FWS/OBS- 84/09

May 1984

THE ECOLOGY OF DELTA MARSHES OF COASTAL LOUISIANA: A Community Profile

Fish and Wildlife Service

Corps
U.S.

of Engineers

105 L8

U.S.

Department of the

Interior

Department of the Army

G67

:5EJP

Salt marshes along the Mississippi deltaic channels (Photograph by Charles Sasser)
.

coast

characterized

by

extensive

tidal

/05'
FWS/OBS-84/09 May 1984

THE ECOLOGY OF DELTA MARSHES OF COASTAL LOUISIANA:

A COMMUNITY PROFILE

by
James G. Gosselink Center for WefTahd Resources Louisiana State University Baton Rouge, LA 70803

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Project Officer Edward C. Pendleton National Coastal Ecosystems Team U.S. Fish and Wildlife Service 1010 Gause Boulevard Slidell, LA 70458

Performed 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

The findings in this report are not to be construed as an official U.S. Fish and Wildlife Service position unless so designated by other authorized documents.

Library of Congress Card Number 34-601047.

This report should be cited:
1984. The ecology of delta marshes of coastal Louisiana: Gosselink, J.G. U.S. Fish Wildl. Serv. FWS/OBS-84/09. 134 pp. profile.
a

community

PREFACE
report is one of a series of and Wildlife Service Community Profiles synthesizing the available literature for selected critical ecosystems into comprehensive and definitive referThe this ence sources. of objective particular account is to review the information available on the marshes of the Plain. The River Deltaic Mississippi in North river system is the largest It drains an area of 3,344,560 America. km^. Over the past 5,000 years the river has built a delta onto the continental shelf of the Gulf of Mexico covering about This low land is primarily 23,900 km^. marshes and represents about 22 percent of the total coastal wetland area of the 48 The delta is conterminous United States. notable for its high primary productivity, its valuable fishery and fur industry, and the recreational fishing and hunting it supports.
This
U.S.
Fish

marsh degradation due to a complex mixture of natural processes and human activities include worldwide sea-level that rise; and extractive subsidence; navigation control flood industry canal dredging; measures that channel the and river; pollution from domestic sewage, exotic organic chemicals, and heavy metals.
The future of the marshes in this region is in jeopardy, and if they are to be saved, it is important to know how they function and what measures can be taken to arrest the present trends.
questions or comments about this publication or requests for the report be should directed to the following address.
Any

same time, the Mississippi At the River Deltaic Plain marshes are subject to the unique problem of extremely rapid

Information Transfer Specialist National Coastal Ecosystems Team U.S. Fish and Wildlife Service NASA/SI idell Computer Complex 1010 Gause Boulevard SI idell, LA 70458

CONVERSION TABLE

Metric to U.S. Customary

Mul tiply

BiL

millimeters (mm)

CONTENTS
Page

PREFACE CONVERSION TABLE F IGURES TABLES ACKNOWLEDGMENTS

i i 1

iv
vi
i

xii
xi v

INTRODUCTION MAN IN THE MISSISSIPPI RIVER DELTA HISTORY OF DELTA RESEARCH WETLAND DEFINITIONS, TYPES, LOCATION, AND EXTENT
THE REGIONAL SETTING CHAPTER ONE. THE CLIMATE, THE OCEAN, AND THE RIVER Insolation Temperature Water Balance GEOLOGICAL PROCESSES Pleistocene sea levels Modern Mississippi Delta

1 l

4
5

9 9 9

10 10 14
14 18
IN

CHAPTER TWO. TEMPORAL AND SPATIAL GRADIENTS MARSHES TEMPORAL GRADIENTS SPATIAL GRADIENTS Flooding Soils
Sal t

DELTA
28 28 32 33 34 35
36

Nutrients Vegetation
Soil

37

ECOLOGICAL PROCESSES IN DELTA MARSHES... CHAPTER THREE. PRIMARY PRODUCTION Emergent Vascular Plants Epiphytic Algae Benthic Microflora in Marsh Ponds Submerged Grasses in Marsh Ponds DECOMPOSI TION CONSUMERS Benthos Nekton Wildlife Carbon Budget NUTRIENT CYCLES Nitrogen Phosphorus Sulfur STORMS

43 43 44 54 55 55 55 60 60 61 63
71

74 74
76

77 77

CHAPTER FOUR. THE MARSH IN THE COASTAL BASIN COUPLINGS AMONG ECOSYSTEMS Intra-Basin Couplings Extra-Basin Couplinqs Intercontinental Couplings TEMPORAL USE OF MARSHES

79 79 79

80
81 81

CHAPTER FIVE. WETLAND VALUES, HUMAN IMPACTS, AND MANAGEMENT WETLAND VALUES Wetland Harvest Environmental Quality Esthetics Confl icting Values WETLAND EVALUATION WETLAND MANAGEMENT Marsh Loss and Salt Intrusion Habitat Qual ity Water Quality
REFERENCES

84 84 84 85 86 86 87

89 89
93 96

100

APPENDICES 1. Plant species composition of salinity zones in the Louisiana coastal marshes 2. Marsh plant decomposition rates, Mississippi River delta marshes 3. Fishes of the Mississippi River Deltaic Plain that are found in marshes and associated water bodies.. 4. Representative vertebrate species of marsh habitats in the Mississippi River Deltaic Plain

116
116

118 120
^28

VI

FIGURES

Number
Salt marshes along the Mississippi deltaic coast characterized Frontispiece. by extensive tidal channels
1

Page
iMcinp

COVER
2

The groves of trees in the middle of this broad expanse of marsh identify the site of old Indian villages The oil storage facility for the nations' s only superport is constructed in a salt dome in the middle of a Mississippi delta The maze of pipes is the primary aboveground brackish marsh. An old oilfield also sits atop this submerged salt dome expression. as shown by the network of tree-lined oilwell access canals

Across this expanse of marsh and swamp looms the Mew Orleans skyline through the haze, a reminder of the proximity of heavy industries and concentrated popul ations Louisiana oil and gas production
Map of the Mississippi units

4
5

4
5

River Deltaic Plain showing the hydrologic

7

The seasonal variation of insolation at various latitudes. The computation assumes a transmission coefficient of 0.7 throughout
Mean monthly air temperature at New Orleans, Louisiana

9 10 10

8
9

Generalized water budget for the Mississippi delta marshes

Average water budget for the upper Barataria basin, 1914-1978. P=precipitation, PE=potential evapotranspiration, AE=actual evapotranspi ration

11

10

All

Freshwater inflows to the Mississippi Delta. Discharges are in cumecs. discharges are for water year 1978 except Mississippi River, which is a long-term mean representing the combined average annual discharge above the confluence of the lower Mississippi (10400 cumecs) and the Atchafalaya (5000 cumecs) Rivers

12

11

Water level trends in delta marshes:, c) daily

a)

long term; b) seasonal;

13

12

Tide levels at Shell Beach, in the Pontchartrain-Lake Borgne basin, associ ated wi th ni ne major storms
vii

14

Number
13

Page
The relationship of glacial advance and retreat to continental exposurs and sedimentation during the Late Quarternary

shelf
25

14

Location of major buried river channels formed during the Wisconsin gl acial period
The position of major delta lobes on the gulf coast during the 20,000 yr B. P. (A) Late Wisconsin, 25,000 previous 25,000 years. (C) Early Holocene, 12,000 (B) Late Wisconsin, 15,000 yr B. P. P 1,000 yr B. 10,000 yr B. P. (D) Present, 5,000

15

15

17

If)

Deltaic lobes of Mississippi River deltas

18

17

Satellite image of the Mississippi Delta Region showing delta lobes of different ages
Six subdeltas of the modern Mississippi Balize Delta recognized Dates indicate year of crevasse maps and sediment analysis. opening
Plan view and cross sections through A-A' and B-B' deposition in a crevasse
from

19

13

20

19

of environments of
21

20
21

Sequential development of Cubits Gap subdelta

22

Linear, areal subdelta

and volume growth curves for the Cubits Gao

23

22

Composite subaerial growth curve, Mississippi River subdeltas. Total subaerial land determined from averages at 10-yr intervals
The accelerating wetland loss rate in the Mississippi

24

23

Delta

24

24

Conputerized re-creation of the west side of Barataria Bay showing the Black is open change in wetlands between 1945 (a) and 1980 (b). water; marshes are shown as varying shades of grey
Environmental
Mineral

25

25 26

succession of an idealized delta cycle

29

arranged
27

content of marsh soils in Mississippi delta hydrologic units, in order of increasing age

30

Marsh soil salinity and percent fresh marsh in Mississippi delta Soil marshes by hydrologic unit, arranged in order of increasing age. is a mean for the whole basin weighted by ares of each marsh salinity zone. The fresh marsh is percent of total marsh area

30

28

Marsh edge length :area ratio and total marsh edge length for delta The units are arranged in order of increasing hydrologic units. age
Net primary production and fishery yield of Mississippi River Deltaic Plain hydrologic units. Production calculated from average production of each habitat type and its area in the hydrologic unit. Shrimp data from Barrett and Gillespie (1975). Basins are, in order of increasing Pontchartrain-Lake Borgne, II - Balize, III -Barataria, IV aye: I

31

29

Terrebonne,

V -

Atchafalaya

VI

-Vermilion
viii

31

Number
30
31

Pag*?

Seasonal

salt marsh inundation patterns

34

Variation in soil density and soil carbon content with distance inland from the strean edge in a salt marsh in the Barataria basin
(A) Mean seasonal Sedimentation rates on the Barataria saline marsh. Mean seasonal sedimentation sedimentation 1975 - 78. (B) 1975 - 79. Sedimentation rates were highest during the winters Hurricane Bob and tropical storm Claudette of 1975 - 78. passed through the area during the summer of 1979, resulting in very high deposition rates

34

32

34

33

The decrease in free soil water salinity (mg/g) of chenier plain marshes with distance (km) from the gulf

36

34

Concentrations of available Na, Ca, zones

Vegetation zones
in

K,

Mg,

P,

and N in different marsh

37

35

the Mississippi

River delta marshes

^g

36

Note the "hummocky" appearance which A deltaic plain brackish marsh. The birds with black-tipped is typical of Spartina patens stands. wi ngs are white pel icans , the smal ler ones ducks , mostly teal
A diverse deltaic plain fresh marsh scene. Species are: Sagittaria falcata (foreground), Typha sp. (right edge), mixed grasses and vines, Myrica shrubs in rear

40

37

41

38

Vegetation zonation in an intermediate marsh transition zone in the Factors arise from statistical clustering techniques Barataria basin. and are identified by the dominant species
Effects of substrate drainage conditions on the dry weight accumulation by (A) Spartina alterni flora and (B) S^. cynosuroides
A conceptual

41

39

42

40

model of

typical wetland ecosystem, showing major

coTiponents and processes

41

44
45

Monthly growth rates of Panicum hemitomon and Spartina alterniflora

Seasonal changes in live and dead biomass of Phragmites austral is and Spartina patens during 1973 - 1975

42

47

43

Production of intertidal S. alterniflora vs. mean tide range for Different symbols represent various Atlantic coastal marshes. di f ferent data sources
Variation in total aboveground biomass and height of Spartina alterniflora with distance inland from the marsh edge in a Barataria basin sal t marsh

49

44

50

45

Gulf-inland variations in live and total al terni flora marshes

Effects of NaCl

biomass in Spartina
50

46

concentration

in

the root medium on the rate of Rb

ix

Number

Page

absorption by excised root tissue of and D. spicata

47

S^.

a1

terni flora
51

This "key" intermediate in Metabolic conversions of pyruvic acid. metabolism can be converted to a variety of end products, depending on the organism and the electron acceptors available

52 53

48

Marsh soil
Seasonal
a

transfonnations that result from tidal

flooding
in

49

changes in various physical, chemical, and biotic factors Barataria basin salt marsh

54

50

Net epiphytic production on stens of Spartina a1 terni flora collected at the water's edge and inland 1.5 m with the averages, extremes, and fitted curve for the water's edge production superimposed

55

51

Number of shore-line epiphytic diatans/cm culm surface area of Results are pooled averages for four stations Spartina al terni f1 o ra. and height classes

55

52

Disappearance of S^. patens Pontchartrain-Borgne basi n

litter from litter bags

in

the

58

53

Decomposition rates (mg/g/day) of S^. al terni flora in 2 -mm mesh bags in different locations
Major pathways of organic energy flow in salt marsh and associated water bodies
a

litter incubated
59

54

Mississippi River deltaic

51
in

55

Length class frequency of qulf menhaden captured Pontchartrai n

and near Lake

52

56

Density of vegetation, detritus, and consumers at the edge of the salt marsh
Pelt production from marsh zones in coastal
Annual muskrat harvest from in the Mississippi Delta
a

53
55

57

Louisiana
ol

53

52,200-ha brackish Scirpus

neyi

marsh
55

59

Ground plan of surface trail s

typical

muskrat house with underground runways and

55

60

A muskrat "eat-out"

in the brackish marsh in the Barataria basin. Note the high densi ty of muskrat houses
a

57

61

Carbon dioxide flux measurements in marsh communi ty

deltaic salt
72

62

Carbon budget of a Mississippi River deltaic salt marsh (see Table 29 Rates (g C/m^/yr) are from CO2 flux measurements, for sources). numbers in parentheses, which are fran other sources except
A schematic outline of the redox zones in a submerged soil showing The aerobic layer has been some of the N transfonnations. In reality it is seldom over 1-2 mm in drawn thick for clarity.

72

63

flooded marshes
X

75

Number
64

Page

Nitrogen and phosphorus budgets for salt marsh

Mississippi deltaic
75

65

Conceptual diagram illustrating the coupling of delta marshes to other ecosystems

79

66

Patterns of estuarine use by nektonic organisms basin, Louisiana

The life cycle of the brown shrimp

in

the Barataria

80
81

67

63
69
70

Major duck migration corridors to gulf coast marshes

Seasonal
use of wetlands by migratory birds, shellfish, and

82
fish

83

The The increase in open water in natural and impounded wetlands. of greater wetland loss in impoundments is consistent in both pattern fall, when water levels are low, and winter when impoundments are f 1 ooded

92 94

71
72

Wildlife management areas

in

the Mississippi Delta

The strong flow of water across A weir in the deltaic plain marshes. These the weir is an indication of the effectiveness of the barrier.

structures are favorite sport fishing spots

73

95

Cumulative number of days per year that ponds in the study area will Based on equal or exceed certain percentages of bottom exposure. contours of 48 ponds and 20 years of tide data on the central depth Louisiana coast
The percentage of different types of vegetation in impoundments
in

96

74

the
97

Rockefeller State Wildlife Refuge

75

Habitat type, vegetative cover, and fish and wildlife values achieved with water management programs operating on the Rockefeller Refuge

98

XI

TABLES

Nu"iber
1

Page

Salinity values (ppt) recorded by various investigators for delta marshes

Classification of coastal marshes of the Mississippi River Delta, and area of different marsh zones in 1978
Average coastal

subnergence on the U.S. east and gulf coasts

13

Land-use changes along the northwest edge of the Barataria basin, on the Bayou Lafourche natural levee
Land use changes,
in

26

hectares, in the Mississippi Delta, 1955-78

27

Regression analyses relating net primary production (NPP) and inshore shrimp production (1955-74) in hydrologic units to various physical parameters
The annual duration and frequency of inundation of marshes in the Barataria basin, Louisiana

32

33

Marsh accretion rates (mm/yr) 1963 ^''Cs fallout peak

in Louisiana delta

marshes, based on the

35

Concentration (C) and accumulation rates (A) of organic carbon, nitrogen, phosphorus, iron, and manganese in Louisiana delta marsh
soil s 35

10

Multiple linear regression models of soil control their distribution

ions showing what factors 36

11

The ratio of the major cations to the chloride ion in normal seawater and in the saline, brackish, inte mediate, and fresh marshes of Louisiana

37

12

Percent cover of the dominant plant species in major marsh zones of the Louisiana coast

38
....

13

Production of marsh vascular plant species

in

the Mississippi

Delta

45

14

Belowground biomass of Mississippi delta marsh plant species

Production estimates for different techniques
a

48

15

Spartina alterniflora stand based on

48
xi
i

Number
16

^age

Year-to-year variations in peak live biomass of Spartina alterni flora at a single site in the Barataria basin
Spartina alterni flora root alcohol dehydrogenase (ADH) activity, adenosine triphosphate (ATP) and ethanol concentrations, and soil in a Louisiana sal t marsh
Percentage of marsh community ;netabolism by Spartina alterniflora
Submerged aquatic plant species composition of ponds and lakes by marsh zone along the Louisiana coast
Range and mean loss rates (mg/g/day) of litter from different marsh plant species (summarized from Appendix 2)

49

17

Eh

52

18
19

54

55

20

59

21

Monthly occurrence and abundance of the fish species collected in smal 1 sal t marsh ponds

62

22

Wildlife species richness (number of species) in the chenier plain marshes

Muskrat house-building activity in 10-ha brackish and salt marsh areas in Barataria basin Density of waterfowl basin in 1980-81

64

23

66

24

(number/100 ha) by marsh zone in the Barataria

68

25

Density of ponds and lakes of various size classes in marsh zones along the Louisiana coast, August, 1968
The percent of the area of ponds and lakes covered with submerged vegetation in August, 1968 by size and marsh zone

69

26

70

27

Density of wading birds and pelicans (number/100 ha) by marsh zone, in the Barataria basin, 1980-81
Birds of the Mississippi Deltaic Plain on the Audubon Society "Blue List," indicating that their populations are declining

70

28

71

29

Estimates of different components of the carbon budget of Mississippi deltaic salt marsh community

73

30

Influence of Spartina alterniflora plants on recovery of ^^N-ammonium added over 18 weeks to soil cores The estimated economic value of harvests from the Barataria basin, Louisiana

76

31

85

32

Estimates of the economic value of Louisiana's coastal wetlands comparing willingness-to-pay approaches with energy analysis approaches

89

33
34

Major wetland issues and human impacts

in

Mississippi delta wetlands ....

89

Impacts of canals in Louisiana coastal marshes leading to habitat loss, and mechanisms and management practices to minimize these impacts
xiii

90

ACKNOWLEDGMENTS
the to profile is dedicated graduate students in Marine Sciences and Fisheries at Louisiana State University (LSU), Baton Rouge, who carried out much of the original field research upon which I have been privithis profile is based. to work closely with thirteen of leged them. Fifty- six literature citations in the profile are authored or coauthored by students. They have made a major contribution toward unravelling the ecology of our coastal ecosystems.

This

have been fortunate to enjoy a coworkers for the last 15 years group and introduced me to the marshes who participated with me in the intellectual Their research. of wetland stiiiiul ation contributions are cited throughout this John Len work. Bahr, Day, They are Roland Parrondo, Jim Charles Hopkinson, Stone, Gene Turner, and more recently Bob Costanza, Flora Wang, Bob Baumann, Deborah Fuller, Gary Paterson, and Charles Sasser.
of

Mendelssohn (LSU), Gerry Bodin [U.S. Fish Wildlife Service (FWS), and Lafayette, La.], Ed Pendleton (FWS National Coastal and Steve Matheis, Team), Ecosystems and John Weber (U.S. Army Suzanne Hawes Corps of Engineers, New Orleans District) reviewed an early draft and provided many it. for improving suggestions helpful and Susan Thanks to Lyster Kathryn EcoCoastal National Lauritzen (FWS edited and did the systems Team); they for the respectively. profile, layout Dawnlyn M. Harris provided word processing Diane Baker, as usual, did a assistance. She the job figures. drafting superb created the cover. My wife, Jean, showed great forbearance and understanding while word the hours over I labored long
,

processor.
Much of the research cited in this manuscript was supported by the National U.S. and the Grant Sea Army Program New The Orleans. District, Engineers been this profile has preparation of sponsored by the Office of the Chief of Engineers in association with the WaterLower the Station, Experiment ways Mississippi Valley Division, and the New Orleans District, Anny Corps of Engineers, and the Fish and Wildlife Service.

Special thanks are extended to Jim Coleman, who drafted the geology section and of this report, and to Linda Deegan Jean and Walt Sikora for information and advice on the benthos and nekton sections, Irv and John Day respectively.

XIV

INTRODUCTION

The history of the marshes of the River Delta is inextricably Mississippi intertwined with the history of the river Like some ancient god, it broods itself. over the coastal plain, implacable in its With its power, its purpose inscrutable. verdant sediment it spawns the flat, marshes of the delta, nourishes them with and finally abandons them its nutrients, under the influence of to senesce slowly the time and subsidence, while it renews cycle elsewhere along the coast.

marshes. The idea of energy flow in ecological systems is still only a guiding principle; the complex details of in molecular the marsh biochemistry of and the the substrate complexity meiofaunal food chain are still largely unexplored.
the coastal
the human This monograph details and to understand, through struggle understanding to manage the Mississippi I will emphasize what we delta marshes. know - and that is considerable - but I hope that the presentation of technical detail does not obscure the large areas of how to the about manage uncertainty Above all I hope that it does system. to cold not reduce the delta marshes statistics; for understanding, I believe, is heightened by emotional involvement.

This community profile deals with the facts and the quantitative analysis of But the cold numbers often this cycle. How much is defy our comprehension. 15,400 cubic meters per second (cLnecs) the average discharge of the Mississippi River? How large is 0.2 y, the size of a And what does it mean to say bacterium? that there are one thousand million of then in a cubic centimeter of marsh soil? are almost These scales unimaginably different, yet understanding a natural ecosystem demands the ability to deal with both.
,

MAN IN THE MISSISSIPPI RIVER DELTA

the technical details like a coastal marsh, the system complexity becomes increasingly apparent, and the cold, technical analysis breaks down more and more often into a sense of wonder at the system's sophistication and the delicate interplay of parts that make whole. the Migratory waterfowl's up to respond to subtle environmental ability cues and navigate thousands of miles from Alaskan prairie potholes to the Louisiana our most rivals marshes coastal sophisticated inertial guidance systems. After years of study we still have little understanding of how passively floating shrimp larvae in the Gulf of Mexico find their way through estuarine passes into
As

one examines

of

When de Soto found and named the Rio Esperitu Santo, now the Mississippi had been in the Indians 1543, River, on the coast for 12,000 years. living They preferred the easy living of the marshes to the uplands because food was abundant and easy to harvest. Oysters and the Ranqia clam were in nearly endless Fish, turtles, and edible plants supply. were plentiful. The tribes now known as Tchefuncte, Marksville, Troyville, Coles and Creek, Caddoan, Mississippian, the settled on slightly Plaquemine banks of river distributaries elevated where they literally ate themselves up out As they ate oysters and of the water. clams, the shells accumulated beneath them. The evidence of these prehistoric villages now dots the marshes as small groves of trees on slightly elevated shell mounds in an otherwise treeless vista (Figure 1).
del

De Soto approached the river from the It was 140 years Peninsula. Florida next before the LaSalle, European, in the coast 1682, explored having He claimed the approached from upriver. river for basin drained by the great France and in 1584 led an expedition to establish a colony at the mouth of the in this he failed river. Although he was and lost his life, attempt, followed by Iberville, who explored and river and by Bienville, who the mapped established New Orleans in 1718.

the main channel) levees for agriculture. Rice, indigo, tobacco, corn, cotton, and later sugarcane were the large plantation crops, but many other crops brought in from Europe and elsewhere were also grown. During this period Germans settled part of In the coast, beginning in about 1720. an 1760 influx of French refugees from Eastern Canada began. These poor farmers, trappers, and fishermen brought with them a strong culture still characteristic of the coastal villages (Kane 1943). One hundred years ago Louisiana had only about 900,000 inhabitants (Kniffen to led the 1968). Many developments industrialized state. The present construction of the levees along

settlenent phase that began the in of the resulted development distributary (a diversion near the mouth water out of of a river that distributes
Thus
a

The groves of trees in the middle of this broad expanse of marsh identify the Figure 1. site of old Indian villages (Photograph courtesy of Louisiana State University Museum of Geosciences, Robert Newman, curator).

Mississippi River did much to develop a and of sense encourage permanence also The levees industrial expansion. waterborne by transportation promoted channelling the Mississippi River and its to distributaries, deepen Dredging became new ones and create channels more fostered These commonpl ace. further and stimulated transportation commercial expansion.

New industries developed based on The late Louisiana's coastal resources. 1800' s and early 1900' s were a time of widespread harvesting of the extensive The fishing cypress forests of the coast. But and fur-trapping industries expanded. the most significant event in the state's life was the discovery of oil in Jennings
in

1901.

are in reserves Louisiana Oil concentrated around salt domes that occur

across the coastal wetlands and on the continental shelf. The inland fields were developed first. An enormous expansion of petroleum demand began in the war years of This 1941-45. resulted in dredging thousands of miles of canals through the coastal wetlands for access to drilling and sites for pipelines, constructing enormous refineries and petrochemical and facilities, processing secondarily stimulating many other industries (Fi-gures 2 and 3). As oil and gas reserves were in the inland marshes, production depleted moved offshore. This shift increased pressure for more and deeper navigation canals to link the offshore rigs with land-based facilities. Production of oil and gas reached its peak in 1971 and has since been declining (Figure 4). However, search for new oil the continues, and no has means modification wetland by Louisiana's wetland management stopped. problems continue to be related to its

The oil storage facility for the nation's only superport is constructed in a Figure 2. The maze of pipes is the salt dome in the middle of a Mississippi delta brackish marsh. An old oilfield also sits atop this submerged salt dome primary aboveground expression. access canals (Photograph by Robert as shown by the network of tree-lined oilwell

Abernathy)

major
and

coastal industries fossil fuel extraction,

transportation

HISTORY OF DELTA RESEARCH

of and Investigations geological of the aspects biological Mississippi Delta both followed the same historic trend from descriptive accounts to greater In emphasis on functional processes. geology early studies are typified by that of Lerch et al. (1892), who carried out a fairly inclusive preliminary survey of Louisiana that included geology, soils, and Davis' (1899) groundwater. physiographic interpretation ushered in the "golden age" of coastal geomorphol ogy (Fisk 1939, 1944; Fisk and McFarlan 1955; Russell 1967; Kolb and Van Lopik 1936, and many others). This was a 1958; period of deciphering the geomorphol ogy of the delta on a scale and regional

the major qualitatively documenting i^ormative In the last 20 processes. the has shifted to years emphasis at intensive usually investigation, specific locations, of nrocess-response rel ationships
.

In the arena early biological comments on delta biota were common, at first emphasizing economically important animals such as furbearers. De Montigny (1753, as quoted in Gowanloch 1933), who spent 25 years in Louisiana, and Le Page and fish du Pratz observed (1758) terrestrial animals in the coastal zone. a the 1800' s In Rafinesque, early at University, professor Transylvania Lexington, Kentucky, described many fish species of the South (Gowanloch 1933). Wilson and Alexander John J. Audubon described Louisiana birds in the early 1800' s. George E. Beyer published "The

Across this expanse of marsh and swamp looms the New Orleans skyline through Figure 3, the haze, a reminder of the proximity of heavy industries and concentrated populations (Photograph by Charles Sasser).

beneficial effect of the Mississippi River on water and nutrients aquatic understood was generally productivity (Gunter 1938; Viosca 1927; Riley 1937). Also during this decade articles devoted were to marsh plants specifically and Penfound 1936; (Brown published These were soon followed Hathaway 1936). by articles that focused on the relation of environmental factors, particularly to and inundation, plant salinity occurrence (Hathaway and Penfound 1936; Penfound and Hathaway 1938; Brown 1944; Walker 1940).
Since that time the focus of biotic research has shifted to the processes that control the distribution and abundance of of whole to and analyses organisms While this communities and ecosystems. was a national trend, on the Louisiana coast it was seen in a series of studies funded by the Louisiana Sea Grant program in the early 1970' s.

1915

1925

1936

1946

1965

1975

1985

Louisiana oil and 4. Figure gas production (Costanza and Cleveland 1984).

WETLAND EXTENT

DEFINITIONS,

TYPES,

LOCATION,

AND

Avifauna of Louisiana" in 1900, a classic collected A.B. description. Langlois 1,200 plants near Plaquemine in the late and 1800's; Riddill, Hale, Carpenter collaborated between 1839 and 1859 to publish a list of 1,800 names of Louisiana and grasses sedges. plants, excluding Cocks (1907) stated that Langlois' collection was shipped to St. Louis University and that most of the Riddel 1 al et collection was lost. Cocks incorporated their lists into his own list of the flora of the Gulf Biologic Station at Cameron, Louisiana. This station also published studies on oysters pioneering (Kellogg 1905; Cary 1907) and shrimp (Spaulding 1908) during this period.
.

The 1930' s brought a sudden wealth of publications. Noteworthy are a series of the bulletins Louisiana published by Department of Conservation on birds, fur of animals and fishes (La. Dept. Conservation 1931; Gowanloch 1933) that sumnarized the available on knowledge these topics. By the late 1930' s the of life the general history pattern commercially valuable estuarine organisms of the delta had been described, and the

in this marshes considered are classified by Cowardin et al. (1979) as persistent or nonpersistent Most of them lie emergent wetlands. or intertidal estuarine the within of this classification palustrine systems scheme, although some could be construed to be riverine, particularly where the Mississippi and Atchafalaya river flows are not confined by levees. In Louisiana these marshes are further subdivided as or freshwater, intermediate, brackish, based on vegetation associations salt, and Penfound established Hathaway by (1938) and Chabreck (1972), rather than on However, the salinity salinity per se. ranges for these associations have been determined by various investigators (Table 1). They correspond fairly closely with the salinity modifiers - fresh, oligohaand mesosaline line, polysaline - of Cowardin et al (1979) as shown in Table This table also shows the area of 2. each marsh type in the Mississippi Delta

The monograph

region.
In both Figure 5, a map of the delta marshes, and in Table 2 the region is divided into drainage basins, the natural ecosystem units of the delta (Costanza et

Table 1. Salinity values (ppt) recorded Wicker et al. 1982). (from

by various

investigators for

delta

marshes

Investigator

V
STUDY AREA

HYDDOLOGK

UNIT BOuNIIARIE;

fT^no,,:,...

PONTCHARTRAINj
'-^

h^, \

lA ^4;

j^RR^BO^Nrft^^.J

G(/tF
Figure
5.

OF

MEXICO
Plain
showing
the

Map

of

the

Mississippi

River Deltaic

hydrologic

units,

These data and maps are from 1983). recent Fish and Wildlife Service study of the Mississippi Delta Plain Region (Wicker 1980; Wicker et al. 1980a, 1980b). The drainage basins are interdistributary in basins formed shifts the major by of the river. Thus they distributary forni a time series of delta lobes of different ages and allow one to see in space the time sequence of the development and decay of the marshes of a delta lobe.
al.

The active Mississipoi River delta, the Balize Delta, is next youngest. It receives two-thirds of the flow of the River, but it is debouching Mississippi into water at the the deep edge of continental shelf. Most of this basin is fresh also, but there has been marine invasion of abandoned subdelta lobes around the of the main edges distributaries, and the marshes here are brackish.
In succession Barataria, Terrebonne, Vermilion-Cote and the Blanche, Pontchartrain-Lake Borgne basins are of increasing age. They all have extensive marshes with salt and well-developed brackish zones. These six basins together form the Mississippi Delta Plain Region, one of the best-developed deltas in the world. The Mississippi Delta Plain Region is also the largest continuous

the basin is youngest Atchafalaya, which is actively prograding out through the shallow Atchafalaya Bay. It receives one-third of the flow of the combined and Red river Mississippi systems, whose freshwater flows into the shallow the whole basin keep bay fresh or nearly fresh all year. All the marshes in this basin are fresh.

The

wetland systan in the United States with 725,000 ha of marshes, not including the forested wetlands at the inland extremes The delta supports the the basins. of nation's largest fishery, produces more furs than any other area in the United an is and important wintering States, In waterfowl. for iiiigratory ground

addition to these renewable resources the intensive scene of is also the delta mineral extraction; the Mississippi River ports between New Orleans and Baton Rouge handle yreater tonnage than any other port the United States; and dense urban. in and industrial, activity agricultural crowds the distributary levees.

CHAPTER ONE THE REGIONAL SETTING

The
and

unique

characteristics

of

the

its marshes result froin the region forces - the of three interaction the oceanic regime, subtropical climate, the on the river - all and acting physiographic template of the northern

the control The forces coast. gulf that have formed the geomorphic processes also the and delta biological of delta marshes. the characteristics

marsh

plants on the coastal resolve into insolaInsolation, tenperature, and water. tion and temperature determine the potential and the rate, respectively, of biotic Within the constraints set productivity. up by these two parameters water is the major controlling function which makes a wetland wet and determines, directly or It is indirectly, its characteristics. also the most complex of the three parameand Insolation ters. tenperature are determined primarily by latitude, with only minor modification by local circumto the water available stances. But, the depth and duration of floodmarshes, ing, current velocity, and water quality are complex functions of marine energy, fluvial processes, rainfall, and evaporation, operating over an irregular surface.
For individual these forces

earth's surface, depending on latitude. transmission an atmospheric Assuming coefficient of 0.7, Crowe (1971) showed varied insolation how seasonally with In the Mississippi latitude (Figure 5). Delta region, at about 30 north latitude, solar energy reaching the earth's surface varies from about 200 cal/cm^/day during of winter to a peak the nearly 600 During the cal/cm^/day in June and July. summer insolation at this latitude is higher than anywhere else on the globe; it falls off both north toward the Arctic and the toward Therefore, south Equator. midsummer growth potential in terms of solar energy is as high in the Mississippi Delta as it is anywhere on earth.
Cloud cover diminishes the potential irradiance, and on the coast where daytime seabreezes move moisture-laden gulf waters inshore, there are clouds almost every day Consequently the during the hot summer.
CAL'Cm'/ oay

JFMAMJ

eoo 600

400 300
200

THE

CLIMATE,

THE

OCEAN,

AND

THE

RIVER
100

Insolation
J

m|a|m|j|j|a|s|o|n|d
at

no weather is There apparently station in the Mississippi Delta region insolation. records that routinely of this records important Existing parameter are scattered and fragmentary. However, the insolation reaching the top of the atmosphere is a constant that varies seasonally at a particular point on the

Figure 6. insolation

variation of seasonal The The latitudes. various a transmission assumes computation coefficient of 0.7 throughout (Copyright. Reprinted from "Concepts of Climatology," 1971, by P.R. Crowe wi th permi ssion of
Longman Group Ltd., England).

seasonal insolation cjrve for the delta coast is probably skewed to the left with in insolation off peak May, falling in somewhat June and July because of clouds
.

Temperature
one might expect, seasonal air As follow insolation closely. tenperatures Mean monthly temperatures range from a of about 14C to a Decenber/January low midsummer high of about 30C. Temperature at the U, S. Weather Bureau station in New is Orleans 7) fairly (Figure representative of the coast because New and marshes is surrounded Orleans by water. Because of the moderating effect and the of the water bodies high seldom humidities, midday temperatures exceed the low 30's (Celsius) despite the During winter in the high insolation. coastal marshes, freezes are infrequent, and the average number of frost- free days In is about 300. fact, the barrier Grand Isle, was chosen for the island. site of a sugar cane breeding laboratory by the Louisiana State University (LSU) Experiment Station because Agricultural the lack of frost allowed sugar cane fruit Since most of the there. to ripen 1 m deep. less than inshore waters are

90 30-

- 80

UJ

?x z
lU uj tr (T

a <
a.

70

(3

^ < 60 < u. cc UJ w 0. u 2 T 50 UJ
liJ

2 UJ O
03 UJ

t-

20

I-

O a

C o

10

40 30
J
F

M A M

DEFICIT

5
5

E E

LJ

SURPLUS

,.

^"^

spring, fed by melting snow and spring rains in the upper Mississippi watershed River flow can be nearly (Figure 10). independent of local rainfall because of the size of the River Mississippi watershed, but often spring rains along the coast reinforce the river flow.
The older basins of the delta are isolated from direct riverine input by natural and manmade levees. Therefore the rivers debouch the Balize and through units in and Atchafalaya hydrologic extreme floods through the Bonnet Carre control structure into Lake Pontchartrain. Their waters flow on out into the gulf and are carried westward the coast, along freshening the tidal water that moves in and out of the Barataria, Terrebonne, and Vermilion basins. while these Thus, three basins have almost no direct inflow freshwater from local except the salt marshes are never runoff, strongly saline because of the moderated salinities offshore.
In addition to the Mississippi and also Rivers, smaller rivers Atchafalaya feed freshwater into the coastal marshes The Pearl River delivers its (Figure 10). water to the mouth of the Pontchartrain basin, freshening the Lake Borgne marshes and through tidal action the lower Lake Pontchartrain marshes. Other small rivers flow into the northern edge of Lake Pontchartrain. The other basins receive stream flow, negligible however, the interior marshes are maintained as fresh marshes the by precipitation surplus.

