Biological Services Program
FWS/OBS-81/24 January 1982
THE ECOLOGY OF THE MANGROVES OF SOUTH FLORIDA: A COMMUNITY PROFILE
**" ?lE
<m «*
J3ureau of Land Management
5H :ish and Wildlife Service
5 HO
,U5G
J.S. Department of the Interior
f
The Biological Services Program was established within the U.S. Fish and Wildlife Service to supply scientific information and methodologies on key environmental issues that impact fish and wildlife resources and their supporting ecosystems. The mission of the program is as follows:
• To strengthen the Fish and Wildlife Service in its role as a primary source of information on national fish and wild- life resources, particularly in respect to environmental impact assessment.
• To gather, analyze, and present information that will aid decisionmakers in the identification and resolution of problems associated with major changes in land and water use.
t To provide better ecological information and evaluation for Department of the Interior development programs, such as those relating to energy development.
Information developed by the Biological Services Program is intended for use in the planning and decisionmaking process to prevent or minimize the impact of development on fish and wildlife. Research activities and technical assistance services are based on an analysis of the issues, a determination of the decisionmakers involved and their information needs, and an evaluation of the state of the art to identify information gaps and to determine priorities. This is a strategy that will ensure that the products produced and disseminated are timely and useful.
Projects have been initiated in the following areas: coal extraction and conversion; power plants; geothermal , mineral and oil shale develop- ment; water resource analysis, including stream alterations and western water allocation; coastal ecosystems and Outer Continental Shelf develop- ment; and systems inventory, including National Wetland Inventory, habitat classification and analysis, and information transfer.
The Biological Services Program consists of the Office of Biological Services in Washington, D.C., which is responsible for overall planning and management; National Teams, which provide the Program's central scientific and technical expertise and arrange for contracting biological services studies with states, universities, consulting firms, and others; Regional Staffs, who provide a link to problems at the operating level; and staffs at certain Fish and Wildlife Service research facilities, who conduct in-house research studies.
FWS/OBS-81/24 January 1982
THE ECOLOGY OF THE MANGROVES OF SOUTH FLORIDA: A COMMUNITY PROFILE
by
William E. Odum Carole C. Mclvor Thomas J . Smi th , III
Department of Environmental Sciences
University of Virginia
Charlottesville, Virginia 22901
Project Officer
Ken Adams
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
1010 Gause Boulevard
SI idel 1 , Louisiana 70458
Performed for
National Coastal Ecosystems Team
Office of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
Washington, D.C. 20240
and
New Orleans OCS Office Bureau of Land Management U.S. Department of the Interior New Orleans, Louisiana 70130
DISCLAIMER
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.
nth Florida:
ar and
E Biological
RETURNED
Library of Congress Card Number 82-600562
This report should be cited:
Odum, W.E., C.C. Mclvor, and T.J. Smith, III. 1982. The ecology of the mangroves of south Florida: a community profile. U.S. Fish and Wildlife Service, Office of Biological Services, Washington, D.C. FWS/OBS-81/24. 144 pp.
PREFACE
This profile of the mangrove commun- ity of south Florida is one in a series of community profiles which treat coastal and marine habitats important to man. The obvious work that mangrove communities do for man includes the stabilization and protection of shorelines; the creation and maintenance of habitat for a great number of animals, many of which are either endangered or have commercial value; and the provision of the basis of whose final products include smorgasbord of oysters, crabs, shrimp, and fish. Less equally important benefits
a food web a seafood lobsters , tangible but include wilder-
ness, aesthetic ations .
and life support consider-
and Tarpon Springs on the west. Refer- ences are provided for those seeking in-depth treatment of a specific facet of mangrove ecology. The format, style, and level of presentation make this synthesis report adaptable to a diversity of needs such as the preparation of environmental assessment reports, supplementary reading in marine science courses, and the devel- opment of a sense of the importance of this resource to those citizens who control its fate.
Any questions requests for this directed to:
or comments about or publication should be
The information on these pages can give a basic understanding of the mangrove community and its role in the regional ecosystem of south Florida. The primary geographic area covered lies along the coast between Cape Canaveral on the east
Information Transfer Specialist National Coastal Ecosystems Team U.S. Fish and Wildlife Service NASA-SI i del 1 Computer Complex 1010 Gause Boulevard Slidell, Louisiana 70458
m
IV
CONTENTS
Page
PREFACE Ill
FIGURES vi i 1
TABLES vi i i
ACKNOWLEDGMENTS i x
CHAPTER 1 . INTRODUCTION 1
1.1 "Mangrove" Definition 1
1.2 Factors Controlling Mangrove Distribution 1
1.3 Geographical Distribution 2
1.4 Mangrove Species Descriptions 5
1.5 Mangrove Community Types 7
1.6 Substrates 9
1 . 7 Water Qual i ty 11
CHAPTER 2 . AUTECOLOGY OF MANGROVES 12
2.1 Adaptations to Natural Stress - Anaerobic Sediments 12
2.2 Adaptations to Natural Stress - Salinity 12
2.3 Reproductive Strategies 14
2.4 Biomass Partitioning 15
2.5 Primary Production 17
2.6 Herbivory 23
2.7 Wood Borers 24
2.8 Mangrove Diseases 25
CHAPTER 3. ECOSYSTEM STRUCTURE AND FUNCTION 26
3.1 Structural Properties of Mangrove Forests 26
3.2 Zonation, Succession and "Land Building" 26
3.3 Nutrient Cycling 30
3.4 Litter Fall and Decomposition 32
3.5 Carbon Export 34
3.6 Energy Flow 36
CHAPTER 4. COMMUNITY COMPONENTS - MICROORGANISMS 40
CHAPTER 5. COMMUNITY COMPONENTS - PLANTS OTHER THAN MANGROVES 41
5.1 Root and Mud Algae 41
5.2 Phytoplankton 43
5.3 Associated Vascular Plants 43
|
CHAPTER 6 |
|
6.1 |
|
6.2 |
|
6.3 |
|
6.4 |
|
7 |
1 |
|
7 |
2 |
|
7 |
3 |
|
7 |
4 |
|
7 |
5 |
|
7 |
6 |
|
CHAPTER 8. |
CONTENTS (continued)
Pase
COMMUNITY COMPONENTS - INVERTEBRATES 45
Ecological Relationships 45
Arboreal Arthropod Communi ty 47
Prop Root and Associated Mud Surface Community 47
Water Column Community 49
CHAPTER 7. COMMUNITY COMPONENTS - FISHES 50
Basin Mangrove Forests 50
Riverine Forests 52
Fringing Forests along Estuarine Bays and Lagoons 54
Fringing Forests along Oceanic Bays and Lagoons 56
Overwash Mangrove Islands 56
Gradient of Mangrove Community Interactions 57
COMMUNITY COMPONENTS - AMPHIBIANS AND REPTILES 58
CHAPTER 9. COMMUNITY COMPONENTS - BIRDS 61
9.1 Ecological Relationships 61
9.2 Wading Birds 61
9 . 3 Probi ng Shorebi rds 65
9.4 Floating and Diving Water Birds 65
9.5 Aerially-searching Birds 67
9.6 Birds of Prey 67
9.7 Arboreal Birds 68
9.8 Associations between Mangrove Community Types and Birds 70
9.9 Mangroves as Winter Habitat for North American Migrant Land Birds . 71
CHAPTER 10. COMMUNITY COMPONENTS - MAMMALS 72
CHAPTER 1 1 . VALUE OF MANGROVE ECOSYSTEMS TO MAN 74
1.1 Shoreline Stabilization and Storm Protection 74
1.2 Habitat Value to Wildlife 74
1.3 Importance to Threatened and Endangered Species 75
1.4 Value to Sport and Commercial Fisheries 75
1.5 Aesthetics, Tourism and the Intangibles 75
1.6 Economic Products 76
CHAPTER 12. MANAGEMENT IMPLICATIONS 77
12.1 Inherent Vulnerability 77
12.2 Man-induced Destruction 77
12.3 Effects of Oil Spills on Mangroves 80
12.4 Man-induced Modifications 81
12.5 Protective Measures Including Transplanting 84
12.6 Ecological Value of Black vs. Red Mangroves 85
12.7 The Importance of Inter-community Exchange 85
12.8 Management Practices: Preservation 86
vi
CONTENTS (continued)
Page
REFERENCES 88
APPENDIX A: SUMMARY OF SITE CHARACTERISTICS AND SAMPLING METHODOLOGY
FOR FISHES 1°6
APPENDIX B APPENDIX C APPENDIX D APPENDIX E
FISHES OF MANGROVE AREAS HO
AMPHIBIANS AND REPTILES FROM MANGROVE AREAS 127
AVIFAUNA OF MANGROVE AREAS 130
MAMMALS OF MANGROVE AREAS I42
vn
FIGURES Number Page
1 Approximate northern limits for the red mangrove (R), black
mangrove (B), and white mangrove (W) in Florida 3
2a A typical intertidal profile from south Florida showing the
distribution of red and black mangroves 4
2b The pattern of annual sea level change in south Florida 4
3 Three species of Florida mangroves with propagules, flowers and leaves. 6
4 The six mangrove community types 8
5a Aboveground and belowground biomass of a Puerto Rican red
mangrove forest 16
5b Light attenuation in a mangrove canopy; canopy height is 8 m 16
6 The hypothetical relationship between waterway position and
community net primary production of Florida mangrove forests 22
7 The hypothetical relationship between nutrient input, biomass, productivity, and nutrient export from mangrove ecosystems 31
8 Potential pathways of energy flow in mangrove ecosystems 37
9 Vertical distribution of selected algae and invertebrates on
red mangrove prop roots 42
10 Photograph of red mangrove prop root habitat in clear shallow
water with associated animal and plant populations 46
11 Gradient of mangrove-associated fish communities showing representative species 51
12 Aerial photograph of the mangrove belt of southwest Florida
near Whi tewater Bay 53
13 The mangrove water snake, Nerodia fasciata compressicauda ,
curl ed on a red mangrove prop root 59
14 A variety of wading birds feeding in a mangrove-lined pool
near Flamingo, Florida 62
15 Osprey returning to its nest in a red mangrove near Whitewater Bay 69
16 Damaged stand of red and black mangroves near Flamingo, Florida,
as it appeared 7 years after Hurricane Donna 78
17 Mangrove forest near Key West as it appeared in 1981 after
being destroyed by diking and impounding 79
18 Mangrove islands in Florida Bay near Upper Matecumbe Key 87
TABLES Number Page
la Estimates of mangrove production in Florida 20
lb Comparative measurements of photosynthesis in gC/m2/day 21
lc Gross primary production at different salinities 21
2 Aboveground biomass of mangrove forests in the Ten Thousand
Islands region of Florida 27
3 Estimates of litter fall in mangrove forests 33
4 Estimates of particulate carbon export from mangrove forests 35
5 Nesting statistics of wading birds and associated species in
south Florida, 1974-75 64
6 Timing of nesting by wading birds and associated species in
south Florida 66
7 General response of mangrove ecosystems to severe oil spills 82
8 Estimated impact of various stages of oil mining on mangrove ecosystems 83
vm
ACKNOWLEDGMENTS
Many individuals and organizations contributed significantly to the creation of this publication. Most notable was Eric Heald who worked extensively with us on the manuscript and contributed unpub- lished data. We thank Jeffrey Carlton, Edward Conner, Roy R. Lewis, III, Ariel Lugo, Larry Narcisse, Steven Macko, Aaron Mills, Michael Robblee, Martin Roessler, Samuel Snedaker, Durbin Tabb, Mike Wein- stein, and Joseph Zieman for information and helpful advice.
The draft manuscript was reviewed for its scientific content by Armando de la Cruz, Thomas Savage, James Kushlan (bird section), and James P. Ray. Each of these
individuals provided information as well as critical comments. We particularly appreciate the unselfish help of Ken Adams and the staff of the National Coastal Eco- systems Team of the U.S. Fish and Wildlife Service. Janet Ryan spent many hours typ- ing the manuscript.
Factual errors and faulty conclu- sions are the sole responsibility of the authors. Carole Mclvor has taken primary responsibility for Chapter 7, Tom Smith for Chapters 8, 9, and 10, and Bill Odum for the remainder of the publication. Unless otherwise noted, photographs, figures, and the cover were produced by the authors.
IX
CHAPTER 1
INTRODUCTION
1.1 "MANGROVE" DEFINITION
The term "mangrove" expresses two distinctly different concepts. One usage refers to halophytic species of trees and shrubs (halophyte = plant growing in saline soil). In this sense, mangrove is a catch-all, botanically diverse, non- taxonomic expression given to approximate- ly 12 families and more than 50 species (Chapman 1970) of tropical trees and shrubs (see Waisel 1972 for a detailed list). While not necessarily closely related, all these plants are adapted to (1) loose, wet soils, (2) a saline habi- tat, (3) periodic tidal submergence, and (4) usually have degrees of viviparity of propagules (see section 2.3 for discussion of "viviparity" and "propagules").
The second usage of the term mangrove encompasses the entire plant community including individual mangrove species. Synonymous terms include tidal forest, tidal swamp forest, mangrove community, mangrove ecosystem, mangal (Macnae 1968), and mangrove swamp.
For consistency, in this publication we will use the word "mangrove" for indi- vidual kinds of trees; mangrove community, mangrove ecosystem or mangrove forest will represent the entire assemblage of "man- groves".
1.2 FACTORS CONTROLLING MANGROVE DISTRI- BUTION
Four major factors appear to limit the distribution of mangroves and deter- mine the extent of mangrove ecosystem development. These factors include (1) climate, (2) salt water, (3) tidal fluc- tuation, and (4) substrate.
Climate
Mangroves are tropical species and do not develop satisfactorily in regions where the annual average temperature is below 19°C or 66°F (Waisel 1972). Normally, they do not tolerate temperature fluctuations exceeding 10°C (18°F) or
temperatures below freezing for any length of time. Certain species, for example, black mangrove, Avicennia germinans, on the northern coast of the Gulf of Mexico, maintain a semi -permanent shrub form by growing back from the roots after freeze damage.
Lugo and Zucca (1977) discuss the impact of low temperature stress on Flori- da mangroves. They found that mangrove communities respond to temperature stress by decreasing structural complexity (de- creased tree height, decreased leaf area index, decreased leaf size, and increased tree density). They concluded that man- groves growing under conditions of high soil salinity stress are less tolerant of low temperatures. Presumably, other types of stress (e.g., pollutants, diking) could reduce the temperature tolerance of man- groves.
High water temperatures can also be limiting. McMillan (1971) reported that seedlings of black mangrove were killed by temperatures of 39° to 40°C (102° to 104°F) although established seedlings and trees were not damaged. To our knowledge, upper temperature tolerances for adult mangroves are not well known. We suspect that water temperatures in the range 42° to 45°C (107° to 113°F) may be limiting.
Salt Water
Mangroves are facultative halo- phytes, i.e., salt water is not a physical requirement (Bowman 1917; Egler 1948). In fact, most mangroves are capable of growing quite well in freshwater (Teas 1979). It is important to note, however, that mangrove ecosystems do not develop in strictly freshwater environments; salinity is important in reducing competition from other vascular plant species (Kuenzler 1974). See section 2.2 about salinity tolerance of mangrove species.
Tidal Fluctuation
While tidal influence is not a direct physiological requirement for
mangroves, it plays an important indirect role. First, tidal stress (alternate wetting and drying), in combination with salinity, helps exclude most other vascular plants and thus reduces competi- tion. Second, in certain locations, tides bring salt water up the estuary against the outward flow of freshwater and allow mangroves to become established well inland. Third, tides may transport nutrients and relatively clean water into mangrove ecosystems and export accumula- tions of organic carbon and reduced sulfur compounds. Fourth, in areas with high evaporation rates, the action of the tides helps to prevent soil salinities from reaching concentrations which might be lethal to mangroves. Fifth, tides aid in the dispersal of mangrove propagules and detritus.
Because of all of these factors, termed tidal subsidies by E.P. Odum (1971), mangrove ecosystems tend to reach their greatest development around the world in low-lying regions with relatively large tidal ranges. Other types of water fluctuation, e.g., seasonal variation in freshwater runoff from the Florida Ever- glades, can provide similar subsidies.
Substrate and Wave Energy
Mangroves grow best in depositional environments with low wave energy. High wave energy prevents establishment of propagules, destroys the relatively shal- low mangrove root system and prevents the accumulation of fine sediments. The most productive mangrove ecosystems develop along deltaic coasts or in estuaries that have fine-grained muds composed of silt, clay and a high percentage of organic matter. Anaerobic sediments pose no problems for mangroves (see section 2.1) and exclude competing vascular plant species.
1.3 GEOGRAPHICAL DISTRIBUTION
Mangroves dominate approximately 75% of the world's tropical coastline between 25°N and 25°S latitude (McGill 1959). On
the east coast of Africa, in Australia and in New Zealand, they extend 10° to 15° farther south (Kuenzler 1974) and in Japan, Florida, Bermuda, and the Red Sea they extend 5° to 7° farther north. These areas of extended range generally occur where oceanographic conditions move un- usually warm water away from the equator.
Although certain reqions such as the tropical Indo-Pacific have as many as 30 to 40 species of mangroves present, only three species are found in Florida: the red mangrove, Rhizophora mangle, the black mangrove, Avicennia germi nans, and the white mangrove, Laguncularia racemosa. A fourth species, buttonwood, Conocarpus erecta, is not a true mangrove (no ten- dency to vivipary or root modification), but is an important species in the transi- tion zone on the upland edge of mangrove ecosystems (Tomlinson 1980).
The ranges of mangrove species in Florida have fluctuated over the past several centuries in response to relative- ly short-term climatic change. Currently, the situation is as follows (Figure 1). The red mangrove and the white mangrove have been reported as far north as Cedar Key on the west coast of Florida (Rehm 1976) and north of the Ponce de Leon Inlet on the east coast (Teas 1977); both of these extremes lie at approximately 29°10' N latitude. Significant stands lie south of Cape Canaveral on the east coast and Tarpon Springs on the west coast. The black mangrove has been reported as far north as 30°N latitude on the east coast of Florida (Savage 1972) and as scattered shrubs along the north coast of the Gulf of Mexico.
Intertidal Distribution
The generalized distribution of the red and black mangrove in relation to the intertidal zone is shown in Figure 2a. Local variations and exceptions to this pattern occur commonly in response to localized differences in substrate type and elevation, rates of sea level rise, and a variety of other factors (see sec- tion 3.2 for a full discussion of mangrove
R,W
Cedar Key
Tarpon Sprin
Tampa Bay
Port Charlotte Harbor/ Sanibel Island \^
R,W
30° N
Ponce de Leon Inlet
Cape Canaveral Indian River
Rookery Bay Ten Thousand Islands Shark River Vall<
Whitewater Bay/North River
Florida Keys #
Biscayne Bay
25°N
Figure 1. Approximate northern limits for the red mangrove (R), black mangrove (B), and white mangrove (W) in Florida (based on Savage 1972); although not in- dicated in the figure, the black mangrove extends along the northern Gulf of Mex- ico as scattered shrubs.
(A)
|
Mnvv — MLW-- |
— — .— f**r ' ■* ■ |
^PEAT |
|
*"** \ y\ |
, ■ • |
|
|
LOW |
HIGH |
|
|
MARSH |
MARSH |
|
|
(RED MANGROVE |
(BLACK MANGROVE |
|
|
DOMINATED) |
DOMINATED) |
/
(B)
FMAMJJASOND
+ 10 cm r
+ 5 cm -
SEA LEVEL
— 5 cm -
-10 cm L
Figure 2. (a) A typical intertidal profile from south Florida showing the dis- tribution of red and black mangrove (adapted from Provost 1974). (b) The pat- tern of annual sea level change in south Florida (Miami ) (adapted from Provost 1974).
zonation). Furthermore, it is important to recognize that the intertidal zone in most parts of Florida changes seasonally (Provost 1974); there is a tendency for sea level to be higher in the fall than in the spring (Figure 2b). As a result the "high marsh" may remain totally dry during the spring and be continually submerged in the autumn. This phenomenon further com- plicates the textbook concept of the in- tertidal, "low marsh" red mangrove and the infrequently flooded, "high marsh" black mangrove.
Mangrove Acreage in Florida
Estimates of the total acreage occupied by mangrove communities in Florida vary widely between 430,000 acres and over 500,000 acres (174,000 ha to over 202,000 ha). Eric Heald (Tropical Bioindustries, 9869 Fern St., Miami, Fl a.; personal communication 1981) has identified several reasons for the lack of agreement between estimates. These include: (1) inclusion or exclusion in surveys of small bays, ponds and creeks which occur within mangrove forests, (2) incorrect identification of mangrove areas from aerial photography as a result of inadequate "ground-truth" observations, poorly controlled aerial photography, and simple errors of planimetry caused by photography of inadequate scale.
The two most detailed estimates of area covered by mangroves in Florida are provided by the Coastal Coordinating Coun- cil, State of Florida (1974) and Birnhak and Crowder (1974). Considerable dif- ferences exist between the two estimates. The estimate of Birnhak and Crowder (1974), which is limited to certain areas of south Florida, appears to be unreal is- tically high, particularly for Monroe County (Eric Heald, personal communication 1981). Coastal Coordinating Council (1974) estimates a total of 469,000 acres (190,000 ha) within the State and suggests an expected margin of error of 15% (i.e. their estimate lies between 400,000 and 540,000 acres or 162,000 and 219,000 ha).
According to this survey, ninety percent of Florida's mangroves are located in the four southern counties of Lee (35,000 acres or 14,000 ha), Collier (72,000 acres or 29,000 ha), Monroe (234,000 acres or 95,000 ha), and Dade (81,000 acres or 33,000 ha).
Much of the area covered by mangroves in Florida is presently owned by Federal, State or County governments, or by non- profit organizations such as the National Audubon Society. Approximately 280,000 acres (113,000 ha) fall into this category (Eric Heald, personal communication 1981). Most of this acreage is held by the Federal Government as a result of the land being including within the Everglades National Park.
1.4 MANGROVE SPECIES DESCRIPTIONS
The following descriptions come largely from Carlton (1975) and Savage (1972); see these publications for further comments and photographs. For more detailed descriptions of germinating seeds (propagules) see section 2.3. The three species are shown in Figure 3.
The Black Mangrove (Avicennia germinans)
Avicennia germinans is synonymous with A. nitida and is a member of the family Avicenniaceae (formerly classed under Verbenaceae). The tree may reach a height of 20 m (64 ft) and has dark, scaly bark. Leaves are 5 to 10 cm (2 to 4 inches) in length, narrowly elliptic or oblong, shiny green above and covered with short, dense hairs below. The leaves are frequently encrusted with salt. This tree is characterized by long horizontal or "cable" roots with short vertical aerating branches (pneumatophores) that profusely penetrate the substrate below the tree. Propagules are lima-bean shaped, dark green while on the tree, and several centimeters (1 inch) long. The tree flowers in spring and early summer.
Black Mangrove, Avicennia germinans
White Mangrove, Laguncularia racemosa
Red Mangrove, Rhizophora mangle
Figure 3. Three species of Florida mangroves with propagules, flowers, and leaves,
The White Mangrove (Laguncularia racemosa)
The white mangrove is one of 450 species of plants in 18 genera of the family Combretaceae (synonymous with Terminal iaceae). It is a tree or shrub reaching 15 m (49 ft) or more in height with broad, flattened oval leaves up to 7 cm (3 inches) long and rounded at both ends. There are two salt glands at the apex of the petiole. The propagule is very small (1.0 to 1.5 cm or 0.4 to 0.6 inches long) and broadest at its apex. Flowering occurs in spring and early summer.
The Red Mangrove (Rhizophora mangle)
The red mangrove is one of more than 70 species in 17 genera in the family Rhizophoraceae. This tree may reach 25 m (80 ft) in height, has thin grey bark and dark red wood. Leaves may be 2 to 12 cm (1 to 5 inches) long, broad and blunt- pointed at the apex. The leaves are shiny, deep green above and paler below. It is easily identified by its charac- teristic "prop roots" arising from the trunk and branches. The penci 1 -shaped propagules are as much as 25 to 30 cm (10 to 12 inches) long after germination. It may flower throughout the year, but in Florida flowering occurs predominately in the spring and early summer.
1.5 MANGROVE COMMUNITY TYPES
Mangrove forest communities exhibit tremendous variation in form. For example, a mixed scrub forest of black and red mangroves at Turkey Point on Biscayne Bay bears little resemblance to the luxuriant forests, dominated by the same two species, along the lower Shark River.
Lugo and Snedaker (1974) provided a convenient classification system based on mangrove forest physiogomy. They identi- fied six major community types resulting from different geological and hydrological processes. Each type has its own charac- teristic set of environmental variables such as soil type and depth, soil salinity
range, and flushing rates. Each community type has characteristic ranges of primary production, litter decomposition and car- bon export along with differences in nutrient recycling rates, and community components. The community types as shown in Figure 4 are as follows:
(1) Overwash mangrove forests - these islands are frequently overwashed by tides and thus have high rates of organic export. All species of mangroves may be present, but red mangroves usually domi- nate. Maximum height of the mangroves is about 7 m (23 ft).
(2) Fringe mangrove forests - man- groves form a relatively thin fringe along waterways. Zonation is typically as de- scribed by Davis (1940) (see discussion in section 3.2). These forests are best defined along shorelines whose elevations are higher than mean high tide. Maximum height of the mangroves is about 10 m (32 ft).
(3) Riverine mangrove forests - this community type includes the tall flood plain forests along flowing waters such as tidal rivers and creeks. Although a shal- low berm often exists along the creek bank, the entire forest is usually flushed by daily tides. All three species of mangroves are present, but red mangroves (with noticeably few, short prop roots) predominate. Mangroves may reach heights of 18 to 20 m (60 to 65 ft).
(4) Basin mangrove forests - these forests occur inland in depressions chan- neling terrestrial runoff toward the coast. Close to the coast they are in- fluenced by daily tides and are usually dominated by red mangroves. Moving in- land, the tidal influence lessens and dominance shifts to black and white man- groves. Trees may reach 15 m (49 ft) in height.
(5) Hammock forests - hammock man- grove communities are similar to the basin type except that they occur on ground that is slightly elevated (5 to 10 cm or 2 to 4 inches) relative to surrounding areas.
(1) OVERWASH FOREST (2) FRINGE FOREST
(3) RIVERINE FOREST (4) BASIN FOREST
-^ggl ScL £L '
(5) HAMMOCK FOREST (6) SCRUB FOREST
Figure 4. The six mangrove community types (Lugo and Snedaker 1974).
All species of mangroves may be present. Trees rarely exceed 5 m (16 ft) in height.
(6) Scrub or dwarf forests - this community type is limited to the flat coastal fringe of south Florida and the Florida Keys. All three species are present. Individual plants rarely exceed 1.5 m (4.9 ft) in height, except where they grow over depressions filled with mangrove peat. Many of these tiny trees are 40 or more years of age. Nutrients appear to be limiting although substrate (usually limestone marl) must play a role.
Throughout this publication we have attempted to refer to Lugo and Snedaker's classification scheme wherever possible. Without a system of this type, comparisons between sites become virtually meaningless.
1.6 SUBSTRATES
Understanding mangrove-substrate relationships is complicated by the ability of mangroves to grow on many types of substrates and because they often alter the substrate through peat formation and by altering patterns of sedimentation. As a result, mangroves are found on a wide variety of substrates including fine, inorganic muds, muds with a high organic content, peat, sand, and even rock and dead coral if there are sufficient crevices for root attachment. Mangrove ecosystems, however, appear to flourish only on muds and fine-grained sands.
In Florida, the primary mangrove soils are either calcareous marl muds or calcareous sands in the southern part of the State and siliceous sands farther north (Kuenzler 1974). Sediment distribu- tion and, hence, mangrove development, is controlled to a considerable extent by wave and current energy. Low energy shorelines accumulate fine-grained sedi- ments such as mud and silt and usually have the best mangrove growth. Higher energy shorelines (more wave action or higher current velocities) are charac- terized by sandy sediments and less pro- ductive mangroves. If the wave energy
becomes too great, mangroves will not be present. Of the three species of Florida mangroves, white mangroves appear to tolerate sandy substrates the best (per- sonal observation), possibly because this species may tolerate a greater depth to the water table than the other two species.
Mangroves in Florida often modify the underlying substrate through peat deposi- tion. It is not unusual to find layers of mangrove peat several meters thick under- lying well-established mangrove ecosystems such as those along the southwest coast of Florida. Cohen and Spackman (1974) pre- sented a detailed account of peat forma- tion within the various mangrove zones of south Florida and also in areas dominated by black needle rush (Juncus roemerianus), smooth cordgrass (Sparti na al terni fl ora) and a variety of other macrophytes; Cohen and Spackman (1974) also provide descrip- tions and photography to aid in the iden- tification of unknown peat samples.
The following descriptions come from Cohen and Spackman (1974) and from the personal observations of W.E. Odum and E.J. Heald. Red mangroves produce the most easily recognized peat. More recent deposits are spongy, fibrous and composed to a great extent of fine rootlets (0.2 to 3.0 mm in diameter). Also present are larger pieces of roots (3 to 25 mm), bits of wood and leaves, and inorganic materials such as pyrite, carbonate minerals, and quartz. Older deposits are less easily differentiated although they remain somewhat fibrous. Peat which has recently been excavated is reddish-brown although this changes to brown-black after a short exposure to air. Older deposits are mottled reddish-brown; deposits with a high content of carbonates are greyish- brown upon excavation.
Cohen and Spackman (1974) were unable to find deposits of pure black mangrove or white mangrove peat suggesting that these two species may not form extensive depos- its of peat while growing in pure stands. There are, however, many examples of peats which are mixtures of red mangrove material and black mangrove roots. They
suggested that the black mangrove peats identified by Davis (1946) were probably mixtures of peat from several sources.
Throughout south Florida the sub- strate underlying mangrove forests may consist of complicated patterns of calcareous muds, marls, shell, and sand interspersed and overlain by layers of mangrove peat and with limestone bedrock at the bottom. Detailed descriptions of this complex matrix and its spatial varia- tion were given by Davis (1940, 1943, 1946), Egler (1952), Craighead (1964), Zieman (1972) and Cohen and Spackman (1974) among others. Scoffin (1970) dis- cussed the ability of red mangrove to trap and hold sediments about its prop roots. So called "land-building" by man- groves is discussed in section 3.2.
The long-term effect of mangrove peat on mangrove distribution is not entirely clear. Certainly, if there is no change in sea level or if erosion is limited, the accumulation of peat under stands of red mangroves combined with deposition and accumulation of suspended sediments will raise the forest floor sufficiently to lead to domination by black or white man- groves and, ultimately, more terrestrial species. Whether this is a common se- quence of events in contemporary south Florida is not clear. It is clear that peat formtion is a passive process and occurs primarily where and when physical processes such as erosion and sea level rise are of minimal importance (Wanless 1974).
Zieman (1972) presented an inter- esting argument suggesting that mangrove peat may be capable of dissolving under- lying limestone rock, since carbonates may dissolve at pH 7.8. Through this process, shallow depressions might become deeper and the overlying peat layer thicker without raising the surface of the forest floor.
Data on chemical characteristics of Florida mangrove soils and peat are limited. Most investigators have found mangrove substrates to be almost totally anaerobic. Lee (1969) recorded typical Eh
values of -100 to -400 mv in mangrove peats. Such evidence of strongly reducing conditions are not surprising considering the fine-grained, high organic nature of most mangrove sediments. Although man- groves occur in low organic sediments (less than 1% organic matter), typical values for mangrove sediments are 10% to 20% organic matter.
Lee (1969) analyzed 3,000- to 3,500- year-old mangrove peat layers underlying Little Black Water Sound in Florida Bay for lipid carbon content. Peat lipid content varied between 0.6 and 2.7 mg lipid-C/gram of peat (dry wt ) or about 3% of the total organic carbon total. These values usually increased with depth. Long chain fatty acids (C-16 and C-18) were the dominant fatty acids found.
Florida mangrove peats are usually acidic, although the presence of carbonate materials can raise the pH above 7.0. Zieman (1972) found red mangrove peats to range from pH 4.9 to 6.8; the most acid conditions were usually found in the cen- ter of the peat layer. Lee (1969) re- corded a pH range from 5.8 to 6.8 in red mangrove peat at the bottom of a shallow embayment. Although Davis (1940) found a difference between red mangrove peat (5.0 to 5.5) and black mangrove peat (6.9 to 7.2), this observation has not been con- firmed because of the previously mentioned difficulty in finding pure black mangrove peat.
Presumably, the acidic character of mangrove peat results from release of organic acids during anaerobic decomposi- tion and from the oxidation of reduced sulfur compounds if the peat is dried in the presence of oxygen. This last point explains why "reclaimed" mangrove areas often develop highly acidic soils (pH 3.5 to 5.0) shortly after reclamation. This "cat clay" problem has greatly complicated the conversion of mangrove regions to agricultural land in Africa and southeast Asia (Hesse 1961; Hart 1962, 1963; Macnae 1968).
In summary, although current under- standing of mangrove peats and soils is
10
fragmentary and often contradictory, we can outline several generalizations:
(1) Mangroves can grow on a wide variety of substrates including mud, sand, rock, and peat.
(2) Mangrove ecosystems appear to flourish on fine-grained sediments which are usually anaerobic and may have a high organic content.
(3) Mangrove ecosystems which per- sist for some time may modify the under- lying substrate through peat formation. This appears to occur only in the absence of strong physical forces.
(4) Mangrove peat is formed pri- marily by red mangroves and consists pre- dominantly of root material.
(5) Red mangrove peats may reach thicknesses of several meters, have a relatively low pH, and may be capable of dissolving underlying layers of limestone.
(6) When drained, dried, and aerated, mangrove soils usually experience dramatic increases in acidity due to the oxidation of reduced sulfur compounds. This greatly complicates their conversion to agricul ture.
1.7 WATER QUALITY
from virtually fresh water to above 40 ppt (discussed in section 2.2), (2) low macro- nutrient concentrations (particularly phosphorous), (3) relatively low dissolved oxygen concentrations, and (4) frequently increased water color and turbidity. The last three characteristics are most pro- nounced in extensive mangrove ecosystems such as those adjacent to the Everglades and least pronounced in small, scattered forests such as the overwash islands in the Florida Keys.
In general, the surface waters associated with mangroves are charac- terized by (1) a wide range of salinities
The results of oxygen depletion and nutrient removal are (1) dissolved oxygen concentrations below saturation, typically 2 to 4 ppm and often near zero in stagnant locations and after heavy, storm-generated runoff, (2) very low total phosphorus values, frequently below detection limits, and (3) moderate total nitrogen values (0.5 to 1.5 mg/1). In addition, TOC (total organic carbon) may range from 4 to 50 ppm or even higher after rain; Eric Heald (personal communication 1981) has measured DOC (dissolved organic carbon) values as high as 110 ppm in water flowing from mangroves to adjacent bays. Tur- bidity usually falls in the 1 to 15 JTU (Jackson turbity units) range. The pH of the water column in Florida swamps is usually between 6.5 and 8.0 and alkalinity between 100 to 300 mg/1. Obviously, ex- ceptions to all of these trends can occur. Both natural and human disturbance can raise macronutrient levels markedly.
11
CHAPTER 2. AUTECOLOGY OF MANGROVES
2.1 ADAPTATIONS TO NATURAL STRESS - ANAEROBIC SEDIMENTS
Mangroves have a series of remarkable adaptations which enable them to flourish in an environment characterized by high temperatures, widely fluctuating salini- ties, and shifting, anaerobic substrates. In this section we review a few of the most important adaptations.
The root system of mangroves provides the key to existence upon unfriendly sub- strates (see Gill and Tomlinson 1971 for an anatomical review of mangrove roots). Unlike most higher plants, mangroves usually have highly developed aerial roots and modest below-ground root systems. The aerial roots allow atmospheric gases to reach the underground roots which are embedded in anaerobic soils. The red mangrove has a system of stilt or prop roots which extend a meter (3 ft) or more above the surface of the soil and contain many small pores (lenticels) which at low tide allow oxygen to diffuse into the plant and down to the underground roots by means of open passages called aerenchyma (Scholander et al. 1955). The lenticels are highly hydrophobic and prevent water penetration into the aerenchyma system during high tide (Waisel 1972).
The black mangrove does not have prop roots, but does have small air roots or pneumatophores which extend vertically upward from the underground roots to a height of 20 to 30 cm (8 to 12 inches) above the soil. These pneumatophores resemble hundreds of tiny fingers sticking up out of the mud underneath the tree canopy. At low tide, air travels through the pneumatophores into the aerenchyma system and then to all living root tis- sues. The white mangrove usually does not have either prop roots or pneumatophores, but utilizes lenticels in the lower trunk to obtain oxygen for the aerenchyma sys- tem. "Peg roots" and pneumatophores may be present in certain situations (Jenik 1967).
Mangroves achieve structural stabili- ty in at least two ways. Species such as the red mangrove use the system of prop
roots to provide a more or less firm foun- dation for the tree. Even though the prop roots are anchored with only a modest assemblage of underground roots, the hori- zontal extent of the prop root system insures considerable protection from all but the worst of hurricanes. Other man- grove species, including the black man- grove, obtain stability with an extensive system of shallow, underground "cable" roots that radiate out from the central trunk for a considerable distance in all directions; the pneumatophores extend up- ward from these cable roots. As in all Florida mangroves, the underground root system is shallow and a tap root is lacking (Walsh 1974). As Zieman (1972) found, individual roots, particularly of red mangroves, may extend a meter or more downward in suitable soils.
From the standpoint of effectiveness in transporting oxygen to the underground roots, both prop roots and cable roots seem equally effective. From the perspec- tive of stability, the prop roots of red mangroves appear to offer a distinct ad- vantage where wave and current energies are high.
Unfortunately, as pointed out by Odum and Johannes (1975), the same structure which allows mangroves to thrive in an- aerobic soil is also one of the tree's most vulnerable components. Exposed por- tions of the aerial root system are sus- ceptible to clogging by fine suspended material, attack by root borers, and pro- longed flooding (discussed further in section 12.1). Such extended stress on the aerial roots can kill the entire tree.
2.2 ADAPTATIONS TO NATURAL STRESS - SALINITY
Mangroves accommodate fluctuations and extremes of water and soil salinity through a variety of mechanisms, although not all mechanisms are necessarily present in the same species. Scholander et al. (1962) reported experimental evidence for two major methods of internal ion regula- tion which they identified in two dif- ferent groups of mangroves: (1) the salt
12
exclusion species and (2) the salt excre- tion species. In addition, some mangroves utilize succulence and the discarding of salt-laden organs or parts (Teas 1979).
The salt-excluding species, which include the red mangrove, separate freshwater from sea water at the root surface by means of a non-metabolic ultra- filtration system (Scholander 1968). This "reverse osmosis" process is powered by a high negative pressure in the xylem which results from transpiration at the leaf surface. Salt concentration in the sap of salt-excluding mangroves is about 1/70 the salt concentration in sea water, although this concentration is almost 10 times higher than found in normal plants (Scholander et al. 1962).
Salt-secreting species, including black and white mangroves (Scholander 1968), use salt glands on the leaf surface to excrete excess salt. This is probably an enzymatic process rather than a physi- cal process since it is markedly tempera- ture sensitive (Atkinson et al. 1967). The process appears to involve active transport with a requirement for biochemi- cal energy input. As a group, the salt secreters tend to have sap salt concentra- tions approximately 10 times higher (1/7 the concentration of sea water) than that of the salt excluders.
In spite of these two general tenden- cies, it is probably safe to say that individual species utilize a variety of mechanisms to maintain suitable salt balance (Albert 1975). For example, the red mangrove is an effective, but not perfect, salt excluder. As a result this species must store and ultimately dispose of excess salt in leaves and fruit (Teas 1979). Most salt secreters, including white and black mangroves, are capable of limited salt exclusion at the root sur- face. The white mangrove, when exposed to hypersaline conditions, not only excludes some salt and secretes excess salt through its salt glands, but also develops thickened succulent leaves and discards salt during leaf fall of senescent leaves (Teas 1979).
There appears to be some variation in the salinity tolerance of Florida man- groves. The red mangrove is probably limited by soil salinities above 60 to 65 ppt. Teas (1979) recalculated Bowman's (1917) data and concluded that transpira- tion in red mangrove seedlings ceases above 65 ppt. Cintron et al. (1978) found more dead than living red mangrove trees where interstitial soil salinities ex- ceeded 65 ppt.
On the other hand, white and black mangroves, which both possess salt excre- tion and limited salt exclusion mech- anisms, can exist under more hypersaline conditions. Macnae (1968) reported that black mangroves can grow at soil salini- ties greater than 90 ppt. Teas (1979) reported dwarfed and gnarled black and white mangroves occurring in Florida at soil salinities of 80 ppt.
There may be an additional factor or factors involved in salinity tolerance of mangroves. McMillan (1975) found that seedlings of black and white mangroves survived short-term exposures to 80 ppt and 150 ppt sea water if they were grown in a soil with a moderate clay content. They failed to survive these salinities, however, if they were grown in sand. A soil with 7% to 10% clay appeared to be adequate for increased protection from hypersaline conditions.
Vegetation-free hypersaline lagoons or bare sand flats in the center of man- grove ecosystems have been described by many authors (e.g., Davis 1940; Fosberg 1961; Bacon 1970). These features have been variously called salitrals (Holdridge 1940), salinas, salterns, salt flats, and salt barrens. Evidently, a combination of low seasonal rainfall, occasional inunda- tion by sea water, and high evaporation rates results in soil salinities above 100 ppt, water temperatures as high as 45°C (113°F) in any shallow, standing water, and subsequent mangrove death (Teas 1979). Once established, salinas tend to persist unless regular tidal flushing is enhanced by natural or artificial changes in tidal circulation.
13
Although salinas occur frequently in Florida, they are rarely extensive in area. For example, between Rookery Bay and Marco Island (south of Naples, Florida) there are a series of salinas in the black mangrove-dominated zone on the upland side of the mangrove swamps. These hypersaline lagoons occur where the normal flow of fresh water from upland sources has been diverted, presumably resulting in elevated soil salinities during the dry winter months.
In summary, salinity is a problem for mangroves only under extreme hypersaline conditions. These conditions occur natu- rally in Florida in irregularly flooded areas of the "high swamp" above the normal high tide mark and are accompanied by high soil salinities. Florida mangroves, listed in order of increasing salinity tolerance, appear to be red, white, and black.
2.3 REPRODUCTIVE STRATEGIES
As pointed out by Rabinowitz (1978a), virtually all mangroves share two common reproductive strategies: dispersal by means of water (van der Pijl 1972) and vivipary (Macnae 1968; Gill and Tomlinson 1969). Vivipary means that the embryo develops continuously while attached to the parent tree and during dispersal. Since there is uninterrupted development from zygote through the embryo to seedling without any intermediate resting stages, the word "seed" is inappropriate for viviparous species such as mangroves; the term "propagule" is generally used in its place.
While the phenology of black and white mangroves remains sketchy, Gill and Tomlinson (1971) thoroughly described the sequence of flowering in the red mangrove. Flowering in this species may take place at any time of the year, at least in extreme south Florida, but reaches a maxi- mum in the late spring and summer. The flowers open approximately 1 to 2 months after the appearance of buds. The flower remains intact only 1 to 2 days; this
probably accounts for the low fertiliza- tion rate, estimated by Gill and Tomlinson at 0% to 7.2%. Propagule development is slow, ranging from 8 to 13 months. Savage (1972) mentions that on the Florida gulf coast, red mangrove propagules mature and fall from the tree from July to September. Within the Everglades National Park, black mangroves flower from May until July and bear fruit from August until November while white mangroves flower from May to August and bear fruit from July to October (Loope 1980).
The propagules of the three species of Florida mangroves are easy to differen- tiate. The following descriptions all come from Rabinowitz (1978a). White man- grove propagules are small and flattened, weigh less than a gram, are about 2 cm long, are pea-green when they fall from the parent tree, and turn mud-brown in two days or so. The pericarp (wall of the ripened propagule) serves as a float and is not shed until the seedling is estab- lished. During dispersal the radicle (embryonic root) emerges from the propa- gule. This germination during dispersal has led Savage (1972) to refer to the white mangrove as "semi-viviparous".
The propagules of the black mangrove when dropped from the tree are oblong- elliptical (resemble a flattened olive), weigh about 1 g and are about 2 cm long. The pericarp is lost within a few days after dropping from the tree; at this point the cotyledons (primary leaves) unfold and the propagule resembles two butterflies on top of one another.
Propagules of the red mangrove under- go extensive vivipary while on the tree. When propagules fall from the tree they resemble large green beans. They are rod- shaped with pointed ends, about 20 cm long, and weigh an average of 15 g.
Propagules of all three species float and remain viable for extended periods of time. Apparently, there is an obligate dispersal time for all Florida mangroves, i.e., a certain period of time must elapse during dispersal for germination to be
14
complete and after which seedling estab- lishment can take place. Rabinowitz (1978a) estimates the obligate dispersal period at approximately 8 days for white mangroves, 14 days for black, and 40 days for red. She further estimates the addi- tional time for root establishment at 5, 7, and 15 days for white, black, and red mangroves, respectively. Her estimate for viable longevity of the propagules is 35 days for white mangroves and 110 days for black. Davis (1940) reports viable propa- gules of red mangroves that had been kept floating for 12 months.
Rabinowitz (1978a) also concluded that black and white mangroves require a stranding period of 5 days or more above the influence of tides to take hold in the soil. As a result, these two species are usually restricted to the higher portions of the mangrove ecosystem where tidal effects are infrequent.
The elongated red mangrove propagule, however, has the potential to become established in shallow water with tidal influence. This happens in at least two ways: (1) stranding in a vertical posi- tion (they float vertically) or (2) stranding in a horizontal position, rooting and then vertical erection by the plant itself. Lawrence (1949) and Rabino- witz (1978a) felt that the latter was the more common method. M. Walterding (Calif. Acad. Sci., San Francisco; personal com- munication 1980) favors vertical estab- lishment; based upon his observations, surface water turbulence works the propa- gule into the substrate during falling tides.
Mortality of established seedlings seems to be related to propagule size. Working in Panama, Rabinowitz (1978b) found that the mortality rate of mangrove seedlings was inversely correlated with initial propagule size. The white man- grove, which has the smallest propagule, has the highest rate of seedling mortal- ity. The black mangrove has an interme- diate mortality rate while the red man- grove, with the largest propagule, has the lowest seedling mortality rate. She
concluded that species with small propagules establish new cohorts annually but die rapidly, while species such as the red mangroves may have long-lived and often overlapping cohorts.
Propagule size and seedling mortality rates are particularly important in con- siderations of succession and replacement in established mangrove forests. Light is usually the most serious limiting factor underneath existing mangrove canopies. Rabinowitz (1978b) suggested that species with short-lived propagules must become established in an area which already has adequate light levels either due to tree fall or some other factor. In contrast, red mangrove seedlings can become estab- lished under an existing, dense canopy and then, due to their superior embryonic reserves, are able to wait for months for tree fall to open up the canopy and pre- sent an opportunity for growth.
2.4 BIOMASS PARTITIONING
Few investigators have partitioned the total biomass, aboveground and below- ground, contained in a mangrove tree. An analysis of red mangroves in a Puerto Rican forest by Golley et al. (1962) gives some insight into what might be expected in south Florida. Aboveground and below- ground biomass existed in a ratio of 1:1 if fine roots and peat are ignored (Figure 5). In this case, peat and very fine roots (smaller than 0.5 cm diameter) ex- ceeded remaining biomass by 5:1. Lugo et al. (1976) reported the following values for a south Florida red mangrove overwash forest. All values were reported in dry grams per square meter, plus and minus one standard error, and ignoring belowground biomass. They found 710 - 22 q/ml of leaves, 12. 8„ - 15.3 g/mz of propagules,
iles, f of
7043 i 7 g/mz of wood, 4695 ± 711 g/rr prop roots and 1565 - 234.5 g/mz of detri- tus on the forest floor.
Biomass partitioning between dif- ferent species and locations must be highly variable. The age of the forest will influence the amount of wood biomass;
15
(A!
SOIL SURFACE
LEAVES (778)
BRANCHES ( 1 274)
PROP ROOTS ( 1437)
TRUNK (2796)
LARGE ROOTS (997)
SMALL ROOTS (4000)
(B)
HEIGHT (M)
o
5000 10000
FOOT CANDLES
Figure 5. (a) Aboveground and belowground blomass of a Puerto R1can red mangrove forest. Values in parentheses are dry g/m2; large roots = 2 cm+ in diameter, small roots = 0.5 - 1.0 cm. (b) Vertical distribution of light Intensity 1n the same forest; canopy height is 8 m (26 ft) (both figures adapted from Golley et al. 1962).
16
detritus varies enormously from one site to the next depending upon the amount of fluvial transport. The biomass charac- teristics of a scrub forest probably bear little resemblance to those of a fringing forest. At the present time, there is not enough of this type of data available to draw many conclusions. One intriguing point is that red mangrove leaf biomass averages between 700 and 800 g/m at various sites with very different forest morphologies (Odum and Heald 1975a). This may be related to the tendency of mangrove canopies, once they have become estab- lished, to inhibit leaf production at lower levels through self-shading.
Golley et al. (1962) showed that the red mangrove canopy is an extremely effi- cient light interceptor. Ninety— five percent of the available light had been intercepted 4 m (13 ft) below the top of the canopy (Figure 5). As a result, 90% of the leaf biomass existed in the upper 4 m of the canopy. Chlorophyll followed the same pattern of distribution.
The leaf area index (LAI) of mangrove forests tends to be relatively low. Gol- ley et al. (1962) found a LAI of 4.4 for a Puerto Rican red mangrove forest. Lugo et al. (1975) reported a LAI of 5.1 for a Florida black mangrove forest and 3.5 for a Florida fringe red mangrove forest. A different black mangrove forest, in Flori- da, was found to have values ranging from 1 to 4 and an average of 2 to 2.5 (Lugo and Zucca 1977). These values compare with LAI's of 10 to 20 recorded for most tropical forests (Golley et al. 1974). The low leaf area values of mangrove forests can be attributed to at least three factors: (1) effective light inter- ception by the mangrove canopy, (2) the inability of the lower mangrove leaves to flourish at low light intensities, and (3) the absence of a low-light-adapted plant layer on the forest floor.
