The Ecology of the Mangroves of South Florida: Acommunity Profile

FWS/OBS-81/24 January 1982 THE ECOLOGY OF THE MANGROVES OF SOUTH FLORIDA: ACOMMUNITY PROFILE

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Bureau of Land Management Fish and Wildlife Service U.S. Department of the Interior FWS/OBS-81/24 January 1982 Reprinted September 1985

THE ECOLOGY OF THE MANGROVES OF SOUTH FLORIDA: A COMMUNITY PROFILE

by William E. Odum Carole C. t·1cIvor Thomas J. Smith, 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 Slidell, 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 OISCLAmER The findings in this report are not to be construed as an official U.S. Fish and Wildlife Service position unless so designated by other authorized documents.

Library of Congress Card Number 82-600562

This report should be cited:

Odum, W.E., C.C. McIvor, and T.J. Smith, III. 1982. The ecology of the mangroves of south Florida: a conmunity profile. U.S. Fish and Wildlife Service, Office of Biological Services, Washington. D. C. FWS/OBS-81/24. 144 pp. Reprinted September 1985. PREFACE

This profile of the mangrove commun­ and Tarpon Springs on the west. Refer­ ity of south Florida is one in a series ences are provi ded for those seeki ng of community profil es whi ch treat coas ta1 in-depth treatment of a specific facet of and marine habitats important to man. The mangrove ecology. The format, style, and obvious work that mangrove communities do level of presentation make this synthes is for man includes the stabilization and report adaptable to a diversity of needs protection of shorelines; the creation and such as the preparation of environmental maintenance of habitat for a oreat number ass ess IT.ent reports, s upp 1ementa ry readi ng of animals, many of which -are either in marine science courses, and the devel­ enda ngered or ha ve commerci alva1ue; and opment of a sense of the importance of the provis ion of the bas is of a food web this resource to those citizens who whose final products include a seafood control its fate. smorgasbord of oys ters, crabs, lobs ters, shrimp, and fish. Less tangible but Any ques ti ons or comments about or equa lly important benefi ts incl ude wil der­ requests for this publication should be ness, aes thetic and 1ife support cons i der­ directed to: ations.

The information on these pages can Information Transfer Specialist give a basic understanding of the mangrove National Coastal Ecosystems Team community and its role in the regional U.S. Fish and Wildlife Service ecosystem of south Florida. The primary NASA-Slidell Computer Complex geographic area covered lies along the 1010 Gause Boulevard coas t between Cape Canavera1 on the eas t Slidell, Louisiana 70458

iii iv CONTENTS

PREFACE ...... •...... iii FIGURES viii TABLES vi i i ACKNOWLEDGMENTS i x

CHAPTER 1. INTRODUCTION .

1.1 "Mangrove" Definition...... 1 1.2 Factors Controll ing Mangrove Distribution...... 1 1.3 Geographical Distribution 2 1.4 Mangrove Species Descriptions .. , 5 1.5 Mangrove Communi ty Types 7 1. 6 Substrates 9 1.7 Wa ter Quality ...... •...•.... 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. ECOSYSTHl STRUCTURE Arm FUNCTION 26

3.1 Structural Properties of flangrove 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 - PLAinS OTHER THAN t1ANGROVES 41

5.1 Root and Mud Algae 41 5.2 Phytoplankton...... 43 5.3 Associated Vascular Plants 43

v CONTENTS (continued)

Page

CHAPTER 6. COM~IUNITY COI·IPONENTS - INVERTEBRATES 45

6.1 Ecological Relationships ...... •...... 45 6.2 Arboreal Arthropod Community...... 47 6.3 Prop Root and Associated Mud Surface Conrnunity 47 6.4 Water Column Community...... 49

CHAPTER 7. cOr~rt,UNITY COMPONENTS - FISHES 50

7.1 Basin r~angrove Forests...... 50 7.2 Riverine Forests. 52 7.3 Fringing Forests along Estuarine Bays and Lagoons 54 7.4 Fringing Forests along Oceanic Bays and Lagoons 56 7.5 Overwash Mangrove Islands 56 7.6 Gradient of 1·langrove Community Interactions 57

CHAPTER 8. COMMUNITY COMPONENTS - AMPHIBIANS ANO REPTILES ...... •.....•.... 58

CHAPTER 9. COMMUNITY CO~1PONENTS - 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 COf·1PONENTS - MAMMALS ..•...... •.•...... 72

CHAPTER 11. VALUE OF MANGROVE ECOSYSTEMS TO MAN ...... •...... 74

11 .1 Shoreline Stabilization and Storm Protection . 74 11 .2 Habitat Value to Wildlife .. 74 11.3 Importance to Threatened and Endangered Species ...... •...... 75 11. 4 Value to Sport and Commercial Fisheries .. 75 11.5 Aesthetics, Tourism and the Intangibles ...... •...... 75 11.6 Economi c Products ...... •...... •...... 76

CHAPTER 12. MANAGEMENT IMPLICATIONS...... 77

12.1 Inherent Vul nerabil ity . 77 12.2 Man-induced Destruction . 77 12.3 Effects of Oil Spill s 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 ~Ianagement Practices: Preservation .. 86 vi CONTENTS (continued)

Page REFERENCES 88 APPENDIX A: SUMMARY OF SITE CHARACTERISTICS AND SAMPLING METHODOLOGY FOR FISHES 106 APPENDIX B: FISHES OF MANGROVE AREAS 110

APPENDIX C: AMPHIBIANS AND REPTILES FROM MANGROVE AREAS 127 APPENDIX D: AVIFAUNA OF MANGROVE AREAS 130

APPENDI X E: i4AM/o1ALS OF 14ANGROVE AREAS 142

vi i FIGURES Number Approximate northern limits for the red mangrove (R), black mangrove (B), and "hite 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 propagu1es, flowers and leaves. 6 4 The six mangrove community types...... 8 5a Aboveground and be10wground biomass of a Puerto Rican red mangrove forest 16 5b Light attenuation in a mangrove canopy; canopy height is 8m 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, producti vi ty, and nutri ent export from mangrove ecosystems 31 8 Potential pathways of energy flow in mangrove ecosystems 37 g Vertical distribution of selected algae and invertebrates on red mangrove prop roots ...... •...... 42 10 Photograph of red mangrove prop root habitat in clear shallow "ater 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 Whitewater Bay 53 13 The mangrove water snake, Nerodia fasciata compressicauda, curled on a red mangrove prop root 59 14 A variety of wading birds feeding in a mangrove-lined pool near F1 ami ngo, Flori da ...... •.... 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 1·langrove forest near Key West as it appeared in 1981 after being destroyed by diking and impounding 79 18 ~Iangrove 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

viii ACKNOWLEDGMENTS

Many individuals and organizations individuals provided information as well contributed significantly to the creation as critical comments. We particularly of this publication. Most notable was appreciate the unselfish help of Ken Adams Eri c Hea 1d who worked extens i ve1y with us and the staff of the National Coastal Eco­ on the manuscript and contributed unpub­ systems Team of the U.S. Fish and Wildlife lished data. We thank Jeffrey Carlton, Service. Janet Ryan s pent many hours typ­ Edward Conner, Roy R. Lewis, Ill, Ariel ing the manuscript. Lugo, Larry Narcisse, Steven Macko, Aaron Mills, Michael Robb1ee, Martin Roessler, Factual errors and faulty conclu­ Samuel Snedaker, Durbin Tabb, Mike Wein­ sions are the sole reseonsibi1ity of the stein, and Joseph Zieman for information authors. Carole McIvor has taken primary and helpful advice. res pons i bi 1ity for Chapter 7, Tom Smith • for Chapters 8, 9, and 10, and Bi 11 Odum The draft manuscript was reviewed for for the remainder of the publication. its scientific content by Armando de 1a Un 1ess otherwis e noted, photographs, Cruz, Thomas Savage, James Kush1an (bird figures, and the cover were produced by section), and James P. Ray. Each of these the authors.

ix x CHAPTER 1 INTRODUCTION

1.1 "MANGROVE" OEFINITION temperatures below freezing for any length of time. Certain species, for example, The term UmangroveU expresses two black mangrove, Avicennia germinans, on distinctly different concepts. One usage the northern coastoT-the-GiiTIcifMexi co, refers to halophytic species of trees and maintain a semi-permanent shrub form by shrubs (halophyte = plant growing in growing back from the roots after freeze saline soil). In this sense, mangrove is damage. a catch-all, botanically diverse, non­ taxonomic expression given to approximate­ Lugo and Zucca (1977) di scuss the ly 12 families and more than 50 species impact of low temperature stress on Flori­ (Chapman 1970) of tropi cal trees and da mangroves. They found that mangrove shrubs (see Wai sel 1972 for a detai 1ed communities respond to temperature stress list). While not necessarily closely by decreasing structural complexity (de­ related, all these plants are adapted to creased tree height, decreased leaf area (1) loose, wet soils, (2) a saline habi­ index, decreased leaf size, and increased tat, (3) periodic tidal submergence, and tree density). They concluded that man­ (4) usually have degrees of viviparity of groves growing under conditions of high propagules (see section 2.3 for discussion soil salinity stress are less tolerant of of "viviparity" and "propagules"). low temperatures. Presumably, other types of stress (e.g., pollutants, diking) could The second usage of the term mangrove reduce the temperature tolerance of man­ encompasses the entire plant community groves. including individual mangrove species. Synonymous terms include tidal forest, High water temperatures can al so be tidal swamp forest, mangrove community, limiting. McMillan (1971) reported that mangrove ecosystem, man gal (Macnae 1968), seedlings of black mangrove were killed by and mangrove swamp. temeP.eratures of 39 0 to 40 0 C (102 0 to 104 F) although established seedlings and For consistency, in this pub1 ication trees were not damaged. To our knowledge, we wi 11 use the word "mangrove" for i ndi­ upper temperature tolerances for adult vidual kinds of trees; mangrove community, mangroves are not well known. We suspect mangrove ecosystem or mangrove forest will that water temperatures in the range 42 0 represent the entire assemblage of "man­ to 450 C (1070 to 113 0 F) may be limiting. groves".

Salt Water 1.2 FACTORS CONTROLLING MANGROVE DISTRI­ BUTION Mangroves are facultative halo­ phytes, i.e., salt water is not a physical Four major factors appear to limit requirement (Bowman 1917; Egler 1948). In the distribution of mangroves and deter­ fact, most mangroves are capable of mine the extent of mangrove ecosystem growing quite well in freshwater (Teas development. These factors include (1) 1979). It is important to note, however, climate, (2) salt water, (3) tidal fluc­ that mangrove ecosystems do not develop in tuation, and (4) substrate. strictly freshwater environments; salinity is important in reducing competition from other vascular plant species (Kuenzler Climate 1974). See section 2.2 about salinity tolerance of mangrove species. Mangroves are tropical species and do not develop satisfactorily in regions where the annual average temperature is Tidal Fluctuation below IgOC or 660 F (Waisel 1972). Normally, they do not tolerate temperature While tidal influence is not a fluctuations exceeding IOoC (l8°F) or direct physiological requirement for 1 mangroves, it plays an important indirect the east coast of Africa, in Australia and role. Fi rst, tidal stress (alternate in New Zealand, they extend 100 to 150 wetting and drying), in combination with farther south (Kuenzler 1974) and in salinity, helps exclude most other Japan, Florida, Bermuda, and the Red Sea vascular plants and thus reduces competi­ they extend 50 to 70 farther north. These tion. Second, in certain locations, tides areas of extended range generally occur bring salt water UP the estuary against where oceanographic conditions move un­ the outward flow of freshwater and allow usua lly warm water away from the equator. mangroves to become establ i shed well inland. Third, tides may transport Althou9h certai n reqi ons such as the nutrients and relatively clean water into tropical Indo-Pacific have as many as 30 mangrove ecosystems and export accumula­ to 40 species of mangroves present, only tions of organic carbon and reduced sulfur three species are found in Florida: the compounds. Fourth, in areas with high red mangrove, Rhi zophora mangl e, the b1 ad evaporation rates, the action of the tides mangrove, Avicennia germinans, and the helps to prevent soil salinities from white mangrove, Laguncul ari a racemosa. A reaching concentrations which might be fourth species, buttonwood, Conocar us lethal to mangroves. Fifth, tides aid in erecta, is not a true mangrove no ten­ the dispersal of mangrove propagules and dency to vivipary or root modification}, detritus. but is an important species in the transi­ tion zone on the upland edge of mangrove Because of all of these factors, ecosystems (Tom1 inson 1980). termed tidal subsidies by E.P. Odum (1 971), mangrove ecosystems tend to reach The ranges of mangrove speci es in thei r greatest de vel opment around the Florida have fluctuated over the past world in low-lying regions with relatively several centuries in response to relative­ 1arge ti da1 ranges. Other types of water ly short-term climatic change. Currently, fluctuation, e.g., seasonal variation in the situation is as follows (Figure 1). freshwater runoff from the Florida Ever­ The red mangrove and the white mangrove glades, can provide similar subsidies. 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 Substrate and Wave Energy on the east coast (Teas 1977); both of these extremes lie at approximately 2g 0 10' Mangroves grow best in depositional N latitude. Significant stands lie south envi ronments wi th low wave energy. Hi gh of Cape Canaveral on the east coast and wave energy prevents establ i shment of Tarpon Springs on the west coast. The propagules, destroys the relatively shal­ black mangrove has been reported as far low mangrove root system and prevents the north as 30 0 N latitude on the east coast accumulation of fine sediments. The most of Flori da (Savage 1972) and as scattered productive mangrove ecosystems develop shrubs along the north coast of the Gulf along deltaic coasts or in estuaries that of Hexi co. have fi ne-gra i ned muds composed of si It, c1 ay and a hi gh percentage of organic matter. Anaerobic sediments pose no Intertidal Distribution problems for mangroves (see section 2.1) and exclude competing vascular plant The generalized distribution of the species. red and black mangrove in relation to the intertidal zone is shown in Figure 2a. Local variations and exceptions to this 1.3 GEOGRAPHICAL DISTRIBUTION pattern occur commonly in response to localized differences in substrate type Mangroves dominate approximately 75% and elevation, rates of sea level rise, of the world's tropical coastline between and a variety of other factors (see sec­ 25 0 N and 25 0 S 1at itude (McGi 11 1959). On t ion 3.2 for a fu 11 di scuss i on of mangrove

2 --____ I,------. , i I .~ _____ 8

____ R,W R,W ~·· Ponce de Leon Inlet Cedar Key

Tarpon Ri ver

Tampa Bay

Port Charlotte Harbor/ Sanibel Island ;.

.-,. Rookery Bay"'" } .,' I Ten Thousand Islands .. Biscayne Bay Shark River Valley. .' I Whitewater Bay/North Ri ver ::: ,.~'? ,. Flori da Keys •• " .."~ ,

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.

3 (A)

MHW-----

...... :: .. '. '."': :~ :.:. MLW--- PEAT \__-.-• ..._/''- --..,..- ----''...., -.J/ LOW HIGH MARSH MARSH (RED MANGROVE (BLACK MANGROVE

DOMINATED) DOMINATED)

(B)

J FMAMJJASOND

+10 em

+5 em

SEA LEVEL

-5 em

-10 em

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) .

4 zonation). Furthermore, it is important According to this survey, ninety percent to recognize that the intertidal zone in of Flori da's mangroves are located in the most parts of Florida changes seasonally four southern counties of Lee (35,000 (Provost 1974); there is a tendency for acres or 14,000 hal. Collier (72,000 acres sea level to be higher in the fall than in or 29,000 hal, Monroe (234,000 acres or the spring (Figure 2b). As a result the 95,000 hal, and Dade (81,000 acres or "high marsh" may remain totally dry during 33,000 hal. the spring and be continually submerged in the autumn. This phenomenon further com­ Much of the area covered by mangroves plicates the textbook concept of the in­ in Florida is presently owned by Federal, tertidal, "low marsh" red mangrove and the State or County governments, or by non­ infrequently flooded, "high marsh" black profit organizations such as the National mangrove. Audubon Society. Approximately 280,000 acres (113,000 hal fall into this category (Eric Heald, personal communication 1981). Mangrove Acreaqe in Florida Most of this acreage is held by the Federal Governf'lent as a result of the land Estimates of the total acreage being including within the Everglades occupied by mangrove communities in National Park. Florida vary widely between 430,000 acres and over 500,000 acres (174,000 ha to over 202,000 hal. Eric Heald (Tropical Bioindustries, 9869 Fern St., Miami, Fla.; 1.4 MANGROVE SPECIES DESCRIPTIONS personal communication 1981) has identified several reasons for the lack of The following descriptions come agreement between estimates. These largely from Carlton (1975) and Savage include: (1) inclusion or exclusion in (1972); see these publications for further surveys of small bays, ponds and creeks comments and photographs. For more which occur within mangrove forests, (2) detailed descriptions of germinating seeds incorrect identification of mangrove areas (propagules) see section 2.3. The three from aeri al photography as a result of species are shown in Figure 3. inadequate "ground-truth" observations, poorly contro11 ed ae ri a 1 photography, and simple errors of planimetry caused by The Black Mangrove (Avicennia ~erminans) photography of inadequate scale. Avicennia germinans is synonymous The two most detailed estimates of with ---,o;:-iiTITda aiid---rsa- me mbe r 0f the area covered by mangroves in Florida are family Avicenniaceae (formerly classed provi ded by the Coastal Coordi nati ng Coun­ under Verbenaceae). The tree may reach a cil, State of Florida (1974) and 8irnhak height of 20 m (64 ft) and has dark, scaly and Crowder (1974). Considerable dif­ bark. Leaves are 5 to 10 cm (2 to 4 ferences exi st between the two esti mates. inches) in length, narrowly elliptic or The estimate of Birnhak and Crowder oblong, shiny green above and covered with (1974), which is limited to certain areas short, dense hai rs below. The 1eaves are of south Florida, appears to be unrealis­ frequently encrusted with salt. Thi s tree tically high, particularly for Monroe is characteri zed by long hori zontal or County (Eric Heald, personal communication "cabl e" roots with short vertical aerating 1981). Coastal Coordinating Council branches (pneumatophores) that profusely (1974) estimates a total of 469,000 acres penetrate the substrate below the tree. (190,000 hal within the State and suggests Propagul es are 1i ma-bean shaped, dark an expected margin of error of 15% (i.e. green while on the tree, and several thei r esti mate lies between 400,000 and centimeters (1 inch) long. The tree 540,000 acres or 162,000 and 219,000 hal. flowers in spring and early summe~

5 Red Mangrove, Rhizophora mangle

~-~ Black ~~~grove, Avicennia germi nans

White Mangrove, Laguncularia racemosa

Figure 3. Three species of Florida mangroves with propagules, flowers, and leaves.

6 The White Mangrove (Laguncularia racemosa) range, and flushing rates. Each community type has cha racteri s tic ranges of pri rna ry The white mangrove is one of 450 production, litter decomposition and car­ species of plants in 18 genera of the bon export along with differences in family Combretaceae (synonymous with nutri ent recycl i ng rates, and community Terminaliaceae). It is a tree or shrub components. The community types as shown reaching 15 m (49 ft) or more in height in Figure 4 are as follows: with broad, flattened oval leaves up to 7 cm (3 inches) long and rounded at both ends. There are two salt glands at the (1) Overwash mangrove forests apex of the petiole. The propagule is these islands are frequently overwashed by very small (1.0 to 1.5 cm or 0.4 to 0.6 tides and thus have ~igh rates of organic inches long) and broadest at its apex. export. All species of mangroves may be Flowering occurs in spring and early present, but red mangroves usually domi­ summer. nate. Maximum height of the mangroves is about 7 m (23 ft).

The Red Mangrove (Rhizophora mangle) (2) Fri nge mangrove forests - man­ groves form a relatively thin fringe along The red mangrove is one of more than waterways. Zonation is typically as de­ 70 species in 17 genera in the family scribed by Davis (1940) (see discussion in Rhizophoraceae. This tree may reach 25 m section 3.2). These forests are best (80 fti in height, has thin grey bark and defined along shorelines whose elevations dark red wood. Leaves may be 2 to 12 cm are hi gher than mean hi gh ti de. Maxi mum (1 to 5 inches) long, broad and bl unt­ height of the mangroves is about 10 m (32 poi nted at the apex. The 1eaves are ft) • shiny, deep green above and paler below. It is easily identified by its charac­ (3) Ri veri ne mang rove fores t s - th is teristic ·prop roots' arising from the community type includes the tall flood trunk and branches. The pencil-shaped plain forests along flowing waters such as propagules are as much as 25 to 30 cm (10 tidal rivers and creeks. Although a shal­ to 12 inches) long after germination. It low berm often exists along the creek may flower throughout the year, but in bank, the entire forest is usually flushed Florida flowering occurs predominately in by daily tides. All three species of the spring and early summer. mangroves are present, but red mangroves (with noticeably few, short proo roots) predomi nate. Mangroves may reach hei ghts 1.5 MANGROVE COMMUNITY TYPES of 18 to 20 m (60 to 65 ft).

Mangrove forest communities exhibit (4) Basin mangrove forests - these tremendous variation in form. For forests occur inland in depressions chan­ example, a mixed scrub forest of black and neling terrestrial runoff toward the red mangroves at Tu rkey Poi nt on Bi scayne coast. Close to the coast they are in­ 8ay bears little resemblance to the fluenced by daily tides and are usually luxuriant forests, dominated by the same dominated by red mangroves. Moving in­ two species, along the lower Shark River. lanrl, the tidal influence lessens and dominance shifts to hlack and white man­ Lugo and Snedaker (1974) provided a groves. Trees may reach 15 m (49 ft) in convenient classification system based on hei ght. mangrove forest physi ogomy. They i denti­ fied six major community types resulting (5) Hammock forests - hammock man­ from different geological and hydrological grove communities are similar to the basin processes. Each type has its own charac­ type except that they occur on ground that teristic set of environmental variables is slightly elevated (5 to 10 cm or 2 to 4 such as soil type and depth, soil sa1i nity inches) relative to surrounding areas.

7 (1) OVERWASH FOREST (2) FRINGE FOREST

(3) RIVERINE FOREST (4) BASIN FOREST

(5) HAMMOCK FOREST (6) SCRUB FOREST

Figure 4. The six mangrove community types (Lugo and Snedaker 1974).

8 All species of mangroves may be present. becomes too great, mangroves wi 11 not be Trees rarely exceed 5 m (16 ft) in hei ght. present. Of the three species of Florida mangroves, white mangroves appear to (6) Scrub or dwarf forests - this tolerate sandy substrates the best (per­ community type is limited to the flat sonal observation), possibly because this coastal fringe of south Florida and the species may tolerate a greater depth to Florida Keys. All three species are the water table than the other two present. Individual plants rarely exceed species. 1.5 m (4.9 ft) in height, except where they grow over depressions filled with Mangroves in Florida often modify the mangrove peat. Many of these tiny trees underlying substrate through peat deposi­ are 40 or more years of age. Nutrients tion. It is not unusual to find layers of appear to be limiting although substrate mangrove peat several meters thick under­ (usually limestone marl) must playa role. lying well-established mangrove ecosystems such as those along the southwest coast of Throughout this publ ication we have Florida. Cohen and Spackman (1974) pre­ attempted to refer to Lugo and Snedaker's sented a detailed account of peat forma­ classifi cation scheme wherever possi b1e. tion within the various mangrove zones of Without a system of this type, comparisons south Florida and also in areas dominated between sites become virtually by black needle rush (Juncus roemerianus), meaningless. smooth cordgrass (Spartina alterniflora) and a variety of other macrophytes; Cohen and Spackman (1974) also provide descrip­ 1.6 SUBSTRATES tions and photography to aid in the iden­ tification of unknown peat samples. Understand i ng mangrove-substrate relationships is complicated by the The following descriptions come from ability of mangroves to grow on many types Cohen and Spackman (1974) and from the of substrates and because they often alter personal ohservations of W.E. Odum and the substrate through peat formation and LJ. Heald. Red mangroves produce the by alterin9 patterns of sedimentation. As most easily recogni zed peat. More recent a result, mangroves are found on a wide deposits are spongy, fi brous and composed variety of substrates including fine, to a great extent of fi ne root1ets (0.2 to inorganic muds, muds with a high organic 3.0 mm in diameter). Also present arp. content, peat, sand, and even rock and larger pieces of roots (3 to 25 mm), hits dead coral if there are sufficient of wood and leaves, and inorganic crevices for root attachment. Mangrove matp.rials such as pyrite, carbonate ecosystems, however, appear to fl ouri sh minerals, and quartz. Older deposits are only on muds and fine-grained sands. less easily differentiated although they remain somewhat fibrous. Peat which has In Florida, the primary mangrove recent ly been excavated is reddi sh-brown soils are either calcareous marl muds or although this changes to brown-black after calcareous sands in the southern part of a short exposure to air. Older deposits the State and siliceous sands farther are mottled reddish-brown; deposits with a north (Kuenzler 1974). Sediment distribu­ high content of carbonates are greyish­ tion and, hence, mangrove development, is brown upon excavat ion. controlled to a considerable extent by wave and current energy. Low energy Cohen and Spackman (1974) were unable shorelines accumulate fine-grained sedi­ to find deposits of pure black mangrove or ments such as mud and silt and usually white mangrove peat suggesting that these have the best mangrove growth. Higher two species may not form extensive depos­ energy shorelines (more wave action or its of peat whil e growi ng in pu re stands. higher current velocities) are charac­ There are, however, many examples of peats teri zed by sandy sedi ments and 1ess pro­ which are mixtures of red mangrove ductive mangroves. If the wave energy material and black mangrove roots. They

9 suggested that the black mangrove peats values of -100 to -400 mv in mangrove identified hy Davis (1946) were probably peats. Such evidence of strongly reducing mixtures of peat from several sources. conditions are not surprising considering the fine-grained, high organic nature of Throughout south Florida the sub­ most mangrove sediments. Although man­ strate underlying mangrove forests may groves occur in low organic sediments cons; st of compl i cated patterns of (less than 1% organic matter), typical calcareous muds, marls, shell, and sand values for mangrove sediments are 10% to interspersed and overlain by layers of 20% organic matter. mangrove peat and with 1i mestone bedrock at the bottom. Detailed descriptions of Lee (1969) analyzed 3,000- to 3,500­ this complex matrix and its spatial varia­ year-old mangrove peat layers underlying tion were given by Davis (1940, 1943, Little Black Water Sound in Florida Bay 1946), Egler (1952), Craighead (1964), for lipid carbon content. Peat lipid Zieman (1972) and Cohen and Spackman content varied between 0.6 and 2.7 109 (1974) among others. Scoffin (1970) di s­ lipid-C/gram of peat (dry wt) or about 3% cussed the ability of red mangrove to of the total organic carbon total. These trap and hold sediments about its prop values usually increased with depth. Long roots. So called "land-building" by man­ chain fatty acids (C-16 and C-18) were the 9roves is discussed in section 3.2. domi nant fatty aci ds found.

The long-term effect of mangrove peat Florida mangrove peats are usually on mangrove distribution is not entirely acidic, although the presence of carbonate clear. Certainly, if there is no change materials can raise the pH above 7.0. in sea level or if erosion is limited, the Zi em an (1972) found red mangrove peats to accumul ati on of peat under stands of red range from pH 4.9 to 6.8; the most acid mangroves combined with deposition and conditions were usually found in the cen­ accumulation of suspended sedi ments wi 11 ter of the peat 1ayer. Lee (1969) re­ raise the forest floor sufficiently to corded a pH range from 5.8 to 6.8 in red lead to domination by black or white man­ man9rove peat at the bottom of a shallow groves and, ultimately, more terrestrial embayment. Althou9h Davis (1940) found a species. Whether this is a common se­ di fference between red mangrove peat (5.0 quence of events in contemporary south to 5.5) and bl ad mangrove peat (6.9 to Florida is not clear. It is clear that 7.2), this observation has not been con­ peat formtion is a passive process and firmed because of the previously mentioned occurs primari ly where and when physical diffi culty in fi ndi ng pu re black man9rove processes such as erosion and sea level peat. rise are of minimal importance (Wanless 1974) • Presumably, the acidic character of mangrove peat results from release of Zieman (1972) presented an inter­ organic acids during anaerobic decomposi­ esting argument suggesting that mangrove tion and from the oxidation of reduced peat may be capabl e of di ssol vi ng under­ sulfur compounds if the peat is dried in lying limestone rock, since carbonates may the presence of oxygen. This last point dissolve at pH 7.8. Through this process, explains why "reclaimed" mangrove areas shallow depressions might become deeper often develop highly acidic soils (pH 3.5 and the overlyi ng peat layer thicker to 5.0) shortly after reclamation. This without raising the surface of the forest "cat clay" problem has greatly complicated f1 oor. the conversion of mangrove regions to agricultural land in Africa and southeast Data on chemical characteristics of Asia (Hesse 1961; Hart 1962, 1963; r~acnae Florida mangrove soils and peat are 1968) . limited. Most investigators have found man9rove substrates to be almost totally In summary, although current under­ anaerobic. Lee (1969) recorded typical Eh standing of mangrove peats and soil sis

10 fragmentary and often contradictory, we from virtually fresh water to above 40 ppt can outline several generalizations: (discussed in section 2.2), (2) low macro­ nutrient concentrations (particularly (1) Mangroves can grow on a wide phosphorous), (3) relatively low dissolved variety of substrates including mud, sand, oxygen concentrations, and (4) frequently rock, and peat. increased water color and turbidity. The last three characteristics are most pro­ (2) Mangrove ecosystems appear to nounced in extensive mangrove ecosystems flourish on fine-grained sediments which such as those adjacent to the Everglades are usua 11 y anae robi c and may ha ve a hi gh and least pronounced in small, scattered organic content. forests such as the overwash islands in the Flori da Keys. (3) Mangrove ecosystems which per­ si st for some ti me may modify the under­ Walsh (1967), working in a mangrove lying substrate through peat formation. swamp in Hawai i, was one of the fi rst to This appears to occur only in the absence document the tendency of mangrove eco­ of strong physi cal forces. systems to act as a consumer of oxygen and a sink for nutrients such as nitrogen and (4) Mangrove peat is formed pri­ phosphorous. Carter et al. (1973) and mari 1y by red mangroves and consi sts pre­ Lugo et al. (1976) confi rmed these obser­ dominantly of root material. vations for Florida mangrove swamps. Evi­ dently, nutrients are removed and oxygen (5) Red mangrove peats may reach consumed by a combination of periphyton on thicknesses of several meters, have a mangrove prop roots, mud, organic detritus relatively low pH, and may be capable of on the sediment surface, the fine root dissolving underlying layers of limestone. system of the mangroves, small inverte­ brates, benthic and epiphytic algae, and (6) When drained, dried, and bacteria and fun9i on all these surfaces. aerated, mangrove soils usually experience dramatic increases in acidity due to the The results of oxygen depletion and oxidation of reduced sulfur compounds. nutrient removal are (1) dissolved oxygen This greatly complicates their conversion concentrations below saturation, typically to agri cul t ureo 2 to 4 ppm and often near zero in stagnant locations and after heavy, storm-generated runoff, (2) very low total phosphorus 1.7 WATER QUALITY values, frequently below detection limits, and (3) moderate total nitrogen val ues Water quality characteristics of sur­ (0.5 to 1.5 mg/l). In addition, TOC face waters flowing through Florida man­ (total organic carbon) may range from 4 to grove ecosystems exhibit great variation 50 ppm or even hi gher after rai n; Eri c from one location to the next. Proximity Heald (personal communication 1981) has to terrestri al ecosystems, the ocean, and measured DOC (dissolved organic carbon) human activities are all important in values as high as 110 ppm in water flowing determining overall water quality. from mangroves to adjacent bays. Tur­ Equally important is the extent of the bidity usually falls in the 1 to 15 JTU mangrove ecosystem since drastic altera­ (Jackson turbity units) range. The pH of tions in water quality can occur within a the water column in Florida swamps is stand of mangroves. usually between 6.5 and 8.0 and alkalinity between 100 to 300 mg/l. Obviously, ex­ In general, the surface waters ceptions to all of these trends can occu~ associ ated with mangroves are charac­ Both natural and human disturbance can terized by (1) a wide range of salinities raise macronutrient levels markedly.

11 CHAPTER 2. AUTECOLOGY OF MANGROVES

2.1 AOAPTATIONS TO NATURAL STRESS - roots to provide a more or less firm foun­ ANAEROBIC SEDIMENTS dation for the tree. Even though the prop roots are anchored with only a modest Mangroves have a series of remarkable assemblage of underground roots, the hori­ adaptations which enable them to flourish zontal extent of the prop root system in an environment characterized by high insures considerable protection from all temperatures, wi dely f1 uctuatin g sa1i ni ­ but the worst of hurri canes. Other man­ ties, and shifting, anaerobic substrates. grove species, including the black man­ In this section we review a few of the grove, obtai n stabil i ty wi th an extens i ve most important adaptat ions. system of shallow, underground "cable" roots that radiate out from the central The root system of mangroves provides trunk for a considerable distance in all the key to existence upon unfriendly sub­ di rection s; the pneumatophores extend up­ strates (see Gill and Tomlinson 1971 for ward from these cable roots. As in all an anatomical review of mangrove roots). Florida mangroves, the underground root Unlike most higher plants, mangroves system is shallow and a tap root is usually have hi ghly developed aeri al roots lacking (Walsh 1974). As Zieman (1972) and modest below-ground root systems. The found, individual roots, particularly of aerial roots allow atmospheric gases to red mangroves, may extend a meter or more reach the underground roots whi ch are downward in suitable soils. embedded in anaerobic soils. The red mangrove has a system of stilt or prop From the standpoi nt of effecti veness roots which extend a meter (3 ft) or more in transporting oxygen to the underground above the surface of the soil and contain roots, both prop roots and cable roots many small pores (lenticels) which at low seem equally effecti ve. From the perspec­ tide allow oxygen to diffuse into the tive of stability, the prop roots of red plant and down to the underground roots by mangroves appear to offer a distinct ad­ means of open passages called aerenchyma vantage where wave and current energies (Scholander et al. 1955). The lenticels are high. are highly hydrophobic and prevent water penetration into the aerenchyma system Unfortunately, as pointed out by Odum during high tide (Waisel 1972). and Johannes (1975), the same structure which allows mangroves to thri ve in an­ The black mangrove does not have prop aerobic soil is also one of the tree's roots, but does have small air roots or most vulnerable components. Exposed por­ pneumatophores which extend vertically tions of the aerial root system are sus­ upward from the underground roots to a ceptible to clogging by fine suspended hei ght of 20 to 30 cm (8 to 12 inches) material, attack by root borers, and pro­ above the soi 1. These pneumatophores longed flooding (discussed further in resemble hundreds of tiny fingers sticking section 12.1). Such extended stress on up out of the mud underneath the tree the aeri al roots can ki 11 the enti re tree. canopy. At low ti de, ai r travel s through the pneumatophores into the aerenchyma system and then to all living root tis­ 2.2 ADAPTATIONS TO NATURAL STRESS ­ sues. The whi te mangrove usually does not SALINITY have either prop roots or pneumatophores, but utilizes lenticels in the lower trunk Manqroves accommodate fluctuations and to obtai n oxygen for the aerenchyma sys­ extremes of water and soil salinity tem. "Peg roots" and pneumatophores may through a variety of mechanisms, although be present in certain situations (Jenik not all mechanisms are necessarily present 1967). in the same species. Scholander et al. (1962) reported experimental evidence for Mangroves achieve structural stabili­ two major methods of internal ion regula­ ty in at least two ways. Species such as tion which they identified in two dif­ the red mangrove use the system of prop ferent groups of mangroves: (1) the salt 12 exc1 usion species and (2) the salt excre­ There appears to be some variation in tion species. In addition, some mangroves the salinity tolerance of Florida man­ utilize succulence and the discarding of groves. The red mangrove is probah1y salt-laden organs or parts (Teas 1979). limited b.y soil salinities above 60 to 65 PPt. Teas (l979) recalculated Bowman's The salt-excluding species, which (1917) data and conc1 uded that transpi ra­ include the red mangrove, separate tion in red mangrove seedlings ceases freshwater from sea water at the root above 65 ppt. Cintron et al. (1978) found surface by means of a non-metabolic ultra­ more dead than 1i vi ng red mangrove trees filtration system (Scho1ander 1968). This where interstitial soil salinities ex­ "reverse osmosis" process is powered by a ceeded 65 ppt. high negative pressure in the xylem which results from transpiration at the leaf On the other hand, white and black surface. Salt concentration in the sap of mangroves, which both possess salt excre­ salt-excluding mangroves is about 1/70 the tion and limited salt exclusion mech­ salt concentration in sea water, although anisms, can exist under more hypersaline this concentration is almost 10 times conditions. Macnae (1968) reported that higher than found in normal plants b1 ack mangroves can grow at soil sal i ni­ (Scholander et al. 1962). ti es greater than 90 ppt. Teas (1979) reported dwarfed and gnarled black and Salt-secreting species, including white mangroves occurring in Florida at bl ack and white mangroves (Scho1 ander soil salinities of 80 ppt. 1968), use salt glands on the leaf surface to excrete excess salt. This is probably There may be an additional factor or an enzymati c process rather than a phys i­ factors involved in salinity tolerance of cal process since it is markedly tempera­ mangroves. McMi 11 an (1 975) found that ture sensitive (Atkinson et al. 1967). seedlings of black and white mangroves The process appears to i nvol ve acti ve survived short-term exposures to 80 ppt transport with a requirement for biochemi­ and 150 ppt sea water if they were grown cal energy input. As a group, the salt in a soi 1 with a moderate c1 ay content. secreters tend to have sap salt concentra­ They failed to survive these salinities, tions approximately 10 times higher (1/7 however, if they were grown in sand. A the concentration of sea water) than that soil with 7% to 10% clay appeared to be of the salt excluders. adequate for increased protection from hypersaline conditions. In spite of these two general tenden­ cies, it is probably safe to say that Vegetation-free hypersaline lagoons individual species utilize a variety of or bare sand flats in the center of man­ mechanisms to maintain suitable salt grove ecosystems have been described by balance (Albert 1975). For example, the many authors (e.g., Davis 1940; Fosberg red mangrove is an effecti ve, but not 1961; Bacon 1970). These features have perfect, salt excluder. As a result this been variously called sa1itra1s (Holdridge species must store and ultimately dispose 1940), salinas, salterns, salt flats, and of excess salt in leaves and fruit (Teas salt barrens. Evidently, a combination of 1979). Most salt secreters, including low seasonal rai nfa11, occasi ona1 i nunda­ white and black mangroves, are capable of tion by sea water, and high evaporation limited salt exclusion at the root sur­ rates results in soil salinities above 100 face. The white mangrove, when exposed to ppt, water temperatures as high as 450 C hypersaline conditions, not only excludes (l13°F) in any shallow, standing water, some salt and secretes excess salt through and subsequent mangrove death (Teas 1979). its salt glands, but also develops Once estab1i shed, sal i nas tend to persi st thickened succulent leaves and discards unless regular tidal flushing is enhanced sa1t du ri ng 1eaf fall of senescent 1eaves by natural or artificial changes in tidal (Teas 1979). circulation.