">

u
Ill

a o
a.

< B > a uj

PE

'M
'"^^1
,

rn

"1
J

r
A

T~r

MONTH

Figure 9. Average water budget for the upper Barataria basin, 1914-1978 (Sklar 1983). P=precipi tation, PE=potential AE=actual evapotranspiration, evapotranspiration.

Evapotranspiration and rainfall sur The effect of precipitation depends plus. not so much on the absolute amount but on the between rainfall and relationship evaporation from water and plant surfaces. Although apparently no one has recorded evapotranspiration directly in the delta marshes, water balances have been calculated from equations developed by Thornthwaite and Mather (1955), These show that water occur during the surpluses winter months, but the summer during precipitation and evaporation tend to be
fairly closely balanced, with occasional deficits in May through August (Figure 9). Annual rainfall surplus is about 60 cm the northern edge of the delta along marshes (Gagliano et al. 1973), decreasabout 40 cm on the coast. This ing to surplus is important in the total water balance of the marshes that includes riverine inputs and gulf marine water, as will be discussed in the following sections.

Upstream freshwater inflows. The largest source of freshwater to delta marshes is the Mississippi River and its major distributary, the Atchafalaya River. The combined annual flow of these two rivers averages about 15,400 cumecs. The flow is strongly seasonal, peaking in late
11

Marine processes Water fluxes in marshes are driven by the water level differences across the estuary. These change in three time scales: long Since the term, seasonal, and daily. ocean reached its approximate present level about 7,000 years ago, it has been rising relative to the land at a rate measured in centimeters per century. The term "coastal is used to submergence" identify this long-term process, which is due not only to true sea- level rise but also to land subsidence as discussed in the following section on geoiTiorphology.

delta

In the last 20 years the rate of submergence has accelerated. Presently in delta marshes it about a averages

Mississippi

Amife

Tickfaw
Pearl

eo

"^
,

^T^Lwi,

PONTCHARTRAIN.

\^[j^EP[MILIOli_.*

jATCHAFALAyA\

MISSISSIPPI

,'^^v|R DELTA
1

/^

Freshwater inflows to the Mississippi Delta. (Oata from IJSGS 1978). Figure 10. All discharges are for water year 1978 except Mississippi Discharges are in cuinecs. River, which is a long-term mean representing the combined average annual discharge above the confluence of the lower Mississippi (10400 cumecs) and the Atchafalaya (5000 cumecs) Rivers. (mb) during December and centimeter per year (Figure 11a). This is 1,021 millibars double the rate anywhere else along the January and 1,015 mb during early summer have and fall. Several eastern United States coast (Table 3). investigations shown that water level decreases nearly 1 on this long-term trend is a Superimposed increase in barometric for each mb cm variation in mean water level seasonal and Lisitzin Pattullo that itself has an excursion of 20 - 25 pressure (e.g. Thus the expected mean seasonal This bimodal variation (Figure lib) 1951). cm. as a the occurs response to range in water level throughout consistently barometric pressure is approximately 6 cm different salinity zones of the delta, or 25 percent of the total observed range. with peaks in the spring and late sunmer. seasonal In the addition, In the Barataria basin the spring maximum warming increases in an inland direction, that is (expansion) and cooling (contraction) of nearshore waters contribute to a seasonal from salt toward fresh marshes, possibly because of the considerable volume of high in the late summer and a low in January and February. surplus precipitation during this time of the year (Baumann 1980).
The seasonal changes in water level to several attributed are interacting Water level varies inversely factors. with barometric pressure which averages
12 be can events These astronomical modeled and compared to the actual water When this is done (Byrne et al. levels. is a there 1976) always significant

residual which is presumably due to other forces and changes dramatically from year to Dominant among these other year. forces and responsible for the secondary maximum in and the spring following secondary minimun in mid-summer is the seasonally changing, dominant wind regime over the Gulf of Mexico (Chew 1962),. Maximum east and southeast winds in in and fall result an onshore spring transport of water. During winter and in winds summer (southwest westerly summer, northwest in winter) strengthen the Mexican Current and draw a return flow of water from the estuaries (Baumann 1980).

about 30 cm at the coast. Because of the broad, shallow expanse of the coastal estuaries, the tides attenuate in an inland direction. lie shows how the Figure normal tide range decreases from salt to freshwater marshes. In this example tides are still perceptible 50 km inland fran the tidal because of the passes extremely slight slope of the land.
It would be misleading to infer that water levels slavishly follow predictable In daily and seasonal cycles. reality they are modified strongly by stochastic meteorologic events which set up or set down water in the bays and marshes. The

Superimposed level change is

on
a

the seasonal water diurnal tide averaging

effect is clearly shown in Figure lie, where gradually decreasing water levels associated with a "cold front" began on 12 October. Then the water levels suddenly rose on 19-22 October when the wind came around to the south. Typically, "cold fronts" across the coast lower moving water levels dramatically. "Warm fronts" with winds from the southern quadrant set The magnitude up water in the estuaries. of these wind effects is often 40-50 cm, which when combined with astronomic tides can result in water level shifts of over a meter within 12 hours.

Table

3. Average coastal submergence on the U.S. east and gulf coasts (Bruun 1973 compiled by Hicks).

Location

Record yr

Rate

Eastport, Maine Portsmouth, N.H. Woods Hole, Mass. Newport R. I. New London, Conn. New York, N.Y. Sandy Hook, N.J. Baltimore, Md. Washington, D.C. Portsmouth, Va. Charleston, S.C.
,

Airplane Lake
7
krti

Fort Pul aski

'
'

Ga.

(fom GuH
' '

May port, Fla. Miami Beach, Fla.

October

1972

Figure marshes: daily.

11.
a)

Water
long

level

term;

trends in delta b) seasonal; c)

Pensacola, Fla. Eugene Island, La, Galveston, Tex.

These meteorologically driven water changes are common events. Tropical storms are much more unusual When they occur water levels can be dramatically elevated. The water level height/frequency curve for Shell Beach, southeast of New Orleans (Figure 12), shows that wind tides as high as 3.5 m have been recorded, and l.S-m tides occur about once every On a coast with a slope of eight years. about n.2 mm/km a 1976) (Byrne et al 1.5-m tide can cause flooding hundreds of kilometers inland. The ecological effects of such flooding can be dramatic.
level
. .

River brought down by the Mississippi created a major sedimentary basin, and of the subsurface many deposits, especially those that formed in localized centers of deposition, have been prolific hydrocarbon-produci ng reservoi rs
.

GEOLOGICAL PROCESSES
The Mississippi River, the largest river systan in North America, drains an area of 3,344,560 km^ (Coleman 1976). The average discharge of the river at the delta apex is approximately 15,360 cimecs with a maximum and minimim of 57,900 and Sediment 2,830 cijmecs, respectively. discharge is generally about 2.4x10^^ kg The sediments brought down by annually. the river to the delta consist primarily of clay, silt, and sand. The sediments are 70 percent cl ay.

The river has had a pronounced influence on the of the development northern Gulf of Mexico throughout a long In the Tertiary period of geologic time. Period (70 - 1 million years before the present) the large volumes of sediment

In more recent times, geologic changing sea levels associated with the advance and retreat of inland glaciers the Pleistocene Ice have during Ages influenced the strongly sedimentary In order to patterns off the coast. understand the of the development coastal wetlands it is present-day necessary to view the progradation of the delta and its adjacent coastal plains in to several time scales. relationship These scales range from the long periods of geologic time associated with changing sea levels to the changes in the last 100 years in the patterns of minor subdeltas that foniied the most recent deltaic lobe, the Balize Delta. In addition, the heavy sediment load the river deposited by during the last several million years has caused excessive subsidence. This factor has to a the large degree controlled construction rate and the rate of coastal wetland loss throughout much of the recent geologic history.

Pleistocene Sea Levels

During the Pleistocene Epoch, some 2.5 million years long, sea level fluctuated several times. Most authorities agree on at least four major low sea-level stands and four or five high level stands. In addition to these major changes in sea level, numerous more rapid fluctuations took The minor place. in level undoubtedly affected the changes development of the delta marshes, but in the younger Pleistocene deposits it is extremely difficult to document the precise changes. At the lower sea-level stands, the ocean surface was 150 - 200 m below its level. the present During higher stands water surfaces were slightly above or near present sea level. These fluctuations resulted in periodic valley cutting during the low stands and valley filling or terrace formation during the This concept is high sea-level stands. in 13. Fisk's 1944 diagrammed Figure paper should be consulted for details of
1.8
-

<
UJ CO

z <
>
<

CD

100

50

OCCURRENCES/

100

YEARS

Tide levels at Shell Beach, in Figure 12. the Pontchartrain-Lake basin, Borgne associated with nine major storms (Wicker et al. 1982).
14

expanses of coastal wetlands, some 50 - 60 percent larger than present-day wetlands, existed the Louisiana coast. along Borings along the present-day coastline hit and offshore often these buried and freshwater marsh swamp deposits.
of the Late Pleistocene Warming climate returned polar meltwaters to the ocean raised sea and basins, level, the stream decreased progressively gradients and carrying capacities of the As a result, the channels filled rivers. and large expanses of coastal wetlands buried beneath the were present Sedi.nentation could continental shelf. not keep pace with the rising sea level and the rapid subsidence, and a series of deltas were left stranded on the present continental shelf.

Figure 13. advance and exposure and Quarternary

The relationship of glacial retreat to continental shelf sedimentation during the Late (after Fisk 1956).

the relationship of sea level changes to delta and river valley response.
In addition to causing cutting and in sea level valley filling, changes in resulted migration of the site of sediment deposition. During falling sea shifted level, seaward, deposition depositing deltaic sediments at or near the edge of the continental shelf. The progradation of the deltas seaward over thick sequences of shelf clays resulted in of the loading major sedimentary underlying clays, causing rap-id downbowing and subsidence. As sea level began to rise, the delta site shifted landward.

Seismic data and offshore foundation borings have been used to reconstruct the at various times lobes major deltaic during the last major rise of sea level. The positions of these lobes, shown in Figure 15 a through d, illustrate that at different times in the past the area of the coastal wetlands was governed by the locus of deposition of the major deltaic The presence of numerous delta lobe. lobes, now buried beneath the continental shelf deposits, points out the role that submergence plays in controlling the total If submergence area of coastal marshes. did not occur along the Louisiana coast, many of these older deltaic lobes would the be still present-day present, and be much more marshes would coastal extensive.
The latest phase of the Quaternary cycle, characterized by relative stability of climates and relatively small changes in sea level, began approximately 5,000 This sequence involves 5,000 years ago. the modern delta cycles described by Fisk and McFarlan (1955) and Frazier (1967). 16 the illustrates major Figure Mississippi River delta lobes that have Although developed during this period. numerous, slightly differing terminologies have evolved to describe the individual delta and their most systems ages, authorities agree on at least seven delta The result of the building and lobes. subsequent abandonment of the Late Recent delta lobes was construction of a modern
15

The most recent cycle of sea- level and rise to its subsequent lowering present level began about 80,000 years ago This McFarlan 1955). Late (Fisk and Quaternary cycle began in response to Sea level cooling Pleistocene climates. was lowered approximately 150 - 170 m below its present level by withdrawal of water into the expanding Wisconsin-stage Streams along the gulf coast glaciers. extensive and Mississippi River eroded valleys across the shelf and dumped their sediment at or near the present-day shelf The generalized locations of these edge. river channels, now buried beneath the younger deltaic sediments, are shown in this During Figure 14. period large

coastal plain which has a total 28,568 km^ of which 23,900 km^ is exposed above the sea surface (subaerial) (Coleman 1976).

deltaic
area

of

In one of its earlier channels the river built the Sale-Cypremont Delta along of the flanks the western present In Delta Plain. River Mississippi an extensive 1,200 years approximately coastal marshland emerged before the river switched its course to another locus of A the Cocodrie system. deposition, similar sequence of events continued, and with time this site of deposition was abandoned and a new del ta lobe began a This process period of active buildout. each delta completing a continued. has that of requires progradation cycle approximately 1,000 - 1,500 years.

formed the modern birdfoot or Balize Delta The modern delta has nearly (Figure 16). completed its progradation cycle, and in the a new distributary, the recent past Atchafalaya River, began tapping off a River's water portion of the Mississippi A new delta is and sediment discharge. its progradational (Van phase beginning Wells et al. Heerden and Roberts 1980;
1982).
In each phase of the progradational marshes are delta cycle, broad coastal constructed. Scruton (1960) referred to as the constructional this phase. However, once the river begins to abandon its major deposition site, the unconsolidated mass of deltaic sediments is immediately subjected to marine reworking proWaves and coastal cesses and subsidence. currents, and subsidence result in progressive inundation of the marshes, and within a few thousand years the delta lobe

Over approximately the most recent years,

the

last

delta

500 cycle has

Location of major buried river channels fomied during the Wisconsin glacial Figure 14. period (after Fisk 1954).
16

The position of major delta lobes on the gulf coast during the previous Figure 15. 25,000 years. (A) Late Wisconsin, 25,000 - 20,000 yr B. P. (B) Late Wisconsin, 15,000 (C) Early Holocene, 12,000 10,000 yr B. P. (0) Present, 5,000 - 1,000 yr B. yr B. P. SL = relative sea level. P.

sunk beneath the marine waters. Scruton (1960) referred to this stage of the delta as the destructional cycle Thus, in a relatively short period phase. of geologic time both land gain and land loss occur, a function of the stage of the normal delta cycle. The initial phase of delta progradation is characterized by formation of coastal marshes associated with the advancing delta.^ Coastal marshes deteriorate when a delta lobe is abandoned, and a new delta cycle begins elsewhere.
has

that was actively prograding some This delta 3,000 years before present. lobe remained active for approximately coastal 1,200 years, broad, forming a marshland along the eastern deltaic plain.
lobe

Figure 17, a satellite image of the eastern portion of the Mississippi Delta shows in several delta lobes Plain, different of construction and stages The oldest shown on this destruction. image is the St. Bernard Delta, a delta
17

Approximately 1,800 years ago, the Lafourche channel began its progradation. In the St. Bernard Delta, deprived of its sediment marine and load, processes subsidence (primarily compaction) became dominant. The Lafourche distributary increased its sediment yield and gradually within 1,000 years built out a major delta lobe west of the modern or Balize Delta. During this time the St. Bernard Delta continued to be dominated marine by Marine waters processes and subsidence. began to intrude into the formerly freshwater marshes, and marshland deterioration

Deltaic lobes Figure 16. Kolb and Van Lopik 1958).

of

Mississippi

River deltas (modified

from

the increased rapidly. Initially and the interior marshes deteriorated, coastal barrier islands were attached to the ends of the fonner distributaries. Delta Lafourche the system Eventually reached its maximum development and the modern delta lobes (Plaqueinine and Balize) The Lafourche began their progradation. to marine was then Delta subjected and compaction. reworking
the past 800 or so years During subsidence in the St. Bernard Delta has reached a stage in which little or no and the marshes freshwater exist, reworked barrier islands have been separated from the mainland. During this same the Lafourche Delta has lost land, period mainly by saltwater intrusion and opening of the marshland behind a coastal barrier still attached to the fonner distributaries.

favor of the Atchafalaya River course structures at manmade river control limit diversion to not did Simmesport about one-third of the Mississippi River's Even with this limited flow discharge. the modern Atchafalaya River will continue to build its delta onto the continental shelf for the next several hundred years.
in

if

Modern Mississippi Delta

Balize Delta has been The modern constructed during the past 500 years. Because it is relatively young, it offers an opportunity to evaluate the short-term processes responsible for delta building When a break (or deterioration. and crevasse) occurs in the levee of one of river distributaries, water rushing the through the break deposits sediment in the These bay fill deposits adjacent bay. the major coastal marshes of the form subaerial delta. Figure 18 illustrates the bay fill sequences within the modern delta during the past few hundred years. Of the six crevasses shown, four have been
18

Meanwhile, in the modern Balize Delta the river has constructed a major delta The river would abandon this lobe lobe.

dated historically, developnent can be maps.

and much of their traced by historic

of

After an initial break in the levee flood major distributary during stage, flow through the crevasse gradually increases successive floods, through reaches a peak of maximum deposition,
a

As wanes, and is cut off (Coleman 1976). a result of the crevasse compaction, system is inundated by marine waters and reverts to a bay environment, thus comits These pleting sedimentary cycle. crevasse systems are similar to the larger delta lobes but develop faster so that the details of the processes responsible for their fomnation can be adequately evalu-

ated.

Bernard
delta

New

Orleans

Belize delta

Satellite image of the Mississippi Figure 17. different ages (NASA photograph 1973).
19

Delta Region

showing

delta

lobes

of

MODERN

MISSISSIPPI RIVER SUBDELTAS A Dry Cypress Bayou Complex

B

Grand

Liord

Complex

C West Boy Complex

i-_

j^
1

'^i?^

D Cubits Gap Complex E Baplisle Collette Complex ^Garden Island Bay Compli i^f*'''"s*>,j/~F
r,-

^.)i^\?'

^10?/r

In cross section, the prodel ta clays of base the constitute the sequence The lowermost clay marks 19b). (Figure the first introduction of sediment into Above the prodel ta clays are the bay. the coarser-grained silts and sands that These form the delta front environment. sandy deposits are laid down immediately front of the advancing river mouth. in Once active sedimentation ceases in the crevasse system, compaction and retreat For a time marsh growth can dominate. keep pace with compaction, but eventually and the to tend develop, bays large Small retreats shoreline rapidly. near the beaches accumulate major where distributaries coarser-grained is available for sediment reworking. Oyster reefs may find a foothold along the old channel margins of the submerged levee

ridges

u Sect

CSI LSU

Six subdeltas of the modern Figure 18. Mississippi Balize Delta recognized from Dates sediment and analysis. maps indicate year of crevasse opening (Wells et al. 1982).

one of these of Historic maps be used can Cubits crevasses, Gap, to illustrate a cycle of delta building and abandonment. Figure 20 shows the sequential development of the Cubits Gap 1838 map was The crevasse. surveyed prior to the break and shows a narrow, natural levee separating the Mississippi River from the shallow Bay Rondo.
In 1862 a ditch excavated by the daughters of an oyster fishennan na:;ied Cubit to allow passage by shallow draft The boats caused the crevasse break. original ditch was about 120 m wide; the flood of 1862 enlarged the opening, and by 1868 the the break was 740 m wide.

idealized sequence is shown in The crevasse the plan view in Figure 19. initiates as a break in the major distributary levee in the vicinity of point A. During the early formative years coarsegrained sediments are deposited in the With immediate vicinity of the break. bifurcate and time new channels fonn, reunite, forming an intricate pattern of distributaries. Later, some distributarand become inactive. abandoned ies are channel a When pattern systematic front advances develops, the bay fill rapidly into the bay, resulting in the deposition of a sheet of relatively coarse near the sediment locally thickening Seaward of the active channel channels. mouths, fine-grained sediments settle out to referred as in commonly deposits of the Other parts clays. prodel ta crevasse system which have been abandoned or dre deprived of a continuing sediment supply compact rapidly, and many areas to tend open up and revert to shallow marine bays

The

By

1834

the

map

shows

the

initial

of series a of buildout complex distributary channels that had deposited relatively coarse sediment near the break. Note also the shoaling in the bay caused of the deposition subaqueous by The map of 1905 finer-grained deposits. of the that shows major many and that had distributaries developed rapid progradation had taken place in the 11-year period since 1884.

major portion of the crevasse had constructed by 1922; some small bays to were up, open already beginning indicating that some parts of the crevasse system were being deprived of sediments. The 1946 map shows that sedimentation was
A

been

2U

V^-^-riy^^
Bay
(s'

Carlo Secl.i

j;.;:n;;ii;;n;U2Hnin;;::;;;;;:;;:;i;;;;H;H;;;;H;H;;;;;::;

SHALLOW -^ MARINE

Oyster Reefx

Beach
Sea Level

-i:r^^^^Carto

Sect .CSI.LSU-

Figure 19. environments

1964).

Plan view and cross sections through A-A' of deposition in a crevasse (after Coleman

and and

B-B' of Gagliano

21

al.

Figure 20. 1982).

Sequential

development of Cubits Gap subdelta (Wells et

22

primarily taking place at the seaward ends and that distributaries of selected take to was loss marshland beginning
place.
By 1971 a large part of the crevasse marine inundated by being system was was loss and marsh becoming waters, The only deposition was at significant. of the some of ends seaward the distributaries and subaqueously in the bay Note that land loss begins front. fill Here break. crevasse the near first sedimentation is extremely slow, depending only on overbank flooding, whereas higher sedimentation rates are still prevailing near the distal parts of the crevasse the illustrates 21 Figure system. crevasse growth and deterioration.

Figure 22 shows on a single plot the cyclic nature of four of the Mississippi River crevasses; each cycle consisted of deterioration. followed by growth trends of the present-day Projection a life cycle for a crevasse indicates 175 115 lasts that years. system

KO

1800

1820

1840

1860

1880 Date

1900

1920

1940

1960

1980

subaerial 22. curve, growth Composite Figure from land determined subaerial subdeltas. Total internals (Wells et al, 1982).

Mississippi averages at

River 10-yr

"greenhouse" effect that is wanning the earth's surface and melting the polar ice The net affect of both true seacaps. level rise and coastal subsidence has been a change in the coastal submergence rate from about 0.27 cm/yr during 1948 to 1959, to nearly 1.3 on/yr between 1959 and 1971. Although these data are for a gauge at
1.5

Bayou Rigaud in the Barataria basin, the trend is similar along the whole Louisiana coast (Gosselink et al 1979).
.

Years spanned by estimate

1

25Midpoint
1

Reference

01 Adams et at 1976 20ozier 1983 3Gaqliano f. Van Beck 1970

0-5

940

YEAR
The accelerating wetland loss Figure 23. rate in the Mississippi Delta (based on data from Dozier 1983).
24

intertidal In order to remain at elevations marshes must accrete vertically The rapid as rapidly as they are sinking. rate of marsh degradation indicates that they are not doing so, an observation supported by recent research (Delaune et is that the reason One al. 1983). Mississippi River no longer supplies as much sediment to the coast as it has al et Keown (1980) historically. reported that sediment supplies are only about 60 percent of what they used to be, despite the presumed increase in erosion that accompanies forest clearing on the is reduction The watershed. upper presumably due to the construction of dams on the upper reaches of the river and its The dams also remove the tributaries. coarser sediments selectively, so that the sediments reaching the coast are depleted of the sand that is the main foundation This means for delta growth. material that the river can no longer support as
.

Computerized re-creation of the west side of Barataria Bay showing Figure 24. marshes Black is open water; in wetlands between 1945 (a) and 1980 (b). change shown as varying shades of grey (Dozier 1983).

the are

In large a delta as it has historically. addition, channel ing and leveeing the river entrains much of the sediment, preventing spring overbank flooding that nourishes the interdistributary marshes
.

There is now strong evidence that the rate of marsh loss is being accelerated by local human activities in addition to the reduction in the river's sediment load. Canals are the major culprit in this scenario. runoff from Formerly, rain adjacent uplands flowed across wetlands, its load of sediment and dropping Now a network of nourishing the marshes. the marsh-upland drainage canals along interfaces of the delta estuaries carries this runoff directly into estuarine lakes
25

and bays, bypassing the swamps and marshes If runoff flowed (Conner and Day 1982). across the wetlands, the trapped sediment would help minimize wetland subsidence and the quality of the runoff water would be improved before it entered the lakes and of the Instead, the bays. portions estuaries near urban areas are becoming increasingly turbid and eutrophic (Craig et al. 1977).

other end of the estuary, canals, especially those that navigation cross the barrier islands, cause major The canals are disruption of circulation. straight and deep in estuaries that have an average depth of only 1 or 2 m. Therefore from smaller flow they capture
At

the

channels and allow the intrusion of salt Saltwater water deep into the estuary. of fresh and the conversion accelerates intermediate marshes to saline marshes. When increases are sudden, sal t-intolerant vegetation can be killed, and the marsh may erode before other vegetation can be There is also established. some suggestion that the biochemistry of marsh sediments changes with salinity, making more vulnerable to erosion the marsh (Dozier 1983).

Table 4. Land-use changes the along northwest edge of the Barataria basin, on the Bayou Lafourche natural levee (Dozier 1983).

a.

Change in developed land Year Developed land area

(ki)

Rate of increase
(km /yr)

1945 1955 1969 1980

19.27 20.80 39.41 71.69

0.13 1.43 2.93

A network of medium-sized canals that are dredged for access to oil and gas well sites is linking the navigation canals to the inner marsh and to the flood drainage canals. These canals are extensive; their The are canals impacts multiple. themselves act like the navigation canals in combination with them, change and, For circulation patterns extensively. in the Leeville oilfield example, (Terrebonne basin) the density of natural declined as channels channels dredged the flow of water (R. E. Turner, captured LSU Center for Wetland Resources; pers. These canals also allow salt comm.). intrusion. Their spoil banks block the flow of water across marshes, depriving them of sediments and nutrients. This is noticeable where canals especially intersect and their spoil banks interlock to impound or partially impound an area. The effect has not been rigorously quantified, but aerial photographs showing the loss of marsh in these semi-impounded areas are too striking to ignore.

b.

Loss of marsh to indicated category, 1945-80 Area Marsh loss

To canal To development To open water

Analysis of marsh loss rates between 1955 and 1978 (mapped by Wicker 1980) shows a direct linear relationship between canal density and the marsh loss rate The rate of loss (Turner et al . 1982). per unit of canal is higher in recently formed deltas where the sediments are less consolidated than in older deltas (Oeegan et al. It seems to be maximum 1983). where fresh marshes are experiencing salt intrusion (Dozier 1983). Turner et al. (1982) found that the intercept of the regression of marsh loss on canal density (that is where canal density is zero) was always less than 10 percent of the total loss and zero. This usually nearly

Land use changes, Table 5. et al 1980a). (Wicker

,

in

hectares, in

the Mississippi

Delta, 1955-73

Unit

CHAPTER TWO TEMPORAL AND SPATIAL GRADIENTS

marsh is determined constrained the by it on which geologic platform These develops, and by the water regime. create physical gradients that are closely related to variations across the delta in fauna and marsh vegetation, ecological Furthermore, in the Mississippi processes. Delta geologic processes are so rapid that be to the platform cannot be assumed of in the time scale human constant generations
the by regional

IN

DELTA MARSHES

The ecology of biota as

geologic one so that peak productivity occurs the delta lobe's during destructional phase (Figure 25). In order to throw some further this on light interesting hypothesis, t is pertinent to describe the way marshes develop in the context of whole basin systems.
i

As we have seen, a typical delta lobe has a life cycle of about 5,000 years. But the accretionary phase is \/ery rapid. Wells et al. showed subdelta (1982) cycles in the modern birds foot delta of 115 - 175 years. In the Atchafalaya Delta about 20 km^ of new land has appeared And with current subsidence since 1973. of rates about 1 even the cm/yr

destrjctional phase of a delta is rapid; marsh to water is degradation open at a net rate of about 75 km^/yr occurring for the deltaic plain as a whole. As a are not result, the spatial gradients constant but vary with the age of the delta lobe. In this chapter we will consider the and spatial temporal of delta marshes, gradients Mississippi particularly as they control the physical substrate, water and water chemistry, and vegetation.

To do this, have used data from the delta hydrologic units, arranged by age to an instant get snapshot of a basin's This approach is development over time. not The ideal. units are hydrologic nterdistributary, except for the active and thus the active deltas, represent of sedimentation more than one river For exa^nple, the west side distributary. of the Sarataria basin was formed when the the Lafourche distributary was active; east side is strongly influenced by recent River sediments. However, Mississippi biological data have, in general, been collected by hydrologic unit, and a rough six tine of units can be sequence identified, ranging from modern to about 5,000 years old.
! i

first begins to The overwhelmingly riverine. mineral sediment load is high, and water is fresh. As a result, the newly emerged are and sediments the first mineral, marshes to appear are fresh (Figures 25 and 27).
a

When

delta

lobe

form, it

is

TEMPORAL GRADIENTS
and Van Gagliano suggested that the geologic growth, abandonment, and a paralleled by cycle The biotic productivity. Beek (1975) cycle of delta destruction is of biological cycle lags the

As the delta grows, the fresh marshes As described in Chapter 1, the expand. expansion is not uniform; as subdeltas are cut off from stream flow, they become more and tidal more influenced marine by waters. Consequently, sal ini ty increases, and brackish and saline marshes begin to appear.

When the river diverts to another delta site, the periphery of the abandoned
28

BIOLOGICAL PRODUCTIVITY AS A FUNCTION OF THE DELTA CYCLE

SALINITY

Maringouin

/?\ ^ (3) St.

Bernard

W Lafourche
Plaquemines

(D Teche
High

SUBAERIAL

DEVELOPMENT
LENGTH OF LAND WATER INTERFACE
BIOLOGICAL PRODUCTIVITY

Marine

^XD>

z - 2
UJ

(0

= a o
UJ

Low

Subaqueous Growth

Rapid Subaerial

Growth
Time Span

Deterioration

NATURAL ENVIRONMENTS
OPEN BAY

SUBAQUEOUS LEVEES MUDFLATS FRESH MARSH BRACKISH MARSH SALINE MARSH SWAMP
LAKES

OYSTER REEFS MARGINAL BEACHES

BARRIER ISLANDS

Figure 25.

Environmental

succession of an idealized delta cycle (fiagliano and

Van Beek 1975).

delta becomes saline and is modified by marine processes which typically rework the delta edge into a series of barrier reefs and islands that protect the inner Riverine hydraulic energy is estuary. much reduced and sediment loads decline.

marsh turbid

sediments, subject tidal washes, are high in mineral content.

to frequent, always fairly

Further increasingly productivity

is marsh development the controlled by which of the vegetation, This is especially true at forms peat. the landward edge of the basin. Here, too far from the coast to experience much river's the and with tidal activity sediment supply cut off, organic material material produced in situ is the only available for marsh accretion. Thus, as Figure 26 shows, fresh marshes start out as highly mineral, but as the delta lobe Salt ages become increasingly organic.

The general sequence is clear in the figure, but some exceptions deserve comSediment mineral content decreases ment. with distance from the river source (that from fresh toward salt marshes) in is, active deltas (units II and V) but dethe marine creases with distance from sediment source in the abandoned basins. This trend is consistent in all basins. low mineral the to However, compared contents in the recently abandoned basins III and IV, marshes of the older basins I and VI have relatively high mineral conThis probably reflects the centrations. sediment- laden freshwater input continued into these systems.
29

INCREASING AGE

VI

PONCHARTRAIN L BORGNE

VERMILION

100

FRESH
INTERMEDIATE
^""'j BRACKISH
SAl
I

PERCENT
MINERAL

50

CONTENT

Mineral content of marsh soils in Mississippi Figure 26. arranged in order of increasing age (data from Chabreck 1972).

delta

hydro! ogic

units.

Ponchartrain-Lake Borgne basin (Unit I) is fed by a number of small, local streams, by the Pearl River, and periodically by diversion of the MissisCarre the Bonnet River sippi through
The
''-^'-"

Within a hydrologic unit of constant size, wetland area and land:water ratio
a a >
I-

INCREASING

Gg>

The

Vermil ion

<0

O
in

#100(0

X K <
III

(A

II

III

IV

VI

The Pontchartrain-Lake Borgne unit is exceptional in that the mean salinity is high, but so is the proportion of fresh This may be a result of the marshes. The gradient physiography of the system. is compressed into the lower half of the basin by the location of the mouth of the freshwater the Pearl River, primary source, and by the small passes into Lake Pontchartrain which restrain free flow of saline water into the lake.

ATCHAF- MIS8.R. ALAVA DELTA

BARATARIA

TERRE-

BONE

PONTVERCHAR- MILION
TRAIN

Figure percent marshes order of a mean area of

and soil Marsh salinity marsh in Mississippi Delta in by hydrologic unit, arranged Soil salinity is increasing age. for the whole basin weighted by The fresh marsh each marsh zone. is percent of total marsh area (data from Chabreck 1972).
27.

fresh

30

inaxiinuin

increase during active delta growth to a when the distributary is abandoned, and then decrease as marshes subside and The degrade back to open water bodies. of the interface between the marsh length and (the marsh adjoining water bodies in delta lobes is small young edge) because the new marsh is fairly solid. After abandonment, however, the marsh edge increases as marshes open up and more and streams more tidal interfinger through them.
This is reflected in the ratio of marsh edge length to marsh area (m/m^) in different marsh zones. There are no measurements of this ratio available for the delta, but in the neighboring chenier plain's fairly solid fresh and intenriediate marshes the ratio is 15 and 17, As tidal energy increases, respectively. ratio increases to 39 in brackish the marshes and 50 in salt marshes (Gosselink et al. 1979). Applying these ratios to the delta hydrologic units, the mean edge length per unit area of marsh, weighted for the area of different marsh zones in a hydrologic unit, increases with the age of the unit (Figure 28). However, because younger units have more marsh, the total length of the marsh edge (the product of the ratio and the marsh area) is greatest in the recently abandoned Barataria and Terrebonne units (III and IV, Figure 28).

Table 5. (NPP) and inshore shrinp Regression analyses relating net primary production NPP was calcuin hydrologic units to various physical parameters. production (1955-74) each habitat type lated from the mean productivity and area of 19S3). (Costanza et al R is the proportion of the variaShrimp catch is from Barrett and Gillespie (1975). in the independent varifor by variations bility in the dependent variable accounted
.

able.

Independent variabl

(Sasser 1977; Baumann 1980), the land slope The slope of also exceedingly snail. is the water slightly steeper in the Atchafalaya basin because of the enormous river inflow. Generally, across the coast it is so slight that "downhill" changes daily, depending on the astronomical tide stage, wind direction and strength, rainfall, local runoff, and river flow.
is

scale of meters rather than kilometers, a slope also exists on the marsh surface from the edge of tidal streams inland. Water overflowing stream banks on flood tides slows and drops much of its sediment load near the stream edge it moves as inland, creating a slight crest or levee next to the stream. Because of this, water tends to drain away fran streams into small marsh channels that eventually carry the water back The natural through the natural levee. creekbank levee, which is usually measured in centimeters, and the slight marsh surface slope are enough to create a gradient of inundation, water chemistry and biotic These hydraul ical ly mediated activity. gradients dre responsible for much of the observed biotic diversity in the delta marshes.
On
a

smaller

inundation statistics for a number of different species and associations from nearby tide gauge records. Byrne et al. (1976) plotted frequency and duration of flooding at locations in the Barataria basin corresponding to salt, brackish and fresh marshes. They did not measure the elevation of any marshes relative to these Sasser's data. However, by interpolating elevations on the graphs by Byrne et al it is possible to come up with several estimates of marsh inundation (Table 7).
.

F1

ooding

Considering the variability in these that the total estimates, it appears duration of flooding during the year is about constant across the whole marsh from coast to upland. But the regular, daily is of the salt marsh tidal flushing replaced by a more infrequent flooding wind and where tides inland upstream runoff play a much larger role. The delta marshes appear to be flooded about 50 perThe average duration of cent of the time. increases from 12 to 16 hours a flooding at the coast to almost 5 days in fresh the streamside Notice that marshes. marsh, some 10 - 15 an above the inland marsh, is inundated almost as often but for much shorter time periods, so that it flooded only about 12 percent of the is year.

on the Information frequency and of marsh is rather duration flooding Sasser (1977) and Baumann (1980) scarce. measured marsh elevations relative to local mean water levels and calculated

Baumann (1980) showed that inundation are not constant characteristics Flooding throughout the year (Figure 30). frequency does not vary much, but because level varies the water seasonally, the

Table 7. The annual duration and frequency of inundation of marshes in the Barataria indicate the basin, Louisiana. Figures in parentheses percentage of the year inundated.
Marsh zone

Reference

40

80-

Frequency

% Time
60-

/ '*>

40

o o
<

20-

o
<A

r r r 1 r
I I
I

-I

MONTHS
Seasonal salt marsh inundation Figure 30. (Baumann 1980). patterns

5
5

10

15

20

DISTANCE FROM TIDAL STREAM

(m)

water depth over the marsh also varies. a There is sharp peak in duration of flooding in September and October when water levels are highest, During this time the salt marshes are inundated more than 80 percent of the time
Soil s As discussed in the previous section changes in an aging delta lobe, the mineral content of marsh soil is directly related to the hydraulic energy of the In abandoned interdistributary system. means that environments this sediment inland to the marsh decreases delivery frail the coast (Units III, IV, I, and VI in Figure 26) and also into the marsh from the edge of local tidal streams (Figure

Variation in soil density and Figure 31. carbon content with distance inland from the stream edge in a salt marsh in the Barataria basin (Buresh 1978).
soil

3. On

Streamside
Inland
2.5-

2.0-

on

2
a o <

1.5

1.0

31).

