIFERUS 
CALLINECTES SAPIDUS 


1.18 


5.1 


53 


CALLINECTES SAPIDUS 
PENAEUS SETIFERUS 


1.39 


5.2 


54 


CALLINECTES SAPIDUS 
PENAEUS SETIFERUS 


1.11 


11.1 


50 


PALAEMONETES PUGIO 
PENAEUS SETIFERUS 


1.17 


8.4 


41 


LOLLIGUNCULA BREVIS 


3.28 



CALLINECTES SAPIDUS 
PORTUNUS GIRBESI 
ACETES AMERICANUS 



but in July, the highest concentrations of 
the sand seatrout are found at the mouth 
of the Apalachicola River. Distribution 
usually remains relatively unchanged 
during August and September. The 
remaining fish, dwindling in numbers 
during the fall months, soread out 
throughout East Bay and northern 
Apalachicola Bay. By winter or early 
spring, as noted above, no sand seatrout 
are taken. 

Spot have a different pattern of 
distribution (Figure 32). As they move 
into the estuary in Jaunary, spot tend to 
congregate in upper East Bay and around 
Nick's Hole drainage off St. George 
Island. This distribution broadens 
throughout eastern portions of East Bay 
and Apalachicola Bay during February and 
March. Concentrations of spot appear in 
areas of the bay that receive freshwater 



runoff from upland areas. East Bay is a 
particularly important nursery area for 
this species. By summer, remnants of the 
population are found off St. Georae 
Island. 

The spatial distribution of 
postlarval penaeid shrimp in the 
Apalachicola estuary illustrates the 
summer and fall dominance of these species 
(Figure 33). During early summer, they 
are concentrated in East Bay. However, 
during July and August, high numbers of 
penaeids are located at the mouth of the 
Apalachicola River. By fall, although 
still concentrated in East Bay, they tend 
to be more evenly distributed throughout 
the estuary as they move into the open 
gul^ to spawn. Few shrimp are taken 
during the winter months. As with other 
dominant (and commercially important) 
species in the bay, the penaeids appear to 



68 





VWJOCOJ 




i 


p wcbitI i 
eumf -  


-~T'r 


5^ 


Jvm»si 


Ha >^^= 


^mm 


IC 




January 











wMjocau ^^^^^^ 


m 


•^^^Vl 


y^ 


fi"^ a 


**"JKL^^ -v 


L^^ 


ff^E^i 


€^^ 


it-,4K'^ 


Februar) 









( 


^y 






^ 


B? 




AMlJOm.J^'' 


^ 


F^^ 


y 


-^ 




Z j^ 




^^ "^-*— ~ • t^ 


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/ SUM 


s\ 


-v_ . ^--' — tS 


U 




April 











WfcUlDCOL* 


t 


1^ 




FHVB 


c^ 


QL 




Af»L»O«0l»^^" 


P 


V^ 


ST «*aNiy 


i 


=*w 


^-^m 


BlAHt/ 


tfjtLMy^ BAT 




^J"^^^ 


^£i]j^ 


■W^ y-* ^^ 


^- 


/si ixwa 






p 


'' SLwa 






^»ss ^^ 






v= — ^ ^ j^^snJe 


CUT 




July 











( 


^^ 






|K, 




WIL KHCOL^^Lj^ 


Hpi 


ST WOHJp^S 




r^ 




^ vulacmcol* my 


/sTMOBa 
■^ tSlUHO 


wkTv,. 






CJlSS >v^ 


/ . . ..^-"''stS cut 




October 









AMLKMCOLA 


{ 


w 




«Mi»Dcoiy 


fi 


^\ 


ST VMCniA. 


^^.,«.^ 


( 


^^ 


was >t 


)')i^ 


5 CUT 


/si OKIW 
' BUWD 


March 













^ 






*«L«:iC0i^'T 


) 




S1 VMCatir^ 


1../: 


^ 


3^ 


PUS ^MM 


(//^ 


CUI 


/sTawMB 

X SLMC 


IVla> 











< 


^ 




tfWUCMCO* 1 


mfL 




„„^^^B 


■jRl 


/ 


"^"^^^"l 


HH 


ST VMCBIT^ 


^ aML*d«t»« rIi, "^ 


^ 


MIO ^<, 


'~^J_^^i^ 




June 













^^ 




RfVBt 


< 






WAUCHCOllV^ 


/ 


^sii^oJ 


/ 


/^^^ 


( 


Ju 


51 VMCBII/ 

KiAie/ 


/4<» 


^ 


\ 




_rE;U.^--5a 




jTsT aDHM 


Aiijiusi 

















>^ 






AMlJOCOl* 


■i 


"fwLr 






W«1«J«JR^|_^ 




J^ 


n ji 


^ \ 






f^m 


S1A»^S 


-^^V 






\^^^ 


^^ 




C^ji 




Jr'vam 


>wN^ 


= 


) 'fe 


^ 


r BUM 


»MS ^ 


"^^. 


f.^ 


OJI 




Seplem 


ber 











inujDCOui 


( 


^ 








^Sl 


SI VMCfxr/ 
SIM)/ 


t^ 




Kraoha 


«tV_ 

f»SS % 




S CUT 




Novem 


ber 









AMIKOWL* 




i^ 


<^ 


^= 


^ 


^r^^i"^^ 


ST VWUdrXv' ^ 

Bum* / ^V 


JMitCHCftAai 


y-' 


'i>^ 


^7\ 


\ (^i 




^ SVC 




X„-Aai 


CUT 




December 









Anchovies 



l^di^idu^l^ per l\Mi-Miniili' l^l^^l lo^^ In Miinlh 

0.0 - S.n [ ] 20.0 - 50.0 

5.0 - 10.0 ^^ 50.0 - 90.0 

10.0 ~ ^^1 <>0.0 - 



Figure 29. Average monthly distribution of anchovies ( Anchoa mitchilli ) in the 
Apalachicola estuary from 1972 to 1979. 



