FWS/OBS-81/24January 1982

THE ECOLOGY OF THEMANGROVES OF SOUTH FLORIDA: ACOMMUNITY PROFILE

Bureau of Land ManagementFish and Wildlife Service

U.S. Department of the Interior

• • --

FWS/OBS-81/24January 1982

Reprinted September 1985

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

by

William E. OdumCarole C. t·1cIvor

Thomas J. Smith, III

Department of Environmental SciencesUniversity of Virginia

Charlottesville, Virginia 22901

Project Officer

Ken AdamsNational Coastal Ecosystems Team

U.S. Fish and Wildlife Service1010 Gause Boulevard

Slidell, Louisiana 70458

Performed forNational Coastal Ecosystems Team

Office of Biological ServicesFish and Wildlife Service

U.S. Department of the InteriorWashington, D.C. 20240

and

New Orleans OCS OfficeBureau of Land Management

U.S. Department of the InteriorNew Orleans, Louisiana 70130

OISCLAmER

The findings in this report are not to be construed as an officialU.S. Fish and Wildlife Service position unless so designated by otherauthorized 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 ecologyof the mangroves of south Florida: a conmunity profile. U.S. Fishand Wildlife Service, Office of Biological Services, Washington.D. C. FWS/OBS-81/24. 144 pp. Reprinted September 1985.

PREFACE

This profil e of the mangrove commun­ity of south Florida is one in a seriesof community profil es whi ch trea t coas ta 1and marine habitats important to man. Theobvious work that mangrove communities dofor man includes the stabilization andprotection of shorelines; the creation andmaintenance of habitat for a oreat numberof animals, many of which -are eitherenda ngered or ha ve commerci alva 1ue; andthe provis ion of the bas is of a food webwhose final products include a seafoodsmorgasbord of oys ters, crabs, lobs ters,shrimp, and fish. Less tangible butequa lly important benefi ts incl ude wil der­ness, aes thet ic and 1ife support cons i der­ations.

The information on these pages cangive a basic understanding of the mangrovecommunity and its role in the regionalecosystem of south Florida. The primarygeographic area covered lies along thecoas t between Cape Canavera1 on the eas t

iii

and Tarpon Springs on the west. Refer­ences are provi ded for those seek i ngin-depth treatment of a specific facet ofmangrove ecology. The format, style, andlevel of presentation make this synthes isreport adaptable to a diversity of needssuch as the preparation of environmentalass ess IT.ent reports, s upp 1ementa ry readi ngin marine science courses, and the devel­opment of a sense of the importance ofthis resource to those citizens whocontrol its fate.

Any ques ti ons or comments about orrequests for this publication should bedirected to:

Information Transfer SpecialistNational Coastal Ecosystems TeamU.S. Fish and Wildlife ServiceNASA-Slidell Computer Complex1010 Gause BoulevardSlidell, Louisiana 70458

iv

CONTENTS

PREFACE ..........•........................................................... iiiFIGURES viiiTABLES vi i iACKNOWLEDGMENTS i x

CHAPTER 1. INTRODUCTION .

1.1 "Mangrove" Definition.............................................. 11.2 Factors Controll ing Mangrove Distribution.......................... 11.3 Geographical Distribution 21.4 Mangrove Species Descriptions .. , 51.5 Mangrove Communi ty Types 71. 6 Substrates 91 .7 Wa ter Quality .............................................•...•.... 11

CHAPTER 2. AUTECOLOGY OF MANGROVES 12

2.1 Adaptations to Natural Stress - Anaerobic Sediments................ 122.2 Adaptations to Natural Stress - Salinity.. 122.3 Reproductive Strategies............................................ 142.4 Biomass Partitioning............................................... 152.5 Primary Production................................................. 172.6 Herbivory............. 232.7 Wood Borers ...........................•...................•........ 242.8 Mangrove Diseases 25

CHAPTER 3. ECOSYSTHl STRUCTURE Arm FUNCTION 26

3.1 Structural Properties of flangrove Forests.......................... 263.2 Zonation, Succession and "Land Building" 263.3 Nutrient Cycling................................................... 303.4 Litter Fall and Decomposition 323.5 Carbon Export 343.6 Energy Flow 36

CHAPTER 4. COMMUNITY COMPONENTS - MICROORGANISMS 40

CHAPTER 5. COMMUNITY COMPONENTS - PLAinS OTHER THAN t1ANGROVES 41

5.1 Root and Mud Algae 415.2 Phytoplankton...................................................... 435.3 Associated Vascular Plants 43

v

CONTENTS (continued)

Page

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

6.1 Ecological Relationships ...................................•...... 456.2 Arboreal Arthropod Community...................................... 476.3 Prop Root and Associated Mud Surface Conrnunity 476.4 Water Column Community............................................ 49

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

7.1 Basin r~angrove Forests............................................ 507.2 Riverine Forests. 527.3 Fringing Forests along Estuarine Bays and Lagoons 547.4 Fringing Forests along Oceanic Bays and Lagoons 567.5 Overwash Mangrove Islands 567.6 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 619.3 Probi ng Shorebi rds .................................•.............. 659.4 Floating and Diving Water Birds................................... 659.5 Aerially-searching Birds.............................. 679.6 Birds of Prey.. 679.7 Arboreal Birds.................................................... 689.8 Associations between Mangrove Community Types and Birds 709.9 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 .111 .211.311. 411.511.6

Shoreline Stabilization and Storm Protection .Habitat Value to Wildlife ..Importance to Threatened and Endangered Species ........•..........Value to Sport and Commercial Fisheries ..Aesthetics, Tourism and the Intangibles ...................•.......Economi c Products ....................•..............•.............

747475757576

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

12.112.212.312.412.512.612.712.8

Inherent Vul nerabil ity .Man-induced Destruction .Effects of Oil Spill s on Mangroves ..........•.•........••.•.......Man-induced Modifications ...................................•.....Protective Measures Including Transplanting ...•...................Ecological Value of Black vs. Red Mangroves ...•...............•.•.The Importance of Inter-community Exchange .~Ianagement Practices: Preservation ..

vi

7777808184858586

CONTENTS (continued)

Page

REFERENCES 88

APPENDIX A: SUMMARY OF SITE CHARACTERISTICS AND SAMPLING METHODOLOGYFOR 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), blackmangrove (B), and "hite mangrove (W) in Florida 3

2a A typical intertidal profile from south Florida showing thedistribution of red and black mangroves........ 4

2b The pattern of annual sea level change in south Florida 43 Three species of Florida mangroves with propagu1es, flowers and leaves. 64 The six mangrove community types...................................... 85a Aboveground and be10wground biomass of a Puerto Rican red

mangrove forest 165b Light attenuation in a mangrove canopy; canopy height is 8m 166 The hypothetical relationship between waterway position and

community net primary production of Florida mangrove forests 227 The hypothetical relationship between nutrient input, biomass,

producti vi ty, and nutri ent export from mangrove ecosystems 318 Potential pathways of energy flow in mangrove ecosystems 37g Vertical distribution of selected algae and invertebrates on

red mangrove prop roots .....................................•......... 4210 Photograph of red mangrove prop root habitat in clear shallow

"ater with associated animal and plant populations 4611 Gradient of mangrove-associated fish communities showing

representative species................................................ 5112 Aerial photograph of the mangrove belt of southwest Florida

near Whitewater Bay 5313 The mangrove water snake, Nerodia fasciata compressicauda,

curled on a red mangrove prop root 5914 A variety of wading birds feeding in a mangrove-lined pool

near F1 ami ngo, Flori da ...........................................•.... 6215 Osprey returning to its nest in a red mangrove near Whitewater Bay .... 6916 Damaged stand of red and black mangroves near Flamingo, Florida,

as it appeared 7 years after Hurricane Donna 7817 1·langrove forest near Key West as it appeared in 1981 after

being destroyed by diking and impounding 7918 ~Iangrove islands in Florida Bay near Upper Matecumbe Key.............. 87

TABLES

Number Page

la Estimates of mangrove production in Florida ...............•........... 20lb Comparative measurements of photosynthesis in gC/m2/day 21lc Gross primary production at different salinities...................... 212 Aboveground biomass of mangrove forests in the Ten Thousand

Islands region of Florida 273 Estimates of litter fall in mangrove forests 334 Estimates of particulate carbon export from mangrove forests ..•...•... 355 Nesting statistics of wading birds and associated species in

south Florida, 1974-75 646 Timing of nesting by wading birds and associated species in

south Flori da ...................................•..................... 667 General response of mangrove ecosystems to severe oil spills 828 Estimated impact of various stages of oil mining on mangrove ecosystems 83

viii

ACKNOWLEDGMENTS

Many individuals and organizationscontributed significantly to the creationof this publication. Most notable wasEri c Hea 1d who worked extens i ve1y with uson the manuscript and contributed unpub­lished data. We thank Jeffrey Carlton,Edward Conner, Roy R. Lewis, Ill, ArielLugo, Larry Narcisse, Steven Macko, AaronMills, Michael Robb1ee, Martin Roessler,Samuel Snedaker, Durbin Tabb, Mike Wein­stein, and Joseph Zieman for informationand helpful advice.

•The draft manuscript was reviewed for

its scientific content by Armando de 1aCruz, Thomas Savage, James Kush1an (birdsection), and James P. Ray. Each of these

ix

individuals provided information as wellas critical comments. We particularlyappreciate the unselfish help of Ken Adamsand the staff of the National Coastal Eco­systems Team of the U.S. Fish and WildlifeService. Janet Ryan s pent many hours typ­ing the manuscript.

Factual errors and faulty conclu­sions are the sole reseonsibi1ity of theauthors. Carole McIvor has taken primaryres pons i bi 1ity for Chapter 7, Tom Smithfor Chapters 8, 9, and 10, and Bi 11 Odumfor the remainder of the publication.Un 1ess otherwis e noted, photographs,figures, and the cover were produced bythe authors.

x

CHAPTER 1

1.1 "MANGROVE" OEFINITION

The term UmangroveU expresses twodistinctly different concepts. One usagerefers to halophytic species of trees andshrubs (halophyte = plant growing insaline soil). In this sense, mangrove isa catch-all, botanically diverse, non­taxonomic expression given to approximate­ly 12 families and more than 50 species(Chapman 1970) of tropi cal trees andshrubs (see Wai sel 1972 for a detai 1edlist). While not necessarily closelyrelated, all these plants are adapted to(1) loose, wet soils, (2) a saline habi­tat, (3) periodic tidal submergence, and(4) usually have degrees of viviparity ofpropagules (see section 2.3 for discussionof "viviparity" and "propagules").

The second usage of the term mangroveencompasses the entire plant communityincluding individual mangrove species.Synonymous terms include tidal forest,tidal swamp forest, mangrove community,mangrove ecosystem, man gal (Macnae 1968),and mangrove swamp.

For consistency, in this pub1 icationwe wi 11 use the word "mangrove" for i ndi­vidual kinds of trees; mangrove community,mangrove ecosystem or mangrove forest willrepresent the entire assemblage of "man­groves".

1.2 FACTORS CONTROLLING MANGROVE DISTRI­BUTION

Four major factors appear to limitthe distribution of mangroves and deter­mine the extent of mangrove ecosystemdevelopment. These factors include (1)climate, (2) salt water, (3) tidal fluc­tuation, and (4) substrate.

Climate

Mangroves are tropical species anddo not develop satisfactorily in regionswhere the annual average temperature isbelow IgOC or 66 0 F (Waisel 1972).Normally, they do not tolerate temperaturefluctuations exceeding IOoC (l8°F) or

INTRODUCTION

temperatures below freezing for any lengthof time. Certain species, for example,black mangrove, Avicennia germinans, onthe northern coastoT-the-GiiTIcifMexi co,maintain a semi-permanent shrub form bygrowing back from the roots after freezedamage.

Lugo and Zucca (1977) di scuss theimpact of low temperature stress on Flori­da mangroves. They found that mangrovecommunities respond to temperature stressby decreasing structural complexity (de­creased tree height, decreased leaf areaindex, decreased leaf size, and increasedtree density). They concluded that man­groves growing under conditions of highsoil salinity stress are less tolerant oflow temperatures. Presumably, other typesof stress (e.g., pollutants, diking) couldreduce the temperature tolerance of man­groves.

High water temperatures can al so belimiting. McMillan (1971) reported thatseedlings of black mangrove were killed bytemeP.eratures of 39 0 to 40 0 C (102 0 to104 F) although established seedlings andtrees were not damaged. To our knowledge,upper temperature tolerances for adultmangroves are not well known. We suspectthat water temperatures in the range 42 0

to 450 C (1070 to 1130 F) may be limiting.

Salt Water

Mangroves are facultative halo­phytes, i.e., salt water is not a physicalrequirement (Bowman 1917; Egler 1948). Infact, most mangroves are capable ofgrowing quite well in freshwater (Teas1979). It is important to note, however,that mangrove ecosystems do not develop instrictly freshwater environments; salinityis important in reducing competition fromother vascular plant species (Kuenzler1974). See section 2.2 about salinitytolerance of mangrove species.

Tidal Fluctuation

While tidal influence is not adirect physiological requirement for

1

mangroves, it plays an important indirectrole. Fi rst, tidal stress (alternatewetting and drying), in combination withsalinity, helps exclude most othervascular plants and thus reduces competi­tion. Second, in certain locations, tidesbring salt water UP the estuary againstthe outward flow of freshwater and allowmangroves to become establ i shed wellinland. Third, tides may transportnutrients and relatively clean water intomangrove ecosystems and export accumula­tions of organic carbon and reduced sulfurcompounds. Fourth, in areas with highevaporation rates, the action of the tideshelps to prevent soil salinities fromreaching concentrations which might belethal to mangroves. Fifth, tides aid inthe dispersal of mangrove propagules anddetritus.

Because of all of these factors,termed tidal subsidies by E.P. Odum(1 971), mangrove ecosystems tend to reachthei r greatest de vel opment around theworld in low-lying regions with relatively1arge ti da1 ranges. Other types of waterfluctuation, e.g., seasonal variation infreshwater runoff from the Florida Ever­glades, can provide similar subsidies.

Substrate and Wave Energy

Mangroves grow best in depositionalenvi ronments wi th low wave energy. Hi ghwave energy prevents establ i shment ofpropagules, destroys the relatively shal­low mangrove root system and prevents theaccumulation of fine sediments. The mostproductive mangrove ecosystems developalong deltaic coasts or in estuaries thathave fi ne-gra i ned muds composed of si It,c1 ay and a hi gh percentage of organicmatter. Anaerobic sediments pose noproblems for mangroves (see section 2.1)and exclude competing vascular plantspecies.

1.3 GEOGRAPHICAL DISTRIBUTION

Mangroves dominate approximately 75%of the world's tropical coastline between25 0 N and 25 0 S 1at itude (McGi 11 1959). On

the east coast of Africa, in Australia andin New Zealand, they extend 100 to 150

farther south (Kuenzler 1974) and inJapan, Florida, Bermuda, and the Red Seathey extend 50 to 70 farther north. Theseareas of extended range generally occurwhere oceanographic conditions move un­usua lly warm water away from the equator.

Althou9h certai n reqi ons such as thetropical Indo-Pacific have as many as 30to 40 species of mangroves present, onlythree species are found in Florida: thered mangrove, Rhi zophora mangl e, the b1 admangrove, Avicennia germinans, and thewhite mangrove, Laguncul ari a racemosa. Afourth species, buttonwood, Conocar userecta, is not a true mangrove no ten­dency to vivipary or root modification},but is an important species in the transi­tion zone on the upland edge of mangroveecosystems (Tom1 inson 1980).

The ranges of mangrove speci es inFlorida have fluctuated over the pastseveral centuries in response to relative­ly short-term climatic change. Currently,the situation is as follows (Figure 1).The red mangrove and the white mangrovehave been reported as far north as CedarKey on the west coast of Florida (Rehm1976) and north of the Ponce de Leon Inleton the east coast (Teas 1977); both ofthese extremes lie at approximately 2g0 10'N latitude. Significant stands lie southof Cape Canaveral on the east coast andTarpon Springs on the west coast. Theblack mangrove has been reported as farnorth as 30 0 N latitude on the east coastof Flori da (Savage 1972) and as scatteredshrubs along the north coast of the Gulfof Hexi co.

Intertidal Distribution

The generalized distribution of thered and black mangrove in relation to theintertidal zone is shown in Figure 2a.Local variations and exceptions to thispattern occur commonly in response tolocalized differences in substrate typeand elevation, rates of sea level rise,and a variety of other factors (see sec­t ion 3.2 for a fu 11 di scuss i on of mangrove

2

Bay

Ri ver

;.

____R,WPonce de Leon Inlet

_____ 8

Rookery Bay"'"Ten Thousand Islands ..

-..,}

.,' IBiscayne

Shark River Valley. .' I

Whitewater Bay/North Ri ver ::: ,.~'?,.

Flori da Keys •• "~ ,.."

Tarpon

Tampa Bay

Port Charlotte Harbor/Sanibel Island

R,W ~··

Cedar Key

,----____ I

---------. ,i I.~

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 in­dicated in the figure, the black mangrove extends along the northern Gulf of Mex­ico as scattered shrubs.

3

(A)

HIGH

MARSH

...... : : .. '. '."': :~ :.:.

PEAT

\__-.-• ..._/''- --..,..- ----''...., -.J/

LOWMARSH

MLW---

MHW-----

(RED MANGROVE

DOMINATED)

(BLACK MANGROVE

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 dis­tribution of red and black mangrove (adapted from Provost 1974). (b) The pat­tern of annual sea level change in south Florida (Miami)(adapted from Provost1974) .

4

zonation). Furthermore, it is importantto recognize that the intertidal zone inmost parts of Florida changes seasonally(Provost 1974); there is a tendency forsea level to be higher in the fall than inthe spring (Figure 2b). As a result the"high marsh" may remain totally dry duringthe spring and be continually submerged inthe autumn. This phenomenon further com­plicates the textbook concept of the in­tertidal, "low marsh" red mangrove and theinfrequently flooded, "high marsh" blackmangrove.

Mangrove Acreaqe in Florida

Estimates of the total acreageoccupied by mangrove communities inFlorida vary widely between 430,000 acresand over 500,000 acres (174,000 ha to over202,000 hal. Eric Heald (TropicalBioindustries, 9869 Fern St., Miami, Fla.;personal communication 1981) hasidentified several reasons for the lack ofagreement between estimates. Theseinclude: (1) inclusion or exclusion insurveys of small bays, ponds and creekswhich occur within mangrove forests, (2)incorrect identification of mangrove areasfrom aeri al photography as a result ofinadequate "ground-truth" observations,poorly cont ro11 ed ae ri a 1 photography, andsimple errors of planimetry caused byphotography of inadequate scale.

The two most detailed estimates ofarea covered by mangroves in Florida areprovi ded by the Coastal Coordi nati ng Coun­cil, State of Florida (1974) and 8irnhakand Crowder (1974). Considerable dif­ferences exi st between the two esti mates.The estimate of Birnhak and Crowder(1974), which is limited to certain areasof south Florida, appears to be unrealis­tically high, particularly for MonroeCounty (Eric Heald, personal communication1981). Coastal Coordinating Council(1974) estimates a total of 469,000 acres(190,000 hal within the State and suggestsan expected margin of error of 15% (i.e.thei r esti mate lies between 400,000 and540,000 acres or 162,000 and 219,000 hal.

According to this survey, ninety percentof Flori da's mangroves are located in thefour southern counties of Lee (35,000acres or 14,000 hal. Collier (72,000 acresor 29,000 hal, Monroe (234,000 acres or95,000 hal, and Dade (81,000 acres or33,000 hal.

Much of the area covered by mangrovesin Florida is presently owned by Federal,State or County governments, or by non­profit organizations such as the NationalAudubon Society. Approximately 280,000acres (113,000 hal fall into this category(Eric Heald, personal communication 1981).Most of this acreage is held by theFederal Governf'lent as a result of the landbeing including within the EvergladesNational Park.

1.4 MANGROVE SPECIES DESCRIPTIONS

The following descriptions comelargely from Carlton (1975) and Savage(1972); see these publications for furthercomments and photographs. For moredetailed descriptions of germinating seeds(propagules) see section 2.3. The threespecies are shown in Figure 3.

The Black Mangrove (Avicennia ~erminans)

Avicennia germinans is synonymouswith ---,o;:-iiTITda aiid---rsa- me mbe r 0f thefamily Avicenniaceae (formerly classedunder Verbenaceae). The tree may reach aheight of 20 m (64 ft) and has dark, scalybark. Leaves are 5 to 10 cm (2 to 4inches) in length, narrowly elliptic oroblong, shiny green above and covered withshort, dense hai rs below. The 1eaves arefrequently encrusted with salt. Thi s treeis characteri zed by long hori zontal or"cabl e" roots with short vertical aeratingbranches (pneumatophores) that profuselypenetrate the substrate below the tree.Propagul es are 1i ma-bean shaped, darkgreen while on the tree, and severalcentimeters (1 inch) long. The treeflowers 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)

The white mangrove is one of 450species of plants in 18 genera of thefamily Combretaceae (synonymous withTerminaliaceae). It is a tree or shrubreaching 15 m (49 ft) or more in heightwith broad, flattened oval leaves up to 7cm (3 inches) long and rounded at bothends. There are two salt glands at theapex of the petiole. The propagule isvery small (1.0 to 1.5 cm or 0.4 to 0.6inches long) and broadest at its apex.Flowering occurs in spring and earlysummer.

The Red Mangrove (Rhizophora mangle)

The red mangrove is one of more than70 species in 17 genera in the familyRhizophoraceae. This tree may reach 25 m(80 fti in height, has thin grey bark anddark red wood. Leaves may be 2 to 12 cm(1 to 5 inches) long, broad and bl unt­poi nted at the apex. The 1eaves areshiny, deep green above and paler below.It is easily identified by its charac­teristic ·prop roots' arising from thetrunk and branches. The pencil-shapedpropagules are as much as 25 to 30 cm (10to 12 inches) long after germination. Itmay flower throughout the year, but inFlorida flowering occurs predominately inthe spring and early summer.

1.5 MANGROVE COMMUNITY TYPES

Mangrove forest communities exhibittremendous variation in form. Forexample, a mixed scrub forest of black andred mangroves at Tu rkey Poi nt on Bi scayne8ay bears little resemblance to theluxuriant forests, dominated by the sametwo species, along the lower Shark River.

Lugo and Snedaker (1974) provided aconvenient classification system based onmangrove forest physi ogomy. They i denti­fied six major community types resultingfrom different geological and hydrologicalprocesses. Each type has its own charac­teristic set of environmental variablessuch as soil type and depth, soil sa 1i nity

range, and flushing rates. Each communitytype has cha racteri s tic ranges of pri rna ryproduction, litter decomposition and car­bon export along with differences innutri ent recycl i ng rates, and communitycomponents. The community types as shownin Figure 4 are as follows:

(1) Overwash mangrove foreststhese islands are frequently overwashed bytides and thus have ~igh rates of organicexport. All species of mangroves may bepresent, but red mangroves usually domi­nate. Maximum height of the mangroves isabout 7 m (23 ft).

(2) Fri nge mangrove forests - man­groves form a relatively thin fringe alongwaterways. Zonation is typically as de­scribed by Davis (1940) (see discussion insection 3.2). These forests are bestdefined along shorelines whose elevationsare hi gher than mean hi gh ti de. Maxi mumheight of the mangroves is about 10 m (32ft) •

(3) Ri veri ne mang rove fores t s - th iscommunity type includes the tall floodplain forests along flowing waters such astidal rivers and creeks. Although a shal­low berm often exists along the creekbank, the entire forest is usually flushedby daily tides. All three species ofmangroves are present, but red mangroves(with noticeably few, short proo roots)predomi nate. Mangroves may reach hei ghtsof 18 to 20 m (60 to 65 ft).

(4) Basin mangrove forests - theseforests occur inland in depressions chan­neling terrestrial runoff toward thecoast. Close to the coast they are in­fluenced by daily tides and are usuallydominated by red mangroves. Moving in­lanrl, the tidal influence lessens anddominance shifts to hlack and white man­groves. Trees may reach 15 m (49 ft) inhei ght.

(5) Hammock forests - hammock man­grove communities are similar to the basintype except that they occur on ground thatis slightly elevated (5 to 10 cm or 2 to 4inches) relative to surrounding areas.

7

(1) OVERWASH FOREST

(3) RIVERINE FOREST

(5) HAMMOCK FOREST

(2) FRINGE FOREST

(4) BASIN FOREST

(6) SCRUB FOREST

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

8

All species of mangroves may be present.Trees rarely exceed 5 m (16 ft) in hei ght.

(6) Scrub or dwarf forests - thiscommunity type is limited to the flatcoastal fringe of south Florida and theFlorida Keys. All three species arepresent. Individual plants rarely exceed1.5 m (4.9 ft) in height, except wherethey grow over depressions filled withmangrove peat. Many of these tiny treesare 40 or more years of age. Nutrientsappear to be limiting although substrate(usually limestone marl) must playa role.

Throughout this publ ication we haveattempted to refer to Lugo and Snedaker'sclassifi cation scheme wherever possi b1e.Without a system of this type, comparisonsbetween sites become virtuallymeaningless.

1.6 SUBSTRATES

Understand i ng mangrove-substraterelationships is complicated by theability of mangroves to grow on many typesof substrates and because they often alterthe substrate through peat formation andby alterin9 patterns of sedimentation. Asa result, mangroves are found on a widevariety of substrates including fine,inorganic muds, muds with a high organiccontent, peat, sand, and even rock anddead coral if there are sufficientcrevices for root attachment. Mangroveecosystems, however, appear to fl ouri shonly on muds and fine-grained sands.

In Florida, the primary mangrovesoils are either calcareous marl muds orcalcareous sands in the southern part ofthe State and siliceous sands farthernorth (Kuenzler 1974). Sediment distribu­tion and, hence, mangrove development, iscontrolled to a considerable extent bywave and current energy. Low energyshorelines accumulate fine-grained sedi­ments such as mud and silt and usuallyhave the best mangrove growth. Higherenergy shorelines (more wave action orhigher current velocities) are charac­teri zed by sandy sedi ments and 1ess pro­ductive mangroves. If the wave energy

becomes too great, mangroves wi 11 not bepresent. Of the three species of Floridamangroves, white mangroves appear totolerate sandy substrates the best (per­sonal observation), possibly because thisspecies may tolerate a greater depth tothe water table than the other twospecies.

Mangroves in Florida often modify theunderlying substrate through peat deposi­tion. It is not unusual to find layers ofmangrove peat several meters thick under­lying well-established mangrove ecosystemssuch as those along the southwest coast ofFlorida. Cohen and Spackman (1974) pre­sented a detailed account of peat forma­tion within the various mangrove zones ofsouth Florida and also in areas dominatedby black needle rush (Juncus roemerianus),smooth cordgrass (Spartina alterniflora)and a variety of other macrophytes; Cohenand Spackman (1974) also provide descrip­tions and photography to aid in the iden­tification of unknown peat samples.

The following descriptions come fromCohen and Spackman (1974) and from thepersonal ohservations of W.E. Odum andLJ. Heald. Red mangroves produce themost easily recogni zed peat. More recentdeposits are spongy, fi brous and composedto a great extent of fi ne root 1ets (0.2 to3.0 mm in diameter). Also present arp.larger pieces of roots (3 to 25 mm), hitsof wood and leaves, and inorganicmatp.rials such as pyrite, carbonateminerals, and quartz. Older deposits areless easily differentiated although theyremain somewhat fibrous. Peat which hasrecent ly been excavated is reddi sh-brownalthough this changes to brown-black aftera short exposure to air. Older depositsare mottled reddish-brown; deposits with ahigh content of carbonates are greyish­brown upon exca vat ion.

Cohen and Spackman (1974) were unableto find deposits of pure black mangrove orwhite mangrove peat suggesting that thesetwo species may not form extensive depos­its of peat whil e growi ng in pu re stands.There are, however, many examples of peatswhich are mixtures of red mangrovematerial and black mangrove roots. They

9

suggested that the black mangrove peatsidentified hy Davis (1946) were probablymixtures of peat from several sources.

Throughout south Florida the sub­strate underlying mangrove forests maycons; st of compl i cated patterns ofcalcareous muds, marls, shell, and sandinterspersed and overlain by layers ofmangrove peat and with 1i mestone bedrockat the bottom. Detailed descriptions ofthis complex matrix and its spatial varia­tion were given by Davis (1940, 1943,1946), Egler (1952), Craighead (1964),Zieman (1972) and Cohen and Spackman(1974) among others. Scoffin (1970) di s­cussed the ability of red mangrove totrap and hold sediments about its proproots. So called "land-building" by man­9roves is discussed in section 3.2.

values of -100 to -400 mv in mangrovepeats. Such evidence of strongly reducingconditions are not surprising consideringthe fine-grained, high organic nature ofmost mangrove sediments. Although man­groves occur in low organic sediments(less than 1% organic matter), typicalvalues for mangrove sediments are 10% to20% organic matter.

Lee (1969) analyzed 3,000- to 3,500­year-old mangrove peat layers underlyingLittle Black Water Sound in Florida Bayfor lipid carbon content. Peat lipidcontent varied between 0.6 and 2.7 109lipid-C/gram of peat (dry wt) or about 3%of the total organic carbon total. Thesevalues usually increased with depth. Longchain fatty acids (C-16 and C-18) were thedomi nant fatty aci ds found.

Presumably, the acidic character ofmangrove peat results from release oforganic acids during anaerobic decomposi­tion and from the oxidation of reducedsulfur compounds if the peat is dried inthe presence of oxygen. This last pointexplains why "reclaimed" mangrove areasoften develop highly acidic soils (pH 3.5to 5.0) shortly after reclamation. This"cat clay" problem has greatly complicatedthe conversion of mangrove regions toagricultural land in Africa and southeastAsia (Hesse 1961; Hart 1962, 1963; r~acnae

1968) .

Florida mangrove peats are usuallyacidic, although the presence of carbonatematerials can raise the pH above 7.0.Zi em an (1972) found red mangrove peats torange from pH 4.9 to 6.8; the most acidconditions were usually found in the cen­ter of the peat 1ayer. Lee (1969) re­corded a pH range from 5.8 to 6.8 in redman9rove peat at the bottom of a shallowembayment. Althou9h Davis (1940) found adi fference between red mangrove peat (5.0to 5.5) and bl ad mangrove peat (6.9 to7.2), this observation has not been con­firmed because of the previously mentioneddiffi culty in fi ndi ng pu re black man9rovepeat.

The long-term effect of mangrove peaton mangrove distribution is not entirelyclear. Certainly, if there is no changein sea level or if erosion is limited, theaccumul ati on of peat under stands of redmangroves combined with deposition andaccumulation of suspended sedi ments wi 11raise the forest floor sufficiently tolead to domination by black or white man­groves and, ultimately, more terrestrialspecies. Whether this is a common se­quence of events in contemporary southFlorida is not clear. It is clear thatpeat formtion is a passive process andoccurs primari ly where and when physicalprocesses such as erosion and sea levelrise are of minimal importance (Wanless1974) •

Zieman (1972) presented an inter­esting argument suggesting that mangrovepeat may be capabl e of di ssol vi ng under­lying limestone rock, since carbonates maydissolve at pH 7.8. Through this process,shallow depressions might become deeperand the overlyi ng peat layer thickerwithout raising the surface of the forestf1 oor.

Data on chemical characteristics ofFlorida mangrove soils and peat arelimited. Most investigators have foundman9rove substrates to be almost totallyanaerobic. Lee (1969) recorded typical Eh

In summary, although currentstanding of mangrove peats and

10

under­soil sis

fragmentary and often contradictory, wecan outline several generalizations:

(1) Mangroves can grow on a widevariety of substrates including mud, sand,rock, and peat.

(2) Mangrove ecosystems appear toflourish on fine-grained sediments whichare usua 11 y anae robi c and may ha ve a hi ghorganic content.

(3) Mangrove ecosystems which per­si st for some ti me may modify the under­lying substrate through peat formation.This appears to occur only in the absenceof strong physi cal forces.

(4) Mangrove peat is formed pri­mari 1y by red mangroves and consi sts pre­dominantly of root material.

(5) Red mangrove peats may reachthicknesses of several meters, have arelatively low pH, and may be capable ofdissolving underlying layers of limestone.

(6) When drained, dried, andaerated, mangrove soils usually experiencedramatic increases in acidity due to theoxidation of reduced sulfur compounds.This greatly complicates their conversionto agri cul t ureo

1.7 WATER QUALITY

Water quality characteristics of sur­face waters flowing through Florida man­grove ecosystems exhibit great variationfrom one location to the next. Proximityto terrestri al ecosystems, the ocean, andhuman activities are all important indetermining overall water quality.Equally important is the extent of themangrove ecosystem since drastic altera­tions in water quality can occur within astand of mangroves.

In general, the surface watersassoci ated with mangroves are charac­terized by (1) a wide range of salinities

from virtually fresh water to above 40 ppt(discussed in section 2.2), (2) low macro­nutrient concentrations (particularlyphosphorous), (3) relatively low dissolvedoxygen concentrations, and (4) frequentlyincreased water color and turbidity. Thelast three characteristics are most pro­nounced in extensive mangrove ecosystemssuch as those adjacent to the Evergladesand least pronounced in small, scatteredforests such as the overwash islands inthe Flori da Keys.

Walsh (1967), working in a mangroveswamp in Hawai i, was one of the fi rst todocument the tendency of mangrove eco­systems to act as a consumer of oxygen anda sink for nutrients such as nitrogen andphosphorous. Carter et al. (1973) andLugo et al. (1976) confi rmed these obser­vations for Florida mangrove swamps. Evi­dently, nutrients are removed and oxygenconsumed by a combination of periphyton onmangrove prop roots, mud, organic detrituson the sediment surface, the fine rootsystem of the mangroves, small inverte­brates, benthic and epiphytic algae, andbacteria and fun9i on all these surfaces.

The results of oxygen depletion andnutrient removal are (1) dissolved oxygenconcentrations below saturation, typically2 to 4 ppm and often near zero in stagnantlocations and after heavy, storm-generatedrunoff, (2) very low total phosphorusvalues, frequently below detection limits,and (3) moderate total nitrogen val ues(0.5 to 1.5 mg/l). In addition, TOC(total organic carbon) may range from 4 to50 ppm or even hi gher after rai n; Eri cHeald (personal communication 1981) hasmeasured DOC (dissolved organic carbon)values as high as 110 ppm in water flowingfrom mangroves to adjacent bays. Tur­bidity usually falls in the 1 to 15 JTU(Jackson turbity units) range. The pH ofthe water column in Florida swamps isusually between 6.5 and 8.0 and alkalinitybetween 100 to 300 mg/l. Obviously, ex­ceptions to all of these trends can occu~

Both natural and human disturbance canraise macronutrient levels markedly.

11

CHAPTER 2. AUTECOLOGY OF MANGROVES

2.1 AOAPTATIONS TO NATURAL STRESS -ANAEROBIC SEDIMENTS

Mangroves have a series of remarkableadaptations which enable them to flourishin an environment characterized by hightemperatures, wi de ly f1 uctuat in g sa 1i ni ­ties, and shifting, anaerobic substrates.In this section we review a few of themost important adaptat ions.

The root system of mangroves providesthe key to existence upon unfriendly sub­strates (see Gill and Tomlinson 1971 foran anatomical review of mangrove roots).Unlike most higher plants, mangrovesusually have hi ghly developed aeri al rootsand modest below-ground root systems. Theaerial roots allow atmospheric gases toreach the underground roots whi ch areembedded in anaerobic soils. The redmangrove has a system of stilt or proproots which extend a meter (3 ft) or moreabove the surface of the soil and containmany small pores (lenticels) which at lowtide allow oxygen to diffuse into theplant and down to the underground roots bymeans of open passages called aerenchyma(Scholander et al. 1955). The lenticelsare highly hydrophobic and prevent waterpenetration into the aerenchyma systemduring high tide (Waisel 1972).

The black mangrove does not have proproots, but does have small air roots orpneumatophores which extend verticallyupward from the underground roots to ahei ght of 20 to 30 cm (8 to 12 inches)above the soi 1. These pneumatophoresresemble hundreds of tiny fingers stickingup out of the mud underneath the treecanopy. At low ti de, ai r travel s throughthe pneumatophores into the aerenchymasystem and then to all living root tis­sues. The whi te mangrove usually does nothave either prop roots or pneumatophores,but utilizes lenticels in the lower trunkto obtai n oxygen for the aerenchyma sys­tem. "Peg roots" and pneumatophores maybe present in certain situations (Jenik1967).

Mangroves achieve structural stabili­ty in at least two ways. Species such asthe red mangrove use the system of prop

roots to provide a more or less firm foun­dation for the tree. Even though the proproots are anchored with only a modestassemblage of underground roots, the hori­zontal extent of the prop root systeminsures considerable protection from allbut the worst of hurri canes. Other man­grove species, including the black man­grove, obtai n stabil i ty wi th an extens i vesystem of shallow, underground "cable"roots that radiate out from the centraltrunk for a considerable distance in alldi rect ion s; the pneumatophores extend up­ward from these cable roots. As in allFlorida mangroves, the underground rootsystem is shallow and a tap root islacking (Walsh 1974). As Zieman (1972)found, individual roots, particularly ofred mangroves, may extend a meter or moredownward in suitable soils.

From the standpoi nt of effect i venessin transporting oxygen to the undergroundroots, both prop roots and cable rootsseem equally effecti ve. From the perspec­tive of stability, the prop roots of redmangroves appear to offer a distinct ad­vantage where wave and current energiesare high.

Unfortunately, as pointed out by Odumand Johannes (1975), the same structurewhich allows mangroves to thri ve in an­aerobic soil is also one of the tree'smost vulnerable components. Exposed por­tions of the aerial root system are sus­ceptible to clogging by fine suspendedmaterial, attack by root borers, and pro­longed flooding (discussed further insection 12.1). Such extended stress onthe aeri al roots can ki 11 the enti re tree.

