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Biological  Services  Program 


FWS/OBS-81/24 
January  1982 


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


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J3ureau  of  Land  Management 

5H         :ish  and  Wildlife  Service 

5  HO 

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J.S.  Department  of  the  Interior 


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The  Biological  Services  Program  was  established  within  the  U.S.  Fish 
and  Wildlife  Service  to  supply  scientific  information  and  methodologies  on 
key  environmental  issues  that  impact  fish  and  wildlife  resources  and  their 
supporting  ecosystems.  The  mission  of  the  program  is  as  follows: 

•  To  strengthen  the  Fish  and  Wildlife  Service  in  its  role  as 
a  primary  source  of  information  on  national  fish  and  wild- 
life resources,  particularly  in  respect  to  environmental 
impact  assessment. 

•  To  gather,  analyze,  and  present  information  that  will  aid 
decisionmakers  in  the  identification  and  resolution  of 
problems  associated  with  major  changes  in  land  and  water 
use. 

t  To  provide  better  ecological  information  and  evaluation 
for  Department  of  the  Interior  development  programs,  such 
as  those  relating  to  energy  development. 

Information  developed  by  the  Biological  Services  Program  is  intended 
for  use  in  the  planning  and  decisionmaking  process  to  prevent  or  minimize 
the  impact  of  development  on  fish  and  wildlife.  Research  activities  and 
technical  assistance  services  are  based  on  an  analysis  of  the  issues,  a 
determination  of  the  decisionmakers  involved  and  their  information  needs, 
and  an  evaluation  of  the  state  of  the  art  to  identify  information  gaps 
and  to  determine  priorities.  This  is  a  strategy  that  will  ensure  that 
the  products  produced  and  disseminated  are  timely  and  useful. 

Projects  have  been  initiated  in  the  following  areas:  coal  extraction 
and  conversion;  power  plants;  geothermal ,  mineral  and  oil  shale  develop- 
ment; water  resource  analysis,  including  stream  alterations  and  western 
water  allocation;  coastal  ecosystems  and  Outer  Continental  Shelf  develop- 
ment; and  systems  inventory,  including  National  Wetland  Inventory, 
habitat  classification  and  analysis,  and  information  transfer. 

The  Biological  Services  Program  consists  of  the  Office  of  Biological 
Services  in  Washington,  D.C.,  which  is  responsible  for  overall  planning  and 
management;  National  Teams,  which  provide  the  Program's  central  scientific 
and  technical  expertise  and  arrange  for  contracting  biological  services 
studies  with  states,  universities,  consulting  firms,  and  others;  Regional 
Staffs,  who  provide  a  link  to  problems  at  the  operating  level;  and  staffs  at 
certain  Fish  and  Wildlife  Service  research  facilities,  who  conduct  in-house 
research  studies. 


FWS/OBS-81/24 
January  1982 


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


by 


William  E.  Odum 
Carole  C.  Mclvor 
Thomas  J .  Smi  th ,  III 

Department  of  Environmental  Sciences 

University  of  Virginia 

Charlottesville,  Virginia   22901 


Project  Officer 

Ken  Adams 

National  Coastal  Ecosystems  Team 

U.S.  Fish  and  Wildlife  Service 

1010  Gause  Boulevard 

SI idel 1  ,  Louisiana  70458 


Performed  for 

National  Coastal  Ecosystems  Team 

Office  of  Biological  Services 

Fish  and  Wildlife  Service 

U.S.  Department  of  the  Interior 

Washington,  D.C.   20240 

and 

New  Orleans  OCS  Office 
Bureau  of  Land  Management 
U.S.  Department  of  the  Interior 
New  Orleans,  Louisiana   70130 


DISCLAIMER 

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


nth  Florida: 

ar  and 

E  Biological 


RETURNED 


Library  of  Congress  Card  Number  82-600562 


This  report  should  be  cited: 

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


PREFACE 


This  profile  of  the  mangrove  commun- 
ity of  south  Florida  is  one  in  a  series 
of  community  profiles  which  treat  coastal 
and  marine  habitats  important  to  man.  The 
obvious  work  that  mangrove  communities  do 
for  man  includes  the  stabilization  and 
protection  of  shorelines;  the  creation  and 
maintenance  of  habitat  for  a  great  number 
of  animals,  many  of  which  are  either 
endangered  or  have  commercial  value;  and 
the  provision  of  the  basis  of 
whose  final  products  include 
smorgasbord  of  oysters,  crabs, 
shrimp,  and  fish.  Less 
equally  important  benefits 


a  food  web 
a  seafood 
lobsters , 
tangible  but 
include  wilder- 


ness, aesthetic 
ations . 


and  life  support  consider- 


and  Tarpon  Springs  on  the  west.  Refer- 
ences are  provided  for  those  seeking 
in-depth  treatment  of  a  specific  facet  of 
mangrove  ecology.  The  format,  style,  and 
level  of  presentation  make  this  synthesis 
report  adaptable  to  a  diversity  of  needs 
such  as  the  preparation  of  environmental 
assessment  reports,  supplementary  reading 
in  marine  science  courses,  and  the  devel- 
opment of  a  sense  of  the  importance  of 
this  resource  to  those  citizens  who 
control  its  fate. 


Any  questions 
requests  for  this 
directed  to: 


or  comments  about  or 
publication  should  be 


The  information  on  these  pages  can 
give  a  basic  understanding  of  the  mangrove 
community  and  its  role  in  the  regional 
ecosystem  of  south  Florida.  The  primary 
geographic  area  covered  lies  along  the 
coast  between  Cape  Canaveral  on  the  east 


Information  Transfer  Specialist 
National  Coastal  Ecosystems  Team 
U.S.  Fish  and  Wildlife  Service 
NASA-SI i del  1  Computer  Complex 
1010  Gause  Boulevard 
Slidell,  Louisiana  70458 


m 


IV 


CONTENTS 


Page 

PREFACE  Ill 

FIGURES  vi i  1 

TABLES  vi  i  i 

ACKNOWLEDGMENTS  i x 

CHAPTER  1 .   INTRODUCTION  1 

1.1  "Mangrove"  Definition  1 

1.2  Factors  Controlling  Mangrove  Distribution  1 

1.3  Geographical  Distribution  2 

1.4  Mangrove  Species  Descriptions  5 

1.5  Mangrove  Community  Types  7 

1.6  Substrates  9 

1 . 7  Water  Qual  i ty  11 

CHAPTER  2 .  AUTECOLOGY  OF  MANGROVES  12 

2.1  Adaptations  to  Natural  Stress  -  Anaerobic  Sediments  12 

2.2  Adaptations  to  Natural  Stress  -  Salinity  12 

2.3  Reproductive  Strategies  14 

2.4  Biomass  Partitioning  15 

2.5  Primary  Production  17 

2.6  Herbivory  23 

2.7  Wood  Borers  24 

2.8  Mangrove  Diseases  25 

CHAPTER  3.   ECOSYSTEM  STRUCTURE  AND  FUNCTION  26 

3.1  Structural  Properties  of  Mangrove  Forests  26 

3.2  Zonation,  Succession  and  "Land  Building"  26 

3.3  Nutrient  Cycling  30 

3.4  Litter  Fall  and  Decomposition  32 

3.5  Carbon  Export  34 

3.6  Energy  Flow  36 

CHAPTER  4.   COMMUNITY  COMPONENTS  -  MICROORGANISMS  40 

CHAPTER  5.   COMMUNITY  COMPONENTS  -  PLANTS  OTHER  THAN  MANGROVES  41 

5.1  Root  and  Mud  Algae  41 

5.2  Phytoplankton  43 

5.3  Associated  Vascular  Plants  43 


CHAPTER  6 

6.1 

6.2 

6.3 

6.4 

7 

1 

7 

2 

7 

3 

7 

4 

7 

5 

7 

6 

CHAPTER  8. 

CONTENTS  (continued) 

Pase 

COMMUNITY  COMPONENTS  -  INVERTEBRATES  45 

Ecological  Relationships  45 

Arboreal  Arthropod  Communi  ty  47 

Prop  Root  and  Associated  Mud  Surface  Community  47 

Water  Column  Community  49 

CHAPTER  7.  COMMUNITY  COMPONENTS  -  FISHES  50 

Basin  Mangrove  Forests  50 

Riverine  Forests  52 

Fringing  Forests  along  Estuarine  Bays  and  Lagoons  54 

Fringing  Forests  along  Oceanic  Bays  and  Lagoons  56 

Overwash  Mangrove  Islands  56 

Gradient  of  Mangrove  Community  Interactions  57 

COMMUNITY  COMPONENTS  -  AMPHIBIANS  AND  REPTILES  58 

CHAPTER  9.   COMMUNITY  COMPONENTS  -  BIRDS  61 

9.1  Ecological  Relationships  61 

9.2  Wading  Birds  61 

9 . 3  Probi ng  Shorebi  rds  65 

9.4  Floating  and  Diving  Water  Birds  65 

9.5  Aerially-searching  Birds  67 

9.6  Birds  of  Prey  67 

9.7  Arboreal  Birds  68 

9.8  Associations  between  Mangrove  Community  Types  and  Birds  70 

9.9  Mangroves  as  Winter  Habitat  for  North  American  Migrant  Land  Birds  .  71 

CHAPTER  10.  COMMUNITY  COMPONENTS  -  MAMMALS  72 

CHAPTER  1 1  .  VALUE  OF  MANGROVE  ECOSYSTEMS  TO  MAN  74 

1.1  Shoreline  Stabilization  and  Storm  Protection  74 

1.2  Habitat  Value  to  Wildlife  74 

1.3  Importance  to  Threatened  and  Endangered  Species  75 

1.4  Value  to  Sport  and  Commercial  Fisheries  75 

1.5  Aesthetics,  Tourism  and  the  Intangibles  75 

1.6  Economic  Products  76 

CHAPTER  12.  MANAGEMENT  IMPLICATIONS  77 

12.1  Inherent  Vulnerability  77 

12.2  Man-induced  Destruction  77 

12.3  Effects  of  Oil  Spills  on  Mangroves  80 

12.4  Man-induced  Modifications  81 

12.5  Protective  Measures  Including  Transplanting  84 

12.6  Ecological  Value  of  Black  vs.  Red  Mangroves  85 

12.7  The  Importance  of  Inter-community  Exchange  85 

12.8  Management  Practices:  Preservation  86 

vi 


CONTENTS  (continued) 


Page 

REFERENCES  88 

APPENDIX  A:  SUMMARY  OF  SITE  CHARACTERISTICS  AND  SAMPLING  METHODOLOGY 

FOR  FISHES  1°6 


APPENDIX  B 
APPENDIX  C 
APPENDIX  D 
APPENDIX  E 


FISHES  OF  MANGROVE  AREAS  HO 

AMPHIBIANS  AND  REPTILES  FROM  MANGROVE  AREAS  127 

AVIFAUNA  OF  MANGROVE  AREAS  130 

MAMMALS  OF  MANGROVE  AREAS  I42 


vn 


FIGURES 
Number  Page 

1  Approximate  northern  limits  for  the  red  mangrove  (R),  black 

mangrove  (B),  and  white  mangrove  (W)  in  Florida  3 

2a   A  typical  intertidal  profile  from  south  Florida  showing  the 

distribution  of  red  and  black  mangroves  4 

2b   The  pattern  of  annual  sea  level  change  in  south  Florida  4 

3  Three  species  of  Florida  mangroves  with  propagules,  flowers  and  leaves.  6 

4  The  six  mangrove  community  types  8 

5a   Aboveground  and  belowground  biomass  of  a  Puerto  Rican  red 

mangrove  forest  16 

5b   Light  attenuation  in  a  mangrove  canopy;  canopy  height  is  8  m  16 

6  The  hypothetical  relationship  between  waterway  position  and 

community  net  primary  production  of  Florida  mangrove  forests  22 

7  The  hypothetical  relationship  between  nutrient  input,  biomass, 
productivity,  and  nutrient  export  from  mangrove  ecosystems  31 

8  Potential  pathways  of  energy  flow  in  mangrove  ecosystems  37 

9  Vertical  distribution  of  selected  algae  and  invertebrates  on 

red  mangrove  prop  roots  42 

10  Photograph  of  red  mangrove  prop  root  habitat  in  clear  shallow 

water  with  associated  animal  and  plant  populations  46 

11  Gradient  of  mangrove-associated  fish  communities  showing 
representative  species  51 

12  Aerial  photograph  of  the  mangrove  belt  of  southwest  Florida 

near  Whi  tewater  Bay  53 

13  The  mangrove  water  snake,  Nerodia  fasciata  compressicauda , 

curl  ed  on  a  red  mangrove  prop  root  59 

14  A  variety  of  wading  birds  feeding  in  a  mangrove-lined  pool 

near  Flamingo,  Florida  62 

15  Osprey  returning  to  its  nest  in  a  red  mangrove  near  Whitewater  Bay  69 

16  Damaged  stand  of  red  and  black  mangroves  near  Flamingo,  Florida, 

as  it  appeared  7  years  after  Hurricane  Donna  78 

17  Mangrove  forest  near  Key  West  as  it  appeared  in  1981  after 

being  destroyed  by  diking  and  impounding  79 

18  Mangrove  islands  in  Florida  Bay  near  Upper  Matecumbe  Key  87 

TABLES 
Number  Page 

la  Estimates  of  mangrove  production  in  Florida 20 

lb  Comparative  measurements  of  photosynthesis  in  gC/m2/day  21 

lc  Gross  primary  production  at  different  salinities  21 

2  Aboveground  biomass  of  mangrove  forests  in  the  Ten  Thousand 

Islands  region  of  Florida  27 

3  Estimates  of  litter  fall  in  mangrove  forests  33 

4  Estimates  of  particulate  carbon  export  from  mangrove  forests  35 

5  Nesting  statistics  of  wading  birds  and  associated  species  in 

south  Florida,  1974-75  64 

6  Timing  of  nesting  by  wading  birds  and  associated  species  in 

south  Florida  66 

7  General  response  of  mangrove  ecosystems  to  severe  oil  spills  82 

8  Estimated  impact  of  various  stages  of  oil  mining  on  mangrove  ecosystems  83 


vm 


ACKNOWLEDGMENTS 


Many  individuals  and  organizations 
contributed  significantly  to  the  creation 
of  this  publication.  Most  notable  was 
Eric  Heald  who  worked  extensively  with  us 
on  the  manuscript  and  contributed  unpub- 
lished data.  We  thank  Jeffrey  Carlton, 
Edward  Conner,  Roy  R.  Lewis,  III,  Ariel 
Lugo,  Larry  Narcisse,  Steven  Macko,  Aaron 
Mills,  Michael  Robblee,  Martin  Roessler, 
Samuel  Snedaker,  Durbin  Tabb,  Mike  Wein- 
stein,  and  Joseph  Zieman  for  information 
and  helpful  advice. 

The  draft  manuscript  was  reviewed  for 
its  scientific  content  by  Armando  de  la 
Cruz,  Thomas  Savage,  James  Kushlan  (bird 
section),  and  James  P.  Ray.  Each  of  these 


individuals  provided  information  as  well 
as  critical  comments.  We  particularly 
appreciate  the  unselfish  help  of  Ken  Adams 
and  the  staff  of  the  National  Coastal  Eco- 
systems Team  of  the  U.S.  Fish  and  Wildlife 
Service.  Janet  Ryan  spent  many  hours  typ- 
ing the  manuscript. 

Factual  errors  and  faulty  conclu- 
sions are  the  sole  responsibility  of  the 
authors.  Carole  Mclvor  has  taken  primary 
responsibility  for  Chapter  7,  Tom  Smith 
for  Chapters  8,  9,  and  10,  and  Bill  Odum 
for  the  remainder  of  the  publication. 
Unless  otherwise  noted,  photographs, 
figures,  and  the  cover  were  produced  by 
the  authors. 


IX 


CHAPTER   1 


INTRODUCTION 


1.1      "MANGROVE"   DEFINITION 

The  term  "mangrove"  expresses  two 
distinctly  different  concepts.  One  usage 
refers  to  halophytic  species  of  trees  and 
shrubs  (halophyte  =  plant  growing  in 
saline  soil).  In  this  sense,  mangrove  is 
a  catch-all,  botanically  diverse,  non- 
taxonomic  expression  given  to  approximate- 
ly 12  families  and  more  than  50  species 
(Chapman  1970)  of  tropical  trees  and 
shrubs  (see  Waisel  1972  for  a  detailed 
list).  While  not  necessarily  closely 
related,  all  these  plants  are  adapted  to 
(1)  loose,  wet  soils,  (2)  a  saline  habi- 
tat, (3)  periodic  tidal  submergence,  and 
(4)  usually  have  degrees  of  viviparity  of 
propagules  (see  section  2.3  for  discussion 
of  "viviparity"  and  "propagules"). 

The  second  usage  of  the  term  mangrove 
encompasses  the  entire  plant  community 
including  individual  mangrove  species. 
Synonymous  terms  include  tidal  forest, 
tidal  swamp  forest,  mangrove  community, 
mangrove  ecosystem,  mangal  (Macnae  1968), 
and  mangrove  swamp. 

For  consistency,  in  this  publication 
we  will  use  the  word  "mangrove"  for  indi- 
vidual kinds  of  trees;  mangrove  community, 
mangrove  ecosystem  or  mangrove  forest  will 
represent  the  entire  assemblage  of  "man- 
groves". 


1.2      FACTORS    CONTROLLING    MANGROVE    DISTRI- 
BUTION 

Four  major  factors  appear  to  limit 
the  distribution  of  mangroves  and  deter- 
mine the  extent  of  mangrove  ecosystem 
development.  These  factors  include  (1) 
climate,  (2)  salt  water,  (3)  tidal  fluc- 
tuation,   and   (4)   substrate. 


Climate 

Mangroves  are  tropical  species  and 
do  not  develop  satisfactorily  in  regions 
where  the  annual  average  temperature  is 
below  19°C  or  66°F  (Waisel  1972). 
Normally,  they  do  not  tolerate  temperature 
fluctuations    exceeding    10°C     (18°F)    or 


temperatures  below  freezing  for  any  length 
of  time.  Certain  species,  for  example, 
black  mangrove,  Avicennia  germinans,  on 
the  northern  coast  of  the  Gulf  of  Mexico, 
maintain  a  semi -permanent  shrub  form  by 
growing  back  from  the  roots  after  freeze 
damage. 

Lugo  and  Zucca  (1977)  discuss  the 
impact  of  low  temperature  stress  on  Flori- 
da mangroves.  They  found  that  mangrove 
communities  respond  to  temperature  stress 
by  decreasing  structural  complexity  (de- 
creased tree  height,  decreased  leaf  area 
index,  decreased  leaf  size,  and  increased 
tree  density).  They  concluded  that  man- 
groves growing  under  conditions  of  high 
soil  salinity  stress  are  less  tolerant  of 
low  temperatures.  Presumably,  other  types 
of  stress  (e.g.,  pollutants,  diking)  could 
reduce  the  temperature  tolerance  of  man- 
groves. 

High  water  temperatures  can  also  be 
limiting.  McMillan  (1971)  reported  that 
seedlings  of  black  mangrove  were  killed  by 
temperatures  of  39°  to  40°C  (102°  to 
104°F)  although  established  seedlings  and 
trees  were  not  damaged.  To  our  knowledge, 
upper  temperature  tolerances  for  adult 
mangroves  are  not  well  known.  We  suspect 
that  water  temperatures  in  the  range  42° 
to  45°C  (107°  to   113°F)   may  be   limiting. 


Salt  Water 

Mangroves  are  facultative  halo- 
phytes,  i.e.,  salt  water  is  not  a  physical 
requirement  (Bowman  1917;  Egler  1948).  In 
fact,  most  mangroves  are  capable  of 
growing  quite  well  in  freshwater  (Teas 
1979).  It  is  important  to  note,  however, 
that  mangrove  ecosystems  do  not  develop  in 
strictly  freshwater  environments;  salinity 
is  important  in  reducing  competition  from 
other  vascular  plant  species  (Kuenzler 
1974).  See  section  2.2  about  salinity 
tolerance  of  mangrove   species. 


Tidal   Fluctuation 

While    tidal    influence    is    not    a 
direct    physiological     requirement    for 


mangroves,  it  plays  an  important  indirect 
role.  First,  tidal  stress  (alternate 
wetting  and  drying),  in  combination  with 
salinity,  helps  exclude  most  other 
vascular  plants  and  thus  reduces  competi- 
tion. Second,  in  certain  locations,  tides 
bring  salt  water  up  the  estuary  against 
the  outward  flow  of  freshwater  and  allow 
mangroves  to  become  established  well 
inland.  Third,  tides  may  transport 
nutrients  and  relatively  clean  water  into 
mangrove  ecosystems  and  export  accumula- 
tions of  organic  carbon  and  reduced  sulfur 
compounds.  Fourth,  in  areas  with  high 
evaporation  rates,  the  action  of  the  tides 
helps  to  prevent  soil  salinities  from 
reaching  concentrations  which  might  be 
lethal  to  mangroves.  Fifth,  tides  aid  in 
the  dispersal  of  mangrove  propagules  and 
detritus. 

Because  of  all  of  these  factors, 
termed  tidal  subsidies  by  E.P.  Odum 
(1971),  mangrove  ecosystems  tend  to  reach 
their  greatest  development  around  the 
world  in  low-lying  regions  with  relatively 
large  tidal  ranges.  Other  types  of  water 
fluctuation,  e.g.,  seasonal  variation  in 
freshwater  runoff  from  the  Florida  Ever- 
glades,   can  provide  similar  subsidies. 


Substrate  and  Wave  Energy 

Mangroves  grow  best  in  depositional 
environments  with  low  wave  energy.  High 
wave  energy  prevents  establishment  of 
propagules,  destroys  the  relatively  shal- 
low mangrove  root  system  and  prevents  the 
accumulation  of  fine  sediments.  The  most 
productive  mangrove  ecosystems  develop 
along  deltaic  coasts  or  in  estuaries  that 
have  fine-grained  muds  composed  of  silt, 
clay  and  a  high  percentage  of  organic 
matter.  Anaerobic  sediments  pose  no 
problems  for  mangroves  (see  section  2.1) 
and  exclude  competing  vascular  plant 
species. 


1.3      GEOGRAPHICAL    DISTRIBUTION 

Mangroves  dominate  approximately  75% 
of  the  world's  tropical  coastline  between 
25°N  and  25°S  latitude  (McGill   1959).     On 


the  east  coast  of  Africa,  in  Australia  and 
in  New  Zealand,  they  extend  10°  to  15° 
farther  south  (Kuenzler  1974)  and  in 
Japan,  Florida,  Bermuda,  and  the  Red  Sea 
they  extend  5°  to  7°  farther  north.  These 
areas  of  extended  range  generally  occur 
where  oceanographic  conditions  move  un- 
usually warm  water  away  from  the  equator. 

Although  certain  reqions  such  as  the 
tropical  Indo-Pacific  have  as  many  as  30 
to  40  species  of  mangroves  present,  only 
three  species  are  found  in  Florida:  the 
red  mangrove,  Rhizophora  mangle,  the  black 
mangrove,  Avicennia  germi  nans,  and  the 
white  mangrove,  Laguncularia  racemosa.  A 
fourth  species,  buttonwood,  Conocarpus 
erecta,  is  not  a  true  mangrove  (no  ten- 
dency  to  vivipary  or  root  modification), 
but  is  an  important  species  in  the  transi- 
tion zone  on  the  upland  edge  of  mangrove 
ecosystems    (Tomlinson    1980). 

The  ranges  of  mangrove  species  in 
Florida  have  fluctuated  over  the  past 
several  centuries  in  response  to  relative- 
ly short-term  climatic  change.  Currently, 
the  situation  is  as  follows  (Figure  1). 
The  red  mangrove  and  the  white  mangrove 
have  been  reported  as  far  north  as  Cedar 
Key  on  the  west  coast  of  Florida  (Rehm 
1976)  and  north  of  the  Ponce  de  Leon  Inlet 
on  the  east  coast  (Teas  1977);  both  of 
these  extremes  lie  at  approximately  29°10' 
N  latitude.  Significant  stands  lie  south 
of  Cape  Canaveral  on  the  east  coast  and 
Tarpon  Springs  on  the  west  coast.  The 
black  mangrove  has  been  reported  as  far 
north  as  30°N  latitude  on  the  east  coast 
of  Florida  (Savage  1972)  and  as  scattered 
shrubs  along  the  north  coast  of  the  Gulf 
of   Mexico. 


Intertidal   Distribution 

The  generalized  distribution  of  the 
red  and  black  mangrove  in  relation  to  the 
intertidal  zone  is  shown  in  Figure  2a. 
Local  variations  and  exceptions  to  this 
pattern  occur  commonly  in  response  to 
localized  differences  in  substrate  type 
and  elevation,  rates  of  sea  level  rise, 
and  a  variety  of  other  factors  (see  sec- 
tion  3.2   for  a  full    discussion  of  mangrove 


R,W 

Cedar  Key 


Tarpon  Sprin 


Tampa  Bay 


Port  Charlotte  Harbor/ 
Sanibel  Island      \^ 


R,W 


30°  N 


Ponce  de  Leon  Inlet 


Cape  Canaveral 
Indian  River 


Rookery  Bay 
Ten  Thousand  Islands 
Shark  River  Vall< 

Whitewater  Bay/North  River 


Florida  Keys  # 


Biscayne  Bay 


25°N 


Figure  1.  Approximate  northern  limits  for  the  red  mangrove  (R),  black  mangrove 
(B),  and  white  mangrove  (W)  in  Florida  (based  on  Savage  1972);  although  not  in- 
dicated in  the  figure,  the  black  mangrove  extends  along  the  northern  Gulf  of  Mex- 
ico as  scattered  shrubs. 


(A) 


Mnvv  — 
MLW-- 

—  —  .— f**r '  ■*  ■ 

^PEAT 

*"**       \          y\ 

,                    ■  • 

LOW 

HIGH 

MARSH 

MARSH 

(RED   MANGROVE 

(BLACK    MANGROVE 

DOMINATED) 

DOMINATED) 

/ 


(B) 


FMAMJJASOND 


+  10   cm   r 


+  5   cm  - 


SEA    LEVEL 


—  5   cm    - 


-10   cm   L 


Figure  2.      (a)     A  typical    intertidal    profile  from  south  Florida  showing  the  dis- 
tribution of  red  and  black  mangrove   (adapted  from  Provost  1974).      (b)     The  pat- 
tern of  annual    sea  level    change  in  south   Florida   (Miami ) (adapted  from  Provost 
1974). 


zonation).  Furthermore,  it  is  important 
to  recognize  that  the  intertidal  zone  in 
most  parts  of  Florida  changes  seasonally 
(Provost  1974);  there  is  a  tendency  for 
sea  level  to  be  higher  in  the  fall  than  in 
the  spring  (Figure  2b).  As  a  result  the 
"high  marsh"  may  remain  totally  dry  during 
the  spring  and  be  continually  submerged  in 
the  autumn.  This  phenomenon  further  com- 
plicates the  textbook  concept  of  the  in- 
tertidal, "low  marsh"  red  mangrove  and  the 
infrequently  flooded,  "high  marsh"  black 
mangrove. 


Mangrove  Acreage  in  Florida 

Estimates  of  the  total  acreage 
occupied  by  mangrove  communities  in 
Florida  vary  widely  between  430,000  acres 
and  over  500,000  acres  (174,000  ha  to  over 
202,000  ha).  Eric  Heald  (Tropical 
Bioindustries,  9869  Fern  St.,  Miami,  Fl a.; 
personal  communication  1981)  has 
identified  several  reasons  for  the  lack  of 
agreement  between  estimates.  These 
include:  (1)  inclusion  or  exclusion  in 
surveys  of  small  bays,  ponds  and  creeks 
which  occur  within  mangrove  forests,  (2) 
incorrect  identification  of  mangrove  areas 
from  aerial  photography  as  a  result  of 
inadequate  "ground-truth"  observations, 
poorly  controlled  aerial  photography,  and 
simple  errors  of  planimetry  caused  by 
photography   of  inadequate   scale. 

The  two  most  detailed  estimates  of 
area  covered  by  mangroves  in  Florida  are 
provided  by  the  Coastal  Coordinating  Coun- 
cil, State  of  Florida  (1974)  and  Birnhak 
and  Crowder  (1974).  Considerable  dif- 
ferences exist  between  the  two  estimates. 
The  estimate  of  Birnhak  and  Crowder 
(1974),  which  is  limited  to  certain  areas 
of  south  Florida,  appears  to  be  unreal is- 
tically  high,  particularly  for  Monroe 
County  (Eric  Heald,  personal  communication 
1981).  Coastal  Coordinating  Council 
(1974)  estimates  a  total  of  469,000  acres 
(190,000  ha)  within  the  State  and  suggests 
an  expected  margin  of  error  of  15%  (i.e. 
their  estimate  lies  between  400,000  and 
540,000  acres  or  162,000  and  219,000  ha). 


According  to  this  survey,  ninety  percent 
of  Florida's  mangroves  are  located  in  the 
four  southern  counties  of  Lee  (35,000 
acres  or  14,000  ha),  Collier  (72,000  acres 
or  29,000  ha),  Monroe  (234,000  acres  or 
95,000  ha),  and  Dade  (81,000  acres  or 
33,000  ha). 

Much  of  the  area  covered  by  mangroves 
in  Florida  is  presently  owned  by  Federal, 
State  or  County  governments,  or  by  non- 
profit organizations  such  as  the  National 
Audubon  Society.  Approximately  280,000 
acres  (113,000  ha)  fall  into  this  category 
(Eric  Heald,  personal  communication  1981). 
Most  of  this  acreage  is  held  by  the 
Federal  Government  as  a  result  of  the  land 
being  including  within  the  Everglades 
National   Park. 


1.4     MANGROVE   SPECIES  DESCRIPTIONS 

The  following  descriptions  come 
largely  from  Carlton  (1975)  and  Savage 
(1972);  see  these  publications  for  further 
comments  and  photographs.  For  more 
detailed  descriptions  of  germinating  seeds 
(propagules)  see  section  2.3.  The  three 
species  are  shown   in   Figure  3. 


The  Black  Mangrove   (Avicennia   germinans) 

Avicennia  germinans  is  synonymous 
with  A.  nitida  and  is  a  member  of  the 
family  Avicenniaceae  (formerly  classed 
under  Verbenaceae).  The  tree  may  reach  a 
height  of  20  m  (64  ft)  and  has  dark,  scaly 
bark.  Leaves  are  5  to  10  cm  (2  to  4 
inches)  in  length,  narrowly  elliptic  or 
oblong,  shiny  green  above  and  covered  with 
short,  dense  hairs  below.  The  leaves  are 
frequently  encrusted  with  salt.  This  tree 
is  characterized  by  long  horizontal  or 
"cable"  roots  with  short  vertical  aerating 
branches  (pneumatophores)  that  profusely 
penetrate  the  substrate  below  the  tree. 
Propagules  are  lima-bean  shaped,  dark 
green  while  on  the  tree,  and  several 
centimeters  (1  inch)  long.  The  tree 
flowers   in   spring  and  early  summer. 


Black  Mangrove,  Avicennia  germinans 


White  Mangrove,  Laguncularia  racemosa 


Red  Mangrove,  Rhizophora  mangle 


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


The  White  Mangrove  (Laguncularia  racemosa) 

The  white  mangrove  is  one  of  450 
species  of  plants  in  18  genera  of  the 
family  Combretaceae  (synonymous  with 
Terminal iaceae).  It  is  a  tree  or  shrub 
reaching  15  m  (49  ft)  or  more  in  height 
with  broad,  flattened  oval  leaves  up  to  7 
cm  (3  inches)  long  and  rounded  at  both 
ends.  There  are  two  salt  glands  at  the 
apex  of  the  petiole.  The  propagule  is 
very  small  (1.0  to  1.5  cm  or  0.4  to  0.6 
inches  long)  and  broadest  at  its  apex. 
Flowering  occurs  in  spring  and  early 
summer. 


The  Red  Mangrove  (Rhizophora  mangle) 

The  red  mangrove  is  one  of  more  than 
70  species  in  17  genera  in  the  family 
Rhizophoraceae.  This  tree  may  reach  25  m 
(80  ft)  in  height,  has  thin  grey  bark  and 
dark  red  wood.  Leaves  may  be  2  to  12  cm 
(1  to  5  inches)  long,  broad  and  blunt- 
pointed  at  the  apex.  The  leaves  are 
shiny,  deep  green  above  and  paler  below. 
It  is  easily  identified  by  its  charac- 
teristic "prop  roots"  arising  from  the 
trunk  and  branches.  The  penci 1 -shaped 
propagules  are  as  much  as  25  to  30  cm  (10 
to  12  inches)  long  after  germination.  It 
may  flower  throughout  the  year,  but  in 
Florida  flowering  occurs  predominately  in 
the  spring  and  early  summer. 


1.5     MANGROVE  COMMUNITY  TYPES 

Mangrove  forest  communities  exhibit 
tremendous  variation  in  form.  For 
example,  a  mixed  scrub  forest  of  black  and 
red  mangroves  at  Turkey  Point  on  Biscayne 
Bay  bears  little  resemblance  to  the 
luxuriant  forests,  dominated  by  the  same 
two  species,   along  the  lower  Shark  River. 

Lugo  and  Snedaker  (1974)  provided  a 
convenient  classification  system  based  on 
mangrove  forest  physiogomy.  They  identi- 
fied six  major  community  types  resulting 
from  different  geological  and  hydrological 
processes.  Each  type  has  its  own  charac- 
teristic set  of  environmental  variables 
such  as  soil   type  and  depth,    soil    salinity 


range,  and  flushing  rates.  Each  community 
type  has  characteristic  ranges  of  primary 
production,  litter  decomposition  and  car- 
bon export  along  with  differences  in 
nutrient  recycling  rates,  and  community 
components.  The  community  types  as  shown 
in  Figure  4  are  as  follows: 


(1)  Overwash  mangrove  forests  - 
these  islands  are  frequently  overwashed  by 
tides  and  thus  have  high  rates  of  organic 
export.  All  species  of  mangroves  may  be 
present,  but  red  mangroves  usually  domi- 
nate. Maximum  height  of  the  mangroves  is 
about  7  m  (23  ft). 