13 A1thou~h salinas occur frequently in probably accounts for the low fertiliza­ Florida, they are rarely extensive in tion rate, estimated by Gill and Tomlinson area. For example, between Rookery Bay at 0% to 7.2%. Propagule development is and Marco Island (south of Naples, slow, rangi ng from 8 to 13 months. Savage Florida) there are a series of salinas in (1972) mentions that on the Florida gulf the bl ack mangrove-domi nated zone on the coast, red mangrove propagules mature and upland side of the mangrove swamps. These fall from the tree from July to September. hypersaline lagoons occur where the normal Within the Everglades National Park, black flow of fresh water from upland sources ",angroves flower from May unti 1 July and has been diverted, presumably resulting in bear fruit from August until November elevated soil salinities during the dry while white mangroves flower from May to wi nter months. August and bear fruit from July to October (Loope 1980). In summary, sal inity is a problem for mangroves only under extreme hypersa1 i ne The propagu1es of the three species conditions. These conditions occur natu­ of Florida mangroves are easy to differen­ rally in Florida in irregularly flooded tiate. The following descriptions all areas of the "high swamp" above the normal come from Rabinowitz (1978a). ,Ihite man­ high tide mark and are accompanied by high grove propagules are small and flattened, soi 1 sal i nities. Florida mangroves, weigh less than a 9ram, are about 2 em listed in order of increasing salinity long, are pea-green when they fall from tolerance, appear to be red, white, and the parent tree, and turn mud-brown in two black. days or so. The peri carp (wall of the ripened propagu1e) serves as a float and is not shed until the seedling is estab­ 2.3 REPRODUCTIVE STRATEGIES lished. During dispersal the radicle (embryonic root) emerges from the propa­ As pointed out by Rabinowitz (197Ba), gule. This germination during dispersal vi rtually all mangroves share two common has 1ed Savage (1972) to refer to the reproductive strate9ies: dispersal by white mangrove as I'semi-viviparous". means of water (van der Pi j 1 1972) and vivipary (Macnae 1968; Gill and Tomlinson The propagul es of the b1 ack mangrove 1969). Vivipary means that the embryo when dropped from the tree are ob10ng­ develops continuously while attached to elliptical (resemble a flattened olive), the parent tree and during dispersal. wei gh about 1 g and are about 2 em long. Since there is uninterrupted development The peri carp is lost within a few days from zygote through the embryo to seed1 i ng after dropping from the tree; at this without any intermediate resting stages, point the cotyledons (primary leaves) the "ord "seed" is inappropriate for unfold and the propagule resembles two viviparous species such as mangroves; the butterfl ies on top of one another. term "propagu1e" is generally used in its place. Propagules of the red mangrove under­ go extensive vivipary while on the tree. "hi1e the phenology of black and When propagu1es fall from the tree they white mangroves remains sketchy, Gill and resemble large green beans. They are rod­ Tomlinson (1971) thoroughly described the shaped with pointed ends, about 20 em sequence of flowering in the red mangrove. long, and wei gh an average of 15 g. Flowering in this species may take place at any time of the year, at least in Propagules of all three species float extreme south Florida, but reaches a maxi­ and remain viable for extended periods of mum in the late spring and summer. The time. Apparently, there is an obligate flowers open approximately 1 to 2 months di spersal ti me for all Flori da mangroves, after the appearance of buds. The flower i.e., a certain period of time must elapse remains intact only 1 to 2 days; this during dispersal for germination to be

14 complete and after which seedling estab­ concluded that species with small lishment can take place. Rabinowitz propagules establish new cohorts annually (1978a) estimates the obligate dispersal but die rapidly, while species such as the period at approximately 8 days for white red mangroves may have long-lived and mangroves, 14 days for black, and 40 days often overl appi ng cohorts. for red. She further estimates the addi­ tional time for root establishment at 5, Propagule size and seedling mortality 7, and 15 days for white, bl ack, and red rates are particularly important in con­ mangroves, respect i ve 1y. Her esti mate for siderations of succession and replacement viable longevity of the propagules is 35 in established mangrove forests. Light is days for white mangroves and 110 days for usually the most serious limiting factor black. Davis (1940) reports viable propa­ underneath existing mangrove canopies. gul es of red mangroves that had been kept Rabinowitz (1978b) suggested that species floating for 12 months. with short-lived propagules must become establ ished in an area which al ready has Rabinowitz (1978a) al so concluded adequate light levels either due to tree that black and white mangroves require a fall or some other factor. In contrast, stranding period of 5 days or more above red mangrove seedlings can become estab­ the influence of tides to take hold in the lished under an existing, dense canopy and soi 1. As a result, these two species are then, due to thei r superi or embryoni c usually restricted to the higher portions reserves, are able to wait for months for of the mangrove ecosystem where tidal tree fall to open up the canopy and pre­ effects. are infrequent. sent an opportunity for growth.

The elongated red mangrove propagule, however, has the potential to become 2.4 BIOMASS PARTITIONING established in shallow water with tidal influence. This happens in at least two Few investigators have partitioned ways: (1) stranding in a vertical posi­ the total biomass, aboveground and below­ tion (they float vertically) or (2) ground, contained in a mangrove tree. An stranding in a horizontal position, analysis of red mangroves in a Puerto rooting and then vertical erection by the Rican forest by Golley et al. (1962) gives plant itself. Lawrence (1949) and Rabino­ some insight into what might be expected witz (l978a) felt that the latter was the in south Florida. Aboveground and below­ more common method. M. Walterding (Cal if. ground biomass existed in a ratio of 1:1 Acad. Sci., San Francisco; personal com­ if fine roots and peat are ignored (Figure munication 1980) favors vertical estab­ 5). In this case, peat and very fine lishment; based upon his observations, roots (small er than 0.5 cm di ameter) ex­ surface water turbulence works the propa­ ceeded remaining biomass by 5:1. Lugo et gule into the substrate during fallin9 a1. (1976) reported the following values tides. for a south Florida red mangrove overwash forest. All values were reported in dry Mortality of established seedlings grams per square meter, plus and minus one seems to be related to propagule size. standard error, and i gnori ng bel owgr~und Working in Panama, Rabinowitz (1978b) biomass. They found 7~0 ! 22 g/m of found that the mortal ity rate of mangrove leave-.s, l2.8 ! 15.3 g/m of propagu1es, seedlings was inversely correlated with 7043 - 7 g/m2 of wood, 4695 ! ~ll g/m of initial propagule size. The white man­ prop roots and 1565 : 234.5 g/m of detri­ grove, which has the smallest propagule, tus on the forest floor. has the highest rate of seedling mortal­ ity. The black mangrove has an interme­ Biomass partitioning between dif­ diate mortality rate while the red man­ ferent species and locations must be grove, with the largest propagule, has the highly variahle. The age of the forest lowest seedling mortality rate. She will influence the amount of wood biomass;

15 (A)

LEAVES (778)

BRANCHES (1274)

SOIL TRUNK (2796) '------,----;-...,..-;::--::-=--r------J ------SURFACE LARGE ROOTS (997) SMALL ROOTS (4000)

(B)

8

6 HEIGHT

(M) 4

2

OLL...-...... L.-...... ----l_...... ~_...... __J o 5000 10000 FOOT CANDLES

Figure 5. (a) Aboveground and belowground biomass of a Puerto Rican red mangrove forest. Values in parentheses are dry g/m2; large roots = 2 cm+ in diameter, small roots = 0.5 - 1.0 em. (b) Vertical distribution of light intensity in 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 of mangroves in Florida. Since that time to the next dependi ng upon the amou nt of knowledge has accumulated rapidly, but it fluvial transport. The biomass charac­ is still unrealistic to expect more than teristics of a scrub forest probably bear preliminary statements about Florida man­ little resemblance to those of a fringing grove productivity. This deficiency can forest. At the present time, there is not be traced to (1) the difficulties asso­ enough of this type of data available to ciated with measurements of mangrove pro­ draw many conclusions. One intriguing ductivity and (2) the variety of factors point is that red mangrove leaf biopass that affect productivity and the resulting averages between 700 and 800 g/m at variations that exist from site to site. various sites with very different forest morphologies (Odum and Heald 1975a). This Productivity estimates come from may be related to the tendency of mangrove three methods: (1) harvest, (2) gas ex­ canopies, once they have become estab­ change, and (3) 1itter fall. Harvest lished, to inhibit leaf production at methods require extensive manpower and lower levels through self-shading. knowledge of the age of the forest. They are best employed in combination with Go11 ey et al. (1962) showed that the silviculture practices. Since silvicul­ red mangrove canopy is an extremely effi­ ture of south Florida mangroves is practi­ cient light interceptor. Ninety-five cally non-existent, this method has rarely percent of the available light had been been used in Florida. Noakes (1955), intercepted 4 m (13 ft) below the top of Macnae (1968), and Walsh (1974) should be the canopy (Figure 5). As a result, 90% consulted for productivity estimates based of the leaf biomass existed in the upper 4 on this technique in other parts of the m of the canopy. Chlorophyll followed the worl d. same pattern of distribution. Gas exchange methods, based on The leaf area index (LAl) of mangrove measurements of C02. changes, have the forests tends to be relatively low. Go1­ advantage of precislon and response to ley et al. (1962) found a LA! of 4.4 for a short-term changes in light, temperature, Puerto Rican red mangrove forest. Lugo et and flooding. They include both above­ a1. (1975) reported a LA! of 5.1 for a ground and belowground production. On the Florida black mangrove forest and 3.5 for negative side, the necessary equipment is a Flori da fri nge red mangrove forest. A expensive and tricky to operate properly. different black mangrove forest, in Flori­ Moreover, extrapolations from short-term da, was found to have values ranging from measurements to long-term estimates offer 1 to 4 and an average of 2 to 2.5 (Lugo considerable opportunity for error. and Zucca 1977). These values compare Nevertheless, the best estimates of pro­ with LA!'s of 10 to 20 recorded for most ductivity come from this method. tropical forests (Golley et al. 1974). The low leaf area values of mangrove The litter fall technique (annual forests can be attributed to at least 1itter fall x 3 = annual net pri mary pro­ three factors: (1) effective light inter­ duction) was proposed by Teas (1979) and ception by the man9rove canopy, (2) the is based on earl i er papers by Bray and inability of the lower mangrove leaves to Gorham (1964) and Goll ey (1972) for other flourish at low light intensities, and (3) types of forests. This is a quick and the absence of a 1ow-1 i ght-adapted p1 ant di rty method although the 1ack of pre­ layer on the forest floor. ci si on remai ns to be demonstrated for mangroves. An even quicker and dirtier method proposed by Teas (1 979) is to (1) 2.5 PR! MAR Y PRODUCTION estimate leaf standing crop (using various techniques including harvesting or 1i9ht Prior to 1970 virtually no informa­ transmission re1ationsbips) and (2) multi­ tion existed concerning the productivity ply by three. Thi s assumes an annual leaf

17 turnover of one, which is supported by the inputs of toxic compounds or nutrients data of Heald (1969) and Pool et al. from human activities (1977). human influences such as diking, Mangrove producti vity is affected by ditching, and altering patterns of many factors; some of these have been runoff. reco9nized and some remain totally ob­ scure. Carter et al. (1973) propose lumping these factors into two broad cate­ In spite of the difficulties with gories: tidal and water chemistry. We various methods and the interaction of believe that a number of additional cate­ controlling factors, it is possible to 90ries should be considered. make general statements about certai n aspects of mangrove producti vity. For A minimal, though incomplete, list of example, Waisel's (1972) statement that factors controlling mangrove productivity mangroves have low transpiration rates must include the following: seems to be genera lly true in Flori da. Lugo et al. (1975) rep~rted transpiration species composition of the stand rates of 2,500 g H20/m /day for mangrove leaves in a frin~,ng red mangrove forest age of the stand and 1,482 g H20/m /day for b1 ack mangrove leaves. This is approximately one-third presence or absence of competing to one-half the value found in temperate species broad leaf forests on hot dry days, but comparable to tropical rainforests (H.T. ~egree of herbivory Odum and Jordan 1970). The low transpi ra­ tion rates of mangroves are probably re­ presence or absence of disease and lated to the energetic costs of main­ parasi tes taining sap pressures of -35 to -60 atmo­ spheres (Scholander et a1. 1965). depth of substrate Litter fall (leaves, twigs, bark, substrate type fruit, and flowers) of Florida mangrove for'2sts appears to average 2 to 3 dry nutrient content of substrate g/m day in most well-developed mangrove stands (see discussion in section 3.4). nutrient content of overlying water This can be an order of magnitude lower in scrub forests. sa1i nity of soil and overlying water Wood production of mangroves appears transport efficiency of oxygen to root to be hi gh compared to other temperate and system tropical trees, although no measurements from Florida are available. Noakes (1955) amount of tidal flushing estimated that the wood production of an intensively managed Malayan forest was relative wave energy 39.7 metric tons/ha/year. Teas (1979) suggested a wood production estimate of 21 presence or absence of nesting birds metric tons/ha/year for a mature unmanaged red mangrove forest in south Florida. His periodicity of severe stress (hurri­ figure was calculated from a litter/total canes, fire, etc.) biomass relationship and is certainly subject to error. time since last severe stress Representative estimates of gross characteristics of ground water primary producti on (GPP) net pri mary

18 product i on (NPP), and respi ration (R) of These three apparent tendencies have Florida mangroves are given in Table lao led Carter et al. (1973) and Lugo et al. Compared to net pri mary production (NPP) (1976) to propose a fourth tendency, an estimates from other ecosystems, including inverted U-shaped relationship between agricultural systems (E.P. Odum 1971), it waterway position and net mangrove com­ appears that mangroves are among the munity productivity (Figure 6). This world's most productive ecosystems. tendency is best understood by visualizing Healthy mangrove ecosystems appear to be a typi cal gradi ent on the southwest coast more productive than sea grass, marsh of Florida. At the landward end of the grass and most other coastal systems. gradient, salinities are very low, nutrient runoff from terrestrial eco­ Further examination of Table la re­ systems may be high and tidal amplitude is veals several possible tendencies. The minor. At the seaward end, salinities are first hypothetical tendency, as discussed relatively high, tidal amplitude is rela­ by Lugo et al. (1975), is for red man9roves tively great and nutrient concentrations to ha ve the hi ghest tota1 net production, tend to be lower. At either end of the black to have intermediate values and gradient, the energetic costs are high and white the lowest. This conclusion assumes a 1a rge percentage of GPP is used for that the plants occur within the zone for se1 f-maintenance; at the landward end, which they are best adapted (see section competition from freshwater plant species 3.2 for discussion of zonation) and are is high and at the seaward end, salinity not existing in an area with strong limit­ stress may be limiting. In this scenario, ing fa,tors. A scrub red mangrove forest, the highest NPP occurs in the middle for example, growing under stressed condi­ region of the gradient; salinity and tidal tions (high soil salinity or low nutrient amplitude are high enough to limit compe­ supply), has relatively low net produc­ tition while tidal flushing and moderate tivity (Teas 1979). The pre-eminent posi­ nutrient levels enhance productivity. tion of red mangroves is shown by the Hicks and Burns (1975) present data to comparative measurements of photosynthesis support thi s hypothesi S. in Table Ib; measurements were made within canopy leaves of trees growing within their zones of optimal growth. In addition to these hypotheses A second noteworthy tendency is that generated from field data, there have been red mangrove GPP decreases with increasing two significant, published attempts to salinity while GPP of black and white deri ve hypotheses from mathemat i cal si mu­ mangroves increases with increasing lation models of mangroves. The first salinity up to a point. Estimates of Hicks (Mi ller 1972) is a model of primary pro­ and Burns (1975) demonstrate that this may duction and transpi ration of red mangrove be a real tendency (Table Ie). canopies and is based upon equations which utilize field measurements of the energy Data presented by Miller (1972), budgets of individual leaves. This model Carter et al. (1973), Lugo and Snedaker predicts a variety of interesting trends (1974), and Hicks and Burns (1975) sug­ which need to be further field tested. gest a third hypothetical tendency, One interesting hypothesis generated by assuming occurrence of the species within the model is that maximum photosynthesis its adapted zone. It appears that the of red mangrove stands should occur with a black mangrove typically has a much higher leaf area index (LAI) of 2.5 if no accli­ respi ration rate, lower net producti vity, mation to shade within the canopy occurs; and lower GPPjR ratio than the red man­ higher LAI's may lead to decreased produc­ grove. This can be attributed at least tion. Another prediction is that red partially, to the greater salinity stress mangrove production is most affected by under which the black mangrove usually air temperature and humidity and, to a grows; this leads to more osmotic work. lesser degree, by the amount of solar

19 Table lao ~stimates of man9rove production in Florida. All values are gc/m 2/day except annual NPP = metric tons/ha/yr. GPP = gross primary production, NPP = net primary jJroduction, L.F. = annual litter fall X 3, R = red mangrove, W = white mangrove, B = black mangrove. Observations 6 and 7 were on sunny days, 0 and 9 on cloudy days.

Species GPP Respiration NPP Annual NPP 11ethod Reference

Nixed R, 24.0 11 . 4 12 .5 46.0 Gas exchange Hicks & Burns (1975) w, a

B 18.U 12.4 5.6 20.5 Ga s exchange Lugo & Snedaker (1974) a a Mature ~ 8.8 20.5 L. F. Teas (1979) a a Scrub R 1 . a 3.8 L. F. Teas (1979) a a Basin B 2 .4 8.6 L. F. Teas (1979)

N R (June) 12.8 7.3 5.5 20.3 Gas exchange Miller (1972) <:> K (Jan. ) 9.4 5 . 1 4.3 15. 7 Gas exchange Miller (1972) R (June) 10.3 6.8 3.5 12.8 Gas exchange Miller (1972) R (Jan.) 10.2 5. a 5.2 18.8 Gas exchange Miller (1972) Mixed R,W, 13. 9 9. 1 4.8 17.5 Gas exchange Carter et a1 . (1973) B(riverine) Mixed R,W , 11.8 4.3 7.5 27.4 Gas exchange Carter et a1 . (1973) B(basin) B 9.0 6.2 2.8 9.4 Gas exchange Lugo et a 1 . (1976) R 6.3 1 .9 4.4 16 . 1 Gas exchange Lugo et a 1 . (1976)

aMethod does not produce this data. Table lb. Comparative measurements of photosynthesis in gC/m 2/day (Lugo et al. 1975).

Mangrove type Daytime net Nighttime photosynthesis respiration

Red 1 .38 0.23 6.0 81ack 1. 24 0.53 2.3

Wh i te 0.58 0.17 3.4 Red (seedling) 0.31 1 .8 g negative

Table lc. Gross primary production (GPP) at different salinities (Hicks and Burns 1975).

Mangrove type Average surface GPP salinity (ppt) (gC/m 2/day)

Red 7.8 8.0 Red 21 . 1 3. g Red 26.6 1 .6 Black 7.8 2.3 Black 21 . 1 5.7 Black 26.6 7.5 White 21.1 2.2 White 26.6 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 lepidopterans (moths and butterfl ies), and photosynthesis per unit leaf area was orthopterans (grasshoppers and crickets). greater at the top of the tree canopy than at the bottom, although the middle levels Heald (1969) estimated a mean grazing had the greatest production. effect on North Ri ver red mangrove 1eaves of 5.1% of the total leaf area; values Miller (1972) concluded by suggesting from leaf to leaf were highly variable that the canopy distribution of red man­ rangi ng from 0 to 18%. Beever et al. grove leaves is nearly optimal for ef­ (1979) presented a detailed stuny of ficient water utilization rather than grazi ng by the mangrove tree crab. Thi s production. This indicates that the cano­ arboreal grapsid crab feeds on numerous py is adapted to maximizing production items including beetles, crickets, cater­ under conditions of saturated water sup­ pillars, littoral algae, and dead animal ply. matter. In Florida, red mangrove leaves form an important component of the di et. The mangrove ecosystem model reported Beever et al. (1979) measured tree crab by Lugo et al. (1976) provides hypotheses grazing ranging from 0.4% of the total on succession, time to arrive at steady leaf area for a Florida Keys overwash state conditions (see section 3.2)' and forest to 7.1% for a fringing forest at several aspects of productivity. The Pine Island, Lee County, Florida. The model output suggests that the relative researchers also found that tree crab amount of tidal amplitude does not affect grazing rates are related tg crab density. GPP significantly; instead, GPP appears to Low densities (one crab/m ) resulted in be extremely sensitive to inputs of ter­ low leaf area damage (less than 1% of restrial nutrients. It follows that loca­ total l'3af area). High densities (four tions with large amounts of nutrient input crabs/m ) were accompanied by leaf area from terrestrial sources (riverine man­ damage ranging from 4% to 6% (see section grove communi ties) ha ve hi gh rates of 6.2) • mangrove production (see section 3.3). All si mul ation model-generated hypotheses Onuf et al. (1977) investigated in­ need to be field tested with a particular­ sect berbivory in fringing and overwash ly critical eye, since the simplifying red mangrove forests in the Indian River assumptions that are made in constructing estuary near Ft. Pierce, Florida. They the model can lead to overly simplistic found si x major herbi vorous insect answers. species, five lepidopteran larvae and a beetle. Comparisons were made at a bigh Mangrove productivity research re­ nutrient site (input from a bird rookery) mains in an embryonic stage. Certain and a low nutrient site. 80th red man­ preliminary tendencies or hypotheses have grove production and leaf nitrogen were been identified, but much work must be significantly bigher at the high nutrient done before we can conclude that these site. This resulted in a four-fold hypotheses cannot be fa1sifi ed. greater loss to berbivores (26% of total leaf area lost to grazing); this increased grazing rate more than offset the in­ 2.6 HERBIVORY creased leaf production due to nutrient input. Oirect herbivory of mangrove leaves, leaf buds, and propagules is moderately Calculations of leaf area damage may low, but highly variable from one site to underestimate the impact of berbivores on the next. Identified grazers of living mangroves. For example, the larvae of tbe plant parts (other than wood) include the olethreutid moth, Ecdytolopha sp., white-tailed deer, Odocoileus virginianus, develops within red mangrove leaf buds and the mangrove tree crab, Aratus pisonii, causes the loss of entire leaves. All and insects including beetles, larvae of s tag e s 0 f the bee t 1 e , Poe c !..l2.P.~

23 rhizophorae, attack mangrove propagules a result of Sphaeroma infestation; this while-still attached to the parent tree poi nt was not documented. (Onuf et al. 1977). Enri ght (1974) produced a tongue-in­ cheek rebuttal, on behalf of Sphaeroma and li 2.7 WOOO BORERS against the t1terrestrial invacter , red mangroves. Snedaker (1974) contributed a Many people have the mistaken idea more substantial argument in which he that mangrove wood is hi ghly resi stant to pointed out that the isopod infestation marine borers. Whi Ie this may be true to might be an example of a long-term eco­ ali mited extent for certain mangrove system control process. species in other parts of the world, none of the Florida mangroves have borer­ Further arguments against the '~coca­ resistant wood. Southwell and BoHman tastrophe" theory were advanced by Estevez (1971) found that the wood of red, black, and Simon (1975) and Estevez (1978). They and white mangroves has no resistance to provi ded more 1i fe hi story i nformati on for Teredo, Pholad and Simnorid borers; pieces Sphaeroma and suggested a possible ex­ Of-red ma-ngrove wood were completely de­ planation for the apparently destructive stroyed after immersion in ocean water for isopod infestations. They found two 14 months. species of isopods inhabiting red mangrove prop roots, S. terebrans and a sympatri c An interesting controversy surrounds congener, ~.guadridentatum. The latter the ability of the wood boring isopod, does not appear to be a wood borer but Sphaeroma terebrans, to burrow into the utilizes S. terebrans burrows. Neither living prop roots of the red mangrove. species appeared to utilize mangrove ·wood Rehm and Humm (1 973) were the fi rst to as a food source. Estevez and Si man attribute apparently extensi ve damage of (1975) found extensive burrowing into red mangroves stands within the Ten seedlings in addition to prop root damage. Thousand Islands area of southwestern In general, infestations appeared to be Florida to an isopod, Sphaeroma. They patchy and limited to the periphery of found extensive damage throughout mangrove ecosystems. In areas with the southwest Florida, some infestation north highest density of burrows, 23% of all to Tarpon Springs, and a total lack of prop roots were infested. There appeared infestation in the Florida Keys from Key to be more colonization by S. terebrans in Largo south to Key Ilest. The destruct; on regions with full strengtilsea water (30 process was described as follows: the to 35 ppt). adult isopod bored into the prop roots (5­ mm diameter hole); this was followed by The most important finding by Estevez reproduction within the hole and develop­ and Si mon (1975) and Estevez (1978) was ment of juveni 1es withi n the root. Thi s that periods of accelerated activity by S. process, combi ned wi th secondary decompo­ terebrans were related to periods of flu~ sition from fungi and bacteria, frequently tuating and slightly increased salinity. results in prop root severance near the This suggests that fluctuations in isopod mean high tide mark. These authors burrowing may be related to the magnitude attributed loss of numerous prop roots of freshwater runoff from the Everglades. and, in some cases, loss of entire trees These authors a9ree with Snedaker (1974) during storms to isopod damage. and suggest that root and tree loss due to Sphaeroma activity may be beneficial to The extent of damage in the Ten mangrove ecosystems by accelerating pro­ Thousand Islands region led Rehm and Humm duction and root germination. Simber10ff (1973) to term the phenomenon an "eco­ et a1. (1978) amplified this last sugges­ catastrophe" of possibly 9reat importancL tion by showing that root branching, which They fu rther stated that sh ri nk i ng of is beneficial to individual trees, is mangrove areas appeared to be occurring as stimulated bY isopod activity.

24 This ecocatastrophe versus beneficial fungi, Phyllosticta hibiscina and Nigro­ stimulus argument is not completely re­ spora sphaerica. These authors found that solved. Probably, Sphaeroma root destruc­ P. hibiscina caused necrotic lesions and tion, in areas of low isopod density, can death of black mangrove 1eaves. They fe 1t be a beneficial process to hoth the in­ that under conditions of high relative dividual tree and to the entire mangrove humidity coupled with high temperatures, stand. Whether changes in freshwater this fungus could pose a serious threat to runoff have accelerated this process to individual trees, particularly if the tree the point where unnatural and widespread had been weakened by some other natural damage is occurri ng is not cl ear. The agent, such as 1i ghtni ng or wi nd damage. data and research perspective to answer Nigrospora sphaerica was considered to be this question do not exist. As a result, of little danger to black mangroves. we are reduced to providing hypotheses Another fungus, Cylinrocarpon ~~~~~~, which cannot be tested with availahle appears to form galls on the prop roots knowledge. and stems of red mangroves. Olexa and Freeman (1975) noted mortality of red mangroves in areas of hi gh gall infesta­ 2.8 MANGROVE 0[SEASES tions, although a direct causation link was not proven. Publ i shed research on mangrove diseases is rare. The short paper by Further research on mangrove diseases Olexa and Freeman (1975) is the principal is hadly needed. Viral disease must be reference for diseases of Florida man­ investigated. The role of pathogens in groves. They reported that black man­ litter production and as indicators of groves are affected by the pathogenic mangrove stress may be very important.

25 CHAPTER 3. ECOSYSTEM STRUCTURE AND FUNCTION

3.1 STRUCTURAL PROPERTI ES OF MANGROVE tree height to be inversely proportional FORESTS (r = 0.72) to soil salinity in the range 30 to 72 ppt. Above 65 ppt sal inity, dead Published information about the tree basal area was higher than live tree structural aspects of Florida mangrove basal area and above 90 ppt there was no forests is limited; most existing data live tree basal area. have been published since the mid-1970's. This lack of information is unfortunate It should be possible to investigate since quantitative structural data greatly the relationship between a variety of aid understanding of processes such as mangrove structural properties and factors succession and primary production. Even such as flushing frequency, soil depth, more important, the response of mangrove nutrient availability, pollution stress, forests to stress, both climatic and man­ and other measures of human impact. Ulti­ induced, can be followed quantitatively mateI y, thi s shou 1d 1ead to an abil ity to with this type of data. predict the form and structure of mangrove forests resulting from various physical Ball (19BO) contributed substantially conditions or artificial impacts. One to understandin9 the role of competi­ example of this potential tool is Ball's tion in man9rove succession by measurin9 (19BO) documentation of structural changes structural factors such as basal area, in mangrove forests resulting from altera­ tree hei9ht, and tree density. Lugo and tions in the hydrological conditions of Zucca (1977) monitored the response of south Florida. mangrove forests to freezing temperatures by observing changes in structural proper­ ties of the trees. 3.2 ZONATION, SUCCESSION AND "LAND- BUILDING" Baseline studies of forest structure have been published by Lugo and Snedaker Much of the worl d's mangrove litera­ (1975), and Pool, Snedaker and Lugo ture consists of descriptive accounts of (1977). For exampl e, LU90 and Snedaker zonation in mangrove forests and the spe­ (1975) compared a fri ngi ng mangrove forest cies composition within these zones. Al­ and a basin forest at Rookery Bay, near thouqh general agreement has been lacking, Naples, Florida. They found the fringing various hypotheses have been put forth forest, which was dominated by red man­ concerning the possible connection between groves, to have a tree diversi~y of H = zonation, ecological succession, competi­ 1.4B, a basal area of 15.9 m ;Sha, an tion, and the role of physical factors aboveground biomass of 17,932 glm , and a such as soil salinity and tidal amplitude. non-existent litter layer. The nearby In this section we review briefly the basin forest was dominated by black man­ dominant ideas about mangrove zonation and groves, had a tree divers~ty of H = 0.96 succession and present our interpretation and a basal area of 23.4 m Iha. The lit­ of the current status of knowl edge. ter lay~ in the basin forest averaged 550 dry g/m. Tree diversity in a hurricane Davis (1940), working in south Flori­ disturbed sect i on of the Rookery Bay da, was one of the first investigators to forest was 1.62. Si mi 1ar data were pre­ describe distinct, almost monospecific, sented for mangrove forests in the Ten zones within mangrove ecosystems. In what Thousand Islands area (Table 2). has become the classical view, he argued tha+, mangrove zonation patterns were Oata of this type are useful for many equivalent to seral stages in succession. purposes including impact statements, en­ The most seaward zone, domi nated by red vironmental surveys, and basic scientific mangroves, was regarded as the "pioneer questions. Cintron et al. (197B) gave an stage". More landward zones were indication of the di rection in which fu­ dominated by white mangrove, black ture research might proceed. Working in a mangrove, buttonwood and, fi nally, the mangrove stand in Puerto Rico, they found climatic climax, a tropical forest. Since 26 Table 2. Aboveground biomass of mangrove forests in the Ten Thousand Islands region of Florida. Values are based on 25 m2 clearcuts and are expressed in dry kg/ha. Data are from Lugo and Snedaker (1975).

Compartment Scrub Overwash Fringe Riverine mangroves mangroves mangroves mangroves

Site A B A B CA B

Leaves 712 7 ,263 6 ,946 5 ,932 5,843 7,037 3,810 9,510

Fru it & flowers no data 20 236 28 210 131 148 N '" Wood 3,959 70,380 70,480 57 , 96 0 84,270 128,510 79,620 161,330

Prop roots 3 ,197 51 ,980 41,920 22,270 27,200 17,190 14,640 3,060

Litter 1 ,140 17,310 13,990 22,730 60,250 98,410 42,950 33,930 ------To ta1 above- 9,008 146,953 133,572 108,920 177,773 251,278 141,168 207,831 .round biomass these zones were regarded as progressively (1975) found that black mangroves and later stages in succession, the entire white mangroves along with the sa1tmeadow mangrove ecosystem was bel i eved to be cordgrass, Spartina patens, are often the movi ng seaward through a process of sedi­ pioneers on new dredge spoil islands in ment accumulation and colonization. Davis central Florida. On the northern coast of based his argument primarily upon the the Gulf of Mexico, where black mangrove sequence of observed zones and cores which is the only mangrove species present, it showed red mangrove peat underlying black may be preceded by marsh grasses such as mangrove peat which, in turn, occurred saltmarsh cordgrass, 2. patens, smooth under terestrial plant communities. cordgrass, S. alternif10ra, or the black needl e rush,-Juncus roemeri anus. I n Puer­ Unfortunately, this Clementsian in­ to Ri co, we ohserved that whi te mangrove terpretation of mangrove zonation was often pi oneers and dominates sites where widely accepted, but rarely tested. For oceanic overwash of heach sand has oc­ example, Chapman (197D) expanded Davis' curred. All of these ohservati ons detract original successional concept from south from Davis' (1940) original contention Florida to explain zonation in mangrove that red mangroves should be regarded as forests in other parts of the world. the initial colonizer of recently de­ \,a1sh (1974) thoroughly reviewed the man­ posited sediments. It appears that under grove succession/zonation literature. certain conditions, e.g., shallow water depths, suhstrate type, and latitude, Fortunately, not everyone accepted white and black mangroves or marsh grasses Davis' point of view. Egler (1952) and can be effective pioneer species. later Thorn (1967, 1975) argued that man­ 9rove zonation was a response to external The work of Rabinowitz (1975) added a physical forces rather than temporal se­ new perspective to the mangrove zonation quence induced by the pl ants themsel ves. debate. Through carefully designed recip­ E91er (1952) showed that patterns of sedi­ rocal planting experiments in Panamanian ment deposition predicted by Davis' (1940) mangrove forests using species of Rhizo­ theory did not always occur. He also ~'!.' .!:.'!._9.,~r!.cularia, Pelliciera and showed that in some cases mangrove zones Avicennia, she demonstrated that each appeared to be moving landward rather than species could grow well within any of the seaward. Sea level has been rising in mangrove zones. In other words, physical south Florida at the rate of 1 ft (30 ern) and chemi cal factors such as soi 1 sal i nity per 100 to 150 years (Provost 1974). or frequency of tidal inundation, within Spackman et a1. (1966) emphasized the role each zone, were not solely responsible for of sea 1eve1 change in determ i ni ng changes excluding species from that zone. To in mangrove zonation, both through sea explain zonation, Rabinowitz proposed level rise and land subsidence. Both tidal sorting of propagu1es based upon Egler (1952) and Spackman et a1. (1966) propagule si ze, rather than habitat adap­ along with Wanless (1974) and Thom (1967, tation,as the most important mechanism for 1975) suggested that mangroves were zonation control. reacting passively rather than actively to strong geomorphological processes. This The most recent pi ece to be added to implies that mangroves should be regarded the zonation/succession puzzle comes from as "1 and-stabi 1i zers" rather than "1 and­ the work of Ball (1980). Based upon re­ builders". search of mangrove secondary succession patterns adjacent to Biscayne Bay, Flori­ Furthermore, field researchers fre­ da, she made a strong case for the impor­ quent ly noted that red mangroves were not tance of interspecific competition in always the only "pioneer species" on re­ controlling zonation. She found that cently deposited sediment. It is not white mangroves, which grow best in unusual to find seedlings of black, white, intertidal areas, do not occur consis­ and red mangroves growing together on a tently in the intertidal zone of mature new colonization site. Lewis and Dunstan mangrove stands. Instead, white manqroves

28 dominate higher, drier locations above frequently observed white mangroves mean high water where the red mangrove flourishing in small lightning-created does not appear to have a competitive openings in the center of red mangrove advantage. She suggested that competition forests. Fire may also playa role in is not so important during the early limiting the inland spread of manQroves. stages of successi on but becomes criti cal Taylor (1981) pointed out that Everglades as indi vidual trees reach maturity and fires appear to prevent the encroachment requi re more space and other resources. of red and white mangroves into adjacent herbaceous communities. Inherent in Ball's concept of zona­ tion is the differential influence of Finally, Lugo and Snedaker (1974), physical factors (e.g., soil salinity, Cintron et al. (1978) and Lugo (1980) depth to water table) on the competitive suggested that mangrove ecosystems abi 1it i es of the di fferent mangrove function as classical successional systems species. She concluded that succession in areas of rapid sediment deposition or proceed s independent 1y withi n each zone, upon recently colonized sites such as althougn breaks in the forest canopy from offshore islands. They concluded that in lightning strikes or high winds may pro­ most areas mangrove forests are an example duce a mosaic of different successional of steady-state cyclical systems. Concep­ stages withi n a zone. These openi ngs tually, this is synonymous to E. P. Odum's allow species whose seedlings do not com­ (1971) cyclic or catastrophic climax. pete well in shade, such as the white Chapnan (lg76a, b) suggested the idea of mangrove, to become established, at least cyclic succession for a variety of coastal temporarily, within solid zones of red ecosystems. mangroves. [f Florida mangrove ecosystems are Zonation of mangrove species does not cyclic systems, then there should be an appear to be controlled by physical and identifiable perturbation capable of set­ chemical factors di rect1y, but by the ting succession back to an early stage. interplay of these factors with interspe­ Lugo and Snedaker (1974) suggested that cific competition and, possibly, through hurricanes may play this role. They tidal sorting of propagu1es. Once succes­ pointed out (without substantiating data) sion in a mangrove zone reaches an equili­ that major hurricanes occur about every brium state, change is unlikely unless an 20-25 years in south Florida. Coinci­ external perturbation occurs. These per­ dently, mangrove ecosystems appear to turbations range from small-scale distur­ reach their maximum levels of productivity bance (lightning strikes) to large-scale in about the same period of time (Lugo and perturbations (sea level change, hurricane Snedaker 1974). This hypothesis suggests damage) and may cause succession within that succession within many mangrove eco­ zones to regress to an earlier stage. systems may proceed on a cyclical basis There is some evidence in south Florida rather than in the classical fashion. that hurricane perturbations occur on a Possibly other physical perturbations may fairly regular basis, creating a pattern influence mangrove succession including of cyclical succession. incursions of freezing temperatures into central Florida, periodic droughts causing Except for Ball (1980) and Taylor unusually high soil salinities (Cintron et (1980), the importance of fires as an al. 1978), and fire spreading into the i nfl uence on mangrove success i on has been upper zones of mangrove forests from ter­ genera11y ignored. Most fi res in the restrial sources. Florida mangrove zone are initiated by lightning and consist of small circular Although understanding of zonation openings in the mangrove canopy (Taylor and succession in mangrove ecosystems 1980). These openi ngs present an opportu­ remains incomplete, a clearer picture is nity for secondary succession within an emerging, at least for south Florida. established zone. For example, we have Contrary to early suggestions, mangrove