2.5 PRIMARY PRODUCTION
Prior to 1970 virtually no informa- tion existed concerning the productivity
of mangroves in Florida. Since that time knowledge has accumulated rapidly, but it is still unrealistic to expect more than preliminary statements about Florida man- grove productivity. This deficiency can be traced to (1) the difficulties asso- ciated with measurements of mangrove pro- ductivity and (2) the variety of factors that affect productivity and the resulting variations that exist from site to site.
Productivity estimates come from three methods: (1) harvest, (2) gas ex- change, and (3) litter fall. Harvest methods require extensive manpower and knowledge of the age of the forest. They are best employed in combination with silviculture practices. Since silvicul- ture of south Florida mangroves is practi- cally non-existent, this method has rarely been used in Florida. Noakes (1955), Macnae (1968), and Walsh (1974) should be consulted for productivity estimates based on this technique in other parts of the world.
Gas exchange methods, based on measurements of CO? changes, have the advantage of precision and response to short-term changes in light, temperature, and flooding. They include both above- ground and belowground production. On the negative side, the necessary equipment is expensive and tricky to operate properly. Moreover, extrapolations from short-term measurements to long-term estimates offer considerable opportunity for error. Nevertheless, the best estimates of pro- ductivity come from this method.
The litter fall technique (annual litter fall x 3 = annual net primary pro- duction) was proposed by Teas (1979) and is based on earlier papers by Bray and Gorham (1964) and Golley (1972) for other types of forests. This is a quick and dirty method although the lack of pre- cision remains to be demonstrated for mangroves. An even quicker and dirtier method proposed by Teas (1979) is to (1 ) estimate leaf standing crop (using various techniques including harvesting or light transmission relationships) and (2) multi- ply by three. This assumes an annual leaf
17
turnover of one, which is supported by the data of Heald (1969) and Pool et al. (1977).
Mangrove productivity is affected by many factors; some of these have been recognized and some remain totally ob- scure. Carter et al. (1973) propose lumping these factors into two broad cate- gories: tidal and water chemistry. We believe that a number of additional cate- gories should be considered.
A minimal, though incomplete, list of factors controlling mangrove productivity must include the following:
* species composition of the stand
* age of the stand
* presence or absence of competing species
* degree of herbivory
* presence or absence of disease and parasites
* depth of substrate
* substrate type
* nutrient content of substrate
* nutrient content of overlying water
* salinity of soil and overlying water
* transport efficiency of oxygen to root system
* amount of tidal flushing
* relative wave energy
' presence or absence of nesting birds
* periodicity of severe stress (hurri- canes, fire, etc.)
* time since last severe stress
* characteristics of ground water
inputs of toxic compounds or nutrients from human activities
human influences such as diking, ditching, and altering patterns of runoff.
In spite of the difficulties with various methods and the interaction of controlling factors, it is possible to make general statements about certain aspects of mangrove productivity. For example, Waisel's (1972) statement that mangroves have low transpiration rates seems to be generally true in Florida. Lugo et al. (1975) reported transpiration rates of 2,500 g HoO/m /day for mangrove leaves in a fringing red mangrove forest and 1,482 g h^O/m /day for black mangrove leaves. This is approximately one-third to one-half the value found in temperate broad leaf forests on hot dry days, but comparable to tropical rainforests (H.T. Odum and Jordan 1970). The low transpira- tion rates of mangroves are probably re- lated to the energetic costs of main- taining sap pressures of -35 to -60 atmo- spheres (Scholander et al. 1965).
Litter fall (leaves, twigs, bark, fruit, and flowers) of Florida mangrove forests appears to average 2 to 3 dry g/m day in most wel 1 -devel oped mangrove stands (see discussion in section 3.4). This can be an order of magnitude lower in scrub forests.
Wood production of mangroves appears to be high compared to other temperate and tropical trees, although no measurements from Florida are available. Noakes (1955) estimated that the wood production of an intensively managed Malayan forest was 39.7 metric tons/ha/year. Teas (1979) suggested a wood production estimate of 21 metric tons/ha/year for a mature unmanaged red mangrove forest in south Florida. His figure was calculated from a litter/total biomass relationship and is certainly subject to error.
Representative estimates of gross primary production (GPP) net primary
18
production (NPP), and respiration (R) of Florida mangroves are given in Table la. Compared to net primary production (NPP) estimates from other ecosystems, including agricultural systems (E.P. Odum 1971), it appears that mangroves are among the world's most productive ecosystems. Healthy mangrove ecosystems appear to be more productive than sea grass, marsh grass and most other coastal systems.
Further examination of Table la re- veals several possible tendencies. The first hypothetical tendency, as discussed by Lugo et al. (1975), is for red mangroves to have the highest total net production, black to have intermediate values and white the lowest. This conclusion assumes that the plants occur within the zone for which they are best adapted (see section 3.2 for discussion of zonation) and are not existing in an area with strong limit- ing factors. A scrub red mangrove forest, for example, growing under stressed condi- tions (high soil salinity or low nutrient supply), has relatively low net produc- tivity (Teas 1979). The pre-eminent posi- tion of red mangroves is shown by the comparative measurements of photosynthesis in Table lb; measurements were made within canopy leaves of trees growing within their zones of optimal growth.
A second noteworthy tendency is that red mangrove GPP decreases with increasing salinity while GPP of black and white mangroves increases with increasing salinity up to a point. Estimates of Hicks and Burns (1975) demonstrate that this may be a real tendency (Table lc).
Data presented by Miller (1972), Carter et al. (1973), Lugo and Snedaker (1974), and Hicks and Burns (1975) sug- gest a third hypothetical tendency, assuming occurrence of the species within its adapted zone. It appears that the black mangrove typically has a much higher respiration rate, lower net productivity, and lower GPP/R ratio than the red man- grove. This can be attributed at least partially, to the greater salinity stress under which the black mangrove usually grows; this leads to more osmotic work.
These three apparent tendencies have led Carter et al. (1973) and Lugo et al. (1976) to propose a fourth tendency, an inverted U-shaped relationship between waterway position and net mangrove com- munity productivity (Figure 6). This tendency is best understood by visualizing a typical gradient on the southwest coast of Florida. At the landward end of the gradient, salinities are very low, nutrient runoff from terrestrial eco- systems may be high and tidal amplitude is minor. At the seaward end, salinities are relatively high, tidal amplitude is rela- tively great and nutrient concentrations tend to be lower. At either end of the gradient, the energetic costs are high and a large percentage of GPP is used for self-maintenance; at the landward end, competition from freshwater plant species is high and at the seaward end, salinity stress may be limiting. In this scenario, the highest NPP occurs in the middle region of the gradient; salinity and tidal amplitude are high enough to limit compe- tition while tidal flushing and moderate nutrient levels enhance productivity. Hicks and Burns (1975) present data to support this hypothesis.
In addition to these hypotheses generated from field data, there have been two significant, published attempts to derive hypotheses from mathematical simu- lation models of mangroves. The first (Miller 1972) is a model of primary pro- duction and transpiration of red mangrove canopies and is based upon equations which utilize field measurements of the energy budgets of individual leaves. This model predicts a variety of interesting trends which need to be further field tested. One interesting hypothesis generated by the model is that maximum photosynthesis of red mangrove stands should occur with a leaf area index (LAI) of 2.5 if no accli- mation to shade within the canopy occurs; higher LAI's may lead to decreased produc- tion. Another prediction is that red mangrove production is most affected by air temperature and humidity and, to a lesser degree, by the amount of solar
19
|
>!•»-> |
|
|
03 Q) |
|
|
•O c cr. |
|
|
-■■» OJ |
|
|
M II +J -o |
|
|
E ■•- c |
|
|
^Q. .c ra |
|
|
o a. 2 |
|
|
aiz co |
|
|
ii |
|
|
oj •» * |
|
|
1. C3 w |
|
|
03 o >i |
|
|
•r— *• oJ |
|
|
oi +-> a> "o |
|
|
oj o > |
|
|
3 3 O >> |
|
|
r-13 1. C |
|
|
03 o en c |
|
|
> 5- C 3 |
|
|
Q. 03 00 |
|
|
r- E |
|
|
>- >i c |
|
|
< 1-TI O |
|
|
03 ai |
|
|
E i- OJ |
|
|
• •!- S_ |
|
|
itj i- ii a) |
|
|
-o a. 2 |
|
|
•t- or |
|
|
i. to r-~ |
|
|
O ul « |
|
|
i— o ro "a |
|
|
u- s- c |
|
|
cn x os |
|
|
c |
|
|
•i- II r— lO |
|
|
r— |
|
|
CO- « Ul |
|
|
O a. M- c |
|
|
■f- <J3 o |
|
|
4-> 1. •!- |
|
|
<J 0J 4-> |
|
|
3 • +j m |
|
|
T3 S- +-> > |
|
|
O >>••- i. |
|
|
s_ ^.— oj |
|
|
Q. oj ui |
|
|
SZ f— J3 |
|
|
m\ mo |
|
|
> W 3 |
|
|
o c c |
|
|
S- o c • |
|
|
01 4-> oj 0J |
|
|
C > |
|
|
03 (J II O |
|
|
E •■- S- |
|
|
s- . en |
|
|
<♦- +J u. c |
|
|
O CD -03 |
|
|
E — I E |
|
|
Ul |
|
|
0J II - ^ |
|
|
■!-> CO |
|
|
03 O- CD 03 |
|
|
E a- .,-,— |
|
|
•t- Z 4-> JD |
• |
|
+-> U |
l/l |
|
Mr- 3 II |
>> |
|
Ul 03 XJ |
03 |
|
3 O CO |
■o |
|
C S_ |
|
|
• c n. - |
>» |
|
03 03 OJ |
-a |
|
l— >1 > |
=3 |
|
+-> s_ o |
O |
|
CD Q. 03 S. |
|
|
■— 0) E cn |
U |
|
-Q CJ 1- C |
|
|
03 X S- o3 |
c |
|
1— 0) Q. E |
o |
oj cj
c
s_ a> m-
ai or
|
ai |
0) |
|
|
cn |
Ol |
|
|
c |
c |
|
|
ro |
03 |
|
|
sz |
sz |
|
|
-a |
o |
o |
|
o |
X |
X |
|
SZ |
aj |
ai |
|
+-> |
||
|
ai |
Ul |
Ul |
|
s: |
OS |
o3 |
|
C3 |
CO |
|
|
Q. |
||
|
Q_ |
a.
03
cd
a.
O-
cj
00
LT)
cn
3
CO
cn
|
S- |
— |
|||||||
|
a> |
||||||||
|
j* |
^~* |
^^ |
*— » |
^~^ |
■ |
|||
|
03 |
CM |
CM |
CXI |
CM |
i — |
|||
|
T3 |
- — . |
^^ |
^-^ |
r-. |
r^ |
r-^ |
r-~ |
03 |
|
ai |
ai |
cn |
Ol |
Ol |
Ol |
Ol |
Ol |
|
|
SZ |
r^ |
r-~ |
r->. |
r— |
, — |
, — |
r— |
-t-> |
|
oo |
cr> |
cn |
Ol |
"— ' |
— ' |
""-* |
|
OJ |
|
oes |
* — |
^ — |
* — |
S. |
S- |
s_ |
5- |
s_ |
|
ai |
<U |
<D |
CD |
(U |
||||
|
o |
ul |
Ul |
ul |
i — |
1 — |
i — |
i — |
■IJ |
|
cn |
03 |
03 |
03 |
• — |
1 — |
i — |
i — |
s_ |
|
3 |
CD |
ai |
CD |
•t— |
03 |
|||
|
_l |
r— |
i— |
1— |
s: |
sz |
3T |
s: |
c_> |
|
ro |
ro |
||
|
r^ |
r-~ |
- — . |
|
|
Ol |
Ol |
lO |
U3 |
|
i — |
i — |
r— |
r-~ |
|
— |
*■ — |
Ol |
Ol |
<u
+j s.
03 O
o
Ol
3
o cn
|
at |
ai |
ai |
ai |
QJ |
<u |
OJ |
0J |
|
cn |
cn |
cn |
cn |
cn |
cn |
cn |
cn |
|
c |
c |
c |
c |
c |
c |
c |
c |
|
ra |
03 |
03 |
«3 |
03 |
03 |
03 |
03 |
|
-cr |
^r |
SI |
^: |
.c |
SZ |
SZ |
sz |
|
(J |
o |
u |
u |
O |
<J |
<J |
u |
|
X |
X |
X |
X |
X |
X |
X |
X |
|
ai |
OJ |
at |
ai |
a) |
OJ |
0J |
aj |
|
ul |
Ul |
Ul |
Ul |
ul |
Ul |
Ul |
ul |
|
OS |
OS |
03 |
03 |
03 |
03 |
ro |
03 |
|
CD |
CO |
CS |
C3 |
C3 |
CO |
C3 |
C3 |
|
O |
m |
in |
CO |
IO |
CO |
r^ |
CO |
00 |
LT) |
«3- |
|
|
r— |
• |
■ |
|||||||||
|
03 |
IO |
o |
C3 |
ro |
00 |
o |
LT) |
CM |
CO |
r-^ |
r-~ |
|
3 |
^1- |
CM |
CM |
CM |
1 — |
1 — |
i — |
■ — |
CM |
||
|
C |
|||||||||||
|
C |
|||||||||||
|
< |
U3 CM
|
in |
CO |
C3 |
<* |
IT) |
ro |
UT) |
CM |
00 |
|
u-> |
00 |
,— |
CM |
LT) |
>* |
ro |
IT) |
«d- |
IS)
r-~
oo
CM
>* «*
CM
T3 X 03
CM I
i — I
O
00
ro i— r-~ is>
CO
CD ■—
UT) Ol
ro ^1-
CM Ol
U3 r—
I CM
"3-
Ol
ro CD
CM CD
Ol ro
CD Ol
ro
|
* aj |
»« |
|||||||
|
3 c |
3 |
|||||||
|
a: |
oo |
0J |
. |
OJ |
. |
az «_ |
cc c |
|
|
OJ |
c |
c |
c |
c |
OJ |
•r- |
||
|
s_ |
JZ\ |
B |
3 |
03 |
3 |
03 |
"O > |
XJ to |
|
3 |
3 |
•r- |
■"3 |
■■3 |
'D |
""3 |
aj ._ |
CD rg |
|
•t-> |
U |
Ul |
1 ' |
• — |
1 ' |
1 — ■ |
X t. |
x -a |
|
03 |
CJ |
03 |
•»— ^^ |
•^ **—••" |
||||
|
S |
oo |
CO |
a: |
nc |
az |
or |
s: co |
s co |
03
■a
OJ
u
3
■a o
s-
CL +J
o
c
Ul
OJ
o
T3
■a o
20
Table lb. Comparative measurements of photosynthesis in gC/m2/day (Lugo et al . 1975) .
Mangrove type
Daytime net photosynthesi s
Ni ghttime respi ra ti on
Pn/R
Red
Bl ack
Whi te
Red ( seedl i ng )
1 .38 1 .24 0.58 0.31
0.23 0.53 0.17 1 .89
6.0 2.3 3.4 negative
Table lc. Gross primary production (GPP) at different salinities (Hicks and Burns 1975).
Mangrove type
Average surface sal i ni ty (ppt )
GPP (gC/m2/day)
Red Red Red Bl ack Bl ack Black White White
7.8 21 .1 26.6
7.8 21 .1 26.6 21 .1 26.6
8.0 3.9 1 .6 2.3 5.7 7.5 2.2 4.8
21
HIGH
COMMUNITY NET PRIMARY PRODUCTION
LOW
LANDWARD
SEAWARD
WATERWAY POSITION
Figure 6. The hypothetical relationship between waterway position and community net primary production of Florida mangrove forests (based on Carter et al . 1973).
22
radiation within the ambient range. Gross photosynthesis per unit leaf area was greater at the top of the tree canopy than at the bottom, although the middle levels had the greatest production.
Miller (1972) concluded by suggesting that the canopy distribution of red man- grove leaves is nearly optimal for ef- ficient water utilization rather than production. This indicates that the cano- py is adapted to maximizing production under conditions of saturated water sup- ply.
The mangrove ecosystem model reported by Lugo et al. (1976) provides hypotheses on succession, time to arrive at steady state conditions (see section 3.2), and several aspects of productivity. The model output suggests that the relative amount of tidal amplitude does not affect GPP significantly; instead, GPP appears to be extremely sensitive to inputs of ter- restrial nutrients. It follows that loca- tions with large amounts of nutrient input from terrestrial sources (riverine man- grove communities) have high rates of mangrove production (see section 3.3). All simulation model -generated hypotheses need to be field tested with a particular- ly critical eye, since the simplifying assumptions that are made in constructing the model can lead to overly simplistic answers.
Mangrove productivity research re- mains in an embryonic stage. Certain preliminary tendencies or hypotheses have been identified, but much work must be done before we can conclude that these hypotheses cannot be falsified.
2.6 HERBIVORY
Direct herbivory of mangrove leaves, leaf buds, and propagules is moderately low, but highly variable from one site to the next. Identified grazers of living plant parts (other than wood) include the white-tailed deer, Odocoileus virginianus, the mangrove tree crab, Aratus pi soni i , and insects including beetles, larvae of
lepidopterans (moths and butterflies), and orthopterans (grasshoppers and crickets).
Heald (1969) estimated a mean grazing effect on North River red mangrove leaves of 5.1% of the total leaf area; values from leaf to leaf were highly variable ranging from 0 to 18%. Beever et al. (1979) presented a detailed study of grazing by the mangrove tree crab. This arboreal grapsid crab feeds on numerous items including beetles, crickets, cater- pillars, littoral algae, and dead animal matter. In Florida, red mangrove leaves form an important component of the diet. Beever et al. (1979) measured tree crab grazing ranging from 0.4% of the total leaf area for a Florida Keys overwash forest to 7.1% for a fringing forest at Pine Island, Lee County, Florida. The researchers also found that tree crab grazing rates are related to crab density. Low densities (one crab/m ) resulted in low leaf area damage (less than 1% of total leaf area). High densities (four crabs/m ) were accompanied by leaf area damage ranging from 4% to 6% (see section 6.2).
Onuf et al. (1977) investigated in- sect herbivory in fringing and overwash red mangrove forests in the Indian River estuary near Ft. Pierce, Florida. They found six major herbivorous insect species, five lepidopteran larvae and a beetle. Comparisons were made at a high nutrient site (input from a bird rookery) and a low nutrient site. Both red man- grove production and leaf nitrogen were significantly higher at the high nutrient site. This resulted in a four-fold greater loss to herbivores (26% of total leaf area lost to grazing); this increased grazing rate more than offset the in- creased leaf production due to nutrient input.
Calculations of leaf area damage may underestimate the impact of herbivores on mangroves. For example, the larvae of the olethreutid moth, Ecdytol opha sp., develops within red mangrove leaf buds and causes the loss of entire leaves. All stages of the beetle, Poeci 1 i ps
23
rhizophorae, attack mangrove propagules while still attached to the parent tree (Onuf et al. 1977).
2.7 WOOD BORERS
Many people have the mistaken idea that mangrove wood is highly resistant to marine borers. While this may be true to a limited extent for certain mangrove species in other parts of the world, none of the Florida mangroves have borer- resistant wood. Southwell and Boltman (1971) found that the wood of red, black, and white mangroves has no resistance to Teredo, Pholad and Simnorid borers; pieces of red mangrove wood were completely de- stroyed after immersion in ocean water for 14 months.
An interesting controversy surrounds the ability of the wood boring isopod, Sphaeroma terebrans, to burrow into the living prop roots of the red mangrove. Rehm and Humm (1973) were the first to attribute apparently extensive damage of red mangroves stands within the Ten Thousand Islands area of southwestern Florida to an isopod, Sphaeroma. They found extensive damage throughout southwest Florida, some infestation north to Tarpon Springs, and a total lack of infestation in the Florida Keys from Key Largo south to Key West. The destruction process was described as follows: the adult isopod bored into the prop roots (5- mm diameter hole); this was followed by reproduction within the hole and develop- ment of juveniles within the root. This process, combined with secondary decompo- sition from fungi and bacteria, frequently results in prop root severance near the mean high tide mark. These authors attributed loss of numerous prop roots and, in some cases, loss of entire trees during storms to isopod damage.
The extent of damage in the Ten Thousand Islands region led Rehm and Humm (1973) to term the phenomenon an "eco- catastrophe" of possibly great importance. They further stated that shrinking of mangrove areas appeared to be occurring as
a result of Sphaeroma infestation; this point was not documented.
Enright (1974) produced a tongue-in- cheek rebuttal, on behalf of Sphaeroma and against the "terrestrial invader", red mangroves. Snedaker (1974) contributed a more substantial argument in which he pointed out that the isopod infestation might be an example of a long-term eco- system control process.
Further arguments against the "ecoca- tastrophe" theory were advanced by Estevez and Simon (1975) and Estevez (1978). They provided more life history information for Sphaeroma and suggested a possible ex- planation for the apparently destructive isopod infestations. They found two species of isopods inhabiting red mangrove prop roots, S^. terebrans and a sympatric congener, S. quadridentatum. The latter does not appear to be a wood borer but utilizes S. terebrans burrows. Neither species appeared to utilize mangrove wood as a food source. Estevez and Simon (1975) found extensive burrowing into seedlings in addition to prop root damage. In general, infestations appeared to be patchy and limited to the periphery of mangrove ecosystems. In areas with the highest density of burrows, 23% of all prop roots were infested. There appeared to be more colonization by ^. terebrans in regions with full strength sea water (30 to 35 ppt).
The most important finding by Estevez and Simon (1975) and Estevez (1978) was that periods of accelerated activity by ^. terebrans were related to periods of fluc- tuating and slightly increased salinity. This suggests that fluctuations in isopod burrowing may be related to the magnitude of freshwater runoff from the Everglades. These authors agree with Snedaker (1974) and suggest that root and tree loss due to Sphaeroma activity may be beneficial to mangrove ecosystems by accelerating pro- duction and root germination. Simberloff et al. (1978) amplified this last sugges- tion by showing that root branching, which is beneficial to individual trees, is stimulated by isopod activity.
24
This ecocatastrophe versus beneficial stimulus argument is not completely re- solved. Probably, Sphaeroma root destruc- tion, in areas of low isopod density, can be a beneficial process to both the in- dividual tree and to the entire mangrove stand. Whether changes in freshwater runoff have accelerated this process to the point where unnatural and widespread damage is occurring is not clear. The data and research perspective to answer this question do not exist. As a result, we are reduced to providing hypotheses which cannot be tested with available knowledge.
2.8 MANGROVE DISEASES
Published research on mangrove diseases is rare. The short paper by Olexa and Freeman (1975) is the principal reference for diseases of Florida man- groves. They reported that black man- groves are affected by the pathogenic
fungi , Phyl losti eta hi bi scina and Nigro- spora sphaerica. These authors found that P. hi bi sci na caused necrotic lesions and death of black mangrove leaves. They felt that under conditions of high relative humidity coupled with high temperatures, this fungus could pose a serious threat to individual trees, particularly if the tree had been weakened by some other natural agent, such as lightning or wind damage. Nigrospora sphaerica was considered to be of little danger to black mangroves. Another fungus, Cyl i nrocarpon didymum, appears to form galls on the prop roots and stems of red mangroves. Olexa and Freeman (1975) noted mortality of red mangroves in areas of high gall infesta- tions, although a direct causation link was not proven.
Further research on mangrove diseases is badly needed. Viral disease must be investigated. The role of pathogens in litter production and as indicators of mangrove stress may be very important.
25
CHAPTER 3. ECOSYSTEM STRUCTURE AND FUNCTION
3.1 STRUCTURAL PROPERTIES OF MANGROVE FORESTS
Published information about the structural aspects of Florida mangrove forests is limited; most existing data have been published since the mid-1970's. This lack of information is unfortunate since quantitative structural data greatly aid understanding of processes such as succession and primary production. Even more important, the response of mangrove forests to stress, both climatic and man- induced, can be followed quantitatively with this type of data.
Ball (1980) contributed substantially to understanding the role of competi- tion in mangrove succession by measuring structural factors such as basal area, tree height, and tree density. Lugo and Zucca (1977) monitored the response of mangrove forests to freezing temperatures by observing changes in structural proper- ties of the trees.
Baseline studies of forest structure have been published by Lugo and Snedaker (1975), and Pool, Snedaker and Lugo (1977). For example, Lugo and Snedaker (1975) compared a fringing mangrove forest and a basin forest at Rookery Bay, near Naples, Florida. They found the fringing forest, which was dominated by red man- groves, to have a tree diversity of H = 1.48, a basal area of 15.9 mz/ha, an aboveground biomass of 17,932 g/m , and a non-existent litter layer. The nearby basin forest was dominated by black man- groves, had a tree diversity of H = 0.96 and a basal area of 23.4 m /ha. The lit- ter layer in the basin forest averaged 550 dry g/m . Tree diversity in a hurricane disturbed section of the Rookery Bay forest was 1.62. Similar data were pre- sented for mangrove forests in the Ten Thousand Islands area (Table 2).
Data of this type are useful for many purposes including impact statements, en- vironmental surveys, and basic scientific questions. Cintron et al. (1978) gave an indication of the direction in which fu- ture research might proceed. Working in a mangrove stand in Puerto Rico, they found
tree height to be inversely proportional (r = 0.72) to soil salinity in the range 30 to 72 ppt. Above 65 ppt salinity, dead tree basal area was higher than live tree basal area and above 90 ppt there was no live tree basal area.
It should be possible to investigate the relationship between a variety of mangrove structural properties and factors such as flushing frequency, soil depth, nutrient availability, pollution stress, and other measures of human impact. Ulti- mately, this should lead to an ability to predict the form and structure of mangrove forests resulting from various physical conditions or artificial impacts. One example of this potential tool is Ball's (1980) documentation of structural changes in mangrove forests resulting from altera- tions in the hydrological conditions of south Florida.
3.2 Z0NATI0N, SUCCESSION AND "LAND- BUILDING"
Much of the world's mangrove litera- ture consists of descriptive accounts of zonation in mangrove forests and the spe- cies composition within these zones. Al- thouqh general agreement has been lacking, various hypotheses have been put forth concerning the possible connection between zonation, ecological succession, competi- tion, and the role of physical factors such as soil salinity and tidal amplitude. In this section we review briefly the dominant ideas about mangrove zonation and succession and present our interpretation of the current status of knowledge.
Davis (1940), working in south Flori- da, was one of the first investigators to describe distinct, almost monospecific, zones within mangrove ecosystems. In what has become the classical view, he argued that mangrove zonation patterns were equivalent to serai stages in succession. The most seaward zone, dominated by red mangroves, was regarded as the "pioneer stage". More landward zones were dominated by white mangrove, black mangrove, buttonwood and, finally, the climatic climax, a tropical forest. Since
26
|
CD |
|
|
CD |
• |
|
s_ |
ra |
|
sz |
|
|
cn |
-^ |
|
-o |
en |
|
c |
^ |
|
ra |
|
|
1 — |
>> |
|
to |
s- |
|
1-1 |
53 |
|
-o |
C |
|
c |
•i— |
|
rO |
|
|
l/l |
-a |
|
3 |
ai |
|
o |
LO |
|
-C |
i/i |
|
r- |
CD |
|
s- |
|
|
c |
a. |
|
ai |
X |
|
l— |
0) |
|
a> |
a> |
|
.c |
5- |
|
+-> |
1a |
|
ai |
=3 |
|
|
s- |
CJ |
|
|
o 4- |
s- 03 CD |
|
|
<1J |
. — |
|
|
> |
<_> |
|
|
O |
• |
|
|
S-CM |
- — . |
|
|
en E |
LT> |
|
|
c |
r-~ |
|
|
ID |
LO |
en |
|
E 4- |
CVI |
1 |
|
O |
o |
CD |
|
C/l |
-a |
-ii |
|
Ul |
0) |
ra |
|
ra |
i/i |
■a |
|
E |
CO |
CD |
|
o |
.a |
c: |
|
•r— jQ |
CD |
oo |
|
i. |
"O |
|
|
X> |
03 |
c |
|
tz |
ro |
|
|
3 |
in |
|
|
O |
CD |
O |
|
s- |
=i |
en |
|
en |
< — |
r> |
|
CD |
ra |
_i |
|
> |
> |
|
|
O |
E |
|
|
-Q |
o |
|
|
< |
■ |
S- |
|
ra |
4- |
|
|
■a |
||
|
• |
■i — |
CD |
|
CVJ |
S- |
i- |
|
o |
ro |
|
|
CD |
1 — |
|
|
i — |
u. |
ro |
|
.a |
4-> |
|
|
rO |
n- |
ro |
|
1— |
o |
Q |
CD </l
C CD
•w- >
i- O
CD i.
> cn
••- c
or: ra
E
in
CD
<D >
cn o
c i- i- cn
i- c U_ ra
t/>
J= CD
in >
ra o
3 s-
s- cn
<D C
> ra
O E
en
<D
>
-O. O
=J S-
s- cn
o c
OO ra
E
ai E
rO
Q.
o
O
10
cn
|
o |
o |
O 1 |
, — |
|
ro |
10 |
ro I |
CO |
|
ro |
o |
cn i |
00 |
|
•• |
•* |
• i |
•» |
|
r— |
ro |
CO 1 |
r-» |
|
LO |
ro i |
o CVI |
|
o |
co |
o |
o |
CD 1 |
CO |
|
t — |
^> |
CM |
•3- |
LO |
LO |
|
00 |
r— |
LO |
LO |
cn |
r— |
|
* |
- |
*• |
•* |
» |
|
|
CO |
cn |
<3- |
CM |
f— |
|
|
r-^ |
i— |
«3" |
"3" |
|
r-^ |
,— |
O |
O |
O |
00 |
|
ro |
CO |
r— |
ct< |
i — |
f~. |
|
O |
f — |
LCI |
i — |
*r |
CM |
|
n |
it |
•* |
* |
* |
|
|
r-. |
00 |
r-. |
00 |
w— |
|
|
CM |
i — |
CTl |
LO |
||
|
•~ |
CM |
||||
|
CO |
o |
o |
o |
O |
CO |
|
«3" |
r~ |
r» |
o |
LO |
C~. |
|
CO |
CM |
CM |
CM |
CM |
r~ |
|
n |
*> |
« |
* |
•» |
|
|
Lf) |
«3" |
r-^ |
o |
r-^ |
|
|
00 |
CM |
LO |
i-» |
||
|
CM |
00 |
o |
O |
O |
O |
|
CO |
CM |
LO |
r~ |
CO |
CM |
|
cn |
cn |
CM |
r^ |
cn |
|
|
n |
«» |
•» |
* |
■» |
|
|
LO |
r>- |
CM |
CM |
oo |
|
|
LO |
CM |
CM |
o |
|
lO |
LO |
o |
ro |
o |
1 CM |
|
«3- |
co |
CO |
CM |
cy> |
1 — |
|
CTl |
CM |
«3" |
cn |
CTV |
LO |
|
« |
M |
•» |
•* |
•» |
|
|
LO |
o |
■ — |
CO |
CO |
|
|
p~ |
«* |
1 — |
CO |
|
CO |
C3 |
O |
o |
O |
CO |
|
LO |
CM |
CO |
CO |
1 — |
1 LO |
|
CM |
co |
cn |
CO |
i cn |
|
|
m |
■1 |
* |
•» |
i * |
|
|
r^ |
o |
, — |
l~~ |
1 LO |
|
|
r-» |
LO |
1— |
1 -S3- |
|
CM |
4-> |
C31 |
r- |
O |
1 00 |
|
i — |
ro |
LO |
CTl |
«* |
1 o |
|
r^ |
-o |
CTl |
r— |
■— |
1 o |
in
CD
>
CD
in
S- 0)
o
O
o
o o
i-
Q.
o s_
Q_
i.
■1-J ■u
in
in
i ra
<D E
> O
O T-
J3 -Q
-a
■— c
ra 3
■M O
o s- I— m
27
these zones were regarded as progressively later stages in succession, the entire mangrove ecosystem was believed to be moving seaward through a process of sedi- ment accumulation and colonization. Davis based his argument primarily upon the sequence of observed zones and cores which showed red mangrove peat underlying black mangrove peat which, in turn, occurred under terestrial plant communities.
Unfortunately, this Clementsian in- terpretation of mangrove zonation was widely accepted, but rarely tested. For example, Chapman (1970) expanded Davis' original successional concept from south Florida to explain zonation in mangrove forests in other parts of the world. Walsh (1974) thoroughly reviewed the man- grove succession/zonation literature.
Fortunately, not everyone accepted Davis' point of view. Egler (1952) and later Thorn (1967, 1975) argued that man- grove zonation was a response to external physical forces rather than temporal se- quence induced by the plants themselves. Egler (1952) showed that patterns of sedi- ment deposition predicted by Davis' (1940) theory did not always occur. He also showed that in some cases mangrove zones appeared to be moving landward rather than seaward. Sea level has been rising in south Florida at the rate of 1 ft (30 cm) per 100 to 150 years (Provost 1974). Spackman et al. (1966) emphasized the role of sea level change in determining changes in mangrove zonation, both through sea level rise and land subsidence. Both Egler (1952) and Spackman et al. (1966) along with Wanless (1974) and Thorn (1967, 1975) suggested that mangroves were reacting passively rather than actively to strong geomorphologi cal processes. This implies that mangroves should be regarded as "land-stabilizers" rather than "land- builders".
Furthermore, field researchers fre- quently noted that red mangroves were not always the only "pioneer species" on re- cently deposited sediment. It is not unusual to find seedlings of black, white, and red mangroves growing together on a new colonization site. Lewis and Dunstan
(1975) found that black mangroves and white mangroves along with the saltmeadow cordgrass, Spartina patens, are often the pioneers on new dredge spoil islands in central Florida. On the northern coast of the Gulf of Mexico, where black mangrove is the only mangrove species present, it may be preceded by marsh grasses such as saltmarsh cordgrass, S. patens, smooth cordgrass, S. al terni fl ora, or the black needle rush, Juncus roemerianus. In Puer- to Rico, we observed that white mangrove often pioneers and dominates sites where oceanic overwash of beach sand has oc- curred. All of these observations detract from Davis' (1940) original contention that red mangroves should be regarded as the initial colonizer of recently de- posited sediments. It appears that under certain conditions, e.g., shallow water depths, substrate type, and latitude, white and black mangroves or marsh grasses can be effective pioneer species.
The work of Rabinowitz (1975) added a new perspective to the mangrove zonation debate. Through carefully designed recip- rocal planting experiments in Panamanian mangrove forests using species of Rhi zo- phora , Laguncul a ri a , P e 1 1 i c i e r a and Avicennia, she demonstrated that each species could grow well within any of the mangrove zones. In other words, physical and chemical factors such as soil salinity or frequency of tidal inundation, within each zone, were not solely responsible for excluding species from that zone. To explain zonation, Rabinowitz proposed tidal sorting of propagules based upon propagule size, rather than habitat adap- tation,as the most important mechanism for zonation control.
The most recent piece to be added to the zonation/succession puzzle comes from the work of Ball (1980). Based upon re- search of mangrove secondary succession patterns adjacent to Biscayne Bay, Flori- da, she made a strong case for the impor- tance of interspecific competition in controlling zonation. She found that white mangroves, which grow best in intertidal areas, do not occur consis- tently in the intertidal zone of mature mangrove stands. Instead, white mangroves
28
dominate higher, drier locations above mean high water where the red mangrove does not appear to have a competitive advantage. She suggested that competition is not so important during the early stages of succession but becomes critical as individual trees reach maturity and require more space and other resources.
Inherent in Ball's concept of zona- tion is the differential influence of physical factors (e.g., soil salinity, depth to water table) on the competitive abilities of the different mangrove species. She concluded that succession proceeds independently within each zone, although breaks in the forest canopy from lightning strikes or high winds may pro- duce a mosaic of different successional stages within a zone. These openings allow species whose seedlings do not com- pete well in shade, such as the white mangrove, to become established, at least temporarily, within solid zones of red mangroves.
Zonation of mangrove species does not appear to be controlled by physical and chemical factors directly, but by the interplay of these factors with interspe- cific competition and, possibly, through tidal sorting of propagules. Once succes- sion in a mangrove zone reaches an equili- brium state, change is unlikely unless an external perturbation occurs. These per- turbations range from small-scale distur- bance (lightning strikes) to large-scale perturbations (sea level change, hurricane damage) and may cause succession within zones to regress to an earlier stage. There is some evidence in south Florida that hurricane perturbations occur on a fairly regular basis, creating a pattern of cyclical succession.
Except for Ball (1980) and Taylor (1980), the importance of fires as an influence on mangrove succession has been generally ignored. Most fires in the Florida mangrove zone are initiated by lightning and consist of small circular openings in the mangrove canopy (Taylor 1980). These openings present an opportu- nity for secondary succession within an established zone. For example, we have
frequently observed white mangroves flourishing in small lightning-created openings in the center of red mangrove forests. Fire may also play a role in limiting the inland spread of mangroves. Taylor (1981) pointed out that Everglades fires appear to prevent the encroachment of red and white mangroves into adjacent herbaceous communities.
Finally, Lugo and Snedaker (1974), Cintron et al. (1978) and Lugo (1980) suggested that mangrove ecosystems function as classical successional systems in areas of rapid sediment deposition or upon recently colonized sites such as offshore islands. They concluded that in most areas mangrove forests are an example of steady-state cyclical systems. Concep- tually, this is synonymous to E. P. Odum's (1971) cyclic or catastrophic climax. Chapman (1976a, b) suggested the idea of cyclic succession for a variety of coastal ecosystems.
If Florida mangrove ecosystems are cyclic systems, then there should be an identifiable perturbation capable of set- ting succession back to an early stage. Lugo and Snedaker (1974) suggested that hurricanes may play this role. They pointed out (without substantiating data) that major hurricanes occur about every 20-25 years in south Florida. Coinci- dently, mangrove ecosystems appear to reach their maximum levels of productivity in about the same period of time (Lugo and Snedaker 1974). This hypothesis suggests that succession within many mangrove eco- systems may proceed on a cyclical basis rather than in the classical fashion. Possibly other physical perturbations may influence mangrove succession including incursions of freezing temperatures into central Florida, periodic droughts causing unusually high soil salinities (Cintron et al. 1978), and fire spreading into the upper zones of mangrove forests from ter- restrial sources.
Although understanding of zonation and succession in mangrove ecosystems remains incomplete, a clearer picture is emerging, at least for south Florida. Contrary to early suggestions, mangrove
29
species zonation does not appear to repre- sent serai stages of succession except, perhaps, for locations of recent coloniza- tion or where sediment is accumulating rapidly. The role of mangroves in land-bui'lding seems more passive than
active. Geomorphological and hydrological processes appear to be the dominant forces in determining whether mangrove shorelines recede or grow. The role of mangroves is to stabilize sediments which have been deposited by physical processes.
3.3 NUTRIENT CYCLING
Current understanding of nutrient cycles in mangrove ecosystems is far from satisfactory. Sporadic field measurements have been made, but a complete nutrient budget has not been published for any mangrove ecosystem in the world.
Several pioneering field studies were conducted in Florida (Carter et al. 1973; Snedaker and Lugo 1973; Onuf et al. 1977) and one simulation model of mangrove nu- trient cycling has been published (Lugo et al. 1976). Preliminary measurements of nitrogen fixation were made (Zuberer and Silver 1975; Gotto and Taylor 1976; Zuberer and Silver 1978; Gotto et al. 1981). Based on these studies, we present the following preliminary conclusions.
Mangrove ecosystems tend to act as a sink (net accumulator) for various ele- ments including macro nutrients such as nitrogen and phosphorus, trace elements, and heavy metals. As we have discussed in section 1.7, these elements are removed from waters flowing through mangrove swamps by the concerted action of the mangrove prop roots, prop root algae, the associated sediments, the fine root system of the mangrove trees, and the host of small invertebrates and microorganisms attached to all of these surfaces. Al- though the turnover times for these ele- ments in mangrove swamps are not known, it appears that at least a portion may be stored or tied up in wood, sediments, and peat for many years.
Although mangrove ecosystems may tend to accumulate nutrients, there is a con- tinual loss through export of particulate and dissolved substances. If significant nutrient storage and resultant high pri- mary production are to occur, there must be a continual input of nutrients to the mangrove forest from outside the system (Figure 7). Where nutrient influx to the mangrove ecosystem is approximately balanced by nutrient loss in exported organic matter, then nutrient storage will be minimal and mangrove net primary pro- duction will be low. This appears to occur in the scrub mangrove community type and to a lesser extent in the basin and hammock community types.
Carter et al. (1973) and Snedaker and Lugo (1973) have hypothesized that the greatest natural nutrient inputs for man- grove swamps come from upland and terres- trial sources. Apparently for this rea- son, the most luxuriant and productive mangrove forests in south Florida occur in riverine locations or adjacent to signifi- cant upland drainage.
Localized sources of nutrients, such as bird rookeries, can result in greater nutrient storage and higher mangrove pro- ductivity (Onuf et al. 1977). If however, large bird rookeries (or artificial nu- trient inputs) occur in poorly flushed sections of mangrove ecosystems, resultant high nutrient levels may inhibit mangrove growth (R. R. Lewis, III, Hillsborough Community College, Tampa, Fla.; personal communication 1981).
The output from the simulation model of Lugo et al. (1976) suggests that if nutrient input to a mangrove ecosystem is reduced, then nutrient storage levels within the mangrove ecosystem will be reduced and mangrove biomass and produc- tivity will decline. To our knowledge this hypothesis has not been tested in the field.
Nitrogen fixation occurs in mangrove swamps at rates comparable to those measured in other shallow, tropical marine areas (Gotto et al. 1981). Nitrogen
30
SMALL IMPORT
LOW STORAGE LOW BIOMASS LOW PRODUCTIVITY
SMALL EXPORT
LARGE IMPORT
MODERATE EXPORT
HIGH STORAGE
HIGH BIOMASS
HIGH PRODUCTIVITY
Figure 7. The hypothetical relationship between nutrient input (excluding carbon), biomass, primary productivity, and nutrient export (including carbon) from mangrove ecosystems. Top: small nutrient import. Bottom: large nutrient import.
31
fixation has been found in association with mangrove leaves, both living and dead, mangrove sediment surfaces, the litter layer in mangrove swamps, and man- grove root systems (Gotto and Taylor 1976; Zuberer and Silver 1978; Gotto et al. 1981). In virtually all cases, nitrogen fixation appears to be limited by the availability of labile carbon compounds. Perhaps for this reason, the highest rates of mangrove nitrogen fixation have been measured in association with decaying mangrove leaves; presumably, the decaying leaves act as a carbon source and thus accelerate nitrogen fixation. Macko (1981), using stable nitrogen ratio techniques, has indicated that as much as 25% of the nitrogen associated with black mangrove peat in Texas is derived from nitrogen fixation.
Zuberer and Silver (1978) speculated that the nitrogen fixation rates observed in Florida mangrove swamps may be suf- ficient to supply a significant portion of the mangrove's growth requirements. Al- though this hypothesis is impossible to test with present information, it might explain why moderately productive mangrove stands occur in waters which are severely nitrogen depleted.
In summary, knowledge of nutrient cycling in mangrove swamps is highly speculative. These ecosystems appear to act as a sink for many elements, including nitrogen and phosphorus, as long as a modest input occurs. Nitrogen fixation within the swamp may provide much of the nitrogen needed for mangrove growth.
3.4 LITTER FALL AND DECOMPOSITION
Unless otherwise stated, litter fall refers to leaves, wood (twigs), leaf scales, propagules, bracts, flowers, and insect frass (excrement) which fall from the tree. Mangrove leaves are shed con- tinuously throughout the year although a minor peak occurs during the early part of the summer wet season in Florida (Heald 1969; Pool et al. 1975). Sporadic litter fall peaks may follow periods of stress from cold air temperatures, high soil
salinities, and pollution events. Litter fall typically can be partitioned as 68% to 86% leaves, 3% to 15% twigs and 8% to 21% miscellaneous; the latter includes flowers and propagules.
Litter fall is an important ecosystem process because it forms the energy basis for detritus-based foodwebs in mangrove swamps (see sections 3.5 and 3.6). The first measurements of litter fall in man- grove swamps were made by E.J. Heald and W.E. Odum, working in the North River estuary in south Florida in 1966-69. This was subsequently published as Heald (1969), Odum (1970), and Odum and Heald (1975a). They estimated that litter pro- duction from riverine red mangrove forests averaged 2.4 dry g of organic matter/m /day (or 876 g/m /year or 8.8 metric tons/ha/year).
Subsequent studies agreed with this early estimate (Table 3), although varia- tion clearly exists between different types of communities. Scrub forests with scattered, very small trees have the smallest amount of leaf fall. Basin and hammock forests, which appear to be nutrient limited, have intermediate leaf fall values. Not surprisingly, the highest values occur in the highly produc- tive fringing, overwash, and riverine forests. Odum and Heald (1975a) suggested that the relatively uniform litter fall values from productive mangrove forests around the world result from the shade intolerance of the canopy leaves and the tendency for the canopy size to remain the same in spite of increasing height. If detailed information is lacking, red man- grove forests of south Florida, which are not severely limited by lack of nutrients, can be assumed to produce litter fall of 2.0 to 3.0 g/m /day of dry organic matter. Pure stands of black mangroves usually have a lower rate of 1.0 to 1.5 g/nr/day (Lugo et al. 1980).
Decomposition of fallen Florida man- grove leaves has been investigated by a number of researchers including Heald (1969), Odum (1970), Odum and Heald (1975a), Pool et al. (1975), Lugo and Snedaker (1975), Twilley (1980) and Lugo et
32
|
,— |
3 |
|
te |
|
|
<t- |
II |
|
i- 3 |
|
|
0) |
|
|
+-> |
• |
|
4-> |
01 |
|
t- |
> |
|
i — |
o |
|
S- |
|
|
■— |
o> |
|
ro |
c |
|
+J |
<o |
|
o |
E |
|
1— |
|
|
■o |
|
|
at |
|
|
• |
s_ |
|
00 |
|
|
+-> |
ii |
|
to |
|
|
OJ on |
|
|
J- |
|
|
o |
|
|
t- |
• |
|
^ |
|
|
OJ |
s_ |
|
> |
rO |
|
o |
-Q |
|
t- |
|
|
en -a |
|
|
c |
c |
|
rO |
ra |
|
E |
|
|
n |
|
|
C |
t/> |
|
•p- |
{. |
|
ai |
|
|
i — |
3 |
|
■ — |
o |
|
rO |
f— |
|
>+- |
<4- |
|
s- |
« . |
|
oj |
00 0J |
|
*-> |
en > |
|
■t-> |
i- o |
|
•n- |
3 s- |
|
. — |
*-> en |
|
00 |
•r— |
^ |
|
01 |
3 |
CJ |
|
+-> |
s- |
rO |
|
ro |
4- |
1 — |
|
E |
„ |
-Q |
|
+-> |
ol |
II |
|
00 |
<V |
|
|
LLl |
> ia |
CO |
|
cu |
* |
|
|
• |
»— |
OJ |
|
OO |
> |
|
|
00 |
o |
|
|
ai |
OJ |
s- |
|
r— |
T3 |
CT1 |
|
JD |
3 |
C |
|
rO |
i — |
ro |
|
1— |
O |
E |
0J
u
c aj
OJ
on
i- >,
ro
S_ -C
OJ -^ +-> oo
4-> C
•i- o
r— O
fO T-
3 S-
C +->
C 0J
< E
rO
ra
OJ ^ ■MCM
ra i—
o ro I— >»-
|
, — |
>> |
|
« — |
rO |
|
fO |
-o |
|
<4- |
' — . |
|
CM |
|
|
4- |
E |
|
ro |
^■^ |
|
O) |
en |
u a> a.
CO
|
IT) |
in |
tr> |
LO |
|
r~ |
r-~ |
r-~ |
r-^ |
|
en |
Ol |
en |
en |
|
CTl |
i — |
i — |
, — |
i — |
|||
|
U3 |
ra |
rO |
ro |
ro |
G1 |
cn |
en |
|
en |
r>~ |
r-» |
i^ |
||||
|
, — |
■!-> |
■!-> |
4-> |
4-> |
en |
en |
en |
|
OJ |
OJ |
0J |
0J |
• — |
i — |
i — |
|
|
T3 |
00 |
00 |
00 |
||||
|
03 |
o |
O |
o |
o |
ro |
ra |
ro |
|
OJ |
o |
O |
o |
o |
OJ |
OJ |
OJ |
|
zn |
a. |
a. |
a. |
a. |
h- |
1— |
(— |
O O LO
co co r-»
en en en
o r— I— .—
co o
en co
i — ro ro ro
aj >, +j +j +j
c 0J OJ OJ OJ
+J r—
S- r— O O r—
3-1- CJ> cn O
O 3 3 3 O
O I— _J _J Q.
en en
|
ra |
c |
|
3 |
|
|
+-> |
CO |
|
0J |
|
|
oS^-% |
|
|
o a. |
|
|
ro |
+-> OJ |
|
aj |
o s- |
|
nz |
co a. |
r-.
oo
cm en
CM
00 00
O O0
o
CM
|
c |
c |
00 |
*~* |
c |
^-^ |
|||
|
1— |
•f— |
rO |
ai |
•r" |
01 |
^-^ |
. — * |
^^ |
|
i. |
i- |
3 |
en |
00 |
S- |
J3 |
c |
c |
|
OJ |
0) |
(_ |
c |
rO |
3 |
3 |
•f~ |
•f- |
|
> |
> |
01 |
•t— |
-O |
+-> |
s- |
00 |
00 |
|
't— |
•r— |
> |
i- |
|
rO |
o |
ra |
rO |
|
s_ |
s_ |
o |
•*- |
E |
OO |
-Q |
JO |
ai ><
|
OJ |
||
|
a. |
||
|
>> |
||
|
4- |
+j |
t/> ra |
|
o |
>> |
OJ f- |
|
+J |
•i— r— |
|
|
>> |
■r- |
u m |
|
+-> |
c: |
OJ S_ |
|
a> |
3 |
CL4-> |
|
r- |
E |
(/> 00 |
|
i. |
F |
3 |
|
rO |
o |
io <: |
|
;» |
CJ |
CM |
33
al. (1980). Heald and Odum showed that decomposition of red mangrove leaves proceeds most rapidly under marine condi- tions, somewhat more slowly in freshwater, and very slowly on dry substrates. For example, using the litter bag method, they found that only 9% of the original dry weight remained after 4 months in sea water. By comparison, 39% and 54% re- mained at the end of comparable periods in brackish water and freshwater. Under dry conditions, 65% remained. Higher decompo- sition rates in sea water were related to increased activity of shredder organisms, such as crabs and amphipods.