2.2 ADAPTATIONS TO NATURAL STRESS ­SALINITY

Manqroves accommodate fluctuations andextremes of water and soil salinitythrough a variety of mechanisms, althoughnot all mechanisms are necessarily presentin the same species. Scholander et al.(1962) reported experimental evidence fortwo major methods of internal ion regula­tion which they identified in two dif­ferent groups of mangroves: (1) the salt

12

exc1 usion species and (2) the salt excre­tion species. In addition, some mangrovesutilize succulence and the discarding ofsalt-laden organs or parts (Teas 1979).

The salt-excluding species, whichinclude the red mangrove, separatefreshwater from sea water at the rootsurface by means of a non-metabolic ultra­filtration system (Scho1ander 1968). This"reverse osmosis" process is powered by ahigh negative pressure in the xylem whichresults from transpiration at the leafsurface. Salt concentration in the sap ofsalt-excluding mangroves is about 1/70 thesalt concentration in sea water, althoughthis concentration is almost 10 timeshigher than found in normal plants(Scholander et al. 1962).

Salt-secreting species, includingbl ack and white mangroves (Scho1 ander1968), use salt glands on the leaf surfaceto excrete excess salt. This is probablyan enzymat i c process rather than a phys i­cal process since it is markedly tempera­ture sensitive (Atkinson et al. 1967).The process appears to i nvol ve acti vetransport with a requirement for biochemi­cal energy input. As a group, the saltsecreters tend to have sap salt concentra­tions approximately 10 times higher (1/7the concentration of sea water) than thatof the salt excluders.

In spite of these two general tenden­cies, it is probably safe to say thatindividual species utilize a variety ofmechanisms to maintain suitable saltbalance (Albert 1975). For example, thered mangrove is an effecti ve, but notperfect, salt excluder. As a result thisspecies must store and ultimately disposeof excess salt in leaves and fruit (Teas1979). Most salt secreters, includingwhite and black mangroves, are capable oflimited salt exclusion at the root sur­face. The white mangrove, when exposed tohypersaline conditions, not only excludessome salt and secretes excess salt throughits salt glands, but also developsthickened succulent leaves and discardssa1t du ri ng 1eaf fall of senescent 1eaves(Teas 1979).

There appears to be some variation inthe salinity tolerance of Florida man­groves. The red mangrove is probah1ylimited b.y soil salinities above 60 to 65PPt. Teas (l979) recalculated Bowman's(1917) data and conc1 uded that transpi ra­tion in red mangrove seedlings ceasesabove 65 ppt. Cintron et al. (1978) foundmore dead than 1i vi ng red mangrove treeswhere interstitial soil salinities ex­ceeded 65 ppt.

On the other hand, white and blackmangroves, which both possess salt excre­tion and limited salt exclusion mech­anisms, can exist under more hypersalineconditions. Macnae (1968) reported thatb1 ack mangroves can grow at soil sal i ni­ti es greater than 90 ppt. Teas (1979)reported dwarfed and gnarled black andwhite mangroves occurring in Florida atsoil salinities of 80 ppt.

There may be an additional factor orfactors involved in salinity tolerance ofmangroves. McMi 11 an (1 975) found thatseedlings of black and white mangrovessurvived short-term exposures to 80 pptand 150 ppt sea water if they were grownin a soi 1 with a moderate c1 ay content.They failed to survive these salinities,however, if they were grown in sand. Asoil with 7% to 10% clay appeared to beadequate for increased protection fromhypersaline conditions.

Vegetation-free hypersaline lagoonsor bare sand flats in the center of man­grove ecosystems have been described bymany authors (e.g., Davis 1940; Fosberg1961; Bacon 1970). These features havebeen variously called sa1itra1s (Holdridge1940), salinas, salterns, salt flats, andsalt barrens. Evidently, a combination oflow seasonal rai nfa 11, occas i ona1 i nunda­tion by sea water, and high evaporationrates results in soil salinities above 100ppt, water temperatures as high as 45 0 C(l13°F) in any shallow, standing water,and subsequent mangrove death (Teas 1979).Once estab1i shed, sal i nas tend to persi stunless regular tidal flushing is enhancedby natural or artificial changes in tidalcirculation.

13

A1thou~h salinas occur frequently inFlorida, they are rarely extensive inarea. For example, between Rookery Bayand Marco Island (south of Naples,Florida) there are a series of salinas inthe bl ack mangrove-domi nated zone on theupland side of the mangrove swamps. Thesehypersaline lagoons occur where the normalflow of fresh water from upland sourceshas been diverted, presumably resulting inelevated soil salinities during the drywi nter months.

In summary, sal inity is a problem formangroves only under extreme hypersa1 i neconditions. These conditions occur natu­rally in Florida in irregularly floodedareas of the "high swamp" above the normalhigh tide mark and are accompanied by highsoi 1 sal i nities. Florida mangroves,listed in order of increasing salinitytolerance, appear to be red, white, andblack.

2.3 REPRODUCTIVE STRATEGIES

As pointed out by Rabinowitz (197Ba),vi rtually all mangroves share two commonreproductive strate9ies: dispersal bymeans of water (van der Pi j 1 1972) andvivipary (Macnae 1968; Gill and Tomlinson1969). Vivipary means that the embryodevelops continuously while attached tothe parent tree and during dispersal.Since there is uninterrupted developmentfrom zygote through the embryo to seed1 i ngwithout any intermediate resting stages,the "ord "seed" is inappropriate forviviparous species such as mangroves; theterm "propagu1e" is generally used in itsplace.

"hi1e the phenology of black andwhite mangroves remains sketchy, Gill andTomlinson (1971) thoroughly described thesequence of flowering in the red mangrove.Flowering in this species may take placeat any time of the year, at least inextreme south Florida, but reaches a maxi­mum in the late spring and summer. Theflowers open approximately 1 to 2 monthsafter the appearance of buds. The flowerremains intact only 1 to 2 days; this

probably accounts for the low fertiliza­tion rate, estimated by Gill and Tomlinsonat 0% to 7.2%. Propagule development isslow, rangi ng from 8 to 13 months. Savage(1972) mentions that on the Florida gulfcoast, red mangrove propagules mature andfall from the tree from July to September.Within the Everglades National Park, black",angroves flower from May unti 1 July andbear fruit from August until Novemberwhile white mangroves flower from May toAugust and bear fruit from July to October(Loope 1980).

The propagu1es of the three speciesof Florida mangroves are easy to differen­tiate. The following descriptions allcome from Rabinowitz (1978a). ,Ihite man­grove propagules are small and flattened,weigh less than a 9ram, are about 2 emlong, are pea-green when they fall fromthe parent tree, and turn mud-brown in twodays or so. The peri carp (wall of theripened propagu1e) serves as a float andis not shed until the seedling is estab­lished. During dispersal the radicle(embryonic root) emerges from the propa­gule. This germination during dispersalhas 1ed Savage (1972) to refer to thewhite mangrove as I'semi-viviparous".

The propagul es of the b1 ack mangrovewhen dropped from the tree are ob10ng­elliptical (resemble a flattened olive),wei gh about 1 g and are about 2 em long.The peri carp is lost within a few daysafter dropping from the tree; at thispoint the cotyledons (primary leaves)unfold and the propagule resembles twobutterfl ies on top of one another.

Propagules of the red mangrove under­go extensive vivipary while on the tree.When propagu1es fall from the tree theyresemble large green beans. They are rod­shaped with pointed ends, about 20 emlong, and wei gh an average of 15 g.

Propagules of all three species floatand remain viable for extended periods oftime. Apparently, there is an obligatedi spersal ti me for all Flori da mangroves,i.e., a certain period of time must elapseduring dispersal for germination to be

14

complete and after which seedling estab­lishment can take place. Rabinowitz(1978a) estimates the obligate dispersalperiod at approximately 8 days for whitemangroves, 14 days for black, and 40 daysfor red. She further estimates the addi­tional time for root establishment at 5,7, and 15 days for white, bl ack, and redmangroves, respect i ve1y. Her est i mate forviable longevity of the propagules is 35days for white mangroves and 110 days forblack. Davis (1940) reports viable propa­gul es of red mangroves that had been keptfloating for 12 months.

Rabinowitz (1978a) al so concludedthat black and white mangroves require astranding period of 5 days or more abovethe influence of tides to take hold in thesoi 1. As a result, these two species areusually restricted to the higher portionsof the mangrove ecosystem where tidaleffects. are infrequent.

The elongated red mangrove propagule,however, has the potential to becomeestablished in shallow water with tidalinfluence. This happens in at least twoways: (1) stranding in a vertical posi­tion (they float vertically) or (2)stranding in a horizontal position,rooting and then vertical erection by theplant itself. Lawrence (1949) and Rabino­witz (l978a) felt that the latter was themore common method. M. Walterding (Cal if.Acad. Sci., San Francisco; personal com­munication 1980) favors vertical estab­lishment; based upon his observations,surface water turbulence works the propa­gule into the substrate during fallin9tides.

Mortality of established seedlingsseems to be related to propagule size.Working in Panama, Rabinowitz (1978b)found that the mortal ity rate of mangroveseedlings was inversely correlated withinitial propagule size. The white man­grove, which has the smallest propagule,has the highest rate of seedling mortal­ity. The black mangrove has an interme­diate mortality rate while the red man­grove, with the largest propagule, has thelowest seedling mortality rate. She

concluded that species with smallpropagules establish new cohorts annuallybut die rapidly, while species such as thered mangroves may have long-lived andoften overl appi ng cohorts.

Propagule size and seedling mortalityrates are particularly important in con­siderations of succession and replacementin established mangrove forests. Light isusually the most serious limiting factorunderneath existing mangrove canopies.Rabinowitz (1978b) suggested that specieswith short-lived propagules must becomeestabl ished in an area which al ready hasadequate light levels either due to treefall or some other factor. In contrast,red mangrove seedlings can become estab­lished under an existing, dense canopy andthen, due to thei r superi or embryoni creserves, are able to wait for months fortree fall to open up the canopy and pre­sent an opportunity for growth.

2.4 BIOMASS PARTITIONING

Few investigators have partitionedthe total biomass, aboveground and below­ground, contained in a mangrove tree. Ananalysis of red mangroves in a PuertoRican forest by Golley et al. (1962) givessome insight into what might be expectedin south Florida. Aboveground and below­ground biomass existed in a ratio of 1:1if fine roots and peat are ignored (Figure5). In this case, peat and very fineroots (small er than 0.5 cm di ameter) ex­ceeded remaining biomass by 5:1. Lugo eta1. (1976) reported the following valuesfor a south Florida red mangrove over washforest. All values were reported in drygrams per square meter, plus and minus onestandard error, and i gnori ng bel owgr~undbiomass. They found 7~0 ! 22 g/m ofleave-.s, l2.8

2! 15.3 g/m of propagu1es,

7043 - 7 g/m of wood, 4695 ! ~ll g/m ofprop roots and 1565 : 234.5 g/m of detri­tus on the forest floor.

Biomass partitioning between dif­ferent species and locations must behighly variahle. The age of the forestwill influence the amount of wood biomass;

15

(A)

LEAVES(778)

BRANCHES (1274)

SOIL TRUNK (2796)'------,----;-...,..-;::--::-=--r--------J - - - - - - - -

SURFACE LARGEROOTS (997)

SMALL ROOTS (4000)

(B)

100005000

6

OLL...-......... .L.-.........----l_...................~_...........__J

o

4

2

8

HEIGHT(M)

FOOT CANDLES

Figure 5. (a) Aboveground and belowground biomass of a Puerto Rican red mangroveforest. 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 thesame forest; canopy height is 8 m (26 ft) (both figures adapted from Golley et al.1962) .

16

detritus varies enormously from one siteto the next dependi ng upon the amou nt offluvial transport. The biomass charac­teristics of a scrub forest probably bearlittle resemblance to those of a fringingforest. At the present time, there is notenough of this type of data available todraw many conclusions. One intriguingpoint is that red mangrove leaf biopassaverages between 700 and 800 g/m atvarious sites with very different forestmorphologies (Odum and Heald 1975a). Thismay be related to the tendency of mangrovecanopies, once they have become estab­lished, to inhibit leaf production atlower levels through self-shading.

Go11 ey et al. (1962) showed that thered mangrove canopy is an extremely effi­cient light interceptor. Ninety-fivepercent of the available light had beenintercepted 4 m (13 ft) below the top ofthe canopy (Figure 5). As a result, 90%of the leaf biomass existed in the upper 4m of the canopy. Chlorophyll followed thesame pattern of distribution.

The leaf area index (LAl) of mangroveforests tends to be relatively low. Go1­ley et al. (1962) found a LA! of 4.4 for aPuerto Rican red mangrove forest. Lugo eta1. (1975) reported a LA! of 5.1 for aFlorida black mangrove forest and 3.5 fora Flori da fri nge red mangrove forest. Adifferent black mangrove forest, in Flori­da, was found to have values ranging from1 to 4 and an average of 2 to 2.5 (Lugoand Zucca 1977). These values comparewith LA!'s of 10 to 20 recorded for mosttropical forests (Golley et al. 1974).The low leaf area values of mangroveforests can be attributed to at leastthree factors: (1) effective light inter­ception by the man9rove canopy, (2) theinability of the lower mangrove leaves toflourish at low light intensities, and (3)the absence of a 1ow-1 i ght-adapted p1 antlayer on the forest floor.

2.5 PR! MAR Y PRODUCTION

Prior to 1970 virtually no informa­tion existed concerning the productivity

of mangroves in Florida. Since that timeknowledge has accumulated rapidly, but itis still unrealistic to expect more thanpreliminary statements about Florida man­grove productivity. This deficiency canbe traced to (1) the difficulties asso­ciated with measurements of mangrove pro­ductivity and (2) the variety of factorsthat affect productivity and the resultingvariations that exist from site to site.

Productivity estimates come fromthree methods: (1) harvest, (2) gas ex­change, and (3) 1itter fall. Harvestmethods require extensive manpower andknowledge of the age of the forest. Theyare best employed in combination withsilviculture practices. Since silvicul­ture of south Florida mangroves is practi­cally non-existent, this method has rarelybeen used in Florida. Noakes (1955),Macnae (1968), and Walsh (1974) should beconsulted for productivity estimates basedon this technique in other parts of theworl d.

Gas exchange methods, based onmeasurements of C02. changes, have theadvantage of precislon and response toshort-term changes in light, temperature,and flooding. They include both above­ground and belowground production. On thenegative side, the necessary equipment isexpensive and tricky to operate properly.Moreover, extrapolations from short-termmeasurements to long-term estimates offerconsiderable opportunity for error.Nevertheless, the best estimates of pro­ductivity come from this method.

The litter fall technique (annual1itter fall x 3 = annual net pri mary pro­duction) was proposed by Teas (1979) andis based on earl i er papers by Bray andGorham (1964) and Goll ey (1972) for othertypes of forests. This is a quick anddi rty method although the 1ack of pre­ci si on remai ns to be demonstrated formangroves. An even quicker and dirtiermethod proposed by Teas (1 979) is to (1)estimate leaf standing crop (using varioustechniques including harvesting or 1i9httransmission re1ationsbips) and (2) multi­ply by three. Thi s assumes an annual leaf

17

time since last severe stress

nutrient content of substrate

depth of substrate

substrate type

Wood production of mangroves appearsto be hi gh compared to other temperate andtropical trees, although no measurementsfrom Florida are available. Noakes (1955)estimated that the wood production of anintensively managed Malayan forest was39.7 metric tons/ha/year. Teas (1979)suggested a wood production estimate of 21metric tons/ha/year for a mature unmanagedred mangrove forest in south Florida. Hisfigure was calculated from a litter/totalbiomass relationship and is certainlysubject to error.

inputs of toxic compounds or nutrientsfrom human activities

Representative estimates of grossprimary producti on (GPP) net pri mary

human influences such as diking,ditching, and altering patterns ofrunoff.

In spite of the difficulties withvarious methods and the interaction ofcontrolling factors, it is possible tomake general statements about certai naspects of mangrove producti vity. Forexample, Waisel's (1972) statement thatmangroves have low transpiration ratesseems to be genera lly true in Flori da.Lugo et al. (1975) rep~rted transpirationrates of 2,500 g H20/m /day for mangroveleaves in a frin~,ng red mangrove forestand 1,482 g H20/m /day for b1 ack mangroveleaves. This is approximately one-thirdto one-half the value found in temperatebroad leaf forests on hot dry days, butcomparable to tropical rainforests (H.T.Odum and Jordan 1970). The low transpi ra­tion rates of mangroves are probably re­lated to the energetic costs of main­taining sap pressures of -35 to -60 atmo­spheres (Scholander et a1. 1965).

Litter fall (leaves, twigs, bark,fruit, and flowers) of Florida mangrovefor'2sts appears to average 2 to 3 dryg/m day in most well-developed mangrovestands (see discussion in section 3.4).This can be an order of magnitude lower inscrub forests.

competing

nutrient content of overlying water

sa1i nity of soil and overlying water

transport efficiency of oxygen to rootsystem

amount of tidal flushing

presence or absence ofspecies

~egree of herbivory

age of the stand

relative wave energy

characteristics of ground water

presence or absence of nesting birds

periodicity of severe stress (hurri­canes, fire, etc.)

species composition of the stand

A minimal, though incomplete, list offactors controlling mangrove productivitymust include the following:

Mangrove producti vity is affected bymany factors; some of these have beenreco9nized and some remain totally ob­scure. Carter et al. (1973) proposelumping these factors into two broad cate­gories: tidal and water chemistry. Webelieve that a number of additional cate­90ries should be considered.

presence or absence of disease andparasi tes

turnover of one, which is supported by thedata of Heald (1969) and Pool et al.(1977).

18

product i on (NPP), and respi rat ion (R) ofFlorida mangroves are given in Table laoCompared to net pri mary production (NPP)estimates from other ecosystems, includingagricultural systems (E.P. Odum 1971), itappears that mangroves are among theworld's most productive ecosystems.Healthy mangrove ecosystems appear to bemore productive than sea grass, marshgrass and most other coastal systems.

Further examination of Table la re­veals several possible tendencies. Thefirst hypothetical tendency, as discussedby Lugo et al. (1975), is for red man9rovesto ha ve the hi ghest tota 1 net product ion,black to have intermediate values andwhite the lowest. This conclusion assumesthat the plants occur within the zone forwhich they are best adapted (see section3.2 for discussion of zonation) and arenot existing in an area with strong limit­ing fa,tors. A scrub red mangrove forest,for example, growing under stressed condi­tions (high soil salinity or low nutrientsupply), has relatively low net produc­tivity (Teas 1979). The pre-eminent posi­tion of red mangroves is shown by thecomparative measurements of photosynthesisin Table Ib; measurements were made withincanopy leaves of trees growing withintheir zones of optimal growth.

A second noteworthy tendency is thatred mangrove GPP decreases with increasingsalinity while GPP of black and whitemangroves increases with increasingsalinity up to a point. Estimates of Hicksand Burns (1975) demonstrate that this maybe a real tendency (Table Ie).

Data presented by Miller (1972),Carter et al. (1973), Lugo and Snedaker(1974), and Hicks and Burns (1975) sug­gest a third hypothetical tendency,assuming occurrence of the species withinits adapted zone. It appears that theblack mangrove typically has a much higherrespi ration rate, lower net producti vity,and lower GPPjR ratio than the red man­grove. This can be attributed at leastpartially, to the greater salinity stressunder which the black mangrove usuallygrows; this leads to more osmotic work.

These three apparent tendencies haveled Carter et al. (1973) and Lugo et al.(1976) to propose a fourth tendency, aninverted U-shaped relationship betweenwaterway position and net mangrove com­munity productivity (Figure 6). Thistendency is best understood by visualizinga typi cal gradi ent on the southwest coastof Florida. At the landward end of thegradient, salinities are very low,nutrient runoff from terrestrial eco­systems may be high and tidal amplitude isminor. At the seaward end, salinities arerelatively high, tidal amplitude is rela­tively great and nutrient concentrationstend to be lower. At either end of thegradient, the energetic costs are high anda 1a rge percentage of GPP is used forse1 f-maintenance; at the landward end,competition from freshwater plant speciesis high and at the seaward end, salinitystress may be limiting. In this scenario,the highest NPP occurs in the middleregion of the gradient; salinity and tidalamplitude are high enough to limit compe­tition while tidal flushing and moderatenutrient levels enhance productivity.Hicks and Burns (1975) present data tosupport thi s hypothesi S.

In addition to these hypothesesgenerated from field data, there have beentwo significant, published attempts toderi ve hypotheses from mathemat i cal si mu­lation models of mangroves. The first(Mi ller 1972) is a model of primary pro­duction and transpi ration of red mangrovecanopies and is based upon equations whichutilize field measurements of the energybudgets of individual leaves. This modelpredicts a variety of interesting trendswhich need to be further field tested.One interesting hypothesis generated bythe model is that maximum photosynthesisof red mangrove stands should occur with aleaf area index (LAI) of 2.5 if no accli­mation to shade within the canopy occurs;higher LAI's may lead to decreased produc­tion. Another prediction is that redmangrove production is most affected byair temperature and humidity and, to alesser degree, by the amount of solar

19

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

Mature ~a a 8.8 20.5 L. F. Teas (1979)

Scrub R a a 1 . a 3.8 L. F. Teas (1979)

Basin B a a 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 a 1 . (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 ingC/m 2/day (Lugo et al. 1975).

Mangrove type

Red

81ack

Wh i te

Red (seedling)

Daytime netphotosynthesis

1 .38

1. 24

0.58

0.31

Nighttimerespiration

0.23

0.53

0.17

1 .8 g

6.0

2.3

3.4

negative

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

Mangrove type Average surfacesalinity (ppt)

GPP(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

WATERWAY POSITION

SEAWARD

Figure 6. The hypothetical relationship between waterway position and communitynet primary production of Florida mangrove forests (based on Carter et al. 1973).

22

radiation within the ambient range. Grossphotosynthesis per unit leaf area wasgreater at the top of the tree canopy thanat the bottom, although the middle levelshad the greatest production.

Miller (1972) concluded by suggestingthat the canopy distribution of red man­grove leaves is nearly optimal for ef­ficient water utilization rather thanproduction. This indicates that the cano­py is adapted to maximizing productionunder conditions of saturated water sup­ply.

The mangrove ecosystem model reportedby Lugo et al. (1976) provides hypotheseson succession, time to arrive at steadystate conditions (see section 3.2)' andseveral aspects of productivity. Themodel output suggests that the relativeamount of tidal amplitude does not affectGPP significantly; instead, GPP appears tobe extremely sensitive to inputs of ter­restrial nutrients. It follows that loca­tions with large amounts of nutrient inputfrom terrestrial sources (riverine man­grove communi ties) ha ve hi gh rates ofmangrove production (see section 3.3).All si mul ation model-generated hypothesesneed to be field tested with a particular­ly critical eye, since the simplifyingassumptions that are made in constructingthe model can lead to overly simplisticanswers.

Mangrove productivity research re­mains in an embryonic stage. Certainpreliminary tendencies or hypotheses havebeen identified, but much work must bedone before we can conclude that thesehypotheses cannot be fa 1sifi ed.

2.6 HERBIVORY

Oirect herbivory of mangrove leaves,leaf buds, and propagules is moderatelylow, but highly variable from one site tothe next. Identified grazers of livingplant parts (other than wood) include thewhite-tailed deer, Odocoileus virginianus,the mangrove tree crab, Aratus pisonii,and insects including beetles, larvae of

lepidopterans (moths and butterfl ies), andorthopterans (grasshoppers and crickets).

Heald (1969) estimated a mean grazingeffect on North Ri ver red mangrove 1eavesof 5.1% of the total leaf area; valuesfrom leaf to leaf were highly variablerangi ng from 0 to 18%. Beever et al.(1979) presented a detailed stuny ofgrazi ng by the mangrove tree crab. Thi sarboreal grapsid crab feeds on numerousitems including beetles, crickets, cater­pillars, littoral algae, and dead animalmatter. In Florida, red mangrove leavesform an important component of the di et.Beever et al. (1979) measured tree crabgrazing ranging from 0.4% of the totalleaf area for a Florida Keys overwashforest to 7.1% for a fringing forest atPine Island, Lee County, Florida. Theresearchers also found that tree crabgrazing rates are related tg crab density.Low densities (one crab/m ) resulted inlow leaf area damage (less than 1% oftotal l'3af area). High densities (fourcrabs/m ) were accompanied by leaf areadamage ranging from 4% to 6% (see section6.2) •

Onuf et al. (1977) investigated in­sect berbivory in fringing and overwashred mangrove forests in the Indian Riverestuary near Ft. Pierce, Florida. Theyfound si x major herbi vorous insectspecies, five lepidopteran larvae and abeetle. Comparisons were made at a bighnutrient site (input from a bird rookery)and a low nutrient site. 80th red man­grove production and leaf nitrogen weresignificantly bigher at the high nutrientsite. This resulted in a four-foldgreater loss to berbivores (26% of totalleaf area lost to grazing); this increasedgrazing rate more than offset the in­creased leaf production due to nutrientinput.

Calculations of leaf area damage mayunderestimate the impact of berbivores onmangroves. For example, the larvae of tbeolethreutid moth, Ecdytolopha sp.,develops within red mangrove leaf buds andcauses the loss of entire leaves. Alls tag e s 0 f the bee t 1 e , Poe c !..l2.P.~

23

rhizophorae, attack mangrove propaguleswhile-still attached to the parent tree(Onuf et al. 1977).

2.7 WOOO BORERS

Many people have the mistaken ideathat mangrove wood is hi ghly resi stant tomarine borers. Whi Ie this may be true toali mited extent for certain mangrovespecies in other parts of the world, noneof the Florida mangroves have borer­resistant wood. Southwell and BoHman(1971) found that the wood of red, black,and white mangroves has no resistance toTeredo, Pholad and Simnorid borers; piecesOf-red ma-ngrove wood were completely de­stroyed after immersion in ocean water for14 months.

An interesting controversy surroundsthe ability of the wood boring isopod,Sphaeroma terebrans, to burrow into theliving prop roots of the red mangrove.Rehm and Humm (1 973) were the fi rst toattribute apparently extensi ve damage ofred mangroves stands within the TenThousand Islands area of southwesternFlorida to an isopod, Sphaeroma. Theyfound extensive damage throughoutsouthwest Florida, some infestation northto Tarpon Springs, and a total lack ofinfestation in the Florida Keys from KeyLargo south to Key Ilest. The destruct; onprocess was described as follows: theadult isopod bored into the prop roots (5­mm diameter hole); this was followed byreproduction within the hole and develop­ment of juveni 1es withi n the root. Thi sprocess, combi ned wi th secondary decompo­sition from fungi and bacteria, frequentlyresults in prop root severance near themean high tide mark. These authorsattributed loss of numerous prop rootsand, in some cases, loss of entire treesduring storms to isopod damage.

The extent of damage in the TenThousand Islands region led Rehm and Humm(1973) to term the phenomenon an "eco­catastrophe" of possibly 9reat importancLThey fu rther stated that sh ri nk i ng ofmangrove areas appeared to be occurring as

a result of Sphaeroma infestation; thispoi nt was not documented.

Enri ght (1974) produced a tongue-in­cheek rebuttal, on behalf of Sphaeroma andagainst the t1terrestrial invacter li

, redmangroves. Snedaker (1974) contributed amore substantial argument in which hepointed out that the isopod infestationmight be an example of a long-term eco­system control process.

Further arguments against the '~coca­

tastrophe" theory were advanced by Estevezand Simon (1975) and Estevez (1978). Theyprovi ded more 1i fe hi story i nformati on forSphaeroma and suggested a possible ex­planation for the apparently destructiveisopod infestations. They found twospecies of isopods inhabiting red mangroveprop roots, S. terebrans and a sympatri ccongener, ~.guadridentatum. The latterdoes not appear to be a wood borer bututilizes S. terebrans burrows. Neitherspecies appeared to utilize mangrove ·woodas a food source. Estevez and Si man(1975) found extensive burrowing intoseedlings in addition to prop root damage.In general, infestations appeared to bepatchy and limited to the periphery ofmangrove ecosystems. In areas with thehighest density of burrows, 23% of allprop roots were infested. There appearedto be more colonization by S. terebrans inregions with full strengtilsea water (30to 35 ppt).

The most important finding by Estevezand Si mon (1975) and Estevez (1978) wasthat periods of accelerated activity by S.terebrans were related to periods of flu~tuating and slightly increased salinity.This suggests that fluctuations in isopodburrowing may be related to the magnitudeof freshwater runoff from the Everglades.These authors a9ree with Snedaker (1974)and suggest that root and tree loss due toSphaeroma activity may be beneficial tomangrove ecosystems by accelerating pro­duction and root germination. Simber10ffet a1. (1978) amplified this last sugges­tion by showing that root branching, whichis beneficial to individual trees, isstimulated bY isopod activity.

24

This ecocatastrophe versus beneficialstimulus argument is not completely re­solved. Probably, Sphaeroma root destruc­tion, in areas of low isopod density, canbe a beneficial process to hoth the in­dividual tree and to the entire mangrovestand. Whether changes in freshwaterrunoff have accelerated this process tothe point where unnatural and widespreaddamage is occurri ng is not cl ear. Thedata and research perspective to answerthis question do not exist. As a result,we are reduced to providing hypotheseswhich cannot be tested with availahleknowledge.

2.8 MANGROVE 0[SEASES

Publ i shed research on mangrovediseases is rare. The short paper byOlexa and Freeman (1975) is the principalreference for diseases of Florida man­groves. They reported that black man­groves are affected by the pathogenic

fungi, Phyllosticta hibiscina and Nigro­spora sphaerica. These authors found thatP. hibiscina caused necrotic lesions anddeath of black mangrove 1eaves. They fe 1tthat under conditions of high relativehumidity coupled with high temperatures,this fungus could pose a serious threat toindividual trees, particularly if the treehad been weakened by some other naturalagent, such as 1i ghtni ng or wi nd damage.Nigrospora sphaerica was considered to beof little danger to black mangroves.Another fungus, Cylinrocarpon ~~~~~~,

appears to form galls on the prop rootsand stems of red mangroves. Olexa andFreeman (1975) noted mortality of redmangroves in areas of hi gh gall infesta­tions, although a direct causation linkwas not proven.

Further research on mangrove diseasesis hadly needed. Viral disease must beinvestigated. The role of pathogens inlitter production and as indicators ofmangrove stress may be very important.

25

CHAPTER 3. ECOSYSTEM STRUCTURE AND FUNCTION

3.1 STRUCTURAL PROPERTI ES OF MANGROVEFORESTS

Published information about thestructural aspects of Florida mangroveforests is limited; most existing datahave been published since the mid-1970's.This lack of information is unfortunatesince quantitative structural data greatlyaid understanding of processes such assuccession and primary production. Evenmore important, the response of mangroveforests to stress, both climatic and man­induced, can be followed quantitativelywith this type of data.

Ball (19BO) contributed substantiallyto understandin9 the role of competi­tion in man9rove succession by measurin9structural factors such as basal area,tree hei9ht, and tree density. Lugo andZucca (1977) monitored the response ofmangrove forests to freezing temperaturesby observing changes in structural proper­ties of the trees.

Baseline studies of forest structurehave been published by Lugo and Snedaker(1975), and Pool, Snedaker and Lugo(1977). For exampl e, LU90 and Snedaker(1975) compared a fri ngi ng mangrove forestand a basin forest at Rookery Bay, nearNaples, Florida. They found the fringingforest, which was dominated by red man­groves, to have a tree diversi~y of H =1.4B, a basal area of 15.9 m ;Sha, anaboveground biomass of 17,932 glm , and anon-existent litter layer. The nearbybasin forest was dominated by black man­groves, had a tree divers~ty of H = 0.96and a basal area of 23.4 m Iha. The lit­ter lay~ in the basin forest averaged 550dry g/m. Tree diversity in a hurricanedisturbed sect i on of the Rookery Bayforest was 1.62. Si mi 1ar data were pre­sented for mangrove forests in the TenThousand Islands area (Table 2).

Oata of this type are useful for manypurposes including impact statements, en­vironmental surveys, and basic scientificquestions. Cintron et al. (197B) gave anindication of the di rection in which fu­ture research might proceed. Working in amangrove stand in Puerto Rico, they found

tree height to be inversely proportional(r = 0.72) to soil salinity in the range30 to 72 ppt. Above 65 ppt sal inity, deadtree basal area was higher than live treebasal area and above 90 ppt there was nolive tree basal area.

It should be possible to investigatethe relationship between a variety ofmangrove structural properties and factorssuch as flushing frequency, soil depth,nutrient availability, pollution stress,and other measures of human impact. Ulti­mate I y, thi s shou 1d 1ead to an abil ity topredict the form and structure of mangroveforests resulting from various physicalconditions or artificial impacts. Oneexample of this potential tool is Ball's(19BO) documentation of structural changesin mangrove forests resulting from altera­tions in the hydrological conditions ofsouth Florida.

3.2 ZONATION, SUCCESSION AND "LAND-BUILDING"

Much of the worl d's mangrove litera­ture consists of descriptive accounts ofzonation in mangrove forests and the spe­cies composition within these zones. Al­thouqh general agreement has been lacking,various hypotheses have been put forthconcerning the possible connection betweenzonation, ecological succession, competi­tion, and the role of physical factorssuch as soil salinity and tidal amplitude.In this section we review briefly thedominant ideas about mangrove zonation andsuccession and present our interpretationof the current status of knowl edge.

Davis (1940), working in south Flori­da, was one of the first investigators todescribe distinct, almost monospecific,zones within mangrove ecosystems. In whathas become the classical view, he arguedtha+, mangrove zonation patterns wereequivalent to seral stages in succession.The most seaward zone, domi nated by redmangroves, was regarded as the "pioneerstage". More landward zones weredominated by white mangrove, blackmangrove, buttonwood and, fi nally, theclimatic climax, a tropical forest. Since

26

Table 2. Aboveground biomass of mangrove forests in the Ten Thousand Islands regionof 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 Riverinemangroves mangroves mangroves mangroves

Site A B A B C A 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 148N

'"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 ta 1 above- 9,008 146,953 133,572 108,920 177,773 251,278 141,168 207,831.round biomass

these zones were regarded as progressivelylater stages in succession, the entiremangrove ecosystem was bel i eved to bemovi ng seaward through a process of sedi­ment accumulation and colonization. Davisbased his argument primarily upon thesequence of observed zones and cores whichshowed red mangrove peat underlying blackmangrove peat which, in turn, occurredunder terestrial plant communities.

Unfortunately, this Clementsian in­terpretation of mangrove zonation waswidely accepted, but rarely tested. Forexample, Chapman (197D) expanded Davis'original successional concept from southFlorida to explain zonation in mangroveforests in other parts of the world.\,a1sh (1974) thoroughly reviewed the man­grove succession/zonation literature.

Fortunately, not everyone acceptedDavis' point of view. Egler (1952) andlater Thorn (1967, 1975) argued that man­9rove zonation was a response to externalphysical forces rather than temporal se­quence induced by the pl ants themsel ves.E91er (1952) showed that patterns of sedi­ment deposition predicted by Davis' (1940)theory did not always occur. He alsoshowed that in some cases mangrove zonesappeared to be moving landward rather thanseaward. Sea level has been rising insouth Florida at the rate of 1 ft (30 ern)per 100 to 150 years (Provost 1974).Spackman et a1. (1966) emphasized the roleof sea 1eve1 change in determ i ni ng changesin mangrove zonation, both through sealevel rise and land subsidence. BothEgler (1952) and Spackman et a1. (1966)along with Wanless (1974) and Thom (1967,1975) suggested that mangroves werereacting passively rather than actively tostrong geomorphological processes. Thisimplies that mangroves should be regardedas "1 and-stabi 1i zers" rather than "1 and­builders".

Furthermore, field researchers fre­quent ly noted that red mangroves were notalways the only "pioneer species" on re­cently deposited sediment. It is notunusual to find seedlings of black, white,and red mangroves growing together on anew colonization site. Lewis and Dunstan

(1975) found that black mangroves andwhite mangroves along with the sa1tmeadowcordgrass, Spartina patens, are often thepioneers on new dredge spoil islands incentral Florida. On the northern coast ofthe Gulf of Mexico, where black mangroveis the only mangrove species present, itmay be preceded by marsh grasses such assaltmarsh cordgrass, 2. patens, smoothcordgrass, S. alternif10ra, or the blackneedl e rush,- Juncus roemeri anus. I n Puer­to Ri co, we ohserved that whi te mangroveoften pi oneers and dominates sites whereoceanic overwash of heach sand has oc­curred. All of these ohservati ons detractfrom Davis' (1940) original contentionthat red mangroves should be regarded asthe initial colonizer of recently de­posited sediments. It appears that undercertain conditions, e.g., shallow waterdepths, suhstrate type, and latitude,white and black mangroves or marsh grassescan be effective pioneer species.

The work of Rabinowitz (1975) added anew perspective to the mangrove zonationdebate. Through carefully designed recip­rocal planting experiments in Panamanianmangrove forests using species of Rhizo­~'!.' .!:.'!._9.,~r!.cularia, Pelliciera andAvicennia, she demonstrated that eachspecies could grow well within any of themangrove zones. In other words, physicaland chemi cal factors such as soi 1 sal i nityor frequency of tidal inundation, withineach zone, were not solely responsible forexcluding species from that zone. Toexplain zonation, Rabinowitz proposedtidal sorting of propagu1es based uponpropagule si ze, rather than habitat adap­tation,as the most important mechanism forzonation control.

The most recent pi ece to be added tothe zonation/succession puzzle comes fromthe work of Ball (1980). Based upon re­search of mangrove secondary successionpatterns adjacent to Biscayne Bay, Flori­da, she made a strong case for the impor­tance of interspecific competition incontrolling zonation. She found thatwhite mangroves, which grow best inintertidal areas, do not occur consis­tently in the intertidal zone of maturemangrove stands. Instead, white manqroves

28

dominate higher, drier locations abovemean high water where the red mangrovedoes not appear to have a competitiveadvantage. She suggested that competitionis not so important during the earlystages of successi on but becomes criti calas indi vidual trees reach maturity andrequi re more space and other resources.

Inherent in Ball's concept of zona­tion is the differential influence ofphysical factors (e.g., soil salinity,depth to water table) on the competitiveabi 1it i es of the di fferent mangrovespecies. She concluded that successionproceed s independent 1y withi n each zone,althougn breaks in the forest canopy fromlightning strikes or high winds may pro­duce a mosaic of different successionalstages withi n a zone. These openi ngsallow species whose seedlings do not com­pete well in shade, such as the whitemangrove, to become established, at leasttemporarily, within solid zones of redmangroves.