(2)  Fringe  mangrove  forests  -  man- 
groves form  a  relatively  thin  fringe  along 
waterways.  Zonation  is  typically  as  de- 
scribed by  Davis  (1940)  (see  discussion  in 
section  3.2).  These  forests  are  best 
defined  along  shorelines  whose  elevations 
are  higher  than  mean  high  tide.  Maximum 
height  of  the  mangroves  is  about  10  m  (32 
ft). 

(3)  Riverine  mangrove  forests  -  this 
community  type  includes  the  tall  flood 
plain  forests  along  flowing  waters  such  as 
tidal  rivers  and  creeks.  Although  a  shal- 
low berm  often  exists  along  the  creek 
bank,  the  entire  forest  is  usually  flushed 
by  daily  tides.  All  three  species  of 
mangroves  are  present,  but  red  mangroves 
(with  noticeably  few,  short  prop  roots) 
predominate.  Mangroves  may  reach  heights 
of  18  to  20  m  (60  to  65  ft). 

(4)  Basin  mangrove  forests  -  these 
forests  occur  inland  in  depressions  chan- 
neling terrestrial  runoff  toward  the 
coast.  Close  to  the  coast  they  are  in- 
fluenced by  daily  tides  and  are  usually 
dominated  by  red  mangroves.  Moving  in- 
land, the  tidal  influence  lessens  and 
dominance  shifts  to  black  and  white  man- 
groves. Trees  may  reach  15  m  (49  ft)  in 
height. 

(5)  Hammock  forests  -  hammock  man- 
grove communities  are  similar  to  the  basin 
type  except  that  they  occur  on  ground  that 
is  slightly  elevated  (5  to  10  cm  or  2  to  4 
inches)    relative    to    surrounding    areas. 


(1)   OVERWASH    FOREST  (2)    FRINGE    FOREST 


(3)    RIVERINE    FOREST  (4)    BASIN    FOREST 


-^ggl  ScL    £L ' 


(5)    HAMMOCK    FOREST  (6)    SCRUB    FOREST 

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


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

(6)  Scrub  or  dwarf  forests  -  this 
community  type  is  limited  to  the  flat 
coastal  fringe  of  south  Florida  and  the 
Florida  Keys.  All  three  species  are 
present.  Individual  plants  rarely  exceed 
1.5  m  (4.9  ft)  in  height,  except  where 
they  grow  over  depressions  filled  with 
mangrove  peat.  Many  of  these  tiny  trees 
are  40  or  more  years  of  age.  Nutrients 
appear  to  be  limiting  although  substrate 
(usually   limestone   marl)  must  play  a  role. 

Throughout  this  publication  we  have 
attempted  to  refer  to  Lugo  and  Snedaker's 
classification  scheme  wherever  possible. 
Without  a  system  of  this  type,  comparisons 
between  sites  become  virtually 
meaningless. 


1.6     SUBSTRATES 

Understanding  mangrove-substrate 
relationships  is  complicated  by  the 
ability  of  mangroves  to  grow  on  many  types 
of  substrates  and  because  they  often  alter 
the  substrate  through  peat  formation  and 
by  altering  patterns  of  sedimentation.  As 
a  result,  mangroves  are  found  on  a  wide 
variety  of  substrates  including  fine, 
inorganic  muds,  muds  with  a  high  organic 
content,  peat,  sand,  and  even  rock  and 
dead  coral  if  there  are  sufficient 
crevices  for  root  attachment.  Mangrove 
ecosystems,  however,  appear  to  flourish 
only  on  muds  and   fine-grained   sands. 

In  Florida,  the  primary  mangrove 
soils  are  either  calcareous  marl  muds  or 
calcareous  sands  in  the  southern  part  of 
the  State  and  siliceous  sands  farther 
north  (Kuenzler  1974).  Sediment  distribu- 
tion and,  hence,  mangrove  development,  is 
controlled  to  a  considerable  extent  by 
wave  and  current  energy.  Low  energy 
shorelines  accumulate  fine-grained  sedi- 
ments such  as  mud  and  silt  and  usually 
have  the  best  mangrove  growth.  Higher 
energy  shorelines  (more  wave  action  or 
higher  current  velocities)  are  charac- 
terized by  sandy  sediments  and  less  pro- 
ductive  mangroves.      If   the   wave   energy 


becomes  too  great,  mangroves  will  not  be 
present.  Of  the  three  species  of  Florida 
mangroves,  white  mangroves  appear  to 
tolerate  sandy  substrates  the  best  (per- 
sonal observation),  possibly  because  this 
species  may  tolerate  a  greater  depth  to 
the  water  table  than  the  other  two 
species. 

Mangroves  in  Florida  often  modify  the 
underlying  substrate  through  peat  deposi- 
tion. It  is  not  unusual  to  find  layers  of 
mangrove  peat  several  meters  thick  under- 
lying well-established  mangrove  ecosystems 
such  as  those  along  the  southwest  coast  of 
Florida.  Cohen  and  Spackman  (1974)  pre- 
sented a  detailed  account  of  peat  forma- 
tion within  the  various  mangrove  zones  of 
south  Florida  and  also  in  areas  dominated 
by  black  needle  rush  (Juncus  roemerianus), 
smooth  cordgrass  (Sparti  na  al  terni  fl  ora) 
and  a  variety  of  other  macrophytes;  Cohen 
and  Spackman  (1974)  also  provide  descrip- 
tions and  photography  to  aid  in  the  iden- 
tification of  unknown   peat   samples. 

The  following  descriptions  come  from 
Cohen  and  Spackman  (1974)  and  from  the 
personal  observations  of  W.E.  Odum  and 
E.J.  Heald.  Red  mangroves  produce  the 
most  easily  recognized  peat.  More  recent 
deposits  are  spongy,  fibrous  and  composed 
to  a  great  extent  of  fine  rootlets  (0.2  to 
3.0  mm  in  diameter).  Also  present  are 
larger  pieces  of  roots  (3  to  25  mm),  bits 
of  wood  and  leaves,  and  inorganic 
materials  such  as  pyrite,  carbonate 
minerals,  and  quartz.  Older  deposits  are 
less  easily  differentiated  although  they 
remain  somewhat  fibrous.  Peat  which  has 
recently  been  excavated  is  reddish-brown 
although  this  changes  to  brown-black  after 
a  short  exposure  to  air.  Older  deposits 
are  mottled  reddish-brown;  deposits  with  a 
high  content  of  carbonates  are  greyish- 
brown   upon   excavation. 

Cohen  and  Spackman  (1974)  were  unable 
to  find  deposits  of  pure  black  mangrove  or 
white  mangrove  peat  suggesting  that  these 
two  species  may  not  form  extensive  depos- 
its of  peat  while  growing  in  pure  stands. 
There  are,  however,  many  examples  of  peats 
which  are  mixtures  of  red  mangrove 
material  and  black  mangrove  roots.     They 


suggested  that  the  black  mangrove  peats 
identified  by  Davis  (1946)  were  probably 
mixtures  of  peat   from  several    sources. 

Throughout  south  Florida  the  sub- 
strate underlying  mangrove  forests  may 
consist  of  complicated  patterns  of 
calcareous  muds,  marls,  shell,  and  sand 
interspersed  and  overlain  by  layers  of 
mangrove  peat  and  with  limestone  bedrock 
at  the  bottom.  Detailed  descriptions  of 
this  complex  matrix  and  its  spatial  varia- 
tion were  given  by  Davis  (1940,  1943, 
1946),  Egler  (1952),  Craighead  (1964), 
Zieman  (1972)  and  Cohen  and  Spackman 
(1974)  among  others.  Scoffin  (1970)  dis- 
cussed the  ability  of  red  mangrove  to 
trap  and  hold  sediments  about  its  prop 
roots.  So  called  "land-building"  by  man- 
groves   is    discussed    in   section   3.2. 

The  long-term  effect  of  mangrove  peat 
on  mangrove  distribution  is  not  entirely 
clear.  Certainly,  if  there  is  no  change 
in  sea  level  or  if  erosion  is  limited,  the 
accumulation  of  peat  under  stands  of  red 
mangroves  combined  with  deposition  and 
accumulation  of  suspended  sediments  will 
raise  the  forest  floor  sufficiently  to 
lead  to  domination  by  black  or  white  man- 
groves and,  ultimately,  more  terrestrial 
species.  Whether  this  is  a  common  se- 
quence of  events  in  contemporary  south 
Florida  is  not  clear.  It  is  clear  that 
peat  formtion  is  a  passive  process  and 
occurs  primarily  where  and  when  physical 
processes  such  as  erosion  and  sea  level 
rise  are  of  minimal  importance  (Wanless 
1974). 

Zieman  (1972)  presented  an  inter- 
esting argument  suggesting  that  mangrove 
peat  may  be  capable  of  dissolving  under- 
lying limestone  rock,  since  carbonates  may 
dissolve  at  pH  7.8.  Through  this  process, 
shallow  depressions  might  become  deeper 
and  the  overlying  peat  layer  thicker 
without  raising  the  surface  of  the  forest 
floor. 

Data  on  chemical  characteristics  of 
Florida  mangrove  soils  and  peat  are 
limited.  Most  investigators  have  found 
mangrove  substrates  to  be  almost  totally 
anaerobic.      Lee    (1969)    recorded   typical    Eh 


values  of  -100  to  -400  mv  in  mangrove 
peats.  Such  evidence  of  strongly  reducing 
conditions  are  not  surprising  considering 
the  fine-grained,  high  organic  nature  of 
most  mangrove  sediments.  Although  man- 
groves occur  in  low  organic  sediments 
(less  than  1%  organic  matter),  typical 
values  for  mangrove  sediments  are  10%  to 
20%  organic   matter. 

Lee  (1969)  analyzed  3,000-  to  3,500- 
year-old  mangrove  peat  layers  underlying 
Little  Black  Water  Sound  in  Florida  Bay 
for  lipid  carbon  content.  Peat  lipid 
content  varied  between  0.6  and  2.7  mg 
lipid-C/gram  of  peat  (dry  wt  )  or  about  3% 
of  the  total  organic  carbon  total.  These 
values  usually  increased  with  depth.  Long 
chain  fatty  acids  (C-16  and  C-18)  were  the 
dominant   fatty   acids   found. 

Florida  mangrove  peats  are  usually 
acidic,  although  the  presence  of  carbonate 
materials  can  raise  the  pH  above  7.0. 
Zieman  (1972)  found  red  mangrove  peats  to 
range  from  pH  4.9  to  6.8;  the  most  acid 
conditions  were  usually  found  in  the  cen- 
ter of  the  peat  layer.  Lee  (1969)  re- 
corded a  pH  range  from  5.8  to  6.8  in  red 
mangrove  peat  at  the  bottom  of  a  shallow 
embayment.  Although  Davis  (1940)  found  a 
difference  between  red  mangrove  peat  (5.0 
to  5.5)  and  black  mangrove  peat  (6.9  to 
7.2),  this  observation  has  not  been  con- 
firmed because  of  the  previously  mentioned 
difficulty  in  finding  pure  black  mangrove 
peat. 

Presumably,  the  acidic  character  of 
mangrove  peat  results  from  release  of 
organic  acids  during  anaerobic  decomposi- 
tion and  from  the  oxidation  of  reduced 
sulfur  compounds  if  the  peat  is  dried  in 
the  presence  of  oxygen.  This  last  point 
explains  why  "reclaimed"  mangrove  areas 
often  develop  highly  acidic  soils  (pH  3.5 
to  5.0)  shortly  after  reclamation.  This 
"cat  clay"  problem  has  greatly  complicated 
the  conversion  of  mangrove  regions  to 
agricultural  land  in  Africa  and  southeast 
Asia  (Hesse  1961;  Hart  1962,  1963;  Macnae 
1968). 

In  summary,  although  current       under- 
standing of  mangrove     peats     and     soils  is 


10 


fragmentary  and  often  contradictory,   we 
can  outline   several    generalizations: 

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

(2)  Mangrove  ecosystems  appear  to 
flourish  on  fine-grained  sediments  which 
are  usually  anaerobic  and  may  have  a  high 
organic  content. 

(3)  Mangrove  ecosystems  which  per- 
sist for  some  time  may  modify  the  under- 
lying substrate  through  peat  formation. 
This  appears  to  occur  only  in  the  absence 
of   strong   physical    forces. 

(4)  Mangrove  peat  is  formed  pri- 
marily by  red  mangroves  and  consists  pre- 
dominantly of  root  material. 

(5)  Red  mangrove  peats  may  reach 
thicknesses  of  several  meters,  have  a 
relatively  low  pH,  and  may  be  capable  of 
dissolving  underlying  layers   of   limestone. 

(6)  When  drained,  dried,  and 
aerated,  mangrove  soils  usually  experience 
dramatic  increases  in  acidity  due  to  the 
oxidation  of  reduced  sulfur  compounds. 
This  greatly  complicates  their  conversion 
to  agricul  ture. 


1.7     WATER   QUALITY 


from  virtually  fresh  water  to  above  40  ppt 
(discussed  in  section  2.2),  (2)  low  macro- 
nutrient  concentrations  (particularly 
phosphorous),  (3)  relatively  low  dissolved 
oxygen  concentrations,  and  (4)  frequently 
increased  water  color  and  turbidity.  The 
last  three  characteristics  are  most  pro- 
nounced in  extensive  mangrove  ecosystems 
such  as  those  adjacent  to  the  Everglades 
and  least  pronounced  in  small,  scattered 
forests  such  as  the  overwash  islands  in 
the    Florida    Keys. 


In  general,  the  surface  waters 
associated  with  mangroves  are  charac- 
terized  by     (1)  a  wide   range  of  salinities 


The  results  of  oxygen  depletion  and 
nutrient  removal  are  (1)  dissolved  oxygen 
concentrations  below  saturation,  typically 
2  to  4  ppm  and  often  near  zero  in  stagnant 
locations  and  after  heavy,  storm-generated 
runoff,  (2)  very  low  total  phosphorus 
values,  frequently  below  detection  limits, 
and  (3)  moderate  total  nitrogen  values 
(0.5  to  1.5  mg/1).  In  addition,  TOC 
(total  organic  carbon)  may  range  from  4  to 
50  ppm  or  even  higher  after  rain;  Eric 
Heald  (personal  communication  1981)  has 
measured  DOC  (dissolved  organic  carbon) 
values  as  high  as  110  ppm  in  water  flowing 
from  mangroves  to  adjacent  bays.  Tur- 
bidity usually  falls  in  the  1  to  15  JTU 
(Jackson  turbity  units)  range.  The  pH  of 
the  water  column  in  Florida  swamps  is 
usually  between  6.5  and  8.0  and  alkalinity 
between  100  to  300  mg/1.  Obviously,  ex- 
ceptions to  all  of  these  trends  can  occur. 
Both  natural  and  human  disturbance  can 
raise  macronutrient   levels  markedly. 


11 


CHAPTER  2.   AUTECOLOGY  OF  MANGROVES 


2.1       ADAPTATIONS    TO    NATURAL    STRESS    - 
ANAEROBIC  SEDIMENTS 

Mangroves  have  a  series  of  remarkable 
adaptations  which  enable  them  to  flourish 
in  an  environment  characterized  by  high 
temperatures,  widely  fluctuating  salini- 
ties, and  shifting,  anaerobic  substrates. 
In  this  section  we  review  a  few  of  the 
most   important   adaptations. 

The  root  system  of  mangroves  provides 
the  key  to  existence  upon  unfriendly  sub- 
strates (see  Gill  and  Tomlinson  1971  for 
an  anatomical  review  of  mangrove  roots). 
Unlike  most  higher  plants,  mangroves 
usually  have  highly  developed  aerial  roots 
and  modest  below-ground  root  systems.  The 
aerial  roots  allow  atmospheric  gases  to 
reach  the  underground  roots  which  are 
embedded  in  anaerobic  soils.  The  red 
mangrove  has  a  system  of  stilt  or  prop 
roots  which  extend  a  meter  (3  ft)  or  more 
above  the  surface  of  the  soil  and  contain 
many  small  pores  (lenticels)  which  at  low 
tide  allow  oxygen  to  diffuse  into  the 
plant  and  down  to  the  underground  roots  by 
means  of  open  passages  called  aerenchyma 
(Scholander  et  al.  1955).  The  lenticels 
are  highly  hydrophobic  and  prevent  water 
penetration  into  the  aerenchyma  system 
during   high   tide    (Waisel    1972). 

The  black  mangrove  does  not  have  prop 
roots,  but  does  have  small  air  roots  or 
pneumatophores  which  extend  vertically 
upward  from  the  underground  roots  to  a 
height  of  20  to  30  cm  (8  to  12  inches) 
above  the  soil.  These  pneumatophores 
resemble  hundreds  of  tiny  fingers  sticking 
up  out  of  the  mud  underneath  the  tree 
canopy.  At  low  tide,  air  travels  through 
the  pneumatophores  into  the  aerenchyma 
system  and  then  to  all  living  root  tis- 
sues. The  white  mangrove  usually  does  not 
have  either  prop  roots  or  pneumatophores, 
but  utilizes  lenticels  in  the  lower  trunk 
to  obtain  oxygen  for  the  aerenchyma  sys- 
tem. "Peg  roots"  and  pneumatophores  may 
be  present  in  certain  situations  (Jenik 
1967). 

Mangroves  achieve  structural  stabili- 
ty in  at  least  two  ways.  Species  such  as 
the   red   mangrove  use  the  system  of  prop 


roots  to  provide  a  more  or  less  firm  foun- 
dation for  the  tree.  Even  though  the  prop 
roots  are  anchored  with  only  a  modest 
assemblage  of  underground  roots,  the  hori- 
zontal extent  of  the  prop  root  system 
insures  considerable  protection  from  all 
but  the  worst  of  hurricanes.  Other  man- 
grove species,  including  the  black  man- 
grove, obtain  stability  with  an  extensive 
system  of  shallow,  underground  "cable" 
roots  that  radiate  out  from  the  central 
trunk  for  a  considerable  distance  in  all 
directions;  the  pneumatophores  extend  up- 
ward from  these  cable  roots.  As  in  all 
Florida  mangroves,  the  underground  root 
system  is  shallow  and  a  tap  root  is 
lacking  (Walsh  1974).  As  Zieman  (1972) 
found,  individual  roots,  particularly  of 
red  mangroves,  may  extend  a  meter  or  more 
downward  in   suitable  soils. 

From  the  standpoint  of  effectiveness 
in  transporting  oxygen  to  the  underground 
roots,  both  prop  roots  and  cable  roots 
seem  equally  effective.  From  the  perspec- 
tive of  stability,  the  prop  roots  of  red 
mangroves  appear  to  offer  a  distinct  ad- 
vantage where  wave  and  current  energies 
are  high. 

Unfortunately,  as  pointed  out  by  Odum 
and  Johannes  (1975),  the  same  structure 
which  allows  mangroves  to  thrive  in  an- 
aerobic soil  is  also  one  of  the  tree's 
most  vulnerable  components.  Exposed  por- 
tions of  the  aerial  root  system  are  sus- 
ceptible to  clogging  by  fine  suspended 
material,  attack  by  root  borers,  and  pro- 
longed flooding  (discussed  further  in 
section  12.1).  Such  extended  stress  on 
the   aerial    roots   can   kill    the  entire  tree. 


2.2      ADAPTATIONS    TO    NATURAL    STRESS    - 
SALINITY 

Mangroves  accommodate  fluctuations  and 
extremes  of  water  and  soil  salinity 
through  a  variety  of  mechanisms,  although 
not  all  mechanisms  are  necessarily  present 
in  the  same  species.  Scholander  et  al. 
(1962)  reported  experimental  evidence  for 
two  major  methods  of  internal  ion  regula- 
tion which  they  identified  in  two  dif- 
ferent   groups    of   mangroves:      (1)   the    salt 


12 


exclusion  species  and  (2)  the  salt  excre- 
tion species.  In  addition,  some  mangroves 
utilize  succulence  and  the  discarding  of 
salt-laden     organs     or     parts   (Teas  1979). 

The  salt-excluding  species,  which 
include  the  red  mangrove,  separate 
freshwater  from  sea  water  at  the  root 
surface  by  means  of  a  non-metabolic  ultra- 
filtration system  (Scholander  1968).  This 
"reverse  osmosis"  process  is  powered  by  a 
high  negative  pressure  in  the  xylem  which 
results  from  transpiration  at  the  leaf 
surface.  Salt  concentration  in  the  sap  of 
salt-excluding  mangroves  is  about  1/70  the 
salt  concentration  in  sea  water,  although 
this  concentration  is  almost  10  times 
higher  than  found  in  normal  plants 
(Scholander  et  al.  1962). 

Salt-secreting  species,  including 
black  and  white  mangroves  (Scholander 
1968),  use  salt  glands  on  the  leaf  surface 
to  excrete  excess  salt.  This  is  probably 
an  enzymatic  process  rather  than  a  physi- 
cal process  since  it  is  markedly  tempera- 
ture sensitive  (Atkinson  et  al.  1967). 
The  process  appears  to  involve  active 
transport  with  a  requirement  for  biochemi- 
cal energy  input.  As  a  group,  the  salt 
secreters  tend  to  have  sap  salt  concentra- 
tions approximately  10  times  higher  (1/7 
the  concentration  of  sea  water)  than  that 
of  the  salt   excluders. 

In  spite  of  these  two  general  tenden- 
cies, it  is  probably  safe  to  say  that 
individual  species  utilize  a  variety  of 
mechanisms  to  maintain  suitable  salt 
balance  (Albert  1975).  For  example,  the 
red  mangrove  is  an  effective,  but  not 
perfect,  salt  excluder.  As  a  result  this 
species  must  store  and  ultimately  dispose 
of  excess  salt  in  leaves  and  fruit  (Teas 
1979).  Most  salt  secreters,  including 
white  and  black  mangroves,  are  capable  of 
limited  salt  exclusion  at  the  root  sur- 
face. The  white  mangrove,  when  exposed  to 
hypersaline  conditions,  not  only  excludes 
some  salt  and  secretes  excess  salt  through 
its  salt  glands,  but  also  develops 
thickened  succulent  leaves  and  discards 
salt  during  leaf  fall  of  senescent  leaves 
(Teas  1979). 


There  appears  to  be  some  variation  in 
the  salinity  tolerance  of  Florida  man- 
groves. The  red  mangrove  is  probably 
limited  by  soil  salinities  above  60  to  65 
ppt.  Teas  (1979)  recalculated  Bowman's 
(1917)  data  and  concluded  that  transpira- 
tion in  red  mangrove  seedlings  ceases 
above  65  ppt.  Cintron  et  al.  (1978)  found 
more  dead  than  living  red  mangrove  trees 
where  interstitial  soil  salinities  ex- 
ceeded  65    ppt. 

On  the  other  hand,  white  and  black 
mangroves,  which  both  possess  salt  excre- 
tion and  limited  salt  exclusion  mech- 
anisms, can  exist  under  more  hypersaline 
conditions.  Macnae  (1968)  reported  that 
black  mangroves  can  grow  at  soil  salini- 
ties greater  than  90  ppt.  Teas  (1979) 
reported  dwarfed  and  gnarled  black  and 
white  mangroves  occurring  in  Florida  at 
soil    salinities   of  80  ppt. 

There  may  be  an  additional  factor  or 
factors  involved  in  salinity  tolerance  of 
mangroves.  McMillan  (1975)  found  that 
seedlings  of  black  and  white  mangroves 
survived  short-term  exposures  to  80  ppt 
and  150  ppt  sea  water  if  they  were  grown 
in  a  soil  with  a  moderate  clay  content. 
They  failed  to  survive  these  salinities, 
however,  if  they  were  grown  in  sand.  A 
soil  with  7%  to  10%  clay  appeared  to  be 
adequate  for  increased  protection  from 
hypersaline  conditions. 

Vegetation-free  hypersaline  lagoons 
or  bare  sand  flats  in  the  center  of  man- 
grove ecosystems  have  been  described  by 
many  authors  (e.g.,  Davis  1940;  Fosberg 
1961;  Bacon  1970).  These  features  have 
been  variously  called  salitrals  (Holdridge 
1940),  salinas,  salterns,  salt  flats,  and 
salt  barrens.  Evidently,  a  combination  of 
low  seasonal  rainfall,  occasional  inunda- 
tion by  sea  water,  and  high  evaporation 
rates  results  in  soil  salinities  above  100 
ppt,  water  temperatures  as  high  as  45°C 
(113°F)  in  any  shallow,  standing  water, 
and  subsequent  mangrove  death  (Teas  1979). 
Once  established,  salinas  tend  to  persist 
unless  regular  tidal  flushing  is  enhanced 
by  natural  or  artificial  changes  in  tidal 
circulation. 


13 


Although  salinas  occur  frequently  in 
Florida,  they  are  rarely  extensive  in 
area.  For  example,  between  Rookery  Bay 
and  Marco  Island  (south  of  Naples, 
Florida)  there  are  a  series  of  salinas  in 
the  black  mangrove-dominated  zone  on  the 
upland  side  of  the  mangrove  swamps.  These 
hypersaline  lagoons  occur  where  the  normal 
flow  of  fresh  water  from  upland  sources 
has  been  diverted,  presumably  resulting  in 
elevated  soil  salinities  during  the  dry 
winter  months. 

In  summary,  salinity  is  a  problem  for 
mangroves  only  under  extreme  hypersaline 
conditions.  These  conditions  occur  natu- 
rally in  Florida  in  irregularly  flooded 
areas  of  the  "high  swamp"  above  the  normal 
high  tide  mark  and  are  accompanied  by  high 
soil  salinities.  Florida  mangroves, 
listed  in  order  of  increasing  salinity 
tolerance,  appear  to  be  red,  white,  and 
black. 


2.3     REPRODUCTIVE   STRATEGIES 

As  pointed  out  by  Rabinowitz  (1978a), 
virtually  all  mangroves  share  two  common 
reproductive  strategies:  dispersal  by 
means  of  water  (van  der  Pijl  1972)  and 
vivipary  (Macnae  1968;  Gill  and  Tomlinson 
1969).  Vivipary  means  that  the  embryo 
develops  continuously  while  attached  to 
the  parent  tree  and  during  dispersal. 
Since  there  is  uninterrupted  development 
from  zygote  through  the  embryo  to  seedling 
without  any  intermediate  resting  stages, 
the  word  "seed"  is  inappropriate  for 
viviparous  species  such  as  mangroves;  the 
term  "propagule"  is  generally  used  in  its 
place. 

While  the  phenology  of  black  and 
white  mangroves  remains  sketchy,  Gill  and 
Tomlinson  (1971)  thoroughly  described  the 
sequence  of  flowering  in  the  red  mangrove. 
Flowering  in  this  species  may  take  place 
at  any  time  of  the  year,  at  least  in 
extreme  south  Florida,  but  reaches  a  maxi- 
mum in  the  late  spring  and  summer.  The 
flowers  open  approximately  1  to  2  months 
after  the  appearance  of  buds.  The  flower 
remains    intact    only    1    to    2    days;    this 


probably  accounts  for  the  low  fertiliza- 
tion rate,  estimated  by  Gill  and  Tomlinson 
at  0%  to  7.2%.  Propagule  development  is 
slow,  ranging  from  8  to  13  months.  Savage 
(1972)  mentions  that  on  the  Florida  gulf 
coast,  red  mangrove  propagules  mature  and 
fall  from  the  tree  from  July  to  September. 
Within  the  Everglades  National  Park,  black 
mangroves  flower  from  May  until  July  and 
bear  fruit  from  August  until  November 
while  white  mangroves  flower  from  May  to 
August  and  bear  fruit  from  July  to  October 
(Loope  1980). 

The  propagules  of  the  three  species 
of  Florida  mangroves  are  easy  to  differen- 
tiate. The  following  descriptions  all 
come  from  Rabinowitz  (1978a).  White  man- 
grove propagules  are  small  and  flattened, 
weigh  less  than  a  gram,  are  about  2  cm 
long,  are  pea-green  when  they  fall  from 
the  parent  tree,  and  turn  mud-brown  in  two 
days  or  so.  The  pericarp  (wall  of  the 
ripened  propagule)  serves  as  a  float  and 
is  not  shed  until  the  seedling  is  estab- 
lished. During  dispersal  the  radicle 
(embryonic  root)  emerges  from  the  propa- 
gule. This  germination  during  dispersal 
has  led  Savage  (1972)  to  refer  to  the 
white  mangrove   as   "semi-viviparous". 

The  propagules  of  the  black  mangrove 
when  dropped  from  the  tree  are  oblong- 
elliptical  (resemble  a  flattened  olive), 
weigh  about  1  g  and  are  about  2  cm  long. 
The  pericarp  is  lost  within  a  few  days 
after  dropping  from  the  tree;  at  this 
point  the  cotyledons  (primary  leaves) 
unfold  and  the  propagule  resembles  two 
butterflies   on   top  of  one  another. 

Propagules  of  the  red  mangrove  under- 
go extensive  vivipary  while  on  the  tree. 
When  propagules  fall  from  the  tree  they 
resemble  large  green  beans.  They  are  rod- 
shaped  with  pointed  ends,  about  20  cm 
long,    and  weigh  an  average  of  15  g. 

Propagules  of  all  three  species  float 
and  remain  viable  for  extended  periods  of 
time.  Apparently,  there  is  an  obligate 
dispersal  time  for  all  Florida  mangroves, 
i.e.,  a  certain  period  of  time  must  elapse 
during    dispersal    for   germination   to   be 


14 


complete  and  after  which  seedling  estab- 
lishment can  take  place.  Rabinowitz 
(1978a)  estimates  the  obligate  dispersal 
period  at  approximately  8  days  for  white 
mangroves,  14  days  for  black,  and  40  days 
for  red.  She  further  estimates  the  addi- 
tional time  for  root  establishment  at  5, 
7,  and  15  days  for  white,  black,  and  red 
mangroves,  respectively.  Her  estimate  for 
viable  longevity  of  the  propagules  is  35 
days  for  white  mangroves  and  110  days  for 
black.  Davis  (1940)  reports  viable  propa- 
gules of  red  mangroves  that  had  been  kept 
floating  for   12  months. 

Rabinowitz  (1978a)  also  concluded 
that  black  and  white  mangroves  require  a 
stranding  period  of  5  days  or  more  above 
the  influence  of  tides  to  take  hold  in  the 
soil.  As  a  result,  these  two  species  are 
usually  restricted  to  the  higher  portions 
of  the  mangrove  ecosystem  where  tidal 
effects    are    infrequent. 

The  elongated  red  mangrove  propagule, 
however,  has  the  potential  to  become 
established  in  shallow  water  with  tidal 
influence.  This  happens  in  at  least  two 
ways:  (1)  stranding  in  a  vertical  posi- 
tion (they  float  vertically)  or  (2) 
stranding  in  a  horizontal  position, 
rooting  and  then  vertical  erection  by  the 
plant  itself.  Lawrence  (1949)  and  Rabino- 
witz (1978a)  felt  that  the  latter  was  the 
more  common  method.  M.  Walterding  (Calif. 
Acad.  Sci.,  San  Francisco;  personal  com- 
munication 1980)  favors  vertical  estab- 
lishment; based  upon  his  observations, 
surface  water  turbulence  works  the  propa- 
gule into  the  substrate  during  falling 
tides. 

Mortality  of  established  seedlings 
seems  to  be  related  to  propagule  size. 
Working  in  Panama,  Rabinowitz  (1978b) 
found  that  the  mortality  rate  of  mangrove 
seedlings  was  inversely  correlated  with 
initial  propagule  size.  The  white  man- 
grove, which  has  the  smallest  propagule, 
has  the  highest  rate  of  seedling  mortal- 
ity. The  black  mangrove  has  an  interme- 
diate mortality  rate  while  the  red  man- 
grove, with  the  largest  propagule,  has  the 
lowest    seedling    mortality    rate.       She 


concluded  that  species  with  small 
propagules  establish  new  cohorts  annually 
but  die  rapidly,  while  species  such  as  the 
red  mangroves  may  have  long-lived  and 
often   overlapping   cohorts. 

Propagule  size  and  seedling  mortality 
rates  are  particularly  important  in  con- 
siderations of  succession  and  replacement 
in  established  mangrove  forests.  Light  is 
usually  the  most  serious  limiting  factor 
underneath  existing  mangrove  canopies. 
Rabinowitz  (1978b)  suggested  that  species 
with  short-lived  propagules  must  become 
established  in  an  area  which  already  has 
adequate  light  levels  either  due  to  tree 
fall  or  some  other  factor.  In  contrast, 
red  mangrove  seedlings  can  become  estab- 
lished under  an  existing,  dense  canopy  and 
then,  due  to  their  superior  embryonic 
reserves,  are  able  to  wait  for  months  for 
tree  fall  to  open  up  the  canopy  and  pre- 
sent an  opportunity  for  growth. 


2.4     BIOMASS    PARTITIONING 

Few  investigators  have  partitioned 
the  total  biomass,  aboveground  and  below- 
ground,  contained  in  a  mangrove  tree.  An 
analysis  of  red  mangroves  in  a  Puerto 
Rican  forest  by  Golley  et  al.  (1962)  gives 
some  insight  into  what  might  be  expected 
in  south  Florida.  Aboveground  and  below- 
ground  biomass  existed  in  a  ratio  of  1:1 
if  fine  roots  and  peat  are  ignored  (Figure 
5).  In  this  case,  peat  and  very  fine 
roots  (smaller  than  0.5  cm  diameter)  ex- 
ceeded remaining  biomass  by  5:1.  Lugo  et 
al.  (1976)  reported  the  following  values 
for  a  south  Florida  red  mangrove  overwash 
forest.  All  values  were  reported  in  dry 
grams  per  square  meter,  plus  and  minus  one 
standard  error,  and  ignoring  belowground 
biomass.  They  found  710  -  22  q/ml  of 
leaves,    12. 8„  -    15.3   g/mz   of   propagules, 


iles, 
f  of 


7043  i  7  g/mz  of  wood,    4695  ±  711    g/rr 
prop  roots  and   1565  -  234.5  g/mz  of  detri- 
tus  on   the   forest    floor. 