Heald (1969) and Odum (1970) also found increases in nitrogen, protein, and caloric content as mangrove leaves pro- gressively decayed. The nitrogen content of leaves decaying under brackish condi- tions (on an AFDW basis) increased from 1.5% (5.6% protein) to 3.3% (20.6% protein) over a 6-month period. Subse- quent information (Odum et al. 1979b) suggested that the protein increase may not have been this great since some of the nitrogen increase probably included non- protein nitrogen compounds such as amino sugars. Fell and Master (1973), Fell et al. (1980), Fell and Newell (1980), and Fell et al. (1980) have provided more detailed information on red mangrove leaf decomposition, the role of fungi in decom- position (see section 4), and nitrogen changes and nitrogen immobilization during decomposition. Fell et al . (1980) have shown that as much as 50% of weight loss of the leaf during decomposition is in the form of dissolved organic matter (DOM).
Heald et al. (1979), Lugo et al. (1980) and Twilley (1980) discovered that black mangrove leaves decompose more ra- pidly than red mangrove leaves and ap- parently produce a higher percentage of DOM. Pool et al. (1975) have shown that mangrove litter decomposes and is exported most rapidly from frequently flooded riverine and overwash forests. These communities have little accumulation of litter on the forest floor. Communities which are not as well-flushed by the tides, such as the basin and hammock
forests, have slower rates of decomposi- tion and lower export rates.
3.5 CARBON EXPORT
Research from Florida mangrove swamps forms a small portion of the larger con- troversy concerned with the extent to which coastal wetlands export particulate organic carbon (reviewed by Odum et al. 1979a). Available evidence from Florida, Puerto Rico and Australia (Table 4) sug- gests that mangrove swamps tend to be net exporters. The values in Table 4 should be regarded as preliminary, however, since all five studies are based upon simplistic assumptions and methodology.
Golley et al. (1962) based their annual estimate of particulate carbon export from a Puerto Rican forest upon a few weeks of measurements. Odum and Heald's estimates were derived from two or three measurements a month. All investi- gators have ignored the importance of bed load transport and the impact of extreme events. All investigators except Lugo et al. (1980) have failed to measure DOC flux.
It seems relatively clear that man- grove forests do export organic carbon to nearby bodies of water. The magnitude of this export has probably been underesti- mated due to ignoring bedload, extreme events, and DOC.
The value of this carbon input to secondary consumers in receiving waters is not clear. As shown in section 3.6, food webs based primarily upon mangrove carbon do exist. The relative importance of mangrove carbon to Florida coastal ecosys- tems remains speculative. We suspect that mangrove-based food webs are dominant in small bays, creeks and rivers within large mangrove ecosystems such as the North River system studied by Heald (1969) and Odum (1970). In intermediate-sized bodies of water, such as Rookery Bay near Naples, Florida, mangroves are probably important but not dominant sources of organic car- bon. Lugo et al. (1980) estimate that mangroves supply 32% of the organic carbon
34
Table 4. Estimates of particulate carbon export from mangrove forests. Lugo et al . (1976) estimated export from a theoreti- cal, steady state forest using a simulation model. Lugo et al . (1980) measured export from an inland black mangrove forest.
Investigators
Location
Export
g/m /day tonnes/ha/yr
Golley et al . (1962) Heald (1969), Odum (1970)' Lugo and Snedaker (1975) Lugo et al . (1976) Boto and Bunt (1981) Lugo et al. (1980)b
^Estimate only includes carbon of mangrove origin. Estimate includes dissolved and particulate carbon.
|
Puerto Rico |
1.1 |
4.0 |
|
Florida |
0.7 |
2.5 |
|
Florida |
0.5 |
2.0 |
|
Florida |
1.5-1.8 |
5.5 - 6.6 |
|
Australia |
1.1 |
4.0 |
|
Florida |
0.2 |
0.7 |
35
input to Rookery Bay. In very large sys- tems, such as Biscayne Bay near Miami, Florida, mangroves are clearly less impor- tant than any other sources such as algae and sea grasses, although mangrove carbon may be important in localized situations such as the immediate vicinity of fringing and overwash forests. The magnitude of mangrove carbon export to unenclosed coastal waters and offshore remains a mystery.
3.6 ENERGY FLOW
At least seven sources of organic carbon may serve as energy inputs for consumers in mangrove ecosystems (Figure 8). The pathways by which this energy containing material is processed and made available to each consumer species is indeed complex. Not surprisingly, current understanding of energy flow in Florida mangrove ecosystems exists largely in a qualitative sense; quantitative data are scarce and piecemeal. A variety of inves- tigators have contributed information over the past decade including, but not limited to, Heald (1969), Odum (1970), Odum and Heald (1972), Carter et al. (1973), Snedaker and Lugo (1973), Heald et al. (1974), Lugo and Snedaker (1974, 1975), Odum and Heald (1975a, b), and Pool et al. (1977). Probably, the most complete study to date is the investigation of energy flow in the black mangrove zone of Rookery Bay by Lugo et al. (1980).
It is possible at this time to pre- sent a series of hypotheses concerning the relative importance of these energy sources. First, the relative importance of each source can vary from one location to the next. As will be shown in the following discussion, the consumers in certain mangrove forests appear to depend primarily upon mangrove-derived carbon while in other locations inputs from phy- toplankton and attached algae are probably more important.
Our second hypothesis is that energy flow based upon phytopl ankton is most important in overwash mangrove forests and other locations associated with large
bodies of clear, relatively deep water. Conversely, phytoplankton are hypothesized to be relatively unimportant to the energy budgets of the large riverine forest com- munities along the southwest coast of Florida. It should be remembered, how- ever, that even where phytoplankton are quantitatively unimportant, they poten- tially perform an important function as the basis of phytopl ankton-zoopl ankton- larval fish food webs (Odum 1970).
As a third hypothesis, Iver Brook (Rosensteil School of Marine and Atmos- pheric Sciences, Rickenbacker Causeway, Miami, Fla.; personal communication 1979) has suggested that both sea grasses and benthic algae serve as an important energy source for fringing mangrove communities adjacent to large bodies of water such as Biscayne Bay and Whitewater Bay. Although little evidence exists to test this hypo- thesis, observations of extensive deposits of sea grass and macroalgal detritus with- in mangrove forests suggest intuitively that Brook's hypothesis may be correct.
In regions where mangrove shading of the prop roots is not severe, our fourth hypothesis suggests that carbon origina- ting from prop root epiphytes may be sig- nificant to community energy budgets. Lugo et al. (1975) have measured net pro- duction of periphyton in mangroves fringing Rookery Ba^ and found average values of 1.1 gC/m /day. Hoffman and Dawes (1980) found a lower value of 0.14 gC/mVday. Because these values are roughly comparable to average exports of mangrove leaf carbon (section 3.5), its potential importance is obvious.
The fifth hypothesis states that mangrove organic matter, particularly leaf material, is an important energy source for aquatic consumers. This hypothesis was first espoused by Heald (1969) and Odum (1970), who worked together in the riverine mangrove communities between the Everglades and Whitewater Bay. Clearly, mangrove carbon is of great importance within the riverine and basin communities all along the southwest coast of Florida (Odum and Heald 1975b); Carter et al . (1973) and Snedaker and Lugo (1973)
36
|
to >> |
|
|
>,*♦- as |
|
|
nj O 3 |
|
|
3 |
|
|
x: >, o |
|
|
•(-> c c |
|
|
<o UJ |
|
|
a. c |
|
|
e i- |
|
|
0) O |
|
|
.— s- -a |
|
|
X! M- C |
|
|
•»- ro |
|
|
in aj |
|
|
Ul +-> tO |
|
|
o <0 t- |
|
|
ac » |
|
|
•r- as |
|
|
.— cox: |
|
|
■— -r- a. |
|
|
O S- E |
|
|
O CO |
|
|
■!-> |
|
|
O "O S- |
|
|
^ i— o |
|
|
ZJ M- |
|
|
O |
|
|
• O XI |
|
|
to a) |
|
|
Ecu |
|
|
cu o c |
|
|
4-> •!- (O |
|
|
10 -t-> XI |
|
|
>> o c |
|
|
to 3 (I) |
|
|
o -o |
|
|
U 11 11 |
|
|
CO 4. $- |
|
|
\ to |
(0 |
|
» oc |
CD 4- |
|
Ul |
> ZS to |
|
UJ => 3: to o z: |
O <4- >> CO 3 3 |
|
•-i o |
C to XI |
|
:r o |
«J ••-» |
|
E "O « |
|
|
C CL |
|
|
C IO |
|
|
1- T3 |
|
|
to cu |
|
|
S -r- to |
|
|
o to lU |
|
|
i- 11^3 |
|
|
<l- C 1 |
|
|
at aj |
|
|
>> en > |
|
|
CO o o |
|
|
J- c i- |
|
|
0) * Ol |
|
|
c x: c |
|
|
co +-> <a |
|
|
CD S |
|
|
>»- E |
|
|
o |
|
|
to ai t- |
|
|
>>.— CO |
|
|
as CL+J |
|
|
3 E +» • |
|
|
JZ as as OJ |
|
|
■>-> X E <~> |
|
|
O 31 C |
|
|
a. u as |
|
|
S. -i- +J |
|
|
.— o c s- |
|
|
iO <*- as O |
|
|
•r- oia |
|
|
■!-> S- E |
|
|
C •« O T- |
|
|
0) c |
|
|
•4-> 3 <4- CU |
|
|
O nj O > |
|
|
Q. S. ■.- |
|
|
■o to +J |
|
|
cu io |
|
|
• C <J i— |
|
|
00 CO S- CO |
|
|
0) 3 L |
|
|
CO XI o |
|
|
$_ to >> |
|
|
3 CO i— |
|
|
CO > CO Q. |
|
|
i- <o x: E |
37
provided subsequent supportive data. What is not clear, is the relative importance of mangrove carbon to consumers within fringing, overwash, and more isolated manqrove communities.
Our sixth hypothesis involves the assemblage of organisms that graze man- grove leaves directly. A variety of in- sects (see section 6) and the mangrove tree crab, Aratus pi soni i , (Beever et al. 1979) obtain much of their energy directly from living mangrove leaves, even though grazing rarely exceeds 10% of net primary production (Odum and Heald 1975b).
As a seventh hypothesis we suggest that anaerobic decomposition of mangrove tissue, particularly root material, may support an extensive food web based on bacteria associated with methanogenesis or the processing of reduced sulfur com- pounds. Our suggestion of the importance of reduced sulfur comes directly from Howarth and Teal's (1980) discovery of this potentially important energy pathway in temperate Spartina (cordgrass) marshes. They found that anaerobic decomposition is such an incomplete process that if sul- fates are available (from sea water) as much as 75% of the original energy in plant tissues may be converted by sulfur reducing bacteria to reduced sulfur com- pounds such as hydrogen sulfide and py- rite. Subsequently, if these reduced sulfur compounds are moved hydrologically to an oxidized environment (sediment sur- face or creek bank) sulfur -oxidizing bac- teria (e.g., Thiobacillus spp.) may convert the chemically stored energy to bacterial - ly stored energy with an efficiency as great as 50% (Payne 1970). Presumably, deposit-feeding organisms such as grass shrimp (Pal aemonetes) and mullet (Mugil ) are capable of grazing these sulfur- oxidizing bacteria from the sediment surface. If this hypothetical trophic exchange does exist, it may be of con- siderable magnitude and may cause us to reexamine current concepts of energy pro- cessing and export from mangrove ecosystems. Since freshwater contains remarkably little sulfate in comparison to seawater, this energy pathway is probably of little importance in mangrove forests
of very low salinity.
Carbon inputs from terrestrial sources may be important to certain man- grove communities. Carter et al. (1973) have shown that terrestrial carbon can reach coastal ecosystems particularly where man has cut deep channels inland for navigation or drainage purposes. The magnitude of this influx has not been adequately measured although Carter et al. did find that mainland forests (including mangroves) contributed approximately 2,100 metric tons of carbon per year to Fahkahatchee Bay.
Atmospheric inputs from rainfall appear to be minimal in all cases. Lugo et al. (1980) measured throughfall (preci- pitation passing through the tree canopy) in Rookery Bay mangrove forests of 15 to 17 gC/m /year. This would be an overesti- mate of atmospheric input since it con- tains carbon leached from mangrove leaves. The best guess of atmospheric input is between 3 to 5 gC/m /year for south Florida mangrove ecosystems.
Subsequent stages of energy transfer in mangrove community food webs remain largely hypothetical. Odum (1970) and Odum and Heald (1975b) have outlined several pathways whereby mangrove carbon and energy are processed by a variety of organisms (see Figure 8). Apparently, the most important pathway follows the se- quence: mangrove-leaf detritus substrate- microbe-detritus consumer-higher consu- mers. The critical links are provided by the microbes such as bacteria and fungi (see Fell et al. 1975) and by the detritus consumers. The latter group was studied by Odum (1970) and Odum and Heald (1975b) and found to consist of a variety of invertebrates (e.g., caridean shrimp, crabs, mollusks, insect larvae, amphipods) and a few fishes.
Stable carbon studies such as those done by Haines (1976) in Spartina (cordgrass) marshes have not been per- formed in mangrove ecosystems. Mangroves are C3 plants and have 613 values in the range of minus 25 to minus 26 (Macko 1981). According to the same author,
38
mangrove peat has a 613 value of minus 22. Because these values are dramatically different from the values for sea grasses and many algae, the possibilities for using this tool in mangrove ecosystems is excellent. Macko (1981) also suggested the utility of using stable nitrogen ra- tios for future mangrove food web investi- gations; he reported 615 values of plus 6.0 to plus 6.5 for mangrove tissue and plus 5 for mangrove peat.
In reviewing contemporary knowledge of energy flow in mangrove ecosystems, three conclusions emerge.
(1) We have a hypothetical framework of mangrove energy flow of a qualitative
nature. This framework appears to be reasonably accurate although subsequent developments, such as elucidation of the reduced sulfur hypothesis, may require some modification.
(2) Measurements of the relative importance of various carbon sources are generally lacking.
(3) Detailed measurements of energy flow including the relative inputs of different carbon sources are critically needed. Technological difficulties, high costs, and difficulties inherent in transferring findings from one estuary to the next present a major challenge to estuarine ecologists of the future.
39
CHAPTER 4. COMMUNITY COMPONENTS - MICROORGANISMS
The mycoflora (fungi) are the best Drechslera and Gloeosporium. In the lat-
studied component of the microbial com- ter stages of decay the dominant genera
munity of mangrove swamps. Much pio- are Calso, Gliocidium, and Lulworthia. neering work has been carried out in south
Florida. Reviews of the current knowledge Understanding the occurrence and suc-
of mangrove-associated fungi can be found cession of fungi on decaying mangrove
in Kohlmeyer and Kohlmeyer (1979) and Fell leaves is important because of their role
et al . (1980). in energy flow in mangrove swamps. Heald
(1969), Odum (1970) and Odum and Heald
One of the earliest studies of man- (1975b) hypothesized that fungi and bac-
grove mycoflora was published by Kohlmeyer teria are important in converting mangrove
(1969). He discovered large populations leaf organic material into a form that can
of marine fungi on the submerged parts of be digested and assimilated by detriti-
aerial roots, stems, and branches and on vores (see section 3.6). living and dead mangrove leaves. Exten- sive work at the University of Miami by
Fell and his coworkers (e.g., Fell and Our understanding of the role and Master 1973; Fell et al. 1975, 1980) ex- occurrence of bacteria in mangrove swamps plored the role of fungi in the decom- is not as well documented as for fungi, position of mangrove leaves and the im- Casagrande and Given (1975) have suggested mobilization of nitrogen. Newell (1974) that bacteria are important in the early studied the succession of mycoflora on stages of mangrove leaf decomposition and seedlings of red mangrove. A survey of are replaced in the latter stages by funqi the aquatic yeasts occurring in the south which are better equipped to attack re- Florida mangrove zone was published by fractive organic compounds. Unlike the Ahearn et al. (1968). mycoflora, the bacteria are clearly impor- tant in the anaerobic regions of mangrove
One of the most interesting pieces of swamps. Vankatesan and Ramamurthy (unpubl .
information to emerge from this extensive data) found denitrifying bacteria to be
mycoflora research concerns the succession abundant and ubiquitous in mangrove soils,
of orqanisms associated with decaying Zuberer and Silver (1978) have emphasized
leaves (summarized by Fell et al. 1975, the importance of nitrogen-fixing bacteria
1980). Senescent leaves of red mangroves in the zone around mangrove roots. They,
are typically colonized by species of in fact, were able to isolate and count a
Nigrospora, Phyllostica, and Pestalotica. variety of types of bacteria from mangrove
Once the leaf has fallen from the tree and sediments including aerobic heterotrophs,
during the early stages of decay, the anaerobic heterotrophs, nitrogen-fixing
fungal flora is dominated by species of heterotrophs, and sul fate-reducing bac-
Phytophthora and, to a lesser extent, teria.
40
CHAPTER 5. COMMUNITY COMPONENTS
PLANTS OTHER THAN MANGROVES
5.1. ROOT AND MUD ALGAE
The aerial root systems of mangroves provide a convenient substrate for at- tachment of algae. These root algal com- munities are particularly noticeable on red mangrove prop roots but also occur to a lesser extent on black mangrove pneumatophores located in the intertidal zone. Productivity of prop root algal communities can be appreciable if shading by mangroves is not too severe; as dis- cussed in section 3.6, Lugo et al. (1975) found a prop root community net primary production rate of 1.1 gC/m /day, a level comparable to mangrove leaf fall. Biomass of these algae can be as high as 200 to 300 g per prop root (Burkholder and Almodovar 1973). Of course, production of this magnitude only occurs on the edge of the forest and is virtually nil in the center of the swamp. Nevertheless, this algal carbon has considerable potential food value either to direct grazers or detriti vores.
Vertical distribution of prop root algae has been studied by many researchers (Gerlach 1958; Almodovar and Biebl 1962 Biebl 1962; Post 1963; Rutzler 1969 Burkholder and Almodovar 1973; Rehm 1974, Yoshioka 1975); only one of these studies (Rehm 1974) was conducted in Florida. There is a tendency for certain genera of algae to form a characteristic association on mangrove roots around the world (Post 1963). Four phyla tend to dominate: Chlorophyta, Cyanophyta, Phaeophyta, and Rhodophyta; the last is usually the most important in terms of biomass. Of 74 species of marine algae recorded as prop root epiphytes between Tampa and Key Largo, 38 were Rhodophyta, 29 Chlorophyta, 4 Phaeophyta and 3 Cyanophyta (Rehm 1974).
Zonation to be expected on Florida mangroves is shown in Figure 9; this se- quence comes largely from Taylor (1960). Near the high water mark, a green band usually exists which is dominated by spe- cies of Rhi zocl oni urn. Below this is a zone dominated by species of Bostrychia, Catenel 1 a, and Cal ogl ossa. It is this association that most people think of when mangrove prop root algae are mentioned.
Because much mud is often deposited on the Bostrychia-Catenell a-Cal ogl ossa complex, it often has a dingy, gray appearance. There are many other algae found in this zone, but these three genera usually domi- nate. At brackish or nearly freshwater locations, they are replaced by species of Batophora, Chaetomorpha , CI adophora, and P e n i c i 1 1 u s . The pneumatophores of Avicennia, when colonized, are often covered with species of Rhi zocl oni urn, Bostrychia and Monostroma (Taylor 1960). Hoffman and Dawes (1980) found that the Bostrychia binderi -dominated community on the pneumatophores of black mangroyes had -tanding crop of 22 g dry wt/m and a production of 0.14 gC/mz/day.
a s
net
If there is a permanently submerged portion of the prop root, it may be covered with rich growths of Acanthophora, Spyri da, Hypnea, Laurencia, Wrangel i a, Valonia, and Caulerpa (Almodovar and Biebl 1962). Additional genera which may be present below mean high water are: Murrayel 1 a, Polysi phoni a, Centroceras, Wurdemannia , D i c tyota , Hal i m ed a , Lau renci a , and Da sya [Taylor 1960; Burkholder and Almodovar 1973; Yoshioka 1975). In addition, anywhere on the moist sections of the prop roots there are usually epiphytic diatoms and filamentous green and blue-green algae of many genera.
Rehm (1974) found a significant dif- ference in the prop root algae between south and central Florida. South of Tampa Bay the standard Bostrychia-Catenel 1 a- Caloglossa dominates. In the Tampa Bay area, species of the orders Ulotrichales and Cladophorales are dominant.
The mud adjacent to the mangrove root community is often richly populated with a variety of algae. These can include species of CI adophoropsi s, Enteromorpha, Vaucheria, and Boodleopsis (Taylor I960) in addition to a whole host of benthic diatoms and dinofl agel 1 ates (Wood 1965) and other filamentous green and blue-green algae (Marathe 1965).
Adjacent to mangrove areas, on the bottoms of shoals, shallow bays and creeks, there is often a variety of
41
PLANTS
ANIMALS
Rhizi
MHW-
Bostryt Catenei
Caloglt
MLW--
Acanthopi
Caulerpa
Wranglela
Figure 9. Vertical distribution of selected algae and Invertebrates on red mangrove prop roots (compiled from Taylor 1960 and our own observations).
42
tropical algae including species of Caul erpa , Acetabularia, P e n i c i 1 1 u s , G r a c i 1 a r i a , H a 1 i m e d a , Sargassum, Batophora , Udotea, and Dasya. These are discussed at length by Zieman (in prep.). Other pertinent references for mangrove regions include Davis (1940), Taylor (1960), Tabb and Manning (1961), and Tabb et al. (1962).
5.2 PHYT0PLANKT0N
All aspects of phytoplankton, from seasonal occurrence to productivity studies, are poorly studied in mangrove ecosystems. This is particularly true in Florida.
Evidence from Brazil (Teixeira et al. 1965, 1967, 1969; Tundisi 1969) indicates that phytoplankton can be an important component of the total primary production in mangrove ecosystems; just how important is not clear. Generally, standing crops of net phytoplankton in mangrove areas are low (personal observation). The nanno- plankton, which have not been studied at all, appear to be most important in terms of total metabolism (Tundisi 1969). The net plankton are usually dominated by diatoms such as Thai assothri x s p p . , Chaetoceras s p p . , Ni t zsc hi a s p p . , Skeletonema spp., and Rhi zosol eni a spp. (Mattox 1949; Wood 1965; Walsh 1967; Bacon 1970). At times, blooms of dinoflagel- lates such as P e r i d i n i u m spp. and Gymnodi ni urn spp. may dominate (personal observation). In many locations, particu- larly in shallow waters with some turbu- lence, benthic diatoms such as Pleurosigma spp., Mastogloia spp., and Disploneis may be numerically important in the net plank- ton (Wood 1965).
Understanding the mangrove-associated phytoplankton community is complicated by the constant mixing of water masses in mangrove regions. Depending upon the location, the phytoplankton may be domi- nated by oceanic and neritic forms, by true estuarine plankton, and by freshwater plankton. The pattern of dominance may change daily or seasonally depending upon the source of the principal water mass.
Before we can understand the impor- tance (or lack of importance) of phyto- plankton in mangrove regions, some ques- tions must be answered. How productive are the nannoplankton? How does the daily and seasonal shift in phytoplankton domi- nance affect community productivity? Does the generally low standing crop of phyto- plankton represent low productivity or a high grazing rate?
5.3 ASSOCIATED VASCULAR PLANTS
Four species of aquatic grasses occur on bay and creek bottoms adjacent to man- grove forests. Turtle grass, Thai assia testudinum, and manatee grass, Syringodium fi Hi forme, are two tropical sea grasses which occur in waters with average salini- ties above about 20 ppt. Shoal grass, Hal odule wri ghti i , is found at somewhat lower salinities and widgeongrass, Ruppia maritima, is a freshwater grass which can tolerate low salinities. These grasses occur throughout south Florida, often in close juxtaposition to mangroves. Zieman (in prep.) presents a thorough review of sea grasses along with comments about possible energy flow linkages with mangrove ecosystems.
For example, along the southwest coast between Flamingo and Naples, marshes are scattered throughout the mangrove belt and also border the mangroves on the upland side. The estuarine marshes within the mangrove swamps have been extensively
eel 1 ul osa, glass wort, Sal i cornia spp., Gulf cordgrass, Spartina spartinae, sea purslane, Sesuvium portulacastrum, salt wort, Batis maritima, and sea ox-eye, B o r r i c h i a f rutescens. Farther north, above Tampa on the west coast of Florida, marshes populated by smooth cordgrass,
43
Spartina al terni f 1 ora, and black needle rush, Juncus roemerianus, become more extensive and eventually replace mangrove swamps. Even in the Everglades region, the saline marshes are comparable to man- groves in areal extent, although they tend to be some distance from open water. Studies of these marshes, including as- sessment of their ecological value, are almost non-existent. Certainly, they have considerable importance as habitat for small fishes which, in turn, support many of the nesting wading birds in south Florida (see section 9).
Tropical hardwood forests may occur within the mangrove zone in south Florida, particularly where old shorelines or areas of storm sedimentation have created ridges 1 m or more above MSL (mean sea level) (Olmstead et al. 1981). Similar forests or "hammocks" occur to the rear of the mangrove zone on higher ground. Typical trees in both forest types include the fan palm, Thri nax r a d i a t a , buttonwood, Conocarpus erecta, manchineel, Hippomane manci nel 1 a, and, in the past, mahogany, Swi et en i a maha goni . Olmstead et al. (1 981 ) provide a description of these communities.
Freshwater marsh plants, such as the grasses, rushes and sedges that dominate the freshwater Everglades, are not mentioned here, although they are occasionally mixed in with small mangroves
that have become established well inland. See Hofstetter (1974) for a review of literature dealing with these plants.
Finally, a group of somewhat salt- tolerant herbaceous plants is found within stands of mangroves. They usually occur where slight increases in elevation exist and where sufficient light filters through the mangrove canopy. Carter et al. (1973) list the following as examples of members of the mangrove community: leather ferns, Acrostichum aureum and A. danaeifolium; Spanish bayonet, Yucca al oi fol ia; spider lily, Hymenocal 1 i s 1 ati fol ia; sea blite, Suaeda 1 ineari s; chaff flower, Al ternanthera ramosissima; samphire, Philoxerus vermicularis; blood- leaf, Iresine celosia; pricklypear cactus, 0 p u n t i a s t r i c t a ; marsh elder, I va f rutescens; the rubber vine, Rhabdadeni a bi f 1 ora; the lianas, Ipomoea tuba and Hippocratea volubilis; and a variety of bromeliads (Bromel i aceae).
Although the lists of vascular plants which occur in mangrove swamps may seem extensive, the actual number of species in any given location tends to be low compared to totally freshwater environ- ments (see Carlton 1977). Analogous to temperate salt marshes, mangrove swamps possess too many sources of stress, particularly from tidal salt water, to have a high diversity of vascular plant species.
44
CHAPTER 6. COMMUNITY COMPONENTS - INVERTEBRATES
6.1 ECOLOGICAL RELATIONSHIPS bryozoans, and tunicates. The most ob- vious and dominant organisms are usually
The mangrove ecosystem, with its tree barnacles, crabs, oysters, mussels, iso-
canopies, masses of aerial roots, muddy pods, polychaetes, gastropods and, tuni-
substrates, and associated creeks and cates. small embayments, offers many habitat
opportunities for a wide variety of inver- A striking characteristic of most
tebrates. While there are few comparisons mangrove swamps is the pattern of horizon-
of species richness with other types of tal and vertical zonation of invertebrates
coastal ecosystems, mangrove swamps appear (Figure 9). Characteristic vertical zona-
to be characterized by moderately high tion patterns are found on the prop roots
invertebrate species diversity. Abele (Rutzler 1969) and not so obvious horizon-
(1974) compared H' (Shannon Weaver) diver- tal distributions occur as you move back
sity of decapod crustaceans between into the center of the swamp (Warner
various littoral marine communities and 1969). Invertebrate biomass in the red
found mangrove swamps in an intermediate mangrove zone on the edge of the swamp may
position with more decapod species than be very high, often in excess of 100 dry
Sparti na marshes but considerably less g/m of organic matter in many locations
than were associated with rocky substrate (personal observation). In the center of
communities. the swamp, particularly where there is
little flooding, biomass is usually an
There is little doubt that the maze order of magnitude less; Golley et al.
of prop roots and muddy substrates under (1962) found an average of 6.4 g/m of
intertidal mangrove trees provides habitat invertebrates in the center of a Puerto
for a wide range of invertebrates and Rican mangrove swamp, fishes (Figure 10) (see section 7 for the
latter). The nursery value of the prop Mangrove-associated invertebrates can
root complex for juvenile spiny lobsters, be placed in four major categories based
Panulirus argus, is well established on trophic position: (01 sen et al. 1975; Olsen and Koblic 1975;
Little 1977; Witham et al. 1968). Ac- (1) direct grazers - limited to cording to these researchers, the phyl-
losome larvae of spiny lobsters often (a) insects and the mangrove tree
settle among the prop roots and remain crab, Aratus pisonii, all of which feed on
there for much of their juvenile lives. leaves in the mangrove canopy and The prop roots provide protection from
predators and a possible source of food in (b) a group of small invertebrates
the associated populations of small inver- which graze the prop root and mud algae
tebrates. To provide the best habitat, a directly; section of the prop roots should extend
below mean low tide. If conditions are (2) filter feeders - largely sessile
suitable, the juveniles may remain in prop root invertebrates which filter phy-
close association with the prop root com- toplankton and detritus from the water; munity for as much as 2 years until they
reach a carapace length of 60 to 70 mm. (3) deposit feeders - mobile inverte- brates which skim detritus, algae and
In addition to its value as spiny occasional small animals from the surface lobster habitat, mangrove ecosystems also of the mud and forest floor; harbor the following invertebrates: bar- nacles, sponges, polychaete worms, gastro- (4) carnivores - highly mobile inverte- pod mollusks, pelecypod mollusks, isopods, brates which feed upon the three preceding amphipods, mysids, crabs, caridean shrimp, groups in all locations from the tree penaeid shrimp, harpacticoid copepods, canopy (largely insects) to the mud sur- snapping shrimp, ostracods, coelenterates, face. Food sources in mangrove swamps and nematodes, a wide variety of insects, energy flow are discussed in section 3.6.
45
46
6.2. ARBOREAL ARTHROPOD COMMUNITY
A surprising variety of arthropods inhabit the mangrove canopy. Because they are frequently secretive or possess camouflage coloration, their numerical importance often has been overlooked. Beever et al. (1979) pointed out that arboreal arthropods have a variety of ecological roles: (1) direct herbivory on mangrove leaves, (2) predator-prey inter- actions, and (3) biomass export through frass production and leaf defoliation. Direct grazing is typically patchy in distribution. It is not unusual to find extensive stretches of mangroves that have scarcely been grazed. In nearby areas, as much as 80% of the leaves may have some damage (Beever et al. 1979). As a general rule, it is probably safe to state that healthy, unstressed mangrove stands nor- mally have less than 10% of their total leaf area grazed (Heald 1969). In many locations, percent leaf area damaged is on the order of 1% to 2% (Beever et al. 1979). There are exceptions. Onuf et al. (1977) reported biomass loss to arthropod grazers as high as 26% in a mangrove stand where growth and nitrogen content of the leaves had been enhanced by input of nu- trients from a bird rookery.
In terms of numbers of species, the dominant group of arboreal arthropods is insects. The most thorough inventory of mangrove-associated insects was conducted by Simberloff and Wilson to obtain the raw data for their papers on island bio- geography (Simberloff and Wilson 1969; Simberloff 1976). These papers list over 200 species of insects associated with overwash mangrove islands in the Florida Keys. There is no reason to expect lesser numbers in other types of mangrove com- munities, except for the mangrove scrub forests. The most thorough study of in- sect grazing on mangrove leaves is that of Onuf et al. (1977) (see section 2.6).
Although not as numerically impres- sive as the insects, the mangrove tree crab, Aratus pisonii , appears to be poten- tially as important in terms of grazing impact (Beever et al. 1979). The life history of this secretive little crab has
been described by Warner (1967). In Jamaica its numbers range from 11 to 16/m at the edge of fringing swamps to 6/nr in the center of large swamps. Beever et al. (1979) reported typical densities for a variety of sites in south Florida of 1 to 4 crabs/m . These same authors reported some interesting details about the crab: (1) the diet is omnivorous ranging from fresh mangrove leaves to caterpillars, beetles, and various insects; (2) the crab suffers highest predation pressure while in the planktonic larval stage; (3) preda- tion on the crabs while in the arboreal community is low and comes from birds such as the white ibis, raccoons, other man- grove tree crabs and, if the crabs fall in the water, fishes such as the mangrove snapper; and (4) in one location in south Florida (Pine Island Sound) they found in accordance with normal bi ogeographical theory, the highest densities of crabs associated with fringing forests and the lowest densities on distant islands, but at Sugar Loaf Key the unexpl ai nabl e reverse distribution was found.
Other invertebrates may visit the canopy from below either for purposes of feeding or for protection from high tides. Included in this group are the pulmonate gastropods, L i tt o r i na angul i f era , Ceri thidea seal ari formi s, and Mel ampus cof feus, the isopod, Li gea exoti ca, and a host of small crabs.
In summary, with the exception of a half dozen key papers, the arboreal man- grove community has been generally ig- nored. Both insects and the mangrove tree crab play significant ecological roles and may affect mangrove productivity to a greater extent than has been recognized.
6.3 PROP ROOT AND ASSOCIATED MUD SURFACE COMMUNITY
These two somewhat distinct com- munities have been lumped together because of the large number of mobile organisms which move back and forth between tidal cycles. The aerial roots are used as protective habitat and to some extent for feeding while the nearby mud substrates are used principally for feeding.
47
The prop roots support an abundance of sessile organisms. The vertical zonation of both mobile and sessile inver- tebrates has been studied extensively in other parts of the world (Goodbody 1961; Macnae 1968; Rutzler 1969; Coomans 1969; Bacon 1970; Kolehmainen 1973; Sasekumar 1974; Yoshioka 1975). Vertical zonation certainly exists on Florida red mangrove roots. The generalized scheme shown in Figure 9 essentially contains two zones: an upper zone dominanted by barnacles and a lower zone dominated by mussels, oysters and ascidians. Between mean high tide and mean tide, the wood boring isopod, Sphaeroma terebrans (discussed at length in section 2.7) is important, both numeri- cally and through the provision of numerous holes for use by other organisms (Estevez 1978).
The most complete study of the Florida mangrove prop root community is Courtney's (1975) comparison of seawall and mangrove associations. He reported an extensive list of invertebrates from man- grove prop roots at Marco Island, Florida, including: Crassost rea v i r g i n i c a , Li ttori na angul i fera, Crepidula pi ana, Diodora cayenensis, Urosalpinx perrugata, Pisania tincta, Brachidontes exustus, nine species of polychaetes, Sphaeroma terebrans , Pal aem on floridanus , Peri cl i menes longi caudatus, Synalpheus fritzmuelleri, Thor floridanus, Petrol isthes armatus, and at least eight species of crabs. The following species were found only on mangrove roots and not on seawalls: T u r i t e 1 1 a sp., Mel ongena corona , Anachi s semiplicata, Bulla striata, Hypselodoris sp., Area imbricata, Carditamera floridana, Pseudoi rus typica, and Martesia striata.
Tabb et al. (1962) and Odum and Heald (1972) reported a variety of invertebrates associated with prop roots in the White- water Bay region. Although many species coincide with Courtney's (1975) list, there are also significant differences due to the lower salinities in this region. It is probably safe to conclude that prop root communities vary somewhat from site to site in response to a number of factors
including latitude, salinity, and proxi- mity to other communities such as sea grass beds and coral reefs.
Sutherland (1980), working on red mangrove prop root communities in Venezuela, found little change in the invertebrate species composition on indi- vidual prop roots during an 18-month period. The species composition varied greatly, however, between adjacent prop roots, presumably in response to stochas- tic (chance) processes.
The mud flats adjacent to mangroves provide feeding areas for a range of in- vertebrates that scuttle, crawl, and swim out from the cover of the mangrove roots. Some emerge at low tide and feed on algae, detritus, and small invertebrates on the mud flats while they are high and dry. Others emerge while the tide is in, parti- cularly at night, and forage across the flooded flats in search of the same foods plus other invertebrates which have emerged from the mud. In many ways the mangrove-mud flat relationship is analo- gous to the coral reef (refuge) sea grass (feeding area) relationship reviewed by Zieman (in prep.). The net effect is that the impact of the mangrove community may extend some distance beyond the boundaries of the mangrove forest.
In addition to the organisms which move from the mangroves to the mud flats, there is a small group which uses the substrate adjacent to mangroves for both habitat and feeding. In the Whitewater Bay region, four crabs exploit the inter- tidal muds from the safety of burrows: Ilea pugi 1 ator, LL speciosa, U. thayeri , and Euryti urn 1 i mosum (Tabb et al. 1962). In low salinity mangrove forests of south Florida, the crayfish, Procambarus alleni, is a dominant member of the burrowing, benthic community (Hobbs 1942) as is the crab, Rhithropanopeus harrisii (Odum and Heald 1972). Both organisms are found in a remarkable number of fish stomachs.
The benthic fauna and infauna of creek and bay bottoms near mangrove forests are highly variable from one
48
location to the next. Many of these organisms, particularly the deposit and filter feeders, benefit from particulate organic matter originating from mangrove litter fall (Odum and Heald 1972, 1975b). Tabb and Manning (1961) and Tabb et al. (1962) present lists and discussions of many of the benthic invertebrates adjacent to mangrove areas of Whitewater Bay. Weinstein et al. (1977) compared the ben- thic fauna of a mangrove-lined creek and a nearby man-made canal on Marco Island. They found (1) the mangrove fauna to be more diverse than the canal fauna and (2) a higher diversity of organisms at the mouths of mangrove creeks than in the "heads" or upstream ends. Courtney (1975) found the same pattern of upstream decreases in diversity, presumably in response to decreasing oxygen concentra- tions and increasingly finer sediments.
Finally, the irregularly flooded sub- strates in the center of mangrove forests contain a small but interesting assemblage of invertebrates. The litter layer, composed largely of mangrove leaves, evi- dently includes a variety of nematodes. Due to the usual taxonomic difficulties in identifying nematodes, complete species lists do not exist for mangrove forests; however, many species and individuals are associated with the decaying leaves (Hopper et al. 1973). In addition to nematodes, the wetter sections of the swamp floor can contain mosquito and other insect larvae, polychaetes, harpacticoid copepods, isopods, and amphipods. Simberloff (1976) lists 16 species of insects associated with the muddy floor of mangrove forests. Roaming across the forest floor during low tide are several crustaceans including the mangrove tree crab, Aratus pi soni i , crabs of the genus Sesarma, and the pulmonate gastropods, Me 1 a mpu s coeffeus and Cerithidea scalariformis. Both snails clearly have the ability to graze and consume recently fallen leaves (personal observation). With favorable conditions (relatively fre- quent tidal inundation plus the presence of red mangroves) Melampus populations can
exceed 500/m and average 100 to 200/m (Heald, unpublished data). Cerithidea i s found largely in association with black mangroves ayid can reach densities of at least 400/n/.
6.4 WATER COLUMN COMMUNITY
This section is embarrassingly short; the reasons for this brevity are (1) the paucity of research on zooplankton in Florida mangrove-dominated areas and (2) our inability to discover some of the work which undoubtedly has been done. Davis and Williams (1950) are usually quoted as the primary reference on Florida mangrove- associated zooplankton, but their paper only lists zooplankters collected in two areas. Zooplankton near mangroves are probably no different from those found in other shallow, inshore areas in south Florida. Based on Davis and Williams (1950) and Reeve (1964), we can hypothe- size that the community is dominated by copepod species of genus Acartia, particu- larly Acarti a tonsa. In addition, we could expect a few other calanoid cope- pods, arrow worms (Sagi tta spp.), many fish, polychaete and crustacean larvae and eggs. Another component of the "plankton" particularly at night, are benthic amphipods, mysids, and isopods which leave the bottom to feed (personal observation).
Plankton are not the only inverte- brates in the water column. Swimming crabs, such as the blue crab, Callinectes sapidus, are plentiful in most estuarine mangrove regions of south Florida. Other swimming crustaceans include the caridean shrimp (Pal aemonetes spp. and Peri - climenes spp.), the snapping shrimp (Al pheus~spp.), and the penaeid shrimp (Penaeus spp). All of these swimming crustaceans spend considerable time on or in the benthos and around mangrove prop roots. From the economic point of view, the pink shrimp, Penaeus duorarum, is probably the most important species asso- ciated with mangrove areas (see discussion in section 11 ).
49
CHAPTER 7. COMMUNITY COMPONENTS - FISHES
Of the six mangrove community types discussed in section 1.5, fishes are an important component of four: (1) basin forests, (2) riverine forests, (3) fringe forests, and (4) overwash island forests. For convenience we have divided fringe forests into two sub-components: (a) forests which fringe estuarine bays and lagoons and (b) forests which fringe oceanic bays and lagoons. This division is necessary because the fish communities differ markedly.
Mangroves serve two distinct roles for fishes and it is conceptually impor- tant to distinguish between them. First, the mangrove-water interface, generally red mangrove prop roots, afford a rela- tively protected habitat which is particu- larly suitable for juvenile fishes. Secondly, mangrove leaves, as discussed in section 3.6, are the basic energy source of a detritus-based food web on which many fishes are dependent. The habitat value of mangroves can be considered strictly a function of the area of interface between the water and the mangrove prop roots; it is an attribute shared by all four types of mangrove communities. The importance of the mangrove detritus-based food web is dependent on the relative contribution of other forms of energy in a given environ- ment, including phytopl ankton, benthic algae, sea grass detritus, and terrestrial carbon sources. Figure 11 provides a diagrammatic representation of the rela- tive positions along a food web continuum of the four mangrove communities.
Fishes recorded from mangrove habi- tats in south Florida are listed in Appen- dix B. Although the fish communities are discussed separately below, they have been combined into certain categories in Appen- dix B; fishes from mangrove basins and riverine forests have been combined under the heading of tidal streams; fishes from fringing forests along estuarine bays and lagoons are listed under the heading of estuarine bays; fishes from oceanic bays and lagoons have been listed under oceanic bays. Since no surveys have been published specifically relating to over- wash island forests, there is no listing for this community type in Appendix B.
Site characteristics and sampling methods for these community types are summarized in Appendix A. Nomenclature and taxonomic order follow Bailey et al. (1970).
7.1 BASIN MANGROVE FORESTS
The infrequently flooded pools in the black mangrove-dominated zone provide an extreme habitat which few species of fishes can tolerate. The waters are darkly stained with organic acids and tannins leached from the thick layer of leaf litter. Dissolved oxygen is frequently low (1-2 ppm) and hydrogen sulfide is released from the sediments following physical disturbance. Salini- ties are highly variable ranging from totally fresh to hypersaline. The fish families best adapted to this habitat are the euryhaline cyprinodonts (killifishes) and the poeciliids (1 i vebearers). The killifishes include Fundulus confluentus (Heald et al. 1974), Rivulus marmoratus (M. P. Weinstein, Va. Commonwealth Univ., Richmond, Va.; personal communication 1981), Floridichthys ca rpi o, and Cyprinodon variegatus (Odum 1970"). The poeciliids include Poecilia latipinna (Odum 1970) and, the most common, Gambusia affinis (Heald et al. 1974). While the species richness of fishes in this habitat is low, the densities of fish are often very high. Weinstein Qjers. comm.) has recorded up to 38 fish/m .
All of these fishes are permanent residents, completing their life cycles in this habitat. They feed primarily on mosquito larvae and small crustaceans such as amphipods which, in turn, feed on man- grove detritus and algae. These small fishes enter coastal food webs when they are flushed into the main watercourses during high spring tides or following seasonally heavy rains. Here they are eaten by numerous piscivorous fishes in- cluding snook, ladyfish, tarpon, gars, and mangrove snappers. The alternate energy pathway for fishes of the black mangrove basin wetlands occurs when the pools shrink during dry weather, the fishes are concentrated into smaller areas, and are fed-upon by various wading birds including
50
X < fq
C (A
O UJ
5
o
5 o
Q
Z <
UJ
O
a
UJ
i
a o o
in
3
E cc
-I K
3 8
oc
5 S
o
> DC
I !
W
m in
(/>
UJ
§
CC
o
|
o |
OC |
|
O |
|
|
o > |
a. |
|
Z H a. Z |
3 |
|
U. 3 it |
HI > |
|
UJ |
|
|
EC |
cc |
z < o a o
a
i
i a
I
T O
i §
X Z
1SVOD 3S
NOI1V30T
1SWOD MS
0IHdVU9O39
T3 "
-t-> II (US.
O i- cn a)
C l£> <U C +»
> v- in
aj "■ — &- ia
t- +J •.- <4- E
n5 <D LO r—
i— no
.c i— ii o
10 3 lf> .C
i- E O i— U U_ i— (A
-o "
qi « jc a)
• Q. 3 (/) f—
t/1 -I— O 'I— *»-*
OJ i- c 4- c
t- 4-> c oj a> •
O in •!- Q. > in
<u e •!- 3 ai
o. n □.■>-> g:
in "O IO
in otj ii c
OJ OJ OJ
> «.C OCTi O t- -C w C i— 1-
+j in Q.-1- <+-
n] •■- O S. *•'- +J U- OJ <f- >>■!->
c >>.£: <o c
aj -o in H s- <u
1/1 (d CT»i—
ai t— ii <- c u
i- i— t- in
a ii en +J
Oj * m i-
S- «*• « -c O
in m c <4-
cji « aj i- &-
CJCTIt- OJ DO •i— in •■- 2 jz 3 t- m o -i-» X O <t- t- o 3 v- .c t- OJ o "O in i— > TJ in c w— •— OJ OJ
in •■- v- •— II O-
v^ i/i 3 a.
i- ra ao «C
+j .c s- s- i—
•r- in a) o aj
cz $- +-> in • oj 3 a a -c i/>
e e s ii w E a,„r
O II T3 CO <t- •
<J •■- i— •■- 4_
CO 4-> r— O
.C "i— •"->
in » ii £ f id
i- sz w .* e
i»- in CO •!-
•■- "- -O +J
•a t- * OJ OJ c
tl OT) 3«J 01
-t-> +J IO CT> ■*-> C7>
Id i- OJ C O i-
•r- 3 r O Q. OJ
o O" in 4-> in in O in O- T3
n o aiJ^i- ai
Ul E HI D Or- al -C OJ o»i- i ii in jz c oj una) > CM oj -* >
O f— U l*» 3
OlUl Cr- C 3 OJ .Q -II
«J ■— > ■*
E 3 3 II i. i—
> •!-> (O CM
<f- •»- CM -C
O i- II •— in *
ai -•-> ii r — . • cz ■!->
C -C O ITJ
aj r— " in E ••->
•I- IO -r- 0} E
■o S- 4- I— o
io • i- en +J c ai io -r- ii
CJ3 ^- -i-j Q. OJ
IO O IO r—
UEIi-r
• m c
i— c i— - OJ
r— O •>- ■— J= >
+j i>- in 3 Oj 5 « t- "1-j
S- C O .£= «f-
3 2 r- O OJ II
cn io r— s- ■—
•■- j- a) oj i- o u. -a >> Q.**- CM
51
herons, ibis and the wood stork (Heald et al. 1974).
7.2 RIVERINE FORESTS
Tidal streams and rivers, fringed largely by red mangroves, connect the freshwater marshes of south Florida with the shallow estuarine bays and lagoons (Figure 12). Few of these streams have been studied thoroughly. The exception is the North River which flows into White- water Bay and was studied by Tabb (1966) and Odum (1970). Springer and Woodburn (1960) collected fishes in a bayou or tidal pass connecting Boca Ciega Bay and Old Tampa Bay. Carter et al. (1973) reported on the fishes of two tidal streams entering Fahkahatchee and Fahka Union Bays. Nugent (1970) sampled fishes in two streams on the western shore of Biscayne Bay. Characteristics of these areas and sampling gear used by the inves- tigators are summarized in Appendix A.
These tidal streams and associated riverine mangrove forests exhibit extreme seasonal variability in both physical characteristics and fish community compo- sition. Salinity variations are directly related to changes in the make-up of the fish assemblage. During the wet season (June - November), salinities fall throughout the water courses and, at some locations in certain heavy runoff years, become fresh all of the way to the mouth (Odum 1970). Opportunistic freshwater species, which are normally restricted to the sawgrass and black needle rush marshes of the headwaters, invade the mangrove zone. These include the Florida gar, Lepi sosteus pi a ty r h i nc us; several centrarchid sunfishes of the genus Lepomis and the largemouth bass, Micropterus sal moi des ; the freshwater catfishes, Ictalurus natalis and Noturus gyrinus; and the killifishes normally considered freshwater inhabitants such as Lucani a goodei and Ri vulus marmoratus.
During the dry season (December to early May) salinities rise as a result of decreased freshwater runoff and continuing evaporation. Marine species invade the
tidal streams primarily on feeding forays. Examples include the jewfish, Epinephelus ita jara, the stingrays (Dasyatidae), the needlefishes (Belonidae), the jacks (Carangidae), and the barracuda, Sphyraena barracuda. Other seasonal movements of fishes appear to be temperature related. Tabb and Manning (1961) documented move- ments of a number of species from shallow inshore waters to deeper water during times of low temperature stress. The lined sole, the hogchoker, the bighead searobin, and the striped mullet, for example, are much less frequently caught in winter in shallow inshore waters.
A third type of seasonality of fish populations in the tidal rivers is related to life cycles. Many of the fish which utilize the tidal stream habitat do so only as juveniles. Thus, there are peaks of abundance of these species following offshore spawning when larval or juvenile forms are recruited to the mangrove stream habitat. In general, recruitment occurs in the late spring or early summer fol- lowing late winter and spring spawning offshore or in tidal passes (Reid 1954). Numerous species are involved in this life cycle phenomenon including striped mullet, grey snapper, sheepshead, spotted sea trout, red drum, and silver perch.