Zonation of mangrove species does notappear to be controlled by physical andchemical factors di rect1y, but by theinterplay of these factors with interspe­cific competition and, possibly, throughtidal sorting of propagu1es. Once succes­sion in a mangrove zone reaches an equili­brium state, change is unlikely unless anexternal perturbation occurs. These per­turbations range from small-scale distur­bance (lightning strikes) to large-scaleperturbations (sea level change, hurricanedamage) and may cause succession withinzones to regress to an earlier stage.There is some evidence in south Floridathat hurricane perturbations occur on afairly regular basis, creating a patternof cyclical succession.

Except for Ball (1980) and Taylor(1980), the importance of fires as ani nfl uence on mangrove success i on has beengenera 11y ignored. Most fi res in theFlorida mangrove zone are initiated bylightning and consist of small circularopenings in the mangrove canopy (Taylor1980). These openi ngs present an opportu­nity for secondary succession within anestablished zone. For example, we have

frequently observed white mangrovesflourishing in small lightning-createdopenings in the center of red mangroveforests. Fire may also playa role inlimiting the inland spread of manQroves.Taylor (1981) pointed out that Evergladesfires appear to prevent the encroachmentof red and white mangroves into adjacentherbaceous communities.

Finally, Lugo and Snedaker (1974),Cintron et al. (1978) and Lugo (1980)suggested that mangrove ecosystemsfunction as classical successional systemsin areas of rapid sediment deposition orupon recently colonized sites such asoffshore islands. They concluded that inmost areas mangrove forests are an exampleof steady-state cyclical systems. Concep­tually, this is synonymous to E. P. Odum's(1971) cyclic or catastrophic climax.Chapnan (lg76a, b) suggested the idea ofcyclic succession for a variety of coastalecosystems.

[f Florida mangrove ecosystems arecyclic systems, then there should be anidentifiable perturbation capable of set­ting succession back to an early stage.Lugo and Snedaker (1974) suggested thathurricanes may play this role. Theypointed out (without substantiating data)that major hurricanes occur about every20-25 years in south Florida. Coinci­dently, mangrove ecosystems appear toreach their maximum levels of productivityin about the same period of time (Lugo andSnedaker 1974). This hypothesis suggeststhat succession within many mangrove eco­systems may proceed on a cyclical basisrather than in the classical fashion.Possibly other physical perturbations mayinfluence mangrove succession includingincursions of freezing temperatures intocentral Florida, periodic droughts causingunusually high soil salinities (Cintron etal. 1978), and fire spreading into theupper zones of mangrove forests from ter­restrial sources.

Although understanding of zonationand succession in mangrove ecosystemsremains incomplete, a clearer picture isemerging, at least for south Florida.Contrary to early suggestions, mangrove

29

species zonation does not appear to repre­sent seral stages of succession except,perhaps, for locations of recent coloniza­tion or where sediment is accumulatingrapidly. The role of mangroves inland-building seems more passive thanact i ve. Geomo rpho log i ca 1 and hydro1ogi ca 1processes appear to be the dominant forcesin determining whether mangrove shorelinesrecede or grow. The role of mangroves isto stabilize sediments which have beendeposited by physical processes.

3.3 NUTRIENT CYCLING

Current understanding of nutrientcycles in mangrove ecosystems is far fromsatisfactory. Sporadic field measurementshave been made, but a complete nutrientbudget has not been publ i shed for anymangrove ecosystem in the world.

Several pioneering field studies wereconducted in Florida (Carter et al. 1973;Snedaker and Lugo 1973; Onuf et al. 1977)and one simulation model of mangrove nu­trient cycling has been published (LU90 etal. 1976). Preliminary measurements ofnitrogen fixation were made (Zuberer andSilver 1975; Gotto and Taylor 1976;Zuberer and Sil ver 1978; Gotto et al.1981). 8ased on these studies, we presentthe following preliminary conclusions.

Mangrove ecosystems tend to act as asink (net accumulator) for various ele­ments including macro nutrients such asnitrogen and phosphorus, trace elements,and heavy metals. As we have discussed insection 1.7, these elements are removedfrom waters f1 owi ng through mangroveSI,amps by the concerted action of themangrove prop roots, prop root al gae, theassociated sediments, the fine root systemof the mangrove trees, and the host ofsmall invertebrates and microorganismsattached to all of these surfaces. Al­thou9h the turnover times for these ele­ments in mangrove swamps are not known, itappears that at least a portion may bestored or tied up in wood, sediments, andpeat for many years.

Although mangrove ecosystems may tendto accumulate nutrients, there is a con­tinual loss through export of particulateand di ssol ved substances. If si gni fi cantnutrient storage and resultant high pri­mary production are to occur, there musthe a continual input of nutrients to themangrove forest from outside the system(Figure 7). Where nutrient influx to themangrove ecosystem is approximatelybalanced by nutrient loss in exportedorganic matter, then nutrient storage willbe minimal and mangrove net primary pro­duction wi 11 be low. This appears tooccur in the scrub mangrove community typeand to a lesser extent in the basin andhammock community types.

Carter et al. (1 973) and Snedaker andLugo (1973) have hypothesi zed that thegreatest natural nutrient inputs for man­grove swamps come from upland and terres­trial sources. Apparently for this rea­son, the most luxuriant and productivemangrove forests in south Florida occur inriverine locations or adjacent to siqnifi­cant upl and drai nage.

Localized sources of nutrients, suchas bird rookeries, can result in greaternutri ent storage and hi gher mangrove pro­duct i vity (Onuf et a1. 1977). I f however,large bird rookeries (or artificial nu­trient inputs) occur in poorly flushedsections of mangrove ecosystems, resultanthigh nutrient levels may inhibit mangrovegrowth (R. R. Lewis, III, HillsboroughCommunity ColI ege, Tampa, Fl a.; personalcommunication 1981).

The output from the simulation modelof Lugo et al. (1976) suggests that ifnutri ent input to a mangrove ecosystem isreduced, then nutrient storage levelswi thi n the mangrove ecosystem wi 11 bereduced and mangrove biomass and produc­tivity will decline. To our knowledgethis hypothesis has not been tested in thefi e1d.

Nitrogen fixation occurs in mangroveswamps at rates comparable to thosemeasured in other shallow, tropical marineareas (Gotto et al. 1981). Nitrogen

30

SMALL

IMPORT

LARGE

IMPORT

LOW STORAGE

LOW BIOMASS

LOW PRODUCTIVITY

HIGH STORAGE

HIGH BIOMASS

HIGH PRODUCTIVITY

SMALL

EXPORT

MODERATE

EXPORT

Figure 7. The hypothetical relationship between nutrient input (excluding carbon),biomass, primary productivity, and nutrient export (including carbon) from mangroveecosystems. Top: small nutrient import. Bottom: large nutrient import.

31

fixation has heen found in associationwith mangrove leaves, both living anddead, mangrove sediment surfaces, thelitter layer in mangrove swamps, and man­grove root systems (Gotto and Taylor 1976;Zuberer and Silver 1978: Gotto et a1.1981). In virtually all cases, nitrogenfixation appears to be limited by theavailability of labile carbon compounds.Perhaps for this reason, the highest ratesof mangrove nitrogen fixation have beenmeasured in association with decayingmangrove leaves; presumably, the decayingleaves act as a carbon source and thusaccelerate nitrogen fixation. Macko(1981), using stable nitrogen ratiotechniques, has indicated that as much as25% of the nitrogen associated with blackmangrove peat in Texas is derived fromnitrogen fixation.

Zuberer and Silver (1978) speculatedthat the nitrogen fi xati on rates observedin Florida mangrove swamps may be suf­ficient to supply a significant portion ofthe mangrove's growth requi rements. Al­though this hypothesis is impossible totest with present information, it mightexplain why moderately productive mangrovestands occur in waters which are severelynitrogen depleted.

In summary, knowledge of nutrientcycling in mangrove swamps is highlyspecul at i ve. These ecosystems appear to

'act as a sink for many elements, includingnitrogen and phosphorus, as long as amodest input occurs. Nitrogen fixationwithin the swamp may provide much of thenitrogen needed for mangrove growth.

3.4 LITTER FALL AND DECOMPOSITION

Unless otherwise stated, litter fallrefers to leaves, wood (twigs), leafscales, propagules, bracts, flowers, andinsect frass (excrement) which fall fromthe tree. Mangrove leaves are shed con­tinuously throughout the year although aminor peak occurs during the early part ofthe summer wet season in Florida (Heald1969; Pool et al. 1975). Sporadic litterfall peaks may follow periods of stressfrom cold air temperatures, high soil

32

salinities, and pollution events. Litterfall typically can be partitioned as 68%to 86% leaves, 3% to 15% twigs and 8% to21% miscellaneous; the latter includesflowers and propagules.

Litter fall is an important ecosystemprocess because it forms the energy basi sfor detritus-based foodwebs in mangroveswamps (see sections 3.5 and 3.6). Thefirst measurements of litter fall in man­grove swamps were made by E.J. Heald andW.E. Odum, working in the North Riverestua ry in south Flori da in 1966-69.This was subsequently published as Heald(1969), Odum (1970), and Odum and Heald(1975a). They estimated that 1i tter pro­duction from riverine red mangrove forestsaverage! 2.4 dry g 2f organicmatter/m /day (or 876 g/m /year or 8.8metri c tons/ha/year).

Subsequent studi es agreed with thi searly estimate (Table 3), although varia­tion clearly exists between differenttypes of communities. Scrub forests withscattered, very small trees have thesmallest amount of leaf fall. Basin andhammock forests, which appear to benutrient limited, have intermediate leaffall values. Not surprisingly, thehighest values occur in the highly produc­tive fringing, overwash, and riverineforests. Odum and Heald (1975a) suggestedthat the relatively uniform litter fallvalues from productive mangrove forestsaround the world result from the shadeintolerance of the canopy leaves and thetendency for the canopy size to remain thesame in spite of increasing height. Ifdetailed information is lacking, red man­grove forests of south Flori da, whi ch arenot severely limited by lack of nutrients,can be assume~ to produce litter fall of2.0 to 3.0 g/m /day of dry organi c matter.Pure stands of black mangroves us~al1y

have a lower rate of 1.0 to 1.5 g/m /day(Lugo et a1. 1980).

Decomposition of fallen Florida man­grove leaves has been investigated by anumber of researchers including Heald(1969), Odum (1970), Odum and Heald(1975a), 'Pool et a1. (1975), Lugo andSnedaker (1975), Twilley (1980) and Lugo et

Table 3. Estimates of litter fall in mangrove forests. Total litter fall in­cludes leaves, fruits, twigs, flowers, and bark. R = red mangrove, W = whitemangrove, 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 a 1 . 1975

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

R (fringe) 2.7 9.9 Po 0 1 et a 1 . 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 1979w

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 a 1 . 1980

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

B 0.8 2.9 Po 0 1 et a 1 . 1975

Vari ety of 0.8 - 2 . 1 2.9 - 7.7 Heald eta 1 . 1979community types

26 species 2.4 8.8 Boto & Bunt (r'lS. in(Austral i a) prep.)

a1. (1980). Heal d and Odum showed thatdecomposition of red man9rove leavesproceeds most rapidly under marine condi­tions, somewhat more slowly in freshwater,and very slowly on dry substrates. Forexample, using the litter bag method, theyfound that only 9% of the original dryweight remained after 4 months in seawater. By comparison, 39% and 54% re­mained at the end of comparable periods inbrackish water and freshwater. Under dryconditions, 65% remained. Higher decompo­sition rates in sea water were related toincreased acti vity of shredder organisms,such as crabs and amphi pods.

Heald (1969) and Odum (1970) alsofound inc rea ses in nit rogen, protei n, andcaloric content as mangrove leaves pro­gressi vely decayed. The ni trogen contentof leaves decaying under brackish condi­tions (on an AFDI' basis) increased from1.5% (5.6% protei n) to 3.3% (20.6%protein) over a 6-month period. Subse­quent information (Odum et a1. 1979b)suggested that the protein increase maynot have been this great since some of thenitrogen increase probably included non­protein nitrogen compounds such as aminosugars. Fell and Master (1973), Fell eta1. (1980), Fell and Newell (1980), andFell et a1. (1980) have provided moredetailed information on red mangrove leafdecomposition, the role of fungi in decom­position (see section 4), and nitrogenchanges and nitrogen immobilization duringdecomposition. Fell et a1. (1980)have shown that as much as 50% of wei ghtloss of the leaf during decomposition isin the form of dissolved organic matter(DOM) •

Heal d et al. (1979), Lugo et al.(1980) and Twilley (1980) discovered thatbl ack mangrove leaves decompose more ra­pidly than red mangrove leaves and ap­parently produce a higher percentage ofDOM. Pool et a1. (1975) have shown thatmangrove litter decomposes and is exportedmost rapidly from frequently floodedriverine and overwash forests. Thesecommunities have little accumulation oflitter on the forest floor. Communitieswhich are not as well-flushed by thetides, such as the basin and hammock

34

forests, have slower rates of decomposi­t i on and lower export rates.

3.5 CAR80N EXPORT

Research from Florida mangrove swampsforms a small portion of the larger con­troversy concerned with the extent towhi ch coastal wetl ands export parti cul ateorganic carbon (reviewed by Odum et a1.1979a). Available evidence from Florida,Puerto Rico and Austral ia (Table 4) sug­gests that mangrove swamps tend to be netexporters. The values in Table 4 shouldbe regarded as preliminary, however, sinceall five studies are based upon simplisticassumptions and methodology.

Goll ey et al. (1962) based thei rannual estimate of particulate carbonexport from a Puerto Rican forest upon afew weeks of measurements. Odum andHeald's estimates were derived from two orthree measurements a month. All i nvesti­gators have ignored the importance of bedload transport and the impact of extremeevents. All investi gators except Lugo eta1. (1980) have failed to measure DOCfl ux.

It seems relatively clear that man­grove forests do export organic carbon tonearby bodies of water. The magnitude ofthis export has probably been underesti­mated due to ignoring bedload, extremeevent s, and DOC.

The value of this carbon input tosecondary consumers in receiving waters isnot clear. As shown in section 3.6, foodwebs based primarily upon mangrove carbondo exist. The relative importance ofmangrove carbon to Florida coastal ecosys­tems remains speculative. We suspect thatmangrove-based food webs are dominant insmall bays, creeks and rivers within largemangrove ecosystems such as the NorthRiver system studied by Heald (1969) andOdum (1970). In i ntermedi ate-si zed bodi esof water, such as Rookery Bay near Naples,Florida, mangroves are probably importantbut not dominant sources of organic car­bon. Lugo et a1. (1980) estimate thatmangroves supply 32% of the organic carbon

Table 4. Estimates of particulate carbon export from mangroveforests. LU90 et a1. (1976) estimated export from a theoreti­cal, 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 tra1i a 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 sys­tems, such as Biscayne Bay near Miami,Florida, mangroves are clearly less impor­tant than any other sources such as a1 gaeand sea grasses, although mangrove carbonmay be important in localized situationssuch as the immediate vicinity of fringingand overwash forests. The magnitude ofmangrove carbon export to unenclosedcoastal waters and offshore remai ns amystery.

3.6 ENERGY FLOW

At least seven sources of organiccarbon may serve as energy inputs forconsumers in mangrove ecosystems (Figure8). The pathways by which this energycontaining material is processed and madeavailable to each consumer species isindeed complex. Not surprisingly, currentunderstanding of energy flow in Floridamangrove ecosystems exists largely in aqualitative sense; quantitative data arescarce and piecemeal. A variety of inves­tigators have contributed information overthe past decade including, but not limitedto, Heald (1969), Odum (1970), Odum andHeald (1972), Carter et a1. (1973),Snedaker and Lugo (1973), Heald et al.(1 974), Lugo and Snedaker (1974, 1975),Odum and Heald (1975a, b), and Pool et al.(1977). Probably, the most complete studyto date is the investigation of energyflow in the black mangrove zone of RookeryBay by Lugo et al. (1980).

It is possible at this time to pre­sent a series of hypotheses concerning therelative importance of these energysources. Fi rst, the rel ati ve importanceof each source can vary from one locati onto the next. As wi 11 be shown in thefollowing discussion, the consumers incertain mangrove forests appear to dependprimarily upon mangrove-derived carbonwhile in other locations inputs from phy­toplankton and attached algae are probablymore important.

Our second hypothesis is that energyflow based upon phytoplankton is mostimportant in overwash mangrove forests andother locations associated with large

bodies of clear, relatively deep water.Conversely, phytoplankton are hypothesizedto be rel ati vely uni mportant to the energybudgets of the 1arge ri veri ne forest com­munities along the southwest coast ofFlorida. It should be remembered, how­ever, that even where phytoplankton arequantitatively unimportant, they poten­tially perform an important function asthe basis of phytoplankton-zooplankton­1arva1 fi sh food webs (Odum 1970).

As a third hypothesis, Iver Brook(Rosensteil School of Marine and Atmos­pheric Sciences, Rickenbacker Causeway,Miami, Fla.; personal communication 1979)has suggested that both sea grasses andbenthic algae serve as an important energysource for fringing mangrove communiti esadjacent to large bodies of water such asBi scayne Bay and Whitewater Bay. Althoughlittle evidence exists to test this hypo­thesis, observations of extensive depositsof sea grass and macroalgal detritus with­in mangrove forests suggest intuitivelythat Brook's hypothesi s may be correct.

In regions where mangrove shading ofthe prop roots is not severe, our fourthhypothesis suggests that carbon origina­t i ng from prop root epi phytes may be si g­nificant to community energy budgets.Lugo et a1. (1975) have measured net pro­duction of periphyton in mangrovesfringing Rookery B~ and found averageval ues of 1.1 gC/m /day. Hoffman andDawe~ (1980) found a lower val ue of 0.14gC/m /day. Because these val ues areroughly comparable to average exports ofmangrove leaf carbon (section 3.5), itspotential importance is obvious.

The fi fth hypothesi s states thatmangrove organic matter, particularly leafmateri a1, is an important energy sourcefor aquatic consumers. This hypothesiswas first espoused by Heald (1969) andOdum (1970), who worked together in theriverine mangrove communities between theEvergl ades and Whitewater Bay. Clearly,mangrove carbon is of great importancewithin the riverine and basin communitiesa 11 along the southwest coast of Flori da(Odum and Heald 1975b); Carter et al.(1973) and Snedaker and Lugo (1973)

36

SUN _

reduced sul fur

ATMOSPHERI CSOURCES

SULFUR OXIDIZINGBACTERIA

TERRESTRIALSOURCES

MANGROVESSEA GRASSESEPIPHYTICALGAE

01 kECT GRAZING~~~~;D~I;ssjO~L;V~ED~~~~~~piA;RTiI~ciUL~ATEORGANIC MATTER ORGANIC MATTER

"" "", ?v MICROBES

", " /-HI GHERCONSUMERS

PHYTOPLANKTON

Figure 8. Potential pathways of energy flow in mangrove ecosystems. Not all possible pathwayshave been drawn; for example. methanogenesis and sulfur reduction could originate from any ofthe sources of organic matter. ~~ngrove-based pathways are enhanced for emphasis and in no wayimply relative importance.

provided subsequent supportive data. Whatis not clear, is the relative importanceof mangrove carbon to consumers withinfringing, overwash, and more isolatedmanqrove communities.

Our si xth hypothesi s invol ves theassemblage of organisms that graze man­grove leaves di rect1y. A variety of in­sects (see section 6) and the mangrovetree crab, Aratus pi soni i, (Reever et al.1979) obtain much of their energy directlyfrom livin9 mangrove leaves, even though9razing rarely exceeds 10% of net primaryproduction (Odum and Heald 1975b).

As a seventh hypothesis we suggestthat anaerobic decomposition of mangrovetissue, particularly root material, maysupport an extensive food web based onbacteria associated with methanogenesis orthe processing of reduced sulfur com­pounds. Our suggestion of the importanceof reduced sul fur comes di rectly fromHowarth and Teal's (1980) discovery ofthis potentially important energy pathwayin temperate Spartina (cordgrass) marshes.They found that anaerobic decomposition issuch an incomplete process that if sul­fates are available (from sea water) asmuch as 75% of the original energy inplant tissues may be converted by sulfurreducing bacteria to reduced sul fur com­pounds such as hydrogen sulfide and Py­rite. Subsequently, if these reducedsulfur compounds are moved hydrologicallyto an oxidized environment (sediment sur­face or creek bank) sulfur -oxidi zi ng bac­te ri a (e.g., Thi ohac ill us sPp.) may conve rtthe chemically-stored energy to bacterial­ly stored energy with an efficiency asgreat as 50% (Payne 1970). Presumably,deposit-feeding organisms such as grassshrimp (Palaemonetes) and mullet (Mugil)are capable--or-grazi ng these su"""if'Ur­oxidizing bacteria from the sedimentsurface. If this hypothetical trophicexchange does exi st, it may be of con­siderable magnitude and may cause us toreexamine current concepts of energy pro­cessing and export from mangroveecosystems. Since freshwater containsremarkably little sulfate in comparison toseawater, this energy pathway is probablyof little importance in mangrove forests

of very low salinity.

Carbon inputs from terrestrialsources may be important to certain man­grove communities. Carter et al. (1973)have shown that terrestrial carbon canreach coastal ecosystems particularlywhere man has cut deep channels inland fornavi gat i on or drai nage purposes. Themagnitude of this influx has not beenadequately measured although Carter et al.did find that mainland forests (includingmangroves) contributed approximately 2,100metri c tons of carbon per year toFahkahatchee Bay.

Atmospheric inputs from rainfallappear to be minimal in all cases. Lugoet al. (1980) measured throughfall (preci­pitation passing through the tree canopy)in Rookzry Bay mangrove forests of 15 to17 gC/m /year. This would be an overesti­mate of atmospheric input since it con­tains carbon leached from mangrove leaves.The best guess of atm~spheric input isbetween 3 to 5 gC/m /year for southFlorida mangrove ecosystems.

Subsequent stages of energy transferin mangrove community food webs remainlargely hypothetical. Odum (1970) andOdum and Heald (1975b) have outlinedseveral pathways whereby mangrove carbonand energy are processed by a variety oforganisms (see Figure 8). Apparently, themost important pathway follows the se­quence: mangrove-leaf detritus substrate­microbe-detritus consumer-higher consu­mers. The critical links are provided bythe microbes such as bacteria and fungi(see Fell et a1. 1975) and by the detritusconsumers. The latter group was studiedby Odum (1970) and Odum and Heald (1975b)and found to consist of a variety ofinvertebrates (e.g., caridean shrimp,crabs, mollusks, insect larvae, amphipods)and a few fi shes.

Stable carbon studies such as thosedone by Haines (1976) in Spartina(cordgrass) marshes have not been per­formed in mangrove ecosystems. Mangrovesare C3 plants and have 6 13 values in therange of minus 25 to minus 26 (Macko1981). According to the same author,

38

mangrove peat has a 6 13 value of minus22. Because these values are dramaticallydi fferent from the values for sea grassesand many algae, the possibilities forusing this tool in mangrove ecosystems isexcellent. Macko (1981) also suggestedthe utility of using stable nitrogen ra­tios for future mangrove food web investi­gations; he reported 6 15 values of plus6.0 to p1 us 6.5 for mangrove ti ssue andplus 5 for mangrove peat.

In reviewing contemporary knowledgeof energy flow in mangrove ecosystems,th ree conc 1us ions emerge.

(1) We have a hypothetical frameworkof mangrove energy flow of a qual i tati ve

nature. This framework appears to bereasonably accurate although subsequentdevelopments, such as elucidation of thereduced sul fur hypothesi s, may requi resome modification.

(2) Measurements of the relativeimportance of vari OUS ca rbon sou rces aregenerally lacking.

(3) Oetai 1ed measurements of energyflow including the relative inputs ofdifferent carhon sources are criticallyneeded. Technological diffi cult i es, hi ghcosts, and difficulties inherent intransferring findings from one estuary tothe next present a major challenge toestuarine ecologists of the future.

39

CHAPTE R 4. COMMUNITY COMPONENTS - MICROORGANISMS

The mycof10ra (fungi) are the beststudied component of the microbial com­munity of mangrove swamps. t1uch pio­neering work has been carried out in southFlorida. Reviews of the current knowledgeof mangrove -associ ated fungi can be foundin Kohlmeyer and Kohlmeyer (1979) and Fellet a1. (1 980 )•

One of the earliest studies of man­grove mycoflora was published by Kohlmeyer(1969). He discovered large populationsof marine funqi on the submerged parts ofaerial roots, stems, and branches and onliving and dead man9rove leaves. Exten­sive work at the University of Miami byFell and his coworkers (e.g., Fell andMaster 1973; Fell et a1. 1975, 1980) ex­plored the role of fungi in the decom­position of mangrove leaves and the im­mobilization of nitr0gen. Newell (1974)studied the succession of mycoflora onseedlings of red man9rove. A survey ofthe aquatic yeasts occurring in the southFlorida mangrove zone was published byAhearn et a1. (1968).

One of the most interestin9 pieces ofinformation to emerge from this extensivemycof10ra research concerns the successionof orqanisms associated with decaying1eaves (summari zed by Fell et a1. 1975,1980). Senescent leaves of red mangrovesare t.vpically colonized by species ofNigrospora, Phy110stica, and Pesta10tica.Once the leaf has fallen from the tree andduri n9 the early stages of decay, thefungal fl ora is domi nated by speci es of~~l.tophthora and, to a lesser extent,

Orechslera and Gloeosporium. In the lat­ter stages of decay the dominant generaare Calso, G1iocidium, and Lulworthia.

Understanding the occurrence and suc­cessi on of fun9i on decayi ng mangroveleaves is important because of their rolein energy flow in mangrove swamps. Heald(1969), Odum (1970) and Odum and Heald(1975b) hypothesi zed that fun9i and bac­teria are important in convertin9 mangrove1eaf organi c materi ali nto a form that canbe digested and assimilated by detriti­vores (see section 3.6).

Our understandi ng of the role andoccurrence of bacteria in man9rove swampsis not as well documented as for fungi.Casagrande and Given (1975) have suggestedthat bacteri a are important in the earlystages of mangrove leaf decomposition andare replaced in the latter stages by funqiwhich are better equipped to attack re­fractive organic compounds. Unlike themycofl ora, the bacteri a are c1ea rly i mpor­tant in the anaerobic regions of mangroveswamps. Vankatesan and Ramamurthy (unpubl.data) found denitrifying bacteria to beabundant and ubiquitous in mangrove soils.Zuberer and Silver (1978) have emphasizedthe importance of nitrogen-fixing bacteriain the zone around mangrove roots. They,in fact, were able to isolate and count avariety of types of bacteria from mangrovesediments including aerobic heterotrophs,anaerobic heterotrophs, nitrogen-fixingheterotrophs, and sulfate-reducing bac­te ri a.

40

CHAPTER 5. COMMUNITY COMPONENTS - PLANTS OTHER THAN MANGROVES

5.1. ROOT AND MUD ALGAE

The aeri a1 root systems of mangrovesprovide a convenient substrate for at­tachment of al gae. These root al gal com­munities are particularly noticeable onred mangrove prop roots but also occur toa 1esser extent on bl ack mangrovepneumatophores located in the intertidalzone. Productivity of prop root algalcommunities can be appreciable if shadingby mangroves is not too severe; as dis­cussed in section 3.6, Lugo et al. (1975)found a prop root communitt' net primaryproduction rate of 1.1 gC/m /day, a levelcomparable to mangrove leaf fall. Biomassof these algae can be as high as 200 to3DD g per prop root (Burkholder andAlmodovar 1973). Of course, production ofthis magnitude only occurs on the edge ofthe forest and is virtually nil in thecenter of the swamp. Nevertheless, thisalgal carbon has considerable potentialfood value either to direct grazers ordetritivores.

Vertical distribution of prop rootalgae has been studied by many researchers(Gerlach 1958; Almodovar and Riebl 1962;Biebl 1962; Post 1963; Rutzler 1969;Burkholder and Almodovar 1973; Rehm 1974;Yoshioka 1975); only one of these studies(Rehm 1974) was conducted in Florida.There is a tendency for certain genera ofalgae to form a characteristic associationon mangrove roots around the world (Post1963). Four phyla tend to dominate:Chlorophyta, Cyanophyta, Phaeophyta, andRhodophyta; the last is usually the mostimportant in terms of biomass. Of 74species of marine algae recorded as proproot epi phytes between Tampa and KeyLargo, 38 were Rhodophyta, 29 Chlorophyta,4 Phaeophyta and 3 Cyanophyta (Rehm 1974).

Zonation to be expected on Floridaman9roves is shown in Figure g; this se­quence comes largely from Taylor (1960).Near the high water mark, a green bandusually exists which is dominated by spe­cies of Rhizoclonium. Below this is azone dominated by species of Bostrychia,Catenella, and Caloglossa. It is thisassociation that most people think of whenmangrove prop root algae are mentioned.

Because much mud is often deposited on theBostrychia-Catenella-Caloglossa complex,it often has a dingy, gray appearance.There are many other algae found in thiszone, but these three genera usually domi­nate. At brackish or nearly freshwaterlocations, they are replaced by species ofBatophora, Chaetomorpha, CladQ.phora, andPenicillus. The pneumatophores ofAVTCennTa-,- when colonized, are oftencovered with species of Rhizoclonium,Bostrychia and Monostroma (Taylor-T§61J1.Hoffman and Dawes (1980) found that theBostrychia binderi -dominated community onthe pneumatophores of black mangroles hada standing crop of 22 g drt' wt/m and anet production of 0.14 gC/m /day.

If there is a permanently submergedportion of the prop root, it may becovered with rich growths of Acanthophora,Spyrida, Hypnea, Laurencia, IIrangelia,~lonia, and Caulerpa (Almodovar and Biebl19~ Additional genera which may bepresent below mean high water are:Murrayella, POlysiphonia, Centroceras,~urd!~ann~, Dic~la, ~~ll~~~,Laurencia, and Dasya TTaylor 1960;Burkholder and Almodovar 1973; Yoshioka1975). In addition, anywhere on the moistsections of the prop roots there areusually epiphytic diatoms and filamentousgreen and blue-green algae of many genera.

Rehm (1974) found a significant dif­ference in the prop root algae betweensouth and central Florida. South of TampaBay the standard Bostrychia-Catenella­Caloglossa dominates. In the Tampa Bayarea, species of the orders Ulotrichalesand Cladophorales are dominant.

The mud adjacent to the mangrove rootcommunity is often richly populated with avariety of algae. These can includespeci es of Cl adophoropsi s, Enteromor ha,Vaucheria, and Boodleopsis Taylor 1960in addition to a whole host of benthicdiatoms and dinoflagellates (llood 1965)and other filamentous green and blue-greenalgae (r~arathe 1965).

Adjacent to mangrove areas, on thebottoms of shoals, shallow bays andcreeks, there is often a variety of

41

PLANTS ANIMALS

----.

Sphaeromaterebrans

Balanus eburneus

Brachidontes spp.

Narels spp.Bulla spp.

Ascldla nigerCrassos trea

virginica

L ittorina angulifera

Ligea exotica

;\-y.--:..=~- - - - - - -

Rhlzoc/onlum spp.

Acanthophora spp.Caulerpa spp.Wrangleia spp.

Bostrychla spp.Catanella spp.

C" ••,,,., •••. LMLW- - - - - =---=->.

MHW - - - - - ~=-='-=

Figure 9. Vertical distribution of selected algae and invertebrates on redmangrove prop roots (compiled from Taylor 1960 and our own observations).

42

tropical algae including species ofCaulerpa, Acetabularia, Penicillus,Grae i 1 a ria, ~ll'!'e d a, Sa r gas sum,Batophora, Udotea, and Dasya. These arediscussed at length by Zieman (in prep.).Other pertinent references for mangroveregions include Davis (1940), Taylor(1960), Tabb and Manning (1961), and Tabbet a1. (1962).

5.2 PHYTOPLANKTON

All aspects of phytoplankton, fromseasonal occurrence to productivitystudies, are poorly studied in mangroveecosystems. This is particularly true inFlorida.

Evidence from Brazil (Teixeira et al.1965,1967,1969; Tundisi 1969) indicatesthat phytoplankton can be an importantcomponent of the total primary productionin mangrove ecosystems; just how importantis not clear. Generally, standing cropsof net phytoplankton in mangrove areas arelow (personal observation). The nanno­plankton, which have not been studied atall, appear to be most important in termsof total metabolism (Tundisi 1969). Thenet plankton are usually dominated bydiatoms such as Thalassothrix spp.,Chaetoceras spp., Ni tzschi a spp.,Skel etonema spp., and RhlZosOTei1i a spp.(Mattox 1949; Wood 1965; Walsh 1967; Bacon1970). At times, blooms of dinoflagel­lates such as Peridinium spp. andG mnodinium spp. may dominate (personalobservation. In many locations, particu­larly in shallow waters with some turbu­lence, benthic diatoms such as Pleurosigmaspp., Mastogl oi a spp., and Di sp 1onei s maybe numerically important in the net plank­ton (Wood 1965).

Understanding the mangrove-associatedphytoplankton community is complicated bythe constant mixing of water masses inmangrove regions. Depending upon thelocation, the phytoplankton may be domi­nated by oceanic and neritic forms, bytrue estuarine plankton, and by freshwaterplankton. The pattern of dominance maychange daily or seasonally depending uponthe source of the principal water mass.

Before we can understand the impor­tance (or lack of importance) of phyto­plankton in mangrove regions, some ques­tions must be answered. How productiveare the nannoplankton? How does the dailyand seasonal shift in phytoplankton domi­nance affect communi ty producti vi ty? Doesthe generally low standing crop of phyto­pl ankton represent low productivity or ahigh qrazing rate?

5.3 ASSOCIATED VASCULAR PLANTS

Four species of aquatic grasses occuron bay and creek bottoms adj acent to man­grove forests. Turtle grass, Thalassiatestudinum, and manatee grass, Syringodiumfilliforme, are two tropical sea grasseswhich occur in waters with average salini­ties above about 20 ppt. Shoal grass,Halodule wrightii, is found at somewhatlower salinities and widgeongrass, ~uppia

maritima, is a freshwater grass which cantolerate low salinities. These grassesoccur throughout south Florida, often inclose juxtaposition to mangroves. Zieman(in prep~ presents a thorough review ofsea grasses along with comments aboutpossible energy flow linkages withmangrove ecosystems.

There are extensive areas of man­groves in south Florida which are closelyassociated with marshes dominated bv avariety of other salt-tolerant plants.For example, along the southwest coastbetween Flamingo and Naples, marshes arescattered throughout the mangrove belt andalso border the mangroves on the uplandside. The estuarine marshes within themangrove s"amps have been extensivelydescribed by Egler (1952), Carter et a1.(1973), and Olmstead et a1. (1981). Theycontain various salt-tolerant marshspecies including: salt grass, Distichlisspicata, black needle rush, Juncusroemerianus, spike rush, Eleochariscellulosa, 91aSS wort, Salicornia spp.,Gulf cordgrass, Spartina spartinae, seapurslane, Sesuvium portulacastrum, saltwort, Batis maritima, and sea ox-eye,Borric~frutescens. Farther north,above Tampa on the west coast of Flori da,marshes populated by smooth cordgrass,

43

Spartina alterniflora, and black needlerush, Juncus roemerianus, become moreextens i ve and eventua 11 y replace mangroveswamps. Even in the Everglades region,the saline marshes are comparahle to man­groves in areal extent, although theytend to be some distance from open water.Studies of these marshes, including as­sessment of their ecological value, arealmost non-existent. Certainly, they haveconsiderable importance as habitat forsmall fishes which, in turn, support manyof the nesti ng wadi ng bi rds in southFlorida (see section g).

Tropical hardwood forests may occurwithin the mangrove zone in south Florida,partIcularly where old shorelines or areasof storm sedimentation have created ridges1 m or more above MSL (mean sea level)(Olmstead et al. 1981). Similar forestsor "hammocks" occur to the rear of themangrove zone on hi gher ground. Typicaltrees in both forest types include the fanpal m, Thrinax radiata, buttonwood,Conocarpus erecta, manchineel, Hippo~an~mancinella, and, in the past, mahogany,SWTetenia mahagoni. Olmstead et al.lT981Tprovide-a descri pt i on of thesecommunities.

Freshwater marsh pl ants, such as thegrasses, rushes and sedges that dominatethe freshwater Evergl ades, are notmentioned here, although they areoccasionally mixed in with small mangroves

that have become established well inland.See Hofstetter (1974) for a review ofliterature dealing with these plants.

Finally, a group of somewhat salt­tolerant herbaceous plants is foundwithin stands of mangroves. They usuallyoccur where slight increases in elevationexist and where sufficient light filtersthrough the mangrove canopy. Carter etal. (1973) list the following as examplesof members of the mangrove community:1eather ferns, Acrostichum aureum and A.danaeifolium; spanish bayonet, Yuccaaloifolia; spider lily, HymenociillTSlatifolia; sea blite, Suaeda linearis;chaff flower, Alternanthera ramosissima;samphire, Philoxerus vermicularis; blood­leaf, Iresine celosia; pricklypear cactus,Opunt.,--a--srricta; marsh elder, Ivafrutescens; the rubber vine, Rhabdadeniabi fl ora; the 1i anas, Ipomoea tuba andHippocratea volubilis; and a variety ofbrornel i adS\Bromel i aceae).

Although the lists of vascular plantswhich occur in mangrove swamps may seemextensi ve, the actual number of speci es inany given location tends to be lowcompared to totally freshwater envi ron­ments (see Carlton 1977). Analogous totemperate salt marshes, mangrove swampspossess too many sources of stress,particularly from tidal salt water, tohave a high diversity of vascular plantspecies.

44

CHAPTER 6. COMMUNITY COMPONENTS - INVERTEBRATES

6.1 ECOLOGICAL RELATIONSHIPS

The mangrove ecosystem, with its treecanopies, masses of aerial roots, muddysubstrates, and associated creeks andsmall embayments, offers many habitatopportunities for a wide variety of inver­tebrates. Whil e there are few compari sonsof species richness with other types ofcoastal ecosystems, mangrove swamps appearto be characterized by moderately highinvertebrate species diversity. Abele(1 974) compared H' (Shannon Weaver) di ver­sity of decapod crustaceans betweenvarious littoral marine communities andfound mangrove swamps in an intermediateposition with more decapod species thanSpartina marshes but considerably lessthan were associ ated with rocky substratecommunities.