Biomass  partitioning  between  dif- 
ferent species  and  locations  must  be 
highly  variable.  The  age  of  the  forest 
will    influence  the  amount  of  wood  biomass; 


15 


(A! 


SOIL 
SURFACE 


LEAVES 
(778) 


BRANCHES  (  1 274) 


PROP  ROOTS  (  1437) 


TRUNK    (2796) 


LARGE 
ROOTS    (997) 


SMALL     ROOTS    (4000) 


(B) 


HEIGHT 
(M) 


o 


5000  10000 

FOOT  CANDLES 


Figure  5.  (a)  Aboveground  and  belowground  blomass  of  a  Puerto  R1can  red  mangrove 
forest.  Values  in  parentheses  are  dry  g/m2;  large  roots  =  2  cm+  in  diameter, 
small  roots  =  0.5  -  1.0  cm.  (b)  Vertical  distribution  of  light  Intensity  1n  the 
same  forest;  canopy  height  is  8  m  (26  ft)  (both  figures  adapted  from  Golley  et  al. 
1962). 


16 


detritus  varies  enormously  from  one  site 
to  the  next  depending  upon  the  amount  of 
fluvial  transport.  The  biomass  charac- 
teristics of  a  scrub  forest  probably  bear 
little  resemblance  to  those  of  a  fringing 
forest.  At  the  present  time,  there  is  not 
enough  of  this  type  of  data  available  to 
draw  many  conclusions.  One  intriguing 
point  is  that  red  mangrove  leaf  biomass 
averages  between  700  and  800  g/m  at 
various  sites  with  very  different  forest 
morphologies  (Odum  and  Heald  1975a).  This 
may  be  related  to  the  tendency  of  mangrove 
canopies,  once  they  have  become  estab- 
lished, to  inhibit  leaf  production  at 
lower  levels   through   self-shading. 

Golley  et  al.  (1962)  showed  that  the 
red  mangrove  canopy  is  an  extremely  effi- 
cient light  interceptor.  Ninety— five 
percent  of  the  available  light  had  been 
intercepted  4  m  (13  ft)  below  the  top  of 
the  canopy  (Figure  5).  As  a  result,  90% 
of  the  leaf  biomass  existed  in  the  upper  4 
m  of  the  canopy.  Chlorophyll  followed  the 
same  pattern   of  distribution. 

The  leaf  area  index  (LAI)  of  mangrove 
forests  tends  to  be  relatively  low.  Gol- 
ley et  al.  (1962)  found  a  LAI  of  4.4  for  a 
Puerto  Rican  red  mangrove  forest.  Lugo  et 
al.  (1975)  reported  a  LAI  of  5.1  for  a 
Florida  black  mangrove  forest  and  3.5  for 
a  Florida  fringe  red  mangrove  forest.  A 
different  black  mangrove  forest,  in  Flori- 
da, was  found  to  have  values  ranging  from 
1  to  4  and  an  average  of  2  to  2.5  (Lugo 
and  Zucca  1977).  These  values  compare 
with  LAI's  of  10  to  20  recorded  for  most 
tropical  forests  (Golley  et  al.  1974). 
The  low  leaf  area  values  of  mangrove 
forests  can  be  attributed  to  at  least 
three  factors:  (1)  effective  light  inter- 
ception by  the  mangrove  canopy,  (2)  the 
inability  of  the  lower  mangrove  leaves  to 
flourish  at  low  light  intensities,  and  (3) 
the  absence  of  a  low-light-adapted  plant 
layer  on   the   forest   floor. 


2.5     PRIMARY   PRODUCTION 

Prior  to  1970  virtually  no  informa- 
tion existed  concerning  the  productivity 


of  mangroves  in  Florida.  Since  that  time 
knowledge  has  accumulated  rapidly,  but  it 
is  still  unrealistic  to  expect  more  than 
preliminary  statements  about  Florida  man- 
grove productivity.  This  deficiency  can 
be  traced  to  (1)  the  difficulties  asso- 
ciated with  measurements  of  mangrove  pro- 
ductivity and  (2)  the  variety  of  factors 
that  affect  productivity  and  the  resulting 
variations  that  exist  from  site  to   site. 

Productivity  estimates  come  from 
three  methods:  (1)  harvest,  (2)  gas  ex- 
change, and  (3)  litter  fall.  Harvest 
methods  require  extensive  manpower  and 
knowledge  of  the  age  of  the  forest.  They 
are  best  employed  in  combination  with 
silviculture  practices.  Since  silvicul- 
ture of  south  Florida  mangroves  is  practi- 
cally non-existent,  this  method  has  rarely 
been  used  in  Florida.  Noakes  (1955), 
Macnae  (1968),  and  Walsh  (1974)  should  be 
consulted  for  productivity  estimates  based 
on  this  technique  in  other  parts  of  the 
world. 

Gas  exchange  methods,  based  on 
measurements  of  CO?  changes,  have  the 
advantage  of  precision  and  response  to 
short-term  changes  in  light,  temperature, 
and  flooding.  They  include  both  above- 
ground  and  belowground  production.  On  the 
negative  side,  the  necessary  equipment  is 
expensive  and  tricky  to  operate  properly. 
Moreover,  extrapolations  from  short-term 
measurements  to  long-term  estimates  offer 
considerable  opportunity  for  error. 
Nevertheless,  the  best  estimates  of  pro- 
ductivity come  from  this  method. 

The  litter  fall  technique  (annual 
litter  fall  x  3  =  annual  net  primary  pro- 
duction) was  proposed  by  Teas  (1979)  and 
is  based  on  earlier  papers  by  Bray  and 
Gorham  (1964)  and  Golley  (1972)  for  other 
types  of  forests.  This  is  a  quick  and 
dirty  method  although  the  lack  of  pre- 
cision remains  to  be  demonstrated  for 
mangroves.  An  even  quicker  and  dirtier 
method  proposed  by  Teas  (1979)  is  to  (1 ) 
estimate  leaf  standing  crop  (using  various 
techniques  including  harvesting  or  light 
transmission  relationships)  and  (2)  multi- 
ply by  three.     This  assumes  an  annual    leaf 


17 


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

Mangrove  productivity  is  affected  by 
many  factors;  some  of  these  have  been 
recognized  and  some  remain  totally  ob- 
scure. Carter  et  al.  (1973)  propose 
lumping  these  factors  into  two  broad  cate- 
gories: tidal  and  water  chemistry.  We 
believe  that  a  number  of  additional  cate- 
gories  should   be   considered. 

A  minimal,  though  incomplete,  list  of 
factors  controlling  mangrove  productivity 
must   include  the  following: 

*  species  composition  of  the  stand 

*  age  of  the  stand 

*  presence     or     absence     of       competing 
species 

*  degree  of  herbivory 

*  presence     or  absence  of     disease     and 
parasites 

*  depth  of  substrate 

*  substrate  type 

*  nutrient  content  of  substrate 

*  nutrient  content  of  overlying     water 

*  salinity  of  soil    and  overlying  water 

*  transport  efficiency  of  oxygen  to   root 
system 

*  amount   of  tidal    flushing 

*  relative  wave  energy 

'  presence  or  absence  of  nesting  birds 

*  periodicity  of  severe  stress     (hurri- 
canes,  fire,   etc.) 

*  time  since  last   severe  stress 

*  characteristics  of  ground     water 


inputs  of  toxic  compounds  or  nutrients 
from  human  activities 

human       influences     such     as       diking, 
ditching,   and  altering  patterns  of 
runoff. 


In  spite  of  the  difficulties  with 
various  methods  and  the  interaction  of 
controlling  factors,  it  is  possible  to 
make  general  statements  about  certain 
aspects  of  mangrove  productivity.  For 
example,  Waisel's  (1972)  statement  that 
mangroves  have  low  transpiration  rates 
seems  to  be  generally  true  in  Florida. 
Lugo  et  al.  (1975)  reported  transpiration 
rates  of  2,500  g  HoO/m  /day  for  mangrove 
leaves  in  a  fringing  red  mangrove  forest 
and  1,482  g  h^O/m  /day  for  black  mangrove 
leaves.  This  is  approximately  one-third 
to  one-half  the  value  found  in  temperate 
broad  leaf  forests  on  hot  dry  days,  but 
comparable  to  tropical  rainforests  (H.T. 
Odum  and  Jordan  1970).  The  low  transpira- 
tion rates  of  mangroves  are  probably  re- 
lated to  the  energetic  costs  of  main- 
taining sap  pressures  of  -35  to  -60  atmo- 
spheres   (Scholander   et    al.    1965). 

Litter  fall  (leaves,  twigs,  bark, 
fruit,  and  flowers)  of  Florida  mangrove 
forests  appears  to  average  2  to  3  dry 
g/m  day  in  most  wel 1 -devel oped  mangrove 
stands  (see  discussion  in  section  3.4). 
This  can  be  an  order  of  magnitude  lower  in 
scrub  forests. 

Wood  production  of  mangroves  appears 
to  be  high  compared  to  other  temperate  and 
tropical  trees,  although  no  measurements 
from  Florida  are  available.  Noakes  (1955) 
estimated  that  the  wood  production  of  an 
intensively  managed  Malayan  forest  was 
39.7  metric  tons/ha/year.  Teas  (1979) 
suggested  a  wood  production  estimate  of  21 
metric  tons/ha/year  for  a  mature  unmanaged 
red  mangrove  forest  in  south  Florida.  His 
figure  was  calculated  from  a  litter/total 
biomass  relationship  and  is  certainly 
subject    to   error. 

Representative  estimates  of  gross 
primary    production    (GPP)    net    primary 


18 


production  (NPP),  and  respiration  (R)  of 
Florida  mangroves  are  given  in  Table  la. 
Compared  to  net  primary  production  (NPP) 
estimates  from  other  ecosystems,  including 
agricultural  systems  (E.P.  Odum  1971),  it 
appears  that  mangroves  are  among  the 
world's  most  productive  ecosystems. 
Healthy  mangrove  ecosystems  appear  to  be 
more  productive  than  sea  grass,  marsh 
grass  and  most  other  coastal    systems. 

Further  examination  of  Table  la  re- 
veals several  possible  tendencies.  The 
first  hypothetical  tendency,  as  discussed 
by  Lugo  et  al.  (1975),  is  for  red  mangroves 
to  have  the  highest  total  net  production, 
black  to  have  intermediate  values  and 
white  the  lowest.  This  conclusion  assumes 
that  the  plants  occur  within  the  zone  for 
which  they  are  best  adapted  (see  section 
3.2  for  discussion  of  zonation)  and  are 
not  existing  in  an  area  with  strong  limit- 
ing factors.  A  scrub  red  mangrove  forest, 
for  example,  growing  under  stressed  condi- 
tions (high  soil  salinity  or  low  nutrient 
supply),  has  relatively  low  net  produc- 
tivity (Teas  1979).  The  pre-eminent  posi- 
tion of  red  mangroves  is  shown  by  the 
comparative  measurements  of  photosynthesis 
in  Table  lb;  measurements  were  made  within 
canopy  leaves  of  trees  growing  within 
their  zones  of  optimal    growth. 

A  second  noteworthy  tendency  is  that 
red  mangrove  GPP  decreases  with  increasing 
salinity  while  GPP  of  black  and  white 
mangroves  increases  with  increasing 
salinity  up  to  a  point.  Estimates  of  Hicks 
and  Burns  (1975)  demonstrate  that  this  may 
be  a   real    tendency    (Table    lc). 

Data  presented  by  Miller  (1972), 
Carter  et  al.  (1973),  Lugo  and  Snedaker 
(1974),  and  Hicks  and  Burns  (1975)  sug- 
gest a  third  hypothetical  tendency, 
assuming  occurrence  of  the  species  within 
its  adapted  zone.  It  appears  that  the 
black  mangrove  typically  has  a  much  higher 
respiration  rate,  lower  net  productivity, 
and  lower  GPP/R  ratio  than  the  red  man- 
grove. This  can  be  attributed  at  least 
partially,  to  the  greater  salinity  stress 
under  which  the  black  mangrove  usually 
grows;   this   leads  to  more  osmotic  work. 


These  three  apparent  tendencies  have 
led  Carter  et  al.  (1973)  and  Lugo  et  al. 
(1976)  to  propose  a  fourth  tendency,  an 
inverted  U-shaped  relationship  between 
waterway  position  and  net  mangrove  com- 
munity productivity  (Figure  6).  This 
tendency  is  best  understood  by  visualizing 
a  typical  gradient  on  the  southwest  coast 
of  Florida.  At  the  landward  end  of  the 
gradient,  salinities  are  very  low, 
nutrient  runoff  from  terrestrial  eco- 
systems may  be  high  and  tidal  amplitude  is 
minor.  At  the  seaward  end,  salinities  are 
relatively  high,  tidal  amplitude  is  rela- 
tively great  and  nutrient  concentrations 
tend  to  be  lower.  At  either  end  of  the 
gradient,  the  energetic  costs  are  high  and 
a  large  percentage  of  GPP  is  used  for 
self-maintenance;  at  the  landward  end, 
competition  from  freshwater  plant  species 
is  high  and  at  the  seaward  end,  salinity 
stress  may  be  limiting.  In  this  scenario, 
the  highest  NPP  occurs  in  the  middle 
region  of  the  gradient;  salinity  and  tidal 
amplitude  are  high  enough  to  limit  compe- 
tition while  tidal  flushing  and  moderate 
nutrient  levels  enhance  productivity. 
Hicks  and  Burns  (1975)  present  data  to 
support   this    hypothesis. 


In  addition  to  these  hypotheses 
generated  from  field  data,  there  have  been 
two  significant,  published  attempts  to 
derive  hypotheses  from  mathematical  simu- 
lation models  of  mangroves.  The  first 
(Miller  1972)  is  a  model  of  primary  pro- 
duction and  transpiration  of  red  mangrove 
canopies  and  is  based  upon  equations  which 
utilize  field  measurements  of  the  energy 
budgets  of  individual  leaves.  This  model 
predicts  a  variety  of  interesting  trends 
which  need  to  be  further  field  tested. 
One  interesting  hypothesis  generated  by 
the  model  is  that  maximum  photosynthesis 
of  red  mangrove  stands  should  occur  with  a 
leaf  area  index  (LAI)  of  2.5  if  no  accli- 
mation to  shade  within  the  canopy  occurs; 
higher  LAI's  may  lead  to  decreased  produc- 
tion. Another  prediction  is  that  red 
mangrove  production  is  most  affected  by 
air  temperature  and  humidity  and,  to  a 
lesser    degree,    by    the    amount    of    solar 


19 


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20 


Table  lb.   Comparative  measurements  of  photosynthesis  in 
gC/m2/day  (Lugo  et  al .  1975)  . 


Mangrove  type 


Daytime  net 
photosynthesi  s 


Ni  ghttime 
respi  ra ti  on 


Pn/R 


Red 

Bl  ack 

Whi  te 

Red  ( seedl i  ng  ) 


1  .38 
1  .24 
0.58 
0.31 


0.23 
0.53 
0.17 
1  .89 


6.0 
2.3 
3.4 
negative 


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


Mangrove  type 


Average  surface 
sal i  ni  ty  (ppt ) 


GPP 
(gC/m2/day) 


Red 
Red 
Red 
Bl  ack 
Bl  ack 
Black 
White 
White 


7.8 
21  .1 
26.6 

7.8 
21  .1 
26.6 
21  .1 
26.6 


8.0 
3.9 
1  .6 
2.3 
5.7 
7.5 
2.2 
4.8 


21 


HIGH 


COMMUNITY 
NET  PRIMARY 
PRODUCTION 


LOW 


LANDWARD 


SEAWARD 


WATERWAY  POSITION 


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


22 


radiation  within  the  ambient  range.  Gross 
photosynthesis  per  unit  leaf  area  was 
greater  at  the  top  of  the  tree  canopy  than 
at  the  bottom,  although  the  middle  levels 
had   the   greatest   production. 

Miller  (1972)  concluded  by  suggesting 
that  the  canopy  distribution  of  red  man- 
grove leaves  is  nearly  optimal  for  ef- 
ficient water  utilization  rather  than 
production.  This  indicates  that  the  cano- 
py is  adapted  to  maximizing  production 
under  conditions  of  saturated  water  sup- 
ply. 

The  mangrove  ecosystem  model  reported 
by  Lugo  et  al.  (1976)  provides  hypotheses 
on  succession,  time  to  arrive  at  steady 
state  conditions  (see  section  3.2),  and 
several  aspects  of  productivity.  The 
model  output  suggests  that  the  relative 
amount  of  tidal  amplitude  does  not  affect 
GPP  significantly;  instead,  GPP  appears  to 
be  extremely  sensitive  to  inputs  of  ter- 
restrial nutrients.  It  follows  that  loca- 
tions with  large  amounts  of  nutrient  input 
from  terrestrial  sources  (riverine  man- 
grove communities)  have  high  rates  of 
mangrove  production  (see  section  3.3). 
All  simulation  model -generated  hypotheses 
need  to  be  field  tested  with  a  particular- 
ly critical  eye,  since  the  simplifying 
assumptions  that  are  made  in  constructing 
the  model  can  lead  to  overly  simplistic 
answers. 

Mangrove  productivity  research  re- 
mains in  an  embryonic  stage.  Certain 
preliminary  tendencies  or  hypotheses  have 
been  identified,  but  much  work  must  be 
done  before  we  can  conclude  that  these 
hypotheses   cannot   be   falsified. 


2.6      HERBIVORY 

Direct  herbivory  of  mangrove  leaves, 
leaf  buds,  and  propagules  is  moderately 
low,  but  highly  variable  from  one  site  to 
the  next.  Identified  grazers  of  living 
plant  parts  (other  than  wood)  include  the 
white-tailed  deer,  Odocoileus  virginianus, 
the  mangrove  tree  crab,  Aratus  pi  soni  i , 
and  insects  including  beetles,  larvae  of 


lepidopterans    (moths   and  butterflies),    and 
orthopterans    (grasshoppers    and    crickets). 

Heald  (1969)  estimated  a  mean  grazing 
effect  on  North  River  red  mangrove  leaves 
of  5.1%  of  the  total  leaf  area;  values 
from  leaf  to  leaf  were  highly  variable 
ranging  from  0  to  18%.  Beever  et  al. 
(1979)  presented  a  detailed  study  of 
grazing  by  the  mangrove  tree  crab.  This 
arboreal  grapsid  crab  feeds  on  numerous 
items  including  beetles,  crickets,  cater- 
pillars, littoral  algae,  and  dead  animal 
matter.  In  Florida,  red  mangrove  leaves 
form  an  important  component  of  the  diet. 
Beever  et  al.  (1979)  measured  tree  crab 
grazing  ranging  from  0.4%  of  the  total 
leaf  area  for  a  Florida  Keys  overwash 
forest  to  7.1%  for  a  fringing  forest  at 
Pine  Island,  Lee  County,  Florida.  The 
researchers  also  found  that  tree  crab 
grazing  rates  are  related  to  crab  density. 
Low  densities  (one  crab/m  )  resulted  in 
low  leaf  area  damage  (less  than  1%  of 
total  leaf  area).  High  densities  (four 
crabs/m  )  were  accompanied  by  leaf  area 
damage  ranging  from  4%  to  6%  (see  section 
6.2). 

Onuf  et  al.  (1977)  investigated  in- 
sect herbivory  in  fringing  and  overwash 
red  mangrove  forests  in  the  Indian  River 
estuary  near  Ft.  Pierce,  Florida.  They 
found  six  major  herbivorous  insect 
species,  five  lepidopteran  larvae  and  a 
beetle.  Comparisons  were  made  at  a  high 
nutrient  site  (input  from  a  bird  rookery) 
and  a  low  nutrient  site.  Both  red  man- 
grove production  and  leaf  nitrogen  were 
significantly  higher  at  the  high  nutrient 
site.  This  resulted  in  a  four-fold 
greater  loss  to  herbivores  (26%  of  total 
leaf  area  lost  to  grazing);  this  increased 
grazing  rate  more  than  offset  the  in- 
creased leaf  production  due  to  nutrient 
input. 

Calculations  of  leaf  area  damage  may 
underestimate  the  impact  of  herbivores  on 
mangroves.  For  example,  the  larvae  of  the 
olethreutid  moth,  Ecdytol opha  sp., 
develops  within  red  mangrove  leaf  buds  and 
causes  the  loss  of  entire  leaves.  All 
stages     of     the     beetle,      Poeci 1 i  ps 


23 


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


2.7  WOOD  BORERS 

Many  people  have  the  mistaken  idea 
that  mangrove  wood  is  highly  resistant  to 
marine  borers.  While  this  may  be  true  to 
a  limited  extent  for  certain  mangrove 
species  in  other  parts  of  the  world,  none 
of  the  Florida  mangroves  have  borer- 
resistant  wood.  Southwell  and  Boltman 
(1971)  found  that  the  wood  of  red,  black, 
and  white  mangroves  has  no  resistance  to 
Teredo,  Pholad  and  Simnorid  borers;  pieces 
of  red  mangrove  wood  were  completely  de- 
stroyed after  immersion  in  ocean  water  for 
14   months. 

An  interesting  controversy  surrounds 
the  ability  of  the  wood  boring  isopod, 
Sphaeroma  terebrans,  to  burrow  into  the 
living  prop  roots  of  the  red  mangrove. 
Rehm  and  Humm  (1973)  were  the  first  to 
attribute  apparently  extensive  damage  of 
red  mangroves  stands  within  the  Ten 
Thousand  Islands  area  of  southwestern 
Florida  to  an  isopod,  Sphaeroma.  They 
found  extensive  damage  throughout 
southwest  Florida,  some  infestation  north 
to  Tarpon  Springs,  and  a  total  lack  of 
infestation  in  the  Florida  Keys  from  Key 
Largo  south  to  Key  West.  The  destruction 
process  was  described  as  follows:  the 
adult  isopod  bored  into  the  prop  roots  (5- 
mm  diameter  hole);  this  was  followed  by 
reproduction  within  the  hole  and  develop- 
ment of  juveniles  within  the  root.  This 
process,  combined  with  secondary  decompo- 
sition from  fungi  and  bacteria,  frequently 
results  in  prop  root  severance  near  the 
mean  high  tide  mark.  These  authors 
attributed  loss  of  numerous  prop  roots 
and,  in  some  cases,  loss  of  entire  trees 
during  storms  to  isopod  damage. 

The  extent  of  damage  in  the  Ten 
Thousand  Islands  region  led  Rehm  and  Humm 
(1973)  to  term  the  phenomenon  an  "eco- 
catastrophe" of  possibly  great  importance. 
They  further  stated  that  shrinking  of 
mangrove  areas  appeared  to  be  occurring  as 


a   result  of  Sphaeroma   infestation;    this 
point  was  not  documented. 

Enright  (1974)  produced  a  tongue-in- 
cheek  rebuttal,  on  behalf  of  Sphaeroma  and 
against  the  "terrestrial  invader",  red 
mangroves.  Snedaker  (1974)  contributed  a 
more  substantial  argument  in  which  he 
pointed  out  that  the  isopod  infestation 
might  be  an  example  of  a  long-term  eco- 
system  control    process. 

Further  arguments  against  the  "ecoca- 
tastrophe" theory  were  advanced  by  Estevez 
and  Simon  (1975)  and  Estevez  (1978).  They 
provided  more  life  history  information  for 
Sphaeroma  and  suggested  a  possible  ex- 
planation for  the  apparently  destructive 
isopod  infestations.  They  found  two 
species  of  isopods  inhabiting  red  mangrove 
prop  roots,  S^.  terebrans  and  a  sympatric 
congener,  S.  quadridentatum.  The  latter 
does  not  appear  to  be  a  wood  borer  but 
utilizes  S.  terebrans  burrows.  Neither 
species  appeared  to  utilize  mangrove  wood 
as  a  food  source.  Estevez  and  Simon 
(1975)  found  extensive  burrowing  into 
seedlings  in  addition  to  prop  root  damage. 
In  general,  infestations  appeared  to  be 
patchy  and  limited  to  the  periphery  of 
mangrove  ecosystems.  In  areas  with  the 
highest  density  of  burrows,  23%  of  all 
prop  roots  were  infested.  There  appeared 
to  be  more  colonization  by  ^.  terebrans  in 
regions  with  full  strength  sea  water  (30 
to  35  ppt). 

The  most  important  finding  by  Estevez 
and  Simon  (1975)  and  Estevez  (1978)  was 
that  periods  of  accelerated  activity  by  ^. 
terebrans  were  related  to  periods  of  fluc- 
tuating and  slightly  increased  salinity. 
This  suggests  that  fluctuations  in  isopod 
burrowing  may  be  related  to  the  magnitude 
of  freshwater  runoff  from  the  Everglades. 
These  authors  agree  with  Snedaker  (1974) 
and  suggest  that  root  and  tree  loss  due  to 
Sphaeroma  activity  may  be  beneficial  to 
mangrove  ecosystems  by  accelerating  pro- 
duction and  root  germination.  Simberloff 
et  al. (1978)  amplified  this  last  sugges- 
tion by  showing  that  root  branching,  which 
is  beneficial  to  individual  trees,  is 
stimulated   by   isopod  activity. 


24 


This  ecocatastrophe  versus  beneficial 
stimulus  argument  is  not  completely  re- 
solved. Probably,  Sphaeroma  root  destruc- 
tion, in  areas  of  low  isopod  density,  can 
be  a  beneficial  process  to  both  the  in- 
dividual tree  and  to  the  entire  mangrove 
stand.  Whether  changes  in  freshwater 
runoff  have  accelerated  this  process  to 
the  point  where  unnatural  and  widespread 
damage  is  occurring  is  not  clear.  The 
data  and  research  perspective  to  answer 
this  question  do  not  exist.  As  a  result, 
we  are  reduced  to  providing  hypotheses 
which  cannot  be  tested  with  available 
knowledge. 


2.8     MANGROVE   DISEASES 

Published  research  on  mangrove 
diseases  is  rare.  The  short  paper  by 
Olexa  and  Freeman  (1975)  is  the  principal 
reference  for  diseases  of  Florida  man- 
groves. They  reported  that  black  man- 
groves   are    affected    by    the    pathogenic 


fungi ,  Phyl losti  eta  hi  bi  scina  and  Nigro- 
spora  sphaerica.  These  authors  found  that 
P.  hi  bi  sci  na  caused  necrotic  lesions  and 
death  of  black  mangrove  leaves.  They  felt 
that  under  conditions  of  high  relative 
humidity  coupled  with  high  temperatures, 
this  fungus  could  pose  a  serious  threat  to 
individual  trees,  particularly  if  the  tree 
had  been  weakened  by  some  other  natural 
agent,  such  as  lightning  or  wind  damage. 
Nigrospora  sphaerica  was  considered  to  be 
of  little  danger  to  black  mangroves. 
Another  fungus,  Cyl i nrocarpon  didymum, 
appears  to  form  galls  on  the  prop  roots 
and  stems  of  red  mangroves.  Olexa  and 
Freeman  (1975)  noted  mortality  of  red 
mangroves  in  areas  of  high  gall  infesta- 
tions, although  a  direct  causation  link 
was   not   proven. 

Further  research  on  mangrove  diseases 
is  badly  needed.  Viral  disease  must  be 
investigated.  The  role  of  pathogens  in 
litter  production  and  as  indicators  of 
mangrove  stress   may  be  very  important. 


25 


CHAPTER    3.       ECOSYSTEM    STRUCTURE    AND   FUNCTION 


3.1       STRUCTURAL    PROPERTIES    OF    MANGROVE 
FORESTS 

Published  information  about  the 
structural  aspects  of  Florida  mangrove 
forests  is  limited;  most  existing  data 
have  been  published  since  the  mid-1970's. 
This  lack  of  information  is  unfortunate 
since  quantitative  structural  data  greatly 
aid  understanding  of  processes  such  as 
succession  and  primary  production.  Even 
more  important,  the  response  of  mangrove 
forests  to  stress,  both  climatic  and  man- 
induced,  can  be  followed  quantitatively 
with  this  type   of  data. 

Ball  (1980)  contributed  substantially 
to  understanding  the  role  of  competi- 
tion in  mangrove  succession  by  measuring 
structural  factors  such  as  basal  area, 
tree  height,  and  tree  density.  Lugo  and 
Zucca  (1977)  monitored  the  response  of 
mangrove  forests  to  freezing  temperatures 
by  observing  changes  in  structural  proper- 
ties  of  the  trees. 

Baseline  studies  of  forest  structure 
have  been  published  by  Lugo  and  Snedaker 
(1975),  and  Pool,  Snedaker  and  Lugo 
(1977).  For  example,  Lugo  and  Snedaker 
(1975)  compared  a  fringing  mangrove  forest 
and  a  basin  forest  at  Rookery  Bay,  near 
Naples,  Florida.  They  found  the  fringing 
forest,  which  was  dominated  by  red  man- 
groves, to  have  a  tree  diversity  of  H  = 
1.48,  a  basal  area  of  15.9  mz/ha,  an 
aboveground  biomass  of  17,932  g/m  ,  and  a 
non-existent  litter  layer.  The  nearby 
basin  forest  was  dominated  by  black  man- 
groves, had  a  tree  diversity  of  H  =  0.96 
and  a  basal  area  of  23.4  m  /ha.  The  lit- 
ter layer  in  the  basin  forest  averaged  550 
dry  g/m  .  Tree  diversity  in  a  hurricane 
disturbed  section  of  the  Rookery  Bay 
forest  was  1.62.  Similar  data  were  pre- 
sented for  mangrove  forests  in  the  Ten 
Thousand    Islands    area    (Table   2). 

Data  of  this  type  are  useful  for  many 
purposes  including  impact  statements,  en- 
vironmental surveys,  and  basic  scientific 
questions.  Cintron  et  al.  (1978)  gave  an 
indication  of  the  direction  in  which  fu- 
ture research  might  proceed.  Working  in  a 
mangrove   stand   in   Puerto  Rico,    they   found 


tree  height  to  be  inversely  proportional 
(r  =  0.72)  to  soil  salinity  in  the  range 
30  to  72  ppt.  Above  65  ppt  salinity,  dead 
tree  basal  area  was  higher  than  live  tree 
basal  area  and  above  90  ppt  there  was  no 
live  tree   basal    area. 

It  should  be  possible  to  investigate 
the  relationship  between  a  variety  of 
mangrove  structural  properties  and  factors 
such  as  flushing  frequency,  soil  depth, 
nutrient  availability,  pollution  stress, 
and  other  measures  of  human  impact.  Ulti- 
mately, this  should  lead  to  an  ability  to 
predict  the  form  and  structure  of  mangrove 
forests  resulting  from  various  physical 
conditions  or  artificial  impacts.  One 
example  of  this  potential  tool  is  Ball's 
(1980)  documentation  of  structural  changes 
in  mangrove  forests  resulting  from  altera- 
tions in  the  hydrological  conditions  of 
south  Florida. 


3.2       Z0NATI0N,     SUCCESSION    AND    "LAND- 
BUILDING" 

Much  of  the  world's  mangrove  litera- 
ture consists  of  descriptive  accounts  of 
zonation  in  mangrove  forests  and  the  spe- 
cies composition  within  these  zones.  Al- 
thouqh  general  agreement  has  been  lacking, 
various  hypotheses  have  been  put  forth 
concerning  the  possible  connection  between 
zonation,  ecological  succession,  competi- 
tion, and  the  role  of  physical  factors 
such  as  soil  salinity  and  tidal  amplitude. 
In  this  section  we  review  briefly  the 
dominant  ideas  about  mangrove  zonation  and 
succession  and  present  our  interpretation 
of  the  current   status   of  knowledge. 

Davis  (1940),  working  in  south  Flori- 
da, was  one  of  the  first  investigators  to 
describe  distinct,  almost  monospecific, 
zones  within  mangrove  ecosystems.  In  what 
has  become  the  classical  view,  he  argued 
that  mangrove  zonation  patterns  were 
equivalent  to  serai  stages  in  succession. 
The  most  seaward  zone,  dominated  by  red 
mangroves,  was  regarded  as  the  "pioneer 
stage".  More  landward  zones  were 
dominated  by  white  mangrove,  black 
mangrove,  buttonwood  and,  finally,  the 
climatic  climax,   a  tropical    forest.     Since 

26 


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27 


these  zones  were  regarded  as  progressively 
later  stages  in  succession,  the  entire 
mangrove  ecosystem  was  believed  to  be 
moving  seaward  through  a  process  of  sedi- 
ment accumulation  and  colonization.  Davis 
based  his  argument  primarily  upon  the 
sequence  of  observed  zones  and  cores  which 
showed  red  mangrove  peat  underlying  black 
mangrove  peat  which,  in  turn,  occurred 
under  terestrial    plant  communities. 

Unfortunately,  this  Clementsian  in- 
terpretation of  mangrove  zonation  was 
widely  accepted,  but  rarely  tested.  For 
example,  Chapman  (1970)  expanded  Davis' 
original  successional  concept  from  south 
Florida  to  explain  zonation  in  mangrove 
forests  in  other  parts  of  the  world. 
Walsh  (1974)  thoroughly  reviewed  the  man- 
grove  succession/zonation    literature. 

Fortunately,  not  everyone  accepted 
Davis'  point  of  view.  Egler  (1952)  and 
later  Thorn  (1967,  1975)  argued  that  man- 
grove zonation  was  a  response  to  external 
physical  forces  rather  than  temporal  se- 
quence induced  by  the  plants  themselves. 
Egler  (1952)  showed  that  patterns  of  sedi- 
ment deposition  predicted  by  Davis'  (1940) 
theory  did  not  always  occur.  He  also 
showed  that  in  some  cases  mangrove  zones 
appeared  to  be  moving  landward  rather  than 
seaward.  Sea  level  has  been  rising  in 
south  Florida  at  the  rate  of  1  ft  (30  cm) 
per  100  to  150  years  (Provost  1974). 
Spackman  et  al.  (1966)  emphasized  the  role 
of  sea  level  change  in  determining  changes 
in  mangrove  zonation,  both  through  sea 
level  rise  and  land  subsidence.  Both 
Egler  (1952)  and  Spackman  et  al.  (1966) 
along  with  Wanless  (1974)  and  Thorn  (1967, 
1975)  suggested  that  mangroves  were 
reacting  passively  rather  than  actively  to 
strong  geomorphologi cal  processes.  This 
implies  that  mangroves  should  be  regarded 
as  "land-stabilizers"  rather  than  "land- 
builders". 