The only estimate of fish standing crop from tidal stream habitats is that of Carter et al. (1973). They recorded 27 species weighing 65,891 g (wet wt.) from an area of 734 m or about 90 g/m . This is probably an overestimate since an un- known portion of the fish community had moved from the flooded lowlands to the stream on the ebb tide; sampling occurred at low tide in October. Nonetheless, this is an indication of the high fish standing crop which this mangrove-associated habi- tat can support. The number of species reported from individual tidal streams annually ranges from 47 to 60 and the total from all tidal streams in southwest Florida is 111 species (Appendix B).
The food webs in these riverine man- grove ecosystems appear to be predomi- nantly mangrove detritus-based, although the Biscayne Bay stream studied by Nugent
52
Figure 12. Aerial photograph of the mangrove belt of southwest Florida near Whitewater Bay. Note the complex system of pools and small creeks which connect with the tidal river system.
53
(1970) may be an exception. The basic link between the mangrove leaf and higher order consumers is provided by micro- organisms (fungi, bacteria, Protozoa) which colonize the decaying leaf and con- vert them into a relatively rich protein source (Odum 1970; Odum and Heald 1975a). These decaying leaf fragments with asso- ciated microorganisms are fed upon by a group of omnivorous detritivores including amphipods, mysids, cumaceans, ostracods, chironomid larvae, harpacticoid and calanoid copepods, snapping shrimp, caridean and penaeid shrimp, a variety of crabs, filter-feeding bivalves, and a few species of fishes (Odum 1970; Odum and Heald 1972; Odum and Heald 1975b). These detritivores, in turn, are consumed by a number of small carnivorous fishes, which in turn, are consumed by larger piscivorous fishes. The concept of man- grove trophic structure is also discussed in section 3.6. See Appendix B for species specific dietary information.
The tidal creeks studied by Nugent (1970) on the western shore of Biscayne Bay differ from the previously discussed streams in the Everglades estuary. The mouths of the Biscayne Bay creeks have dense growths of sea grasses which con- tribute sea grass detritus. The salini- ties are considerably greater and the streams are located only a few kilometers from coral reefs, which are largely absent on Florida's west coast, at least close to shore. As a result, 23 species listed in Appendix B were captured by Nugent (1970) and are not recorded from riverine man- grove habitat on the west coast of Florida. Examples include several of the grunts (Pomadasyidae), the gray trigger- fish, Balistes capriscus, the barbfish, Scorpaena brasiliensis, the scrawled box- fish, Lactophrys quadricorni s, and the snappers, Lutjanus apodus and L. synagris.
Riverine mangrove communities and associated tidal streams and rivers are typified by the following families of fishes: killifishes (Cypri nodontidae), livebearers (Poeci 1 i idae), silversides (Atherinidae), mojarras (Gerreidae), tar- pon (Elopidae), snook (Centropomidae), snappers (Lut jani dae) , sea catfishes
(Ariidae), gobies (Gobi idae), porgys (Sparidae), mullets (Mugilidae), drums (Sci aenidae), and anchovies (Engraulidae). The mangrove-lined streams and associated pools are important nursery areas for several marine and estuarine species of gamefish. The tarpon, Megalops atlantica, snook, Centropomus undecimalis, and lady- fish, Elops saurus, utilize these areas from the time they reach the estuary as post-larvae, having been spawned offshore. Gray snapper, Lutjanus g r i s e u s , sheepshead, Archosargus probatocephalus, spotted seatrout, Cynoscion nebulosus, and red drum, Sciaenops ocellata, are re- cruited to grass beds of shallow bays and lagoons as post-larvae and enter the mangrove-lined streams for the next sever- al years (Heald and Odum 1970). Of these species, only the spotted seatrout prob- ably spawns in the estuary (Tabb 1966). Other species of commercial or game impor- tance which use the riverine fringing habitat include crevalle jack, gafftopsail catfish, jewfish, striped mojarra, barra- cuda, Atlantic thread herring, and yellow- fin menhaden (Odum 1970).
7.3 FRINGING FORESTS ALONG ESTUARINE BAYS AND LAGOONS
Mangrove-fringed estuarine bays and lagoons are exemplified by the Ten Thousand Islands area and Whitewater Bay. Quantitative fish data are available from Fahkahatchee Bay (Carter et al. 1973; Yokel 1975b; Seaman et al. 1973), Fahka Union Bay (Carter et al. 1973), Rookery Bay (Yokel 1975a), the Marco Island Estuary (Weinstein et al. 1977; Yokel 1975a), and Whitewater Bay (Clark 1970). Individual site characteristics are summarized in Appendix A. All except Fahka Union Bay contain significant amounts of sea grasses. Macroalgae domi- nate the benthic producers of Fahka Union Bay. Studies by Reid (1954) and Kilby (1955) near Cedar Key, Florida, were not included in our summary because mangroves are sparse in this area and no mention of mangrove collecting sites were made by these authors. Studies of Caloosahatchee Bay (Gunter and Hall 1965) and of Charlotte Harbor (Wang and Raney 1971)
54
were omitted because the areas studied have been highly modified and because data from many habitats were pooled in the final presentation.
All of the bays reviewed in our sum- maries are fringed by dense growths of red mangroves and all contain small mangrove islets. Carter et al. (1973), in their studies of Fahkahatchee and Fahka Union bays, estimated that 57% to 80% of the total energy budget of these two bays is supported by exports of particulate and dissolved organic matter from the man- groves within the bays and inflowing tidal streams. Lugo et al. (1980) estimated that the mangroves surrounding Rookery Bay provide 32% of the energy base of the heterotrophic community found in the bay.
Salinities in these bays tend to be higher than in the tidal streams and rivers and the fish assemblages reflect both this feature and the added habitat dimension of sea grass and macro algae beds. Truly freshwater species are rare in these communities and a proportionally greater percentage of marine visitors is present. The dominant fish families of the benthic habitat include drums (Sci aenidae), porgys (Sparidae), grunts (Pomadasyidae), mojarras (Gerreidae), snappers (Lutjanidae), and mullet (Mugili- dae). Other familes with sizeable contri- butions to the benthic fauna include pipe- fishes (Syngnathidae), flounder (Bothi- dae), sole (Soleidae), searobins (Trigli- dae), and toadfishes (Batrachoididae).
Numerically abundant fishes of the mid and upper waters include anchovies (Engraul idae), herrings (Clupeidae) and needlefishes (Belonidae). At all loca- tions studied, the benthic fauna was domi- nated by the pinfish, Lagodon rhomboides, the silver perch, Bairdiella chrysura, the pigfish, Orthopristis chrysoptera. and the mojarras, Euci nostomus gula and E. argenteus. The most common midwater and surface species include the two anchovies, Anchoa mitchilli and A_. hepsetus, and two clupeids, Brevoortia smithi and Harengula pensacol ae. The total number of species recorded in the individual studies ranged from 47 to 89; a total of 117 species was
collected in these mangrove-fringed bays and lagoons (Appendix B).
In none of these studies were the fishes specifically utilizing the fringing mangrove habitat enumerated separately from those collected in the bay as a whole. The collections were most often at open water stations easily sampled by otter trawl. Carter et al. (1973) had two shore seine stations adjacent to mangroves but the data were pooled for publication. Of the four stations in Rookery Bay sam- pled by Yokel (1975a), one was immediately adjacent to the fringing mangrove shore- line and had moderate amounts of sea grasses.
The typical pattern which emerges from many estuarine studies is that rela- tively few fish species numerically domi- nate the catch. This is certainly true in mangrove-fringed estuaries. In Rookery Bay (Yokel 1975a) six species comprised 88% of the trawl -catchable fishes, in Fahkahatchee Bay seven species comprised 97% of the catch from three capture techniques (Carter et al. 1973), and in the Marco Island estuary 25 species com- prised 97% of the trawl -catchable fishes (Weinstein et al. 1977).
Like tidal river and stream communi- ties, these shallow bays serve as nur- series for numerous species of estuarine- dependent fishes that are spawned off- shore. Based on the distribution and abundance of juvenile fishes of all spe- cies in six habitats, Carter et al. (1973) ranked the mangrove-fringed bays as the most important nursery grounds; the tidal streams were a close second. Shallow bays and tidal streams provide safe nurseries due to seasonally abundant food resources and the low frequency of large predators (Carter et al. 1973; Thayer et al. 1978). The relative lack of large predaceous fishes is probably due to their general inability to osmoregulate in waters of low and/or fluctuating salinity.
As in tidal streams, the peak abun- dance of juvenile and larval fishes in the bays is in spring and early summer (Reid 1954). In general, the highest standing
55
crops and the greatest species richness of fishes occur in the late summer and early fall (Clark 1970). Fish densities decline in the autumn and winter as many fishes move to deeper waters.
7.4 FRINGING FORESTS ALONG OCEANIC BAYS AND LAGOONS
Mangrove-fringed "oceanic" bays and lagoons are exemplified by Porpoise Lake in eastern Florida Bay (Hudson et al. 1970), western Florida Bay (Schmidt 1979), southern Biscayne Bay (Bader and Roessler 1971), and Old Rhodes Key Lagoon in eastern Biscayne Bay (Holm 1977). Charac- teristics of these sites are summarized in Appendix A. Compared to the mangrove- fringed bays discussed in the previous section, these environments generally ex- hibit clearer water, sandier substrates, and higher and less variable salinities. Closer proximity to the Florida reef tract, the Atlantic Ocean, and the Gulf of Mexico results in a larger potential pool of fish species. These four locations have produced reports of 156 fish species (Appendix B).
Mangrove fringes make up a relatively small proportion of these environments; accordingly, their contribution to the bay food webs is probably not very large. Bader and Roessler (1972) estimated that the fringing mangrove community contrib- utes approximately 1% of the total energy budget of southern Biscayne Bay; they considered only mainland mangroves and did not include the small area of mangrove islands. The main ecological role of the fringing mangroves in this type of en- vironment is probably twofold. First, they increase the habitat diversity within an otherwise relatively homogeneous bay system. Second, they provide a relatively protected habitat for juvenile fishes (and certain invertebrates) that later move to more open water or coral reef communities. The second role is analogous to one of the ecological roles of sea grass communities (see Zieman, in prep.) although the fish species involved may be different.
Based primarily on habitat designa- tions of Voss et al. (1969), the fishes of Biscayne Bay can be characterized as to preferred habitat. Of the three main habitat types, (1) rock/coral/seawall, (2) grassbed/tidal flat, and (3) mangrove, the grassbed/tidal flat ranked first in fish species occurrences. One hundred and twenty-two of 156 species (79%) are known to occur in this environment. Rock/coral /seawal 1 habitats were fre- quented by 49 species (32%) and mangroves are known to be utilized by 54 species (35%) of the total fish species recorded from this bay.
7.5 OVERWASH MANGROVE ISLANDS
In terms of fish-related research, these communities are the least studied of all mangrove community types in south Florida. They are typified by the low- lying mangrove-covered islands that occur in the Florida Keys and Florida Bay and may be overwashed periodically by the tides. Examples include Shell Key, Cotton Key, and the Cowpens. Islands of this type extend southwest from the Florida mainland through the Marquesas. The Dry Tortugas lack well -developed mangrove com- munities although stunted trees are found (Davis 1942).
These islands are the most oceanic of any of the mangrove communities discussed. They are characterized by relatively clear water (Gore 1977) and are largely free of the freshwater inflow and salinity varia- tions which characterize other Florida mangrove communities to varying degrees. Numerous statements exist in the litera- ture acknowledging the frequent proximity of mangrove islands to coral reefs and sea grass beds (McCoy and Heck 1976; Thayer et al. 1978). Olsen et al. (1973) workinq in the U.S. Virgin Islands, found 74% to 93% overlap in the fish species composition of fringing coral reefs and shallow mangrove- fringed oceanic bays. Voss et al. (1969) listed fish species that were collected from all three types of communities: fringing mangroves, coral reefs and sea
56
grass beds in Biscayne Bay, but there appears to have been no systematic survey of the fish assemblage characteristic of the mangrove-covered or mangrove-fringed Florida Keys. No one has quantified the faunal connections which we hypothesize exist between the mangroves and sea grasses and between the mangroves and coral reefs.
In the absence of published data from the mangrove key communities, only tenta- tive statements can be made. In general, we expect that while mangrove islands serve as a nursery area for juvenile fishes, this function is limited largely to coral reef and marine inshore fishes and not the estuarine-dependent species that we have discussed previously. The latter (juvenile snook, red drum, spotted seatrout) appear to require relatively low salinities not found in association with most of the overwash islands. Casual observation around the edges of these islands suggests that characteristic fishes include the sea bass family (Ser- ranidae), t ri ggerf i shes (Bal istidae), snappers (Lut jam' dae), grunts (Poma- dasyidae), porgies (Span'dae) parrotfishes (Scaridae), wrasses (Labridae), bonefishes (Albulidae), jacks (Carangidae), damsel- fishes (Pomacentridae), and surgeonf ishes (Acanthuridae); many of these fishes occur on or are associated with coral reefs. We also suspect that considerable overlap occurs in the fish assemblage of these mangrove islands and sea grass communi- ties; examples include puffers (Tetrao- dontidae), pipefishes (Syngnathidae) , go- bies (Gobiidae) and scorpionf ishes (Scor- paenidae). Stark and Schroeder (1971) suggested that juvenile gray snapper, which use the fringing mangroves of the keys as shelter during the day, forage in adjacent sea grass beds at night. In the absence of salinity barriers, predatory fishes probably enter the fringes of these
mangrove islands on the rising tide. Included in this group are sharks, tarpon, jacks, snook, bonefish and barracuda.
7.6 GRADIENT OF MANGROVE COMMUNITY INTERACTIONS
Mangrove communities occur under a wide range of conditions from virtually freshwater at the headwaters of tidal streams to nearly oceanic conditions in the Florida Keys. Attempting to present a single list of fish characteristic of mangrove environments (Appendix B) can be misleading. For this reason we presented the concept of a continuum or complex gradient in Figure 11 and have followed that scheme throughout section 7. The gradient stretches from seasonally fresh to oceanic conditions, from highly varia- ble salinities to nearly constant salini- ty, from muddy and limestone substrates to sandy substrates, from dark-stained and sometimes turbid waters to clear waters, and from food webs that are predominantly mangrove detritus-based to food webs based primarily on other energy sources. Clear- ly, there are other gradients as one moves from north to south in the State of Florida. At the northern end of the State, temperatures are more variable and seasonally lower than in the south. Sedi- ments change from predominantly silicious in central and north Florida to predomi- nantly carbonate in extreme south Florida. Nevertheless, the complex gradient shown in Figure 11, while greatly simplified for graphic purposes, suggests that charac- teristic fish assemblages replace one another along a gradient of changing physical and biogeographic conditions. Such a concept is useful in understanding the factors controlling the composition of fish assemblages associated with mangroves of the four major community types in south Florida.
57
CHAPTER 8. COMMUNITY COMPONENTS - AMPHIBIANS AND REPTILES
Food habits and status of 24 species of turtles, snakes, lizards, and frogs of the Florida mangrove region are given in Appendix C. Any of three criteria had to be met before a species was included in this table: (1) a direct reference in the literature to mangrove use by the species, (2) reference to a species as being present at a particular geographical location within the mangrove zone of Florida, and (3) North American species recorded from mangroves in the West Indies or South America, but not from Florida. This last criterion assumes that a species which can utilize mangroves outside of Florida will be able to use them in Florida. Ten turtles are listed of which four (striped mud turtle, chicken turtle, Florida red-bellied turtle, and softshell turtle) are typical of freshwater. Two (mud turtle and the ornate diamondback terrapin) are found in brackish water and the remainder (hawksbill, green, logger- head, and Atlantic ridley) are found in marine waters.
Freshwater species usually occur in the headwater regions of mangrove-lined river systems. All four freshwater species are found in habitats other than mangrove swamps including streams, ponds, and freshwater marshes. The brackish water species are found in salt marshes in addition to mangrove swamps. Mangroves, however, are the principal habitat for the ornate diamondback terrapin (Ernst and Barbour 1972). Cam and Goi n (1955) listed two subspecies of the diamondback: Malaclemys terrapin macrospilota and M. t. rhizophorarum. Malaclemys terrapin macro- spilota inhabits the southwest and south- ern coasts, and M. ;t. rhizophorarum is found in the Florida Keys. The two sub- species intergrade in the region of north- ern Florida Bay.
All four of the marine turtles are associated with mangrove vegetation at some stage of their lives. Loggerhead and green turtles are apparently much less dependent on mangroves than the remaining two, although we strongly suspect that recently hatched loggerheads may use man- grove estuaries as nursery areas. Green turtles are generally believed to feed on
a variety of submerged aquatic plants and sea grasses; recent evidence has shown that they also feed on mangrove roots and leaves (Ernst and Barbour 1972). The Atlantic ridley's preferred habitat is "shallow coastal waters, especially the mangrove-bordered bays of the southern half of the peninsula of Florida" (Carr and Goin 1955). Hawksbill turtles feed on a variety of plant materials including mangrove (especially red mangrove), fruits, leaves, wood, and bark (Ernst and Barbour 1972).
Three species in the genus Anol i s have been reported from Florida mangroves: the green anole, the cuban brown anole, and the Bahaman bank anole. All are arboreal lizards that feed on insects. The green anole is widespread throughout the Southeastern United States and is not at all dependent on mangrove swamps. The other two species have much more restricted distributions in the United States and are found only in south Florida. They also are not restricted to mangrove ecosystems. Of the six species of snakes listed, the mangrove water snake (Figure 13) is most dependent upon mangrove habitats.
Two important species of reptiles found in mangrove swamps are the American alligator and the American crocodile. The alligator is widespread throughout the Southeastern United States and is only incidentally found in low salinity sec- tions of Florida mangrove areas (Kushlan 1980). The American crocodile is rare; historically its distribution was centered in the mangrove-dominated areas of the upper and lower Florida Keys (particularly Key Largo) and the mangrove-lined shore- lines and mud flats along the northern edge of Florida and Whitewater Bays (Kushlan 1980). Mangroves appear to be critical habitat for this species. Its range has shrunk considerably in south Florida since the 1930's, even though Florida Bay was added to Everglades National Park in 1950 (Moore 1953; Ogden 1978). Much of the decrease in range is due to increased human activity in the Florida Keys. The remaining population centers of the American crocodile are in
Figure 13. The mangrove water snake, Nerodja fasciata compressicauda, curled on a red mangrove prop root. Photograph by David Scott.
59
northern Florida Bay and adjacent coastal swamps and the northern end of Key Largo (Ogden 1978; Kushlan 1980). The species uses a variety of habitats for nesting in the Florida Bay region including open hardwood thickets along creek banks, hardwood-shrub thickets at the heads of sand-shell beaches, and thickets of black mangroves behind marl banks (Ogden 1978). On Key Largo the crocodile locates its nests on creek and canal banks in red and black mangrove swamps (Ogden 1978). Man- grove areas thus appear to be important in the breeding biology of this endangered species.
Interestingly, only three species of
amphibians, to our knowledge, have been recorded in Florida mangrove swamps (Ap- pendix C). This is due to two factors: (1) lack of detailed surveys in low sa- linity swamps and (2) the inability of most amphibians to osmoregulate in salt water. No doubt, several additional species occur in the freshwater -dominated hammock and basin mangrove communities inland from the coast. Possible addi- tional species include: the eastern narrow-mouthed toad, Gastrophryne caro- 1 in en si s, the eastern spadefoot toad, Scaphiopus holbrooki, the cricket frog, Acris gryllus, the green tree frog, Hyl a cinerea, and the southern leopard frog, Rana utricularia.
60
CHAPTER 9. COMMUNITY COMPONENTS - BIRDS
9.1 ECOLOGICAL RELATIONSHIPS
Because mangroves present a more diverse structural habitat than most coastal ecosystems, they should harbor a greater variety of birdlife than areas such as salt marshes, mud flats, and beaches (MacArthur and MacArthur 1961). The shallow water and exposed sediments below mangroves are available for probing shorebirds. Longer-legged wading birds utilize these shallow areas as well as deeper waters along mangrove-lined pools and waterways. Surface-feeding and diving birds would be expected in similar areas as the wading birds. The major difference between mangrove swamps and other coastal ecosystems is the availability of the trunks, limbs, and foliage comprising the tree canopy. This enables a variety of passerine and non-passerine birds, which are not found commonly in other wetland areas, to use mangrove swamps. It also allows extensive breeding activity by a number of tree-nesting birds.
The composition of the avifauna com- munity in mangrove ecosystems is, in fact, highly diverse. Cawkell (1964) recorded 45 species from the mangroves of Gambia (Africa). Haverschmidt (1965) reported 87 species of birds which utilized mangroves in Surinam (S. America). Ffrench (1966) listed 94 species from the Caroni mangrove swamp in Trinidad while Bacon (1970) found 137 in the same swamp. In Malaya, Nisbet (1968) reported 121 species in mangrove swamps and Field (1968) observed 76 from the mangroves of Sierra Leone (Africa).
Use of mangrove ecosystems by birds in Florida has not been recorded in de- tail. Ninety -two species have been ob- served in the mangrove habitat of Sanibel Island, Florida (L. Narcisse, J.N. "Ding" Darling Natl. Wildlife Refuge, Sanibel Is., Fla.; personal communication 1981). Robertson (1955) and Robertson and Kushlan (1974) reported on the entire breeding bird fauna of peninsular south Florida, including mangrove regions. Based on limited surveys, these authors reported only 17 species as utilizing mangroves for breeding purposes. Because their studies did not consider migrants or non-breeding
residents, a significant fraction of the avifauna community was omitted.
Based on information gleaned from the literature, we have compiled a list of 181 species of birds that use Florida mangrove areas for feeding, nesting, roosting, or other activities (Appendix D). Criteria for listing these species is the same as that used for listing reptiles and amphi- bians (see Chapter 8 of this volume).
Often references were found stating that a given species in Florida occurred in "wet coastal hammocks", "coastal wet forests" or the like, without a specific reference to mangroves. These species were not included in Appendix D. Thus, this list is a conservative estimate of the avifauna associated with Florida man- grove swamps. Sources for each listing are provided even though many are redun- dant. Food habit data are based on Howell (1932) and Martin et al. (1951). Esti- mates of abundance were derived from bird lists published by the U.S. Fish and Wildlife Service for the J.N. "Ding" Darling National Wildlife Refuge at Sanibel Island, Florida, and by the Ever- glades Natural History Association for Everglades National Park. Frequently, species were recorded from mangrove swamps at one location, but not the other.
We have divided the mangrove avifauna into six groups based on similarities in methods of procuring food. These groups (guilds) are the wading birds, probing shorebirds, floating and diving water- birds, aerially-searching birds, birds of prey, and arboreal birds. This last group is something of a catch-all group, but is composed mainly of birds that feed and/or nest in the mangrove canopy.
9.2 WADING BIRDS
Herons, egrets, ibises, bitterns, and spoonbills are the most conspicuous group of birds found in mangroves (Figure 14) and are by far the most studied and best understood. Eighteen species (and one important subspecies) are reported from south Florida mangroves.
61
Figure 14. A variety of wading birds feeding in a mangrove-lined pool near Flamingo, Florida. Photograph by David Scott.
62
Mangrove swamps provide two functions for wading birds. First, they function as feeding grounds. Two-thirds of these species feed almost exclusively on fishes. Although much of their diet is provided by freshwater and non-mangrove marine areas, all of them feed frequently in mangrove swamps. White ibis feed predominantly on crabs of the genus Uca when feeding in mangroves (Kushlan and Kushlan 1975; Kushlan 1979). Mollusks and invertebrates of the sediments are principal foods of the roseate spoonbill although some fish are eaten (Allen 1942). Yellow-crowned night herons and American bitterns eat crabs, crayfish, frogs, and mice in addi- tion to fishes. Snails of the genus Pomacea are fed upon almost exclusively by the limpkin. The sandhill crane is an anomaly in this group since a majority of its food is vegetable matter, especially roots and rhizomes of Cyperus and Sagittaria. Its use of mangroves is probably minimal, occurring where inland coastal marshes adjoin mangroves (Kushlan, unpubl . data). The remaining 12 species are essentially piscivorous although they differ somewhat in the species and sizes of fishes that they consume.
Mangrove swamps also serve as breeding habitat for wading birds. With the exception of the limpkin, sandhill crane, and the two bitterns, all wading bird species in Appendix D build their nests in all three species of mangrove trees (Maxwell and Kale 1977; Girard and Taylor 1979). The species often aggregate in large breeding colonies with several thousand nesting pairs (Kushlan and White 1977a). The Louisiana heron, snowy egret, and cattle egret are the most numerous breeders in south Florida mangroves (based on data in Kushlan and White 1977a).
In wet years over 90% of the south Florida population of white ibis breed in the interior, freshwater wetlands of the Everglades; during these times the man- groves are apparently unimportant, sup- porting less than 10% of the population (Kushlan 1976, 1977a, b). During drought years, however, production is sustained solely by breeding colonies located in mangroves near the coast (Kushlan 1977a,
b). Mangroves are critically important for the survival of the white ibis popula- tion even though they appear to be utilized to a lesser extent than fresh- water habitats. This pattern of larger but less stable breeding colonies using inland marshes and smaller but more stable colonies using mangroves is also charac- teristic of heron populations (Kushlan and Frohring, in prep.).
Table 5 gives the number of active nests observed in mangrove regions during the 1974-75 nesting season and the percen- tage this represents of the entire south Florida breeding population for the nine most abundant species of waders and three associated species. The dependence of roseate spoonbills, great blue herons, Louisiana herons, brown pelicans, and double-crested cormorants on mangrove regions is evident. Nesting by the red- dish egret was not quantified during this study although Kushlan and White (1977a) indicated that the only nests of this species which they saw were, in fact, in mangroves. Further observations indicate that this species nests in mangroves ex- clusively (Kushlan, pers. comm.). Similar- ly, the great white heron is highly depen- dent upon mangroves for nesting; they use the tiny mangrove islets which abound along the Florida Keys and in Florida Bay (Howell 1932).
During many years the Everglades population of wood storks is known to nest almost solely in mangroves (Ogden et al. 1976); this population comprises approxi- mately one-third of the total south Florida population. Successful breeding of all these mangrove nesters is un- doubtedly correlated with the abundant supply of fishes associated with man- groves. Meeting the energetic demands of growing young is somewhat easier in habi- tats with abundant prey. This is especially important for the wood stork which requires that its prey be concen- trated into small pools by falling water levels during the dry season before it can nest successfully (Kahl 1964; Kushlan et al. 1975; Odgen et al. 1978). Breeding activity by wading birds in mangroves along the southwest and southern Florida
63
Table 5. Nesting statistics of wading birds and associated species in south Florida, 1974-1975 (based on data in Kushlan and White 1977a).
Species
|
% of |
total active |
|
|
Active nests in |
nests in south |
|
|
mangroves |
Florida |
|
|
1914 |
7 |
|
|
500 |
100 |
|
|
1335 |
31 |
|
|
458 |
92 |
|
|
1812 |
39 |
|
|
2377 |
46 |
|
|
71 |
15 |
|
|
3410 |
70 |
|
|
2180 |
13 |
|
|
741 |
100 |
White ibis
Roseate spoonbill
Wood stork
Great blue heron
Great egret
Snowy egret
Little blue heron
Louisiana heron
Cattle egret
Brown pelican
Double-crested cormorant 1744 83
64
coasts takes place throughout the year (Table 6); at least one species of wader breeds during every month. Colonies on the mangrove islands in Florida Bay were noted to be active nesting sites during all months of the year except September and October (Kushlan and White 1977a).
The seasonal movements of wood storks and white ibises between the various south Florida ecosystems were described by Ogden et al. (1978) and Kushlan (1979). Mangrove ecosystems appear to be most heavily used for feeding in summer (white ibis) and early winter (white ibis and wood stork). The remaining species of wading birds appear to use mangrove areas most heavily in the winter months, reflec- ting the influx of migrants from farther north.
Wading birds play an important role in nutrient cycling in the coastal man- grove zone. Mclvor (pers. observ.) has noted increased turbidity, greater algal biomass, and decreased fish abundance around red mangrove islets with nesting frigate birds and cormorants. Onuf et al. (1977) reported results from a small (100 bird) rookery on a mangrove islet on the east coast of Florida. Additions of ammonium-nitrogen from the bird's droppings exceeded 1 g/m /day. Water beneath the mangroves contained five times more ammonium and phosphate than water beneath mangroves without rookeries. Although the wading birds were shown to be a vector for concentrating nutrients, it must be noted that this is a localized phenomenon restricted to the areas around rookeries in the mangrove zone. The effect would be larger around larger rookeries. Onuf et al. (1977) also reported that mangroves in the area of the rookery had increased levels of primary production, higher stem and foliar nitro- gen levels, and higher herbivore grazing impact than mangroves without rookeries. Lewis and Lewis (1978) stated that man- groves in large rookeries may eventually be killed due to stripping of leaves and branches for nesting material and by poisoning due to large volumes of urea and ammonia that are deposited in bird guano. This latter effect would be more
pronounced in rookeries within mangrove regions subject to infrequent tidal flush- ing.
9.3 PROBING SHOREBIRDS
Birds in this group are commonly found associated with intertidal and shal- low water habitats. Wolff (1969) and Schneider (1978) have shown that plovers and sandpipers are opportunistic feeders, taking the most abundant, proper-sized invertebrates present in whatever habitat the birds happen to occupy.
Of the 25 species included in this guild (Appendix D), two are year-round residents (clapper rail and willet), two breed in mangrove areas (clapper rail and black-necked stilt), and the remainder are transients or winter residents. Baker and Baker (1973) indicated that winter was the most crucial time for shorebirds, in terms of survival. Coi ncidental ly, winter is the time when most shorebirds use mangrove areas. The invertebrate fauna (mollusks, crustaceans, and aquatic insects) which occur on the sediments under intertidal mangroves forms the principal diet of these species. Willets and greater yellowlegs eat a large amount of fishes, especially Fundulus, in addition to inver- tebrates. Many of the species listed in this guild obtain a significant portion of their energy requirements from other habi- tats, particularly sandy beaches, marshes, and freshwater prairies. Of the species in this guild, the clapper rail is prob- ably most dependent on mangroves for survival in south Florida (Robertson 1955), although in other geographical locations they frequent salt and brackish marshes.
9.4 FLOATING AND DIVING WATER BIRDS
Twenty-nine species of ducks, grebes, loons, cormorants, and gallinules were identified as populating mangrove areas in south Florida (Appendix D). Eight species are year-round residents while the remainder are present only during migra- tion or as winter visitors.
65
Table 6. Timing of nesting by wading birds and associated species in south Florida. Adapted from data in Kushlan and White (1977a), Kushlan and McEwan (in press).
White ibis
Wood stork
Roseate spoonbill
Great blue/white heron
Great egret
Little blue heron
Cattle egret
Double-crested cormorant
Brown pelican
Months
Species SONDJFMAMJJA
66
From the standpoint of feeding, mem- bers of this guild are highly hetero- geneous. Piscivorous species include the cormorant, anhinga, pelicans, and mergan- sers. Herbivorous species include the pintail, mallard, wigeon, mottled duck, and teals. A third group feeds primarily on benthic mollusks and invertebrates. Scaup, canvasback, redhead, and gallinules belong to this group. The ducks in this last group also consume a significant fraction of plant material.
Species of this guild are permanent residents and usually breed in mangrove swamps. As shown in Table 5, the brown pelican and double-crested cormorant are highly dependent upon mangroves for nesting in south Florida even though both will build nests in any available tree in other geographical regions. It seems that when mangroves are available, they are the preferred nesting site. The anhinga breeds in mangrove regions but is more commonly found inland near freshwater (J. A. Kushlan, So. Fla. Res. Ctr., Everglades Natl. Park, Homestead, Fla.; personal communication 1981). For the other species listed in this guild, mangrove swamps provide a common but not a required habi- tat; all of these species utilize a variety of aquatic environments.
Kushlan et al. (in prep.) provide recent data on the abundance and distribu- tion of 22 species of waterfowl and the American coot in south Florida estuaries. The American coot is by far the most abun- dant species, accounting for just over 50% of the total population. Six species of ducks were responsible for more than 99% of the individuals seen: blue-winged teal (41%), lesser scaup (24%), pintail (18%), American wigeon (9%), ring-necked duck (5%), and shoveler (3%). The major habi- tats included in these authors' surveys were coastal prairie and marshes, mangrove forests, and mangrove-lined bays and waterways of the Everglades National Park.
From these data it appears that waterfowl and coots are most abundant in regions where mangrove, wet coastal prairies, marshes, and open water are interspersed. Overall, the Everglades
estuaries support from 5% to 10% of the total wi nteri ng waterfowl population in Florida (Goodwin 1979; Kushlan et al. in prep.). As Kushlan et al. point out, however, the Everglades are not managed for single species or groups of species as are areas of Florida supporting larger waterfowl populations. Although the importance of south Florida's mangrove estuaries to continental waterfowl popula- tions may be small, the effect of 70,000 ducks and coots on these estuaries probably is not (Kushlan et al. in prep.).
Kushlan (personal communication) thinks that the estuaries of the Ever- glades have an important survival value for some segments of the American white pelican population. In winter, approxi- mately 25% of the white pelicans are found in Florida Bay and 75% in the Cape Sable region. They feed primarily in freshwater regions of coastal marshes and prairies and use mangroves where they adjoin this type of habitat.
9.5 AERIALLY-SEARCHING BIRDS
Gulls, terns, the kingfisher, the black skimmer, and the fish crow comprise this guild of omnivorous and piscivorous species (Appendix D). These birds hunt in ponds, creeks, and waterways adjacent to mangrove stands. Many fishes and inverte- brates upon which they feed come from mangrove-based food webs. Only six of the 14 species are year-round residents of south Florida. The least tern is an abun- dant summer resident and the remainder are winter residents or transients.
Only the fish crow actually nests in mangroves. Gulls and terns prefer open sandy areas for nesting (Kushlan and White 1977b) and use mangrove ecosystems only for feeding. All of the species in this guild are recorded from a variety of coastal and inland wetland habitats.
9.6 BIRDS OF PREY
This guild is composed of 20 species of hawks, falcons, vultures, and owls
67
which utilize mangrove swamps in south are common inhabitants of mangrove areas.
Florida (Appendix D). The magnificant This could also be true for the merlin,
frigatebird has been included in this which like the peregrine falcon, feeds on
group because of its habit of robbing many waterfowl and shorebirds. The remaining
of these birds of their prey. Prey con- species in this guild are probably not so
sumed by this guild includes snakes, dependent on mangroves; although they may
lizards, frogs (red-shouldered hawk, be common in mangrove ecosystems, they
swallow-tailed kite), small birds (short- utilize other habitats as well, tailed hawk), waterfowl (peregrine falcon, great-horned owl), fishes (osprey, bald
eagle), and carrion (black and turkey 9.7 ARBOREAL BIRDS vultures).
This guild is the largest (71
Eleven of these species are permanent species) and most diverse group inhabiting
residents, one a summer resident, and the mangrove forests. Included are pigeons,
remainder are winter residents. Their use cuckoos, woodpeckers, flycatchers,
of mangrove areas varies greatly. The thrushes, vireos, warblers, blackbirds,
magnificent frigatebird, which occurs and sparrows. We have lumped this diverse
principally in extreme southern Florida group together because they utilize man-
and the Florida Keys, utilizes small over- grove ecosystems in remarkably similar
wash mangrove islands for both roosts and ways. Invertebrates, particularly
nesting colonies. Both species of vul- insects, make up a significant portion of
tures are widely distributed in south most of these birds' diets, although the
Florida mangrove regions; large colonial white-crowned pigeon, mourning dove, and
roosts can be found in mangrove swamps many of the fringilids (cardinal, townee)
near the coast. Swallow-tailed kites are eat a variety of seeds, berries, and
common over the entire Florida mangrove fruits, region (Robertson 1955; Snyder 1974).
Snyder (1974) reports extensively on the As the name given this guild implies,
breeding biology of the swallow-tailed these birds use the habitat provided by
kites in south Florida. The nests he the mangrove canopy. Many birds also use
observed were all located in black man- the trunk, branches, and aerial roots for
groves although they do nest in other feeding. Several different types of
habitats. searching patterns are used. Hawking of
insects is the primary mode of feeding by
The bald eagle, osprey (Figure 15), the cuckoos, chuck-wi 1 1 s-wi dows, the
and peregrine falcon are dependent upon kingbirds, and the flycatchers. Gleaning
mangrove ecosystems for their continued is employed by most of the warblers,
existence in south Florida. Both the bald Woodpeckers and the prothonotary warbler
eagle and osprey feed extensively on the are classic probers, wealth of fishes found associated with
mangrove ecosystems. Additionally, man- Several of the birds in this guild groves are used as roosts and support are heavily dependent upon mangrove areas, structures for nests. Nisbet (1968) indi- The prairie warbler and the yellow warbler cated that in Malaysia the most important are subspecies of more widespread North role of mangroves for birds may be as American species (see Appendix D for wintering habitat for palaearctic mi- scientific names). They are found largely grants, of which the peregrine falcon is within mangrove areas (Robertson and one. Kushlan (pers. comm.) stated that Kushlan 1974). The white-crowned pigeon, recent surveys have shown falcons to mangrove cuckoo, gray kingbird, and black- winter in mangroves, particularly along whiskered vireo are of recent West Indian the shore of Florida Bay where they estab- origin. They first moved into the lish feeding territories. They forage on mangrove-covered regions of south Florida concentrations of shorebirds and water- from source areas in the islands of the fowl. These prey species of the peregrine Caribbean. Confined at first to mangrove
68
Figure 15. Osprey returning to its nest in a red mangrove tree near Whitewater Bay. Photograph by David Scott.
69
swamps, all but the mangrove cuckoo have expanded their range in peninsular Florida by using non-mangrove habitat. In this vein it is interesting to note that many species of rare and/or irregular occur- rence in south Florida are of West Indian origin and use mangroves to a considerable extent. These include the Bahama pintail, masked duck, Caribbean coot, loggerhead kingbird, thick-billed vireo, and stripe- headed tanager (Robertson and Kushlan 1974).
Twenty-four of the species in this guild are permanent residents, 27 are win- ter, and 6 are summer residents. Fourteen species are seen only during migrations.
9.8 ASSOCIATIONS BETWEEN COMMUNITY TYPES AND BIRDS
MANGROVE
Estimating the degree of use of mangrove swamps by birds as we have done (Appendix D) is open to criticism because of the paucity of information upon which to base judgements. Estimating which mangrove community types (see section 1, Figure 4) are used by which birds is open to even more severe criticism. For this reason the following comments should be regarded as general and preliminary.
In terms of utilization by avifauna, the scrub mangrove swamps are probably the least utilized mangrove community type. Because the canopy is poorly developed, most of the arboreal species are absent, although Emlen (1977) recorded the red- winged blackbird, hairy woodpecker, north- ern waterthrush, yellow-rumped warbler, common yel 1 owth roat , orange-crowned warbler, palm warbler, yellow warbler, mourning dove, and gray kingbird in scrub mangroves on Grand Bahama Island. Of 25 different habitats surveyed by Emlen (1977), the yellow warbler and gray kingbird were found in the scrub mangroves only. Aerially-searching and wading birds might use scrub mangroves if fishes are present.
Overwash mangrove islands are utilized in a variety of ways by all of the bird guilds. Most of the wading birds
plus the magnificent f ri gatebi rd, the anhinga, the cormorant, and the brown pelican use overwash islands for nesting (Kushlan and White 1977a). Wading and aerially-searching birds commonly feed in close proximity to overwash islands. A variety of migrating arboreal and probing species use the islands for feeding and roosting. Yellow and palm warblers are common around mangrove islands in Florida Bay as are the black-bellied plover, ruddy turnstone, willet, dunlin, and short- billed dowitcher. Rafts of ducks are common near the inshore islands and birds of prey such as the osprey, the bald eagle, and both vultures use mangrove islands for roosting and nesting.
Fringe and riverine mangrove com- munities are important feeding areas for wading and probing birds. Floating and diving and aerially-searching birds use the lakes and waterways adjacent to these mangrove communities for feeding. Many of the wading birds nest in fringe and riverine forests. For example, when the wood ibis nests in coastal areas, it uses these mangrove communities almost exclu- sively (Kushlan, personal communication). Most of the arboreal birds and birds of prey associated with mangroves are found in these two types of communities. This is not surprising since the tree canopy is extremely wel 1 -devel oped and offers roosting, feeding and nesting opportuni- ties.
Hammock and basin mangrove communi- ties are so diverse in size, location, and proximity to other communities that it is difficult to make many general statements about their avifauna. Since there often is little standing water in hammock forests, wading and diving birds probably are not common. Proximity to terrestrial communities in some cases may increase the diversity of arboreal species in both hammock and basin forests; proximity to open areas may increase the likelihood of bi rds of prey.
It seems safe to conclude that each of the six mangrove community types has some value to the avifauna. This value differs according to community type and
70
kind of bird group under consideration. Certainly, more information is needed, particularly concerning the dependence of rare or endangered species on specific community types.
9.9 MANGROVES AS WINTER HABITAT FOR NORTH AMERICAN MIGRANT LAND BIRDS
An interesting observation based on the data in this chapter is the seemingly important role that mangrove ecosystems play in providing wintering habitat for migrants of North American origin. Lack and Lack (1972) studied the wintering warbler community in Jamaica. In four natural habitats including mangrove forest, lowland dry limestone forest, mid- level wet limestone forest, and montane cloud forest, a total of 174, 131, 61, and 49 warblers (individuals) were seen, respectively. When computed on a per hour of observation basis, the difference is more striking with 22 warblers per hour seen in mangroves and only 1, 2, and 1 seen in the other forest habitats, respec- tively. For all passerines considered together, 26 passerines/hour were seen in mangroves with 5, 13, and 3 respectively in the other forest habitats. On a
species basis only 9 were recorded from mangroves whereas 19, 13, and 16 species, respectively, were seen in the other habi- tats. This large number of species from the other habitats appears to result from the sighting of rare species after many hours of observation. Only 9 hours were spent by Lack and Lack (1972) in the man- groves whereas between 30 and 86 hours were spent in other habitats. More time in the mangrove zone would have undoubted- ly resulted in more species (and in- dividuals) observed (Preston 1979).
Hutto (1980) presented extensive data concerning the composition of migratory land bird communities in Mexico in winter for 13 habitat types. Mangrove areas tended to have more migrant species than most natural habitats (except gallery forests) and also had a greater density of individuals than other habitats (again except for gallery forests). In both Lack and Lack's and Hutto's studies, disturbed and edge habitats had the highest number of species and greatest density of individuals. The percentage of the avifauna community composed of migrants was highest in mangrove habitats, however. From this we can infer the importance of mangroves in the maintenance of North American migrant land birds.
71
CHAPTER 10. COMMUNITY COMPONENTS - MAMMALS
Thirty-six native and nine introduced (Layne 1974). Hamilton and Whittaker species of land mammals occur in the south (1979) state that it is the coastal ham- Florida region (Layne 1974; Hamilton and mocks of south Florida, including mangrove Whittaker 1979). Of these, almost 50% (18 areas, which serve to preserve this species) are found in the mangrove zone species in the Eastern United States. (Layne 1974). In addition, two species of Shemnitz (1974) reported that most of the marine mammals are known from mangrove remaining panthers were found in the areas. Data on the abundance and food southwest portion of Florida along the habits of these 20 species are summarized coast and in the interior Everglades in Appendix E. All are permanent resi- regions, dents. The criteria for inclusion in this
table are similar to those used for the The extent to which other carnivores
avifauna. Sight records in mangroves or use mangrove areas varies widely among
locality data from known mangrove areas species. Schwartz (1949) states that
were required before a species was in- mink, although rare, prefer mangroves to
eluded. This has produced a conservative other coastal habitats in Florida. Layne
estimate of the mammal species that uti- (1974, see his figure 1) gives a disjunct
lize mangrove areas. distribution for this species in Florida,
with the major geographical range being
Several mammals do not appear in the southwest coast. River otters also
Appendix E because they have not been utilize mangrove habitat heavily. Otters
recorded from mangrove swamps in south have been found even far from shore on
Florida; however, they occur so widely small mangrove overwash islands in Florida
that we suspect they will be found in this Bay (Layne 1974). Gray fox are not depen-
habitat in the future. This group dent upon mangroves, although they occa-
includes the cotton mouse, Peromyscus sionally use this habitat. Less than 20%
gossypinus, the hispid cotton rat, Si g- of all sightings of this species in Ever-
modon hispidus, the round-tailed muskrat, glades National Park were from mangroves
Neof iber a! leni , the house mouse, Mus (Layne 1974). Bobcat are found in almost
musculus, the least shrew, Cryptoti s all habitats in south Florida from pine-
parva, and the short -tailed shrew, Blarina lands to dense mangrove forests. The
brevicauda. preponderance of recent sightings, how- ever, has been made from the mangrove
Few rodents and no bats are included zone, particularly on offshore mangrove in Appendix E. Compared to the rest of overwash islands (Layne 1974). Black bear the State, the south Florida region is are apparently most abundant in the Big deficient in these two groups (Layne Cypress Swamp of Collier County (Shemnitz 1974). Although we have no confirmative 1974) and are rare in the remainder of field data, we suspect that mangrove south Florida, swamps along the central and north Florida coasts contain more mammal species, par- ticularly rodents and bats. The small mammal fauna of the man- grove zone of south Florida are predomi-
A number of medium-sized and large nately arboreal and terrestrial species
carnivores, including panther, gray fox, which are adapted to periodic flooding,
bobcat, striped skunk, raccoon, mink, Opossum, marsh rabbits, cotton rats, and
river otter, and black bear, appear to rice rats are commonly found in mangrove
utilize south Florida mangrove areas. swamps. The Cudjoe Key rice rat is a
Only three of these species (striped newly described species found only on
skunk, raccoon, and bobcat) are common in Cudjoe Key in the Florida Keys. This
mangroves, but several of the rarer species appears to be closely associated
species seem to be highly dependent on with stands of white mangroves (Hamilton
mangrove swamps. Of 18 recent sightings and Whittaker 1979). of the panther in Everglades National
Park, 15 were from mangrove ecosystems White-tailed deer are common in
72
Florida mangrove swamps, although they utilize many other habitats. The key deer, a rare and endangered subspecies, is restricted to the Big Pine Key group in the Florida Keys, although it ranged onto the mainland in historical times. Al- though this little deer makes use of pine uplands and oak hammocks, it extensively exploits mangrove swamps for food and cover.
Two marine mammals, the bottlenose porpoise and the manatee, frequent mangrove-lined waterways. The bottlenose porpoise feeds on mangrove-associated fishes such as the striped mullet, Mugi 1 cephal us. Although the manatee feeds
primarily upon sea grasses and other submerged aquatic plants, it is commonly found in canals, coastal rivers, and embayments close to mangrove swamps.
Except for the Cudjoe Key rice rat, none of the mammals found in Florida man- groves are solely dependent upon mangrove ecosystems; all of these species can utilize other habitats. The destruction of extensive mangrove swamps would, how- ever, have deleterious effects on almost all of these species. Populations of panther, key deer, and the river otter would probably be the most seriously affected, because they use mangrove habi- tat extensively.
73
CHAPTER 11
VALUE OF MANGROVE ECOSYSTEMS TO MAN
Mangrove swamps are often hot, fetid, mosquito-ridden, and almost impenetrable. As a consequence, they are frequently held in low regard. It is possible that more acres of mangrove, worldwide, have been obliterated by man in the name of "recla- mation" than any other type of coastal environment. Reclamation, according to Webster's, means "to claim back, as of wasteland". Mangrove swamps are anything but wasteland, however, and it is impor- tant to establish this fact before a valuable resource is lost. We can think of six major categories of mangrove values to man; no doubt, there are more.
11.1 SHORELINE STABILIZATION AND STORM PROTECTION
The ability of all three Florida mangroves to trap, hold and, to some extent, stabilize intertidal sediments has been demonstrated repeatedly (reviewed by Scoffin 1970; Carlton 1974). The contem- porary view of mangroves is that they function not as "land builders" as hypo- thesized by Davis (1940) and others, but as "stabilizers" of sediments that have been deposited largely by geomorphological processes (see section 3.2).
Gill (1970), Savage (1972), Teas (1977), and others have emphasized that land stabilization by mangroves is pos- sible only where conditions are relatively quiescent and strong wave action and/or currents do not occur. Unfortunately, no one has devised a method to predict the threshold of physical conditions above which mangroves are unable to survive and stabilize the sediments. Certainly, this depends to some extent on substrate type; mangroves appear to withstand wave energy best on solid rock substrates with many cracks and crevices for root penetration. From our own experience, we suspect that mangroves on sandy and muddy substrates cannot tolerate any but the lowest wave energies, tidal currents much above 25 cm/s, or heavy, regular boat wakes.
The concept that the red mangrove is the best land stabilizer has been ques-
tioned by Savage (1972), Carlton (1974), and Teas (1977). These authors argue that the black mangrove (1) is easier to transplant as a seedling, (2) establishes its pneumatophore system more rapidly than the red mangrove develops prop roots, (3) has an underground root system that is better adapted to holding sediments (Teas 1977), (4) is more cold-hardy, and (5) can better tolerate "artificial" substrates such as dredge-spoil, finger fills, and causeways. Generally, the white mangrove is regarded as the poorest land stabilizer of the Florida mangroves (Hanlon et al. 1975).
Although mangroves are susceptible to hurricane damage (see section 12.1), they provide considerable protection to areas on their landward side. They cannot prevent all flooding damage, but they do mitigate the effects of waves and breakers. The degree of this protection is roughly proportional to the width of the mangrove zone. Very narrow fringing forests offer minimal protection while extensive stands of mangroves not only prevent wave damage, but reduce much of the flooding damage by damping and holding flood waters. Fosberg (1971) suggested that the November 1970 typhoon and accom- panying storm surge that claimed between 300,000 and 500,000 human lives in Bangladesh might not have been so destruc- tive if thousands of hectares of mangrove swamps had not been replaced with rice paddies.
11.2 HABITAT VALUE TO WILDLIFE
Florida mangrove ecosystems are important habitat for a wide variety of reptiles, amphibians, birds, and mammals (see sections 8, 9, and 10). Some of these animals are of commercial and sport importance (e.g., white-tailed deer, sea turtles, pink shrimp, spiny lobster, snook, grey snapper). Many of these are important to the south Florida tourist industry including the wading birds (e.g., egrets, wood stork, white ibis, herons) which nest in the mangrove zone.