69 





»p»lm:miloi» 


iV^ 




"V^ 


/cowl 




""flJ^Ss^ 


\/M 


^ 




^^'l 


SI ncEsi£-~~ 


% ^^ 


-^ \ 


ctwe/T^^^ 


^r;55-^ 


i/sTOORa 




/"^ > 


** isiwe 


«3T%i3-> 






PASS >; 






January 







Atlantic 
Croaker 





Rrvpi 


C 


^ 


x;; 


"^ 




P\ 


SI VHUNll ^ 


^(™rAffltoi,«_a«. 


^ 


■^ 1SIU« 


P*SS >v^ 


. ^-^ 


d 




Februarx 











*/'»L*CMC0li1 






^ 




*P»L»£MICOl^fi 






^n 


ST vwcai/'^ 


^R 




1 


IP 


P*SS >t 


/f -W 


^* 


^ 




April 


--_LL„--'^™r 


lu 











1 


^y 




VAUOCOUt 


J 


^P 




/Sml 




APALW-flCIXO 


\ 


1 


ST nesfj^- 


AfWJiM«LOlA UA 


) 


y 


PASS >^ 

July 


. ^ 


Ts cui 


' istwc 





AfAUWHCOU 






*ML«ijc.ai\ 


\ 


IT ^HScJ 




\ 


Bi**/ 


JMLweoi/Mr 


six ^m 


•bt'>^=- 


^/ 


'*'^ 


PWI ^^.^^ 


_,^'-'t^[ 




October 









ap«l«:mcol4 


Q 


w 




"% 


jtf^ 


M^; 




»f»l»cmkoia5u 


i 


HLj|| 




^^^^^^P^^ 


n 


e^^PSa 


S' VHlKjU. 


w^B/^ 


•^ 


^^1 


>si«m/"^ 


~oiS*w' 




\^^^^ 


c^ 


c^^ 


) 


.1 .-iWM 


»*ss ^v 


. / 


y 




March 











c 

AP*t*CMCOl» / 


^ 




ApftL «:iiK:Di^yx5*vA 


IKA 


y. 


^""■"'"T^^^s 


^l^s 


^^ 


^v.  ' JjfflW. 


^^^^^i 


aiMBf  


■»<ij^_^^--v^ 


^^ 


wcNI:;-"- 


^^■"■.■;::_\e^ 


' Kl*M) 


May 


v;^- ' ;■-.■ ;■ ',>'''s«iB CUT 







APALACKOl* 
RIVIB 


^ 


L»5i a*v /ui 




apalacmcoiA!^ 




^'^l/j 




C^O 




■<^M 


ST vnanfl 




< 


\^P 


June 


N,^ __.>-^'''srie 


CUI 


^ »l*« 





( 


.^ 'if 




APAltDClRA 

BTVDl ^_/ 


/P«N1 


S1 IfHCENI/ 

slam/ 


apalachkolO 

AP«^«.H^uli> W 


3 


rfE 


. ^ 


j/ ISLAND 


PASS ^^= 

AugusI 





^ 




v»iJDca* ^ --^ 




^^>^--:"=i-^"----^K)«n 


^ 


APAL«C«ca?^ '\^ \ 


ST tr*C8)T/ 
Slum/ 


v.^-*rt«ju&w jif^^ 


PASS >L 


J*' ELA« 


Seplemhcr 





AfAUCWOLA 


^ /p(WL 


*«iLJ«o*ca.ir^.-* 


SiS 


,-^,--='^, 


.^x-^ 




ST ncB.'/ 


\ 


"SiAw/ i«iic«;iX' a'' 


y^ 




--j' SiJM) 




CUI 


November 






December 



lndiMdu:)N pi-r I«oAlinii(v Irjwt lii« b> Miinlh 



0.0 - 5.0 
5.0 - 10.0 
rO.O - 20.0 



20.0 - .10.0 
,10.0 50.0 
50,0 - W.O 



Figure 30. Average monthly distribution of Atlantic croaker ( Micropogonias 
undulatus) from 1972 to 1979. 



70 







^ ^ 








F=?=^ 




APUJOCOUt 


Ara*;!,, 


^^^/^''^ 


/«<i 


, 


^^..-J 


\ 


St nKBiiK 
BUM)/ 


V*u«ia*Dt^ a^T 


^^ SUWD 


PASS %. 


__,^ 




Januar> 







Sand 
Seatrout 





VUJOKDU 


^ ^ 


/^J-v 


^ 


MtiicMcaA: 


I 


5T V»CB.I/ 

esLiwe/ 


v«L.«CJitfflJ a'J' 


i 




_^ 


j^ suwe 


Februar> 









»P«U«CMCOU 


^7" 




RFVBl 








\ 


SI «CENT^^ 


aSSESE 


\ 


61 Wc/ 


V^LMrtCai.^ BAT 


/S~ GtWIU 

^ tsuwo 


•^Nsi. 






f«5 >^ 


»^ .^-'-SiT-, 




April 







( 


^w^ 


««llJO«0U 1 


^ii*' 


».uo^^^p 


"S^M^tui 




2^ 1 


BU«J^^^ JT" ^.J-^^^ 


jfj^ ^=?ga:>*_2. ^"^ 


j""''^ 




' SlMCI 


^^s.. il ^j-'-'itS fjjr 




July 







4PW.ACHC0L" 
RIVER 


^ 


/^VVwwi 


y 


u>ij.ȣnKao 




Vjv/^ 


SI «<f nJ^v 


v»L»DflCOL»>*r^ 


:ir 




»Ici^ 

f»SS >i 


/ .^'-^ 


CUT 




Oclober 













e^ 




HHrtR 






UPAucMca^ 


^ 


psiwe/ 


V»LACMKC(.* Wr 


^ 


WE '"v. 