There is little doubt that the mazeof prop roots and muddy substrates underi ntert i dal mangrove trees provi des habitatfor a wi de range of invertebrates andfishes (Figure 10) (see section 7 for thelatter). The nursery value of the proproot complex for juvenile spiny lobsters,Panu1irus argus, is well established(Olsen et ar:-l"975; Olsen and Koblic 1975;Little 1977; Witham et al. 1968). Ac­cording to these researchers, the phyl­losome larvae of spiny lobsters oftensettle among the prop roots and remainthere for much of their juvenile lives.The prop roots provide protection frompredators and a possible source of food inthe associated populations of small inver­tebrates. To provide the best habitat, asection of the prop roots should extendbelow mean low tide. If conditions aresuitable, the juveniles may remain inclose association with the prop root com­muni ty for as much as 2 years unti 1 theyreach a carapace length of 60 to 70 mm.

In addition to its value as spinylobster habitat, mangrove ecosystems alsoharbor the following invertebrates: bar­nacles, sponges, polychaete worms, gastro­pod mollusks, pelecypod mollusks, isopods,amphipods, mysids, crabs, caridean shrimp,penaeid shrimp, harpacticoid copepods,snapping shrimp, ostracods, coelenterates,nematodes, a wide variety of insects,

bryozoans, and tunicates. The most ob­vious and dominant organisms are usuallybarnacles, crabs, oysters, mussels, ;50­pods, polychaetes, gastropods and, tuni­cates.

A striking characteristic of mostmangrove swamps is the pattern of horizon­tal and vertical zonation of invertehrates(Fi gure 9). Characteri stic verti cal zona­tion patterns are found on the prop roots(Rutzler 1969) and not so obvious horizon­tal distributions occur as you move backinto the center of the swamp (Warner1969). Invertebrate biomass in the redmangrove zone on the edge of the swamp maybe ~ery high, often in excess of 100 dryg/m of organic matter in many locations(persona1 observat ion). In the center ofthe swamp, particularly where there islittle flooding, biomass is usually anorder of magnitude less; Golley et

2al.

(1 962) found an average of 6.4 g/m ofinvertebrates in the center of a PuertoRican mangrove swamp.

Mangrove-associated invertebrates canbe placed in four major categories basedon trophic position:

(1) di rect grazers - 1i mited to

(a) insects and the mangrove treecrab, Aratus pisonii, all of which feed onleaves in the mangrove canopy and

(b) a group of small invertebrateswhich graze the prop root and mud algaedi rect1y;

(2) filter feeders - largely sessileprop root invertebrates whi ch fi Her phy­toplankton and detritus from the water;

(3) deposit feeders - mobile inverte­brates which ski m detritus, a1 gae andoccasional small animals from the surfaceof the mud and forest floor;

(4) carnivores - highly mobile inverte­brates which feed upon the three precedinggroups in all locations from the treecanopy (largely insects) to the mud sur­face. Food sources in man9rove swamps andener9Y flow are di scussed insect ion 3.6.

45

Figure 10. Photograph of red mangrove prop root habitat in clear shallow water with associated animaland plant populations. Photograph is by Bianca Lavies (copyright, National Geographic Society).

6.2. ARBOREAL ARTHROPOD COMMUNITY

A surprising variety of arthropodsinhabit the mangrove canopy. Because theyare frequently secretive or possesscamouflage coloration, their numericalimportance often has been overlooked.Beever et al. (1 979) poi nted out thatarboreal arthropods have a variety ofecological roles: (1) direct herbivory onmangrove 1eaves, (2) predator-prey i nter­actions, and (3) biomass export throughfrass production and leaf defoliation.Oirect grazing is typically patchy indistribution. It is not unusual to findextensive stretches of mangroves that havescarcely been grazed. In nearby areas, asmuch as 80% of the leaves may have somedamage (Beever et al. 1979). As a generalrule, it is probably safe to state thathealthy, unstressed mangrove stands nor­mally have less than 10% of their totalleaf area grazed (Heald 1969). In manylocations, percent leaf area damaged is onthe order of I% to 2% (Beever et al.1979). There are excepti ons. Onuf et a1.(1977) reported biomass loss to arthropodgrazers as high as 26% in a mangrove standwhere growth and nitrogen content of theleaves had been enhanced by input of nu­t ri ents from a bi rd rookery.

In terms of numbers of species, thedominant group of arboreal arthropods isinsects. The most thorough inventory ofmangrove-associated insects was conductedby Simberloff and Wilson to obtain the rawdata for their papers on island bio­geography (Simberloff and Wilson 1969;Si mber10ff 1976). These papers 1ist over200 species of insects associated withoverwash mangrove islands in the FloridaKeys. There is no reason to expect lessernumbers in other types of mangrove com­munities, except for the mangrove scrubforests. The most thorough study of i n­sect grazing on mangrove leaves is that ofOnuf et al. (1977) (see section 2.6).

Although not as numerically impres­sive as the insects, the man9rove treecrab, Aratus pisonii, appears to be poten­tially as important in terms of 9razingimpact (Beever et al. 1979). The lifehistory of this secretive little crab has

47

been described by Warner (1967). 12Jamaica its numbers range from 11 to l~/m

at the edge of fri ngi ng sl<amps to 6/m inthe center of large swamps. Beever et al.(1979) reported typical densities for avariety o~ sites in south Florida of 1 to4 crabs/m. These same authors reportedsome interestin9 details about the crab:(1) the diet is omnivorous rangin9 fromfresh mangrove leaves to caterpillars,beetles, and various insects; (2) the crabsuffers highest predation pressure whilein the planktonic larval stage; (3) preda­tion on the crabs while in the arhorealcommunity is low and comes from bi rds suchas the white ibis, raccoons, other man­grove tree crabs and, if the crabs fall inthe water, fi shes such as the mangrovesnapper; and (4) in one location in southFlorida (Pine Island Sound) they found inaccordance with normal biogeographicaltheory, the highest densities of crabsassociated with fringing forests and thelowest densities on distant islands, butat Sugar Loaf Key the unexplainablereverse distribution was found.

Other invertebrates may visit thecanopy from below either for purposes offeeding or for protection from high tides.Included in this group are the pulmonategastropods, Li ttori~ angu1 i fera,Cerithidea scalariformis, and Me1ampuscoffeus, the isopod, Ligea~)(otica, and ahost of small crabs.

In summary, with the exception of ahal f dozen key papers, the arboreal man­grove community has been generally ig­nored. Both insects and the mangrove treecrab play significant ecological roles andmay affect mangrove producti vity to agreater extent than has heen recognized.

6.3 PROP ROOT AND ASSOC IATED MUD SURFACECOMMUNITY

These two somewhat di sti nct com­munities have been lumped together becauseof the large number of mobile organismswhich move back and forth between tidalcycles. The aerial roots are used asprotecti ve habitat and to some extent forfeeding while the nearby mud substratesare used principally for feeding.

The prop roots support an abundanceof sessile organisms. The verticalzonation of both mobile and sessile inver­tebrates has been studied extensively inother parts of the world (Good body 1961;Macnae 1968; Rutz1er 1969; Coomans 1969;Bacon 1970; Ko1ehmainen 1973; Sasekumar1974; Yoshioka 1975). Vertical zonationcertainly exists on Florida red mangroveroots. The general i zed scheme shown inFiqure 9 essentially contains two zones:an upper zone dominanted by barnacles anda lower zone dominated by mussels, oystersand ascidians. Between mean high tide andmean tide, the wood boring isopod,~haeroma terebrans (discussed at lengthin section 2.7) is important, both numeri­cally and through the provision ofnumerous holes for use by other organisms(Estevez 1978).

The most compl ete study of theFlorida mangrove prop root community isCourtney's (1975) comparison of seawalland mangrove associations. He reported anextensive list of invertebrates from man­grove prop roots at Marco Island, Florida,including: Crassostrea virginica,Litt:.~r:!..'!.~ angulifera, Crepidula ~,Jiodora cayenensis, Urosalpinx perrugata,Pisania tincta, Brachidontes exustus,nine species of polychaetes, ~erom~terebrans, Palaemon floridanus,PeMelT ,!!~es longieaudatu~-Synarpheusfritzmuel1eri, Thor f10ridanus,pet"rofisthesarmatu.;;-a;ld .itleast ei ghtspecies of crabs. The following specieswere found only on mangrove roots and noton seawalls: Turitel1a sp., Me10ngenacoro~, Anachis semip1icata, Bullastriata, Hypselodoris sp., Area imbricata,Carditamera fl ori dana, Pseudoi rus !l.P.!..ca,and Martesia striata.

Tabb et a1. (1962) and Odum and Heald(1972) reported a variety of invertebratesassociated with prop roots in the Hhite­water Bay region. Although many speciescoincide with Courtney's (1975) list,there are also significant differences dueto the lower salinities in this region.It is probably safe to conclude that proproot commun it i es vary somewhat from siteto site in response to a number of factors

48

including latitude, salinity, and proxi­mi ty to other communi ties such as seagrass beds and coral reefs.

Sutherland (1980), working on redmangrove prop root communities inVenezuela, found little change in theinvertebrate species composition on indi­vidual prop roots during an 18-monthperiod. The species composition variedgreatly, however, between adjacent proproots, presumably in response to stochas­tic (chance) processes.

The mud flats adjacent to mangrovesprovide feeding areas for a range of in­vertebrates that scuttle, crawl, and swimout from the cover of the mangrove roots.Some emerge at low tide and feed on algae,detritus, and small invertebrates on themud f1 ats whi 1e they are hi gh and dry.Others emerge while the tide is in, parti­cularly at night, and forage across theflooded flats in search of the same foodsplus other invertebrates which haveemerged from the mud. In many ways themangrove-mud flat relationship is analo­gous to the coral reef (refuge) sea grass(feeding area) relationship reviewed byZieman (in prep.). The net effect is thatthe impact of the mangrove community mayextend some distance beyond the boundariesof the mangrove forest.

In addition to the organisms whichmove from the mangroves to the mud flats,there is a small qroup which uses thesubstrate adjacent to mangroves for bothhabitat and feeding. In the WhitewaterBay region, four crabs exploit the inter­tidal muds from the safety of burrows:Uca pugi1ator, Q. speciosa, Q. thayeri,and Eurytium limosum (Tahb et a1. f962).In low sal inity mangrove forests of southFlorida, the crayfish, Procambarus alleni,is a dominant member of the burrowing,benthic community (Hobbs 1942) as is thecrab, Rhithropanopeus harrisii (Odum andHeald 1972). Both organisms are found ina remarkable number of fi sh stomachs.

The benthic fauna and infauna ofcreek and bay bottoms near mangroveforests are highly variable from one

location to the next. Many of theseorganisms, particularly the deposit andfilter feeders, benefit from particulateorganic matter originating from mangrovelitter fall (Odum and Heald 1972, 1975b).Tabb and Manning (1961) and Tabb et a1.(1962) present lists and discussions ofmany of the benthic invertebrates adjacentto mangrove areas of Whitewater Bay.Weinstein et al. (1977) compared the ben­thic fauna of a mangrove-lined creek and anearby man-made canal on Marco Island.They found (1) the mangrove fauna to bemore di verse than the canal fauna and (2)a higher diversity of organisms at themouths of mangrove creeks than in the"heads" or upstream ends. Courtney (1975)found the same pattern of upstreamdecreases in diversity, presumably inresponse to decreasing oxygen concentra­tions and increasingly finer sediments.

Finally, the irregularly flooded sub­strates in the center of mangrove forestscontain a small but interesting assemblageof invertebrates. The litter layer,composed largely of mangrove leaves, evi­dently includes a variety of nematodes.Due to the usual taxonomic difficulties inidentifying nematodes, complete speciesI ists do not exist for mangrove forests;however, many species and individuals areassociated with the decaying leaves(Hopper et al. 1973). In addition tonematodes, the wetter sections of theswamp floor can contain mosquito and otherinsect larvae, po1ychaetes, harpacticoidcopepods, i sopods, and amphi pods.Simberloff (1976) lists 16 species ofinsects associated with the muddy floor ofmangrove forests. Roaming across theforest floor during low tide are severalcrustaceans including the mangrove treecrab, Aratus pisonii, crabs of the genusSesarma, and the pulmonate gastropods,~~l~~£~~ coeffeus and Cerithideasca1ariformis. Both snails clearly havethe ability to graze and consume recentlyfallen leaves (personal observation).With favorable conditions (relatively fre­quent tidal inundation plus the presenceof red mangroves) Me1ampus populations can

49

exceed 500/m 2 and average 100 to 200/m 2

(Heald, unpublished data). Cerithidea isfound largely in association with blackmangroves ~d can reach densities of at1east 400/m •

6.4 WATER COLUMN COMMUNITY

This section is embarrassingly short;the reasons for thi s brevity are (I) thepaucity of research on zoopl ankton inFlorida mangrove-dominated areas and (2)our i nabi 1i ty to ·di scover some of the workwhich undoubtedly has been done. Davisand WilliaMS (1950) are usually quoted asthe primary reference on Florida mangrove­associated zooplankton, but their paperonly lists zoop1ankters collected in twoareas. Zooplankton near mangroves areprobably no di fferent from those found inother shallow, inshore areas in southFlorida. Based on Davis and Williams(1950) and Reeve (1964), we can hypothe­size that the community is dominated bycopepod species of genus Acartia, particu­larly Acartia tonsa. In addition, wecould expect a few other calanoid cope­pods, arrow worms (Sagitta spp.), manyfish, polychaete and crustacean larvae andeggs. Another component of the "pl ankton~'particularly at night, are benthicamphipods, mysids, and isopods which leavethe bottom to feed (personal observati on).

Plankton are not the only inverte­brates in the water column. Swimmingcrabs, such as the blue crab, Callinectessapidus, are plentiful in most estuarinemangrove regi ons of south Flori da. Otherswimming crustaceans include the carideanshrimp (Pa1aemonetes spp. and Peri­c 1 ; menes-spp-:T, -the-snappi n g s h r; mprAlphe~ spp.), and the penaeid shrimp(Penaeus spp). All of these swimmingcrustaceans spend considerable time on orin the benthos and around mangrove proproots. From the economic point of view,the pi nk shri mp, Penaeus duorarum, isprobably the most important species asso­ciated with mangrove areas (see discussionin section 11).

CHAPTER 7. COMMUNITY COMPONENTS - FISHES

Of the si x mangrove community typesdiscussed in section 1.5, fishes are animportant component of four: (1) basinforests, (2) riverine forests, (3) fringeforests, and (4) overwash island forests.For convenience we have divided fringeforests into two sub-components: (a)forests which fringe estuarine bays andlagoons and (b) forests which fringeoceanic bays and lagoons. This divisionis necessa ry because the fi sh communit i esdi ffer markedl y.

Mangroves serve two distinct rolesfor fishes and it is conceptually impor­tant to distinguish between them. First,the mangrove-water interface, generallyred mangrove prop roots, afford a rela­tively protected habitat which is particu­larly suitable for juvenile fishes.Secondly, mangrove leaves, as discussed insection 3.6, are the basic energy sourceof a detritus-based food web on which manyfi shes are dependent. The habitat val ueof mangroves can be considered strictly afunction of the area of interface betweenthe water and the mangrove prop roots; itis an attribute shared by all four typesof mangrove communities. The importanceof the mangrove detritus-based food web isdependent on the re 1at i ve contri but i on ofother forms of energy in a given environ­ment, including phytoplankton, benthicalgae, sea grass detritus, and terrestrialcarbon sources. Figure 11 provides adi agrammat i c representation of the rela­tive positions along a food web continuumof the four mangrove communities.

Fishes recorded from mangrove habi­tats in south Florida are listed in Appen­dix B. Although the fish communities arediscussed separately below, they have beencombined into certain categories in Appen­dix B; fishes from mangrove basins andriverine forests have been combined underthe heading of tidal streams; fishes fromfringing forests along estuarine bays andlagoons are listed under the heading ofestuarine bays; fishes from oceanic baysand lagoons have been listed under oceanicbays. Si nce no surveys have beenpublished specifically relating to over­wash island forests, there is no 1 istingfor this community type in Appendix B.

Site characteristics and sampling methodsfor these community types are summarizedin Appendix A. Nomenclature and taxonomicorder follow Bailey et a!. (1970).

7.1 BASIN MANGROVE FORESTS

The infrequently flooded pools in theblack mangrove-dominated zone provide anextreme habitat which few species offi shes can tolerate. The waters aredarkly stained with organic acids andtannins leached from the thick layer ofleaf litter. Dissolved oxygen isfrequently low (1-2 ppm) and hydrogensulfide is released from the sedimentsfollowing physical disturbance. Salini­ties are highly variable ranging fromtotally fresh to hypersaline. The fishfamilies best adapted to this habitat arethe euryhaline cyprinodonts (killifishes)and the poeciliids (livebearers). Thekillifishes include Fundulus confluentus(Heald et al. 1974), Rivulus marmoratus(M. P. Weinstein, Va. Commonwealth Univ.,Richmond, Va.; personal communication19B1), Floridichthys carpio, andCyprinodon variegatus (Odum 1970). Thepoecillids include Poecilia latipinna(Odum 1970) and, the most common, Gambusiaaffinis (Heald et a!. 1974). While thespecies richness of fishes in this habitatis low, the densities of fish are oftenvery hi gh. Weinstein \fers. comm.) hasrecorded up to 3B fish/m •

A11 of these fi shes a re permanentresidents, completing their life cycles inthis habitat. They feed primarily onmosquito larvae and small crustaceans suchas amphipods which, in turn, feed on man­grove detritus and algae. These smallfi shes enter coastal food webs when theyare flushed into the main watercoursesduring high spring tides or followingseasonally heavy rains. Here they areeaten by numerous pi sci vorous fi shes i n­cluding snook, ladyfish, tarpon, gars, andmangrove snappers. The alternate energypathway for fi shes of the bl ad mangrovebasin wetlands occurs when the poolsshri nk duri ng dry weather, the fi shes areconcent rated into sma 11 er a rea s, and arefed-upon by various wading birds including

50

/f"":<0>;­21 ~'i,P(/ /:;__ ,

--.,- -'-'-"""":":

"

"

- ,,- - IJ

- '-,---:... ~--,7"::-' ,.

.,

"

------ .',-,'~....o---...<'_~,-_/

...'"..0

z u0 w~ '"...u9

u~0.....II:~ ...0 '"..w 0~ u

;<

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BLACK MANGROVEBASIN FOREST

RIVERINE FRINGINGCOMMUNITY

ESTUARINE BAYFRINGING COMMUNITY

OCEANIC BAYFRINGING COMMUNITY

OVERWASH ISLAND.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 notdrawn 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 = silverperch, 11 = pigfish, 12 = b1ackcheek tonguefish, 13 = scrawled cowfish, 14 = fringed pipefish, 15 = fringedfilefish, 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 eta1. 1974).

7.2 RIVERINE FORESTS

Tidal streams and rivers, fringedlargely by red mangroves, connect thefreshwater marshes of south Florida withthe shallow estuarine bays and lagoons(Figure 12). Few of these streams havebeen studied thoroughly. The exception isthe North River which flows into White­water Bay and was studied by Tabb (1966)and Odum (1970). Springer and Woodburn(1960) collected fishes in a bayou ortidal pass connecting Boca Ciega Bay andOld Tampa Bay. Carter et a1. (1973)reported on the fishes of two tidalstreams entering Fahkahatchee and FahkaUnion Bays. Nugent (1970) sampled fishesin two streams on the western shore ofBiscayne Bay. Characteristics of theseareas and sampling gear used by the inves­tigators are summarized in Appendix A.

These tidal streams and associatedriverine mangrove forests exhibit extremeseasonal variability in both physicalcharacteristics and fish community compo­sition. Salinity variations are directlyrelated to changes in the make-up of thefish assemblage. During the wet season(June - November), salinities fallthroughout the water courses and, at somelocations in certain heavy runoff years,become fresh a 11 of the way to the mouth(Odum 1970). Opportuni sti c freshwaterspecies, which are normally restricted tothe sawgrass and black needle rush marshesof the headwaters, invade the mangrovezone. These include the Florida gar,Lepisosteus Ql~!.lrhincus; severalcentrarchid sunfishes of the genus Lepomisand the largemouth bass, Micropterussa1moides; the freshwater catfishes,Ictalurus nata1is and Noturus gyri nus; andthe ki1lifishes normally consideredfreshwater inhabitants such as Lucaniagoodei and Rivulus marmoratus.

During the dry season (December toearly May) salinities rise as a result ofdecreased freshwater runoff and continuingevaporation. Marine species invade the

tidal streams primarily on feeding forays.Examples include the jewfish, E ine helusitajara, the stingrays (Dasyatidae , theneed1efishes (Be10nidae), the jacks(Carangidae), and the barracuda, Sphyraenabarracuda. Other seasonal movements offishes appear to be temperature related.Tabb and Manning (1961) documented move­ments of a number of species from shallowinshore waters to deeper water duringtimes of low temperature stress. Thelined sole, the hogchoker, the bigheadsearobin, and the striped mullet, forexample, are much less frequently caughtin winter in shallow inshore waters.

A third type of seasonality of fishpopulations in the tidal rivers is relatedto life cycles. Many of the fish whichutilize the tidal stream habitat do soonly as juveniles. Thus, there are peaksof abundance of these species followingoffshore spawning when larval or juvenileforms are recruited to the mangrove streamhabitat. In general, recruitment occursin the late spring or early summer fol­lowing late winter and spring spawningoffshore or in tidal passes (Reid 1954).Numerous species are involved in this lifecycle phenomenon including striped mullet,grey snapper, sheepshead, spotted seatrout, red drum, and silver perch.

The only estimate of fish standingcrop from ti dal stream habitats is that ofCarter et a1. (1973). They recorded 27species weighin~ 65,B91 g (wet w~.) froman area of 734 m or about 90 gfm. Thisis probably an overestimate since an un­known portion of the fish community hadmoved from the flooded lowlands to thestream on the ebb tide; sampling occurredat low tide in October. Nonetheless, thisis an indication of the high fish standingcrop which this mangrove-associated habi­tat can support. The number of speciesreported from individual tidal streamsannually ranges from 47 to 60 and thetotal from all tidal streams in southwestFlorida is 111 species (Appendix B).

The food webs in these riverine man­grove ecosystems appear to be predomi­nantly mangrove detritus-based, althoughthe Biscayne Bay stream studied by Nugent

52

Figure 12. Aerial photograph of the mangrove belt of southwest Florida nearWhitewater Bay. Note the complex system of pools and small creeks which connectwith the tidal river system.

53

(1970) may be an exception. The basiclink between the mangrove leaf and higherorder consumers is provided by micro­organisms (fungi, bacteria, Protozoa)which colonize the decaying leaf and con­vert them into a relatively rich proteinsource (Odum 1970; Odum and Heald 1975a).These decaying leaf fragments with asso­ciated microorganisms are fed upon by agroup of omnivorous detritivores includingamphipods, mysids, cumaceans, ostracods,chironomid larvae, harpacticoid andcalanoid copepods, snapping shrimp,caridean and penaeid shrimp, a variety ofcrabs, filter-feeding bivalves, and a fewspeci es of fi shes (Odum 1970; Odum andHeald 1972; Odum and Heald 1975b). Thesedetritivores, in turn, are consumed by anumber of small carnivorous fishes, whichin turn, are consumed by largerpiscivorous fishes. The concept of man­9rove trophic structure is a1 so discussedin section 3.6. See Appendix B forspecies specific dietary information.

The tidal creeks studied by Nugent(1970) on the western shore of BiscayneBay differ from the previously discussedstreams in the Everglades estuary. Themouths of the Biscayne Bay creeks havedense growths of sea grasses which con­tribute sea grass detritus. The salini­ties are considerably greater and thestreams are located only a few kilometersfrom coral reefs, whi ch are 1argely absenton Florida's west coast, at least close toshore. As a result, 23 species listed inAppendi x B were captured by Nugent (1 970)and are not recorded from riverine man­grove habitat on the west coast ofFlorida. Examples include several of thegrunts (Pomadasyidae), the gray trigger­fish, Ba1istes capriscus, the barbfish,Scorpaena brasiliensis, the scrawled box­fish, Lactophrys guadricornis, and thesnappers, Lutjanus apodus and I. synagri s.

Riverine mangrove communities andassociated tidal streams and rivers aretypified by the following families offishes: kil1ifishes (Cyprinodontidae),1ivebearers (Poeciliidae), si1versides(Atheri ni dae), moj arras (Gerreidae), tar­pon (Elopidae), snook (Centropomidae),snappers (Lutjanidae), sea catfishes

(Ariidae), gobies (Gobiidae), porgys(Sparidae), mullets (Mugi1idae), drums(Sciaenidae), and anchovies (Engrau1idae).The mangrove-lined streams and associatedpool s are important nursery areas forseveral marine and estuarine species ofgamefish. The tarpon, Megalops atlantica,snook, Centropomus undecima1is, and 1ady­fish, Elops saurus, utilize these areasfrom the time they reach the estuary aspost-larvae, having been spawned offshore.Gray snapper, Lutj anus .9.ri seus,sheep shead, Archosargus probatocephalus,spotted seat rout, Cvnoscion nebulosus, andred drum, Sciaenops uce1lata, are re­cruited to grass beds of shallow bays andlagoons as post-larvae and enter themangrove-lined streams for the next sever­al years (Heald and Odum 1970). Of thesespecies, only the spotted seatrout prob­ably spawns in the estuary (Tabb 1966).Other species of commercial or game impor­tance which use the riverine fringinghabitat include creval1e jack, gaff topsailcatfish, jewfish, striped mojarra, barra­cuda, Atlantic thread herring, and yel10w­fin menhaden (Odum 1970).

7.3 FRINGING FORESTS ALONG ESTUARINE BAYSANO LAGOONS

Mangrove-fringed estuarine bays andlagoons are exemplified by the TenThousand Is1 ands area and Whitewater Bay.Quantitative fish data are available fromFahkahatchee Bay (Carter et al. 1973;Yokel 1975b; Seaman et a1. 1973), FahkaUnion Bay (Carter et a1. 1973), RookeryBay (Yokel 1975a), the Marco IslandEstuary (Weinstein et a1. 1977; Yokel1975a), and Whitewater Bay (Clark 1970).Individual site characteristics aresummarized in Appendix A. All exceptFahka Union Bay contain significantamounts of sea 9rasses. Macroa19ae domi­nate the benthic producers of Fahka UnionBay. Studi es by Reid (1954) and Ki 1by(1955) near Cedar Key, F10rida,were notincluded in our summary because man9rovesare sparse in this area and no mention ofman9rove collecting sites were made bythese authors. Studies of Ca100sahatcheeBay (Gunter and Hall 1965) and ofCharlotte Harbor (Wang and Raney 1971)

54

were omitted because the areas studiedhave been highly modified and because datafrom many habitats were pooled in thefinal presentation.

All of the bays reviewed in our sum­maries are fringed by dense growths of redmangroves and all contain small mangroveislets. Carter et al. (1973), in theirstudies of Fahkahatchee and Fahka Unionbays, esti mated that 57% to 80% of thetotal energy budget of these two bays issupported by exports of particulate anddissolved organic matter from the man­groves within the bays and inflowing tidalstreams. lugo et al. (1 980) esti matedthat the mangroves surrounding Rookery Bayprovi de 32% of the energy base of theheterotrophic community found in the bay.

Salinities in these bays tend to behigher than in the tidal streams andrivers and the fish assemblages reflectboth this feature and the added habitatdimension of sea grass and macro algaebeds. Truly freshwater species are rarein these communi ties and a propo rt i ona llygreater percentage of marine visitors ispresent. The dominant fish families ofthe benthic habitat include drums(Sciaenidae), porgys (Sparidae), grunts(Pomadasyidae), mojarras (GerreidaeJ,snappers (lutjani dae), and mull et (Mugi 1i­dae). Other fami1es with sizeable contri­butions to the benthic fauna include pipe­fishes (Syngnathidae), flounder (Bothi­dae), sole (Soleidae), searobins (Trig1i­dae), and toadfishes (Batrachoididae).

Numerically abundant fishes of themid and upper waters include anchovies(Engrau1idae), herrings (Clupeidae) andneed1efishes (Belonidae). At all loca­tions studied, the benthic fauna was domi­nated by the pinfish, lagodon rhomboides,the silver perch, Bairdie11a chrysura, thepigfish, Orthopristis chrysoptera, and themojarras, Eucinostomus ~ and I.argenteus. The most common ml dwater andsurface species include the two anchovies,Anchoa mitchilli and ~. hepsetus, and twoclupeids, Brevoortia smithi and Harengu1apensaco1ae. The total number of speciesrecorded in the individual studies rangedfrom 47 to 89; a total of 117 speci es was

collected in these mangrove-fringed baysand lagoons (Appendix B).

In none of these studies were thefishes specifically utilizing the fringingmangrove habitat enumerated separatelyfrom those collected in the bay as awhole. The collections were most often atopen water stations easily sampled byotter trawl. Carter et al. (1973) had twoshore seine stations adjacent to mangrovesbut the data were pooled for pub1i cati on.Of the four stations in Rookery Bay sam­pled by Yokel (lg75a), one was immediatelyadjacent to the fringing mangrove shore­1i ne and had moderate amounts of seagrasses.

The typical pattern which emergesfrom many estuarine studies is that rela­tively few fish species numerically domi­nate the catch. This is certainly true inmangrove-fringed estuaries. In RookeryBay (Yokel Ig75a) six species comprised88% of the trawl-catchable fishes, inFahkahatchee Bay seven species comprised97% of the catch from three capturetechniques (Carter et al. 1973), and inthe Marco Island estuary 25 species com­prised 97% of the trawl-catchable fishes(Weinstein et al. 1977).

like tidal river and stream communi­ties, these shallow bays serve as nur­series for numerous species of estuarine­dependent fishes that are spawned off­shore. Based on the distribution andabundance of juveni Ie fishes of all spe­cies in six habitats, Carter et al. (1973)ranked the mangrove-fringed bays as themost important nursery grounds; the ti dalstreams were a close second. Shallow baysand tidal streams provide safe nurseriesdue to seasonally abundant food resourcesand the low frequency of large predators(Carter et al. 1973; Thayer et al. 1978).The relative lack of large predaceousfishes is probably due to their generalinability to osmoregu1ate in waters of lowand/or fl uctuating sal inity.

As in tidal streams, the peak abun­dance of juvenile and larval fishes in thebays is in spring and early summer (Reid1954). In general, the highest standing

55

crops and the greatest species richness offishes occur in the late summer and earlyfall (Clark 1970). Fish densities declinein the autumn and winter as many fishesmove to deeper waters.

7.4 FRINGING FORESTS ALONG OCEANIC BAYSAND LAGOONS

Mangrove-fri nged "oceani c" bays andlagoons are exemplified by Porpoise Lakein eastern Florida Bay (Hudson et al.1970), western Florida Bay (Schmidt 1979),southern Biscayne Bay (Bader and Roessler1971), and Old Rhodes Key Lagoon ineastern Biscayne Bay (Holm 1977). Charac­teristics of these sites are summarized inAppendix A. Compared to the mangrove­fringed bays discussed in the previoussect ion, these envi ronments generally ex­hibit clearer water, sandier substrates,and higher and less variable salinities.Closer proximity to the Florida reeftract, the Atlantic Ocean, and the Gulf ofMexico results in a larger potential poolof fish species. These four locationshave produced reports of 156 fi sh speci es(Appendix B).

Mangrove fringes make up a relativelysmall proportion of these environments;accordingly, their contribution to the bayfood webs is probably not very 1arge.Bader and Roessler (1972) esti mated thatthe fringing mangrove community contrib­utes approximately 1% of the total energybudget of southern Biscayne Bay; theyconsidered only mainland mangroves and didnot include the small area of mangroveislands. The main ecological role of thefringing mangroves in this type of en­vironment is probably twofold. First,they increase the habitat diversity withinan otherwise relatively homogeneous baysystem. Second, they provide a relativelyprotected habitat for juvenile fishes (andcertain invertebrates) that later move tomore open water or coral reef communities.The second role is analogous to one of theecological roles of sea grass communities(see Zi eman, in prep.) a 1though the fi shspeci es invo1 ved may be different.

Based pri mari lyon habitat desi gna­tions of Voss et al. (1969), the fishes ofBiscayne Bay can be characterized as topreferred habitat. Of the three mainhabitat types, (1) rock/coral/seawall, (2)grassbed/tida1 flat, and (3) mangrove, thegrassbed/tida1 flat ranked first in fishspeci es occurrences. One hundred andtwenty- two of 156 speci es (79%) are knownto occur in this environment.Rock/coral/seawall habitats were fre­quented by 49 species (32%) and mangrovesare known to be utilized by 54 species(35%) of the total fish species recordedfrom this bay.

7.5 OVER WASH MANGROVE ISLANDS

In terms of fish-related research,these communities are the least studied ofall mangrove community types in southFlorida. They are typified by the 10w­1yi ng mangrove-covered is 1ands that occurin the Florida Keys and Florida Bay andmay be overwashed periodically by thetides. Examples include Shell Key, CottonKey, and the Cowpens. Islands of thistype extend southwest from the Floridamai n1 and through the Marquesas. The DryTortugas lack well-developed mangrove com­munities although stunted trees are found(Davi s 1942).

These islands are the most oceanic ofany of the mangrove communiti es di scussed.They are characterized by relatively clearwater (Gore 1977) and are largely free ofthe freshwater inflow and salinity varia­tions which characterize other Floridamangrove communities to varying degrees.Numerous statements exist in the litera­tu re acknow1 edgi ng the frequent proxi mityof mangrove islands to coral reefs and seagrass beds (McCoy and Heck 1976; Thayer eta1.1g7B). Olsen et al. (1973) working inthe U.S. Virgin Islands, found 74% to 93%overlap in the fish species composition offringing coral reefs and shallow mangrove­fringed oceanic bays. Voss et al. (1969)listed fish species that were collectedfrom all three types of communities:fringing mangroves, coral reefs and sea

56

grass beds in Biscayne Bay, but thereappears to have been no systematic surveyof the fish assemblage characteristic ofthe mangrove-covered or mangrove-fringedFlorida Keys. No one has quantified thefaunal connections which we hypothesizeexist between the mangroves and seagrasses and between the mangroves andcoral reefs.

In the absence of published data fromthe mangrove key communities, only tenta­tive statements can be made. In general,we expect that while mangrove islandsserve as a nursery area for juvenilefishes, this function is limited largelyto coral reef and marine inshore fishesand not the estuarine-dependent speciesthat we have discussed previously. Thelatter (juvenile snook, red drum, spottedseat rout) appear to require relatively lowsalinities not found in association withmost of the overwash is 1ands. Casu a 1observation around the edges of theseislands suggests that characteristicfishes include the sea bass family (Ser­ranidae), triggerfishes (Balistidae),snappers (Lutjanidae), grunts (Poma­dasyidae), porgies (Sparidae)\parrotfishes(Scaridae), wrasses \LabridaeJ, bonefishes(Albulidae), jacks (Carangidae), damsel­fishes (Pomacentridae), and surgeonfi shes(Acanthuri dae); many of these fi shes occuron or are associ ated with coral reefs. Wealso suspect that considerable overlapoccurs in the fish assemblage of thesemangrove islands and sea grass communi­ties; examples include puffers (Tetrao­dontidae), pipefishes (Syngnathidae), go­bies (Gobiidae) and scorpionfishes (Scor­paenidae). Stark and Schroeder (1971)suggested that juveni le gray snapper,which use the fringing mangroves of thekeys as shelter during the day, forage inadjacent sea grass beds at night. In theabsence of salinity barriers, predatoryfishes probably enter the fringes of these

57

mangrove islands on the rlSlng tide.Included in this group are sharks, tarpon,jacks, snook, bonefish and barracuda.

7.6 GRADIENT OF MANGROVE COMMUNITYINTERACTIONS

Mangrove commun i ties occur under awide range of conditions from virtuallyfreshwater at the headwaters of tidalstreams to nearly oceanic conditions inthe Florida Keys. Attempting to present asingle list of fish characteristic ofmangrove environments (Appendix B) can bemisleading. For this reason we presentedthe concept of a continuum or complexgradient in Figure 11 and have followedthat scheme throughout section 7. Thegradient stretches from seasonally freshto oceanic conditions, from highly varia­ble salinities to nearly constant salini­ty, from muddy and 1i mestone subs t rates tosandy substrates, from dark-stained andsometimes turbid waters to clear waters,and from food webs that are predomi nantlymangrove detritus-based to food webs basedpri mari lyon other energy sources. Cl ear­ly, there are other gradients as one movesfrom north to south in the State ofFlorida. At the northern end of theState, temperatures are more variable andseasonally lower than in the south. Sedi­ments change from predominantly siliciousin central and north Florida to predomi­nant ly ca rhonate in extreme south Flori da.Nevertheless, the complex gradient shownin Figure 11, while greatly simplified forgraphic purposes, suggests that charac­teristic fish assemblages replace oneanother along a 9radi ent of changi ngphysical and biogeographic conditions.Such a concept is useful in understandingthe factors controlling the composition offish assemblages associated with mangrovesof the four major community types in southFlorida.

CHAPTER B. COMMUNITY COMPONENTS - AMPHIBIANS AND REPTILES

Food habits and status of 24 speciesof turtles, snakes, lizards, and frogs ofthe Florida mangrove region are given inAppendix C. Any of three criteria had tobe met before a species was included inthis table: (1) a di rect reference inthe literature to mangrove use by thespecies, (2) reference to a species asbeing present at a particular geographicallocation within the mangrove zone ofFlorida, and (3) North American speciesrecorded from mangroves in the West Indiesor South America, but not from Florida.This last criterion assumes that a specieswhich can utilize mangroves outside ofFlorida will be able to use them inFlorida. Ten turtles are listed of whichfour (striped mud turtle, chicken turtle,Florida red-bellied turtle, and softshellturtle) are typical of freshwater. Two(mud turtle and the ornate diamondbackterrapin) are found in brackish water andthe remai nder (hawksbi 11, green, logger­head, and Atlantic ridley) are found inrna ri ne waters.