Furthermore,  field  researchers  fre- 
quently noted  that  red  mangroves  were  not 
always  the  only  "pioneer  species"  on  re- 
cently deposited  sediment.  It  is  not 
unusual  to  find  seedlings  of  black,  white, 
and  red  mangroves  growing  together  on  a 
new   colonization    site.      Lewis   and   Dunstan 


(1975)  found  that  black  mangroves  and 
white  mangroves  along  with  the  saltmeadow 
cordgrass,  Spartina  patens,  are  often  the 
pioneers  on  new  dredge  spoil  islands  in 
central  Florida.  On  the  northern  coast  of 
the  Gulf  of  Mexico,  where  black  mangrove 
is  the  only  mangrove  species  present,  it 
may  be  preceded  by  marsh  grasses  such  as 
saltmarsh  cordgrass,  S.  patens,  smooth 
cordgrass,  S.  al terni  fl ora,  or  the  black 
needle  rush,  Juncus  roemerianus.  In  Puer- 
to Rico,  we  observed  that  white  mangrove 
often  pioneers  and  dominates  sites  where 
oceanic  overwash  of  beach  sand  has  oc- 
curred. All  of  these  observations  detract 
from  Davis'  (1940)  original  contention 
that  red  mangroves  should  be  regarded  as 
the  initial  colonizer  of  recently  de- 
posited sediments.  It  appears  that  under 
certain  conditions,  e.g.,  shallow  water 
depths,  substrate  type,  and  latitude, 
white  and  black  mangroves  or  marsh  grasses 
can   be  effective   pioneer   species. 

The  work  of  Rabinowitz  (1975)  added  a 
new  perspective  to  the  mangrove  zonation 
debate.  Through  carefully  designed  recip- 
rocal planting  experiments  in  Panamanian 
mangrove  forests  using  species  of  Rhi  zo- 
phora ,  Laguncul a  ri  a ,  P e 1 1 i  c i  e r a  and 
Avicennia,  she  demonstrated  that  each 
species  could  grow  well  within  any  of  the 
mangrove  zones.  In  other  words,  physical 
and  chemical  factors  such  as  soil  salinity 
or  frequency  of  tidal  inundation,  within 
each  zone,  were  not  solely  responsible  for 
excluding  species  from  that  zone.  To 
explain  zonation,  Rabinowitz  proposed 
tidal  sorting  of  propagules  based  upon 
propagule  size,  rather  than  habitat  adap- 
tation,as  the  most  important  mechanism  for 
zonation  control. 

The  most  recent  piece  to  be  added  to 
the  zonation/succession  puzzle  comes  from 
the  work  of  Ball  (1980).  Based  upon  re- 
search of  mangrove  secondary  succession 
patterns  adjacent  to  Biscayne  Bay,  Flori- 
da, she  made  a  strong  case  for  the  impor- 
tance of  interspecific  competition  in 
controlling  zonation.  She  found  that 
white  mangroves,  which  grow  best  in 
intertidal  areas,  do  not  occur  consis- 
tently in  the  intertidal  zone  of  mature 
mangrove  stands.     Instead,  white  mangroves 


28 


dominate  higher,  drier  locations  above 
mean  high  water  where  the  red  mangrove 
does  not  appear  to  have  a  competitive 
advantage.  She  suggested  that  competition 
is  not  so  important  during  the  early 
stages  of  succession  but  becomes  critical 
as  individual  trees  reach  maturity  and 
require  more  space  and   other   resources. 

Inherent  in  Ball's  concept  of  zona- 
tion  is  the  differential  influence  of 
physical  factors  (e.g.,  soil  salinity, 
depth  to  water  table)  on  the  competitive 
abilities  of  the  different  mangrove 
species.  She  concluded  that  succession 
proceeds  independently  within  each  zone, 
although  breaks  in  the  forest  canopy  from 
lightning  strikes  or  high  winds  may  pro- 
duce a  mosaic  of  different  successional 
stages  within  a  zone.  These  openings 
allow  species  whose  seedlings  do  not  com- 
pete well  in  shade,  such  as  the  white 
mangrove,  to  become  established,  at  least 
temporarily,  within  solid  zones  of  red 
mangroves. 

Zonation  of  mangrove  species  does  not 
appear  to  be  controlled  by  physical  and 
chemical  factors  directly,  but  by  the 
interplay  of  these  factors  with  interspe- 
cific competition  and,  possibly,  through 
tidal  sorting  of  propagules.  Once  succes- 
sion in  a  mangrove  zone  reaches  an  equili- 
brium state,  change  is  unlikely  unless  an 
external  perturbation  occurs.  These  per- 
turbations range  from  small-scale  distur- 
bance (lightning  strikes)  to  large-scale 
perturbations  (sea  level  change,  hurricane 
damage)  and  may  cause  succession  within 
zones  to  regress  to  an  earlier  stage. 
There  is  some  evidence  in  south  Florida 
that  hurricane  perturbations  occur  on  a 
fairly  regular  basis,  creating  a  pattern 
of   cyclical    succession. 

Except  for  Ball  (1980)  and  Taylor 
(1980),  the  importance  of  fires  as  an 
influence  on  mangrove  succession  has  been 
generally  ignored.  Most  fires  in  the 
Florida  mangrove  zone  are  initiated  by 
lightning  and  consist  of  small  circular 
openings  in  the  mangrove  canopy  (Taylor 
1980).  These  openings  present  an  opportu- 
nity for  secondary  succession  within  an 
established  zone.     For  example,   we  have 


frequently  observed  white  mangroves 
flourishing  in  small  lightning-created 
openings  in  the  center  of  red  mangrove 
forests.  Fire  may  also  play  a  role  in 
limiting  the  inland  spread  of  mangroves. 
Taylor  (1981)  pointed  out  that  Everglades 
fires  appear  to  prevent  the  encroachment 
of  red  and  white  mangroves  into  adjacent 
herbaceous  communities. 

Finally,  Lugo  and  Snedaker  (1974), 
Cintron  et  al.  (1978)  and  Lugo  (1980) 
suggested  that  mangrove  ecosystems 
function  as  classical  successional  systems 
in  areas  of  rapid  sediment  deposition  or 
upon  recently  colonized  sites  such  as 
offshore  islands.  They  concluded  that  in 
most  areas  mangrove  forests  are  an  example 
of  steady-state  cyclical  systems.  Concep- 
tually, this  is  synonymous  to  E.  P.  Odum's 
(1971)  cyclic  or  catastrophic  climax. 
Chapman  (1976a,  b)  suggested  the  idea  of 
cyclic  succession  for  a  variety  of  coastal 
ecosystems. 

If  Florida  mangrove  ecosystems  are 
cyclic  systems,  then  there  should  be  an 
identifiable  perturbation  capable  of  set- 
ting succession  back  to  an  early  stage. 
Lugo  and  Snedaker  (1974)  suggested  that 
hurricanes  may  play  this  role.  They 
pointed  out  (without  substantiating  data) 
that  major  hurricanes  occur  about  every 
20-25  years  in  south  Florida.  Coinci- 
dently,  mangrove  ecosystems  appear  to 
reach  their  maximum  levels  of  productivity 
in  about  the  same  period  of  time  (Lugo  and 
Snedaker  1974).  This  hypothesis  suggests 
that  succession  within  many  mangrove  eco- 
systems may  proceed  on  a  cyclical  basis 
rather  than  in  the  classical  fashion. 
Possibly  other  physical  perturbations  may 
influence  mangrove  succession  including 
incursions  of  freezing  temperatures  into 
central  Florida,  periodic  droughts  causing 
unusually  high  soil  salinities  (Cintron  et 
al.  1978),  and  fire  spreading  into  the 
upper  zones  of  mangrove  forests  from  ter- 
restrial  sources. 

Although  understanding  of  zonation 
and  succession  in  mangrove  ecosystems 
remains  incomplete,  a  clearer  picture  is 
emerging,  at  least  for  south  Florida. 
Contrary  to  early  suggestions,  mangrove 


29 


species  zonation  does  not  appear  to  repre- 
sent serai  stages  of  succession  except, 
perhaps,  for  locations  of  recent  coloniza- 
tion or  where  sediment  is  accumulating 
rapidly.  The  role  of  mangroves  in 
land-bui'lding     seems  more  passive  than 

active.  Geomorphological  and  hydrological 
processes  appear  to  be  the  dominant  forces 
in  determining  whether  mangrove  shorelines 
recede  or  grow.  The  role  of  mangroves  is 
to  stabilize  sediments  which  have  been 
deposited   by   physical    processes. 


3.3      NUTRIENT    CYCLING 

Current  understanding  of  nutrient 
cycles  in  mangrove  ecosystems  is  far  from 
satisfactory.  Sporadic  field  measurements 
have  been  made,  but  a  complete  nutrient 
budget  has  not  been  published  for  any 
mangrove  ecosystem  in  the  world. 

Several  pioneering  field  studies  were 
conducted  in  Florida  (Carter  et  al.  1973; 
Snedaker  and  Lugo  1973;  Onuf  et  al.  1977) 
and  one  simulation  model  of  mangrove  nu- 
trient cycling  has  been  published  (Lugo  et 
al.  1976).  Preliminary  measurements  of 
nitrogen  fixation  were  made  (Zuberer  and 
Silver  1975;  Gotto  and  Taylor  1976; 
Zuberer  and  Silver  1978;  Gotto  et  al. 
1981).  Based  on  these  studies,  we  present 
the  following  preliminary  conclusions. 

Mangrove  ecosystems  tend  to  act  as  a 
sink  (net  accumulator)  for  various  ele- 
ments including  macro  nutrients  such  as 
nitrogen  and  phosphorus,  trace  elements, 
and  heavy  metals.  As  we  have  discussed  in 
section  1.7,  these  elements  are  removed 
from  waters  flowing  through  mangrove 
swamps  by  the  concerted  action  of  the 
mangrove  prop  roots,  prop  root  algae,  the 
associated  sediments,  the  fine  root  system 
of  the  mangrove  trees,  and  the  host  of 
small  invertebrates  and  microorganisms 
attached  to  all  of  these  surfaces.  Al- 
though the  turnover  times  for  these  ele- 
ments in  mangrove  swamps  are  not  known,  it 
appears  that  at  least  a  portion  may  be 
stored  or  tied  up  in  wood,  sediments,  and 
peat    for   many  years. 


Although  mangrove  ecosystems  may  tend 
to  accumulate  nutrients,  there  is  a  con- 
tinual loss  through  export  of  particulate 
and  dissolved  substances.  If  significant 
nutrient  storage  and  resultant  high  pri- 
mary production  are  to  occur,  there  must 
be  a  continual  input  of  nutrients  to  the 
mangrove  forest  from  outside  the  system 
(Figure  7).  Where  nutrient  influx  to  the 
mangrove  ecosystem  is  approximately 
balanced  by  nutrient  loss  in  exported 
organic  matter,  then  nutrient  storage  will 
be  minimal  and  mangrove  net  primary  pro- 
duction will  be  low.  This  appears  to 
occur  in  the  scrub  mangrove  community  type 
and  to  a  lesser  extent  in  the  basin  and 
hammock  community  types. 

Carter  et  al.  (1973)  and  Snedaker  and 
Lugo  (1973)  have  hypothesized  that  the 
greatest  natural  nutrient  inputs  for  man- 
grove swamps  come  from  upland  and  terres- 
trial sources.  Apparently  for  this  rea- 
son, the  most  luxuriant  and  productive 
mangrove  forests  in  south  Florida  occur  in 
riverine  locations  or  adjacent  to  signifi- 
cant   upland    drainage. 

Localized  sources  of  nutrients,  such 
as  bird  rookeries,  can  result  in  greater 
nutrient  storage  and  higher  mangrove  pro- 
ductivity (Onuf  et  al.  1977).  If  however, 
large  bird  rookeries  (or  artificial  nu- 
trient inputs)  occur  in  poorly  flushed 
sections  of  mangrove  ecosystems,  resultant 
high  nutrient  levels  may  inhibit  mangrove 
growth  (R.  R.  Lewis,  III,  Hillsborough 
Community  College,  Tampa,  Fla.;  personal 
communication    1981). 

The  output  from  the  simulation  model 
of  Lugo  et  al.  (1976)  suggests  that  if 
nutrient  input  to  a  mangrove  ecosystem  is 
reduced,  then  nutrient  storage  levels 
within  the  mangrove  ecosystem  will  be 
reduced  and  mangrove  biomass  and  produc- 
tivity will  decline.  To  our  knowledge 
this  hypothesis  has  not  been  tested  in  the 
field. 

Nitrogen  fixation  occurs  in  mangrove 
swamps  at  rates  comparable  to  those 
measured  in  other  shallow,  tropical  marine 
areas    (Gotto    et    al.    1981).      Nitrogen 


30 


SMALL 
IMPORT 


LOW    STORAGE 
LOW   BIOMASS 
LOW   PRODUCTIVITY 


SMALL 
EXPORT 


LARGE 
IMPORT 


MODERATE 
EXPORT 


HIGH   STORAGE 

HIGH   BIOMASS 

HIGH    PRODUCTIVITY 

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


31 


fixation  has  been  found  in  association 
with  mangrove  leaves,  both  living  and 
dead,  mangrove  sediment  surfaces,  the 
litter  layer  in  mangrove  swamps,  and  man- 
grove root  systems  (Gotto  and  Taylor  1976; 
Zuberer  and  Silver  1978;  Gotto  et  al. 
1981).  In  virtually  all  cases,  nitrogen 
fixation  appears  to  be  limited  by  the 
availability  of  labile  carbon  compounds. 
Perhaps  for  this  reason,  the  highest  rates 
of  mangrove  nitrogen  fixation  have  been 
measured  in  association  with  decaying 
mangrove  leaves;  presumably,  the  decaying 
leaves  act  as  a  carbon  source  and  thus 
accelerate  nitrogen  fixation.  Macko 
(1981),  using  stable  nitrogen  ratio 
techniques,  has  indicated  that  as  much  as 
25%  of  the  nitrogen  associated  with  black 
mangrove  peat  in  Texas  is  derived  from 
nitrogen  fixation. 

Zuberer  and  Silver  (1978)  speculated 
that  the  nitrogen  fixation  rates  observed 
in  Florida  mangrove  swamps  may  be  suf- 
ficient to  supply  a  significant  portion  of 
the  mangrove's  growth  requirements.  Al- 
though this  hypothesis  is  impossible  to 
test  with  present  information,  it  might 
explain  why  moderately  productive  mangrove 
stands  occur  in  waters  which  are  severely 
nitrogen  depleted. 

In  summary,  knowledge  of  nutrient 
cycling  in  mangrove  swamps  is  highly 
speculative.  These  ecosystems  appear  to 
act  as  a  sink  for  many  elements,  including 
nitrogen  and  phosphorus,  as  long  as  a 
modest  input  occurs.  Nitrogen  fixation 
within  the  swamp  may  provide  much  of  the 
nitrogen  needed   for  mangrove  growth. 


3.4      LITTER    FALL   AND   DECOMPOSITION 

Unless  otherwise  stated,  litter  fall 
refers  to  leaves,  wood  (twigs),  leaf 
scales,  propagules,  bracts,  flowers,  and 
insect  frass  (excrement)  which  fall  from 
the  tree.  Mangrove  leaves  are  shed  con- 
tinuously throughout  the  year  although  a 
minor  peak  occurs  during  the  early  part  of 
the  summer  wet  season  in  Florida  (Heald 
1969;  Pool  et  al.  1975).  Sporadic  litter 
fall  peaks  may  follow  periods  of  stress 
from    cold    air    temperatures,    high    soil 


salinities,  and  pollution  events.  Litter 
fall  typically  can  be  partitioned  as  68% 
to  86%  leaves,  3%  to  15%  twigs  and  8%  to 
21%  miscellaneous;  the  latter  includes 
flowers  and   propagules. 

Litter  fall  is  an  important  ecosystem 
process  because  it  forms  the  energy  basis 
for  detritus-based  foodwebs  in  mangrove 
swamps  (see  sections  3.5  and  3.6).  The 
first  measurements  of  litter  fall  in  man- 
grove swamps  were  made  by  E.J.  Heald  and 
W.E.  Odum,  working  in  the  North  River 
estuary  in  south  Florida  in  1966-69. 
This  was  subsequently  published  as  Heald 
(1969),  Odum  (1970),  and  Odum  and  Heald 
(1975a).  They  estimated  that  litter  pro- 
duction from  riverine  red  mangrove  forests 
averaged  2.4  dry  g  of  organic 
matter/m  /day  (or  876  g/m  /year  or  8.8 
metric  tons/ha/year). 

Subsequent  studies  agreed  with  this 
early  estimate  (Table  3),  although  varia- 
tion clearly  exists  between  different 
types  of  communities.  Scrub  forests  with 
scattered,  very  small  trees  have  the 
smallest  amount  of  leaf  fall.  Basin  and 
hammock  forests,  which  appear  to  be 
nutrient  limited,  have  intermediate  leaf 
fall  values.  Not  surprisingly,  the 
highest  values  occur  in  the  highly  produc- 
tive fringing,  overwash,  and  riverine 
forests.  Odum  and  Heald  (1975a)  suggested 
that  the  relatively  uniform  litter  fall 
values  from  productive  mangrove  forests 
around  the  world  result  from  the  shade 
intolerance  of  the  canopy  leaves  and  the 
tendency  for  the  canopy  size  to  remain  the 
same  in  spite  of  increasing  height.  If 
detailed  information  is  lacking,  red  man- 
grove forests  of  south  Florida,  which  are 
not  severely  limited  by  lack  of  nutrients, 
can  be  assumed  to  produce  litter  fall  of 
2.0  to  3.0  g/m  /day  of  dry  organic  matter. 
Pure  stands  of  black  mangroves  usually 
have  a  lower  rate  of  1.0  to  1.5  g/nr/day 
(Lugo  et  al.  1980). 

Decomposition  of  fallen  Florida  man- 
grove leaves  has  been  investigated  by  a 
number  of  researchers  including  Heald 
(1969),  Odum  (1970),  Odum  and  Heald 
(1975a),  Pool  et  al.  (1975),  Lugo  and 
Snedaker   (1975),  Twilley  (1980)   and   Lugo  et 


32 


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al.  (1980).  Heald  and  Odum  showed  that 
decomposition  of  red  mangrove  leaves 
proceeds  most  rapidly  under  marine  condi- 
tions, somewhat  more  slowly  in  freshwater, 
and  very  slowly  on  dry  substrates.  For 
example,  using  the  litter  bag  method,  they 
found  that  only  9%  of  the  original  dry 
weight  remained  after  4  months  in  sea 
water.  By  comparison,  39%  and  54%  re- 
mained at  the  end  of  comparable  periods  in 
brackish  water  and  freshwater.  Under  dry 
conditions,  65%  remained.  Higher  decompo- 
sition rates  in  sea  water  were  related  to 
increased  activity  of  shredder  organisms, 
such   as   crabs  and  amphipods. 

Heald  (1969)  and  Odum  (1970)  also 
found  increases  in  nitrogen,  protein,  and 
caloric  content  as  mangrove  leaves  pro- 
gressively decayed.  The  nitrogen  content 
of  leaves  decaying  under  brackish  condi- 
tions (on  an  AFDW  basis)  increased  from 
1.5%  (5.6%  protein)  to  3.3%  (20.6% 
protein)  over  a  6-month  period.  Subse- 
quent information  (Odum  et  al.  1979b) 
suggested  that  the  protein  increase  may 
not  have  been  this  great  since  some  of  the 
nitrogen  increase  probably  included  non- 
protein nitrogen  compounds  such  as  amino 
sugars.  Fell  and  Master  (1973),  Fell  et 
al.  (1980),  Fell  and  Newell  (1980),  and 
Fell  et  al.  (1980)  have  provided  more 
detailed  information  on  red  mangrove  leaf 
decomposition,  the  role  of  fungi  in  decom- 
position (see  section  4),  and  nitrogen 
changes  and  nitrogen  immobilization  during 
decomposition.  Fell  et  al .  (1980) 
have  shown  that  as  much  as  50%  of  weight 
loss  of  the  leaf  during  decomposition  is 
in  the  form  of  dissolved  organic  matter 
(DOM). 

Heald  et  al.  (1979),  Lugo  et  al. 
(1980)  and  Twilley  (1980)  discovered  that 
black  mangrove  leaves  decompose  more  ra- 
pidly than  red  mangrove  leaves  and  ap- 
parently produce  a  higher  percentage  of 
DOM.  Pool  et  al.  (1975)  have  shown  that 
mangrove  litter  decomposes  and  is  exported 
most  rapidly  from  frequently  flooded 
riverine  and  overwash  forests.  These 
communities  have  little  accumulation  of 
litter  on  the  forest  floor.  Communities 
which  are  not  as  well-flushed  by  the 
tides,     such    as    the    basin    and    hammock 


forests,    have    slower    rates    of    decomposi- 
tion and  lower  export    rates. 


3.5     CARBON    EXPORT 

Research  from  Florida  mangrove  swamps 
forms  a  small  portion  of  the  larger  con- 
troversy concerned  with  the  extent  to 
which  coastal  wetlands  export  particulate 
organic  carbon  (reviewed  by  Odum  et  al. 
1979a).  Available  evidence  from  Florida, 
Puerto  Rico  and  Australia  (Table  4)  sug- 
gests that  mangrove  swamps  tend  to  be  net 
exporters.  The  values  in  Table  4  should 
be  regarded  as  preliminary,  however,  since 
all  five  studies  are  based  upon  simplistic 
assumptions  and  methodology. 

Golley  et  al.  (1962)  based  their 
annual  estimate  of  particulate  carbon 
export  from  a  Puerto  Rican  forest  upon  a 
few  weeks  of  measurements.  Odum  and 
Heald's  estimates  were  derived  from  two  or 
three  measurements  a  month.  All  investi- 
gators have  ignored  the  importance  of  bed 
load  transport  and  the  impact  of  extreme 
events.  All  investigators  except  Lugo  et 
al.  (1980)  have  failed  to  measure  DOC 
flux. 

It  seems  relatively  clear  that  man- 
grove forests  do  export  organic  carbon  to 
nearby  bodies  of  water.  The  magnitude  of 
this  export  has  probably  been  underesti- 
mated due  to  ignoring  bedload,  extreme 
events,    and    DOC. 

The  value  of  this  carbon  input  to 
secondary  consumers  in  receiving  waters  is 
not  clear.  As  shown  in  section  3.6,  food 
webs  based  primarily  upon  mangrove  carbon 
do  exist.  The  relative  importance  of 
mangrove  carbon  to  Florida  coastal  ecosys- 
tems remains  speculative.  We  suspect  that 
mangrove-based  food  webs  are  dominant  in 
small  bays,  creeks  and  rivers  within  large 
mangrove  ecosystems  such  as  the  North 
River  system  studied  by  Heald  (1969)  and 
Odum  (1970).  In  intermediate-sized  bodies 
of  water,  such  as  Rookery  Bay  near  Naples, 
Florida,  mangroves  are  probably  important 
but  not  dominant  sources  of  organic  car- 
bon. Lugo  et  al.  (1980)  estimate  that 
mangroves   supply  32%  of  the  organic  carbon 


34 


Table  4.  Estimates  of  particulate  carbon  export  from  mangrove 
forests.  Lugo  et  al .  (1976)  estimated  export  from  a  theoreti- 
cal, steady  state  forest  using  a  simulation  model.  Lugo  et  al . 
(1980)  measured  export  from  an  inland  black  mangrove  forest. 


Investigators 


Location 


Export 


g/m  /day     tonnes/ha/yr 


Golley  et  al .  (1962) 
Heald  (1969),  Odum  (1970)' 
Lugo  and  Snedaker  (1975) 
Lugo  et  al .   (1976) 
Boto  and  Bunt  (1981) 
Lugo  et  al.  (1980)b 


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


Puerto  Rico 

1.1 

4.0 

Florida 

0.7 

2.5 

Florida 

0.5 

2.0 

Florida 

1.5-1.8 

5.5  -  6.6 

Australia 

1.1 

4.0 

Florida 

0.2 

0.7 

35 


input  to  Rookery  Bay.  In  very  large  sys- 
tems, such  as  Biscayne  Bay  near  Miami, 
Florida,  mangroves  are  clearly  less  impor- 
tant than  any  other  sources  such  as  algae 
and  sea  grasses,  although  mangrove  carbon 
may  be  important  in  localized  situations 
such  as  the  immediate  vicinity  of  fringing 
and  overwash  forests.  The  magnitude  of 
mangrove  carbon  export  to  unenclosed 
coastal  waters  and  offshore  remains  a 
mystery. 


3.6     ENERGY   FLOW 

At  least  seven  sources  of  organic 
carbon  may  serve  as  energy  inputs  for 
consumers  in  mangrove  ecosystems  (Figure 
8).  The  pathways  by  which  this  energy 
containing  material  is  processed  and  made 
available  to  each  consumer  species  is 
indeed  complex.  Not  surprisingly,  current 
understanding  of  energy  flow  in  Florida 
mangrove  ecosystems  exists  largely  in  a 
qualitative  sense;  quantitative  data  are 
scarce  and  piecemeal.  A  variety  of  inves- 
tigators have  contributed  information  over 
the  past  decade  including,  but  not  limited 
to,  Heald  (1969),  Odum  (1970),  Odum  and 
Heald  (1972),  Carter  et  al.  (1973), 
Snedaker  and  Lugo  (1973),  Heald  et  al. 
(1974),  Lugo  and  Snedaker  (1974,  1975), 
Odum  and  Heald  (1975a,  b),  and  Pool  et  al. 
(1977).  Probably,  the  most  complete  study 
to  date  is  the  investigation  of  energy 
flow  in  the  black  mangrove  zone  of  Rookery 
Bay  by  Lugo  et  al.  (1980). 

It  is  possible  at  this  time  to  pre- 
sent a  series  of  hypotheses  concerning  the 
relative  importance  of  these  energy 
sources.  First,  the  relative  importance 
of  each  source  can  vary  from  one  location 
to  the  next.  As  will  be  shown  in  the 
following  discussion,  the  consumers  in 
certain  mangrove  forests  appear  to  depend 
primarily  upon  mangrove-derived  carbon 
while  in  other  locations  inputs  from  phy- 
toplankton  and  attached  algae  are  probably 
more  important. 

Our  second  hypothesis  is  that  energy 
flow  based  upon  phytopl ankton  is  most 
important  in  overwash  mangrove  forests  and 
other    locations    associated    with    large 


bodies  of  clear,  relatively  deep  water. 
Conversely,  phytoplankton  are  hypothesized 
to  be  relatively  unimportant  to  the  energy 
budgets  of  the  large  riverine  forest  com- 
munities along  the  southwest  coast  of 
Florida.  It  should  be  remembered,  how- 
ever, that  even  where  phytoplankton  are 
quantitatively  unimportant,  they  poten- 
tially perform  an  important  function  as 
the  basis  of  phytopl ankton-zoopl ankton- 
larval    fish   food   webs   (Odum  1970). 

As  a  third  hypothesis,  Iver  Brook 
(Rosensteil  School  of  Marine  and  Atmos- 
pheric Sciences,  Rickenbacker  Causeway, 
Miami,  Fla.;  personal  communication  1979) 
has  suggested  that  both  sea  grasses  and 
benthic  algae  serve  as  an  important  energy 
source  for  fringing  mangrove  communities 
adjacent  to  large  bodies  of  water  such  as 
Biscayne  Bay  and  Whitewater  Bay.  Although 
little  evidence  exists  to  test  this  hypo- 
thesis, observations  of  extensive  deposits 
of  sea  grass  and  macroalgal  detritus  with- 
in mangrove  forests  suggest  intuitively 
that    Brook's   hypothesis   may  be   correct. 

In  regions  where  mangrove  shading  of 
the  prop  roots  is  not  severe,  our  fourth 
hypothesis  suggests  that  carbon  origina- 
ting from  prop  root  epiphytes  may  be  sig- 
nificant to  community  energy  budgets. 
Lugo  et  al.  (1975)  have  measured  net  pro- 
duction of  periphyton  in  mangroves 
fringing  Rookery  Ba^  and  found  average 
values  of  1.1  gC/m  /day.  Hoffman  and 
Dawes  (1980)  found  a  lower  value  of  0.14 
gC/mVday.  Because  these  values  are 
roughly  comparable  to  average  exports  of 
mangrove  leaf  carbon  (section  3.5),  its 
potential    importance   is  obvious. 

The  fifth  hypothesis  states  that 
mangrove  organic  matter,  particularly  leaf 
material,  is  an  important  energy  source 
for  aquatic  consumers.  This  hypothesis 
was  first  espoused  by  Heald  (1969)  and 
Odum  (1970),  who  worked  together  in  the 
riverine  mangrove  communities  between  the 
Everglades  and  Whitewater  Bay.  Clearly, 
mangrove  carbon  is  of  great  importance 
within  the  riverine  and  basin  communities 
all  along  the  southwest  coast  of  Florida 
(Odum  and  Heald  1975b);  Carter  et  al . 
(1973)    and    Snedaker    and    Lugo    (1973) 


36 


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37 


provided  subsequent  supportive  data.  What 
is  not  clear,  is  the  relative  importance 
of  mangrove  carbon  to  consumers  within 
fringing,  overwash,  and  more  isolated 
manqrove  communities. 

Our  sixth  hypothesis  involves  the 
assemblage  of  organisms  that  graze  man- 
grove leaves  directly.  A  variety  of  in- 
sects (see  section  6)  and  the  mangrove 
tree  crab,  Aratus  pi  soni  i ,  (Beever  et  al. 
1979)  obtain  much  of  their  energy  directly 
from  living  mangrove  leaves,  even  though 
grazing  rarely  exceeds  10%  of  net  primary 
production    (Odum   and   Heald   1975b). 

As  a  seventh  hypothesis  we  suggest 
that  anaerobic  decomposition  of  mangrove 
tissue,  particularly  root  material,  may 
support  an  extensive  food  web  based  on 
bacteria  associated  with  methanogenesis  or 
the  processing  of  reduced  sulfur  com- 
pounds. Our  suggestion  of  the  importance 
of  reduced  sulfur  comes  directly  from 
Howarth  and  Teal's  (1980)  discovery  of 
this  potentially  important  energy  pathway 
in  temperate  Spartina  (cordgrass)  marshes. 
They  found  that  anaerobic  decomposition  is 
such  an  incomplete  process  that  if  sul- 
fates are  available  (from  sea  water)  as 
much  as  75%  of  the  original  energy  in 
plant  tissues  may  be  converted  by  sulfur 
reducing  bacteria  to  reduced  sulfur  com- 
pounds such  as  hydrogen  sulfide  and  py- 
rite.  Subsequently,  if  these  reduced 
sulfur  compounds  are  moved  hydrologically 
to  an  oxidized  environment  (sediment  sur- 
face or  creek  bank)  sulfur -oxidizing  bac- 
teria (e.g.,  Thiobacillus  spp.)  may  convert 
the  chemically  stored  energy  to  bacterial - 
ly  stored  energy  with  an  efficiency  as 
great  as  50%  (Payne  1970).  Presumably, 
deposit-feeding  organisms  such  as  grass 
shrimp  (Pal  aemonetes)  and  mullet  (Mugil ) 
are  capable  of  grazing  these  sulfur- 
oxidizing  bacteria  from  the  sediment 
surface.  If  this  hypothetical  trophic 
exchange  does  exist,  it  may  be  of  con- 
siderable magnitude  and  may  cause  us  to 
reexamine  current  concepts  of  energy  pro- 
cessing and  export  from  mangrove 
ecosystems.  Since  freshwater  contains 
remarkably  little  sulfate  in  comparison  to 
seawater,  this  energy  pathway  is  probably 
of  little  importance   in   mangrove   forests 


of  very  low  salinity. 

Carbon  inputs  from  terrestrial 
sources  may  be  important  to  certain  man- 
grove communities.  Carter  et  al.  (1973) 
have  shown  that  terrestrial  carbon  can 
reach  coastal  ecosystems  particularly 
where  man  has  cut  deep  channels  inland  for 
navigation  or  drainage  purposes.  The 
magnitude  of  this  influx  has  not  been 
adequately  measured  although  Carter  et  al. 
did  find  that  mainland  forests  (including 
mangroves)  contributed  approximately  2,100 
metric  tons  of  carbon  per  year  to 
Fahkahatchee  Bay. 

Atmospheric  inputs  from  rainfall 
appear  to  be  minimal  in  all  cases.  Lugo 
et  al.  (1980)  measured  throughfall  (preci- 
pitation passing  through  the  tree  canopy) 
in  Rookery  Bay  mangrove  forests  of  15  to 
17  gC/m  /year.  This  would  be  an  overesti- 
mate of  atmospheric  input  since  it  con- 
tains carbon  leached  from  mangrove  leaves. 
The  best  guess  of  atmospheric  input  is 
between  3  to  5  gC/m  /year  for  south 
Florida  mangrove  ecosystems. 

Subsequent  stages  of  energy  transfer 
in  mangrove  community  food  webs  remain 
largely  hypothetical.  Odum  (1970)  and 
Odum  and  Heald  (1975b)  have  outlined 
several  pathways  whereby  mangrove  carbon 
and  energy  are  processed  by  a  variety  of 
organisms  (see  Figure  8).  Apparently,  the 
most  important  pathway  follows  the  se- 
quence: mangrove-leaf  detritus  substrate- 
microbe-detritus  consumer-higher  consu- 
mers. The  critical  links  are  provided  by 
the  microbes  such  as  bacteria  and  fungi 
(see  Fell  et  al.  1975)  and  by  the  detritus 
consumers.  The  latter  group  was  studied 
by  Odum  (1970)  and  Odum  and  Heald  (1975b) 
and  found  to  consist  of  a  variety  of 
invertebrates  (e.g.,  caridean  shrimp, 
crabs,  mollusks,  insect  larvae,  amphipods) 
and  a    few    fishes. 