_--^ 


/s" GfOflGE 


March 









( 


!# 




"^^/^ 


M^b: 




^C^^^M^ 


^^ 




\^K 


'^XJ 


SLMiO Av,^ 


. "" '(? 


^)^ 


«stV^ 


y^ 




P»SS \^^ 


<_ ,,--''t^cui 




Ma> 









RIVER 


i 


^ 


^ 


*P»lACMICOl Offlft 


1 


s 


Si»>*d/  


\;™^^J3C 


% 


' isuue 


PASS nA 


■^^ . ^,f'''^^\i 


OJI 




June 














w»i*o«a» 


1 
^ 




>< 


<^ 


tfajiOgaxjSgl 




^^ 


SI VfCfM/ 


I 


S 


;5^ 


t^^ 




/ 


__.,-i3i 


CUT 




Sepiemb 


er 

















/^ 


™^^ ^ 


y 




""'"""■ ^----^ 


WE5I%. 
P«iS >, 




^^ Sl_«Nt 


Novem 


ber 









^^ 






'^^^^^ 


APALACMCOm 


^ 


prm^ 


WalacmciaVj 


\ 


\ 


„ .^^j'^'^ L 


/ 


\ 


'■"""'/ 6P4i«d.H|i.uu wi- 




^^ 


p*ss ^*^E - ,---'^ 


S CUI 


fvti*^ 


December 







lndiMdii;jK pi-r iMii-MJntik- l^u^^l \tn\ h\ Monlli 



0.0 - 2.0 
2.0 - 5.0 
5.0 - lO.O 



10.0 - 15.0 
15.0 - 25.0 
25.0 - .W.O 



Figure 31. Average monthly distribution of sand seatrout ( Cynoscion arenarius ) 
in the Apalachicola estuary from 1972 to 1979. 



71 





WW tcMCDi a 


^ 




\i/^ 


'^W' w/mi 




MUlKDCOlp 


<7| 


SI nCEMT/ 

tsinw/ 


i^iJ^IaJ a 


^ 








I '^ 


^ U«NC 


WBI*^ 






WSS >v 


( ^-'ISTl^ 




January 







July 




Spot 




Ki'briiar> 







■f"!: 


r 

1*11 




««U.«D«0lO%.,^__, 


I s 


a 


SI VICBd/S. 


V^ V*i Aca^ wfl 




jCi 


KSS >^ 








April 











W*l<C«COl» 


>7 




^s^- 


sT^ 


^^ 




\ 


SI V«C»T/ 

Elmo/ 


v^LWuauiw 






,-^ 




Oclober 









u>*i lotr.n « 


o» 




WWH 


.^^^. 




WW (CHcni >S -^\ 


v^ 


/ 


r^ 


^pl 


51 nKiHif 




pp^ 








March 









WWKMCni 


0^^ 




"¥^ 


•'^./piwi 




•«n«o«)i?5[_,,J Z' 


V^ 


^ 


.^-^--^^U 


"^^ \ 


SI ivatif 


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Figure 32. Average monthly distribution of spot ( Leiostomus xanthurus ) in the 
Apalachicola estuary from 1972 to 1979. 



72 





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20.0 - 30.0 
JO.O - 70.0 



Figure 33. Average monthly distribution of penaeid shrimp ( Penaeus spp.) in 
the Apalachicola estuary from 1972 to 1979. 



73 





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Blue Crabs 






Figure 34. Average monthly distribution of blue crabs ( Callinectes sapidus ) 
in the Apalachicola estuary from 1972 to 1979. 



74 



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 communi- 
ties) 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 (soft- 
sediment) 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 litter- 
associated 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 litter- 
associated 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: 

UJ 

m 

3 



< 

Q 

Z 

m 
< 

_i 
< 
o 

tr 
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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 





30- 




• station 5a ( ohgoholine ) 
 Stotion 3 ( oligoholine ) 
' ' Station IX ( mesohcline ) 








25- 






o 




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CO 
LlI 

o 

UJ 

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5 10 15 20 25 30 35 

SALINITY 

Figure 36. Regression of numbers of 
species of litter-associated 
macroinvertebrates on salinity at three 
stations in the Apalachicola estuary. 
Samples were taken over a 12-month period 
in oligohaline (stations 5A, 3) and 
mesohaline (station IX) areas. 



anaerobic or microaerophi 1 ic bacteria. 
The data siiggest that distinct populations 
may choose different microbes. The 
component energy linkages are poorly 
understood, however. Little is known 
concerning the protozoan components of 
litter associations, although preliminary 
analyses in East Bay indicate that 
ciliates constitute the dominant protozoan 
inhabitants of the litter assemblages (D. 
Cairns, pers. comm.). 

In summary, phvsical/chemical 
features such as temperature and salinity 
influence the spatial-temporal 
distribution of litter-associated 
macroinvertebrates in the estuary. Such 
distribution is also determined by 
productivity trends and the biochemical 
features of the microbial communities. 
The detri ti vorous macroinvertebrates serve 
as a link between the microbial producers 
and important estuarine fishes and 
invertebrates that feed on these species 
(Laughlin 1979; Livingston et al. 1977; 
Sheridan 1978, 1979; Sheridan and 
Livingston 197Q). 