0.5-

According to Baumann (1980), most of the sediment is deposited during frequent winter storms and rare summer tropical disturbances, probably by redistribution of sediment from bay bottoms (Figure 32). As expected, the sediment size fraction also varies with the hydraulic energy. There is hardly any sand in delta marshes, but the fraction of clays increases inland
with decreasing hydraulic 1977). (Gosselink et al
.

W SU

SP
F

SEASONS
(1975-1978)

SEASONS
(1975-1979)

energy

Rates of sediment rather well known, both (Cs) profiles and from laid down on the surface time (Hatton 1981, Table

deposition are ^" Cesium marker horizons and tracked over Streams ide 8).
from 34

Sedimentation rates on the Barataria saline marsh. (A) Mean seasonal Mean 1975 78. (B) sedimentation 1975 79. sedimentation seasonal Sedimentation rates were highest during Hurricane Bob the winders of 1975 - 78. Claudette storm and passed tropical through the area during the summer of 1979, resulting in very high desposition rates (Baumann 1980).
Figure 32.

1.4 while about rates cm/yr, average is accretion in inland marshes lower, the Table 9 shows about 0.75 cm/yr. deposition rate of certain soil components Mineral Hatton as (1981). by given deposition, which is directly proportional to bulk density, is much faster in salt Even though the than in fresh marshes. fresh marshes are much more organic than the saline marshes, the rate of deposition of organic carbon is no faster in these It only marshes. appears to be because is not the organic material deposited as much mineral matter. "diluted" by

Salt

One component of the mineral sediment Despite the earlier discussion of discrete marsh vegetation zones, the salt gradient is horizontally stratified. decreases sediment Rather, salinity from the coast inland (Figure gradually There does not seem to be much of a 33). gradient from the edge of a stream into In the marsh interior. many marshes increases elsewhere, salinity actually inland as elevation increases, and the flooded soils in salts infrequently because accumulate evaporation exceeds rain. But in these delta marshes this In fact, impounded marshes does not occur. typically become less saline as surplus rain gradually leaches out the sediment
is

salt.

sal ts.

Table 8. Louisiana

in Accretion rates (mm/yr) delta marshes, based on the 1963 ^''Cs fallout peak (S=streamside, I=inland; Hatton 1981).

Marsh zone

10

20

30

DISTANCE FROM GULF (km)

in free soil The decrease Figure 33. water salinity (mg/g) of chenier plain marshes with distance (km) from the gul f (Rainey 1979).

closely related to surface water salinity since it is a major component of sea water. However most available K and Mg are held in the soil exchange complex. Therefore, available K and Mg are strongly influenced by the adsorptive capacity of the soil mineral component as indicated by their high regression coefficients with bulk density in Table 10. Phosphorus distribution is also strongly related to the mineral component of the soil. The major source of phosphorus to the marsh is probably from mineral sediment deposits.
Neither total nitrogen (N) nor calcium (Ca) (either soluble or exchangeable) are closely related to salinity or to bulk Unlike the other soluble cations, density. Ca is abundant in freshwater, and runoff from the surrounding upland areas into the fresh marsh contains high quantities of Ca. This explains the high Ca/Cl ratios

Soil

Nutrients

The nutrient content of delta marshes well known from a comprehensive taken of surface sediment samples set across the whole coast by R. H. Chabreck, LSU, in 1968 and analyzed by Brupbacker et al. (1973). Rainey (1979) used the same data set to draw a number of conclusions sediment about the factors controlling Because the nutrient concentrations. density of marsh soils varied from 0.05 to in Chabreck's data set, a 20-fold 0.97 Rainey converted all nutrient conrange, to a basis as volumetric centrations Boelter and Blake (1954), recommended by and Mehl ich Clarke and Hannon (1967),
is

quite

Multiple linear regression what factors ions showing Louisiana their distribution in control marshes (Rainey 1979). For each nutrient the model soil factor entering the first value. The total is shown with its R accounted of the variability proportion for when salinity, bulk density and orentered in the model ganic matter are all In general, one factor is also shown. accounts for most of the variability.
Table 10. models of
soil

(1972, 1973).

Soil

nutrient

When analyzed on a volumetric basis (dry mass/volume wet soil), the distribution of nutrients across the marshes falls into a predictable pattern. As one would expect, the soluble ions associated with sea water [sodium {Ha), chloride (CI), (K), magnesium (Mg), and total potassium soluble salts] are closely controlled by the surface water salinity (Table 10). This is also shown in Table 11, which compares the ratio of soluble nutrients to chloride in seawater and in the different marsh zones. Sodium, K, and Mg ratios in the marsh are never more than twice the seawater ratio.

Compared to the soluble ions, some of the total available ions (the soluble plus the exchangeable fractions) behave someTotal available Na is what differently.

Calfound in fresh marshes (Table 11). cium is tightly bound to organic material. (However, on a volumetric basis neither Ca nor organic content shows a wide range of values, and as a result the statistical is not association strong). Nitrogen It is distribution is similarly affected. relatively constant in organic material (C:N = 16.5; Chabreck 1972), and most of the N in the sediment is tied up in organic form.

Vegetation
and I have discussed the physical chemical traits of the vegetation zones in It is time delta marshes in some detail. now to consider the vegetation itself. from classification on a Based early studies by Penfound and Hathaway (1938), classified the and Chabreck surveyed I Louisiana marshes in 1968 and 1978.

Sulfate distribution is interesting because the major source is presumably seawater, but the concentration in marsh sediments is as much as four times that expected from the sulfate:chloride ratio in seawater. However, the biochemistry of sulfur (S) in anaerobic soils is complex; sulfates are reduced to insoluble sulfides that can accumulate in the soil and later be re-oxidized to sulfate.
of the distribution Summarizing, nutrient elements in the delta marsh zones (Figure 34) is understandable in light of the source of each and its soil chemistry. The ions Na , K, and Mg, associated with sea water, decrease from salt to fresh marshes as salinity decreases. Phosphorus also but for a different decreases, reason; it is carried into the marsh with sediment and sedimentation rates decrease Calcium increases inland since it inland. is derived mostly from upland runoff. Nitrogen is fairly constant across the marshes since it is closely associated with organic matter.

.'Aa)

S 2.5V

2-

o
; c z
1-5

u I o

111

'

.5-

S B

SB

S B

S B

SB
P

im
S B
I

Na

MG
Mg,

Ca

Figure
Na,

34.
K,

Ca,

Concentrations of available P, and N in different marsh

.

zones (Rainey 1979)

The ratio of the major cations to the chloride ion in Table 11. nomal seawater and in the saline, brackish, intermediate, and fresh marshes of Louisiana (Rainey 1979).

Cation

Percent cover of the dominant plant species Table 12, marsh zones of the Louisiana coast (Chabreck 1972).
Species

in

major

VEGETATION TYPES:

^1- FRESH MARSHES

INTERMEDIATE

MARSHES
SALINE

BRACKISH MARSHES

MARSHES

NL>N-MARSH AREAS

Vegetation Figure 35. Linscombe 1978).

zones

in

the

Mississippi

River

Delta

marshes

(Chabreck

and

Chabreck': data are for the coastal There is some marshes of the whole state. difference in the species found in the western chenier plain compared to the More impordelta, but these are minor. tant is that the species list is a comNo one posite from many different, sites. site would be expected to contain all the species, especially in the intermediate Each major zone is and fresh marshes. actually a complex mosaic of many subThe primary zones are, as associations. the names the indicate, determined by Within salinity tolerance of the plants. each zone detailed mosaics result from much more complex factors including soil nutrients and elevation (hence flooding frequency and duration).
example, a 90-km^ site in the intermediate marsh in the Barataria basin and from aerial was imagery, mapped intensive ground surveys were conducted. associations were identified Six plant statistical techniques clustering using (Figure 38), and even more complex visual patterns are seen in the aerial imagery.
For

The observed patterns seem to result from the interaction of brackish water entering and east and south, from the the marsh fresh upland runoff from the west, combined with slight elevation differences (Sasser et al . 1982).

Vegetation studies in the Atchafalaya basin fresh marshes show the importance of elevation and exposure to direct river flow versus stagnating backwater flooding in controlling the species distribution (Johnson et al., LSU Center for Wetland Greenhouse Resources; unpublished). studies on salt marsh species from the in the delta clearly show differences ability of different species to tolerate In these 1978). flooding (Parrondo et al studies, although S^. al terniflora and S. cynosuroides appeared to be equally weTl adapted to salt, the latter was far less The tolerant of flooding (Figure 39). studies quantify qualitative greenhouse observations that S^. cynosuroides is found in the locations in slightly elevated marsh.
.

39

S*2J**^^^Ii^'

'^
^.V^.
is A deltaic plain brackish marsh. Note the "hunmocky" appearance wh ich Figure 36. white are of Spartina patens stands. The birds with black-tipped wings typical pelicans, the smaller ones ducks, mostly teal (Photograph by Robert Abernathy)
.

The roles of chance and competition marsh plant distribution have not been extensively studied in the delta marshes. We usually assume that seed sources are abundant so that a supply of propagules does not limit invasion by a species and not of one the presence species does from another species adapted prevent In fact, competition is probably invading. With a very strong distribution factor. the exception of a few true obligate halophytes (represented on the gulf coast by Batis mari tima and several species of Sal icornia ), the salt-tolerant species will in fresh or nearly fresh all grow well Since these species are not substrates. found in salt-free areas, presumably they are confined to saline areas because they marsh with fresh cannot compete well Another species in a fresh environment. example of competition is the observation that the thick layer of dead vegetation covering a stand of the perennial grass ^. and annual excludes olneyi patens S_.
in

It is common to burn S_. patens grasses. stands to encourage these other species which are nore desirable as food for ducks and muskrats (Hoffpauir 1968).
In early literature on delta marsh plants it was assumed that the vegetation modified the landscape so that the environment was changed, allowing other speFor example, Penfound and cies to invade. Hathaway (1938) outlined a successional sequence fran saline through fresh marshes The sequence was based to upland forests. on the idea that marsh plants, by producpeat, could elevate the sites they ing grew on until upland species could invade This idea of autogenic and survive there. succession arose before we understood the rapidity of subsidence on the gulf coast. It is clear now, I think, that most vegetation changes in the delta marshes occur In a because of allogenic processes. sense, the most the biota can do is resist and slow down the inevitable change from

40

K-

A diverse deltaic plain fresh marsh scene. Species are: Sagittaria Figure 37. (foreground), Typha sp. (right edge), mixed grasses and vines, Myrica shrubs (Photograph by Charles Sasser).

in

falcata rear

F.ciof
I

High WIregrass
High Bulllongue

H^H
'

F.clor 2

High Salt Grass and Oystsr Grass Medium Salt Grass and Oyster Grass

HighSpikerush
High'Mixed Fresh'

dor
I

6
I

E::n;;::H

^^H

Medium

Builtongua

High Coast Bacopa

Figure 38. Vegetation zonation in an intermediate marsh transition zone in the Barataria basin (Sasser et al. Factors arise from statistical clustering 1982). techniques and are identified by the dominant species.
41

fresh to saline conditions associated with the processes. geomorphic overriding

Flooded Sediments
[rrprrirrfi

Perhaps one exception to this generalization is the fresh floating marsh. This marsh is a thick (up to 1 m) mat of roots interwoven peat binding decaying into a platform that floats on the water. a It supports diverse flora of emergent species dominated by Panicum hemitomon. The origins of these mats is not known, Russell (1942) suggested that they arise by growing out into lakes from the shoreO'Neil line. (1949) thought that they began as anchored marshes that broke loose from their substrate during a high-water period because of the bouyant force of the The fresh floating marshes are in mat. self-controlled. respects highly many Since they float they are never deeply flooded, but by the same token the water is level always near the marsh surface. The production of organic matter maintains Thus the vagaries of the floating mat. water supply are effectively controlled, the and hydrologic environment of the floating marsh is nearly constant.

SSSSy Drained Sediment

B.

k
c
re

^l

I g
III

k
h
t*

> a. o

^
L3.

R C S ALTERNI FLORA
R- Roots C- Culms P - Plant

P S

CYNOSUROIDES

Effects of substrate drainage Figure 39. conditions on the dry weight accumulation and (B) S^. (A) Spartina alterniflora by Bot. from cynosuroides (reproduced J.G. 1978 by R.T. Parrando, Gazette, Gosselink, and C.S. Hopkinson with permission of The University of Chicago).

42

CHAPTER THREE ECOLOGICAL PROCESSES IN DELTA MARSHES

I In the chapter, previous considered marsh changes across spatial gradients and also those temporal changes that are measured in hundreds or thousands But within of any fairly years. homogeneous patch of marsh, many complex interacting processes occur and reoccur in and in are measured that days cycles In order to understand the seasons. to it is marsh necessary ecosystem, understand how these processes operate and how they interact. However, it is not One can clear how best to study them. the individual components of the analyze to these from and attempt systan Or conversely, it reconstruct the whole. is possible to examine the system from a "macroscopic" point of view, almost as an as an independent organism which acts Both approaches individual. integrated The have their strengths and weaknesses. has been latter approach "systems" emphasized in Mississippi delta marshes in studies supported by the Louisiana Sea Grant program, and I will draw heavily on In addition, much them in this chapter. excellent research has also focused on individual fish, especially species, these Without birds. and mammals, studies it would not have been possible to draw as complete a picture as we now have.

emphasizes the importance of (1) primary and its (2) control, production decomposition, detritus, and the role of micro-organi sns, (3) the benthos, (4) the food chain to vertebrates - fish, waterfowl, and fur animals, and (5) nutrient cycles.

Throughout this discussion the role This hydrology will be emphasized. makes wetlands unique. Nearly property everything that happens in wetlands is influenced by the flooding properties of Some of these the site. flooding dynamics, chemical and physical properties of the substrate, vegetation zones - have In addition, already been considered. each of the five groups of processes emphasized in Figure 40 is influenced by The extent of hydrology's hydrology. influence should become increasingly clear in the following discussion.
of

PRIMARY PRODUCTION
to consider marsh It is convenient plants in four different groups. (1) The most extensively studied are the emergent vascular plants, most of them grasses for most inarsh are which responsible Almost always (2) photosynthesis. associated with the emergent plants on the mud surface, and especially on the lower parts of the vascular plant stems, is an active community of epiphytic filamentous with diatoms and many along algae The benthic (3) microscopic consumers. community in marsh ponds, almost algal surface rich is a submerged, always coating of diatoms and other unicellular (4) Finally, green and blue-green algae. in many marsh ponds submerged macrophytes Eleocharis maritima as such Ruppia
,

the systems approach one often In relies heavily on ecosystem models which conceptually organize and simplify the Although more ecosystem under study. of models quantitative sophisticated, delta marshes have been published (Day al. et 1977; 1973; Hopkinson and Day Costanza et al. 1983), I will use a simple to focus the reader's conceptual model attention on the most important coiTiponents in the marsh and ecosystem. processes Each of these will then be considered This model further. 40) (Figure

43

ADJACENT

WATER

O
and
A conceptual Figure 40, processes.

DEPOSITS IN DEEP SEDIMENTS

model

of a

typical

wetland ecosystem,

showing

major

components

Cha ra parvula spp. are found.

,

vulgaris

and

Potomageto n

Emergent Vascular Plants

The energent vascular plants are by far the most intensively studied of these biomass Much four plant groups. infonnation about delta marsh species has the past decade. been generated during Seven studies of marsh grass productivity covering nine plant species have been performed (Table 13)
.

considerably, but most estimates are very in to studies other high compared localities in the temperate zone. This is because production generally increases with decreasing latitude (Turner 1976).
The seasonal growth of marsh plants Louisiana shows two patterns (Figure is characteristic of annual One 41). and many species witii perennial plants roots that die to the ground every winter. These species have a single, smooth growth in from near zero curve which builds January to a peak sometime between July Each year almost all of and September. the new stems anerge at once when growth In Figure 41 P_. commences in the spring. For austral is this illustrates group. like this, peak biomass represents species net about 40 - 60 percent of annual The rest is accounted for by production. shedding of leaves during the spring and some continued growth into the fall that is masked by mortality after the peak is attained. appears to Sagittaria falcata but same follow the pattern, growth leaves of this actually the individual have a short lifespan and are species replaced constantly throughout the year.
in

most common infonnation related is end-of-season peak production In iiore northerly climates where biomass. are and the all plants growth ceases the killed to the ground winter, during is often an excellent estimate of this
The
to
in net But true the production. subtropical climate of the gulf coast peak biomass has been shown to underestimate production by a factor of 1.6 to over 4, even in those species that have a single growth cycle each year (Hopkinson et al a As result, one must interpret 1978a). Table 13 peak bianass data with caution. estimates shows production vary
.

44

Table 13.
(g dv;/m^

Production of marsh vascular plant species biomass and g dw/mVyr production).

in

the Mississippi Delta

Species

Tabl e 13.

Concluded,

bpecies

Site

Yr

Peak 1 ive biomass

Production Different 3est estimate techniques

2,541 2,487 3,075 3,192 2,861 3,595
a

Ref.

Goose Point 78

2,130

b+
a

3,075

Irish Bayou 78

2,466

b+

3,595

Interiiediate narsh

Phragmites communis Sagi ttaria falcata

Barataria Terrebonne

74-5 74-5

990

648

Terrebonne
Fresh marsh
Sci rpus val idus

74

360

2,364 1,402 2,310 1,113 700 508

1,261 a 1,700 b 1,810 f

2,364

2,310 608
1,261

2 5

Panicum hemitomon

Terrebonne Barataria

74 80

800
1,150

1,700

Techniques a - Smalley 1958 b - Wiegert and Evans 1964 b+- Wiegert and Evans 1954, modified 1980 c - Mortality, Hopkinson et al d - Williams and Murdoch 1972 e - Lomnicki et al . 1968 f - Density and longevity, Sasser
:

Reference
1

2 3

4 5
-

6 7

rby and Gosselink 1976 1980 Hopkinson et al Kaswadji 1982 White et al 1978 Payonk 1975 Cramer and Day 1980 Sasser et al. 1982
Ki
.

et al.

1982

the other extreme, Spartina patens example of a species that grows throughout the year, continuously adding foliage and losing it through death in a Biomass fluctuates kind of steady state. widely around a mean, and there is little For species like if any seasonal pattern. these, peak biomass tells almost nothing about annual production, which is three to alterni flora four times higher. S_. It two extremes. falls between these continues to grow slowly during the winter and always has some green foliage, but is a distinct on this superimposed
At
is

an

o
.J.

a P.
..

hemitomon

STREAMSIDE
INLAND

]-

tllmrntllora

500

seasonal

cycle.

42 contrasts the monthly Figure with growth pattern of S_. al terniflora that of the fresh marsh species Panicum The latter has a broad peak in hemitomon its growth rate during the spring; growth
.

rates of 41. Monthly growth Figure Panicum hemitomon (Sasser et al. 1982) and Spartina alterni flora (Kirby 1971).

46

p. australis

1600

E
X

-8

a
4000
3500
|-

DEAD

3i00r

DEAD

2600
3000

-k.

1974

2500 2000

'-I-.

A972

2100

1975,'

/V'''.'\

1600
1100
1973

1500

I'

1000
J
J J

600
Tim, monthj

Tim, months

Seasonal changes in live and dead biomass of Phragmite s au stral is and Spartina Figure 42. 1975 (Copyright. patens during 1973 Reprinted " from "Aboveground production of seven in Ecology coastal marsh plant species in coastal Louisiana 1978, by C.S. Hopkinson, J.G. Gosselink, and R.T. Parrondo with permission of Ecological Society of America).
,

gradually tapers into the fall with a resurgence after the hottest months, and the plants die to the ground each winter. al terni fl ora maintains active growth S^. throughout the year, with a maximum rate The pattern of during the early summer. streamside and inland plants is similar, but the inland rates are lower.
All the production data reported so far have been for aboveground growth. Root production is difficult to measure because it is difficult to determine, in a all substrate that is root nearly Table material, which roots are living. 14 lists reports of root biomass from a The number of studies in the delta. reported biomass varies widely, partly as a result of differences in techniques. in Fresh and marsh brackish species

established, highly organic marshes have enormous belowground biomass, whereas the same (for species Sagittaria example, spp.. Table 14) in the mineral sediments of the Delta few Atchafalaya produce
roots.

Outside of the del ta, root production measurements have been almost as variable. Good et al, (1982) reported S_. al terni flora root production estimates ranging from 220 to 3500 g/mVyr for tall form (streamside) locations and 420 to G200 g/m^/yr for short form (inland) locations. root:shoot ratios have been conHigh sidered indicative of unfavorable soil conditions requiring greater root surface area to support a unit of aboveground This material (Shaver and Billings 1975). in seems to hold marshes relationship
47

Table 14. Belowground biomass of Mississippi Delta narsh plant species (g dw/m^).
Species

is a there still deal of real good variation in the productivity of a single This species in different environments. is best shown by differences in peak bioinass, which although not equivalent to production are a pretty good index of These differences relative production. are temporal as well as At spatial. Airplane Lake in the Barataria basin, peak biomass has varied by over 300 g/m^ from

year to year (Table 16).

Turner (1979) found a positive relationship between biomass and potential evaporation (which is in turn related to the average air tenperature) during the difgrowing season. By implication, ferences in biomass among years at one location should be related to annual differences in the accumulated potential While this kind of evaporation. relationship has been confirmed for many agricultural crops, it has not been studied in marshes, perhaps because long-term data sets are not available.
variations in biomass have Spatial been the subject of many investigations, both to determine the correlation of biomass with environmental variables and to identify the physiological mechanisms of adaptation to the marsh environment. Figures 43, 44, and 45 show three typical examples of spatial variations in marsh biomass. It is instructive to examine them because they throw light on the physiological responses of plants.
The first of these is the "tidal subsidy", discussed by Oduin and Fanning (1973) as a reason for the high producof coastal marshes. Tides tivity

Table
1

16.

Year-to-year variation

in

peak

ive biomass of Spartina al terniflora at a

singl

Plan Biomass
Plant Heighl

70

50

O
z <

-30

500
5

ID

15

20

10

DISTANCE FROM STREAM (m)

Variation in total aboveground Figure 44. of biomass and Spartina height a1 terni flora with distance inland from the marsh edge in a Barataria basin salt marsh (Buresh 1978).

cell's ready energy triphosphate (ATP).

currency

adenosine

D. spicata

o CO
O o
(A a>

the absence of In oxygen, cell metabolism is incomplete; less energy is from an released equivalent amount of sugar (1 mole of glucose yields 2 moles of ATP under anaerobic conditions compared to 36 moles under aerobic conditions); and organic "waste products" like ethanol and lactic acid accumulate because they cannot be oxidized to carbon dioxide (Figure 47).
In the surrounding root medium, when oxygen is depleted, other materials act as electron acceptors, almost always through some microbial intermediary rather than chemical strictly inorganic through transfonnations. Many ionic species are The reduced form of metallic reduced. ions such as manganese and iron is more soluble than the oxidized form, and the At ions can accumulate to toxic levels. very low reduction potentials, sulfate is reduced to the toxic sulfide. highly Since the substrate is largely organic and micro-organisms are active, organic toxins such as ethylene can also potentially be produced.

3
z g a cc O
CO OQ

<
cc

NaCI (mM/l)

Effects of NaCl concentration Figure 46. of Rb root medium on the rate in the absorption by excised root tissue of S. alterniflora and D. spicata (1 mM Rb; 2 mM Ca; reprinted from Bot. Gazette, 1981, by R.T. Parrando, J.G. Gosselink, and C.S. Hopkinson with permission of The University of Chicago)
.

concentration of other ions such as (Smart and Barko 1978). Finally, the plant leaves have secretory glands called hydathodes which selectively secrete certain ions. All this regulatory activity requires extra energy expenditure by the It is not surprising then that the plant. growth rate decreases as the external salt concentration increases.
the
K

The problem of anoxia is complex because it affects not only the plant Itself but also the microblally mediated biochemical reactions that occur in the soil around the roots. Oxygen is required as an electron acceptor in aerobic cell Its the allows respiration. presence efficient oxidation of organic sugars to carbon dioxide and water to produce high energy-reduced organic compounds and the
51

Marsh plant species have developed a number of adaptations to cope with anoxia, with but even these the are plants stressed effects of sublethal by anaerobiosis (Mendelssohn and McKee 1982). One of the main adaptations of nearly all wetland plant species is the extensive development of aerenchyna tissues in the leaves, stems, and roots, which allow the diffusion of oxygen from aerial plant parts into the roots (Etherington 1975, Teal and Kanwisher There is 1966). that this source is evidence oxygen to the root nomially enough satisfy metabolic requirements of wetland plants, In addi tion, diffusion of oxygen out of the roots can buffer the effect of soil anoxia by creating a thin, oxidized layer in the Mendelssohn and Postek rhizosphere. denonstrated (1982) eloquently through x-ray scanning electron microscopy and microanalysis that the brown precipitate often seen surrounding S_. al terni flora in indeed enriched roots is highly iron (Fe) and manganese oxidized (Mn).

to anoxia

Another adaptation of wetland plants is the evolution of the ability

This "key" Metabolic conversions of pyruvic acid. Figure 47. intemiediate in metabolism can be converted to a variety of end products, depending on the organism and the electron acceptors available (Nester et al. 1973).

anaerobic to aerobic In one study, (fennentation) metabolism. alcohol (ADH) dehydrogenase enzymatic a measure of the cells' ability activity, to convert acetaldehyde to ethanol during alcoholic fermentation, was much higher in reduction sites where the soil inland than in a was intense nearby potential less-reduced streamside marsh (Table 17). Alcohol did not accumulate in inland plant
to

shift

from

tissues in spite of the high ADH activity, indicating that it was able to diffuse out of the roots.
In spite of these adaptations marsh plants in highly reduced environments are stressed, as shown by reduced growth rates, and in severe cases, death. Comparison of streamside to inland sites in the salt of the marsh examples good provides intensity of the stressing agents, their relationship to tidal flooding, and their effects on plant growth. Figure 48 shows schematically a few of the transfonnations that result from tidal action, and their When the tide effects on plant growth. both carries it minerals, rises particulate and dissolved, onto the marsh. Because the water slows as it crosses the natural levee, most of the sediment is deposited close to the stream bank, less same time, At the inland 9). (Table flooding water reduces the diffusion rate The result of oxygen into the marsh soil. is usually anoxic soils, especially where The is concentration high. organic streamside dred is flooded as regularly as

Table

17. Spartina alcohol dehydrogenase adenosine triphosphate

al

terni

fl

ora

root

(ADH) activity, (ATP) and ethanol

a
al

concentrations, and soil Eh in et salt marsh (Mendelssohn

Variable
Unit
Strediaside

Louisiana
.

1982).

Location
Inl

and

ADH

ATP Ethanol
Eh

t 9 nmoles NADH oxi- 36 dized/g fw/hr 218 -23 umoles/g dw 1.17t .07 umoles/g fw

325

^71"

mV

174

t30

-25 248 .08 1.10* ^22 -131

Meantstandard error of mean.

52

WIND AND LUNAR TIDES

0,
^

EXCHANGE,

SOIL

REDOX

POTENTIAL

SEDIMENT AND NUTRIENT

INFLUX

J ^ "nIO"

MICROBIAL

ANAEROBIC METABOLISM

levels of these toxins are potential higher in inland marshes than streamside marshes, increasing the stress on inland plants. to Finally, referring again Figure 48, the direct flushing of marsh soils and the leaching of olant leaves can dilute toxic materials, their reducing activity. Flushing occurs more readily in streamside sites, reducing the potential for accumulation of toxins. With all these potential effects it is not surprising that plant production is higher along streams than inland.

'"flONS

Marsh soil Figure 48. transformations that result from tidal flooding.

inland, but for shorter periods of time (Table 7), and the inland floodwaters are more slowly exchanged. Furthermore, the streamside marshes drain better on falling tides because their sediments are coarser. They also contain more reducible mineral ions to buffer redox changes. All these factors lead to stronger reducing in inland marshes than potentials streamside.

analyses can, at times, mislead. has been found example, it that ammonium in marsh soil interstitial water is more concentrated inland than streamside. This is not expected, considering the higher rates of ammonium production in streamside areas. however, Apparently, the interstitial water concentration is controlled by the rate of plant root uptake. The concentration is maintained at low levels by streamside plants; it accumulates in inland sites because the less robust inland plants are unable to use all the ammonium available to them.
For

Soil

The chemistry of many minerals is influenced the redox strongly by a potential Phosphorus, key plant nutrient, is much more soluble (and hence availabl e to plants) under reduced than oxidized conditions (Delaune et al. 1981).
.

Figure 49 summarizes typical seasonal patterns for various physical and biological processes in marsh soils. Soil water salinity is highest during the summer but probably does not reach levels that ax-& biologically limiting for the euryhaline marsh species. The low winter and early spring salinities correspond with winter rains and low transpiration rates, indicating flushing of the marsh by rainwater.

Inorganic nitrogen, the primary limiting nutrient in marshes, is reduced to the aimonium ion which is readily absorbed by More nutrients are delivered plant roots. to streamside than to inland sites; this should favor streamside plant growth rates. also more Organic nitrogen is mineralized to ammonium in rapidly streamside sites (Brannon 1973).

Other minerals may be transformed to toxins or accumulate in toxic concentrations (for example, sulfide) (Hollis 1967). Toxic byproducts of anaerobic microbial metabolism may accumulate. In general, the
53

Soil-reducing potential (Eh) is least (least negative the anaerobic) during winter, but even during this period it is too low to support any free oxygen. The seasonal Eh curve is the inverse of the tenperature curve - the soil becomes more and more reduced as temperatures rise and increases. biological Soils activity begin to become less anoxic in late summer as temperature drops, even though the marsh is flooded almost all the time Free sulfide follows during these months. the redox curve closely. It is generally Extracthighest when the Eh is lowest. able is an of a manganese example metal ion that is fairly easily reduced. The substrate is always anoxic enough to reduce the manganic ion and the reduced

Free ammoniuin present year round. only forn of inorganic nitrogen to in reduced available these plants In marshes it is soils. streamside naintained at a low level of 1 - 2 pg/ml plant uptake during the spring and by when plant suiii'iier, building up in the fall growth tapers off.
fonii
is

is

the

Epiphytic Algae

Where emergent grasses and algae grow is probably nearly together the grass always the dominant producer. Certainly

develops the largest biomass, but this not a good criterion for comparison because the turnover rate of algae is much In a study in faster than that of grass. which the carbon dioxide uptake of both of these groups was measured simultaneously et al. the (Gosselink algal 1977), community was responsible for only 4-11 percent of the photosynthesis but 61 - 76 percent of the total respiration (Table It has not been possible to separate 18). from the the out respiratory plants activi ty the associated with active consumers - bacteria, fungi, protozoans, and other invertebrates - found in this communi ty
it
is
.

Stowe (1972) found that only along edges of the marsh where adequate light penetrated did photosynthesis exceed He estimated respiration (Figure 50). that net carbon (C) fixation amounted to about 60 g C/m^ annually at the water's The edge, compared to -18 g C/m^ inland. inland was more community consuming Nearly organic carbon than it produced. all of the photosynthetic activity was associated with organisms growing on the base of S_. al terniflora culms rather than on the sediment surface.
the

was Filamentous production algal dominated by the genera Enteromorpha and in the winter and Bostrichia Ectocarpus in The and the summer. Polysiphonia diatom community was also abundant; the cells clustered on the intertidal portion of the culms, decreasing in concentration upward into the drier environment (Figure 51). Although quantitatively the algal rather to be appears community insignificant, the cells are much higher

Table 18. Percentage of marsh community al terniflora metabolism by Spartina (Gosselink et al 1977),
.

December
1975

March 1976

May 1976

Gross photosynthes 89 6 3611 Respiration

Seasonal changes Figure 49. physical, chemical, and biotic a Barataria basin salt marsh.
in

926 365

96+3

249

various factors in

Meanstandard deviation.
54

'^

these are usually considered grazing food chains from phytopl ankters to herbaceous consumers. to larger zooplankton In Terrestrial grasslands are another. are these, the large mammals, grazers that are able to digest the ruminants rather refractory cellulose that is the in material structural plants major harbor tracts their because digestive bacteria and protozoa that can break it down to simpler compounds.

in that terrestrial large counterparts The microbial mammals are not common. equivalents to the digestive microflora of in are bound the the ruminants up decomposing grass and sediment on and in Much research has been devoted the marsh. to elucidation of this pathway of energy I will and in to flow marshes, try the major current ideas about sui'iinarize how it works
.

Marshes grasslands, but

are

called often differ from they

wet their

Before decomposition, considering however, let us pause to consider whether herbivory is really as insignificant as it

Table 19. plant species composition of ponds and Submerged aquatic lakes by marsh zone along the Louisiana coast (August 1968, Chabreck
1971a).

Plant species

The idea is usually considered to be. that herbivory is not important in marshes from our stems qualitative partially that we do not see cows, observations deer, buffalo, and other large grazers in the marsh very often.

Smalley (1950) quantified energy flow the (Orchel imum grasshopper through and concluded that it grazed f idici nium ) less than 10 percent of the net production Parsons al terni flora of its host, S_. Cruz estimated that and de la (1980) in a by grasshoppers consumption Mississippi coast marsh was only about 5.4 have Other investigators g/m^/yr. identified a broad diversity of insects in marshes but little is known about their importance in controlling the flow of
.

indicated (1949) Similarly, O'Neil that dense concentrations of nutria and/or a marsh area. muskrats can "eat out" These mammals are attracted to stands of Sci rpus olneyi , Typha spp . , _P. hemi tomon and other species. They are reported to eat up to one-third of their weight per day (O'Neil 1949) and destroy much more vegetation than they eat.
can be locally Although grazing in marshes, most discussions of important marsh processes ignore it and assume that is as a whole it marsh the over of the The bulk organic negligible. the matter emergent by produced macrophytes dies and falls to the marsh this of The surface. decomposition material can be divided into two phases: an initial rapid loss of easily soluble organic compounds, followed by a longer, slower decomposition rate.

organic energy.

Common invertebrates of the Louisiana have been enumerated (Gosselink et al. 1979), but quantitative studies of productivity and consumption are lacking. Invertebrates other than grasshoppers may ingest significant amounts of live grass tissue, even though this is an accidental For example, the component of their diets. marsh snail (Littorina irrorata ) grazes up and down S_. al terniflora stems, skimming and material dead the off organic It also scrapes off living epiphytes. Alexander grass tissue in this process. (1976) estimated that about 4 percent of the marsh snail's diet is living tissue, which amounts to less than 1 percent of In fresh the production of that plant. marshes insect herbivory is thought to be in than salt more marshes, important because there appear to be more insects in that environment. However, no supporting data are available in the delta.
coast
the delta marshes larger consumers such as snow geese, muskrats, and nutria probably are responsible for more grass consumption or destruction than insects. For example. Smith (1982) reported that snow in Atlantic coast geese grazing marshes can reduce the plant cover by two-thirds where they concentrate and virtually destroy the plants by digging up their roots. This results in significant in changes plant composition the next
In

The first phase takes only about 2 of release The weeks. easily rapid soluble metabolites from the grass tissue and the continuous leaching of organic live grass from the (Turner compounds flow of 1978) represent a significant - 30 organic energy, perhaps as much as 20 of aboveground primary production percent The fate of this material (Teal 1983). in coast been studied has not gulf marshes, but a number of investigations in were conducted Georgia (Pomeroy and of the There, much 1981). Wiegert in the dissolved organic carbon (DOC) column is water refractory, probably released from later stages of decay of the is likely that the It marsh detritus. soluble compounds released when readily the grass cells die are easily metabol i zed by micro-organisms and disappear rapidly from the water column.
a recent review article Ducklow assembled evidence that bacterial production in the ocean is not only high but is also a significant food supply for and ciliates. planktonic zooflagellates Most of these bacteria are apparently using DOC as an energy source since they with are not associated particulate We need to know much more about matter. this pathway of energy flow in coastal If Ducklow' s model for the ocean marshes. and continental shelf is any guide, the food chain from grass to DOC to bacteria

In

(1983)

year.
57

to iiiicrozooplankton and

filter
may

eventually to such feeders as mollusks and menhaden be more significant than has been

real ized.

The second phase of decomposition often takes a year or more, depending on the environment and the plant species At the end of this et al. 1982). (Valiela period about 10 percent of the original detrital biomass may remain as refractory organic compounds.