29 species zonation does not appear to repre­ Although mangrove ecosystems may tend sent seral stages of succession except, to accumulate nutrients, there is a con­ perhaps, for locations of recent coloniza­ tinual loss through export of particulate tion or where sediment is accumulating and di ssol ved substances. If si gni fi cant rapidly. The role of mangroves in nutrient storage and resultant high pri­ land-building seems more passive than mary production are to occur, there must act i ve. Geomo rphologi ca 1 and hydro1ogi ca1 he a continual input of nutrients to the processes appear to be the dominant forces mangrove forest from outside the system in determining whether mangrove shorelines (Figure 7). Where nutrient influx to the recede or grow. The role of mangroves is mangrove ecosystem is approximately to stabilize sediments which have been balanced by nutrient loss in exported deposited by physical processes. organic matter, then nutrient storage will be minimal and mangrove net primary pro­ duction wi 11 be low. This appears to occur in the scrub mangrove community type 3.3 NUTRIENT CYCLING and to a lesser extent in the basin and hammock community types. Current understanding of nutrient cycles in mangrove ecosystems is far from Carter et al. (1 973) and Snedaker and satisfactory. Sporadic field measurements Lugo (1973) have hypothesi zed that the have been made, but a complete nutrient greatest natural nutrient inputs for man­ budget has not been publ i shed for any grove swamps come from upland and terres­ mangrove ecosystem in the world. trial sources. Apparently for this rea­ son, the most luxuriant and productive Several pioneering field studies were mangrove forests in south Florida occur in conducted in Florida (Carter et al. 1973; riverine locations or adjacent to siqnifi­ Snedaker and Lugo 1973; Onuf et al. 1977) cant upl and drai nage. and one simulation model of mangrove nu­ trient cycling has been published (LU90 et Localized sources of nutrients, such al. 1976). Preliminary measurements of as bird rookeries, can result in greater nitrogen fixation were made (Zuberer and nutri ent storage and hi gher mangrove pro­ Silver 1975; Gotto and Taylor 1976; ducti vity (Onuf et a1. 1977). If however, Zuberer and Sil ver 1978; Gotto et al. large bird rookeries (or artificial nu­ 1981). 8ased on these studies, we present trient inputs) occur in poorly flushed the following preliminary conclusions. sections of mangrove ecosystems, resultant high nutrient levels may inhibit mangrove Mangrove ecosystems tend to act as a growth (R. R. Lewis, III, Hillsborough sink (net accumulator) for various ele­ Community ColI ege, Tampa, Fl a.; personal ments including macro nutrients such as communication 1981). nitrogen and phosphorus, trace elements, and heavy metals. As we have discussed in The output from the simulation model section 1.7, these elements are removed of Lugo et al. (1976) suggests that if from waters f1 owi ng through mangrove nutri ent input to a mangrove ecosystem is SI,amps by the concerted action of the reduced, then nutrient storage levels mangrove prop roots, prop root al gae, the wi thi n the mangrove ecosystem wi 11 be associated sediments, the fine root system reduced and mangrove biomass and produc­ of the mangrove trees, and the host of tivity will decline. To our knowledge small invertebrates and microorganisms this hypothesis has not been tested in the attached to all of these surfaces. Al­ fi e1d. thou9h the turnover times for these ele­ ments in mangrove swamps are not known, it Nitrogen fixation occurs in mangrove appears that at least a portion may be swamps at rates comparable to those stored or tied up in wood, sediments, and measured in other shallow, tropical marine peat for many years. areas (Gotto et al. 1981). Nitrogen

30 SMALL SMALL

IMPORT EXPORT LOW STORAGE

LOW BIOMASS

LOW PRODUCTIVITY

LARGE MODERATE

IMPORT 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 heen found in association salinities, and pollution events. Litter with mangrove leaves, both living and fall typically can be partitioned as 68% dead, mangrove sediment surfaces, the to 86% leaves, 3% to 15% twigs and 8% to litter layer in mangrove swamps, and man­ 21% miscellaneous; the latter includes grove root systems (Gotto and Taylor 1976; flowers and propagules. Zuberer and Silver 1978: Gotto et a1. 1981). In virtually all cases, nitrogen Litter fall is an important ecosystem fixation appears to be limited by the process because it forms the energy basi s availability of labile carbon compounds. for detritus-based foodwebs in mangrove Perhaps for this reason, the highest rates swamps (see sections 3.5 and 3.6). The of mangrove nitrogen fixation have been first measurements of litter fall in man­ measured in association with decaying grove swamps were made by E.J. Heald and mangrove leaves; presumably, the decaying W.E. Odum, working in the North River leaves act as a carbon source and thus estua ry in south Flori da in 1966-69. accelerate nitrogen fixation. Macko This was subsequently published as Heald (1981), using stable nitrogen ratio (1969), Odum (1970), and Odum and Heald techniques, has indicated that as much as (1975a). They estimated that 1i tter pro­ 25% of the nitrogen associated with black duction from riverine red mangrove forests mangrove peat in Texas is derived from average! 2.4 dry g 2f organic nitrogen fixation. matter/m /day (or 876 g/m /year or 8.8 metri c tons/ha/year). Zuberer and Silver (1978) speculated that the nitrogen fi xati on rates observed Subsequent studi es agreed with thi s in Florida mangrove swamps may be suf­ early estimate (Table 3), although varia­ ficient to supply a significant portion of tion clearly exists between different the mangrove's growth requi rements. Al­ types of communities. Scrub forests with though this hypothesis is impossible to scattered, very small trees have the test with present information, it might smallest amount of leaf fall. Basin and explain why moderately productive mangrove hammock forests, which appear to be stands occur in waters which are severely nutrient limited, have intermediate leaf nitrogen depleted. fall values. Not surprisingly, the highest values occur in the highly produc­ In summary, knowledge of nutrient tive fringing, overwash, and riverine cycling in mangrove swamps is highly forests. Odum and Heald (1975a) suggested specul at i ve. These ecosystems appear to that the relatively uniform litter fall 'act as a sink for many elements, including values from productive mangrove forests nitrogen and phosphorus, as long as a around the world result from the shade modest input occurs. Nitrogen fixation intolerance of the canopy leaves and the within the swamp may provide much of the tendency for the canopy size to remain the nitrogen needed for mangrove growth. same in spite of increasing height. If detailed information is lacking, red man­ grove forests of south Flori da, whi ch are 3.4 LITTER FALL AND DECOMPOSITION not severely limited by lack of nutrients, can be assume~ to produce litter fall of Unless otherwise stated, litter fall 2.0 to 3.0 g/m /day of dry organi c matter. refers to leaves, wood (twigs), leaf Pure stands of black mangroves us~al1y scales, propagules, bracts, flowers, and have a lower rate of 1.0 to 1.5 g/m /day insect frass (excrement) which fall from (Lugo et a1. 1980). the tree. Mangrove leaves are shed con­ tinuously throughout the year although a Decomposition of fallen Florida man­ minor peak occurs during the early part of grove leaves has been investigated by a the summer wet season in Florida (Heald number of researchers including Heald 1969; Pool et al. 1975). Sporadic litter (1969), Odum (1970), Odum and Heald fall peaks may follow periods of stress (1975a), 'Pool et a1. (1975), Lugo and from cold air temperatures, high soil Snedaker (1975), Twilley (1980) and Lugo et

32 Table 3. Estimates of litter fall in mangrove forests. Total litter fall in­ cludes leaves, fruits, twigs, flowers, and bark. R = red mangrove, W = white mangrove, B = black mangrove.

Species Leaf fall Total lit~er Annual litter Reference (g/m 2/day) fall (g/m /day) fall (metric tons/ha/yr)

R (riverine) 1 . 3 2.4 8.8 Heald 1969

R (riverine) 3.6 12.8 Po 0 1 et a1 . 1975

R (overwash) 2 . 7 9.9 Po 0 1 et a 1 . 1975

R (fringe) 2.7 9.9 Po 0 1 et a1 . 1975

R,B (basin) 2.0 7.3 Po 0 1 et a 1 . 1975 R (mature) 2.2 2.9 10.6 Teas 1979 w R (scrub) 0.2 0.4 1 .3 Teas 1979 w B (basin) 0.7 0.8 2.9 Tea s 1979 B (basin) 2.2 8.0 Courtney 1980 B 1 . 3 4.9 Twilley 1380 B 1.3 4.8 Lugo et a1 . 1980

Mixed R, B t W 2.5 9.0 Lugo et a 1 . 1980

B 0.8 2.9 Po 0 1 et a1 . 1975 Vari ety of 0.8 - 2 . 1 2.9 - 7.7 Heald eta1 . 1979 community types

26 species 2.4 8.8 Boto & Bunt (r'lS. in (Austral i a) prep.) a1. (1980). Heal d and Odum showed that forests, have slower rates of decomposi­ decomposition of red man9rove leaves t i on and lower export rates. proceeds most rapidly under marine condi­ tions, somewhat more slowly in freshwater, and very slowly on dry substrates. For 3.5 CAR80N EXPORT example, using the litter bag method, they found that only 9% of the original dry Research from Florida mangrove swamps weight remained after 4 months in sea forms a small portion of the larger con­ water. By comparison, 39% and 54% re­ troversy concerned with the extent to mained at the end of comparable periods in whi ch coastal wetl ands export parti cul ate brackish water and freshwater. Under dry organic carbon (reviewed by Odum et a1. conditions, 65% remained. Higher decompo­ 1979a). Available evidence from Florida, sition rates in sea water were related to Puerto Rico and Austral ia (Table 4) sug­ increased acti vity of shredder organisms, gests that mangrove swamps tend to be net such as crabs and amphi pods. exporters. The values in Table 4 should be regarded as preliminary, however, since Heald (1969) and Odum (1970) also all five studies are based upon simplistic found inc rea ses in nit rogen, protei n, and assumptions and methodology. caloric content as mangrove leaves pro­ gressi vely decayed. The ni trogen content Goll ey et al. (1962) based thei r of leaves decaying under brackish condi­ annual estimate of particulate carbon tions (on an AFDI' basis) increased from export from a Puerto Rican forest upon a 1.5% (5.6% protei n) to 3.3% (20.6% few weeks of measurements. Odum and protein) over a 6-month period. Subse­ Heald's estimates were derived from two or quent information (Odum et a1. 1979b) three measurements a month. All i nvesti­ suggested that the protein increase may gators have ignored the importance of bed not have been this great since some of the load transport and the impact of extreme nitrogen increase probably included non­ events. All investi gators except Lugo et protein nitrogen compounds such as amino a1. (1980) have failed to measure DOC sugars. Fell and Master (1973), Fell et fl ux. a1. (1980), Fell and Newell (1980), and Fell et a1. (1980) have provided more It seems relatively clear that man­ detailed information on red mangrove leaf grove forests do export organic carbon to decomposition, the role of fungi in decom­ nearby bodies of water. The magnitude of position (see section 4), and nitrogen this export has probably been underesti­ changes and nitrogen immobilization during mated due to ignoring bedload, extreme decomposition. Fell et a1. (1980) events, and DOC. have shown that as much as 50% of wei ght loss of the leaf during decomposition is The value of this carbon input to in the form of dissolved organic matter secondary consumers in receiving waters is (DOM) • not clear. As shown in section 3.6, food webs based primarily upon mangrove carbon Heal d et al. (1979), Lugo et al. do exist. The relative importance of (1980) and Twilley (1980) discovered that mangrove carbon to Florida coastal ecosys­ bl ack mangrove leaves decompose more ra­ tems remains speculative. We suspect that pidly than red mangrove leaves and ap­ mangrove-based food webs are dominant in parently produce a higher percentage of small bays, creeks and rivers within large DOM. Pool et a1. (1975) have shown that mangrove ecosystems such as the North mangrove litter decomposes and is exported River system studied by Heald (1969) and most rapidly from frequently flooded Odum (1970). In i ntermedi ate-si zed bodi es riverine and overwash forests. These of water, such as Rookery Bay near Naples, communities have little accumulation of Florida, mangroves are probably important litter on the forest floor. Communities but not dominant sources of organic car­ which are not as well-flushed by the bon. Lugo et a1. (1980) estimate that tides, such as the basin and hammock mangroves supply 32% of the organic carbon

34 Table 4. Estimates of particulate carbon export from mangrove forests. LU90 et a1. (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.

Export

Investi ga tors Location tonnes/ha/yr

Golley et a1. (1962) Puerto Rico 1.1 4.0 Heald (1969), Odum (1970)a Florida 0.7 2.5 Lugo and Snedaker (1975) Flori da 0.5 2.0 LU90 et al. (1976) Flori da 1.5 - 1.8 5.5 - 6.6 Boto and Bunt (1981) Aus tra1ia 1.1 4.0 Lugo et al. (1980)b Florida 0.2 0.7

~Estimate only includes carbon of mangrove origin. Estimate includes dissolved and particulate carbon.

35 input to Rookery Bay. In very large sys­ bodies of clear, relatively deep water. tems, such as Biscayne Bay near Miami, Conversely, phytoplankton are hypothesized Florida, mangroves are clearly less impor­ to be rel ati vely uni mportant to the energy tant than any other sources such as a1 gae budgets of the 1arge ri veri ne forest com­ and sea grasses, although mangrove carbon munities along the southwest coast of may be important in localized situations Florida. It should be remembered, how­ such as the immediate vicinity of fringing ever, that even where phytoplankton are and overwash forests. The magnitude of quantitatively unimportant, they poten­ mangrove carbon export to unenclosed tially perform an important function as coastal waters and offshore remai ns a the basis of phytoplankton-zooplankton­ mystery. 1arva1 fi sh food webs (Odum 1970).

As a third hypothesis, Iver Brook 3.6 ENERGY FLOW (Rosensteil School of Marine and Atmos­ pheric Sciences, Rickenbacker Causeway, At least seven sources of organic Miami, Fla.; personal communication 1979) carbon may serve as energy inputs for has suggested that both sea grasses and consumers in mangrove ecosystems (Figure benthic algae serve as an important energy 8). The pathways by which this energy source for fringing mangrove communiti es containing material is processed and made adjacent to large bodies of water such as available to each consumer species is Bi scayne Bay and Whitewater Bay. Although indeed complex. Not surprisingly, current little evidence exists to test this hypo­ understanding of energy flow in Florida thesis, observations of extensive deposits mangrove ecosystems exists largely in a of sea grass and macroalgal detritus with­ qualitative sense; quantitative data are in mangrove forests suggest intuitively scarce and piecemeal. A variety of inves­ that Brook's hypothesi s may be correct. tigators have contributed information over the past decade including, but not limited In regions where mangrove shading of to, Heald (1969), Odum (1970), Odum and the prop roots is not severe, our fourth Heald (1972), Carter et a1. (1973), hypothesis suggests that carbon origina­ Snedaker and Lugo (1973), Heald et al. t i ng from prop root epi phytes may be si g­ (1 974), Lugo and Snedaker (1974, 1975), nificant to community energy budgets. Odum and Heald (1975a, b), and Pool et al. Lugo et a1. (1975) have measured net pro­ (1977). Probably, the most complete study duction of periphyton in mangroves to date is the investigation of energy fringing Rookery B~ and found average flow in the black mangrove zone of Rookery val ues of 1.1 gC/m /day. Hoffman and Bay by Lugo et al. (1980). Dawe~ (1980) found a lower val ue of 0.14 gC/m /day. Because these val ues are It is possible at this time to pre­ roughly comparable to average exports of sent a series of hypotheses concerning the mangrove leaf carbon (section 3.5), its relative importance of these energy potential importance is obvious. sources. Fi rst, the rel ati ve importance of each source can vary from one locati on The fi fth hypothesi s states that to the next. As wi 11 be shown in the mangrove organic matter, particularly leaf following discussion, the consumers in materi a1, is an important energy source certain mangrove forests appear to depend for aquatic consumers. This hypothesis primarily upon mangrove-derived carbon was first espoused by Heald (1969) and while in other locations inputs from phy­ Odum (1970), who worked together in the toplankton and attached algae are probably riverine mangrove communities between the more important. Evergl ades and Whitewater Bay. Clearly, mangrove carbon is of great importance Our second hypothesis is that energy within the riverine and basin communities flow based upon phytoplankton is most a 11 along the southwest coast of Flori da important in overwash mangrove forests and (Odum and Heald 1975b); Carter et al. other locations associated with large (1973) and Snedaker and Lugo (1973)

36 SUN _

PHYTOPLANKTON EPIPHYTIC SEA GRASSES MANGROVES TERRESTRIAL ATMOSPHERI C ALGAE SOURCES SOURCES GRAZING~~~~;D~I;ssjO~L;V~ED~~~~~~piA;RTiI~ciUL~ATE reduced sul fur 01 kECT ORGANIC MATTER ORGANIC MATTER

"" , ? v ""MICROBES SULFUR OXIDIZING BACTERIA ", " /- HI GHER CONSUMERS

Figure 8. Potential pathways of energy flow in mangrove ecosystems. Not all possible pathways have been drawn; for example. methanogenesis and sulfur reduction could originate from any of the sources of organic matter. ~~ngrove-based pathways are enhanced for emphasis and in no way imply relative importance. provided subsequent supportive data. What of very low salinity. is not clear, is the relative importance of mangrove carbon to consumers within Carbon inputs from terrestrial fringing, overwash, and more isolated sources may be important to certain man­ manqrove communities. grove communities. Carter et al. (1973) have shown that terrestrial carbon can Our si xth hypothesi s invol ves the reach coastal ecosystems particularly assemblage of organisms that graze man­ where man has cut deep channels inland for grove leaves di rect1y. A variety of in­ navi gat i on or drai nage purposes. The sects (see section 6) and the mangrove magnitude of this influx has not been tree crab, Aratus pi soni i, (Reever et al. adequately measured although Carter et al. 1979) obtain much of their energy directly did find that mainland forests (including from livin9 mangrove leaves, even though mangroves) contributed approximately 2,100 9razing rarely exceeds 10% of net primary metri c tons of carbon per year to production (Odum and Heald 1975b). Fahkahatchee Bay.

As a seventh hypothesis we suggest Atmospheric inputs from rainfall that anaerobic decomposition of mangrove appear to be minimal in all cases. Lugo tissue, particularly root material, may et al. (1980) measured throughfall (preci­ support an extensive food web based on pitation passing through the tree canopy) bacteria associated with methanogenesis or in Rookzry Bay mangrove forests of 15 to the processing of reduced sulfur com­ 17 gC/m /year. This would be an overesti­ pounds. Our suggestion of the importance mate of atmospheric input since it con­ of reduced sul fur comes di rectly from tains carbon leached from mangrove leaves. Howarth and Teal's (1980) discovery of The best guess of atm~spheric input is this potentially important energy pathway between 3 to 5 gC/m /year for south in temperate Spartina (cordgrass) marshes. Florida mangrove ecosystems. They found that anaerobic decomposition is such an incomplete process that if sul­ Subsequent stages of energy transfer fates are available (from sea water) as in mangrove community food webs remain much as 75% of the original energy in largely hypothetical. Odum (1970) and plant tissues may be converted by sulfur Odum and Heald (1975b) have outlined reducing bacteria to reduced sul fur com­ several pathways whereby mangrove carbon pounds such as hydrogen sulfide and Py­ and energy are processed by a variety of rite. Subsequently, if these reduced organisms (see Figure 8). Apparently, the sulfur compounds are moved hydrologically most important pathway follows the se­ to an oxidized environment (sediment sur­ quence: mangrove-leaf detritus substrate­ face or creek bank) sulfur -oxidi zi ng bac­ microbe-detritus consumer-higher consu­ te ri a (e.g., Thi ohac ill us sPp.) may conve rt mers. The critical links are provided by the chemically-stored energy to bacterial­ the microbes such as bacteria and fungi ly stored energy with an efficiency as (see Fell et a1. 1975) and by the detritus great as 50% (Payne 1970). Presumably, consumers. The latter group was studied deposit-feeding organisms such as grass by Odum (1970) and Odum and Heald (1975b) shrimp (Palaemonetes) and mullet (Mugil) and found to consist of a variety of are capable--or-grazi ng these su"""if'Ur­ invertebrates (e.g., caridean shrimp, oxidizing bacteria from the sediment crabs, mollusks, insect larvae, amphipods) surface. If this hypothetical trophic and a few fi shes. exchange does exi st, it may be of con­ siderable magnitude and may cause us to Stable carbon studies such as those reexamine current concepts of energy pro­ done by Haines (1976) in Spartina cessing and export from mangrove (cordgrass) marshes have not been per­ ecosystems. Since freshwater contains formed in mangrove ecosystems. Mangroves 13 remarkably little sulfate in comparison to are C3 plants and have 6 values in the seawater, this energy pathway is probably range of minus 25 to minus 26 (Macko of little importance in mangrove forests 1981). According to the same author,

38 mangrove peat has a 6 13 value of minus nature. This framework appears to be 22. Because these values are dramatically reasonably accurate although subsequent di fferent from the values for sea grasses developments, such as elucidation of the and many algae, the possibilities for reduced sul fur hypothesi s, may requi re using this tool in mangrove ecosystems is some modification. excellent. Macko (1981) also suggested the utility of using stable nitrogen ra­ (2) Measurements of the relative tios for future mangrove food web investi­ importance of vari OUS ca rbon sou rces are gations; he reported 6 15 values of plus generally lacking. 6.0 to p1 us 6.5 for mangrove ti ssue and plus 5 for mangrove peat. (3) Oetai 1ed measurements of energy flow including the relative inputs of In reviewing contemporary knowledge different carhon sources are critically of energy flow in mangrove ecosystems, needed. Technological diffi cult i es, hi gh th ree conc1us ions emerge. costs, and difficulties inherent in transferring findings from one estuary to (1) We have a hypothetical framework the next present a major challenge to of mangrove energy flow of a qual i tati ve estuarine ecologists of the future.

39 CHAPTE R 4. COMMUNITY COMPONENTS - MICROORGANISMS

The mycof10ra (fungi) are the best Orechslera and Gloeosporium. In the lat­ studied component of the microbial com­ ter stages of decay the dominant genera munity of mangrove swamps. t1uch pio­ are Calso, G1iocidium, and Lulworthia. neering work has been carried out in south Florida. Reviews of the current knowledge Understanding the occurrence and suc­ of mangrove -associ ated fungi can be found cessi on of fun9i on decayi ng mangrove in Kohlmeyer and Kohlmeyer (1979) and Fell leaves is important because of their role et a1. (1 980 )• in energy flow in mangrove swamps. Heald (1969), Odum (1970) and Odum and Heald One of the earliest studies of man­ (1975b) hypothesi zed that fun9i and bac­ grove mycoflora was published by Kohlmeyer teria are important in convertin9 mangrove (1969). He discovered large populations 1eaf organi c materi ali nto a form that can of marine funqi 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 man9rove leaves. Exten­ sive work at the University of Miami by Fell and his coworkers (e.g., Fell and Our understandi ng of the role and Master 1973; Fell et a1. 1975, 1980) ex­ occurrence of bacteria in man9rove 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 nitr0gen. Newell (1974) that bacteri a are important in the early studied the succession of mycoflora on stages of mangrove leaf decomposition and seedlings of red man9rove. 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 a1. (1968). mycofl ora, the bacteria are c1ea rly i mpor­ tant in the anaerobic regions of mangrove One of the most interestin9 pieces of swamps. Vankatesan and Ramamurthy (unpubl. information to emerge from this extensive data) found denitrifying bacteria to be mycof10ra research concerns the succession abundant and ubiquitous in mangrove soils. of orqanisms associated with decaying Zuberer and Silver (1978) have emphasized 1eaves (summari zed by Fell et a1. 1975, the importance of nitrogen-fixing bacteria 1980). Senescent leaves of red mangroves in the zone around mangrove roots. They, are t.vpically colonized by species of in fact, were able to isolate and count a Nigrospora, Phy110stica, and Pesta10tica. variety of types of bacteria from mangrove Once the leaf has fallen from the tree and sediments including aerobic heterotrophs, duri n9 the early stages of decay, the anaerobic heterotrophs, nitrogen-fixing fungal fl ora is domi nated by speci es of heterotrophs, and sulfate-reducing bac­ ~~l.tophthora and, to a lesser extent, te ri a.

40 CHAPTER 5. COMMUNITY COMPONENTS - PLANTS OTHER THAN MANGROVES

5.1. ROOT AND MUD ALGAE Because much mud is often deposited on the Bostrychia-Catenella-Caloglossa complex, The aeri a1 root systems of mangroves it often has a dingy, gray appearance. provide a convenient substrate for at­ There are many other algae found in this tachment of al gae. These root al gal com­ zone, but these three genera usually domi­ munities are particularly noticeable on nate. At brackish or nearly freshwater red mangrove prop roots but also occur to locations, they are replaced by species of a 1esser extent on bl ack mangrove Batophora, Chaetomorpha, CladQ.phora, and pneumatophores located in the intertidal Penicillus. The pneumatophores of zone. Productivity of prop root algal AVTCennTa-,- when colonized, are often communities can be appreciable if shading covered with species of Rhizoclonium, by mangroves is not too severe; as dis­ Bostrychia and Monostroma (Taylor-T§61J1. cussed in section 3.6, Lugo et al. (1975) Hoffman and Dawes (1980) found that the found a prop root communitt' net primary Bostrychia binderi -dominated community on production rate of 1.1 gC/m /day, a level the pneumatophores of black mangroles had comparable to mangrove leaf fall. Biomass a standing crop of 22 g drt' wt/m and a of these algae can be as high as 200 to net production of 0.14 gC/m /day. 3DD g per prop root (Burkholder and Almodovar 1973). Of course, production of If there is a permanently submerged this magnitude only occurs on the edge of portion of the prop root, it may be the forest and is virtually nil in the covered with rich growths of Acanthophora, center of the swamp. Nevertheless, this Spyrida, Hypnea, Laurencia, IIrangelia, algal carbon has considerable potential ~lonia, and Caulerpa (Almodovar and Biebl food value either to direct grazers or 19~ Additional genera which may be detritivores. present below mean high water are: Murrayella, POlysiphonia, Centroceras, Vertical distribution of prop root ~urd!~ann~, Dic~la, ~~ll~~~, algae has been studied by many researchers Laurencia, and Dasya TTaylor 1960; (Gerlach 1958; Almodovar and Riebl 1962; Burkholder and Almodovar 1973; Yoshioka Biebl 1962; Post 1963; Rutzler 1969; 1975). In addition, anywhere on the moist Burkholder and Almodovar 1973; Rehm 1974; sections of the prop roots there are Yoshioka 1975); only one of these studies usually epiphytic diatoms and filamentous (Rehm 1974) was conducted in Florida. green and blue-green algae of many genera. There is a tendency for certain genera of algae to form a characteristic association Rehm (1974) found a significant dif­ on mangrove roots around the world (Post ference in the prop root algae between 1963). Four phyla tend to dominate: south and central Florida. South of Tampa Chlorophyta, Cyanophyta, Phaeophyta, and Bay the standard Bostrychia-Catenella­ Rhodophyta; the last is usually the most Caloglossa dominates. In the Tampa Bay important in terms of biomass. Of 74 area, species of the orders Ulotrichales species of marine algae recorded as prop and Cladophorales are dominant. root epi phytes between Tampa and Key Largo, 38 were Rhodophyta, 29 Chlorophyta, The mud adjacent to the mangrove root 4 Phaeophyta and 3 Cyanophyta (Rehm 1974). community is often richly populated with a variety of algae. These can include Zonation to be expected on Florida speci es of Cl adophoropsi s, Enteromor ha, man9roves is shown in Figure g; this se­ Vaucheria, and Boodleopsis Taylor 1960 quence comes largely from Taylor (1960). in addition to a whole host of benthic Near the high water mark, a green band diatoms and dinoflagellates (llood 1965) usually exists which is dominated by spe­ and other filamentous green and blue-green cies of Rhizoclonium. Below this is a algae (r~arathe 1965). zone dominated by species of Bostrychia, Catenella, and Caloglossa. It is this Adjacent to mangrove areas, on the association that most people think of when bottoms of shoals, shallow bays and mangrove prop root algae are mentioned. creeks, there is often a variety of 41 PLANTS ANIMALS

Ligea exotica Rhlzoc/onlum spp. L ittorina angulifera MHW --- -- ~=-='-= ;\-y.--:..=~------Balanus eburneus Brachidontes spp. Bostrychla spp. Catanella spp. Narels spp. Bulla spp. C" ••,,,., •••. L Ascldla niger Crassos trea virginica MLW- - --- =---=->. Sphaeroma Acanthophora spp. terebrans Caulerpa spp. Wrangleia spp. ----.

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 Before we can understand the impor­ Caulerpa, Acetabularia, Penicillus, tance (or lack of importance) of phyto­ Grae i 1 a ria, ~ll'!'e d a, Sa r gas sum, plankton in mangrove regions, some ques­ Batophora, Udotea, and Dasya. These are tions must be answered. How productive discussed at length by Zieman (in prep.). are the nannoplankton? How does the daily Other pertinent references for mangrove and seasonal shift in phytoplankton domi­ regions include Davis (1940), Taylor nance affect communi ty producti vi ty? Does (1960), Tabb and Manning (1961), and Tabb the generally low standing crop of phyto­ et a1. (1962). pl ankton represent low productivity or a high qrazing rate?

5.2 PHYTOPLANKTON 5.3 ASSOCIATED VASCULAR PLANTS All aspects of phytoplankton, from seasonal occurrence to productivity Four species of aquatic grasses occur studies, are poorly studied in mangrove on bay and creek bottoms adj acent to man­ ecosystems. This is particularly true in grove forests. Turtle grass, Thalassia Florida. testudinum, and manatee grass, Syringodium filliforme, are two tropical sea grasses Evidence from Brazil (Teixeira et al. which occur in waters with average salini­ 1965,1967,1969; Tundisi 1969) indicates ties above about 20 ppt. Shoal grass, that phytoplankton can be an important Halodule wrightii, is found at somewhat component of the total primary production lower salinities and widgeongrass, ~uppia in mangrove ecosystems; just how important maritima, is a freshwater grass which can is not clear. Generally, standing crops tolerate low salinities. These grasses of net phytoplankton in mangrove areas are occur throughout south Florida, often in low (personal observation). The nanno­ close juxtaposition to mangroves. Zieman plankton, which have not been studied at (in prep~ presents a thorough review of all, appear to be most important in terms sea grasses along with comments about of total metabolism (Tundisi 1969). The possible energy flow linkages with net plankton are usually dominated by mangrove ecosystems. diatoms such as Thalassothrix spp., Chaetoceras spp., Ni tzschi a spp., There are extensive areas of man­ Skel etonema spp., and RhlZosOTei1i a spp. groves in south Florida which are closely (Mattox 1949; Wood 1965; Walsh 1967; Bacon associated with marshes dominated bv a 1970). At times, blooms of dinoflagel­ variety of other salt-tolerant plants. lates such as Peridinium spp. and For example, along the southwest coast G mnodinium spp. may dominate (personal between Flamingo and Naples, marshes are observation. In many locations, particu­ scattered throughout the mangrove belt and larly in shallow waters with some turbu­ also border the mangroves on the upland lence, benthic diatoms such as Pleurosigma side. The estuarine marshes within the spp., Mastogl oi a spp., and Di sp1onei s may mangrove s"amps have been extensively be numerically important in the net plank­ described by Egler (1952), Carter et a1. ton (Wood 1965). (1973), and Olmstead et a1. (1981). They contain various salt-tolerant marsh Understanding the mangrove-associated species including: salt grass, Distichlis phytoplankton community is complicated by spicata, black needle rush, Juncus the constant mixing of water masses in roemerianus, spike rush, Eleocharis mangrove regions. Depending upon the cellulosa, 91aSS wort, Salicornia spp., location, the phytoplankton may be domi­ Gulf cordgrass, Spartina spartinae, sea nated by oceanic and neritic forms, by purslane, Sesuvium portulacastrum, salt true estuarine plankton, and by freshwater wort, Batis maritima, and sea ox-eye, plankton. The pattern of dominance may Borric~frutescens. Farther north, change daily or seasonally depending upon above Tampa on the west coast of Flori da, the source of the principal water mass. marshes populated by smooth cordgrass,

43 Spartina alterniflora, and black needle that have become established well inland. rush, Juncus roemerianus, become more See Hofstetter (1974) for a review of extens i ve and eventua11 y replace mangrove literature dealing with these plants. swamps. Even in the Everglades region, the saline marshes are comparahle to man­ Finally, a group of somewhat salt­ groves in areal extent, although they tolerant herbaceous plants is found tend to be some distance from open water. within stands of mangroves. They usually Studies of these marshes, including as­ occur where slight increases in elevation sessment of their ecological value, are exist and where sufficient light filters almost non-existent. Certainly, they have through the mangrove canopy. Carter et considerable importance as habitat for al. (1973) list the following as examples small fishes which, in turn, support many of members of the mangrove community: of the nesti ng wadi ng bi rds in south 1eather ferns, Acrostichum aureum and A. Florida (see section g). danaeifolium; spanish bayonet, Yucca aloifolia; spider lily, HymenociillTS Tropical hardwood forests may occur latifolia; sea blite, Suaeda linearis; within the mangrove zone in south Florida, chaff flower, Alternanthera ramosissima; partIcularly where old shorelines or areas samphire, Philoxerus vermicularis; blood­ of storm sedimentation have created ridges leaf, Iresine celosia; pricklypear cactus, 1 m or more above MSL (mean sea level) Opunt.,--a--srricta; marsh elder, Iva (Olmstead et al. 1981). Similar forests frutescens; the rubber vine, Rhabdadenia or "hammocks" occur to the rear of the bi fl ora; the 1i anas, Ipomoea tuba and mangrove zone on hi gher ground. Typical Hippocratea volubilis; and a variety of trees in both forest types include the fan brornel i adS\Bromel i aceae). pal m, Thrinax radiata, buttonwood, Conocarpus erecta, manchineel, Hippo~an~ Although the lists of vascular plants mancinella, and, in the past, mahogany, which occur in mangrove swamps may seem SWTetenia mahagoni. Olmstead et al. extensi ve, the actual number of speci es in lT981Tprovide-a descri pt i on of these any given location tends to be low communities. compared to totally freshwater envi ron­ ments (see Carlton 1977). Analogous to Freshwater marsh pl ants, such as the temperate salt marshes, mangrove swamps grasses, rushes and sedges that dominate possess too many sources of stress, the freshwater Evergl ades, are not particularly from tidal salt water, to mentioned here, although they are have a high diversity of vascular plant occasionally mixed in with small mangroves 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, ;50­ 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. Whil e there are few compari sons mangrove swamps is the pattern of horizon­ of species richness with other types of tal and vertical zonation of invertehrates coastal ecosystems, mangrove swamps appear (Fi gure 9). Characteri stic verti cal 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 (1 974) compared H' (Shannon Weaver) di ver­ 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 ~ery high, often in excess of 100 dry Spartina marshes but considerably less g/m of organic matter in many locations than were associ ated with rocky substrate (persona1 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 (1 962) found an average of 6.4 g/m 2 of i nterti dal mangrove trees provi des habitat invertebrates in the center of a Puerto for a wi de 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 Panu1irus argus, is well established on trophic position: (Olsen et ar:-l"975; Olsen and Koblic 1975; Little 1977; Witham et al. 1968). Ac­ (1) di rect grazers - 1i mited 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 di rect1y; 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 whi ch fi Her phy­ close association with the prop root com­ toplankton and detritus from the water; muni ty for as much as 2 years unti 1 they reach a carapace length of 60 to 70 mm. (3) deposit feeders - mobile inverte­ brates which ski m detritus, a1 gae 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 man9rove swamps and nematodes, a wide variety of insects, ener9Y flow are di scussed insection 3.6. 45 Figure 10. Photograph of red mangrove prop root habitat in clear shallow water with associated animal and plant populations. Photograph is by Bianca Lavies (copyright, National Geographic Society). 6.2. ARBOREAL ARTHROPOD COMMUNITY been described by Warner (1967). 12 Jamaica its numbers range from 11 to l~/m A surprising variety of arthropods at the edge of fri ngi ng slPine Island Sound) they found in healthy, unstressed mangrove stands nor­ accordance with normal biogeographical mally have less than 10% of their total theory, the highest densities of crabs leaf area grazed (Heald 1969). In many associated with fringing forests and the locations, percent leaf area damaged is on lowest densities on distant islands, but the order of I% to 2% (Beever et al. at Sugar Loaf Key the unexplainable 1979). There are excepti ons. Onuf et a1. reverse distribution was found. (1977) reported biomass loss to arthropod grazers as high as 26% in a mangrove stand Other invertebrates may visit the where growth and nitrogen content of the canopy from below either for purposes of leaves had been enhanced by input of nu­ feeding or for protection from high tides. t ri ents from a bi rd rookery. Included in this group are the pulmonate gastropods, Li ttori~ angu1 i fera, In terms of numbers of species, the Cerithidea scalariformis, and Me1ampus dominant group of arboreal arthropods is coffeus, the isopod, Ligea~)(otica, and a insects. The most thorough inventory of host of small crabs. mangrove-associated insects was conducted by Simberloff and Wilson to obtain the raw In summary, with the exception of a data for their papers on island bio­ hal f dozen key papers, the arboreal man­ geography (Simberloff and Wilson 1969; grove community has been generally ig­ Si mber10ff 1976). These papers 1ist over nored. Both insects and the mangrove tree 200 species of insects associated with crab play significant ecological roles and overwash mangrove islands in the Florida may affect mangrove producti vity to a Keys. There is no reason to expect lesser greater extent than has heen recognized. numbers in other types of mangrove com­ munities, except for the mangrove scrub 6.3 PROP ROOT AND ASSOC IATED MUD SURFACE forests. The most thorough study of i n­ COMMUNITY sect grazing on mangrove leaves is that of Onuf et al. (1977) (see section 2.6). These two somewhat di sti nct com­ munities have been lumped together because Although not as numerically impres­ of the large number of mobile organisms sive as the insects, the man9rove tree which move back and forth between tidal crab, Aratus pisonii, appears to be poten­ cycles. The aerial roots are used as tially as important in terms of 9razing protecti ve habitat and to some extent for impact (Beever et al. 1979). The life feeding while the nearby mud substrates history of this secretive little crab has are used principally for feeding.