78 



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 oyster- 
associated 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 long- 
term 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 soft- 
sediment 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 




Factor 1 
(49.0% of 
variance) 


Factor 2 
(22.3% of 
variance) 


Factor 3 
(17.9% of 
variance) 


Factor 4 
(10.8% of 
variance) 


River flow 




-0.82 


-0.08 


-0.07 


-0.08 


Local rainfall 




-0.04 


-0.30 


-0.09 


0.20 


Tide (incoming 


or outgoing) 


0.26 


0.61 


-0.68 


0.06 


Tide (high or 


low) 


0.09 


0.39 


0.61 


-0.37 


Wind direction 


(E-W) 


-0.02 


0.09 


0,36 


0.37 


Wind direction 


(N-S) 


0.10 


-0.20 


0.22 


0.31 


Secchi 




0.57 


-0.07 


-0.17 


0.24 


Color 




-0.80 


0.33 


0.01 


0.07 


Turbidity 




-0.73 


0.54 


0.08 


0.23 


Temperature 




0.38 


0.15 


0.02 


-0.18 


Salinity 




0.68 


0.21 


0.23 


-0.02 


Chlorophyll _a 




0.47 


0.51 


0.09 


0.31 



benthic macroinvertebrates taken 
(seasonally) in litter baskets at 
different stations (3, 5A, IX) along a 
salinity gradient (Figure 36) (F = 30.4, 
r^ = 0.45, with S as the dependent 
variable). Numbers of species taken 
during a season vary directly with 
salinity rather than with station-specific 
characteristics. Similarity coefficients 
of species composition at the sampled 
stations are closest during fall periods 
of high salinity. These results indicate 
that quantitative and qualitative species 
representation, regardless of location, 
are closely related to salinity. 

Similar trends are found for phyto- 
plankton (Estabrook 1*^73), zooplankton 
(Edmisten 1979), infaunal 



macroinvertebrates (Livingston unpublished 
data), and epibenthic fishes and 
invertebrates (Livingston 1979). 
Livingston (1979) showed that salinity is 
directly related to species richness and 
diversity of estuarine nekton. Stations 
characterized by low salinity are 
associated with high numbers of 
individuals, high relative dominance, and 
low species richness (Table 20). Outer 
bay stations, with higher salinities, are 
defined by relatively low dominance, high 
species richness and low numerical 
abundance. High densities of organisms 
that use the bay as a nursery, such as 
penaeid shrimp, blue crabs and various 
finfishes are not usually found in areas 
having stable patterns of relatively high 
salinity (Livingston 1984a). 



il 



Table 22. Correlation coefficients of linear regressions of nitrate, orthophosphate, 
silicate, and ammonia on salinity (from Livingston et a1. 1974). 



Date 




NO3 


PO4 


Si03 


NH3 


Oct. 14 1972 


T 
B 


-0.70 
+0.12 


-0.73 
-0.14 






Dec. 2 1972 


T 
B 


-0.88 
-0.75 


-0.20 
-0.55 


-0.98 
-0.85 




Jan. 6 1973 


T 
B 


-0.55 
-0.84 


-0.89 
-0.82 


-0.P9 
-0.87 




Feb. 17 1973 


T 
B 


+0.00 
+0.58 


-0.Q5 
-0.11 


-0.33 
-0.002 


-0.02 
-0.15 


Mar. 19 1973 


T 
B 


-0.95 
-0.97 


-0.78 
-0.60 


-0.98 
-0.998 


-0.85 
-0.45 


Apr. 22 1973 


T 
B 


-0.76 
-0.62 


-0.77 
-0.62 


-0.03 
-0.80 


-0.67 
-0.93 


May 19 1973 


T 
B 


-0.88 
-0.96 


-0.54 
-0.65 


-0.998 
-0.99 


-0.48 
-0.81 


Jun. 11 1973 


T 
B 


-0.60 
-0.94 


-0.01 
-0.61 


-0.995 
-0.93 


-0.55 
+0.06 


Jul. 12 1973 


T 
B 


-0.82 
-0.80 


-0.10 
+0.42 


-0.97 
-0.93 


-0.82 
+0.03 


Aug. 22 1973 


T 
B 


-0.90 
-0.91 


+0.04 
-0.84 


-0.95 
-0.94 


-0.50 
-0.91 


Sep. 10 1973 


T 
B 


-0.99 
-0.^8 


-0.29 
+0.15 


-0.995 
-0.^9 


-0.83 
-0.98 



Species richness and diversity of 
nekton are directly associated with areas 
of high environmental stability but low 
secondary productivity. Infaunal 
macroinvertebrates show the same general 
response to salinity (Livingston 1983d). 
Within a given area of low salinity, 
however, species richness may increase in 
areas of relatively high primary 
productivity and detritus availability. 
In this way, the influence of salinity may 
be modified by ambient habitat conditions. 