50

100

150

200

250
20)
2/25

DAYS FROM START OF INCUBATION {JUNE

common way to investigate the loss by enclosing dead plant material in litter bags (small nylon mesh bags with 2 to 5-mm holes), suspending the bags in at them and the marsh, retrieving amount of examine the to intervals material remaining. Decomposition is not the only thing measured by this technique. As soon as the plant fragments become small enough to escape from the bags, they nay be lost by the flushing action of In addition, usually flooding water. larvae of many invertebrates find their way into the bags and prosper on the Their action in fragmenting the detritus. detritus is undoubtedly important in the
^

7/10

8/7

9/15 10/15 11/7

12/6

1/18

rates

is

DATE

patens Disappearance of Figure 52. S_. in from litter litter the bags basin from (data Pontchartrain-Borgne Cramer and Day 1980)
.

loss rate.
A number of decomposition studies carried out in the delta are summarized in In this Appendix and the figAppendix 2. ures and tables that fol low, decotnposition rates have been standardized by assuming an exponential decay rate (Wiegert and The data are reported as Evans 1964). loss rates, _r [mg dry weight (dw) lost/g dw detritus/day], defined as [ln(initial mass/final mass)]/time interval.

these factors are probably reThe histogram this graph. showing the changing rate for each successive interval of time indicates that the initial rapid rate was declining as early as August before air tenperature dropped This implies a change in significantly. On the kind of material being decomposed. the other hand, the rate began to increase again at the end of the experiment when be most the remaining materials would refractory; this coincided with the early spring increase in the ambient temperatures.
Both of flected
in

These studies support results found elsewhere: the three main factors controlling decomposition are tenperature, location in the intertidal zone, and the plant Nutrient levels and the presence species. that shred the of macro-invertebrates detritus are also important.
Figure 52 shows that the decomposition rate of S^. patens detritus decreases This could happen for two with time. reasons, "^irst, this study was initiated in June, and the rate declined as the air Second, one would temperature declined. the more easily decomposed material expect to the more first, disappear leaving refractory, slowly decomposing compounds.
58

Figure 53 shows mean loss rates of detritus from litterbags terniflora S_. but susoended off the bottom in submerged a tidal stream, on the surface of a streamside marsh, and on the marsh surface further inland. Decomposition was fastest tidal where second in water, flowing was vigorous, and slowest where flushing the bags tended to be submerged most of The figure the time in stagnant water. also demonstrates the temperature (seasonal) effect.
al

the 20 summarizes Table Finally, the decomposition of species-dependency rate. Variability is high, hut I believe the means are fairly reliable indicators of the relative rates of decomposition of different species. S^. al terniflora is the most easily broken down of the grasses, but they all tend to be fairly fibrous and

to filainentous yeasts fungi in different predictably depending on their history.

changed wetlands

packaging, and inoculating the with bacteria in fecal pellets.

It

detritus

Mixed with these decomposers on the soil surface is an active community of autotrophic algae, chiefly diatoms, that enter the food web at the same level as the decomposers and may be an important additional source. Most energy investigators, however, are concerned more with the biochemical activity mediated by with microbiota than the species identification. They are satisfied to get some relative index of microbial bioTiass like that afforded by total ATP activity, or to characterize the microbiota by their chemical activity (White et al. 1979).
of The decomposition underground biomass has been studied very little. No studies are available from the Louisiana The best infonnation on delta marshes. the subject comes from studies in Atlantic coast salt marshes summarized by Valiela et al. (1982), Teal (1983), and Howarth and Hobbie (1982).

appears now that bacterial density low on most detrital material to provide a sufficient food source for the macro-benthos (Wiebe and Pomeroy 1972). This change in viewpoint is reflected in The the trophic diagram of Figure 54. meiofauna are seen to have a crucial role in energy transfer (1 in Figure 54). They are from macrofauna distinguished Both are found in or primarily by size. on the substrate during all or part of are life Meiobenthos their cycles. generally microscopic; macrobenthos are larger and include such taxonomic groups as snails, mussels, and crabs.
is

too

Sikora

et

al

(1977)

found

that

Since the soil environment is anoxic, most of the decomposition must be anaeroThe leaching phase of decomposition bic. is the same as aboveground, but subseof the disappearance organic quently material is slower. Nitrogen stimulates the decomposition rate, indicating that it is limiting belowground as well as in an One reason is that aerobic environment. nitrate may control the metabolic rate by as the an electron acceptor in acting absence of oxygen. Most underground production, however, is decomposed through reduction the fennentation and sulfate pathways (Howarth and Teal 1979).

meiobenthic nematodes account for 70 - 90 percent of ttie sediment ATP, indicating that nearly all living biomass in anoxic is not marsh sedi;nents meiofaunal, These organisms are thought bacterial. to be siiial enough to graze the bacteria efficiently and "package" that organic energy supply in bite-sized portions for macrobenthic deposit slightly larger
1

feeders (3 in Figure 54).

Sikora (1977) showed that the chelae of the grass shrimp (Pal eomonetes spp.) are about to the size right capture nematodes and speculated that grass shrimp are more likely to use this food than

detritus.
this

CONSUMERS
Benthos
energy transfer it is microflora act as the between the intermediary organic production of the higher plants and the At first levels. higher trophic that the macroscopic investigators thought feeders were deposit ingesting bacteria-laden the detritus; skimming from bacteria and it; fragmenting.
In

terms

of

assumed

that' the

study (1980) supports found that meiobenthic polychaete and copepod densities increased in caged exclosures that reduced macrofaunal Gut seldom predation. analyses turn up nematodes, the dominant meiofaunal taxon, but this is probably because their soft bodies are dissolved rapidly. Macrobenthic deposit feeders are thus ingesting and using as an energy source meiofauna, which in turn have been cropping bacteria. The deposit feeders themselves are prey for the many small fish, shellfish, and birds that use the marsh, marsh creeks, and small marsh ponds (3 and 4, Figure 54). Although apparently each step in this can be transfer quite energy efficient - net growth efficiencies up to 50 percent for bacteria (Payne 1970), 38 and for nematodes (Marchant percent Nicholas 1974) - the trophic pathway from to meiofauna to microbes to detritus
idea.

Bell's
She

60

Bacteria
Fungi

Protozoa

Nematodes Turbellarians Gastrotrichs Polychaete larvae Harpacticoid copepods Ostracods

f
I

Detmus

^F~)
j'-f
I

Microbes

*
J
1

Clapper rail Sora Belted kingfisher Fish crow Black duck Least bittern Northern shoveler Hooded merganser American avocet Western sandpiper Solitary sandpiper Wil son' s phalarope Common snipe
Dunl in

Macro Denthos
-

MetobenTfTos

Polychaetes Amphipods 01 igochaetes Tenaiads Isopods Melampus sp. Caridean shrimp Fiddler crabs Small blue crabs Littorina snails Neritina snails Carol ina marsh clam
Speckled trout Gizzard shad Hogchoker Pinfish (juvenile)
Spot Tidewater silverside Atlantic croaker

Piping plover
Kill deer

\/^^

Penaeid shrimp
Blue crab
Sea catfish Blue catfish Channel catfish Largemouth bass Black drum Red drum Striped mullet Silver perch Spotted gar Alligator gar Yellow bass

American alligator Snapping turtle Mississippi mud turtle Red-eared turtle Graham's water snake Western ribbon snake

Major pathways of organic energy Figure 54. marsh and associated water bodies.

and
as

migrating species that use the marsh for a Many of nursery. juveniles and feeders are benthic these species represent the next link in the benthic in the described chain food previous section.

of what comprises a marsh definitions Nevertheless, much evidence points pond. to heavy use of the marsh by nekton for Ruebsamen (1972) both food and shelter. fish in the the small found only As they grew they intertidal marsh ponds. from the samples. disappeared usually

Ruebsamen (1972) studied the stomach of fish captured by seine in contents small, shallow intertidal marsh ponds in Of the the Barataria basin (Table 21). were abundant most nine species, six described as feeding on benthic infauna such as copepods, amphipods, ostracods, tendipedid polychaetes, mysidaceans, worms. and annelid nematodes, larvae, Two were described as detritus eaters, (which probably means that they were using The small the meiofauna in the sediment). marsh ponds are frequented primarily by resident fish, while migratory fish are In creeks. in the deeper marsh found of small marsh ponds, Ruebsamen' s study Leiostomus xan thurus was the only spot migratory species found in large numbers.
(
)

found 20 to Hinchee (1977) the of menhaden edges along as Ponchartrain, they apparently into the estuary from the gulf. small juveniles moved into the marsh

25-mm

Lake moved These where they reached about 50 they stayed until which their after mm, they began emigration back out through the lake to the open gulf (Figure 55).

When conditions permit, many nektonic organisms move up into the marsh itself. Sikora (1977) found this true for the grass shrimp in Georgia, and Wernie (1981) the silverside of 30 found percent (Menidia menidia ) and mummichog ( Fundulus heterocl itus ) in a north Atlantic estuary up in the marsh at high tide.
(1965) sampled fish in marsh In this active Balize Delta. ponds nearly freshwater area he found mullet and blue catfish the most abundant, but he also croaker, spot, plentiful reported and sand seatrout, seatrout, spotted that is It menhaden. interesting

Variation in the particular species is often to use marsh ponds reported related to differences in gear used and

Kelley

in the

and Table 21. occurrence Monthly abundance of the fish species collected in small salt marsh ponds (Ruebsamen 1972).
Species

freshwater coastal marsh/ aquatic systems represented by the Balize and Atchafalaya Deltas are found to function in very much the same way as saline estuaries, with the same suite of marine/estuarine fish and In addi tion, freshwater species shellfish. like gars (Lepisosteus spp.), gizzard shad Dorosoma cepedianum ), and blue catfish Ictalurus furcatus are common (Kelley 1965, Thompson and Deegan 1983).
)

Even when they are seldom found up in the marsh itself or in the small marsh ponds, other species concentrate along the marsh edges where food is abundant and shelter is available in the streamside For example, Peterson (LSU; grass stans pers. comm.) was unsuccessful in capturing larval spotted sea trout until he began to seine along the very edge of marshes as to more compared open aquatic environments. Spotted sea trout are just one example of the concentration of both the food supply and the aquatic organisms that depend on it.

Biological activity is concentrated the marsh For edge (Figure 56). reasons already discussed, pi ant production is highest along the marsh edge. Finely detritus from the decomposed previous year's plant crop is flushed from the marsh during the winter and accumulates the marsh edge in deep deposits along known to local as "coffee shrimpers Nematode numbers are highest grounds." here as are the concentrations of small It is no wonder that deposit feeders. larger invertebrates - shrimp and crabs and larval and juvenile fish are also attracted to this feast. Virtually every kind of organism enumerated has been found to concentrate along marsh edges.
at

56. of Figure Density vegetation detritus and consumers at the edge of the salt marsh.
,

The importance of this energy flow in marshes can be seen pathway qualitatively by comparing the list of nektonic species in Figure 54 that use the benthic pathway predominantly with those that use the planktonic pathway. Of the abundant species only the gulf menhaden, the bay anchovy, and the juvenile Atlantic croaker are filter feeders. Crabs, shrimp, drum, gar, mullet and nearly all the small resident marsh fish are benthic feeders.

This benthic food is the pyramid dominant one in salt marshes. Meiofauna, particularly nematodes, graze the bacteria on decomposing grass, are ingested in turn by deposit feeders which are a major source of food to nektonic fish, shellfish and birds. The marsh-dependent fish, the \/ery small ones, graze and especially shelter up in the marsh when it is flooded and lie in the small marsh ponds and along the edges of fine feeder creeks at other times. As they grow they frequent deeper, more open water.

Wildlife
Wildlife species that use Mississippi delta marshes are abundant. Table 22 summarizes the of different species taxonomic groups that are likely to be found in different marsh zones in the chenier of southwestern plain region Louisiana The deltaic plain has about the same In species. general, species richness is highest in the fresh marsh, No amphibidecreasing into saline areas. ans and only 4 reptile species are found
63

richness Wildlife 22. Table species (number of species) in the chenier plain marshes (Gosselink et al 1979).
.

Wildlife

Crawfish, and in brackish areas blue but crabs, are major alligator foods, alligators are also reported to eat birds, fiddler crabs, fish, insects, nuskrats, and nutria, turtles, snails, shrimp, In the Florida grasses (Chabreck 1971b). Everglades they make "wallows" that are ecologically important for fish during the dry season, but this has not been reported in delta marshes.
Muskrat and nutria The muskrat (Ondatra zibethicus and the ..._ nutria , (Myocastor coypus ), both herbivores, are the dominant mammals in the delta marshes. The nutria is an introduced species. It is debatable whether muskrats are native or not. O'Neil stated that (1949) although early surveyors' records provide an unconfirmed record of high density muskrat in the Baratariapopulations Lafitte area in 1840, fur harvesting did not begin until the first years of the twentieth century, and old-time trappers all claimed that no "rats" were seen much prior to that time. However, Arthur (1931) , in a Louisiana Department of Conservation from the of Bulletin, quotes journal Father Jacques Gravier describing travels down the Mississippi River. He described the of the Tunica dress Indians in a November, 1700 entry:
. )

fresh (Figure 57), the nutria prefers marsh and swamp forests and often ventures into nearby ricefields to feed. There is some evidence that the 1974) (Lowery present muskrat distribution results from the invasion of fresh marshes by the more robust nutria which displace muskrats into less desirable brackish areas. Although both species often exist side-by-side in the same area, they appear to have very much the same food habits, and it has been noted that when nutria are heavily trapped, the muskrat population can soar (Evans 1970).

Muskrats often seem to be the primary agents in a 10- to 14-year cycle of marsh growth and collapse (Figure 58). They

MUSKRAT
< o o o
700-

TO 6,478

600500400-

o 3 Q O
oc Q.

300-

200100-

"Most of the men have long hair and have no dress but a wretched deerskin. Sometimes they, as well as the women, also have mantels of turkey feathers or muskrat skins well woven and worked."

UJ a.

0-

1920

1945

If water levels are low food, crashes. for a year or two to allow regrowth of the vegetation, the marsh may recover (and the muskrat population with it), but often the damage extends so deeply into the marsh that recovery is poor at best. Severe storms may reset this cycle by destroying nests and burrows and the drowning predatory disease organisms they harbor. The muskrat population often comes back after these storms (O'Neill strongly 1949).

Annual muskrat harvest from a Figure 58. 52,200-hd brackish Scirpus ojneyi marsh in the Delta (O'Neil 1949). Mississippi

kill much for the vegetation digging In roots. addition, their preferred house-building activity, underground runs, and surface trails (Figure 59) destroy much more marsh than is directly eaten. For example, in a ID-ha brackish marsh area that contained 24 active and 30 in inactive houses 31 new 19S2, April houses were built and 10 "refurbished" during the next year (Table 23). Sixty of the active houses and 57 percent of the inactive ones percent simply disappeared.

It is interesting that "eat-outs" are seldan found outside of brackish marshes and are always attributed to muskrats, not nutria (O'Neil 1949). The nutria has a much longer gestation period (130 days compared to 28 days for the muskrat) so its that for to response potential environmental change is much slower than the muskrat' s. its Consequently, is more stable. Muskrat population in "eat-outs" fresh marshes have been recorded (O'Neil 1949) but the preference for brackish marsh makes this a more "Eat-outs" are much rarer likely site. today than in the 20's and 30's because the trapping keeps population down to nondamaging levels.
In of the local light apparent importance of plant-eating furbearers and the relative the earlier discussion of in lack of is marshes, it herbivory informative to reconsider the importance

When muskrat populations are dense, all this activity can decimate a marsh, creating large "eat-outs" especially in the favored brackish marsh three-corner (Sci rpus olneyi ) 60). (Figure grass Subsequently the local popul ation, with no

Table 23. Muskrat house-building activity in 10-ha brackish and salt marsh areas in basin Barataria (Sasser et al. 1982).
status

Ground of a Figure 59. plan typical muskrat house with underground runways and surface trails (barred lines) (Arthur
1931).

A muskrat "eat-out" in the brackish marsh in the Barataria Figure 60. high density of muskrat houses (Photograph by Robert Abernathy).

basin.

Note

the

Huskrats are reported to herbivory. one-third of their weight per day (O'Neil 1949), and a nutria consumes 1.5 2 kg of vegetation each day (Lowery 1974). The average population of nutrias and Chien Wildlife muskrats from Point au Area in the delta, from 1973 to Management was 1.2 and 0.8/ha, respectively 1981, (from Sasser et al. 1982, assuming the population is double the catch (O'Neil
of

eat

building nests, and digging for roots was 10 times greater than ingestion, it is easy to see that a significant portion of the vegetation would be destroyed.
of one-third Deer Although Louisiana's white-tailed deer (Odocoileus to population is reported virqinianus marshes coastal in the live (which comprise only 13 percent of the state) (St. Amant 1959), very few studies have been made of their feeding and habitat environment. this in requirements are preferred Apparently, fresh marshes almost to the exclusion of brackish and sal ine marshes
. )
.

1949).

0.3

nutria eats 2 kg/day, a muskrat muskrat weighs about a (a kg/day 20 is and the vegetation kilogram), percent dry weight, then their combined intake is about 150 kg/ha/yr, compared to about 30,000 of a productivity plant Direct grazing is thus less kg/ha/yr. O'Neil than 1 percent of production. (1949) reported a peak harvest of 46 muskrats/ha in a brackish marsh (Figure With the same assumptions, that many 59). animals would eat as much as 7 percent of If damage from burrowing. the vegetation.
If
a

Based upon data gathered over 20 Wildlife and Kidd B. (La. years, J. Fisheries Commission), in a 1972 letter (as reported in Self 1975), estimated that the "potential" density of deer by marsh type was one deer per 12 ha in the fresh marsh, 1 per 330 ha in the brackish marsh, This and 1 per 2900 ha in the salt marsh.
67

assessment of carrying capacity for fresh marsh agrees well with observations by Jessie Fontenot (Morgan City, La., 1983; pers. comm.) about the deer density in his 1600-ha hunting lease in a fresh marsh in unit. He the hydrologic Atchafalaya reported 180 deer (about one per 9 ha) on his lease, which he said was overstocked.
areas deer White-tailed prefer elevated above the marsh such as slightly natural levees and spoil banks which can be used for travel, bedding, and fawning. From a browse study made on spoil levees in the fresh marsh in the Rockefeller Wildlife Refuge in the chenier plain of Louisiana, and from rumen analyses of deer Self in that killed (1975) area. determined that deer ate nearly any plants that were succulent and green.

Table

of waterfowl Density zone in marsh the ha) (nuiiiber/100 by Barataria basin in 1980-31 (total for 13 fl ights; Sasser et al. 1982).
24.

Species^

were

Important food plants during the fall Al ternanthera philoxeri odes Bacopa hal imifol ia Vigna luteol a, Sal i x nig ra, wal teri , Echinochloa monnieri B_. vi rginica Leptochloa Kosteletzkya Panic um dicotomiflorum , and fascicul aris During the spring and Paspalum vaginatum summer the same species and Phragmi tes and austral is . Iva annua, Cyperus vi rens All browsed. angustifol ia were Typh a and in fresh these species are found The brackish marsh intermediate marshes. in was grazed paten s Spartina grass to its abundance but was not a proportion preferred species.
,
,

Waterfowl, coots, and wading birds Functionally, birds that use Louisiana's delta marshes can be divided into dabbling or puddle ducks and coots, diving ducks, geese, wading birds, birds of prey, and The birds other marsh 4). (Appendix waterfowl and coots are by far the most winter are abundant. They mostly residents that migrate as far north as the Arctic Circle each summer. Of this group, only the mottled duck breeds in Louisiana Duck with marshes any regularity. in are variable populations highly censuses because of their mobility, but peak populations in the deltaic plain are Table 24 usually over 2 million birds. common of the most shows the density transects through Barataria species along Gadwall basin. strepera ) (Anas and mallard blue-winged teal (A. discors ) platyrhynchosT were the most common (A_.
. , ,

from a few centimeters to one-half meter deep. are They primarily herbivores, and good stands of submerged grasses improve the quality of habitat. maritima the Ruppia (widgeongrass) is the preferred food in brackish Potamogeton pu s i 1 1 u s ponds; Najas quadalupensis (naiad), (pondweed), in freshwater and Lemna spp. (duckweed) In brackish marshes Scirpus olneyi ponds. (three-cornered grass), Bacopa monnieri Eleochari s parvul a (water hyssop), and are desirable foods. (dwarf spikerush) Echinochloa walteri (wild millet), fascicularis Leptochloa (sprangletop) Panicum sp and other (fal 1 panicum) annuals that produce abundant seeds are The succulent good fresh marsh foods.
(less

than 5 ha)

about

duck foods, the saline marshes are tively poor puddle duck habitat.

rela-

roots and tubers of species such as S^. and Sagittaria platyphyl a (delta olneyi duck potato) are also favorite foods, for geese. especially
1

It

is

easy

to

see

why

fresh

and

brackish marshes in the delta support so There are thousands many dabbling ducks. of small marsh ponds in all salinity zones (Table 25), and the dominant plant species in brackish to fresh ponds are considered excellent duck food. Ponds 0.4 - 4 ha in size have the best growth of submerged because wind-induced grasses, possibly turbulence is low in these small ponds. Saline ponds are poorly vegetated (Table Because of this and because the 26). plant species of this marsh zone make poor

Much attention has been focused on habitat conditions of arctic and subarctic nesting grounds and their influence on the growth of duck populations. Much less attention has been directed toward the importance of wintering grounds for reproductive success. A recent study and Fredrickson by Heitneyer (1931), however, emphasized this important aspect of wintering grounds. They found a direct linear relationshio between winter precipitation in the Mississippi delta riparian hardwoods (an index of pond number and hence habitat quality) and reproductive success of mallards as measured by the ratio of young to mature mallards. In their multiple regression models both the wintering ground quality index and the numbers of ponds in the nesting area in May and June were significantly positively related to mallard age ratios. The study that the quality of deltaic plain implies marshes may also be important in duck reproductive success.
the
In contrast to puddle ducks, diving ducks usually prefer deep water. They are carnivores, diving to depths of over 10 meters in some cases to obtain their food. Because of this preference they are usualfound in open water and ly along the nearshore zone. However, they are also known to feed on the vegetation of shallow

Table 25. Density of ponds and lakes of various size classes in marsh zones along the Louisiana coast in August, 1968 (Chabreck
1971a).
Pond and lake size class

Salt
(acres) 0.01

Marsh zone Brackish Intermediate

(number per 100,000 acres)

Fresh

0.01-0.10 0.10-1.0 1.0-10 10-80 80-640 640-3,200 3,200-16,000 16,000-32,000 64,000

27,700.2 16,749.0 4,702.6 700.0

32.2

118,841.7 62,162.2 14,139.0 1,376.1

55,952.2 45,024.0 10,432.8

759.1

59,181.2 47,637.4 9,796.8 1,070.5

The percent of the area of ponds and lakes covered with Table 25. submerged vegetation in August, 1968 (Chabreck 1971a).
Pond and lake size

tailed grackle (Quiscalus mexicanus ) and blackbird the ( Agel aius red-winged phoeni ceus ) are the most numerous of the The other marsh birds. latter two are abundant during species, especially, the spring season. breeding They are and are absent the during migratory winter. Northern harriers are also seen

number of consumer groups. However, I consider the overall input-output budget without this detail. Unfortunately, several key flows in the budget are still not quantified. As a result, any carbon balance inust be considered tentative even today.
a

will

frequently

in

all

marsh

environments.

Some of these species are endangered or rare (Table 28). The beautiful brown pelican, in particular, has been almost lost from the delta (King et al 1977). It has been reintroduced from Florida and is found in two nesting colonies on mangroves on Queen Bess Island in Barataria Island just west of the Bay and North Chandeleur Island chain.
.

Day et al (1973) published the first It was budget for a delta salt marsh. based almost on entirely aboveground benthic primary production, commmunity respiration, and calculated energy flow Loss to through the abundant consumers. deep sediments was assumed to come from root production, and both were ignored in the balance. These authors concluded that 50 percent of net production was exported from the marsh. It has not been possible to measure this organic export directly.
.

Carbon Budget One v/ay of summarizing quantitatively the productivity and trophic relations discussed is with a C budget. Most C budgets are primarily input-output budgets that treat the ecosystem under study as a black box so that internal details of the trophic structure are ignored, and metabolism of all consumers is lumped as commuIn particular, higher nity respiration. consumers contribute little to community Both respiration and are usually ignored. and Costanza et al . (1973) Day et al. (1983) are exceptions to this generalization; they calculated metabolic rates for

calculated the Happ et al (1977) export of total organic carbon (TOC) from the Barataria estuary to the nearshore gulf from the gradient of decreasing TOC across the passes and an estimate of the turnover rate of water. bay They estimated that the export of TOC was about Since 150 g/m^/yr. aquatic primary production and community respiration in the bay appear to be about equal (Allen 1975), this export from the estuary must reflect marsh export. It amounted to about one-half of the Day et al. estimate.
.

Table Birds of the 28. Mississippi Deltaic Plain on the Audubon Society "Blue List," indicating that their populations are declining (Mabie 1976).

Brown Pel ican (Pelecanus occidental is American White Pel ican (P. erythrorhynchos Reddish Egret (Egretta rufescens ) White- faced Ibis (PI egad is chihi White Ibis (Eudocimus albusl Black-crowned Night Heron (Nycticorax nycticorax Red-shouldered Hawk (Buteo 1 ineatus Northern Harrier (Circus cyaneus Osprey (Pandion hal iaetusT Bl ac k vul ture (Coragyps atratus ) Loggerhead Shrike (Lanius ludovicianus )
) ) ) ) )

Hopkinson et al published additional in salt marsh data 1978. respiration Since then Smith et al (1982) published an incomplete carbon budget for the same area which includes estimates of methane evolution and new data on CO2 I evolution. have attempted to create a new budget from all this information and some direct carbon dioxide flux measurements of that photosynthesis include root production (Gosselink et al The weakest links in all these 1977). budgets are the paucity of root production infonnation and our inability to measure marsh export directly.
.
.

Endangered species,
71

Figure 51 shows measurements of CO2 through a S_. al terni flora stand at different seasons. The cuvette used to collect these data enclosed 0.07 m^ of marsh, including sediment and aboveground vegetation, so the data should represent the whole community. Notice that nearly
flux

Spt*lb('7S

il Is I

4,680

is due to g C/mVyr, most of which the einerjent grass. Net primary production was 3,670 g/m^/yr. There are no other figures comparable to these from direct measurement.

Net aboveground production from clip plot studies is only about 850 g/m^/yr.

leaving an estimated 2,800 g/mVyr underThat is not impossible ground production. but is certainly very high. Community respiration was about 3,150 g/m^/yr, which is not too different from the estimates of Day et al (1973) and Hopkinson and Day of around 3000 g/mVyr; but in (1977) their studies 90 percent of this was plant
.

Table 29. Estimates of different components of the carbon budget of Mississippi deltaic salt marsh community (g C/m^/yr).
Carbon flux
Technique

Reference

Input Gross community primary

production

4,680

CO2 flux
II

Net plant primary production (above and belowground) 3,670 793 Aboveground einergents 578 871

a
b

CI ip
11

plot
II II

M
II

c d

1,158

II

Mean Belowground production

850 2,820

II

II

Mean

Difference (3,670-850)

Output Community respiration

3.150 3,081
1,010 2,750

Emergent plant respiration

CO2 flux Sed. oxygen flux & calc. pi ant resp. CO2 flux calculated from

a
f ,g a

other studies
Consumers
2,140 302-316
140
5

f,g
a

CO2 difference

Leaching from live plants Methane production Lost to deep sediments

265

Oxygen flux & calc. for large consumers Leaching studies Methane flux Subsidence rate X sed. C content

f,g
i

Balance (export and unaccounted) Net community production 1,260 300

300 150

from CO 2 from organic balance from N bal ance from estuary export & bay P:R ratio

f,g
j

References
a c
-

b -

Gossel ink et al Kirby 1971 Kaswadji 1982

1977 and unpubl.

e
f

Hopkinson et al. 1978 White et al . 1978 Hopkinson and Day 1977

from literature respiration (calculated In the CO; flux studies, twovalues). is thirds associated with the aufwuchs The expericoinmuni ty and the sediments. detennined data for consumer mentally respiration are 2,140 g/m^/yr from CO2 flux measurements and about 300 g/m^/yr The CO2 flux was determined from O2 flux. with the marsh unflooded, the Oj flux when About the marsh surface was submerged.
140 g/m'^/yr may be lost through leaching, 265 g/m^ /yr are lost to deep sediments, and another 5 g/m-^/yr are lost as methane.

Its understood. is production poorly than the salt marsh. high, probably higher Because flushing energy is low, export is This suggests expected to be low also. that respiration must be very high, but 1978) decomposition studies (White et al rates than in show slower loss salt marshes.
.

NUTRIENT CYCLES
coastal marsh ecosystems, as in In other types, organic productivity depends on the availability of inorganic nutrients in the right proportions at the right to both times. Growth limitation due nutrient limitation and toxicity can and probably do occur in marshes. However, of the 12 inorganic minerals known to be required by plants, only N appears to be regularly limiting to marsh plant growth.
Iron limitations have been reported (Adams 1963), but subsequent studies have not supported this observation (Haines and In fact Fe and Mn are much Dunn 1976). more likely to be in toxic concentrations in i.iarsh soils because of their increased availability under anaerobic conditions. For example, Fe is found in marsh plant tissues in concentrations up to 1,800 ppm (Haines and Dunn 1976), which is well over 10 times the concentration in most agricultural crops.

Over the whole community the net the for balance unaccounted (that is, C available for export) is 1,120 organic Export of all the aboveground g/m^/yr. Hopkinproduction would not equal this. son's estimate of about 300 g exported/m^/ yr is also the balance left over when all other inputs and outputs are considered. It is a reasonable figure in that it the estimate of et al matches Happ (1977). Furthermore, the H budget (see Nutrient Cycling), which is derived from different and measurements, assumptions also makes a value of about 300 g C reasonable, assuming that the exported N is all organic with a C:N ratio of 21.6 (Delaune et al. 1981),
.

The discrepancy between 300 and 1,120 is large. The best that can be said for the C balance in deltaic salt marshes at present is that there appears to be a large amount of organic production for which the fate is unknown. Part of it is certainly exported, but we do not know how much. differences Methodological certainly contribute to the uncertainty.

g/m^/yr

We know even less about C balances in other than the salt marsh. Burial of C in deep sediments does not vary much from salt to fresh marshes. However, as

zones

sulfate decreases, methane availability The annual loss of production increases. C as methane increases from 5 g/m^ in salt marshes to 73 g/m^ in brackish marshes and 160 g/m^ in fresh marshes (Smith et al
.

Marshes are open systems, and the absorption and release of nutrients can have strong effects on adjacent waters. Marshes have been said to reduce eutrofrom nutrients by removing phication these water bodies and, conversely, to be a that source of nutrients supplements The evidence for aquatic production. is that Mississippi delta salt marshes they are sinks for all nutrients, that they absorb inorganic N and release part it as reduced ammonia and organic of and that fonns, export organic they the most C. important Ecologically nutrients in the marsh are N, P, and S.
Ni

1982a).
the other hand, because flushing energies are lower than in salt marshes one would expect waterborne organic export to decrease toward fresh areas. The
On

trogen

brackish

marsh,

in

particular,

is

very
74

Nitrogen, as mentioned earlier, has been found to limit growth in most marshes 1982). (see Mendelssohn et al Nitrogen chemistry in anoxic soils is extremely complex and is made even more so by the
.

proximity of aerobic and anaerobic layers In the in marsh sediments (Figure 53). aerobic layer, oxidation of ammonium to nitrate occurs. This is an extremely thin layer in most delta marshes because the into the rate of diffusion of oxygen flooded soil is not fast enough to supply microbial the demand the by large The nitrate can diffuse down population. it is zone where the anaerobic into reduced to nitrous oxide and nitrogen gas and lost from the marsh ecosystem.
Nitrate can also be reduced all the to ammonium, and perhaps as much as 50 way percent of it is reduced to this form under the environmental conditions of a delta salt marsh (Smith et al 1932a).
.

zone to another ions.

the zones migrate to the

Either the oxidized nitrate or the reduced ammonium can be taken up by the emergent grasses, but free nitrate is present in Undoubtedly, only the thin aerobic layer. nearly all the N absorbed by the marsh The nitrification of plants is ammonium. ammonium and its subsequent denitrification to N2 is facilitated by the vertical movement of the aerobic-anaerobic interThe face as the tide rises and falls. ions do not even have to diffuse from one

Most of the N in the substrate is organic; mineralization (the decomposition of organic material and release of inof this material nutrients) organic all of the ammonium yields nearly available for and for absorption nitrification (Patrick 1982). As much as 3.8 yg N/ml soil/week (inland) to 11.1 is mineralized (streamside) ug/ml/week under optimum conditions (Brannon 1973). This compares to a peak demand by S_. al terni flora of about 2.1 ug/ml/week based on the Tiaximimum growth rates detennined by Ki rby (1971). Kirby's estimate does not include root production so it is an underestimate, but the indication is that mineralization can provide nearly all the takes inorganic N that the plant up. Delaune and Patrick (1979) came to the same conclusion based on average annual rates.
It is likely, for two reasons, that plant uptake tracks mineralization closely the active part of the growing during season: (1) Nitrogen is limiting plant growth so the plants would be expected to take it up as it became available. (2) During the active growing season, sediment ammo ni urn- N remains at a low very concentration of less than 1 ug/ml , increasing to higher levels of 6 - 7 ug/ml during October and November when the plant growth demand is much reduced (Brannon 1973).

NH,

*
ji.

^s oa"^

LAYER t NITRIFICATION NAEROBIC ..,.,. UPWARD SOIL DIFFUSION LAYER

|^|_|t

NH:-N+N0-2-N+ NOi-Nj
^^'Ji

X DIFFUSION
O"*^*""- "

DOWNWARD

DIFFUSION MINERALIZATION
DENITRIFICATION LEACHING ..^

Nj.N^O

-NOa'-N

A schematic outline of the Figure 63. redox zones in a submerged soil showing some of the N transformations (Copyright. Reprinted from "Nitrogen in Agricultural with of the Soils," 1982, permission American of The Society Agronomy). aerobids layer has been drawn thick for In reality, it is seldom over 1-2 clarity. mm in flooded marshes.

Ammonium not taken up by plants is likely to be lost through deni trif ication. Vegetated marsh plots retained percent of added labelled ammonium-N in the plant and soil, whereas in soil cores without plants only 56 percent of the N labelled was recovered (Table 30). However, denitrif ication and other gaseous losses of N are reported to be low in delta salt because marshes, probably plants absorb ammonium before it can be denitrified. Smith et al. (1982a) reported that only about 50 mg N/m^/yr are released as N2O, and estimated that about 5 g N/m^/yr is released as Nj through deni trif ication. is Nitrogen fixation also relatively minor. Casselman et al (1981) measured fixation rates of 15 and 4.5 N/m^/yr in a streamside and an g inland marsh, respectively.

93-94

75

Table
al

30.

terni flora

'^N- ammonium cores (Buresh et al. 1982)

Influence of Spartina on of plants recovery added over 18 weeks to soil

.

Recovery of added
Soil

Aboveground tissue

Total

Soil

core with

plants
Bare soil

core

so no volatile has forms, Phosphorus losses must occur through and sources Studies in water flow across the marsh. Georgia salt marshes have shown that P in estuarine accuTiulates sediments, reservoir of many fonning an enormous years supply (Pomeroy and Wiegert 1981).
In aerobic soils P rapidly becomes unavailable because it is tied up with But aluminum (Al). under Ca and Fe, anoxic conditions the ferric phosphates are reduced to the more soluble ferrous anions can form, exchange phosphate between clay and organic anions, sulfides in ferric can phosphate replace phosphates, and hydrolysis of phosphate compounds can occur.

of the net belowground primary productivApparently ity of the dominant grasses. most of the stored sulfides are reoxidized annually, by oxygen diffusing into the substrate from the marsh roots grass (Howarth and Teal 1979), but there is a of soluble sulfides possibility being flushed from the marsh to become a source In of biological energy elsewhere. the marsh cited above, Howarth et al (1983) estimated that 2.5 to 5.3 moles of reduced are water by pore S/m^/yr exported with creeks. This adjacent exchange amounts to about percent of the S reduced in the sediment, and as much as 20 - 40 percent of net aboveground pro.

3-7

duction.
No

The P budget for a delta salt marsh in Extractable is presented Figure 64. P averages (and presumably available) between 4 and 3 g/m^ in the sediment over Since the annual the year (Brannon 1973). demand for P by the emergent plants is only about 2.6 g/m^ there does not seem to About be any lack of P for plant growth. 2.3 g/m^ is brought in with sediments, and This 1.7 g/m^ is lost to deep sediments. leaves a balance of 0.6 g P/m^ exported, again probably as organic P.
Sul fur

export of
cant
in

one has investigated whether the reduced S compounds is signifi-

marshes. delta Mississippi the total S measured (1973) of salt marsh sediments (Figure 49) and found the same kind of seasonal al et Howarth variation by reported A crude estimate of the amount of (1983). reduced S lost to deep sediments by marsh subsidence shows it to be in the neighborThis is hood of 1 g (0.3 mol)/m^/yr. about the same amount of S deposited by in southeastern forests precipitation We have no idea of (Swank et al 1984). the reduced S flux from the marsh.
Brannon content
. .