47 The prop roots support an abundance including latitude, salinity, and proxi­ of sessile organisms. The vertical mi ty to other communi ties such as sea zonation of both mobile and sessile inver­ grass beds and coral reefs. tebrates has been studied extensively in other parts of the world (Good body 1961; Sutherland (1980), working on red Macnae 1968; Rutz1er 1969; Coomans 1969; mangrove prop root communities in Bacon 1970; Ko1ehmainen 1973; Sasekumar Venezuela, found little change in the 1974; Yoshioka 1975). Vertical zonation invertebrate species composition on indi­ certainly exists on Florida red mangrove vidual prop roots during an 18-month roots. The general i zed scheme shown in period. The species composition varied Fiqure 9 essentially contains two zones: greatly, however, between adjacent prop an upper zone dominanted by barnacles and roots, presumably in response to stochas­ a lower zone dominated by mussels, oysters tic (chance) processes. and ascidians. Between mean high tide and mean tide, the wood boring isopod, The mud flats adjacent to mangroves ~haeroma terebrans (discussed at length provide feeding areas for a range of in­ in section 2.7) is important, both numeri­ vertebrates that scuttle, crawl, and swim cally and through the provision of out from the cover of the mangrove roots. numerous holes for use by other organisms Some emerge at low tide and feed on algae, (Estevez 1978). detritus, and small invertebrates on the mud f1 ats whi 1e they are hi gh and dry. The most compl ete study of the Others emerge while the tide is in, parti­ Florida mangrove prop root community is cularly at night, and forage across the Courtney's (1975) comparison of seawall flooded flats in search of the same foods and mangrove associations. He reported an plus other invertebrates which have extensive list of invertebrates from man­ emerged from the mud. In many ways the grove prop roots at Marco Island, Florida, mangrove-mud flat relationship is analo­ including: Crassostrea virginica, gous to the coral reef (refuge) sea grass Litt:.~r:!..'!.~ angulifera, Crepidula ~, (feeding area) relationship reviewed by Jiodora cayenensis, Urosalpinx perrugata, Zieman (in prep.). The net effect is that Pisania tincta, Brachidontes exustus, the impact of the mangrove community may nine species of polychaetes, ~erom~ extend some distance beyond the boundaries terebrans, Palaemon floridanus, of the mangrove forest. PeMelT ,!!~es longieaudatu~-Synarpheus fritzmuel1eri, Thor f10ridanus, In addition to the organisms which pet"rofisthesarmatu.;;-a;ld .itleast ei ght move from the mangroves to the mud flats, species of crabs. The following species there is a small qroup which uses the were found only on mangrove roots and not substrate adjacent to mangroves for both on seawalls: Turitel1a sp., Me10ngena habitat and feeding. In the Whitewater coro~, Anachis semip1icata, Bulla Bay region, four crabs exploit the inter­ striata, Hypselodoris sp., Area imbricata, tidal muds from the safety of burrows: Carditamera fl ori dana, Pseudoi rus !l.P.!..ca, Uca pugi1ator, Q. speciosa, Q. thayeri, and Martesia striata. and Eurytium limosum (Tahb et a1. f962). In low sal inity mangrove forests of south Tabb et a1. (1962) and Odum and Heald Florida, the crayfish, Procambarus alleni, (1972) reported a variety of invertebrates is a dominant member of the burrowing, associated with prop roots in the Hhite­ benthic community (Hobbs 1942) as is the water Bay region. Although many species crab, Rhithropanopeus harrisii (Odum and coincide with Courtney's (1975) list, Heald 1972). Both organisms are found in there are also significant differences due a remarkable number of fi sh stomachs. to the lower salinities in this region. It is probably safe to conclude that prop The benthic fauna and infauna of root commun iti es vary somewhat from site creek and bay bottoms near mangrove to site in response to a number of factors forests are highly variable from one

48 location to the next. Many of these exceed 500/m2 and average 100 to 200/m2 organisms, particularly the deposit and (Heald, unpublished data). Cerithidea is filter feeders, benefit from particulate found largely in association with black organic matter originating from mangrove mangroves ~d can reach densities of at litter fall (Odum and Heald 1972, 1975b). 1east 400/m • Tabb and Manning (1961) and Tabb et a1. (1962) present lists and discussions of many of the benthic invertebrates adjacent 6.4 WATER COLUMN COMMUNITY to mangrove areas of Whitewater Bay. Weinstein et al. (1977) compared the ben­ This section is embarrassingly short; thic fauna of a mangrove-lined creek and a the reasons for thi s brevity are (I) the nearby man-made canal on Marco Island. paucity of research on zoopl ankton in They found (1) the mangrove fauna to be Florida mangrove-dominated areas and (2) more di verse than the canal fauna and (2) our i nabi 1i ty to ·di scover some of the work a higher diversity of organisms at the which undoubtedly has been done. Davis mouths of mangrove creeks than in the and WilliaMS (1950) are usually quoted as "heads" or upstream ends. Courtney (1975) the primary reference on Florida mangrove­ found the same pattern of upstream associated zooplankton, but their paper decreases in diversity, presumably in only lists zoop1ankters collected in two response to decreasing oxygen concentra­ areas. Zooplankton near mangroves are tions and increasingly finer sediments. probably no di fferent from those found in other shallow, inshore areas in south Finally, the irregularly flooded sub­ Florida. Based on Davis and Williams strates in the center of mangrove forests (1950) and Reeve (1964), we can hypothe­ contain a small but interesting assemblage size that the community is dominated by of invertebrates. The litter layer, copepod species of genus Acartia, particu­ composed largely of mangrove leaves, evi­ larly Acartia tonsa. In addition, we dently includes a variety of nematodes. could expect a few other calanoid cope­ Due to the usual taxonomic difficulties in pods, arrow worms (Sagitta spp.), many identifying nematodes, complete species fish, polychaete and crustacean larvae and I ists do not exist for mangrove forests; eggs. Another component of the "pl ankton~' however, many species and individuals are particularly at night, are benthic associated with the decaying leaves amphipods, mysids, and isopods which leave (Hopper et al. 1973). In addition to the bottom to feed (personal observati on). nematodes, the wetter sections of the swamp floor can contain mosquito and other Plankton are not the only inverte­ insect larvae, po1ychaetes, harpacticoid brates in the water column. Swimming copepods, i sopods, and amphi pods. crabs, such as the blue crab, Callinectes Simberloff (1976) lists 16 species of sapidus, are plentiful in most estuarine insects associated with the muddy floor of mangrove regi ons of south Flori da. Other mangrove forests. Roaming across the swimming crustaceans include the caridean forest floor during low tide are several shrimp (Pa1aemonetes spp. and Peri­ crustaceans including the mangrove tree c 1 ; menes-spp-:T,-the-snappi n g s h r; mp crab, Aratus pisonii, crabs of the genus rAlphe~ spp.), and the penaeid shrimp Sesarma, and the pulmonate gastropods, (Penaeus spp). All of these swimming ~~l~~£~~ coeffeus and Cerithidea crustaceans spend considerable time on or sca1ariformis. Both snails clearly have in the benthos and around mangrove prop the ability to graze and consume recently roots. From the economic point of view, fallen leaves (personal observation). the pi nk shri mp, Penaeus duorarum, is With favorable conditions (relatively fre­ probably the most important species asso­ quent tidal inundation plus the presence ciated with mangrove areas (see discussion of red mangroves) Me1ampus populations can in section 11).

49 CHAPTER 7. COMMUNITY COMPONENTS - FISHES

Of the si x mangrove community types Site characteristics and sampling methods discussed in section 1.5, fishes are an for these community types are summarized important component of four: (1) basin in Appendix A. Nomenclature and taxonomic forests, (2) riverine forests, (3) fringe order follow Bailey et a!. (1970). forests, and (4) overwash island forests. For convenience we have divided fringe forests into two sub-components: (a) 7.1 BASIN MANGROVE FORESTS forests which fringe estuarine bays and lagoons and (b) forests which fringe The infrequently flooded pools in the oceanic bays and lagoons. This division black mangrove-dominated zone provide an is necessa ry because the fi sh communit i es extreme habitat which few species of di ffer markedl y. fi shes can tolerate. The waters are darkly stained with organic acids and Mangroves serve two distinct roles tannins leached from the thick layer of for fishes and it is conceptually impor­ leaf litter. Dissolved oxygen is tant to distinguish between them. First, frequently low (1-2 ppm) and hydrogen the mangrove-water interface, generally sulfide is released from the sediments red mangrove prop roots, afford a rela­ following physical disturbance. Salini­ tively protected habitat which is particu­ ties are highly variable ranging from larly suitable for juvenile fishes. totally fresh to hypersaline. The fish Secondly, mangrove leaves, as discussed in families best adapted to this habitat are section 3.6, are the basic energy source the euryhaline cyprinodonts (killifishes) of a detritus-based food web on which many and the poeciliids (livebearers). The fi shes are dependent. The habitat val ue killifishes include Fundulus confluentus of mangroves can be considered strictly a (Heald et al. 1974), Rivulus marmoratus function of the area of interface between (M. P. Weinstein, Va. Commonwealth Univ., the water and the mangrove prop roots; it Richmond, Va.; personal communication is an attribute shared by all four types 19B1), Floridichthys carpio, and of mangrove communities. The importance Cyprinodon variegatus (Odum 1970). The of the mangrove detritus-based food web is poecillids include Poecilia latipinna dependent on the re1ati ve contri buti on of (Odum 1970) and, the most common, Gambusia other forms of energy in a given environ­ affinis (Heald et a!. 1974). While the ment, including phytoplankton, benthic species richness of fishes in this habitat algae, sea grass detritus, and terrestrial is low, the densities of fish are often carbon sources. Figure 11 provides a very hi gh. Weinstein \fers. comm.) has di agrammat i c representation of the rela­ recorded up to 3B fish/m • tive positions along a food web continuum of the four mangrove communities. A11 of these fi shes a re permanent residents, completing their life cycles in Fishes recorded from mangrove habi­ this habitat. They feed primarily on tats in south Florida are listed in Appen­ mosquito larvae and small crustaceans such dix B. Although the fish communities are as amphipods which, in turn, feed on man­ discussed separately below, they have been grove detritus and algae. These small combined into certain categories in Appen­ fi shes enter coastal food webs when they dix B; fishes from mangrove basins and are flushed into the main watercourses riverine forests have been combined under during high spring tides or following the heading of tidal streams; fishes from seasonally heavy rains. Here they are fringing forests along estuarine bays and eaten by numerous pi sci vorous fi shes i n­ lagoons are listed under the heading of cluding snook, ladyfish, tarpon, gars, and estuarine bays; fishes from oceanic bays mangrove snappers. The alternate energy and lagoons have been listed under oceanic pathway for fi shes of the bl ad mangrove bays. Si nce no surveys have been basin wetlands occurs when the pools published specifically relating to over­ shri nk duri ng dry weather, the fi shes are wash island forests, there is no 1 isting concent rated into sma 11 er a rea s, and are for this community type in Appendix B. fed-upon by various wading birds including 50 ., - , ..... ,- - IJ '"0 z u " 0 w ~ '" /f"":<0>;­ ... " 21 ~'i,P(/ /:;__ , u --.,- -'- 9 '-"""":": u ~ 0.. ------.',-, - '-,---:.. ... '~....o---...<'_~,-_/ . ~--,7"::-' ,. II: ~ ... 0 w ..'"0 " ~ u ;< '"

BLACK MANGROVE RIVERINE FRINGING ESTUARINE BAY OCEANIC BAY OVERWASH ISLAND. BASIN FOREST COMMUNITY FRINGING COMMUNITY FRINGING COMMUNITY FLORIDA KEYS

HIGH SALINITY VARIABILITY LOW

HIGH RELATIVE IMPORTANCe of MANGROVE DETRITUS in FOOD WEB LOW

MUDDY, HIGH ORGANIC SUBSTRATE LARGELY SANDY

GRADIENTS

Figure 11. Gradient of mangrove-associated fish communities showing representative species. Fish are not drawn to scale. 1 = rivu1us, 2 = mosquitofish, 3 = marsh killifish, 4 = ladyfish, 5 = striped mullet, 6 = yel10wfin mojarra, 7 = juvenile sheepshead, 8 = tidewater si1versides, 9 = sheepshead minnow, 10 = silver perch, 11 = pigfish, 12 = b1ackcheek tonguefish, 13 = scrawled cowfish, 14 = fringed pipefish, 15 = fringed filefish, 16 = lemon shark, 17 = goldspotted killifish, 18 = southern stingray, 19 = juvenile schoolmaster, 20 = juvenile tomtate, 21 = juvenile sergent major. See Appendix B for scientific names. herons, ibis and the wood stork (Heald et tidal streams primarily on feeding forays. a1. 1974). Examples include the jewfish, E ine helus itajara, the stingrays (Dasyatidae , the need1efishes (Be10nidae), the jacks 7.2 RIVERINE FORESTS (Carangidae), and the barracuda, Sphyraena barracuda. Other seasonal movements of Tidal streams and rivers, fringed fishes appear to be temperature related. largely by red mangroves, connect the Tabb and Manning (1961) documented move­ freshwater marshes of south Florida with ments of a number of species from shallow the shallow estuarine bays and lagoons inshore waters to deeper water during (Figure 12). Few of these streams have times of low temperature stress. The been studied thoroughly. The exception is lined sole, the hogchoker, the bighead the North River which flows into White­ searobin, and the striped mullet, for water Bay and was studied by Tabb (1966) example, are much less frequently caught and Odum (1970). Springer and Woodburn in winter in shallow inshore waters. (1960) collected fishes in a bayou or tidal pass connecting Boca Ciega Bay and A third type of seasonality of fish Old Tampa Bay. Carter et a1. (1973) populations in the tidal rivers is related reported on the fishes of two tidal to life cycles. Many of the fish which streams entering Fahkahatchee and Fahka utilize the tidal stream habitat do so Union Bays. Nugent (1970) sampled fishes only as juveniles. Thus, there are peaks in two streams on the western shore of of abundance of these species following Biscayne Bay. Characteristics of these offshore spawning when larval or juvenile areas and sampling gear used by the inves­ forms are recruited to the mangrove stream tigators are summarized in Appendix A. habitat. In general, recruitment occurs in the late spring or early summer fol­ These tidal streams and associated lowing late winter and spring spawning riverine mangrove forests exhibit extreme offshore or in tidal passes (Reid 1954). seasonal variability in both physical Numerous species are involved in this life characteristics and fish community compo­ cycle phenomenon including striped mullet, sition. Salinity variations are directly grey snapper, sheepshead, spotted sea related to changes in the make-up of the trout, red drum, and silver perch. fish assemblage. During the wet season (June - November), salinities fall The only estimate of fish standing throughout the water courses and, at some crop from ti dal stream habitats is that of locations in certain heavy runoff years, Carter et a1. (1973). They recorded 27 become fresh a 11 of the way to the mouth species weighin~ 65,B91 g (wet w~.) from (Odum 1970). Opportuni sti c freshwater an area of 734 m or about 90 gfm. This species, which are normally restricted to is probably an overestimate since an un­ the sawgrass and black needle rush marshes known portion of the fish community had of the headwaters, invade the mangrove moved from the flooded lowlands to the zone. These include the Florida gar, stream on the ebb tide; sampling occurred Lepisosteus Ql~!.lrhincus; several at low tide in October. Nonetheless, this centrarchid sunfishes of the genus Lepomis is an indication of the high fish standing and the largemouth bass, Micropterus crop which this mangrove-associated habi­ sa1moides; the freshwater catfishes, tat can support. The number of species Ictalurus nata1is and Noturus gyri nus; and reported from individual tidal streams the ki1lifishes normally considered annually ranges from 47 to 60 and the freshwater inhabitants such as Lucania total from all tidal streams in southwest goodei and Rivulus marmoratus. Florida is 111 species (Appendix B). During the dry season (December to The food webs in these riverine man­ early May) salinities rise as a result of grove ecosystems appear to be predomi­ decreased freshwater runoff and continuing nantly mangrove detritus-based, although evaporation. Marine species invade the 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 (Ariidae), gobies (Gobiidae), porgys link between the mangrove leaf and higher (Sparidae), mullets (Mugi1idae), drums order consumers is provided by micro­ (Sciaenidae), and anchovies (Engrau1idae). organisms (fungi, bacteria, Protozoa) The mangrove-lined streams and associated which colonize the decaying leaf and con­ pool s are important nursery areas for vert them into a relatively rich protein several marine and estuarine species of source (Odum 1970; Odum and Heald 1975a). gamefish. The tarpon, Megalops atlantica, These decaying leaf fragments with asso­ snook, Centropomus undecima1is, and 1ady­ ciated microorganisms are fed upon by a fish, Elops saurus, utilize these areas group of omnivorous detritivores including from the time they reach the estuary as amphipods, mysids, cumaceans, ostracods, post-larvae, having been spawned offshore. chironomid larvae, harpacticoid and Gray snapper, Lutj anus .9.ri seus, calanoid copepods, snapping shrimp, sheep shead, Archosargus probatocephalus, caridean and penaeid shrimp, a variety of spotted seatrout, Cvnoscion nebulosus, and crabs, filter-feeding bivalves, and a few red drum, Sciaenops uce1lata, are re­ speci es of fi shes (Odum 1970; Odum and cruited to grass beds of shallow bays and Heald 1972; Odum and Heald 1975b). These lagoons as post-larvae and enter the detritivores, in turn, are consumed by a mangrove-lined streams for the next sever­ number of small carnivorous fishes, which al years (Heald and Odum 1970). Of these in turn, are consumed by larger species, only the spotted seatrout prob­ piscivorous fishes. The concept of man­ ably spawns in the estuary (Tabb 1966). 9rove trophic structure is a1 so discussed Other species of commercial or game impor­ in section 3.6. See Appendix B for tance which use the riverine fringing species specific dietary information. habitat include creval1e jack, gafftopsail catfish, jewfish, striped mojarra, barra­ The tidal creeks studied by Nugent cuda, Atlantic thread herring, and yel10w­ (1970) on the western shore of Biscayne fin menhaden (Odum 1970). Bay differ from the previously discussed streams in the Everglades estuary. The mouths of the Biscayne Bay creeks have 7.3 FRINGING FORESTS ALONG ESTUARINE BAYS dense growths of sea grasses which con­ ANO LAGOONS tribute sea grass detritus. The salini­ ties are considerably greater and the Mangrove-fringed estuarine bays and streams are located only a few kilometers lagoons are exemplified by the Ten from coral reefs, whi ch are 1argely absent Thousand Is1 ands area and Whitewater Bay. on Florida's west coast, at least close to Quantitative fish data are available from shore. As a result, 23 species listed in Fahkahatchee Bay (Carter et al. 1973; Appendi x B were captured by Nugent (1 970) Yokel 1975b; Seaman et a1. 1973), Fahka and are not recorded from riverine man­ Union Bay (Carter et a1. 1973), Rookery grove habitat on the west coast of Bay (Yokel 1975a), the Marco Island Florida. Examples include several of the Estuary (Weinstein et a1. 1977; Yokel grunts (Pomadasyidae), the gray trigger­ 1975a), and Whitewater Bay (Clark 1970). fish, Ba1istes capriscus, the barbfish, Individual site characteristics are Scorpaena brasiliensis, the scrawled box­ summarized in Appendix A. All except fish, Lactophrys guadricornis, and the Fahka Union Bay contain significant snappers, Lutjanus apodus and I. synagri s. amounts of sea 9rasses. Macroa19ae domi­ nate the benthic producers of Fahka Union Riverine mangrove communities and Bay. Studi es by Reid (1954) and Ki 1by associated tidal streams and rivers are (1955) near Cedar Key, F10rida,were not typified by the following families of included in our summary because man9roves fishes: kil1ifishes (Cyprinodontidae), are sparse in this area and no mention of 1ivebearers (Poeciliidae), si1versides man9rove collecting sites were made by (Atheri ni dae), moj arras (Gerreidae), tar­ these authors. Studies of Ca100sahatchee pon (Elopidae), snook (Centropomidae), Bay (Gunter and Hall 1965) and of snappers (Lutjanidae), sea catfishes Charlotte Harbor (Wang and Raney 1971)

54 were omitted because the areas studied collected in these mangrove-fringed bays have been highly modified and because data and lagoons (Appendix B). from many habitats were pooled in the final presentation. In none of these studies were the fishes specifically utilizing the fringing All of the bays reviewed in our sum­ mangrove habitat enumerated separately maries are fringed by dense growths of red from those collected in the bay as a mangroves and all contain small mangrove whole. The collections were most often at islets. Carter et al. (1973), in their open water stations easily sampled by studies of Fahkahatchee and Fahka Union otter trawl. Carter et al. (1973) had two bays, esti mated that 57% to 80% of the shore seine stations adjacent to mangroves total energy budget of these two bays is but the data were pooled for pub1i cation. supported by exports of particulate and Of the four stations in Rookery Bay sam­ dissolved organic matter from the man­ pled by Yokel (lg75a), one was immediately groves within the bays and inflowing tidal adjacent to the fringing mangrove shore­ streams. lugo et al. (1 980) esti mated 1i ne and had moderate amounts of sea that the mangroves surrounding Rookery Bay grasses. provi de 32% of the energy base of the heterotrophic community found in the bay. The typical pattern which emerges from many estuarine studies is that rela­ Salinities in these bays tend to be tively few fish species numerically domi­ higher than in the tidal streams and nate the catch. This is certainly true in rivers and the fish assemblages reflect mangrove-fringed estuaries. In Rookery both this feature and the added habitat Bay (Yokel Ig75a) six species comprised dimension of sea grass and macro algae 88% of the trawl-catchable fishes, in beds. Truly freshwater species are rare Fahkahatchee Bay seven species comprised in these communi ties and a propo rti ona lly 97% of the catch from three capture greater percentage of marine visitors is techniques (Carter et al. 1973), and in present. The dominant fish families of the Marco Island estuary 25 species com­ the benthic habitat include drums prised 97% of the trawl-catchable fishes (Sciaenidae), porgys (Sparidae), grunts (Weinstein et al. 1977). (Pomadasyidae), mojarras (GerreidaeJ, snappers (lutjani dae), and mull et (Mugi 1i­ like tidal river and stream communi­ dae). Other fami1es with sizeable contri­ ties, these shallow bays serve as nur­ butions to the benthic fauna include pipe­ series for numerous species of estuarine­ fishes (Syngnathidae), flounder (Bothi­ dependent fishes that are spawned off­ dae), sole (Soleidae), searobins (Trig1i­ shore. Based on the distribution and dae), and toadfishes (Batrachoididae). abundance of juveni Ie fishes of all spe­ cies in six habitats, Carter et al. (1973) Numerically abundant fishes of the ranked the mangrove-fringed bays as the mid and upper waters include anchovies most important nursery grounds; the ti dal (Engrau1idae), herrings (Clupeidae) and streams were a close second. Shallow bays need1efishes (Belonidae). At all loca­ and tidal streams provide safe nurseries tions studied, the benthic fauna was domi­ due to seasonally abundant food resources nated by the pinfish, lagodon rhomboides, and the low frequency of large predators the silver perch, Bairdie11a chrysura, the (Carter et al. 1973; Thayer et al. 1978). pigfish, Orthopristis chrysoptera, and the The relative lack of large predaceous mojarras, Eucinostomus ~ and I. fishes is probably due to their general argenteus. The most common ml dwater and inability to osmoregu1ate in waters of low surface species include the two anchovies, and/or fl uctuating sal inity. Anchoa mitchilli and ~. hepsetus, and two clupeids, Brevoortia smithi and Harengu1a As in tidal streams, the peak abun­ pensaco1ae. The total number of species dance of juvenile and larval fishes in the recorded in the individual studies ranged bays is in spring and early summer (Reid from 47 to 89; a total of 117 speci es was 1954). In general, the highest standing

55 crops and the greatest species richness of Based pri mari lyon habitat desi gna­ fishes occur in the late summer and early tions of Voss et al. (1969), the fishes of fall (Clark 1970). Fish densities decline Biscayne Bay can be characterized as to in the autumn and winter as many fishes preferred habitat. Of the three main move to deeper waters. habitat types, (1) rock/coral/seawall, (2) grassbed/tida1 flat, and (3) mangrove, the grassbed/tida1 flat ranked first in fish 7.4 FRINGING FORESTS ALONG OCEANIC BAYS speci es occurrences. One hundred and AND LAGOONS twenty- two of 156 speci es (79%) are known to occur in this environment. Mangrove-fri nged "oceani c" bays and Rock/coral/seawall habitats were fre­ lagoons are exemplified by Porpoise Lake quented by 49 species (32%) and mangroves in eastern Florida Bay (Hudson et al. are known to be utilized by 54 species 1970), western Florida Bay (Schmidt 1979), (35%) of the total fish species recorded southern Biscayne Bay (Bader and Roessler from this bay. 1971), and Old Rhodes Key Lagoon in eastern Biscayne Bay (Holm 1977). Charac­ teristics of these sites are summarized in 7.5 OVER WASH MANGROVE ISLANDS Appendix A. Compared to the mangrove­ fringed bays discussed in the previous In terms of fish-related research, section, these envi ronments generally ex­ these communities are the least studied of hibit clearer water, sandier substrates, all mangrove community types in south and higher and less variable salinities. Florida. They are typified by the 10w­ Closer proximity to the Florida reef 1yi ng mangrove-covered is1ands that occur tract, the Atlantic Ocean, and the Gulf of in the Florida Keys and Florida Bay and Mexico results in a larger potential pool may be overwashed periodically by the of fish species. These four locations tides. Examples include Shell Key, Cotton have produced reports of 156 fi sh speci es Key, and the Cowpens. Islands of this (Appendix B). type extend southwest from the Florida mai n1 and through the Marquesas. The Dry Mangrove fringes make up a relatively Tortugas lack well-developed mangrove com­ small proportion of these environments; munities although stunted trees are found accordingly, their contribution to the bay (Davi s 1942). food webs is probably not very 1arge. Bader and Roessler (1972) esti mated that These islands are the most oceanic of the fringing mangrove community contrib­ any of the mangrove communiti es di scussed. utes approximately 1% of the total energy They are characterized by relatively clear budget of southern Biscayne Bay; they water (Gore 1977) and are largely free of considered only mainland mangroves and did the freshwater inflow and salinity varia­ not include the small area of mangrove tions which characterize other Florida islands. The main ecological role of the mangrove communities to varying degrees. fringing mangroves in this type of en­ Numerous statements exist in the litera­ vironment is probably twofold. First, tu re acknow1 edgi ng the frequent proxi mity they increase the habitat diversity within of mangrove islands to coral reefs and sea an otherwise relatively homogeneous bay grass beds (McCoy and Heck 1976; Thayer et system. Second, they provide a relatively a1.1g7B). Olsen et al. (1973) working in protected habitat for juvenile fishes (and the U.S. Virgin Islands, found 74% to 93% certain invertebrates) that later move to overlap in the fish species composition of more open water or coral reef communities. fringing coral reefs and shallow mangrove­ The second role is analogous to one of the fringed oceanic bays. Voss et al. (1969) ecological roles of sea grass communities listed fish species that were collected (see Zi eman, in prep.) a 1though the fi sh from all three types of communities: speci es invo1 ved may be different. fringing mangroves, coral reefs and sea

56 grass beds in Biscayne Bay, but there mangrove islands on the rlSlng tide. appears to have been no systematic survey Included in this group are sharks, tarpon, of the fish assemblage characteristic of jacks, snook, bonefish and barracuda. the mangrove-covered or mangrove-fringed Florida Keys. No one has quantified the faunal connections which we hypothesize 7.6 GRADIENT OF MANGROVE COMMUNITY exist between the mangroves and sea INTERACTIONS grasses and between the mangroves and coral reefs. Mangrove commun i ties occur under a wide range of conditions from virtually In the absence of published data from freshwater at the headwaters of tidal the mangrove key communities, only tenta­ streams to nearly oceanic conditions in tive statements can be made. In general, the Florida Keys. Attempting to present a we expect that while mangrove islands single list of fish characteristic of serve as a nursery area for juvenile mangrove environments (Appendix B) can be fishes, this function is limited largely misleading. For this reason we presented to coral reef and marine inshore fishes the concept of a continuum or complex and not the estuarine-dependent species gradient in Figure 11 and have followed that we have discussed previously. The that scheme throughout section 7. The latter (juvenile snook, red drum, spotted gradient stretches from seasonally fresh seatrout) appear to require relatively low to oceanic conditions, from highly varia­ salinities not found in association with ble salinities to nearly constant salini­ most of the overwash is 1ands. Casu a 1 ty, from muddy and 1i mestone subst rates to observation around the edges of these sandy substrates, from dark-stained and islands suggests that characteristic sometimes turbid waters to clear waters, fishes include the sea bass family (Ser­ and from food webs that are predomi nantly ranidae), triggerfishes (Balistidae), mangrove detritus-based to food webs based snappers (Lutjanidae), grunts (Poma­ pri mari lyon other energy sources. Cl ear­ dasyidae), porgies (Sparidae)\parrotfishes ly, there are other gradients as one moves (Scaridae), wrasses \LabridaeJ, bonefishes from north to south in the State of (Albulidae), jacks (Carangidae), damsel­ Florida. At the northern end of the fishes (Pomacentridae), and surgeonfi shes State, temperatures are more variable and (Acanthuri dae); many of these fi shes occur seasonally lower than in the south. Sedi­ on or are associ ated with coral reefs. We ments change from predominantly silicious also suspect that considerable overlap in central and north Florida to predomi­ occurs in the fish assemblage of these nantly ca rhonate in extreme south Flori da. mangrove islands and sea grass communi­ Nevertheless, the complex gradient shown ties; examples include puffers (Tetrao­ in Figure 11, while greatly simplified for dontidae), pipefishes (Syngnathidae), go­ graphic purposes, suggests that charac­ bies (Gobiidae) and scorpionfishes (Scor­ teristic fish assemblages replace one paenidae). Stark and Schroeder (1971) another along a 9radi ent of changi ng suggested that juveni le gray snapper, physical and biogeographic conditions. which use the fringing mangroves of the Such a concept is useful in understanding keys as shelter during the day, forage in the factors controlling the composition of adjacent sea grass beds at night. In the fish assemblages associated with mangroves absence of salinity barriers, predatory of the four major community types in south fishes probably enter the fringes of these Florida.

57 CHAPTER B. COMMUNITY COMPONENTS - AMPHIBIANS AND REPTILES

Food habits and status of 24 species a variety of submerged aquatic plants and of turtles, snakes, lizards, and frogs of sea grasses; recent evidence has shown the Florida mangrove region are given in that they al so feed on mangrove roots and Appendix C. Any of three criteria had to 1eaves (Ernst and Barbour 1972). The be met before a species was included in Atlantic ridley's preferred habitat is this table: (1) a di rect reference in "shallow coastal waters, especially the the literature to mangrove use by the mangrove-bordered bays of the southern species, (2) reference to a species as half of the peninsula of Florida" (Carr being present at a particular geographical and Goin 1955). Hawksbill turtles feed on location within the mangrove zone of a variety of plant materials including Florida, and (3) North American species mangrove (especially red mangrove), recorded from mangroves in the West Indies fruits, leaves, wood, and bark (Ernst and or South America, but not from Florida. Barbour 1972). This last criterion assumes that a species which can utilize mangroves outside of Three species in the genus Anolis Florida will be able to use them in have been reported from Florida mangroves: Florida. Ten turtles are listed of which the green anole, the cuban brown anole, four (striped mud turtle, chicken turtle, and the Bahaman bank anole. All are Florida red-bellied turtle, and softshell arboreal lizards that feed on insects. turtle) are typical of freshwater. Two The green anole is widespread throughout (mud turtle and the ornate diamondback the Southeastern United States and is not terrapin) are found in brackish water and at all dependent on mangrove swamps. The the remai nder (hawksbi 11, green, logger­ other two species have much more head, and Atlantic ridley) are found in restricted distributions in the United rna ri ne waters. States and are found only in south Florida. They also are not restricted Freshwater species usually occur in to mangrove ecosystems. Of the six the headwater regions of mangrove-lined species of snakes listed, the mangrove river systems. All four freshwater water snake (Figure 13) is most dependent species are found in habitats other than upon mangrove habitat~ mangrove swamps including streams, ponds, and freshwater marshes. The brackish Two important speci es of repti 1es water species are found in salt marshes in found in mangrove swamps are the American addition to mangrove swamps. Mangroves, all i gator and the Ameri can crocodi 1e. The however, are the principal habitat for the alligator is widespread throughout the ornate diamondback terrapin (Ernst and Southeastern United States and is only Barbour 1972). Carr and Goi n (1955) incidentally found in low salinity sec­ listed two subspecies of the diamondback: tions of Florida mangrove areas (Kush1 an Malaclemys terrapin macrospilota and ~. 1. 1980). The American crocodile is rare; rhizophorarum. Malaclemys terrapin macro­ historically its distribution was centered spilota inhabits the south\~est and south­ in the mangrove-dominated areas of the ern coasts, and ~. 1. rhizophorarum is upper and lower Florida Keys (particularly found in the Florida Keys. The two sub­ Key Largo) and the mangrove-l ined shore­ species intergrade in the region of north­ lines and mud flats along the northern ern Florida Bay. edge of Florida and Whitewater Bays (Kushlan 19BO). Mangroves appear to be All four of the marine turtles are critical habitat for this species. Its associated with mangrove vegetation at range has shrunk considerably in south some stage of their lives. Loggerhead and Florida since the 1930's, even though green turtles are apparently much less Florida Bay was added to Everglades dependent on mangroves than the remai ni ng National Park in 1950 (Moore 1953; Ogden two, although we strongly suspect that 1978). Much of the decrease in range is recently hatched loggerheads may use man­ due to increased human activity in the grove estuaries as nursery areas. Green Florida Keys. The remaining population turtles are generally believed to feed on centers of the American crocodile are in 58 Figure 13. The mangrove water snake. Nerodia fasciata compressicauda. curled on a red mangrove prop root. Photograph by David Scott.

59 northern Flori da Bay and adjacent coasta1 amphibians, to our knowledge, have been swamps and the northern end of Key Largo recorded in Florida mangrove swamps (Ap­ (Ogden 1978; Kushlan 1980). The species pendi x C). Thi sis due to two factors: uses a variety of habitats for nesting in (1) lack of detailed surveys in low sa­ the Florida Bay region including open l i nit.y swamps and (2) the i nabi 1i ty of hardwood thickets along creek banks, most amphibians to osmoregulate in salt hardwood-shrub thickets at the heads of water. No doubt, several additional sand-shell beaches, and thickets of black speci es occur in the freshwater -domi nated mangroves behind marl banks (Ogden 1978). hammock and basin mangrove communities On Key Largo the crocodile locates its inland from the coast. Possible addi­ nests on creek and canal banks in red and tional species include: the eastern black man9rove swamps (Ogden 1978). Man­ narrow-mouthed toad, Gastrophryne caro­ grove areas thus appear to be important in linensis, the eastern spadefoot toad, the breeding biology of this endangered Scaphiopus holbrooki, the cricket frog, species. Acris gryllus, the green tree frog, ~ cinerea, and the southern leopard frog, Interestingly, only three species of Rana utricularia.