In low-salinity estuaries, species 
diversity indices tend to reflect the 
effects of salinity on recruitment of 



dominant populations. Within a given 
habitat (such as an oyster bar, 
unvegetated soft-sediment area, or 
seagrass bed), the spatial distribution of 
organisms at any given time may depend on 
gradients of productivity and salinity. 
The regulating features may change their 
relative importance through any given 
seasonal succession. Temperature and 
other physical features seasonally modify 
the productivity-salinity association. 
Among the phytoplankton, water temperature 
is the primary limiting factor, although 
river discharge, nutrients (mainly phos- 
phorus), turbidity, and light inhibition 
may control phytoplankton productivity at 



82 



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 poly- 
chaetes, 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 water- 
column 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 relation- 
ships 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 dis- 
tribution, 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 poly- 
chaetes, shrimp, and crabs. Spot tend to 
exploit smaller organisms, such as nema- 
todes, harpacticoid copepods, juvenile 
bivalves, and smaller forms of poly- 
chaetes. 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 



CALANOI D COPEP QDS 
-^ 

OTHER _iO0PL^NK.T0N_ 




, INVERTBBRATES 

 ANCHpyiES 




\ ^ ANCHOVIES 

PENAElp SHRIM P 



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 subsur- 
face), 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 trawl- 
susceptible catch, five ( Peaneus 
setiferus , Palaemonetes pugio , Callinect es 
sapid us, Penaeus azte cus 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, soecies- 
specific recruitment patterns, predator- 
prey 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 assem- 



blages (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 long- 
term 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-to- 
year 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 ff iiiiii|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 




Size 2 


Size 3 


Salinity 


Temperature 


River flow 


Rainfall 


Size 1 
(1-30 mm) 


r 
T 


0. 
0. 


,323 
,156 


-0.690* 
-0.554* 


-0.616* 
-0.351 


-0.774** 
-0.534** 


-0.450 
-0.260 


-0.070 
-0.040 


Size 2 
(31-60 mm) 


r 

T 






0.147 
-0.015 


-0.526 
-0.325 


-0.212 
-0.294 


-0.570 
-n.387* 


0.340 
-0.236 


Size 3 
(60 mm) 


r 
T 








0.172 


0.690* 
0.656** 


-0.017 
0.040 


0.135 
0.108 


Salinity 


r 

T 










0.586* 
0.330 


-O.QIS** 
-0.697* 


0.306 
0.060 


*p < 0.05 
**p < 0.01 



















Table 25. Multiple stepwise regression of seasonal variations of 
frequencies of blue crabs from three size groups and abiotic 
variables (N=12 months). Variables represent mean monthly averages 
using 7-year data. Salinity and temperature means are baywide over 
14 stations in the Apalachicola estuary (from Laughlin and 
Livingston unpubl ). 



A. 


De^en 
Step 


dent 


variable 
Variables entered 


Size (< 30 

r2 


mm 


carapace width) 
Significance 


B. 


1 
2 
3 
4 

Depen 
Step 


dent 


temperature 

rainfall 

size 2(31-60 mm) 

size 3( 60 mm) 

variable 

Variables entered 


0.559 
0.800 
0.890 
0.Q08 

Size 2 (31- 


-60 


0.003 
0.001 
0.0001 
0.001 

mm) 
Significance 


C. 


1 
2 
3 

Dependent 
Step 


ri verf low 

size 3( 60 mm) 

size 1 (1-30 mm) 

variable 

Variable entered 


0.323 
0.348 
0.430 

Size 3 ( 

R? 


60 


0.054 

0.146 (N.S) 
0.191 (N.S) 

mm) 
Significance 




1 
2 
3 

4 




temperature 
ri verf 1 ow 
size 1(1-30 mm) 
size 2(31-60 mm) 


0.478 
0.570 
0.650 
0.704 




0.013 
0.022 
0.028 
0.048 



'iZ 



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, long- 
term 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 winter- 
spring 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 

I- 

Li_ 

o 

h- 

UJ 

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 long- 
term 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 short- 
term 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 pa fr onus 
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 gulf- 
coast 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 channel- 
maintenance 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 attri- 
butes 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). 









County 






Total 
(square mi 




Land use 


Gulf 


Liberty 


Calhoun 


Jackson 


Gadsden 


Franklin 


les) 


Low density 
residential 


2.50 


0.25 


13.50 


7.00 


2.00 


1.00 


26.25 




Medium density 
residential 


0.?5 


— 


1.00 


1.50 


1.00 


1.00 


4.75 




Commercial 


1.50 


0.25 


4.50 


2.25 


0.25 


0.-'5 


11.00 




Industrial 


— 


0.25 


0.50 


1.50 


0.25 


1.00 


3.50 




Recreational^ 


38.00 


194.50 


146.00 


22.00 


-- 


58.00 


458.50 




Marshes and 
flood lands 


06.00 


83.00 


29.00 


16.00 


9.00 


45.00 


288.00 




Agriculture 


12.00 


12.00 


73.00 


399.00 


20.00 


2.00 


518.00 




Forestry 


175.00 


32.00 


314.00 


114.00 


46.00 


21.00 


702.00 




Water 


14.50 


0.50 


3.50 


33.00 


0.50 


0.50 


52.50 




Total 


349.75 


322.75 


586.00 


5^6.75 


79.75 


12P.75 


2,064.50 





^Includes Apalachicola National Forest. 



chain reaction throuqhout the local 
economy because direct and indirect 
purchases generate income, the so-called 
"multiplier" effect. Recent estimates 
indicate that the forestry and fisheries 
"export" values are even more important 
than previous studies indicated since 
oractically all such production is sold 
outside the region. The total current 
value of fisheries in the drainage system 
and associated coastal areas exceeds '^'^^3 
million. Colberg et al. (1%8) projected 
a value of S34.2 million for commercial 
fishing and tourism by the year 2000 if 
water quality and natural productivity are 
maintained. Value added as a "multiplier" 
effect would increase this estimate to 
almost S67 million. Thus, the as yet 
undiminished natural resources in the 
Apalachicola valley provide an important 
economic base for the local area, and such 
natural industries have a direct influence 
on the region throuqh export and 
respending. 