Freshwater species usually occur inthe headwater regions of mangrove-linedriver systems. All four freshwaterspecies are found in habitats other thanmangrove swamps including streams, ponds,and freshwater marshes. The brackishwater species are found in salt marshes inaddition to mangrove swamps. Mangroves,however, are the principal habitat for theornate diamondback terrapin (Ernst andBarbour 1972). Carr and Goi n (1955)listed two subspecies of the diamondback:Malaclemys terrapin macrospilota and ~. 1.rhizophorarum. Malaclemys terrapin macro­spilota inhabits the south\~est and south­ern coasts, and ~. 1. rhizophorarum isfound in the Florida Keys. The two sub­species intergrade in the region of north­ern Florida Bay.

All four of the marine turtles areassociated with mangrove vegetation atsome stage of their lives. Loggerhead andgreen turtles are apparently much lessdependent on mangroves than the remai ni ngtwo, although we strongly suspect thatrecently hatched loggerheads may use man­grove estuaries as nursery areas. Greenturtles are generally believed to feed on

a variety of submerged aquatic plants andsea grasses; recent evidence has shownthat they al so feed on mangrove roots and1eaves (Ernst and Barbour 1972). TheAtlantic ridley's preferred habitat is"shallow coastal waters, especially themangrove-bordered bays of the southernhalf of the peninsula of Florida" (Carrand Goin 1955). Hawksbill turtles feed ona variety of plant materials includingmangrove (especially red mangrove),fruits, leaves, wood, and bark (Ernst andBarbour 1972).

Three species in the genus Anolishave been reported from Florida mangroves:the green anole, the cuban brown anole,and the Bahaman bank anole. All arearboreal lizards that feed on insects.The green anole is widespread throughoutthe Southeastern United States and is notat all dependent on mangrove swamps. Theother two species have much morerestricted distributions in the UnitedStates and are found only in southFlorida. They also are not restrictedto mangrove ecosystems. Of the sixspecies of snakes listed, the mangrovewater snake (Figure 13) is most dependentupon mangrove habitat~

Two important speci es of rept i 1esfound in mangrove swamps are the Americanall i gator and the Ameri can crocodi 1e. Thealligator is widespread throughout theSoutheastern United States and is onlyincidentally found in low salinity sec­tions of Florida mangrove areas (Kush1 an1980). The American crocodile is rare;historically its distribution was centeredin the mangrove-dominated areas of theupper and lower Florida Keys (particularlyKey Largo) and the mangrove-l ined shore­lines and mud flats along the northernedge of Florida and Whitewater Bays(Kushlan 19BO). Mangroves appear to becritical habitat for this species. Itsrange has shrunk considerably in southFlorida since the 1930's, even thoughFlorida Bay was added to EvergladesNational Park in 1950 (Moore 1953; Ogden1978). Much of the decrease in range isdue to increased human activity in theFlorida Keys. The remaining populationcenters of the American crocodile are in

58

Figure 13. The mangrove water snake. Nerodia fasciata compressicauda. curled ona red mangrove prop root. Photograph by David Scott.

59

northern Flori da Bay and adjacent coasta 1swamps and the northern end of Key Largo(Ogden 1978; Kushlan 1980). The speciesuses a variety of habitats for nesting inthe Florida Bay region including openhardwood thickets along creek banks,hardwood-shrub thickets at the heads ofsand-shell beaches, and thickets of blackmangroves behind marl banks (Ogden 1978).On Key Largo the crocodile locates itsnests on creek and canal banks in red andblack man9rove swamps (Ogden 1978). Man­grove areas thus appear to be important inthe breeding biology of this endangeredspecies.

Interestingly, only three species of

amphibians, to our knowledge, have beenrecorded in Florida mangrove swamps (Ap­pendi x C). Thi sis due to two factors:(1) lack of detailed surveys in low sa­l i nit.y swamps and (2) the i nabi 1i ty ofmost amphibians to osmoregulate in saltwater. No doubt, several additionalspeci es occur in the freshwater -domi natedhammock and basin mangrove communitiesinland from the coast. Possible addi­tional species include: the easternnarrow-mouthed toad, Gastrophryne caro­linensis, the eastern spade foot toad,Scaphiopus holbrooki, the cricket frog,Acris gryllus, the green tree frog,~cinerea, and the southern leopard frog,Rana utricularia.

60

CHAPTER 9. COMMUNITY COMPONENTS - BIRDS

9.1 ECOLOG ICAL RELA11 ONSH I PS

Because mangroves present a moredi verse st ructural habitat than mostcoastal ecosystems, they should harbor agreater variety of bird1ife than areassuch as salt marshes, mud fl ats, andbeaches (MacArthur and MacArthur 1961).The shallow water and exposed sedimentsbelow mangroves are available for probingshorebirds. Longer-legged wading birdsutilize these shallow areas as well asdeeper waters along mangrove-lined poolsand waterways. Surface-feeding and divingbirds would be expected in similar areasas the wadi ng bi rds. The major di fferencebetween mangrove swamps and other coastalecosystems is the availability of thetrunks, limbs, and foliage comprising thetree canopy. This enables a variety ofpasserine and non-passerine birds, whichare not found commonly in other wetlandareas, to use mangrove swamps. It a1 soallows extensive breeding activity by anumber of tree-nesti ng bi rds.

The composition of the avifauna com­munity in mangrove ecosystems is, in fact,highly diverse. Cawkell (1964) recorded45 species from the mangroves of Gambia(Africa). Haverschmidt (1965) reported 87species of birds which utilized mangrovesin Surinam (S. America). Ffrench (1966)listed 94 species from the Caroni mangroveswamp in Trinidad while Bacon (1970) found137 in the same swamp. In Malaya, Nisbet(1968) reported 121 speci es in mangroveswamps and Field (1968) observed 76 fromthe mangroves of Sierra Leone (Africa).

Use of mangrove ecosystems by bi rdsin Florida has not been recorded in de­tai 1. Ninety-two species have been ob­served in the mangrove habitat of SanibelIsland, Florida (L. Narcisse, J.N. "Ding"Darling Nat1. Wildlife Refuge, SanibelIs., Fla.; personal communication 1981).Robertson (1955) and Robertson and Kush1an(1974) reported on the entire breedingbird fauna of peninsular south Florida,including mangrove regions. Based on1i mi ted su rveys, these authors reportedonly 17 species as utilizing mangroves forbreeding purposes. Because their studiesdid not consider migrants or non-breeding

residents, a significant fraction of theavifauna communi ty was omi tted.

Based on information 91eaned from theliterature, we have compiled a list of 181species of birds that use Florida mangroveareas for feeding, nesting, roosting, orother activities (Appendix D). Criteriafor listing these species is the same asthat used for listing reptiles and amphi­bians (see Chapter 8 of this volume).

Often references were found statingthat a given species in Florida occurredin "wet coastal hammocks", "coastal wetforests" or the 1 ike, without a specificreference to mangroves. These specieswere not included in Appendix D. Thus,this list is a conservative estimate ofthe avifauna associ ated with Flori da man­grove swamps. Sources for each listingare provided even though many are redun­dant. Food habit data are based on Howell(1932) and Martin et a1. (1951). Esti­mates of abundance were derived from birdlists published by the U.S. Fish andWildlife Service for the J.N. "Ding"Darling National Wildlife Refuge atSanibel Island, Florida, and by the Ever­glades Natural History Association forEverglades National Park. Frequently,species were recorded from mangrove swampsat one location, but not the other.

We have divided the mangrove avifaunainto six groups based on similarities inmethods of procuri ng food. These groups(guilds) are the wading birds, probingshorebirds, floating and diving water­birds, aerially-searching birds, birds ofprey, and a rborea1 bi rds. Thi s 1ast groupis something of a catch-all group, but iscomposed mai n1y of bi rds that feed and/ornest in the mangrove canopy.

9.2 WAD I NG BIRDS

Herons, egrets, ibises, bitterns, andspoonbills are the most conspicuous groupof birds found in mangroves (Figure 14)and are by far the most studied and bestunderstood. Eighteen species (and oneimportant subspecies) are reported fromsouth Flori da mangroves.

61

Figure 14. A variety of wading birds feeding in a Inan9rove-lined pool nearFlamingo, Florida. Photograph by David Scott.

62

Mangrove swamps provide two functionsfor wadi ng bi rds. Fi rst, they funct i on asfeedi ng grounds. Two-thi rds of thesespecies feed almost exclusively on fishes.Although much of their diet is provided byfreshwater and non-mangrove mari ne areas,all of them feed frequently in mangroveswamps. White ibis feed predominantly oncrabs of the genus Uca when feeding inmangroves (Kushlanand Kushlan 1975;Kush1an 1979). Mollusk sand i nve rtebratesof the sedi ments are pri nci pal foods ofthe roseate spoonbi 11 although some fishare eaten (Allen 1942). Yellow-crownednight herons and American bitterns eatcrabs, crayfi sh, frogs, and mi ce in addi­tion to fishes. Snails of the genusPomacea are fed upon al most excl usi vely bythe limpkin. The sandhill crane is ananomaly in this group since a majority ofits food is vegetable matter, especiallyroots and rhi zomes of Cyperus andSagi ttari a. Its use of mangroves isprobably minimal, occurring where inlandcoastal marshes adjoin mangroves (Kushlan,unpubl. data). The remaining 12 speciesare essentially piscivorous although theydiffer somewhat in the species and sizesof fishes that they consume.

Mangrove swamps al so serve asbreeding habitat for wading bi rds. Withthe exception of the limpkin, sandhillcrane, and the two bitterns, all wadingbird species in Appendix D build theirnests in all three species of mangrovetrees (Maxwell and Kale 1977; Gi rard andTaylor 1979). The speci es often aggregatein large breeding colonies with severalthousand nesting pairs (Kushlan and White1977a). The Louisiana heron, snowy egret,and cattle egret are the most numerousbreeders in south Florida mangroves (basedon data in Kushlan and White 1977a).

In wet years over 90% of the southFlorida population of white ibis breed inthe i nteri or, freshwater wet 1ands of theEverglades; during these times the man­groves are apparently unimportant, sup­porting less than 10% of the population(Kushlan 1976, 1977a, b). During droughtyears, however, production is sustainedsolely by breeding colonies located inmangroves near the coast (Kushl an 1977a,

b). Mangroves are critically importantfor the survival of the white ibis popula­tion even though they appear to beutilized to a lesser extent than fresh­water habitats. This pattern of largerbut less stable breeding colonies usinginland marshes and smaller but more stablecolonies using mangroves is also charac­teristic of heron populations (Kushlan andFrohring, in prep.).

Table 5 gives the number of activenests observed in mangrove regi ons du ri ngthe 1974-75 nesting season and the percen­tage this represents of the enti re southFlorida breeding population for the ninemost abundant species of waders and threeassociated species. The dependence ofroseate spoonbills, great blue herons,Louisiana herons, brown pelicans, anddouble-crested cormorants on mangroveregions is evident. Nesting by the red­dish egret was not quantified during thisstudy although Kushlan and White (1977a)indicated that the only nests of thisspecies which they saw were, in fact, inmangroves. Fu rther observat ions i ndi catethat this species nests in mangroves ex­clusively (Kushlan, pers. comm.). Similar­ly, the great white heron is highly depen­dent upon mangroves for nesting; they usethe tiny mangrove islets which aboundalong the Florida Keys and in Florida Bay(Howell 1932).

During many years the Evergladespopulation of wood storks is known to nestalmost solely in mangroves (Ogden et al.1976); this population comprises approxi­mately one-thi rd of the total southFlorida population. Successful breedingof all these mangrove nesters is un­doubtedly correlated with the abundantsupply of fishes associated with man­groves. Meeting the energetic demands ofgrowing young is somewhat easier in habi­tats with abundant prey. Thi sisespecially important for the wood storkwhich requires that its prey be concen­trated into small pool s by fall i ng waterlevels during the dry season before it cannest successfully (Kahl 1964; Kushlan eta1. 1975; Odgen et al. 1978). Breedingacti vity by wadi ng bi rds in mangrovesalong the southwest and southern Florida

63

Table 5. Nesting statistics of wading birds and associatedspecies in south Florida, 1974-1975 (based on data inKushlan and White 1977a).

% of total act i veAct i 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-crestedcormorant 1744 83

64

coasts takes place throughout the year(Table 6); at least one species of waderbreeds during every month. Colonies onthe mangrove islands in Florida Bay werenoted to be active nesting sites duringall months of the year except Septemberand October (Kush1an and White 1977a).

The seasonal movements of wood storksand white ibises between the various southFlorida ecosystems were described byOgden et a1. (197B) and Kushl an (1979).Mangrove ecosystems appear to be mostheavily used for feeding in summer (whiteibis) and early winter (white ibis andwood stork). The remai ni n9 speci es ofwading birds appear to use mangrove areasmost heavily in the winter months, reflec­ting the influx of migrants from farthernorth.

Wading birds play an important rolein nutrient cycling in the coastal man­grove zone. Mcivor (pers. observ.) hasnoted increased turbidity, greater algalbiomass, and decreased fish abundancearound red mangrove islets with nestingfri gate bi rds and cormorants. Onuf et a1.(1977) reported resu 1ts f rom a sma 11 (100bi rd) rookery on a mangrove isl et on theeast coast of Flori da. Addi t ions ofammonium-nitrogen fro~ the bird'sdroppings exceeded 1 glm Iday. Waterbeneath the mangroves contained five timesmore ammonium and phosphate than waterbeneath mangroves without rookeries.Although the wading birds were shown to bea vector for concentrating nutrients, itmust be noted that this is a localizedphenomenon restricted to the areas aroundrookeri es in the mangrove zone. Theeffect would be larger around largerrookeries. Onuf et al. (1977) alsoreported that mangroves in the area of therookery had increased levels of primaryproduction, higher stem and foliar nitro­gen levels, and hi9her herbivore grazingimpact than mangroves without rookeries.Lewis and Lewis (1978) stated that man­groves in large rookeries may eventuallybe killed due to stripping of leaves andbranches for nesting material and bypoisoning due to large volumes of urea andammonia that are deposited in bird guano.This latter effect would be more

pronounced in rookeries within mangroveregions subject to infreguent tidal f1ush­i ng.

9.3 PROBING SHOREBIROS

Bi rds in this group are commonlyfound associated with intertidal and shal­low water habitats. Wo1 ff (1969) andSchnei der (1978) have shown that ploversand sandpi pers are opportuni sti c feeders,takin9 the most abundant, proper-sizedinvertebrates present in whatever habitatthe bi rds happen to occupy.

Of the 25 species included in thisguild (Appendix DJ, two are year-roundresidents (clapper rail and willet), twobreed in man9rove areas (clapper rail andblack-necked stilt), and the remainder aretransients or winter residents. Baker and8aker (1973) indicated that winter was themost crucial time for shorebirds, in termsof survival. Coincidentally, winter isthe time when most shorebirds use man9roveareas. The invertebrate fauna (moll usks,crustaceans, and aquatic insects) whichoccur on the sediments under intertidalmangroves forms the pri nci pal di et ofthese species. Willets and greateryellowlegs eat a large amount of fishes,especially Fundulus, in addition to inver­tebrates. Many of the speci es 1 i sted inthis guild obtain a significant portion oftheir energy requirements from other habi­tats, particularly sandy beaches, marshes,and freshwater prai ri es. Of the speci esin this guild, the clapper rail is prob­ably most dependent on mangroves forsurvi val in south Florida (Robertson1955), although in other geographicallocations they frequent salt and brackishmarshes.

9.4 FLOATING AND DIVING WATER BIRDS

Twenty-nine species of ducks, grebes,loons, cormorants, and gallinules wereidentified as populating mangrove areas insouth Flori da (Appendi x D). Ei ght speci esare year-round residents while theremainder are present only during migra­tion or as winter visitors.

65

Table 6. Timing of nesting by wading birds and associatedspec i es in south Flori da. Adapted from data in Kush 1an andWhite (1977a). Kushlan and McEwan (in press).

Species

White ibis

Wood stork

Roseate spoonbill

Great bl ue/whiteheron

Great egret

Little blue heron

Cattle egret

Double-crestedcormorant

Brown pelican

MonthsSON 0 J F M A M J J A

66

From the standpoint of feeding, mem­bers of this guild are highly hetero­geneous. Piscivorous species include thecormorant, anhinga, pelicans, and mergan­sers. Herbivorous species include thepintail, mallard, wigeon, mottled duck,and tea 1s. A th i rd group feeds pri ma ri 1yon benthic mollusks and invertebrates.Scaup, canvasback, redhead, and gallinulesbelong to this group. The ducks in thislast group al so consume a significantfraction of plant material.

Species of this guild are permanentresidents and usually breed in mangroveswamps. As shown in Table 5, the brownpelican and double-crested cormorant arehighly dependent upon mangroves fornestin~ in south Florida even though bothwill build nests in any available tree inother geographical regions. It seems thatwhen m~ngroves are available, they are thepreferred nesting site. The anhingabreeds in mangrove regions but is morecommonly found inland near freshwater (J.A. Kush1an, So. Fla. Res. Ctr., EvergladesNat1. Park, Homestead, Fla.; ·personalcommunication 1981). For the other specieslisted in this guild, mangrove swampsprovi de a common but not a requi red habi­tat; all of these species utilize avariety of aquatic envi ronments.

Kush1an et a1. (in prep.) providerecent data on the abundance and distribu­tion of 22 species of waterfowl and theAmerican coot in south Florida estuaries.The American coot is by far the most abun­dant species, accounting for just over 50%of the total popu1 at ion. Si x spec i es ofducks were responsible for more than 99%of the individuals seen: blue-winged teal(41%), lesser scaup (24%), pintaii (18%),American wigeon (9%), ring-necked duck(5%), and shoveler (3%). The major habi­tats included in these authors' surveyswere coastal prairie and marshes, mangroveforests, and mangrove-1 i ned bays andwaterways of the Everglades National Park.

From these data it appears thatwaterfowl and coots are most abundant inregions where mangrove, wet coastalprairies, marshes, and open water areinterspersed. Overall, the Ever~lades

estuaries support from 5% to 10% of thetotal wintering waterfowl population inFlorida (Goodwin 1979; Kush1an et a1. inprep.). As Kush1an et a1. point out,however, the Everglades are not managedfor single species or ~roups of species asare areas of Florida supporting largerwaterfowl populations. Although theimportance of south Flori da's mangroveestuaries to continental waterfowl popula­tions may be small, the effect of 70,000ducks and coots on these estuariesprobably is not (Kush1an et a1. in prep.).

Kushlan (personal communication)thinks that the estuaries of the Ever­glades have an important survival valuefor some segments of the Ameri can whitepel i can popu1 ati on. In wi nter, approxi­mate1y 25% of the white pelicans are foundin Florida Bay and 75% in the Cape Sableregion. They feed pri marily in freshwaterregions of coastal marshes and prairiesand use mangroves where they adjoi n thi stype of habitat.

9.5 AERIALLY-SEARCHING BIRDS

Gulls, terns, the kingfisher, theblack skimmer, and the fish crow comprisethis 9ui1d of omnivorous and piscivorousspecies (Appendix D). These bi rds hunt inponds, creeks, and waterways adjacent to·man9rove stands. Many fishes and inverte­brates upon which they feed come frommangrove-based food webs. Only six of the14 species are year-round residents ofsouth Florida. The least tern is an abun­dant summer resident and the remainder arewinter residents or transients.

Only the fish crow actually nests inmangroves. Gulls and terns prefer opensandy areas for nesting (Kushlan and White1977b) and use mangrove ecosystems onlyfor feeding. All of the species in thisguild are recorded from a variety ofcoastal and inland wetland habitats.

9.6 BIRDS OF PREY

This guild is composed of 20 speciesof hawks, falcons, vultures, and owls

67

which utilize mangrove swamps in southFlorida (Appendix D). The magnificantfrigatebird has been included in thisgroup because of its habit of robbing manyof these birds of their prey. Prey con­sumed by this guild includes snakes,lizards, frogs (red-shouldered hawk,swallow-tailed kite), small birds (short­tail ed ha wk), waterfowl (pe regri ne fa 1con,great-horned owl), fishes (osprey, baldeagle), and carrion (black and turkeyvultures) •

Eleven of these species are permanentresidents, one a summer resident, and therema i nder a re wi nter res i dents. Thei ruseof mangrove areas varies greatly. Themagni fi cent fri gatebi rd, whi ch occu rsprincipally in extreme southern Floridaand the Florida Keys, utilizes small over­wash mangrove islands for both roosts andnesting colonies. Both species of vul­tures are widely distributed in southFlorida mangrove regions; large colonialroosts can be found in mangrove swampsnear the coast. Swallow-tailed kites arecommon over the enti re Florida mangroveregion (Robertson 1955; Snyder 1974).Snyder (1974) reports extensively on thebreeding biology of the swallow-tailedkites in south Flori da. The nests heobserved were all located in black man­groves although they do nest in otherhabitats.

The bald eagle, osprey (Figure 15),and peregrine falcon are dependent uponmangrove ecosystems for thei r cont i nuedexistence in south Florida. Both the baldeagle and osprey feed extensively on thewealth of fishes found associated withmangrove ecosystems. Additionally, man­groves are used as roosts and supportstructures for nests. Nisbet (1968) indi­cated that in Ma 1ays i a the most importantrole of mangroves for bi rds may be aswintering habitat for palaearctic mi­grants, of which the peregrine falcon isone. Kushl an (pers. comm.) stated thatrecent surveys have shown falcons towinter in mangroves, particularly alongthe shore of Florida Bay where they estab­lish feeding territories. They forage onconcentrations of shorebirds and water­fowl. These prey speci es of the peregri ne

are common inhabitants of mangrove areas.This could also be true for the merlin,which like the peregrine falcon, feeds onwaterfowl and shorebirds. The remainingspecies in this gui ld are probably not sodependent on mangroves; although they maybe common in mangrove ecosystems, theyutilize other habitats as well.

9.7 ARBOREAL BIRDS

This guild is the largest (71species) and most diverse group inhabitingmangrove forests. Included are pigeons,cuckoos, woodpeckers, flycatchers,thrushes, vireos, warblers, blackbirds,and sparrows. We have lumped this diversegroup together because they utilize man­grove ecosystems in remarkably similarways. Invertebrates, particularlyinsects, make up a significant portion ofmost of these birds' diets, although thewhite-crowned pigeon, mourning dove, andmany of the fringilids (cardinal, towhee)eat a variety of seeds, berries, andfruits.

As the name given this guild implies,these birds use the habitat provided bythe mangrove canopy. Many bi rds al so usethe trunk, branches, and aerial roots forfeedi ng. Several di fferent types ofsearching patterns are used. Hawking ofinsects is the primary mode of feeding bythe cuckoos, chuck-wi 11 s-widows, thekingbi rds, and the flycatchers. Gleaningis employed by most of the warblers.Woodpeckers and the prothonotary warblerare classic probers.

Several of the bi rds in this guildare heavi ly dependent upon mangrove areas.The prairie warbler and the yellow warblerare subspecies of more widespread NorthAmerican species (see Appendix 0 forscientific names). They are found largelywithin mangrove areas (Robertson andKushlan 1974). The white-crowned pigeon,mangrove cuckoo, gray kingbird, and b1ack­whi skered vi reo are of recent West Indianor i gin. They fir s t m0 ve din tot hemangrove-covered regions of south Floridafrom source areas in the islands of theCaribbean. Confined at fi rst to mangrove

68

Figure 15. Osprey returning to its nest in a red mangrove tree near WhitewaterBay. Photograph by David Scott.

69

swamps, all but the mangrove cuckoo haveexpanded their range in peninsular Floridaby using non-mangrove habitat. ln thisvein it is interesting to note that manyspecies of rare and/or irregular occur­rence in south Florida are of West Indianorigin and use mangroves to a considerableextent. These include the Bahama pintail,masked duck, Caribbean coot, loggerheadkingbird, thick-billed vireo, and stripe­headed tanager (Robertson and Kushl an1974).

Twenty-four of the species in thisguild are permanent residents, 27 are win­ter, and 6 are summer residents. Fourteenspecies are seen only during migrations.

9.B ASSOCIATIONS BETWEEN MANGROVECOMMUNITY TYPES AND BIRDS

Estimating the degree of use ofmangrove swamps by bi rds as we have done(Appendi x D) is open to crit i cis m becau seof the pauc i ty of in format i on upon whi chto base judgements. Estimating whichmangrove community types (see section 1,Figure 4) are used by which birds is opento even more severe criticism. For thisreason the foll owi ng comments shou1 d beregarded as general and pre1 i mi nary.

In terms of utilization by avifauna,the scrub mangrove swamps are probably theleast utilized mangrove community type.Because the canopy is poorly developed,most of the arboreal species are absent,although Emlen (1977) recorded the red­winged blackbird, hairy woodpecker, north­ern waterthrush, yellow-rumped warbler,common yell owth roat, orange-crownedwarbler, palm warbler, yellow warbler,mourning dove, and gray kingbird in scrubmangroves on Grand Bahama Island. Of 25di fferent habitats surveyed by Em1en(1977), the yellow warbler and graykingbird were found in the scrub mangrovesonly. Aerially-searching and wading birdsmight use scrub mangroves if fishes arepresent.

Overwash mangrove i sl ands areutilized in a variety of ways by all ofthe bi rd guil ds. Most of the wadi ng bi rds

plus the magnificent frigatebird, theanhinga, the cormorant, and the brownpelican use overwash islands for nesting(Kushlan and White 1977a). Wading andaerially-searching birds commonly feed inclose proximity to overwash islands. Avariety of migrating arboreal and probingspecies use the islands for feeding androosting. Yellow and palm warblers arecommon around mangrove islands in FloridaBay as are the black-bellied plover, ruddyturnstone, willet, dunlin, and short­bi lled dowitcher. Rafts of ducks arecommon near the inshore islands and birdsof prey such as the osprey, the baldeagle, and both vultures use mangroveislands for roosting and nesting.

Fringe and riverine mangrove com­munities are important feeding areas forwading and probing birds. Floating anddiving and aerially-searching birds usethe lakes and waterways adjacent to thesemangrove communities for feeding. Many ofthe wading birds nest in fringe andri veri ne forests. For examp1 e, when thewood ibis nests in coastal areas, it usesthese mangrove communities almost exclu­sivel.y (Kushlan, personal communication).Most of the arboreal bi rds and bi rds ofprey associated with mangroves are foundin these two types of communi ties. Thi sis not surprising since the tree canopy isextremely well-developed and offersroosting, feeding and nesting opportuni­ties.

Hammock and basi n mangrove communi­ties are so diverse in size, location, andproximity to other communities that it isdi ffi cult to make many general statementsabout thei r avi fauna. Si nce there oftenis little standing water in hammockforests, wading and diving birds probablyare not common. Proxi mity to terrestri alcommunities in some cases may increase thedi versity of arboreal speci es in bothhammock and basin forests; proximity toopen areas may increase the likelihood ofbi rds of prey.

It seems safe to conclude that eachof the six mangrove community types hassome value to the avifauna. This valuedi ffers accordi ng to community type and

70

kind of bird group under consideration.Certainly, more information is needed,particularly concerning the dependence ofrare or endangered species on specificcommuni ty types.

g.g MANGROVES AS WINTER HABITAT FOR NORTHAMERICAN MIGRANT LAND BIRDS

An interestin9 observation based onthe data in this chapter is the seeminglyimportant rol e that mangrove ecosystemsplay in providing wintering habitat formi grants of North Ameri can ori gi n. Lackand Lack (1972) studi ed the wi nteri ngwarbler community in Jamaica. In fournatural habitats including mangroveforest, lowland dry limestone forest, mid­I evel wet 1i mestone forest, and montanecloud forest,a total of 174,131,61, and49 warblers (individuals) were seen,respectively. When computed on a per hourof observation basis, the difference ismore striking with 22 warblers per hourseen in mangroves and only I, 2, and 1seen in the other forest habitats, respec­tively. For all passerines consideredtogether, 26 passerines/hour were seen inmangroves with 5, 13, and 3 respectivelyin the other forest habitats. On a

species basis only 9 were recorded frommangroves whereas 19, 13, and 16 species,respectively, were seen in the other habi­tats. This large number of species fromthe other habitats appears to result fromthe sighting of rare species after manyhours of observation. Only 9 hours werespent by Lack and Lack (1972) in the man­groves whereas between 30 and 86 hourswere spent in other habitats. More ti mein the mangrove zone would have undoubted­ly resulted in more species (and in­dividuals) observed (Preston 1979).

Hutto (1980) presented extensive dataconcerning the composition of migratoryland bird communities in Mexico in winterfor 13 habitat types. Mangrove areastended to have more migrant species thanmost natura 1 habi tats (except gall eryforests) and also had a greater density ofindividuals than other habitats (againexcept for gallery forests). In both Lackand Lack's and Hutto's studies, disturbedand edge habitats had the hi ghest numberof species and greatest density ofi ndi vi dual s. The percentage of theavi fauna communi ty composed of mi grantswas hi ghest in mangrove habitats, however.From this we can infer the importance ofmangroves in the ma i ntenance of NorthAmerican migrant land birds.

71

CHAPTER 10. COMMUNITY COMPONENTS - MAMMALS

Thirty-six native and nine introducedspecies of land mammals occur in the southFlorida region (Layne 1974; Hamilton andWhittaker 1979). Of these, almost 50'; (18species) are found in the mangrove zone(Layne 1974). In addition, two species ofmarine mammals are known from mangroveareas. Oata on the abundance and foodhabits of these 20 species are summarizedin Appendix E. All are permanent resi­dents. The criteria for inclusion in thistable are similar to those used for theavifauna. Sight records in mangroves orlocality data from known mangrove areaswere required before a species was in­cluded. This has produced a conservativeest i mate of the mamma 1 speci es that uti­1ize mangrove areas.

Several mammal s do not appear inAppendix E because they have not beenrecorded from mangrove swamps in southFlorida; however, they occur so widelythat we suspect they will be found in thishabitat in the future. This groupincludes the cotton mouse, Peromyscusgossypinus, the hispid cotton rat, Sig­modon hispidus, the round-tailed muskrat,Neofiber alleni, the house mouse, Musmusculus,-fh-e-least shrew, CryptotlSparva, and the short-tailed shrew, BlarinabfeVTcauda.

Few rodents and no bats are includedin Appendix E. Compared to the rest ofthe State, the south Flori da regi on isdefi ci ent in these two groups (Layne1974). Although we have no confirmativefield data, we suspect that mangroveswamps along the central and north Floridacoasts contain more mammal species, par­ticularly rodents and bats.

A number of medium-sized and largecarnivores, including panther, gray fox,bobcat, striped skunk, raccoon, mink,river otter, and black bear, appear tout i 1i ze south Flori da mangrove areas.Only three of these species (stripedskunk, raccoon, and bobcat) are common inmangroves, but several of the rarerspecies seem to be highly dependent onmangrove swamps. Of 18 recent sightingsof the panther in Everglades NationalPark, 15 were from mangrove ecosystems

72

(Layne 1974). Hamilton and Whittaker(1979) state that it is the coastal ham­mocks of south Florida, including mangroveareas, which serve to preserve thisspeci es in the Eastern Uni ted States.Shemnitz (1974) reported that most of theremaining panthers were found in thesouthwest portion of Florida along thecoast and in the interior Evergladesregions.

The extent to which other carnivoresuse mangrove areas varies widely amongspecies. Schwartz (1949) states thatmink, although rare, prefer mangroves toother coastal habitats in Florida. Layne(1974, see his figure 1) gives a disjunctdistribution for this species in Florida,with the major geographical range beingthe southwest coast. River otters alsoutil i ze mangrove habitat heavily. Ottershave been found even far from shore onsmall mangrove overwash islands in FloridaBay (Layne 1974). Gray fox are not depen­dent upon mangroves, although they occa­sionally use this habitat. Less than 20';of all sightings of this species in Ever­glades National Park were from mangroves(Layne 1974). Bobcat are found in almostall habitats in south Florida from pine­lands to dense mangrove forests. Thepreponderance of recent sightings, how­ever, has been made from the mangrovezone, particularly on offshore mangroveoverwash islands (Layne 1974). Black bearare apparently most abundant in the BigCypress Swamp of Collier County (Shemnitz1974) and are rare in the remainder ofsouth Florida.

The small mammal fauna of the man­grove zone of south Florida are predomi­nately arboreal and terrestrial specieswhich are adapted to periodic flooding.Opossum, marsh rabbits, cotton rats, andri ce rats are commonly found in mangroveswamps. The Cudjoe Key rice rat is anewly described species found only onCudjoe Key in the Florida Keys. Thisspecies appears to be closely associatedwith stands of white mangroves (Hamiltonand Whittaker 1979).

White-tailed deer are common in

Florida mangrove swamps, although theyutilize many other habitats. The keydeer, a rare and endangered subspecies, isrestricted to the Big Pine Key group inthe Florida Keys, although it ranged ontothe mainland in historical times. Al­though this little deer makes use of pineuplands and oak hammocks, it extensivelyexploits mangrove swamps for food andcover.

Two marine mammals, the bottlenoseporpoi se and the manatee, frequentmangrove-l i ned waterways. The bott1enoseporpoi se feeds on mangrove-associ atedfishes such as the striped mullet, Mugilcephal us. Although the manatee -reeas

pri mari ly upon sea grasses and othersubmerged aquatic plants, it is commonlyfound in canals, coastal rivers, andembayments close to mangrove swamps.

Except for the Cudjoe Key rice rat,none of the mammals found in Fl or; da man­groves are sol ely dependent upon mangroveecosystems; all of these speci es canutilize other habitats. The destructionof extensive mangrove swamps would, how­ever, have deleterious effects on almostall of these species. Populations ofpanther, key deer, and the ri ver otterwould probably be the most seriouslyaffected, because they use mangrove habi­tat extensi vely.

73

CHAPTER 11. VALUE OF MANGROVE ECOSYSTEMS TO MAN

Mangrove swamps are often hot, fetid,mosqui to-ri dden, and a1most i mpenet rab1e.As a consequence, they are frequently heldin low regard. It is possible that moreacres of mangrove, worldwide, have beenobI iterated by man in the name of "recla­mation" than any other type of coastalenvironment. Reclamation, according toWebster's, means "to claim back, as ofwaste1and". Mangrove swamps are anythi ngbut wasteland, however, and it is impor­tant to establish this fact before avaluable resource is lost. We can thinkof six major categories of mangrove valuesto man; no doubt, there are more.

11.1 SHORELINE STABILIZATION AND STORMPROTECTION

The ability of all three Floridamangroves to trap, hold and, to someextent, stabilize intertidal sediments hasbeen demonstrated repeatedly (revi ewed byScoffi n 1970; Ca r 1ton 197 4). The contem­pora ry vi ew of mangroves is that theyfunction not as "land builders" as hypo­thesized by Davis (1940) and others, butas "stabilizers" of sediments that havebeen deposited largely by geomorphologicalprocesses (see section 3.2).

Gill (1970), Savaqe (1972), Teas(1977), and others have emphasized thatland stabilization by mangroves is pos­sible only where conditions are relativelyquiescent and strong wave action and/orcurrents do not occur. Unfortunately, noone has devised a method to predict thethreshold of physical conditions abovewhich mangroves are unable to survive andstabilize the sediments. Certainly, thisdepends to some extent on substrate type;mangroves appear to withstand wave energybest on solid rock substrates with manycracks and crevices for root penetration.From our own experience, we suspect thatmangroves on sandy and muddy substratescannot tolerate any but the lowest waveenergies, tidal currents much above 25cm/s, or heavy, regul ar boat wakes.

The concept that the red mangrove isthe best land stabilizer has been ques-

tioned by Savage (1972), Carlton (1974),and Teas (1 977). These authors argue thatthe black mangrove (1) is easier totransplant as a seedling, (2) establishesits pneumatophore system more rapidly thanthe red mangrove develops prop roots, (3)has an underground root system that isbetter adapted to holding sediments (Teas1977), (4) is more cold-hardy, and (5) canbetter tolerate "artificial" substratessuch as dredge-spoi 1, fi nger fi 11 s, andcauseways. Generally, the white mangroveis regarded as the poorest land stabilizerof the Florida mangroves (Hanlon et al.1975) •

Although mangroves are susceptible tohurricane damage (see section 12.1), theyprovide considerable protection to areason thei r 1andward si de. They cannotprevent all flooding damage, but they domitigate the effects of waves andbreakers. The degree of this protectionis roughly proportional to the width ofthe mangrove zone. Very narrow fringingforests offer minimal protection whileextensive stands of mangroves not onlyprevent wave damage, but reduce much ofthe flooding damage by damping and holdingflood waters. Fosberg (1971) suggestedthat the November 1970 typhoon and accom­panying storm surge that claimed between300,000 and 500,000 human lives inBangladesh might not have been so destruc­tive if thousands of hectares of mangroveswamps had not been replaced with ricepaddies.

11.2 HABITAT VALUE TO WILDLIFE

Florida mangrove ecosystems areimportant habitat for a wide variety ofreptiles, amphibians, birds, and mammals(see sections 8, 9, and 10). Some ofthese animals are of commercial and sportimportance (e.g., white-tailed deer, seaturtles, pink shrimp, spiny lobster,snook, grey snapper). Many of these areimportant to the south Florida touristindustry including the wading birds (e.g.,egrets, wood stork, white ibis, herons)which nest in the mangrove zone.