60 CHAPTER 9. COMMUNITY COMPONENTS - BIRDS

9.1 ECOLOG ICAL RELA11 ONSH I PS residents, a significant fraction of the avifauna communi ty was omi tted. Because mangroves present a more di verse st ructural habitat than most Based on information 91eaned from the coastal ecosystems, they should harbor a literature, we have compiled a list of 181 greater variety of bird1ife than areas species of birds that use Florida mangrove such as salt marshes, mud fl ats, and areas for feeding, nesting, roosting, or beaches (MacArthur and MacArthur 1961). other activities (Appendix D). Criteria The shallow water and exposed sediments for listing these species is the same as below mangroves are available for probing that used for listing reptiles and amphi­ shorebirds. Longer-legged wading birds bians (see Chapter 8 of this volume). utilize these shallow areas as well as deeper waters along mangrove-lined pools Often references were found stating and waterways. Surface-feeding and diving that a given species in Florida occurred birds would be expected in similar areas in "wet coastal hammocks", "coastal wet as the wadi ng bi rds. The major di fference forests" or the 1 ike, without a specific between mangrove swamps and other coastal reference to mangroves. These species ecosystems is the availability of the were not included in Appendix D. Thus, trunks, limbs, and foliage comprising the this list is a conservative estimate of tree canopy. This enables a variety of the avifauna associ ated with Flori da man­ passerine and non-passerine birds, which grove swamps. Sources for each listing are not found commonly in other wetland are provided even though many are redun­ areas, to use mangrove swamps. It a1 so dant. Food habit data are based on Howell allows extensive breeding activity by a (1932) and Martin et a1. (1951). Esti­ number of tree-nesti ng bi rds. mates of abundance were derived from bird lists published by the U.S. Fish and The composition of the avifauna com­ Wildlife Service for the J.N. "Ding" munity in mangrove ecosystems is, in fact, Darling National Wildlife Refuge at highly diverse. Cawkell (1964) recorded Sanibel Island, Florida, and by the Ever­ 45 species from the mangroves of Gambia glades Natural History Association for (Africa). Haverschmidt (1965) reported 87 Everglades National Park. Frequently, species of birds which utilized mangroves species were recorded from mangrove swamps in Surinam (S. America). Ffrench (1966) at one location, but not the other. listed 94 species from the Caroni mangrove swamp in Trinidad while Bacon (1970) found We have divided the mangrove avifauna 137 in the same swamp. In Malaya, Nisbet into six groups based on similarities in (1968) reported 121 speci es in mangrove methods of procuri ng food. These groups swamps and Field (1968) observed 76 from (guilds) are the wading birds, probing the mangroves of Sierra Leone (Africa). shorebirds, floating and diving water­ birds, aerially-searching birds, birds of Use of mangrove ecosystems by bi rds prey, and a rborea1 bi rds. Thi s 1ast group in Florida has not been recorded in de­ is something of a catch-all group, but is tai 1. Ninety-two species have been ob­ composed mai n1y of bi rds that feed and/or served in the mangrove habitat of Sanibel nest in the mangrove canopy. Island, Florida (L. Narcisse, J.N. "Ding" Darling Nat1. Wildlife Refuge, Sanibel Is., Fla.; personal communication 1981). 9.2 WAD I NG BIRDS Robertson (1955) and Robertson and Kush1an (1974) reported on the entire breeding Herons, egrets, ibises, bitterns, and bird fauna of peninsular south Florida, spoonbills are the most conspicuous group including mangrove regions. Based on of birds found in mangroves (Figure 14) 1i mi ted su rveys, these authors reported and are by far the most studied and best only 17 species as utilizing mangroves for understood. Eighteen species (and one breeding purposes. Because their studies important subspecies) are reported from did not consider migrants or non-breeding south Flori da mangroves. 61 Figure 14. A variety of wading birds feeding in a Inan9rove-lined pool near Flamingo, Florida. Photograph by David Scott.

62 Mangrove swamps provide two functions b). Mangroves are critically important for wadi ng bi rds. Fi rst, they funct i on as for the survival of the white ibis popula­ feedi ng grounds. Two-thi rds of these tion even though they appear to be species feed almost exclusively on fishes. utilized to a lesser extent than fresh­ Although much of their diet is provided by water habitats. This pattern of larger freshwater and non-mangrove mari ne areas, but less stable breeding colonies using all of them feed frequently in mangrove inland marshes and smaller but more stable swamps. White ibis feed predominantly on colonies using mangroves is also charac­ crabs of the genus Uca when feeding in teristic of heron populations (Kushlan and mangroves (Kushlanand Kushlan 1975; Frohring, in prep.). Kush 1an 1979). Mollusk sand i nve rtebrates of the sedi ments are pri nci pal foods of Table 5 gives the number of active the roseate spoonbi 11 although some fish nests observed in mangrove regi ons du ri ng are eaten (Allen 1942). Yellow-crowned the 1974-75 nesting season and the percen­ night herons and American bitterns eat tage this represents of the enti re south crabs, crayfi sh, frogs, and mi ce in addi­ Florida breeding population for the nine tion to fishes. Snails of the genus most abundant species of waders and three Pomacea are fed upon al most excl usi vely by associated species. The dependence of the limpkin. The sandhill crane is an roseate spoonbills, great blue herons, anomaly in this group since a majority of Louisiana herons, brown pelicans, and its food is vegetable matter, especially double-crested cormorants on mangrove roots and rhi zomes of Cyperus and regions is evident. Nesting by the red­ Sagi ttari a. Its use of mangroves is dish egret was not quantified during this probably minimal, occurring where inland study although Kushlan and White (1977a) coastal marshes adjoin mangroves (Kushlan, indicated that the only nests of this unpubl. data). The remaining 12 species species which they saw were, in fact, in are essentially piscivorous although they mangroves. Fu rther observations i ndi cate differ somewhat in the species and sizes that this species nests in mangroves ex­ of fishes that they consume. clusively (Kushlan, pers. comm.). Similar­ ly, the great white heron is highly depen­ Mangrove swamps al so serve as dent upon mangroves for nesting; they use breeding habitat for wading bi rds. With the tiny mangrove islets which abound the exception of the limpkin, sandhill along the Florida Keys and in Florida Bay crane, and the two bitterns, all wading (Howell 1932). bird species in Appendix D build their nests in all three species of mangrove During many years the Everglades trees (Maxwell and Kale 1977; Gi rard and population of wood storks is known to nest Taylor 1979). The speci es often aggregate almost solely in mangroves (Ogden et al. in large breeding colonies with several 1976); this population comprises approxi­ thousand nesting pairs (Kushlan and White mately one-thi rd of the total south 1977a). The Louisiana heron, snowy egret, Florida population. Successful breeding and cattle egret are the most numerous of all these mangrove nesters is un­ breeders in south Florida mangroves (based doubtedly correlated with the abundant on data in Kushlan and White 1977a). supply of fishes associated with man­ groves. Meeting the energetic demands of In wet years over 90% of the south growing young is somewhat easier in habi­ Florida population of white ibis breed in tats with abundant prey. Thi sis the i nteri or, freshwater wet 1ands of the especially important for the wood stork Everglades; during these times the man­ which requires that its prey be concen­ groves are apparently unimportant, sup­ trated into small pool s by fall i ng water porting less than 10% of the population levels during the dry season before it can (Kushlan 1976, 1977a, b). During drought nest successfully (Kahl 1964; Kushlan et years, however, production is sustained a1. 1975; Odgen et al. 1978). Breeding solely by breeding colonies located in acti vity by wadi ng bi rds in mangroves mangroves near the coast (Kushl an 1977a, 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).

% of total acti ve Acti ve nests in nests in south Species mangroves Florida ------White ibi s 1914 7

Roseate spoonbill 500 100

Wood stork 1335 31

Great blue heron 458 92

Great egret 1812 39

Snowy egret 2377 46

Little blue heron 71 15

Loui si ana heron 3410 70

Cattle egret 2180 13

Brown pelican 741 100

Double-crested cormorant 1744 83

64 coasts takes place throughout the year pronounced in rookeries within mangrove (Table 6); at least one species of wader regions subject to infreguent tidal f1ush­ breeds during every month. Colonies on i ng. the mangrove islands in Florida Bay were noted to be active nesting sites during 9.3 PROBING SHOREBIROS all months of the year except September and October (Kush1an and White 1977a). Bi rds in this group are commonly found associated with intertidal and shal­ The seasonal movements of wood storks low water habitats. Wo1 ff (1969) and and white ibises between the various south Schnei der (1978) have shown that plovers Florida ecosystems were described by and sandpi pers are opportuni sti c feeders, Ogden et a1. (197B) and Kushl an (1979). takin9 the most abundant, proper-sized Mangrove ecosystems appear to be most invertebrates present in whatever habitat heavily used for feeding in summer (white the bi rds happen to occupy. ibis) and early winter (white ibis and wood stork). The remai ni n9 speci es of Of the 25 species included in this wading birds appear to use mangrove areas guild (Appendix DJ, two are year-round most heavily in the winter months, reflec­ residents (clapper rail and willet), two ting the influx of migrants from farther breed in man9rove areas (clapper rail and north. black-necked stilt), and the remainder are transients or winter residents. Baker and Wading birds play an important role 8aker (1973) indicated that winter was the in nutrient cycling in the coastal man­ most crucial time for shorebirds, in terms grove zone. Mcivor (pers. observ.) has of survival. Coincidentally, winter is noted increased turbidity, greater algal the time when most shorebirds use man9rove biomass, and decreased fish abundance areas. The invertebrate fauna (moll usks, around red mangrove islets with nesting crustaceans, and aquatic insects) which fri gate bi rds and cormorants. Onuf et a1. occur on the sediments under intertidal (1977) reported resu1ts f rom a sma 11 (100 mangroves forms the pri nci pal di et of bi rd) rookery on a mangrove isl et on the these species. Willets and greater east coast of Flori da. Addi t ions of yellowlegs eat a large amount of fishes, ammonium-nitrogen fro~ the bird's especially Fundulus, in addition to inver­ droppings exceeded 1 glm Iday. Water tebrates. Many of the speci es 1 i sted in beneath the mangroves contained five times this guild obtain a significant portion of more ammonium and phosphate than water their energy requirements from other habi­ beneath mangroves without rookeries. tats, particularly sandy beaches, marshes, Although the wading birds were shown to be and freshwater prai ri es. Of the speci es a vector for concentrating nutrients, it in this guild, the clapper rail is prob­ must be noted that this is a localized ably most dependent on mangroves for phenomenon restricted to the areas around survi val in south Florida (Robertson rookeri es in the mangrove zone. The 1955), although in other geographical effect would be larger around larger locations they frequent salt and brackish rookeries. Onuf et al. (1977) also marshes. reported that mangroves in the area of the rookery had increased levels of primary production, higher stem and foliar nitro­ gen levels, and hi9her herbivore grazing 9.4 FLOATING AND DIVING WATER BIRDS impact than mangroves without rookeries. Lewis and Lewis (1978) stated that man­ Twenty-nine species of ducks, grebes, groves in large rookeries may eventually loons, cormorants, and gallinules were be killed due to stripping of leaves and identified as populating mangrove areas in branches for nesting material and by south Flori da (Appendi x D). Ei ght speci es poisoning due to large volumes of urea and are year-round residents while the ammonia that are deposited in bird guano. remainder are present only during migra­ This latter effect would be more tion or as winter visitors.

65 Table 6. Timing of nesting by wading birds and associated spec i es in south Florida. Adapted from data in Kush 1an and White (1977a). Kushlan and McEwan (in press).

Months Species SON 0 J FMAM JJ A

White ibis Wood stork Roseate spoonbill Great bl ue/white heron Great egret Little blue heron Cattle egret Double-crested cormorant Brown pelican

66 From the standpoint of feeding, mem­ estuaries support from 5% to 10% of the bers of this guild are highly hetero­ total wintering waterfowl population in geneous. Piscivorous species include the Florida (Goodwin 1979; Kush1an et a1. in cormorant, anhinga, pelicans, and mergan­ prep.). As Kush1an et a1. point out, sers. Herbivorous species include the however, the Everglades are not managed pintail, mallard, wigeon, mottled duck, for single species or ~roups of species as and tea1s. A th i rd group feeds pri ma ri 1y are areas of Florida supporting larger on benthic mollusks and invertebrates. waterfowl populations. Although the Scaup, canvasback, redhead, and gallinules importance of south Flori da's mangrove belong to this group. The ducks in this estuaries to continental waterfowl popula­ last group al so consume a significant tions may be small, the effect of 70,000 fraction of plant material. ducks and coots on these estuaries probably is not (Kush1an et a1. in prep.). Species of this guild are permanent residents and usually breed in mangrove Kushlan (personal communication) swamps. As shown in Table 5, the brown thinks that the estuaries of the Ever­ pelican and double-crested cormorant are glades have an important survival value highly dependent upon mangroves for for some segments of the Ameri can white nestin~ in south Florida even though both pel i can popu1 ati on. In wi nter, approxi­ will build nests in any available tree in mate1y 25% of the white pelicans are found other geographical regions. It seems that in Florida Bay and 75% in the Cape Sable when m~ngroves are available, they are the region. They feed pri marily in freshwater preferred nesting site. The anhinga regions of coastal marshes and prairies breeds in mangrove regions but is more and use mangroves where they adjoi n thi s commonly found inland near freshwater (J. type of habitat. A. Kush1an, So. Fla. Res. Ctr., Everglades Nat1. Park, Homestead, Fla.; ·personal communication 1981). For the other species 9.5 AERIALLY-SEARCHING BIRDS listed in this guild, mangrove swamps provi de a common but not a requi red habi­ Gulls, terns, the kingfisher, the tat; all of these species utilize a black skimmer, and the fish crow comprise variety of aquatic envi ronments. this 9ui1d of omnivorous and piscivorous species (Appendix D). These bi rds hunt in Kush1an et a1. (in prep.) provide ponds, creeks, and waterways adjacent to recent data on the abundance and distribu­ ·man9rove stands. Many fishes and inverte­ tion of 22 species of waterfowl and the brates upon which they feed come from American coot in south Florida estuaries. mangrove-based food webs. Only six of the The American coot is by far the most abun­ 14 species are year-round residents of dant species, accounting for just over 50% south Florida. The least tern is an abun­ of the total popu1 at ion. Si x spec i es of dant summer resident and the remainder are ducks were responsible for more than 99% winter residents or transients. of the individuals seen: blue-winged teal (41%), lesser scaup (24%), pintaii (18%), Only the fish crow actually nests in American wigeon (9%), ring-necked duck mangroves. Gulls and terns prefer open (5%), and shoveler (3%). The major habi­ sandy areas for nesting (Kushlan and White tats included in these authors' surveys 1977b) and use mangrove ecosystems only were coastal prairie and marshes, mangrove for feeding. All of the species in this forests, and mangrove-1 i ned bays and guild are recorded from a variety of waterways of the Everglades National Park. coastal and inland wetland habitats.

From these data it appears that waterfowl and coots are most abundant in 9.6 BIRDS OF PREY regions where mangrove, wet coastal prairies, marshes, and open water are This guild is composed of 20 species interspersed. Overall, the Ever~lades 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 gui ld 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. tail ed ha wk), waterfowl (pe regri ne fa1con, 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, rema i nder are wi nter resi dents. Thei ruse cuckoos, woodpeckers, flycatchers, of mangrove areas varies greatly. The thrushes, vireos, warblers, blackbirds, magni fi cent fri gatebi rd, whi ch occu rs 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, towhee) near the coast. Swallow-tailed kites are eat a variety of seeds, berries, and common over the enti re 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 Flori da. The nests he the mangrove canopy. Many bi rds al so use observed were all located in black man­ the trunk, branches, and aerial roots for groves although they do nest in other feedi ng. Several di fferent 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 11 s-widows, the and peregrine falcon are dependent upon kingbi rds, and the flycatchers. Gleaning mangrove ecosystems for thei r cont i nued 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 bi rds in this guild groves are used as roosts and support are heavi ly dependent upon mangrove areas. structures for nests. Nisbet (1968) indi­ The prairie warbler and the yellow warbler cated that in Ma 1aysi a the most important are subspecies of more widespread North role of mangroves for bi rds may be as American species (see Appendix 0 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. Kushl an (pers. comm.) stated that Kushlan 1974). The white-crowned pigeon, recent surveys have shown falcons to mangrove cuckoo, gray kingbird, and b1ack­ winter in mangroves, particularly along whi skered vi reo are of recent West Indian the shore of Florida Bay where they estab­ or i gin. They fir s t m0 ve dintot he 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 speci es of the peregri ne Caribbean. Confined at fi rst 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 plus the magnificent frigatebird, the expanded their range in peninsular Florida anhinga, the cormorant, and the brown by using non-mangrove habitat. ln this pelican use overwash islands for nesting vein it is interesting to note that many (Kushlan and White 1977a). Wading and species of rare and/or irregular occur­ aerially-searching birds commonly feed in rence in south Florida are of West Indian close proximity to overwash islands. A origin and use mangroves to a considerable variety of migrating arboreal and probing extent. These include the Bahama pintail, species use the islands for feeding and masked duck, Caribbean coot, loggerhead roosting. Yellow and palm warblers are kingbird, thick-billed vireo, and stripe­ common around mangrove islands in Florida headed tanager (Robertson and Kushl an Bay as are the black-bellied plover, ruddy 1974). turnstone, willet, dunlin, and short­ bi lled dowitcher. Rafts of ducks are Twenty-four of the species in this common near the inshore islands and birds guild are permanent residents, 27 are win­ of prey such as the osprey, the bald ter, and 6 are summer residents. Fourteen eagle, and both vultures use mangrove species are seen only during migrations. islands for roosting and nesting.

Fringe and riverine mangrove com­ 9.B ASSOCIATIONS BETWEEN MANGROVE munities are important feeding areas for COMMUNITY TYPES AND BIRDS wading and probing birds. Floating and diving and aerially-searching birds use Estimating the degree of use of the lakes and waterways adjacent to these mangrove swamps by bi rds as we have done mangrove communities for feeding. Many of (Appendi x D) is open to criti cis m becau se the wading birds nest in fringe and of the pauc i ty of in format i on upon whi ch ri veri ne forests. For examp1 e, when the to base judgements. Estimating which wood ibis nests in coastal areas, it uses mangrove community types (see section 1, these mangrove communities almost exclu­ Figure 4) are used by which birds is open sivel.y (Kushlan, personal communication). to even more severe criticism. For this Most of the arboreal bi rds and bi rds of reason the foll owi ng comments shou1 d be prey associated with mangroves are found regarded as general and pre1 i mi nary. in these two types of communi ties. Thi s is not surprising since the tree canopy is In terms of utilization by avifauna, extremely well-developed and offers the scrub mangrove swamps are probably the roosting, feeding and nesting opportuni­ least utilized mangrove community type. ties. Because the canopy is poorly developed, most of the arboreal species are absent, Hammock and basi n mangrove communi­ although Emlen (1977) recorded the red­ ties are so diverse in size, location, and winged blackbird, hairy woodpecker, north­ proximity to other communities that it is ern waterthrush, yellow-rumped warbler, di ffi cult to make many general statements common yell owth roat, orange-crowned about thei r avi fauna. Si nce there often warbler, palm warbler, yellow warbler, is little standing water in hammock mourning dove, and gray kingbird in scrub forests, wading and diving birds probably mangroves on Grand Bahama Island. Of 25 are not common. Proxi mity to terrestri al di fferent habitats surveyed by Em1en communities in some cases may increase the (1977), the yellow warbler and gray di versity of arboreal speci es in both kingbird were found in the scrub mangroves hammock and basin forests; proximity to only. Aerially-searching and wading birds open areas may increase the likelihood of might use scrub mangroves if fishes are bi rds of prey. present. It seems safe to conclude that each Overwash mangrove i sl ands are of the six mangrove community types has utilized in a variety of ways by all of some value to the avifauna. This value the bi rd guil ds. Most of the wadi ng bi rds di ffers accordi ng to community type and

70 kind of bird group under consideration. species basis only 9 were recorded from Certainly, more information is needed, mangroves whereas 19, 13, and 16 species, particularly concerning the dependence of respectively, were seen in the other habi­ rare or endangered species on specific tats. This large number of species from communi ty types. 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 g.g MANGROVES AS WINTER HABITAT FOR NORTH groves whereas between 30 and 86 hours AMERICAN MIGRANT LAND BIRDS were spent in other habitats. More ti me in the mangrove zone would have undoubted­ An interestin9 observation based on ly resulted in more species (and in­ the data in this chapter is the seemingly dividuals) observed (Preston 1979). important rol e that mangrove ecosystems play in providing wintering habitat for Hutto (1980) presented extensive data mi grants of North Ameri can ori gi n. Lack concerning the composition of migratory and Lack (1972) studi ed the wi nteri ng land bird communities in Mexico in winter warbler community in Jamaica. In four for 13 habitat types. Mangrove areas natural habitats including mangrove tended to have more migrant species than forest, lowland dry limestone forest, mid­ most natura 1 habi tats (except gall ery I evel wet 1i mestone forest, and montane forests) and also had a greater density of cloud forest,a total of 174,131,61, and individuals than other habitats (again 49 warblers (individuals) were seen, except for gallery forests). In both Lack respectively. When computed on a per hour and Lack's and Hutto's studies, disturbed of observation basis, the difference is and edge habitats had the hi ghest number more striking with 22 warblers per hour of species and greatest density of seen in mangroves and only I, 2, and 1 i ndi vi dual s. The percentage of the seen in the other forest habitats, respec­ avi fauna communi ty composed of mi grants tively. For all passerines considered was hi ghest in mangrove habitats, however. together, 26 passerines/hour were seen in From this we can infer the importance of mangroves with 5, 13, and 3 respectively mangroves in the ma i ntenance of North in the other forest habitats. On a 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 speci es in the Eastern Uni ted 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. Oata 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 cluded. This has produced a conservative other coastal habitats in Florida. Layne esti mate of the mamma 1 species that uti­ (1974, see his figure 1) gives a disjunct 1ize mangrove areas. distribution for this species in Florida, with the major geographical range being Several mammal s do not appear in the southwest coast. River otters also Appendix E because they have not been util i ze 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, Sig­ of all sightings of this species in Ever­ modon hispidus, the round-tailed muskrat, glades National Park were from mangroves Neofiber alleni, the house mouse, Mus (Layne 1974). Bobcat are found in almost musculus,-fh-e-least shrew, CryptotlS all habitats in south Florida from pine­ parva, and the short-tailed shrew, Blarina lands to dense mangrove forests. The bfeVTcauda. 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 Flori da regi on is are apparently most abundant in the Big defi ci ent 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 ri ce rats are commonly found in mangrove ut i 1i ze south Flori da 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 pri mari ly upon sea grasses and other utilize many other habitats. The key submerged aquatic plants, it is commonly deer, a rare and endangered subspecies, is found in canals, coastal rivers, and restricted to the Big Pine Key group in embayments close to mangrove swamps. the Florida Keys, although it ranged onto the mainland in historical times. Al­ Except for the Cudjoe Key rice rat, though this little deer makes use of pine none of the mammals found in Fl or; da man­ uplands and oak hammocks, it extensively groves are sol ely dependent upon mangrove exploits mangrove swamps for food and ecosystems; all of these speci es can cover. utilize other habitats. The destruction of extensive mangrove swamps would, how­ Two marine mammals, the bottlenose ever, have deleterious effects on almost porpoi se and the manatee, frequent all of these species. Populations of mangrove-l i ned waterways. The bott1enose panther, key deer, and the ri ver otter porpoi se feeds on mangrove-associ ated would probably be the most seriously fishes such as the striped mullet, Mugil affected, because they use mangrove habi­ cephal us. Although the manatee -reeas tat extensi vely.

73 CHAPTER 11. VALUE OF MANGROVE ECOSYSTEMS TO MAN

Mangrove swamps are often hot, fetid, tioned by Savage (1972), Carlton (1974), mosqui to-ri dden, and a1most i mpenet rab1e. and Teas (1 977). These authors argue that As a consequence, they are frequently held the black mangrove (1) is easier to in low regard. It is possible that more transplant as a seedling, (2) establishes acres of mangrove, worldwide, have been its pneumatophore system more rapidly than obI iterated by man in the name of "recla­ the red mangrove develops prop roots, (3) mation" than any other type of coastal has an underground root system that is environment. Reclamation, according to better adapted to holding sediments (Teas Webster's, means "to claim back, as of 1977), (4) is more cold-hardy, and (5) can waste1and". Mangrove swamps are anythi ng better tolerate "artificial" substrates but wasteland, however, and it is impor­ such as dredge-spoi 1, fi nger fi 11 s, and tant to establish this fact before a causeways. Generally, the white mangrove valuable resource is lost. We can think is regarded as the poorest land stabilizer of six major categories of mangrove values of the Florida mangroves (Hanlon et al. to man; no doubt, there are more. 1975) •

Although mangroves are susceptible to 11.1 SHORELINE STABILIZATION AND STORM hurricane damage (see section 12.1), they PROTECTION provide considerable protection to areas on thei r 1andward si de. They cannot The ability of all three Florida prevent all flooding damage, but they do mangroves to trap, hold and, to some mitigate the effects of waves and extent, stabilize intertidal sediments has breakers. The degree of this protection been demonstrated repeatedly (revi ewed by is roughly proportional to the width of Scoffi n 1970; Ca r 1ton 197 4). The contem­ the mangrove zone. Very narrow fringing pora ry vi ew of mangroves is that they forests offer minimal protection while function not as "land builders" as hypo­ extensive stands of mangroves not only thesized by Davis (1940) and others, but prevent wave damage, but reduce much of as "stabilizers" of sediments that have the flooding damage by damping and holding been deposited largely by geomorphological flood waters. Fosberg (1971) suggested processes (see section 3.2). that the November 1970 typhoon and accom­ panying storm surge that claimed between Gill (1970), Savaqe (1972), Teas 300,000 and 500,000 human lives in (1977), and others have emphasized that Bangladesh might not have been so destruc­ land stabilization by mangroves is pos­ tive if thousands of hectares of mangrove sible only where conditions are relatively swamps had not been replaced with rice quiescent and strong wave action and/or paddies. 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 11.2 HABITAT VALUE TO WILDLIFE stabilize the sediments. Certainly, this depends to some extent on substrate type; Florida mangrove ecosystems are mangroves appear to withstand wave energy important habitat for a wide variety of best on solid rock substrates with many reptiles, amphibians, birds, and mammals cracks and crevices for root penetration. (see sections 8, 9, and 10). Some of From our own experience, we suspect that these animals are of commercial and sport mangroves on sandy and muddy substrates importance (e.g., white-tailed deer, sea cannot tolerate any but the lowest wave turtles, pink shrimp, spiny lobster, energies, tidal currents much above 25 snook, grey snapper). Many of these are cm/s, or heavy, regul ar boat wakes. important to the south Florida tourist industry including the wading birds (e.g., The concept that the red mangrove is egrets, wood stork, white ibis, herons) the best land stabilizer has been ques- which nest in the mangrove zone.

74 11.3 IMPORTANCE TO THREATENED AND ENDAN­ of these connections were discussed by GERED SPECIES Heald (1969), Odum (1970), Heald and Odum (1970), and Odum and Heald (1975b) in The mangrove forests of south Florida terms of support for commercial and sport are important habitat for at least seven fisheries. endangered species, five endangered sub­ species, and three threatened species A minimal list of mangrove-associated (Federal Regi ster 1980). The endangered organi sms of commerci al or sport val ue species include the American crocodile, includes oysters, blue crabs, spiny the hawksbill sea turtle, the Atlantic lobsters, pink shrimp, snook, mullet, ridley sea turtle, the Florida manatee, menhaden, red drum, spotted sea trout, the bald eagle, the American peregrine gray and other snapper, tarpon, falcon, and the brown pelican. The endan­ sheepshead, ladyfish, jacks, gafftopsail gered subspecies are the key deer catfi sh, and the jewfi sh. Heald and Odum (Odocoileus vir inianus clavium), the (1970) pointed out that the commercial Florida panther Felis concolor coryi), fisheries catch, excluding shrimp, in the the Barbados yellow warbler (Dendroica area from Naples to Florida Bay was 2.7 etechia petechia), the Atlantic saltmarsh million pounds in 1965. Almost all of the snake Nerodia fasciata taeniata) and the fish and shellfish which make up this eastern indigo snake (Drymarchon corais catch utilize the mangrove habitat at some couperi). Threatened species include the point during their life cycles. In addi­ American alligator, the green sea turtle tion, the Tortugas pink shrimp fishery, and tM loggerhead sea turtle. Although which produces in excess of 11 million all of these animals utilize mangrove pounds of shrimp a year (Idyll 1965a), is habitat at times in their life histories, closely associated with the Everglades species that would be most adversely estuary and its mangrove-lined bays and affected by widespread mangrove destruc­ ri verso tion are the American crocodi le, the Florida panther, the American peregrine fal con, the brown pel i can, and the 11.5 AESTHETICS, TOURISM AND THE Atlantic ridley sea turtle. The so-called INTANGIBLES mangrove fox sgui rrel (Sciurus niger avicennia) is widely believed to be a One va 1ue of the mangrove ecosystem, mangrove-dependent endangered species. which is difficult to document in dollars This is not the case since it is currently ·or pounds of meat, is the aesthetic value regarded as 'Irare'~ not endan~eredt and, to man. Admittedly, not all individuals further, there is some question whether find visits to mangrove swamps a pleasant or not this is a legitimate sub-species experience. There are many others, how­ (Hall 1981). As a final note, we should ever, who place a great deal of value on point out that the red wolf (Canis rufus), the extensive vistas of mangrove canopies, which is believed to be extinct in waterways, and associated wildlife and Florida, at one time used mangrove habitat fishes of south Florida. In a sense, this in addition to other areas in south mangrove belt along with the remaining Florida. sections of the freshwater Everglades and Hi g Cypress Swamp are the only remai ni ng wilderness areas in this part of the 11.4 VALUE TO SPORT AND COMMERCIAL Uni ted States. FISHERIES Hundreds of thousands of visitors The fish and invertebrate fauna of each year visit the Everglades National mangrove waterways are closely 1inked to Park; part of the reason for many of these mangrove trees through (a) the habitat visits includes hopes of catching snook or va 1ue of the aeri a1 root structu re and (b) gray snappers in the mangrove-lined water­ the mangrove leaf detritus-based food web ways, seeing exotic wading birds, croco­ (see sections 6 and 7). The impl ications diles, or panthers, or simply discovering

75 what a tropical mangrove forest looks makes a durable and water resistant timber like. The National Park Service,in an which has been used successfully for resi­ attempt to accommodate this last wish, dential buildings, boats, pilings, maintains extensive boardwalks and canoe hogsheads, fence posts, and furniture trai 1s through the mangrove forests near (Kuenzler 1974; Hanlon et al. 1975). In Flamingo, Florida. In other, more Southeast Asia mangrove wood is widely developed parts of the State, small stands used for high quality charcoal. of mangroves or manqrove islands provide a feeling of wilderness in proximity to the Morton (1965) mentions that red man­ rapidly burqeoning urban areas. A variety grove fru i ts are somt i mes eaten by humans of tourist attractions including Fairchild in Central America, but only by popula­ Tropical Gardens near Miami and Tiki tions under duress and subject to starva­ Gardens near St. Petersburg utilizes the tion. Mangrove leaves have variously been exotic appearance of mangroves as a key used for teas, medicinal purposes, and ingredient in an attractive landscape. livestock feeds. Manqrove teas must be Clearly, mangroves contribute intangibly drunk in small quantities and mixed with by diversifying the appearance of south milk because of the high tannin content Florida. (Morton 1962); the mil k bi nds the tanni ns and makes the beverage more pal atab1e.

11.6 ECONOMIC PRODUCTS As a final note, we should point out that mangrove trees are responsible for Elsewhere in the world, mangrove contributing directly to one commercial forests serve as a renewable resource for product in Florida. The flowers of black many valuable products. For a full dis­ mangroves are of considerable importance cussion of the potential uses of mangrove to the three million dollar (1965 figures) products, see de la Cruz (i n press a), Flori da honey i ndust ry (Morton 1964). Morton (1965) for red mangrove products, and Moldenke (1967) for black mangrove Other than the honey industry, most products. of these economic uses are somewhat destructive. There are many cases in In many countries the bark of man­ which clear-cut mangrove forests have groves is used as a source of tanni ns and failed to regenerate successfully for many dyes. Si nce the ba rk is 20'; to 30'; tanni n years because of lack of propagule on a dry wei ght basi s, it is an excell ent dispersal or increased soil salinities source (Hanlon et al. 1975). Silviculture (Teas 1979). We believe that the best use (forestry) of mangrove forests has been of Florida mangrove swamps will continue practiced extensively in Africa, Puerto to be as preserved areas to support Rico, and many parts of Southeast Asia wildlife, fishing, shoreline stabiliza­ (Holdridge 1940; Noakes 1955; Macnae 1968; tion, endangered species, and aesthetic Wal sh 1974; Teas 1977). Mangrove wood values.

76 CHAPTER 12. MANAGEMENT IMPLICATIONS

12.1 INHERENT VULNERABILITY by oXYgen depletion due to accumulations of decomposin9 organic matter (Tabb and Manqroves have evolved remarkable Jones 1962). physiological and anatomical adaptations enabling them to flourish under conditions Hurricane Betsy in 1965 did little of high temperatures, widely fluctuating damage to mangroves in south Florida; salinities, high concentrations of heavy there was also little deposition of silt metal s (Wal sh et a1. 1979), and anaerobic and marl within man9rove stands from this soi 1s. Unfortunately, one of these adap­ minimal storm (Alexander 1967). Lugo et tations, the aerial root system, is also a1. (1976) have hypothesized that severe one of the plant's most vulnerable compo­ hurricanes occur in south Florida and nents. Odum and Johannes (1 975) have Puerto Rico on a time interval of 25 to 30 referred to the aerial roots as the man­ years and that mangrove ecosystems are grove's Achi 11 es' heel because of thei r adapted to reach maximum biomass and pro­ susceptibility to clogging, prolonged ductivity on the same time cycle. fl oodi ng, and bori ng damage from i sopods and other invertebrates (see section 6 for a discussion of the latter). This means 12.2 MAN-INOUCED DESTRUCTION that any process, natu ra1 or man -i nduced, which coats the aerial roots with fine Destruction of mangrove forests in sediments or covers them with water for Florida has occurred in various ways extended periods has the potential for including outright destruction and land mangrove destruction. Bacon (1970) men­ filling, dikin9 and flooding (Fi9ure 17), tions a case in Trinidad where the Caroni through introduction of fine particulate River inundated the adjacent Caroni material, and pollution damage, par­ Mangrove Swamp during a flood and ticularly oil spills. To our knowledge deposited a layer of fine red marl in a there are no complete, published docu­ large stand of black mangroves which sub­ mented estimates of the amount of mangrove sequently died. Many examp1 es of damage forests in Florida which have been to mangrove swamps from human activities destroyed by man in this century. Our have been documented (see secti on 12.2). conclusion is that total loss statewide is not too great, probably in the range of 3 One of the few natural processes that to 5% of the ori9ina1 area covered by causes periodic and extensive damage to mangroves in the 19th century, but that mangrove ecosystems is 1a rge hu rri canes losses in specific areas, particularly (Fi gure 16). Crai ghead and Gi 1bert (1962) urban areas, are appreciable. This con­ and Tabb and Jones (1962) have documented clusion is based on four pieces of infor­ the impact of Hurricane Oonna in 1960 on mat ion. (1) Linda11 and Sal oman (1977) parts of the man9rove zone of south have estimated that the total loss of Florida. Crai9head and Gilbert (1962) vegetated intertidal marshes and mangrove found extensive damage over an area of swamps in Florida due to dredge and fill 100,000 acres (40,000 hal. Loss of trees is 23,521 acres (9,522 hal; remember that ranqed from 25% to 100%. Oamage occurred there are between 430,000 and 500,000 in three ways: (1) wind shearing of the acres (174,000 to 202,000 hal of mangroves trunk 6 to 10 ft (2 to 3 m) above 9round, in Florida (see section 1.3). (2) (2) overwash man9rove islands being swept Birnhak and Crowder (1974) estimate a loss clean, and (3) trees dying months after of approximately 11 ,000 acres (4,453 hal the storm, apparently in response to of mangroves between 1943 and 1970 in damage to the prop roots from coati ngs by three counties (Co11 i er, Monroe, and marl and fi ne organic matter. The 1atter Dade). (3) An obvious loss of mangrove type of damage was most widespread, but forests has occurred in Tampa Bay, around rarely occurred in intertidal forests, Marco Island, in the Florida Keys, and presumably because the aerial roots were along the lower east coast of Florida. flushed and cleaned by tidal action. Fish For example, Lewis et a1. (1979) estimated and invertebrates were adversely affected 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 , ....

• _ .f",t.

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 Although mangroves commonly occur in estuary has been destroyed during the past a reas of rapi d sedi mentat i on, they cannot 100 yea rs. (4) Hea 1d (unpub1i shed MS.) survive heavy loads of fine, floculent has estimated a loss of 2,000 acres (810 materials which coat the prop roots. The hal of mangroves within the Florida Keys instances of mangrove death from these (not considered by Bi rnhak and Crowder substances have been bri efly revi ewed by 1974). So while loss of mangrove ecosys­ Odum and Johannes (1975). Mangrove deaths tems throughout Flori da is not over­ from fi ne muds and marl, ground bauxite whelming, losses at specific locations and other ore wastes, sugar cane wastes, have been substantial. pulp mill effluent, sodium hydroxide wastes from bauxi te processing, and from Diking, impounding, and long-term intrusion of large quantities of beach flooding of mangroves with standing water sand have been documented from various can cause mass mortality, especially when areas of the worl d. prop roots and pneumatophores are covered (Breen and Hi 11 1969; Odum and Johannes 12.3 EFFECTS OF OIL SPILLS ON MANGROVES 1975; Patterson-Zucca 1978; Lugo 1981). In south Florida, E. Heald (pers. comm.) There is little doubt that petroleum has observed that permanent impoundment by and petroleum byproducts can be extremely diking which prevents any tidal exchange harmful to mangroves. Oamage from oil and raises water levels significantly spill s has been reviewed by Odum and during the wet season will kill all adult Johannes (1975), Carlberg (1980), Ray (in red and black mangrove trees. If condi­ press), and de la Cruz (in press, b). tions behind the dike remain relatively Over 100 references detailing the effects dry, the mangroves may survive for many of oi 1 spi 11 s on mangroves and mangrove­ years until replaced by terrestrial vege­ associated biota are included in these tation. revi ews.