74
11.3 IMPORTANCE TO THREATENED AND ENDAN- GERED SPECIES
The mangrove forests of south Florida are important habitat for at least seven endangered species, five endangered sub- species, and three threatened species (Federal Register 1980). The endangered species include the American crocodile, the hawksbill sea turtle, the Atlantic ridley sea turtle, the Florida manatee, the bald eagle, the American peregrine falcon, and the brown pelican. The endan- gered subspecies are the key deer (Odocoi leus vi rginianus cl avi urn), the Florida panther (Felis concolor coryi ), the Barbados yellow warbler (Dendroica petechia petechia), the Atlantic saltmarsh snake (Nerodia fasciata taeniata) and the eastern indigo snake (Drymarchon corais couperi). Threatened species include the American alligator, the green sea turtle and the loggerhead sea turtle. Although all of these animals utilize mangrove habitat at times in their life histories, species that would be most adversely affected by widespread mangrove destruc- tion are the American crocodile, the Florida panther, the American peregrine falcon, the brown pelican, and the Atlantic ridley sea turtle. The so-called mangrove fox squirrel (Sciurus ni ger avicennia) is widely believed to be a mangrove-dependent endangered species. This is not the case since it is currently regarded as "rare", not endangered, and, further, there is some question whether or not this is a legitimate sub-species (Hall 1981). As a final note, we should point out that the red wolf (Cam's rufus), which is believed to be extinct in Florida, at one time used mangrove habitat in addition to other areas in south Florida.
11.4 VALUE TO SPORT AND COMMERCIAL FISHERIES
The fish and invertebrate fauna of mangrove waterways are closely linked to mangrove trees through (a) the habitat value of the aerial root structure and (b) the mangrove leaf detritus-based food web (see sections 6 and 7). The implications
of these connections were discussed by Heald (1969), Odum (1970), Heald and Odum (1970), and Odum and Heald (1975b) in terms of support for commercial and sport fisheries.
A minimal list of mangrove-associated organisms of commercial or sport value includes oysters, blue crabs, spiny lobsters, pink shrimp, snook, mullet, menhaden, red drum, spotted sea trout, gray and other snapper, tarpon, sheepshead, ladyfish, jacks, gafftopsail catfish, and the jewfish. Heald and Odum (1970) pointed out that the commercial fisheries catch, excluding shrimp, in the area from Naples to Florida Bay was 2.7 million pounds in 1965. Almost all of the fish and shellfish which make up this catch utilize the mangrove habitat at some point during their life cycles. In addi- tion, the Tortugas pink shrimp fishery, which produces in excess of 11 million pounds of shrimp a year (Idyll 1965a), is closely associated with the Everglades estuary and its mangrove-lined bays and rivers.
11.5 AESTHETICS, INTANGIBLES
TOURISM AND THE
One value of the mangrove ecosystem, which is difficult to document in dollars or pounds of meat, is the aesthetic value to man. Admittedly, not all individuals find visits to mangrove swamps a pleasant experience. There are many others, how- ever, who place a great deal of value on the extensive vistas of mangrove canopies, waterways, and associated wildlife and fishes of south Florida. In a sense, this mangrove belt along with the remaining sections of the freshwater Everglades and Big Cypress Swamp are the only remaining wilderness areas in this part of the United States.
Hundreds of thousands of visitors each year visit the Everglades National Park; part of the reason for many of these visits includes hopes of catching snook or gray snappers in the mangrove-lined water- ways, seeing exotic wading birds, croco- diles, or panthers, or simply discovering
75
what a tropical mangrove forest looks like. The National Park Service, in an attempt to accommodate this last wish, maintains extensive boardwalks and canoe trails through the mangrove forests near Flamingo, Florida. In other, more developed parts of the State, small stands of mangroves or mangrove islands provide a feeling of wilderness in proximity to the rapidly burgeoning urban areas. A variety of tourist attractions including Fairchild Tropical Gardens near Miami and Tiki Gardens near St. Petersburg utilizes the exotic appearance of mangroves as a key ingredient in an attractive landscape. Clearly, mangroves contribute intangibly by diversifying the appearance of south Florida.
11.6 ECONOMIC PRODUCTS
Elsewhere in the world, mangrove forests serve as a renewable resource for many valuable products. For a full dis- cussion of the potential uses of mangrove products, see de la Cruz (in press a), Morton (1965) for red mangrove products, and Moldenke (1967) for black mangrove products.
In many countries the bark of man- groves is used as a source of tannins and dyes. Since the bark is 20% to 30% tannin on a dry weight basis, it is an excellent source (Hanlon et al. 1975). Silviculture (forestry) of mangrove forests has been practiced extensively in Africa, Puerto Rico, and many parts of Southeast Asia (Holdridge 1940; Noakes 1955; Macnae 1968; Walsh 1974; Teas 1977). Mangrove wood
makes a durable and water resistant timber which has been used successfully for resi- dential buildings, boats, pilings, hogsheads, fence posts, and furniture (Kuenzler 1974; Hanlon et al. 1975). In Southeast Asia mangrove wood is widely used for high quality charcoal.
Morton (1965) mentions that red man- grove fruits are somtimes eaten by humans in Central America, but only by popula- tions under duress and subject to starva- tion. Mangrove leaves have variously been used for teas, medicinal purposes, and livestock feeds. Mangrove teas must be drunk in small quantities and mixed with milk because of the high tannin content (Morton 1962); the milk binds the tannins and makes the beverage more palatable.
As a final note, we should point out that mangrove trees are responsible for contributing directly to one commercial product in Florida. The flowers of black mangroves are of considerable importance to the three million dollar (1965 figures) Florida honey industry (Morton 1964).
Other than the honey industry, most of these economic uses are somewhat destructive. There are many cases in which clear-cut mangrove forests have failed to regenerate successfully for many years because of lack of propagule dispersal or increased soil salinities (Teas 1979). We believe that the best use of Florida mangrove swamps will continue to be as preserved areas to support wildlife, fishing, shoreline stabiliza- tion, endangered species, and aesthetic values.
76
CHAPTER 12. MANAGEMENT IMPLICATIONS
12.1 INHERENT VULNERABILITY
Mangroves have evolved remarkable physiological and anatomical adaptations enabling them to flourish under conditions of high temperatures, widely fluctuating salinities, high concentrations of heavy metals (Walsh et al. 1979), and anaerobic soils. Unfortunately, one of these adap- tations, the aerial root system, is also one of the plant's most vulnerable compo- nents. Odum and Johannes (1975) have referred to the aerial roots as the man- grove's Achi 1 1 es' heel because of their susceptibility to clogging, prolonged flooding, and boring damage from isopods and other invertebrates (see section 6 for a discussion of the latter). This means that any process, natural or man-induced, which coats the aerial roots with fine sediments or covers them with water for extended periods has the potential for mangrove destruction. Bacon (1970) men- tions a case in Trinidad where the Caroni River inundated the adjacent Caroni Mangrove Swamp during a flood and deposited a layer of fine red marl in a large stand of black mangroves which sub- sequently died. Many examples of damage to mangrove swamps from human activities have been documented (see section 12.2).
One of the few natural processes that causes periodic and extensive damage to mangrove ecosystems is large hurricanes (Figure 16). Craighead and Gilbert (1962) and Tabb and Jones (1962) have documented the impact of Hurricane Donna in 1960 on parts of the mangrove zone of south Florida. Craighead and Gilbert (1962) found extensive damage over an area of 100,000 acres (40,000 ha). Loss of trees ranged from 25% to 100%. Damage occurred in three ways: (1) wind shearing of the trunk 6 to 10 ft (2 to 3 m) above ground, (2) overwash mangrove islands being swept clean, and (3) trees dying months after the storm, apparently in response to damage to the prop roots from coatings by marl and fine organic matter. The latter type of damage was most widespread, but rarely occurred in intertidal forests, presumably because the aerial roots were flushed and cleaned by tidal action. Fish and invertebrates were adversely affected
by oxygen depletion due to accumulations of decomposing organic matter (Tabb and Jones 1962).
Hurricane Betsy in 1965 did little damage to mangroves in south Florida; there was also little deposition of silt and marl within mangrove stands from this minimal storm (Alexander 1 967). Lugo et al. (1976) have hypothesized that severe hurricanes occur in south Florida and Puerto Rico on a time interval of 25 to 30 years and that mangrove ecosystems are adapted to reach maximum biomass and pro- ductivity on the same time cycle.
12.2 MAN-INDUCED DESTRUCTION
Destruction of mangrove forests in Florida has occurred in various ways including outright destruction and land filling, diking and flooding (Figure 17), through introduction of fine particulate material, and pollution damage, par- ticularly oil spills. To our knowledge there are no complete, published docu- mented estimates of the amount of mangrove forests in Florida which have been destroyed by man in this century. Our conclusion is that total loss statewide is not too great, probably in the range of 3 to 5% of the original area covered by mangroves in the 19th century, but that losses in specific areas, particularly urban areas, are appreciable. This con- clusion is based on four pieces of infor- mation. (1) Lindall and Saloman (1977) have estimated that the total loss of vegetated intertidal marshes and mangrove swamps in Florida due to dredge and fill is 23,521 acres (9,522 ha); remember that there are between 430,000 and 500,000 acres (174,000 to 202,000 ha) of mangroves in Florida (see section 1.3). (2) Birnhak and Crowder (1974) estimate a loss of approximately 11,000 acres (4,453 ha) of mangroves between 1943 and 1970 in three counties (Collier, Monroe, and Dade). (3) An obvious loss of mangrove forests has occurred in Tampa Bay, around Marco Island, in the Florida Keys, and along the lower east coast of Florida. For example, Lewis et al. (1979) estimated that 44% of the intertidal vegetation
77
Figure 16. Damaged stand of red and black mangroves near Flamingo, Florida, as it appeared 7 years after Hurricane Donna.
78
Figure 17. Mangrove forest near Key West as it appeared in 1981 after being destroyed by diking and impounding.
79
including mangroves in the Tampa Bay estuary has been destroyed during the past 100 years. (4) Heald (unpublished MS.) has estimated a loss of 2,000 acres (810 ha) of mangroves within the Florida Keys (not considered by Birnhak and Crowder 1974). So while loss of mangrove ecosys- tems throughout Florida is not over- whelming, losses at specific locations have been substantial.
Diking, impounding, and long-term flooding of mangroves with standing water can cause mass mortality, especially when prop roots and pneumatophores are covered (Breen and Hill 1969; Odum and Johannes 1975; Patterson-Zucca 1978; Lugo 1981). In south Florida, E. Heald (pers. comm.) has observed that permanent impoundment by diking which prevents any tidal exchange and raises water levels significantly during the wet season will kill all adult red and black mangrove trees. If condi- tions behind the dike remain relatively dry, the mangroves may survive for many years until replaced by terrestrial vege- tation.
Mangroves are unusually susceptible to herbicides (Walsh et al. 1973). At least 250,000 acres (100,000 ha) of man- grove forests were defoliated and killed in South Viet Nam by the U.S. military. This widespread destruction has been docu- mented by Tschirley (1969), Orians and Pfeiffer (1970), Westing (1971), and a committee of the U.S. Academy of Sciences (Odum et al. 1974). In many cases these forests were slow to regenerate; observa- tions by de Sylva and Michel (1974) indi- cated higher rates of siltation, greater water turbidity, and possibly lower dis- solved oxygen concentrations in swamps which sustained the most damage. Teas and Kelly (1975) reported that in Florida the black mangrove is somewhat resistant to most herbicides but the red mangrove is extremely sensitive to herbicide damage. He hypothesized that the vulnerability of the red mangrove is related to the small reserves of viable leaf buds in this tree. Following his reasoning, the stress of a single defoliation is sufficient to kill the entire tree.
Although mangroves commonly occur in areas of rapid sedimentation, they cannot survive heavy loads of fine, floculent materials which coat the prop roots. The instances of mangrove death from these substances have been briefly reviewed by Odum and Johannes (1975). Mangrove deaths from fine muds and marl, ground bauxite and other ore wastes, sugar cane wastes, pulp mill effluent, sodium hydroxide wastes from bauxite processing, and from intrusion of large quantities of beach sand have been documented from various areas of the world.
12.3 EFFECTS OF OIL SPILLS ON MANGROVES
There is little doubt that petroleum and petroleum byproducts can be extremely harmful to mangroves. Damage from oil spills has been reviewed by Odum and Johannes (1975), Carlberg (1980), Ray (in press), and de la Cruz (in press, b). Over 100 references detailing the effects of oil spills on mangroves and mangrove- associated biota are included in these reviews.
Petroleum and its byproducts injure and kill mangroves in a variety of ways. Crude oil coats roots, rhizomes, and pneu- matophores and disrupts oxygen transport to underground roots (Baker 1971). Various reports suggest that the critical concentration for crude oil spills which may cause extensive damage is between 100 and 200 ml/m of swamp surface (Odum and Johannes 1975). Petroleum is readily absorbed by lipophylic substances on sur- faces of mangroves. This leads to severe metabolic alterations such as displacement of fatty molecules by oil hydrocarbons leading to destruction of cellular permea- bility and/or dissolution of hydrocarbons in lipid components of chloroplasts (Baker 1971).
As with other intertidal communities, many of the invertebrates, fishes, and plants associated with the mangrove com- munity are highly susceptible to petroleum products. Widespread destruction of organisms such as attached algae, oysters, tunicates, crabs, and gobies have been reported in the literature (reviewed by de
80
la Cruz in press, b; Ray in press).
12.4 MAN-INDUCED MODIFICATIONS
Damage from oil spills follows a predictable pattern (Table 7) which may require years to complete. It is impor- tant to recognize that many of the most severe responses, including tree death, may not appear for months or even years after the spill.
In Florida, Chan (1977) reported that red mangrove seedlings and black mangrove pneumatophores were particularly sensitive to an oil spill which occurred in the Florida Keys. Lewis (1979a, 1980b) has followed the long-term effects of a spill of 150,000 liters (39,000 gal) of bunker C and diesel oil in Tampa Bay. He observed short-term (72-hour) mortality of inverte- brates such as the gastropod Mel ongena corona and the polychaete Laeonerei s cul veri. Mortality of all three species of mangroves began after three weeks and continued for more than a year. Sub- lethal damage included partial defoliation of all species and necrosis of black mangrove pneumatophores; death depended upon the percentage of pneumatophores affected.
In addition to the damage from oil spills, there are many adverse impacts on mangrove forests from the process of oil exploration and drilling (Table 8). This type of damage can often be reduced through careful management and monitoring of drilling sites.
Although little is known concerning ways to prevent damage to mangroves once a spill has occurred, protection of aerial roots seems essential. Prop roots and pneumatophores must be cleaned with com- pounds which will not damage the plant tissues. Dispersants commonly used to combat oil spills are, in general, toxic to vascular plants (Baker 1971). If pos- sible, oil laden spray should not be allowed to reach leaf surfaces. Damage during clean-up (e.g., trampling, compac- tion, bulldozing) may be more destructive than the untreated effects of the oil spill (de la Cruz in press, b).
In south Florida, man has been re- sponsible for modifications which, while not killing mangroves outright, have al- tered components of the mangrove ecosys- tem. One of the most widespread changes involves the alteration of freshwater runoff. Much of the freshwater runoff of the Florida Everglades has been diverted elsewhere with the result that salinities in the Everglades estuary are generally higher than at the turn of the century. Teas (1977) points out that drainage in the Miami area has lowered the water table as much as 2 m (6 ft).
Interference with freshwater inflow has extensive effects on estuaries (Odum 1970). Florida estuaries are no excep- tion; the effects on fish and invertebrate species along the edge of Biscayne and Florida Bays have been striking. The mismanagement of freshwater and its effects on aquatic organisms have been discussed by Tabb (1963); Idyll (1965a,b); Tabb and Yokel (1968) and Idyll et al. (1968). In addition, Estevez and Simon (1975) have hypothesized that the impact of the boring isopod, Sphaeroma terebrans, may be more severe when freshwater flows from the Everglades are altered.
One generally unrecognized side effect of lowered freshwater flow and salt water intrusion has been the inland expan- sion of mangrove forests in many areas of south Florida. There is documented evi- dence that the mangrove borders of Biscayne Bay and much of the Everglades estuary have expanded inland during the past 30 to 40 years (Reark 1975; Teas 1979; Ball 1980).
Sections of many mangrove forests in south Florida have been replaced by filled residential lots and navigation canals. Although these canal systems have not been studied extensively, there is some evi- dence, mostly unpublished, that canals are not as productive in terms of fishes and invertebrates as the natural mangrove- lined waterways which they replaced.
81
Table 7. General response of mangrove ecosystems to severe oil spills (from Lewis 1980b)
Stage
Observed impact
Acute
0 to 15 days
15 to 30 days
Deaths of birds, turtles, fishes, and invertebrates
Defoliation and death of small mangroves, loss of aerial root community
Chronic
30 days to 1 year
1 year to 5 years
1 year to 1 0 years ( ? )
10 to 50 years (?)
Defoliation and death of medium-sized mangroves (1-3 m), tissue damage to aerial roots
Death of large mangroves (greater than 3 m), loss of oiled aerial roots, and regrowth of new roots (often deformed)
Recol oni zati on of oil-damaged areas by new seedlings
Reduction in litter fall, reduced re- production, and reduced survival of seedl 1 ngs
Death or reduced growth of young trees colonizing spill site (?)
Increased insect damage (?)
Complete recovery
82
|
1 |
C= 1 |
||||||||||||||||
|
TJ |
i |
1 |
o c |
||||||||||||||
|
ai |
>, |
+-> |
i |
•r- IO |
|||||||||||||
|
to |
X o |
IO |
IO |
-c: U |
+-> +J 1- to |
||||||||||||
|
4- |
3 |
3 |
LO XI |
||||||||||||||
|
o |
-a |
QJ |
CL |
CO |
O 3 |
||||||||||||
|
to |
cu |
en c |
O |
CL IO |
|||||||||||||
|
10 |
UJ |
to to |
> |
io o |
CL |
CO |
E |
||||||||||
|
e |
c |
ai |
>> cu |
r- |
c: i- |
cu |
o o |
||||||||||
|
0) |
o |
S- |
IO +J |
O |
•i- +■> |
i — |
to |
O 1- |
|||||||||
|
+J |
■f" |
<o |
3 io |
to |
■O (J |
iO |
CO |
CU X |
|||||||||
|
to |
+J |
x: s_ |
to |
S- 3 |
E |
CD |
-a o |
||||||||||
|
>> |
IO |
-a |
+-> |
•r- |
-o S- |
•i— |
o |
+-> |
|||||||||
|
to |
+J |
a) |
iO s- |
-o |
LO |
+-> |
ll |
c |
o |
•o |
|||||||
|
O |
ai |
XI |
Q_ CD |
d) |
+-> -o to |
CU |
IO |
i- |
c +j |
||||||||
|
O |
en |
s. |
x: |
-o |
O |
n3 c C |
o |
CL |
IO c |
||||||||
|
<V |
cu |
c to |
3 |
3 en |
ai |
>> c |
ai |
>>•<-> |
c |
■o |
cu |
||||||
|
> |
o cu |
+-> |
O •■- |
s_ |
+J |
ia |
M- |
+j |
-.- ■— O |
IO |
c |
E |
c +j |
||||
|
0) |
•r- ai |
to |
■ — x: |
aj |
to |
f— |
+-> |
•i— |
■f |
XI 4J |
-l-> |
IO |
CD |
o to |
|||
|
> |
cn+J > |
•r— |
4- |
s |
s- |
T3 |
to |
■ — |
-a |
rd CU CD |
to |
■u |
•f— -i— |
||||
|
o |
c |
o ai |
T3 |
* |
o |
a> |
•i— |
-Q |
■o |
•i— |
x: 3 c |
XI |
4-> |
LO |
+J to |
||
|
J- |
•r- |
(0 i— |
s- >> |
. — |
+j |
XI |
3 |
^~ |
XI |
•i- |
3 |
c |
>> |
o $- |
|||
|
en |
s_ |
a. |
C |
(V +J |
IO |
S- |
to |
•r- |
t- |
■— c: i— |
to |
IO |
CO |
3 ai |
|||
|
c |
IO |
E ■— |
t— |
■!-> •■- |
■o |
5 |
3 |
3 |
3 |
IO •■- <U |
i — |
O |
-a cl |
||||
|
IO |
ai |
o <o |
iO XJ |
c |
4-> |
«J |
■»-> |
C CL |
u |
CL |
U |
o |
CO |
||||
|
E |
(_ |
o s_ |
+-> |
s -^ |
la |
>> |
t— |
t»- |
O IO 1— |
f— |
CD |
S_ 4- |
|||||
|
u |
3 |
10 |
XI |
XI |
x: |
X |
o |
x: |
•■- 0) Q. |
X |
M- |
CL o |
1— |
||||
|
C <-» |
J* ■!-> |
4-> |
4- S_ |
•» |
s- |
en o |
cn+j en |
o |
O |
4- |
o |
||||||
|
O XI |
•o |
o ia |
•r- |
O 3 |
c |
10 |
•1— |
•!-> |
+-> |
•i— |
•^ c E |
-u |
O |
>1 c |
LO |
||
|
* |
c |
(O c |
XI |
4-> |
o |
a> |
x: |
c |
x: |
T3 IO O |
c |
s- o |
|||||
|
en m |
io |
5- |
IO |
C |
•1 — |
c |
t|- |
0) |
■o x: s_ <<- to |
o |
C |
IO -i- |
o |
||||
|
C to |
■!-> O |
x: |
o -o |
+-> |
•a |
o |
E |
T3 |
IO O <4- |
O i— |
•1— |
o |
E 4-> |
+-> |
|||
|
1- Q) |
en |
+J |
•r- <U |
rV |
c |
ai |
cu |
01 |
1 — |
+-> |
■r" |
•i- o |
a |
||||
|
c s. |
10 |
c |
ai |
4- |
4-> tO |
*-> |
-i— |
3 |
0) |
u |
3 |
4- s_ to |
cu ••- |
u </> |
•!-> |
L. 3 |
•i — |
|
■r- CL |
+-> |
•■— |
t— ai |
o |
IO IO |
C |
C |
to |
10 |
C |
O CU c |
LO CL |
3 c |
iO |
cl-o |
||
|
E |
(J |
x: |
o en |
S- 0) |
ai |
c |
•^ |
IO |
1 — |
■!"• |
X S. |
IO to |
S- O |
1- |
o |
to |
|
|
c |
US |
to |
•■- to |
to |
ai s_ |
E |
<D |
+J |
ai |
a. |
+J |
to +J <U |
cu |
4-> •^> |
CD |
to s. |
CD |
|
i— •?- |
a. |
3 |
x; E |
to |
■u o |
•i — |
en |
C |
• — |
to |
c |
VI LP |
f^ r— |
to *-> |
+-> |
10 4J |
CJ |
|
-r- |
E |
S- |
0) it) |
o |
i— c |
o |
CD |
•i— |
o |
O 3 |
CU i- |
0) |
i — |
c |
|||
|
O N |
t— i |
O |
> Q |
-J <XL >— |
o a: |
a |
o |
—1 u_ |
oc o |
Q |
< |
1— 1 |
|||||
|
3 |
|||||||||||||||||
|
4- S_ |
|||||||||||||||||
|
o o |
ai |
1 |
|||||||||||||||
|
</> a |
+j |
to |
to |
1 |
CO |
cu |
|||||||||||
|
0) >— |
to |
t- |
E |
cu |
cu cu |
CO |
!_ |
||||||||||
|
en |
ai |
c |
to |
s_ |
c |
3 CL |
CD |
3 |
|||||||||
|
ia ai |
c |
o |
o |
•i- -o |
•a -■- |
■ — |
+J |
||||||||||
|
4J "0 |
•f- = |
•r— |
+J |
cn |
1*- |
i— c: |
CL |
CD |
a. |
||||||||
|
10 |
■ — to |
■l-> |
iO |
c |
+J |
cu en io |
to |
S- |
3 |
||||||||
|
-a |
01 |
■r" |
•1— |
IO |
CL S= |
1 — •» |
ia |
S- LO |
|||||||||
|
<o c |
>> c |
to |
>. |
1 — |
1 — |
•f- •<- to |
1— +-> |
CJ |
CD |
||||||||
|
3 «J |
CU -f- |
o c |
■u |
. — |
a. |
Q. cn^^ |
■!-> |
1- 3 |
CU 1- |
||||||||
|
o |
> 1— |
c |
a. o |
■i— |
•r— |
T3 C |
tz |
CL O |
• |
cn +J |
|||||||
|
•^ 00 |
</> |
$_ |
o |
<D 1- |
> |
(. |
<4- |
1- 41 10 |
cu |
to | |
cu |
i~ t- |
|||||
|
s_ f~. |
>> |
3 +J |
•f— |
•a 4-> |
•i— |
TJ |
O |
o s_ +> |
E |
3 |
CD |
iO > |
|||||
|
(O CT> |
<D |
to O |
■!-> |
o |
+-> |
TJ |
CL |
■o o |
ia |
XI T- |
|||||||
|
> I— |
> |
x: |
iO |
r— 3 |
o |
o |
c |
C 4- |
•1— |
C i— |
^ |
■l-» |
|||||
|
t- |
4- (/> |
> |
•1- J- |
■o |
-!-> |
o |
O CD O |
3 |
IO XI |
iO |
■a o |
||||||
|
•1- • |
3 |
O = |
(O |
O -M |
•f- |
•i- CJ |
cr |
CD |
C IO |
||||||||
|
o •— |
LO |
(J |
Q. to |
-a |
-o |
■■-> |
■!-> C +J |
cu |
LO 1 — |
S- |
IO |
||||||
|
lO |
>> |
en en |
X |
to C |
at |
cu |
o |
O IO c |
J* •— |
-Q |
CL |
||||||
|
•!-» |
+j |
o |
c c |
ai |
O |
to |
•!-> |
3 |
3 C CU |
C |
IO CD |
" 3 |
|||||
|
O 4J |
•r- |
■i— |
•I— I- |
ai o |
•o |
<o |
S_ |
S- 01 E |
cu |
0) 3 |
CU |
L0 1 |
|||||
|
<a aj |
> |
E |
$- i— |
r— |
en |
01 |
1 — |
■!-> |
+-> 4-> CU |
x: |
1 — |
cc |
LO C |
||||
|
a. |
•f— |
LO |
IO i— |
IO |
-a -a |
s- |
CD |
to |
to C CJ |
■♦-> |
O |
•r— |
CU aj |
||||
|
E >> |
+-> |
•r- |
OJ •■- |
c |
01 IO |
o |
t- |
C |
C T- IO |
o |
■— +J |
i — |
CZ CD |
||||
|
•i- a) |
o |
ai |
I— S_ |
(O |
i- o |
c |
o |
O Bi- |
tm |
^~ |
|||||||
|
i — |
«£ |
co |
<_> Q |
t_) |
q a: |
»— 1 |
uosa |
o |
t_) |
||||||||
|
"O en |
|||||||||||||||||
|
ai c |
|||||||||||||||||
|
+-> o |
|||||||||||||||||
|
ITS -1 |
|||||||||||||||||
|
E |
|||||||||||||||||
|
t- E |
|||||||||||||||||
|
■u o |
|||||||||||||||||
|
to $_ |
|||||||||||||||||
|
UJ <*- |
|||||||||||||||||
|
-o |
c |
||||||||||||||||
|
• 0> |
c |
o |
|||||||||||||||
|
00 -r- |
o |
■f™ |
|||||||||||||||
|
4- |
•r- |
■•-> |
|||||||||||||||
|
0) i- |
+-> |
IO |
|||||||||||||||
|
f— T3 |
IO |
s- |
|||||||||||||||
|
x> O |
S- |
IO |
c |
to |
|||||||||||||
|
* E |
o |
a. |
o |
r— |
|||||||||||||
|
1-^-' |
ai en nj +J |
a. X 01 1 ai S- Q. |
ai S- a. ai CO |
en c •f— s. Q |
•r- +-> o 3 "0 o t- |
•r— CL LO •r— O |
83
Weinstein et al. (1977) found that artifi- cial canals had lower species diversity of benthic infauna and trawl -captured fishes and generally finer sediments than the natural communities. Courtney (1975) reported a number of mangrove-associated invertebrates which did not occur in the artificial channels.
Mosquito production is a serious problem in black mangrove-dominated swamps in Florida (Provost 1969). The salt marsh mosquitos, Aedes taeniorhynchus and A. sol 1 icitans, do not reproduce below the mean high tide mark and for this reason are not a serious problem in the inter- tidal red mangrove swamps. Mosquitos lay their eggs on the damp soil of the irregu- larly flooded black mangrove zone; these eggs hatch and develop when flooded by spring tides, storm tides or heavy rains. As with the "high marsh" of temperate latitudes, there have been some attempts to ditch the black mangrove zone so that it drains rapidly after flooding. Although properly designed ditching does not appear to be particularly harmful to mangrove swamps (other than the area destroyed to dig the ditch and receive the spoil), it is an expensive practice and for this reason is not widely practiced. Properly managed diking can be an effec- tive mosquito control approach with mini- mal side effects to black mangroves (Provost 1969). Generally, ditching or diking of the intertidal red mangrove zone is a waste of money.
Mangrove swamps have been proposed as possible tertiary treatment areas for sewage (see discussion by Odum and Johannes 1975). To our knowledge, this alternate use is not currently practiced in south Florida. Until more experimental results are available on the assimilative capacities and long-term changes to be expected in mangrove forests receiving heavy loads of secondary treated sewage, it would be an environmental risk to use mangrove forests for this purpose.
In many areas of the world mangrove swamps have been converted to other uses such as aquaculture and agriculture (see de la Cruz, in press, a). Although some
of the most productive aquaculture ponds in Indonesia and the Philippines are located in former mangrove swamps, there is some question whether the original natural system was not equally productive in terms of fisheries products at no cost to man (Odum 1974). Conversion to aquaculture and agriculture is cursed with a variety of problems including subsequent land subsidence and the "cat clay" problem. The latter refers to the drastically lowered soil pH which often occurs after drainage and has been traced to oxidation of reduced sulfur compounds (Dent 1947; Tomlinson 1957; Hesse 1961; Hart 1962, 1963; Moorman and Pons 1975). Experience in Africa, Puerto Rico, and Southeast Asia confirms that mangrove forests in their natural state are more valuable than the "reclaimed" land.
12.5 PROTECTIVE MEASURES INCLUDING TRANSPLANTING
Protection of mangroves includes (1) prevention of outright destruction from dredging and filling; (2) prevention of drainage, diking and flooding (except for carefully managed mosquito control); (3) prevention of any alteration of hydrologi- cal circulation patterns, particularly i nvol ving tidal exchange; (4) prevention of introduction of fine-grained materials which might clog the aerial roots, such as clay, and sugar cane wastes; (5) preven- tion of oil spills and herbicide spray driftage; and (6) prevention of increased wave action or current velocities from boat wakes, and sea walls.
Where mangroves have been destroyed, they can be replanted or suitable alter- nate areas can be planted, acre for acre, through mitigation procedures (see Lewis et al. 1979). An extensive body of literature exists concerning mangrove planting techniques in Florida (Savage 1972; Carlton 1974; Pulver 1976; Teas 1977; Goforth and Thomas 1979; Lewis 1979b). Mangroves were initially planted in Florida at least as early as 1917 to protect the overseas railway in the Florida Keys (Teas 1977).
Both red and black mangroves have
84
been used in transplanting. As we men- tioned in section 11, black mangroves seem to have certain advantages over red man- groves. Properly designed plantings are usually 75% to 90% successful, although the larger the transplanted tree, the lower its survival rate (Teas 1977). Pruning probably enhances survival of trees other than seedlings (Carlton 1974). Important considerations (Lewis 1979b; Teas 1977) in transplanting mangroves are: (1) to plant in the intertidal zone and avoid planting at too high or too low an elevation, (2) to avoid planting where the shoreline energy is too great, (3) to avoid human vandalism, and (4) to avoid accumulations of dead sea grass and other wrack.
Costs of transplanting have been variously estimated. Teas (1977) suggests $462 an acre ($l,140/ha) for unrooted propagules planted 3 ft (0.9 m) apart, $1,017 an acre ($2,500/ha) for established seedlings planted 3 ft (0.9 m) apart and $87,500 ($21 6,130/ha) for 3 year-old nur- sery trees planted 4 ft (1.2 m) apart. Lewis (1979b) criticized Teas' costs as unrealistically low and reported a project in Puerto Rico which used established seedlings at a cost of $5,060 an acre ($12,500/ha); he did suggest that this cost could be cut in half for larger projects.
12.6 ECOLOGICAL VALUE OF BLACK VS. RED MANGROVES
One unanswered question of current interest in Florida concerns the ecologi- cal value of black mangrove forests com- pared to intertidal red mangrove forests. In many respects, this is identical to the "high marsh" versus "low marsh" debate in temperate wetlands. One hypothetical argument which has been presented fre- quently in court cases during the past decade suggests that black mangrove forests have less ecological value than red mangrove forests to both man and coastal ecosystems. This argument is based on an apparent lack of substantial particulate detritus export from black mangrove forests above mean high tide and
the generally perceived lack of organisms, particularly gamefishes, which use black mangrove forests as habitat.
The counter argument states that black mangrove forests are important for the support of wildlife and the export of substantial quantities of dissolved organic matter (DOM). Lugo et al. (1980) provide evidence that black mangrove forests do, in fact, export large quanti- ties of DOM. They point out that (1 ) black mangrove leaves decompose more rapidly than red mangrove leaves and thus produce relatively more DOM and (2) abso- lute export of carbon from these forests, on a statewide scale, is equal or greater than from red mangrove forests.
12.7 THE IMPORTANCE OF INTER-COMMUNITY EXCHANGE
From previous discussions (sections 6 and 7.5 and Appendices B, C, D and E) it is clear that many species of fishes, invertebrates, birds, and mammals move between mangrove forest communities and other habitats including sea grass beds, coral reefs, terrestrial forests, and the freshwater Everglades. For example, the gray snapper, Lut janus gri seus, spends part of its juvenile life in sea grass beds, moves to mangrove-lined bays and rivers, and then migrates to deeper water and coral reefs as an adult (Croaker 1962; Starck and Schroeder 1971). The pink shrimp, Penaeus duorarum, spends its juve- nile life in mangrove-lined bays and rivers before moving offshore to the Tortugas grounds as an adult. During its juvenile period it appears to move back and forth from mangrove-dominated areas to sea grass beds. The spiny lobster, Panulirus argus, as a juvenile frequently uses mangrove prop root communities as a refuge; when nearing maturity this species moves to deeper water in sea grass and coral reef communities (see discussion section 6.1). Many of the mammals (sec- tion 10) and birds (section 9) move back and forth between mangrove communities and a variety of other environments.
These are only a few of many
85
examples. Clearly, mangrove ecosystems are linked functionally to other south Florida ecosystems through physical pro- cesses such as water flow and organic carbon flux. As a result, the successful management and/or preservation of many fishes, mammals, birds, reptiles, and amphibians depends on proper understanding and management of a variety of ecosystems and the processes that link them. Saving mangrove stands may do the gray snapper little good if sea grass beds are destroyed. Pink shrimp populations will be enhanced by the preservation of sea grass beds and mangrove-lined waters, but shrimp catches on the Tortugas grounds will decline if freshwater flow from the Everglades is not managed carefully (Idyll et al. 1968). Successful management of south Florida mangrove ecosystems, including their valuable resources, will depend on knowledgeable management of a number of other ecosystems and the processes which link them.
12.8 MANAGEMENT PRACTICES: PRESERVATION
Based on years Florida and based
of research in south on the information
reviewed for this publication, we have concluded that the best management prac- tice for all types of Florida mangrove ecosystems is preservation. Central to this concept is the preservation of adjacent ecosystems that are linked signi- ficantly by functional processes. The continued successful functioning of the mangrove belt of southwest Florida is highly dependent on the continual exis- tence of the Everglades and Big Cypress Swamp in an ecologically healthy condi- tion.
At no cost to man, mangrove forests provide habitat for valuable birds, mam- mals, amphibians, reptiles, fishes, and invertebrates and protect endangered species, at least partially support exten- sive coastal food webs, provide shoreline stability and storm protection, and generate aesthetically pleasing experi- ences (Figure 18). In situations where overwhelming economic pressures dictate mangrove destruction, every effort should be made to ameliorate any losses either through mitigation or through modified development as described by Voss (1969) and Tabb and Heald (1973) in which canals and seawalls are placed as far to the rear of the swamp as possible.
86
Figure 18. Mangrove islands in Florida Bay near Upper Matecumbe Key. Note the extensive stands of seedling red mangroves which have become established (1981) after a long period without major hurricanes. Mangrove islands in the Florida Keys tend to expand during storm- free intervals.
87
REFERENCES
Abele, L.G. 1974. Species diversity of decapod crustaceans in marine habi- tats. Ecology 55: 156-161.
Ahearn, D.G., F.J. Roth, and S.P. Meyers. 1968. Ecology and characterization of yeasts from aquatic regions of south Florida. Mar. Biol. 1: 291- 308.
Albert, R. 1975. Salt regulation in halophytes. Oecologia 21: 57-71.
Alexander, T.R. 1967. Effect of Hurri- cane Betsy on the southeastern Ever- glades. Q. J. Fla. Acad. Sci. 30: 10-24.
Allen, R.P. 1942. Res. Rep. 2. ety, New York,
The roseate spoonbill. National Audubon Soci-
Almodovar, L.R., and R. Biebl. 1962. Osmotic resistance of mangrove algae around La Parguera, P.R. Rev. Algol. (N.S.) 6(3): 203-208.
Atkinson, M.R., G.P. Findlay, A.B. Hope, M.G. Pitman, H.D.W. Saddler, and H.R. West. 1967. Salt regulation in the mangroves Rhizophora mangle Lam. and Aerialitis annulata R. Aust. J. Biol. Sci. 20: 589-599.
Austin, H. 1971. A survey of the ich- thyofauna of the mangroves of western Puerto Rico during December 1967 - Caribb. J. Sci. 11:
August 27-39.
1968.
Austin, H., and S. Austin. 1971. The feeding habitats of some juvenile marine fishes from the mangroves in western Puerto Rico. Caribb. J. Sci. 11: 171-178.
Bacon, P.R. 1970. The ecology of Caroni Swamp. Special Publication, Central Statistical Office, Trinidad. 68 pp.
Bader, R.G., and M.A. Roessler. 1971. Progress report on an ecological study of south Biscayne Bay and Card Sound. Rosenstiel School of Marine and Atmospheric Science, Univ. of Miami, to U.S. Atomic Energy Commis- sion and Florida Power and Light Company.
Bader, R.G., and M.A. Roessler. 1972. Progress report on an ecological study of south Biscayne Bay and Card Sound. Rosenstiel School of Marine and Atmospheric Science, Univ. of Miami, to U.S. Atomic Energy Commis- sion and Florida Power and Light Company.
Bailey, R.M., J.E. Fitch, E.S. Heald, E.A. Lackner, C.C. Lindsey, C.R. Robins, and W.B. Scott. 1970. A list of common and scientific names of fishes from the United States and Canada. 3rd ed. Spec. Publ. 6, Am. Fish. Soc. , Washington, D.C. 150 pp.
Baker, J.M. 1971. The effects of oil on plant physiology. Pages 88-98 j[n E.B. Cowell, ed. Ecological effects of oil pollution. Applied Science Publishers, London.
Baker, M.C., and E.M. Baker. 1973. Niche relationships among six species of shorebirds on their wintering and breeding ranges. Ecol. Monogr. 43: 193-212.
Ball, M.C. 1980. succession in south Florida.
Patterns of secondary a mangrove forest in Oecologia 44: 226-235.
88
Beever, J.W., D. Simberloff, and L.L. King. 1979. Herbivory and predation by the mangrove tree crab, Aratus pisonii. Oecologia 43: 317-328.
Bennett, S.E. 1980. Interspecific compe- tition and the niche of the American redstart (Setophaga ruticilla) in wintering and breeding communities. Pages 319-335 jn A. Keast and E.S. Morton, eds. Birds in the neotrop- ics : ecology, behavior, distribu- tion, and conservation. Smithsonian Inst. Press, Washington, D.C.
Biebl, R. 1962. Protoplasmatischokolog- ische Untersuchungen an Mangrovealgen von Puerto Rico. Protoplasma 55: 572-606.
Birdsong, R.S. 1981. A review of the gob i i d fish genus Microgobius Poey. Bull. Mar. Sci . 31: 267-306.
Birnhak, B.I., and J. P. Crowder. 1974. Evaluation of the extent of vegeta- tive habitat alteration in south Florida, 1943-1970. Appendix to south Florida ecological study. Pre- pared by U.S. Department of the Interior, Bureau of Sport Fish and Wildlife, Atlanta, Ga.
Bohlke, J.E., and C.C.6. Chaplin. 1968. Fishes of the Bahamas and adjacent tropical waters. Academy of Natural Sciences of Philadelphia, Livingston Publ. Co., Wynnewood, Pa. 771 pp.
Boto, K.G., and J.S. Bunt. 1981. Tidal export of particulate organic matter from a northern Australian mangrove system. Estuarine Coastal Shelf Sci. 13: 247-257.
Bowman, H.H.M. 1917. Ecology and physio- logy of red mangroves. Proc. Am. Philos. Soc. 56: 589-672.
Bray, J.R., and E. Gorham. 1964. Litter production in forests of the world. Adv. Ecol. Res. 2: 101-157.
Breder, CM. 1946. An analysis of the deceptive resemblance of fishes to plant parts, with critical remarks on protective coloration, mimicry and adaptation. Bull. Bingham Oceanogr. Collect. 10: 1-49.
Breen, CM., and B.J. Hill. 1969. A mass mortality of mangroves in the Kosi estuary. Trans. R. Soc. S. Afr. 38: 285-303.
Brook, I. 1975. Some aspects of the trophic relationships among the higher consumers in a seagrass com- munity (Thalassia testudinum) in Card Sound, Florida. Dissertation. Univ. of Miami, Coral Gables, Fla. 133 pp.
Burkholder, P.R., and L.R. Almodovar. 1973. Studies on mangrove algal com- munities in Puerto Rico. Fla. Sci. 36: 50-66.
Carlberg, S.R. 1980. Oil pollution of the marine environment—with an emphasis on estuarine studies. Pages 367-402 in E. Olausson and I. Cato, eds. Chemistry and biogeochemistry of estuaries. John Wiley and Sons, New York.
Carlton, J.M. 1974. Land-building and stabilization by mangroves. Environ. Conserv. 1: 285-294.
Carlton, J.M. 1975. A guide to common Florida salt marsh and mangrove vege- tation. Fla. Mar. Res. Publ. 6.
Carlton, J.M. 1977. A survey of coastal vegetation communities of Florida. Fla. Mar. Res. Publ. 30.
Carr, A.W., and C.J. Goin. 1955. Rep- tiles, amphibians and freshwater fishes of Florida. Univ. of Florida, Gainesville. 341 pp.
Carr, W.E.S., and C.A. Adams. 1973. Food habits of juvenile marine fishes occupying seagrass beds in the estua- rine zone near Crystal River, Flor- ida. Trans. Am. Fish. Soc. 102: 511-540.
Carter, M.R., L.A. Burus , T.R. Cavinder, K.R. Dugger, P.L. Fore, D.B. Hicks, H.L. Revells, and T.W. Schmidt. 1973. Ecosystem analysis of the Big Cypress Swamp and estuaries. U.S. Environmental Protection Agency Region IV, Atlanta, Ga.
Casagrande, D.J., and P.H. Given. 1975. Geochemistry of amino acids in some
89
Florida peat accumulations. I. Analytical approach and total amino acid concentration. Geochim. Cosmo- chim. Acta 38: 419-434-
Cawkell, E.M. mangroves 106: 251-
1964. The utilization of by African birds. Ibis 253.
Chan, E.I. 1977. Oil pollution and trop- ical littoral communities: biologi- cal effects of the 1975 Florida Keys oil spill. Pages 539-542 jn Pro- ceedings 1977 oil spill conference. American Petroleum Institute, Envi- ronmental Protection Agency and U.S. Coast Guard: 539-542.
Chapman, V.J. 1970. ciology. Trop.
Mangrove phytoso- Ecol. 11: 1-19.
Chapman, V.J. tion. J.
1976a. Mangrove vegeta- Cramer, Germany. 447 pp.
Chapman, V.J. 1976b. Pergamon Press,
Coastal vegetation. New York. 292 pp.
Cintron, G., A.E. Lugo, D.J. Pool, and G. Morris. 1978. Mangroves of arid environments in Puerto Rico and adja- cent islands. Biotropica 10: 110- 121.
Clark, E., and K. Sharks of the Florida. Bull,
von Schmidt. 1965.
central gulf coast of
Mar. Sci. 15: 13-83.
Clark, S.H. 1570. Factors affecting the distribution of fishes in Whitewater Bay, Everglades National Park, Flor- ida. Dissertation. Univ. of Miami, Coral Gables, Fla.
Clough, B.F. 1981. in Australia. Press, Australia
Mangrove ecosystems National University , 300 pp.
Coastal Coordinating Council, State of Florida. 1974. Florida coastal zone management atlas. Coastal Coordinat- ing Council, Tallahassee, Fla.
Cohen, A.D., and W. Spackman. 1974. The petrology of peats from the Ever- glades and coastal swamps of southern Florida. Miami Geol. Soc. Mem. 2: 233-255.
Coomans, H.E. 1969. Biological aspects of mangrove mollusks in the West Indies. Malacologia 9: 79-84.
Courtney, CM. 1975. Mangrove and sea- wall oyster communities at Marco Island, Florida. Bull. Am. Malacol. Union Inc. 41: 29-32.
Courtney, CM. 1980. Production and decomposition in an impounded black mangrove forest. Fla. Sci. 43 (sup- plement): 23.
Craighead, F.C 1964. and hurricanes. Gard. Bull. 19: 5-
Land, mangroves Fairchild Trop.
32.
Craighead, F.C, and V.C Gilbert. 1962. The effects of Hurricane Donna on the vegetation of southern Florida. Q. J. Fla. Acad. Sci. 25: 1-28.
Croaker, R.A. 1962. Growth and food of the gray snapper, Lutjanus qriseus, in Everglades National Park. Trans. Am. Fish. Soc. 91: 379-383.
Darnell, R.M. 1958. Food habits of fishes and larger invertebrates of Lake Pontchartrain, Louisiana, an estuarine community. Publ. Inst. Mar. Sci. Univ. Tex. 5: 353-416.
Darnell, R.M. 1961. Fishes of the Rio Tamesi and related coastal lagoon in east-central Mexico. Publ. Inst. Mar. Sci. Univ. Tex. 8: 299-365.
Davis, C.C., and R.H. Williams. 1950. Brackish water plankton of mangrove areas in southern Florida. Ecology 31: 519-531.
Davis, J.H., Jr. 1940. The ecology and geologic role of mangroves in Flor- ida. Carnegie Institute, Washington, D.C Publ. 517. Tortugas Lab. Pap. 32: 303-412.
Davis, J.H., Jr. 1942. The ecology of the vegetation and topography of the Sand Keys of Florida. Tortugas Lab. Pap. 3: 113-195.
Davis, J.H., Jr. 1943. The natural fea- tures of south Florida, especially
90
the vegetation and the Everglades. Fla. Geol. Surv. Bull. 25: 1-333.
Davis, J.H., Jr. 1946. Peat deposits of Florida, their occurrence, develop- ment and uses. Fla. Geol. Surv. Bull. 30.
de
la Cruz, A. A. (in press, a) Economic evaluations and ecological implica- tions of alternative uses of mangrove swamps in Southeast Asia. Proceed- ings of the Asia symposium on man- grove environment: research and man-
agement. Univ. Lampur, Malaysia.
of Malaya, Kuala
de
la Cruz, A. A. (in press, b) The impact of crude oil and oil-related activities on coastal wetlands--a review. Proceedings of the interna- tional wetlands conference, Delhi, India.
Dent, J.M. 1947. Some soil problems in
empoldered rice lands in Sierra
Leone. Emp. J. Exp. Agric. 15: 206-212.
Desselle, W.J., M.A. Poirrier, J.S. Rogers r.achnor. 1978. A discrim- analysis of habits and in the
and R.C. Cashner inant functions omi s )
sunfish
feeding
Lake Pont-
(Lepomis) food habits and feeding niche segregation in the Lake Pont- chartrain, Louisiana estuary. Trans. Am. Fish. Soc. 107: 713-710.
de Sylva, D.P., and H.B. Michel. 1974. Effects of mangrove defoliation on the estuarine ecology of South Viet Nam. Pages 710-728 j_n G. Walsh, S. Snedaker and H. Teas, eds. Proceed- ings of the international symposium on the biology and management of mangroves. Univ. of Florida, Gaines- ville.
Dunson, W.A. 1979. Occurrence of par- tially striped forms of the mangrove snake, Nerodia fasciata compress i- cauda Kennicott, and comments on the status of N_. f. taeniata Cope. Fla. Sci. 42: 102-112.
Eaton, S.W. ing in 174.
1953. Wood warblers winter- Cuba. Wilson Bull. 65: 169-
Egler, F.E. 1948. The dispersal and establishment of the red mangrove, Rhizophora, in Florida. Caribb. For. 9: 299-310.
Egler, F.E. 1952. Southeast saline Ever- glades vegetation, Florida, and its management. Vegetatio 3: 213-265.
Emlen, J.T. 1977. Land bird communities of Grand Bahama Island: the struc- ture and dynamics of an avifauna. A.O.U. Ornithol. Monogr. 24.
Enright, J.T. and the 1037.
1974. Mangroves, isopods ecosystem. Science 183:
Ernst, C.H., and R.W. Barbour. 1972. Turtles of the United States. Univ. of Kentucky Press, Lexington. 299 pp.
Estevez, E.D. 1978. Ecology of Sphaeroma terebrans Bate, a wood boring isopod, in a Florida mangrove forest. Ph.D. Dissertation. Univ. of South Flor- ida, Tampa. 168 pp.
Estevez, E.D., and J.L. Simon. 1975. Systematics and ecology of Sphaeroma (Crustacea: Isopoda) in the mangrove habitats of Florida. Pages 286-304 in G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the international symposium on the biology and manage- ment of mangroves. Univ. of Florida, Gainesville.