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11.3 IMPORTANCE TO THREATENED AND ENDAN­GERED SPECIES

The mangrove forests of south Floridaare important habitat for at least sevenendangered species, five endangered sub­species, and three threatened species(Federal Regi ster 1980). The endangeredspecies include the American crocodile,the hawksbill sea turtle, the Atlanticridley sea turtle, the Florida manatee,the bald eagle, the American peregrinefalcon, and the brown pelican. The endan­gered subspecies are the key deer(Odocoileus vir inianus clavium), theFlorida panther Felis concolor coryi),the Barbados yellow warbler (Dendroicaetechia petechia), the Atlantic saltmarsh

snake Nerodia fasciata taeniata) and theeastern indigo snake (Drymarchon coraiscouperi). Threatened species include theAmerican alligator, the green sea turtleand tM loggerhead sea turtle. Althoughall of these animals utilize mangrovehabitat at times in their life histories,species that would be most adverselyaffected by widespread mangrove destruc­tion are the American crocodi le, theFlorida panther, the American peregrinefal con, the brown pel i can, and theAtlantic ridley sea turtle. The so-calledmangrove fox sgui rrel (Sciurus nigeravicennia) is widely believed to be amangrove-dependent endangered species.This is not the case since it is currentlyregarded as 'Irare'~ not endan~eredt and,further, there is some question whetheror not this is a legitimate sub-species(Hall 1981). As a final note, we shouldpoint out that the red wolf (Canis rufus),which is believed to be extinct inFlorida, at one time used mangrove habitatin addition to other areas in southFlorida.

11.4 VALUE TO SPORT AND COMMERCIALFISHERIES

The fish and invertebrate fauna ofmangrove waterways are closely 1inked tomangrove trees through (a) the habitatva 1ue of the aeri a1 root st ructu re and (b)the mangrove leaf detritus-based food web(see sections 6 and 7). The impl ications

of these connections were discussed byHeald (1969), Odum (1970), Heald and Odum(1970), and Odum and Heald (1975b) interms of support for commercial and sportfisheries.

A minimal list of mangrove-associatedorgani sms of commerci al or sport val ueincludes oysters, blue crabs, spinylobsters, pink shrimp, snook, mullet,menhaden, red drum, spotted sea trout,gray and other snapper, tarpon,sheepshead, ladyfish, jacks, gafftopsailcatfi sh, and the jewfi sh. Heald and Odum(1970) pointed out that the commercialfisheries catch, excluding shrimp, in thearea from Naples to Florida Bay was 2.7million pounds in 1965. Almost all of thefish and shellfish which make up thiscatch utilize the mangrove habitat at somepoint during their life cycles. In addi­tion, the Tortugas pink shrimp fishery,which produces in excess of 11 millionpounds of shrimp a year (Idyll 1965a), isclosely associated with the Evergladesestuary and its mangrove-lined bays andri verso

11.5 AESTHETICS, TOURISM AND THEINTANGIBLES

One va 1ue of the mangrove ecosystem,which is difficult to document in dollars

·or pounds of meat, is the aesthetic valueto man. Admittedly, not all individualsfind visits to mangrove swamps a pleasantexperience. There are many others, how­ever, who place a great deal of value onthe extensive vistas of mangrove canopies,waterways, and associated wildlife andfishes of south Florida. In a sense, thismangrove belt along with the remainingsections of the freshwater Everglades andHi g Cypress Swamp are the only remai ni ngwilderness areas in this part of theUni ted States.

Hundreds of thousands of visitorseach year visit the Everglades NationalPark; part of the reason for many of thesevisits includes hopes of catching snook orgray snappers in the mangrove-lined water­ways, seeing exotic wading birds, croco­diles, or panthers, or simply discovering

75

what a tropical mangrove forest lookslike. The National Park Service,in anattempt to accommodate this last wish,maintains extensive boardwalks and canoetrai 1s through the mangrove forests nearFlamingo, Florida. In other, moredeveloped parts of the State, small standsof mangroves or manqrove islands provide afeeling of wilderness in proximity to therapidly burqeoning urban areas. A varietyof tourist attractions including FairchildTropical Gardens near Miami and TikiGardens near St. Petersburg utilizes theexotic appearance of mangroves as a keyingredient in an attractive landscape.Clearly, mangroves contribute intangiblyby diversifying the appearance of southFlorida.

11.6 ECONOMIC PRODUCTS

Elsewhere in the world, mangroveforests serve as a renewable resource formany valuable products. For a full dis­cussion of the potential uses of mangroveproducts, see de la Cruz (i n press a),Morton (1965) for red mangrove products,and Moldenke (1967) for black mangroveproducts.

In many countries the bark of man­groves is used as a source of tanni ns anddyes. Si nce the ba rk is 20'; to 30'; tanni non a dry wei ght basi s, it is an excell entsource (Hanlon et al. 1975). Silviculture(forestry) of mangrove forests has beenpracticed extensively in Africa, PuertoRico, and many parts of Southeast Asia(Holdridge 1940; Noakes 1955; Macnae 1968;Wal sh 1974; Teas 1977). Mangrove wood

makes a durable and water resistant timberwhich has been used successfully for resi­dential buildings, boats, pilings,hogsheads, fence posts, and furniture(Kuenzler 1974; Hanlon et al. 1975). InSoutheast Asia mangrove wood is widelyused for high quality charcoal.

Morton (1965) mentions that red man­grove fru i ts are somt i mes eaten by humansin Central America, but only by popula­tions under duress and subject to starva­tion. Mangrove leaves have variously beenused for teas, medicinal purposes, andlivestock feeds. Manqrove teas must bedrunk in small quantities and mixed withmilk because of the high tannin content(Morton 1962); the mil k bi nds the tanni nsand makes the beverage more pal atab 1e.

As a final note, we should point outthat mangrove trees are responsible forcontributing directly to one commercialproduct in Florida. The flowers of blackmangroves are of considerable importanceto the three million dollar (1965 figures)Flori da honey i ndust ry (Morton 1964).

Other than the honey industry, mostof these economic uses are somewhatdestructive. There are many cases inwhich clear-cut mangrove forests havefailed to regenerate successfully for manyyears because of lack of propaguledispersal or increased soil salinities(Teas 1979). We believe that the best useof Florida mangrove swamps will continueto be as preserved areas to supportwildlife, fishing, shoreline stabiliza­tion, endangered species, and aestheticvalues.

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CHAPTER 12. MANAGEMENT IMPLICATIONS

12.1 INHERENT VULNERABILITY

Manqroves have evolved remarkablephysiological and anatomical adaptationsenabling them to flourish under conditionsof high temperatures, widely fluctuatingsalinities, high concentrations of heavymetal s (Wal sh et a1. 1979), and anaerobicsoi 1s. Unfortunately, one of these adap­tations, the aerial root system, is alsoone of the plant's most vulnerable compo­nents. Odum and Johannes (1 975) havereferred to the aerial roots as the man­grove's Achi 11 es' heel because of thei rsusceptibility to clogging, prolongedfl oodi ng, and bori ng damage from i sopodsand other invertebrates (see section 6 fora discussion of the latter). This meansthat any process, natu ra1 or man -i nduced,which coats the aerial roots with finesediments or covers them with water forextended periods has the potential formangrove destruction. Bacon (1970) men­tions a case in Trinidad where the CaroniRiver inundated the adjacent CaroniMangrove Swamp during a flood anddeposited a layer of fine red marl in alarge stand of black mangroves which sub­sequently died. Many examp1 es of damageto mangrove swamps from human activitieshave been documented (see secti on 12.2).

One of the few natural processes thatcauses periodic and extensive damage tomangrove ecosystems is 1a rge hu rri canes(Fi gure 16). Crai ghead and Gi 1bert (1962)and Tabb and Jones (1962) have documentedthe impact of Hurricane Oonna in 1960 onparts of the man9rove zone of southFlorida. Crai9head and Gilbert (1962)found extensive damage over an area of100,000 acres (40,000 hal. Loss of treesranqed from 25% to 100%. Oamage occurredin three ways: (1) wind shearing of thetrunk 6 to 10 ft (2 to 3 m) above 9round,(2) overwash man9rove islands being sweptclean, and (3) trees dying months afterthe storm, apparently in response todamage to the prop roots from coati ngs bymarl and fi ne organic matter. The 1attertype of damage was most widespread, butrarely occurred in intertidal forests,presumably because the aerial roots wereflushed and cleaned by tidal action. Fishand invertebrates were adversely affected

by oXYgen depletion due to accumulationsof decomposin9 organic matter (Tabb andJones 1962).

Hurricane Betsy in 1965 did littledamage to mangroves in south Florida;there was also little deposition of siltand marl within man9rove stands from thisminimal storm (Alexander 1967). Lugo eta1. (1976) have hypothesized that severehurricanes occur in south Florida andPuerto Rico on a time interval of 25 to 30years and that mangrove ecosystems areadapted to reach maximum biomass and pro­ductivity on the same time cycle.

12.2 MAN-INOUCED DESTRUCTION

Destruction of mangrove forests inFlorida has occurred in various waysincluding outright destruction and landfilling, dikin9 and flooding (Fi9ure 17),through introduction of fine particulatematerial, and pollution damage, par­ticularly oil spills. To our knowledgethere are no complete, published docu­mented estimates of the amount of mangroveforests in Florida which have beendestroyed by man in this century. Ourconclusion is that total loss statewide isnot too great, probably in the range of 3to 5% of the ori9ina1 area covered bymangroves in the 19th century, but thatlosses in specific areas, particularlyurban areas, are appreciable. This con­clusion is based on four pieces of infor­mat ion. (1) Linda 11 and Sal oman (1977)have estimated that the total loss ofvegetated intertidal marshes and mangroveswamps in Florida due to dredge and fillis 23,521 acres (9,522 hal; remember thatthere are between 430,000 and 500,000acres (174,000 to 202,000 hal of mangrovesin Florida (see section 1.3). (2)Birnhak and Crowder (1974) estimate a lossof approximately 11 ,000 acres (4,453 halof mangroves between 1943 and 1970 inthree counties (Co11 i er, Monroe, andDade). (3) An obvious loss of mangroveforests has occurred in Tampa Bay, aroundMarco Island, in the Florida Keys, andalong the lower east coast of Florida.For example, Lewis et a1. (1979) estimatedthat 44% of the intertidal vegetation

77

Figure 16. Damaged stand of red and black mangroves near Flamingo, Florida, asit appeared 7 years after Hurricane Donna.

78

• _ .f",t.

, ....

Figure 17. Mangrove forest near Key West as it appeared in 1981 after beingdestroyed by diking and impounding.

79

including mangroves in the Tampa Bayestuary has been destroyed during the past100 yea rs. (4) Hea 1d (unpub 1i shed MS.)has estimated a loss of 2,000 acres (810hal of mangroves within the Florida Keys(not considered by Bi rnhak and Crowder1974). So while loss of mangrove ecosys­tems throughout Flori da is not over­whelming, losses at specific locationshave been substantial.

Diking, impounding, and long-termflooding of mangroves with standing watercan cause mass mortality, especially whenprop roots and pneumatophores are covered(Breen and Hi 11 1969; Odum and Johannes1975; Patterson-Zucca 1978; Lugo 1981).In south Florida, E. Heald (pers. comm.)has observed that permanent impoundment bydiking which prevents any tidal exchangeand raises water levels significantlyduring the wet season will kill all adultred and black mangrove trees. If condi­tions behind the dike remain relativelydry, the mangroves may survive for manyyears until replaced by terrestrial vege­tation.

Mangroves are unusually susceptibleto herbicides (Walsh et al. 1973). Atleast 250,000 acres (100,000 hal of man­grove forests were defoliated and killedin South Viet Nam by the U.S. military.This widespread destruction has been docu­mented by Tschirley (1969), Orians andPfeiffer (1970), Westing (1971), and acommittee of the U.S. Academy of Sciences(Odum et al. 1974). In many cases theseforests were slow to regenerate; observa­tions by de Sylva and Michel (1974) indi­cated higher rates of siltation, greaterwater turbidity, and possibly lower dis­solved oxygen concentrations in swampswhich sustained the most damage. Teas andKelly (1975) reported that in Florida theblack mangrove is somewhat resistant tomost herbicides but the red mangrove isextremely sensitive to herbicide damage.He hypothes i zed that the vul ne rabil i ty ofthe red mangrove is related to the smallreserves of viable leaf buds in this tree.Following his reasoning, the stress of asingle defoliation is sufficient to killthe enti re tree.

Although mangroves commonly occur ina reas of rapi d sedi mentat i on, they cannotsurvive heavy loads of fine, floculentmaterials which coat the prop roots. Theinstances of mangrove death from thesesubstances have been bri efly revi ewed byOdum and Johannes (1975). Mangrove deathsfrom fi ne muds and marl, ground bauxiteand other ore wastes, sugar cane wastes,pulp mill effluent, sodium hydroxidewastes from bauxi te proces sing, and fromintrusion of large quantities of beachsand have been documented from variousareas of the worl d.

12.3 EFFECTS OF OIL SPILLS ON MANGROVES

There is little doubt that petroleumand petroleum byproducts can be extremelyharmful to mangroves. Oamage from oilspill s has been reviewed by Odum andJohannes (1975), Carlberg (1980), Ray (inpress), and de la Cruz (in press, b).Over 100 references detailing the effectsof oi 1 spi 11 s on mangroves and mangrove­associated biota are included in theserevi ews.

Petroleum and its byproducts injureand ki 11 mangroves in a variety of ways.Crude oil coats roots, rhizomes, and pneu­matophores and disrupts oxygen transportto underground roots (Baker 1971).Various reports suggest that the criticalconcentration for crude oil spills whichmay cause extznsive damage is between 100and 200 ml /m of swamp surface (Odum andJohannes 1975). Petroleum is readilyabsorbed by 1i pophyl i c substances on su r­faces of mangroves. This leads to severemetabolic alterations such as displacementof fatty molecul es by oil hydrocarbonsleading to destruction of cellular permea­bility and/or dissolution of hydrocarbonsin lipid components of chloroplasts (Baker1971).

As with other intertidal communities,many of the invertebrates, fishes, andplants associated with the mangrove com­munity are highly susceptible to petroleumproducts. Widespread destruction oforganisms such as attached algae, oysters,tunicates, crabs, and gobies have beenreported in the 1iterature (reviewed by de

80

la Cruz in press, b; Ray in press).

Damage from oil spills follows apredictable pattern (Table 7) which mayrequire years to complete. It is impor­tant to recognize that many of the mostsevere responses, including tree death,may not appear for months or even yearsafter the spill.

In Florida, Chan (1977) reported thatred mangrove seedl i ngs and black mangrovepneumatophores were particularly sensitiveto an oi 1 spi 11 whi ch occurred in theFlorida Keys. Lewis (1979a, 1980b) hasfollowed the long-term effects of a spi 11of 150,000 liters (39,000 gal) of bunker Cand diesel oil in Tampa Bay. He observedshort-term (72-hour) mortality of inverte­brates such as the gastropod Melongenacorona and the polychaete Laeonereisculveri. Mortal ity of all three speciesof mangroves began after three weeks andcontinued for more than a year. Sub­lethal damage included partial defoliationof all species and necrosis of blackmangrove pneumatophores; death dependedupon the percentage of pneumatophoresaffected.

In addition to the damage from oilspi 11 s, there are many adverse impacts onmangrove forests from the process of oi 1exploration and drilling (Table 8). Thistype of damage can often be reducedthrough careful management and monitori ngof dri 11 i ng sites.

Although little is known concerningways to prevent damage to mangroves once aspill has occurred, protection of aerialroots seems essential. Prop roots andpneumatophores must be cl eaned with com­pounds whi ch wi 11 not damage the pl anttissues. Dispersants commonly used tocombat oil spills are, in general, toxicto vascular plants (8aker 1971). If pos­sible, oil laden spray should not beallowed to reach leaf surfaces. Damageduring clean-up (e.g., trampling, compac­tion, bulldozing) may be more destructivethan the unt reated effects of the oi 1spill (de la Cruz in press, b).

12.4 MAN-INDUCED MODIFICATIONS

In south Florida, man has been re­sponsible for modifications which, whilenot ki 11 i ng mangroves outri ght, have al­tered components of the mangrove ecosys­tem. One of the most widespread changesinvolves the alteration of freshwaterrunoff. Much of the freshwater runoff ofthe Flori da Evergl ades has been di vertedelsewhere with the result that salinitiesin the Everglades estuary are generallyhigher than at the turn of the century.Teas (1977) points out that drainage inthe Miami area has lowered the water tableas much as 2 m (6 ft).

Interference with freshwater inflowhas extensi ve effects on estuaries (Odum1970). Florida estuaries are no excep­tion; the effects on fish and invertebratespecies along the edge of Biscayne andFlorida Bays have been striking. Themismanagement of freshwater and itseffects on aquatic organisms have beendiscussed by Tabb (1963); Idyll (1965a,b);Tabb and Yokel (1968) and Idyll et al.(1968). In addition, Estevez and Simon(1975) have hypothesized that the impactof the boring isopod, Sphaeroma terebrans,may be more severe when freshwater f1 owsfrom the Everglades are altered.

One generally unrecognized sideeffect of lowered freshwater flow and saltwater intrusion has been the inland expan­sion of mangrove forests in many areas ofsouth Flori da. There is documented evi­dence that the mangrove borders ofBiscayne Bay and much of the Evergladesestuary have expanded inland during thepast 30 to 40 years (Reark 1975; Teas1979; Ball 1980).

Sections of many mangrove forests insouth Florida have been replaced by filledresidential lots and navigation canals.Although these canal systems have not beenstudied extensively, there is some evi­dence, mostly unpublished, that canals arenot as productive in terms of fishes andinvertebrates as the natural mangrove­lined waterways which they replaced.

81

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

Stage

Acute

o to 15 days

15 to 30 days

Chronic

30 days to 1 year

1 year to 5 years

1 year to 10 years (?)

10 to 50 years (?)

Observed impact

Deaths of birds, turtles, fishes, andinvertebrates

Defoliation and death of small mangroves,loss of aerial root community

Defoliation and death of medium-sizedmangroves (1 - 3 m), tissue damage toaeri a 1 roots

Death of large mangroves (greater than3 m), loss of oiled aerial roots, andregrowth of new roots (often deformed)

Recolonization of oil-damaged areas bynew seedl i ngs

Reduction in litter fall, reduced re­production, and reduced survival ofseedl ings

Death or reduced growth of young treescolonizing spill site (?)

Increased insect damage (?)

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).

cow

Stage

Pre-exploration

Site preparation

Drilling

Production

Oil spills

Activity

Seismic surveysClearing of survey linesDrilling "shot lines'

Canal excavationDredge spoil depositionRoad construction

Increased activity at siterelated to drilling

Construction of platformsConstruction of pipelinesMaintenance dredgingPlacement of tanks and

other equipment

Oil leaks and spills dueto well blow-out, pipe­line breakage, careless­ness, and barge rupture

Clean-up activities

Impacts

Crushing and clearing vegetationVehicle track compactionDamage to natural levees

Loss of habitat in disturbed areasAlteration of water flow pathwaysIncreased turbidity, higher rates of sed-

imentation, and lowered dissolved oxy­gen in nearby waters

Continued high turbidityRelease of toxic substancesDisplacement of wildlife

Continued high turbidityLoss of additional habitatFurther changes in wetland drainage pat­

terns from pipeline constructionRelease of toxic substancesOil spills

Destruction of plant and animal popula­tions

Alteration of ecosystem processes suchas primary production and decomposition

Introduction of persistent toxic substan­ces into soil s

Weinstein et al. (1977) found that artifi­cial canals had lower species diversity ofbenthic infauna and trawl-captured fishesand generally finer sediments than thenatural communities. Courtney (1975)reported a number of mangrove-associatedinvertebrates which did not occur in theartificial channels.

Mosquito production is a seriousproblem in black mangrove-dominated swampsin Florida (Provost 1969). The salt marshmosquitos, Aedes taeniorhynchus and ~.

sollicitans, do not reproduce below themean high tide mark and for this reasonare not a serious problem in the inter­tidal red mangrove swamps. Mosquitos laytheir eggs on the damp soil of the irregu­larly flooded black mangrove zone; theseeggs hatch and develop when flooded byspring tides, storm tides or heavy rains.As with the "hi gh marsh" of temperatelatitudes, there have been some attemptsto ditch the black mangrove zone so thatit drains rapidly after flooding.Although properly designed ditchinq doesnot appear to be particularly harmful tomangrove swamps (other than the areadestroyed to dig the ditch and receive thespoil), it is an expensive practice andfor this reason is not widely practiced.Properly managed diking can be an effec­tive mosquito control approach with mini­mal side effects to black mangroves(Provost 1969). Generally, ditching ordiking of the intertidal red mangrove zoneis a waste of money.

Mangrove swamps have been proposed aspossible tertiary treatment areas forsewage (see discussion by Odum andJohannes 1975). To our knowledge, thisalternate use is not currently practicedin south Florida. Until more experimentalresults are available on the assimilativecapacities and long-term changes to beexpected in mangrove forests receivingheavy loads of secondary treated sewage,it would be an environmental risk to usemangrove forests for this purpose.

In many areas of the world mangroveswamps have been converted to other usessuch as aquaculture and agriculture (seede la Cruz, in press, a). Although some

of the most product i ve aquaculture pondsin Indonesia and the Philippines arelocated in former mangrove swamps, thereis some question whether the originalnatura 1 system was not equa 11 y product i vein terms of fi sheri es products at no costto man (Odum 1974). Conversion toaquaculture and agriculture is cursed witha variety of problems including subsequent1and subsi dence and the "cat cl ay"problem. The latter refers to thedrastically lowered soil pH which oftenoccurs after drai nage and has been tracedto oxidation of reduced sulfur compounds(Dent 1947; Tomlinson 1957; Hesse 1961;Hart 1962, 1963; Moorman and Pons 1975).Experience in Africa, Puerto Rico. andSoutheast Asia confi rms that mangroveforests in their natural state are morevaluable than the "reclaimed" land.

12.5 PROTECTIVE MEASURES INCLUDINGTRANSPLANT! NG

Protect i on of mangroves inc 1udes (l)prevention of outright destruction fromdredging and filling; (2) prevention ofdrainage, diking and flooding (except forcarefully managed mosquito control); (3)prevention of any alteration of hydrologi­cal ci rculation patterns, particularlyinvolving tidal exchange; (4) preventionof introduction of fine-grained materialswhich might clog the aerial roots. such ascl ay. and sugar cane wastes; (5) preven­tion of oil spills and herbicide spraydriftage; and (6) prevention of increasedwave action or current velocities fromboat wakes, and sea wall s.

Where mangroves have been destroyed,they can be replanted or suitable alter­nate areas can be planted, acre for acre,through mitigation procedures (see Lewiset al. 1979). An extensive body ofliterature exists concerning mangroveplanting techniques in Florida (Savage1972; Carlton 1974; Pulver 1976; Teas1977; Goforth and Thomas 1979; Lewis1979b). Mangroves were initially plantedin Florida at least as early as 1917 toprotect the overseas railway in theFlori da Keys (Teas 1977).

Both red and b1 ack mangroves have

84

been used in transplanting. As we men­tioned in section 11, black mangroves seemto have certain advantages over red man­groves. Properly designed plantings areusually 75% to 90% successful, althoughthe larger the transplanted tree, thelower its survival rate (Teas 1977).Pruning probably enhances survival oftrees other than seedl i ngs (Carl ton 1974).Important consi derati ons (Lewi s 1979b;Teas 1977) in transplanting mangroves are:(1) to plant in the intertidal zone andavoid plantin9 at too high or too low anelevation, (2) to avoid planting where theshoreline ener9Y is too great, (3) toavoid human vandalism, and (4) to avoidaccumulations of dead sea grass and otherwrack •

Costs of transplanting have beenvariously estimated. Teas (1977) suggests$462 an acre ($1, 140/ha) for un rootedpropagules planted 3 ft (0.9 m) apart,$1,017 an acre ($2,500/ha) for establishedseedlings planted 3 ft (0.9 m) apart and$87,500 ($216,130/ha) for 3 year-old nur­sery trees planted 4 ft (1.2 m) apart.Lewis (1979b) criticized Teas' costs asunrealistically low and reported a projectin Puerto Rico which used establishedseedl ings at a cost of $5,060 an acre($12,500/ha); he did suggest that thiscost could be cut in half for largerprojects.

12.6 ECOLOGICAL VALUE OF BLACK VS. REDMANGROVES

One unanswered question of currentinterest in Florida concerns the ecologi­cal value of black mangrove forests com­pared to intertidal red mangrove forests.In many respects, this is identical to thelIhi gh rna rs h" versus III ow rna rsh" debate intemperate wetl ands. One hypothet i calargument which has been presented fre­quently in court cases during the pastdecade suggests that black mangroveforests have less ecological value thanred mangrove forests to both man andcoastal ecosystems. This argument isbased on an apparent lack of substantialparticulate detritus export from blackmangrove forests above mean high tide and

the generally perceived lack of organisms,particularly gamefishes, which use blackmangrove forests as habitat.

The counter argument states thatblack mangrove forests are important forthe support of wil dl ife and the export ofsubstantial quantities of dissolvedorganic matter (DOM). Lugo et al. (1980)provide evidence that black man9roveforests do, in fact, export large quanti­ties of DOM. They poi nt out that (1)black man9rove leaves decompose morerapidly than red mangrove leaves and thusproduce rel ati vely more DOM and (2) abso­1ute export of carbon from these forests,on a statewide scale, is equal or greaterthan from red man9rove forests.

12.7 THE IMPORTANCE OF INTER-COMMUNITYEXCHANGE

From previous discussions (sections 6and 7.5 and Appendi ces B, C, 0 and E) itis clear that many species of fishes,invertebrates, bi rds, and mammal s movebetween mangrove forest communities andother habitats including sea grass beds,coral reefs, terrestrial forests, and thefreshwater Everglades. For example, thegray snapper, Lutjanus griseus, spendspart of its juveni le 1ife in sea grassbeds, moves to mangrove-lined bays andrivers, and then migrates to deeper waterand coral reefs as an adult (Croaker 1962;Starck and Schroeder 1971). The pi nkshrimp, Penaeus duorarum, spends its juve­nile life in mangrove-lined bays andri vers before movi ng offshore to theTortugas grounds as an adult. During itsjuvenile period it appears to move backand forth from mangrove-dominated areas tosea grass beds. The spi ny lobster,Panul i rus argus, as a juveni le frequentlyuses mangrove prop root communities as arefuge; when nearing maturity this speciesmoves to deeper water in sea grass andcoral reef communities (see discussionsection 6.1). Many of the mammals (sec­tion 10) and birds (section g) move backand forth bet ween mangrove communi ties anda variety of other environments.

These are only a few of many

85

examples. Clearly, mangrove ecosystemsare linked functionally to other southFlorida ecosystems through physical pro­cesses such as water flow and organiccarbon flux. As a result, the successfulmanagement and/or preservation of manyfishes, mammals, birds, reptiles, andamphibians depends on proper understandingand management of a variety of ecosystemsand the processes that link them. Savingmangrove stands may do the gray snapper1itt 1e good if sea grass beds aredestroyed. Pink shri mp populations willbe enhanced by the preservation of seagrass beds and mangrove-lined waters, butshrimp catches on the Tortugas groundswill decline if freshwater flow from theEverglades is not managed carefully (Idyllet al. 1968). Successful management ofsouth Flori da mangrove ecosystems,including their valuable resources, willdepend on knowledgeable management of anumber of other ecosystems and theprocesses which link them.

12.8 MANAGEMENT PRACTICES: PRESERVATION

8ased on years of research in southFlorida and based on the information

reviewed for this publication, we haveconcluded that the best management prac­tice for all types of Florida mangroveecosystems is preservation. Central tothis concept is the preservation ofadjacent ecosystems that are linked signi­ficantly by functional processes. Thecontinued successful functioning of themangrove belt of southwest Florida ishighly dependent on the continual exis­tence of the Everglades and Big CypressSwamp in an ecologically healthy condi­tion.

At no cost to man, mangrove forestsprovide habitat for valuable birds, mam­mals, amphibians, reptiles, fishes, andinvertebrates and protect endangeredspecies, at least partially support exten­sive coastal food webs, provide shorelinestability and storm protection, andgenerate aesthetically pleasing experi­ences (Figure 18). In situations whereoverwhelming economic pressures dictatemangrove destruction, every effort shouldbe made to ameliorate any losses eitherthrough mitigation or through modifieddevelopment as described by Voss (1969)and Tabb and Heald (1973) in which canalsand seawalls are placed as far to the rearof the swamp as possible.

86

1•. If I...j

Fi gu re 18. Mangrove is 1ands in Flori da Bay nea r Upper 11a tecumbe Key. Note theextensive stands of seedling red mangroves which have become established (19Bl)after a long period without major hurricanes. Mangrove islands in the FloridaKeys tend to expand during storm-free intervals.

87

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Bacon, P.R. 1970. The ecology of CaroniSwamp. Special Publication, CentralStatistical Office, Trinidad. 68 pp.

Bader, R.G., and M.A. Roessler. 1971.Progress report on an eco1ogi ca1study of south Biscayne Bay and CardSound. Rosenstiel School of Marineand Atmospheric Science, Univ. ofMiami, to U.S. Atomic Energy Commis­sion and Florida Power and LightCompany.

Bader, R.G., and M.A. Roessler. 1972.Progress report on an eco1ogi ca1study of south Biscayne Bay and CardSound. Rosenstiel School of Marineand Atmospheric Science, Univ. ofMiami, to U.S. Atomic Energy Commis­sion and Florida Power and LightCompany.

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Ahearn, D.G., F.J. Roth, and S.P. Meyers.1968. Eco109Y and characteri za t i onof yeasts from aquatic regions ofsouth Florida. Mar. Biol. 1: 291­308.

Albert, R. 1975. Salt regulation inhalophytes. Oecologia 21: 57-71.

Alexander, T.R. 1967. Effect of Hurri-cane Betsy on the southeastern Ever­9lades. Q. J. Fla. Acad. Sci. 30:10-24.

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104

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105

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APPENDIX A. Summary of the site characteristics and samplingmethodology for fishes in: A-l - mangrove-fringedtidal streams and rivers, A-2 - mangrove-linedestuarine bays and lagoons, and A-3 - mangrove­lined oceanic bays and lagoons.

106

Table A-l. Site characteristics and sampling methodology for fishesin mangrove-fringed tidal streams and rivers.

Meandepth; Number

Salinity Temperature tidal speciesLoca ti on range range range Subs tra te Benthic vegetation Sampling methods Frequency recorded

North Ri ver; 0-27 0/00 15.40 _33.20 C 1 ffi; largely exposed Scattered~ Bag seine. throw r~onthly. Sept. 55Tabb 1966, 0.5 m 1imestone. and maritima near mouth nets, dip nets, 1965 throughOdum 1970 sand banks; traps, pound net, Sept. 1966

undercut man- fish poison, rod & (Tabb)grove peat reel, trammel net.

set lines

Cross Bayou 3.2-29.8 0/00 13.00 _31.50 C Max. Hard muddy sand Sparse Bag seine; minnow Monthly, Sept. 60..... Canal (Boca depth Enteromorpha se; ne 1957 through0 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. 47stream, 1 m; range black net &rote- 1972 throughstream enterin9 not given none for single Dec. 1972Fahka Union standing cropCana 1; Carter estimateet al. 1973

Unnamed streams 16-32 0/00 13.20 _37.loC 1.1 m; Thick organic Dense Thalassia Gill nets; hoop Weekly, gill 52near Turkey 0.5 m mud-gi 11 net & testudinum at nets; traps nets; bimonthly,Point, Biscayne trap sites; mouth others; AugustBay; Nugent culvert at hoop 1968 through1970 net sites Dec. 1969

Total IIIOnly taken in SE Fla. -23

88

Table A-2. Site characteristics and sampling methodology for fishesin mangrove-lined estuarine bays and lagoons.

­aco

location

FahkahatcheeBay, 740 ha;Yokel 1975b,Carter et a 1.1973

Fahka UnionBay, 186 ha;Carter et al.1973

Rookery Bay,419 ha;Yokel 1975a

Marco IslandEstuary; Yokel1975b; Wcin­steinetal.1977

Whlte\·/aterBay, Clark1970

Salinityrange

Average ­15-37 0/00,low of 1 0/000recorded byYoke1, Sept.1971

5-35 0/00subject tosporadicmassivefreshwaterinp~ts fromGAC drai nagecana 1s

8.9-38 5 0/00

19 0/00 Sept.1971; other­wi se over a4-yr period29-39 0/00

2.9-29.3 0/00

Tempera turerange

21 0 _31 oC(18 ke1 6;

23.5 -32 C(Carteret al.)

~'ean

depth;tidalrange

1. 2 m;

0.9 m;.55 m

Not given

Shallowstations­1 Ill, deepstalions­0.8-1.0 m;0.6 m

Substrate

Generallymuddy, somesand & shell

Muddy,occasionalsandy areaor oys terbac

Hud, sand,shell

Mud, muddysand. shellysand

Peat, silt,marl P. shell,sand & shell

Benthic vegetation

Extensive areas ofHalodule wrightii ,lhalassia testudinumin northern portion

Little seagrass,high standing cropof green algae

Halodule wrightii,ThaTassia testudinum,l~al.1WJ.!.iJa engelmannli

Hillodulc wr{9htitbedS in sha lowback-bays of man­grove complex;Thalassia not welldeveloped

Halodule ~ti~,Udotea conylutinata,Chara hornC'rllanOl,Qa_sy~ p-e-J{c'cfla ta,Gracilaria sp.,Ha10phila baillonis

Sampling methods

Vegeta ted, mud,sand/shell bottomssampled by ottertrawl (Yokel); 2bay seines, ottertrawl, surfacetrawl (Carteret a1. )

2 bag seines,otter trawl,surface trawl

Vegeta tM, mud,sand/shell bottomssampled by othertrawl

vegetuted, mud,sand/shell bottomssamplpd by ottertrawl (Yokel);otter trawl (Wein­stein et al.)

Roller frame trawl

Frequency

Monthly, July1971 throughJuly 1972(lokel);Monthly, Jan.1972 throughDec. 1972(Carter et a1.)

Month ly, Jan.1972 throughDec. 1972

Monthly, June1970 throughJuly 1972

~1onthly, July1971 throughJuly, 1972(Yokel);Monthly, July1971 throughJan. 1975(Weinsteinet a1. )

Monthly - 8s ta ti onsSept. 1968throughNov. 1969

Numberspeciesrecorded

47(Yokel)

89'

64

59(Yokel)

82­(Wein~stein)

67

a89 species in rahkahatc.:hee and rahka Union Bays combined.bGulf American Corporation.

Total 117

Table A-3. Site characteristics and sampling methodology for fishesin mangrove-lined oceanic bays and lagoons.

Meandepth; Number

Sal inHy Temperature tidal speciesLocation range range range Subs tra te Benthic vegetation Sampling methods Frequency recorded

._--~

Old Rhodes Average Apr; 1- Average for 0.61 m; Seagrasscs: Visual counts. Monthly, 1973 31Key lagoon. June 1973 - Apr. -June

oO.S m Thalassia testudinum traps. hool< and

Holm 1977 37 0/00 1973 - 28 C range 1ine

Porpoise lak.e. 27.8-49.6 0/00 16.60 -32.20C f1ax. Carbonate Seagrasses: exten- Suction sampler, Monthly, Apr;l 64Florida Bay, depth mud & shell sive Thalassia slednet. pushnet, 1965 throughHudson et a1. 2.1 m; fragments testudlnum. sparse beach seine, cast Jan. 19681970 Italodule wrigh_'tiJ.. net. roller-frame

trawls, hook &line

~ Southern 5.0-43.8 0/00 13.50 _38.70C' Range Mud. sand. Seagrasses: Otter trawl ~lonth1y. Ju ly 750 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 109Florid~ Oay author sta tes each balloon otter 1973 throu9h315 km , representative trawl June 1974 atSchmidt 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 byhabitat type. Key to numbered references appearsat the end of the table. Diet items listed inorder of decreasing Lrnportance.

110

:a~ily and Species

orcctolobidae -carp€t snar},sGinqlynosto~ cirratC3nurse sha:.-k

Carcha:.-hinicaerequi::::. shari:s

Carcharhinus leucasbull shark

Carcharhinus li~ tusblac;;tiI) shark

!ieca::.rion breviros'trislc=n sha.d:

Sphyrnicac - h~-erhead

sharksSpn"lrna t i huro ­honnethcad

?ris'tidaa - sawfishes?ristis oectinaUis=alltooth sawfish

R.~ino~tid<le ­guit.arCishes

Rhinobatos lent i­cino~us ntlanticgui.Uirfisil

+

Refercnc,-,

1, 5, 7

8

II

-i. 5, 7

2, 5

5, 1S

Olt1t

Fish. cEr;halopocs, :-:olluscs,s~i~?, sea urchins

J ...·J~nile3' fi::;n (~~,:'ol:noco:'i'Js, :-:'.15i1 cE:l:'al•.ls,5r~~oa~~ia ca't~o~us.