Petroleum and its byproducts injure Mangroves are unusually susceptible and ki 11 mangroves in a variety of ways. to herbicides (Walsh et al. 1973). At Crude oil coats roots, rhizomes, and pneu­ least 250,000 acres (100,000 hal of man­ matophores and disrupts oxygen transport grove forests were defoliated and killed to underground roots (Baker 1971). in South Viet Nam by the U.S. military. Various reports suggest that the critical This widespread destruction has been docu­ concentration for crude oil spills which mented by Tschirley (1969), Orians and may cause extznsive damage is between 100 Pfeiffer (1970), Westing (1971), and a and 200 ml /m of swamp surface (Odum and committee of the U.S. Academy of Sciences Johannes 1975). Petroleum is readily (Odum et al. 1974). In many cases these absorbed by 1i pophyl i c substances on su r­ forests were slow to regenerate; observa­ faces of mangroves. This leads to severe tions by de Sylva and Michel (1974) indi­ metabolic alterations such as displacement cated higher rates of siltation, greater of fatty molecul es by oil hydrocarbons water turbidity, and possibly lower dis­ leading to destruction of cellular permea­ solved oxygen concentrations in swamps bility and/or dissolution of hydrocarbons which sustained the most damage. Teas and in lipid components of chloroplasts (Baker Kelly (1975) reported that in Florida the 1971). black mangrove is somewhat resistant to most herbicides but the red mangrove is As with other intertidal communities, extremely sensitive to herbicide damage. many of the invertebrates, fishes, and He hypothesi zed that the vul ne rabil i ty of plants associated with the mangrove com­ the red mangrove is related to the small munity are highly susceptible to petroleum reserves of viable leaf buds in this tree. products. Widespread destruction of Following his reasoning, the stress of a organisms such as attached algae, oysters, single defoliation is sufficient to kill tunicates, crabs, and gobies have been the enti re tree. reported in the 1iterature (reviewed by de

80 la Cruz in press, b; Ray in press). 12.4 MAN-INDUCED MODIFICATIONS

Damage from oil spills follows a In south Florida, man has been re­ predictable pattern (Table 7) which may sponsible for modifications which, while require years to complete. It is impor­ not ki 11 i ng mangroves outri ght, have al­ tant to recognize that many of the most tered components of the mangrove ecosys­ severe responses, including tree death, tem. One of the most widespread changes may not appear for months or even years involves the alteration of freshwater after the spill. runoff. Much of the freshwater runoff of the Flori da Evergl ades has been di verted In Florida, Chan (1977) reported that elsewhere with the result that salinities red mangrove seedl i ngs and black mangrove in the Everglades estuary are generally pneumatophores were particularly sensitive higher than at the turn of the century. to an oi 1 spi 11 whi ch occurred in the Teas (1977) points out that drainage in Florida Keys. Lewis (1979a, 1980b) has the Miami area has lowered the water table followed the long-term effects of a spi 11 as much as 2 m (6 ft). of 150,000 liters (39,000 gal) of bunker C and diesel oil in Tampa Bay. He observed Interference with freshwater inflow short-term (72-hour) mortality of inverte­ has extensi ve effects on estuaries (Odum brates such as the gastropod Melongena 1970). Florida estuaries are no excep­ corona and the polychaete Laeonereis tion; the effects on fish and invertebrate culveri. Mortal ity of all three species species along the edge of Biscayne and of mangroves began after three weeks and Florida Bays have been striking. The continued for more than a year. Sub­ mismanagement of freshwater and its lethal damage included partial defoliation effects on aquatic organisms have been of all species and necrosis of black discussed by Tabb (1963); Idyll (1965a,b); mangrove pneumatophores; death depended Tabb and Yokel (1968) and Idyll et al. upon the percentage of pneumatophores (1968). In addition, Estevez and Simon affected. (1975) have hypothesized that the impact of the boring isopod, Sphaeroma terebrans, In addition to the damage from oil may be more severe when freshwater f1 ows spi 11 s, there are many adverse impacts on from the Everglades are altered. mangrove forests from the process of oi 1 exploration and drilling (Table 8). This One generally unrecognized side type of damage can often be reduced effect of lowered freshwater flow and salt through careful management and monitori ng water intrusion has been the inland expan­ of dri 11 i ng sites. sion of mangrove forests in many areas of south Flori da. There is documented evi­ Although little is known concerning dence that the mangrove borders of ways to prevent damage to mangroves once a Biscayne Bay and much of the Everglades spill has occurred, protection of aerial estuary have expanded inland during the roots seems essential. Prop roots and past 30 to 40 years (Reark 1975; Teas pneumatophores must be cl eaned with com­ 1979; Ball 1980). pounds whi ch wi 11 not damage the pl ant tissues. Dispersants commonly used to Sections of many mangrove forests in combat oil spills are, in general, toxic south Florida have been replaced by filled to vascular plants (8aker 1971). If pos­ residential lots and navigation canals. sible, oil laden spray should not be Although these canal systems have not been allowed to reach leaf surfaces. Damage studied extensively, there is some evi­ during clean-up (e.g., trampling, compac­ dence, mostly unpublished, that canals are tion, bulldozing) may be more destructive not as productive in terms of fishes and than the unt reated effects of the oi 1 invertebrates as the natural mangrove­ spill (de la Cruz in press, b). lined waterways which they replaced.

81 Table 7. General response of mangrove ecosystems to severe oil spills (from Lewis 1980b)

Stage Observed impact

Acute o to 15 days Deaths of birds, turtles, fishes, and invertebrates 15 to 30 days Defoliation and death of small mangroves, loss of aerial root community Chronic 30 days to 1 year Defoliation and death of medium-sized mangroves (1 - 3 m), tissue damage to aeri a 1 roots 1 year to 5 years Death of large mangroves (greater than 3 m), loss of oiled aerial roots, and regrowth of new roots (often deformed) Recolonization of oil-damaged areas by new seedl i ngs

1 year to 10 years (?) Reduction in litter fall, reduced re­ production, and reduced survival of seedl ings Death or reduced growth of young trees colonizing spill site (?)

Increased insect damage (?)

10 to 50 years (?) Complete recovery

82 Table 8. Estimated impact of various stages of oil mlnlng on mangrove ecosystems (modified from Longley et al. 1978 and de la Cruz in press,b).

Stage Activity Impacts

Pre-exploration Seismic surveys Crushing and clearing vegetation Clearing of survey lines Vehicle track compaction Drilling "shot lines' Damage to natural levees Site preparation Canal excavation Loss of habitat in disturbed areas Dredge spoil deposition Alteration of water flow pathways Road construction Increased turbidity, higher rates of sed- imentation, and lowered dissolved oxy­ gen in nearby waters Drilling Increased activity at site Continued high turbidity related to drilling Release of toxic substances Displacement of wildlife co w Production Construction of platforms Continued high turbidity Construction of pipelines Loss of additional habitat Maintenance dredging Further changes in wetland drainage pat­ Placement of tanks and terns from pipeline construction other equipment Release of toxic substances Oil spills Oil spills Oil leaks and spills due Destruction of plant and animal popula­ to well blow-out, pipe­ tions line breakage, careless­ Alteration of ecosystem processes such ness, and barge rupture as primary production and decomposition Clean-up activities Introduction of persistent toxic substan­ ces into soil s Weinstein et al. (1977) found that artifi­ of the most product i ve aquaculture ponds cial canals had lower species diversity of in Indonesia and the Philippines are benthic infauna and trawl-captured fishes located in former mangrove swamps, there and generally finer sediments than the is some question whether the original natural communities. Courtney (1975) natura1 system was not equa 11 y producti ve reported a number of mangrove-associated in terms of fi sheri es products at no cost invertebrates which did not occur in the to man (Odum 1974). Conversion to artificial channels. aquaculture and agriculture is cursed with a variety of problems including subsequent Mosquito production is a serious 1and subsi dence and the "cat cl ay" problem in black mangrove-dominated swamps problem. The latter refers to the in Florida (Provost 1969). The salt marsh drastically lowered soil pH which often mosquitos, Aedes taeniorhynchus and ~. occurs after drai nage and has been traced sollicitans, do not reproduce below the to oxidation of reduced sulfur compounds mean high tide mark and for this reason (Dent 1947; Tomlinson 1957; Hesse 1961; are not a serious problem in the inter­ Hart 1962, 1963; Moorman and Pons 1975). tidal red mangrove swamps. Mosquitos lay Experience in Africa, Puerto Rico. and their eggs on the damp soil of the irregu­ Southeast Asia confi rms that mangrove larly flooded black mangrove zone; these forests in their natural state are more eggs hatch and develop when flooded by valuable than the "reclaimed" land. spring tides, storm tides or heavy rains. As with the "hi gh marsh" of temperate 12.5 PROTECTIVE MEASURES INCLUDING latitudes, there have been some attempts TRANSPLANT! NG to ditch the black mangrove zone so that it drains rapidly after flooding. Protect i on of mangroves inc1udes (l) Although properly designed ditchinq does prevention of outright destruction from not appear to be particularly harmful to dredging and filling; (2) prevention of mangrove swamps (other than the area drainage, diking and flooding (except for destroyed to dig the ditch and receive the carefully managed mosquito control); (3) spoil), it is an expensive practice and prevention of any alteration of hydrologi­ for this reason is not widely practiced. cal ci rculation patterns, particularly Properly managed diking can be an effec­ involving tidal exchange; (4) prevention tive mosquito control approach with mini­ of introduction of fine-grained materials mal side effects to black mangroves which might clog the aerial roots. such as (Provost 1969). Generally, ditching or cl ay. and sugar cane wastes; (5) preven­ diking of the intertidal red mangrove zone tion of oil spills and herbicide spray is a waste of money. driftage; and (6) prevention of increased wave action or current velocities from Mangrove swamps have been proposed as boat wakes, and sea wall s. possible tertiary treatment areas for sewage (see discussion by Odum and Where mangroves have been destroyed, Johannes 1975). To our knowledge, this they can be replanted or suitable alter­ alternate use is not currently practiced nate areas can be planted, acre for acre, in south Florida. Until more experimental through mitigation procedures (see Lewis results are available on the assimilative et al. 1979). An extensive body of capacities and long-term changes to be literature exists concerning mangrove expected in mangrove forests receiving planting techniques in Florida (Savage heavy loads of secondary treated sewage, 1972; Carlton 1974; Pulver 1976; Teas it would be an environmental risk to use 1977; Goforth and Thomas 1979; Lewis mangrove forests for this purpose. 1979b). Mangroves were initially planted in Florida at least as early as 1917 to In many areas of the world mangrove protect the overseas railway in the swamps have been converted to other uses Flori da Keys (Teas 1977). such as aquaculture and agriculture (see de la Cruz, in press, a). Although some Both red and b1 ack mangroves have

84 been used in transplanting. As we men­ the generally perceived lack of organisms, tioned in section 11, black mangroves seem particularly gamefishes, which use black to have certain advantages over red man­ mangrove forests as habitat. groves. Properly designed plantings are usually 75% to 90% successful, although The counter argument states that the larger the transplanted tree, the black mangrove forests are important for lower its survival rate (Teas 1977). the support of wil dl ife and the export of Pruning probably enhances survival of substantial quantities of dissolved trees other than seedl i ngs (Carl ton 1974). organic matter (DOM). Lugo et al. (1980) Important consi derati ons (Lewi s 1979b; provide evidence that black man9rove Teas 1977) in transplanting mangroves are: forests do, in fact, export large quanti­ (1) to plant in the intertidal zone and ties of DOM. They poi nt out that (1) avoid plantin9 at too high or too low an black man9rove leaves decompose more elevation, (2) to avoid planting where the rapidly than red mangrove leaves and thus shoreline ener9Y is too great, (3) to produce rel ati vely more DOM and (2) abso­ avoid human vandalism, and (4) to avoid 1ute export of carbon from these forests, accumulations of dead sea grass and other on a statewide scale, is equal or greater wrack • than from red man9rove forests.

Costs of transplanting have been variously estimated. Teas (1977) suggests 12.7 THE IMPORTANCE OF INTER-COMMUNITY $462 an acre ($1,140/ha) for un rooted EXCHANGE propagules planted 3 ft (0.9 m) apart, $1,017 an acre ($2,500/ha) for established From previous discussions (sections 6 seedlings planted 3 ft (0.9 m) apart and and 7.5 and Appendi ces B, C, 0 and E) it $87,500 ($216,130/ha) for 3 year-old nur­ is clear that many species of fishes, sery trees planted 4 ft (1.2 m) apart. invertebrates, bi rds, and mammal s move Lewis (1979b) criticized Teas' costs as between mangrove forest communities and unrealistically low and reported a project other habitats including sea grass beds, in Puerto Rico which used established coral reefs, terrestrial forests, and the seedl ings at a cost of $5,060 an acre freshwater Everglades. For example, the ($12,500/ha); he did suggest that this gray snapper, Lutjanus griseus, spends cost could be cut in half for larger part of its juveni le 1ife in sea grass projects. beds, moves to mangrove-lined bays and rivers, and then migrates to deeper water and coral reefs as an adult (Croaker 1962; 12.6 ECOLOGICAL VALUE OF BLACK VS. RED Starck and Schroeder 1971). The pi nk MANGROVES shrimp, Penaeus duorarum, spends its juve­ nile life in mangrove-lined bays and One unanswered question of current ri vers before movi ng offshore to the interest in Florida concerns the ecologi­ Tortugas grounds as an adult. During its cal value of black mangrove forests com­ juvenile period it appears to move back pared to intertidal red mangrove forests. and forth from mangrove-dominated areas to In many respects, this is identical to the sea grass beds. The spi ny lobster, lIhi gh rna rs h" versus III ow rna rsh" debate in Panul i rus argus, as a juveni le frequently temperate wetl ands. One hypothet i cal uses mangrove prop root communities as a argument which has been presented fre­ refuge; when nearing maturity this species quently in court cases during the past moves to deeper water in sea grass and decade suggests that black mangrove coral reef communities (see discussion forests have less ecological value than section 6.1). Many of the mammals (sec­ red mangrove forests to both man and tion 10) and birds (section g) move back coastal ecosystems. This argument is and forth between mangrove communi ties and based on an apparent lack of substantial a variety of other environments. particulate detritus export from black mangrove forests above mean high tide and These are only a few of many

85 examples. Clearly, mangrove ecosystems reviewed for this publication, we have are linked functionally to other south concluded that the best management prac­ Florida ecosystems through physical pro­ tice for all types of Florida mangrove cesses such as water flow and organic ecosystems is preservation. Central to carbon flux. As a result, the successful this concept is the preservation of management and/or preservation of many adjacent ecosystems that are linked signi­ fishes, mammals, birds, reptiles, and ficantly by functional processes. The amphibians depends on proper understanding continued successful functioning of the and management of a variety of ecosystems mangrove belt of southwest Florida is and the processes that link them. Saving highly dependent on the continual exis­ mangrove stands may do the gray snapper tence of the Everglades and Big Cypress 1itt 1e good if sea grass beds are Swamp in an ecologically healthy condi­ destroyed. Pink shri mp populations will tion. be enhanced by the preservation of sea grass beds and mangrove-lined waters, but At no cost to man, mangrove forests shrimp catches on the Tortugas grounds provide habitat for valuable birds, mam­ will decline if freshwater flow from the mals, amphibians, reptiles, fishes, and Everglades is not managed carefully (Idyll invertebrates and protect endangered et al. 1968). Successful management of species, at least partially support exten­ south Flori da mangrove ecosystems, sive coastal food webs, provide shoreline including their valuable resources, will stability and storm protection, and depend on knowledgeable management of a generate aesthetically pleasing experi­ number of other ecosystems and the ences (Figure 18). In situations where processes which link them. overwhelming economic pressures dictate mangrove destruction, every effort should be made to ameliorate any losses either through mitigation or through modified 12.8 MANAGEMENT PRACTICES: PRESERVATION development as described by Voss (1969) and Tabb and Heald (1973) in which canals 8ased on years of research in south and seawalls are placed as far to the rear Florida and based on the information of the swamp as possible.

86 ...•. If I 1 j

Fi gu re 18. Mangrove is1ands in Flori da Bay nea r Upper 11a tecumbe Key. Note the extensive stands of seedling red mangroves which have become established (19Bl) after a long period without major hurricanes. Mangrove islands in the Florida Keys tend to expand during storm-free intervals.

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Sutherland, J.P. 1980. Dynamics of the Taylor, D.L. 1981. Fire history and fire epibenthic community on roots of the records for Everglades National Park mangrove, Rhizophora mangle, at Bahia IS48-1979. Report T-619, Everglades de Buche, Venezuela. Mar. Biol. 58: National Park, South Florida Research 75-84. Center. 121 pp.

Tabb, D.C. 1963. A summary of existin9 Taylor, W.R. 1960. Marine algae of the information on the freshwater and eastern tropical and subtropical marine ecol09Y of the Florida Ever­ coasts of the Americas. Univ. of glades region in relation to fresh­ ~lichigan Press, Ann Arbor. 879 pp. water needs of Everglades National Park. Report to the Superintendent Teas, H. ISn. Ecology and restoration of Everglades National Park from the of mangrove shorelines in Florida. University of Miami Marine Lab. Environ. Conserv. 4: 51-57. 153 pp. Teas, H. 1979. Silviculture with saline Tabb, D.C. 1966. The estuary as a habi- water. Pages 117-161 in A. Hollaen­ tat for spotted seatrout, Cynoscion der, ed. The bi osa1i ne concept. nebulosus. Am. Fish. Soc. Spec. Plenum Publ. Corp. Publ. 3: 59-67. Teas, H., and J. Kelly. 1975. Effects of Tabb, D.C., and E.J. Heald. 1973. The herbicides on mangroves of S. Vietnam coastal interceptor waterway. Univ. and Florida. Pages 719-728 .!.-". G. Miami Sea Grant Coastal Zone Manage. Walsh, S. Snedaker and H. Teas, eds. Bull. 4. ID pp. Proceedings of the international sym­ posium on the biology and management Tabb, D.C., and A.C. Jones. 1962. Effect of mangroves. Univ. of Florida, of Hurricane Donna on the aquatic Gainesville. fauna of North Florida Bay. Trans. Am. Fish. Soc. 91: 375-378. Teixeira, C., J. Tundisi, and M.B. Kutner. 1965. Plankton studies in a mangrove Tabb, D.C., and R.B. Manning. 1961. A environment. II. The standing stock checkl ist of the flora and fauna of and some ecological factors. Biol. northern Florida Bay and adjacent Inst. Ocean. Sao Paulo, Publ. 14: brackish waters of the Florida main­ 13-41. land collected during the period July 1957 through September 1960. Bull. Teixeira, C., J. Tundisi, and J.S. Ycaza. Mar. Sci. 11: 552-649. 1967. Plankton studies in a mangrove environment. IV. Size fractionation Tabb, D.C., and B.J. Yokel. 1968. Report of the phytoplankton. BioI. Inst. to the Conservation Foundation: pre­ Ocean. Sao Paulo, Publ. 16: 39-42. liminary ecological study of Rookery Bay Sanctuary, Naples, Florida. Teixeira, C., J. Tundisi, and J.S. Ycaza. Univ. of Miami, Inst. Mar. Sci. 1969. Plankton studies in a mangrove 26 pp. environment. VI. Primary production, zooplankton standing-stock and some Tabb, D.C., D.L. Dubrow, and R.B. Manning. envi ronmenta1 factors. I nt. Rev. 1962. The ecology of northern Flor­ Gesamten. Hydrobiol. 54: 289-301. ida Bay and adjacent estuaries. Fla. Board Conserv. Tech. Ser. 39. 79 pp. Terborgh, J.W., and J.R. Faaborg. 1980. Factors affecting the distribution Taylor, D.L. 1980. Fire history and man­ and abundance of North American induced fi re prob1ems in subtropi ca 1 migrants in the eastern Caribbean south Florida. Proc. fire history region. Pages 145-155 .!.-". A. Keast workshop. Tech. Rep. RM-81, Rocky and E.S. Morton, eds. Migrant birds

103 in the neotroRics: ecology, behav­ Voss, G.L., F.M. Bayer, C.R. Robins, M. lor, dlstnbutlOn, and conservation. Gomon, and L I. LaRoe. 1969. A Smithsonian Inst. Press, \~ashington, report to the National Park Service, D.C. Department of the Interior, on the marine ecol09Y of the Biscayne Thayer, G.W., H.H. Stuart, W.J. Kenworthy, National Monument. Inst. of Marine J.F. Ustach, and A.B. Hall. 1978. and Atmospheric Sciences, Univ. of Habitat value of salt marshes, man­ Miami, Fla. 128 pp. groves and seagrasses for aquatic organisms. Pages 235-247 in Wetland Wade, R.A. 1962. The biology of the functions and values: thestate of tarpon, Megalops atlanticus, and our understanding. Am. Water Resour. the ox-eye, Megalops cyprinoides, Assoc., r1inneapolis, Minn. with emphasis on larval development. Bull. Mar. Sci. 13: 545-622. Thorn, B.G. 1967. Mangrove ecology and deltaic geomorphology: Tabasco, Waisel, Y. 1972. Biology of halophytes. Mexico. J. Ecol. 55: 301-343. Academic Press, New York. 395 pp.

Thom, B.G. 1975. ~Iangrove ecology from a Walsh, G.E. 1967. An ecological study geomorph i c vi ewpoi nt. Pages 469-481 of a Hawaiian mangrove swamp. Pages i.!! G. Walsh, S. Snedaker and H. Teas, 420-431 in G.H. Lauff, ed. Estuaries. eds. Proceedings of the interna­ American~ssociationAdvancement Sci­ tional symposium on the biology and ence Publ. B3. flashington, D.C. management of mangroves. Univ. of Florida, Gainesville. Walsh, G.E. 1974. ~angroves: a review. Pages 51-174 ln R. Reimhold and Tomlinson, I.E. 1957. Relationship W. Queen, eds. -!::cology of halophy­ between ma ngrove vegetati on, soil tes. Academic Press, New York. texture and reacti on of surface soil after empoldering saline swamps in Walsh, G.E., R. Barrett, G.H. Cook, and Sierra Leone. Trop. Agri c. 34: T.A. Hollister. 1973. Effects of 41-50. herbicides on seedlings of the red ma~9rove, Rhizophora !!Jangle L. Bio­ Tomlinson, P.B. 1980. The biology of SClence 23: 361-364. trees native to tropical Florida. Harvard Univ. Press, Cambridge, Mass. Walsh, G.E., K.A. Ainsworth, and R. Rigby. 1979. Resistance of red mangrove Tschirley, F.H. 1969. Defoliation in (Rhizophora mangle L.) seedlings to Vietnam. Science 163: 779-786. lead, cadmium, and mercury. Biotrop­ i ca 11: 22-27. Tundisi, J. 1969. Plankton studies in a mangrove environment -- its biology Wang, J.C., and LC. Raney. 1971. Distri­ and primary production. Pages 485­ bution and fluctuations in the fish 494 i.!! Lagunas Costeras, un simposio. fauna of the Charlotte Harbor estu­ UNAM-UNESCO. Mexico, O.F. ary, Florida. Mote Marine Labora­ tory, Sarasota, Fla. 56 pp. Twilley, R. 1980. Organic exports from b1ad man9rove fores ts in south Wanless, H. 1974. Mangrove sedimentation Florida. Ph.O. Dissertation. Univ. in geological perspective. ~:iami of Florida, Gainesville. Geol. Soc. f1em. 2: 190-200. van der Pijl, L. 1972. Principles of Warner, G.F. 1967. The life history of dispersal in higher plants. Springer­ the man9rove tree crab, Aratus Verlag, New York. pisoni. J. Zool. 153: 321-335. Voss, G. 1969. Preserving Florida's Warner, G. F. 1969. The occurrence and estuaries. Fla. Nat. 19: 2-4. distribution of crabs in a Jamaican

104 mangrove swamp. J. Anim. Eco1. 38: Es tuar i ne bi 0 logy. The Conservat ion 379-389. Foundation, Washington, D.C. 112 pp.

Weinstein, M.P., C.M. Courtney, and J.C. Yokel, B.J. 1975b. A comparison of ani­ Kinch. 1977. The Marco Island estu­ mal abundance and distribution in ary: a summa ry of phys i cochemi ca1 similar habitats in Rookery Bay, and biological parameters. Fla. Sci. flarco Is land and Fakahatchee on the 40: 97-124. southwest coast of Florida. Prelim. Rep. to Deltona Corp. Rosenstiel Westing, A.H. 1971. Fores try and the war School of Marine and Atmospheric in South Vietnam. J. For. 69: 777- Science, Univ. of Miami. 162 pp. 784. Yoshioka, P.M. 1975. Mangrove root com­ Witham, R.R., R.M. Ingle, and LA. Joyce, munities in Jobos Bay. Pages 50-65 Jr. 1968. Physiological and ecolog­ in Final report on Aguirre environ­ ical studies of £.. argus from St. mental studies. Puerto Rico Nuclear Lucie estuary. Fla. 80ard Conserv. Center. Tech. Ser. 53. 31 pp. Zieman, J.C., Jr. 1972. Origin of circu­ Wolff, W.J. 1969. Distribution of non- lar beds of Thalassia testudinum in breedi ng waders in an es tuari ne area south Biscayne Bay, Fl or i da, and in relation to the distribution of their relationship to mangrove ham­ their food organisms. Ardea 57: mocks. Bull. Mar. Sci. 22: 559-574. 1-28. Zieman, J.C .• Jr. (in prep.) A community Wolffenden, G.E., and R.W. Schreiber. profile: the ecology of the seagrass 1973. The common birds of the saline ecosystem of south Florida. U.S. habitats of the eastern Gulf of Mex­ Fish and ,iildlife Service, Office of ico: their distribution, seasonal Biol09ical Services, Washington. D.C. status, and feeding ecology. Pages 1-22 in J. Jones, R.E. Ring, M.O. Zuberer, D.A., and W.S. Silver. 1975. Rinkel and R.E. Smith, eds. A sum­ flangrove-associ ated nitrooen fixa­ mary of knowledge of the eastern Gulf tion. Pages 643-653 j.!!. G.-Walsh, S. of ~:exico. Inst. of Oceanography, Snedaker and H. Teas, eds. Proceed­ St. Petersburg, Fla. 604 pp. ings of the international symposium on the bi 0logy and management of ,lan­ Wood, E.J.F. 1965. Marine microbial groves. Univ. of Florida, Gaines­ ecology. Rei nho I d Pub 1. Corp., New ville. York. 243 pp. Zuberer, D.A., and W.S. Silver. 1978. Yokel, B.J. 1975a. Rookery 8ay land Biological nitrogen fixation (acety- use s tudi es. Envi ronmenta I pI an- lene reduction) associated with ni ng s trateg i es for the deve1opment Flori da mangroves. App I. Envi ron. of a mangrove shorel ine. Study 5, ~licrobiol. 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 Table A-l. Site characteristics and sampling methodology for fishes in mangrove-fringed tidal streams and rivers.

Mean depth; Number Salinity Temperature tidal species Loca ti on range range range Subs trate Benthic vegetation Sampling methods Frequency recorded

0 0 North Ri ver; 0-27 0/00 15.4 _33.2 C 1 ffi; largely exposed Scattered ~ Bag seine. throw r~onthly. Sept. 55 Tabb 1966, 0.5 m 1imestone. and maritima near mouth nets, dip nets, 1965 through Odum 1970 sand banks; traps, pound net, Sept. 1966 undercut man- fish poison, rod & (Tabb) grove peat reel, trammel net. set lines 0 0 Cross Bayou 3.2-29.8 0/00 13.0 _31.5 C Max. Hard muddy sand Sparse Bag seine; minnow Monthly, Sept. 60 ..... Canal (Boca depth Enteromorpha se; ne 1957 through 0 Ciega Bay to 1.5 ffi; Dec. 1958 " Old Tampa Bay); 0.9 m Springer & Woodburn 1960

Fahkahatchee 2-36 0/00 22° _29°C Less than Not given Not given Seines routinely; Monthly, Jan. 47 stream, 1 m; range black net &rote- 1972 through stream enterin9 not given none for single Dec. 1972 Fahka Union standing crop Cana 1; Carter estimate et al. 1973 o Unnamed streams 16-32 0/00 13.20 _37.l C 1.1 m; Thick organic Dense Thalassia Gill nets; hoop Weekly, gill 52 near Turkey 0.5 m mud-gi 11 net & testudinum at nets; traps nets; bimonthly, Point, Biscayne trap sites; mouth others; August Bay; Nugent culvert at hoop 1968 through 1970 net sites Dec. 1969

Total III Only taken in SE Fla. -23 88 Table A-2. Site characteristics and sampling methodology for fishes in mangrove-lined estuarine bays and lagoons.

~'ean depth; Number Salinity Tempera ture tidal species location range range range Substrate Benthic vegetation Sampling methods Frequency recorded

0 o Fahkahatchee Average ­ 21 _31 C 1. 2 m; Generally Extensive areas of Vegeta ted, mud, Monthly, July 47 Bay, 740 ha; 15-37 0/00, (18ke1 6; muddy, some Halodule wrightii , sand/shell bottoms 1971 through (Yokel) Yokel 1975b, low of 1 0/000 23.5 -32 C sand & shell lhalassia testudinum sampled by otter July 1972 Carter et a 1. recorded by (Carter in northern portion trawl (Yokel); 2 (lokel); 89' 1973 Yoke 1, Sept. et al.) bay seines, otter Monthly, Jan. 1971 trawl, surface 1972 through trawl (Carter Dec. 1972 et a1. ) (Carter et a1.)

Fahka Union 5-35 0/00 Muddy, Little seagrass, 2 bag seines, Month ly, Jan. Bay, 186 ha; subject to occasional high standing crop otter trawl, 1972 through Carter et al. sporadic sandy area of green algae surface trawl Dec. 1972 1973 massive or oys ter freshwater bac a inp~ts from ­co GAC drai nage cana 1s

Rookery Bay, 8.9-38 5 0/00 0.9 m; Hud, sand, Halodule wrightii, Vegeta tM, mud, Monthly, June 64 419 ha; .55 m shell ThaTassia testudinum, sand/shell bottoms 1970 through Yokel 1975a l~al.1WJ.!.iJa engelmannli sampled by other July 1972 trawl

Marco Island 19 0/00 Sept. Not given Mud, muddy Hillodulc wr{9htit vegetuted, mud, ~1onthly, July 59 Estuary; Yokel 1971; other­ sand. shelly bedS in sha low sand/shell bottoms 1971 through (Yokel) 1975b; Wcin­ wi se over a sand back-bays of man­ samplpd by otter July, 1972 82­ steinetal. 4-yr period grove complex; trawl (Yokel); (Yokel); (Wein~ 1977 29-39 0/00 Thalassia not well otter trawl (Wein­ Monthly, July stein) developed stein et al.) 1971 through Jan. 1975 (Weinstein et a1. )

~ti~, Roller frame trawl Monthly - 8 67 Whlte\·/ater 2.9-29.3 0/00 Shallow Peat, silt, Halodule stations­ marl P. shell, Udotea conylutinata, s ta ti ons Bay, Clark Sept. 1968 1970 1 Ill, deep sand & shell Chara hornC'rllanOl, stalions­ Qa_sy~ p-e-J{c'cfla ta, through 0.8-1.0 m; Gracilaria sp., Nov. 1969 0.6 m Ha10phila baillonis Total 117

a89 species in rahkahatc.:hee and rahka Union Bays combined. bGulf American Corporation. Table A-3. Site characteristics and sampling methodology for fishes in mangrove-lined oceanic bays and lagoons.

Mean depth; Number Sal inHy Temperature tidal species Location range range range Subs trate Benthic vegetation Sampling methods Frequency recorded

._--~

Old Rhodes Average Apr; 1- Average for 0.61 m; Seagrasscs: Visual counts. Monthly, 1973 31 Key lagoon. June 1973 - Apr. -June O.S m Thalassia testudinum traps. hool< and o Holm 1977 37 0/00 1973 - 28 C range 1ine 0 0 Porpoise lak.e. 27.8-49.6 0/00 16.6 -32.2 C f1ax. Carbonate Seagrasses: exten- Suction sampler, Monthly, Apr;l 64 Florida Bay, depth mud & shell sive Thalassia slednet. pushnet, 1965 through Hudson et a1. 2.1 m; fragments testudlnum. sparse beach seine, cast Jan. 1968 1970 Italodule wrigh_'tiJ.. net. roller-frame trawls, hook &line

0 0 ~lonth ~ Southern 5.0-43.8 0/00 13.5 _38.7 C' Range Mud. sand. Seagrasses: Otter trawl 1y. Ju ly 75 0 Bi scayne Bay, 1.0-2.5 m coarse Thalassia testudinum, 1968 through '" Bader and sand & shell Halod~l~ ~htii, June 1970 Roessler fragments red algue-Lilurencia. 1971 Oigenia Wes tern Not given Not given Not given Not given Not given though Gag seine. semi- Monthly, May 109 Florid~ Oay author sta tes each balloon otter 1973 throu9h 315 km , representative trawl June 1974 at Schmidt 1979 benthic habi tat 12 stations was sampled Tota 1 155

aSome sampling stations were within the area of the thermal plume from the Turkey Point power plant. temperature elevation up to 5.2oC above ambient. P~PENDIX 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 Lrnportance.

110 Did :a~ily and Species Refercnc,-, Olt1t :tei.,.fc:nc<::'

orcctolobidae - carp€t snar},s Ginqlynosto~ cirratC3 1, 5, 7 Fish. cEr;halopocs, :-:olluscs, ?.ar:dall 1967 nurse sha:.-k s~i~?, sea urchins Cl.:;.r:: s; -.-on :::c1-.:::lid'C 1965 Eohl}:€: S C1Hpl:.n l'?ES

Carcha:.-hinicae requi::::. shari:s 3 Carcharhinus leucas 8 J ...·J~nile3' fi::;n (~~, cc.= 1971 bull shark :'ol:noco:'i'Js, :-:'.15i1 cE:l:'al•.ls, 5r~~oa~~ia ca't~o~us. :";icro'::o:;on unOulatus). crus­ taceans incluuing ~E~~E:C shri=?, blu~ c~abs

Carcharhinus li~ tus II :"ish (Car~!'..:·: sp.• Ccn'tro:.o=c;s clar;: <; blac;;tiI) shark und~i-~1i5, Chilo "lc~Lrus \-on Sch:::liet. scho&~fi, nrius ~, ~cto­ i

!ieca::.rion breviros'tris -i. 5, 7 YounG: crustac:ans. fish ~!lcall 1967 lc=n sha.d: hcults' ~ish, crustaceans C1

Sphyrnicac - h~-erhead sharks Spn"lrna t i huro ­ 2, 5 ~antis shri=?, s:'ri=p, lSOpoCS, ?ohl:':e S honnethcad bar~ac1cs, oi~alve oo11uscs, Cha?11n 1968 cep:'alopoCs, fi~:'

?ris'tidaa - sawfishes ?ristis oectinaUi 5, 1S Fish. benthic c:.-ustaceans 3Oh1ke to s=alltooth sawfish Chaplln 1968

R.~ino~tid

To::?£:cinidae - electric raJ's !larcinc: brasiliensis­ + + I, !1. 18 lesser electric re.'l

?..ajidac - SJ.""...atC5 CruStacCA. ri5h, annelids R~id 195" ?..aja~ rOlrleel sJ.:":li:e

Das'latidae - stingrays D5s"latis a=cricana ­ + + + :2,4, 5, 7 'ishes. sipunculid anc pol:;- ;::a~call 1967 southam stingray chaste ~o~. crab~, bivalves, shr~p. =antis shricp

Das1atis sabina 2. B. 13, 3cnthic invertebrates inc1u- Darnell 1958 ;;t1a:ltic stingray 17 ding bivalves. xanthid a~d port~~id crabs, silri~ps, a~~hipods, annelids. chiro~o- ~irl 1arv.1e G\'r:!llura cicrura ­ + 17 ?i5h, l::olluscs. annelids, ?eterson • s=ooth butterfly shrio? ot..l1er s::::.a1l peterson 1979 ,ay crusUlceans

Uroloohus ia::.aicensis­ + ! ?.robaDl·", s;:.all burro-"'ing BOhH:e , ¥e11a~ stingray inver1:ebrates Chaplin 1908

8!his and all subsequenL Odu::J 1971 cital:ions refer 1:0 lo'.E. Odu::::; 1971.