Evink, G.L. 1975. Macrobenthos compari- sons in mangrove estuaries. Pages 256-285 in G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the international symposium on the bio- logy and management of mangroves. Univ. of Florida, Gainesville.
Federal Register. 33781.
1980. 45(99): 33768-
Fell, J.W. 1975. Phycomycetes associated with degrading mangrove leaves. Can. J. Bot. 53: 2908-2922.
Fell, J.W. 1980. The association and potential role of fungi in mangrove detrital systems. Bot. Mar. 23: 257-263.
91
Fell, J.W., and I.M. Master. 1973. Fungi associated with the degradation of mangrove (R_. mangle) leaves in south Florida. Pages 455-466 in H.L. Stevenson and R.R. Colwell, eds. Estuarine microbial ecology. Univ. of South Carolina Press, Columbia.
Fell, J.W., and S.Y. Newell. 1980. Role of fungi in carbon flow and nitrogen immobilization in coastal marine plant litter systems. J£ D.T. Wick- low and G.C. Carroll, eds. The fungal community: its organization and role in the ecosystem. Marcel Dekker, New York.
Fell, J.W., R.C. Cefalu, I.M. Master, and A.S. Tallman. 1975. Microbial ac- tivities in the mangrove (Rhizophora mangle) leaf detrital system. Pages 661-679 in G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the international symposium on the biol- ogy and management of mangroves. Univ. of Florida, Gainesville.
Fell, J.W., I.M. Master, and S.Y. Newell. 1980. Laboratory model of the poten- tial role of fungi (Phytophthora spp. ) in the decomposition of red mangrove (Rhizophora mangle) leaf litter. JrPTX Tenore and B.C. Coull, eds. Marine benthic dynamics. Univ. of South Carolina Press, Colum- bia.
Ffrench, G.D. 1966. The utilization of mangroves by birds in Trinidad. Ibis 108: 423-424.
Field, G.D. 1968. Utilization of man- groves by birds on the Freetown Peninsula, Sierra Leone. Ibis 110: 354-357.
Fosberg, F.R. 1961. Vegetation-free zones of dry mangrove coasts. U.S. Geo!. Surv. Prof. Pap. 365: 216-218.
Fosberg, F.R. 1971. Mangroves versus tidal waves. Biol. Conserv. 4: 38-39.
Gerlach, S. 1958. Die Mangroveregion tropischer Duesten als Lebensraum. Z. Morphol. Oekol. Tiere 46: 636- 730.
Gilfillan, E.S., R.P. Gefber, and D.S. Page. 1981. Effects of the Zoe Colocotroni oil spill on infaunal communities associated with man- groves. Pages 71-87 j_n R.C. Carey, P.S. Markovits and J.B. Kirkwood, eds. Proceedings U.S. Fish and Wild- life Service workshop on coastal eco- systems of the Southeastern United States. U.S. Fish and Wildlife Ser- vice, Office of Biological Services, Washington, D.C. FWS/0BS-80/59.
Gill, A.M. 1970. The mangrove fringe on the eastern Pacific. Fairchild Trop. Gard. Bull. 25: 7-11.
Gill, A.M., and P.B. Tomlinson. 1969. Studies on the growth of red mangrove (Rhizophora mangle L.). I. Habitat and general morphology. Biotropica 1: 1-9.
Gill, A.M., and P.B. Tomlinson. 1971. Studies on the growth of red mangrove (Rhizophora mangle L.). II. Growth and differentiation of aerial roots. Biotropica 3: 63-77.
Gill, A.M., and P.B. Tomlinson. 1977. Studies on the growth of red mangrove (Rhizophora mangle L.). IV. The adult root system. Biotropica 9: 145-155.
Girard, G.T., and W.K. Taylor. 1979. Re- productive parameters for nine avian species at Moore Creek, Merritt Island National Wildlife Refuge, Florida. Fla. Sci. 42: 94-102.
Goforth, H.W., and J.R. Thomas. 1979. Plantings of red mangroves (Rhizo- phora mangle) for stabilization of marl shorelines in the Florida Keys. Pages 207-230 jm Proceedings 6th annual conference on the restoration and creation of wetlands. Hills- borough Community College, Tampa, Fla.
Golley, F.B. 1972. Tropical ecology with an emphasis on organic productivity. Pages 407-413 j_n F.B. Golley and R. Misra, eds. Tropical ecology. Univ. of Georgia, Athens.
Golley, F.B., H.T. Odum, and R.F. Wilson. 1962. The structure and metabolism
92
of a Puerto Rican red mangrove forest in May. Ecology 43: 1-19.
Golley, F.B., J.T. McGinnis, R.G. Cle- ments, G.I. Child, and M.J. Duever. 1974. Mineral cycling in a tropical moist forest ecosystem. Univ. of Georgia Press, Athens.
Goodbody, I. 1961. Inhibition of the development of a marine sessile com- munity. Nature 190: 282-283.
Goodwin, T.M. 1979. Waterfowl management practices employed in Florida and their effectiveness on native and migratory waterfowl populations. Fla. Sci. 42: 123-129.
Gore, R. 1977. The tree nobody liked. Natl. Geogr. Mag. 151: 669-689.
Gotto, J.W., and B.F. Taylor. 1976. N fixation associated with decaying leaves of the red mangrove, Rhizo- phora mangle. Appl. Environ. Micro- biol. 31: 781-783.
Gotto, J.W., F.R. Tabita, and C.V. Baalen. 1981. Nitrogen fixation in inter- tidal environments of the Texas gulf coast. Estuarine Coastal Shelf Sci. 12: 231-235.
Gunter, G. 1956. A revised list of eury- haline fishes of North and middle America. Am. Midi. Nat. 56: 345-354.
Gunter, G., and G.E. Hall. 1965. A bio- logical investigation of the Caloosa- hatchee estuary of Florida. Gulf Res. Rep. 2: 1-71.
Haines, E. 1976. Stable carbon isotope ratios in the biota, soils and tidal water of a Georgia salt marsh. Estu- arine Coastal Mar. Sci. 4: 609-616.
Hall, E.R. 1981. The mammals of North America. Vol. 1. 2nd ed. John Wiley and Sons, New York. 600 pp.
Hamilton, W.J., Jr., and J.O. Whittaker, Jr. 1979. Mammals of the Eastern United States. 2nd ed. Cornell Univ. Press, Ithaca, N.Y. 345 pp.
Hanlon, R. , F. Bayer, and G. Voss. 1975. Guide to the mangroves, buttonwood
and poisonous shoreline trees of Florida, the Gulf of Mexico and the Caribbean Region. Univ. of Miami Sea Grant Field Guide Series 3. 29 pp.
Hart, M.G.R. 1962. Observations on the source of acid in empoldered mangrove soils. I. Formation of elemental sulphur. Plant Soil 17: 87-98.
Hart, M.G.R. 1963. Observations on the source of acid in empoldered mangrove soils. II. Oxidation of soil poly- sulphides. Plant Soil 19: 106-114.
Haverschmidt, F. 1965. The utilization of mangroves by South American birds. Ibis 107: 540-542.
Heald, E.J. 1969. The production of organic detritus in a south Florida estuary. Ph.D. Dissertation. Univ. of Miami , Fla. 110 pp.
Heald, E.J. 1971. The production of organic detritus in a south Florida estuary. Univ. of Miami Sea Grant Tech. Bull. 6. 110 pp.
Heald, E.J., and W.E. Odum. 1970. The contribution of mangrove swamps to Florida fisheries. Proc. Gulf Caribb. Fish. Inst. 22: 130-135.
Heald, E.J., W.E. Odum, and D.C. Tabb. 1974. Mangroves in the estuarine food chain. Miami Geol. Soc. Mem. 2: 182-189.
Heald, E.J., M.A. Roessler, and G.L. Beards ley. 1979. Litter production in a southwest Florida black mangrove community. Pages 24-33 jj^ Proceed- ings Florida antimosquito association 50th meeting.
Hesse, P.R. 1961. The decomposition of organic material in a mangrove swamp soil. Plant Soil 14: 249-263.
Hesse, P.R. 1962. Phosphorous fixation in mangrove swamp muds. Nature 193: 295-296.
Hicks, D.B., and L.A. Burns. 1975. Man- grove metabolic response to alter- ations of natural freshwater drain- age to southwestern Florida estu- aries. Pages 238-255 jn G. Walsh,
93
S. Snedacker and H. Teas, eds . Pro- ceedings of the international sympo- sium on the biology and management of mangroves. Univ. of Florida, Gaines- ville.
Hobbs, H.H., Jr. 1942. The crayfishes of Florida. Univ. Fla. Biol. Sci. Ser. 3: 1-179.
Hoffman, W.E., and C.J. Dawes. 1980. Photosynthetic rates and primary pro- duction by two Florida benthic red algal species from a salt marsh and a mangrove community. Bull. Mar. Sci. 30: 358-364.
Hofstetter, R.H. 1974. The effect of fire on the pineland and sawgrass communities of southern Florida. Miami Geol. Soc. Mem. 2: 201-209.
Holdridge, L.R. 1940. Some notes on the mangrove swamps of Puerto Rico. Ca- ribb. For. 1: 19-29.
Holm, R.F. fishes Fla. Sci
1977. The standing crop of in a tropical marine lagoon. : 258-261.
Hopper, B.E., J.W. Fell, and R.C. Cefalu. 1973. Effect of temperature on life cycles of nematodes associated with the mangrove (R. mangle) detrital system. Mar. Biol. 23: 293-296.
Howarth, R.W., and J.M. Teal. 1980. Energy flow in a salt marsh ecosys- tem: the role of reduced inorganic sulfur compounds. Am. Nat. 116: 862-872.
Howell, A.H. 1932. Florida bird life.
Florida Department of Game and Fresh
Water Fisheries. Cowan-McCann, Inc., New York. 579 pp.
Hudson, H.J., D.M. Allen, and T.J. Cos- tello. 1970. The flora and fauna of a basin in central Florida Bay. U.S. Fish Wildl. Serv. Spec. Sci. Rep. Fish. 604. Washington, D.C. 14 pp.
Hutto, R.L. 1980. Winter habitat distri- bution of migratory land birds in Western Mexico, with special refer- ence to small foliage-gleaning insec- tivores. Pages 181-203 in A. Keast and E.S. Morton, eds. Migrant birds
in the neotropics: ecology, behavior, distribution, and conservation. Smithsonian Inst. Press, Washington, D.C.
Idyll, C.P. 1965a. Shrimp need freshwa- ter, too. Natl. Parks Mag. 39: 14-15.
Idyll, C.P. 1965b. The freshwater re- quirements of Everglades National Park. Univ. of Miami Institute of Marine Science (mimeo). 3 pp.
Idyll, C.P., D.C. Tabb, and B. Yokel. 1968. The value of estuaries to shrimp. Pages 83-90 vn J.D. Newsom, ed. Marsh and estuary management symposium. Louisiana State Univer- sity Press, Baton Rouge.
Jenik, J. 1967. Root adaptations in west African trees. J. Linn. Soc. Lond. Bot. 60: 126-140.
Kahl, M.P., Jr. 1964. Food ecology of the wood stork (Mycteria americana) in Florida. Ecol. Monogr. 34: 97-117.
Keast, A. 1980. Spatial relationships between migratory parulid warblers and their ecological counterparts in the neotropics. Pages 109-130 jn^ A. Keast and E.S. Morton, eds. Migrant birds in the neotropics: ecology, behavior, distribution, and conserva- tion. Smithsonian Inst. Press, Wash- ington, D.C.
Kilby, J.D. 1955. The fishes of two gulf coastal marsh areas of Florida. Tu- lane Stud. Zool . 2: 176-247.
Kohlmeyer, J. 1969. Ecological notes on fungi in mangrove forests. Trans. Brit. Mycol. Soc. 53: 237-250.
Kohlmeyer, J., and E. Kohlmeyer. 1979. Marine mycology. The higher fungi. Academic Press, New York. 690 pp.
Kolehmainen, S.E. 1973. Ecology of ses- sile and free-living organisms on mangrove roots in Jobos Bay. Pages 141-173 jn Aguirre Power Project environmental studies 1972, annual report. Puerto Rico Nuclear Center.
Kolehmainen, S.E., and W.K. Hildner. 1975. Zonation of organisms in Puerto Rican
94
red mangrove (Rhizophora mangle L.) swamps. Pages 357-369 Irf G-E- Walsh, S.C. Snedaker and H.J. Teas, eds. Proceedings of the international sym- posium on the biology and management of mangroves. Univ. of Florida, Gainesville.
Kolehmainen, S.E., T.O. Morgan, and R. Castro. 1974. Mangrove root commun- ities in a thermally altered area in Guayanilla Bay, Puerto Rico. In J.W. Gibbons and R.R. Scharitz, eds. Ther- mal ecology. U.S. AEC Conf. 730505.
Kuenzler, E.J. 1974. Mangrove swamp sys- tems. Pages 346-371 J_n H.T. Odum, B.J. Copeland and E.A. McMahon, eds. Coastal ecological systems. Vol. 1. Conservation Foundation, Washington, D.C.
Kushlan, J. A. 1976. Site selection for
nesting colonies of white ibis, Eudo-
cimus albus, in Florida. Ibis 118: 590-593.
Kushlan, J. A. getics of 114-122.
1977a. Population ener- the white ibis. Auk 94:
Kushlan, J. A. 1977b. of the white ibis. 342-345.
Foraging behavior Wilson Bull. 89:
Kushlan, J. A. 1979. Feeding ecology and prey selection in the white ibis. Condor 81: 376-389.
Kushlan, J. A. 1980. The status of croco- dilians in south Florida. I. U.C.N. Crocodile Specialists Group. Unpub- lished MS.
Kushlan, J. A., and P.C. Frohring. (in prep.) Colonial waterbird use of the Everglades estuaries. Proceedings conference on estuarine birds. Univ. of South Carolina, Columbia.
Kushlan, J. A. Food of Florida.
, and M.S. Kushlan. 1975.
the white ibis in southern
Fla. Field Nat. 3: 31-38.
Kushlan, J. A., and T.E. Lodge. 1974. Ecological and distributional notes on the freshwater fish of southern Florida. Fla. Sci. 37: 110-128.
Kushlan, J. A., and L.C. McEwan. (in press) Nesting phenology of the double-crested cormorant. Wilson Bull.
Kushlan, J. A., and L.C. McEwan. (in prep.) The brown pelican population in southern Florida.
Kushlan, J. A., and L.C. McEwan. (in prep.) The sandhill crane in the Everglades.
Kushlan, J. A., and W.B. Robertson, Jr. 1977. White ibis nesting in the lower Florida Keys. Fla. Field Nat. 5: 127-131.
Kushlan, J. A., and D.A. White. 1977a. Nesting wading bird populations in southern Florida. Fla. Sci. 40: 65-72.
Kushlan, J. A., and D.A. White. 1977b. Laughing gull colonies in extreme southern Florida. Fla. Field Nat. 5: 123-126.
Kushlan, J. A., J.C. Ogden, and A.L. Higer. 1975. Relation of water level and fish availability to wood stork reproduction in the southern Ever- glades, Florida. Open File Rep. 75-434. U.S. Department of the Interior, Geological Survey. Talla- hassee, Fla. 56 pp.
Kushlan, J. A., O.L. Bass, Jr., and L.C. McEwan. (in prep.) Wintering water- fowl in the Everglades estuaries.
Lack, D., and P. Lack, warblers in Jamaica. 129-153.
1972. Wintering Living Bird 11:
LaHunt, D.E., and G.W. Cornwell. 1970. Habitat preference and survival of Florida duck broods. Proc. Annu. Conf. Southeast. Assoc. Game Fish. Comm. 24: 117-121.
Lawrence, D.B. 1949. Self-erecting habit of seedling red mangroves (Rhizophora mangle L.). Am. J. Bot. 36: 426-427.
Layne, J.N. 1974. south Florida. 2: 386-413.
the land mammals of Miami Geol. Soc. Mem.
95
Lee, C.C. 1969. The decomposition of organic matter in some shallow water, calcareous sediments of Little Black Water Sound, Florida Bay. Disserta- tion. Univ. of Miami, Fla. 106 pp.
Lewis, R.R., III. 197Sa. Oil and man- grove forests: the aftermath of the Howard Starr oil spill (abstract). Fla. Sci. (supplement) 42: 26.
Lewis, R.R., III. 1979b. Large scale mangrove restoration on St. Croix, V.I. Pages 231-242 jm Proceedings 6th conference on restoration and creation of wetlands. Hillsborough Community College, Tampa, Fla.
Lewis, R.R., III. 1980a. Oil and man- grove forests: observed impacts 12 months after the Howard Starr oil spill (abstract). Fla. Sci . (supple- ment) 43: 23.
Lewis, R.R., III. 1980b. Impact of oil spills on mangrove forests. Page 36 in Second international symposia on the biology and management of man- groves and tropical shallow water communities. Port Moresby, Madang, Papau, New Guinea.
Lewis, R.R., III. 1981. Economics and feasibility of mangrove restoration. Pages 88-103 in R.C. Carey, P.S. Markovits and J.B. Kirkwood, eds. Proceedings U.S. Fish and Wildlife Service workshop on coastal ecosys- tems of the Southeastern United States. U.S. Fish and Wildlife Ser- vice, Office of Biological Services, Washington, D.C. FWS/OBS-80/59.
Lewis, R.R., III, and F.M. Dunstan. 1975. Use of spoil islands in reestablished mangrove communities in Tampa Bay, Florida. Pages 766-775 jn G.E. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the international sym- posium on the biology and management of mangroves. Univ. of Florida, Gainesville.
Lewis, R.R., III, and C.S. Lewis. 1978. Colonial bird use and plant succes- sion on dredge material islands in Florida. U.S. Army Corps of Engi- neers Tech. Rep. D-78-14.
Lewis, R.R., III, C.S. Lewis, W.K. Fehring, and J. A. Rodgers. 1979. Coastal habitat mitigation in Tampa Bay, Florida. Pages 136-140 j[n Proceedings mitigation symposium. Gen. Tech. Rep. RM-65. U.S. Depart- ment of Agriculture, Fort Collins, Col.
Lindall, W.N., and C.H. Saloman. 1977. Alteration and destruction of estua- ries affecting fishery resources of the Gulf of Mexico. Mar. Fish. Rev. 39, Pap. 1262. 7 pp.
Little, E.J. 1977. Observations on recruitment of postlarval spiny lobster, Panulirus argus , to the south Florida coast. Fla. Mar. Res. Publ. 29. 35 pp.
Longley, W.L., R. Jackson, and B. Snyder. 1978. Managing oil and gas activi- ties in coastal environments. U.S. Fish and Wildlife Service, Office of Biological Services. FWS/OBS-78/54.
Loope, L.L. 1980. Phenology of flowering and fruiting in plant communities of Everglades National Park and Biscayne National Monument, Florida. Rep. T-593, Everglades National Park, South Florida Research Center. 121 pp.
Mangrove ecosystems: or steady state? Bio- tropica. 12(2): 65-73.
Lugo, A.E. 1980. successional
Lugo, A.E. 1981. Mangrove issue debates in courtrooms. Pages 48-60 in R.C. Carey, P.S. Markovits and J.B. Kirk- wood, eds. Proceedings U.S. Fish and Wildlife Service workshop on coastal ecosystems of the Southeastern United States. U.S. Fish and Wildlife Service, Office of Biological Ser- vices, Washington, D.C. FWS/OBS- 80/59.
Lugo, A.E., and G. Cintron. 1975. The mangrove forests of Puerto Rico and their management. Pages 825-846 jn G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the international symposium on the biology and manage- ment of mangroves. Univ. of Florida, Gainesville.
96
Lugo, A.E., and S.C. Snedaker. 1974. The ecology of mangroves. Annu. Rev. Ecol. Syst. 5: 39-64.
Lugo, A.E., and S.C. Snedaker. 1975. Properties of a mangrove forest in southern Florida. Pages 170-211 jm G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the interna- tional symposium on the biology and management of mangroves. Univ. of Florida, Gainesville.
Lugo, A.E., and C.P. Zucca. 1977. The impact of low temperature stress on mangrove structure and growth. Trop. Ecol. 18: 149-161.
Lugo, A.E., G. Evink, K.M. Brinson, A. Broce, and S.C. Snedaker. 1975. Diurnal rates of photosynthesis, respiration and transpiration in man- grove forests in south Florida. Pages 335-350 jn F. Golley and G. Medina, eds. Tropical ecological systems. Springer-Verlag, New York.
Lugo, A.E., M. Sell, and S.C. Snedaker. 1976. Mangrove ecosystem analysis. Pages 113-145 jji B.C. Patten, ed. Systems analysis and simulation in ecology. Academic Press, New York.
Lugo, A.E., R.R. Twilley, and C. Patter son-Zucca. 1980. The role of blacK mangrove forests in the productivity of coastal ecosystems in south Flor-
1980. The role of black
Jl UVC I U I CJ L.O 111 L n 6?
coastal ecosystems in south Flor- ida. Report to E.P.A. Corvallis Environmental Research Laboratory, Corvallis, Oregon. 281 pp.
MacArthur, R.H., and J.W. MacArthur. 1961. On bird species diversity. Ecology 42: 594-598.
Macko, S. 1981. Stable nitrogen isotopes as tracers of organic geochemical processes. Ph.D. Dissertation. Univ. of Texas, Austin.
Macnae, W. 1968. A general account of the fauna and flora of mangrove swamps and forests in the Indo-West- Pacific region. Adv. Mar. Biol. 6: 73-270.
Mahmoud, I.Y. 1968. Feeding behavior of kinosternoid turtles. Herpetologia 24: 300-305.
Marathe, K.V. 1965. A study of the sub- terranean algae flora of some man- grove swamps. J. Indian Soc. Soil Sci. 13: 81-84.
Martin, A.C., H.S. Zin, and A.L. Nelson.
1951. American wildlife and plants.
A guide to wildlife food habits.
Dover Publ., Inc., New York. 500 pp.
Mattox, N.T. 1949. Studies on the biol- ogy of the edible oyster, 0s trea rhizophorae Guilding, in Puerto Rico. Ecol. Monogr. 19: 339-356.
Maxwell, G.R., III, and H.W. Kale, II. 1977. Breeding biology of five spe- cies of herons in coastal Florida. Auk 94: 689-700.
McCoy, E.D., and K.L. Heck, Jr. 1976. Biogeography of corals, seagrasses and mangroves: an alternative to the center of origin concept. Syst. Zool. 25: 201-210.
McGill, J.T. 1959. Coastal classifica- tion maps. Pages 1-22 ±n R.J. Rus- sell, ed. Second coastal geography conference. Coastal Studies Inst., Louisiana State Univ., Baton Rouge.
McMillan, C. 1971. Environmental factors affecting seedling establishment of the black mangrove on the central Texas coast. Ecology 52: 927-930.
McMillan, C. 1975. Interaction of soil texture with salinity tolerance of black mangrove (Avicennia) and white mangrove (Laguncularia) from North America. Pages 561-566 in G. Walsh, S. Snedaker and H. Teas, eds. Pro- ceedings of the international sympo- sium on the biology and management of mangroves. Univ. of Florida, Gaines- ville.
McNulty, J.K., W.N. Lindall, and J.E. Sykes. 1972. Cooperative Gulf of Mexico estuarine inventory and study, Florida. Phase I, area description. N0AA Tech. Rep. NMFS Circ-368. 127 pp.
Miller, P.C. 1972. Bioclimate, leaf tem- perature, and primary production in red mangrove canopies in south Flor- ida. Ecology 53: 22-45.
97
Miller, P.C. 1974. The potential use of vegetation to enhance water cooling in holding ponds. Pages 610-627 j_n J.W. Gibbons and R.R. Scharitz, eds. Thermal ecology. U.S. AEC Conf. 730505.
Miller, P.C. 1975. Simulation of water relations and net photosynthesis in mangroves in southern Florida. Pages 615-631 jm G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the international symposium on the bio- logy and management of mangroves. Univ. of Florida, Gainesville.
Moldenke, H.N. 1967. Additional notes on the genus Avicennia. Phytologia 14: 301-336.
Moore, J.C. 1953. The crocodile in the Everglades National Park. Copeia 1953: 54-59.
Moorman, F.R., and L.J. Pons. 1975. Characteristics of mangrove soils in relation to their agricultural land use and potential. Pages 529-547 j[n_ G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the international symposium on the biology and manage- ment of mangroves. Univ. of Florida, Gainesville.
Morton, E.S. 1980. Adaptations to sea- sonal changes by migrant land birds in the Panama Canal Zone. Pages 437- 453 j£ A. Keast and E.S. Morton, eds. Migrant birds in the neotropics: ecology, behavior, distribution, and conservation. Smithsonian Inst. Press, Washington, D.C.
Morton, J.F. 1962. Wild plants for sur- vival in south Florida. Southeastern Printing Co., Stuart, Fla. 80pp.
Morton, J.F. 1964. south Florida. Hortic. Soc. 77:
Honeybee plants of Proc. Fla. State 415-436.
Morton, J F. 1965. Can the red mangrove provide food, feed and shelter? Econ. Bot. 19: 113-123.
Newell, S.Y. 1974. The succession in the mycoflora of red mangrove (Rhizophora mangle) seedlings. Ph.D. Disserta- tion. Univ. of Miami, Fla.
Nisbet, I.C.T. 1968. The utilization of mangroves by Malayan birds. Ibis 110: 348-352.
Noakes, D.S.P. 1955. Methods of increas- ing growth and obtaining natural regeneration of the mangrove type in Malaya. Malays. For. 18: 23-30.
Nugent, R.S. 1970. The effects of ther- mal effluent on some of the macro- fauna of a subtropical estuary. Ph.D. Dissertation. Univ. of Miami, Fla. 198 pp.
Odum, E.P. 1971. Fundamentals of ecology. Saunders Publ. Co., Philadelphia, Pa. 574 pp.
Odum, H.T., and C.F. Jordan. 1970. Metabolism and evapotranspiration of the lower forest in a giant plastic cylinder. Pages 1-165 - 1-189 in H.T. Odum and R.F. Pidgeon, eds. A tropical rain forest. Div. of Tech. Info. U.S.A.E.C, Oak Ridge, Tenn.
Odum, H.T., M. Sell, M. Brown, J. Zuc- chetto, C. Swallows, J. Browder, T. Ahlstom, and L. Peterson. 1974. The effects of herbicides in South Viet- nam: models of herbicide, mangroves, and war in Vietnam. U.S. National Academy of Sciences - National Research Council, Washington, D.C.
Odum, W.E. 1970. Pathways of energy flow in a south Florida estuary. Ph.D. Dissertation. Univ. of Miami, Fla. 162 pp.
Odum, W.E. 1971. Pathways of energy flow in a south Florida estuary. Univ. of Miami Sea Grant Bull. 7. 162 pp.
Odum, W.E. 1974. Potential effects of aquaculture on inshore coastal waters. Environ. Conserv. 1: 225- 230.
Odum, W.E. 1976. Ecological guidelines for tropical coastal development. IUCN Publ. 42. IUCN, Morges, Swit- zerland. 60 pp.
Odum, W.E., and E.J. Heald. 1972. Trophic analyses of an estuarine mangrove community. Bull. Mar. Sci. 22: 671- 738.
98
Odum, W.E., and E.J. Heald. 1975a. Man- grove forests and aquatic productiv- ity. Chapter 5 jn An introduction to land-water interactions. Springer- Verlag Ecological Study Series, New York.
Odum, W.E., and E.J. Heald. 1975b. The detritus-based food web of an estu- arine mangrove community. Pages 265- 286 in Estuarine research. Academic Press, New York.
Odum, W.E., and R.E. Johannes. 1975. The response of mangroves to man-induced environmental stress. Pages 52-62 in E.J.F. Wood and R.E. Johannes, eds. Tropical marine pollution. Elsevier (Oceanography series), Amsterdam, Netherlands.
Odum, W.E., J.S. Fisher, and J. Pickral. 1979a. Factors controlling the flux of particulate organic carbon from estuarine wetlands. Pages 69-80 in R.J. Livingston, ed. Ecological processes in coastal and marine systems. Plenum Press, No. 10 in the Ecological Study Series.
Odum, W.E., P.W. Kirk, and J. Zieman. 1979b. Non-protein nitrogen com- pounds associated with particles of vascular plant detritus. Oikos 32: 363-367.
Ogden, J.C. 1969. Checklist of birds of Everglades National Park. Everglades Natural History Association. 30 pp.
Ogden, J.C. 1978. Status and nesting biology of the American crocodile, Crocodylus acutus, in Florida. J. Herpetol. 12: 183-196.
Ogden, J.C, J. A. Kushlan, and J.T. Til- mont. 1976. Prey selectivity by the wood stork. Condor 78: 324-330.
Ogden, J.C, J. A. Kushlan, and J.T. Til- mont. 1978. The food habits and nesting success of wood storks in Everglades National Park, 1974. U.S. Dep. Inter. Natl. Park Serv. Natl. Res. Rep. 16. 25 pp.
Olexa, M.T., and T.E. Freeman. 1975. Oc- currence of three unrecorded diseases
on mangrove in Florida. Pages 688- 692 in G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the inter- national symposium on the biology and management of mangroves. Univ. of Florida, Gainesville.
Olmstead, I.C, L.L. Loope, and R.P. Russell. 1981. Vegetation of the southern coastal region of the Ever- glades National Park between Flamingo and Joe Bay. Rep. T-620, Everglades National Park, South Florida Research Center. 18 pp.
Olsen, D.A., and I.G Koblic. 1975. Popu- lation dynamics, ecology and behavior of spiny lobsters, Panulirus argus, of St. John, V.I. II. Growth and mortality. Nat. Hist. Mus. Los Ang. Cty. Sci. Bull. No. 20: 17-21.
Olsen, D.A., A.E. Dammann, J.F. Hess, J.R. Sylvester, and J. A. Untema. 1973. The ecology of fishes in two mangrove lagoons in the U.S. Virgin Islands. Rep. Puerto Rico Int. Underseas Lab. 42 pp. (unpublished).
Olsen, D.A., W.F. Herrnkind, and R.A. Cooper. 1975. Population dynamics, ecology and behavior of spiny lob- ster, Panulirus argus, of St. John, V.I. Nat. Hist. Mus. Los Ang. Cty. Sci. Bull. No. 20: 11-16.
Onuf, C.P., J.M. Teal, and I. Valiela. 1977. Interactions of nutrients, plant growth and herbivory in a man- grove ecosystem. Ecology 58: 514- 526.
Orians, G.H. 1969. The number of bird species in some tropical forests. Ecology 50: 783-801.
Orians, G.H., and E.W. Pfeiffer. 1970. Ecological effects of the war in Vietnam. Science 168: 544-554.
Patterson-Zucca, C 1978. The effects of road construction on a mangrove eco- system. M.S. Thesis. Univ. of Puerto Rico at Rio Piedras. 77 pp.
Payne, W.J. 1970. Energy yields and growth of heterotrophs. Annu. Rev. Microbiol. 24: 17-51.
99
Peterson, C.H., and N.M. Peterson. 1979. The ecology of intertidal flats of North Carolina: a community profile. U.S. Fish and Wildlife Service, Office of Biological Services. FWS/ OBS-79/39. 73 pp.
Pool, D.J., A.E. Lugo, and S.C. Snedaker. 1975. Litter production in mangrove forests of southern Florida and Puerto Rico. Pages 213-237 in G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the international sym- posium on the biology and management of mangroves. Univ. of Florida, Gainesville.
Pool, D.J., S.C. Snedaker, and A.E. Lugo. 1977. Structure of mangrove forests in Florida, Puerto Rico, Mexico and Central America. Biotropica 9: 195- 212.
Post, E. 1963. Systematische und pflan- zen-Geographische Notizen zur Bos- trichia-Caloglossa Assoziation. Rev. Algol. 91 UW.
Preston, F.W. 1979. The invisible birds. Ecology 60: 451-454.
Provost, M.W. 1959. Impounding salt marshes for mosquito control and its effects on bird life. Fla. Nat. 32: 163-170.
Provost, M.W. 1969. Ecological control of salt marsh mosquitos with side benefits to birds. Proceedings Tall Timbers conference on ecological animal control. Tallahassee, Fla.
Provost, M.W. 1974. Mean high water mark and use of tidelands in Florida. Fla. Sci. 36: 50-66.
Pulver, T.R. 1976. Transplant techniques for sapling mangrove trees, Rhizo- phora mangle, Avicennia germinans and Laguncularia racemosa. Fla. Mar. Res. Publ. 22.
Rabinowitz, D. 1975. Planting experi- ments in mangrove swamps of Panama. Pages 385-393 vn G. Walsh, S. Sne- daker and H. Teas, eds. Proceedings of the international symposium on the biology and management of mangroves. Univ. of Florida, Gainesville.
Rabinowitz, D. 1978a. Dispersal proper- ties of mangrove propagules. Bio- tropica 10: 47-57.
Rabinowitz, D. 1978b. Early growth of mangrove seedlings in Panama, and an hypothesis concerning the relation- ship of dispersal and zonation. J. Biogeog. 5: 113-133.
Rabinowitz, D. 1978c. Mortality and ini- tial propagule size in mangrove seed- lings in Panama. J. Ecol. 66: 45-51.
Ramos, M.A. , and D.W. Warner. 1980. Analysis of North American subspecies of migrant birds wintering in Los Tustlas, southern Veracruz, Mexico. Pages 173-180 vn A. Keast and E.S. Morton, eds. Migrant birds in the neotropics: ecology, behavior, distribution, and conservation. Smithsonian Inst. Press, Washington, D.C.
Randall, J.E. 1967. Food habits of reef fishes of the West Indies. Pages 665-847 jm Studies in tropical ocean- ography. Inst, of Mar. Sci. Univ. of Miami, Fla.
Ray, J. P. (in press) Fate and effects of petroleum hydrocarbons on mangrove ecosystems.
Reark, J.B. 1975. A history of the colonization of mangroves on a tract of land on Biscayne Bay, Fla. Pages 776-804 jn G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the international symposium on the bio- logy and management of mangroves. Univ. of Florida, Gainesville.
Recher, H.F. 1966. Some aspects of the ecology of migrant shorebirds. Eco- logy 47: 393-407.
Reeve, M.R. 1964. Studies on the seasonal variation of the zooplankton in a marine subtropical in-shore environ- ment. Bull. Mar. Sci. 14: 103-122.
Rehm, A.E. 1974. A study of the marine algae epiphytic on the prop roots of Rhizophora mangle L. from Tampa to Key Largo, Florida. Dissertation. Univ. of South Florida, Tampa. 183 pp.
100
Rehm, A.E. 1976. The effects of the wood-boring isopod, Sphaeroma tere- brans, on the mangrove communities of Florida. Environ. Conserv. 3: 47-57.
Rehm, A.E., and H.J. Humm. 1973. Sphae- roma terebrans: a threat to the man- groves of southeastern Florida. Science 182: 173-174.
Reid, G.K., Jr. 1954. An ecological study of the Gulf of Mexico fishes in the vicinity of Cedar Key, Florida. Bull. Mar. Sci. 4: 73-94.
Rivas, L.R. 1962. The Florida fishes of the genus Centropomus, commonly known as snook. Q. J. Fla. Acad. Sci. 25: 53-64.
Rivas, L.R. 1969. A revision of the poeciliid fishes of the Gambusia punctata species group, with descrip- tions of two new species. Copeia 4: 778-795.
Robertson, W.B., Jr. 1955. An analysis of the breeding-bird populations of tropical Florida in relation to vege- tation. Ph.D. Dissertation. Univ. of Illinois, Urbana.
Robertson, W.B., Jr., and J. A. Kushlan. 1974. The southern Florida avifauna. Miami Geol. Soc. Mem. 2: 414-452.
Russell, S.M. 1980. Distribution and abundance of North American migrants in the lowlands of northern Columbia. Pages 249-252 in A. Keast and E.S. Morton, eds. Migrant birds in the neotropics: ecology, behavior, dis- tribution, and conservation. Smith- sonian Inst. Press, Washington, D.C.
Rutzler, K. 1969. The mangrove community, aspects of its structure, faunistics and ecology. Pages 515-536 jm Lagunas Costeras, un simposio. UNAM-UNESCO. Mexico, D.F.
Rutzler, K. 1970. Oil pollution: damage observed in tropical communities along the Atlantic seaboard of Panama. Bioscience 20: 222-224.
Saenger, P., and C.C. Mclvor. 1975. Water quality and fish populations in a
mangrove estuary modified by residen- tial canal developments. Pages 753- 765 in G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the inter- national symposium on the biology and management of mangroves. Univ. of Florida, Gainesville.
Sasekumar, A. macrofauna shore. J. Anim
1974. Distribution of
on a Malayan mangrove
Ecol. 43: 51-69.
Savage, T. 1972. Florida mangroves as shoreline stabilizers. Fla. Dep. Nat. Resour. Prof. Pap. 19. 46 pp.
Schmidt, T.W. 1979. Seasonal biomass estimates of marine and estuarine fishes within the western Florida Bay portion of Everglades National Park, May 1973 to July 1974. Pages 665-672 in R.M. Linn, ed. Proceedings 1st conference on scientific resources in the national parks. U.S. Dep. Inter., Natl. Park Serv. Trans. Proc. Ser. 5.
Schneider, D. 1978. Equalization of prey numbers by migratory shorebirds. Nature 271: 353-354.
Scholander, P.F. 1964. Hydrostatic pres- sure and osmotic potential in leaves of mangrove and some other plants. Proc. Natl. Acad. Sci. U.S.A. 52: 119-125.
Scholander, P.F. 1968. How mangroves desalinate seawater. Physiol. Plant. 21: 258-268.
Scholander, P.F., L. van Dam, and S.I.
Scholander. 1955. Gas exchange in
the roots of mangroves. Am. J. Bot. 42: 92-98.
Scholander, P.F., H.T. Hammel, E. Hemming- sen, and W. Cary. 1962. Salt balance in mangroves. Plant Physiol. 37: 722-729.
Scholander, P.F., H.T. Hammel, E.D. Brad- street, and E.A. Hemmingsen. 1965. Sap pressure in vascular plants. Science 148: 339-346.
Scholander, P.F., E.D. Bradstreet, H.T. Hammel, and E.A. Hemmingsen. 1966. Sap concentrations in halophytes and
101
some other plants. 41: 529-532.
Plant Physiol,
Scholl, D.W. 1965. High interstitial water chlorinity in estuarine man- grove swamps, Florida. Nature 207: 284-285.
Schwartz, A. 1949. A second specimen of Mustela vison evergladensis Hami 1 ton . J. Mammal. 30: 315-316.
Schwartz, P. 1964. The northern water- thrush in Venezuela. Living Bird 3: 169-184.
Scoffin, T.P. 1970. The trapping and binding of subtidal carbonate sedi- ment by marine vegetation in Bimini Lagoon, Bahamas. J. Sediment Petrol. 40: 249-273.
Seaman, W. , C.A. Adams, and S.C. Snedaker. 1973. Biomass determinations in shallow estuaries; technique evalua- tion and preliminary data. Pages J1-J23 In S.C. Snedaker and A.E. Lugo, eds. The role of mangrove ecosystems in the maintenance of environmental quality and a high productivity of desirable fish- eries. Final Rep. 14-16-008-606 to the Bureau of Sport Fisheries and Wildlife.
Shemnitz, S.D. 1974. Populations of bear, panther, alligator and deer in the Florida Everglades. Fla. Sci. 37: 157-167.
Shines, J.E. 1979. Distribution of mangrove communities: State of Florida. U.S.E.P.A. Final Contract 68-03-2636.
Simberloff, D. 1976. Experimental zoo- geography of islands: effects of island size. Ecology 57: 629-648.
Simberloff, D., and E.0. Wilson. 1969. Experimental zoogeography of islands: the colonization of empty islands. Ecology 50: 278-296.
Simberloff, D. , B.J. Brown, and S. Lowrie. 1978. Isopod and insect root borers may benefit Florida mangroves. Science 201: 630-632.
Snedaker, S.C. 1974. Mangroves, isopods and the ecosystem. Science 183: 1036-1037.
Snedaker, S.C. 1978. Mangroves: their value and perpetuation. Nat. Resour. 14: 6-13. UNESCO, Paris.
Snedaker, S.C. 1981. Mangrove misconcep- tions and regulatory guidelines. Pages 61-67 in R.C. Carey, P.S. Mark- ovits and J.B. Kirkwood, eds. Pro- ceedings U.S. Fish and Wildlife Ser- vice workshop on coastal ecosystems of the Southeastern United States. U.S. Fish and Wildlife Service, Office of Biological Services, Wash- ington, D.C. FWS/OBS-80/59.
Snedaker, S.C, and A.E. Lugo. 1973. The role of mangrove ecosystems in the maintenance of environmental quality and a high productivity of desirable fisheries. Final Rep. on Contract 14-16-008-606 to the Bureau of Sport Fisheries and Wildlife. 381 pp.
Snyder, N.F.R. 1974. Breeding biology of swallow- tailed kites in Florida. Living Bird 13: 73-97.
Southwell, C.R., and J.D. Boltman. 1971. Marine borer resistances of untreated woods over long periods of immersion in tropical waters. Biotropica 3: 81-107.
Spackman, W. , D.W. Scholl, and W.H. Taft. 1964. Field guide book to environ- ments of coal formation in southern Florida. Geol. Soc. Am. 67 pp.
Spackman, W., C.P. Dolsen, and W. Riegal. 1966. Phytogenic organic sediments and sedimentary environments in the Everglades-mangrove complex. I. Evi- dence of a transgressing sea and its effect on environments of the Shark River area of southwest Florida. Paleotropica 117: 135-152.
Springer, V.G., and K.D. Woodburn. 1960. An ecological study of the fishes of the Tampa Bay area. Fla. Board Con- serv. Prof. Pap. Ser. 1. 104 pp.
Stark, W.A., II, and R.E. Schroeder. 1971. Investigations on the gray
102
snapper, Lutjanus qriseus. Univ. of Miami Press, Miami, Fla. 224pp.
Sutherland, J. P. 1980. Dynamics of the epibenthic community on roots of the mangrove, Rhizophora mangle, at Bahia de Buche, Venezuela. Mar. Biol. 58: 75-84.
Tabb, D.C. 1963. A summary of existing information on the freshwater and marine ecology of the Florida Ever- glades region in relation to fresh- water needs of Everglades National Park. Report to the Superintendent of Everglades National Park from the University of Miami Marine Lab. 153 pp.
Tabb, D.C. 1966. The estuary as a habi- tat for spotted seatrout, Cynoscion nebulosus. Am. Fish. Soc. Spec. Publ. 3: 59-67.
Tabb, D.C, and E.J. Heald. 1973. The coastal interceptor waterway. Univ. Miami Sea Grant Coastal Zone Manage. Bull. 4. 10 pp.
Tabb, D.C, and A.C Jones. 1962. Effect
of Hurricane Donna on the aquatic
fauna of North Florida Bay. Trans. Am. Fish. Soc. 91: 375-378.
Tabb, D.C, and R.B. Manning. 1961. A checklist of the flora and fauna of northern Florida Bay and adjacent brackish waters of the Florida main- land collected during the period July 1957 through September 1960. Bull. Mar. Sci. 11: 552-649.
Tabb, D.C, and B.J. Yokel. 1968. Report to the Conservation Foundation: pre- liminary ecological study of Rookery Bay Sanctuary, Naples, Florida. Univ. of Miami, Inst. Mar. Sci. 26 pp.
Tabb, D.C, D.L. Dubrow, and R.B. Manning. 1962. The ecology of northern Flor- ida Bay and adjacent estuaries. Fla. Board Conserv. Tech. Ser. 39. 79 pp.
Taylor, D.L. 1980. Fire history and man- induced fire problems in subtropical south Florida. Proc. fire history workshop. Tech. Rep. RM-81, Rocky
Mt. Forest and Range Exp. Stn., U.S.D.A.: 63-68.
Taylor, D.L. 1981. Fire history and fire records for Everglades National Park 1948-1979. Report T-619, Everglades National Park, South Florida Research Center. 121 pp.
Taylor, W.R. 1960. Marine algae of the eastern tropical and subtropical coasts of the Americas. Univ. of Michigan Press, Ann Arbor. 879 pp.
Teas, H. 1977. Ecology and restoration of mangrove shorelines in Florida. Environ. Conserv. 4: 51-57.
Teas, H. 1979. Silviculture with saline water. Pages 117-161 JL A. Hollaen- der, ed. The biosaline concept. Plenum Publ. Corp.
Teas, H., and J. Kelly. 1975. Effects of herbicides on mangroves of S. Vietnam and Florida. Pages 719-728 j_n G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the international sym- posium on the biology and management of mangroves. Univ. of Florida, Gainesville.
Teixeira, C , J. Tundisi, and M.B. Kutner. 1965. Plankton studies in a mangrove environment. II. The standing stock and some ecological factors. Biol. Inst. Ocean. Sao Paulo, Publ. 14: 13-41.
Teixeira, C, J. Tundisi, and J.S. Ycaza. 1967. Plankton studies in a mangrove environment. IV. Size fractionation of the phytoplankton. Biol. Inst. Ocean. Sao Paulo, Publ. 16: 39-42.
Teixeira, C, J. Tundisi, and J.S. Ycaza. 1969. Plankton studies in a mangrove environment. VI. Primary production, zooplankton standing-stock and some environmental factors. Int. Rev. Gesamten. Hydrobiol. 54: 289-301.
Terborgh, J.W., and J.R. Faaborg. 1980. Factors affecting the distribution and abundance of North American migrants in the eastern Caribbean region. Pages 145-155 j_n A. Keast and E.S. Morton, eds. Migrant birds
103
ecology, behav- and conservation.
in the neotropics:
ior, distribution,
Smithsonian Inst. Press, Washington,
D.C.
Thayer, G.W., H.H. Stuart, W.J. Kenworthy, J.F. Ustach, and A.B. Hall. 1978. Habitat value of salt marshes, man- groves and seagrasses for aquatic organisms. Pages 235-247 jm Wetland functions and values: the state of our understanding. Am. Water Resour. Assoc, Minneapolis, Minn.
Thorn, B.G. 1967. Mangrove ecology and deltaic geomorphology: Tabasco, Mexico. J. Ecol. 55: 301-343.
Thorn, B.G. 1975. Mangrove ecology from a geomorphic viewpoint. Pages 469-481 J_n G. Walsh, S. Snedaker and H. Teas, eds. Proceedings of the interna- tional symposium on the biology and management of mangroves. Univ. of Florida, Gainesville.
Tomlinson, T.E. 1957. Relationship between mangrove vegetation, soil texture and reaction of surface soil after empoldering saline swamps in Sierra Leone. Trop. Agric. 34: 41-50.
Tomlinson, P.B. 1980. The biology of trees native to tropical Florida. Harvard Univ. Press, Cambridge, Mass.
Tschirley, F.H. 1969. Vietnam. Science
Defoliation in 163: 779-786.
Tundisi, J. 1969. Plankton studies in a mangrove environment -- its biology and primary production. Pages 485- 494 2D. Lagunas Costeras, un simposio. UNAM-UNESCO. Mexico, D.F.
Twilley, R. 1980. Organic exports from black mangrove forests in south Florida. Ph.D. Dissertation. Univ. of Florida, Gainesville.
Voss, G.L., F.M. Bayer, C.R. Robins, M. Gomon, and E.T. LaRoe. 1969. A report to the National Park Service, Department of the Interior, on the marine ecology of the Biscayne National Monument. Inst, of Marine and Atmospheric Sciences, Univ. of Miami, Fla. 128 pp.
Wade, R.A. 1962. The biology of the tarpon, Megalops atlanticus, and the ox-eye, Megalops cyprinoides, with emphasis on larval development. Bull. Mar. Sci. 13: 545-622.
Waisel, Y. 1972. Biology of halophytes. Academic Press, New York. 395 pp.
Walsh, G.E. 1967. An ecological study of a Hawaiian mangrove swamp. Pages 420-431 jn G.H. Lauff, ed. Estuaries. American Association Advancement Sci- ence Publ. 83. Washington, D.C.
Walsh, G.E. 1974. Mangroves: a review. Pages 51-174 jn. R- Reimhold and W. Queen, eds. Ecology of halophy- tes. Academic Press, New York.
Walsh, G.E., R. Barrett, G.H. Cook, and T.A. Hoi lister. 1973. Effects of herbicides on seedlings of the red mangrove, Rhizophora mangle L. Bio- science 23: 361-364.
Walsh, G.E., K.A. Ainsworth, and R. Rigby. 1979. Resistance of red mangrove (Rhizophora mangle L. ) seedlings to lead, cadmium, and mercury. Biotrop- ica 11: 22-27.
Wang, J.C., and E.C. Raney. 1971. Distri- bution and fluctuations in the fish fauna of the Charlotte Harbor estu- ary, Florida. Mote Marine Labora- tory, Sarasota, Fla. 56 pp.
Wanless, H. 1974. Mangrove sedimentation in geological perspective. Miami Geol. Soc. Mem. 2: 190-200.
van der Pijl, L. 1972. Principles of dispersal in higher plants. Springer- Verlag, New York.
Voss, G. 1969. estuaries.
Preserving Florida's Fla. Nat. 19: 2-4.
Warner, G.F. 1967. The the mangrove tree pisoni . J. Zool.
life history of
crab, Aratus
153: 321-335.
Warner, G.F. 1969. The occurrence and distribution of crabs in a Jamaican
104
mangrove swamp. 379-389.
J. Anim. Ecol. 38:
Weinstein, M.P. , CM. Courtney, and J.C. Kinch. 1977. The Marco Island estu- ary: a summary of physicochemical and biological parameters. Fla. Sci. 40: 97-124.
Westing, A.H. 1971. Forestry and the war in South Vietnam. J. For. 69: 777- 784.
Witham, R.R., R.M. Ingle, and E.A. Joyce, Jr. 1968. Physiological and ecolog- ical studies of P_. argus from St. Lucie estuary. Fla. Board Conserv. Tech. Ser. 53. 31 pp.
Wolff, W.J. 1969. Distribution of non- breeding waders in an estuarine area in relation to the distribution of their food organisms. Ardea 57: 1-28.