Stable  carbon  studies  such  as  those 
done  by  Haines  (1976)  in  Spartina 
(cordgrass)  marshes  have  not  been  per- 
formed in  mangrove  ecosystems.  Mangroves 
are  C3  plants  and  have  613  values  in  the 
range  of  minus  25  to  minus  26  (Macko 
1981).       According    to    the    same    author, 


38 


mangrove  peat  has  a  613  value  of  minus 
22.  Because  these  values  are  dramatically 
different  from  the  values  for  sea  grasses 
and  many  algae,  the  possibilities  for 
using  this  tool  in  mangrove  ecosystems  is 
excellent.  Macko  (1981)  also  suggested 
the  utility  of  using  stable  nitrogen  ra- 
tios for  future  mangrove  food  web  investi- 
gations; he  reported  615  values  of  plus 
6.0  to  plus  6.5  for  mangrove  tissue  and 
plus   5   for  mangrove  peat. 

In  reviewing  contemporary  knowledge 
of  energy  flow  in  mangrove  ecosystems, 
three  conclusions   emerge. 

(1)  We  have  a  hypothetical  framework 
of  mangrove  energy  flow  of  a  qualitative 


nature.  This  framework  appears  to  be 
reasonably  accurate  although  subsequent 
developments,  such  as  elucidation  of  the 
reduced  sulfur  hypothesis,  may  require 
some  modification. 

(2)  Measurements  of  the  relative 
importance  of  various  carbon  sources  are 
generally  lacking. 

(3)  Detailed  measurements  of  energy 
flow  including  the  relative  inputs  of 
different  carbon  sources  are  critically 
needed.  Technological  difficulties,  high 
costs,  and  difficulties  inherent  in 
transferring  findings  from  one  estuary  to 
the  next  present  a  major  challenge  to 
estuarine  ecologists  of  the  future. 


39 


CHAPTER  4.   COMMUNITY  COMPONENTS  -  MICROORGANISMS 

The   mycoflora    (fungi)    are   the   best  Drechslera    and   Gloeosporium.      In   the    lat- 

studied  component  of  the  microbial    com-  ter  stages  of  decay  the  dominant   genera 

munity    of   mangrove    swamps.      Much    pio-  are  Calso,   Gliocidium,    and  Lulworthia. 
neering  work  has  been  carried  out   in   south 

Florida.     Reviews  of  the  current   knowledge  Understanding  the  occurrence  and  suc- 

of  mangrove-associated   fungi    can    be   found  cession    of    fungi    on    decaying    mangrove 

in  Kohlmeyer  and  Kohlmeyer   (1979)  and  Fell  leaves   is   important   because  of   their    role 

et  al .    (1980).  in   energy   flow   in   mangrove  swamps.     Heald 

(1969),    Odum    (1970)    and    Odum    and    Heald 

One  of  the  earliest  studies  of  man-  (1975b)  hypothesized  that  fungi   and  bac- 

grove  mycoflora  was  published  by  Kohlmeyer  teria  are  important  in  converting  mangrove 

(1969).     He  discovered  large  populations  leaf  organic   material    into  a  form  that  can 

of  marine   fungi    on   the   submerged   parts   of  be  digested  and  assimilated  by  detriti- 

aerial    roots,  stems,  and  branches  and  on  vores  (see  section  3.6). 
living  and  dead  mangrove  leaves.     Exten- 
sive work  at  the  University  of  Miami   by 

Fell  and  his  coworkers  (e.g.,  Fell  and  Our  understanding  of  the  role  and 
Master  1973;  Fell  et  al.  1975,  1980)  ex-  occurrence  of  bacteria  in  mangrove  swamps 
plored  the  role  of  fungi  in  the  decom-  is  not  as  well  documented  as  for  fungi, 
position  of  mangrove  leaves  and  the  im-  Casagrande  and  Given  (1975)  have  suggested 
mobilization  of  nitrogen.  Newell  (1974)  that  bacteria  are  important  in  the  early 
studied  the  succession  of  mycoflora  on  stages  of  mangrove  leaf  decomposition  and 
seedlings  of  red  mangrove.  A  survey  of  are  replaced  in  the  latter  stages  by  funqi 
the  aquatic  yeasts  occurring  in  the  south  which  are  better  equipped  to  attack  re- 
Florida  mangrove  zone  was  published  by  fractive  organic  compounds.  Unlike  the 
Ahearn  et  al.  (1968).  mycoflora,  the  bacteria  are  clearly  impor- 
tant   in   the   anaerobic   regions   of   mangrove 

One  of  the  most   interesting  pieces  of  swamps.  Vankatesan  and  Ramamurthy  (unpubl . 

information  to  emerge   from  this  extensive  data)   found     denitrifying     bacteria     to  be 

mycoflora   research  concerns  the  succession  abundant  and  ubiquitous  in  mangrove  soils, 

of    orqanisms    associated    with    decaying  Zuberer  and  Silver  (1978)   have     emphasized 

leaves    (summarized   by  Fell    et   al.    1975,  the  importance  of  nitrogen-fixing  bacteria 

1980).      Senescent    leaves    of    red   mangroves  in  the  zone  around  mangrove  roots.       They, 

are    typically    colonized    by    species    of  in  fact,     were  able  to  isolate  and  count  a 

Nigrospora,  Phyllostica,    and   Pestalotica.  variety  of  types  of  bacteria  from  mangrove 

Once  the  leaf  has   fallen   from  the  tree  and  sediments  including  aerobic  heterotrophs, 

during    the    early    stages    of    decay,  the  anaerobic   heterotrophs,   nitrogen-fixing 

fungal    flora   is  dominated  by  species  of  heterotrophs,  and  sul  fate-reducing  bac- 

Phytophthora    and,    to    a    lesser    extent,  teria. 


40 


CHAPTER    5.       COMMUNITY    COMPONENTS 


PLANTS    OTHER    THAN    MANGROVES 


5.1.      ROOT  AND   MUD   ALGAE 

The  aerial  root  systems  of  mangroves 
provide  a  convenient  substrate  for  at- 
tachment of  algae.  These  root  algal  com- 
munities are  particularly  noticeable  on 
red  mangrove  prop  roots  but  also  occur  to 
a  lesser  extent  on  black  mangrove 
pneumatophores  located  in  the  intertidal 
zone.  Productivity  of  prop  root  algal 
communities  can  be  appreciable  if  shading 
by  mangroves  is  not  too  severe;  as  dis- 
cussed in  section  3.6,  Lugo  et  al.  (1975) 
found  a  prop  root  community  net  primary 
production  rate  of  1.1  gC/m  /day,  a  level 
comparable  to  mangrove  leaf  fall.  Biomass 
of  these  algae  can  be  as  high  as  200  to 
300  g  per  prop  root  (Burkholder  and 
Almodovar  1973).  Of  course,  production  of 
this  magnitude  only  occurs  on  the  edge  of 
the  forest  and  is  virtually  nil  in  the 
center  of  the  swamp.  Nevertheless,  this 
algal  carbon  has  considerable  potential 
food  value  either  to  direct  grazers  or 
detriti vores. 

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

Zonation  to  be  expected  on  Florida 
mangroves  is  shown  in  Figure  9;  this  se- 
quence comes  largely  from  Taylor  (1960). 
Near  the  high  water  mark,  a  green  band 
usually  exists  which  is  dominated  by  spe- 
cies of  Rhi  zocl oni  urn.  Below  this  is  a 
zone  dominated  by  species  of  Bostrychia, 
Catenel 1  a,  and  Cal ogl ossa.  It  is  this 
association  that  most  people  think  of  when 
mangrove  prop  root  algae  are  mentioned. 


Because  much  mud  is  often  deposited  on  the 
Bostrychia-Catenell  a-Cal  ogl  ossa  complex, 
it  often  has  a  dingy,  gray  appearance. 
There  are  many  other  algae  found  in  this 
zone,  but  these  three  genera  usually  domi- 
nate. At  brackish  or  nearly  freshwater 
locations,  they  are  replaced  by  species  of 
Batophora,  Chaetomorpha ,  CI adophora,  and 
P e n i  c i 1 1 u s .  The  pneumatophores  of 
Avicennia,  when  colonized,  are  often 
covered  with  species  of  Rhi  zocl oni  urn, 
Bostrychia  and  Monostroma  (Taylor  1960). 
Hoffman  and  Dawes  (1980)  found  that  the 
Bostrychia  binderi -dominated  community  on 
the  pneumatophores  of  black  mangroyes  had 
-tanding  crop  of  22  g  dry  wt/m  and  a 
production    of  0.14   gC/mz/day. 


a  s 

net 


If  there  is  a  permanently  submerged 
portion  of  the  prop  root,  it  may  be 
covered  with  rich  growths  of  Acanthophora, 
Spyri  da,  Hypnea,  Laurencia,  Wrangel i  a, 
Valonia,  and  Caulerpa  (Almodovar  and  Biebl 
1962).  Additional  genera  which  may  be 
present  below  mean  high  water  are: 
Murrayel 1  a,  Polysi  phoni  a,  Centroceras, 
Wurdemannia ,  D  i  c tyota ,  Hal i  m  ed  a , 
Lau  renci  a ,  and  Da  sya  [Taylor  1960; 
Burkholder  and  Almodovar  1973;  Yoshioka 
1975).  In  addition,  anywhere  on  the  moist 
sections  of  the  prop  roots  there  are 
usually  epiphytic  diatoms  and  filamentous 
green  and  blue-green  algae  of  many   genera. 

Rehm  (1974)  found  a  significant  dif- 
ference in  the  prop  root  algae  between 
south  and  central  Florida.  South  of  Tampa 
Bay  the  standard  Bostrychia-Catenel 1 a- 
Caloglossa  dominates.  In  the  Tampa  Bay 
area,  species  of  the  orders  Ulotrichales 
and  Cladophorales   are  dominant. 

The  mud  adjacent  to  the  mangrove  root 
community  is  often  richly  populated  with  a 
variety  of  algae.  These  can  include 
species  of  CI  adophoropsi  s,  Enteromorpha, 
Vaucheria,  and  Boodleopsis  (Taylor  I960) 
in  addition  to  a  whole  host  of  benthic 
diatoms  and  dinofl agel  1  ates  (Wood  1965) 
and  other  filamentous  green  and  blue-green 
algae    (Marathe    1965). 

Adjacent  to  mangrove  areas,  on  the 
bottoms  of  shoals,  shallow  bays  and 
creeks,     there      is     often    a    variety    of 


41 


PLANTS 


ANIMALS 


Rhizi 

MHW- 


Bostryt 
Catenei 

Caloglt 


MLW-- 

Acanthopi 

Caulerpa 

Wranglela 


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


42 


tropical  algae  including  species  of 
Caul erpa ,  Acetabularia,  P  e  n  i  c  i 1 1 u  s , 
G  r  a  c  i 1  a  r  i  a  ,  H  a  1  i  m  e  d  a  ,  Sargassum, 
Batophora ,  Udotea,  and  Dasya.  These  are 
discussed  at  length  by  Zieman  (in  prep.). 
Other  pertinent  references  for  mangrove 
regions  include  Davis  (1940),  Taylor 
(1960),  Tabb  and  Manning  (1961),  and  Tabb 
et   al.   (1962). 


5.2     PHYT0PLANKT0N 

All  aspects  of  phytoplankton,  from 
seasonal  occurrence  to  productivity 
studies,  are  poorly  studied  in  mangrove 
ecosystems.  This  is  particularly  true  in 
Florida. 

Evidence  from  Brazil  (Teixeira  et  al. 
1965,  1967,  1969;  Tundisi  1969)  indicates 
that  phytoplankton  can  be  an  important 
component  of  the  total  primary  production 
in  mangrove  ecosystems;  just  how  important 
is  not  clear.  Generally,  standing  crops 
of  net  phytoplankton  in  mangrove  areas  are 
low  (personal  observation).  The  nanno- 
plankton,  which  have  not  been  studied  at 
all,  appear  to  be  most  important  in  terms 
of  total  metabolism  (Tundisi  1969).  The 
net  plankton  are  usually  dominated  by 
diatoms  such  as  Thai assothri  x  s  p  p . , 
Chaetoceras  s  p  p . ,  Ni  t  zsc  hi  a  s  p  p . , 
Skeletonema  spp.,  and  Rhi zosol eni a  spp. 
(Mattox  1949;  Wood  1965;  Walsh  1967;  Bacon 
1970).  At  times,  blooms  of  dinoflagel- 
lates  such  as  P  e  r  i  d  i  n  i  u  m  spp.  and 
Gymnodi  ni  urn  spp.  may  dominate  (personal 
observation).  In  many  locations,  particu- 
larly in  shallow  waters  with  some  turbu- 
lence, benthic  diatoms  such  as  Pleurosigma 
spp.,  Mastogloia  spp.,  and  Disploneis  may 
be  numerically  important  in  the  net  plank- 
ton   (Wood    1965). 

Understanding  the  mangrove-associated 
phytoplankton  community  is  complicated  by 
the  constant  mixing  of  water  masses  in 
mangrove  regions.  Depending  upon  the 
location,  the  phytoplankton  may  be  domi- 
nated by  oceanic  and  neritic  forms,  by 
true  estuarine  plankton,  and  by  freshwater 
plankton.  The  pattern  of  dominance  may 
change  daily  or  seasonally  depending  upon 
the  source  of  the  principal    water  mass. 


Before  we  can  understand  the  impor- 
tance (or  lack  of  importance)  of  phyto- 
plankton in  mangrove  regions,  some  ques- 
tions must  be  answered.  How  productive 
are  the  nannoplankton?  How  does  the  daily 
and  seasonal  shift  in  phytoplankton  domi- 
nance affect  community  productivity?  Does 
the  generally  low  standing  crop  of  phyto- 
plankton represent  low  productivity  or  a 
high  grazing  rate? 


5.3     ASSOCIATED   VASCULAR    PLANTS 

Four  species  of  aquatic  grasses  occur 
on  bay  and  creek  bottoms  adjacent  to  man- 
grove forests.  Turtle  grass,  Thai assia 
testudinum,  and  manatee  grass,  Syringodium 
fi  Hi  forme,  are  two  tropical  sea  grasses 
which  occur  in  waters  with  average  salini- 
ties above  about  20  ppt.  Shoal  grass, 
Hal odule  wri  ghti  i ,  is  found  at  somewhat 
lower  salinities  and  widgeongrass,  Ruppia 
maritima,  is  a  freshwater  grass  which  can 
tolerate  low  salinities.  These  grasses 
occur  throughout  south  Florida,  often  in 
close  juxtaposition  to  mangroves.  Zieman 
(in  prep.)  presents  a  thorough  review  of 
sea  grasses  along  with  comments  about 
possible  energy  flow  linkages  with 
mangrove  ecosystems. 


For  example,  along  the  southwest  coast 
between  Flamingo  and  Naples,  marshes  are 
scattered  throughout  the  mangrove  belt  and 
also  border  the  mangroves  on  the  upland 
side.  The  estuarine  marshes  within  the 
mangrove    swamps    have    been    extensively 


eel  1 ul osa,  glass  wort,  Sal i  cornia  spp., 
Gulf  cordgrass,  Spartina  spartinae,  sea 
purslane,  Sesuvium  portulacastrum,  salt 
wort,  Batis  maritima,  and  sea  ox-eye, 
B o r  r i  c h i  a  f rutescens.  Farther  north, 
above  Tampa  on  the  west  coast  of  Florida, 
marshes    populated    by   smooth    cordgrass, 


43 


Spartina  al terni f 1 ora,  and  black  needle 
rush,  Juncus  roemerianus,  become  more 
extensive  and  eventually  replace  mangrove 
swamps.  Even  in  the  Everglades  region, 
the  saline  marshes  are  comparable  to  man- 
groves in  areal  extent,  although  they 
tend  to  be  some  distance  from  open  water. 
Studies  of  these  marshes,  including  as- 
sessment of  their  ecological  value,  are 
almost  non-existent.  Certainly,  they  have 
considerable  importance  as  habitat  for 
small  fishes  which,  in  turn,  support  many 
of  the  nesting  wading  birds  in  south 
Florida    (see    section    9). 

Tropical  hardwood  forests  may  occur 
within  the  mangrove  zone  in  south  Florida, 
particularly  where  old  shorelines  or  areas 
of  storm  sedimentation  have  created  ridges 
1  m  or  more  above  MSL  (mean  sea  level) 
(Olmstead  et  al.  1981).  Similar  forests 
or  "hammocks"  occur  to  the  rear  of  the 
mangrove  zone  on  higher  ground.  Typical 
trees  in  both  forest  types  include  the  fan 
palm,  Thri  nax  r  a  d  i  a  t  a  ,  buttonwood, 
Conocarpus  erecta,  manchineel,  Hippomane 
manci  nel 1  a,  and,  in  the  past,  mahogany, 
Swi  et en  i  a  maha  goni .  Olmstead  et  al. 
(1 981 )  provide  a  description  of  these 
communities. 

Freshwater  marsh  plants,  such  as  the 
grasses,  rushes  and  sedges  that  dominate 
the  freshwater  Everglades,  are  not 
mentioned  here,  although  they  are 
occasionally  mixed  in  with  small   mangroves 


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

Finally,  a  group  of  somewhat  salt- 
tolerant  herbaceous  plants  is  found 
within  stands  of  mangroves.  They  usually 
occur  where  slight  increases  in  elevation 
exist  and  where  sufficient  light  filters 
through  the  mangrove  canopy.  Carter  et 
al.  (1973)  list  the  following  as  examples 
of  members  of  the  mangrove  community: 
leather  ferns,  Acrostichum  aureum  and  A. 
danaeifolium;  Spanish  bayonet,  Yucca 
al oi  fol ia;  spider  lily,  Hymenocal  1  i  s 
1  ati  fol ia;  sea  blite,  Suaeda  1 ineari  s; 
chaff  flower,  Al ternanthera  ramosissima; 
samphire,  Philoxerus  vermicularis;  blood- 
leaf,  Iresine  celosia;  pricklypear  cactus, 
0  p  u  n  t  i  a  s  t  r  i  c  t  a  ;  marsh  elder,  I  va 
f rutescens;  the  rubber  vine,  Rhabdadeni  a 
bi f 1 ora;  the  lianas,  Ipomoea  tuba  and 
Hippocratea  volubilis;  and  a  variety  of 
bromeliads  (Bromel  i aceae). 

Although  the  lists  of  vascular  plants 
which  occur  in  mangrove  swamps  may  seem 
extensive,  the  actual  number  of  species  in 
any  given  location  tends  to  be  low 
compared  to  totally  freshwater  environ- 
ments (see  Carlton  1977).  Analogous  to 
temperate  salt  marshes,  mangrove  swamps 
possess  too  many  sources  of  stress, 
particularly  from  tidal  salt  water,  to 
have  a  high  diversity  of  vascular  plant 
species. 


44 


CHAPTER    6.       COMMUNITY    COMPONENTS    -    INVERTEBRATES 

6.1      ECOLOGICAL   RELATIONSHIPS  bryozoans,    and   tunicates.     The  most  ob- 
vious and  dominant  organisms   are  usually 

The  mangrove  ecosystem,  with  its  tree  barnacles,  crabs,  oysters,  mussels,  iso- 

canopies,   masses  of  aerial    roots,   muddy  pods,  polychaetes,  gastropods  and,  tuni- 

substrates,     and    associated    creeks    and  cates. 
small    embayments,    offers    many    habitat 

opportunities  for  a  wide  variety  of  inver-  A    striking    characteristic    of    most 

tebrates.     While  there  are  few  comparisons  mangrove  swamps  is  the  pattern  of  horizon- 

of  species   richness  with  other  types   of  tal    and  vertical    zonation  of  invertebrates 

coastal  ecosystems,  mangrove  swamps  appear  (Figure  9).      Characteristic   vertical    zona- 

to   be   characterized   by   moderately    high  tion    patterns    are    found   on   the   prop   roots 

invertebrate    species    diversity.      Abele  (Rutzler  1969)   and   not   so  obvious   horizon- 

(1974)   compared  H'    (Shannon    Weaver)    diver-  tal   distributions  occur  as  you  move  back 

sity    of    decapod    crustaceans    between  into    the    center    of    the    swamp    (Warner 

various  littoral  marine  communities  and  1969).      Invertebrate   biomass   in   the   red 

found    mangrove    swamps    in    an    intermediate  mangrove  zone  on  the  edge  of  the  swamp  may 

position  with  more  decapod  species  than  be  very  high,   often  in  excess  of  100  dry 

Sparti  na    marshes    but    considerably   less  g/m     of   organic    matter    in    many    locations 

than   were   associated   with   rocky   substrate  (personal    observation).      In    the    center    of 

communities.  the   swamp,    particularly   where   there    is 

little   flooding,    biomass    is    usually   an 

There  is  little  doubt  that  the  maze  order    of    magnitude    less;    Golley    et    al. 

of  prop  roots  and  muddy  substrates  under  (1962)    found    an    average    of    6.4    g/m      of 

intertidal    mangrove  trees  provides  habitat  invertebrates   in  the   center  of  a   Puerto 

for    a    wide    range    of    invertebrates    and  Rican  mangrove  swamp, 
fishes  (Figure  10)  (see  section  7  for  the 

latter).     The  nursery  value  of  the   prop  Mangrove-associated  invertebrates  can 

root    complex    for   juvenile   spiny   lobsters,  be  placed  in  four  major  categories  based 

Panulirus    argus,    is    well    established  on   trophic   position: 
(01  sen    et   al.    1975;  Olsen   and   Koblic   1975; 

Little    1977;    Witham    et    al.    1968).      Ac-  (1)  direct  grazers   -  limited   to 
cording  to   these   researchers,   the  phyl- 

losome    larvae    of    spiny    lobsters    often  (a)    insects    and   the    mangrove   tree 

settle   among   the   prop    roots    and   remain  crab,  Aratus   pisonii,   all    of  which   feed  on 

there   for  much   of  their  juvenile   lives.  leaves   in  the  mangrove  canopy  and 
The    prop    roots    provide   protection    from 

predators  and  a  possible  source  of  food   in  (b)  a  group  of  small    invertebrates 

the  associated  populations  of  small    inver-  which   graze  the  prop  root  and  mud  algae 

tebrates.     To  provide  the  best  habitat,  a  directly; 
section   of  the   prop   roots  should  extend 

below  mean   low  tide.      If  conditions  are  (2)    filter    feeders    -    largely    sessile 

suitable,    the    juveniles    may    remain    in  prop   root    invertebrates    which    filter   phy- 

close   association   with  the  prop   root   com-  toplankton   and  detritus   from  the  water; 
munity  for  as  much  as  2  years  until   they 

reach  a  carapace  length  of  60  to  70  mm.  (3)   deposit   feeders   -    mobile   inverte- 
brates   which    skim    detritus,    algae    and 

In  addition  to  its  value  as  spiny  occasional  small  animals  from  the  surface 
lobster  habitat,  mangrove  ecosystems  also  of  the  mud  and  forest  floor; 
harbor  the  following  invertebrates:  bar- 
nacles, sponges,  polychaete  worms,  gastro-  (4)  carnivores  -  highly  mobile  inverte- 
pod  mollusks,  pelecypod  mollusks,  isopods,  brates  which  feed  upon  the  three  preceding 
amphipods,  mysids,  crabs,  caridean  shrimp,  groups  in  all  locations  from  the  tree 
penaeid  shrimp,  harpacticoid  copepods,  canopy  (largely  insects)  to  the  mud  sur- 
snapping  shrimp,  ostracods,  coelenterates,  face.  Food  sources  in  mangrove  swamps  and 
nematodes,     a    wide    variety    of    insects,  energy   flow   are  discussed   in   section   3.6. 

45 


46 


6.2.  ARBOREAL  ARTHROPOD  COMMUNITY 

A  surprising  variety  of  arthropods 
inhabit  the  mangrove  canopy.  Because  they 
are  frequently  secretive  or  possess 
camouflage  coloration,  their  numerical 
importance  often  has  been  overlooked. 
Beever  et  al.  (1979)  pointed  out  that 
arboreal  arthropods  have  a  variety  of 
ecological  roles:  (1)  direct  herbivory  on 
mangrove  leaves,  (2)  predator-prey  inter- 
actions, and  (3)  biomass  export  through 
frass  production  and  leaf  defoliation. 
Direct  grazing  is  typically  patchy  in 
distribution.  It  is  not  unusual  to  find 
extensive  stretches  of  mangroves  that  have 
scarcely  been  grazed.  In  nearby  areas,  as 
much  as  80%  of  the  leaves  may  have  some 
damage  (Beever  et  al.  1979).  As  a  general 
rule,  it  is  probably  safe  to  state  that 
healthy,  unstressed  mangrove  stands  nor- 
mally have  less  than  10%  of  their  total 
leaf  area  grazed  (Heald  1969).  In  many 
locations,  percent  leaf  area  damaged  is  on 
the  order  of  1%  to  2%  (Beever  et  al. 
1979).  There  are  exceptions.  Onuf  et  al. 
(1977)  reported  biomass  loss  to  arthropod 
grazers  as  high  as  26%  in  a  mangrove  stand 
where  growth  and  nitrogen  content  of  the 
leaves  had  been  enhanced  by  input  of  nu- 
trients  from  a   bird   rookery. 

In  terms  of  numbers  of  species,  the 
dominant  group  of  arboreal  arthropods  is 
insects.  The  most  thorough  inventory  of 
mangrove-associated  insects  was  conducted 
by  Simberloff  and  Wilson  to  obtain  the  raw 
data  for  their  papers  on  island  bio- 
geography  (Simberloff  and  Wilson  1969; 
Simberloff  1976).  These  papers  list  over 
200  species  of  insects  associated  with 
overwash  mangrove  islands  in  the  Florida 
Keys.  There  is  no  reason  to  expect  lesser 
numbers  in  other  types  of  mangrove  com- 
munities, except  for  the  mangrove  scrub 
forests.  The  most  thorough  study  of  in- 
sect grazing  on  mangrove  leaves  is  that  of 
Onuf  et  al.  (1977)  (see  section  2.6). 

Although  not  as  numerically  impres- 
sive as  the  insects,  the  mangrove  tree 
crab,  Aratus  pisonii ,  appears  to  be  poten- 
tially  as  important  in  terms  of  grazing 
impact  (Beever  et  al.  1979).  The  life 
history   of   this    secretive    little  crab  has 


been  described  by  Warner  (1967).  In 
Jamaica  its  numbers  range  from  11  to  16/m 
at  the  edge  of  fringing  swamps  to  6/nr  in 
the  center  of  large  swamps.  Beever  et  al. 
(1979)  reported  typical  densities  for  a 
variety  of  sites  in  south  Florida  of  1  to 
4  crabs/m  .  These  same  authors  reported 
some  interesting  details  about  the  crab: 
(1)  the  diet  is  omnivorous  ranging  from 
fresh  mangrove  leaves  to  caterpillars, 
beetles,  and  various  insects;  (2)  the  crab 
suffers  highest  predation  pressure  while 
in  the  planktonic  larval  stage;  (3)  preda- 
tion on  the  crabs  while  in  the  arboreal 
community  is  low  and  comes  from  birds  such 
as  the  white  ibis,  raccoons,  other  man- 
grove tree  crabs  and,  if  the  crabs  fall  in 
the  water,  fishes  such  as  the  mangrove 
snapper;  and  (4)  in  one  location  in  south 
Florida  (Pine  Island  Sound)  they  found  in 
accordance  with  normal  bi ogeographical 
theory,  the  highest  densities  of  crabs 
associated  with  fringing  forests  and  the 
lowest  densities  on  distant  islands,  but 
at  Sugar  Loaf  Key  the  unexpl ai nabl e 
reverse  distribution  was   found. 

Other  invertebrates  may  visit  the 
canopy  from  below  either  for  purposes  of 
feeding  or  for  protection  from  high  tides. 
Included  in  this  group  are  the  pulmonate 
gastropods,  L  i  tt  o  r i  na  angul i  f era  , 
Ceri thidea  seal  ari  formi  s,  and  Mel  ampus 
cof feus,  the  isopod,  Li  gea  exoti  ca,  and  a 
host  of  small    crabs. 

In  summary,  with  the  exception  of  a 
half  dozen  key  papers,  the  arboreal  man- 
grove community  has  been  generally  ig- 
nored. Both  insects  and  the  mangrove  tree 
crab  play  significant  ecological  roles  and 
may  affect  mangrove  productivity  to  a 
greater  extent  than   has  been  recognized. 

6.3     PROP   ROOT   AND   ASSOCIATED    MUD    SURFACE 
COMMUNITY 

These  two  somewhat  distinct  com- 
munities have  been  lumped  together  because 
of  the  large  number  of  mobile  organisms 
which  move  back  and  forth  between  tidal 
cycles.  The  aerial  roots  are  used  as 
protective  habitat  and  to  some  extent  for 
feeding  while  the  nearby  mud  substrates 
are  used   principally   for   feeding. 


47 


The  prop  roots  support  an  abundance 
of  sessile  organisms.  The  vertical 
zonation  of  both  mobile  and  sessile  inver- 
tebrates has  been  studied  extensively  in 
other  parts  of  the  world  (Goodbody  1961; 
Macnae  1968;  Rutzler  1969;  Coomans  1969; 
Bacon  1970;  Kolehmainen  1973;  Sasekumar 
1974;  Yoshioka  1975).  Vertical  zonation 
certainly  exists  on  Florida  red  mangrove 
roots.  The  generalized  scheme  shown  in 
Figure  9  essentially  contains  two  zones: 
an  upper  zone  dominanted  by  barnacles  and 
a  lower  zone  dominated  by  mussels,  oysters 
and  ascidians.  Between  mean  high  tide  and 
mean  tide,  the  wood  boring  isopod, 
Sphaeroma  terebrans  (discussed  at  length 
in  section  2.7)  is  important,  both  numeri- 
cally and  through  the  provision  of 
numerous  holes  for  use  by  other  organisms 
(Estevez   1978). 

The  most  complete  study  of  the 
Florida  mangrove  prop  root  community  is 
Courtney's  (1975)  comparison  of  seawall 
and  mangrove  associations.  He  reported  an 
extensive  list  of  invertebrates  from  man- 
grove prop  roots  at  Marco  Island,  Florida, 
including:  Crassost  rea  v i  r g i  n  i  c a  , 
Li ttori  na  angul i  fera,  Crepidula  pi  ana, 
Diodora  cayenensis,  Urosalpinx  perrugata, 
Pisania  tincta,  Brachidontes  exustus, 
nine  species  of  polychaetes,  Sphaeroma 
terebrans ,  Pal aem  on  floridanus , 
Peri  cl i  menes  longi  caudatus,  Synalpheus 
fritzmuelleri,  Thor  floridanus, 
Petrol isthes  armatus,  and  at  least  eight 
species  of  crabs.  The  following  species 
were  found  only  on  mangrove  roots  and  not 
on  seawalls:  T u r i t e 1 1  a  sp.,  Mel ongena 
corona ,  Anachi  s  semiplicata,  Bulla 
striata,  Hypselodoris  sp.,  Area  imbricata, 
Carditamera  floridana,  Pseudoi  rus  typica, 
and    Martesia    striata. 

Tabb  et  al.  (1962)  and  Odum  and  Heald 
(1972)  reported  a  variety  of  invertebrates 
associated  with  prop  roots  in  the  White- 
water Bay  region.  Although  many  species 
coincide  with  Courtney's  (1975)  list, 
there  are  also  significant  differences  due 
to  the  lower  salinities  in  this  region. 
It  is  probably  safe  to  conclude  that  prop 
root  communities  vary  somewhat  from  site 
to  site  in   response  to  a  number  of  factors 


including  latitude,  salinity,  and  proxi- 
mity to  other  communities  such  as  sea 
grass   beds   and   coral    reefs. 

Sutherland  (1980),  working  on  red 
mangrove  prop  root  communities  in 
Venezuela,  found  little  change  in  the 
invertebrate  species  composition  on  indi- 
vidual prop  roots  during  an  18-month 
period.  The  species  composition  varied 
greatly,  however,  between  adjacent  prop 
roots,  presumably  in  response  to  stochas- 
tic (chance)  processes. 

The  mud  flats  adjacent  to  mangroves 
provide  feeding  areas  for  a  range  of  in- 
vertebrates that  scuttle,  crawl,  and  swim 
out  from  the  cover  of  the  mangrove  roots. 
Some  emerge  at  low  tide  and  feed  on  algae, 
detritus,  and  small  invertebrates  on  the 
mud  flats  while  they  are  high  and  dry. 
Others  emerge  while  the  tide  is  in,  parti- 
cularly at  night,  and  forage  across  the 
flooded  flats  in  search  of  the  same  foods 
plus  other  invertebrates  which  have 
emerged  from  the  mud.  In  many  ways  the 
mangrove-mud  flat  relationship  is  analo- 
gous to  the  coral  reef  (refuge)  sea  grass 
(feeding  area)  relationship  reviewed  by 
Zieman  (in  prep.).  The  net  effect  is  that 
the  impact  of  the  mangrove  community  may 
extend  some  distance  beyond  the  boundaries 
of  the   mangrove  forest. 

In  addition  to  the  organisms  which 
move  from  the  mangroves  to  the  mud  flats, 
there  is  a  small  group  which  uses  the 
substrate  adjacent  to  mangroves  for  both 
habitat  and  feeding.  In  the  Whitewater 
Bay  region,  four  crabs  exploit  the  inter- 
tidal  muds  from  the  safety  of  burrows: 
Ilea  pugi  1  ator,  LL  speciosa,  U.  thayeri , 
and  Euryti  urn  1  i  mosum  (Tabb  et  al.  1962). 
In  low  salinity  mangrove  forests  of  south 
Florida,  the  crayfish,  Procambarus  alleni, 
is  a  dominant  member  of  the  burrowing, 
benthic  community  (Hobbs  1942)  as  is  the 
crab,  Rhithropanopeus  harrisii  (Odum  and 
Heald  1972).  Both  organisms  are  found  in 
a  remarkable  number  of  fish   stomachs. 