:";icro'::o:;on unOulatus). crus­taceans incluuing ~E~~E:C

shri=?, blu~ c~abs

:"ish (Car~!'..:·: sp .• Ccn'tro:.o=c;sund~i-~1i5, Chilo "lc~Lrus

scho&~fi, nrius ~, ~cto­

rhrys trigonnus L:~oCon

rho~ic€s), cra:'s

YounG: crustac:ans. fishhcults' ~ish, crustaceans

~antis shri=?, s:'ri=p, lSOpoCS,bar~ac1cs, oi~alve oo11uscs,cep:'alopoCs, fi~:'

Fish. benthic c:.-ustaceans

Did:tei.,.fc:nc<::'

?.ar:dall 1967Cl.:;.r:: s; -.-on

:::c1-.:::lid'C 1965Eohl}:€: S

C1Hpl:.n l'?ES

cc.= 19713

clar;: <;

\-on Sch:::liet.i<?6:'

~!lcall 1967C1<lrk & ':on5c~..=id~ 1965

?ohl:':e SCha?11n 1968

3Oh1ke toChaplln 1968

To::?£:cinidae - electricraJ's

!larcinc: brasiliensis­lesser electric re.'l

?..ajidac - SJ.""...atC5

?..aja~

rOlrleel sJ.:":li:e

+ + I, !1. 18

CruStacCA. ri5h, annelids R~id 195"

Das'latidae - stingraysD5s"latis a=cricana ­southam stingray

Das1atis sabina;;t1a:ltic stingray

+ + + :2,4, 5, 7

2. B. 13,17

'ishes. sipunculid anc pol:;- ;::a~call 1967chaste ~o~. crab~, bivalves,shr~p. =antis shricp

3cnthic invertebrates inc1u- Darnell 1958ding bivalves. xanthid a~d

port~~id crabs, silri~ps,

a~~hipods, annelids. chiro~o-

~irl 1arv.1e

G\'r:!llura cicrura ­s=ooth butterfly,ay

Uroloohus ia::.aicensis­¥e11a~ stingray

+ 17 ?i5h, l::olluscs. annelids, ?eterson •shrio? ot..l1er s::::.a1l peterson 1979crusUlceans

+ ! ?.robaDl·", s;:.all burro-"'ing BOhH:e ,inver1:ebrates Chaplin 1908

.-

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

111

:a=ily and Spect~s Refer.:onceDid

R.,i ~ r~nc" C~nl

~iliooaticae - eagle rays';etoDatus na.rina.ri ­spotted eagle ray

2 3.ohlr.o:1 !O

!!.3plln 19&3

LepisosteidaeLepisosteusPlorida gar

garsolatZJ::hineus - I" 2. 1. 13,

1SFish (~eiliids. cyprinodonts. Od~

sr"",ll cenrrarchids), crustaceans(caricean 5hrl~?}.inseer lar;a~

1971

Elopidae - tarf'O:'ls::10p5~­lac]!ish

?~~ale~s atl~~t1ca ­t.L"1'On

;'lbuli::h.e - hone fishes~ vulP$$­

bonefi:Jh

h...,guillidae - eels~"9uilla rostratar..::.e.rican eel

O?hichthidac - sn..1o;.eeels

~(ro:;his cunctatusspecklel:! WOr.:! eel

aascanichthy~ scuti­~ Ahip eels

Onhichthus 900esi ­Shri::;> eel

Clupeidae - herringsBrevoortia 5~thi ­yellv"'fin s~

•

•

2, J, 7,8. 13, 15

7, e. 13,15

4, S

~, 13

2, 3. n,18

)

3, 17

2, S. 17

< ~5 ~: zoo?la~to:'l. c~aeto­

Sna~~$. ?olychaet~

·...o~.....s> ~5 ~: caridea-~ 5 pe~aeid

8hri~, various s=all:hh

< ~5 ~: ?la.~kto~ (~lC!o;oid

co~ 's)

j~~eni1e5: fish (G~~ia,

;~"dulus ~cteroclit~. ~~;il

ce~halus), crus~ce~"5 (ostra­c -$, carlc!eil~ s;~.ri=;))

adults: ~id~ ~arlety o~ tis~,

c~a~~, shri=? ctenophcres,insec=

c1a~, snails, shri~, s~ll

fish

50-200 ~, ~?hi~s. isop0d3180-472 ~: xanthic crabs.caridea~ shri~? fish(Loohosooius ~luri~oi~e5)

?o1ychaetes, 3ranchio5~

car~~~, s~~d cr~hs

~= 1971Austin I>';'.1stb 1971

O:l:c. 1971':'\l.!O,:i:l &

.:.us~in 1971

30hlke toCh...plin1968

Cdu::. 1971

;;prin;er '"\l:oodk-':J...~

1960,Reid 195':

O:>liSil::'e. air:tIrea to":", r"S. .Juv­enilell inh..!;)! ~sha1lc~ orac~ish

?DOls lcr~ in oxyqe.n,often containing~:S (nac~ 1962)

Y~rs of thisfa.:lily b=ro-...in ::::;"Jd or sa."ld.u.,eersa=pled ~7

=cst :rti:.ods(Eohlke & Chaplin1908)

Darnell1958

3re~~rtia patronus ­Gulf ~hacen

12 38-~8 ~: phyto?1ankton, zoo­?1a."'l1:to~, plant frao;=oc:n=.d!'! ritus85-103 =: organic :::att.er, silt.ciato~, for~ni!era~s, C0?epoC5

112

Family and Species

Habitat rype

Reference DietDiet

Reference Comments

Harengula pensacolaescaled sardine

Opi~thonema oglinum ­Atlantic thread herring

Sardinella anchoviaSpanish sardine

Engraulidae - anchoviesAnchoa cubana -Cuban anchovy

Anchoa hepsetus ­striped anchovy

Anchoa lamnrotaenia ­bIgeye anchovy

+

+

+

+ +

+ +

+

+

+

+

2, 3, 8,13

2, 3, 5,13, 17

17

2, 16

2, 3, 13,16, 17

5

30 m~: planktonic copepods,zoea, naupli!, larval fish64-96 mm: amphipods,harpacticoid copepods, isopods,mysids, chironomid larvae

Copepods, polychaetes, shrimp,fishes, crab larvae, mysids

Ostracods, copepods

32-114 rom: copepods, isopods,mysids, caridean shrimp, smallbivalves

Odum 1971

Odum 1971

Springer &\.o:oodburn1960

Springer &~,loodburn

1960

Anchoa mitchillibay anchovy

+ + + 1, 2, 3,5, 7, 8,13, 16-18

<25 rom: microzooplankton Odum 197131-62 mm: amphipods, zooplank-ton, mysids, ostracods, plantdetritus, copepods, sw~ll molluscs,chironomid larvae

Synodontidae ~

lizardfishesSyoodus .~etens

inshore lizardfish

Catostomidae ~ suckersJ~zson sucetta ­lak-e- -c1iUbslicKC'r-

ictnluriJae - freshwatercatfish

Ictalutus natalis ­yellow bll1lhead

Noturus zy!inus ­tadpole :uadtom

+

+

+

+

+ + 1-3, 5, 8, Small fish, crabs, shri~p,

17, 18 polychaete worn.s

14

14

Odum 1971

A freshwaterstray

A freshwaterstray

A freshl\'uterstray

Arriidae - sea catfishesArlus feli_~ - sea catfish

Bagre ~rinus ­gaff topsail catfish

+

+

+

+

+ 2, 3, 5,7, 8, 13,17

2, 8, 17

100 W~: copepods, zooplanktonamphipods, mysids, chi ronomidlarvae, isopods, small crabs100-200 rom: benthic inverte­brates200-330 rom: crabs, anphipods,mysids, fishes, bark, crayfish,catidean and penaeid shrimp

262-445 nun; blue ctabs, smallfishes

Odum 1971

Odum 1971

Battachoididae ­Opsanus beta ­Gulf toadfisil

toad fishes+ + + 1-3, 5,

7, 12, 13,15, 17, 18

18-60 m:n: amphipods, chironomidlarvae, mysids, isopods, few fish>60 mm: caridean shrimp, xanthidcrabs, snapping shrimp, mussels,fish, mangrove hark

113

Odum 1971 Salinities10 0100 >(Odum 1971)

Family and Species DietDiet

Reference Comments

Porichthys porosissimusAtlantic midshipman

+ 3, 18

Gobiesoeiclae - elingfishesGobiesox strumosusskl11etflsh

Ogcocephalldae - bat fishesOgcocephalus nasutusshortnose batfish

Ogeocephalus radiatuspolka-clot batfish

Gadidae - codfishesUrophyciS floridanusSouthern hake

Ophidiidae - cusk-eels,brotulas

Gunterlchthys longipenisgold brotula

Ogilbia eayorumkey brotula

Ophidion holbrookibank cusk eel

+ + + Z. ] . 5. 8 10-32 mm: amphipods, isopods, Odurn 1971chlronomid larvae

+ + 18 50a11 bivalves, gastropods, Reid 1954polyehaetes

+ Z. 11. 17,18

+ 12 Amphlpods, isopods, mysids, Springer & A species moredecapod shrimp, polyehaetcs, \~oodbu[n common at ="insect larvae, fishes (Lagodon 1960 northerlyrhonbo1des, Paralichthys latitudesalbigutea)

+ 17

+ + I, ]

+ ]

Exocoetidae - flying­fishes, ha1fbeaks

Chriodorus atherinoideshardhead ha1fbeak

Hyporhamphus unifasciatushalfbeak

•

•

•

5

2, 3, 5 ju~eni1es: zooplankton including Carr ~

crab megalops, ve1igers, cope- hda~s 1973pods130-199 IIUII; epiphytic algae,detritus, seagras8

Belonidae - needle fishesStron9ylura marinaAtlantic needle fish

Strongy1ura notataredfin needle::ish

Strongylura timucutimucu

Tylosurus crocodilus­houndfish

•

•

•

•

•

•

•

•

•

2, 7, 15

2, 3, 5,8, 13

2, 3, 11

11

357-475 nffi: s~all fis~es,

insects, shrimp, small amolli~ts

of vascular plant material andalgae

In grassbe::lsJuveniles: po1yc~aete worms,cumaceans, fishAdults, fish, pri~ri1y

ntnC'rinids

159-37B ~: anchovies. shrim?

250-1320 rom: fishes, shrimp

114

Darnell1958

Brook1975

Ra:ldall1967

Randall1967

Primarily ins~ore

species~freely

enters fresh­",ater (Randall1967)

Open water andinshore surfacewater inhabitant(Voss et al.1969)

Family and Species Reference DietDiet

Reference Comments

Cy?rinodontidae - killi­fishes

Adinia xenica ­~d killifish

+ 2.6,13-15 Plant detritus, diatoms,amphipods, harpacticoidcopepods, insects

Odur.l 1971

Cyprinodon varicqatussheelJshead cinna...

floridichthys carpiogoldspotted ~illifish

+ , , 2, ,, ., Plant detritus, algae, O<I~, 197113-15 nell'latodcs, small c.,-:ustaceans

, 2, J, 8, ""..nphipods, ostracods, isoporls, 0<1= 197113 copcpods, chironoll'.id larvae,

nerr.atodes, plant detritus, algae

Fund~lus conflucntusnarsh ~illifish

Fundulus chrysotusgolden topni:lno....'

+

+

2,8,13-15 Caride.:an shrimp, small fish, Odum 1971(Gambusia affinisl, ~phipods,

isopods, a~ult & larval insects,copcpods, mysids, ostracoCs,algal filaments

14. 15 Rare in mangrovezone, headwaterpools on,ly

Fur~ulus grandi~

CuI f killifish2,8,13-15 I\mphipods, isopoos, xanthid

crabs, chirononid larvae,terrestrial insects, snails,algae, small fish (poeciliidsl

Odwn 1971

fundulus hctcrocli~us

11ummichog------~+ ,

14-1 "i

11-15

Sm.Jll crustaceans l~phipods,

isopods, ostracods, t2n~ids,

cope?Ods), detritus, polychaeteworms, insects, snails, inver­tcb.,-:ate eggs

Peterson &

Peterson1979

Pri..m.l.rily afreshwater form,nead....·ater poolsonly

Primarily fresh­water, cammon inpools in headwaterregions

~3E?2..~btucfin killifish

+ 2,8. )3·1', Snall crustac-edns bOpCpoC5, Od\ll:l 1971cladocerans, ostracods), insectlarvae

lleadwatcr poolsand channel

~parva

rainwater kill:fi~h

Ri·..ulus A'larll"JOra~us

rivIOlus

+

+

+ 1-3. 1;" 8.U-1S" 17

3,8,13,15

<20 mm: ?lanktonic c0?epods21-37 ~: amphipods, rr~sids,

chiror-o~d larvae, ostracods,molluscs, plant detritus

OC:um 1971

Pocciliidae - livebearersGrm:usia a:finismos<:.uitof~

Gambusia rnizophoraemangrove gambusia

+

2, 3, 7,13-15

6, 9

A versatile feeder: dITlphipods, Cd\ll:l 1971chironomid l~rvae, hydracarina,harpacticoid copepods. s~ils,

ants, adult insects, ?Qlychaetc\oIOrns, ostracods, mosq",ito pupae,algae

115

Fresh and braCkishwater in Rhizophoraswamps, nOrthernCuba, southeasternFlorida

Fanily and Species Rercrence DietDiet

Reference Coments

Hctcrandria fornosaleast killi:~

poc~ilia latipinnasailfin rlOlly

Atherinidae - silv~rsldes

AIIanptta harringtonensisreef siIvE"Isi::lf!'

8. 14, 15

S, 7, 8.13-1S

Chironomid larvae, narpacticoid Odum 1971and ?lankton1C copepods, clado-cerans, terrestrial inse~ts,

algae, diatolf.S

Plant detrItus, al~ae, diatons Cdum 1971

39-60 mn: copppods, fish larvae, Randallpolyc:,aete larvae 1%7

Mcmbras cartinicaro~qh Si:VE"fSlde

~enidia bcryll1r.atid~~ater silv~rsidE"

Sy::lqnathidile ­pipefis~p~. s~a~ors~s

('orythoic~!:m.

albirostris~~~~~ipc~~sh

lI..i....P.pocampu~~~

11 ned s(>ahor~p

h i ':)E~campu~ ~.!:'E~e

d....·ur"" s ...ahor!>p

Mlc~ccnd:h~s cri::liger~s

fringpd pl?cfish

§}'n:F-:3t~ floric.aed'Jsky pi Fcf. sh

5vngnathus louisianaec::'air. ?ipef:..sh

2. 5. 11

2, J. a.11. 1:1..n. 1,.

1.

~, 11

1 • 2. ~. 11.,

1J

\, 2. ). c'.11. 16. 1.7.

'8

1.. 5, IJ

1-3,~. 1:

1-). 11.16-:5

S:na 11 zooplankton ~rustaceans,

juvenile &- larval fishes,insects. detritus, snails

Insects. COpCpod3, chironomidlarv~p. ~ysids, an?hipods

C,2-R2 mm: l'Opq::oc.s. micro­crust;;cea:ls

Ca!"lc.car. shri'TIp, a:np::i?Qds.tanai::!s, iso;x>:]~

:opepods, ~~~hipods, sm~ll

'i:':.Ti:"ltl

Peterson &­

Peterson19B

Od..lT.l 1971

Rei:] 19')4

Brook 1975

~e:d 1954

Associated withvegetated areas {Tabbfl !·lann1n':j 1961)

Intimately associatedwith unattached algae('ral::b 6: Manning1961), or grassyareas (Springer &"O'Joodburn 1960)

Inh,1bit grassyflats (Springer &

\.;oodturn 1960)

~ynqnathus s=cVE":l:G'J:f r:ir-e:·.ah

Synqna:h~s ~Frin9pri

~.:: 1 ~jl-'f>flsh

syncna:~~s cur.ck~~

~ugr.osp ?locfish

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

Cer.:r0I'0:--idJoc - snoo:<5ce~tropo~us ~aral:eIus

~at sr.cok

Centroponus ectinatustarpon snook

1-3. ;;. Il­l F.-I ...

:2. I:'

11

7, 3. 13

',mr-':':ipoCs. is:)~:15, rancHes.COPE'"P.:cs, 'tir.y ~aric.ean

ShCIFO. q~st~cp~s (Ei~t;un.

14itrell':') ----

116

Brook 1975Sprin']cl" '"\·:ooob'..:rn 131;0Reia }QS4

i\ssociated 'withvegetated a!"eas(:abb & :.talming1901J

Fanily as a wholeshows preferencefor estuarine :nan­grove ho1bi tat(Rivas 1962)

Habitat Tvpe

Family and Species Refetence IlietDiet

Reference COQlf'tents

Centropomus undeciroalissnook

Serranidae - sea bassesCentropristis striatablack seabass

Diplectrum formos~

sand perch

Epinephelus itajarajewfish

EPinephelus nlOri<:·red grouper

Epineehelus striatusNassau grouper

Hypoplectrus puellabarred hamlet

+

+

+

+

+

+

+

+

2,5,7.8.11, 13, 14

11

2, 3, 11,16-18

2.5, 7,8,11, 13, 15

11

4, 11

11

JuveniJes, caridean shrimp,small cyprinodont fisr.es.gcbies, nojarras

Adults: fish, crabs, penaeidshrimp, crayfish, snappingshrimp

Family in general carnivorouson fish, crustaceans

Caridean &. pen"eid s::trilllp,cO?epods, crabs, :ish

Juveniles: penaeic shrimp,xanthid crabs

226-)40 mm: crustaceans,crabs. fishes

170-686 mro, fish, crabs,sto~topods, cerha1opocis,shrimp, spiny lobsters,gastropods, bivalves, isopods

54-98 mm, sna?ping shrimp.crabs, fis~, mysids, sto~ato~

pods;, isopods

OCum 1971Austin &

;'ustin1971

Randall1%7

Reid 1954

1971

Randall1961

RandallL9E7

Randall1967

iOy far mostab\;ndant ofthree species(Rivas 1962)

The most abundantof t:1.e sca::>a:o:sesi~ "~n9rOve habitats

Mycteroperca ~rolepis

gag

Centrarchidae - sunfishesElassoma evergladeiEvorglades pygmy sunfish +

Lepomis .auritusredbreast sunfish

+ + L 2, 5, 1.1,17, 18

14

14

71.-100 rrun, peflileid shrimp,fi!:lh

Reid 1954

Family is primarilyfresnwater, fis~

occasionally enterhcad'~ater areaof mangcova­fri.nged stre,l;:!

LeDOnis qulosuswarmouth

~ macrochirusblueqil.1

Lepomis microlophusredear sunfish

+

2, 13, 15

2, 15

2, 13-15

Shr imp (Palaemonetesl, =ish{Cobiosoma besd, LepomismacrochiT~sl, detritus,Vallisneria, am?hipods, xan­thid crabs, blue crabs

~phipods, blue crab (Cal-l inectes sapid'J.s), xanthidcrabs, detritus, Vallisneria,cla>:ls (Rall9ia cuneata}.sponge (Ephydatia fluviatilis),barnacles, insect larvae

Chironomid larvae, amphipoes,xanthid crabs, clam (Ranqiacuneatal, sponge (Ephvdatiafluviatilis), detritus

Desselle eta1. 1978

Dessell< eta1. 1978

Desselle etal. 197a

Diet from LakePontchartrainsa lin i ties J _6­4.1 0/00

Diet fron LakePontchartrainsalinities 1.6­4.10/00

Diet from LakePontchartrainsalin1 ties 1.6­4.1 0/00

Lepomis punctatusspotted sunfish

+ 6, 14, 15 Cladocerans, small crabs, Odum 1971mysids, chironomids, ~phipods,

insects, molluscs, isopods,fish, algae

117

Salini ties < 15 0/00(OdUl':l 1971)

Family and Species

Habitat Type

~ Iu';j ~ ~ 'd.." ... 3 >./ ~ >,;::: t: :l 2 :s ~ Re[erence Diet

DietReferenee Comments

"icropterus salmoideslargemouth bass

Apogonidae - cardInalfishesAstrapogon alutusbronze cardinalfish

Astrapogon stellatusconchfish

+

+ +

+

13-15

" 3

1

Caridean shrimp, small blue Darnell 1958crabs, crayfish, xanthid crabs.2S species of fiSh, Vallisneris,Cladophora

Pomatomidae - bluefishesPomato~us saltatrixbluefish

+ 11 Young; mainly fishes (anchovies,silversides, killifishes, cen­haden, shad, spotted scatrout),shrinp, crabs, other smallcrustaceans, annelids, snails

Peterson &Peterson 1979

Rachycentridae - cobiasRachycentron canadumcobia

+ +5,7,11 fish, crabs Randall 1967

Echeneidae - remorasEcheneis neucratoideswhite fin sharksucker

Remora~

remora

+

+

+ + 2, 11

7

Fish, lsopods, other crustacea

58-175 ~m; copcpoda, isopods,vertebrate musclc tlSStie, crablarvae, fish remains, crusta­ceans, amphipods

Randall 1967 Members of thlsfa-oily attach tosharks and largebony fishes(Randall 1967)

Randall 1967

Carangidae - jacks, pOmpanosCaranx~ - blue +runner

+ + 2, 4, 5,7, 11

Family of s ....ift­swimmln~, carnh-­orous fishes,often running inschools, -.'idc­ranging (Randall1967)

~hippos

crevalle jack+ + + 2, 5, 7, fishes, crustaceans

8, II, 13Odum 1971

CaratU: ruberbar jac-k--

+ 4. 11 160-547 mm; fish, shrimp, myslds, Randall 1967stonstopods, gastropods

Chloroscombrus chrysurusAtlantic bu~per

+ + 2, II, 17,18

Ollgoplites saurusleatherjacket

Trachinotus carolinusFlorida pompano

Trachiootus falcatuspermit

+

+

+ +

+

+

2, J, 5,8, 11, 13

11

7, 11

Snapping shrimp, penneid shrimp,larval anchovies, ladyfish,harracticoid copepods

sardines (Hnrcngula sp.),mole crabs (~sp.),

bivalves (Donax sp.)

15-70 mm; OOy81d8, shri~p,

anchovies, silversides, crabs,snails

Tabb &!-tanning 1961

Spr:lnRer 6­,,"oodburn1960

Carr & ,'dams1973

CODDlon over Inudbottoll'-s (Randall1967)

Hore apt to occurover sandy bot tonsthan T. carolinas(Randall 1967)-

Selene \'OI:1erlookdown---

+ + + 2, 3, 7,11

Young; shrimp and othercrustac~ans, small molluscs

118

Peterson &Peterson 1979

Family and Species

Hel:licaranxamblyrhynchus ­bluntnose jack

Habitat Type•.~"".::; >.:r. ~

~"'"

+

Reference

17

DietDiet

Reference COMents

Caranx latushorse-eye jack

Lutjanidae - snappersLut1anus analtsmut.ton snapper

+

+

12

I, 4, 11

Predaceous on other fishes

204-620 mn: crabs, fish,gastropods, octopods, hentit.crabs, penaeid shri~p. spinylobsrer, stomatopods

Darnell 1958 Considered byGunter (1956) tobe eur)'hal1ne

Randall 1967 COlmlonly foundover sand, sea­grass, rubble,coral reefs(:l.anda11 1967)

Lut1anus apodusschoolmaster

+ + I, 4, 5,7. 11

Crustaceans (shrimp, snappingshrimp, blue crabs. xanthincrabs. grapsid crabs). fisr.

Nugent 1970

Lut1anus griseusgray snapper

+ + + 1-3. 7,8, 11-13,15-18

<50:nm: reside in grassbeds Odurr. 1971feeding on s~all crustaceans,insect larvae95-254 mm: reside in mangrovecreeks feeding on crustaceans(snapping shrimp, xanthid crabs,penaeld shrimp, crayfish, carideanshrimp), fish including gobles,anchovies, poeciliids, eels,k11lif1shes

By far the flOSt

abundant snapperin mangrovehabitats

Lutjanus iocudog snapper

Lut1anus synagrislane snapper

Gerreidae - mojarrasDiapterus olisthostomusIrish pompano

Diapterus plumier!striped mojarra

Euciaostomus argenteusspotfin lIlojarra

Eucinostomus gulasilver jenny

Eucinostomus lefroyiPlOttled IDOjarra

Gerres cinereusyellowfio mojarra

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

1

I, 2, ),5, 7, II,

16-18

2

2. 7. 8,11-13.15. 18

1-5, 7,8, 11-LJ,16-18

1-3, 5.7, 8,11-13,16-18

10

2. 7, 11

190-630 mm: fish, crabs,octopods, spiny lobster,gastropods

snapping shrimp, crabs,anchovies. annelids, nolluscs

110-116 mm: green algae(Enteromorpha flexuosa,Cladophora), Ruppia n:aritina.bluc-green algae (Lyngbama1uscula)

36-172 ~: mysids. amphipods,harpacticoid copepods,chironomid larvae, ostracods,bivalves. plant detritus

19-63 rem: ~phipods,

chironomids. harpacticoidcopepods. ostracods. mys~ds,

molluscs, plant detritus

19-70 nm: amphipods, chironomidlarvae, harpacticoid copepods,molluscs. mysids. ostracods,plaot. detritus

Crabs. bivalves, gastropods,polychaete WOrDS, shrimp,ostracods

Randall 1967

Stark bSchroeder1970

.\ustin GAustin 1971

Odum 1971

Odurn 1971

Odum 1971

Randall 1967,Austin &Austin 1971

Known fron brackishwater to depths of

220 fathoms(Randall 1967)

A permanentresident (OdUD1971)

Pomadasyldae - grunts Family carnivorous though rarely Randall 1967piscivorous

119

Most. shelter oncoral reef byday, feed ongrassy flats bynight (Randall1967)

Family and Species

Anisotremus ::5rginic~2­

porkfish

Haemulon aurolineatumtomtate

Ilaemulon carbo:1driumCaesar grunt

Haernulon flavolineatum-~~- --------French gr'..lnt

Haernulon "arra~

sailor's choice

+

+

+

Reference

11

1, 4, 11

" 7

Diet

112-264 mm: brittle stars, crabs,shrimp, po1ychaetes, isopods,bivalves, stomatopods, gastropods

97-170 rom: shrimp & shrimp lar­vae, polychaetes, hermit crabs,amphipods, copepods, gastropods,bivalves

156-273 rom: crabs, gastropods,sea urchins, chi tons , poly­chaetes, brittle stars, sipun­culid 'NOrms, shrimp

113-228 rom: polychaetes, crabs,sir-unculid worms, chitons,ho1oth~rians, isopods, shrimp,bivalves

Benthic invertebrates includingshrimp, crabs, a~phipods, gas­tropods, polychaete worms,bivalves

DietReference

Randall1967

Randall1967

Randall1967

Randall1967

Randall1967

Comments

Haernulcn plurnieriwhite grunt

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

Haenulon albummargate

Haemulon sciurusbluestriped grunt

Orthopristis chrysop­

=pigfish

Sparidae - porgiesArchosargus probatocepha­1u,sheepshead

Archosargus rhomboidal issea brea.'1l

+

+

+ +

+

+

+

1, 3, 5,7, 11

1-3, 5,11, 16-18

2, 3, 5, 7,8, 11-13,17-18

5, 11

Benthic invertebrates includingcrabs, shrimp, polychaete worms,amphipods, copepods, snails,bivalves

Benthic invertebrates includingcrustaceans, molluscs, annelidworms

Juveniles: 16-30 mm: planktonincluding copepods, mysids,?Ostlarva1 shrimp

>30 ~m: polychaetes, shrimp,arnphipods

<40 mm: in grassbeds - copepods,amphipods, chironomid larvae,mysids, algae, molluscs

>40 rom: in mangrove creeks ­mussels, false mussels, crabs,snapping shrimp, crayfish,hydrazoans, algae, plantdetritus

32-85 rom: in puerto Rico man­groves - lOO~ blue-greenalgae (~mojuscula)

105-220 rom: seagrasses Cyn,odocea& Tha1assia, algae~crabs, gas­tropods, invertebrate eggs,bivalves

Randall1967

Randall1967

carr &

Adams1973

Odurn 1971Austin &

Austin1971

Randall1967

Strong preferencefor vegetated sub­strate in bayareas (Weinsteinet al. 1977)

usually seen inmangrove sloughs,rare on reefs(Randall 1967)

Calamus arctifronsgrass porg;,'

+ + 11, 17 Copepods, amphipods, mysids, Reidshrimp, bivalves, gastropods(Mitrella, Bittium), polychaetes

1954 Associated withgrassy flats (Tabb &Manning 1961)

~calamus

saucereye porgy+ 1 190-250 mm: polychaetes, brittle

stars, bivalves, hermit crabs,sea urchins, gastropods, chi tons

120

Randall1967

F"a:::Jily and S;Jecies Referenc.e DietDiet

Referenc.e CollliDents

Lagodon rhomboidespbfish

5ciae~idae - drums3air1iella batabanaI::::'.:e croaker

+ +

+

+

+

1-3,5,7,8,n, 12, 16­

1.

3, 11

In :·'angro\'e <;;reek - scorc:ted:nussel, mysids, a:n,hipods,false 1:11.:sselIn IOhitewater Bay - lao' plantmaterial

0<1=Reid

19711954

Strong preferencefor vegetated sub­strate in bay areas(Weinstein et al.1977)

Bairdiella chrvsurasllv!:!r perch

Cvr.oscion arcnariussan::] seatrout

+

+

+ ::'-1,8,11­13, 16-18

2, 12, 17,1.

Larvae: copepods, larval fish(~ beryllina)127-181 mm: fish (Anchoamitchil1i) , mysids

~ostly fish, caridean shrinp,nysids, anphipods, crab zoea

OdW'J 1971

S?ringer &

Woodburn1960

Cynoscio~ nebulos~s

5Fott~d seatroc!t+ + + 1-3,5,7,

8,11-13,:5, 1';', 18

<50 :nD: copepods, planktonic Odu:n 1971crllstacea

50-275 :mu: fish (MugU cephalus,Lagodon rhomboidcs, ~ucino­

StomU5 quIa, E. arqenteus,Cyprinodon varieqatus,Gobicsoma robust=, Anchoa::Iitchilli)

Leiost~ xa~th~

spot+ 2, 7, 12,

17-18<40 mm: ?lanktonic organisms>40 mm: filar:tentous algae,

desmids, forams, amphipods,~ysids, copepods, ostracods,isopods, chaetoqnaths, bi­valves, s~ils, polychaete'....OrT.\S

Spr inger &

Woodburn1960

~0nticir=hus anerican~s + + 2, )1-12,so~ther~':;q~~-'- 17-15

L1enticirrhus litt~ 2, IIGulf kinqfish

t-:icropogon undu1at:u~ + 11, 12Atlant~c croaker

pog(lnia~~ + + + 2, 7, 11.black drum 12, l'

rish, benthic crustaceans

Polychaetes, bivalves (Donax) ,sand crab It:merital, razor clams

Juveniles: copepods, ~ysids,

carid~an Shrimp, polychaeteworms, insect larvae, iso­pods, s~all bivalves

<100 ~: molluscs, xanthidcrabs

>100 mm: bivalves, amphipods,blue crabs, pcnaeid shrUnp,c-aridean shrimp

Springer I;

I\oodburn1960

Springer &

Woodburn1960

Spri:lger '"\ol"oodburn1960

Darnell.1958

"'.ost COl:llllon off sandybeaches <springer &

woodburn 1960)

Sciaenoos ocellatared drum

E3u.ct'..Is ~cuminatus

high-hat+

2, 3, S, 8,11-13, IS,

17

11

<10 ron: planktonic organisms(copepods, crab zoca, larvalfish)

34-42 nm: mysids, amphlpods,ca r idea n Shrimp

>50 rnrn: xanthid & portunidcrabs, pcnaeid shrimp,small fish

308-403 ~: xanthid crabs

68-152 mrn: shrimp & shrinplarvae, isopods, stomatopodlarvae, copepods, anphipods

121

Odum 1971

Randall1967

Characteristic ofcoral reefs(Randall 1967)

?ani 1y and Species

Er-r.ipp~da~ - spadcfis~p.s

chactod~~~crus faber.A"t\-a--;:;-ili;pad\"fish--

r-or..a::ef,cridae ­dans~::i,,;,es

Abudefd~f saxatilis;;;rge;'.;-tllId~-

Habitat T\·pc

! I ~~ ~ .""

:s~~~I;~!-o u:. I=.l OQ 0::0 Reference

+ 2. 3. :;,11, 16-13

Diet

Wonns. crustaceans, debris

101-135 mm: COpepoc.s. algae,fi~~ ~qqs. flSh. sr.rimp larvae.po1ych':ll"rc"i>

DietReference

Jarnell1961

Rand;.ll1967

COIIIDents

Juveniles (7-12 mm)

inhabit very shallownearshore sandybeaches. Bear adeceptive resemblanceto infertile redmangrove seed pods(Breder 1946)

Characteristic familyof coral reefs (Ran­dall 1967)

A habitat generalist:reefs, grassbeds,rock piles, wharfs(Bohlke S Chaplin1968)

l."'lo::-lc.ae - wr:J,;<sesllalic::oer~s biv:'ttatull;llpp<>ry dlC-k-----

Scal:"id"-i:! - p,-,xl:"ctfishestlicnolsin,-, ust<aemp.calc pdl:"rotfis~

SEar isoma ct,rysopteru.!!:.redtail parrctfish

SF,-,ris~ma rubriElnneredfln ?drrot:ish

searisclIla_ ~id(!

st~plight parrot~ish

+

+

,

I , 2, 11. If!

4

11

1J

f>7-153 mill: crabs. Si!'3 .lrchins,polyc~detes. gastropods, brlttlestars. bivalves, shri~? fish,!:erm~t crnba

FamIly heorbivorous, feedingpr~m~rily on alqae grOwingon l:ard substri1tes, >'lecondari1yon scaq(aSS~!:l

Randall1967

Randall1%7

Shallow water patchreefs, sand bottoms,grassbeds (Randall1967)

Family characteris­tic of coral reefs,ranging into grass­beds

Requires near marinesalinities (Tabb &:!'Iannin9 19611

~~gilidae - ~ullcts

MUCll (;eE:tal:.;~

striped l:ullel:

+ + 2.3,5,7, a, Inorganic sedil"lents, fine11-13, 15 c:et::-~tus, micro-algae

Odum 1971

Muqil ~~white mullet.

Hugil trich<)dcl1~il~

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

1971

• • 2,7,11.12

sphyracnidae - barracudasSFnyracna barracudagreat ba::-racuda

+ + 1-5, 7, 8,J 1, l3

135-369 mm: fish {Eucinostomus Odum 1971quIa, Menidia ber}'llina,~sarsus probatoccEhalus}

Salinities >10 0/00(OQUIlI 1971)

epistognathida;) - ja""fishcs~oistognathus maxillosus~ttled jawfish

+ 53-110 mm: shri~p, isopods.fishes, po1ychaetes, mysids,copepods

122

Randall1967

Fanily lives inburrows in sediment.often in vicinityof reefs (Randall1967)

Habitat Ty..£..l:......

•.5 uE '"' ..-l.-l 0;1 <0 c

~~3>.:::>.__F_a_m_'_'_y_a_nd_s_pe_c_'c_' c;:~:;:=cc:l~~~oooc~=__R_e_f_e_re.n:.:. _ Oiet

DietReference Co:vr.ents

Clinidae - cl1nidsChae:lOpsis oc(!llatablue throat pi~cblcnny

Paracli:lus ~rnoratus

JIlarbled J:>lli'n:l}'

Paracllnus fasciatcsbanded J:>lli'nny

Stat~notus he.phill~

b1ackbcl1y blfOnr;y

"amily ~pr-"~r'.; to =X;, car-n~ .....orous R3.ooa:'10:1 tenthi.: irwec:::ebrate:-, 19E::

1. ~. Ii

1

Inshcre on ~:Jck,

cor31 ur rubblesuostrat.cs (Rdn­c.aL. :'~67:,

B1enniidae - co~too~h

blennlcsChasmodes saburraeFlorida bl;;ny---

Blennius marmor~u~

seaweed blc:lr.y

Ble:miu!; nicholsihiqhfin blenny

Callionymidae - drago~et5

~ionym\l!; rallcirildi;;;tll,,_spotted dragonot

l-L 11. 17.,.

1. 5. 11

21-2~ mm, an~h~~5

2S-~~ mM: aD~h~~J~~, d~trltus.

pol ych<o"'t" ...~ snilll S

AI<::"c, o~':la:lic cf'L:-itllS,br:c-;::fO star;;, pr~ly"ha02t0s.

hyc.p):ds

Carr ..Adi!.ms]')73

:<.and"'ll19(,1

Cr;rnt1On brilckish~at.er ~len:lY (~atb

~ ~dr.nlnq 196:')

Elcotridae - sleepersDormitator maculatu~

fat sl eepe'C

Gobiidae - gobie<~~hygobi~ aor-oratorfrillfin goby

1), 1. 5

2, 3, B, 11, Car-idean shrimp, ::::,iron::.:mid17 larv<le, <H:lp~ipo::l.,;

('dol:"! 1971

Fresh.....a':cr a:1c.':'010' sa1:"nity area3iD3rncll 19611

Gobionellus ;,astatus +sharptai 1 qo~y

Gobionellus shufe1dci +fresh'",atcr goby

Gobionellus smaraJ::Ius +emerald qoby

Gobiose>::l<l bosci + +naked ""by

Gobioscraa loncipalatwosca 1e goby

GobiOSOl:la robustuncode qoby

12

2. 17, !R

3, a, 10, 11.15

2. 12

17

1-3. '>. ti,11.16-18

F'ilancntous algae l~nterc­

IflOrphal. ostraC:Jds. :::opepod~,

i 1l';,",Ct. larval.'

Small crllst~ceans includlngamphipods, annelids. £:s::,f:'~h e"qs

Amphipods, mysids, chironomidlarvae

Spr lnger Ii.

I-:oodburn19bJ

Petersoll 6­

Pet.erson1919

Odum 1971

Lopho90bius cyprlno~des

crested qoby1-3. 7, S,

13A vers~tilc feeder, am~hipods, O::l~ 1971mangrove detritus. =ilanentousalgae, "ysids, caridean &pcnaeid shrimp. polychaete~~rms, ostracods, bivalves,chironomid larvae. harpacticoidcopcpods. Lsopods, x3nthidcrabs, snails

123

Family and Species Diet.Diet.