Wolffenden, G.E., and R.W. Schreiber. 1973. The common birds of the saline habitats of the eastern Gulf of Mex- ico: their distribution, seasonal status, and feeding ecology. Pages 1-22 j_n J. Jones, R.E. Ring, M.O. Rinkel and R.E. Smith, eds. A sum- mary of knowledge of the eastern Gulf of Mexico. Inst, of Oceanography, St. Petersburg, Fla. 604 pp.
Wood, E.J.F. 1965. Marine microbial ecology. Reinhold Publ. Corp., New York. 243 pp.
Yokel, B.J. 1975a. Rookery Bay land use studies. Environmental plan- ning strategies for the development of a mangrove shoreline. Study 5,
Estuarine biology. The Conservation Foundation, Washington, D.C. 112 pp.
Yokel, B.J. 1975b. A comparison of ani- mal abundance and distribution in similar habitats in Rookery Bay, Marco Island and Fakahatchee on the southwest coast of Florida. Prelim. Rep. to Deltona Corp. Rosenstiel School of Marine and Atmospheric Science, Univ. of Miami. 162 pp.
Yoshioka, P.M. 1975. Mangrove root com- munities in Jobos Bay. Pages 50-65 in Final report on Aguirre environ- mental studies. Puerto Rico Nuclear Center.
Zieman, J.C, Jr. 1972. Origin of circu- lar beds of Thalassia testudinum in south Biscayne Bay, Florida, and their relationship to mangrove ham- mocks. Bull. Mar. Sci. 22: 559-574.
Zieman, J.C, Jr. (in prep.) A community profile: the ecology of the seagrass ecosystem of south Florida. U.S. Fish and Wildlife Service, Office of Biological Services, Washington, D.C.
Zuberer, D.A., and W.S. Silver. 1975. Mangrove-associated nitrogen fixa- tion. Pages 643-653 2D. S. Walsh, S. Snedaker and H. Teas, eds. Proceed- ings of the international symposium on the biology and management of man- groves. Univ. of Florida, Gaines- ville.
Zuberer, D.A., and W.S. Silver. 1978. Biological nitrogen fixation (acety- lene reduction) associated with Florida mangroves. Appl. Environ. Microbiol. 35: 567-575.
105
APPENDIX A. Summary of the site characteristics and sampling
methodology for fishes in: A-l - mangrove- fringed tidal streams and rivers, A-2 - mangrove-lined estuarine bays and lagoons, and A-3 - mangrove- lined oceanic bays and lagoons .
106
V)
0>
|
s- |
|
|
o 4- |
|
|
>> |
a |
|
Ol |
CO |
|
o |
s- |
|
t— |
01 |
|
o |
> |
|
■o |
•r— |
|
o |
t- |
|
JZ |
|
|
+J-o |
|
|
0) |
C |
|
B |
*> |
|
O) 10 |
|
|
c |
E |
|
•I— |
<0 |
|
^- |
0) |
|
Q. S. |
|
|
E |
■»-> |
|
TJ |
CO |
|
co |
|
|
r— |
|
|
TJ |
ia |
|
c -o |
|
|
fO |
•1— |
|
+j |
|
|
M |
|
|
O X |
|
|
•r— |
o< |
|
•!-> |
Ol |
|
CO |
c |
|
•r- |
•1— |
|
J_ |
i- |
|
at «»- |
|
|
■!-> |
1 |
|
o |
cv |
|
<T3 |
> |
|
S_ |
o |
|
ITS |
s_ |
|
.C |
Ol |
|
u |
c |
|
(0 |
|
|
a) |
E |
I
5
U IA 01 0) (D-O .O -r- S_ E O O 3 0) U
z a. ai
|
>* |
O.SZ |
|
u |
CU C7> |
|
c |
I/) 3iO |
|
cu |
O <X> |
|
3 |
- i- a> |
|
cr |
>,-C .-* |
|
cu |
I— +J *~* |
|
L. |
x: »-0 |
|
Ll_ |
+-> LT) +J X) |
|
C l£> Q. TO |
|
|
OCJl Q)H |
|
|
s: i-h t/> — - |
c jc r- a>
TO +-> TO CT> CU Q.-0 C
|
+■> cu |
ro |
|
TO C7> |
CO |
|
i- c |
i |
|
<u m |
o |
|
Q. i- |
**- |
|
i |
|
|
s |
in |
i-h nko
|
Q.J= |
||
|
OJ |
Cn |
|
|
L/l |
=5 |
|
|
O CO |
||
|
L. |
in |
|
|
>,-C cd |
||
|
w |
||
|
£ |
||
|
4-J |
r-- |
|
|
c |
lT> |
u |
|
o |
en |
QJ |
|
c |
£ |
|
|
■a |
3 |
|
|
O CM |
||
|
S- r-. |
||
|
>,X: C7> |
||
|
4-> |
. — i |
|
|
£ |
||
|
-t-> CM |
||
|
C |
r-» |
o |
|
o en |
a> |
|
|
3E |
• — i |
a |
|
x: to |
+-> |
r~ |
|
i— -M 3 X: |
o |
u_ |
|
.— c o> en |
1— |
|
|
i- O 3 3 |
Ul |
|
|
en E <C o en |
</) |
|
|
■.- S- <X) |
||
|
*xi • *> x: en |
c |
|
|
>> (/> *-> »— 1 |
||
|
.— •* S- |
||
|
J* L0 CU CO ■ |
c |
|
|
ai ■*-> jz kr> u |
0) |
|
|
CU 0) ■*-> CT» Q> |
-X. |
|
|
3 C O -H O |
TO |
|
+-> |
2 |
||
|
5 +JT3 a) |
o |
||
|
O * CU o c |
c |
||
|
S- i/l C S- |
c |
||
|
JC ■•-> i— |
|||
|
■m cu -o * 0) |
E |
||
|
c c c £ |
|||
|
* 3 O E |
■ M |
||
|
CU Q. O t/> TO |
V) |
<u |
|
|
c i- a.*!- t- |
CU |
c |
|
|
T- T3 O +■* |
c |
||
|
a> »a |
CU |
||
|
(/)»(/) • |
(/) |
CU |
|
|
i/i a.x: r— |
c |
||
|
CT> +-» TO (/> CU |
*J |
CT» |
(— |
|
TO CU J- •<- CU |
QJ |
A3 |
CD |
|
CO C -M M- S- |
C/J |
CO |
l/l |
>, (U i— +-> CU CU o .—
c s_ en a. ■^ CO 4-> oS -.- t_
3 W U
o ■•->
'- QJ i- CJ) O)
c o c ■■ *
c: o
V)
TO
•O E
C U CU C -r-
•■- fo c m +-> cu i — o +■> i/>
i^jj C in d)
TO O
CU TO t- E CU "f-
■o at T3
a
S-
o
E CU o (/> s- I- CU
' TO
X .yi EP
CU CU _
c c +->
>> O <0 3
f— -(-> JD U
<U c/l S-
CT CU "O CU
1- E C "D
<o 1- <o c
_l .— </> 3
E E rt in
x CLin cti 03 cu * •
ZTJHO
C ID •* TO C W +-> C7> CU TO l/>
S- .— +J> CU
O ■ •- +■> +■>
•i- W> S- -^
^1 CT> CU </)
U I Q. >
*r- "O TO •— ■*->
x: 3 s- 3 a; i— E ■•-> a c
E E i-i ir> ^-1 O
o
CO
|
>> o |
•i- S- |
TO >> |
|||
|
O TO <X) |
cu |
S- CU |
CU 03 |
||
|
s- |
3 TO ■*-* CO CJ^ |
CU |
cu c 4-> ro |
S- >> O -M |
|
|
cu • |
O O ©0 i-h |
x: |
-»-> o i- r-* |
+-> CU cn C |
|
|
> vo o |
>> O >> TO |
o |
c ••- to en |
M ._£ *r- 0) |
|
|
•rlfiN |
TO CO TO O- 1- C |
•M |
(U CUH |
S- CO CD |
|
|
q; en oi |
00* — CO E CU S- |
TO |
ZD |
TD 3 3 |
|
|
1— 1 i-H |
TO CD 3 |
x: |
E |
E — • |
CU h- "ZT |
|
JC |
Wl i — TO h- C XI |
TO |
TO |
TO TO i— i— |
E +> |
|
4-> XI e |
(/I TO C7> ■•— "O |
-^ |
QJ |
QJ Jit' TO TO |
TO i~ c ••> o |
|
S- X) 3 |
o c cu -o S- O |
-C |
E. |
_. -C C |
C TO »r- >>r--. |
|
O TO TJ |
S- TO •>- ^ Q. O |
TO |
+J |
•*-> TO TO •*-> |
C CU O TO CT> |
|
ZHO |
UUUOW3 |
Li. |
Ul |
CO Li_ CJ CU |
13 c a. co ^< |
107
i- o
>> •
0> 00
o c
i— o
o o
0 (O
j= i— +j
01 -o
|
ro |
|
|
en |
|
|
c |
00 |
|
•r— |
> |
|
r— |
(D |
|
a.£t |
|
|
E |
|
|
10 |
0) |
|
oo |
C |
|
•i — |
|
|
■a |
S- |
|
c |
fD |
|
TO |
^ |
|
+-> |
|
|
oo |
00 |
|
u |
ai |
|
•r— |
|
|
+-> |
-o |
|
00 |
0) |
|
•1 — |
c |
|
s_ |
•1 — |
|
aj |
r— |
|
■t-> |
1 |
|
o |
<u |
|
n3 |
> |
|
S- |
o |
|
ro |
i- |
|
-C |
Dl |
|
u |
c |
|
r0 |
|
|
aj |
E |
|
+-> |
|
|
•i — |
c |
|
CO |
•^ |
<\J I
u o
ai u
a. o>
en
r-* a* ai -* -* CO
|
X C -C |
a |
c XI |
|
<d en |
||
|
3 -3 3 |
u |
*-3 3 |
|
O CM O CM |
cd |
O CM |
|
i_ r^. •* * t. r^ |
• i~ r-. |
|
|
X: CI—- >, JT cn |
$- |
>»J= Cn |
|
4-) .-t f— .— ■!-> .-1 |
01 |
i— -t-J i-H |
|
ai -c |
■t-> |
JC |
|
r-H >>-* ■*-> CM ■ |
s- |
■<-> CM • |
|
r-» i— o c r*. o |
n3 |
CN U |
|
C71 3> OOl ai |
O |
O Cn CU |
|
^ho^^h -<cj |
*•— ■* |
s: — i a |
|
O 3 |
•~D 3 CM -D 3 C |
CO |
|
O CM |
O r^- O 1ft *r- |
1 \£) C~i |
|
•> i_ r-. |
* i_ cn •- -J-r-cu |
m cn <x> |
|
>>.c cn |
>>x: .-h.— ■ >>x: cn ■*->-—*• |
>, c .-h x: en |
|
f— w «-« |
.— -M r— ,— +J ,-H tf> • |
•— o cn^-t |
|
j= |
x: » cu x: c r— |
-C -i- -3 |
|
■•-> o >> |
■*-> r-t >^J* +J .-■ • -i- Id |
P+Jp o • |
|
c r-. .— |
cr^<— ocrs-ccu |
C id CL J- > |
|
o en 3 |
OCT) 3 >- OCn ITJ3-P |
O -*-> CU XI O |
|
z: ^h -3 |
s: •-* t— ' Z •-» *"3 — - ai |
z wyi+JZ |
1_ CM 0)
» +-» CD
i (J —
rO Oj
CU "4- -•-'
*r- >>J* W L I.
TJ ' — -O O CU 3 id
cu cu >- c wu
■*-» x: -a-—- -r- * --^
is i/i a> cu * *
4->--*,i — I — Oil — i — i—
ai-a a j 3 2 m cn c: e id cn id id
uiaifli-iautP
>U1 in -4-> XI +-> -t-> ai
|
E |
|
|
3 |
|
|
>*- * C |
|
|
O 1- |
|
|
T3 |
|
|
W *-J |
3 |
|
id -C |
*-J |
|
cu en |
LO |
|
i- -^ |
<U |
|
T3 1- |
■M |
|
S |
|
|
cu |
a |
|
> cu |
|
|
^ |
|
|
VI 3 |
en |
|
c -o |
id |
|
0) o |
|
|
+J 1— |
id |
|
X (O |
-C |
|
uj m |
1- |
cni. uj *a CU **-
j- c en
cn-i- r—
HIT3 rtJ
ai c
in nj c
-*-> cu
QJ i/l CD
r— t.
P£ CT ■M cn
|
O 1- |
E C O J- ••- |
|
|
- -♦-> ai |
- -4-3 cu a> |
|
|
■a ■*-» x: |
■O -M ■<-> --3 |
|
|
3 O 4-> |
3 O +■»— - .•^-'^-* |
|
|
E -O O |
E X) 0<— |
|
|
*.— >> |
-^- >,J£ 2 *a |
|
|
-o^- X) |
"D >— XJ O ra |
|
|
ai cu |
CU CU >- S- •*-> |
|
|
■u x: -o |
■*-> x: -a — - -*-* <u |
|
|
US VI CU |
id m cu |
|
|
+-* ^ •— |
■!-> --s..— t— i- c |
|
|
ait) as |
cu -o CL 3 cu -r- |
|
|
en c e |
<d |
cn c E *o ■*-» cu |
|
CD id m |
L. |
CU «3 ro S- -4-> +-> |
|
> v» in |
■*-> |
> m in +J o in |
|
. |
.^ |
|
|
3 |
c |
|
|
- C |
c |
|
|
T3 |
||
|
T3 |
E |
|
|
-t-J |
=3 |
|
|
x: |
4J |
CD |
|
cn |
LO |
cn |
|
a> |
C |
|
|
U |
■M |
CD |
|
J |
||
|
id |
<T3 |
|
|
CD |
||
|
tn |
||
|
3 |
m |
x: |
|
-a |
id |
CL |
|
o |
O |
|
|
"3 |
||
|
D |
x: |
■D |
|
a: |
h- |
31 |
<o 5 E •* X -t-J
r— 4- CU O
|
L/> |
\S> |
E |
id |
|
>. O |
|||
|
c |
rd |
O |
on |
|
-Q |
L^ |
||
|
1 |
CD |
id |
|
|
-X |
> |
||
|
T3 |
U |
O |
n |
|
CU |
<D |
s. |
x: |
|
a |
is |
cnl— |
|
CD |
, — |
|
|
E |
CD |
|
|
>. O |
x: |
|
|
IA |
(/) |
|
|
id |
°c |
|
|
t. |
>i |
|
|
CD |
XJ |
TJ |
|
c |
■Q |
C |
3 u ta i. id S o in o Xl
E
in oi ir>
|
-*-■ a> |
01 |
||
|
f— x: |
x: |
||
|
•t- Wl |
t/) |
||
|
in |
|||
|
o0 |
oO |
||
|
4-» ^~ |
-o |
||
|
<d i- |
c |
||
|
cu id |
D |
||
|
o- E |
(/» |
||
|
i |
CL |
, |
E |
|
m |
CD |
in |
|
|
3 c |
CD |
c o |
|
|
o o |
X) |
o |
|
|
rH E |
|||
|
f— +J |
*-> |
I |
|
|
ta (o |
£i |
D |
CO >x> |
|
x: -P |
4-> |
||
|
</> in |
•— • |
moo |
|
"^-. |
i— CM |
|
P0 |
0) ro |
|
1 |
jh: i |
|
D |
CX) |
|
■— 1 |
>- LO |
I O CU
cu o cu
cn **- xj -
*d r«* o i- i—
i- cn o OlrH
cu i 5 u ^ r^.
> m o cu o cn
oi id xj n x: x: in
u r*. 4-»
■*-> o cn cu
id «3- rH
x: r- l. »d •— o* ^: * <u ■*-> m x: >,^£ i_ r-.
id it) o id cn
U- CO >- c_) ^-<
O ■*-> 0J 1- -r-
O (J 4-> »*- ft)
n+J1"- ai «j i-
OU"0 > Jm-o m
<u id -i- x: 4-> »—
in ■!-> i_ in m 3X1 ««
ro xj o i/* cu a.c_>
i 3 Q. io "
in in in e '
Q. i- id "O cu cu o o
m x: s- -r- o
■m cu s- -^
o o > cu o
o t-t cu t. m
cz er «J
■ ■•- c3 u
x: >> i- r-. <d id id cn
u_ aa cj *-*
|
cn |
w .- 3 *-> |
|
>, .-l-H |
k- >> 0J |
|
5- Id |
i~ •• |
|
cux^ |
O id X} c |
|
j* 0) |
u 3 in -r- r*. |
|
o cn ^£ |
t. +Jt^ aj k |
|
Ot-tO |
id m cn +j cn |
|
cc <a- >- |
ZUH WM |
|
id |
|
|
5 O |
|
|
cu |
|
|
+J |
-O |
|
>>r- |
|
|
x: |
id cn |
|
3CQH |
cu -3
108
<L>
o
>1
cn • o i/>
i— c
o o -o o o o>
-C rd 4-> i—
0)
E "O
c
CO <TJ
c
•t- Ul
I— >>
Q. 03 E XI
to
</> CJ ■r—
-o c
C <0
03 QJ
o
10 O
o
i- -o
i/i c
•r— *r— 4- >— QJ I
+-> <D O > (O O i. $- rO CD
-C C O ITJ E 01
■!-> C
n i
•a:
x>
r0
|
s_ |
00 CU |
|
QJ |
OJ "D |
|
-Q |
•i- S- |
|
E |
U O |
|
3 |
d) {_> |
|
■zl |
Q. QJ |
3
o
|
CL |
en |
|
|
<=c |
3 |
|
|
oco |
||
|
s- |
vo |
|
|
>>-c: |
cn |
|
|
4-> |
i— « |
|
|
jt |
||
|
*J |
in |
|
|
C |
-X) |
c |
|
O |
cn |
IT) |
|
?: |
•— i |
'•a |
oo » s_ r»*
>> sz cn
|
>,j=: +■> |
|
|
fl Olfl |
1/1 |
|
s: 3 |
c |
|
O «=T |
o |
|
* s- r--. |
|
|
>,J= CTi |
-*-> |
|
»—■*-> i— i |
iq |
|
x: |
4-> |
|
+-> CO CU |
V) |
|
c r-. c |
|
|
O Cn 3 |
CM |
|
S ^H -TJ |
. — i |
|
•> +J CD «r- |
||
|
*+J go E i— |
||
|
■o |
'. iU .11 'Tl |
|
|
- c |
QJ C U I- oO |
|
|
4-> |
•— -C 4- CL 00 * 1 ^ |
|
|
C _* |
E 3 OJ S- O |
|
|
3 O |
ro Q. C OJ O |
|
|
O O |
ui •>- <— .c: |
|
|
U -C |
- OJ r— |
|
|
o cu i- on |
||
|
fO i/) |
1- C -C »— |
|
|
3 Q. |
QJ |
4-> -o u • 5 |
|
00 03 |
d |
u oi rd +J fo |
|
r- S- |
3 «— QJ QJ S- |
|
|
> +J |
VI i/)J3 C+J |
|
to |
l/l |
|
ro |
(/ |
|
L. |
T3 |
|
CD |
|
|
T3 |
IC |
|
OJ |
</i |
.^. |
|
4-> fc. -r- |
||
|
X <o +-» |
||
|
OJ (O |
CLJZ |
|
|
00 O" |
||
|
00 |
||
|
• • 00 |
" i. |
|
|
U0 A3 |
E |
3 |
|
0J r— |
3 |
|
|
UO 03 |
C |
OJ |
|
00 J= |
||
|
fO h- |
■a |
3 |
|
5- 2 |
■o |
|
|
cn qj ■*-> |
o |
|
|
A3 > 00 |
||
|
cu •.- qj |
n3 |
|
|
CO 00 |
+j |
31 |
QJ ns
00 -r-
oo oo
A3 oo
|
oO O |
^_ |
|||
|
^~ |
3 |
|||
|
CT)r- |
ra |
|||
|
fO fO |
s- |
|||
|
CO .o |
■*-> |
|||
|
<T5 |
||||
|
JC |
||||
|
u |
u |
|||
|
c |
.C (D |
|||
|
■M |
cu |
CD QJ |
+J |
|
|
JC |
S- |
3 |
aj ro |
|
|
cn |
3 |
O oO |
> +j |
|
|
ro |
J= QJ |
|||
|
L. |
_l |
■l~> +J |
+JJ3-0 |
|
|
5 |
I |
(O |
(O «3 QJ |
|
|
OJ |
C +-> |
+-> J= t— |
||
|
QJ |
<o |
QJ 00 |
C O. |
|
|
p — |
CD TJ |
> |
Q)OE |
|
|
3 |
•F- S_ |
00 -r- fO |
||
|
■o |
fO C |
cn o |
QJ -C 00 |
|
|
o |
QJ |
JC |
S- +-> |
|
|
■o cn |
-M +J |
Q. C 00 |
||
|
id |
QJ -r- |
O 3 |
0J QJ r0 |
|
|
IC |
s. |
o |
S (T3 |
S- ^l 2 |
|
QJ QJ 00 |
•o |
JZ |
(^ |
|
4-> ^ +J |
c |
00 |
4-> |
|
fO 00 C |
fO |
c |
|
|
c a> |
OO |
aj oo |
aj |
|
O oO E |
00 |
E |
|
|
-Q cn |
S- TD |
cn |
|
|
S- -p « |
■o |
*D C |
03 |
|
fO 3 V- |
3 |
o <o |
i- |
E o s-
<4-
|
C -C i— |
QJ |
.«. |
1 |
LD |
|
|
(O -IJ fO |
cn |
E |
|||
|
OJ CL-O |
c |
E |
CU |
-c E |
OJ <NJ |
|
zoit- |
(d |
cn |
• -M 1 |
cn i |
|
|
-a -m |
t_ |
■£> LD |
c |
X CL<-H 1 |
c o |
|
It) |
03 QJ • 1 |
iT3 • |
|||
|
O O |
s- |
S-OCM |
CC r-^ |
|
o |
o |
|
u- |
CD CO |
|
C CM |
|
|
OJ |
3 |
|
enrs i |
|
|
<o |
1 |
|
S- |
* n |
|
aj |
i_ r^. |
|
> |
Q.cn |
|
< «i -* |
o
00
i- 0J o OJ c
> 3N
<c r^ cn
|
aj on |
CO +-> |
■D |
CO i-( |
|
|
"O O f-v |
<U OJ |
c: |
QJ C C |
|
|
o cna> |
00 (T3 |
s_ |
C A3 CU |
C n3CM +J |
|
j: <o<h |
•rTJC |
QJ |
>1 r- |
S--DET3 |
|
or _i |
O «i- O |
x: |
fl3 S- 00 |
OJ -i- ^^ -r- |
|
E |
as. wo |
+j |
U OJ oO t— i |
4J S- E |
|
TD >>^ |
1- OT3N |
3 |
oo -a cu r^ |
w OiO£ |
|
r- ai o |
Or- 3Cft |
O |
■1- <T5 O CTt |
QJr-H U |
|
o ^ X |
n. u_ x i-h |
CO |
CO CO CtL r— 1 |
3 Lu CO CO |
109
APPENDIX B. Fishes of mangrove areas of Florida tabulated by habitat type. Key to numbered references appears at the end of the table. Diet items listed in order of decreasing importance.
110
Family and Species
Habitat Type
B
h a
*J >i at >,
Reference
Diet
Diet Reference
Comments
Orectolobidae - carpet sharks
Ginglymostoma cirratum
nurse shark
5, 7
Fish, cephalopods, molluscs, shrimp, sea urchins
Randall 1967 Clark & von Schmidt 1965 Bohlke & Chaplin 1968
Carcharhinidae - requiem sharks
Carcharhinus leucas
bull shark
Carcharhinus limbatus
blacktip shark
Negaprion brevirostris lemon shark
11
4, 5, 7
Juveniles: fish (Arius felis, Lophoqobius , Mugil cephalus , Brevoortia patronus, Micropogon undulatus) , crus- taceans including penaeid shrimp, blue crabs
Fish (Caranx sp., Centropomus undecimalis, Chilomycterus schoepfi, Arius felis, Lacto- phrys trigonnus Lagodon rhomboides) , crabs
Young: crustaceans, fish Adults: fish, crustaceans
Odum 1971
Clark & von Schmidt 1965
Randall 1967 Clark & von Schmidt 1965
Sphyrnidae - hammerhead sharks
Sphyrna tiburo -
bonnethead
+ + 2, 5
Mantis shrimp, shrimp, isopods, Bohlke s barnacles, bivalve molluscs, Chaplin 1968 cephalopods, fish
Pristidae - sawfishes Pristis pectinata smalltooth sawfish
+ 5, 15
Fish, benthic crustaceans
Bohlke s Chaplin 1968
Rhinobatidae - guitarfishes
Rhinobatos lenti-
guitarfish
Torpedinidae - electric rays
Narcine brasiliensis •
+ 1
+ + 1, 17, 18
lesser electric ray
Rajidae - skates Raja texana roundel skate
Crustacea, fish, annelids
Reid 1954
Dasyatidae - stingrays Dasyatis americana - southern stingray
Dasyatis sarjina Atlantic stingray
+ + 2, 4, 5, 7
2, 8, 13,
17
Fishes, sipunculid and poly- chaete worms , crabs , bivalves shrimp , mantis shrimp
Benthic invertebrates inclu- ding bivalves , xanthid and portunid crabs , shrimps , amphipods, annelids, chirono- mid larvae
Randall 1967
Darnell 1958
Gymnura micrura - smooth butterfly ray
Urolophus jamaicensis ■ yellow stingray
17 Fish, molluscs, annelids, shrimp, other small crustaceans
1 Probably small burrowing invertebrates
Peterson s Peterson 1979
Bohlke s Chaplin 1968
This and all subsequent Odum 1971 citations refer
to W.E. Odum 1971. Ill
Family and Species
Habitat Type
*-> > <y >.
Reference
Diet Reference
Comments
Myliobatidae - eagle rays Aetobatus narinari - Spotted eagle ray
Lepisosteidae - gars
Lepisosteus platyrhincus - + Florida gar
2, 1,
15
13,
Clams, oysters
Boh Ike & Chaplin 1968
Fish (poeciliids, cyprinodonts , Odum 1971 small centrarchids) , crustaceans (caridean shrimp), insect larvae
Elopidae - tarpons El ops saurus - ladyfish
2, 3, 7,
8, 13, 15
< 45 mm: zooplankton, chaeto-
gnaths, polychaete
worms > 45 mm: caridean & penaeid
shrimp, various small
fish
Odum 1971 Austin & Austin 1971
Mega lops atlantica - tarpon
, 8, 13, < 45 mm: plankton (cyclopoid 15 copepods)
juveniles: fish (Gambusia, Fundulus heteroclitus , Mugil cephalus) , crustaceans (ostra- cods, caridean shrimp) adults: wide variety of fish, crabs , shrimp , ctenophores , insects
Odum 1971 Austin & Austin 1971
Obligate air breathe rs . Juv- eniles inhabit shallow brackish pools low in oxygen, often containing H,S (Wade 1962)
Albulidae - bone fishes Albula vulpes - bonefish
Anguillidae - eels Anguilla rostrata - American eel
Ophichthidae - snake
eels
Myrophis punctatus speckled worm eel
4, 5
8, 13
2, 3, 17, 18
Clams, snails, shrimp, small fish
50-200 mm: amphipods, isopods 180-472 mm: xanthid crabs,, caridean shrimp, fish ( Lophogobi us cyprinoides)
Polychaetes, Branch iostoma caribaeum, sand crabs
Bohlke &
Chaplin
1968
Odum 1971
Springer & Woodburn 1960, Reid 1954
Members of this family burrow in mud or sand, undersampled by most methods (Bohlke & Chaplin 1968)
Bascanichthys scuti- caris - whip eels
Ophichthus gome si - shrimp eel
Clupeidae - herrings Brevoortia smithi - yellowfin sardine
3, 17
2, 5, 17
Brevoortia patronus Gulf menhaden
38-48 mm: phy toplankton , zoo- Darnell
plankton, plant fragments , 1958
detritus
85-103 mm: organic matter, silt,
diatoms, foraminiferans, copepods
112
Family and Species
Habitat Type
a
Reference
Diet
Diet
Reference
Harengula pensacolae scaled sardine
2, 3, 8, 30 mm: planktonic copepods, 13 zoea, nauplii, larval fish 64-96 mm: amphipods, harpactlcoid copepods, isopods mysids, chironomid larvae
Odum 1971
Opisthonema oglinum - Atlantic thread herring
2, 3, 5, 13, 17
Copepods, polychaetes, shrimp, fishes, crab larvae, mysids
Odum 1971
Sardinella anchovia Spanish sardine
17
Engraulidae - anchovies Anchoa cub ana - Cuban anchovy
2, 16 Ostracods, copepods
Springer &
Woodburn
1960
Anchoa hepsetus striped anchovy
2, 3, 13, 32-114 mm: copepods, isopods, Springer & 16, 17 mysids, caridean shrimp, small Woodburn bivalves 1960
Anchoa lampro taenia bigeye anchovy
Anchoa mitchilli - bay anchovy
1, 2, 3, <25 mm: microzooplankton Odum 1971
5, 7, 8, 31-62 mm: amphipods, zooplank- 13, 16-18 ton, mysids, ostracods, plant
detritus, copepods, small molluscs,
chironomid larvae
Synodontidae -
lizardfishes
Synodus foetens - inshore lizardfish
1-3, 5, 8, Small fish, crabs, shrimp, 17, 18 polychaete worms
Odum 1971
Catostomidae - suckers Erimyzson sucetta " lake chubsucker
A freshwater stray
Ictaluridae - freshwater catfish
Ictalurus natal is -
yellow bullhead
14
A freshwater stray
Noturus gyrinus tadpole mad torn
14
A freshwater stray
Arriidae - sea catfishes
Arius felis - sea catfish +
2, 3, 5, 100 mm: copepods, zooplankton 7, 8, 13, amphipods, mysids, chironomid 17 larvae, isopods, small crabs 100-200 mm: benthic inverte- brates
200-330 mm: crabs, amphipods, mysids, fishes, bark, crayfish, caridean and penaeid shrimp
Odum 1971
Bagre marinus - gafftopsail catfish
2, 8, 17
262-445 mm: blue crabs, small fishes
Odum 1971
Batrachoididae - toadfishes Opsanus beta - Gulf toadfish
1-3, 5, 18-60 mm: amphipods, chironomid 7, 12, 13, larvae, mysids, isopods, few fish 15, 17, 18 >60 mm: caridean shrimp, xanthid
crabs, snapping shrimp, mussels,
fish, mangrove bark
Odum 1971
Salinities 10 o/oo > (Odum 1971)
113
Family and Species
at Type
Htn ma o <o
Reference
Diet Reference
Comments
Porlchthys porosi6simus Atlantic midshipman
3, 18
Gobiesocidae - clingfishes Gobiesox strumosus skilletfish
2, 3, 5, 8 10-32 mm: amphipods, isopods, chironomid larvae
Odum 1971
Ogcocephalidae - batfishes Ogcocephalus nasutus shortnose batfish
Ogcocephalus radiatus polka-dot batfish
18
Small bivalves, gastropods, polychaetes
2, 11, 17, 18
Reid 1954
Gadidae - codfishes Urophycis floridanus Southern hake
Ophidiidae - cusk-eels, brotulas
Gunterichthys longipenis
gold brotula
Ogilbia cayorum key brotula
Ophidion holbrooki bank cusk-eel
12
17
1, 3
3
Amphipods, isopods, mysids, Springer & A species more
decapod shrimp, polychaetes, Woodburn common at more
insect larvae, fishes (Lagodon 1960 northerly rhomboides, Paralichthys latitudes
alblgutta)
Exocoetidae - flying- fishes , halfbeaks
Chriodorus atherinoides
hardhead halfbeak
Hyporhamphus unifasciatus halfbeak
3, 5 juveniles: zooplankton including crab megalops, veligers, cope- pods
130-199 mm: epiphytic algae, detritus, seagrass
Carr £ Adams
1973
Belonidae - needlefishes Strongylura marina Atlantic needlefish
Strongylura notata redfin needlefish
7, 15
2, 3, 5, 8, 13
357-475 mm: small fishes, Darnell insects, shrimp, small amounts 1958 of vascular plant material and algae
In grassbeds - Brook
Juveniles: polychaete worms, 1975 cumaceans , fish Adults: fish, primarily atherinids
Strongylura timucu timucu
Tylosurus crocodilus ■ houndf ish
2, 3, 11 159-378 mm: anchovies, shrimp
11 250-1320 mm: fishes, shrimp
|
Randall |
Primarily inshore |
|
1967 |
species, freely |
|
enters fresh- |
|
|
water (Randall |
|
|
1967) |
|
|
Randall |
Open water and |
|
1967 |
inshore surface |
|
water inhabitant |
|
|
{Voss et al . |
|
|
1969) |
114
Habitat Type
Family and Species
Reference
Diet
Diet Reference
Comments
Cyprinodontidae - killi-
fishes
Adinia xenica - diamond killifish
Cyprinodon variegatus sheepshead minnow
Floridichthys carpio goldspotted killifish
Fundulus confluentus marsh killifish
Fundulus chrysotus golden topminnow
Fundulus grandis Gulf killifish
Fundulus heteroclitus Mummichog
Fundulus seminolis Seminole killifish
Jordanella floridae flagfish
Lucania goodei bluefin killifish
Lucania parva rainwater killifish
Rivulus marmoratus rivulus
2,8, 13-15 Plant detritus, diatoms, amphipods , harpacticoid copepods, insects
2, 7, 8, 13-15
2, 3, 8, 13
Plant detritus, algae, nematodes, small crustaceans
Odum 1971
Odum 1971
Amphipods, ostracods, isopods, Odum 1971 copepods, chironomid larvae, nematodes, plant detritus, algae
2, 8, 13-15 Caridean shrimp, small fish,
(Gambusia aff inis) , amphipods , isopods, adult & larval insects, copepods, mysids, ostracods, algal filaments
14, 15
Odum 1971
Rare in mangrove zone, headwater pools only
Odum 1971
2,8, 13-15 Amphipods, isopods, xanthid crabs, chironomid larvae, terrestrial insects, snails, algae, small fish (poeciliids)
7 Small crustaceans (amphipods, Peterson & isopods, ostracods, tanaids, Peterson copepods) , detritus, polychaete 1979 worms, insects, snails, inver- tebrate eggs
14-15
13-15
2, 8, 13-15 Small crustaceans (copepods,
cladocerans, ostracods), insect larvae
1-3, 5, 8, 13-15, 17
3, 8, 13, 15
<20 mm: planktonic copepods 21-37 nun: amphipods, mysids, chironomid larvae, ostracods, molluscs, plant detritus
Odum 1971
Odum 1971
Primarily a freshwater form, headwater pools only
Primarily fresh- water , common in pools in headwater regions
Headwater pools and channel
Poeciliidae - livebearers Gambusia affinis mosquitofish
Gambusia rhizophorae mangrove gambusia
2, 3, 7, 13-15
6, 9
A versatile feeder: amphipods, chironomid larvae, hydracarina, harpacticoid copepods, snails, ants, adult insects, polychaete worms, ostracods, mosquito pupae algae
Odum 1971
Fresh and brackish water in Rhizophora swamps , northern Cuba, southeastern Florida
115
Family and Species
Habitat Type
Reference
Diet Reference
Heterandria forroosa least killifish
Poecilia latipinna sailfin molly
Atherinidae - silversides Allanetta harringtonensis reef silver side
Membras martinica rough silverside
Menidia beryl ling tidewater silverside
8, 14, 15 Chironomid larvae, harpacticoid Odum 1971 and planktonic copepods, clado- cerans, terrestrial insects, algae, diatoms
5, 7, 8, Plant detritus, algae, diatoms Odum 1971 13-15
39-60 mm: copepods, fish larvae, Randall polychaete larvae 1967
2, 5, 11
2, 3, 8,
11, 12,
13, 17,
18
Small zooplankton crustaceans, Peterson &
juvenile & larval fishes, Peterson
insects, detritus, snails 1979
Insects, copepods, chironomid Odum 1971 larvae , mysids , amphipods
Syngnathidae - pipefishes , seahorses
Corythoichthys
albirostris
whitenose pipefish
Hippocampus erectus lined seahorse
Hippocampus zoster ae dwarf seahorse
Micrognathus crinigerus fringed pipefish
Syngnathus f loridae dusky pipefish
Syngnathus louisianae chain pipefish
1, 11
|
, 2, 3. |
11. |
|
17 |
|
|
. 2, 3, |
5, |
|
1, 16, |
17 |
|
18 |
1, 5, 10 52-82 mm: copepods, micro- crustaceans
1-3, 5, 11 Caridean shrimp, amphipods, tanaids, isopods
1-3, 11 , Copepods, amphipods, small 16-18 shrimp
Reid 1954
Associated with vegetated areas (Tabb & Manning 1961)
Intimately associated with unattached algae (Tabb & Manning 1961) , or grassy areas (Springer & woodburn 1960)
Brook 1975
Reid 1954
Inhabit grassy flats (Springer & Woodburn 1960)
Syngnathus scovelli Gulf pipefish
Syngnathus springe ri bull pipefish
Syngnathus dunckeri Pugnose pipefish
1-3, 5, 11, Amphipods, isopods, tanaids, 16-18 copepods, tiny caridean
shrimp, gastropods (Bittium, Mitrella)
2, 17
11
Brook 197 5 Springer & Woodburn 1960 Reid 1954
Associated with vegetated areas (Tabb S> Manning 1961)
Syngnathus pelagicus sargassum pipefish
Centropomidae - snooks C e n t r opojmus parallelus fat snook
Family as a whole shows preference for estuarine man- grove habitat (Rivas 1962)
Centropomus pectinatus tarpon snook
7, 8, 13
116
Habitat Type
Family and Species
u « Reference
Diet
Diet Reference
Centropomus undecimalis + snook
2, 5, 7 , 8, Juveniles: caridean shrimp, Odum 1971
11, 13, 14 small cyprinodont fishes, Austin &
gobies, mojarras Austin
Adults: fish, crabs, penaeid 1971
shrimp, crayfish, snapping
shrimp
By far most abundant of three species (Rivas 1962)
Serranidae - sea basses Centropristis striata black seabass
Diplectrum formosum sand perch
Epinephelus itajara jewfish
Epinephelus morio red grouper
2, 3, 11,
16-18
2, 5, 7, 8, 11, 13, 15
Family in general carnivorous on fish, crustaceans
Caridean & penaeid shrimp, copepods, crabs, fish
Juveniles: penaeid shrimp, xanthid crabs
228-340 mm: crustaceans, crabs, fishes
|
Randall |
|
|
1967 |
|
|
Re id |
1954 |
|
Odum |
1971 |
The most abundant
of the seabasses
in mangrove habitats
Randall 1967
Epinephelus striatus Nassau grouper
11 170-686 mm: fish, crabs, stomatopods , cephalopods , shrimp, spiny lobsters, gastropods, bivalves, isopods
Randall 1967
Hypoplectrus puella barred hamlet
11 54-98 mm: snapping shrimp, Randall crabs, fish, mysids, stomato- 1967 pods , isopods
Mycteroperca microlepis gag
Centrarchidae - sunfishes Elassoma evergladei Everglades pygmy sunfish +
1, 2, 5, 11, 71-100 mm: 17, 18 fish
penaeid shrimp.
Reid 1954
Family is primarily freshwater , fish occasionally enter headwater area of mangrove- fringed stream
Lepomis auritus redbreast sunfish
Lepomis gulosus warmouth
Shrimp (Palaemonetes), fish ( Gob io soma bosci , Lepomis macrochirus) , detritus, Vallisneria, amphipods , xan- thid crabs, blue crabs
Desselle et al. 1978
Diet from Lake Pontchartrain salinities 1.6- 4 . 1 o/oo
Lepomis macrochirus bluegill
2, 15 Amphipods, blue crab (Cal- linectes sapidus) , xanthid crabs, detritus, Vallisneria, clams (Rangia cuneata) , sponge (Ephydatia fluviatilis) , barnacles, insect larvae
Desselle et al. 1978
Diet from Lake Pontchartrain salinities 1.6- 4.1 o/oo
Lepomis microlophus redear sunfish
2, 13-15 Chironomid larvae, amphipods, xanthid crabs, clam (Rangia cuneata) , sponge (Ephydatia fluviatilis) , detritus
Desselle et al. 1978
Diet from Lake Pontchartrain salinities 1.6- 4 . 1 o/oo
Lepomis punctatus spotted sunfish
8, 14, 15 Cladocerans, small crabs, odum 1971 mysids , chironomids , amphipods , insects, molluscs, isopods, fish, algae
Salinities < 15 o/oo (Odum 1971)
117
Family and Species
Habitat Type
■32
3
3 4 to
o « Reference
Diet
Diet Reference
Mlcropterus salmoldes largemouth bass
Apogonldae - cardlnalflshes Astrapogon alutus bronze cardinalf ish
13-15
1. 3
Caridean shrimp, small blue crabs, crayfish, xanthld crabs, 25 species of fish, Vallisnerla, Cladophora
Darnell 1958
Astrapogon stellatus conchfish
Pomatomldae - blueflshes Pomatomus saltatrlx bluefish
11 Young: mainly fishes (anchovies, sllversldes, killifishes, men- haden, shad, spotted seatrout), shrimp, crabs, other small crustaceans, annelids, snails
Peterson & Peterson 1979
Rachycentrldae - cobias Rachycentron canadum cobia
+ 5, 7, 11 Fish, crabs
Randall 1967
Echeneldae - remoras Echenels neucratoides vhltefln sharksucker
2, 11 Fish, isopods, other Crustacea Randall 1967
Members of this family attach to sharks and large bony fishes (Randall 1967)
Remora remora
58-175 mm: copepods, Isopods, vertebrate muscle tissue, crab larvae, fish remains, crusta- ceans, amphipods
Randall 1967
Carangidae - jacks, pompanos Caranx crysos - blue + runner
+ 2, 4, 5,
7. 11
Family of swift- svlmmlng, carniv- orous fishes, often running in schools, wide- ranging (Randall 1967)
Caranx hippos crevalle Jack
Caranx ruber bar jack
Chloroscombrus chrysurus Atlantic bumper
Ollgoplltes 8aurua leatherjacket
Trachlnotus carolinus Florida pompano
Trachlnotus falcatus permit
+ 2, 5, 7, Fishes, crustaceans 8, 11, 13
+ 4, 11 160-547 mm: fish, shrimp, myslds, stomatopods, gastropods
+ 2, 11, 17, 18
+ 2, 3, 5, Snapping shrimp, penaeld shrimp, 8, 11, 13 larval anchovies, lady fish, harpactlcoid copepods
+ 11 sardines (Harengula sp.) , mole crabs (Hlppa sp . ) , bivalves (Donax sp.)