The  benthic  fauna  and  infauna  of 
creek  and  bay  bottoms  near  mangrove 
forests    are    highly    variable    from    one 


48 


location  to  the  next.  Many  of  these 
organisms,  particularly  the  deposit  and 
filter  feeders,  benefit  from  particulate 
organic  matter  originating  from  mangrove 
litter  fall  (Odum  and  Heald  1972,  1975b). 
Tabb  and  Manning  (1961)  and  Tabb  et  al. 
(1962)  present  lists  and  discussions  of 
many  of  the  benthic  invertebrates  adjacent 
to  mangrove  areas  of  Whitewater  Bay. 
Weinstein  et  al.  (1977)  compared  the  ben- 
thic fauna  of  a  mangrove-lined  creek  and  a 
nearby  man-made  canal  on  Marco  Island. 
They  found  (1)  the  mangrove  fauna  to  be 
more  diverse  than  the  canal  fauna  and  (2) 
a  higher  diversity  of  organisms  at  the 
mouths  of  mangrove  creeks  than  in  the 
"heads"  or  upstream  ends.  Courtney  (1975) 
found  the  same  pattern  of  upstream 
decreases  in  diversity,  presumably  in 
response  to  decreasing  oxygen  concentra- 
tions  and   increasingly  finer  sediments. 

Finally,  the  irregularly  flooded  sub- 
strates in  the  center  of  mangrove  forests 
contain  a  small  but  interesting  assemblage 
of  invertebrates.  The  litter  layer, 
composed  largely  of  mangrove  leaves,  evi- 
dently includes  a  variety  of  nematodes. 
Due  to  the  usual  taxonomic  difficulties  in 
identifying  nematodes,  complete  species 
lists  do  not  exist  for  mangrove  forests; 
however,  many  species  and  individuals  are 
associated  with  the  decaying  leaves 
(Hopper  et  al.  1973).  In  addition  to 
nematodes,  the  wetter  sections  of  the 
swamp  floor  can  contain  mosquito  and  other 
insect  larvae,  polychaetes,  harpacticoid 
copepods,  isopods,  and  amphipods. 
Simberloff  (1976)  lists  16  species  of 
insects  associated  with  the  muddy  floor  of 
mangrove  forests.  Roaming  across  the 
forest  floor  during  low  tide  are  several 
crustaceans  including  the  mangrove  tree 
crab,  Aratus  pi  soni  i ,  crabs  of  the  genus 
Sesarma,  and  the  pulmonate  gastropods, 
Me  1  a  mpu  s  coeffeus  and  Cerithidea 
scalariformis.  Both  snails  clearly  have 
the  ability  to  graze  and  consume  recently 
fallen  leaves  (personal  observation). 
With  favorable  conditions  (relatively  fre- 
quent tidal  inundation  plus  the  presence 
of  red  mangroves)  Melampus  populations  can 


exceed   500/m     and  average  100  to  200/m 
(Heald,  unpublished   data).     Cerithidea  i  s 
found  largely  in   association  with  black 
mangroves  ayid  can   reach  densities   of   at 
least    400/n/. 


6.4     WATER  COLUMN  COMMUNITY 

This  section  is  embarrassingly  short; 
the  reasons  for  this  brevity  are  (1)  the 
paucity  of  research  on  zooplankton  in 
Florida  mangrove-dominated  areas  and  (2) 
our  inability  to  discover  some  of  the  work 
which  undoubtedly  has  been  done.  Davis 
and  Williams  (1950)  are  usually  quoted  as 
the  primary  reference  on  Florida  mangrove- 
associated  zooplankton,  but  their  paper 
only  lists  zooplankters  collected  in  two 
areas.  Zooplankton  near  mangroves  are 
probably  no  different  from  those  found  in 
other  shallow,  inshore  areas  in  south 
Florida.  Based  on  Davis  and  Williams 
(1950)  and  Reeve  (1964),  we  can  hypothe- 
size that  the  community  is  dominated  by 
copepod  species  of  genus  Acartia,  particu- 
larly Acarti  a  tonsa.  In  addition,  we 
could  expect  a  few  other  calanoid  cope- 
pods,  arrow  worms  (Sagi tta  spp.),  many 
fish,  polychaete  and  crustacean  larvae  and 
eggs.  Another  component  of  the  "plankton" 
particularly  at  night,  are  benthic 
amphipods,  mysids,  and  isopods  which  leave 
the   bottom  to   feed    (personal    observation). 

Plankton  are  not  the  only  inverte- 
brates in  the  water  column.  Swimming 
crabs,  such  as  the  blue  crab,  Callinectes 
sapidus,  are  plentiful  in  most  estuarine 
mangrove  regions  of  south  Florida.  Other 
swimming  crustaceans  include  the  caridean 
shrimp  (Pal  aemonetes  spp.  and  Peri  - 
climenes  spp.),  the  snapping  shrimp 
(Al  pheus~spp.),  and  the  penaeid  shrimp 
(Penaeus  spp).  All  of  these  swimming 
crustaceans  spend  considerable  time  on  or 
in  the  benthos  and  around  mangrove  prop 
roots.  From  the  economic  point  of  view, 
the  pink  shrimp,  Penaeus  duorarum,  is 
probably  the  most  important  species  asso- 
ciated with  mangrove  areas  (see  discussion 
in  section  11 ). 


49 


CHAPTER  7.   COMMUNITY  COMPONENTS  -  FISHES 


Of  the  six  mangrove  community  types 
discussed  in  section  1.5,  fishes  are  an 
important  component  of  four:  (1)  basin 
forests,  (2)  riverine  forests,  (3)  fringe 
forests,  and  (4)  overwash  island  forests. 
For  convenience  we  have  divided  fringe 
forests  into  two  sub-components:  (a) 
forests  which  fringe  estuarine  bays  and 
lagoons  and  (b)  forests  which  fringe 
oceanic  bays  and  lagoons.  This  division 
is  necessary  because  the  fish  communities 
differ   markedly. 

Mangroves  serve  two  distinct  roles 
for  fishes  and  it  is  conceptually  impor- 
tant to  distinguish  between  them.  First, 
the  mangrove-water  interface,  generally 
red  mangrove  prop  roots,  afford  a  rela- 
tively protected  habitat  which  is  particu- 
larly suitable  for  juvenile  fishes. 
Secondly,  mangrove  leaves,  as  discussed  in 
section  3.6,  are  the  basic  energy  source 
of  a  detritus-based  food  web  on  which  many 
fishes  are  dependent.  The  habitat  value 
of  mangroves  can  be  considered  strictly  a 
function  of  the  area  of  interface  between 
the  water  and  the  mangrove  prop  roots;  it 
is  an  attribute  shared  by  all  four  types 
of  mangrove  communities.  The  importance 
of  the  mangrove  detritus-based  food  web  is 
dependent  on  the  relative  contribution  of 
other  forms  of  energy  in  a  given  environ- 
ment, including  phytopl ankton,  benthic 
algae,  sea  grass  detritus,  and  terrestrial 
carbon  sources.  Figure  11  provides  a 
diagrammatic  representation  of  the  rela- 
tive positions  along  a  food  web  continuum 
of  the  four  mangrove  communities. 

Fishes  recorded  from  mangrove  habi- 
tats in  south  Florida  are  listed  in  Appen- 
dix B.  Although  the  fish  communities  are 
discussed  separately  below,  they  have  been 
combined  into  certain  categories  in  Appen- 
dix B;  fishes  from  mangrove  basins  and 
riverine  forests  have  been  combined  under 
the  heading  of  tidal  streams;  fishes  from 
fringing  forests  along  estuarine  bays  and 
lagoons  are  listed  under  the  heading  of 
estuarine  bays;  fishes  from  oceanic  bays 
and  lagoons  have  been  listed  under  oceanic 
bays.  Since  no  surveys  have  been 
published  specifically  relating  to  over- 
wash  island  forests,  there  is  no  listing 
for    this    community   type   in   Appendix   B. 


Site  characteristics  and  sampling  methods 
for  these  community  types  are  summarized 
in  Appendix  A.  Nomenclature  and  taxonomic 
order   follow  Bailey  et    al.    (1970). 


7.1      BASIN   MANGROVE   FORESTS 

The  infrequently  flooded  pools  in  the 
black  mangrove-dominated  zone  provide  an 
extreme  habitat  which  few  species  of 
fishes  can  tolerate.  The  waters  are 
darkly  stained  with  organic  acids  and 
tannins  leached  from  the  thick  layer  of 
leaf  litter.  Dissolved  oxygen  is 
frequently  low  (1-2  ppm)  and  hydrogen 
sulfide  is  released  from  the  sediments 
following  physical  disturbance.  Salini- 
ties are  highly  variable  ranging  from 
totally  fresh  to  hypersaline.  The  fish 
families  best  adapted  to  this  habitat  are 
the  euryhaline  cyprinodonts  (killifishes) 
and  the  poeciliids  (1 i vebearers).  The 
killifishes  include  Fundulus  confluentus 
(Heald  et  al.  1974),  Rivulus  marmoratus 
(M.  P.  Weinstein,  Va.  Commonwealth  Univ., 
Richmond,  Va.;  personal  communication 
1981),  Floridichthys  ca  rpi  o,  and 
Cyprinodon  variegatus  (Odum  1970").  The 
poeciliids  include  Poecilia  latipinna 
(Odum  1970)  and,  the  most  common,  Gambusia 
affinis  (Heald  et  al.  1974).  While  the 
species  richness  of  fishes  in  this  habitat 
is  low,  the  densities  of  fish  are  often 
very  high.  Weinstein  Qjers.  comm.)  has 
recorded   up  to  38  fish/m  . 

All  of  these  fishes  are  permanent 
residents,  completing  their  life  cycles  in 
this  habitat.  They  feed  primarily  on 
mosquito  larvae  and  small  crustaceans  such 
as  amphipods  which,  in  turn,  feed  on  man- 
grove detritus  and  algae.  These  small 
fishes  enter  coastal  food  webs  when  they 
are  flushed  into  the  main  watercourses 
during  high  spring  tides  or  following 
seasonally  heavy  rains.  Here  they  are 
eaten  by  numerous  piscivorous  fishes  in- 
cluding snook,  ladyfish,  tarpon,  gars,  and 
mangrove  snappers.  The  alternate  energy 
pathway  for  fishes  of  the  black  mangrove 
basin  wetlands  occurs  when  the  pools 
shrink  during  dry  weather,  the  fishes  are 
concentrated  into  smaller  areas,  and  are 
fed-upon  by  various  wading  birds  including 


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herons,  ibis  and  the  wood  stork  (Heald  et 
al.    1974). 


7.2     RIVERINE    FORESTS 

Tidal  streams  and  rivers,  fringed 
largely  by  red  mangroves,  connect  the 
freshwater  marshes  of  south  Florida  with 
the  shallow  estuarine  bays  and  lagoons 
(Figure  12).  Few  of  these  streams  have 
been  studied  thoroughly.  The  exception  is 
the  North  River  which  flows  into  White- 
water Bay  and  was  studied  by  Tabb  (1966) 
and  Odum  (1970).  Springer  and  Woodburn 
(1960)  collected  fishes  in  a  bayou  or 
tidal  pass  connecting  Boca  Ciega  Bay  and 
Old  Tampa  Bay.  Carter  et  al.  (1973) 
reported  on  the  fishes  of  two  tidal 
streams  entering  Fahkahatchee  and  Fahka 
Union  Bays.  Nugent  (1970)  sampled  fishes 
in  two  streams  on  the  western  shore  of 
Biscayne  Bay.  Characteristics  of  these 
areas  and  sampling  gear  used  by  the  inves- 
tigators are  summarized  in  Appendix  A. 

These  tidal  streams  and  associated 
riverine  mangrove  forests  exhibit  extreme 
seasonal  variability  in  both  physical 
characteristics  and  fish  community  compo- 
sition. Salinity  variations  are  directly 
related  to  changes  in  the  make-up  of  the 
fish  assemblage.  During  the  wet  season 
(June  -  November),  salinities  fall 
throughout  the  water  courses  and,  at  some 
locations  in  certain  heavy  runoff  years, 
become  fresh  all  of  the  way  to  the  mouth 
(Odum  1970).  Opportunistic  freshwater 
species,  which  are  normally  restricted  to 
the  sawgrass  and  black  needle  rush  marshes 
of  the  headwaters,  invade  the  mangrove 
zone.  These  include  the  Florida  gar, 
Lepi  sosteus  pi  a ty  r  h  i  nc  us;  several 
centrarchid  sunfishes  of  the  genus  Lepomis 
and  the  largemouth  bass,  Micropterus 
sal moi  des  ;  the  freshwater  catfishes, 
Ictalurus  natalis  and  Noturus  gyrinus;  and 
the  killifishes  normally  considered 
freshwater  inhabitants  such  as  Lucani  a 
goodei    and  Ri vulus   marmoratus. 

During  the  dry  season  (December  to 
early  May)  salinities  rise  as  a  result  of 
decreased  freshwater  runoff  and  continuing 
evaporation.      Marine  species   invade  the 


tidal  streams  primarily  on  feeding  forays. 
Examples  include  the  jewfish,  Epinephelus 
ita jara,  the  stingrays  (Dasyatidae),  the 
needlefishes  (Belonidae),  the  jacks 
(Carangidae),  and  the  barracuda,  Sphyraena 
barracuda.  Other  seasonal  movements  of 
fishes  appear  to  be  temperature  related. 
Tabb  and  Manning  (1961)  documented  move- 
ments of  a  number  of  species  from  shallow 
inshore  waters  to  deeper  water  during 
times  of  low  temperature  stress.  The 
lined  sole,  the  hogchoker,  the  bighead 
searobin,  and  the  striped  mullet,  for 
example,  are  much  less  frequently  caught 
in  winter  in  shallow  inshore  waters. 

A  third  type  of  seasonality  of  fish 
populations  in  the  tidal  rivers  is  related 
to  life  cycles.  Many  of  the  fish  which 
utilize  the  tidal  stream  habitat  do  so 
only  as  juveniles.  Thus,  there  are  peaks 
of  abundance  of  these  species  following 
offshore  spawning  when  larval  or  juvenile 
forms  are  recruited  to  the  mangrove  stream 
habitat.  In  general,  recruitment  occurs 
in  the  late  spring  or  early  summer  fol- 
lowing late  winter  and  spring  spawning 
offshore  or  in  tidal  passes  (Reid  1954). 
Numerous  species  are  involved  in  this  life 
cycle  phenomenon  including  striped  mullet, 
grey  snapper,  sheepshead,  spotted  sea 
trout,    red  drum,    and  silver  perch. 

The  only  estimate  of  fish  standing 
crop  from  tidal  stream  habitats  is  that  of 
Carter  et  al.  (1973).  They  recorded  27 
species  weighing  65,891  g  (wet  wt.)  from 
an  area  of  734  m  or  about  90  g/m  .  This 
is  probably  an  overestimate  since  an  un- 
known portion  of  the  fish  community  had 
moved  from  the  flooded  lowlands  to  the 
stream  on  the  ebb  tide;  sampling  occurred 
at  low  tide  in  October.  Nonetheless,  this 
is  an  indication  of  the  high  fish  standing 
crop  which  this  mangrove-associated  habi- 
tat can  support.  The  number  of  species 
reported  from  individual  tidal  streams 
annually  ranges  from  47  to  60  and  the 
total  from  all  tidal  streams  in  southwest 
Florida    is    111    species    (Appendix   B). 

The  food  webs  in  these  riverine  man- 
grove ecosystems  appear  to  be  predomi- 
nantly mangrove  detritus-based,  although 
the  Biscayne  Bay   stream   studied   by   Nugent 


52 


Figure  12.  Aerial  photograph  of  the  mangrove  belt  of  southwest  Florida  near 
Whitewater  Bay.  Note  the  complex  system  of  pools  and  small  creeks  which  connect 
with  the  tidal  river  system. 


53 


(1970)  may  be  an  exception.  The  basic 
link  between  the  mangrove  leaf  and  higher 
order  consumers  is  provided  by  micro- 
organisms (fungi,  bacteria,  Protozoa) 
which  colonize  the  decaying  leaf  and  con- 
vert them  into  a  relatively  rich  protein 
source  (Odum  1970;  Odum  and  Heald  1975a). 
These  decaying  leaf  fragments  with  asso- 
ciated microorganisms  are  fed  upon  by  a 
group  of  omnivorous  detritivores  including 
amphipods,  mysids,  cumaceans,  ostracods, 
chironomid  larvae,  harpacticoid  and 
calanoid  copepods,  snapping  shrimp, 
caridean  and  penaeid  shrimp,  a  variety  of 
crabs,  filter-feeding  bivalves,  and  a  few 
species  of  fishes  (Odum  1970;  Odum  and 
Heald  1972;  Odum  and  Heald  1975b).  These 
detritivores,  in  turn,  are  consumed  by  a 
number  of  small  carnivorous  fishes,  which 
in  turn,  are  consumed  by  larger 
piscivorous  fishes.  The  concept  of  man- 
grove trophic  structure  is  also  discussed 
in  section  3.6.  See  Appendix  B  for 
species   specific   dietary   information. 

The  tidal  creeks  studied  by  Nugent 
(1970)  on  the  western  shore  of  Biscayne 
Bay  differ  from  the  previously  discussed 
streams  in  the  Everglades  estuary.  The 
mouths  of  the  Biscayne  Bay  creeks  have 
dense  growths  of  sea  grasses  which  con- 
tribute sea  grass  detritus.  The  salini- 
ties are  considerably  greater  and  the 
streams  are  located  only  a  few  kilometers 
from  coral  reefs,  which  are  largely  absent 
on  Florida's  west  coast,  at  least  close  to 
shore.  As  a  result,  23  species  listed  in 
Appendix  B  were  captured  by  Nugent  (1970) 
and  are  not  recorded  from  riverine  man- 
grove habitat  on  the  west  coast  of 
Florida.  Examples  include  several  of  the 
grunts  (Pomadasyidae),  the  gray  trigger- 
fish,  Balistes  capriscus,  the  barbfish, 
Scorpaena  brasiliensis,  the  scrawled  box- 
fish,  Lactophrys  quadricorni  s,  and  the 
snappers,    Lutjanus   apodus   and  L.    synagris. 

Riverine  mangrove  communities  and 
associated  tidal  streams  and  rivers  are 
typified  by  the  following  families  of 
fishes:  killifishes  (Cypri nodontidae), 
livebearers  (Poeci 1 i idae),  silversides 
(Atherinidae),  mojarras  (Gerreidae),  tar- 
pon (Elopidae),  snook  (Centropomidae), 
snappers    (Lut jani dae) ,     sea    catfishes 


(Ariidae),  gobies  (Gobi idae),  porgys 
(Sparidae),  mullets  (Mugilidae),  drums 
(Sci  aenidae),  and  anchovies  (Engraulidae). 
The  mangrove-lined  streams  and  associated 
pools  are  important  nursery  areas  for 
several  marine  and  estuarine  species  of 
gamefish.  The  tarpon,  Megalops  atlantica, 
snook,  Centropomus  undecimalis,  and  lady- 
fish,  Elops  saurus,  utilize  these  areas 
from  the  time  they  reach  the  estuary  as 
post-larvae,  having  been  spawned  offshore. 
Gray  snapper,  Lutjanus  g  r  i  s  e  u  s , 
sheepshead,  Archosargus  probatocephalus, 
spotted  seatrout,  Cynoscion  nebulosus,  and 
red  drum,  Sciaenops  ocellata,  are  re- 
cruited to  grass  beds  of  shallow  bays  and 
lagoons  as  post-larvae  and  enter  the 
mangrove-lined  streams  for  the  next  sever- 
al years  (Heald  and  Odum  1970).  Of  these 
species,  only  the  spotted  seatrout  prob- 
ably spawns  in  the  estuary  (Tabb  1966). 
Other  species  of  commercial  or  game  impor- 
tance which  use  the  riverine  fringing 
habitat  include  crevalle  jack,  gafftopsail 
catfish,  jewfish,  striped  mojarra,  barra- 
cuda, Atlantic  thread  herring,  and  yellow- 
fin  menhaden  (Odum  1970). 


7.3     FRINGING   FORESTS   ALONG   ESTUARINE   BAYS 
AND   LAGOONS 

Mangrove-fringed  estuarine  bays  and 
lagoons  are  exemplified  by  the  Ten 
Thousand  Islands  area  and  Whitewater  Bay. 
Quantitative  fish  data  are  available  from 
Fahkahatchee  Bay  (Carter  et  al.  1973; 
Yokel  1975b;  Seaman  et  al.  1973),  Fahka 
Union  Bay  (Carter  et  al.  1973),  Rookery 
Bay  (Yokel  1975a),  the  Marco  Island 
Estuary  (Weinstein  et  al.  1977;  Yokel 
1975a),  and  Whitewater  Bay  (Clark  1970). 
Individual  site  characteristics  are 
summarized  in  Appendix  A.  All  except 
Fahka  Union  Bay  contain  significant 
amounts  of  sea  grasses.  Macroalgae  domi- 
nate the  benthic  producers  of  Fahka  Union 
Bay.  Studies  by  Reid  (1954)  and  Kilby 
(1955)  near  Cedar  Key,  Florida,  were  not 
included  in  our  summary  because  mangroves 
are  sparse  in  this  area  and  no  mention  of 
mangrove  collecting  sites  were  made  by 
these  authors.  Studies  of  Caloosahatchee 
Bay  (Gunter  and  Hall  1965)  and  of 
Charlotte    Harbor    (Wang    and    Raney    1971) 


54 


were  omitted  because  the  areas  studied 
have  been  highly  modified  and  because  data 
from  many  habitats  were  pooled  in  the 
final   presentation. 

All  of  the  bays  reviewed  in  our  sum- 
maries are  fringed  by  dense  growths  of  red 
mangroves  and  all  contain  small  mangrove 
islets.  Carter  et  al.  (1973),  in  their 
studies  of  Fahkahatchee  and  Fahka  Union 
bays,  estimated  that  57%  to  80%  of  the 
total  energy  budget  of  these  two  bays  is 
supported  by  exports  of  particulate  and 
dissolved  organic  matter  from  the  man- 
groves within  the  bays  and  inflowing  tidal 
streams.  Lugo  et  al.  (1980)  estimated 
that  the  mangroves  surrounding  Rookery  Bay 
provide  32%  of  the  energy  base  of  the 
heterotrophic  community  found  in  the  bay. 

Salinities  in  these  bays  tend  to  be 
higher  than  in  the  tidal  streams  and 
rivers  and  the  fish  assemblages  reflect 
both  this  feature  and  the  added  habitat 
dimension  of  sea  grass  and  macro  algae 
beds.  Truly  freshwater  species  are  rare 
in  these  communities  and  a  proportionally 
greater  percentage  of  marine  visitors  is 
present.  The  dominant  fish  families  of 
the  benthic  habitat  include  drums 
(Sci aenidae),  porgys  (Sparidae),  grunts 
(Pomadasyidae),  mojarras  (Gerreidae), 
snappers  (Lutjanidae),  and  mullet  (Mugili- 
dae).  Other  familes  with  sizeable  contri- 
butions to  the  benthic  fauna  include  pipe- 
fishes (Syngnathidae),  flounder  (Bothi- 
dae),  sole  (Soleidae),  searobins  (Trigli- 
dae),    and    toadfishes    (Batrachoididae). 

Numerically  abundant  fishes  of  the 
mid  and  upper  waters  include  anchovies 
(Engraul idae),  herrings  (Clupeidae)  and 
needlefishes  (Belonidae).  At  all  loca- 
tions studied,  the  benthic  fauna  was  domi- 
nated by  the  pinfish,  Lagodon  rhomboides, 
the  silver  perch,  Bairdiella  chrysura,  the 
pigfish,  Orthopristis  chrysoptera.  and  the 
mojarras,  Euci  nostomus  gula  and  E. 
argenteus.  The  most  common  midwater  and 
surface  species  include  the  two  anchovies, 
Anchoa  mitchilli  and  A_.  hepsetus,  and  two 
clupeids,  Brevoortia  smithi  and  Harengula 
pensacol ae.  The  total  number  of  species 
recorded  in  the  individual  studies  ranged 
from  47  to  89;  a  total    of  117  species     was 


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

In  none  of  these  studies  were  the 
fishes  specifically  utilizing  the  fringing 
mangrove  habitat  enumerated  separately 
from  those  collected  in  the  bay  as  a 
whole.  The  collections  were  most  often  at 
open  water  stations  easily  sampled  by 
otter  trawl.  Carter  et  al.  (1973)  had  two 
shore  seine  stations  adjacent  to  mangroves 
but  the  data  were  pooled  for  publication. 
Of  the  four  stations  in  Rookery  Bay  sam- 
pled by  Yokel  (1975a),  one  was  immediately 
adjacent  to  the  fringing  mangrove  shore- 
line and  had  moderate  amounts  of  sea 
grasses. 

The  typical  pattern  which  emerges 
from  many  estuarine  studies  is  that  rela- 
tively few  fish  species  numerically  domi- 
nate the  catch.  This  is  certainly  true  in 
mangrove-fringed  estuaries.  In  Rookery 
Bay  (Yokel  1975a)  six  species  comprised 
88%  of  the  trawl -catchable  fishes,  in 
Fahkahatchee  Bay  seven  species  comprised 
97%  of  the  catch  from  three  capture 
techniques  (Carter  et  al.  1973),  and  in 
the  Marco  Island  estuary  25  species  com- 
prised 97%  of  the  trawl -catchable  fishes 
(Weinstein   et    al.    1977). 

Like  tidal  river  and  stream  communi- 
ties, these  shallow  bays  serve  as  nur- 
series for  numerous  species  of  estuarine- 
dependent  fishes  that  are  spawned  off- 
shore. Based  on  the  distribution  and 
abundance  of  juvenile  fishes  of  all  spe- 
cies in  six  habitats,  Carter  et  al.  (1973) 
ranked  the  mangrove-fringed  bays  as  the 
most  important  nursery  grounds;  the  tidal 
streams  were  a  close  second.  Shallow  bays 
and  tidal  streams  provide  safe  nurseries 
due  to  seasonally  abundant  food  resources 
and  the  low  frequency  of  large  predators 
(Carter  et  al.  1973;  Thayer  et  al.  1978). 
The  relative  lack  of  large  predaceous 
fishes  is  probably  due  to  their  general 
inability  to  osmoregulate  in  waters  of  low 
and/or    fluctuating    salinity. 

As  in  tidal  streams,  the  peak  abun- 
dance of  juvenile  and  larval  fishes  in  the 
bays  is  in  spring  and  early  summer  (Reid 
1954).      In  general,   the  highest  standing 


55 


crops  and  the  greatest  species  richness  of 
fishes  occur  in  the  late  summer  and  early 
fall  (Clark  1970).  Fish  densities  decline 
in  the  autumn  and  winter  as  many  fishes 
move  to  deeper  waters. 


7.4     FRINGING  FORESTS  ALONG  OCEANIC  BAYS 
AND   LAGOONS 

Mangrove-fringed  "oceanic"  bays  and 
lagoons  are  exemplified  by  Porpoise  Lake 
in  eastern  Florida  Bay  (Hudson  et  al. 
1970),  western  Florida  Bay  (Schmidt  1979), 
southern  Biscayne  Bay  (Bader  and  Roessler 
1971),  and  Old  Rhodes  Key  Lagoon  in 
eastern  Biscayne  Bay  (Holm  1977).  Charac- 
teristics of  these  sites  are  summarized  in 
Appendix  A.  Compared  to  the  mangrove- 
fringed  bays  discussed  in  the  previous 
section,  these  environments  generally  ex- 
hibit clearer  water,  sandier  substrates, 
and  higher  and  less  variable  salinities. 
Closer  proximity  to  the  Florida  reef 
tract,  the  Atlantic  Ocean,  and  the  Gulf  of 
Mexico  results  in  a  larger  potential  pool 
of  fish  species.  These  four  locations 
have  produced  reports  of  156  fish  species 
(Appendix  B). 

Mangrove  fringes  make  up  a  relatively 
small  proportion  of  these  environments; 
accordingly,  their  contribution  to  the  bay 
food  webs  is  probably  not  very  large. 
Bader  and  Roessler  (1972)  estimated  that 
the  fringing  mangrove  community  contrib- 
utes approximately  1%  of  the  total  energy 
budget  of  southern  Biscayne  Bay;  they 
considered  only  mainland  mangroves  and  did 
not  include  the  small  area  of  mangrove 
islands.  The  main  ecological  role  of  the 
fringing  mangroves  in  this  type  of  en- 
vironment is  probably  twofold.  First, 
they  increase  the  habitat  diversity  within 
an  otherwise  relatively  homogeneous  bay 
system.  Second,  they  provide  a  relatively 
protected  habitat  for  juvenile  fishes  (and 
certain  invertebrates)  that  later  move  to 
more  open  water  or  coral  reef  communities. 
The  second  role  is  analogous  to  one  of  the 
ecological  roles  of  sea  grass  communities 
(see  Zieman,  in  prep.)  although  the  fish 
species   involved   may  be  different. 


Based  primarily  on  habitat  designa- 
tions of  Voss  et  al.  (1969),  the  fishes  of 
Biscayne  Bay  can  be  characterized  as  to 
preferred  habitat.  Of  the  three  main 
habitat  types,  (1)  rock/coral/seawall,  (2) 
grassbed/tidal  flat,  and  (3)  mangrove,  the 
grassbed/tidal  flat  ranked  first  in  fish 
species  occurrences.  One  hundred  and 
twenty-two  of  156  species  (79%)  are  known 
to  occur  in  this  environment. 
Rock/coral /seawal  1  habitats  were  fre- 
quented by  49  species  (32%)  and  mangroves 
are  known  to  be  utilized  by  54  species 
(35%)  of  the  total  fish  species  recorded 
from  this   bay. 


7.5     OVERWASH   MANGROVE    ISLANDS 

In  terms  of  fish-related  research, 
these  communities  are  the  least  studied  of 
all  mangrove  community  types  in  south 
Florida.  They  are  typified  by  the  low- 
lying  mangrove-covered  islands  that  occur 
in  the  Florida  Keys  and  Florida  Bay  and 
may  be  overwashed  periodically  by  the 
tides.  Examples  include  Shell  Key,  Cotton 
Key,  and  the  Cowpens.  Islands  of  this 
type  extend  southwest  from  the  Florida 
mainland  through  the  Marquesas.  The  Dry 
Tortugas  lack  well -developed  mangrove  com- 
munities although  stunted  trees  are  found 
(Davis  1942). 

These  islands  are  the  most  oceanic  of 
any  of  the  mangrove  communities  discussed. 
They  are  characterized  by  relatively  clear 
water  (Gore  1977)  and  are  largely  free  of 
the  freshwater  inflow  and  salinity  varia- 
tions which  characterize  other  Florida 
mangrove  communities  to  varying  degrees. 
Numerous  statements  exist  in  the  litera- 
ture acknowledging  the  frequent  proximity 
of  mangrove  islands  to  coral  reefs  and  sea 
grass  beds  (McCoy  and  Heck  1976;  Thayer  et 
al.  1978).  Olsen  et  al.  (1973)  workinq  in 
the  U.S.  Virgin  Islands,  found  74%  to  93% 
overlap  in  the  fish  species  composition  of 
fringing  coral  reefs  and  shallow  mangrove- 
fringed  oceanic  bays.  Voss  et  al.  (1969) 
listed  fish  species  that  were  collected 
from  all  three  types  of  communities: 
fringing  mangroves,   coral    reefs  and  sea 


56 


grass  beds  in  Biscayne  Bay,  but  there 
appears  to  have  been  no  systematic  survey 
of  the  fish  assemblage  characteristic  of 
the  mangrove-covered  or  mangrove-fringed 
Florida  Keys.  No  one  has  quantified  the 
faunal  connections  which  we  hypothesize 
exist  between  the  mangroves  and  sea 
grasses  and  between  the  mangroves  and 
coral    reefs. 

In  the  absence  of  published  data  from 
the  mangrove  key  communities,  only  tenta- 
tive statements  can  be  made.  In  general, 
we  expect  that  while  mangrove  islands 
serve  as  a  nursery  area  for  juvenile 
fishes,  this  function  is  limited  largely 
to  coral  reef  and  marine  inshore  fishes 
and  not  the  estuarine-dependent  species 
that  we  have  discussed  previously.  The 
latter  (juvenile  snook,  red  drum,  spotted 
seatrout)  appear  to  require  relatively  low 
salinities  not  found  in  association  with 
most  of  the  overwash  islands.  Casual 
observation  around  the  edges  of  these 
islands  suggests  that  characteristic 
fishes  include  the  sea  bass  family  (Ser- 
ranidae),  t ri ggerf i shes  (Bal  istidae), 
snappers  (Lut  jam' dae),  grunts  (Poma- 
dasyidae),  porgies  (Span'dae)  parrotfishes 
(Scaridae),  wrasses  (Labridae),  bonefishes 
(Albulidae),  jacks  (Carangidae),  damsel- 
fishes  (Pomacentridae),  and  surgeonf ishes 
(Acanthuridae);  many  of  these  fishes  occur 
on  or  are  associated  with  coral  reefs.  We 
also  suspect  that  considerable  overlap 
occurs  in  the  fish  assemblage  of  these 
mangrove  islands  and  sea  grass  communi- 
ties; examples  include  puffers  (Tetrao- 
dontidae),  pipefishes  (Syngnathidae) ,  go- 
bies (Gobiidae)  and  scorpionf ishes  (Scor- 
paenidae).  Stark  and  Schroeder  (1971) 
suggested  that  juvenile  gray  snapper, 
which  use  the  fringing  mangroves  of  the 
keys  as  shelter  during  the  day,  forage  in 
adjacent  sea  grass  beds  at  night.  In  the 
absence  of  salinity  barriers,  predatory 
fishes  probably  enter  the  fringes  of  these 


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


7.6     GRADIENT  OF    MANGROVE   COMMUNITY 
INTERACTIONS 

Mangrove  communities  occur  under  a 
wide  range  of  conditions  from  virtually 
freshwater  at  the  headwaters  of  tidal 
streams  to  nearly  oceanic  conditions  in 
the  Florida  Keys.  Attempting  to  present  a 
single  list  of  fish  characteristic  of 
mangrove  environments  (Appendix  B)  can  be 
misleading.  For  this  reason  we  presented 
the  concept  of  a  continuum  or  complex 
gradient  in  Figure  11  and  have  followed 
that  scheme  throughout  section  7.  The 
gradient  stretches  from  seasonally  fresh 
to  oceanic  conditions,  from  highly  varia- 
ble salinities  to  nearly  constant  salini- 
ty, from  muddy  and  limestone  substrates  to 
sandy  substrates,  from  dark-stained  and 
sometimes  turbid  waters  to  clear  waters, 
and  from  food  webs  that  are  predominantly 
mangrove  detritus-based  to  food  webs  based 
primarily  on  other  energy  sources.  Clear- 
ly, there  are  other  gradients  as  one  moves 
from  north  to  south  in  the  State  of 
Florida.  At  the  northern  end  of  the 
State,  temperatures  are  more  variable  and 
seasonally  lower  than  in  the  south.  Sedi- 
ments change  from  predominantly  silicious 
in  central  and  north  Florida  to  predomi- 
nantly carbonate  in  extreme  south  Florida. 
Nevertheless,  the  complex  gradient  shown 
in  Figure  11,  while  greatly  simplified  for 
graphic  purposes,  suggests  that  charac- 
teristic fish  assemblages  replace  one 
another  along  a  gradient  of  changing 
physical  and  biogeographic  conditions. 
Such  a  concept  is  useful  in  understanding 
the  factors  controlling  the  composition  of 
fish  assemblages  associated  with  mangroves 
of  the  four  major  community  types  in  south 
Florida. 