The S is not cycle interesting S because has been to limit reported plant growth in marshes, but because of its important role in energy transfer. and still not This is a new fully When oxygen and nitrate understood role. are depleted in flooded soil s, sul fate can act as a terminal electron acceptor and is reduced to sulfide in the process. (This gives the marsh its characteristic rotten egg odor).
In anoxic salt marsties sulfate is a In fresh marshes major electron acceptor. where the supply of sulfate is limited, C The is reduced to methane instead. sulfide radical is a form of stored energy that can be tapped by S bacteria in the of or other oxidants presence oxygen

STORIES

The role of severe storms on marshes received little attention, mostly because their occurrence is unpredictable and their immediate effects difficult to Storms occur with remarkable document. A 1.5-m frequency on the delta plain. wind tide occurs about every 8 years. (Figure 12), and smaller storms are annual Most of the sediment is deposited events. in the coastal marshes during these high water periods or during winter storms (Figure 32).
has

(Howarth et al. 1983).

the energy
S

northeast Atlantic coast marsh flow through reduced inorganic compounds was equivalent to 70 percent
In
a

that et al (1977) reported Day Hurricane Carmen in 1974 defoliated swamp forests in its path two months earlier A large amount of than nornial leaf fall. and P was flushed from the organic C, N, swamp to the fresh, brackish, and salt the marshes of the lower estuary by Part of rains. torrential accompanying this material undoubtedly resulted from
.

77

the early defoliation, but visual evidence pointed to thorough flushing of stored detritus froui the swamp floor which would weather nonnal under out wash not conditions
.

On the other hand, a survey of salt in Barataria and the biomass marsh Terrebonne basins in progress at the time of the same hurricane (Gosselink et al 1977) showed no evidence that dead biomass collected from the marsh surface was any before the in different sampled plots hurricane than after.
.

of Hurricane effects Short-term Camille on species composition in fresh and brackish marshes near the mouth of the described River were Mississippi by Chabreck and Palmisano (1973). They found that an increase in salinity caused by the The major hurricane tide was ephemeral. effect seened to be widespread destruction of vegetation, especially woody species, and water which uprooted and by wind Recovery ripped apart stands of plants. of most species was rapid so that prehurricane levels of abundance were approached lakes and In the small within a year. ponds, however, the submerged and floating vegetation was slow to recover.

Probably the most dramatic alteration documented in marshes is that described by Valentine (1977) in the chenier plain of southwestern Louisiana. One hundred sixty CI adium ha of thousand jamai cense (sawgrass) were killed by the saline tide in The 1957. of Hurricane Audrey following year 86 percent of this area was During the drought years of open water. the early 50' s annual grasses and sedges 1972 abundant. became Sagittaria By falcata (bulltongue) occupied 74 percent of the area and Nymphaea odorata (white 11 jamaicense percent. water-lily) C_. in itself never reestablished any because seed extensive areas, oerhaps Secondary effects viability was very low. on duck these of changes vegetation Prior to feeding habits were dramatic. an were seeds 1959 jamaicense C^. In the important component of duck diets. years immediately following the hurricane, stomachs contained primarily rice duck dependence on seeds, heavy indicating agricultural areas outside the marshes. During succeeding drought years, when the of marshes quantities produced large annual grass seeds, large numbers of both ducks and geese were attracted to these seems likely, It habitats. therefore, that hurricanes are major forces on gulf coast marshes, initiating changes that can have significant consequences for years following the storm.

78

CHAPTER FOUR THE MARSH IN THE COASTAL BASIN

that is, they are not isolated islands out of touch with their the Quite surroundings. contrary, the main reason that they are of particular interest to environmentalists and conservationists is because they are with strongly coupled surrounding In Chapter 2 we say that the ecosystems. main physical driving forces for marshes are the upstream river and the downstream ocean. Both are outside the marsh, but the annual variation in river flow, the and periodic switching of its channel thereby its nutrients and sediment, and the periodic variation in the gulf water level and all determine the salinity character of the marsh. Similarly,
flarshes are open ecosystems;
marshes are biotically they open contribute to other biologically many ecosystems. Figure 65 illustrates these marsh couplings with other ecosystems: zone to marsh zone; marsh to estuary; marsh/estuary to gulf, river and adjacent uplands; and intercontinental couplings.

COUPLINGS AMONG ECOSYSTEMS

Intra-Basin Couplings
The coastal basin can be viewed as a set of coupled subsystems, for indeed the marshes, bays and streams in the basin are A basin is tightly coupled. typical organized by the internal freshwater-salt

MARSH ZONE TO MARSH ZONE

Figure 65. Conceptual other ecosystems.

diagram

illustrating

the

coupling

of

delta

marshes

to

79

water gradient. We take the organization for jranted, but brackish areas are always between fresh and salt areas. The marshes next to the uplands are usually fresher than marshes in the interior of the basin because they receive rain runoff; salt marshes are more naturally dissected by channels than fresh marshes because they receive stronger tidal so energy, and forth.
biotic are Similarly, assemblages We have organized along these gradients. seen that one of the chief consumer groups in the marsh, the waterfowl, partitions itself within the different marsh zones according to the tolerance of individual and for salt for species preference available foods, marsh ponds, and water But these preferences are only depths. On any single aerial bird ones. average census, individual flocks may be found in fresh marsh or in salt marsh. They inove the different marsh zones, freely among taking advantage of favorably changing The increased conditions. waterfowl when marshes changed from sawgrass density to in the annuals, mentioned previous chapter, is an example of the mobility of the fauna among marsh zones. The possible displacement of muskrats toward saline marshes by the invading nutria is another.

FISH & SHELLFISH

WHirr iiHRiMP

TEAL & PINTAIL

WAGING BIRDS

BLACKBIRDS
&

CRACKLES

Patterns of estuarine use by Figure 66. nektonic organisms in the Barataria basin, Louisiana (Chambers 1980).
are, from an economic point of view, the most important group of consumers that frequent the coastal marshes. Typically they spawn on the continental shelf, move into estuaries as juveniles, and return to the Gulf of Mexico as adults to continue the cycle. Nearly all the commercially important nektonic species on the gulf are coast (Gunter estuarine-dependent Within the estuary marsh habitat 1957). is crucial for these species. For example. Turner (1977) showed that both along the gulf coast and worldwide, the commercial shrimp harvest is directly related to the The marsh area in the inshore nursery. relationship is to the total marsh area not just salt marsh; the relationship of yield to the inshore open water area is

provide organisms particularly good examples of the use of coastal multiple subsystems within the basin 56). Many year-round (Figure residents of the estuary are euryhaline move and the basin. freely throughout Such species as the bay anchovy, mullet, alligator gar, rainwater killifish, and tidewater silverside are found from salt to freshwater, many of them in the small creeks that border the marshes. Others, like the threadfin shad, the blue and channel catfish, and the river shrimp move down basin during the fall and winter as brackish areas freshen. The marinespawned croaker, menhaden, and blue crab use the whole estuary as a nursery area, penetrating all the way through salt and brackish zones to fresh marshes in their migrations.
Extra-Basin Couplings
The marine-spawned, estuarine-dependand shellfish mentioned above
80

Nektonic

poor.

The brown
cal

shrimp

life

cycle

is

typi-

estuarine-dependent species in their juvenile Early (Figure 67). stage they can be found deep in the marsh
in

for these

small
in

ent

fish

crease

As they inbayous and ponds. move slowly out into size, they

Spfi

Figure 67.

The life cycle of the brown shrimp (Gosselink 1980),

The birds then spread out shallow ponds. Deer and forage elsewhere at night. other mammals may also venture out into marshes to forage from upland resting areas (Schitoskey and Linder 1979).
to

larger, deeper water bodies which they for appear to use as "staging areas" occur These anigrations emigration. primarily at night and are keyed to the the lunar tidal cycle, with phase of movement during periods of highgreatest est tides (Blackmon 1974).
the Mississippi Delta there appear fish species that spawn in fresh water and move to the ocean as they in But other locations these mature. species make extensive use of the marshes through which they pass on these migrations.
In

Intercontinental

Couplings

to be

no

A different kind of migratory use of marshes is that of numerous bird species which move daily in and out of the marshes to feed. Wading birds, for example, may nest in adjacent upland areas and along beach rims but feed along the marsh edges Their and in marsh ponds during the day. One daily travels may cover many miles. manber of this group, the white ibis, has been reported to travel as much as 80 km its from (Lowery nesting site to feed In a similar vein,Tamasier (1976) 1960). teal and found green-winged wintering pintail resting during the day on large.

dramatic most The inter-ecosystem couplings are those of the migratory birds that link Canadian and Alaskan pothole wetlands to coast marshes. The gulf delta wetlands are at the Mississippi southern extreme of the major duck and (Figure 68). goose migration corridors Many songbird species winter further south and are found moving through the delta marshes only during fall and spring migraAs mentioned earlier, we have wery tions. poor information about the importance of winter-habitat quality of birds that nest in the far north, but all indications are that it is extremely important for nesting success. TEMPORAL USE OF MARSHES
It is to interesting different migrating species wetlands at different times.

observe
use

how coastal

(Figure 69).

81

Major duck migration Figure 58. coast marshes (Bellrose 1980).

Bird populations sre largest during the winter when ducks and geese are abundant. all these to It is misleading group on as sone migrate however, species, to South America, as shown for the through and teals. These two species pintail reach peak abundance late in the year and again in the spring, apparently because a large proportion of the population moves in across the mid-winter. south gulf

corridors

to

gulf

About 60 species the summer (Mabie 1975). of land birds, mostly songbirds, migrate through the delta to South America each They do not use the marsh extenyear. However, sively, but usually fly over it.

Wading bird densities in the marsh peak during the summer. Although they are year-round residents, they appear to be much more active in marshy areas during
82

during northward spring migrations they frequently encounter strong head winds and take refuge on the first landing sites, the cheniers and slightly elevated marsh occasions their these During ridges. the and be can densities high, very marshes can be important for their surSome of these songbirds, like the vival. red-winged blackbird and the great-tailed grackle, nest in the coastal marshes in

MARINE-SPAWNED JUVENILES
(EURYHALINE)

o o
X HI s

O 3

u.

CHAPTER FIVE WETLAND VALUES, HUMAN IMPACTS, AND MANAGEMENT

terai The "value" an imposes orientation on the anthropocentric The term can be discussion of marshes. used in an ecological sense to refer to functional processes, for example, when we speak of the "value" of primary production in providing the food energy that drives the ecosysten or the "value" of a predator in the size of herbivore controlling But it is to populations. important distinguish this use of the term from its ordinary use which refers to the services wetlands perfonn for man
.

WETLAND VALUES
Wetl and Harves
t

The reasons that wetlands are legally protected have to do with their value to society, not with any abstruse ecological processes that proceed therein; this is the sense in which "value" is used in this These perceived values arise out chapter. of the functional ecological processes described in the previous chapters, but are detennined also by the location of a particular marsh, the human population pressures on it, and the extent of the resource.

The easiest wetland value to discuss is the harvest of animals on it. Aside from the important fur animals, most commercially important species associated with wetlands are in habitats migratory, requiring addition to marsh to complete their life This includes all cycles. group commercially important fish and shellfish, recreational fish and hunted species, waterfowl. Qual tatively, t is clear that delta marshes are important habitats for these species, and the completion of their normal life cycles depends on the marshes.
and quantify that depend
i i

The of in extent the marsh, particular, has been one factor that has lowered the value of gulf coast marshes in human eyes. There is so much marsh that a few acres for losing any specific project has not been seen to be of much In this chapter I will first consequence. review the services natural wetland systems provide for society, then discuss the problems of trying to compare the values of natural ecosystems with more conventional economic systems. Finally, I will outline what appear to me to be the major management issues in Mississippi del ta marshes .
84

This has been the dependence rationale for imputing the whole economic value of the harvest to the marsh, although this is not without problems from an economist's of view. The point Louisiana coast fishery harvest is the largest in poundage in the country, and wild fur the harvest is also without and recreational equal. Sport fishing The hunting generate comparable revenues. per acre dollar value of these harvests has been detennined a number of by individuals. The figures in Table 31 for the Barataria basin are representative. Cited values usually range from $50 to $200/ha/yr, depending on the geographic area and the Other assumptions made. measures of wetland value for harvested species would be the weight of harvested or animals the number of hides and carcasses. These measures would not be to subject year-to-year variability in prices, but from an economic point of view they are not much good for comparison to other commodities.

The estimated economic value of Table 31. the Barataria from harvests basin, Louisiana (Mumphrey et al 1978).
.

Ac t i V

ty

the abundance of is masked effect hy sulfate in seawater, so perhaps sulfide accumulation in freshwater wetlands is a In better index oF atmospheric input. delta fresh marshes about 20 mq S/mVyr in is sulfide as deep sequestered This is more or sediments (Hatton 1981). less pennanently removed from circulation
in

the

geography physical (Mclntire 1959).

of

the

delta

the S cycle.

also valuable because are Marshes act as giant water reservoirs during they The vegetation may provide some floods. resistance to the flow of water, slowing it down and thus protecting inland areas, but most of the benefit is probably its This is best seen on storage capacity. rivers where large riparian areas store storm waters and decrease the river stage downstream, reducing flood damage.
the Charles River in nassachusetts, role was deemed effective enough by that U.S. they Engineers Army river flood plain rather the purchased flood-control build than expensive structures to protect Boston (D.S. Army coastal The broad, 1972). Engineers expanse of the Mississippi Delta acts more Its value has to be as a storm buffer. seen in the context of marsh conservation a The full vs. fury of development. coastal storm hits the barrier islands and marshes first and it attenuated as it crosses them, damaging little property of other and value. societal Buildings are zone coastal in this structures vulnerable to the same storms, and damage is often high. Inevitably the public pays this damage much of the cost of through
On

Smardon (1979) described wetlands as and rich educationally visually environments because of their ecological Their complexity interest and diversity. for wetlands excellent sites makes research. Many artists have been drawn to tiiem, notably the Georgia poet Sidney Lanier, the painters John Constable and the Louisiana John Singer Sargent, and Lockwood, many photographer Clyde of lesser artists other public Each year thousands of these recognition. I artists paint and photograph marshes. suspect that many wetland visitors use hunting and fishing only as excuses to wildness and its solitude, experience frontier that pioneering expressing instinct that may lurk in us all.
Confl icting Values

this the

With this long list of marsh values one might expect marsh conservation to be issue that an support. everyone would This is not so, and the reason is simple. tract a marsh owner of The private few of from very benefits financially In Louisiana land can be these services. leased to trappers and hunters for perhaps of LSU School (Chabreck, $25/ha/yr Forestry and Wildlife Management; pers. The owner has no monopoly on, and comm.). cannot sell, the fishery resources and the improved air and water quality associated with the marshes.
To the owner the wetland is valuable drainage for primarily for development construction or agriculture, or dredging subsurface mineral for and drilling resources - that can bring in thousands of This hectare dollars annually. per conflict between private ownership and public services is becoming more intense as density population everywhere it is but increases, particularly several for in wetlands impassioned reasons. First, population density and development pressure are particularly high are marshes on open coasts; second, in considered be that cannot systems and third, marsh development is isolation; essentially irreversible.

for taxes relief, rebuilding public services such as roads and utilities, and federally guaranteed insurance.

Esthetics
A very real but difficult aspect of its to the marsh esthetic capture is value, often hidden under the dry tenn "nonconsumpti ve use values", which simply in out means that people enjoy being delta marshes marshes. The Mississippi are a rich source of information on our remains of The cultural heritage. Indian villages, mounds of prehistoric shells or middens, have contributed to our understanding of both their culture and

86

Recognizing the value of wetlands and educating the public and public officials to these values are important milestones that have led to legislation (particularly Section 404 of the Clean Water Act of 1977) protecting marshes from unconsidered modification. Wetland management did not with this legislation, hut certainly begin the Clean Water Act has focused attention Some of these on many wetland issues. those that relate Issues, particularly delta marshes, directly to Mississippi in will be discussed the rest of this chapter.

instruments Probably the most used for ecological evaluations in general are the U.S. Fish and Wildlife Service Habitat Evaluation Procedures (HEP, USFWS 1980) and the U. S. Habitat Army Engineers Evaluation System (HES, USAE 1980). Both HES has were developed for upland sites. not been adapted for wetlands, and HEP wetland applications are still evolving. These procedures are most valuable when used to compare two different areas or to compare an area before modification to the expected state afterward.

WETLAND EVALUATION

One important component of wetland management is the evaluation of proposed actions in wetlands. Under Section 404 of the Clean Water Act of 1977 a perniit is required for wetland activities that might affect water quality. For activities that an environmental impact statement require (as required by the National Environmental two different kinds of Policy Act) evaluation are involved. the First, ecological value of the area in question is determined - that is, the quality of the site as compared to other similar sites or its suitability for supporting wildlife. Second, the ecological value of the habitat is compared to the economic value of some proposed activity that would destroy or modify the habitat - in other Both prowords, a benefit:cost analysis. cedures are fraught with difficulties. Both require an evaluation of the relative values of different commodities, like comAbove all, both paring apples and oranges. about require numerous value judgments what is ecologically desirable.

probably the aiore illustrates both the potential and the problems of evaluation. In this procedure the suitability of a site is evaluted for a number of different game species, commercially important species, and of interest for species special structure or function. For each ecosystem species, habitat suitability is evaluated on a scale of 1.0 for a number of habitat characteristics. These Habitat Suitability Indices (HSI's) are multipl ied by the area of each species' habitat under consideration to Habitat Units yield Thus both habitat quality and (HU's). area are combined in one number. (1979) listed the Schamberger et al. (1) habitat assumptions of the system: value can be habitat (2) quantified; suitability for a species of concern can be evaluated from habitat characteristics; overall can habitat value be (3) determined by assessing suitability for selected species; (4) habitat quantity and quality are directly related to animal numbers. It is the that apparent community HSI's depend on the species selected for evaluation.

The HEP procedure,

detailed,

Essentially all proceaures now in use assess the relative value of wildlife Lonard et al habitat. (1981) evaluated 20 different wetland valuation systems. The in all of them was enphasis overwhelmingly on the evaluation of the ecological habitat function of wetlands. Hydrology functions are poorly documeted Evaluation of and difficult to quantify. and recreation silviculture, heritage, functions are also considered open for improvement (Lonard et al 1981).
. .

The result of the HEP analysis is a set of HU's for individual species for the site or sites in question. The HU's can be compared within a site or among sites for determining best management scenarios. The values can be used to help make a management decision about the site, as for instance, project impacts offsetting In this case, sites through mitigation. with equal value in terms of HU's are created or set aside for use by the species in question. 87

This or any other evaluation system nust play off bewildering detail against simplifying integrations to facilitate the The evaluator decisionmaking process. must integrate mentally the information about a number of different individual species in order to make the decision. The ideal solution is a compromise between extremes to allow a simple enough decision to be made, but detailed enough for the decisionmaker to feel confident about it.

values related to clean air and water and other "life support" functions.

One controversial approach uses the idea that energy flow through an ecosystem or the similar concept "embodied energy" (the total energy required to produce the commodity, Costanza 1980) is a valid index of the totality of ecosystem functions; and that index is furthermore, this Thus applicable to human systems as well. natural and human systems can be evaluated of on the basis one common currency:

All to make procedures developed decisions about wetlands are based on human values and human judgments about what is good and what is not. They reflect what humans think is important, and that fact is a basic ingredient in all In the case of management. HEP, the have been procedures standardized, individuals can be trained and certified to carry them out, and reproducibility is These facts often make us quite good. forget the value-laden nature of the whole enterprise.

"embodied energy." is a (Since there linear between embodied relationship energy and dollars, that more familiar currency can also be used.)

When habitat values are monetized for benefit :cost analyses, a whole new set of are on the assumptions superimposed do not intend to ecological evaluation. discuss these because are well they covered by several other authors (Shabman and Batie 1978; McAllister igS'"). The has evolved from economic methodology theory that assumes that in a free economy the market price reflects the value of a (the commodity willingness-to-pay approach).
I

The general response to this kind of approach is probably fairly summed up by and "Certain Reppert (1979): Sigleo aspects of the evaluation structure .... are too theoretical and unsubstantiated to be considered for application, general particularly those involving the analysis of energy flows and the conversion of values to values." energy monetary recent both the However, in years theoretical base and the methodology have been much improved
.

This leads to real in problems monetizing nonmarket commodities like pure water and air, and in pricing marshes whose monetary value in the marketplace is determined by their value as real estate, not their "free services" to society. Consequently, attempts to monetize marsh values have the generally emphasized commercial "crops" from marshes - fish, shellfish, furs, and recreational fishing and for which hunting pricing are available. As Odum methodologies (1979) pointed out, this kind of pricing ignores ecosystem-level values related to and hydrology and productivity, global
88

better assumptions, Costanza Using showed that the economist's and willingness-to-pay approach energy analysis converge to a surprising degree. In Table 32 the average gross benefits arrived at by summing the gross economic value of marsh different resources are roughly equivalent to ($342/acre/yr) the latest value arrived at from the embodied energy of biological productivity This convergence suggests ($300/acre/yr). an integrated methodological framework for The approach has the real evaluation. merit of being equally applicable to both natural and human systems, but like e^ery other approach it simplifies by converting everything into one currency.
(1933)

Since the purpose of the exercise is to compare apples to oranges or oil wells to marshes, some kind of equivalence must to me it seems be but established, lose of the real to dangerous sight

Estimates of the economic value Table 32. of Louisiana's coastal wetlands comparing willingness-to-pay approaches with energy 1983). (Costanza approaches analysis Approach
Shadow value*
1979 $/acre/yr)
Reference

I will discuss oriented (Table 33). issue or each major problem, briefly bringing in the role of the various human activities as they apply. Since habitat is far the most loss (marsh loss) by pressing problem, it will receive the major emphasis.

Marsh Loss and Salt Intrusion

As discussed in Chapter 1 (Figure 23), the rate of marsh loss to open water has been accelerating over the past 50 years to the present rate of about 1.5 percent of the delta marshes being lost annually. Although the circumstances leading to this natural and involve loss are complex there is processes beyond human control good evidence that a significant part of of human result is a the problem modification of the Mississippi River and This discussion will the deltaic plain. be limited to these latter factors, that is, those which man can hope to manage on a regional scale.
,

Willingness-to-pay approaches 155 Consumer surplus 241 Gross benefits

352 544 231

Average of gross benefits Net benefits Replacement value

342 237
25 ,662
3

d b
d

,120

Energy Analysis approaches Biological productivity

7,374 300
in

*Price that would prevail market.

References
a
:

perfect

Humphrey et

al

b
c

Gossel ink et al Vora 1974 Costanza 1983

1978 1974

the development activites listed All Table 33 contribute to marsh loss. Reclamation does so because it impounds and drains wetlands, essentially turning them into upland habitat. Although marsh "reclamation" is still occurring, the pace of development is much slower than it was early in this century (Gossel ink et al of cost the and impounding, 1979), an area is and maintaining draining economics that so prohibitive becoming
in
.

One compromise has structures involved. been suggested by Lichfield et al. (1975), who used a planning balance sheet to list the major commodities exchanged and to identify the recipients of the cost and This procedure ensures that the benefits. the important factors in the benefit:cost analysis are explicitly recognized rather than being lumped into a single dollar value.

Table 33. Major wetland in Mississippi impacts

issues and human delta wetlands.

ENVIRONMENTAL ISSUES

MINERAL EXTRACTON

WETLAND MANAGEMENT
River Deltaic Mississippi Plain the major wetland management issues are marsh loss, salt intrusion, and the maintenance of habitat and water quality. These are interrelated problems. They are affected by a number of human activities, can be grouped as the major ones but or conservationeither development
In

FLOOD CONTROL
NAVIGATION

the

RECLAMATION

_
+

HABITAT
PROTECTION
&

ENHANCNG
HABITAT

CREATION

89

dictates
purposes
.

against

this

practice

for

;iiost

extraction, The impact of mineral flood control, and navigation on marsh loss occurs primarily through the canals Table 34 dredged for these operations. of effects the major ecological lists canals in the deltaic marshes, the kinds of iiiechanisns that should minimize these and the specific impacts, ecological practices that are being used management to used be could or implement these Because mechanisms. experimental good often lacking, many of the is evidence are and effects mitigation procedures those document will I inferred. But statanents that can be documented. many are merely reasonable extrapolations from what is known.

operate, but the result is often death of the plants and, as the roots die, loss of If the their capacity. peat-binding that the so rapidly changes salinity are not immediately by replaced plants salt-tolerant the more species, often underlying peat rapidly erodes and large, These shallow lakes appear (Dozier 1933). and biocheinical to linked are changes microbial changes in the peat associated with salt intrusion (Dozier 1983).
also Canals hydrologic change patterns that inodify a marsh independently of any salt effect. Straight, deep canals and marshes shallow in lakes, bays, the natural flow, depriving capture channels of water (L. Gosselink 1934; are Canals comm.). Turner, pers. efficient, allowing more hydrological ly the fresh water than runoff of rapid a As sinuous channels. normal result, water levels fluctuate more rapidly than in unmodified marshes, and minimum levels Sheet flow of are lowered (Light 1976). water across the marsh surface is reduced by the spoil banks that almost always line a canal. Consequently, the sediment supply to the marsh is reduced, and the water on the marsh is more likely to stagnate than when freely flooded.

salt

Canals alter marshes by accelerating intrusion, changing hydrology, and affecting benthic and aquatic organisms. Salt intrusion is closely tied to changes when occurs It in deep, hydrology. connect channels low-salinity straight zones. to areas Large high-salinity channels that link the marshes navigation directly to the gulf are particularly intrusion salt in efficient allowing but a channel et al 1979), (Gosselink from a saline bay into a less saline marsh also allows salt intrusion.
.

fresh and into intrusion Salt the stresses marshes ntennediate We do not know exactly how vegetation. in the salinity fairly subtle changes
i

Since canals change the marsh water budget, the salt budget, and the sediment supply, any mechanisms that can influence these three factors might be useful ways of canals. effects the of minimizing mechanisms. several 34 lists Table Generally, an increased freshwater supply

Table 34. Impacts of canals in Louisiana coastal marshes leading to habitat loss, and mechanisms and management practices to minimize these impacts.

Type of impact

Mechanisms to minimize impacts

Management practices

1.

Salt

intrusion

to a marsh also increases the sediment load since rain runoff and river water are

Mechanisms both generally quite turbid. shallow maintain sinuous, that slow, natural channels and overland flow will generally also reduce salt intrusion and stabilize water levels. They may also reduce the sediment-carrying capacity of the water, but this has to be balanced flow. overland increased the against
are already A number of practices being used or are potentially useful to minimize marsh loss (Table 34). They can be grouped as those that build new marshes those that to replace those lost and minimize the loss of existing marshes.

for There are also possibilities better design of canals. Where possible, channels in follow natural they should order to maintain natural circulation are Spoil deposits usually patterns. on both sides of the canal, placed the from canal the adjacent isolating marsh. Any design that breaks the spoil barrier to allow better exchange with the marsh would probably be an improvenent. Unfortunately, there are no studies upon which to base detailed recommendations.
It is common practice to require that when canals cross natural streams and other canals, they must be blocked to minimize the danger that the new canal will capture the flow of the other channels Some fairly and/or allow salt intrusion. work is needed straightforward engineering to improve the design of these barriers. or rock are usually Earth fill, shell, These materials have densities much used. greater than the organic marsh, and their weight tends to settle and load down the As a result, the barriers adjacent marsh. are constantly breaching, especially at It would seem that an inert their ends. plastic material of the same density as the surrounding marsh, perhaps anchored into place with a mininum number of pilings, could be more effective.
- cerMany canals can be backfilled all those dredged for pipelines and tainly also many that lead to dry or depleted know little about the we Yet wells. relative value of backfilling compared to in Work canals. (Menprogress open delssohn, Sikora and Turner, Center for to the Wetland Resources, LSU) points canals of effectiveness backfilling because the practice removes spoil banks and also raises the bottom of the canal (although it seldom fills it completely because of the oxidation and dissipation of sediments when they are exposed in spoil banks) to a depth where the water column does not stratify. Oxygen is then available to the sediments, and a healthy In addition, benthic infauna can grow. there is some evidence that these shallow ditches, if left open in areas where marsh the can is circulation improve poor, Such marshes. of adjacent quality research on canals can yield major benefits to the State by providing practical marsh means of degradation. reducing

assessed the (1982) Day and Craig potential for reduction in wetland loss by several techniques. They mitigation concluded that diversion of fresh water to baild new marshes could only create the km^ of marsh a year, and Atchafalaya had the potential of building about 18 The largest potential for saving km^/yr. marshlands (30 - 40 km^/yr), therefore, was of new strict control by regulatory

1-3

canal s.

We little have experimental experience on which to outline the best Prohibition against canal ing technology. new canals would be the best solution, but barrier against crossing prohibition islands, connecting basin interiors to the periphery, and creating canals that shunt upland runoff around marshes would be partial solutions.
is a wellDirectional drilling established that would technology eliminate the need to dredge canals for It has not been used many well heads. often in the coastal marshes, and good studies the extra cost of comparing the directional drilling against environmental of the canal cost are

needed.

Another technology that needs to be of is the use air cushion explored These vehicles to traverse the marshes. are used in the tundra and might provide a to well sites and even way approach transport drilling rigs without damaging the marsh extensively and without the need for canal dredging.
91

Recently some permits for dredging in the delta marshes have included requirements for marsh improvement elsewhere to mitigate the damage in the permit area. This is a creative mechanism for conservmarsh, although at the expense of ing other marsh tracts. Unfortunately, the methodology for assessing the true environmental cost of canals is rudimentary, so the relationship between the canal damage and the mitigation effort is somewh a t
a

common in the chenier plain than in the because the firner delta, primarily levee substrate in the cheniers makes construction much less expensive and more effective.

rb i

ra ry

If environmental costs of development wetlands are to be internalized by the developer, we need much better information In a about how to assess these costs. recent article /Vnft et al (in review) present a methodology and make a benefitrcost assessment of an oil well access canal in the chenier plain. Based on their suggest that a methodology, they conservative estimate of the environmental is cost for a typical exploratory well $380,000 (1981 dollars) per kilometer of access canal
in
.
.

The idea behind these impoundments is prevent salt intrusion and thus retard marsh loss. Unfortunately, there is little evidence to show that they are effective, and some evidence to suggest that they are not. Baumann, Conner, and Gosselink (LSU Center for Wetland Resources; unpubl, MS.) analyzed marsh loss rates in impoundments compared to adjacent unimpounded areas, ivere loss rates and concluded that in than impoundments actually higher Wicker et al, outside them (Figure 70). (1983) also measured marsh loss rates in different kinds of impoundments in the Wildlife Rockefeller Although Refuge. they presented no comparative data, it is apparent from their maps that marsh degradation is occurring in all the impoundments except perhaps those with pumps for water level control
to
.

A word needs to be said about some current practices that do not seen to One of effectively retard marsh loss. these is channelizing upland runoff. In to fairness, this practice is not used minimize marsh loss, but it is a common flood control measure. The impact on marshes is negative because it shunts the sediments of rivers and runoff away from both rivers to marshes, by leveeing prevent overbank flooding and by digging to deliver flood deep-dredged channels water through and around marshes instead of over them. This is a case of conflicting interests in the coastal zone. Until interests recently, flood control took ascendancy over marsh loss concerns. A more balanced of evaluation this "solution" to flooding is needed.

The is that problem, think, sediment input is a key element in the ability of a marsh to accrete fast enough to keep up with subsidence. Impounding
I

/f"-4
NATimAL tUfiSH

IMPOUNDED MARSH
,

K"

z
111

o
o

a.

Another common is the practice construction of levees and impoundments to In recent years, all prevent marsh loss. over the deltaic and the chenier plain marshes small levees no nore than a meter high have been thrown up by private land owners. Marsh are also impound-nents in and wildlife common State Federal management areas where they were created to improve habitat for waterfowl and fur These animals. levees are much more
92

1962

YEAR

YEAR

The increase in open water in Figure 70. The wetlands. natural and impounded in wetland loss of pattern greater impoundments is consistent in both fall, when water levels ire low, and winter, when impoundments are flooded (W. Conner and R. Center for Wetland Resources, Baumann, Louisiana State University; pers. comm.).

In sediment off the supply. nterdistributary basins which have very little surface fresh water input, most of action. the sediments come from tidal to these circumstances Under attempts also retard salt intrusion restrict sediment input.

cuts
i

erosion by keeping the marsh from drying out and oxidizing. Weirs seem fairly effective for stabilizing water levels (Figure 73) and for promoting growth of submerged aquatic plants (Chabreck 1968).
other hand, the evidence from (1976; see study of Steever et al Figure 43) that marsh plant biomass is directly proportional to tide range makes it that marsh is likely productivity reduced by these structures. As far as erosion prevention is considered, there is no evidence that weirs are effective. Weirs are the cheapest kind of marsh Because of the increase in management. behind submerged vegetation, the ponds attract more wintering waterfowl weirs than unwei red ponds (Spiller 1975). They also improve conditions for fur animals.
On

the

the

addition to marsh loss caused by intrusion and hydrologic changes, salt canals also directly change benthic and The nektonic habitat quality (Table 34). deep canals are depauperate in benthic organisms because, at least in bulkheaded the lower part of the water channels, column and the sediments are anoxic most of the year (W. Sikora, LSU Center for Wetland Resources; pers. comm.).
In

On the other hand, canals might enable nektonic to penetrate marsh organisms areas where they previously had no access, although the presence of spoil banks would Fish can use the cancel this benefit. deep water of canals as a refuge during cold when the shallow natural spells streams become almost as cold as the air above them.

Habitat Qua! ity

In the wildlife management areas of delta (Figure 71) several kinds of are to modifications marsh practiced Generally this improve habitat quality. means improved quality for waterfowl and fur animals, sometimes at the expense of But in recent years the fishery species. aim has been a diversified habitat that will support a broad range of species.

The next level of control device is the flap gate and/or variable level dam in a The flap completely Impounded marsh. gate allows water to flow one way through the control structure. Modern ones are reversible, but in Louisiana, with its high rainfall, they are usually set to allow freshwater to flow out of the impoundment and to prevent saltwater from moving in. Because of the all rainfall, surplus impounded areas become fresher with time.

the

is level Where water management active, the opening and closing of water control structures is timed to increase the availability of the managed area to migratory fish and shellfish species. The structure is the weir simplest control 72); this Is a common device found (Figure all over the coastal zone, especially in areas State or Federal managed by authorities. It is a dam placed in tidal creeks to maintain a minimum water level In the marshes drained the creek. by Usually the top of the weir Is about 15 cm below the average marsh surface. The purpose of the weir is to stabilize water levels to the of encourage growth submerged aquatic plants and reduce marsh

The variable height device, which is often incorporated in the same structure, allows the manager to set minimum water levels behind the weir. With this "gravity drainage" systen, if the weather cooperates it is possible to draw down the water In the spring to allow seeds of annual It can emergents to germinate. raised in the winter to make then be shallow ponds for ducks. The most sophisticated water level control Is obtained by pumping water out of or into the impoundment (forced drainThe effectiveness of these manageage). ment measures can be judged by the kinds and of diversity produced vegetation and the use of the (habitat quality) impoundment by birds, fur animals, fish, and shellfish.

Wicker et al (1983) summarized the in effectiveness of the impoundments Annual Wildlife Rockefeller Refuge. vegetation surveys carried out since 1958
.

93

STUDY AREA

--

HYDROLOGIC UNIT BOUNDARIES

GULF
1

OF

MEXICO
10WISNER STATE WMA
1 1

PAUL J. RAINEY WILDLIFE REFUGE

ST.

TAMMANY STATE WMA

2 LOUISIANA STATE WILDLIFE

REFUGE
3 RUSSELL SAGE FOUNDATION WILDLIFE REFUGE

12 PEARL RIVER

WMA

13 BILOXI

WMA WMA

14 BOHEMIA STATE

4 ATCHAFALAYA
5

WMA SALVADOR STATE WMA

15 DELTA NATIONAL WILDLIFE

REFUGE
16 PASS A LOUTRE STATE

6 JEAN LAFITTE NATIONAL HISTORICAL PARK

7 JOYCE

WMA

17 BRETON NATIONAL WILDLIFE

WMA

REFUGE

8 MANCHAC STATE

WMA

9 POINTE-AU-CHIEN STATE

WMA
Figure 71.