111 .- Did :a=ily and Spect~s Refer.:once R.,i ~ r~nc" C~nl

~iliooaticae - eagle rays ';etoDatus na.rina.ri ­ 2 3.ohlr.o:1 !O spotted eagle ray !!.3plln 19&3

Lepisosteidae gars (~eiliids. Od~ Lepisosteus olatZJ::hineus - I" 2. 1. 13, Fish cyprinodonts. 1971 Plorida gar 1S sr"",ll cenrrarchids), crustaceans (caricean 5hrl~?}.inseer lar;a~

Elopidae - tarf'O:'ls ::10p5 ~­ • 2, J, 7, < ~5 ~: zoo?la~to:'l. c~aeto­ ~= 1971 lac]!ish 8. 13, 15 Sna~~$. ?olychaet~ Austin I> ·... o~.....s ';'.1stb 1971 > ~5 ~: caridea-~ 5 pe~aeid 8hri~, various s=all :hh

?~~ale~s atl~~t1ca ­ 7, e. 13, < ~5 ~: ?la.~kto~ (~lC!o;oid O:l:c. 1971 O:>liSil::'e. air :tIreato":", r"S. .Juv­ t.L"1'On 15 co~ 's) ':'\l.!O,:i:l & j~~eni1e5: fish (G~~ia, .:.us~in 1971 enilell inh..!;)! ~ ;~"dulus ~cteroclit~. ~~;il sha1lc~ orac~ish ce~halus), crus~ce~"5 (ostra­ ?DOls lcr~ in oxyqe.n, c -$, carlc!eil~ s;~.ri=;)) often containing adults: ~id~ ~arlety o~ tis~, ~:S (nac~ 1962) c~a~~, shri=? ctenophcres, insec=

;'lbuli::h.e - hone fishes ~ vulP$$­ 4, S c1a~, snails, shri~, s~ll 30hlke to bonefi:Jh fish Ch ...plin 1968 h...,guillidae - eels ~"9uilla rostrata ~, 13 50-200 ~, ~?hi~s. isop0d3 Cdu::. 1971 r..::.e.rican eel 180-472 ~: xanthic crabs. caridea~ shri~? fish (Loohosooius ~luri~oi~e5)

O?hichthidac - sn..1o;.e eels ~(ro:;his cunctatus • 2, 3. n, ?o1ychaetes, 3ranchio5~ ;;prin;er '" Y~rs of this specklel:! WOr.:! eel 18 car~~~, s~~d cr~hs \l:oodk-':J...~ fa.:lily b=ro-... 1960, in ::::;"Jd or sa."ld. Reid 195': u.,eersa=pled ~7 =cst :rti:.ods (Eohlke & Chaplin 1908)

aascanichthy~ scuti­ ) ~ Ahip eels

Onhichthus 900esi ­ 3, 17 Shri::;> eel

Clupeidae - herrings Brevoortia 5~thi ­ 2, S. 17 yellv"'fin s~

3re~~rtia patronus ­ 12 38-~8 ~: phyto?1ankton, zoo­ Darnell Gulf ~hacen ?1a."'l1:to~, plant frao;=oc:n=. 1958 d!'! ritus 85-103 =: organic :::att.er, silt. ciato~, for~ni!era~s, C0?epoC5

112 Habitat rype

Diet Family and Species Reference Diet Reference Comments

Harengula pensacolae + + + 2, 3, 8, 30 m~: planktonic copepods, Odum 1971 scaled sardine 13 zoea, naupli!, larval fish 64-96 mm: amphipods, harpacticoid copepods, isopods, mysids, chironomid larvae

Opi~thonema oglinum ­ + + + 2, 3, 5, Copepods, polychaetes, shrimp, Odum 1971 Atlantic thread herring 13, 17 fishes, crab larvae, mysids

Sardinella anchovia + 17 Spanish sardine

Engraulidae - anchovies Anchoa cubana - + 2, 16 Ostracods, copepods Springer & Cuban anchovy \.o:oodburn 1960

Anchoa hepsetus ­ + + 2, 3, 13, 32-114 rom: copepods, isopods, Springer & striped anchovy 16, 17 mysids, caridean shrimp, small ~,loodburn bivalves 1960

Anchoa lamnrotaenia ­ + 5 bIgeye anchovy

Anchoa mitchilli + + + 1, 2, 3, <25 rom: microzooplankton Odum 1971 bay anchovy 5, 7, 8, 31-62 mm: amphipods, zooplank- 13, 16-18 ton, mysids, ostracods, plant detritus, copepods, sw~ll molluscs, chironomid larvae

Synodontidae ~ lizardfishes Syoodus .~etens + + + 1-3, 5, 8, Small fish, crabs, shri~p, Odum 1971 inshore lizardfish 17, 18 polychaete worn.s

Catostomidae ~ suckers J~zson sucetta ­ + A freshwater lak-e- -c1iUbslicKC'r- stray ictnluriJae - freshwater catfish Ictalutus natalis ­ + 14 A freshwater yellow bll1lhead stray

Noturus zy!inus ­ + 14 A freshl\'uter tadpole :uadtom stray

Arriidae - sea catfishes Arlus feli_~ - sea catfish + + + 2, 3, 5, 100 W~: copepods, zooplankton Odum 1971 7, 8, 13, amphipods, mysids, chi ronomid 17 larvae, isopods, small crabs 100-200 rom: benthic inverte­ brates 200-330 rom: crabs, anphipods, mysids, fishes, bark, crayfish, catidean and penaeid shrimp

Bagre ~rinus ­ + + 2, 8, 17 262-445 nun; blue ctabs, small Odum 1971 gafftopsail catfish fishes

Battachoididae ­ toadfishes Opsanus beta ­ + + + 1-3, 5, 18-60 m:n: amphipods, chironomid Odum 1971 Salinities Gulf toadfisil 7, 12, 13, larvae, mysids, isopods, few fish 10 0100 > 15, 17, 18 >60 mm: caridean shrimp, xanthid (Odum 1971) crabs, snapping shrimp, mussels, fish, mangrove hark

113 Diet Family and Species Diet Reference Comments

Porichthys porosissimus + 3, 18 Atlantic midshipman

Gobiesoeiclae - elingfishes Gobiesox strumosus + + + Z. ] . 5. 8 10-32 mm: amphipods, isopods, Odurn 1971 skl11etflsh chlronomid larvae

Ogcocephalldae - batfishes Ogcocephalus nasutus + + 18 50a11 bivalves, gastropods, Reid 1954 shortnose batfish polyehaetes

Ogeocephalus radiatus + Z. 11. 17, polka-clot batfish 18

Gadidae - codfishes UrophyciS floridanus + 12 Amphlpods, isopods, mysids, Springer & A species more Southern hake decapod shrimp, polyehaetcs, \~oodbu[n common at =" insect larvae, fishes (Lagodon 1960 northerly rhonbo1des, Paralichthys latitudes albigutea)

Ophidiidae - cusk-eels, brotulas Gunterlchthys longipenis + 17 gold brotula

Ogilbia eayorum + + I, ] key brotula

Ophidion holbrooki + ] bank cusk eel

Exocoetidae - flying­ fishes, ha1fbeaks Chriodorus atherinoides • 5 hardhead ha1fbeak Hyporhamphus unifasciatus • • 2, 3, 5 ju~eni1es: zooplankton including Carr ~ halfbeak crab megalops, ve1igers, cope- hda~s 1973 pods 130-199 IIUII; epiphytic algae, detritus, seagras8

Belonidae - needlefishes Stron9ylura marina • • 2, 7, 15 357-475 nffi: s~all fis~es, Darnell Atlantic needlefish insects, shrimp, small amolli~ts 1958 of vascular plant material and algae Strongy1ura notata • • • 2, 3, 5, In grassbe::ls Brook redfin needle::ish 8, 13 Juveniles: po1yc~aete worms, 1975 cumaceans, fish Adults, fish, pri~ri1y ntnC'rinids Strongylura timucu • • • 2, 3, 11 159-37B ~: anchovies. shrim? Ra:ldall Primarily ins~ore timucu 1967 species~freely enters fresh­ ",ater (Randall 1967)

Tylosurus crocodilus­ • 11 250-1320 rom: fishes, shrimp Randall Open water and houndfish 1967 inshore surface water inhabitant (Voss et al. 1969)

114 Diet Family and Species Reference Diet Reference Comments

Cy?rinodontidae - killi­ fishes Adinia xenica ­ + 2.6,13-15 Plant detritus, diatoms, Odur.l 1971 ~d killifish amphipods, harpacticoid copepods, insects

Cyprinodon varicqatus sheelJshead cinna... + , , 2, ,, ., Plant detritus, algae, O

floridichthys carpio , 2, J, 8, ""..nphipods, ostracods, isoporls, 0<1= 1971 goldspotted ~illifish 13 copcpods, chironoll'.id larvae, nerr.atodes, plant detritus, algae

Fund~lus conflucntus + 2,8,13-15 Caride.:an shrimp, small fish, Odum 1971 narsh ~illifish (Gambusia affinisl, ~phipods, isopods, a~ult & larval insects, copcpods, mysids, ostracoCs, algal filaments

Fundulus chrysotus + 14. 15 Rare in mangrove golden topni:lno....' zone, headwater pools on,ly

Fur~ulus grandi~ 2,8,13-15 I\mphipods, isopoos, xanthid Odwn 1971 CuI f killifish crabs, chirononid larvae, terrestrial insects, snails, algae, small fish (poeciliidsl

fundulus hctcrocli~us + , Sm.Jll crustaceans l~phipods, Peterson & 11ummichog------~ isopods, ostracods, t2n~ids, Peterson cope?Ods), detritus, polychaete 1979 worms, insects, snails, inver­ tcb.,-:ate eggs

14-1 "i Pri..m.l.rily a freshwater form, nead....·ater pools only

11-15 Primarily fresh­ water, cammon in pools in headwater regions

~3E?2..~ + 2,8. )3·1', Snall crustac-edns bOpCpoC5, Od\ll:l 1971 btucfin killifish lleadwatcr pools cladocerans, ostracods), insect and channel larvae

~parva + + 1-3. 1;" 8. <20 mm: ?lanktonic c0?epods rainwater kill:fi~h OC:um 1971 U-1S" 17 21-37 ~: amphipods, rr~sids, chiror-o~d larvae, ostracods, molluscs, plant detritus

A'larll"JOra~us Ri·.. ulus + 3,8,13,15 rivIOlus

Pocciliidae - livebearers Grm:usia a:finis 2, 3, 7, mos<:.uitof~ A versatile feeder: dITlphipods, Cd\ll:l 1971 13-15 chironomid l~rvae, hydracarina, harpacticoid copepods. s~ils, ants, adult insects, ?Qlychaetc \oIOrns, ostracods, mosq",ito pupae, algae

Gambusia rnizophorae + 6, 9 mangrove gambusia Fresh and braCkish water in Rhizophora swamps, nOrthern Cuba, southeastern Florida

115 Diet Fanily and Species Rercrence Diet Reference Coments

Hctcrandria fornosa 8. 14, 15 Chironomid larvae, narpacticoid Odum 1971 least ki lli:~ and ?lankton1C copepods, clado- cerans, terrestrial inse~ts, algae, diatolf.S

poc~ilia latipinna S, 7, 8. Plant detrItus, al~ae, diatons Cdum 1971 sailfin rlOlly 13-1S

Atherinidae - silv~rsldes AIIanptta harringtonensis 39-60 mn: copppods, fish larvae, Randall reef siIvE"Isi::lf!' polyc:,aete larvae 1%7

Mcmbras cartinica 2. 5. 11 S:na 11 zooplankton ~rustaceans, Peterson &­ ro~qh Si:VE"fSlde juvenile &- larval fishes, Peterson insects. detritus, snails 19B

~enidia bcryll1r.a 2, J. a. Insects. COpCpod3, chironomid Od..lT.l 1971 tid~~ater silv~rsidE" 11. 1:1.. larv~p. ~ysids, an?hipods n. 1,. 1.

Sy::lqnathidile ­ pipefis~p~. s~a~ors~s ('orythoic~!:m. ~, 11 albirostris ~~~~~ipc~~sh Associated with lI..i....P.pocampu~ ~~ 1 • 2. ~. 11., vegetated areas {Tabb 11 ned s(>ahor~p 1J fl !·lann1n':j 1961)

h i ':)E~campu~ ~.!:'E~e \, 2. ). c'. Intimately associated d ....·ur"" s ...ahor!>p 11. 16. 1.7. with unattached algae '8 ('ral::b 6: Manning 1961), or grassy areas (Springer & "O'Joodburn 1960)

Mlc~ccnd:h~s cri::liger~s 1.. 5, IJ C,2-R2 mm: l'Opq::oc.s. micro­ Rei:] 19')4 fringpd pl?cfish crust;;cea:ls

§}'n:F-:3t~ floric.ae 1-3,~. 1: Ca!"lc.car. shri'TIp, a:np::i?Qds. Brook 1975 d'Jsky pi Fcf. sh tanai::!s, iso;x>:]~

5vngnathus louisianae 1-). 11. :opepods, ~~~hipods, sm~ll ~e:d 1954 Inh,1bit grassy c::'air. ?ipef:..sh 16-:5 'i:':.Ti:"ltl flats (Springer & \.;oodturn 1960)

~ynqnathus s=cVE":l: 1-3. ;;. Il­ ',mr-':':ipoCs. is:)~:15, rancHes. Brook 1975 G'J:f r:ir-e:·.ah l F.-I ... COPE'"P.:cs, 'tir.y ~aric.ean Sprin']cl" '" ShCIFO. q~st~cp~s (Ei~t;un. \·:ooob'..:rn 131;0 14itrell':') ---- Reia }QS4

Synqna:h~s ~Frin9pri :2. I:' ~.:: 1 ~jl-'f>flsh

syncna:~~s cur.ck~~ 11 i\ssociated 'with ~ugr.osp ?locfish vegetated a!"eas (:abb & :.talming 1901J

~~ pcl,lJi:::us ~arqa~su~ ?ipc:ish

Cer.:r0I'0:--idJoc - snoo:<5 ce~tropo~us ~aral:eIus Fanily as a whole ~at sr.cok shows preference for estuarine :nan­ grove ho1bi tat (Rivas 1962)

Centroponus ectinatus 7, 3. 13 tarpon snook 116 Habitat Tvpe

Diet Family and Species Refetence Iliet Reference COQlf'tents

Centropomus undeciroalis + + + 2,5,7.8. JuveniJes, caridean shrimp, OCum 1971 iOy far most snook 11, 13, 14 small cyprinodont fisr.es. Austin & ab\;ndant of gcbies, nojarras ;'ustin three species Adults: fish, crabs, penaeid 1971 (Rivas 1962) shrimp, crayfish, snapping shrimp

Serranidae - sea basses Centropristis striata 11 Family in general carnivorous Randall black seabass on fish, crustaceans 1%7

Diplectrum formos~ + 2, 3, 11, Caridean &. pen"eid s::trilllp, Reid 1954 sand perch 16-18 cO?epods, crabs, :ish

Epinephelus itajara + + + 2.5, 7,8, Juveniles: penaeic shrimp, 1971 The most abundant jewfish 11, 13, 15 xanthid crabs of t:1.e sca::>a:o:ses i~ "~n9rOve habitats

EPinephelus nlOri<:· 11 226-)40 mm: crustaceans, Randall red grouper crabs. fishes 1961

Epineehelus striatus 4, 11 170-686 mro, fish, crabs, Randall Nassau grouper sto~topods, cerha1opocis, L9E7 shrimp, spiny lobsters, gastropods, bivalves, isopods

Hypoplectrus puella + 11 54-98 mm, sna?ping shrimp. Randall barred hamlet crabs, fis~, mysids, sto~ato~ 1967 pods;, isopods

Mycteroperca ~rolepis + + L 2, 5, 1.1, 71.-100 rrun, peflileid shrimp, Reid 1954 gag 17, 18 fi!:lh Centrarchidae - sunfishes Elassoma evergladei Evorglades pygmy sunfish + 14 Family is primarily fresnwater, fis~ occasionally enter hcad'~ater area of mangcova­ fri.nged stre,l;:!

Lepomis .auritus 14 redbreast sunfish

LeDOnis qulosus 2, 13, 15 Shrimp (Palaemonetesl, =ish Desselle et Diet from Lake warmouth {Cobiosoma besd, Lepomis a1. 1978 Pontchartrain macrochiT~sl, detritus, salini ties J _6­ Vallisneria, am?hipods, xan­ 4.1 0/00 thid crabs, blue crabs

Diet fron Lake ~ macrochirus 2, 15 ~phipods, blue crab (Cal- Dessell< et Pontchartrain blueqil.1 l inectes sapid'J.s), xanthid a1. 1978 crabs, detritus, Vallisneria, salinities 1.6­ cla>:ls (Rall9ia cuneata}. 4.10/00 sponge (Ephydatia fluviatilis), barnacles, insect larvae

Lepomis microlophus + 2, 13-15 Chironomid larvae, amphipoes, Desselle et Diet from Lake redear sunfish xanthid crabs, clam (Ranqia al. 197a Pontchartrain cuneatal, sponge (Ephvdatia salin1 ties 1.6­ fluviatilis), detritus 4.1 0/00

Lepomis punctatus + 6, 14, 15 Cladocerans, small crabs, Odum 1971 Salinities < 15 0/00 spotted sunfish mysids, chironomids, ~phipods, (OdUl':l 1971) insects, molluscs, isopods, fish, algae

117 Habitat Type ~ Iu ';j ~ ~ 'd ~ Diet Family and Species .." ... 3 >./ >, ;::: t: :l 2 :s ~ Re[erence Diet Referenee Comments

"icropterus salmoides + 13-15 Caridean shrimp, small blue Darnell 1958 largemouth bass crabs, crayfish, xanthid crabs. 2S species of fiSh, Vallisneris, Cladophora Apogonidae - cardInalfishes Astrapogon alutus + + bronze cardinalfish " 3

Astrapogon stellatus + 1 conchfish

Pomatomidae - bluefishes Pomato~us saltatrix + 11 Young; mainly fishes (anchovies, Peterson & bluefish silversides, killifishes, cen­ Peterson 1979 haden, shad, spotted scatrout), shrinp, crabs, other small crustaceans, annelids, snails Rachycentridae - cobias Rachycentron canadum + +5,7,11 fish, crabs Randall 1967 cobia

Echeneidae - remoras Echeneis neucratoides + + + 2, 11 Fish, lsopods, other crustacea Randall 1967 Members of thls whitefin sharksucker fa-oily attach to sharks and large bony fishes (Randall 1967)

Remora~ + 7 58-175 ~m; copcpoda, isopods, Randall 1967 remora vertebrate musclc tlSStie, crab larvae, fish remains, crusta­ ceans, amphipods

Carangidae - jacks, pOmpanos Caranx ~ - blue + + + 2, 4, 5, Family of s ....ift­ runner 7, 11 swimmln~, carnh-­ orous fishes, often running in schools, -.'idc­ ranging (Randall 1967)

~hippos + + + 2, 5, 7, fishes, crustaceans Odum 1971 crevalle jack 8, II, 13

CaratU: ruber + 4. 11 160-547 mm; fish, shrimp, myslds, Randall 1967 bar jac-k-- stonstopods, gastropods

Chloroscombrus chrysurus + + 2, II, 17, Atlantic bu~per 18

Ollgoplites saurus + + + 2, J, 5, Snapping shrimp, penneid shrimp, Tabb & leatherjacket 8, 11, 13 larval anchovies, ladyfish, !-tanning 1961 harracticoid copepods

Trachinotus carolinus + 11 sardines (Hnrcngula sp.), Spr:lnRer 6­ CODDlon over Inud Florida pompano mole crabs (~sp.), ,,"oodburn bottoll'-s (Randall bivalves (Donax sp.) 1960 1967)

Trachiootus falcatus + + 7, 11 15-70 mm; OOy81d8, shri~p, Carr & ,'dams Hore apt to occur permit anchovies, silversides, crabs, 1973 over sandy bottons snails than T. carolinas (Randall 1967)-

Selene \'OI:1er + + + 2, 3, 7, Young; shrimp and other Peterson & lookdown--- 11 crustac~ans, small molluscs Peterson 1979

118 Habitat Type • .~ " .::; >. Diet ":r. ~ Family and Species ~"'" Reference Diet Reference COMents

Hel:licaranx + 17 amblyrhynchus ­ bluntnose jack

Caranx latus + 12 Predaceous on other fishes Darnell 1958 Considered by horse-eye jack Gunter (1956) to be eur)'hal1ne

Lutjanidae - snappers Lut1anus analts + I, 4, 11 204-620 mn: crabs, fish, Randall 1967 COlmlonly found mut.ton snapper gastropods, octopods, hentit. over sand, sea­ crabs, penaeid shri~p. spiny grass, rubble, lobsrer, stomatopods coral reefs (:l.anda11 1967)

Lut1anus apodus + + I, 4, 5, Crustaceans (shrimp, snapping Nugent 1970 schoolmaster 7. 11 shrimp, blue crabs. xanthin crabs. grapsid crabs). fisr.

Lut1anus griseus + + + 1-3. 7, <50:nm: reside in grassbeds Odurr. 1971 By far the flOSt gray snapper 8, 11-13, feeding on s~all crustaceans, abundant snapper 15-18 insect larvae in mangrove 95-254 mm: reside in mangrove habitats creeks feeding on crustaceans (snapping shrimp, xanthid crabs, penaeld shrimp, crayfish, caridean shrimp), fish including gobles, anchovies, poeciliids, eels, k11lif1shes

Lutjanus iocu + 1 190-630 mm: fish, crabs, Randall 1967 dog snapper octopods, spiny lobster, gastropods

Lut1anus synagris + + + I, 2, ), snapping shrimp, crabs, Stark b Known fron brackish lane snapper 5, 7, II, anchovies. annelids, nolluscs Schroeder water to depths of 16-18 1970 220 fathoms (Randall 1967)

Gerreidae - mojarras Diapterus olisthostomus + 2 110-116 mm: green algae .\ustin G Irish pompano (Enteromorpha flexuosa, Austin 1971 Cladophora), Ruppia n:aritina. bluc-green algae (Lyngba ma1uscula)

Diapterus plumier! + + + 2. 7. 8, 36-172 ~: mysids. amphipods, Odum 1971 A permanent striped mojarra 11-13. harpacticoid copepods, resident (OdUD 15. 18 chironomid larvae, ostracods, 1971) bivalves. plant detritus

Euciaostomus argenteus + + + 1-5, 7, 19-63 rem: ~phipods, Odurn 1971 spotfin lIlojarra 8, 11-LJ, chironomids. harpacticoid 16-18 copepods. ostracods. mys~ds, molluscs, plant detritus

Eucinostomus gula + + + 1-3, 5. 19-70 nm: amphipods, chironomid Odum 1971 silver jenny 7, 8, larvae, harpacticoid copepods, 11-13, molluscs. mysids. ostracods, 16-18 plaot. detritus

Eucinostomus lefroyi + + 10 PlOttled IDOjarra

Gerres cinereus + + 2. 7, 11 Crabs. bivalves, gastropods, Randall 1967, yellowfio mojarra polychaete WOrDS, shrimp, Austin & ostracods Austin 1971

Pomadasyldae - grunts Family carnivorous though rarely Randall 1967 Most. shelter on piscivorous coral reef by day, feed on grassy flats by 119 night (Randall 1967) Diet Family and Species Reference Diet Reference Comments

Anisotremus ::5rginic~2­ + 11 112-264 mm: brittle stars, crabs, Randall porkfish shrimp, po1ychaetes, isopods, 1967 bivalves, stomatopods, gastropods

Haemulon aurolineatum 1, 4, 11 97-170 rom: shrimp & shrimp lar­ Randall tomtate vae, polychaetes, hermit crabs, 1967 amphipods, copepods, gastropods, bivalves

Ilaemulon carbo:1drium + 156-273 rom: crabs, gastropods, Randall Caesar grunt sea urchins, chitons , poly­ 1967 chaetes, brittle stars, sipun­ culid 'NOrms, shrimp

Haernulon flavolineatum 113-228 rom: polychaetes, crabs, Randall -~~------French gr'..lnt sir-unculid worms, chitons, 1967 ho1oth~rians, isopods, shrimp, bivalves

"arra~ Randall Haernulon + " 7 Benthic invertebrates including sailor's choice shrimp, crabs, a~phipods, gas­ 1967 tropods, polychaete worms, bivalves

Haernulcn plurnieri + + 1,2,11,18 130-279 rom: crabs, polychaete Randall white grunt "'"Drms, sea urchins, sipuncu1 id 1967 WOrr.1S, gastropods, shrimp, britt.le Reid stars; juveniles: copepods, rr,ysids 1954

Haenulon album + Benthic invertebrates including Randall margate crabs, shrimp, polychaete worms, 1967 amphipods, copepods, snails, bivalves

Haemulon sciurus + + 1, 3, 5, Benthic invertebrates including Randall bluestriped grunt 7, 11 crustaceans, molluscs, annelid 1967 worms

Orthopristis chrysop­ + 1-3, 5, Juveniles: 16-30 mm: plankton carr & Strong preference 11, 16-18 including copepods, mysids, Adams for vegetated sub­ =pigfish ?Ostlarva1 shrimp 1973 strate in bay >30 ~m: polychaetes, shrimp, areas (Weinstein arnphipods et al. 1977)

Sparidae - porgies Archosargus probatocepha­ + + 2, 3, 5, 7, <40 mm: in grassbeds - copepods, Odurn 1971 1u, 8, 11-13, amphipods, chironomid larvae, Austin & sheepshead 17-18 mysids, algae, molluscs Austin >40 rom: in mangrove creeks ­ 1971 mussels, false mussels, crabs, snapping shrimp, crayfish, hydrazoans, algae, plant detritus 32-85 rom: in puerto Rico man­ groves - lOO~ blue-green algae (~mojuscula)

Archosargus rhomboidalis + 5, 11 105-220 rom: seagrasses Cyn,odocea Randall usually seen in sea brea.'1l & Tha1assia, algae~crabs, gas­ 1967 mangrove sloughs, tropods, invertebrate eggs, rare on reefs bivalves (Randall 1967)

Calamus arctifrons + + 11, 17 Copepods, amphipods, mysids, Reid 1954 Associated with grass porg;,' shrimp, bivalves, gastropods grassy flats (Tabb & (Mitrella, Bittium), polychaetes Manning 1961)

~calamus + 1 190-250 mm: polychaetes, brittle Randall saucereye porgy stars, bivalves, hermit crabs, 1967 sea urchins, gastropods, chitons

120 Diet F"a:::Jily and S;Jecies Referenc.e Diet Referenc.e CollliDents

Lagodon rhomboides + + + 1-3,5,7,8, In :·'angro\'e <;;reek - scorc:ted 0<1= 1971 Strong preference pbfish n, 12, 16­ :nussel, mysids, a:n,hipods, Reid 1954 for vegetated sub­ 1. false 1:11.:ssel strate in bay areas In IOhitewater Bay - lao' plant (Weinstein et al. material 1977)

5ciae~idae - drums 3air1iella batabana + + 3, 11 I::::'.:e croaker

Bairdiella chrvsura + + ::'-1,8,11­ Larvae: copepods, larval fish OdW'J 1971 sllv!:!r perch 13, 16-18 (~ beryllina) 127-181 mm: fish (Anchoa mitchil1i) , mysids

Cvr.oscion arcnarius + 2, 12, 17, ~ostly fish, caridean shrinp, S?ringer & san::] seatrout 1. nysids, anphipods, crab zoea Woodburn 1960

Cynoscio~ nebulos~s + + + 1-3,5,7, <50 :nD: copepods, planktonic Odu:n 1971 5Fott~d seatroc!t 8,11-13, crllstacea :5, 1';', 18 50-275 :mu: fish (MugU cephalus, Lagodon rhomboidcs, ~ucino­ StomU5 quIa, E. arqenteus, Cyprinodon varieqatus, Gobicsoma robust=, Anchoa ::Iitchilli)

Leiost~ xa~th~ + 2, 7, 12, <40 mm: ?lanktonic organisms Springer & spot 17-18 >40 mm: filar:tentous algae, Woodburn desmids, forams, amphipods, 1960 ~ysids, copepods, ostracods, isopods, chaetoqnaths, bi­ valves, s~ils, polychaete '....OrT.\S

~0nticir=hus anerican~s + + 2, )1-12, rish, benthic crustaceans Springer I; so~ther~':;q~~-'- 17-15 I\oodburn 1960

Polychaetes, bivalves (Donax) , Springer & "'.ost COl:llllon off sandy L1enticirrhus litt~ 2, II Gulf kinqfish sand crab It:merital, razor clams Woodburn beaches

pog(lnia~ ~ + + + 2, 7, 11. <100 ~: molluscs, xanthid Darnell. black drum 12, l' crabs 1958 >100 mm: bivalves, amphipods, blue crabs, pcnaeid shrUnp, c-aridean shrimp

Sciaenoos ocellata 2, 3, S, 8, <10 ron: planktonic organisms Odum 1971 red drum 11-13, IS, (copepods, crab zoca, larval 17 fish) 34-42 nm: mysids, amphlpods, ca r idea n Shrimp >50 rnrn: xanthid & portunid crabs, pcnaeid shrimp, small fish 308-403 ~: xanthid crabs

E3u.ct'..Is ~cuminatus + 11 68-152 mrn: shrimp & shrinp Randall Characteristic of high-hat larvae, isopods, stomatopod 1967 coral reefs larvae, copepods, anphipods (Randall 1967)

121 Habitat T\·pc ! I ~ ~ ~ ."" :s~~~I;~ Diet ?ani 1y and Species !-o u:. I=.l OQ 0::0 Reference Diet Reference COIIIDents

Er-r.ipp~da~ - spadcfis~p.s chactod~~~crus faber + 2. 3. :;, Wonns. crustaceans, debris Jarnell Juveniles (7-12 mm) .A"t\-a--;:;-ili;pad\"fish-- 11, 16-13 1961 inhabit very shallow nearshore sandy beaches. Bear a deceptive resemblance to infertile red mangrove seed pods (Breder 1946) Characteristic family r-or..a::ef,cridae ­ of coral reefs (Ran­ dans~::i,,;,es dall 1967) Abudefd~f saxatilis 101-135 mm: COpepoc.s. algae, Rand;.ll ;;;rge;'.;-tllId~- fi~~ ~qqs. flSh. sr.rimp larvae. 1967 A habitat generalist: po1ych':ll"rc"i> reefs, grassbeds, rock piles, wharfs (Bohlke S Chaplin 1968) l."'lo::-lc.ae - wr:J,;7-153 mill: crabs. Si!'3 .lrchins, Randall Shallow water patch ;llpp<>ry dlC-k----- polyc~detes. gastropods, brlttle 1967 reefs, sand bottoms, stars. bivalves, shri~? fish, grassbeds (Randall !:erm~t crnba 1967)

Scal:"id"-i:! - p,-,xl:"ctfishes tlicnolsin,-, ust'lecondari1y ranging into grass­ on scaq(aSS~!:l beds

4

SEarisoma ct,rysopteru.!!:. + 11 redtail parrctfish Requires near marine SF,-,ris~ma rubriElnne salinities (Tabb & redfln ?drrot:ish :!'Iannin9 19611 searisclIla_ ~id(! 1J st~plight parrot~ish

~~gilidae - ~ullcts 1971 MUCll (;eE:tal:.;~ + + 2.3,5,7, a, Inorganic sedil"lents, fine Odum striped l:ullel: 11-13, 15 c:et::-~tus, micro-algae

, 25-73 ~, plant detritus, blue- r.ustin Muqil ~~ • 2, S, 7, • white mullet. 11-12 green algae \Lynchya majuscula) :.ustin 1971

Hugil trich<)dcl1 • • 2,7,11.12 ~il~

Salinities >10 0/00 sphyracnidae - barracudas + + 1-5, 7, 8, 135-369 mm: fish {Eucinostomus Odum 1971 (OQUIlI 1971) SFnyracna barracuda J 1, l3 quIa, Menidia ber}'llina, ~ great ba::-racuda sarsus probatoccEhalus} Fanily lives in epistognathida;) - ja""fishcs shri~p, Randall burrows in sediment. ~oistognathus maxillosus + 53-110 mm: isopods. 1967 often in vicinity ~ttled jawfish fishes, po1ychaetes, mysids, copepods of reefs (Randall 1967)

122 Habitat Ty..£..l:...... • .5 u E '"' ..-l .-l 0;1 <0 c ~~3>.:::>. Diet __F_a_m_'_'_y_a_nd_s_pe_c_'c_' c;:~:;:=cc:l~~~oooc~=__R_e_f_e_re.n:.:. _ Oiet Reference Co:vr.ents

Clinidae - cl1nids "amily ~pr-"~r'.; to =X;, car-n~ .....orous R3.ooa:'1 Inshcre on ~:Jck, Chae:lOpsis oc(!llata 0:1 tenthi.: irwec:::ebrate:-, 19E:: cor31 ur rubble bluethroat pi~cblcnny suostrat.cs (Rdn­ c.aL. :'~67:,

Paracli:lus ~rnoratus 1. ~. Ii JIlarbled J:>lli'n:l}'

Paracllnus fasciatcs banded J:>lli'nny

Stat~notus he.phill~ 1 b1ackbcl1y blfOnr;y

B1enniidae - co~too~h blennlcs Chasmodes saburrae l-L 11. 17. 21-2~ mm, an~h~~5 Carr .. Cr;rnt1On brilckish Florida bl;;ny--- ,. 2S-~~ mM: aD~h~~J~~, d~trltus. Adi!.ms ~at.er ~len:lY (~atb pol ych

Blennius marmor~u~ AI<::"c, o~':la:lic cf'L:-itllS, :<.and"'ll seaweed blc:lr.y br:c-;::fO star;;, pr~ly"ha02t0s. 19(,1 hyc.p):ds

Ble:miu!; nicholsi hiqhfin blenny

Callionymidae - drago~et5 ~ionym\l!; rallcirildi;;;tll,,_ 1. 5. 11 spotted dragonot

Elcotridae - sleepers Dormitator maculatu~ 1), 1. 5 Fresh.....a':cr a:1c. fat sl eepe'C ':'010' sa1:"nity area3 iD3rncll 19611 Gobiidae - gobie< ~~hygobi~ aor-orator 2, 3, B, 11, Car-idean shrimp, ::::,iron::.:mid ('dol:"! 1971 frillfin goby 17 larv

Gobionellus ;,astatus + 12 F'ilancntous algae l~nterc­ Sprlnger Ii. sharptai 1 qo~y IflOrphal. ostraC:Jds. :::opepod~, I-:oodburn i 1l';,",Ct. larval.' 19bJ

Gobionellus shufe1dci + 2. 17, !R fresh'",atcr goby

Gobionellus smaraJ::Ius + 3, a, 10, 11. emerald qoby 15

Gobiose>::l

Gobioscraa loncipala 17 twosca1e goby

GobiOSOl:la robustun 1-3. '>. ti, Amphipods, mysids, chironomid Odum 1971 code qoby 11.16-18 larvae

Lopho90bius cyprlno~des 1-3. 7, S, A vers~tilc feeder, am~hipods, O::l~ 1971 crested qoby 13 mangrove detritus. =ilanentous algae, "ysids, caridean & pcnaeid shrimp. polychaete ~~rms, ostracods, bivalves, chironomid larvae. harpacticoid copcpods. Lsopods, x3nthid crabs, snails

123 Diet. Family and Species Diet. Reference Comments

MicrogObius 9ulosus 2,5,8, lI­ Amphipods, copepods, chironomid odum 1971 clown goby 13.15, 17, larvae Ie

Microgobius microlepis 5 Planktonic organisms Birdsong banner goby 1981

Microgobius thalassinus 2, 3. 12 Small crustaceans including Peterson ... green goby amphipods, other invertebrates Peterson 1979

Scombridae - mackerels, tunas Sco~romorus maculat.us 2.11.n. Adults feeding on penaeid Tabb & Spanish mackerel 15 shrimp nigratinq from tidal "".anning stream 1961

ScomberOI:lOrU$ cavalla 11 3S0-1022 mm, fish Randall king mackerel 1%7

Scorpaenidae - scorpio~­ fishes Scoroaena bn'lsi liensis • • 7, 11 Shrim?, other crustaceans. Randall barbfish fish 1967

Scorpaena grandlcornis 37-102 1l1tI'l, shrL"p. fish, Randa 11 ~ost o~ten found plumed scorpionfish unidentifi~d crustaceans 1967 in seagr;1Ss

Triglidae - searobins Prionotus salmonicolor blackwing searcbin "

Prionotus scirulus + + + 1-3, 11, Small molluscs, shrimp, crabs Peterson & leopard searob!n 16-18 fish, small crustaceans I'cterson 1979 (ostracods, cumaceans)

Prionotus tribulus + + + 1-3, ll-lJ, Shrimp, crabs, fishes, a:nphi- Peter;.;on , bighead searobin 17, 1S pods, copepods, annelids, Peterson 1979 bivalves, sea urchtns

Rothidae - lefteye flounders Bathus ocellatU9 + 1, 11 68-130 1!lIIl: fish, crabs, shrimp, Randall 1967 eyed flounder amphipods

Citharichthys ~crops + 1 spotted whiff

Cit.harichrhys + + 1, 17, 1S Mainly mysids, also shri:np, Peterson , Recorded fron spilopterus crabs. copepods, amphipods, Peterson salinity ranRe. bay whiff fishes, annelids 1979 2.5-36.7 0/00 (Darnell 1961) Etropus crossotus + + 3, 11, 16 Calanoid copepods, cumaceans, Peterson • fringed flounder a.11Iphipods, mysids, shrimp, Peterson crabs, illOpods, annelids, 1979 molluscs, fisheR

Paralichthys albigutta + + + 1-3, 7, 11, <45 rem: s~a11 crustaceans, Springer (;. Gulf flounder 12, 11, 18 including amphipods, small Woodburn fish 1960; Reid >45 mm: fish (pigfish, pinftsh, 1954 l1zardfish, bay anchovy, labrids), crustaceans

Paralichthys lethostigma ­ + Mainly fishes (mullet, ~n~a­ ?cterson 6­ Southern flounder den, shad, anchovies, pinfish, Peterson mojarras, croakers), crabs, 1979 mysids, molluscs, penaeid shrimp, amphLpods

124 Habitat Type I~ ~m'l!'~ ~ ~ Diet Fa~ily and Species ! 51 Reference Diet Reference Comments _.. ~~---_.- --~~- - -~~------Sya~i~n pa~illos~n 1 Gusky flour_Ger

Soleidae - sales Acr-.:'..rJ.s lineatus + + 1-3,5,8, 32-74 ;urn: chironomid larvae, Odum 1971 ~ole 11-13, 17­ polychaete worms, foraminiferans 18

Trinec~€s inscript~ + 1 scr:;;.wled sale

'.rr~n0c:;es maculctu~ + 2, 3, 8, 14-110 om: amphipods, mysids Odum 1971 togchoker 11-13, 17, 16

Cynoglcsslda~ ~ tonqu0­ fish"'" ~Gph~rus plagius~ Dlackcheck tonguefish 1, 3, 11, 35-102 mID: polychaete worms, Austin & 12, 16-18 ostracods, portunid crabs, Austin RUF~ia and Ilalodule plant 1971 tips

Balistidae - trigs"'rfishes & filefishes Aluterus schoepfi + I, 11 Seaqr30sses, algae, hermit Randall Associated with ~-ril~:ish crabs, gastropods 1967 grassbeds, sponge/sea fan habitats (Ran­ dall 1967, Voss et 211. 1969)