57 


CHAPTER  8.   COMMUNITY  COMPONENTS  -  AMPHIBIANS  AND  REPTILES 


Food  habits  and  status  of  24  species 
of  turtles,  snakes,  lizards,  and  frogs  of 
the  Florida  mangrove  region  are  given  in 
Appendix  C.  Any  of  three  criteria  had  to 
be  met  before  a  species  was  included  in 
this  table:  (1)  a  direct  reference  in 
the  literature  to  mangrove  use  by  the 
species,  (2)  reference  to  a  species  as 
being  present  at  a  particular  geographical 
location  within  the  mangrove  zone  of 
Florida,  and  (3)  North  American  species 
recorded  from  mangroves  in  the  West  Indies 
or  South  America,  but  not  from  Florida. 
This  last  criterion  assumes  that  a  species 
which  can  utilize  mangroves  outside  of 
Florida  will  be  able  to  use  them  in 
Florida.  Ten  turtles  are  listed  of  which 
four  (striped  mud  turtle,  chicken  turtle, 
Florida  red-bellied  turtle,  and  softshell 
turtle)  are  typical  of  freshwater.  Two 
(mud  turtle  and  the  ornate  diamondback 
terrapin)  are  found  in  brackish  water  and 
the  remainder  (hawksbill,  green,  logger- 
head, and  Atlantic  ridley)  are  found  in 
marine  waters. 

Freshwater  species  usually  occur  in 
the  headwater  regions  of  mangrove-lined 
river  systems.  All  four  freshwater 
species  are  found  in  habitats  other  than 
mangrove  swamps  including  streams,  ponds, 
and  freshwater  marshes.  The  brackish 
water  species  are  found  in  salt  marshes  in 
addition  to  mangrove  swamps.  Mangroves, 
however,  are  the  principal  habitat  for  the 
ornate  diamondback  terrapin  (Ernst  and 
Barbour  1972).  Cam  and  Goi  n  (1955) 
listed  two  subspecies  of  the  diamondback: 
Malaclemys  terrapin  macrospilota  and  M.  t. 
rhizophorarum.  Malaclemys  terrapin  macro- 
spilota inhabits  the  southwest  and  south- 
ern coasts,  and  M.  ;t.  rhizophorarum  is 
found  in  the  Florida  Keys.  The  two  sub- 
species intergrade  in  the  region  of  north- 
ern Florida  Bay. 


All  four  of  the  marine  turtles  are 
associated  with  mangrove  vegetation  at 
some  stage  of  their  lives.  Loggerhead  and 
green  turtles  are  apparently  much  less 
dependent  on  mangroves  than  the  remaining 
two,  although  we  strongly  suspect  that 
recently  hatched  loggerheads  may  use  man- 
grove estuaries  as  nursery  areas.  Green 
turtles    are   generally   believed   to   feed   on 


a  variety  of  submerged  aquatic  plants  and 
sea  grasses;  recent  evidence  has  shown 
that  they  also  feed  on  mangrove  roots  and 
leaves  (Ernst  and  Barbour  1972).  The 
Atlantic  ridley's  preferred  habitat  is 
"shallow  coastal  waters,  especially  the 
mangrove-bordered  bays  of  the  southern 
half  of  the  peninsula  of  Florida"  (Carr 
and  Goin  1955).  Hawksbill  turtles  feed  on 
a  variety  of  plant  materials  including 
mangrove  (especially  red  mangrove), 
fruits,  leaves,  wood,  and  bark  (Ernst  and 
Barbour  1972). 

Three  species  in  the  genus  Anol i  s 
have  been  reported  from  Florida  mangroves: 
the  green  anole,  the  cuban  brown  anole, 
and  the  Bahaman  bank  anole.  All  are 
arboreal  lizards  that  feed  on  insects. 
The  green  anole  is  widespread  throughout 
the  Southeastern  United  States  and  is  not 
at  all  dependent  on  mangrove  swamps.  The 
other  two  species  have  much  more 
restricted  distributions  in  the  United 
States  and  are  found  only  in  south 
Florida.  They  also  are  not  restricted 
to  mangrove  ecosystems.  Of  the  six 
species  of  snakes  listed,  the  mangrove 
water  snake  (Figure  13)  is  most  dependent 
upon   mangrove  habitats. 

Two  important  species  of  reptiles 
found  in  mangrove  swamps  are  the  American 
alligator  and  the  American  crocodile.  The 
alligator  is  widespread  throughout  the 
Southeastern  United  States  and  is  only 
incidentally  found  in  low  salinity  sec- 
tions of  Florida  mangrove  areas  (Kushlan 
1980).  The  American  crocodile  is  rare; 
historically  its  distribution  was  centered 
in  the  mangrove-dominated  areas  of  the 
upper  and  lower  Florida  Keys  (particularly 
Key  Largo)  and  the  mangrove-lined  shore- 
lines and  mud  flats  along  the  northern 
edge  of  Florida  and  Whitewater  Bays 
(Kushlan  1980).  Mangroves  appear  to  be 
critical  habitat  for  this  species.  Its 
range  has  shrunk  considerably  in  south 
Florida  since  the  1930's,  even  though 
Florida  Bay  was  added  to  Everglades 
National  Park  in  1950  (Moore  1953;  Ogden 
1978).  Much  of  the  decrease  in  range  is 
due  to  increased  human  activity  in  the 
Florida  Keys.  The  remaining  population 
centers  of  the  American  crocodile  are  in 


Figure  13.  The  mangrove  water  snake,  Nerodja  fasciata  compressicauda,  curled  on 
a  red  mangrove  prop  root.  Photograph  by  David  Scott. 


59 


northern  Florida  Bay  and  adjacent  coastal 
swamps  and  the  northern  end  of  Key  Largo 
(Ogden  1978;  Kushlan  1980).  The  species 
uses  a  variety  of  habitats  for  nesting  in 
the  Florida  Bay  region  including  open 
hardwood  thickets  along  creek  banks, 
hardwood-shrub  thickets  at  the  heads  of 
sand-shell  beaches,  and  thickets  of  black 
mangroves  behind  marl  banks  (Ogden  1978). 
On  Key  Largo  the  crocodile  locates  its 
nests  on  creek  and  canal  banks  in  red  and 
black  mangrove  swamps  (Ogden  1978).  Man- 
grove areas  thus  appear  to  be  important  in 
the  breeding  biology  of  this  endangered 
species. 

Interestingly,    only   three    species    of 


amphibians,  to  our  knowledge,  have  been 
recorded  in  Florida  mangrove  swamps  (Ap- 
pendix C).  This  is  due  to  two  factors: 
(1)  lack  of  detailed  surveys  in  low  sa- 
linity swamps  and  (2)  the  inability  of 
most  amphibians  to  osmoregulate  in  salt 
water.  No  doubt,  several  additional 
species  occur  in  the  freshwater -dominated 
hammock  and  basin  mangrove  communities 
inland  from  the  coast.  Possible  addi- 
tional species  include:  the  eastern 
narrow-mouthed  toad,  Gastrophryne  caro- 
1  in en  si  s,  the  eastern  spadefoot  toad, 
Scaphiopus  holbrooki,  the  cricket  frog, 
Acris  gryllus,  the  green  tree  frog,  Hyl a 
cinerea,  and  the  southern  leopard  frog, 
Rana  utricularia. 


60 


CHAPTER    9.       COMMUNITY    COMPONENTS    -    BIRDS 


9.1      ECOLOGICAL    RELATIONSHIPS 

Because  mangroves  present  a  more 
diverse  structural  habitat  than  most 
coastal  ecosystems,  they  should  harbor  a 
greater  variety  of  birdlife  than  areas 
such  as  salt  marshes,  mud  flats,  and 
beaches  (MacArthur  and  MacArthur  1961). 
The  shallow  water  and  exposed  sediments 
below  mangroves  are  available  for  probing 
shorebirds.  Longer-legged  wading  birds 
utilize  these  shallow  areas  as  well  as 
deeper  waters  along  mangrove-lined  pools 
and  waterways.  Surface-feeding  and  diving 
birds  would  be  expected  in  similar  areas 
as  the  wading  birds.  The  major  difference 
between  mangrove  swamps  and  other  coastal 
ecosystems  is  the  availability  of  the 
trunks,  limbs,  and  foliage  comprising  the 
tree  canopy.  This  enables  a  variety  of 
passerine  and  non-passerine  birds,  which 
are  not  found  commonly  in  other  wetland 
areas,  to  use  mangrove  swamps.  It  also 
allows  extensive  breeding  activity  by  a 
number  of  tree-nesting  birds. 

The  composition  of  the  avifauna  com- 
munity in  mangrove  ecosystems  is,  in  fact, 
highly  diverse.  Cawkell  (1964)  recorded 
45  species  from  the  mangroves  of  Gambia 
(Africa).  Haverschmidt  (1965)  reported  87 
species  of  birds  which  utilized  mangroves 
in  Surinam  (S.  America).  Ffrench  (1966) 
listed  94  species  from  the  Caroni  mangrove 
swamp  in  Trinidad  while  Bacon  (1970)  found 
137  in  the  same  swamp.  In  Malaya,  Nisbet 
(1968)  reported  121  species  in  mangrove 
swamps  and  Field  (1968)  observed  76  from 
the   mangroves  of  Sierra   Leone   (Africa). 

Use  of  mangrove  ecosystems  by  birds 
in  Florida  has  not  been  recorded  in  de- 
tail. Ninety -two  species  have  been  ob- 
served in  the  mangrove  habitat  of  Sanibel 
Island,  Florida  (L.  Narcisse,  J.N.  "Ding" 
Darling  Natl.  Wildlife  Refuge,  Sanibel 
Is.,  Fla.;  personal  communication  1981). 
Robertson  (1955)  and  Robertson  and  Kushlan 
(1974)  reported  on  the  entire  breeding 
bird  fauna  of  peninsular  south  Florida, 
including  mangrove  regions.  Based  on 
limited  surveys,  these  authors  reported 
only  17  species  as  utilizing  mangroves  for 
breeding  purposes.  Because  their  studies 
did   not   consider   migrants   or   non-breeding 


residents,  a  significant  fraction  of  the 
avifauna  community  was  omitted. 

Based  on  information  gleaned  from  the 
literature,  we  have  compiled  a  list  of  181 
species  of  birds  that  use  Florida  mangrove 
areas  for  feeding,  nesting,  roosting,  or 
other  activities  (Appendix  D).  Criteria 
for  listing  these  species  is  the  same  as 
that  used  for  listing  reptiles  and  amphi- 
bians  (see  Chapter  8  of  this   volume). 

Often  references  were  found  stating 
that  a  given  species  in  Florida  occurred 
in  "wet  coastal  hammocks",  "coastal  wet 
forests"  or  the  like,  without  a  specific 
reference  to  mangroves.  These  species 
were  not  included  in  Appendix  D.  Thus, 
this  list  is  a  conservative  estimate  of 
the  avifauna  associated  with  Florida  man- 
grove swamps.  Sources  for  each  listing 
are  provided  even  though  many  are  redun- 
dant. Food  habit  data  are  based  on  Howell 
(1932)  and  Martin  et  al.  (1951).  Esti- 
mates of  abundance  were  derived  from  bird 
lists  published  by  the  U.S.  Fish  and 
Wildlife  Service  for  the  J.N.  "Ding" 
Darling  National  Wildlife  Refuge  at 
Sanibel  Island,  Florida,  and  by  the  Ever- 
glades Natural  History  Association  for 
Everglades  National  Park.  Frequently, 
species  were  recorded  from  mangrove  swamps 
at   one   location,    but  not  the  other. 

We  have  divided  the  mangrove  avifauna 
into  six  groups  based  on  similarities  in 
methods  of  procuring  food.  These  groups 
(guilds)  are  the  wading  birds,  probing 
shorebirds,  floating  and  diving  water- 
birds,  aerially-searching  birds,  birds  of 
prey,  and  arboreal  birds.  This  last  group 
is  something  of  a  catch-all  group,  but  is 
composed  mainly  of  birds  that  feed  and/or 
nest   in  the  mangrove  canopy. 


9.2     WADING   BIRDS 

Herons,  egrets,  ibises,  bitterns,  and 
spoonbills  are  the  most  conspicuous  group 
of  birds  found  in  mangroves  (Figure  14) 
and  are  by  far  the  most  studied  and  best 
understood.  Eighteen  species  (and  one 
important  subspecies)  are  reported  from 
south   Florida   mangroves. 


61 


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


62 


Mangrove  swamps  provide  two  functions 
for  wading  birds.  First,  they  function  as 
feeding  grounds.  Two-thirds  of  these 
species  feed  almost  exclusively  on  fishes. 
Although  much  of  their  diet  is  provided  by 
freshwater  and  non-mangrove  marine  areas, 
all  of  them  feed  frequently  in  mangrove 
swamps.  White  ibis  feed  predominantly  on 
crabs  of  the  genus  Uca  when  feeding  in 
mangroves  (Kushlan  and  Kushlan  1975; 
Kushlan  1979).  Mollusks  and  invertebrates 
of  the  sediments  are  principal  foods  of 
the  roseate  spoonbill  although  some  fish 
are  eaten  (Allen  1942).  Yellow-crowned 
night  herons  and  American  bitterns  eat 
crabs,  crayfish,  frogs,  and  mice  in  addi- 
tion to  fishes.  Snails  of  the  genus 
Pomacea  are  fed  upon  almost  exclusively  by 
the  limpkin.  The  sandhill  crane  is  an 
anomaly  in  this  group  since  a  majority  of 
its  food  is  vegetable  matter,  especially 
roots  and  rhizomes  of  Cyperus  and 
Sagittaria.  Its  use  of  mangroves  is 
probably  minimal,  occurring  where  inland 
coastal  marshes  adjoin  mangroves  (Kushlan, 
unpubl .  data).  The  remaining  12  species 
are  essentially  piscivorous  although  they 
differ  somewhat  in  the  species  and  sizes 
of  fishes  that  they  consume. 

Mangrove  swamps  also  serve  as 
breeding  habitat  for  wading  birds.  With 
the  exception  of  the  limpkin,  sandhill 
crane,  and  the  two  bitterns,  all  wading 
bird  species  in  Appendix  D  build  their 
nests  in  all  three  species  of  mangrove 
trees  (Maxwell  and  Kale  1977;  Girard  and 
Taylor  1979).  The  species  often  aggregate 
in  large  breeding  colonies  with  several 
thousand  nesting  pairs  (Kushlan  and  White 
1977a).  The  Louisiana  heron,  snowy  egret, 
and  cattle  egret  are  the  most  numerous 
breeders  in  south  Florida  mangroves  (based 
on   data   in   Kushlan   and   White  1977a). 

In  wet  years  over  90%  of  the  south 
Florida  population  of  white  ibis  breed  in 
the  interior,  freshwater  wetlands  of  the 
Everglades;  during  these  times  the  man- 
groves are  apparently  unimportant,  sup- 
porting less  than  10%  of  the  population 
(Kushlan  1976,  1977a,  b).  During  drought 
years,  however,  production  is  sustained 
solely  by  breeding  colonies  located  in 
mangroves  near  the  coast  (Kushlan  1977a, 


b).  Mangroves  are  critically  important 
for  the  survival  of  the  white  ibis  popula- 
tion even  though  they  appear  to  be 
utilized  to  a  lesser  extent  than  fresh- 
water habitats.  This  pattern  of  larger 
but  less  stable  breeding  colonies  using 
inland  marshes  and  smaller  but  more  stable 
colonies  using  mangroves  is  also  charac- 
teristic of  heron  populations  (Kushlan  and 
Frohring,    in    prep.). 

Table  5  gives  the  number  of  active 
nests  observed  in  mangrove  regions  during 
the  1974-75  nesting  season  and  the  percen- 
tage this  represents  of  the  entire  south 
Florida  breeding  population  for  the  nine 
most  abundant  species  of  waders  and  three 
associated  species.  The  dependence  of 
roseate  spoonbills,  great  blue  herons, 
Louisiana  herons,  brown  pelicans,  and 
double-crested  cormorants  on  mangrove 
regions  is  evident.  Nesting  by  the  red- 
dish egret  was  not  quantified  during  this 
study  although  Kushlan  and  White  (1977a) 
indicated  that  the  only  nests  of  this 
species  which  they  saw  were,  in  fact,  in 
mangroves.  Further  observations  indicate 
that  this  species  nests  in  mangroves  ex- 
clusively (Kushlan,  pers.  comm.).  Similar- 
ly, the  great  white  heron  is  highly  depen- 
dent upon  mangroves  for  nesting;  they  use 
the  tiny  mangrove  islets  which  abound 
along  the  Florida  Keys  and  in  Florida  Bay 
(Howell    1932). 

During  many  years  the  Everglades 
population  of  wood  storks  is  known  to  nest 
almost  solely  in  mangroves  (Ogden  et  al. 
1976);  this  population  comprises  approxi- 
mately one-third  of  the  total  south 
Florida  population.  Successful  breeding 
of  all  these  mangrove  nesters  is  un- 
doubtedly correlated  with  the  abundant 
supply  of  fishes  associated  with  man- 
groves. Meeting  the  energetic  demands  of 
growing  young  is  somewhat  easier  in  habi- 
tats with  abundant  prey.  This  is 
especially  important  for  the  wood  stork 
which  requires  that  its  prey  be  concen- 
trated into  small  pools  by  falling  water 
levels  during  the  dry  season  before  it  can 
nest  successfully  (Kahl  1964;  Kushlan  et 
al.  1975;  Odgen  et  al.  1978).  Breeding 
activity  by  wading  birds  in  mangroves 
along  the  southwest   and   southern   Florida 


63 


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


Species 


%  of 

total  active 

Active  nests  in 

nests  in  south 

mangroves 

Florida 

1914 

7 

500 

100 

1335 

31 

458 

92 

1812 

39 

2377 

46 

71 

15 

3410 

70 

2180 

13 

741 

100 

White  ibis 

Roseate  spoonbill 

Wood  stork 

Great  blue  heron 

Great  egret 

Snowy  egret 

Little  blue  heron 

Louisiana   heron 

Cattle  egret 

Brown   pelican 

Double-crested 
cormorant  1744  83 


64 


coasts  takes  place  throughout  the  year 
(Table  6);  at  least  one  species  of  wader 
breeds  during  every  month.  Colonies  on 
the  mangrove  islands  in  Florida  Bay  were 
noted  to  be  active  nesting  sites  during 
all  months  of  the  year  except  September 
and   October   (Kushlan   and   White   1977a). 

The  seasonal  movements  of  wood  storks 
and  white  ibises  between  the  various  south 
Florida  ecosystems  were  described  by 
Ogden  et  al.  (1978)  and  Kushlan  (1979). 
Mangrove  ecosystems  appear  to  be  most 
heavily  used  for  feeding  in  summer  (white 
ibis)  and  early  winter  (white  ibis  and 
wood  stork).  The  remaining  species  of 
wading  birds  appear  to  use  mangrove  areas 
most  heavily  in  the  winter  months,  reflec- 
ting the  influx  of  migrants  from  farther 
north. 

Wading  birds  play  an  important  role 
in  nutrient  cycling  in  the  coastal  man- 
grove zone.  Mclvor  (pers.  observ.)  has 
noted  increased  turbidity,  greater  algal 
biomass,  and  decreased  fish  abundance 
around  red  mangrove  islets  with  nesting 
frigate  birds  and  cormorants.  Onuf  et  al. 
(1977)  reported  results  from  a  small  (100 
bird)  rookery  on  a  mangrove  islet  on  the 
east  coast  of  Florida.  Additions  of 
ammonium-nitrogen  from  the  bird's 
droppings  exceeded  1  g/m  /day.  Water 
beneath  the  mangroves  contained  five  times 
more  ammonium  and  phosphate  than  water 
beneath  mangroves  without  rookeries. 
Although  the  wading  birds  were  shown  to  be 
a  vector  for  concentrating  nutrients,  it 
must  be  noted  that  this  is  a  localized 
phenomenon  restricted  to  the  areas  around 
rookeries  in  the  mangrove  zone.  The 
effect  would  be  larger  around  larger 
rookeries.  Onuf  et  al.  (1977)  also 
reported  that  mangroves  in  the  area  of  the 
rookery  had  increased  levels  of  primary 
production,  higher  stem  and  foliar  nitro- 
gen levels,  and  higher  herbivore  grazing 
impact  than  mangroves  without  rookeries. 
Lewis  and  Lewis  (1978)  stated  that  man- 
groves in  large  rookeries  may  eventually 
be  killed  due  to  stripping  of  leaves  and 
branches  for  nesting  material  and  by 
poisoning  due  to  large  volumes  of  urea  and 
ammonia  that  are  deposited  in  bird  guano. 
This     latter     effect     would     be     more 


pronounced  in  rookeries  within  mangrove 
regions  subject  to  infrequent  tidal  flush- 
ing. 

9.3      PROBING   SHOREBIRDS 

Birds  in  this  group  are  commonly 
found  associated  with  intertidal  and  shal- 
low water  habitats.  Wolff  (1969)  and 
Schneider  (1978)  have  shown  that  plovers 
and  sandpipers  are  opportunistic  feeders, 
taking  the  most  abundant,  proper-sized 
invertebrates  present  in  whatever  habitat 
the  birds   happen  to  occupy. 

Of  the  25  species  included  in  this 
guild  (Appendix  D),  two  are  year-round 
residents  (clapper  rail  and  willet),  two 
breed  in  mangrove  areas  (clapper  rail  and 
black-necked  stilt),  and  the  remainder  are 
transients  or  winter  residents.  Baker  and 
Baker  (1973)  indicated  that  winter  was  the 
most  crucial  time  for  shorebirds,  in  terms 
of  survival.  Coi ncidental ly,  winter  is 
the  time  when  most  shorebirds  use  mangrove 
areas.  The  invertebrate  fauna  (mollusks, 
crustaceans,  and  aquatic  insects)  which 
occur  on  the  sediments  under  intertidal 
mangroves  forms  the  principal  diet  of 
these  species.  Willets  and  greater 
yellowlegs  eat  a  large  amount  of  fishes, 
especially  Fundulus,  in  addition  to  inver- 
tebrates. Many  of  the  species  listed  in 
this  guild  obtain  a  significant  portion  of 
their  energy  requirements  from  other  habi- 
tats, particularly  sandy  beaches,  marshes, 
and  freshwater  prairies.  Of  the  species 
in  this  guild,  the  clapper  rail  is  prob- 
ably most  dependent  on  mangroves  for 
survival  in  south  Florida  (Robertson 
1955),  although  in  other  geographical 
locations  they  frequent  salt  and  brackish 
marshes. 


9.4     FLOATING   AND  DIVING  WATER  BIRDS 

Twenty-nine  species  of  ducks,  grebes, 
loons,  cormorants,  and  gallinules  were 
identified  as  populating  mangrove  areas  in 
south  Florida  (Appendix  D).  Eight  species 
are  year-round  residents  while  the 
remainder  are  present  only  during  migra- 
tion or  as   winter  visitors. 


65 


Table  6.  Timing  of  nesting  by  wading  birds  and  associated 
species  in  south  Florida.  Adapted  from  data  in  Kushlan  and 
White  (1977a),  Kushlan  and  McEwan  (in  press). 


White  ibis 

Wood  stork 

Roseate  spoonbill 

Great  blue/white 
heron 

Great  egret 

Little  blue  heron 

Cattle  egret 

Double-crested 
cormorant 

Brown  pelican 


Months 

Species  SONDJFMAMJJA 


66 


From  the  standpoint  of  feeding,  mem- 
bers of  this  guild  are  highly  hetero- 
geneous. Piscivorous  species  include  the 
cormorant,  anhinga,  pelicans,  and  mergan- 
sers. Herbivorous  species  include  the 
pintail,  mallard,  wigeon,  mottled  duck, 
and  teals.  A  third  group  feeds  primarily 
on  benthic  mollusks  and  invertebrates. 
Scaup,  canvasback,  redhead,  and  gallinules 
belong  to  this  group.  The  ducks  in  this 
last  group  also  consume  a  significant 
fraction  of  plant  material. 

Species  of  this  guild  are  permanent 
residents  and  usually  breed  in  mangrove 
swamps.  As  shown  in  Table  5,  the  brown 
pelican  and  double-crested  cormorant  are 
highly  dependent  upon  mangroves  for 
nesting  in  south  Florida  even  though  both 
will  build  nests  in  any  available  tree  in 
other  geographical  regions.  It  seems  that 
when  mangroves  are  available,  they  are  the 
preferred  nesting  site.  The  anhinga 
breeds  in  mangrove  regions  but  is  more 
commonly  found  inland  near  freshwater  (J. 
A.  Kushlan,  So.  Fla.  Res.  Ctr.,  Everglades 
Natl.  Park,  Homestead,  Fla.;  personal 
communication  1981).  For  the  other  species 
listed  in  this  guild,  mangrove  swamps 
provide  a  common  but  not  a  required  habi- 
tat; all  of  these  species  utilize  a 
variety  of  aquatic  environments. 

Kushlan  et  al.  (in  prep.)  provide 
recent  data  on  the  abundance  and  distribu- 
tion of  22  species  of  waterfowl  and  the 
American  coot  in  south  Florida  estuaries. 
The  American  coot  is  by  far  the  most  abun- 
dant species,  accounting  for  just  over  50% 
of  the  total  population.  Six  species  of 
ducks  were  responsible  for  more  than  99% 
of  the  individuals  seen:  blue-winged  teal 
(41%),  lesser  scaup  (24%),  pintail  (18%), 
American  wigeon  (9%),  ring-necked  duck 
(5%),  and  shoveler  (3%).  The  major  habi- 
tats included  in  these  authors'  surveys 
were  coastal  prairie  and  marshes,  mangrove 
forests,  and  mangrove-lined  bays  and 
waterways  of  the  Everglades   National    Park. 

From  these  data  it  appears  that 
waterfowl  and  coots  are  most  abundant  in 
regions  where  mangrove,  wet  coastal 
prairies,  marshes,  and  open  water  are 
interspersed.       Overall,  the    Everglades 


estuaries  support  from  5%  to  10%  of  the 
total  wi nteri ng  waterfowl  population  in 
Florida  (Goodwin  1979;  Kushlan  et  al.  in 
prep.).  As  Kushlan  et  al.  point  out, 
however,  the  Everglades  are  not  managed 
for  single  species  or  groups  of  species  as 
are  areas  of  Florida  supporting  larger 
waterfowl  populations.  Although  the 
importance  of  south  Florida's  mangrove 
estuaries  to  continental  waterfowl  popula- 
tions may  be  small,  the  effect  of  70,000 
ducks  and  coots  on  these  estuaries 
probably  is  not  (Kushlan  et  al.  in  prep.). 

Kushlan  (personal  communication) 
thinks  that  the  estuaries  of  the  Ever- 
glades have  an  important  survival  value 
for  some  segments  of  the  American  white 
pelican  population.  In  winter,  approxi- 
mately 25%  of  the  white  pelicans  are  found 
in  Florida  Bay  and  75%  in  the  Cape  Sable 
region.  They  feed  primarily  in  freshwater 
regions  of  coastal  marshes  and  prairies 
and  use  mangroves  where  they  adjoin  this 
type   of   habitat. 


9.5     AERIALLY-SEARCHING   BIRDS 

Gulls,  terns,  the  kingfisher,  the 
black  skimmer,  and  the  fish  crow  comprise 
this  guild  of  omnivorous  and  piscivorous 
species  (Appendix  D).  These  birds  hunt  in 
ponds,  creeks,  and  waterways  adjacent  to 
mangrove  stands.  Many  fishes  and  inverte- 
brates upon  which  they  feed  come  from 
mangrove-based  food  webs.  Only  six  of  the 
14  species  are  year-round  residents  of 
south  Florida.  The  least  tern  is  an  abun- 
dant summer  resident  and  the  remainder  are 
winter   residents  or  transients. 

Only  the  fish  crow  actually  nests  in 
mangroves.  Gulls  and  terns  prefer  open 
sandy  areas  for  nesting  (Kushlan  and  White 
1977b)  and  use  mangrove  ecosystems  only 
for  feeding.  All  of  the  species  in  this 
guild  are  recorded  from  a  variety  of 
coastal    and  inland  wetland   habitats. 


9.6     BIRDS   OF   PREY 

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


67 


which   utilize  mangrove   swamps    in   south  are   common    inhabitants   of  mangrove   areas. 

Florida    (Appendix    D).      The    magnificant  This  could   also  be  true   for  the   merlin, 

frigatebird    has    been    included    in    this  which    like  the   peregrine   falcon,    feeds  on 

group  because  of  its  habit  of  robbing  many  waterfowl   and  shorebirds.     The   remaining 

of  these  birds  of  their  prey.     Prey  con-  species    in   this    guild   are   probably  not   so 

sumed    by    this    guild    includes    snakes,  dependent   on   mangroves;    although   they  may 

lizards,    frogs    (red-shouldered    hawk,  be   common   in   mangrove  ecosystems,    they 

swallow-tailed   kite),    small    birds    (short-  utilize  other  habitats   as  well, 
tailed  hawk),   waterfowl    (peregrine  falcon, 
great-horned  owl),   fishes   (osprey,   bald 

eagle),     and    carrion    (black    and    turkey  9.7     ARBOREAL    BIRDS 
vultures). 

This     guild     is     the     largest     (71 

Eleven  of  these  species  are  permanent  species)  and  most  diverse  group   inhabiting 

residents,   one  a   summer   resident,   and  the  mangrove  forests.     Included  are  pigeons, 

remainder  are  winter  residents.     Their  use  cuckoos,     woodpeckers,      flycatchers, 

of   mangrove   areas   varies    greatly.      The  thrushes,   vireos,  warblers,   blackbirds, 

magnificent    frigatebird,     which    occurs  and  sparrows.     We  have  lumped  this  diverse 

principally  in  extreme  southern  Florida  group  together  because  they  utilize  man- 

and  the  Florida  Keys,  utilizes  small   over-  grove  ecosystems   in   remarkably   similar 

wash   mangrove   islands   for  both   roosts   and  ways.        Invertebrates,      particularly 

nesting  colonies.     Both   species   of  vul-  insects,    make   up  a   significant   portion    of 

tures    are    widely    distributed    in    south  most  of  these  birds'   diets,  although  the 

Florida   mangrove   regions;  large  colonial  white-crowned    pigeon,    mourning    dove,    and 

roosts   can   be   found    in   mangrove    swamps  many   of   the    fringilids    (cardinal,    townee) 

near  the   coast.     Swallow-tailed   kites   are  eat    a    variety    of    seeds,     berries,     and 

common  over  the  entire  Florida  mangrove  fruits, 
region    (Robertson    1955;    Snyder    1974). 

Snyder  (1974)  reports  extensively  on  the  As  the  name  given  this  guild  implies, 

breeding   biology  of   the  swallow-tailed  these  birds  use  the  habitat  provided  by 

kites    in    south    Florida.      The    nests    he  the   mangrove  canopy.      Many  birds  also  use 

observed  were  all    located  in  black  man-  the  trunk,    branches,    and  aerial    roots   for 

groves    although    they    do    nest    in    other  feeding.       Several    different    types    of 

habitats.  searching  patterns  are  used.     Hawking  of 

insects   is   the  primary  mode  of   feeding  by 

The   bald  eagle,   osprey  (Figure  15),  the   cuckoos,    chuck-wi  1 1  s-wi  dows,  the 

and  peregrine  falcon  are  dependent   upon  kingbirds,    and    the    flycatchers.      Gleaning 

mangrove  ecosystems  for  their  continued  is    employed    by    most    of    the    warblers, 

existence  in  south  Florida.     Both  the   bald  Woodpeckers   and   the   prothonotary  warbler 

eagle  and  osprey  feed  extensively  on  the  are  classic   probers, 
wealth   of   fishes    found   associated   with 

mangrove  ecosystems.  Additionally,  man-  Several  of  the  birds  in  this  guild 
groves  are  used  as  roosts  and  support  are  heavily  dependent  upon  mangrove  areas, 
structures  for  nests.  Nisbet  (1968)  indi-  The  prairie  warbler  and  the  yellow  warbler 
cated  that  in  Malaysia  the  most  important  are  subspecies  of  more  widespread  North 
role  of  mangroves  for  birds  may  be  as  American  species  (see  Appendix  D  for 
wintering  habitat  for  palaearctic  mi-  scientific  names).  They  are  found  largely 
grants,  of  which  the  peregrine  falcon  is  within  mangrove  areas  (Robertson  and 
one.  Kushlan  (pers.  comm.)  stated  that  Kushlan  1974).  The  white-crowned  pigeon, 
recent  surveys  have  shown  falcons  to  mangrove  cuckoo,  gray  kingbird,  and  black- 
winter  in  mangroves,  particularly  along  whiskered  vireo  are  of  recent  West  Indian 
the  shore  of  Florida  Bay  where  they  estab-  origin.  They  first  moved  into  the 
lish  feeding  territories.  They  forage  on  mangrove-covered  regions  of  south  Florida 
concentrations  of  shorebirds  and  water-  from  source  areas  in  the  islands  of  the 
fowl.    These  prey   species   of   the   peregrine  Caribbean.      Confined   at    first    to   mangrove 

68 


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


69 


swamps,  all  but  the  mangrove  cuckoo  have 
expanded  their  range  in  peninsular  Florida 
by  using  non-mangrove  habitat.  In  this 
vein  it  is  interesting  to  note  that  many 
species  of  rare  and/or  irregular  occur- 
rence in  south  Florida  are  of  West  Indian 
origin  and  use  mangroves  to  a  considerable 
extent.  These  include  the  Bahama  pintail, 
masked  duck,  Caribbean  coot,  loggerhead 
kingbird,  thick-billed  vireo,  and  stripe- 
headed  tanager  (Robertson  and  Kushlan 
1974). 