Wildlife mdnagement areas

94

in

the Mississippi

Delta.

lb

A weir in the deltaic plain marshes. The strong flow of water across Figure 72. weir is an indication of the effectiveness of the barrier. These structures favorite sport fishing spots (Photograph by Robert Chabreck).

the are

show that the production of the desired emergent annuals and aquatic plants was Even with pumps it was variable. not to control water level in very possible rainy years like 1973, and the level of control decreased as the sophistication of the control devices decreased. In general, the better the water level management, the greater the diversity and desirability of the vegetation (Figure 74).

with the marsh is not surrounding to is off) completely cut comparable unmanaged marshes (Chabreck et al. 1975; Davidson and Chabreck 1983).
The value of freshwater impoundments for species other than ducks is not as fur animals, clear; geese, and marine (Chabreck organisms are not benefitted crawfish can be 1975). However, in raised impoundments successfully managed for ducks (Perry et al. 1970). Brackish marsh impoundments seen to yield excellent crops of marine shellfish and fish if the control gates are managed to allow the juvenile organisms access during their immigration periods (Davidson and Chabreck 1983). Figure 75 summarizes the effectiveness of impoundments.

Water level in the management Rockefeller Wildlife Refuge is credited with increasing waterfowl use from a peak population of about 75,000 ducks in 1951 1952 to over 400,000 dabbling ducks, 40,000 coots and 10,000 diving ducks in 1958 - 1959 when the control structures were put into use (Chabreck 1951). The freshwater impoundments attract the most ducks; use of brackish water impoundments (usually areas in which water exchange
95

Marshes, inside are often burned as

impoundments and out, management practice.

(0

> < Q

source of delta sediments, the Mississippi River itself, is heavily polluted with exotic chenicals which become incorporated in the sediments of any marshes created.
From here they can be magnified into the food chain, leading to the kind of on individual that effects species occurred with the brown pelican. That from the del ta was extirpated s|-iecies of of chlorinated the effect because on shell pesticides egg hydrocarbon been only recently strength; it has reintroduced from Florida (Blus et al 1975).

cr
III

03

z
UJ

>

3 O
20
40

60

80

100

BOTTOM EXPOSURE
Figure 73. year that

(%)

number of days per study area will certain percentages of or exceed equal Based on depth contours bottom exposure, of 43 ponds and 20 years of tide data on coast Louisiana the central (Chabreck

Cumulative
in

ponds

the

1979

Chabreck (1975) questioned the value of he effort. most of this However, acknowledged that burning can be useful to remove a heavy vegetation thatch to allow annual species to germinate and to give an three-cornered earlier start grass during the growing season. Burning is widely practiced to attract snow geese to an area. Trappers find burned areas inuch easier walking, and animal trails are much and more noticeable. However^ nutria raccoon often move from a burned marsh because of the lack of adequate cover.
Water Qual

runoff from urban and a is also serious areas agricultural problem. Seaton and Day (1979), Seaton and Day (1980), and Kemp (1973) documented the runoff from the New of urban effects Orleans area into the Sarataria basin and and Hopkinson Gael Lake Pontchartrai n. showed that eutrophi cation of water (1979) is accelerated by canals which bodies shunt the water around marshes instead of have over them. High coliform counts resulted in oyster bed closures in much of the estuarine area soutli of New Orleans In all and east of the Mississippi River. the primary concern has these examples been with the quality of water in the If 'iiore runoff coastal lakes and bays. to flow across the allowed water was marshes instead of bypassing it through flood drainage canals, it is likely that water quality would improve significantly.
Local

gas production is surprising that activity wetlands, so little is known about the effect of In the delta only oilspills on wetlands. This one group of studies is available. research showed that chronic, low-level oilspills resulted in fairly high levels of hydrocarbons in marsh sediments (Bishop et al 1976) in the Leeville oilfield.
in it
.

With

all

the

oil

and

ty

Water quality is a major issue in Louisiana wetlands as in inany other areas it has received of the but country, little attention, probably relatively because the much more pressing issue of marsh loss has taken the spotlight. The
96

concentrations are These high in aromatic the reflected hydrocarbon in of benthic tissues concentration organisms such as oysters and mussels. The free-swimming emergent grasses and organisms such as the grass shrimp and of killifish had concentrations high unresolved hydrocarbon components (Milan The influence of this and Whelan 1979). pollution on biota could not be separated

AQUATIC PLANTS

80

i;

'|;|

EMERGENT ANNUALS
EMERGENT PERENNIALS

80'

^M
-

Dnll'D
60 66
70

Id
74

:|liiillliU
158

MANAGEMENT
EVENTS

1955-LEVEES BUILT METAL FLAP -GATE

FLAP-GATE CULVERTS LFAKING

MANAGEMENT
EVENTS
MARSH BURN FOLLOWED
BY HIGH SALINE TIDES SEPT 1965 SEVERE

CULVERTS
INSTALLED

CONCRETE VARIABLE
CREST REVERSIBLE LEVEE REPAIRS FLAP-GATE STRUCTURES WEST SIDE i SOME FLAP-GATE ALL FLAP-GATE CULVERTS CULVERTS REMOVED PLUGGED

CAUSED MARSH LOSS

80-

^iufliuil
1965-

iUMil 1
VEAR METAL FLAP-GATE CULVERTS

LEVEES
FLAP-GATE

CONSTRUCTED

MANAGEMENT METAL
EVENTS

CULVERTS
INSTALLED

VEAR FmP- GATE CULVERTS LEAKING WOODEN BOX FLAPGATES INSTALLED CONCRETE VARIABLE
-

MANAGEMENT
EVENTS

MAINTAINED AS

LEVEE REPAIRS

CREST REVERSIBLE FLAP GATE STRUCTURES'

ALL FLAP-GATE CULVERTS

PERMANTLV FLOODED FRESHWATER IMPOUNDMENT

BEGAN SPRING AND SUMMER DRAW-DOWN

DROUGHT FLOODED IN SUMMER FOR MOTTLED DUCK

NESTING

DOUBLE DIVERGENT
PUMPING UNIT

EASTSIOE SOME FLAP-GATE

CULVERTS PLUGGED

REMOVED. WOODEN BOX FLAP-GATE CULVERTS

PLUGGED

The percentage of different types of vegetation Figure 74. Rockefeller State Wildlife Refuge (Wicker et al 1983).
.

in

impoundments

in

the

from the effect of the associated dense network of canals and spoil banks, but the density of marsh grass culms and average height was lower than in control areas (R. E. Turner; pers. comm.).
total and crustaceans, Amphipods, total benthic organisms were reduced 50 percent compared to non-oilfield control areas Killifish (Lindstedt 1978). abundance was substantially less in oilfield marsh ponds than at control sites, although not statistically so because of the large confidence limits. However, the fecundity of Fundulus grandis in oilfield marshes was significantly lower than at control sites, especially the condition

index of females 61-80 mm long (May 1977). It is apparent that we need to know much more about the of effects chronic low-level oilspills.

From a management point of view, water pollution is a good example of the need to Water manage on many different levels. quality of the Mississippi River must be This is a problem national in improved. river's enormous scope because of the watershed.

The control of urban runoff in the delta itself is a regional problem that affects marshes and estuaries in the New Orleans area more than other delta
97

TARGET HABITAT TYPE

PASSIVE ESTUARINE

WATER MANAGEMENT PROGKAMi

CONTROLLED ESTUARINE
Concrete Variable Crest Reversible Flap-Gates Concrete Radial Lift Gates

GRAVITY DRAINAGE
36-in and i8-in Flap-Gates Concrete Variable Crest Reversible Flap-Gates

FORCED DRAINAGE
Pumps

UNCONTROLLED

Wakefield Weirs at -0.5 ft MSL

Nonexisting or Non operable Structures

EMERGENT PERENNIAL VEGETATION:

Ve-H Du-P, Mu-P, Nu-P
.

Intermediate

Ve-A

Local marsh management cannot wetlands. solve that problem. The recommendation to route upland runoff across wetlands rather than around than in order to take advantage of marshes' ability to intercept pola lutants is basin-level problem that involves local, State and Federal manage-

inent

agencies. Finally, closer control of oilspills, oxidation ponds or drilling mud in wetlands are problens that disposal involve not only the local, State and Federal enforcement but also agencies industries in site-specific single problens.

99

REFERENCES

1963. Adams, D. A, vascular plant Carolina salt 44:445-456.

Factors influencing in zonation North marshes. Ecology

Adams,

R. D. B. B. J. fl. Barrett, B. and W. W. G. ackmon, Gane, Mc In tire, Barataria basin: 1976. and framework. processes geologic Wetland Center for Resources, Louisiana State Baton University, Sea Grant Publ LSU-T-76-006. Rouge.
,

Bl

1976. Alexander, S. K. Relationship of macraphyte detritus to the salt marsh Lit t orina rrorata Say. periwinkle, Ph.D. Thesis. Louisiana State Baton 104 pp. University, Rouge.
i

1950. Allan, P. F. Ecological gases for land use in planning gulf coast marshlands. J. Soil Water Conserv. 5:57-52, 85.

1975. Allen, R. Aquatic primary productivity in various marsh environments in Louisiana. M.S. Thesis. Louisiana State Baton University, Rouge. 50 pp.

American

Union. 1983. Ornithologists' 0. U. Check-list of North American Birds. 6th ed Allen Inc. Kansas. Press, Lawrence,
The
A.
.

877 pp.

Amft

3.

Gossel ink, J. M. C. A. Teague, H, Cowan, and C. A. An environreview. evaluation a of

G.

in

izing a Coastal

Cameron Parish, cost-benefit Zone Manage.

Bishop, W,, Whelan.

Bishop, M. Hood, and T. Hydrocarbons. Vol. 3, R. G. J. 6.2 ji_n RosseTink, App. Miller, M. Hood, and L. M. Bahr, eds. Louisiana offshore oil port: environCenter for baseline study. mental State Louisiana Wetland Resources, Baton Prepared Rouge. University, Oil Offshore Louisiana Port, for New Orleans Inc.
F.

1976.

Ph.D. Thesis. Louisiana State marsh. University, Baton Rouge.

Buresh, R. J Patrick, Spartina

1

OSS Amer. J. 45:660-551,

DeLaune, and W. H. 1981. Influence of al ternifl ora on nitrogen Soil Sci. Soc. from marsh soil
R.

D.

Jr.

Observations on 1974. Blackinon, J. H. the emigration of the brown shrimp, Penaeus aztecus , through a tidal
pass in the Caminada bay, Louisiana Louisiana State M.S. Thesis. area. Baton Rouge. University,

P M. Borengasser, G. Drew, R. Byrne, Muller, B. L. Smith, Jr., and C. Wax. Barataria basin: 1976. hydrologic Center and processes. climatologic Louisiana for Wetland Resources, Sea State University, Baton Rouge. 175 pp. Grant Publ. LSU-T-76-012.

Cary,

L.

R.

1907.

The

Blus,

L. R.

and

Brown pelicans certain environmental pollutants Bull. Environ. Contam. in Louisiana. Toxicol. 13:646-555.

J., T. Joanen, A. 1975. M. Prouty.

A.

Bel isle,

and

in Louisiana. oysters Stn., Cameron, La. Bull.

cultivation of Gulf Biol.

8.

56 pp.

1964. Boelter, D. H., and G. R. Blake. of volumetric expression Importance of water contents of organic soils. Soil Sci. Soc. /Vn. Proc. 28:176-178.

Casselman, M. E., W. H. Patrick, Jr., and 1981. Nitrogen fixaR. 0. DeLaune. in a tion gulf coast salt marsh. Soil Sci. Soc. Amer. J. 45:51-56.
H. R. Chabreck, impoundments

Brannon,
of

Seasonal variation 1973. M. and nutrients physicochemical properties in the salt marsh soils of M.S. Louisiana. Barataria bay, Louisiana State University, Thesis. 130 pp. Baton Rouge.
J.

marsh Coastal 1961. for ducks in Louisiana. Southeast. Conf. Annu. 14th Proc. Assoc. Game Fish Comm. 14:24-29.

1936. Brown, C. A. Indian mounds,

The vegetation of the middens, and marshes Bernard St. and in Plaquemines Conserv. Bull. La. Dep. parishes.

1968. Weirs, plugs and Chabreck, R. H. artificial potholes for the management of wildlife in coastal marshes. J. D. Newsom, ed. Pages 178-192 ^n and of marsh estuary Proceedings Division of management symposium. Continuing Education, Louisiana State University, Baton Rouge.

8:423-440.
Brown,

on

Historical commentary 1944. A, the distribution of vegetation in Louisiana and some recent observaProc. La. Acad. Sci. 8:35-47. tions.
C.

Ponds and lakes 1971a. Chabreck, R. H. of the Louisiana coastal marshes and their value to fish and wildlife. Southeast. Conf. 25th Annu. Proc. Assoc. Game Fish Comm. 25:206-215.

Brupbacher,

R. H., J. E. Sedberry, Jr., The coastal 1973. and W, H. Willis. chemical of Louisiana: marshlands materials. properties of the soil La. Agric. Exp. Stn., Baton Rouge. Bull. 672.

The foods and 1971b. R. H. Chabreck, from feeding habits of alligators in environments saline and fresh Annu. Conf. 25th Proc. Louisiana. Game Fish Comm. Assoc. Southeast. Unpubl. MS.

Chabreck,
and

R.H.
soil

1972. Vegetation, water of the characteristics

1978. Buresh, R. J. Nitrogen transformations and utilization by Spartina salt in a Louisiana alterniflora
101

Louisiana coastal Exp. Stn., Baton

72 pp.

region. Rouge.

La. Agric. Bull. 664.

R. Chabreck, wetlands

H.
for
.

1975.

Management of wildlife habitat

Cocks, R. S.

3rd Proc. Bienn. improve.nent Conf. Estuarine Res, Fed. Galveston, Tex.

1907. The flora of the Gulf Gulf Biol. Stn. Biologic Station. Cameron, La. Bull. 7.

Chabreck,

R.H., The on Camille

and

A.

W.

1973.

Mississippi 54:1118-li:n

effects of the marshes River delta.

Palmisano. Hurricane
of

the

1976. Deltas: processes Coleman, J. M. of deposition and models for exploration. Continuing Education PublishInc. ing Company, Champaign, 111. 102 pp.

Ecology
Coleman, J. M., and S. M. Gagliano. 1964. Cyclic sedimentation in the MissisRiver deltaic Gulf plain. sippi Coast Assoc. Geol Soc. Trans. 14:67-80.
.

Chabreck,

Yancey, and L. Duck of usage in the Louisiana management units coatal marsh. Proc. 28th Annu. Conf. Assoc. Game Fish Comm. Southeast.
R.

fIcNease.

R. M., 1975.

K.

Col

23:507-516.
and G. Linscombe. 1978. Chabreck, R. H. Vegetative type map of the Louisiana Louisiana Departcoastal marshes. ment of Wildlife and Fisheries, New
,

ins , J. T., R. Conant, J. E. Huheey, J. L. Kn light, E. M. Rundquist, and H. M. Smith. 1. 1982. Standard common and

Or! eans

current scientific names for North American and amphibians reptiles. 2nd ed. Society for the Study of Amphibians and Reptiles, Department of Zoology, Miami University, Oxford, Ohio.
Conner,
The the
W.
H.
,

Chabreck, R. H., R. J. Hoar, and W. D. 1979. Soil and water Larrick, Jr. characteristics of coastal marshes influenced by weirs. Pages 129-145 J_n J. W. Day, Jr., 0. D. Culley, Jr., R. E. Turner, and A. J. Mumphrey, Proc. 3rd Coastal Marsh Jr., eds. Division of Estuary Manage. Symp. Continuing Education, Louisiana State Baton 511 pp. University, Rouge.
An 1980. Chambers, D. G. analysis of in nekton communities the upper Barataria Louisiana. M.S. basin, Thesis. Louisiana State University, 285 pp. Baton Rouge.

and J.

W.

Day,

Jr.

1932.

ecology of forested wetlands in southeastern United States. 59-87 R. E. Pages Gopal ui RR. G. and D. F. Turner, Wetzel, Wetlands ecology and Whigham, eds. National Institute of management. and International Science Ecology Publications. Jaipur, India.
,

1980. Embodied energy and Costanza, R. economic valuation. Science 210: 1219-1224.
R. 1983. Natural resource Costanza, valuation and management: toward an Center for approach. integrated Wetland Resources. Louisiana State University, Baton Rouge. Unpublished

Clarke,

L.,

and

H.

J.

Harmon.

1967.

The

mangrove swamp and salt marsh communities of the Sydney district. I. soils and climate. J. Vegetation, Ecol. 55:753-771.
Chew,
Sea level changes along the northern coast of the Gulf of Mexico. Trans. Am. Geophys. Union
F.

MS.

1962.

and C. J. Cleveland. 1984. Costanza, R. Ultimate recoverable hydrocarbons in a net energy approach. Louisiana: Scient in press) .
, (

45:272-280.
Costanza, R.,
1972. Coates, D.R., ed. Environmental geomorphol ogy and landscape conservation. Vol 1. Page 350 j_n Benchmark papers in geology. Dowden, Hutchinson and Ross, Stroudsburg, Pa.
R.
C. Neill, S.G. Leibowitz, J. Fruci , L. M. Bahr, Jr., and J. W. 1983. Day, Jr. Ecological models of the Mississippi deltaic plain region: data collection and presentation. U.S. Fish Wildl. Serv. Div. Biol.

102

Sen/., 82/68.

Washington,
342 pp.

D.C.

FWS/OBS-

Coward in, L. M. V. Carter, F. C. Golet, and E. T. LaRoe. 1979. Classification of wetlands and deepwater habitats of the United States. U. S. Fish Wildl. Oiv. Biol. Serv., Serv., 103 FWS/OBS-79/31 Washington, O.C.
.

Conner. 1977. and Productivity nutrient export studies in a cypress swamp and lake system in Louisiana. vn 255-269 M. ed. Pages Wiley, Estuarine Vol 2. processes. Academic Press, New York.
Day,
J.

W.,

Jr.,

pp.

Comparison management

Craig,

J., J. W. Day, Jr., A. Seaton, G. Sniith. 1977, Kemp, and W. Eutrophication. Pages 1-97 ^n N. J. CumuCraig and J. W. Day, Jr., eds. lative impact studies in the Louisiana coastal zone. Part I. Center for Wetland Resources, Louisiana State University, Baton Rouge.
N.
P.

effectiveness of options for wetland loss in the coastal zone of Louisiana. Pages 232-239 in D. F. Boesch, ed. Proceedings of a conference on coastal erosion and wetland modification in Louisiana: causes, consequences, and options. U. S. Fish Wildl. Serv. Biol. Serv. Program, Washington, D.C. 256 pp. FWS/OBS-82/59.
S. Hopkinson, and W. An 1982. analysis of environmental factors regulating fisheries community metabolism and
J.

and N. of the

J.

Craig.

1982.

Day,
1980. Cramer, G. W., and J. W. Day, Jr. of the and Productivity swamps Lake Pontcharmarshes surrounding train, La. Pages 593-545 j_n Stone, J. H., ed. Environmental analysis of Lake its Pontchartain, Louisiana, selected surrounding wetlands, and land uses. Vol 2. Coastal Ecology Lab., Center for Wetland Resources, Louisiana State Baton University, for U.S. Rouge. Prepared Army New Orleans. District, Engineers Contract DACW 29-77-C-0253.

H.

W., Jr., Conner.

C.

production in a Louisiana estuary. Pages 121-136 j_n V. S. Kennedy, ed. Academic Estuarine comparisons.
Press,
New York.
709 pp.

Crowe,

P.

R.

tol ogy.

1971. Concepts of cl imaLongman Group Ltd., London.

589 pp.

L. H. M. A., Kennedy, and R. Deegan, 1983. Factors contributCostanza. ing to marsh land loss in Louisiana's coastal zone. Pages 915-920 j_n W. K. Lauenroth, G. V. Skogerboe, and M. Analysis of ecological Flug, eds. systems: state of the art in ecologiElsevier cal modeling. Publishing New York. 992 pp. Co,

Davidson,

and R. R. H. Chabreck. B., 1983. Fish, wildlife and recreational values of brackish marsh impound-

ments. Proceedings of water quality and wetland management conference. New Orleans, La. Aug. 4, 1983.
1899. Davis, W. M. cycle. Geogr. J.

The

geographical

1984. Deegan, L. A., and B. A. Thompson. The ecology of fish communities of River Deltaic Plain the Mississippi In Yanez-Arancibia A., ed. region. Fish community ecology in estuaries and coastal Universidad lagoons. Nacional Autonoma De Mexico, Mexico City, Mexico, (in press).
,

14:4.81-504.

Deevey,
Day,

E.

S.

Jr.

1970.

Mineral

cycles.

W., Jr., W. G. Smith, P. Wagner, and W. Stowe. 1973. Community structure and carbon budget in a salt marsh and shallow estuarine bay in Louisiana. Center for system

J.

Sci.

Am.
R.

223:148-158.
D.,

DeLaune,

Wetland Louisiana Resources, Sea University, Baton Rouge. Publ. 30 pp. LSU-SG-72-04.
Day,
J. W.

State Grant

Patrick, Jr. Nitrogen phosphorus in a cycling gulf coast salt marsh. 143-151 in Proc. 5th Bienn. Pages Int. Estuarine Res. Conf. Jekyll Ga. October 7-12, 1979. Island,
W.

and

H.

1979.

and

Jr.

T.

J.

Butler, and W.

H.

Delaune,
103

R.

D.

R.

J.

Buresh,

and

W.

H.

1979. Patrick, Jr. Relationship of to properties standing crop biomass of Spartina al terniflora in a Estuarine Coastal Louisiana marsh. Mar. Sci. 8:477-487.
soil

in slopes 2:181-199.

Louisiana.

Geonoroh,

Fisk,

H.

tion

N., 1944. Geological investigaof the alluvial valley of the

S.
'\r'iiy

DeLaune, R. D., C. N. Reddy, and U. H. 1981. Accumulation of Patrick, Jr. nutrients and heavy metals plant through sedimentation processes and in a accretion Louisiana marsh. Estuaries 4:328-334.
DeLaune, R. D., R. H. Baumann, and J. G. Gosselink. 1983. Relationships vertical accretion, apparent among sea level rise and land loss in a Louisiana coast marsh. J. gulf Sediment. Petrol. 53:147-157.

lower Mississippi River. U. Engineers, Mississippi Commission, Vickshurg, Miss.

Fisk,
H. N.

River

1952. Geological investigathe Atchafalaya basin and problens of Mississippi River diversion. U. S. Engineers, Army

tions

of

River Mississippi 145 po. Vickshurg, Miss.

Commission,

1955. Nearshore sediments of Fisk, H. N, the continental shelf off Louisiana. Pages 1-23 j_n Proc. 8th Tex. Conf. Soil Mech. Found. Fng.
Fisk,

Delwiche, cycle.

C.

C.

1970.
Am.

Sci.

The nitrogen 223:137-146.

and E. McFarlan, Jr. 1955. quarternary deltaic deposits of the Mississippi River. Crust of the
H.

N.,

Late

1983. Assessment of change Dozier, M. D. in the marshes of southwestern Barataria basin, Louisiana, using historical aerial photographs and a information system. M. S. spatial Thesis. Louisiana State University, Baton Rouge. 102 pp.
1983. Production and fate Ducklow, H. W. of bacteria in the oceans. Bioscience 33:494-501.
R. M., E. A. Antie, and W. H. Engler, 1976. Effect of disPatrick, Jr. solved oxygen on redox potential and nitrate removal in flooded swamp and marsh soils. J. Environ. Qual 5:230-235.
.

earth. Geol 62:279-302.

Soc.

Am.

Spec.

Pao.

D. F. 1957. Recent deltaic Frazier, deposits of the Mississippi River, their and developnent chronology. Gulf Coast Assoc. Geol. Soc. Trans. 17:287-315.

Gael,

1975. Environment and Ethrington, J. R. John Wiley and Sons, plant ecology. 347 pp. London.

and C. S. Hopkinson. B. T. 1979. land-use and Drainage density, in Barataria eutrophication basin, Louisiana. Pages 147-163 j_n J. W. Day, Jr., 0. D. Culley, Jr., R. E. Turner, and A. J. Humphrey, Jr., eds. Environmental in conditions the coastal 3rd zone. Proc. Coastal Marsh Estuary Manage. Symp. Division of Louisiana Education, Continuing Baton State University, Rouge.
,

511 pp.

Gagliano,
About nutria and Evans, control. U.S. Bur. Sport Wildl. Denver, Colo. 65 pp.
J.

S.

M,,

and

J.

L.

Van

Beek.

1970.

their Fish.

Fenchel

T. 1970. Studies on the decomposition of organic detritus derived from the turtle Thai assia grass testudinum Limnol. Oceanogr. 15:14-20.
,
.

and Geologic geonorphic of deltaic aspects processes, Mississippi delta system. Hydrologic and of studies coastal geologic Center for Wetland Louisiana. Louisiana State UniverResources, sity, Baton Rouge. Rep. 1.
1970.

Gagliano,
1975.

S.

M.,
An

and

J.

L.

Van

Beek.

Fisk,

H.

N.

1939.

Oepositional

terrace
104

management

to multiuse approach in the Mississippi delta

223-238 system. Pages Broussard, ed Deltas, Houston exploration. Society. Houston, Tex.
.

jji

M.
s

L.

nodel

for

Geological

marsh. Center for Wetland Resources, Louisiana State Baton University, Sea Grant Publ LSU-SG-74Rouge.
.

03.

Gagliano,
P.

P. Culley, D. W. Earle, Latiolais, P. Light, A. Rowland, R. Shleinon, and J. L. Van Beek. 1973. Environmental atlas and multiuse management plan for southcentral Louisiana. Center for Wetland Louisiana State Resources, Baton Coastal University, Rouge. Resour. Unit Rep. 18, Vol. 2.

S.

M.,

Gosselink,
and

King,

C.

C. S. Hopkinson, Jr., Parrondo. 1977. Common narsh plant species of the gulf coast area. Center for Wetland Resources, Louisiana State Baton University, Rouge. Prepared for Dredged Material Research U. S. Program, Army Engineers Waterways Experiment Station, Vicksburg, Miss. Tech. Rep. D-77-44.

J.

G.,

R.

T.

Gagliano,
K.

M., K. J. Meyer-Arendt, and Wicker. 1981. Land loss in the Mississippi River Deltaic Plain. Trans. Gulf Coast Assoc. Geol. Soc. 31:295-300.
S.

M.

1982. Nutrient dynamics of Gaudet, J. J. papyrus swamps. Pages 305-319 vn_ B. Gopal, R. E, Turner, R. G. Wetzel, and D. F. eds. Wetlands Whigham, and National ecology management. Institute of InternaEcology and tional Science Publications. Jaipur,
India.

Gosselink, J. G., C. L. Cordes, and J. W. Parsons. 1979. An ecological characterization study of the Chenier Plain coastal ecosysten of Louisiana and Texas. 3 vols. U.S. Fish Wildl. Biol. Serv. Serv., Program, D.C. FWS/0BS-7B/9 Washington, through 78/11.
1984. Gosselink, L. Hydrology and the effects of canals in Western Terrebonne parish marsh. La. M.S. Thesis, Louisiana State Baton University, Rouge.

514 pp

L. W. 1966. RelationshiD of Giles, vegetation to salinity in a southeast Louisiana coastal marsh. Bureau of Fisheries and Sport Wildlife, Division of River Basin Studies. 26 pp. Miss. Vicksburg,

Gowanloch,
ing
in

H.

N.

1933.

Fishes
La.

Louisiana.
23.

Dep

and fishConserv.

Bull.

638 pp.

Good, R. E., N. F. Good, and B. R. Frasco. A review of primary production 1982. and decomposition dynamics of the belowground marsh component. Pages 139-157 V. S. ed. Kennedy, jn Estuarine Academic comparisons. 709 pp. Press, New York.
1980. Tidal marshes Gosselink, J. G. the boundary between land and ocean. U. S. Fish Wildl. Serv., Biol. Serv. FWS/OBSProgram, Washington, D.C. 13 pp. 80/15.
1974. Gosselink, J. G., and C. J. Kirby. Decomposition of salt marsh grass, al terni flora , Loisel. Spartina Limnol. Oceanogr. 19:825-832.

1938. Seasonal variations in Gunter, G. abundance of certain estuarine and marine in fishes with Louisiana, reference to life particular histories. 8. Monogr. Ecology 3:313-346.

Some relationships of Gunter, G. 1967. estuaries to the fisheries of the Gulf of Mexico. Pages 621-638 lO. G. H. Lauff, ed. Estuaries. Am. Assoc.
Adv.

Sci.

Publ.

83.

Haines, E. Growth

1976. B., and E. L. Dunn. and resource allocation reof al terniflora Spartina sponses Loisel. to three levels of NH -N, Fe and NaCl in solution culture. Bot. Gaz. 137:224-230.

Happ,

Gosselink,
Pope.

J.

G.

E.

1974.

Odum, and R. M. The value of the tidal

P.

G., J. G. 1977. Jr.

of
105

Gosselink, and J. W. Day, The seasonal distribution in a Louisiana organic carbon

J. Estuarine estuary. Sci. 5:695-705.

Coastal

Mar.

Hatriaway, 1936.

E.

S.,

and

W.

T.

relation of in communities plant Proc. La. Louisiana. 3:68-71.

The

Penfound. salinity to southeastern

Acad.
Sci.

Hopkinson, C. S., J. W. Day, Jr., and B. 1978a. T. Gael. Respiration studies Louisiana salt marsh. in a An. Y Mar. Limnol. Centro. Cienc. del Univ. Auton. Mexico 5:225-238.

Hopkinson,
and

1981. Aspects of marsh Hatton, R. S. accretion and 'jeochemistry Barataria Thesis. Louisiana. M.S. basin, Baton State Louisiana University, 115 pp. Rouye.
:

Jr., J. G. Gosselink, Parrondo. 1978b. Abovemarsh of seven production ground in coastal Louisiana. plant species Ecology 59:750-769.
C.

S.,

R.

T.

Heitmeyer,
1981.

and L. H. Fredrickson. wetland conditions in the Mississippi delta hardwoods influence mallard recruitment? Pages 44-57 j_n Wildl. Nat. 46th N. Am. Trans.
M.

E.,

Do

J. G. Gosselink, and R. Hopkinson, C. S. Production of 1980. Parrondo. T. coastal Louisiana marsh plants calcufrom lated phenometric techniques. Ecology 61:1091-1098.
,

Resour.

Conf.
1978. level
J.

M. Teal. 1979. Howarth, R. W., and J Sulfate reduction in a New England Limnol salt marsh. Oceanogr. 24:999-1013.

Hicks,

S.

n.

tial

sea

States. 1379.

An average geopotenseries for the United Geophys. Res. 83:1377-

Howarth,

Hinchee,

Selected aspects of E. 1977. biology of Lake Pontchartrain , Louisiana M.S. Thesis. Louisiana. State University, Baton Rouge. 75 pp.
R.

the

and J. E. Hobbie. 1982. W. regulation of decomposition and heterotrophic microbial activity in salt marsh soils: a review. Pages ed . V. S. 183-207 Kennedy, vn_ Estuarine Acadenic comparisons. Press, New York.
R.
,

The

Hoffpauir,

for C. M. 1968. Burning marsh coastal management. Pages Marsh 134-139 _i_n J. D. Newsom, ed. and estuary management symposium. Baton State Louisiana University,

Howarth, R. W,, A. Giblin, J. Gale, B. J. Peterson, and G. W. Luther III. 1983. Reduced sjlfur canpounds in the pore waters of a New England salt marsh. Bull. Ecol. (Stockholm) 35:135-152.
and L. McNease. 1972. Joanen, T. Popuof lation distribution alligators the reference to with special Louisiana coastal marsh zones. Symof American the Alligator posium Council, Lake Charles, La. Unpubl.
,

Rouge.

Mollis, J.
rice. 514.

P.

1967.
La.

Agric. Baton Rouge.

Toxicant diseases of Bull. Stn. Exp.

MS.

Hood,

1971. A., and A. R. Colmer. Seasonal bacterial studies in Barafor VJetland Center taria bay.
M.

Johnson,

Louisiana Resources, Baton Rouge. sity, Bull. 6:16-26.

State UniverCoastal Stud.

Hopkinson,
1977.

Jr., and J. W. Day, Jr. model of the Barataria bay salt marsh ecosystem. Pages 235-265 in C. A. S. Hall and J. W. Day, Jr., eds. Ecosystem modeling in theory John Wiley and Sons, and practice. 684 pp. New York.
C.

S.,

and J. G. Gosselink. W. B., Wetland loss directly associ1982. ated with dredging in the Louisiana coastal zone. Pages 60-72 i_n D. F. Boesch, ed. Proceedings of a conference on coastal erosion and wetin Louisiana: land modification and causes, options. consequences, U.S. Fish Wildl. Serv., Biol. Serv. FWS/OBSProgram, Washington, D.C. 256 pp. 82/59.
J.

Jones,
106

Jr.

D.

C.

Carter,

and

H.

checklist

197 5. Revised Genoways. of North Anerican mammals Occas. north of Mexico. Pap. Mus. No. 28. 14 pp. Tech. Univ. Tex. H.

mental

U.S. Laboratory, Army Engineers Waterways Experiment Station., Vicksburg, Miss. Prepared for U.S. Army Engineers District, New Orleans, La. Potamology Invest. Rep.
221.

Kadlec,

for 1979. Wetlands H. R. Pages 490-504 j_n tertiary treatment. P. E. Greeson, J. R. Clark, and J. E. Wetland functions and Clark, eds. values: the state of our understandWater Resources American ing. Minn. Association, Minneapolis, 674 pp.

King,

K. A., E. L. Flickinger, and M. H. Hildebrand. 1977. The decline of brown pelicans on the Louisiana and Texas gulf coast. Southwest. Nat. 2:417-431.

Kirby,

Kadlec, Wetlands Pages 436-456 in P. E. R. Greeson, J. Clark, and J. E. Clark, eds. Wetland functions and values: the state of American Water our understanding Resources Association, Minneapolis, 674 pp. Minn.
,

R.

H.

and J. A. Kadlec, and water qual i ty.

1979.

The annual net pridecomposition of mary the salt marsh grass Spartina al ter ni flora Loisel. in the Barataria bay Ph.D. Thesis. estuary of Louisiana. Louisiana State Baton University, 74. pp. Rouge.
C.
J.

1971.

production and

Kane,

H.

T.

1943.

Louisiana. New York.

William
341 pp.

The of bayous Morrow and Co.,

1972. Circulation and Kjerfve, B. J. salinity distribution in a marsh-lake system in coastal Louisiana. Center for Wetland Louisiana Resources, Sea State University, Baton Rouge. 54 pp. Grant Publ. LSU-SG-72-06.

Kaswadji,

primary different

The estimation of 1982. five production using in a methods Spartina salt marsh in Barataria al terni flora Louisiana. M.S. Thesis. bay, State Raton Louisiana University,
R.
F.

Kniffen,
and

Louisiana, its land Louisiana State Univerpeople. sity Press, Baton Rouge.
F.

B.

1968.

Kolb,

Rouge.

39 pp. R.

Kelley,

A taxonomic survey 1965. Delta fishes of National Wildlife Refuge with emphasis upon distribution and abundance. M.S. Louisiana State University, Thesis. Baton Rouge. 133 pp.
.3.

of

the

and J. R. Van Lopik. 1958. of the Mississippi deltaic U.S. plain - southeastern Louisiana. Army Eng. Waterways Exp. Stn. Tech.
C.

R.,

Geology

Rep.

2:3-482.
R.

Lemaire,
of

Kellogg,

J. L. 1905. Upon the marine food mollusks of Louisiana. Gulf 43 Biol. Stn., Cameron, La. Bull. 3. pp.
G. P. 1978. Agricultural runoff and nutrient dynamics of a swamp forest in Louisiana. M.S. Thesis. Louisiana State Baton University,

Kemp,

J. 1960. Preliminary report marsh vegetation study, Mississippi River-Gulf Outlet navigation project, Orleans and St. Bernard U.S. Army Engineers, Parishes, La. Branch of River Basin Studies, Prepared for U.S. Vicksburg, Miss. Fish and Wildlife Service, Bureau of Sport Fisheries and Wildlife, Region 35 pp. 4, Atlanta, Ga.

Le

Rouge. Keown, M. P., E. A. Dardeau, and E. M. 1980. Characterization of Causey. and the suspended-sediment regime bed-material of the gradation River basin. EnvironMississippi
107

Histoire 1758. Page du Pratz, A. S. de la contenant la Louisiane, 3 vols. decoverte de ce vaste pays. Paris
.

Lerch, 0., W. W. Clendenin, and W. C. Stubbs. 1892. Geology and agriculture of Louisiana. Louisiana Agri-

cjl tural

Experiment
295 pp.

Station

Baton

Rouye.

State Louisiana University 565 pp. Baton Rouge.

Mabie,
R.'