Balistes vetula + 11 130-480 rnrn, sea urchins, crabs, Randall Solitary reef fish queen triggerfish bivalves, brittle stars, poly­ 1967 ranging into grass­ chaetes, hermit crabs, gastro­ bed, pods, algae

Monacanthus ciliatus + I, 11, 17 47-97 mm: Algae, organic detri­ Randall, Closely associated Wnged fi.le~- tus, seagrass, copepods, shrimp 1967 with vegetated areas ;:, shrimp larvae, amphipods, Springer I> (Tabb I> "lanning tanaids, polychaetes, molluscs woodburn 1961) 1960 ~onacanthus hispidus 1-3, 11, Detritus, bryozoans, annelids, Peterson I> Associated with planehead filefish 16-18 harpacticoid copepods, amphi­ Peterson vegetated areas (Tabb pods, hemit crabs, molluscs, 1979 & Manning 1961) algae, soa urchins

Balistes capriscus + gray trig~erfish

Ostraciidae - boxfishes Lactouhrys cuadracornis + + + 1,2, 5,7, Vegetation, algae, bivalves Reid 1954 Young mimic sea­ scrawled cowfish 11,16-18 grass blades (Bohlke & Chaplin 1968)

Lactophrys trigonus + 1, 4, 11 109-395 mm: crabs, bivalves, Randall Primarily a resident trunkfish polychaetes, sea urchins, algae, 1967 of seagrass (Randall seagrass, gastropods, amphipods 1967)

Lactophr~ trigueter + 1 93-250 mm, po1ychaetes, sipun­ Randall Primarily a reef smooth trunkfish culid worms, crabs, shrimp, 1967 species (Randall gastropods, hermit crabs, sea 1967) urchins, bivalves

Tetraodontidae - puffers Sphoeroides nephelus + + 1-3,5,11, Juveniles; detritus, fecal Carr I> sout.hern puffer 16-18 pellets, zooplankton, poly­ Adams chaetes, gastropods, crabs, 1973 shrimp Adults, small crabs, bivalves

125 Diet Family and Species Reference Diet Reference Comments

Sphoeroides spengleri + + I, 7, 11 Crabs, bivalves, snails, Randall Inhabits sea­ band ta U puff er polychactes, amphlpods, 1967 grass, reef, shrimp rubble, man­ ~rovcs (Randall 1967; Voss et a1. 1969)

Sphoeroides testudineus + + 1, 7 85-92 rom: portunid ncga10ps Austin & checkered puffer larvae, gastropods !'1ustin 1971

Diodontidae - porcupine­ fishes Chilomycterus antennatus + 11 Ga;ltropods, hemit crabs, Randall Reefs and grass­ bridled burrfish isopods, crabs, shrimp 1967 beds (Voss ct 81. 1969)

Ch110mycterus antillarum + 2 web burrfish

Chiloaycterus schoepfi + + 1-3, 5, Gastropods, barnacles, crabs, Springer <\ Associated with striped burrfish 11, 16-18 amphipods Woodburn grassbcds (voss 1960 et a1. 1969) Salinities >25 0/00 (Springer <\ Woodburn 1960)

Reference ~umbers Key

1. Bader & Roessler 1971 10. Seaman at al. 1973 2. Carter et a1. 1973 11. Schmidt 1979 3. Clark 1970 12. Springer &Woodburn 1960 4. Holm 1977 13. Tabb 1966 5. Hudson et a1. 1970 L4. Tabb, Dubro\ol & Hanning 1962 6. Kush1an & Lodge 1974 15. Tabb & ~ning 1961 7. Nugent 1970 16. Weinstein et a1. 1977 8. Odum 1971 17. Yokel 1975a 9. Rivas 1969 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 Abundant Insects, crustaceans, (Kinosternon subrubruml mollusks

Striped Mud Turtle Common Algae, snails, dead (Kinosternon bauri) fish

Ornate Diamondback Terrapin Uncommon Littorina, Melampus, ~, (Malaclemys terrapin Anomalocardia macrospilota and ~.~. rhizophorarum)

Florida Red-bellied Turtle Rare - Uncomroon Sagittaria, Lerona, Naias (Chrysemys nelsoni)

Chicken Turtle Unconunon Crayfish, insects, Nuphar (Deirochelys reticularia)

Green Turtle Uncommon Mangrove roots and leaves, (Chelonia mydas) seagrasses

Hawksbill Rare Rhizophora: fruits, leaves (Eretmochelys imbricata) wood, bark

Loggerhead Common Crabs, jellyfish, tuni­ (Caretta caretta) cates

Atlantic Ridley Uncommon Snails, crabs, clams CLepidochelys kempii)

Florida Softshell Cornman Snails, crayfish, mussels, (Trionyx ferox) frogs, fish, waterfowl

Green Anole Common Insects (Anolis carolinensis>

Cuban Brown Anole Common Insects (Anolis sagrei)

Bahaman Bank Anale Unconunon Insects CAnalis distichus)

Green Water Snake Carrunon Fish (Nerodia cyclopion)

Mangrove Water Snake Corranon Fish, invertebrates CNerodia fasciata compress1cauda)

128 M~HIBIANS AND REPTILES OF FLORIDA'S MANGROVES (concluded)

Species Status Food Habits

Striped Swamp Snake Uncommon Crayfish, sirens, frogs (Liodytes alieni)

Eastern Indigo Snake Uncommon Small mammals, birds, (Drymarchon corais) frogs

Rat Snake Uncommon Small mammals, birds (Elaphe obsoleta)

Eastern Cottonmouth Unconunon Fish, frogs, snakes, (Agkistrodon piscivorus) birds, small mammals

American Alligator Common Fish, waterbirds (Alligator mississippiensis)

American Crocodile Rare Fish, waterbirds (Crocodylus acutus)

Giant Toad Common Invertebrates (aufo marinus)

Squirrel Treefrog Abundant Insects (Hyla squirella)

Cuban Treefrog Corrrrnon Insects, frogs, toads, (Hyla septentrionalis) lizards

References: Carr and Gain 1955; Ernst & Barbour 1972;

Mahmuud 1965; L. Narcisse, R.N. IlDing" Darling Fed. Wildlife Refuge, Sanibell Is., Fla.; personal communication (1981).

129 APPENDIX D. Avifauna of south Florida mangrove swamps.

130 WADING BIRDS

Cnll'tlOn ;~.1"·_· ~;~a!;Ol\ .:>f ("J.lin 11.\1<''' ) Almndil .... cc {~currcncca ~1"~Lin'J a FOOtl Hilbit.5

-~~------

Great Egret Co~n v, V Fish Ho....el1 1932 (cas..nerodius albus) Kushlan I> White 1977a

Sno'''''Y Egret COllIllOn Y< Y f'ish Ho....e11 19]2 (Egretta Chula) Kush1an I> White 1977a Ffrcnch 1966

Cattle Egret Common Y< Y Fish Howell 1932 (Bubulcus lb ls) Kush1an & "''nite 1977a

Great White Heron Ra,. Y< V Fish Howell 1932 (Ardea herod1as Kushlan & "''bite 1977a occidentalisJ

Great Blue Heron Common Y< Y Fish Ho...'e1l 1932 (~ herodias) Kushlan & \·nti te 1977a

Reddish Egret U:lcOtfl:nOn Yr V ?ish Howell 1932 (Dichrolllanassa Kushlan &'..lhite 1977a rufes::ens)

Louisiana Heeon Common y, Y Fish Kushlan & \1hite 1977a (Hydranassa tricolor) I'\axwell .. Kale 1977 Girard & Taylor 1919

Little Blue Heron Com:non Y< V Fish Kushlan I> White 1911a (Florida caerulea) />taxwell ,<; Kale 1977 Girard & Taylor 1919

Green Heron Co:nmon Y< Y Fish Robertson & Kush1an 1974 (Butorides strlntus) jl\ax....·ell , Kale 1977 Girard ,<; Taylor 1979

Black-cro...med Night Co:r:unon y, Y Fish, crustaceans, pfrench 1966 Heron frogs, mice .:1ax ....'e11 I> Kdle 1917 (Nycticorax Girard & Taylor 1979 nvcticoraxl

Yello""-crowned Night COllClOn V, V Fish, crayfish. pfrench 1366 Heron crabs Girarc I> Tdylor 1979 (Nyctanassa violaceal

Lea,t Bittern Uncom.~n Y< V ('ish Pfrc:lch 1966 (Ixobrychus exilis)

American 3ittern Cneol'lV'"..on \1,1' V Crayfisr.. froqs. (Botauru.s sll'all fishes lentiginosus)

W~d Stork CO:nr:Jon Y Fish Kahl 1964 ntycteria americanal ( locally " Ogden et al. 1976 abundant) Kush1an 1919

Glossy Ibis Uncommon FiSh Bacon 1910 (Plegadis falci- " Howell 1932 nellus

\1hi ':.e Ibis Abundant y, V F'ish, erabs (~l Kushlan 1,}79 (Eudocimus albus) Kushlan & Kushlan 1915 Giraed £ Tayloe 1979

Roseate Spoonbill Rare to Y< V Shring. fish, Kush1an & White 1971a (Ajaia ajaja) UnCOllTllon aqul:I.tic vegetation lIo....ell 1932

Sandhill crane Rare Vr 'Roots, rhizo::nes of Ogcen 1969 (Grus canadensis) Cyoerus &- Sa9it- Ho....ell 1932 caria

Lu?kin Rare Y< V Snails {~l H

131 PROBING SHORE BIRDS

COllllllOO Name Season of a (Latin name) Abundance Occurrence8 Nesting Food Habits References

King Rail Yr Beetles, grass­ Narcise. pets. comm. (Rallus eigans) hoppers. aquatic Martin et a1. 1951 bugs

Clapper Rail t:ncolllI!lOn­ y, Y Crabs, shricp Howell 1932 (Rallus 100g1ro­ COllmon Ffrench 1966 stds) Racon 1970

Virginia Rail Rue Beetles, snails, Marclsse, pers. C~. (Rallus limicola) spiders Martin et al. 1951

SOTa UnCOtmlOn to Insects, seeds of Howell 1932 (Porzana carolina) locally emergent aquatic Bacon 1970 abundant plants

Black Rail Rare Beetles, snails Narcisse, pets. co~m. (LateralIus jaoaicensis)

Semlpalmated Plover Locally Crus taceans, Ffrench 1966 (Charadrius semi­ connon mollusks Bacon 1970 palmatus) -- Baker 6. Baker 1973

"'ilson I s Plover Locally Crabs, shrimp, Howell 1932 (Charadrius wilsonia) common crayfish Bacon 1970

Black-bellied Plover Common Crabs, mollusks Howell 1932 (Pluvialis Bacon 1970 sguatarola) FEre-nch 1966

Ruddy Turnstone Common Insects, crus­ O~den 1969 CArcoaTis interpres) taceans, mollusks Ho\~ell 1932

Common SnLpe Uncommon W,T Molillsks, insects, HOI,'ell 1932 (.f2pella gallinago) worms Bacon 1970

Long-billed CurIel,' Rare-uncomnon Crustaceans, Ogden 1969 (Numenius americanus) insects Io.'himbrel Uncommon l~ollusks, crus­ (Numeniu8 phaeopus) Ogden 1968 taceans, wor:ns, Howell 1932 insects

Spotted Sandpiper Abundant. W,1' Mollusks, crus­ (Actitis macularia) Ffrench 1966 taceans Bacon 1970 Russel 1980 Solitary Sandpiper COllIIl.on W,T Crustaceans, aquattc Howell 1932 (Tringa solitada) insects, small frogs Bacon 1970 Willet COllDOn Yr Crabs, crayfishes, Howell 1932 (Catoptrophorus killifishes seIJl1palJlat.us) Bacon 1970

Greater Yellowlegs CO!:lIllOD W,T Fishes, crabs, Howell 1932 (Tringa crustaceans Ffrench 1966 I:I.elanoleucas) Bacon 1970 Lesser Yell~'legs Common W,T Snails, mollusks, ~'french 1966 (Tringa flavipes) crahs Bacon 1970 Baker & Baker 1973 Red Knot. Uncomnon W, r :-t.:lrine ...!orms, Howell 1932 (Calidris ~) crustaceans Ogden 1964

Dunlin CO!lllllOn W Narinc worms, (Calidris alpjna) Ogden 196t. mollusks 8aker 6- Baker 1973 t"Thit:e-rumped Sandpiper Rare T Chironomids, snails (Calldris fU8cicollis) Howell 1932 Bacon 1970

132 PROBING SHOREBIRDS (concludedj

Comoon Name Season of (Latin name) Abundance Occurrence8 Nesting8 Food Habits References

Least Sandpiper Conunon "101, T Pupae of beetles Bacon 1970 (Calidris rninutilla) and flies Baker & Baker 1973

Short-billed Dowitcher COflIllon W,T Mollusks. Bacon 1970 (Limnodromus griseus) crustaceans Baker & Baker 1973

Stilt Sandpip.er Rare-uncommon W,T Chironomids Howell 1932 (Micropalama Bacon 1970 himantopus)

Semipalmated Sandpiper Common- \-/, T :iollusks. insects Bacon 1970 (Calidris pusil~) abundant Baker & Baker 1973

Western Sandpiper Common- H,T Chironomids Howell 1932 (Cal1dris mauri) abundant Bacon 1970

Harbled Godwit Rare-connnon W Crustaceans, Hm,'ell 1932 (Limosa fedoa) mollusks. seeds of emergent aquatic plants

American Avocet Uncommon "",t Marine worms, Ogden 1969 (Recurvirostra aquatic insects americana)

Black-necked Stilt Common S Aquatic beetles Howell 1932 (Himantopus mexicanus) Bacon 1970

133 SURFACE AND DIVING BIRDS

COlillnon Name 5eafiOil of (Latin !lamp) f.bundancc Oc:::uT::"cncea tlcsting a Food ;!abits References

COllTl\on Loon occasiona 1 Fish, crabs, rr~llusks Narcisse, pers. co:nm. (Cavia immer) "

Horned Gre1>e UnCOll"fllon Fish, aq~atic insects, c~den 1969 (Podiceps aurltus) " mollusks

pied-billed ~rebe UncannQn- ':'r Crayfish, fish, Narcisse, pers. COIJ[ll. (Podilynbus CO!mlon ~ollusks podiceps)

White Pelican Ra'" 5 Fish Nar::;isse, pers. COIml. (Peleca!lU5 COllUlcn erythrorhynchos) "

Bral'm Pelican Corrmon Fish Ffrench 1966 (?elecanus " Bacon 1970 occidentalis)

Double-crested Cor.JIlO:l V, y Fish Kush!an & ~hite 1977a cormorant (Phalacrocorax auritus)

Anhin<;ld CQrJllO:'l. y, Fish F:renc:: 1966 ("nhings anhinga)

Fulvous ~.lhist:lLng Duck Uncanmon Ogeen 10369 (Dendrocygr.a " Smith, pers. o~s. biealor)

:-laUard UJ'lc=on '.~, 'I' Wi.dgeon grass ::Jgde:l.. 1969 (,\oas platyrhynchos) Kus:-:lan et al., in prep.

Black Duck \':,T Moll~sks, cr~stD­ C·gden 1969 (~ rubeipes) ceans, ~idgcon grass

,,;ott.led Duck Unconmon y, v ?olvqonum, snai~s, LaHunt & Cornwell 1970 (Anas Culvigula) Rup'O:'a Kushlan et al., in prep.

Gaawall l:ncOIIOOn '.~,T ;<.uE'Oia, ~, Ogeen 1969 (~ streoera) n.ollusks

Pintail Abundant ~'1 ,T Saggitaria, mollusks, Narcisse, pers. co~. (~ acuta) cype~ K·.... shlan et al., in prep.

Green-winged Teal Abundant \I',T Ruppia, Zoster;).. ~arcisSQ, pers. cumru. (A:1as ~ carolinensis) aquatic insects Kushlan ct al., in prep.

Blue-winged Toal rJ:JUndant V, ~, snails, Karcisse, pers. comm. (~ discorsl insects, crustacoans Ffrenc:t 1966

American Wigeon COInIJlon W,T Ruppia, ~, Narcisse. pers. comm. l~ a.'l\ericana) mollusks Kushlan et al.• in peep.

Korthern Shoveler COmll'lOn .... ,:: mollusks, aq~atic Narcisso, pers. cocm . l~ dvpeata) insects, Rupnia, ~

Wood Duck Race ~uts, seeds Ogden 1969 (~ sponsal "

Redhead Race Snails, clams, a~uatic Ogden 1969 (Aythya americana) " insects, RUPEia, Zos- tera

Ring-necked Duck Abundant Polygonum, Reppia, Ogaen 1969 (Aythya collarisJ " crayfish, snails Kushlan et al., io prep.

canvasback Uncommon w Vallisneria, Ruppia, Ogden 1969 (Aythya valisineriaJ ~ Kushlan et al., io prep.

134 SURFACE AND DIVING BIRDS (concluded)

Common Name Season of (Latin name) Abundance Occurrence a Nesting a Food Habits References

Lesser Scaup COI:IIDQn- W Mollusks, Ruppia Narcisse, pers. coorn. (Aythya aHiuis) abundant. Ogden 1969 Kushlan et al., in prep.

Bufflehead Rare W Gastropods, crabs. Ogden 1969 (Bucephala alheala) crustaceans Kushlan et a!.. in prep.

Ruddy Duck Common W Pot.anogeton. ~ajas. Ogden 1969 (Oxyura jamaicensis) Zostera, Ruppia, Kushlan et al.• in prep. nollusks t100ded Merganser Rare-unconoon W Fish Ogden 1969 (Lophodytes cucullatus)

Red-breasted Merganser Camocn !\I.T Fish Narcisse. pers. carom. (~ergus serrstor)

Purple Gallinule Rare Yr V ,'\quatic insects, Narcisse. pers. comm. (Porphyrula mollusks. Ffrench 1966 martinica) Eleocharis. Paspalum

Common Gallinule Common Yr y Seeds, aquatic Narcisse, pcrs. comn.. (Gallinula chloropus) insects fo'french 1966

American Coot Abundant w,r Ruppia, Najas. Narcisse, pers. camm. (Fuliea americana) Potamogeton, aquatic insects

135 AERIALLY SEAHCHING

Comrr,on Na:me Season of a (Latin name) Abundance Occurr.ence Nest:.inif" Food Habits References

Herring Gull Uncommon W Fish, mollusks, Narcisse, pers. corom. (~ arqento.tus) crustaceans Ogden 1969

Ring-billed Gull Common ..i,T Fish, insects, Narcisse, pers. corom. (Larus delawarensis) mollusks Ogden 1969

Laughing Gull Corrnnon Yr Fish, shrimp, crabs N:arcisse, pers. c_. (Larus atricilla) Ogden 1969

Bonaparte's Gull Uncorrunon Fish, insects Ogden 1969 (Larus philadelphia) "

Gull-billed Tern Uncommon Yr l>layflies I dragonflies Ogden 1969 (Gelochelidon nilotica)

Forster's Tern Uncorrunon- W Fish Narcisse, pers. carom. (Sterna fosteri) COUlmon Ogden 1969

Common Tern Uncorrunon Fish Ogden 1969 (Sterna hirundol '"

Least Tern Common S Fish Narcisse, [-ers. CO!1UTl. (Sterna albifrons) Ogden 1969

Royal Tern Cornman -.-I,T Fish Ogden 1969 (Thalasseus maxima)

Sand·....ich Tern Uncommon Yr Fish Narcisse, pers. comm. (Sterna sand- Ogden 1969 vicensisl

Caspian Tern Unconunon " Fish Ogden 1969 (?terna caspi;~)

Black SkirnmelC Comro.on Yr Fish Ogden 1969 (Rynchops niryra)

Belted :

Fish Crow Common Yr Y Fish Narcisse, pers_ corom. (Corvus ossifragus)

136 BIRDS OF PREY

::,;,,! ·"11 :) r Occ:urr

TUYkey vul tun~ Common y Carrion Narcisse, pers. corom. (Cathartes aura) Orians 1969

Black Vultun, Corrunon Carrion Robertson & Kushlan (Coragyp~ atratus) 1974 Orians 1969

S'i.'allow-tailc:d Ki tc Conmon y Snakes, lizards, Howell 1932 (Elanoides :orfica­ frogs Snydc.r 1974 t.us)

Sharp-shinned Hawk Unco:nmon Smaller passerines Howell 1932 (~cipiter striatusl

Cooper's Hawk Uncommon y l~rger passerines Howell 1932 (M:Tipi~~ ~~r:ii)

Red-tailed ;'d\·..k Uncattunon Small mammals, birds I1m"ell 1932 (Buteo jarnaic€rlsis)

Red-shouldered l!

Broad-winged 1M"'': lIr:comnO[l Insects, small Ho ..... ell 1932 (Buteo platypt;,.erus) maT'1Il\als

SWflinson's Hawk Il.are Small manmals, grass- Howell 1932 (Buteo swai:lsoni) hoppers

Short-tailed Hawk UnCCITlTlon ':1 SmaIl birds 1I0·...ell 1932 (~tea £=ach~..::.'::.:'~}

Billd Ed'jle Rare-locally y Fishes Ilo'.".'e 11 1932 (Haliaeetus common [Fla. !..,ucocephalus) 3ay)

Mill':"sh Ilawk Unco:nmon Small mammals, shore- Rowell 1932 (Circus £..Zar.eusl birds

Os?rpy ::onmon y, Fishes Howell 1932 (!:.:"J.ndi0r:. :Jaliaetus)

Peregrine Falcon Very rilre­ Waterfowl, shorebirds Nisbet 1968 (!::3.1co -:Jeregr~r.~ 100a II.,. comnon Ogden 1969 {Fla. Bay] Howell 1932 l·le-r1in Uncommon Small birds, share- Howell 1932 ("'a1~ co1umbari;.ts) birds

AmerIcan Kestrel Common w Tnsects Howell 1932 (~l:£<2 ~arverius)

Barn Owl Cncor:unon y Sma 11 n1i'1lml\als Howell 1932 (~1~.£ alba]

Great lIorned 0·,011 Uncomnon y ~;aterfowl , small Howell 1932 (Bu~ virainianus) mammals

3arred Owl lI:lCOIT\ll\On y, y Small mammals, frogs, Howell 1932 (Strix varia) snakes

137 ARBOREAL BIRDS

Common Name Season of (Latin name) Abundance Occurrences Nestin~ Food Habits References f-lourning Dove Uncommon Y Y Seeds Emle-n 1977 (Zenaidura macroura)

\Vhite-crowned Pigeon UncomlllOu Yr Y Berries. seeds, Howell 1932 (Columba fruits Robertson & Kushlan 1974 ~pha1a)

Mangrove Cuckoo Uncommon Yr Y Ca terpillars. Howell 1932 (COCCy;z:us minor) mantids Ffrench 1966 Robertson & Kush1an 1974 Martin et al. 1951

Yellow-billed Cuckoo Common s y Caterpillars, Howell 1932 (Coccyzus arnericanus) beetles Ffrench 1966 Martin et al. 1951

Smooth-billed Ani Y, Y Insects HO\o,'ell 1932 (Crotophaga ani) Ffrench 1966

Chuck~will's-~idow Uncommon y, Y Mosquitos, moths Martin et al. 1951 (Caprimulgus Karcisse, pers. comm. carolinensis)

Common Flicker Uncommon Y, Y Ants, beetles, Karcisse, pers. corum. (Colaptes auratus) fruits in winter Martin et al. 1951

Pileated Woodpecker Unconunon Yr y Beetles. berries. Howell 1932 (DryocopuS pileatus) fruits Robertson 1955 Robertson & Kushlan 1974

Red-bellied Woodpecker Common y, y Beetles, ants, Narcisse, pers. comm. (Helane~ carolinus) grasshoppers, ~artin et al. 1951 crickets

Red-headed Woodpecker Rare Y, Y Beetles, ants, Narcis5e, pers. comm. (t-lelanerpes grasshoppers, ~lartin et a1. 1951 ervthrocephalus) caterpillars

Yellow-hellied Uncorrnnon ',;I.T Beetles, Llnts-, ",arcissc, pers. corum. Sapsucker caterpillars Martin et al. 1951 (Sphyrapicus varius)

Hairy Woodpecker Rare p Insects, beetle I:mlen 1977 (Picoides larvae ~)

Eastern Kingbird Uncommon S,T y Ants, WLISPS, Narcisse. pers. comm. (Iyrannus tyrannus) grasshoppers Martin et al. 1951

Gray Kingbird Common S,T y Bees, wasps, HO\,'ell 1932 (Tyrannus beetles, dragon Robertson ~ Kush1an 1974 dominicensis)

Western Kingbird Rare W,T Bees, 'Wasps. Narcisse, pers. comm. (Tyrannus verticalus) grasshoppers Martin 8t al. 1951

Great Crested Unconn:Jon Yr Y Insects, berries Howell 1932 Flycatcher (coTlUnon S) Robertson 1955 (Myiarchus crinitus)

Acadian Flycatcher Rare T Small flying insects Morton 1980 (Empidonax virescens)

Eastern Phoebe Common Bees, wasps, ants Narcisse, pers. cornm. (Sayomis phoebe) ~artin et al. 1951

Eastern Wood Pewee Rare-uncommon s,r Bees, wasps, ants, Narcisse, pers. comm. (Contopus virens) moths 1l00,'ell 1932

138 ARBOREAL BIRDS (continued)

Cormon N

Barn Swallow Locally cOllUllOn lnsects Howell 1932 (Hirundo rusHea) Bacon 1970

Blue Jay Uncommon Yr Grasshoppers, cater­ Narcisse, pers_ comm. (cyanocitta cristata) pillars, beetles Hartin et a1. 1951

Tufted titmouse Very rare­ w Caterpillars, wasps, Howell 1932 (Parus bicolor) rare bees Robertson &0 Kushlan 1974

Carolina Wren Uncommon Yr Y Ants, flies, milli­ !>'arcisse, pers. COOlIn.· (i'hryothorus peds r~rtin et al. 1951 ludovicianusl

"'.ock ingbird Abundant Yr Y Fruits, berries Robertson 1955 (t-lil!'lus polyglottos)

Catbird COllllllOn \oI,T Fruits, insects Narcisse, pers. comm. (Dumetella ~­ Martin et a1. 1951. linensisl

Brown Thrasher Uncoll'lllOn Yr Y Beetles Narcisse, pers. comm. (Toxost:.oma rufulll) ~tartin et al. 1951

An::erican Robin Abundant Worms, berries, Narcisse, pers. corom. (Turdus migratoriusJ insects Martin et al.. 1951

~lue-gray Gnatcatcher Uncollll':'tOn W,T Insects, especially Narcisse, pers. comm.. (Polioptila eaerulea) Hymenopterans Howell 1932

Ruby-crowned Kinglet Uncommon W,T Wasps, ants ~arcisse, pers. carnm. (Regulus calendula) Howell 1932

White-eyed Vireo Unconunon S,T Y Butterflies, moths Robertson 1955 l~ griseusl

Black-~hiskcred Vireo Uncommon Yr Y Spiders, caterpillars lIowell 1932 (Vireo altiloquus) Robertson & Kushlan. 1974

Red-eyed Vireo Uncommon S,T Y Caterpillars, beetles Narcisse, pers. comm. (~ 01 i vaceus I Howell 1932

Yellow-throated Vireo uncommon W Butterflies, moths, ~k>rton 1980 (Vireo flavj£rons)

Black-and4"'hite Fairly W,T Wood boring insects Lack and Lack 1972 Warbler COITIon Keast 1990 (Mniotilta varia) Ogden 1969

"~orm-eatin9 "~arbler Uncommon w caterpillars, spiders Ogden. 1969 (lIellllitheros vermi- Kush1an, pers. comm. c ~)

Prothonotary Warbler Unco~n T Insects Ffrench 1966 {Protonotaria citrea) Russel. 1980

Yellow-throated Common Beetles, moths, Morton 1980 \~arbler spiders (Dendroica dominica)

Yellow Warbler conmon yr y Insects Haverschmidt 1965 (Dendroica petechial Ffrench 1966 Orians ~969 Terborgh & Faaborg 1980 Yellow-:etlll'\ped Warbler Abundant W,T Dipterans, bayberries Narcisse, pers. comm. (Dendroica coronata)

Prairie Warbler Uncommon Yr MOths, beetles, flies Lack & Lack 1972 (Dendroica discolor) Robertson & Kush1an 1974

Palm Warbler A~ndant W,T Insects Lack & Lack 1972 (Dendroica palmarurol Etll1en. 1977 139 ARBOREAL BIRDS (continued)

ConU7101l N"nHC! Season o[ (T~"ll:iJl :lilITH,,) A)"JIlC];lnce Occnrrctlco;! a Nc:::till(j a References

Blackpoll Warbler Uncommon T Insects Ffrench 1966 (Dendroica striata)

Bay-breasted Warbler Rare T Insects -"1orton 1980 (Dendroica castanea)

Black-throated Gree~ Uncommon Aphids, leaf-rollers, Ogden 1969 \'larbler and other insects Kush1an, pers. comm. (Dendroica virens)

Chestnut-sided Warbler Rare T Insects Morton 1980 (Dendroica Densyl- v

Cape May Warbler Uncorunon H Ogden 1969 (Dendroica tigrinaJ Common T

Black-throated Gray Rare Insects Ogden 1969 \~arbler Kush1an, pers. conrn. (Dendroica nigrescens) Hutto 1980

Black-throated Blue Uncommon Beetles, flies, ants Kush1an, pers. corum. \"Iarbler Common Ogden 1969 (Cendroica caeru- lescens)

:-Jorthern Waterti'.rush 1'.bundant Insects Schwartz 1964 (Seiurus novebol::a­ Rare Ffrench 1966 censis) Racon 1970 Russell 1980

Ye11owthroat Common Yr y Grasshoppers, ~rickets, Narcisse, pers. comm. (Geothlypus trichas) ants, wasps HowelJ 1932 Lack & Lack 1972

American Redstart Common T Caterpillars Bennett 1980 (Setophaaa rutici1la) Ffrench 1966 Bacon 1970

Tennessee Warbler Unco~n T Insects :-tortan 1980 (Vermivora "::lereqrina)

Nasheville 'warbler Rare T Insects Hutto 1980 (Ve:rmivora rufi- capilla) --

Oranqe-cr~~ned Warbler common Insects Hutto 1980 (vermivora celata)

Golden-winged Warbler Rare T Insects ~\orton 1980 (Vermivara chrysop- tera)

Northern Parula Common Hymenoptera Lack and Lack 1972 (Farula americana)

Ovenbird Common Beetles, crickets, Lack and Lack 1972 (Seiurus aurocapil­ grasshoppers Ius)

Kentucky Warbler Rare-uncommon T Beetles, caterpillars, Morton 1980 (Oporornis formosus) ants

Mourning Warbler Rare T Insects Morton 1980 (Ooorornis philadel- phia)

Yellow-breasted Chat Common Hymenoptera Hutt0. 1980 (~virens)

140 ARBOREAL BIRDS (concluded)

C..,nWllOIl 1I,1m... Season or ('.,It in 11,111:(') OCelll"I"r-ncea Food I I"..., i,U:

fJlJttu I.']HO \~ilson' 5 \'i<)([,ler R,lce-unCOIIU1l0n '1' Tnsects 1~11lQ<; !

Red-..inqed 1l1ackbird Comraon Yr Seeds, insects nowell 1932 {A9alaiu~ phoeniccusl Hohcrlson 1955

Bo~t-tailed GracxLe ~ncommon y Crayfish, cr:-abs, Robcrtsofl 1955 (Qutscalus major) shr hop G~rard • Taylor 1979

common Grackle UncommOn y Insects, cater-- Howell 1':.1)2 IQuiscalu!'> 9.ulscula.) ?iilacs I

Card i na.i Common y In!iccts, sced~ Roherts')ll 1955 (CardinaUs c

Orchard Oriole Rare T Grnsshoppers. beetles ~brt.Oll 1900 (Icterus spurius)

Indigo Bunting Uncol"JlToOn \".'1' Grasshoppers, cater­ Narcisse. pers. COTlt'll. IPasJ'>erina cyanea) pillal"s Howp.ll 1~32

SUIlVl(lI" Tanager Uncommon T IlYl:lenoptera Horton 1990 (P~.ran9a rubral

Dlckci:-:sel Uncomr.\On Caterpillars. beetles Bacon 1970 (Spiza al"erici'lna) Hartin ct al. 1951

Rufous-sided 1uwhee Common Yr y Caterpillars. b~y­ Narcisse, pers. COr.'ill. (PiFilo erythroph­ barries, fruit!> Howell j 932 ~)

S....amp Sparrow Common 'fI.T Narcisl:le. pers. COntIno (Melospiz~ 9corqiana) Howell 1932

3.yr .. year round resident 5 '" summer resident W• winter resident T• transient, present only during spring and fall migration Y = species breeds in mangroves b L. ~a[cisse, R.N. "Ding" Darling Fed. Wildlife Refuge, Sanibel Island, Fla. (1981). cJ •A• Kushlan, So. Fla. Res. Ctr., Everglades Katl. Park, Homestead, Fla.

141 APPENDIX E. Mammals of south Florida mangrove swamps.

142 MAMMALS OF FLORIDA MANGROVES

Species Status Food Habits

Virginia Opossum Abundant Fruits, berries, insects, (DidelphlS virginiana) frogs, snakes, small birds and mammals

Short-tailed Shrew Uncommon Insects (Blarina brevicauda)

Marsh Rabbit Abundant Emergent aquatics (Sylvilagus ?alustris)

Gray Squirrel Occasional Fruits, berries, mast, (Sciurus carolinensis) seeds

Fox Squirrel Rare Fruits, berries, mast (Sciurus niger)

Marsh Rice Rat Uncommon Seeds of emergent plants, (Oryzomys palustris) insects, crabs

Cudjoe Key Rice Rat Rare Seeds, insects, crabs (Oryzomys argentatusl

Cotton Rat Abundant Sedges, grasses, cray­ (Sigmodon hispidus) fish, crabs, insects

Gray Fox Uncommon Small mammals, birds (Urocyon cinereoargenteus)

Black Bear Rare Fruits, berries, fish, (Ursus americanus) mice

Raccoon Abundant Crayfish, frogs, fish (Procyon lotor)

Mink Rare Small mammals, fish, (Mustela vison) frogs, snakes, aquatic insects

Striped Skunk Common Bird eggs and young (Mephitis mephitis) frogs, mice, larger invertebrates

River Otter Uncommon Crayfish, fish, mussels u..utra canadensis)

143 ~~LS or rLORIDA ~~NGROVES (concluded)

Species Status Food Habits

Panther Very rare Deer, rabbits, mice, (Felis cancolor) birds

Bobcat Common Rabbits, squirrels, (relis rufus) birds

White-tailed Deer Common Emergent aquatics, nuts, (Odocoileus virginianus) acorns, occasionally mangrove leaves

Key Deer Common on cer­ Emergent aquatics and (Q.. ~. clavium) tain Florida Keys other vegetation (no longer on mainland)

Black Rat Common (Rattus rattus)

Bottle-nosed Dolphin Uncommon Fish (Tursiops truncatus)

West Indian Manatee Uncommon Submerged aquatics, (Trichechus manatus) Zostera, Ruppia, Halodule, Syringodium, Cyrnodocea, Thalassia

References: Layne 1974; Hamilton and Whittaker 1979; L. Narcisse, R.N. "DingII Darling Fed. Wildlife Refuge, Sanibel Island, Fla.; personal commu­ nication.

144 .50271-101 REPORT DOCUMENTATION ( .... REPORT NO. J. RKipient's Accession No. PAGE FWS/OBS-81/24 T' ... Title lind Subtitle 50 Rel'Ort Olte THE ECOLOGY OF THE MANGROVES OF SOUTH FLORIOA: A COMMUNITY January 1982 * PROFILE 6. - 7. Author(s) 8. Performing OrllJniution Rept. No. Will iam E. Odum, Carole C. McIvor, Thomas J. Smi th, III 9. Address of Authors 10. ProjectfTask/Work Unit No.

Department of Environmental Sciences 11. Contrael(C) or Grant(Gl No. University of Virginia 'C) Charlottesville, Virginia 22901 (0)

12. Sl'Onsoring Orllilniution Nllme lind Address 13. Type of RepOrt & Period Covered Office of Bio109ica1 Services New Orleans OCS Office Fish and Wildlife Service Bureau of Land Management U.S. Department of the Interior U.S. Department of the Interior Washington, D.C. 20240 New Orleans, Louisiana 70130 ". - 15. Supprementllry Notes *Reprinted September 19B5 --- ·16. Ab'stAct ~Limit: h wo,ds) etai ed 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 di seases to aspects of reproduction, bi omass partitioning, and adaptations to stress. Mangrove ecosystems are considered in terms of zonation, successi on, 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, i nvertebra tes, fishes, repti 1es, amphibians, birds, and mamma 1s. Finally, bm sections 5urrmarize 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 hal in Florida, are a resource of great value and should be protected and preserved wherever possible.

17. Document Analysi. II. Descriptors

Eco logy, value, management, fauna

b. Identifiers/Open.Ended Terms

Mangroves, halophytes, coasta1 wetlands, ecosystem, South Florida

c. COSATI field/Group

IS. AVllilllbility Stiltemerrt 19. Security Class (This Rel'Ort) 21. No. of Pages Unlimited "nr'---'~'-" 154 20. Seoeurity Class (This Pille) 22. Price

(See ANSI-239.18) OPT,ONAL fORM 212 (4-11) (Fo'merly NTIS-JS) Oepllrtment of Commen:e DEPARTMENT OF THE INTERIOR u.s. FISH AND WILDLIFE SERVICE

As the Nation's principal conservation agency. the Department of the Interior has respon­ sibility for most of our .nationally owned public lands and natural resources. This includes fostering the wisest use of our land and water resources, protecting our fish and wildlife, preserving 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 territones under . u.s. administration.