+ 7, 11 15-70 mm: myslds, shrimp,
anchovies, sllversldes, crabs, snails
Odum 1971
Randall 1967
Tabb & Manning 1961
Springer &
Woodburn
1960
Common over mud bottoms (Randall 1967)
Carr 6 Adams More apt to occur 1973 over sandy bottoms than T. carolinas (Randall 1967)
Selene vomer lookdown
+ +2,3, 7, Young: shrimp and other
11 crustaceans, small molluscs
Peterson 6 Peterson 1979
118
|
Habitat Type |
Reference |
Diet |
Diet Reference |
||
|
Family and Species |
■o u ■H U |
01 q M 3 U 03 |
u <H d u (0 O CQ |
Comments |
Hemlcaranx amblyrhynchus - bluntnose Jack
Caranx latus horse-eye Jack
Lutjanidae - snappers Lut Janus ana lis mutton snapper
Lut J anus apod us schoolmaster
Lut Janus grlseus gray snapper
+ +
+ + +
17
12
1, 4. 11
Predaceous on other fishes
204-620 mm: crabs, fish, gastropods, octopods, hermit crabs, penaeid shrimp, spiny lobster, stomatopods
Darnell 1958
Randall 1967
1, 4, 5, Crustaceans (shrimp, snapping 7, 11 shrimp, blue crabs, xanthid crabs, grapsid crabs), fish
Nugent 1970
1-5, 7, <50 mm: reside in grassbeds Odum 1971
8, 11-13, feeding on small crustaceans,
15-18 insect larvae
95-254 mm: reside in mangrove creeks feeding on crustaceans (snapping shrimp, xanthid crabs, penaeid shrimp, crayfish, caridean shrimp), fish including gobies, anchovies, poeciliids, eels, killiflshes
Considered by Gunter (1956) to be euryhaline
Commonly found over sand, sea- grass, rubble, coral reefs (Randall 1967)
By far the most abundant snapper in mangrove habitats
Lut Janus Jocu dog snapper
Lutjanus synagris lane snapper
Gerreidae - moj arras
Diapterus olisthostomus Irish pompano
Diapterus plumleri + + + striped mojarra
Eucinostomus argenteus + + + spotfin mojarra
Eucinostomus gula silver Jenny
Eucinostomus lefroyi mottled mojarra
Gerres clnereus yellowfin mojarra
Pomadasyidae - grunts
+ + +
1 190-630 mm: fish, crabs, octopods, spiny lobster, gastropods
1, 2, 3, snapping shrimp, crabs, 5, 7, 11, anchovies, annelids, molluscs 16-18
2 110-116 mm: green algae (Enteromorpha flexuosa, Cladophora) , Ruppia maritima, blue-green algae (Lyngba majuscula)
2, 7, 8, 36-172 mm: mysids, amphipods, 11-13, harpacticoid copepods, 15, 18 chironomid larvae, ostracods, bivalves, plant detritus
1-5, 7, 19-63 mm: amphipods, 8, 11-13, chironomids, harpacticoid 16-18 copepods, ostracods, mysids, molluscs, plant detritus
1-3, 5, 7, 8, 11-13, 16-18
10
2, 7, 11
19-70 mm: amphipods, chironomid larvae, harpacticoid copepods, molluscs, mysids, ostracods, plant detritus
Crabs, bivalves, gastropods, polychaete worms, shrimp, ostracods
Family carnivorous though rarely piscivorous
Randall 1967
Stark &
Schroeder
1970
Austin & Austin 1971
Odum 1971
Odum 1971
Odum 1971
Known from brackish water to depths of 220 fathoms (Randall 1967)
A permanent resident (Odum 1971)
Randall 1967, Austin & Austin 1971
Randall 1967
Most shelter on coral reef by day, feed on grassy flats by night (Randall 1967)
119
Habitat Type
Family and Species
o co Reference
Diet
Diet Reference
Comments
Amsotremus virginicus porkfish
Haemulon aurolineatum
11 112-264 mm: brittle stars, crabs, Randall shrimp, polychaetes , isopods , 1967 bivalves, stomatopods, gastropods
1, 4, 11 97-170 mm: shrimp & shrimp lar- Randall vae, polychaetes, hermit crabs, 1967 amphipods , copepods, gastropods, bivalves
Haemulon carbonarium Caesar grunt
Haemulon flavolineatum
French grunt
156-273 mm: crabs, gastropods, Randall sea urchins, chitons, poly- 1967 chaetes, brittle stars, sipun- culid worms, shrimp
113-228 mm: polychaetes, crabs, Randall sipunculid worms, chitons, 1967 holothurians, isopods, shrimp, bivalves
Haemulon parrai sailor's choice
1, 7 Benthic invertebrates including shrimp , crabs , amphipods , gas- tropods , polychaete worms, bivalves
Randall 1967
Haemulon plumieri white grunt
Haemulon album
1, 2, 11, 18
margate
Haemulon sciurus bluestriped grunt
Orthopristis chrysop-
tera
pigfish
Spaxidae - porgies
Archosargus pr oba toe eph a -
lus
sheepshead
Archosargus rhomboidalis sea bream
Calamus arctifrons
grass porgy
Calamus calamus saucereye porgy
I, 3, 5, 7, 11
1-3, 5, 11, 16-18
130-279 mm: crabs, polychaete Randall worms, sea urchins, sipunculid 1967 worms, gastropods, shrimp, brittle Reid stars; juveniles: copepods, mysids 1954
Benthic invertebrates including Randall crabs, shrimp, polychaete worms, 1967 amphipods , copepods , snails , bivalves
Benthic invertebrates including crustaceans , molluscs , annelid worms
Juveniles: 16-30 mm: plankton including copepods, mysids, postlarval shrimp
>30 mm: polychaetes, shrimp, amphipods
2, 3, 5, 7, <40 mm: in grassbeds - copepods , 8, 11-13 , amphipods , chironomid larvae , 17-18 mysids , algae, molluscs
>40 mm: in mangrove creeks - mussels, false mussels, crabs, snapping shrimp, crayfish, hydrazoans , algae , plant detritus 32-85 mm: in Puerto Rico man- groves - 100% blue-green algae (Lyngbya mojuscula)
5, 11 105-220 mm: seagrasses Cymodocea Randall & Thalassia, algae, crabs, gas- 1967 tropods, invertebrate eggs, bivalves
|
Randall |
|
|
1967 |
|
|
Carr & |
Strong preference |
|
Adams |
for vegetated sub |
|
1973 |
strate in bay |
|
areas (Weinstein |
|
|
et al. 1977) |
|
|
Odum 1971 |
|
|
Austin & |
|
|
Austin |
|
|
1971 |
Usually seen in mangrove sloughs, rare on reefs (Randall 1967)
Copepods , amphipods , mysids , Reid shrimp, bivalves, gastropods (Hitrella, Bittium) , polychaetes
1954 Associated with
grassy flats (Tabb s. Manning 1961)
190-250 mm: polychaetes, brittle Randall stars, bivalves, hermit crabs, 1967 sea urchins, gastropods, chitons
120
Habitat Type
Family and Species
|
01 |
|
|
a |
|
|
■H |
|
|
« OJ |
3 |
|
TT U |
|
|
i-( U |
on a |
|
H W |
Diet
Diet Reference
Lagodon rhomboides pinfish
1-3, 5, 7, B, In mangrove creek - scorched 11, 12, 16- mussel, mysids, amphipods, 18 false mussel
In Whitewater Bay - 100% plant
material
Odum 1971 Strong preference Reid 1954 for vegetated sub- strate in bay areas (Weinstein et al. 1977)
Sciaenidae - drums Balrdiella batabana blue croaker
Balrdiella chrysura silver perch
Cynoscion arenarlus sand seatrout
Cynoscion nebulosus spotted seatrout
3, 11
Lelostomus xanthurus spot
Menticirrhus americanus Southern kingfish
Menticirrhus littoralis Gulf kingfish
Micropogon undulatus Atlantic croaker
Pogonlas cromis black drum
Sciaenops ocellata red drum
|
+ 1-3, 8, Il- ls, 16-18 |
Larvae: copepods, larval fish (Henidia beryllina) |
Odum 1971 |
|
127-181 mm: fish (Anchoa mitchilli) , mysids |
||
|
2, 12, 17, 18 |
Mostly fish, caridean shrimp, mysids, amphipods, crab zoea |
Springer & Woodburn 1960 |
|
+ 1-3, 5, 7, 8, 11-13, 15, 17, 18 |
<50 mm: copepods, planktonic Crustacea 50-275 mm: fish (Mugil cephalus, Lagodon rhomboides, Eucino- stomus gula, E. argenteus, Cyprinodon varieqatus, Gobiosoma robustum, Anchoa mitchilli) |
Odum 1971 |
|
2, 7, 12, 17-18 |
<40 mm: planktonic organisms >40 mm: filamentous algae, desmids, forams, amphipods. |
Springer & Woodburn 1960 |
2, 11-12, 17-18
11, 12
+ 2, 7, 11, 12, 15
2, 3j 5, 8,
11-13, 15,
17
Equetus acuminatus high-hat
mysids, copepods, ostracods, isopods , chaetognaths , bi- valves, snails, polychaete worms
Fish, benthic crustaceans
Polychaetes , bivalves (Dona*) , sand crab Emerita), razor clams
Juveniles: copepods, mysids, caridean shrimp, polychaete worms, insect larvae, iso- pods, small bivalves
<100 mm: molluscs, xanthid
crabs >100 mm: bivalves, amphipods,
blue crabs, penaeid shrimp,
caridean shrimp
<10 mm: planktonic organisms
(copepods, crab zoea, larval
fish) 34-42 mm: mysids, amphipods,
caridean shrimp >50 mm: xanthid & portunid
crabs, penaeid shrimp,
small fish 308-403 mm: xanthid crabs
68-152 mm: shrimp S shrimp larvae, isopods, stoma topod larvae , copepods , amphipods
Springer &
Woodburn
1960
Springer & Most common off sandy Woodburn beaches (Springer & 1960 woodburn 1960)
Springer &
Woodburn
1960
Darnell 1958
Odum 1971
Randall 1967
Characteristic of coral reefs (Randall 1967)
121
Habitat Type
Family and Species
|
a |
||
|
•S |
||
|
H |
a |
3 |
|
a |
||
|
■d |
u |
|
|
*-> |
CO « |
|
|
H |
■s> |
u oa |
Diet Reference
Ephippidae - spadefishes Chaetodipterus faber ■Atlantic spadef ish
Pomacentridae - damsel fishes
Abudefduf saxatilis
sergeant major
2, 3, 5,
11, 16-18
Worms, crustaceans, debris Darnell
1961
101-135 mm: copepods, algae, Randall fish eggs, fish, shrimp larvae, 1967 polychaetes
Juveniles (7-12 mm) inhabit very shallow nearshore sandy beaches. Bear a deceptive resemblance to infertile red mangrove seed pods (Breder 1946)
Characteristic family of coral reefs (Ran- dall 1967)
A habitat generalist: reefs , grassbeds , rock piles, wharfs (Bohlke & Chaplin 1968)
Labridae - wrasses
Halichoeres bivittatus slippery dick
67-153 mm: crabs, sea urchins, Randall polychaetes , gastropods , brittle 1967 stars, bivalves , shrimp, fish, hermit crabs
Shallow water patch reefs, sand bottoms, gr a s sbeds ( Randa 1 1 1967)
Scaridae - parrotfishes Nicholsina usta emerald parrotfish
+ 1, 2, 11, 18
Family herbivorous, feeding primarily on algae growing
on hard substrates, on seagrasses
secondarily
Randal] Family characteris- 1967 tic of coral reefs, ranging into grass- beds
Scarus coeruleus blue parrotfish
Scarus croicensis
striped parrotfish
Spar i soma chrysopterum redtail parrotfish
Sparisoma rubripinne redfin parrotfish
Sparisoma viride stoplight parrotfish
Mugilidae - mullets Mugil cephalus striped mullet
Mugil curema white mullet
11
+ 2, 3, 5, 7,
11-13, 15
+ 2, 5, 7, 11-12
Inorganic sediments, fine detritus , micro-algae
25-73 mm: plant detritus, blue- green algae (Lyngbya majuscula)
Odum 1971
Austin &
Austin
1971
Requires near marine salinities {Tabb & Manning 1961)
Mugil trichodon fantail mullet
Sphyraenidae - barracudas Sphyraena barracuda great barracuda
Opistognathidae - jawfishes Opistognathus maxillosus mottled jawfish
+ 2, 7, 11, 12
1-5, 7, 8, 11, 13
135-369 mm: fish (Eucinostomus quia, Menidia beryllina, Archo- sargus probatocephalus)
53-110 mm: shrimp, isopods, fishes, polychaetes, mysids, copepods
Odum 1971
Randa 1 1 1967
Salinities >10 o/oo (Odum 1971)
Family lives in burrows in sediment, often in vicinity of reefs (Randall 1967)
122
Family and Species
Habitat Type
|
0} |
|
|
3 |
|
|
1 |
|
|
0) |
3 |
|
M |
*-> >> |
|
u |
<n cu |
|
03 |
Cx3 32 |
o « Reference
Diet Reference
Clinidae - clinids Chaenopsis ocellata bluethroat pikeblenny
Family appears to be carnivorous Randall on benthic invertebrates 1967
Inshore on rock, coral or rubble substrates (Ran- dall 1967)
Paraclinus marmoratus marbled blenny
Paraclinus fasciatus banded blenny
Stathmonotus hemphilli blackbelly blenny
+ 1, 5, 11
Blenniidae - combtooth blennies
Chasmodes saburrae
Florida blenny
Blennius marmoreus
seaweed blenny
1-3, 11, 17, 21-25 mm: amphipods 18 25-60 mm: amphipods, polychaetes , snails
detritus,
Algae, organic detritus, brittle stars, polychaetes, hydro ids
Carr &
Adams
1973
Randall 1967
Common brackish water blenny (Tabb & Manning 1961)
Blennius nicholsi highfin blenny
Callionymidae - dragonets Callionymus pauciradiatus spotted dragonet
Eleotridae - sleepers Dormitator maculatus fat sleeper
Gobiidae - gobies
Bathygobius sopor a tor frillfin goby
Gobionellus hastatus sharptail goby
13, 15
2, 3, 8,11, Caridean shrimp, chironomid 17 larvae, amphipods
Filamentous algae (Entero- morpha) , ostracods, copepods, insect larvae
Odum 1971
Springer &
Woodburn
1960
Freshwater and low salinity areas (Darnell 1961)
Gobionellus shufeldti freshwater goby
Gobionellus smaragdus emerald goby
2, 17, 18
3, 8, 10, 11, 15
Gobiosoma bosci naked goby
2, 12 Small crustaceans including Petersons amphipods, annelids, fish, Peterson fish eggs 1979
Gobiosoma longipala twoscale goby
Gobiosoma macrodon tiger goby
Gobiosoma robustum code goby
Lophogobius cyprinoides crested goby
1-3, 5, 8, Amphipods, mysids, chironomid 11, 16-18 larvae
1-3, 7, 8, A versatile feeder: amphipods, 13 mangrove detritus, filamentous algae, mysids, caridean & penaeid shrimp, polychaete worms , ostracods , bivalves , chironomid larvae, harpacticoid copepods, isopods, xanthid crabs , snails
Odum 1971
Odum, 1971
123
Family and Species
|
Habitat |
rype |
|
|
a |
||
|
u |
||
|
a |
h |
•H |
|
■ |
a |
|
|
(Q (U |
9 |
1 |
|
4J > |
41 >» |
|
|
■H 4J |
a> Q |
O Q |
|
H v. |
W 03 |
O 03 |
Diet Reference
Comments
Microgobius gulosus clown goby
Microgobius microlepis banner goby
Microgobius thalassinus green goby
2,5,8, 11- Amphipods, copepods, chironomid Odum 1971 13, 15, 17, larvae
18
5
3, 12
Planktonic organisms
Bird song 1981
Small crustaceans including Peterson & amphipods, other invertebrates Peterson
1979
Scombridae - mackerels, tunas
Scomberomorus maculatus
Spanish mackerel
2, 11, 12, Adults feeding on penaeid Tahb & 15 shrimp migrating from tidal Manning stream 1961
Scomberomorus cavalla king mackerel
11
350-1022
fish
Randall 1967
Scorpaenidae - scorpion- fishes
Scorpaena brasiliensis
barbfish
Scorpaena grandicornis plumed scorpionfish
11 Shrimp, other crustaceans, fish
1 37-102 ram: shrimp, fish, unidentified crustaceans
|
1967 |
|
|
Randall |
Most often found |
|
1967 |
in seagrass |
Triglidae - searobins Prionotus salmonicolor blackwing searobin
Prionotus scitulus leopard searobin
Prionotus tribulus blghead searobin
+ 1-3, 11, Small molluscs, shrimp, crabs 16-18 fish, small crustaceans (ostracods , cumaceans)
+ 1-3, 11-13, Shrimp, crabs, fishes, amphi- 17 i 18 pods, copepods, annelids, bivalves, sea urchins
Peterson & Peterson 1979
Peterson & Peterson 1979
Bothidae - lefteye flounders
Bo thus ocellatus
eyed flounder
Citharichthys macrops spotted whiff
Citharichthys
spllopterus bay whiff
i, n
1, 17, 18
68-130 ram: fish, crabs, shrimp, amphipods
Mainly myslds, also shrimp, crabs, copepods, amphipods, fishes, annelids
Randall 1967
Peterson & Recorded from Peterson salinity range 1979 2.5-36.7 o/oo (Darnell 1961)
Etropus crossotus fringed flounder
Farallchthys albigutta Gulf flounder
Parallchthys lethostlgma Southern flounder
3, 11, 16
1-3, 7, 11,
12, 17, 18
Calanoid copepods, cumaceans, Peterson &
amphipods, myslds, shrimp, Peterson
crabs, isopods, annelids, 1979 molluscs, fishes
<A5 mm: small crustaceans. Springer &
including amphipods, small Woodburn
fish I960-, Reid
>45 mm: fish (pigflsh, plnfish, 1954 lizardflsh, bay anchovy, labrids), crustaceans
Mainly fishes (mullet, menha- Peterson &
den, shad, anchovies, plnfish, Peterson
mojarras, croakers), crabs, 1979 mysids, molluscs, penaeid shrimp, amphipods
124
Family and Species
Habitat Type
Reference
Diet
Diet Reference
Comments
Syacium papillosum dusky flounder
Soleidae - soles Achirus lineatus lined sole
1-3, 5, 8,
11-13, 17-
18
32-74 mm: chironomid larvae, polychaete worms, foraminiferans
Odum 1971
Trinectes inscriptus scrawled sole
Trinectes maculatus hogchoker
2, 3, 8, 11-13, 17, 18
14-110 mm: amphipods , mysids
Odum 1971
Cynoglossidae - tongue- fishes
Symphurus plagiusa blackcheek tonguefish
Balistidae - triggerfishes & filefishes
Aluterus schoepfi
orange filefish
Balistes vetula
queen triggerfish
Monacanthus ciliatus fringed filefish
Monacanthus hispidus planehead filefish
Balistes capriscus gray triggerfish
1, 3, 11, 12, 16-18
1, 11
35-102 mm: polychaete worms, ostracods, portunid crabs, Ruppia and Halodule plant tips
Seagrasses, algae, hermit crabs, gastropods
11 130-480 mm: sea urchins, crabs, bivalves, brittle stars, poly- chaetes, hermit crabs, gastro- pods, algae
1, 11, 17 47-97 mm: Algae, organic detri- tus , seagrass , copepods, shrimp & shrimp larvae, amphipods, tanaids, polychaetes, molluscs
1-3, 11, Detritus, bryozoans, annelids, 16-18 harpacticoid copepods , amphi- pods, hermit crabs, molluscs, algae, sea urchins
Austin Austin 1971
Randall 1967
Randall 1967
Randall, 1967
Springer & Woodburn 1960
Peterson &
Peterson
1979
Associated with grassbeds , sponge/sea fan habitats (Ran- dall 1967, Voss et al. 1969)
Solitary reef fish ranging into grass- beds
Closely associated with vegetated areas (Tabb & Manning 1961)
Associated with vegetated areas {Tabb & Manning 1961)
Ostraciidae - boxfishes Lactophrys quadracornis scrawled cowfish
Lactophrys trigonus trunkfish
Lactophrys triqueter smooth trunkfish
Tetraodontidae - puffers Sphoeroides nephelus southern puffer
I, 2, 5, 7,
II, 16-18
1, 4, 11
1-3, 5, 11, 16-18
Vegetation , algae , bivalves
109-395 mm: crabs, bivalves, polychaetes, sea urchins, algae, seagrass, gastropods, amphipods
93-250 mm: polychaetes, sipun- culid worms, crabs, shrimp, gastropods, hermit crabs, sea urchins, bivalves
Juveniles : detritus , fecal pellets, zooplankton, poly- chaetes, gastropods, crabs, shrimp Adults : small crabs , bivalves
Reid 1954 Young mimic sea- grass blades (Bohlke & Chaplin 1968)
|
Randall |
Primarily a resident |
|
1967 |
of seagrass (Randall |
|
1967) |
|
|
Randall |
Primarily a reef |
|
1967 |
species (Randall |
|
1967) |
Carr &
Adams
1973
125
Habitat Type
Family and Species
M 3, Reference
Diet
Diet Reference
Comments
Sphoeroides spengleri bandtall puffer
Sphoeroides testudineus + checkered puffer
Diodontidae - porcupine- fishes
Chilomycterus antenna tus
bridled burrfish
Chilomycterus ant 11 la rum web burrfish
1» 7, 11 Crabs, bivalves, snails, polychaetes, amphipods, shrimp
1, 7 85-92 mm: portunid megalops larvae, gastropods
11 Gastropods, hermit crabs, isopods, crabs, shrimp
|
Randall |
Inhabits sea- |
|
1967 |
grass, reef, |
|
rubble, man- |
|
|
groves (Randall |
|
|
1967; Voss et al |
|
|
1969) |
|
|
Austin & |
|
|
Austin 1971 |
|
Randall |
Reefs and grass |
|
1967 |
beds (Voss |
|
et al. 1969) |
Chilomycterus schoepf 1 striped burrfish
1-3, 5, Gastropods, barnacles, crabs, 11, 16-18 amphipods
Springer &
Uoodburn
1960
Associated with
grassbeds (Voss
et al. 1969)
Salinities
>25 o/oo (Springer
& Woodburn 1960)
Reference Numbers Key
1. Bader & Roessler 1971
2. Carter et al. 1973
3. Clark 1970
4. Holm 1977
5. Hudson et al. 1970
6. Rush Ian & Lodge 1974
7. Nugent 1970
8. Odum 1971
9. Rivas 1969
10. Seaman et al. 1973
11. Schmidt 1979
12. Springer 6, Woodburn 1960
13. Tabb 1966
14. Tabb, Dubrow & Manning 1962
15. Tabb & Manning 1961
16. Weinstein et al. 1977
17. Yokel 1975a
18. Yokel 1975b
126
APPENDIX C. Amphibians and reptiles recorded from south Florida mangrove swamps .
127
AMPHIBIANS AND REPTILES OF FLORIDA'S MANGROVES
Species
Status
Food Habits
Mud Turtle
(Kinosternon subrubrum)
Abundant
Insects, crustaceans, mollusks
Striped Mud Turtle (Kinosternon bauri )
Common
Algae, snails, dead fish
Ornate
Diamondback Terrapin (Malaclemys terrapin macrospilota and M.t_. rhizophorarum)
Florida Red-bellied Turtle (Chrysemys nelsoni)
Chicken Turtle
(Deirochelys reticularia)
Green Turtle
(Chelonia mydas)
Hawksbill
(Eretmochelys imbricata)
Loggerhead
(Caretta caretta)
Atlantic Ridley
(Lepidochelys kempii)
Florida Softshell (Trionyx ferox)
Green Anole
(Anolis carolinensis)
Cuban Brown Anole (Anolis sagrei)
Bahaman Bank Anole (Anolis distichus)
Green Water Snake (Nerodia cyclopion)
Mangrove Water Snake (Nerodia fasciata compressicauda)
Uncommon
Littorina, Melampus, Uca . Anomalocardia
Rare - Uncommon Sagittaria, Lemna, Naias
Uncommon
Uncommon
Rare
Common
Uncommon
Common
Common
Common
Uncommon
Common
Common
Crayfish, insects, Nuphar
Mangrove roots and leaves , seagrasses
Rhizophora: fruits, leaves wood , bark
Crabs, jellyfish, tuni- cates
Snails, crabs, clams
Snails, crayfish, mussels, frogs, fish, waterfowl
Insects
Insects
Insects
Fish
Fish, invertebrates
128
AMPHIBIANS AND REPTILES OF FLORIDA'S MANGROVES (concluded)
Species
Status
Food Habits
Striped Swamp Snake Uncommon
(Liodytes alleni)
Eastern Indigo Snake Uncommon
(Drymarchon corais)
Rat Snake Uncommon
(Elaphe obsoleta)
Eastern Cottonmouth Uncommon
(Agkistrodon piscivorus)
American Alligator Common
(Alligator mississippiensis)
American Crocodile Rare
(Crocodylus acutus)
Giant Toad Common
(Bufo marlnus)
Crayfish, sirens, frogs
Small mammals, birds, frogs
Small mammals , birds
Fish, frogs, snakes, birds , small mammals
Fish, waterbirds
Fish, waterbirds
Invertebrates
Squirrel Treefrog (Hyla squirella)
Cuban Treefrog
(Hyla septentrionalis)
Abundant
Common
Insects
Insects, frogs, toads, lizards
References :
Carr and Goin 1955; Ernst & Barbour 1972; Mahmuud 1965; L. Narcisse, R.N. "Ding" Darling Fed. Wildlife Refuge, Sanibell Is., Fla. ; personal communication (1981) .
129
APPENDIX D. Avifauna of south Florida mangrove swamps.
130
WADING BIRDS
Common Name (Latin name)
Season of
Occurrence3 Nestinya Food Habits
References
Great Egret
(Casmerodius albus)
Howell, 1932
Kushlan & White 1977a
Snowy Egret
(Egretta thula)
Fish
Howell 1932
Kushlan 6 White 1977a
Ffrench 1966
Cattle Egret
(Bubulcus ibis)
Fish
Howell 1932
Kushlan & White 1977a
Great White Heron (Ardea herodias occidentalis)
Fish
Howell 1932
Kushlan & White 1977a
Great Blue Heron (Ardea herodias)
Howell 1932
Kushlan & White 1977a
Reddish Egret (Dichromanassa ruf escens)
Howell 1932
Kushlan & White 1977a
Louisiana Heron Common (Hydranassa tricolor)
Kushlan s. White 1977a Maxwell & Kale 1977 Girard & Taylor 1979
Little Blue Heron (Florida caerulea)
Kushlan & White 1977a Maxwell & Kale 1977 Girard & Taylor 1979
Green Heron Common
(Butorides striatus)
Fish
Robertson & Kushlan 1974 Maxwell & Kale, 1977 Girard & Taylor 1979
Black-crowned Night Heron
(Nycticorax nycticorax)
Fish , crustaceans , frogs, mice
Ffrench 1966 Maxwell & Kale 1977 Girard & Taylor 1979
Yellow-crowned Night Heron
(Nyctanassa violacea)
Fish, crayfish, crabs
Ffrench 1966
Girard & Taylor 1979
Least Bittern
(Ixobrychus exilis)
Fish
Ffrench 1966
American Bittern (Botaurus lentiginosus)
W,T
Crayfish, frogs, small fishes
Narcisse , pers . comm .
Wood Stork
(Mycteria americana)
Common (locally abundant)
Kahl 1964
Ogden et al. 1976
Kushlan 1979
Glossy Ibis
(Plegadis falci- nellus
Uncommon
Fish
Bacon 1970 Howell 1932
White Ibis
(Eudocimus albus)
Fish, crabs (Uca)
Kushlan 1979
Kushlan & Kushlan 1975
Girard & Taylor 1979
Roseate Spoonbill (Ajaia ajaja)
Sandhill crane
(Grus canadensis)
Rare to Uncommon
Shrimp, fish, aquatic vegetation
Roots, rhizomes of Cyperus & Sagit-
taria
Kushlan & White 1977a Howell 1932
Ogden 1969 Howell 1932
Limpkin
(Aramus guar a una)
Snails (Pomacea)
Howell 1932 Bacon 1970
131
PROBING SHORE BIRDS
Common Name (Latin name)
Abundance
Season of Occurrence3
Nesting*
Food Habits
References
King Rail
(Rail us elgans)
Clapper Rail
(Rallus longiro- stris)
Virginia Rail
(Rallus limicola)
Sora
(Porzana Carolina)
Black Rail (Laterallus jamaicensis)
Semipalmated Plover (Charadrius semi - palmatus)
Wilson's Plover
(Charadrius wilsonia)
Black-bellied Plover (Pluvialis squatarola)
Ruddy Turnstone
(Arenaria lnterpres)
Common Snipe
(Capella gallinago)
Common
Uncommon- common
Rare
Uncommon to
locally
abundant
Rare
Locally common
Locally common
Yr
W,T
Uncommon
W,T
Beetles, grass- hoppers, aquatic bugs
Crabs, shrimp
Beetles, snails, spiders
Insects, seeds of emergent aquatic plants
Beetles, snails
Crustaceans, mollusks
Crabs, shrimp, crayfish
Crabs , mollusks
Insects, crus- taceans , mollusks
Mollusks, insects, worms
Narcise, pers. comm. Martin et al. 1951
Howell 1932 Ffrench 1966 Bacon 1970
Marcisse, pers. coram. Martin et al. 1951
Howell 1932 Bacon 1970
Narcisse, pers- coram.
Ffrench 1966
Bacon 1970
Baker & Baker 1973
Howell 1932 Bacon 1970
Howell 1932 Bacon 1970 Ffrench 1966
Ogden 1969 Howell 1932
Howell 1932 Bacon 1970
Long-billed Curlew
(Numenlus americanus)
Whimbrel
(Numenius phaeopus)
Spotted Sandpiper (Ac tit is macularia)
Solitary Sandpiper (Tringa solitaria)
Willet
(Catoptrophorus semipalmatus)
Greater Yellowlegs (Tringa melanoleucas)
Lesser Yellowlegs (Tringa flavipes)
Rare-uncommon WtT
Abundant
Common
Common
Red Knot
(Calidrls canutus)
Dunlin
(Calidris alplna)
White-rumped Sandpiper Rare (Calidris fusclcollls)
Common
Uncommon Common
W,T
W,T
Yr
W,T
W,T
W,T
Crustaceans, insects
Mollusks, crus- taceans , worms , insects
Mollusks, crus- taceans
Ogden 1969
Ogden 1968 Howell 1932
Ffrench 1966 Bacon 1970 Russel 1980
Crustaceans, aquatic Howell 1932 insects, small frogs Bacon 1970
Crabs, crayfishes, Howell 1932 killifishes Bacon 1970
Fishes, crabs , crustaceans
Snails, mollusks, crabs
Marine worms , crustaceans
Marine worms, mollusks
Howell 1932 Ffrench 1966 Bacon 1970
Ffrench 1966
Bacon 1970
Baker & Baker 1973
Howell 1932 Ogden 1964
Ogden 1964
Baker & Baker 1973
Chironomids, snails Howell 1932 Bacon 1970
132
PROBING SHOREBIRDS (concluded)
|
Common Name (Latin name) |
Abundance |
Season of Occurrence3 |
|
Least Sandpiper (Calidris minutilla) |
Common |
W,T |
|
Short-billed Dowitcher (Limnodromus griseus) |
Common |
W,T |
|
Stilt Sandpiper (Micropalama himantopus) |
Rare-uncommon |
W,T |
|
Semipalmated Sandpiper (Calidris pusilla) |
Common- abundant |
W,T |
|
Western Sandpiper (Calidris mauri) |
Common- abundant |
W,T |
|
Marbled Godwit (Limosa fedoa) |
Rare-common |
W |
Nesting3
Food Habits
References
American Avocet (Recurvlrostra americana)
Uncommon
W,T
Pupae of beetles and flies
Mollusks, crustaceans
Chironomids
Mollusks, insects
Chironomids
Crustaceans, mollusks, seeds of emergent aquatic plants
Marine worms, aquatic insects
Bacon 1970
Baker & Baker 1973
Bacon 1970
Baker 6. Baker 1973
Howell 1932 Bacon 1970
Bacon 1970
Baker & Baker 1973
Howell 1932 Bacon 1970
Howell 1932
Ogden 1969
Black-necked Stilt Common (Himantopus mexicanus)
Aquatic beetles
Howell 1932 Bacon 1970
133
SURFACE AND DIVING BIRDS
|
Common Name |
Season of |
|
|
(Latin name) |
Abundance |
Occurrence' |
|
Common Loon |
Occasional |
w |
|
(Gavia immer) |
||
|
Horned Grebe |
Uncommon |
w |
|
(Podiceps auritus) |
||
|
Pied-billed Grebe |
Uncommon- |
Yr |
|
(Podilymb'us |
common |
|
|
podiceps) |
||
|
White Pelican |
Rare |
S |
|
(Pelecanus |
Common |
w |
|
ery throrhy nchos ) |
||
|
Brown Pelican |
Common |
Yr |
|
(Pelecanus |
||
|
occidentalia) |
||
|
Double-crested |
Common |
Yr |
|
cormorant |
||
|
(Phalacrocorax |
||
|
auritus) |
||
|
Anhinga |
Common |
Yr |
|
(Anhinga anhinga) |
||
|
Fulvous Whistling Duck |
Uncommon |
w |
|
(Dendrocygna |
||
|
bicolor) |
||
|
Mallard |
Uncommon |
W,T |
|
(Anas platyrhynchos) |
||
|
Black Duck |
Rare |
W,T |
|
(Anas rubripes) |
||
|
Mottled Duck |
Uncommon |
Yr |
|
(Anas fulvigula) |
||
|
Gad wall |
Uncommon |
W,T |
|
(Anas strepera) |
||
|
Pintail |
Abundant |
W,T |
|
(Anas acuta) |
||
|
Green-winged Teal |
Abundant |
W,T |
|
(Anas crecca carollnensis) |
||
|
Blue-winged Teal |
Abundant |
Yr |
|
(Anas discors) |
Common |
|
|
American Wigeon |
W,T |
|
|
(Anas americana) |
||
|
Northern Shoveler |
Common |
W,T |
|
(Anas clypeata) |
||
|
Wood Duck |
Rare |
W |
|
(Aix sponsa) |
||
|
Redhead |
Rare |
W |
|
(Aythya americana) |
Nesting'
Food Habits
References
Fish, crabs, mollusks Narcisse, pers. co
Fish, aquatic insects, Ogden 1969 mollusks
Crayfish, fish, mollusks
Widgeon grass
Mollusks, crusta- ceans, widgeon grass
Polygonum, snails, Ruppia
Ruppia , Zostera, mollusks
Saggitaria, mollusks, Cyperus
Ruppia , Zostera.- aquatic insects
Cyperus , snails, insects, crustaceans
Ruppia , Zostera, mollusks
mollusks, aquatic insects, Ruppia, Zostera
Nuts, seeds
Narcisse, pare, coram.
Narcisse, pers. conm.
Ffrench 1966 Bacon 1970
Kushlan & White 1977a
Ffrench 1966
Ogden 1969 Smith, pers. obs .
Ogden 1969
Kushlan et al., in prep.
Ogden 1969
LaHunt & Cornwell 1970 Kushlan et al., in prep.
Ogden 1969
Narcisse, pers. comm. Kushlan et al.,in prep.
Narcisse, pers. comm.
Kushlan et al., in prep.
Narcisse, pers. coram. Ffrench 1966
Narcisse, pers. comm.
Kushlan et al. , in prep.
Narcisse, pers. comm.
Ogden 1959
Snails, clams, aquatic Ogden 1969
insects, Ruppia, Zos-
tera
Ring-necked Duck Abundant
(Aythya collar is)
Polygonum, Ruppia , crayfish, snails
Ogden 1969
Kushlan et al., in prep.
Canvasback Uncommon
(Aythya valiaineria)
Vallisneria, Ruppia , Zostera
Ogden 1969
Kushlan et al., in prep.
134
SURFACE AND DIVING BIRDS (concluded)
Common Name (Latin name)
Abundance
Season of Occurrence
Nesting '
Food Habits
References
Lesser Scaup Common-
(Aythya affinls) abundant
Mollusks, Ruppia
Narcisse, pers. coram.
Ogden 1969
Kushlan et al., In prep.
Bufflehead
(Bucephala albeola)
Rare
Ruddy Duck Common W
(Oxyura jamaicensis)
Gastropods, crabs, Ogden 1969
crustaceans Kushlan et al., in prep.
Potamogeton, Najas, Ogden 1969 Zostera, Ruppia, Kushlan et al. mollusks
in prep.
Hooded Merganser (Lophodytes cucullatus)
Red-breasted Merganser (Mergus serrator )
Purple Gallinule (Porphyrula martlnica)
Common Gallinule
(Gallinula chloropus)
American Coot
(Fulica americana)
Rare-uncommon
Common
Rare
Common
Abundant
W.T
Yr
Yr
W,T
Fish
Fish
Ogden 1969
Narcisse, pers. coram.
Aquatic insects, Narcisse, pers. coram, mollusks, Ffrench 1966
Eleocharis, Paspalum
Seeds, aquatic insects
Ruppia, Na j as , Potamogeton, aquatic insects
Narcisse, pers. coram. Ffrench 1966
Narcisse, pers. coram.
135
AERIALLY SEARCHING
Common Name (Latin name)
Abundance
Season of Occurrence
Nesting" Food Habits
References
Herring Gull
(Larus argentatus)
Ring-billed Gull
(Larus delawarensis)
Common
Fish, mollusks, crustaceans
Fish, insects, mollusks
Narcisse, pers. comm. Ogden 1969
Narcisse, pers. comm. Ogden 1969
Laughing Gull Common
(Larus atricilla)
Bonaparte's Gull Uncommon (Larus Philadelphia)
Gull-billed Tern Uncommon (Gelochelidon nilotica)
Forster's Tern Uncommon- ( Sterna fosteri) common
Fish, shrimp, crabs
Fish, insects
Narcisse, pers. comm. Ogden 1969
Ogden 1969
Mayflies, dragonflies Ogden 1969
Fish
Narcisse, pers. comm. Ogden 1969
Common Tern
(Sterna hirundo)
Fish
Ogden 1969
Least Tern
(Sterna albifrons)
Fish
Narcisse, pers. comm. Ogden 1969
Royal Tern
(Thalasseus maxima)
Common
W,T
Fish
Ogden 1969
Sandwich Tern (Sterna sand- vicensis)
Uncommon
Yr
Fish
Narcisse, pers. comm. Ogden 1969
Caspian Tern
(Sterna caspia)
Uncommon
Fish
Ogden 1969
Black Skimmer Common
(Rynchops nigra)
Belted Kingfisher Common (Megaceryle alcyon)
Fish Crow Common
(Corvus ossifragus)
Yr
Fish
Fish
Fish
Ogden 1969
Narcisse, pers. comm.
Narcisse, pers. comm.
136
BIRDS OF PREY
Common Name (Latin name)
Season of Abundance Occurrence* Nesting3 Food Habits
References
Magnificent Frigate- Common S bird Uncommon W
(Fregata magnificens)
Turkey Vulture (Cathartes aura)
Common
Fish
Narcisse,
pera. com.
Smith, pers. obs.
Narcisse, pers. comm. Orians 1969
Black Vulture Common
(Coragyps atratus)
Swallow-tailed Kite Common (Elanoides forf ica- tus)
Sharp-shinned Hawk Uncommon (Accipiter striatus)
Cooper's Hawk Uncommon
(Accipiter cooper ii)
Red-tailed Hawk Uncommon (Buteo jamaicensis)
Red-shouldered Hawk Common (Buteo lineatus)
Broad-winged Hawk (Buteo platypterus)
Swainson's Hawk (Buteo swainsoni)
Short-tailed Hawk (Buteo brachyurus)
Bald Eagle (Haliaeetus leucocephalus)
Marsh Hawk
(Circus cyaneus)
Osprey
(Pandion haliaetus)
Peregrine Falcon (Falco peregrinus)
Merlin
(Falco columbarius)
American Kestrel Common (Falco sparverius)
Barn Owl Uncommo
(Tyto alba)
Great Horned Owl Uncommo (Bubo virginianus)
Barred Owl Uncommo
(Strix varla)
Rare- locally common (Fla. Bay)
Uncommon
Very rare- locally common (Fla. Bay)
Uncommon
Carrion
Snakes, lizards, frogs
Robertson & Kushlan
1974 Orians 1969
Howell 1932 Snyder 1974
Smaller passerines Howell 1932 Larger passerines Howell 1932 Small mammals, birds Howell 1932
Snakes, frogs, lizards. Insects
Insects, small mammals
Howel] 1932 Robertson & Kushlan. 1974
Howell 1932
Small mammals, grass- Howell 1932 hoppers
Small birds
Fishes
Howell 1932
Howell 1932
Small mammals, shore- Howell 1932 birds
Fishes
Howell 1932
Waterfowl, shorebirds Nisbet 1968 Ogden 1969 Howell 1932
Small birds, shore- Howell 1932 birds
Insects
Small mammals
Waterfowl , small mammals
Hovel) 1932
Howell 1932
Howell 1932
Y Small mammals, frogs, Howell 1932 snakes
137
ARBOREAL BIRDS
Common Name (Latin name)
Mourning Dove
(Zenaidura macroura)
White-crowned Pigeon (Columba leucocephala)
Mangrove Cuckoo (Coccyzus minor)
Abundance
Season of Occurrence3
Nesting3 Food Habits
Uncommon
Yr
Seeds
Berries, seeds fruits
Caterpillars, man t ids
References
Emlen 1977
Howell 1932
Robertson & Kushlan 1974
Howell 1932 Ffrench 1966
Robertson & Kushlan 1974 Martin et al. 1951
Yellow-billed Cuckoo Common (Coccyzus americanus)
Caterpillars, beetles
Howell 1932 Ffrench 1966 Martin et al. 1951
Smooth-billed Anl Rare Yr
(Crotophaga anl)
Chuck-will 's-widow Uncommon Yr (Caprimulgus carolinensis)
Common Flicker Uncommon Yr
(Colaptes auratus)
Pileated Woodpecker Uncommon Yr (Dryocopus pileatus)
Mosqultos, moths
Ants, beetles, fruits in winter
Beetles, berries, fruits
Howell 1932 Ffrench 1966
Martin et al. 1951 Narcisse, pers. comm.
Narcisse, pers. coram. Martin et al. 1951
Howell 1932 Robertson 1955 Robertson 4 Kushlan 1974
Red-bellied Woodpecker Common (Melanerpes carolinus)
Beetles, ants,
grasshoppers,
crickets
Narcisse, pers. coram. Martin et al. 1951
Red-headed Woodpecker (Melanerpes erythrocephalus)
Yellow-bellied Sapsucker
(Sphyraplcus varius)
Hairy Woodpecker (Picoides vlllosus)
Eastern Kingbird
(Tyrannus tyrannus)
Gray Kingbird (Tyrannus dominicensis)
Western Kingbird
(Tyrannus vertlcalus)
Great Crested Flycatcher
(Mylarchus crlnitus)
Acadian Flycatcher
(Empldonax virescens)
Eastern Phoebe
(Sayornis phoebe)
Eastern Wood Pewee (Contopus vlrens)
Uncommon
Common
Uncommon (common S)
Rare
Common
Rare-uncommon
Yr
W,T
S,T
W,T
Beetles, ants,
grasshoppers,
caterpillars
Beetles, ants-, caterpillars
Insects, beetle larvae
Ants, wasps, grasshoppers
Bees, wasps, beetles, dragon
Bees, wasps, grasshoppers
Insects, berries
Narcisse, pers. comm. Martin et al. 1951
Narcisse, pers. coram. Martin et al. 1951
Emlen 1977
Narcisse, pers. comm. Martin et al. 1951
Howell 1932
Robertson & Kushlan 1974
Narcisse, pers. comm. Martin et al. 1951
Howell 1932 Robertson 1955
Small flying insects Morton 1980
Bees, wasps, ants Narcisse, pers. comm. Martin et al. 1951
Bees, wasps, ants, Narcisse, pers. comm. moths Howell 1932
138
ARBOREAL BIRDS (continued)
Common *<ame (Latin name)
Season of Occurrence*
Nesting
Food Habits
References
Barn Swallow
(Hirundo rustica)
Blue Jay
(Cyanocitta crlstata)
Tufted titmouse (Parus bicolor)
Carolina Wren (Thryothorus ludovicianus)
Mockingbird
(Mimus polyglottos)
Catbird
(Dumetella caro- linensis)
Locally common
Very rare- rare
W,T
Grasshoppers, cater- pillars, beetles
Caterpillars , wasps , bees
Ants, flies, milli- peds
Fruits , berries
Fruits, insects
Howell 1932 Bacon 1970
Narcisse, pers. comm. Martin et al. 1951
Howell 1932
Robertson & Kushlan 1974
Narcisse , pers . comm . Martin et al. 1951
Robertson 1955
Narcisse , pers . comm . Martin et al. 1951
Brown Thrasher
(Toxo stoma rufum)
Narcisse, pers. comm. Martin et al. 1951
American Robin Abundan (Turdus migratorius)
Blue-gray Gnatcatcher Uncommo (Polioptila caerulea)
Ruby-crowned Kinglet Uncommo (Regulus calendula)
White-eyed Vireo Uncommo (Vireo griseus)
Black-whiskered vireo Uncommo (vireo altiloquus)
Red-eyed Vireo Uncommo (Vireo olivaceus)
Yellow-throated Vireo Uncommo (Vireo f lavifrons)
Black-and-white Fairly Warbler common
(Mniotilta varia)
Worm-eating Warbler Uncommo (Helmitheros vermi- vorus)
W,T
Worms , berries, insects
Insects , especially Hymenopterans
Wasps, ants Butterflies, moths
Spiders , caterpillars Caterpillars , beetles Butterflies , moths , Wood boring insects
Caterpillars, spiders
Narcisse, pers. comm. Martin et al. . 1951
Narcisse , pers . comm. Howell 1932
Narcisse, pers. comm. Howell 1932
Robertson 1955
Howell 1932
Robertson & Kushlan, 1974
Narcisse, pers . comm. Howell. 1932
Morton 1980
Lack and Lack 1972 Keast 1980 Ogden 1969
Ogden 1969
Kushlan , pers . comm .
Prothonotary Warbler (Protonotaria citrea)
Ffrench. 1966 Russel 1980
Yel low-throated Warbler
(Pendroica dominica)
Beetles, moths, spiders
Morton 1980
Yellow Warbler
(Pendroica petechia)
Y e 1 1 ow- rumpe d
Warbler Abundant
(Pendroica corona ta)
Prairie Warbler Uncommon (Pendroica discolor)
Palm Warbler Abundant
(Pendroica palmar urn)
yr
W,T
Pipterans , bayberries Moths, beetles, flies Insects
Haverschmidt 1965
Ffrench 1966
Orians i969
Terborgh & Faaborg 1980
Narcisse, pers . comm.
Lack & Lack 1972 Robertson & Kushlan 1974
Lack & Lack 1972 Emlen, 1977
139
ARBOREAL BIRDS (continued)
Common Name (Latin name)
Abundance
Season of Occurrence a
Nesting* Food Habits
References
Blackpoll Warbler Uncommon (Dendroica striata)
Ffrench 1966
Bay-breasted Warbler Rare (Dendroica castanea)
Morton 1980
Black-throated Green Uncommon Warbler
(Dendroica virens)
Aphids , leaf-rollers and other insects
Ogden 1969
Kushlan , pers . comm .
Chestnut-sided Warbler Rare (Dendroica pensyl- yanica)
Cape May Warbler Uncommon (Dendroica tigrina) Common
Black-throated Gray Rare Warbler
(Dendroica nigrescens)
Black-throated Blue Uncommon Warbler Common
(Dendroica caeru- lescens)
Beetles, flies, ants
Morton 1980
Ogden 1969
Ogden 1969 Kushlan, pers. comm. Hutto 1980
Kushlan, pers. comm. Ogden 1969
Northern Waterthrush Abundant (Seiurus novebora- Rare censis)
Schwartz 1964 Ffrench 1966 Bacon 1970 Russell 1980
Yellowthroat Common
(Geothlypus trichas)
Grasshoppers , crickets , Narcisse, pers . comm. ants, wasps Howell 1932
Lack & Lack 1972
American Redstart Common (Setophaga ruticilla)
Caterpillars
Bennett 1980 Ffrench 1966 Bacon 1970
Tennessee Warbler Uncommon (Vermivora peregrina)
Nasheville Warbler Rare (Vermivora ruf i- capllla)
Orange-crowned Warbler Common (Vermivora celata)
Golden-winged Warbler Rare (Vermivora chrysop- tera)
Northern Parula
(Parula americana)
Ovenbird
(Seiurus aurocapll- lus)
Kentucky Warbler
(Oporornis formosus)
Mourning Warbler
(Oporornis Philadel- phia)
Yellow-breasted Chat (Icteria virens)
Rar e - un co mmo n
Hymenoptera
Morton 1980
Hutto 1980
Hutto 1980
Morton 1980
Lack and Lack 1972
Beetles, crickets, Lack and Lack 1972 grasshoppers
Beetles, caterpillars, Morton 1980 ants
Hymenoptera
Morton 1980
Hutto. 1980
140
ARBOREAL BIRDS (concluded)
Common Nnmc (Latin name)
Abundance
Season of Occurrence3
Wilson's Warbler (Wilsonia pusilla)
Rar e - uncommon
Red-winged Blackbird Common (Agelaius phoeniceus)
Boat-tailed Grackle Uncommo (Quiscalus major)
Common Grackle Uncommo
(Quiscalus guiscula)
Card inal Common
(Cardinalis cardinalis)
Orchard Oriole Rare
(Icterus spurius)
Indigo Bunting Uncommo
(Passerina cyanea)
Summer Tanager (Piranga rubra)
Dickcissel
(Spiza americana)
Rufous-sided Townee (Pipllo erythroph- thalmus )
Swamp Sparrow
(Melospiza georgiana)
Uncommon
Uncommon
Yr
W,T
Yr
W,T
Nesting3 Food Habits
Seeds, insects
Crayfish, crabs, shrimp
Insects, cater- pillars
Insects, seeds
References
Hutto 1980
Ramos and Warner 1980
Howel] 1932 Robertson 1955
Robertson 1955 Girard & Taylor 1979
Howell 1932 Robertson 1955
Robertson 1955
Grasshoppers, beetles Morton 1980
Grasshoppers , cater- pillars
Hymenoptera
Narcisse, pers. comm. Howell 1932
Morton 1980
Caterpillars, beetles Bacon 1970
Martin et al. 1951
Caterpillars, bay- berries, fruits
Ants, flies, seeds
Narcisse, pers. comm. Howell 1932
Narcisse, pers. comm. Howell 1932
Tr = year round resident S = summer resident W = winter resident
T = transient, present only during spring and fall migration Y = species breeds in mangroves
bL. Narcisse, R.N. "Ding" Darling Fed. Wildlife Refuge, Sanibel Island, Fla. (1981). J. A. Kushlan, So. Fla. Res. Ctr. , Everglades Natl. Park, Homestead, Fla.
141
APPENDIX E. Mammals of south Florida mangrove swamps.
142
MAMMALS OF FLORIDA MANGROVES
Species
Status
Food Habits
Virginia Opossum
(Didelphis virginiana)
Abundant
Fruits, berries, insects, frogs, snakes, small birds and mammals
Short-tailed Shrew (Blarina brevicauda)
Uncommon
Insects
Marsh Rabbit
(Sylvilagus palustris)
Gray Squirrel
(Sciurus carolinensis)
Abundant
Occasional
Emergent aquatics
Fruits, berries, mast, seeds
Fox Squirrel
(Sciurus niger)
Marsh Rice Rat
(Oryzomys palustris)
Cudjoe Key Rice Rat (Oryzomys argentatus)
Rare
Uncommon
Rare
Cotton Rat Abundant
(Sigmodon hispidus)
Gray Fox Uncommon
(Urocyon ciner eoar genteus )
Black Bear Rare
(Ursus americanus)
Fruits, berries, mast
Seeds of emergent plants, insects, crabs
Seeds, insects, crabs
Sedges, grasses, cray- fish, crabs, insects
Small mammals, birds
Fruits, berries, fish, mice
Raccoon
(Procyon lotor)
Mink
(Mustela vison)
Abundant
Rare
Crayfish, frogs, fish
Small mammals, fish, frogs , snakes , aquatic insects
Striped Skunk
(Mephitis mephitis)
River Otter
(Lutra canadensis)
Common
Uncommon
Bird eggs and young frogs, mice, larger invertebrates
Crayfish, fish, mussels
143
MAMMALS OF FLORIDA MANGROVES (concluded)
Species
Status
Food Habits
Panther
(Felis concolor)
Very rare
Deer, rabbits, mice, birds
Bobcat
(Felis rufus)
Common
Rabbits, squirrels, birds
White-tailed Deer
(Odocoileus virglnianus)
Common
Emergent aquatics, nuts, acorns, occasionally mangrove leaves
Key Deer
(O.v. clavium)
Common on cer- Emergent aquatics and tain Florida Keys other vegetation (no longer on mainland)
Black Rat
(Rattus rattus)
Common
Bottle-nosed Dolphin (Tursiops truncatus)
West Indian Manatee (Trichechus manatus)
Uncommon
Uncommon
Fish
Submerged aquatics, Zostera, Ruppia , Halodule, Syringodium, Cymodocea , Thalassia
References: Layne 1974; Hamilton and Whittaker 1979;
L. Narcisse, R.N. "Ding" Darling Fed. Wildlife Refuge, Sanibel Island, Fla.; personal commu- nication.
144
50272 -101
REPORT DOCUMENTATION PAGE
l._REPORT NO.
FWS/OBS-81/24
3. Recipient's Accession No.
4. Title and Subtitle
THE ECOLOGY OF THE MANGROVES OF SOUTH FLORIDA: PROFILE
A COMMUNITY
5. Report Date
January 1982
7. Author(s)
William E. Odum, Carole C. Mclvor, Thomas J. Smith, III
8. Performing Organization Rept. No.
s. Address of Authors
Department of Environmental Sciences University of Virginia Charlottesville, Virginia 22901
10. Proiect/Task/Work Unit No.
11. Contract(C) or Grant(G) No. (C)
(G)
12. Sponsoring Organization Name and Address
Office of Biological Services Fish and Wildlife Service U.S. Department of the Interior Washington, D.C. 20240
New Orleans OCS Office Bureau of Land Management U.S. Department of the Interior New Orleans, Louisiana 70130
13. Type of Report & Period Covered
15. Supplementary Notes
16. Abstract (Limit: 200 words)
A detailed description is given of the community structure and ecosystem processes of the mangrove forests of south Florida. This description is based upon a compilation of data and hypotheses from published and unpublished sources.
Information covered ranges from details of mangrove distribution, primary production, and diseases to aspects of reproduction, biomass partitioning, and adaptations to stress. Mangrove ecosystems are considered in terms of zonation, succession, litter fall and decomposition, carbon export, and energy flow.
Most of the components of mangrove communities are cataloged and discussed; these include microorganisms, plants other than mangroves, invertebrates, fishes, reptiles, amphibians, birds, and mammals.
Finally, two sections summarize the value of mangrove ecosystems to man and present ways to manage this type of habitat. It is concluded that mangrove forests, which cover between 430,000 and 500,000 acres (174,000 - 202,000 ha) in Florida, are a resource of great value and should be protected and preserved wherever possible.
17. Document Analysis a. Descriptors
Ecology, value, management, fauna
b. Identifiers/Open Ended Terms
Mangroves, halophytes, coastal wetlands, ecosystem, South Florida
c. COSATI Field/Group
18. Availability Statement
Unlimited
19. Security Class (This Report)
Unr.lassifipH
20. Security Class (This Page)
21. No. of Pages
154
22. Price
(See ANSI-Z39.18)
See Instructions on Reverse
OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) Department of Commerce
«U.S. GOVERNMENT PRINTING OFFICE: 1982-572-346
ft
©■©
Headquarters - Office of Biological Services, Washington, D.C.
National Coastal Ecosystems Team,
Slidell. La.
Regional Offices
U.S. FISH AND WILDLIFE SERVICE REGIONAL OFFICES
REGION 1
Regional Director
U.S. Fish and Wildlife Service
Lloyd Five Hundred Building. Suite 1692
500 N.E. Multnomah Street
Portland. Oregon 97232
REGION 2
Regional Director
U.S. Fish and Wildlife Service
P.O.Box 1306
Albuquerque. New Mexico 87 1 03
REGION 3
Regional Director U.S. Fish and Wildlife Service Federal Building, Foit Snelling Twin Cities. Minnesota 55111
REGION 4
Regional Director U.S. Fish and Wildlife Service Richard B. Russell Building 75 Spring Street, S.W. Atlanta, Georgia 30303
REGION 5
Regional Director
U.S. Fish and Wildlife Service
One Gateway Center
Newton Corner, Massachusetts 02 1 58
REGION 6
Regional Director
U.S. Fish and Wildlife Service
P.O. Box 25486
Denver Federal Center
Denver, Colorado 80225
REGION 7
Regional Directoi U.S. Fish and Wildlife Service 101 1 E.Tudor Road Anchorage, Alaska 99503
DEPARTMENT OF THE INTERIOR
U.S. FISH AND WILDLIFE SERVIDE
As the Nation's principal conservation agency, the Department of the Interior has respon- sibility for most of our nationally owned public lands and natural resources. This includes fostering the wisest use of our land and water resources, protecting our fish and wildlife, preserving the-environmental and cultural values of our national parks and historical places, and providing for the enjoyment of life through outdoor recreation. The Department as- sesses our energy and mineral resources and works to assure that their development is in the best interests of all our people. The Department also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S. administration.