Press,

and M. P. Kettle, N., Lichfield, Evaluation in the 1975. Whitbread. Press, Pergainon planning process. Oxford.

0.

U.
6,

App.
R.

1976. Sec. 12
M.

Avifauna.
rn_

Vol.

3,

J.

G.

Gosselink,

Light,

197G. Ch. ?j_nJ. Hydrology. Gosselink, R. R. Miller, M. Hood, Louisiana and L. M. Bahr, Jr., eds. offshore oil environmental port: Technical Vol. II. baseline study. for Center Wetland Appendices. Louisiana State UniverResources, for Baton Prepared sity, Rouge. Louisiana Offshore Oil Port, Inc., New Orleans.
P.

G.

A. Hood, and L. M. Louisiana offshore Jr., eds. oil environmental baseline port: Center for Wetland Resources, study. Louisiana State Baton University,

Miller,

Bahr,

for Louisiana Prepared Rouge. Offshore Oil Port, Inc., New Orleans.

Maltby, E. nunbers

Lindstedt, tenn

long on operations oil-recovery inacrobenthic communities near marshM.S. Thesis. estuarine creek banks. Baton Louisiana State University,
Rouge.
67 pp
E.,
.

0.

M.

1978.

Effects

of

Changes in microbial from alternative resulting managenent strategies in wetland and in related habitats southern 477-506 Louisiana. B. Pages J_n R. E. Turner, R. G. Wetzel, Gopal and D. F. Wetlands Whigham, eds. and National ecology management. Institute of Ecology and International Science Publications, Jaipur, India. 514 pp.
1982.
, ,

Lisitzin,

1961. Pattullo. The principal factors influencing the oscillation of sea level. seasonal J. Geophys. Res. 56:345-852.

and

J.

G.

and W. L. Nicholas. 1974. Marchant, R. An energy budget for the free-living Pel odora nematode (Rhabdi tidae) Oecologia 16:237-252.
.

May,

Lomincki

Bandola, and J. JankowaModification of ska. 1968. the Wi egert-Evans method for estimation of net primary production. Ecology 49:147-149.
,

A.,

E.

The effects of oilL. N. 1977. recovery operations on the biology in a of killifishes and ecology M.S. Thesis. Louisiana salt narsh. Baton State Louisiana University, Rouge. 30 pp.

Lonard, R.,
J.

E.

J.

Clairain,

R.

T.

Huffman,

P. L. 0. E. Brown, Hardy, 1981. Watts. W. Ballard, and J. Analysis of methodologies used for the assessment of wetlands values. Environmental Laboratory, U.S. Army Engineers Waterways Experiment Station, Vicksburg, Miss. Prepared for U.S. Water Resource Council, Washing68 pp. ton, D.C.

W.

Evaluation in 1932. McAllister, D. M. Massaenvironmental planning. of Institute chusetts Technology Press, Cambridge.
Methods of cor1959. Mclntire, W. G. cultural remains vvith stages relating of coastal development. Pages 340359
in

2nd

Coastal

Geogr.

Conf.

CoastaT Studies Institute, Center for

Louisiana Wetland Resources, University, Baton Rouge.
Mehlich,
A.

State

Louisiana

of Conservation. Department The birds of Louisiana. 1931. Dep. 598 pp. Conserv; Bull. 20.

Louisiana 1960 Lowery, G. H. State Unive Louisiana ^rsity 567 pp Baton Rouge
1974. G. H. Lowery, Louisiana and its

birds, Press,

1972. Uniformity of expressA case for test results. on a volume results calculating Commun. Soil Sci. Plant Anal. basis.
ing
soil

3:417-424.

of The mammals adjacent waters.

Mehlich,
test
108

A.

197 3.

results

as

Uniformity of soil influenced by volume

Commun. weight. Anal. 4:475-486.

So

i 1

Sc

Plant

Engineers District, New Orleans. Mumphrey,

J,, J. S. Brooks, T. D. Fox, Fromherz, R. J. Marak, and J. D. Wilkinson. 1978. The valuation of wetlands in the Barataria basin. Urban Studies Institute, University of New Orleans. Prepared for Louisiana Departnent of Transportation and Development, Baton Rouge.
C.
A.

Mendelssohn,

A., Postek. 198?. Elenental analysis of deposits on the roots of Spartina al terni fl ora Loisel. Am. J. Bot. 69:904-912.
M.

E.

and

T.

3.

^1endelssohn, Postek.

A., K. L. McKee, and M. T. 1982. Sublethal stresses al terni flora Spartina controlling
1.

Pages 223-242 ht_ B. Turner, R. G. Wetzel and D. F. Whigha-n, eds. Wetlands ecology and management. National Institute of Ecology and International Science 514 pp. Publications, Jaipur, India.

productivity.
Gopal
,

National
of

R.

E.

Research Council. 1982. Impacts trends on emerging agricultural fish and wildlife habitat. National Academic Press, Washington, D.C. 244

pp.

National
Meo,
M., J. W. Ford. Day, Jr., and T. 1975. Overland flow in the Louisiana coastal zone. Center for Wetland

1979. annual
D.C.

Atmospheric Agency. imatological data, U.S. summary for Louisiana. Departnent of Commerce, Washington,
local
cl

Oceanic

and

Louisiana State UniverResources, Sea Grant Publ sity, Baton Rouge.

.

LSU-SG-T-75-04.
Meyers, S. J., D. G. Ahearn, and P. C. Miles. 1971. Characterization of Center for Wetland Resourcyeasts. es, Louisiana State University, Baton
Rouge.
Coastal
S.
,

E. C. E. Nester, W., B. J. Roberts, McCarthy, and N. N. Pearsall. 1973. microbes Microbiology. Molecules, and man. Holt, Rinehart and Winston, Inc. New York. 719 pp.

Stud.

Bull.

6:7-15.

Milan,

C.

Accumulation
bons
in
a

exposed to Pages 65-37 j_n J. W. Day, Jr., D. D. Culley, Jr., R. E. Turner, and A. J. eds. Proc. 3rd Jr., Humphrey, Coastal Marsh Estuary Manage. Symp. Division of Continuing Education, Louisiana State Baton University, 511 pp. Rouge.

T. Whelan. 1979. petroleum hydrocarsalt marsh ecosysten steady state oil input.

and of

1982. The ecology of New Nixon, S. W. England high salt marshes: a commuU. S. Fish nity profile. Wildl. Serv., Biol. Serv. Program, WashingD.C. 70 pp. ton, FWS/OBS-81/55.

Odum,

P. 1979. The value of wetlands: hierarchical approach. Pages 16-25 in P. E. Greeson, J. R. Clark, and J. E. Clark, eds. Wetland functions and values: the state of our understandAnerican Water ing. Resources 674 Association, Minneapolis, Minn. pp.

E.

Milner, C, and R. E. Hughes. 1968. Method for the measurement of the of primary production grasslands. Int. Biol. Handb. 6. Programme Blackwell Scientific Publications, 70 pp. Oxford, England.
1983. Filamentous algal Moncreiff, C. A. mat communities in the Atchafalaya River delta. M.S. Thesis. Louisiana State University, Baton Rouge. 129

Odum,

E. and M. E. Fanning. P., 1973. Comparison of the productivity of Spartina al terni flora and Spartina in cynosuroides coastal Georgia marshes. Ga. Acad. Sci. Bull. 31:1-12.

Odum,

pp.

and R. C. Pinkerton. 1955. speed regulator: the optimmi efficiency for maximum power output in physical and biological systems. Am. Sci. 43:331-343.
H.

T.,

Time's

Montz, G. N. Reports

1976. to the

Botanical elements. U. S. file, Army

109

O'Neil,

T.

1949.

The

muskrat

in

the

Louisiana Louisiana coastal marsh, Department of Wildlife and Fisheries,

New Orl eans.
IS 2 pp.

Plant distribution 1944. Penfound, W. T. in relation to the geology of Louisi8:25-34. ana. Proc. La. Acad. Sci
.

Commercial Jr., 1971. Palmisano, A. W. wildlife work unit report to Wildlife Fisheries Commission, study of and the Louisiana coast at the AtchafaLouisiana Wildlife and basin. 1 aya 180 Fisheries Commission, Vol. 1.
?P-

Penfound,

S. and E. T., Hathaway. the of Plant communities marshes. Louisiana southeastern 3:61-52. Proc. La. Acad. Sci

W.

1936.

Penfound,

Palmisano, A. W. 1972. Habitat preference of waterfowl and fur animals in the northern gulf coast marshes. Pages H. ed. R. 153-190 Chabreck, ji_n Coastal marsh and estuary management Division of symposium proceedings. Continuing Education and School of Forestry, Louisiana State University, 315 pp. Baton Rouge.
Parrondo, R. T., J. G. Gosselink, and C.S. Effects of salin1978. Hopkinsnn. and drainage on the growth of ity Rot. Gaz. three salt marsh grasses. 139:102-107.

S. E. Hathaway. in the communities marshlands of southeastern Louisiana. Ecol Monogr 8: 1-56.

W.

1933.
.

T., Plant
.

and

Perry,

W. Joanen, and L. G., Jr., T. 1970. Crawfish-waterfowl , a McNease. multiple use concept for impounded Conf. 24th Annu. Proc. marshes. Southeast. Assoc. Game Fish Comm. 23: 178-183.

Pomeroy,

and R. G. Wiegert, eds. ecology of a salt marsh. 271 Springer-Verl ag. Inc., New York.
L.

R.,

1981.

The

pp.
1977. Nesting colonies of Portnoy, J. W. seabirds and wading birds in coastal Louisiana, Mississippi and Alabama. U.S. Fish Wildl. Serv. Biol. Serv. FWS/OBSProgram, Washington, D.C. 77/07.

Parrondo, R. T., j. G. Gosselink, and C. 1981. Infuence of S. Hopkinson, Jr. on the absorption of rubidisalinity al terni flora and un Spartina by Ga z Bot. Pi stichl is spicata 142:402-407.
. .

Parsons,
1980.

K.

A.,

and

A.

A.

de

la

Cruz.

and flow grazing Energy behaviour of conocephal ine grasshoppers in a Juncus roemerianus marsh.

1979. Factors affecting Rainey, G. B. nutrient chemistry distribution in M.S. marshes. coastal Louisiana Louisiana State University, Thesis. 103 pp. Baton Rouge.

Ecology 51:1045-1050
Jr. 1982. W. H., Patrick, Nitrogen transformations in submerged soils. Pages 449-455 j_n F. J. Stevenson, ed. Am. Nitrogen in agricultural soils. 22. American Soc. Agron. flonogr. Society of Agronomy, Madison, Wis.

Reddy,

R. W. H. Patrick, Jr., and R. Evaluation of 1980. Phillips. selected processes controlling nitroSoil Sci. gen loss in flooded soil. Soc. Am. J. 44:1241-1246.
K.
,

E.

1970. Energy Payne, W. J. growth of heterotrophs. Microbiol. 24:17-52.

yields and Annu. Rev.

1975. The response of three Payonk, P. M. of marsh to macrophytes species enrichment at artificial Dulac, Louisiana Louisiana. M.S. Thesis. 122 State University, Baton Rouge. pp.

1979. 17/3. cinu W. R. w. r\. Jiyicu. Reppert, R. T., and -''--*- Sigleo. for wetlands Concepts and methods evaluation under development by the U. S. Army Corps of Engineers. Pages 57-52 i_n P. E. Greeson, J. R. Clark, Wetland funcand J. F. Clark, eds. tions and values: the state of our Water American understanding. Resources Association, Minneapolis, Minn. 674 pp. 1937. Riley, G. Mississippi

The significance of the River drainage for bio-

110

logical

conditions

(Dama

dam a )

nippon]"!

and sika deer (Cervus , M.S. Thesis. Louisiana

Siialley,

A.

E.

1960.

State University, Baton Rouge.

1978. Shabman, L. A., and S. S. Batie. coastal Economic value of natural wetlands: a critique. Coastal Zone Manage. J. 4:231-247.
1975. Shaver, G. R., and W. n. Billings. Root production and root turnover in tundra a wet Barrow, ecosystem, Alaska. Ecology 55:401-409.

salt marsh 41:785-790.

Energy flow of a grasshopper. Ecology

R. C. 1979. Visual-cultural Smardon, values of wetlands. Pages 535-544 j_n P. E. Greeson, J. R. Clark, and J. E. Wetland functions and Clark, eds. values: the state of our understandAmerican Water Resources Assoing. ciation, Minneapolis, Minn. 674 pp.

Snart, R. M., Influence

Shew,

D. M., R. A. Linthurst, and E. 0. 1981. Seneca. Comparison of production computation methods in a southCarol ina North eastern Spartina a1 terni flora salt marsh. Estuaries 4:97-109.

and J. W. Barko. 1978. of sediment salinity and nutrients on the physiological ecolosalt marsh plants. gy of selected Estuarine Coastal Sci . Mar. 7:487495.

J. W. B. C. W. Sikora, P., Sikora, 1977. Erkenbrecher, and B. C. Coull. and of carbon, ATP, Significance caloric content of meiobenthic nemain todes partitioning benthic biomass. Mar. Biol 44:7-14.
.

Smith, C. J., R. 0. DeLaune, and W. H. 1982a. Nitrate reducPatrick, Jr. tion in Spartina a1 terni flora marsh soil. Soil Sci. Soc. Am. J. 46:748750.

The ecology of W. B. 1977. Sikora, Paleomonetes pugio in a southeastern salt marsh ecosystem with particular emphasis on production and trophic Ph.D. Thesis. Univerrelationships. Columbia. Carolina, sity of South 122 pp.
1982. Sikora, W. B., and J. P. Sikora. Ecological implications of the vertical distribution of meiofauna in salt marsh sediments. Pages 269-282 i_n V. Estuarine compariS. Kennedy, ed. sons. Academic Press, New York. 709

Smith, C. J., R. D. DeLaune, and W. H. 1982b. Carbon and Jr. Patrick, in a Spartina nitrogen cycling al terni flora salt marsh. Pages 97-104 jji J- R- Freney, and I. E. The cycling of carGalvally, eds. bon, nitrogen, sulfur and phosphorus in terrestrial and ecoaquatic systems. Springer-Verl ag. New York.
T. III. 1982. HerbivoreSnith, J., induced changes in salt marsh plant community structure. Pages 131-149 in B. R. E. Turner, R. G. Gopal and D. F. eds. Wetzel, Whigham, Wetlands and ecology management. National Institute of Ecology and International Science Publications, Jaipur, India. 514 pp.
,

pp.

1983. Water budget, benthoSklar, F. characterization, and simulalogical tion of aquatic material flows in a Louisiana freshwater Ph.D. swamp. Louisiana State University, Thesis. Baton Rouge.

M. H. 1908. Spaulding, Preliminary report on the life history and habits of the "Lake shrimp" (Pennaeus Gulf Biol. setiferus ) Stn., Bui 1 11. Cameron, La.
. , .

Smalley, invertebrate populations, Littorina and Orchel ium f idicinium , in i rrorata the energy flow of a salt marsh Ph.D. Thesis. University ecosysten. of Georgia, Athens.
112

A.

E.

1958.

The

role

of

two

A comparison of 1975. Spiller, S. F. wildlife abundance between areas influenced weirs and control by areas. M.S. Thesis. Louisiana State 94 Baton University, Rouge. pp.
St.

Anant,

L.

S.

1959.

Louisiana wild-

life inventory and management plan. Louisiana Wildlife and Fisheries New Orleans. 329 Commission, pp.
Steever, E. 7.., R. S. Warren, and W. A. Tidal energy subsidy 1976. Niering. and of standing crop production al terni flora Estuarine Spartina Coastal Mar. Sci 4:473478.
. .

duction: a review. Sci 20:47-63.

.

Contrib.

Mar.

Turner,

R. E, 1977. Intertidal vegetation and comiiercial yield of penaeid Trans. Am. Fish. Soc. 105: shri'iip. 411-416.

Stowe,

C. 1972. Community structure productivity of the epiphytic algae in the Barataria bay area of Louisiana. Ph.D. Thesis. Louisiana State University, Baton Rouge. 84

W.

and

1978. Turner, R. E. Community plankton respiration in a salt marsh estuary and the importance of macrophytic leachates. Linnol. Oceanogr. 23:442451.

Turner,

pp.
1982. Diatoms epiphytic on Stowe, W. C. the emergent grass Spartina alternifl ora in a Louisiana sal t marsh, Trans Am, Microsc. Soc. 101:162-173.

A simple model R. E. 1979. of the seasonal of growth Spartina al terni flora and Spartina patens Contrib. Mar. Sci. 22:137147.
.

Turner,

Swank,

W. T. , J, W. Fitzgerald , and J. T. Ash. 1984. Microbial transformation of sulfate in forest soils. Science

223:182184.
1976. Diurnal activities of Tamisier, A. yreen-winged teal and pintail winterWildfowl 27:19-32. ing in Louisiana.

R. Costanza, and W. Scaife. Canals and wetland erosion in coastal Louisiana. Pages 73-84 iji D- F- Boesch, ed. Proceedof a conference on coastal ings erosion and wetland modification in Louisiana: causes, consequences, and U. S. Fish Wildl. options. Serv., Off. Biol. Serv., Washington. D.C. 256 pp. FWS/OBS-82/59. R.

E.,

19S2. rates

U.S.

Teal,

J. M. 1984. Tidal salt marshes of eastern North America: the ecology of the low salt marsh. U.S. Fish Wildl. Serv., Office Biol. Serv. Washington, D.C. FWS/OBS- (in press).

1972. Charles River Army Engineers. Mass. New watershed, England 65 Mass. Division, Waltham, pp.
S.

U.

Teal,

and J. W. Kanwisher. 1966. transport in the marsh grass, J. Exp. Bot. Spartina al terni flora 17:355-361.
J.

M.,

Gas

1974. Army Engineers. Pages 11-19 to 11-36 in draft environmental statement, deep draft access to the ports of New Orleans and Baton Rouge, Louisiana. New Orleans District.

U.

S.

The 1983. Thompson, B., and L. Deegan. River delta: a new Atchafalaya fishery nursery, with recommendations for management. Pages 217-239 jjn Proc. 10th Annu. Conf. Wetlands Restoration Creation.

A habitat 1980. Army Engineers. evaluation for water system (HES) resources Lower planning. Mississippi Valley Division, Vicksburg. Miss.

U.

S.

Department of Agriculture. No date. Gulf coast wetlands handbook. Soil Conservation Service, Alexandria, La.
Fish

Thornthwaite,
1955. tions
in

C.

The

Institute
phia, Pa.
Turner, R. tions
E.

and J. R. Mather. balance. PublicaDrexel Climatology 8(1). of PhiladelTechnology, 86 pp.

W.,

U.

S.

and

water

Habitat manual.

Wildlife Service. 1980. evaluation proceduress (HEP) Off. Biol. Serv., Washing-

ton, D.C.

U.S.
in

1976. Geographic variasalt marsh macrophyte pro113

1978. Water Geological Survey. resources data for Louisiana. Water Vol. 3. Coastal Louisiyear 1978.

ana.

Water Resources Division, Baton Surv. WaterU.S. Geol Rouge, La. Data Rep. LA-78-3.
.

Plant succession 1977. Valentine, J. M. after saw-grass mortality in southwestern Louisiana, Proc. Conf. Game Southeast. Assoc. Fish Comm. 30:534-540.
R. I., Valiela, Howes, R. Howarth, A. Giblin, K. Foreman, J. Teal, and J. The regulation of 1982. E. Hobbie. production and decomposition primary a in salt marsh ecosystem. Pages 151-168 i_n B. Gopal R. E. Turner, R. and D. F. Whigham, eds. G. Wetzel Wetlands and ecology inanagement. Institute of Ecology and National Science Publications, International 514 pp. Jaipur, India.
, ,

water marshes for wildlife. Pages 267-284 j_nR. E. Good, D. F. Whigham, and R. L. Simpson, eds. Freshwater wetlands: and ecological processes Academic potential. management 378 pp. Press, New York.
Wells, J. T., Coleman.

Chinburg, and J. M. Development of the River deltas: Atchafalaya generic Coastal Studies Institute, analysis. Center for Wetland Resources, Louisiana State University, Baton Rouge. for U.S. Prepared Anny Engineers Waterways Experiinent Station, Vicks91 pp. burg. Miss.
S. J.

1982.

Wenne,
in

C.

E.

1981.

Resource partitioning

the salt marsh fish community. Ph.D. Boston Thesis. University, 125 pp. Boston, Mass.
1981. White, D. A., and J. M. Trapani Factors influencing disappearance of a1 terni flora from litterSpartina bags. Ecology 63:242-245.
.

LI 1983. Deltaic sediVan Heerden, nentation in eastern Atchafalaya bay, Louisiana Louisiana. Ph.D. Thesis. State University, Baton Rouge.
I
. .

Van

Heerden,
1980.

I.

LI.,

and

H.

The

progradation retreating Louisiana).

225-240.

Atchafalaya a along traditionally coast (south-central Z. Geomorph. N.F. 34:

Roberts. delta: rapid

H.

White,

D. A., T. E. Weiss, J. M. Trapani, and L. B. Thien. 1978. Productivity and decomposition of the dominant

salt marsh plants Ecology 59:751-759.

in

Louisiana.

1927. Flood control in the Viosca, P. Mississippi valley and its relation to Louisiana fisheries. Trans. Am. Fish. Soc. 57:49-51.

R. J. White, D. C, Livingston, R. J. S. Nickels. 1979. Bobbie, and J. Effects of surface composition, water column chemistry, and time of exposure on the of the composition detrital microflora and associated

Vora,

S.

M.

1974.

Optimization

natural

Louisiana Rouge.

Ph.D. systems. State University,

of Thesis.

Baton

macrofauna in Apalachicola Bay, Pages 83-116 j_n R. .]. Livingston, Ecological processes in coastal marine systems. Plenum Press, 548 pp. York.

Fla. ed. and

New

Walker,
a

1940. Biotic succession in R. E. coastal salt marsh. Proc. Okla. Acad. Sci. 20:95-97.

Wax,

C. L., M. J. Borengasser, and R. A, Barataria basin: Muller. 1978. synoptic weather types and environCenter for Wetland mental response. Louisiana State UniverResources, Sea Grant Publ. sity, Baton Rouge. LSU-T-78-Q01.

1980. Wicker, K. M. Mississippi deltaic plain region ecological characterizaA a habitat mapping study. tion: user's guide to the habitat maps. Fish Wildl. Serv., Off. Biol. U.S. D.C. Serv., FWS/OBSWashington,

79/07.

Weller, M. W.

1978.

Management of fresh114

The K et al 1980a. M. Wicker, deltaic plain region Mississippi habitat mapping study:habitat area data tapes. U.S. Fish Wildl. Serv.,
.

Off.

Biol.

Serv.

Washington,

D.C,

Wiebe,

W.

FWS/OBS-79/07.
The M., Wicker, deltaic plain region Mississippi 464 maps. habitat mapping study. Fish Wildl. Serv., Off. Biol. U.S. n.C. FWS/OBSServ., Washington, 79/07.
K.

^1icroorganis;ns
witii

et

al

1980b,

sea: 1st.

R. 1972. Pomeroy. and their association aggregates and detritus in the a Mem. study. microscopic Ital. 29 Idrobiol. (suppl.):

J.,

and

L.

325-352.
and F. C. Evans. 1964. Wiegert, R. G. Primary production and the disappearance of dead vegetation in an old field in southwestern Michigan. 45:49-63. Ecology
,

Wicker,

Castille, D. .]. Davis, Gagliano, D. W. Roberts, S. D. Sabin, and R. A. Weinstein. 1982. St. Bernard parish: a study in Coastal Environwetland management. ments, Inc., Baton Rouge, La.
K.

M.,

G.

C.

S.

M.

1972. Williams, R. 8., and M. B. Murdoch. Conpartmental analysis of the production of Juncus roemerianus in a North Carol ina sal t marsh. Chesapeake Sci 13:69-79.
1960. J. Sperry, and D. Huss. Wright, R. Vegetation type mapping studies of southeastern marshes of the Louisiana. Texas A and M Research Fd., College Station. Project 191. Prepared for U.S. Bur. Sport Fish. Contract Inter. Wildl., U.S. Dep. 42 pp. 14-16-003-538.
,

Wicker,

D. Davis, and D. Roberts. Wildlife State Rockefeller and Game Preserve: evaluation Refuge of wetland techniques. management Raton Coastal Inc., Environments, La. for Coastal Prepared Rouge, Management Section, Louisiana DepartBaton of Natural ment Resources,
K.

M.

1983.

Rouge.

115

Plant species composition of salinity zones in the Louisiana coastal Appendix 1. marshes (Chabreck 1972). Scientific names conform with the National List of Scientific Plant Names (Soil Conservation Service 1982).

Appendix

1.

Concluded.

Appendix marshes .

2.

Marsh plant

decomposition rates,

Mississippi

River delta

Species
Distichli

Month initiated
s

Loss rate

Comment

Citation

(mg/g/day)

spicata
5-mm mesh bags on marsh
3

Open plots in marsh

5-mm mesh bags on marsh

Open plots in marsh

3 4

Open plots in marsh

Open plots in marsh

5-mm mesh bags on marsh

2-mm mesh bags in bayou

2-mm mesh bags, streamside marsh

2-mm mesh bags, inland marsh

5-mm mesh bags on marsh

Open

pi

ots

in

marsh

Open

pi

ots on marsh

(Continued)

Appendix

2.

Concluded.

Species

Month initiated

Loss rate

Comment

Citation

Spartina patens June

Summer Winter

4.6
11.9 9.1

5-mm mesh bags on marsh

Open plots in marsh

June
Ci

2.8-3.0

2-mm mesh bags on marsh

tations:
-

2 3

White and Trapani 1982 Kirby 1971 White et al 1978

.

4
5

Hopkinson et al. 1978 Cramer and Day 1980

119

Appendix 3. Fishes of the Mississippi River Deltaic Plain that are found in marshes and associated water bodies (compiled by Gosselink et al 1979; Deegan and Thompson 1984; see Scientific and common names conform to Robins et these documents for original sources).
,

al.

(1980),

Ecological

affinity

Trophic relations

Local

distribution

Relative and seasonal abundance

Economic imoortance

FAMILY DASYATIDAE STINGRAYS

Dasyatis sabina (Lesueur)

Atlantic Stingray
FAMILY LEPISOSTEIDAE GARS

Carnivore; predator
on meiofauna

Broadly euryhallne; to freshwater; widespread

Abundant, especially in open bay areas, larger canal s

None

Lepisosteus oculatus (Winchell)

Spotted Gar

Carnivore; predator/ scavenger on fishes, macroinvertebrates

Fresh to brackish areas, principally in protected areas; swamps, bayous, canals

Locally abundant, esoecially

in fresh swamps, hayous, canal s

Limited value as cofmercial fish tranmel nets) ; much less important than other gars
(

lepisosteus osseus (Linnaeus)

Longnose Gar

Carnivore; predator
on fishes, macroand micro-fauna

Broadly euryhaline; widespread hut mainly in freshwater areas, rivers, canals, akes
, 1

Moderately abundant in akes rivers, canal s,

1

Minor value as coimerclal fish (tratmel nets)

Lepisosteus

spatul a

Lacepede

All igator Gar

Carnivore; predator/ scavenger on fishes, larger invertebrates

See longnose gar entry; less rheophil ic than L. osseus

Moderately abundant in upper bays, canals, lakes, bayous

Moderate value as
fish (trammel nets) (most important of gars)
coiTiTiercial

FAMILY AMIIDAE BOWFINS

Atnia

caW a

(Linnaeus)

Bov/f in

Carnivore; predator/ scavenger on fishes, amphibians, macroinvertebrates

Fresh to si ightly brackish ared^ only; mainly In quiet water, swamps, canals, ditches, bayous, fresh lakes

Locally abundant

Limited value as gamefish

FAMILY ELOPIOAE TARPONS

Elops saurus (Linnaeus)

Ladyfish

Adults

Carnivore; predator fishes, invertebrates, zooplankton

on small

Pelagic; mainly in high salinity areas; lower passes

Locally abundant

None

Ladyflsh

Young

Same as adu1 ts

Pelagic; broadly euryhaline; to fresh areas; larvae and juveniles widespread in inland open-water areas

Moderately abundant along marsh edges, AprilJune

None

FAMILY ANGUILLIOAE FRESHWATER EELS

Angullla rostrata (Lesueur)

Anerican Eel
-

HA

on fishes

Adults

Carnivorous; predators macroInvertebrates

,

Demersal broadly euryhaline but mainly In brackish to fresh areas except during spawning migration; river channel, upper bay, larger bayous
;

Sparse; very cryptic; occasionally taken in trawls,

seines

hook and

ine

Anerican Eel

Young

Planktonic larvae mainly offshore; demersal elvers widespread in bays, bayous , lakes

Sparse; very cryotic; occasionally taken by trawls,

seines

FAMILY CLUPEIDAE HERRINGS

Alosa chrysochlorls (Rafinesque) Skipjack Herring

-

Adults

Carnivore; predator invertebrates, -forage species

on fishes,

Broadly euryhaline, hut mainly in fresher areas; river channels, upper

bays,
fresh lakes

Very cyclic; year-class strengths seem to fluctuate radically; can be moderately abundant in some years
See above entry; in "good" years larvae moderately abundant April - July; juveniles moderately abundant June - October

Limited value as baltfish (dipcrawfish ines) traps

1 ,

Skipjack Herring

Young

-forage species

Platonic larvae mainly in rivers

None

(Continued)

120

Appendix

3.

Continued,

Ecological affinity

Trophic relations

Local

distribution

Relative and seasonal abundance

Economic importance

Brevoortia patronus Goode

Gulf Menhaden

Fil ter feeder on plankton, suspended benthic algae, and detritus

Euryhaline; juveniles
found from fresh to
sal ine marshes

Very abundant

Dorosoma ceped an jn (Lesueur)

i

fil ter Omni vore: feeder of plankton

Broadly euryhaline, hut

Tiainly in fresher areas,

bundant, locally

*\Dderflte v-ilue

in

Gizzard Shad

Adults

detritus, benthic algae

where very widespread

soring dipnet fishery for bait,

troutl ines
,

and

crawfish traps
Gizzard Shad
-

Young

-forage species

Planktonic larvae mainly in rivers

Larvae abundant late March - June; juveniles moderately ahundant June - October
Same as gizzard shad

None

Dorosoma petenense (Gunther)

Onnivore; strainer of plankton, detritus* benthic algae

Same as gizzard shad

Limited value as

haitfish

Threadfin Shad Threadfin Shad

Adults
Young

-forage species -forage species

Same as gizzard shad

Larvae abundant May September; juveniles abundant June - November

FAMILY ENGRAULIDAE ANCHOVIES

Anchoa mitchill

(Valenciennes)
-

Bay Anchovy

Adults

Carnivore; predator invertebrates -forage species

on fishes,

Pelagic; broadly euryhal ine to fresh water; widespread

Abundant;
in

in

increasingly so ijsually taken seines, trawls, cast-nets

suiTimer;

Mone

Bay Anchovy

Young

-forage species

Planktonic larvae widespread; juveniles as adults

JVbundant year-round, oeak usually in early sumner

None

FAMILY CYPRINIDAE MINNOWS AND CARPS

Cyprinus carpio Linnaeus

Carp

Oinivore; grazer/ sucker-type feeder on plants, benthic inyertebrates,. detritus, carrion Onnivore; midwater and surface grazer/predator on zoopl ankton, filamentous algae, periphyton, fouling invertebrates -forage species

Fresh to brackish areas; widespread larvae planktonic; post larvae and juveniles mainly in temporarily flooded areas
,

Moderately abundant fresh areas; young ate March abundant

1

Minor co'iponent of freshwater hoopnet fisherv

through

suriTier

NoteiTii _qonus

crysoleucas (Mitchill)

Fresh to brackish areas; widespread

Locally abundant

Golden Shiner

"Jone; fthose sold as bait brought in fron ni nnow farms outside the

area)

FAMILY

liJALDRIDAE BULLHEAD CATFISHES

ftrinivore; mainly carnivorous; predator/ grazer on fishes macro-invertebrates carrion
, ,

Ictalurus furcatus (Lesueur)

Blue Catfish
-

Adults

Fresh to moderate sal ini ty areas; mainly in fresh and brackish

Abundant; often taken in trawls, coimercial nets , hook and 1 ine

f^opular Ta'iefish

major conponent
of inland troutline, hoopnet , trammel net catches; used in local fish culture

areas; river channel bayous, uoper bay, Tiarsh lakes

Blue Catfish

Young

Omnivore; sifnilar to adults but using more insect larvae, smaller invertebrates, detritus Omnivore, predator/ grazer on benthic invertebrates carrion, detritus
,

Essential ly as adul ts but preferring fresh areas; river channel

Locally abundant; see habitat entry

None

Ictalurus natal is (Lesueur)

Yellow Bullhead

Fresh to si ightly brackish; swanps, bayous, canals, ditches

Locally abundant, esnecially in small canals, ditches, swamps

Ictalurus punctatus (Rafinesque)

Channel

See blue catfish entry

Catfish

Adults

See blue catfish entry; this species si ightly less salt-tolerant and tends to prefer quieter water areas than K

See blue catfish entry; tends to predominate in fresher areas

furcatus
(Continued)

See blue catfish entry; this snecies tends ti oredQininate in *"resher areas and laore henthic situations

121

Appendix

3.

Continued,

Ecological af f ini ty

Trophic relations

Local

distribution

Relative and seasonal abundance

Economic
imooT-tance

Pylodictis

ol

ivaris (Rafmesque)

Flathead Carfish

Carnivore; predator on f isnes macroinvertebrates

,

Fresh to brackish areas; nainly in river channel

Sparse

^opul ar qamefish; minov

cnmponpnt n*" inland hoopnet

and
trotl ine

catch
FAMILY ARIIDAc SEA CATFISHES
Aru|_s

fe_Hs (Linnaeus)

Hardhead Catfish

Onnivore; grazer/ scavenger on carrion, detritus, macro- and meio-benthos

ESM

Broadly euryhaline, hut mainly in high to moderate salinity areas;

Local ly abundant , -nainly during warm months

fi-in% of

industrial bottomfish catch

^sqre marinus (Mitchill)

Gafftopsail Catfish

Omnivore; grazer/ scavenger on carrion, detritus, macro- and meio-henthos

To moderate sal ini ty areas; -nainly limited to high sal ini ty; lower bays, passes

Sparse; found in and around marshes in warn months only

Minor coaponent of bottomfish catch; not distinguished fron Sea Catfish

FW\U

GOHIESOCIDAE CLINGFISHES
Carnivore; feeds on macro- and meiobenthos
High to moderate salinity areas; mainly near reefs, pilings, letties
Sparse; occasionally taken in trawls, dredges; larvae in plankton near reefs, late winter, spring
None

Gobiesox strunosus Cope

Skilletfish

FAMILY HELONIOAE

NEEDLEFISHES
Strongylura marina (Walbaum)
Atlantic Needlefish
ESM

(jn

Carnivore; predator fishes , macroinvertebrates

Broadly euryhaline; to freshwater; widespread

Moderately abundant but seldom concentrated; often taken in seine, castnets

FAMILY CYPRINOOONTIDAE KILLFISHES

Adinia xenica (Jordan and Gilbert)

Diamond Ki!
1

if ish

Omnivore; mainly herbivorous; grazer on algae, periphyton, detritus

Broadly euryhal ine; to freshwater, but mainly in high to moderate salinities; mainly along edges of protected areas (marshes); ponds, ditches, canals Broadly euryhaline; widespread along shores and in protected marsh waters

Locally abundant, esnecially in winter and spring

Cyprinodon variegatus Lacepede

Sheepshead Minnow

ftnnivore; primarily

herbivorous; grazer on algae detri tus benthic invertebrates periphyton -forage species
, , ,

Abundant, peaks observed in winter and soring

Minor value as haitfish

Fundulus chrysotus (Gunther)

Golden Topninnow

-forage species

Fresh to si ightly brackish areas; mainly in fresh swamps, ditches, canal s, borrow pits
See sheepshead minnow entry

Locally abundant; esnecially quiet marshy areas

Fundulus 'jrandi

Baird and Gi ra rd

Gulf Killifish

Omnivore; mainly carnivorous; predator/ grazer on small invertebrates, fishes, detritus -forage species

See sheepshead

ninnow entrv

Minor value as haitfish

Fu ndulus

Jenkins

(Evennann)

-forage species

Broadly euryhaline; in protected marsh areas

^are, occasionally seined in marsh ditches, ponds

Saltmarsh Topininnow

Fundulus pul vereus (Evennann)

Bayou
Ki
1
1

ES

if ish

Carnivore; predator/ grazer on smal invertebrates

1

Broadly euryhaline; in protected marsh areas; bayous, canals, ditches, ponds

Broadly euryhaline but greatest concentrations in moderate to high salinities; along beaches, edges of marsh lakes, bayous
(Continued)

Locally abundant, winter through spring

Fundulus simil

Bai rd

and Gi rard)

ES

Longnose

Kil

ifish

Omnivore; predator/ grazer on benthic invertebrates , detritus

Locally abundant; lower bavs, high salinity marshes

122

Appendix

3.