Reference Comments

MicrogObius 9ulosusclown goby

2,5,8, lI­13.15, 17,

Ie

Amphipods, copepods, chironomidlarvae

odum 1971

Microgobius microlepisbanner goby

Microgobius thalassinusgreen goby

Scombridae - mackerels,tunasSco~romorus maculat.usSpanish mackerel

ScomberOI:lOrU$ cavallaking mackerel

5

2, 3. 12

2.11.n.15

11

Planktonic organisms

Small crustaceans includingamphipods, other invertebrates

Adults feeding on penaeidshrimp nigratinq from tidalstream

3S0-1022 mm, fish

Birdsong1981

Peterson ...Peterson1979

Tabb &"".anning1961

Randall1%7

Scorpaenidae - scorpio~­

fishesScoroaena bn'lsi liensisbarbfish

Scorpaena grandlcornisplumed scorpionfish

Triglidae - searobinsPrionotus salmonicolorblackwing searcbin

Prionotus scirulusleopard searob!n

Prionotus tribulusbighead sea robin

Rothidae - lefteyeflounders

Bathus ocellatU9eyed flounder

Citharichthys ~crops

spotted whiff

Cit.harichrhysspilopterusbay whiff

Etropus crossotusfringed flounder

• • 7, 11 Shrim?, other crustaceans. Randallfish 1967

37-102 1l1tI'l, shrL"p. fish, Randa 11 ~ost o~ten foundunidentifi~d crustaceans 1967 in seagr;1Ss

"+ + + 1-3, 11, Small molluscs, shrimp, crabs Peterson &

16-18 fish, small crustaceans I'cterson 1979(ostracods, cumaceans)

+ + + 1-3, ll-lJ, Shrimp, crabs, fishes, a:nphi- Peter;.;on ,17, 1S pods, copepods, annelids, Peterson 1979

bivalves, sea urchtns

+ 1, 11 68-130 1!lIIl: fish, crabs, shrimp, Randall 1967amphipods

+ 1

+ + 1, 17, 1S Mainly mysids, also shri:np, Peterson , Recorded froncrabs. copepods, amphipods, Peterson salinity ranRe.fishes, annelids 1979 2.5-36.7 0/00

(Darnell 1961)

+ + 3, 11, 16 Calanoid copepods, cumaceans, Peterson •a.11Iphipods, mysids, shrimp, Petersoncrabs, illOpods, annelids, 1979molluscs, fisheR

Paralichthys albiguttaGulf flounder

+ + + 1-3, 7, 11,12, 11, 18

<45 rem: s~a11 crustaceans,including amphipods, smallfish

>45 mm: fish (pigfish, pinftsh,l1zardfish, bay anchovy,labrids), crustaceans

Springer (;.Woodburn1960; Reid1954

Paralichthys lethostigma ­Southern flounder

+ Mainly fishes (mullet, ~n~a­

den, shad, anchovies, pinfish,mojarras, croakers), crabs,mysids, molluscs, penaeidshrimp, amphLpods

124

?cterson 6­Peterson1979

Fa~ily and Species

Habitat Type

m'l!' I~! ~ ~ 51 ~ ~ Reference_.. ~~---_.---~~- --~~---------------

DietDiet

Reference Comments

Sya~i~n pa~illos~n

Gusky flour_Ger

Soleidae - salesAcr-.:'..rJ.s lineatus~ole

Trinec~€s inscript~

scr:;;.wled sale

+ +

+

1

1-3,5,8,11-13, 17­

18

1

32-74 ;urn: chironomid larvae, Odum 1971polychaete worms, foraminiferans

'.rr~n0c:;es maculctu~

tog choker

Cynoglcsslda~ ~ tonqu0­fish"'"~Gph~rus plagius~

Dlackcheck tonguefish

+ 2, 3, 8,11-13, 17,

16

1, 3, 11,12, 16-18

14-110 om: amphipods, mysids

35-102 mID: polychaete worms,ostracods, portunid crabs,RUF~ia and Ilalodule planttips

Odum 1971

Austin &Austin1971

Balistidae - trigs"'rfishes& filefishes

Aluterus schoepfi~-ril~:ish

Balistes vetulaqueen triggerfish

Monacanthus ciliatusWnged fi.le~-

+

+

+

I, 11 Seaqr30sses, algae, hermitcrabs, gastropods

11 130-480 rnrn, sea urchins, crabs,bivalves, brittle stars, poly­chaetes, hermit crabs, gastro­pods, algae

I, 11, 17 47-97 mm: Algae, organic detri­tus, seagrass, copepods, shrimp;:, shrimp larvae, amphipods,tanaids, polychaetes, molluscs

Randall1967

Randall1967

Randall,1967Springer I>

woodburn1960

Associated withgrassbeds, sponge/seafan habitats (Ran­dall 1967, Vosset 211. 1969)

Solitary reef fishranging into grass­bed,

Closely associatedwith vegetated areas(Tabb I> "lanning1961)

~onacanthus hispidusplanehead filefish

Balistes capriscusgray trig~erfish

+

1-3, 11,16-18

Detritus, bryozoans, annelids,harpacticoid copepods, amphi­pods, hemit crabs, molluscs,algae, soa urchins

Peterson I> Associated withPeterson vegetated areas (Tabb1979 & Manning 1961)

Ostraciidae - boxfishesLactouhrys cuadracornisscrawled cowfish

+ + + 1,2, 5,7, Vegetation, algae, bivalves11,16-18

Reid 1954 Young mimic sea­grass blades(Bohlke & Chaplin1968)

Lactophrys trigonustrunkfish

Lactophr~ triguetersmooth trunkfish

+

+

1, 4, 11

1

109-395 mm: crabs, bivalves,polychaetes, sea urchins, algae,seagrass, gastropods, amphipods

93-250 mm, po1ychaetes, sipun­culid worms, crabs, shrimp,gastropods, hermit crabs, seaurchins, bivalves

Randall1967

Randall1967

Primarily a residentof seagrass (Randall1967)

Primarily a reefspecies (Randall1967)

Tetraodontidae - puffersSphoeroides nephelussout.hern puffer

+ + 1-3,5,11, Juveniles; detritus, fecal16-18 pellets, zooplankton, poly­

chaetes, gastropods, crabs,shrimpAdults, small crabs, bivalves

125

Carr I>

Adams1973

Family and Species Reference DietDiet

Reference Comments

Sphoeroides spengleriband ta U puf f er

+ + I, 7, 11 Crabs, bivalves, snails,polychactes, amphlpods,shrimp

Randall1967

Inhabits sea­grass, reef,rubble, man­~rovcs (Randall1967; Voss et a1.1969)

Sphoeroides testudineus +checkered puffer

Diodontidae - porcupine­fishes

Chilomycterus antennatusbridled burrfish

Ch110mycterus antillarumweb burrfish

Chiloaycterus schoepfistriped burrfish

Reference ~umbers Key

+

+

+

+

+

1, 7

11

2

1-3, 5,11, 16-18

85-92 rom: portunid ncga10pslarvae, gastropods

Ga;ltropods, hemit crabs,isopods, crabs, shrimp

Gastropods, barnacles, crabs,amphipods

Austin &!'1ustin 1971

Randall1967

Springer <\Woodburn1960

Reefs and grass­beds (Vossct 81. 1969)

Associated withgrassbcds (vosset a1. 1969)Salinities>25 0/00 (Springer<\ Woodburn 1960)

1. Bader & Roessler 19712. Carter et a1. 19733. Clark 19704. Holm 19775. Hudson et a1. 19706. Kush1an & Lodge 19747. Nugent 19708. Odum 19719. Rivas 1969

10. Seaman at al. 197311. Schmidt 197912. Springer &Woodburn 196013. Tabb 1966L4. Tabb, Dubro\ol & Hanning 196215. Tabb & ~ning 196116. Weinstein et a1. 197717. Yokel 1975a18. Yokel 1975b

126

APPENDIX C. Amphibians and reptiles recorded from south Florida mangroveswamps.

127

AMPHIBIANS AND REPTILES OF FLORIDA'S MANGROVES

Species

Mud Turtle(Kinosternon subrubruml

Striped Mud Turtle(Kinosternon bauri)

OrnateDiamondback Terrapin(Malaclemys terrapinmacrospilota and~.~. rhizophorarum)

Flor ida Red-bellied Turtle(Chrysemys nelsoni)

Chicken Turtle(Deirochelys reticularia)

Green Turtle(Chelonia mydas)

Hawksbill(Eretmochelys imbricata)

Loggerhead(Caretta caretta)

Atlantic RidleyCLepidochelys kempii)

Florida Softshell(Trionyx ferox)

Green Anole(Anolis carolinensis>

Cuban Brown Anole(Anolis sagrei)

Bahaman Bank AnaleCAnalis distichus)

Green Water Snake(Nerodia cyclopion)

Mangrove Water SnakeCNerodia fasciatacompress1cauda)

Status

Abundant

Common

Uncommon

Rare - Uncomroon

Unconunon

Uncommon

Rare

Common

Uncommon

Cornman

Common

Common

Unconunon

Carrunon

Corranon

128

Food Habits

Insects, crustaceans,mollusks

Algae, snails, deadfish

Littorina, Melampus, ~,Anomalocardia

Sagittaria, Lerona, Naias

Crayfish, insects, Nuphar

Mangrove roots and leaves,seagrasses

Rhizophora: fruits, leaveswood, bark

Crabs, jellyfish, tuni­cates

Snails, crabs, clams

Snails, crayfish, mussels,frogs, fish, waterfowl

Insects

Insects

Insects

Fish

Fish, invertebrates

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

Species

Striped Swamp Snake(Liodytes alieni)

Eastern Indigo Snake(Drymarchon corais)

Rat Snake(Elaphe obsoleta)

Eastern Cottonmouth(Agkistrodon piscivorus)

American Alligator(Alligator mississippiensis)

American Crocodile(Crocodylus acutus)

Giant Toad(aufo marinus)

Squirrel Treefrog(Hyla squirella)

Cuban Treefrog(Hyla septentrionalis)

Status

Uncommon

Uncommon

Uncommon

Unconunon

Common

Rare

Common

Abundant

Corrrrnon

Food Habits

Crayfish, sirens, frogs

Small mammals, birds,frogs

Small mammals, birds

Fish, frogs, snakes,birds, small mammals

Fish, waterbirds

Fish, waterbirds

Invertebrates

Insects

Insects, frogs, toads,lizards

References: Carr and Gain 1955; Ernst & Barbour 1972;Mahmuud 1965; L. Narcisse, R.N. IlDing" DarlingFed. Wildlife Refuge, Sanibell Is., Fla.;personal communication (1981).

129

APPENDIX D. Avifauna of south Florida mangrove swamps.

130

Pfrc:lch 1966

Bacon 1910Howell 1932

19321970

19691932

OgcenHo....ell

H<Y...e11Bacon

Kush1an & White 1971alIo....ell 1932

Kushlan 1,}79Kushlan & Kushlan 1915Giraed £ Tayloe 1979

pfrench 1366

Girarc I> Tdylor 1979

Kahl 1964Ogden et al. 1976Kush1an 1919

Ho....el1 1932Kushlan I> White 1977a

Ho....e11 19]2Kush1an I> White 1977aFfrcnch 1966

Howell 1932Kush1an & "''nite 1977a

Howell 1932Kushlan & "''bite 1977a

Ho...'e1l 1932Kushlan & \·nti te 1977a

Howell 1932Kushlan & '..lhite 1977a

Kushlan & \1hite 1977aI'\axwell .. Kale 1977Girard & Taylor 1919

Kushlan I> White 1911a/>taxwell ,<; Kale 1977Girard & Taylor 1919

Robertson & Kush1an 1974jl\ax....·ell , Kale 1977Girard ,<; Taylor 1979

pfrench 1966.:1ax....'e11 I> Kdle 1917Girard & Taylor 1979

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(cas..nerodius albus)

Sno'''''Y Egret COllIllOn Y< Y f'ish(Egretta Chula)

Cattle Egret Common Y< Y Fish(Bubulcus lb ls)

Great White Heron Ra,. Y< V Fish(Ardea herod1asoccidentalisJ

Great Blue Heron Common Y< Y Fish(~ herodias)

Reddish Egret U:lcOtfl:nOn Yr V ?ish(Dichrolllanassarufes::ens)

Louisiana Heeon Common y, Y Fish(Hydranassa tricolor)

Little Blue Heron Com:non Y< V Fish(Florida caerulea)

Green Heron Co:nmon Y< Y Fish(Butorides strlntus)

Black-cro...med Night Co:r:unon y, Y Fish, crustaceans,Heron frogs, mice

(Nycticoraxnvcticoraxl

Yello""-crowned Night COllClOn V, V Fish, crayfish.Heron crabs

(Nyctanassa violaceal

Lea,t Bittern Uncom.~n Y< V ('ish

(Ixobrychus exilis)

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

W~d Stork CO:nr:Jon " Y Fish

ntycter ia americanal ( locallyabundant)

Glossy Ibis Uncommon " FiSh(Plegadis falci-nellus

\1hi ':.e Ibis Abundant y, V F'ish, erabs (~l

(Eudocimus albus)

Roseate Spoonbill Rare to Y< V Shring. fish,(Ajaia ajaja) UnCOllTllon aqul:I.tic vegetation

Sandhill crane Rare Vr 'Roots, rhizo::nes of(Grus canadensis) Cyoerus &- Sa9it-

caria

Lu?kin Rare Y< V Snails {~l(Aramus quarauna)

131

PROBING SHORE BIRDS

COllllllOO Name(Latin name)

King Rail(Rallus eigans)

Clapper Rail(Rallus 100g1ro­stds)

Virginia Rail(Rallus limicola)

SOTa(Porzana carolina)

Black Rail(LateralIusjaoaicensis)

Semlpalmated Plover(Charadrius semi­palmatus) --

Abundance

t:ncolllI!lOn­COllmon

Rue

UnCOtmlOn tolocallyabundant

Rare

Locallyconnon

Season ofOccurrence8

Yr

y,

Nestinga

Y

Food Habits

Beetles, grass­hoppers. aquaticbugs

Crabs, shricp

Beetles, snails,spiders

Insects, seeds ofemergent aquaticplants

Beetles, snails

Crus taceans,mollusks

References

Narcise. pets. comm.Martin et a1. 1951

Howell 1932Ffrench 1966Racon 1970

Marclsse, pers. C~.Martin et al. 1951

Howell 1932Bacon 1970

Narcisse, pets. co~m.

Ffrench 1966Bacon 1970Baker 6. Baker 1973

"'ilson I s Plover Locally(Charadrius wilsonia) common

Crabs, shrimp,crayfish

Howell 1932Bacon 1970

Black-bellied Plover(Pluvialissguatarola)

Common Crabs, mollusks Howell 1932Bacon 1970FEre-nch 1966

Ruddy Turnstone CommonCArcoaTis interpres)

Common SnLpe Uncommon

(.f2pella gallinago)

Long-billed CurIel,' Rare-uncomnon(Numenius americanus)

W,T

Insects, crus­taceans, mollusks

Molillsks, insects,worms

Crustaceans,insects

O~den 1969Ho\~ell 1932

HOI,'ell 1932Bacon 1970

Ogden 1969

Io.'himbrel(Numeniu8 phaeopus)

Spotted Sandpiper(Actitis macularia)

Uncommon

Abundant. W,1'

l~ollusks, crus­taceans, wor:ns,insects

Mollusks, crus­taceans

Ogden 1968Howell 1932

Ffrench 1966Bacon 1970Russel 1980

Solitary Sandpiper(Tringa sol itada)

COllIIl.on W,T Crustaceans, aquattc Howell 1932insects, small frogs Bacon 1970

Willet(CatoptrophorusseIJl1palJlat.us)

Greater Yellowlegs(TringaI:I.elanoleucas)

Lesser Yell~'legs

(Tringa flavipes)

COllDOn

CO!:lIllOD

Common

Yr

W,T

W,T

Crabs, crayfishes,killifishes

Fishes, crabs,crustaceans

Snails, mollusks,crahs

Howell 1932Bacon 1970

Howell 1932Ffrench 1966Bacon 1970

~'french 1966Bacon 1970Baker & Baker 1973

Red Knot. Uncomnon(Calidris~)

Dunlin CO!lllllOn(Calidris alpjna)

t"Thit:e-rumped Sandpiper Rare(Calldris fU8cicollis)

W, r

W

T

132

:-t.:lrine ...!orms,crustaceans

Narinc worms,mollusks

Chironomids, snails

Howell 1932Ogden 1964

Ogden 196t.8aker 6- Baker 1973

Howell 1932Bacon 1970

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 1970himantopus)

Semipalma ted 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 aquaticplants

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

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

133

SURFACE AND DIVING BIRDS

COlillnon Name(Latin !lamp) f.bundancc

5eafiOil of

Oc:::uT::"cncea tlcsting a Food ;!abi ts References

Fish, aq~atic insects, c~den 1969mollusks

COllTl\on Loon(Cavia immer)

Horned Gre1>e(Podiceps aurltus)

pied-billed ~rebe

(Podilynbuspodiceps)

White Pelican(Peleca!lU5erythrorhynchos)

Bral'm Pelican(?elecanusoccidentalis)

Double-crestedcormorant

(Phalacrocorax

auritus)

occasiona 1 "UnCOll"fllon "UncannQn- ':'r

CO!mlon

Ra'" 5COllUlcn "Corrmon "

Cor.JIlO:l V, y

Fish, crabs, rr~llusks

Crayfish, fish,~ollusks

Fish

Fish

Fish

Narcisse, pers. co:nm.

Narcisse, pers. COIJ[ll.

Nar::;isse, pers. COIml.

Ffrench 1966Bacon 1970

Kush!an & ~hite 1977a

Ogeen 1969

Narcisso, pers. cocm .

C·gden 1969

LaHunt & Cornwell 1970Kushlan et al., in prep.

::Jgde:l.. 1969Kus:-:lan et al., in prep.

F:renc:: 1966

~arcisSQ, pers. cumru.Kushlan ct al., in prep.

Ogeen 10369Smith, pers. o~s.

Narcisse, pers. co~.

K·....shlan et al., in prep.

Karcisse, pers. comm.Ffrenc:t 1966

Ogden 1969Kushlan et al., io prep.

Ogaen 1969Kushlan et al., io prep.

Narcisse. pers. comm.Kushlan et al .• in peep.

Wi.dgeon grass

Moll~sks, cr~stD­

ceans, ~idgcon grass

~uts, seeds Ogden 1969

Vallisneria, Ruppia,

~

Ruppia, ~,mollusks

Snails, clams, a~uatic Ogden 1969insects, RUPEia, Zos-tera

mollusks, aq~atic

insects, Rupnia,

~

Fish

Ruppia, Zoster;)..aquatic insects

~, snails,insects, crustacoans

?olvqonum, snai~s,

Rup'O:'a

Saggitaria, mollusks,cype~

;<.uE'Oia, ~,n.ollusks

Polygonum, Reppia,crayfish, snails

v

y,

"

'.~, 'I'

\':,T

y,

'.~,T

~'1 ,T

\I',T

V,

W,T

.... ,::

"

"

"w

134

Race

Race

COmll'lOn

CQrJllO:'l.

Abundant

Abundant

l:ncOIIOOn

Unconmon

rJ:JUndant

COInIJlon

Korthern Shovelerl~ dvpeata)

Redhead(Aythya americana)

Pintail(~ acuta)

Black Duck(~ rube ipes)

,,;ott.led Duck

(Anas Culvigula)

Wood Duck(~ sponsal

Blue-winged Toal(~ discorsl

American Wigeonl~ a.'l\ericana)

Gaawall(~ streoera)

Green-winged Teal(A:1as~ carolinensis)

Fulvous ~.lhist:lLng Duck Uncanmon(Dendrocygr.abiealor)

:-laUard UJ'lc=on

(,\oas platyrhynchos)

canvasback Uncommon(Aythya valisineriaJ

Ring-necked Duck Abundant(Aythya collarisJ

Anhin<;ld("nhings anhinga)

SURFACE AND DIVING BIRDS (concluded)

Common Name(Latin name) Abundance

Season ofOccurrence a Nesting a Food Habits References

V ,'\quatic insects, Narcisse. pers. comm.mollusks. Ffrench 1966Eleocharis. Paspalum

y Seeds, aquatic Narcisse, pcrs. comn..insects fo'french 1966

Ruppia, Najas. Narcisse, pers. camm.Potamogeton,aquatic insects

Lesser Scaup(Aythya aHiuis)

Bufflehead(Bucephala alheala)

Ruddy Duck(Oxyura jamaicensis)

t100ded Merganser(Lophodytescucullatus)

Red-breasted Merganser(~ergus serrstor)

Purple Gallinule(Porphyrulamartinica)

Common Gallinule(Gallinula chloropus)

American Coot(Fuliea americana)

COI:IIDQn- Wabundant.

Rare W

Common W

Rare-unconoon W

Camocn !\I.T

Rare Yr

Common Yr

Abundant w,r

135

Mollusks, Ruppia

Gastropods, crabs.crustaceans

Pot.anogeton. ~ajas.

Zostera, Ruppia,nollusks

Fish

Fish

Narcisse, pers. coorn.Ogden 1969Kushlan et al., in prep.

Ogden 1969Kushlan et a!.. in prep.

Ogden 1969Kushlan et al .• in prep.

Ogden 1969

Narcisse. pers. carom.

AERIALLY SEAHCHING

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

Herring Gull Uncommon W Fish, mollusks, Narcisse, pers. corom.(~ arqen to. 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(Gelochelidonnilotica)

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 1969vicensisl

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

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

Belted :<ingfisher Common Yr Fish Narcisse, pers. corom.

(r·legaceryle alcyon)

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

136

BIRDS OF PREY

::,;,,! ·"11 :) rOcc:urr<Cl1e"a Hc:£erenccs

------------ ----------------~ugnificent Frigate- Common Sbit"d Uncommon l"I

(?reqat~ magnificens)

y Fish Narcisse, pers. comm.Smith, pers. obs.

TUYkey vul tun~

(Cathartes aura)

Black Vultun,(Coragyp~ atratus)

Common

Corrunon

y Carrion

Carrion

Narcisse, pers. corom.Orians 1969

Robertson & Kushlan1974

Orians 1969

S'i.'allow-tailc:d Ki tc(Elanoides :orfica­t.us)

Conmon y Snakes, lizards,frogs

HowellSnydc.r

19321974

Sharp-shinned Hawk Unco:nmon

(~cipiter striatusl

Cooper's Hawk Uncommon(M:Tipi~~ ~~r:ii)

y

Smaller passerines

l~rger passerines

Howell 1932

Howell 1932

Red-tailed ;'d\·..k

(Buteo jarnaic€rlsis)

Red-shouldered l!<r,.,k(Buteo !2:neatus)

Broad-winged 1M"'':(Buteo platypt;,.erus)

SWflinson's Hawk(Buteo swai:lsoni)

Short-tailed Hawk

(~tea £=ach~..::.'::.:'~}

Billd Ed'jle

(Haliaeetus

!..,ucocephalus)

Mill':"sh Ilawk

(Circus £..Zar.eusl

Os?rpy(!:.:"J.ndi0r:. :Jaliaetus)

Peregrine Falcon(!::3.1co -:Jeregr~r.~

l·le-r1in("'a1~ co1umbari;.ts)

AmerIcan Kestrel(~l:£<2 ~arverius)

Barn Owl

(~1~.£ alba]

Great lIorned 0·,011

(Bu~ virainianus)

3arred Owl(Strix varia)

Uncattunon

Canmon

lIr:comnO[l

Il.are

UnCCITlTlon

Rare-locallycommon [Fla.3ay)

Unco:nmon

::onmon

Very rilre­100a II.,. comnon{Fla. Bay]

Uncommon

Common

Cncor:unon

Uncomnon

lI:lCOIT\ll\On

Yr

':1

y,

w

y,

137

Small mammals, birds I1m"ell 1932

Sna:"es, frogs, Howel] 1932li zards, insects Robertson , :<.ushlan

1974

Insects, small Ho.....ell 1932maT'1Il\als

Small manmals, grass- Howell 1932hoppers

SmaIl birds 1I0·...ell 1932

y Fishes Ilo'.".'e 11 1932

Small mammals, shore- Rowell 1932birds

Fishes Howell 1932

Waterfowl, shorebirds Nisbet 1968Ogden 1969Howell 1932

Small birds, share- Howell 1932birds

Tnsects Howell 1932

y Sma 11 n1i'1lml\als Howell 1932

y ~;aterfowl , small Howell 1932mammals

y Small mammals, frogs, Howell 1932snakes

ARBOREAL BIRDS

Common Name(Latin name)

f-lourning Dove(Zenaidura macroura)

\Vhite-crowned Pigeon(Columba~pha1a)

Abundance

Uncommon

UncomlllOu

Season ofOccurrences

Y

Yr

Nestin~ Food Habits

Y Seeds

Y Berries. seeds,fruits

References

Emle-n 1977

Howell 1932Robertson & Kushlan 1974

Mangrove Cuckoo(COCCy;z:us minor)

Uncommon Yr Y Ca terpillars.mantids

Howell 1932Ffrench 1966Robertson & Kush1an 1974Martin et al. 1951

Yellow-billed Cuckoo Common(Coccyzus arnericanus)

Smooth-billed Ani(Crotophaga ani)

s

Y,

y

Y

Caterpillars,beetles

Insects

Howell 1932Ffrench 1966Martin et al. 1951

HO\o,'ell 1932Ffrench 1966

Chuck~will's-~idow

(Caprimulguscarolinensis)

Common Flicker(Colaptes auratus)

Pileated Woodpecker(DryocopuS pileatus)

Uncommon

Uncommon

Unconunon

y,

Y,

Yr

Y

Y

y

Mosquitos, moths

Ants, beetles,fruits in winter

Beetles. berries.fruits

Martin et al. 1951Karcisse, pers. comm.

Karcisse, pers. corum.Martin et al. 1951

Howell 1932Robertson 1955Robertson & Kushlan 1974

Red-bellied Woodpecker Common(Helane~ carolinus)

Red-headed Woodpecker Rare(t-lelanerpeservthrocephalus)

y,

Y,

y

Y

Beetles, ants,grasshoppers,crickets

Beetles, ants,grasshoppers,caterpillars

Narcisse, pers. comm.~artin et al. 1951

Narcis5e, pers. comm.~lartin et a1. 1951

Yellow-helliedSapsucker

(Sphyrapicus varius)

Hairy Woodpecker(Picoides~)

Eastern Kingbird(Iyrannus tyrannus)

Gray Kingbird(Tyrannusdominicens is)

Uncorrnnon

Rare

Uncommon

Common

',;I.T

p

S,T

S,T

y

y

Beetles, Llnts-,caterpillars

Insects, beetlelarvae

Ants, WLISPS,

grasshoppers

Bees, wasps,beetles, dragon

",arcissc, pers. corum.Martin et al. 1951

I:mlen 1977

Narcisse. pers. comm.Martin et al. 1951

HO\,'ell 1932Robertson ~ Kush1an 1974

Western Kingbird Rare(Tyrannus verticalus)

W,T Bees, 'Wasps.grasshoppers

Narcisse, pers. comm.Martin 8t al. 1951

Great CrestedFlycatcher

(Myiarchus crinitus)

Unconn:Jon(coTlUnon S)

Yr Y Insects, berries Howell 1932Robertson 1955

Acadian Flycatcher Rare(Empidonax virescens)

T Small flying insects Morton 1980

Eastern Phoebe(Sayomis phoebe)

Eastern Wood Pewee(Contopus virens)

Common

Rare-uncommon s,r

138

Bees, wasps, ants

Bees, wasps, ants,moths

Narcisse, pers. cornm.~artin et al. 1951

Narcisse, pers. comm.1l00,'ell 1932

ARBOREAL BIRDS (continued)

Cormon N<lmc

(Latin n<1mc) AbundanceScas~n ofOccurr.enc~ Food H.:loits Ref!'.::rcnc:e5

Barn Swallow(Hirundo rusHea)

Locally cOllUllOn lnsects HowellBacon

19321970

Blue Jay Uncommon(cyanocitta cristata)

Yr Grasshoppers, cater­pillars, beetles

Narcisse, pers_ comm.Hartin et a1. 1951

Tufted titmouse(Parus bicolor)

Carolina Wren(i'hryothorus

ludovicianusl

"'.ock ingbird(t-lil!'lus polyglottos)

Catbird(Dumetella ~­linensisl

Brown Thrasher(Toxost:.oma rufulll)

Very rare­rare

Uncommon

Abundant

COllllllOn

Uncoll'lllOn

w

Yr

Yr

\oI,T

Yr

Y

Y

Y

Caterpillars, wasps,bees

Ants, flies, mill i­peds

Fruits, berries

Fruits, insects

Beetles

Howell 1932Robertson &0 Kushlan 1974

!>'arcisse, pers. COOlIn.·

r~rtin et al. 1951

Robertson 1955

Narcisse, pers. comm.Martin et a1. 1951.

Narcisse, pers. comm.~tartin et al. 1951

An::erican Robin Abundant(Turdus migratoriusJ

~lue-gray Gnatcatcher Uncollll':'tOn(Polioptila eaerulea)

Ruby-crowned Kinglet Uncommon(Regulus calendula)

W,T

W,T

Worms, berries,insects

Insects, especiallyHymenopterans

Wasps, ants

Narcisse, pers. corom.Martin et al .. 1951

Narcisse, pers. comm..Howell 1932

~arcisse, pers. carnm.Howell 1932

White-eyed Vireol~ griseusl

Black-~hiskcred Vireo(Vireo altiloquus)

Red-eyed Vireo(~ 01 i vaceus I

Yellow-throated Vireo(Vireo flavj£rons)

Black-and4"'hiteWarbler

(Mniotilta varia)

Unconunon

Uncommon

Uncommon

uncommon

FairlyCOITIon

S,T

Yr

S,T

W

W,T

Y

Y

Y

Butterflies, moths

Spiders, caterpillars

Caterpillars, beetles

Butterflies, moths,

Wood boring insects

Robertson 1955

lIowell 1932Robertson & Kushlan. 1974

Narcisse, pers. comm.Howell 1932

~k>rton 1980

Lack and Lack 1972Keast 1990Ogden 1969

"~orm-eatin9 "~arbler Uncommon(lIellllitheros vermi-~)

Prothonotary Warbler Unco~n

{Protonotar ia ci trea)

Yellow-throated Common\~arbler

(Dendroica dominica)

Yellow Warbler conmon(Dendroica petechial

Yellow-:etlll'\pedWarbler Abundant

(Dendroica coronata)

Prairie Warbler Uncommon(Dendroica discolor)

Palm Warbler A~ndant

(Dendroica palmarurol

w

T

yr

W,T

Yr

W,T

y

139

caterpillars, spiders

Insects

Beetles, moths,spiders

Insects

Dipterans, bayberries

MOths, beetles, flies

Insects

Ogden. 1969Kush1an, pers. comm. c

Ffrench 1966Russel. 1980

Morton 1980

Haverschmidt 1965Ffrench 1966Orians ~969

Terborgh & Faaborg 1980

Narcisse, pers. comm.

Lack & Lack 1972Robertson & Kush1an 1974

Lack & Lack 1972Etll1en. 1977

ARBOREAL BIRDS (continued)

ConU7101l N"nHC!

(T~"ll:iJl :lilITH,,) A)"JIlC];lnce

Season o[Occnrrctlco;! a Nc:::till(j a References

Blackpoll Warbler Uncommon(Dendroica striata)

Bay-breasted Warbler Rare(Dendroica castanea)

T

T

Insects

Insects

Ffrench 1966

-"1orton 1980

Black-throated Gree~

\'larbler(Dendroica virens)

Uncommon Aphids, leaf-rollers,and other insects

Ogden 1969Kush1an, pers. comm.

Chestnut-sided Warbler Rare(Dendroica Densyl-v<lnica)

T Insects Morton 1980

Cape May Warbler(Dendroica tigrinaJ

Uncorunon

CommonHT

Ogden 1969

Black-throated Gray Rare\~arbler

(Dendroica nigrescens)

Black-throated Blue Uncommon\"Iarbler Common

(Cendroica caeru-lescens)

Insects

Beetles, flies, ants

Ogden 1969Kush1an, pers. conrn.Hutto 1980

Kush1an, pers. corum.Ogden 1969

:-Jorthern Waterti'.rush(Seiurus novebol::a­censis)

Ye11owthroat(Geothlypus trichas)

1'.bundant

Rare

Common Yr y

Insects

Grasshoppers,ants, wasps

Schwartz 1964Ffrench 1966Racon 1970Russell 1980

~rickets, Narcisse, pers. comm.HowelJ 1932Lack & Lack 1972

American Redstart Common(Setophaaa rutici1la)

Tennessee Warbler Unco~n

(Vermivora "::lereqrina)

Nasheville 'warbler Rare(Ve:rmivora rufi-capilla) --

Oranqe-cr~~ned Warbler common(vermivora celata)

Golden-winged Warbler Rare(Vermivara chrysop-tera)

T

T

T

T

Caterpillars

Insects

Insects

Insects

Insects

Bennett 1980Ffrench 1966Bacon 1970

:-tortan 1980

Hutto 1980

Hutto 1980

~\orton 1980

Northern Parula(Farula americana)

Common Hymenoptera Lack and Lack 1972

Ovenbird(Seiurus aurocapil­Ius)

Common Beetles, crickets,grasshoppers

Lack and Lack 1972

Kentucky Warbler Rare-uncommon(Oporornis formosus)

Mourning Warbler Rare(Ooorornis philadel-phia)

Yellow-breasted Chat Common(~virens)

T

T

140

Beetles, caterpillars, Morton 1980

ants

Insects Morton 1980

Hymenoptera Hutt0. 1980

ARBOREAL BIRDS (concluded)

C..,nWllOIl 1I,1m...

('.,It in 11,111:(')

Season orOCelll"I"r-ncea Food I I"..., i,U:

\~ilson' 5 \'i<)([,ler R,lce-unCOIIU1l0n(~il<;onid lZ.l.<si11 al

Red-.. inqed 1l1ackbird Comraon{A9alaiu~ phoeniccusl

Bo~t-tailed GracxLe ~ncommon

(Qutscalus major)

common Grackle UncommOnIQuiscalu!'> 9.ulscula.)

Card i na.i Common(CardinaUsc<H'din~lis)

'1'

Yr

Tnsects fJlJttu I.']HO

1~11lQ<; .:tnd Warner !<JHO

Seeds, insects nowell 1932Hohcrlson 1955

y Crayfish, cr:-abs, Robcrtsofl 1955

shr hop G~rard • Taylor 1979

y Insects, cater-- Howell 1':.1)2

?iilacs I<obcrtson J955

y In!iccts, sced~ Roherts')ll 1955

Orchard Oriole(Icterus spur ius)

Indigo BuntingIPasJ'>erina cyanea)

SUIlVl(lI" Tanager(P~.ran9a rubral

Dlckci:-:sel(Spiza al"erici'lna)

Rufous-sided 1uwhee(PiFilo erythroph­~)

Rare

Uncol"JlToOn

Uncommon

Uncomr.\On

Common

T

\".'1'

T

Yr y

Grnsshoppers. beetles

Grasshoppers, cater­pillal"s

IlYl:lenoptera

Caterpillars. beetles

Caterpillars. b~y­

barries, fruit!>

~brt.Oll 1900

Narcisse. pers. COTlt'll.

Howp.ll 1~32

Horton 1990

Bacon 1970Hartin ct al. 1951

Narcisse, pers. COr.'ill.

Howell j 932

S....amp Sparrow Common(Melospiz~ 9corqiana)

'fI.T Narcisl:le. pers. COntIno

Howell 1932

3.yr .. year round resident5 '" summer residentW • winter residentT • transient, present only during spring and fall migrationY = species breeds in mangroves

b L. ~a[cisse, R.N. "Ding" Darling Fed. Wildlife Refuge, Sanibel Island, Fla. (1981).

cJ •A• Kushlan, So. Fl Ra. es. Ctr., Everglades Katl. Park, Homestead, Fla.

141

APPENDIX E. Mammals of south Florida mangrove swamps.

142

Species

MAMMALS OF FLORIDA MANGROVES

Status Food Habits

Virginia Opossum(DidelphlS virginiana)

Short-tailed Shrew(Blarina brevicauda)

Marsh Rabbit(Sylvilagus ?alustris)

Gray Squirrel(Sciurus carolinensis)

Fox Squirrel(Sciurus niger)

Marsh Rice Rat(Oryzomys palustris)

Cudjoe Key Rice Rat(Oryzomys argentatusl

Cotton Rat(Sigmodon hispidus)

Gray Fox(Urocyon cinereoargenteus)

Black Bear(Ursus americanus)

Raccoon(Procyon lotor)

Mink(Mustela vison)

Striped Skunk(Mephitis mephitis)

River Otteru..utra canadensis)

Abundant

Uncommon

Abundant

Occasional

Rare

Uncommon

Rare

Abundant

Uncommon

Rare

Abundant

Rare

Common

Uncommon

143

Fruits, berries, insects,frogs, snakes, smallbirds and mammals

Insects

Emergent aquatics

Fruits, berries, mast,seeds

Fruits, berries, mast

Seeds of emergent plants,insects, crabs

Seeds, insects, crabs

Sedges, grasses, cray­fish, crabs, insects

Small mammals, birds

Fruits, berries, fish,mice

Crayfish, frogs, fish

Small mammals, fish,frogs, snakes, aquaticinsects

Bird eggs and youngfrogs, mice, largerinvertebrates

Crayfish, fish, mussels

~~LS or rLORIDA ~~NGROVES (concluded)

Species

Panther(Felis cancolor)

Bobcat(relis rufus)

White-tailed Deer(Odocoileus virginianus)

Key Deer(Q..~. clavium)

Black Rat(Rattus rattus)

Bottle-nosed Dolphin(Tursiops truncatus)

West Indian Manatee(Trichechus manatus)

Status

Very rare

Common

Common

Common on cer­tain Florida Keys(no longer onmainland)

Common

Uncommon

Uncommon

Food Habits

Deer, rabbits, mice,birds

Rabbits, squirrels,birds

Emergent aquatics, nuts,acorns, occasionallymangrove leaves

Emergent aquatics andother vegetation

Fish

Submerged aquatics,Zostera, Ruppia, Halodule,Syringodium, Cyrnodocea,Thalassia

References: Layne 1974; Hamilton and Whittaker 1979;L. Narcisse, R.N. "Ding II Darling Fed. WildlifeRefuge, Sanibel Island, Fla.; personal commu­nication.

144

OPT ONAL 2 ((Fo'merly NTIS-JS)Oepllrtment of Commen:e

(See ANSI-239.18)

.50271-101

REPORT DOCUMENTATION ( .... REPORT NO. T' J. RKipient's Accession No.

PAGE FWS/OBS-81/24... Title lind Subtitle 50 Rel'Ort OlteTHE 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, I I I9. 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 OfficeFish and Wildlife Service Bureau of Land ManagementU.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 processesof the mangrove forests of south Florida. This description is based upon a compilation ofdata 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 anddecomposition, carbon export, and energy flow.

Most of the components of mangrove communities are cataloged and discussed; theseinclude 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 presentways to manage this type of habitat. It is concluded that mangrove forests, which coverbetween 430,000 and 500,000 acres (174,000 - 202,000 hal in Florida, are a resource ofgreat 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'---'~'-" 15420. Seoeurity Class (This Pille) 22. Price

, fORM 12 4-11)

DEPARTMENT OF THE INTERIORu.s. FISH AND WILDLIFE SERVICE

As the Nation's principal conservation agency. the Department of the Interior has respon­sibility for most of our .nationally owned public lands and natural resources. This includesfostering 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 as­sesses our energy and mineral resources and works to assure that their development is inthe best interests of all our people. The Department also has a major responsibility forAmerican Indian reservation communities and for people who live in island territones under

. u.s. administration.