Rockwood et al. (1973) and Rockwood 
and Leitman (1977) provided an in-depth 
analysis of the socioeconomic basis of the 
Aoalachicola oyster industry. The 
potential for oyster production has yet to 
be reached; greater production will be 
necessary if the relatively low per-capita 
income is to be increased and more 
employment is to be provided for young 
people in the area. Tn terms of general 
determinants of regional growth, Franklin 
County is rich in natural resources on 
which it is almost entirely dependent. 
Recent historic trends have contributed to 
the insularity of the community. The 
development of strong clan ties of the 
English and Scotch-Irish inhabitants adds 
to the geographic isolation of the region. 
Independence and individualism are hall- 
marks of this society and have led to the 
view that outside intervention by 
government agencies or large corporations 
has a negative influence on the community. 
The oyster industry is based on 



102 



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 free- 
flowing 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, toxa- 
phene), 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 Developme nt 

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 consi- 
derable 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 rela- 



tively 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 



and Florida yew 

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 long- 
term 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 

an 

be 



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, includ- 
ing actual and proposed purchases accord- 
ing 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 physico- 
chemical 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 


1°, 


265 


9 


1.23 


27 


4.2 X 


K 


3200 


19 


1.46 


166 


3.8 X 


lo" 


390 


6 


1.42 


3 


3.3 X 


1942 


10 


1.52 


97 


1.1 X 


10^ 


11500 


7 


0.73 


56 


2.2 X 


loi 


122 


5 


0.43 


51 


3.8 X 


10 

lo; 

lo" 


1251 


6 


0.46 


45 


1.1 X 


305 


3 


0.15 


26 


4.4 X 


210 


2 


0.55 


6 




lof 


1070 


3 


0.41 


12 


4.0 X 


176 


2 


0.30 




1.6 X 


10^ 


1240 


2 


1.5 


107 






445 


4 


1.3 








490 


6 


1.7 


320 


9; 


J 


32 


8 


0.43 


2 



"Approximate annual 
freshwater input 



al mean hydraulic residence time. The 
to Barataria Bay has not been reported. 
"Below Smyrna River. 
■-Area includes mud flats, mean depth = 6 m excluding flats. 



Livingston (1981b), in a comparison 
of the distribution of various sciaenids 
in estuaries alonq the northeast Gulf of 
Mexico, found that the Apalachicola 
estuary is extremely productive in terms 
of fish populations (Table 31). Prime 
habitats include the mud flats of East Bay 
and the mouth of the Apalachicola River 
and the grass beds in Apalachicola Bay off 
St. George Island. The unpolluted, highly 
turbid estuary, with its high plankton 
productivity and abundant allochthonous 
detritus, presents an optimal environment 
for benthic omnivores (such as croaker and 
spot) and epibenthic carnivores (such as 
silver perch and sand seatrout). The 
Econfina estuary is a relatively clear, 
unpolluted system dominated by benthic 
plants (macrophytes), which provide the 
major source of productivity and habitat 
features for other organisms in the area. 
This estuary, which receives considerably 
less overland runoff than the Apalachicola 
system, is dominated by fishes associated 
with the extensive seaqrass beds in the 
area. Although fish productivity is 
relatively high, the sciaenids are not as 
well represented and account for only 
about ?0% percent of the total fish 
catches over the 9-year sampling period. 

The Fenholloway estuary, polluted for 
over 20 years by pulpmill effluents, is 



Table 28. Estimates of particulate pri- 
mary production in various estuaries in 
the United States (after Nixon 1983). 



Location 



Primary production 
g C m"'' y"l 



Mid Narragansett Bay 


310 


Mid Long Island Sound 


205 


Lower New York Bay 


483 


Lower Delaware Bay^ 


206 


Mid Chesapeake Bay*^ 


445 


Patuxent Estuary 


210 


Pamlico Estuary 


200-SOO 


Apalachicola Bay 


360 


Barataria Bay^ 


360 


San Francicsco Bay 




Suisun Bay 


95 


San Pablo Bay 


100-130 


South Bay 


150 


Kaneohe Bay 


165 



^Below Leipsic River, 80^ of total bay 
production. 
''Four-year mean (1«574-1Q77). 
•^Phytoplankton 165, Benthos 195. 



largely devoid of benthic plants and has 
an increase in phytoplankton productivity 
and associated planktonic food webs. 
Relatively high levels of phytoplankton 
productivity (derived from anthropoaenic 
input of nutrients) are correlated with 
increased reoresentation by fishes 
associated with planktonic food webs. 
Overall fish productivity has been 
severely reduced because of the impact of 
the pulpmill effluents on the biological 
organization of the estuary. Although the 
overall abundance is low, sciaenids are 
well represented in terms of numbers of 



113 



Table 29. Approximate land use distribution and population density surrounding the 
estuarine study areas (Nixon 1983). 











Population 


Study area 


Developed 


Aqriculture 


Other 


density 




{%) 


{%) 


{%) 


(people/acre) 


Narragansett Ray 


37 


6 


57 


l.S 


Long Island Sound 


?9 


3 


68 


1.1 


New York Bay 


40 


— 


60 


3.? 


Raritan Bay 


39 


14 


47 


1.^ 


Delaware Bay 


27 


35 


38 


0.? 


Chesapeake Ray 


27 


24 


49 


1.2 


Patuxent Estuary 


36 


21 


43 


0.4 


Potomac Estuary 


27 


22 


51 


0.1 


Pamlico Estuary 


3 


21 


76 


0.02 


Apalachicola Bay 


1 


21 


77 


0.3 


Mobile Ray 


13 


15 


73 


1.=^ 


Barataria Bay 


in 


41 


49 


2.3 


San Francisco Bay 


18 


22 


60 


4.6 


Kaneohe Bay 


32 


10 


58 





species in the Fenholloway estuary. This 
phenomenon can be attributed to the fact 
that the pollution altered the natural 
habitat in such a way as to induce a 
superficial resemblance to the 
Apalachicola estuary. This altered 
habitat favored plankton-feeding and mud- 
flat species as part of an unstable 
succession of adventitious populations in 
the polluted estuary (Livingston l^^RPb). 