Continued.

Ecoloyical affinity

rel

Trophic ations

Local

distribution

Relative and seasonal abundance

economic
rtport-incf*

Lucam

parva (Baird)

"Rainwa'ttjr Kill if ish

Onnivore; priinarily carnivorous; predator/ qrdier on invertebrates, detritus -forage species

Sa^ne as

sheepshead ininnnw

Locally -jhijndant, nea-:s

SuniTier

in

FAfllLr

^'0^:CILIIDA

LlVtRBEAREtlS

Gdmbu s_i_d af f im

(Baird and Gir.jrd)

Mosqji tof isn

Onnivore; primarily carnivorojs; predator/ grazer on invertebrates -forage species

Broadly euryhaline,
Tiainly
in

^>st

fresh to

Loral y abundant, in fresh areas

1

brackisH areas; along edges of protected areas; swanps, marshes, canals, ditches, bayo'js, ponds
Fresh and brackish areas only; swanps, ditches, borrow pits; usually in larshy areas Rare; occasionally \a^en in di tchps borrow oits
,

Heterandria fornosa Agassi z

Least Kill ifish

Herbivore; -jrdZBr on hentb ic ep ip^ytes algae -forage species

,

Poecil ia

atipinna (Lesueur)

Sailfin Molly

Herbivore; -jrazer on epiphytes, benthic algae, detritus

Rroadly euryhaline to freshwater; widespread along protected shores, open beaches, bayous, ditches, canals, nonds

Locally abundant vear-round

FAMILY ATHERINIUAE

SILVERSIDES
Labidesthes sicculus
Brook
(

Cope

on

Silverside

Carnivore; predator neustomc invertebrates, zooplankton -forage soecies

Carnivore, predator invertebrates -forage species

small

Fresh areas onl streais

Locally ^hundant areas

in

fro^h

Membr a^ martinica (Valenciennes) Rough

Si
1

on small

vers ide

Broadly euryhaline; to freshwater; ^nainly along marshy shores of bays, lakes, lar^jp ranals, bayous
Broadly euryhal ^iread
i

t.ocally abundant during suimier

Memd

beryl

l_i_n_a

(Cope)

Inland Silverside

Carnivore, predator/ grazer on zooplanktnn other smal invertebrates snecies -forage

1

ne

wide-

Abundant,

riealrs

in

sum^ier

FAMILY SYNGNATHIDAE PIPEFISHES AND SEAHORSES

Synqnathus louisianae Gunther

on small

Carnivore; predator invertebrates

Cham Pipefish

High to noderate salinity areas; nainly associated with vegetation

Rare; occasionally taken by seines in higher salinity

narsh nonds, ditchps

Syngnathus scovel

(Evermann and Kendall)

on small

Carnivore; predator invertebrates

Gulf Pipefish

Broadly euryhaline; to freshwater; widespread along edges and areas having dense vegetation; ditches, canals, nonds

Local

abundant

FAMILY PERCICHTHYIDAE TEMPERATE BASSES

Mo rone chrysops (Raf inesgue)

Carnivore, predator :iai nly on fishes

White Bass

"broadly euryhaline but nainly in fresh and brackish areas; pelagic in open waters of river channel, large bayous, canals, lakes, upper hays

Lor^l
in

y abundant fresher areas

1

"i nor val UP ga ipf isb

'i'

Mo rone mississippiensis Jordan and Eigeninann Yellow Bass

Carnivore; predator mainly on fishes

See white bass entry; this fonn slightly "tore salt tolerant and nore common in snail er water

Locally ibundant; nainly in fresh areas , f"! vpr channel , swa^ips

Minor valije as gaief ish

bodi es Mo rone saxatil is (Walbaum)

Larmvore; voracious
predator on smal
1

Mai nly

in

nl

and waters

Striped Bass

fisb

Rare; occasionally caudbt ^v book -ind traiTinel nets

Lviited valyo IS gamp^is^

(Continued)

123

Appendix

3.

Continued.

Ecological af f ini ty
FAMILY CENTRARCHIDAE

Trophic relations

Local

distribution

Relative and seasonal abundance

ficonoinic

imoortance

SUNFISHES

Centrarchus macropterus (Lacepede)

Flier

Carnivore; predator on fishes macrosinal invertebrates

1 ,

Fresh to si ightly brackish areas; swamps, marshes, hayous, sluggish streams

Fresh to brackish areas; backwaters of streams, swamps, ditches, canals

Sparse

Limited value as qamefish

Leponis cyanel lus Rafinesque

Green Sunfish

FW

Carnivore; predator on fishes macroinvertebrates

,

Sparse

None

Lepomis gulosus (Cuvier)

Wa mouth

on fishes

Carnivore; predator , macroinvertebrates

Omnivore; predator/ grazer on invertebrates, algae

Fresh to brackish areas; swamps, borrow pits, canals, bayous Fresh to brackish areas; widespread in fresh habitats

Local ly abundant;

especially in swamps

Minor value as gamef ish

Lepomis macrochirus Rafinesque

Sluegill

Locally abundant

Minor value as qanefish

Lepomis marginatus (Holbrook) Dollar Sunfish

Lepomis megalotis (Rafinesque)

Fresh to brackish areas; especially swamps, borrow pits

Locally abundant fresh areas

in

Longear Sunfish

Carnivore; predator/ qrdzer on invertebrates, especially insects

Omnivore; primarily carnivorous; predator/ grazer on invertebrates mainly mollusks
,

Fresh areas only; mainly n rivers , creeks

i

Sparse

None

Lepomis microlophus (Gunther)

mainly

Redear Sunfish

Fresh to brackish areas; in swamps, borrow pits, canals, bayous, lakes

Fresh areas only; mainly in swamps

Moderately abundant in fresh lakes, ponds, borrow pits

Minor value as gamef ish

Lepomis punctatus (Valenciennes)

Spotted Sunfish

See redear sunfish

entry

Minor value as gamef ish

Lepomis symmetricus Forbes

Bantam Sunfish

Fresh to brackish areas; common in swamps, borrow pits, ditches

Local ly abundant

Micropterus salmoides (Lacepede)

Largemouth Bass
-

Adults

Carnivore; predator mainly on fishes, macroinvertebrates

Fresh to brackish; widespread in lentic situations, especial ly in areas of low turbidity

Abundant in lentic habitats, sluggish streams, canals, bayous

pQDular qamefish; large quantities caunht in marsh impoundponds ments

,

Largefnouth Bass

Toung

Carnivore; predator
on zooplankton, later Insects, small fishes

Minimally in fresh areas; shallow marginal zones of swamps, stream backwaters

Fresh to brackish; widespread In low turbid lentic situations

Moderately abundant In lentk freshwater areas, April through summer

Pomox 1

nigromacuTatus (Lesueur)
Black Crapple

FH

Carnivore; predator
on fishes, macro-

Invertebrates; larvae feed an zooplankton

Moderately abundant in fresh areas, esoeclally quiet, weedy areas

Popular gameflsh

FAMILY CARANGIDAE JACKS

Oligopl

Ues saurus (Schneider)

Leatherjacket
-

ESM

on small

Young

Carnivore; predator fishes. Invertebrates

Broadly euryhal Ine; to freshwater, but mainly moderate to high salinity areas; bay shores, bayous, marsh lakes

Moderately abundant during warn months

FAMILY GERREIDAE

HOJARRAS

Eucinostanus argenteus Balrd

Spotfln Mojarra
-

E94

Young

Carnivore; predator/ grazer on benthlc invertebrates

Broadly euryhallne, hut mainly In moderate to high salinities; widespread

Moderately ahundant In shore seines during warm months

FAt4ILY

SPARIDAE PORGIES
E9^

Archosarqus probatocephalus (Walbaum)

Sheepshead
-

Adults

Omnivore; grazer/ predator on perlphyton, macroinvertebrates.

Mainly In high salinity areas, lower bays, tidal passes; near pll Inqs, reefs

Moderately abundant, year-round; often taken by anglers, trairmel nets

Minor value as comnercial fish (trarmel net); popular gameflsh

(Continued)

124

Appendix

3.

Continued-

Ecological affinity

Trophic relations

Local

dtstributlon

Relative and seasonal abundance

Economic importance

Sheepshead

Young

Broadly euryhaline; widespread In protected waters, marsh bayous, canals lakes

,

ftoderately abundant, mainly

spring, early sunmer

Lagodon rhoinboides (Linnaeus) Pinfish

-

Adults

Oinnivore; predator/ grazer on fishes , detritus, invertebrates, algae

Broadly eurytiallne, but Moderately abundant, mainly in high to moderate especially during warm months salinity areas; lower bays, bayous
Broadly eurytiallne; to freshwater; widespread along shores and in marsh bayous, ditches, ponds
Abundant, late winter through suntner

Pinfish

Young

FAMILY SCIAENIOAE DRUMS

Aplodinotus grunniens Raf Inesque Freshwater Dru

-

Adults

Carnivore; predator/ grazer on benthic Invertebrates espe,

Fresh to brackish areas; especially river channel

Locally abundant yearround

Major component
of Inland hoopnet catch;

cial ly mol lusks

and

minor gamefish

fishes

Freshwater Drun

Young

Chinlvore; larvae predators on zooplankton; juveniles grazers on benthtc invertebrates, detritus

larvae planktonic In river, upper bays, demersal, especially over soft mud/detritus bottoms

Locally abundant, Hay through early fal

1

Bairdlella chrysoura (Lacepede)

Silver Perch

ESM

Carnivore; adults predatory on small fishes, benthic invertebrates

Broadly euryhaline but nainly 1n inoderate to high salinity; widespread

Locally abundant especially as postlarval and early juveniles, April through early sunmer
,

Cynosc 1 on arenarlus Ginsburg

on fishes

Sand Seatrout

Adults

Carnivore; predator macroinvertebrates

,

Moderate to high salinity areas; widespread In bays, marsh lakes, bayous

Moderately abundant, declining In cold months

Ponular gamefish; minor comnonent of inland

tranmel

net

catch
Sand Seatrout
-

Young

Broadly euryhaline; widespread; very smal juveniles prefer protected marsh waters
1

Abundant, April early Fall

through

None

Cynosclon nebulosus (Cuvler)

Spotted Seatrout

ESM

Carnivore; predator on fishes and macroInvertebrates

Abundant schooling fish Abundant year-round, except In saline and brackish winter areas, often found in marsh bayous and shallow lakes, especially juveniles

Popular sportfish

Leiostomus xanthurus Lacepede

Spot
-

Young

Oinnivore; primarily carnivorous on zooplankton; grazer on detritus

Broadly euryhaline, but mainly In moderate to high salinity areas; postlarvae and early juveniles mainly in protected marsh waters; older juveniles widespread
Adults move offshore in fall

Abundant, especially late spring through sifimer

5-71 of industrial bototmfish catch in spring and sumner; moderately valuable as qamefish

Spot

Adul ts

Graze on benthic invertebrates and detri tus

ESM

MIcropogonUs undulatus {Linnaeus)

Atlantic Croalter

Onnivores; grazers on benthic Invertebrates detritus, small fishes; young subsist on zooplankton

Euryhaline, preferring salinity areas around marshes as juveniles, moving to saline areas wl th maturity

Very abundant; moving offshore in winter

More than S of

Industrial hottomflsh catch

Poqonlas cromis (Linnaeus)

ESM

Black Drun

Adults

Carnivore predator/ grazer on benthic invertebrates, especially blvalue tnol lusks

Broadly euryhaline, but mainly in high to moderate

lower passes; mainly near reefs
sal tnity areas;

Moderately abundant, often taken by tranmel nets, hook

and line

Same value as sportfish and and cornnerclal fish

Black Orim

Young

Predatory on small benthic Invertebrates

Larvae mainly In offshore areas; postlarvae and Juveniles occasionally entering bays lower marshes
,

Sparse; occasionally taken in seines

None

(Continued)

125

Appendix

3.

Continued,

Ecological affinity

Trophic relations

Local

distribution

Relative and seasonal abundance

Economic importance

ScUenops occl latus (Linnaeus)

Red Drun

Carnivores; predators
on fishes and crustaceans

Widespread in saMne and brackish areas, often In shal low marsh, ponds, and streams
Mainly In high salinity areas; lower bays, passes

Abundant especially In fall and early winter

Valuable gamefish

Stel

tfer lanceoUtus (Holbrook) Star

DrijTi

Sparse; occasionally taken In trawls

FAMILY EPHIPPIDAE

SPADEflSHES Chaetodipterus faber

Atlantic Spadefish
fAJ^JLY

ttnnivore; grazer on attached algae,

Mainly In high sal Inlty areas, near tidal passes

Moderately abundant, locally, especially during sunrier and fall

Young

foul log

Invertebrates

MUCILIOAE MULLETS
Onni vore; primarl ly herbivorous; -forage species

Mu^il cephalus Linnaeus

Broadly euryhaline; to freshwater;

Abundant

v^ar-round

Striped **]llet
Striped HjI let

Adults

Young

Onnlvore; primarily herbivorous

Broadly euryhaline; to freshwater; widespread; planktonic larvae offshore

Abundant, especially late winter, early spring

None

FAMILY ELEOTRIOAE

SLEEPERS

Dormitator maculatus (Bloch)

fat Sleeper
El

ES

on fishes

Carnivore; predator macroInvertebrates

,

Broadly eurytial ine; mainly In ditches,

canals, bayous

Moderately abundant,
local ly

eotrls pisonis (Gmel

In)

ES

Same as fat sleeper

Splnycheek Sleeper
FAMILY GOBIIDAE

Broadly euryhaline; hut mainly In fresh or brackish areas; canals, ditches

Very rare

GOBIES
Evorthodus lyricus (Girard)
Lyre Goby
ES

Broadly euryhal Ine; but mainly in moderate to high sal Inl ty areas; marsh dl tches , canal s
,

Local ly abundant

ponds

Gobioides broussoneti Lacepede

Violet Goby

ES

Broadly euryhaline; but mainly In high salinity areas; open bays, bayous, marsh lakes
Broadly euryhal Ine; widespread

Sparse; occasionally taken In trawls

Goblonel lus boleosoma (Jordan and Gilbert) Darter Goby

ES

Local ly abundant, esfieclally during cold

months
ES

GobloncUus hastatus Girard

Sharptail Goby

Onnlvore; grazer on algae, benthlc invertebrates

Broadl y euryhal Ine; widespread

Sparse; occasionally taken In trawls

Goblonel lus

s^kj

feldtl

(Jordan and Elgenmann)

ES

Freshwater Goby

Broadly eurytial Ine, hut mainly in fresh to brackish areas, where widespread
Carnivore; predator/ scavenger on benthlc Invertebrates, carrion
Carnivore; predator/ grazer on benthlc Invertebrates

Locally abundant

Goblosawa bosci (Lacepede)

Naked Goby

ES

Broadly euryhaline, widespread

Locally abundant, on reefs, marsh ponds, ditches

Gobiosoma robustuw Glnsburg

Code Goby

ES

Broadly euryhaline. hut mainly In moderate to high salinities; mainly associated with vegetation Broadly euryhal ine, widespread; mainly near vegetation

Sparse, occasionally taken In seines

Microqoblus gulosus (Girard)

Clown Goby

ES

Onnlvore; predator/ qraier on benthic Invertebrates, algae

Sparse; occasionally taken in trawls, seines

Mlcroqobius thalassinus {Jordan and Gilbert) Green Goby

ES

Broadly euryhaline, but mainly In high salinity areas; near vegetation

(Continued)

Very rare; occasionally taken In seines

126

Appendix

3,

Concluded.

Ecologtcal affinity

Trophic relations

Local

distribution

Relative and seasonal abundance

Economic importance

FWILY BOTHIOAE
LEFTEYE FLOUNDERS
CI

tharlchth^s macrops Dresel

Spotted Whiff

on small

Carnivore; predator crustaceans

Limited to high sal Inlty areas; lower bays, passes

Rare; occasionally taken in trawls

Paral Ichthys lethostiqma Jordan and Gilbert

E^

Southern Flounder

Carnivore; predator on small fishes, macro! nvertebrates

Euryhallne; juveniles
and adults
found
in

from
f

Fairly ahundant, esoecially during warm months

freshwater to gul

Valuable snort and connercial fish

saMnitles.

tidal

channels and shallow lakes; larvae offshore

FWILY SOLEIOAE
SOLES
Achirus
1

ineatus (Linnaeus)

Broadly euryhal Ine.

but mai nly in high to moderate sal Inlty;

Moderately abundant,
late sumner, fall

Lined Sole

widespread
Trinectes maculatus (Bloch and Schneider) Hogchoker - Adul ts
Grazer on melo- and macro-benthos detrl tus
.

Broadly euryhal Ine; to freshwater, but mainly In brackish to high sal Inity
Broadly euryhallne, but mainly In moderate to high salinity; widespread

Abundant, mainly spring and summer

None

Syrnphurus plaglusa

(Linnaeus)

BlacVcheek Tongueflsh

Carnivore; predator on benthk invertebrates

Abundant
in

mai nly

None

snring

FW MA ES ESM

= = =
=

freshwater marine estuarine estuarine-marine (migratory)

127

Representative vertebrate species of marsh habitats in the Mississippi River Appendix 4. Deltaic Plain (compiled by Mabie, 1976 and Gosselink et al. 1979; see these documents for = Scientific Fresh, I = Intermediate, B = Brackish, S = Saline). original sources) (F and common names of amphibians and reptiles conform to Collins et al. (1982); birds to American Ornithologists' Union (1983); and mammals to Jones et al (1975).
.

Species

Seasonal peaks of abundance or activi ty

AHPHIBIANS
Anbystoind opacun

Marbled salamander

Appendix

4.

Continued.

species

Food

Seasonal peaks of abundance or activity

Remarks

Pseudemjs conci nna River cooter

Pseudemys floridana Missouri si ider Pseudemys pi eta Southern painted turtle Pseudemys scrip ta Red-eared turtle
Fie

Largely aquatic vegetation

Largely aquatic vegetation

Juvenile:
Adult:

13i plant, 85% animal 881 plant, 10% animal

30% plant, 70% animal (e.g., amphipods) Juvenile: Adult: 89% plant, 11% animal (e.g., crayfish)

Deirochelys reticularia Chicken turtle

Graptemys kohni Mississippi map turtle
i

FIB

Tadpoles, crayfish, plant material

Graptemys pseudogeographica Sabine map turtle

Trionyx spini ferus Spiny sof tshel 1
Anol is carol ini ens
is

Carnivorous

Insects and spiders

Coluber constrictor Racer

Farancia abacura

FIB

Insects, frogs, snakes, young birds

Breeds: May Hatch: July- Sept.

/Vnphiunna

Si

ren , frogs

Lampropel tis getulus Speckl ed ki ng snake

Me rod
1

FIB

Other snakes, small birds, lizards, mice, rats

cycl op ion

C^mbusia (77.6%); other fish (18.6%); tadpoles

(3.5%)
i

Mar. -Oct.

Green water snake

Ne rod i a fasciata cl arki Gul f sal t marsh snake BS

Fish,

fiddler crab

fterodia fasci ata conf luens

FIB

Broad-banded water snake

Fish (86.9%); (4.3%)

Fish (92.7%); (6.1%)

frogs and toads (6.4%);

tadpoles

Mar. -Sept.

Nerodia rhombi fera Diamondback water snake

Reqina grahami i Graham's crayfish snake

frogs and toads (1.0%};

tadpoles

Mar. -Oct.

Crayfish (inOi)

f^r.-Sept.

Regina rigida Glossy crayfish snake

FIB

Si ren

fish, crayfish

Storerla dekayi Brown snake

Earthworms, snails. Insects, small

frogs, fish

Thamnophis proximus Western ribbon snake

Thamnophis sirtal is Comnon garter snake
Aqki strodon plsci vorus
FI

Insects, fish, frogs, salamanders, mice, toads

Earthworms, moUusks, insects, fish, salamanders, toads, frogs, small manmals, small birds
Fish, salamanders, frogs, reptiles, birds, mammals

FIB

Cottonmoutn

BIRDS

GREBES & WATERFOML

Podilymbus podiceps Pied-billed grebe
Podiceps nigricol Eared grebe
1

FIBS

Mostly animal:
small

frogs and fish,

aquatic worms and Insects, snails, seeds and soft parts plants:

Oct. -Apr.

is

Insects, shrimp, some water plants, feathers

Oct. -Nay

Dendrocygna bicolor Fulvous whistling-duck

Mostly seeds of grasses and weeds; also grasses, grain

(Continued)

Apr. -Sept.

129

Appendix

4.

Continued.

species

Food

Seasonal peaks of abundance or activity

Insects
Nov. -Mar.

Remarks

Anser albifrons Greater white- fronted yoose

Anas strepera Gadl^aTT
Anas americana

FIBS

Grain, tender shoots, occasional

FIBS

Principally plants

Oct. -Mar.

FIBS

90% plant. 101 animal

(from Sept. -Apr.)

Oct. -Apr.

American wujeon
Aythya col lans Ring-necked duck
Ay thy a

FIBS

19% animal: Insects, mollusks; 81% plant: plants, sedges, grasses, smartweeds

aquatic

Oct. -Apr.

aff mis Lesser scaup

FIBS

Similar to A. marl

Oct. -Apr.

Bucephala albeola bufflehead

Lophodytes cucul latus
Hoocfed inerganser

FIBS

79% animal: insects, crustaceans, mollusks, fish; 21% plant: pondweeds , misc.

Nov. -Mar.

FI

Mostly insects; also small fish, frogs, mollusks, crayfish, roots of aquatic plants, seeds, grain
72% plant: 28% animal:

Nov. -Apr.

Qxyura jamaicensis Ruddy duck

Porphyrul a martirnca Purple gal 1 inule
Ga

FIBS

aquatic plants, grasses, sedges; Insects, mollusks, crustaceans

Nov, -Apr.

Rice, other seeds, worms, mollusks

Apr. -Sept.

inula chloropus Common inoortien

1 1

Seeds, roots, soft parts of aquatic plants, snails insects, worms FIB Leaves, fronds, seeds and roots of aquatic plants; wild celery, algae; worms, snails. Insects, small fish, tadpoles
Almost wholly plants: grain, roots and culms of grasses; some insects, mol lusks Almost wholly plants: aquatic plants, marsh grasses sedges; some mollusks, crustaceans
10% animal: Insects, mollusks, crustaceans 90% plant: sedges, pondweeds and grasses (62%); other (28%)

Apr. -Nov.

Ful

Kd djnencana

Sept. -Apr.

Ainerican coot

Chen caerulescens Snow goose Branta canadensis Canada goose

Anas crecca 5reen-w1ng ed

FIBS

Oct. -Apr.

FIBS

Oct. -Feb.

FIBS
teal

Oct, -Mar.

Anas njbripes American black duck Anas ful vigula Mottled duck Anas platyrtiynchos

FIBS

Mast, grain, mollusks, crustaceans

Oct. -Mar.

FIBS

40% animal: mollusks. Insects, crayfish, small fish; 60% plant: inostly grasses (plants and seeds)

FIBS

Mallard

90% plant: sedges, grasses, smartweeds, pondweeds, duckweeds, tubers, mast; 10% animal: Insects, crustaceans, mollusks, fish
13% animal: mollusks, crustaceans. Insects 87% plant: pondweed sedges and grasses (60%); other (27%)
,

Anas acuta

FIBS
1

Oct. -Mar.

Northern pintal

Ana_s

discors B1ue-Hlnged teal

FIBS

30% animal: worms, mollusks, insects, tadpoles 70% plant: sedges, pondweeds and grasses (43,6%); other (26.4%)

Feb. -Apr.; Sept. -Nov.

Anas c lypeatd Northern shovel er

FIBS

Animal: worms, small mollusks, insects, shrimp, small fish, small frogs. Plant: buds and young shoots of rushes and other aquatics; grasses

Oct. -Apr.

WADING 81RDS
Botdurus lentiglnosus 'Vnerican bittern
FIB

Mollusks, crayfish. Insects, small lizards, small snakes, mice

fish, frogs,

Oct. -May

"Blue List" Natl. Aud. Soc. (1976)

Ixubrychus exil is Least bittern

Ardea herodias Great blue heron
Casjnerodius albus Great egret

FIBS

Slugs, leeches, insects, small frogs, lizards, small mammals

fish, tadpoles, small

Apr. -Sept.

FIBS

Mostly fish; also crustaceans. Insects, frogs, llzarxjs, snakes, birds, small manmals
Small

Year- Round

FIBS

fish, snails, fiddlers. Insects, frogs, lizards, small snakes, mice, some plant material

(Continued)

130

Appendix

4.

Continued.

Speci es

Seasonal peaks of abundance or activity FIBS

Remarks

Egretta thuJa Snowy egret Egretta caerulea Little blue heron

Shrimp, small fish, fiddlers, snails, insects. crayfish, small lizards, small frogs, small snakes

Mar, -Oct.

BS

Crayfish, small crabs, insects, fish, frogs, lizards

Mar. -Oct.

Egretta tricolor Tricolored heron

Egretta rufescens Reddish egret

FIBS

Slugs, snails, crayfish, insects, small izards, frogs

1

fish,

Mar. -Nov.

BS

Mar, -Oct.

"Blue List" Natl. Aud. See. (1976)

Bubulcus ibis Cattle egret

Butorides striatus Green-backed heron
Small

fish, earthworms. Insects, tadpoles, snakes, small maimials

frogs.

Mar. -Oct.

Nycticorax nycticorax Black-crowned night-heron

Mostly fish (alive or dead), worms, crustaceans. Insects

Mar. -Sept.

"Blue List" Natl. Aud. Soc. (1976)

Nycticorax violaceus Yel low-crowned night heron

Eudociinus albus White ibis

FIBS

Snails, crayfish, crabs, fish, small mamnals and birds

reptiles, small

Mar. -Sept.

FBS

Mostly crayfish; also other crustaceans, slugs snails, small snakes, insects

Mar. -Sept.

"Blue List" Natl. Aud. Soc. (1976)

Plegadis falcinel lus Glossy ibis Plegadis chihi White-face3"ibis

FIBS

Insects, crayfish, young snakes

Earthworms, crayfish, mollusks, insects, small fish and frogs, newts, leeches

"Blue List" Natl. Aud. Soc. (1976) Jun.-Sept.

"Blue List" Natl. Aud. Soc. (1976)

Hycteria americana Wood stork

Fish, aquatic reptiles, insects

SHORE BIROS
Pluvial
is

Bl ac k-

squatarola bellied plove r

Marine worms, s<ndl 1 mollusks, crustaceans, insects, some plant material Worms, small mollusks, crusteans, insects

Sept. -May

Charadrius sanipalinatus Semipalmated plover

Sept. -May

Himantopus mexlcanus Black-necked stilt

FIBS

Recurvi ros tra americana American avocet

FIBS

Appendix

4.

Continued,

Species
Cal idris maurt

Food

Seasonal peaks of abundance or activity

FIBS

Insects, marine wonns, snail

snails

Auq.-f^y

Western sandpiper
Cal idris

Least sandpiper
Cal idris ba
i

minuplla
rd
i i

FIBS

Mostly insects; also small

crustaceans, wonns

AuQ.-Apr.

FIBS

Insects, amphipods, algae

Mar. -May; July-Oct.

Bdird's sandpiper
Cdl idris alpina Ounl in
Cal

FIBS

Small mollusks, small crustaceans. Insects, marine worms, occasionally seeds

Oct. -May

idru
.

FIBS

Animal

{70*):

small

worms, mol lusks,

insects

Ap f

f^ y

Stilt

Plant (301):
FIBS

seeds

Limnodroinus qriseus

Short-billed dowitcher
Limnodroinus scolopaceus Long-bil led dowitcher
Gal

Wonns, insects, fish egys, small mol lusks, seeds and roots of aquatic plants
Insect larvae, some plant material

f^r.-May; Sept. -Nov

FIBS

Oct.-^by

ina^o gal iraij o Common snipe

1
i

FIBS

Mostly earthwcjnni, also other lyonns, insects, some seeds of marsh plants
Aquatic insects and their larvae; amphipods; seeds of aquatic plants

Oct. -Apr.

Phalaropus tricolor Wilson's phalarope

FIBS

Aor.-May, July- Sept.

FISHING BIROS

Pelecanus erythrorhynchus Aflierican white pelican

Larus atrici
1

as

Sent. -May

la
1

IBS

Lauqhing qui

Hnsfly small refuse

fish;

also eqqs of other seahirds,

Year-Round

Sterna nilot ica Gull-billed tern

IBS

Oct. -Apr.

"Blue List"
Natl
.

Aurt.

Soc.

(19;M
Sterna caspia Caspian tern Sterna forsteri Forster's tern
EOS

Almost wholly small fish; also shrimp and other surface-swiiiminq aquatic life
Insects,

Year-Round

floating carrion

Year- Round

Childonias ni^er Black tern

C eryle

F!

snail

fish, insects

\)r.-Sept. (nonbreeding)
Sept .-Aor.

alcyon

Belted kingfisher

Almost wholly fish; also insects, crustaceans, mollusks, amphibians, small reptiles, birds, .nice, herries

BIROS
Ci

Of

PREY
Small mammals, herons, ducks, coots, rails. shorebirds, songbirds

reus cyaneus Northern harrier

Sept.-Aor.

"Blue List" Natl. Aud. Soc. (1976)

Falco sparser lus American kestrel

FIBS

Insects, amphibians, reptiles, bi rds

marinals

Sept. -May

Hatl

"Blue List" Aud. Soc. (19761

.

Falco coljnbarius
Merl in

Mostly birds: green-winged teal, shorebirds. small chickens, various songbirds; also insects, sniders, reptiles, mice, pocket gophers, squirrels, liats
IBS

Sept. -May

"Blue List" Natl. Aud. Soc. (19/6)

Fdlco pereqrinus Per egrTiT? rSTco n

Primarily birds, also small maminals. insects

Sept. -May

Endangered

Asio

fl

ammeus

Mostly small mammals, also small binds, ins*icts

fht.-May

"Blue List"
Natl
.

Short-e.ired owl

Aud. Soc.

(1976)

(Continued)

132

Appendix

4.

Continued,

Species

Food

Seasonal peaks of abundance or activity

OTHER HARSH BIRDS

Chordeiles minor tornmon nighthawk
Insects, mostly flying
Apr. -Oct.

"Blue List" Natl. Aud. See. (1976)

Coturniccys noveboracensis

FIBS

Oct. -Hay

Laterallus jamaicens Black rail

Ra 1
1 u s lonyirostris Clapper rail

is

FIBS

Nov. -Apr.

BS

Ra 1

us

elegans

FIB

Grass seeds,

insects, slugs, leeches, tadpoles,

Year-Round

King rail

crayfish
FIBS

Rallus

1 imicola Virginia rail

Earthworms, crayfish, insects, snails, small some grass seeds

Small mollusks,

fish,

Oct. -Apr.

Porzana carol ina Sora

Tach_ycineta bicolor Tree swallow

FIBS

insects, seeds

Sept. -May

FIBS

81% animal: insects and spiders 2\% plant: seeds and berries
Insects

Sept. -Hay

Riparia riparia Banic swallow

Hi rundo pyrr^ionota
CI

FIBS

Apr. -May; July-Oct.

IBS

Apr. -June

iff swallow

Hi

rundo rustica Barn swallow

FIBS

99% animal:

insects; some spiders and snails

Ma r .

f^ y ;

Au g . - Nov

CorvLis

ossifragus

FIBS

Fish crow

Carrion, crustaceans, fish, bird eggs, insects; berries, tree fruits, seeds, some grain
Insects, spiders

Year-Round

Cistothorus platens Sedge wren

is

FIBS

Oct. -Mar.

Cistothorus palustns Harsh wren

Anthus spinoletta Water pipit

FIBS

Insects; especially Coleoptera and Oiptera

Year-Round

FIBS

Nov .-Mar.

Geothlypis trichas Common yellowthroat

FIBS

Hostty insects,

few seeds

Mar. -Oct.

Passerculus sandwichens Savannah sparrow

Ammodrainus caudacutus

is

FIBS

92% plant: (winter)

seeds; 8% animal:

mostly insects

Oct. -Apr.

BS

Sharp-tailed sparrow

81% animal: insects, amphipods, spiders, snails 19% plant: grasses, seeds

Nov.-mr.

Ammodramus maritimus Seaside sparrow

f'telospi za
t

Marine worms, crustaceans, insects, spiders, mollusks, weed and grass seeds
5S% insects; 45% seeds

Year-Round

jeor ^ iana

FI

Sept. -Hay

Swamp sparrow

Dolichonyx oryzivorus
Bobol ink

FIBS

57% animal: insects, spiders, myriapods; 43% plant: weed seeds, grain

Hay

Agelaius phoeniceus Red-wi nged bVackbi rd

Quiscalus major Boat-tailed grackle

FIBS

73% plant: weed seeds, grain, fruit; 27% animal: mostly insects and spiders Insects, spiders, small
fish, tadpoles

Year-Round

FIBS

Year-Round

Didelphis virgin! ana Virginia opossum

FIBS

Insects, birds, carrion, plant material

Breeds

in Jan. -Feb,

(Continued)

133

Appendix

4.

Concluded.

species

Food

Seasonal peaks of abundance or activity

Remarks

Myotis austroripanus Southeastern myotis

Las lurus boreal is Red bat

Insects

Active year-round in warm weather; mating in spring Active year-round in warm weather; young born MayJune
Active year-round in warn weather; young born In June
Breeds in July-Auq.

Insects

Las iurus seninolus Semi'nole bat

Insects

Dasypus novemcinc tus Nine-t>anded armadillo

FIBS

Insects, plant material

Sylvllagus aquaticus Swamp rabKit

Oryzomys palustns ryzomys p
Harsfi
ri ice

FIBS

Green plants

Breeds Jan. -Sept.

FIBS

rat

Plant material, insects, crustaceans, bird eggs and young

Breeds Mar. -Oct.

Ondatra zibethicus Common musk rat

FIBS

61% crayfish; also crabs, birds, fish, Insects

Active year-round; breeding peaks Nov and Har.

.

Hyocastor coypus
Nutria

FI

Aquatic vegetation

Proc^ on lot or Northern raccoon

Hu stela vison Hink
Lut ra canadens
is

FIBS

Animals and plant material

Breeds Oec.-Jan.

FIB

Crayfish, rodents, birds, fish, crabs, frogs

Active year-round, young born In early spring Breeds

in

FIBS

Crabs, crayfish, fish, frogs, turtles, snakes

late fall

^T7er otter

Ddqco lcus yirginianus Uhite-tailed deer

i

FIB

Plant material

Breeds

in

Sept. -Mar.

134

i0?7?-10l

REPORT DOCUMENTATION
PAGE

w^

..r^

-^

Headquarters, Division

of Biological

Services, Washington,

DC

Eastern Energy and Land Use Teai

Leetown

WV

National Coastal Ecosystems

Slideii.

Team

LA

Western Energy and Land Use Team

Ft

Collins,

CO

Locations of Regional Offices

REGION
RegHiriiil

REGION
6^2
P.O.

REGION

Director

Regional Director
U.S. Fish and Wildlife Service

U.S. Fish and Wildlile Service

Lloyd Five Hundred Building, Suite 500 N.E. Multnomah Street

Portland, Oregon

Box 1306

Regional Director U.S. Fish and Wildlife Service Federal Building. Fort Snelhng

Alhuquerque,

New Mexico X7 103

Twin

Cities,

Minnesota 55111

97232

REGION

REGION

REGION
P.O.

Regional Director
U.S. Fish and Wildlife Service

Regional Director
U.S. Fish and Wildlife Service

Regional Director
U.S. Fish and Wildlile Sei^ice

Richard B. Russell Building 75 Spring Street, S.W.

Atlanta, Georgia

One Gateway Center Newton Corner, Massachusetts 0215X

Box 25486

30303

Denver Federal Center Denver, Colorado 80225

REGION

Regional Direcloi U.S. Fish and Wildlife Service 1011 b. Tudor Road

Anchorage, Alaska 9^503

DEPARTMENT OF THE INTERIOR

U.S.

FISH

AND WILDLIFE SERVICE

sibility for

principal conservation agency, the Department of the Interior has responof our nationally owned public lands and natural resources. This includes fostering the wisest use of our land and wator resources, protecting our fish and wildlife, preserving the environmental and cultural values oi our national parks and historical places, and pro-

As the Nation's
most

life through outdoor recreation. The Department assesses 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.

viding for the enjoyment of