Compared with other estuaries, the 
Apalachicola system has relatively low 
finfish landings, while blue crab landings 
are moderately high (Nixon 1983). 
However, in terms of oyster yield per unit 
area, the Apalachicola estuary was the 
second highest of those systems surveyed 
(Nixon 1'383). Although the connection 
between fishery yields and primary 
production remains largely undetermined in 
a quantitative sense, the importance of 



the response of individual species to 
varying sets of environmental conditions 
probably plays a considerable role in the 
form and direction of secondary production 
in any given system. Also, socioeconomic 
factors are important in the definition 
and use of a qiven Fishery resource. 

It is clear that relatively little 
has been done to compare various 
ecological characteristics of different 
estuaries. °art of the problem lies in 
the difficulty of carryina out 
simultaneous long-term studies in separate 
estuaries usinq comparable methods o^" data 
collection. The organization, funding, 
and execution of studies on more than one 
such system is difficult (Nixon 1983). It 
is clear that more comoarative studies 
will be necessary if we are to understand 
the significance of the driving 
environmental features of any qiven 
estuary. 



114 



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 


Sewage 


Total 


Percent 
sewage 


R60 
60 


390 
40 


950 
100 


41 


130 
10 


270 
20 


400 
30 


67 


5,700 
800 


26,230 
3,750 


31,930 
4,550 


82 


200 

50 


1,260 
280 


1,460 
330 


86 


650 
70 


650 
70 


1,300 
140 


50 


340 
50 


170 
30 


510 
80 


33 


310 
60 


290 
50 


600 

110 


48 


420 
80 


390 
60 


810 
140 


48 


860 
250 


minor 


860 
250 


< 1 


550 
210 


10 
3 


560 
213 


2 


1,206 
370 


80 
30 


1,280 
400 


7 


570 

^'go 


minor 


570 
290 


< 1 


1,100 
160 


910 
130 


2,010 
290 


45 


minor 


1,600 
310 


1,600 
310 


100 


50 
10 


180 
30 


230 
40 


78 



Narragansett Bay 

Long Island Sound 

New York Bay 

Raritan Bay 

Delaware Bay 

Chesapeake Bay 

Patuxent Estuary 

Potomac Estuary 

Pamlico Estuary 

Apalachicola Bay 

Mobile Bay 

Barataria Bay 

Northern San Francisco Bay 

South San Francisco Bay 

Kaneohe Bay 



(continued) 
115 



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 study 
The 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 


320 


68 


46 


68 




263 


263 


100 


50 


50 

22 
3 





^Data rounded to the nearest 10 units, compiled and calculated for various years 
from different sources. 



116 



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 
41 
37 
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 ( 


56.3) 


338 


13.4) 


156 


6.2) 


59 


2.3) 


53 


2.1) 


50 


2.0) 


47 


1.9) 


44 


1.8) 


42 


1.7) 


40 


' 1.6) 


3' 


' 1.3) 


28 


' 1.1) 


23 


' 0.9) 



Centropristis melana 
Paracl inus fasciatus 
Syngnathus scovel i 
Chasmodes saburr ae 
Cynoscion nebulosu s 
Lucani a parva 
Microgobius gulosus 
Chilomycterus schoepf i 
Urophycis flo r idanus 
Anchoa mitchil 1 i 
Haemulon plumieri 
Sphoeroides nephelus 



Fenholloway estua ry 



Anchoa mitchi 1 1 i 
Leiostomus xanthur us 
Lagodon rhomboides 
Bairdiella chry suFa* 
Anchoa hepsetus 
Orthopristis chrysoptera 
Eucinostomus gula 
Eucinostromus argenteus 
Gobiosoma robustum 
Paracl inus fasciatus 
Chilomyceterus schoepf i 



Micropogonia s undul atus 
Syngnathus scovel li" 
Urophycis floridanus 
Cyno scion arenarius 
Opsanus beta 
Stephanolepis hispidu s 
Micrognathus cri nigerus 
Sphoeroides nephelus 
Polydactylus octonemus 
Cynos cion nebulosus 
Monacanthus ci 1 i atus 
Centrop r i s tis melana" 
Syngnathu's floridae 
Etropus crossotus 



19 ( 


0.8) 


18 ( 


0.7) 


18 ( 


0.7) 


16 ( 


0.6) 


13 ( 


0.5) 


13 ( 


0.5) 


9 ( 


0.3) 


9 ( 


0.3) 


8 


0.3) 


8 


0.3) 


8 


0.3) 


7 


0.2) 


231 


26.3) 


228 


'25.9) 


95 


'10.8) 


53 


' 6.1) 


36 


' 4.1) 


26 


' 2.9) 


23 


[ 2.6) 


19 


( 2.^) 


15 


' 1.7) 


12 


( 1.4) 


in 


[ 1.2) 


9 


' 1.1) 


8 


( 1.0) 


8 


( 1.0) 


7 


( 0.9) 


6 


[ 0.7) 


6 


( 0.7) 


5 


( 0.6) 


5 


( 0.6) 


5 


( 0.6) 


5 


( 0.6) 


4 


( 0.5) 


4 


( 0.5) 


4 


( 0.5) 


4 


( 0.5) 



117 



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Movement, microhabitat, exploitation, 
and management of Gulf of Mexico 
sturgeon, Apalachicola River, Florida 
(in review). 

Wooley, C. M., P. A. Moon, and E. J. 

Crateau. 1982. A larval Gulf of Mexico 
sturgeon ( Acipenser oxyrhynchus 

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

Yerger, R. W. 1977. Fishes of the 
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 pre- 
sent. 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 ash- 
free 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 repeti- 
tive 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 ash- 
free 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, inver- 
tebrate, and plant populations are assembled and punched on standard 80-column 
cards or entered directly via a computer terminal. Upon completion of a preli- 
minary 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 tele- 
type 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 data- 
base 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 two- 

137 



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 







C 






B 




C 










C 






R 


R 


C,R 


R 


R 






C 







ROWS 





1 Al 1 


1 1 • 1  1 1 












~ P 






P 
P 






— 



Matrix 

two-dimensional form 

(table) 



PLANES 



Matrix 
three-dimensional form 



In the above diagrams, each cell in a sample column vector has been 
labelled with a "C", each cell of a sample row vector with an "R," and each cell 
of a plane vector with a "P." 



An individual 
numbers are, by con 
tions indicated by 
with "C" above are 
always be referred 
tion by row, column 
can be described as 
The three numbers c 
dimensional, as in 
simply comprise a _s 
(2,1,1), where all 
cell in a single-ce 



row, column, or plane may be referred to by a number, and 
vention, assigned in order (starting with 1) in the direc- 
the arrows in the diagrams above. Thus all the cells labeled 
contained in column "3." An individual cell in a matrix can 
to by a unigue set of three numbers, one each for its posi- 
and plane. Thus the locus for the cell labelled "A" above 
row 1, column 2, and plane 3, or alternatively, "(1,?,3)." 
an always be assigned, even if the matrix is effectively two- 
a table, or even one-dimensional (e.g., a "matrix" might 
ingle cell ). The point "B" above could be located by 
the cells in a table would be assigned plane number 1. A 
11 "matrix" would therefore be located at (1,1,1). 



b. Rationale for the MATRIX System 



There are 
First, many an 
Easy Graphing, 
either in row 
points to be u 
files contain 
dispersed thro 
sis. Data poi 
raw data files 
if, over a Ion 
ferent format) 



two underlying reasons for the de 
alytical program packages such as 

and the SYMAP spatial mapping sys 
and column form or in some other s 
tilized occur together (and seguen 
data points that, for a certain de 
ughout the file; they must be "bro 
nts to be analyzed together might 
. This dispersion of data points 
g period of time, many different k 

are collected and entered as comp 



velopment of the MATRIX system. 
SPSS, BMDP, MINITAB, PLOT-10 
tem reguire input data that is 
pecial form in which all data 
tially). Second, many raw data 
sired analysis, are in some way 
ught together" prior to analy- 
even be scattered over several 
can be especially troublesome 
inds of data (each with a dif- 
uter data files. 



The above conditions result in what could be called a "format gap." There 
are two aspects of this gap: one is that the raw-data format is not suitable 
for direct entry of the data into an analytic routine; the other is that data 
points required for an analysis do not occur together. The MATRIX program 



138 



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 manipula- 
tion 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 manipula- 
tive 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 higher- 
level 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 func- 
tion 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 for- 
mat 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 spe- 
cies. 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 subana- 
lyses 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 


NIND (no. of indiv.) 


DATE (YYMMDD) 


BIOMASS 


STATION 


NSAMP (no. of samples) 


SPECIES 




TOO 




SEX (invertebrates only) 




AREA 


NIND 


YRMON 


BIOMASS 


STATION 


NSAMP 


SPECIES 




TOD 




SEX (invertebrates only) 




AREA 


NIND 


MONTH 


BIOMASS 


STATION 


NSAMP 


SPECIES 




TOD 




SEX 




AREA 


DRY WT (dry weight) 


DATE (YYMMDD) 


WET WT (wet weight) 


STATION 


NSAMP 


GENSPE 




TOD 




AREA 


DRY WT 


YRMON 


WET WT 


STATION 


NSAMP 


SPECIES 




TOD 




AREA 


DEPTH CHL A 


DATE 


SECCHI RIVFLOW 


STATION 


DISS02 RAINFALL 


TOD 


COLOR NITRATE 


DEPTHCODE 


TURBIDITY PHOSPHATE 



TEMP 

SALINITY 

PH 



PRDCTVTY 
AMMONIA 



(continued] 
142 



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 


CHL A 


SECCHI 


RIVFLOW 


DISS02 


RAINFALL 


COLOR 


NITRATE 


TURBIDITY 


PHOSPHATE 


TEMP 


PRDCTVTY 


SALINITY 


AMMONIA 


PH 




BRILL DIV 


DAP 


SHANN DIV 


MACl 


BRILL EVEN 


MAC2 


SHANN EVEN 


HURLBERT 


SIMPSON 


TOTNIND 


MARGALEF 


NSPECIES 



large data base will only be read once. The SLECT command can then be used to 
create smaller load/sort files, which contain the data for specific analyses. 
When these smaller files are input to MATRIX, GENERATE will only have to read 
in relevant data points and the collapse specifications will be easy to enter. 

Currently, all SPECS commands have been placed within the MATRIX operating 
system, and the SPECS system has been reduced to a data access system. 



143 



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 multi- 
disciplinary array of scientists, and a permanent staff of post-doctoral fellows 
and full-time personnel, R. J. Livingston has put together a series of multi- 
disciplinary 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 macroinver- 
tebrates 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, epi- 
benthic 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 Profile 



S. 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 impor- 
processes 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 respon- 
sibility 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.