Twenty-four  of  the  species  in  this 
guild  are  permanent  residents,  27  are  win- 
ter, and  6  are  summer  residents.  Fourteen 
species  are  seen  only  during   migrations. 


9.8       ASSOCIATIONS     BETWEEN 
COMMUNITY  TYPES  AND  BIRDS 


MANGROVE 


Estimating  the  degree  of  use  of 
mangrove  swamps  by  birds  as  we  have  done 
(Appendix  D)  is  open  to  criticism  because 
of  the  paucity  of  information  upon  which 
to  base  judgements.  Estimating  which 
mangrove  community  types  (see  section  1, 
Figure  4)  are  used  by  which  birds  is  open 
to  even  more  severe  criticism.  For  this 
reason  the  following  comments  should  be 
regarded  as  general    and   preliminary. 

In  terms  of  utilization  by  avifauna, 
the  scrub  mangrove  swamps  are  probably  the 
least  utilized  mangrove  community  type. 
Because  the  canopy  is  poorly  developed, 
most  of  the  arboreal  species  are  absent, 
although  Emlen  (1977)  recorded  the  red- 
winged  blackbird,  hairy  woodpecker,  north- 
ern waterthrush,  yellow-rumped  warbler, 
common  yel 1 owth roat ,  orange-crowned 
warbler,  palm  warbler,  yellow  warbler, 
mourning  dove,  and  gray  kingbird  in  scrub 
mangroves  on  Grand  Bahama  Island.  Of  25 
different  habitats  surveyed  by  Emlen 
(1977),  the  yellow  warbler  and  gray 
kingbird  were  found  in  the  scrub  mangroves 
only.  Aerially-searching  and  wading  birds 
might  use  scrub  mangroves  if  fishes  are 
present. 

Overwash  mangrove  islands  are 
utilized  in  a  variety  of  ways  by  all  of 
the  bird  guilds.     Most  of  the  wading  birds 


plus  the  magnificent  f ri gatebi rd,  the 
anhinga,  the  cormorant,  and  the  brown 
pelican  use  overwash  islands  for  nesting 
(Kushlan  and  White  1977a).  Wading  and 
aerially-searching  birds  commonly  feed  in 
close  proximity  to  overwash  islands.  A 
variety  of  migrating  arboreal  and  probing 
species  use  the  islands  for  feeding  and 
roosting.  Yellow  and  palm  warblers  are 
common  around  mangrove  islands  in  Florida 
Bay  as  are  the  black-bellied  plover,  ruddy 
turnstone,  willet,  dunlin,  and  short- 
billed  dowitcher.  Rafts  of  ducks  are 
common  near  the  inshore  islands  and  birds 
of  prey  such  as  the  osprey,  the  bald 
eagle,  and  both  vultures  use  mangrove 
islands   for   roosting  and   nesting. 

Fringe  and  riverine  mangrove  com- 
munities are  important  feeding  areas  for 
wading  and  probing  birds.  Floating  and 
diving  and  aerially-searching  birds  use 
the  lakes  and  waterways  adjacent  to  these 
mangrove  communities  for  feeding.  Many  of 
the  wading  birds  nest  in  fringe  and 
riverine  forests.  For  example,  when  the 
wood  ibis  nests  in  coastal  areas,  it  uses 
these  mangrove  communities  almost  exclu- 
sively (Kushlan,  personal  communication). 
Most  of  the  arboreal  birds  and  birds  of 
prey  associated  with  mangroves  are  found 
in  these  two  types  of  communities.  This 
is  not  surprising  since  the  tree  canopy  is 
extremely  wel 1 -devel oped  and  offers 
roosting,  feeding  and  nesting  opportuni- 
ties. 

Hammock  and  basin  mangrove  communi- 
ties are  so  diverse  in  size,  location,  and 
proximity  to  other  communities  that  it  is 
difficult  to  make  many  general  statements 
about  their  avifauna.  Since  there  often 
is  little  standing  water  in  hammock 
forests,  wading  and  diving  birds  probably 
are  not  common.  Proximity  to  terrestrial 
communities  in  some  cases  may  increase  the 
diversity  of  arboreal  species  in  both 
hammock  and  basin  forests;  proximity  to 
open  areas  may  increase  the  likelihood  of 
bi  rds    of   prey. 

It  seems  safe  to  conclude  that  each 
of  the  six  mangrove  community  types  has 
some  value  to  the  avifauna.  This  value 
differs  according  to  community  type  and 


70 


kind  of  bird  group  under  consideration. 
Certainly,  more  information  is  needed, 
particularly  concerning  the  dependence  of 
rare  or  endangered  species  on  specific 
community  types. 


9.9     MANGROVES  AS  WINTER  HABITAT  FOR  NORTH 
AMERICAN  MIGRANT  LAND  BIRDS 

An  interesting  observation  based  on 
the  data  in  this  chapter  is  the  seemingly 
important  role  that  mangrove  ecosystems 
play  in  providing  wintering  habitat  for 
migrants  of  North  American  origin.  Lack 
and  Lack  (1972)  studied  the  wintering 
warbler  community  in  Jamaica.  In  four 
natural  habitats  including  mangrove 
forest,  lowland  dry  limestone  forest,  mid- 
level  wet  limestone  forest,  and  montane 
cloud  forest, a  total  of  174,  131,  61,  and 
49  warblers  (individuals)  were  seen, 
respectively.  When  computed  on  a  per  hour 
of  observation  basis,  the  difference  is 
more  striking  with  22  warblers  per  hour 
seen  in  mangroves  and  only  1,  2,  and  1 
seen  in  the  other  forest  habitats,  respec- 
tively. For  all  passerines  considered 
together,  26  passerines/hour  were  seen  in 
mangroves  with  5,  13,  and  3  respectively 
in    the    other    forest    habitats.      On    a 


species  basis  only  9  were  recorded  from 
mangroves  whereas  19,  13,  and  16  species, 
respectively,  were  seen  in  the  other  habi- 
tats. This  large  number  of  species  from 
the  other  habitats  appears  to  result  from 
the  sighting  of  rare  species  after  many 
hours  of  observation.  Only  9  hours  were 
spent  by  Lack  and  Lack  (1972)  in  the  man- 
groves whereas  between  30  and  86  hours 
were  spent  in  other  habitats.  More  time 
in  the  mangrove  zone  would  have  undoubted- 
ly resulted  in  more  species  (and  in- 
dividuals)  observed  (Preston  1979). 

Hutto  (1980)  presented  extensive  data 
concerning  the  composition  of  migratory 
land  bird  communities  in  Mexico  in  winter 
for  13  habitat  types.  Mangrove  areas 
tended  to  have  more  migrant  species  than 
most  natural  habitats  (except  gallery 
forests)  and  also  had  a  greater  density  of 
individuals  than  other  habitats  (again 
except  for  gallery  forests).  In  both  Lack 
and  Lack's  and  Hutto's  studies,  disturbed 
and  edge  habitats  had  the  highest  number 
of  species  and  greatest  density  of 
individuals.  The  percentage  of  the 
avifauna  community  composed  of  migrants 
was  highest  in  mangrove  habitats,  however. 
From  this  we  can  infer  the  importance  of 
mangroves  in  the  maintenance  of  North 
American  migrant   land  birds. 


71 


CHAPTER  10.   COMMUNITY  COMPONENTS  -  MAMMALS 

Thirty-six  native  and  nine  introduced  (Layne  1974).  Hamilton  and  Whittaker 
species  of  land  mammals  occur  in  the  south  (1979)  state  that  it  is  the  coastal  ham- 
Florida  region  (Layne  1974;  Hamilton  and  mocks  of  south  Florida,  including  mangrove 
Whittaker  1979).  Of  these,  almost  50%  (18  areas,  which  serve  to  preserve  this 
species)  are  found  in  the  mangrove  zone  species  in  the  Eastern  United  States. 
(Layne  1974).  In  addition,  two  species  of  Shemnitz  (1974)  reported  that  most  of  the 
marine  mammals  are  known  from  mangrove  remaining  panthers  were  found  in  the 
areas.  Data  on  the  abundance  and  food  southwest  portion  of  Florida  along  the 
habits  of  these  20  species  are  summarized  coast  and  in  the  interior  Everglades 
in  Appendix  E.  All  are  permanent  resi-  regions, 
dents.      The  criteria   for  inclusion   in  this 

table  are  similar  to  those  used   for  the  The   extent   to   which  other  carnivores 

avifauna.     Sight  records  in  mangroves  or  use  mangrove  areas   varies   widely   among 

locality  data  from  known  mangrove  areas  species.       Schwartz    (1949)    states    that 

were    required   before   a   species    was   in-  mink,  although  rare,  prefer  mangroves  to 

eluded.      This    has   produced   a   conservative  other   coastal    habitats    in    Florida.      Layne 

estimate   of   the   mammal    species   that    uti-  (1974,  see  his  figure  1)  gives  a  disjunct 

lize  mangrove  areas.  distribution    for    this    species    in    Florida, 

with  the  major  geographical    range  being 

Several     mammals    do    not    appear    in  the   southwest   coast.     River  otters  also 

Appendix    E    because    they    have    not    been  utilize   mangrove   habitat   heavily.      Otters 

recorded   from   mangrove  swamps   in   south  have   been   found   even   far   from   shore   on 

Florida;    however,    they   occur   so   widely  small  mangrove  overwash  islands  in  Florida 

that  we  suspect  they  will  be  found  in  this  Bay  (Layne  1974).     Gray  fox  are  not  depen- 

habitat     in    the    future.       This    group  dent  upon  mangroves,  although  they  occa- 

includes    the    cotton    mouse,     Peromyscus  sionally   use   this    habitat.      Less    than    20% 

gossypinus,   the  hispid  cotton    rat,    Si g-  of  all    sightings   of  this    species    in    Ever- 

modon   hispidus,    the    round-tailed    muskrat,  glades    National    Park    were    from    mangroves 

Neof iber    a! leni ,    the    house    mouse,    Mus  (Layne  1974).     Bobcat  are  found  in  almost 

musculus,     the    least    shrew,     Cryptoti  s  all   habitats  in  south  Florida  from  pine- 

parva,  and  the  short -tailed  shrew,  Blarina  lands    to    dense    mangrove    forests.       The 

brevicauda.  preponderance  of  recent   sightings,  how- 
ever, has      been  made   from   the   mangrove 

Few   rodents   and  no  bats   are   included  zone,  particularly  on  offshore  mangrove 
in  Appendix   E.     Compared  to  the   rest   of  overwash   islands    (Layne   1974).     Black   bear 
the    State,    the   south    Florida    region    is  are   apparently  most   abundant   in  the  Big 
deficient     in    these    two    groups     (Layne  Cypress   Swamp  of  Collier  County    (Shemnitz 
1974).     Although  we  have  no  confirmative  1974)    and    are    rare    in   the    remainder   of 
field    data,     we    suspect    that    mangrove  south  Florida, 
swamps  along  the  central   and  north  Florida 
coasts    contain    more   mammal    species,    par- 
ticularly   rodents   and   bats.  The  small    mammal    fauna  of  the  man- 
grove zone  of  south  Florida  are  predomi- 

A  number  of  medium-sized  and   large  nately  arboreal   and  terrestrial    species 

carnivores,  including  panther,  gray  fox,  which  are  adapted  to  periodic   flooding, 

bobcat,    striped    skunk,     raccoon,     mink,  Opossum,  marsh  rabbits,  cotton  rats,  and 

river    otter,    and   black   bear,    appear   to  rice    rats    are   commonly    found    in    mangrove 

utilize    south    Florida    mangrove    areas.  swamps.      The    Cudjoe    Key    rice    rat    is    a 

Only    three    of    these    species    (striped  newly    described    species    found    only    on 

skunk,    raccoon,    and   bobcat)   are  common   in  Cudjoe    Key    in    the    Florida    Keys.       This 

mangroves,    but    several    of    the    rarer  species  appears  to  be  closely  associated 

species    seem   to   be   highly   dependent    on  with    stands    of    white    mangroves    (Hamilton 

mangrove    swamps.      Of    18    recent    sightings  and   Whittaker    1979). 
of    the    panther    in    Everglades    National 

Park,    15   were   from  mangrove  ecosystems  White-tailed    deer    are    common    in 

72 


Florida  mangrove  swamps,  although  they 
utilize  many  other  habitats.  The  key 
deer,  a  rare  and  endangered  subspecies,  is 
restricted  to  the  Big  Pine  Key  group  in 
the  Florida  Keys,  although  it  ranged  onto 
the  mainland  in  historical  times.  Al- 
though this  little  deer  makes  use  of  pine 
uplands  and  oak  hammocks,  it  extensively 
exploits  mangrove  swamps  for  food  and 
cover. 

Two  marine  mammals,  the  bottlenose 
porpoise  and  the  manatee,  frequent 
mangrove-lined  waterways.  The  bottlenose 
porpoise  feeds  on  mangrove-associated 
fishes  such  as  the  striped  mullet,  Mugi  1 
cephal us.       Although    the    manatee    feeds 


primarily  upon  sea  grasses  and  other 
submerged  aquatic  plants,  it  is  commonly 
found  in  canals,  coastal  rivers,  and 
embayments  close  to  mangrove  swamps. 

Except  for  the  Cudjoe  Key  rice  rat, 
none  of  the  mammals  found  in  Florida  man- 
groves are  solely  dependent  upon  mangrove 
ecosystems;  all  of  these  species  can 
utilize  other  habitats.  The  destruction 
of  extensive  mangrove  swamps  would,  how- 
ever, have  deleterious  effects  on  almost 
all  of  these  species.  Populations  of 
panther,  key  deer,  and  the  river  otter 
would  probably  be  the  most  seriously 
affected,  because  they  use  mangrove  habi- 
tat extensively. 


73 


CHAPTER  11 


VALUE  OF  MANGROVE  ECOSYSTEMS  TO  MAN 


Mangrove  swamps  are  often  hot,  fetid, 
mosquito-ridden,  and  almost  impenetrable. 
As  a  consequence,  they  are  frequently  held 
in  low  regard.  It  is  possible  that  more 
acres  of  mangrove,  worldwide,  have  been 
obliterated  by  man  in  the  name  of  "recla- 
mation" than  any  other  type  of  coastal 
environment.  Reclamation,  according  to 
Webster's,  means  "to  claim  back,  as  of 
wasteland".  Mangrove  swamps  are  anything 
but  wasteland,  however,  and  it  is  impor- 
tant to  establish  this  fact  before  a 
valuable  resource  is  lost.  We  can  think 
of  six  major  categories  of  mangrove  values 
to  man;    no  doubt,   there  are  more. 


11.1      SHORELINE   STABILIZATION   AND   STORM 
PROTECTION 

The  ability  of  all  three  Florida 
mangroves  to  trap,  hold  and,  to  some 
extent,  stabilize  intertidal  sediments  has 
been  demonstrated  repeatedly  (reviewed  by 
Scoffin  1970;  Carlton  1974).  The  contem- 
porary view  of  mangroves  is  that  they 
function  not  as  "land  builders"  as  hypo- 
thesized by  Davis  (1940)  and  others,  but 
as  "stabilizers"  of  sediments  that  have 
been  deposited  largely  by  geomorphological 
processes    (see   section  3.2). 

Gill  (1970),  Savage  (1972),  Teas 
(1977),  and  others  have  emphasized  that 
land  stabilization  by  mangroves  is  pos- 
sible only  where  conditions  are  relatively 
quiescent  and  strong  wave  action  and/or 
currents  do  not  occur.  Unfortunately,  no 
one  has  devised  a  method  to  predict  the 
threshold  of  physical  conditions  above 
which  mangroves  are  unable  to  survive  and 
stabilize  the  sediments.  Certainly,  this 
depends  to  some  extent  on  substrate  type; 
mangroves  appear  to  withstand  wave  energy 
best  on  solid  rock  substrates  with  many 
cracks  and  crevices  for  root  penetration. 
From  our  own  experience,  we  suspect  that 
mangroves  on  sandy  and  muddy  substrates 
cannot  tolerate  any  but  the  lowest  wave 
energies,  tidal  currents  much  above  25 
cm/s,   or  heavy,    regular  boat  wakes. 

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


tioned by  Savage  (1972),  Carlton  (1974), 
and  Teas  (1977).  These  authors  argue  that 
the  black  mangrove  (1)  is  easier  to 
transplant  as  a  seedling,  (2)  establishes 
its  pneumatophore  system  more  rapidly  than 
the  red  mangrove  develops  prop  roots,  (3) 
has  an  underground  root  system  that  is 
better  adapted  to  holding  sediments  (Teas 
1977),  (4)  is  more  cold-hardy,  and  (5)  can 
better  tolerate  "artificial"  substrates 
such  as  dredge-spoil,  finger  fills,  and 
causeways.  Generally,  the  white  mangrove 
is  regarded  as  the  poorest  land  stabilizer 
of  the  Florida  mangroves  (Hanlon  et  al. 
1975). 

Although  mangroves  are  susceptible  to 
hurricane  damage  (see  section  12.1),  they 
provide  considerable  protection  to  areas 
on  their  landward  side.  They  cannot 
prevent  all  flooding  damage,  but  they  do 
mitigate  the  effects  of  waves  and 
breakers.  The  degree  of  this  protection 
is  roughly  proportional  to  the  width  of 
the  mangrove  zone.  Very  narrow  fringing 
forests  offer  minimal  protection  while 
extensive  stands  of  mangroves  not  only 
prevent  wave  damage,  but  reduce  much  of 
the  flooding  damage  by  damping  and  holding 
flood  waters.  Fosberg  (1971)  suggested 
that  the  November  1970  typhoon  and  accom- 
panying storm  surge  that  claimed  between 
300,000  and  500,000  human  lives  in 
Bangladesh  might  not  have  been  so  destruc- 
tive if  thousands  of  hectares  of  mangrove 
swamps  had  not  been  replaced  with  rice 
paddies. 


11.2     HABITAT   VALUE  TO   WILDLIFE 

Florida  mangrove  ecosystems  are 
important  habitat  for  a  wide  variety  of 
reptiles,  amphibians,  birds,  and  mammals 
(see  sections  8,  9,  and  10).  Some  of 
these  animals  are  of  commercial  and  sport 
importance  (e.g.,  white-tailed  deer,  sea 
turtles,  pink  shrimp,  spiny  lobster, 
snook,  grey  snapper).  Many  of  these  are 
important  to  the  south  Florida  tourist 
industry  including  the  wading  birds  (e.g., 
egrets,  wood  stork,  white  ibis,  herons) 
which  nest   in  the  mangrove  zone. 


74 


11.3      IMPORTANCE    TO   THREATENED   AND    ENDAN- 
GERED  SPECIES 

The  mangrove  forests  of  south  Florida 
are  important  habitat  for  at  least  seven 
endangered  species,  five  endangered  sub- 
species, and  three  threatened  species 
(Federal  Register  1980).  The  endangered 
species  include  the  American  crocodile, 
the  hawksbill  sea  turtle,  the  Atlantic 
ridley  sea  turtle,  the  Florida  manatee, 
the  bald  eagle,  the  American  peregrine 
falcon,  and  the  brown  pelican.  The  endan- 
gered subspecies  are  the  key  deer 
(Odocoi leus  vi  rginianus  cl  avi  urn),  the 
Florida  panther  (Felis  concolor  coryi ), 
the  Barbados  yellow  warbler  (Dendroica 
petechia  petechia),  the  Atlantic  saltmarsh 
snake  (Nerodia  fasciata  taeniata)  and  the 
eastern  indigo  snake  (Drymarchon  corais 
couperi).  Threatened  species  include  the 
American  alligator,  the  green  sea  turtle 
and  the  loggerhead  sea  turtle.  Although 
all  of  these  animals  utilize  mangrove 
habitat  at  times  in  their  life  histories, 
species  that  would  be  most  adversely 
affected  by  widespread  mangrove  destruc- 
tion are  the  American  crocodile,  the 
Florida  panther,  the  American  peregrine 
falcon,  the  brown  pelican,  and  the 
Atlantic  ridley  sea  turtle.  The  so-called 
mangrove  fox  squirrel  (Sciurus  ni  ger 
avicennia)  is  widely  believed  to  be  a 
mangrove-dependent  endangered  species. 
This  is  not  the  case  since  it  is  currently 
regarded  as  "rare",  not  endangered,  and, 
further,  there  is  some  question  whether 
or  not  this  is  a  legitimate  sub-species 
(Hall  1981).  As  a  final  note,  we  should 
point  out  that  the  red  wolf  (Cam's  rufus), 
which  is  believed  to  be  extinct  in 
Florida,  at  one  time  used  mangrove  habitat 
in  addition  to  other  areas  in  south 
Florida. 


11.4       VALUE    TO    SPORT    AND    COMMERCIAL 
FISHERIES 

The  fish  and  invertebrate  fauna  of 
mangrove  waterways  are  closely  linked  to 
mangrove  trees  through  (a)  the  habitat 
value  of  the  aerial  root  structure  and  (b) 
the  mangrove  leaf  detritus-based  food  web 
(see  sections  6  and  7).    The  implications 


of  these  connections  were  discussed  by 
Heald  (1969),  Odum  (1970),  Heald  and  Odum 
(1970),  and  Odum  and  Heald  (1975b)  in 
terms  of  support  for  commercial  and  sport 
fisheries. 

A  minimal  list  of  mangrove-associated 
organisms  of  commercial  or  sport  value 
includes  oysters,  blue  crabs,  spiny 
lobsters,  pink  shrimp,  snook,  mullet, 
menhaden,  red  drum,  spotted  sea  trout, 
gray  and  other  snapper,  tarpon, 
sheepshead,  ladyfish,  jacks,  gafftopsail 
catfish,  and  the  jewfish.  Heald  and  Odum 
(1970)  pointed  out  that  the  commercial 
fisheries  catch,  excluding  shrimp,  in  the 
area  from  Naples  to  Florida  Bay  was  2.7 
million  pounds  in  1965.  Almost  all  of  the 
fish  and  shellfish  which  make  up  this 
catch  utilize  the  mangrove  habitat  at  some 
point  during  their  life  cycles.  In  addi- 
tion, the  Tortugas  pink  shrimp  fishery, 
which  produces  in  excess  of  11  million 
pounds  of  shrimp  a  year  (Idyll  1965a),  is 
closely  associated  with  the  Everglades 
estuary  and  its  mangrove-lined  bays  and 
rivers. 


11.5        AESTHETICS, 
INTANGIBLES 


TOURISM     AND     THE 


One  value  of  the  mangrove  ecosystem, 
which  is  difficult  to  document  in  dollars 
or  pounds  of  meat,  is  the  aesthetic  value 
to  man.  Admittedly,  not  all  individuals 
find  visits  to  mangrove  swamps  a  pleasant 
experience.  There  are  many  others,  how- 
ever, who  place  a  great  deal  of  value  on 
the  extensive  vistas  of  mangrove  canopies, 
waterways,  and  associated  wildlife  and 
fishes  of  south  Florida.  In  a  sense,  this 
mangrove  belt  along  with  the  remaining 
sections  of  the  freshwater  Everglades  and 
Big  Cypress  Swamp  are  the  only  remaining 
wilderness  areas  in  this  part  of  the 
United  States. 

Hundreds  of  thousands  of  visitors 
each  year  visit  the  Everglades  National 
Park;  part  of  the  reason  for  many  of  these 
visits  includes  hopes  of  catching  snook  or 
gray  snappers  in  the  mangrove-lined  water- 
ways, seeing  exotic  wading  birds,  croco- 
diles, or  panthers,  or  simply  discovering 


75 


what  a  tropical  mangrove  forest  looks 
like.  The  National  Park  Service,  in  an 
attempt  to  accommodate  this  last  wish, 
maintains  extensive  boardwalks  and  canoe 
trails  through  the  mangrove  forests  near 
Flamingo,  Florida.  In  other,  more 
developed  parts  of  the  State,  small  stands 
of  mangroves  or  mangrove  islands  provide  a 
feeling  of  wilderness  in  proximity  to  the 
rapidly  burgeoning  urban  areas.  A  variety 
of  tourist  attractions  including  Fairchild 
Tropical  Gardens  near  Miami  and  Tiki 
Gardens  near  St.  Petersburg  utilizes  the 
exotic  appearance  of  mangroves  as  a  key 
ingredient  in  an  attractive  landscape. 
Clearly,  mangroves  contribute  intangibly 
by  diversifying  the  appearance  of  south 
Florida. 


11.6     ECONOMIC   PRODUCTS 

Elsewhere  in  the  world,  mangrove 
forests  serve  as  a  renewable  resource  for 
many  valuable  products.  For  a  full  dis- 
cussion of  the  potential  uses  of  mangrove 
products,  see  de  la  Cruz  (in  press  a), 
Morton  (1965)  for  red  mangrove  products, 
and  Moldenke  (1967)  for  black  mangrove 
products. 

In  many  countries  the  bark  of  man- 
groves is  used  as  a  source  of  tannins  and 
dyes.  Since  the  bark  is  20%  to  30%  tannin 
on  a  dry  weight  basis,  it  is  an  excellent 
source  (Hanlon  et  al.  1975).  Silviculture 
(forestry)  of  mangrove  forests  has  been 
practiced  extensively  in  Africa,  Puerto 
Rico,  and  many  parts  of  Southeast  Asia 
(Holdridge  1940;  Noakes  1955;  Macnae  1968; 
Walsh   1974;      Teas   1977).     Mangrove  wood 


makes  a  durable  and  water  resistant  timber 
which  has  been  used  successfully  for  resi- 
dential buildings,  boats,  pilings, 
hogsheads,  fence  posts,  and  furniture 
(Kuenzler  1974;  Hanlon  et  al.  1975).  In 
Southeast  Asia  mangrove  wood  is  widely 
used   for  high   quality  charcoal. 

Morton  (1965)  mentions  that  red  man- 
grove fruits  are  somtimes  eaten  by  humans 
in  Central  America,  but  only  by  popula- 
tions under  duress  and  subject  to  starva- 
tion. Mangrove  leaves  have  variously  been 
used  for  teas,  medicinal  purposes,  and 
livestock  feeds.  Mangrove  teas  must  be 
drunk  in  small  quantities  and  mixed  with 
milk  because  of  the  high  tannin  content 
(Morton  1962);  the  milk  binds  the  tannins 
and  makes  the  beverage  more  palatable. 

As  a  final  note,  we  should  point  out 
that  mangrove  trees  are  responsible  for 
contributing  directly  to  one  commercial 
product  in  Florida.  The  flowers  of  black 
mangroves  are  of  considerable  importance 
to  the  three  million  dollar  (1965  figures) 
Florida    honey   industry   (Morton   1964). 

Other  than  the  honey  industry,  most 
of  these  economic  uses  are  somewhat 
destructive.  There  are  many  cases  in 
which  clear-cut  mangrove  forests  have 
failed  to  regenerate  successfully  for  many 
years  because  of  lack  of  propagule 
dispersal  or  increased  soil  salinities 
(Teas  1979).  We  believe  that  the  best  use 
of  Florida  mangrove  swamps  will  continue 
to  be  as  preserved  areas  to  support 
wildlife,  fishing,  shoreline  stabiliza- 
tion, endangered  species,  and  aesthetic 
values. 


76 


CHAPTER    12.       MANAGEMENT    IMPLICATIONS 


12.1      INHERENT  VULNERABILITY 

Mangroves  have  evolved  remarkable 
physiological  and  anatomical  adaptations 
enabling  them  to  flourish  under  conditions 
of  high  temperatures,  widely  fluctuating 
salinities,  high  concentrations  of  heavy 
metals  (Walsh  et  al.  1979),  and  anaerobic 
soils.  Unfortunately,  one  of  these  adap- 
tations, the  aerial  root  system,  is  also 
one  of  the  plant's  most  vulnerable  compo- 
nents. Odum  and  Johannes  (1975)  have 
referred  to  the  aerial  roots  as  the  man- 
grove's Achi 1 1 es'  heel  because  of  their 
susceptibility  to  clogging,  prolonged 
flooding,  and  boring  damage  from  isopods 
and  other  invertebrates  (see  section  6  for 
a  discussion  of  the  latter).  This  means 
that  any  process,  natural  or  man-induced, 
which  coats  the  aerial  roots  with  fine 
sediments  or  covers  them  with  water  for 
extended  periods  has  the  potential  for 
mangrove  destruction.  Bacon  (1970)  men- 
tions a  case  in  Trinidad  where  the  Caroni 
River  inundated  the  adjacent  Caroni 
Mangrove  Swamp  during  a  flood  and 
deposited  a  layer  of  fine  red  marl  in  a 
large  stand  of  black  mangroves  which  sub- 
sequently died.  Many  examples  of  damage 
to  mangrove  swamps  from  human  activities 
have   been   documented   (see   section   12.2). 

One  of  the  few  natural  processes  that 
causes  periodic  and  extensive  damage  to 
mangrove  ecosystems  is  large  hurricanes 
(Figure  16).  Craighead  and  Gilbert  (1962) 
and  Tabb  and  Jones  (1962)  have  documented 
the  impact  of  Hurricane  Donna  in  1960  on 
parts  of  the  mangrove  zone  of  south 
Florida.  Craighead  and  Gilbert  (1962) 
found  extensive  damage  over  an  area  of 
100,000  acres  (40,000  ha).  Loss  of  trees 
ranged  from  25%  to  100%.  Damage  occurred 
in  three  ways:  (1)  wind  shearing  of  the 
trunk  6  to  10  ft  (2  to  3  m)  above  ground, 
(2)  overwash  mangrove  islands  being  swept 
clean,  and  (3)  trees  dying  months  after 
the  storm,  apparently  in  response  to 
damage  to  the  prop  roots  from  coatings  by 
marl  and  fine  organic  matter.  The  latter 
type  of  damage  was  most  widespread,  but 
rarely  occurred  in  intertidal  forests, 
presumably  because  the  aerial  roots  were 
flushed  and  cleaned  by  tidal  action.  Fish 
and   invertebrates   were   adversely   affected 


by  oxygen  depletion  due  to  accumulations 
of  decomposing  organic  matter  (Tabb  and 
Jones   1962). 

Hurricane  Betsy  in  1965  did  little 
damage  to  mangroves  in  south  Florida; 
there  was  also  little  deposition  of  silt 
and  marl  within  mangrove  stands  from  this 
minimal  storm  (Alexander  1 967).  Lugo  et 
al.  (1976)  have  hypothesized  that  severe 
hurricanes  occur  in  south  Florida  and 
Puerto  Rico  on  a  time  interval  of  25  to  30 
years  and  that  mangrove  ecosystems  are 
adapted  to  reach  maximum  biomass  and  pro- 
ductivity on  the  same  time  cycle. 


12.2     MAN-INDUCED   DESTRUCTION 

Destruction  of  mangrove  forests  in 
Florida  has  occurred  in  various  ways 
including  outright  destruction  and  land 
filling,  diking  and  flooding  (Figure  17), 
through  introduction  of  fine  particulate 
material,  and  pollution  damage,  par- 
ticularly oil  spills.  To  our  knowledge 
there  are  no  complete,  published  docu- 
mented estimates  of  the  amount  of  mangrove 
forests  in  Florida  which  have  been 
destroyed  by  man  in  this  century.  Our 
conclusion  is  that  total  loss  statewide  is 
not  too  great,  probably  in  the  range  of  3 
to  5%  of  the  original  area  covered  by 
mangroves  in  the  19th  century,  but  that 
losses  in  specific  areas,  particularly 
urban  areas,  are  appreciable.  This  con- 
clusion is  based  on  four  pieces  of  infor- 
mation. (1)  Lindall  and  Saloman  (1977) 
have  estimated  that  the  total  loss  of 
vegetated  intertidal  marshes  and  mangrove 
swamps  in  Florida  due  to  dredge  and  fill 
is  23,521  acres  (9,522  ha);  remember  that 
there  are  between  430,000  and  500,000 
acres  (174,000  to  202,000  ha)  of  mangroves 
in  Florida  (see  section  1.3).  (2) 
Birnhak  and  Crowder  (1974)  estimate  a  loss 
of  approximately  11,000  acres  (4,453  ha) 
of  mangroves  between  1943  and  1970  in 
three  counties  (Collier,  Monroe,  and 
Dade).  (3)  An  obvious  loss  of  mangrove 
forests  has  occurred  in  Tampa  Bay,  around 
Marco  Island,  in  the  Florida  Keys,  and 
along  the  lower  east  coast  of  Florida. 
For  example,  Lewis  et  al.  (1979)  estimated 
that    44%    of    the    intertidal    vegetation 


77 


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


78 


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


79 


including  mangroves  in  the  Tampa  Bay 
estuary  has  been  destroyed  during  the  past 
100  years.  (4)  Heald  (unpublished  MS.) 
has  estimated  a  loss  of  2,000  acres  (810 
ha)  of  mangroves  within  the  Florida  Keys 
(not  considered  by  Birnhak  and  Crowder 
1974).  So  while  loss  of  mangrove  ecosys- 
tems throughout  Florida  is  not  over- 
whelming, losses  at  specific  locations 
have   been   substantial. 

Diking,  impounding,  and  long-term 
flooding  of  mangroves  with  standing  water 
can  cause  mass  mortality,  especially  when 
prop  roots  and  pneumatophores  are  covered 
(Breen  and  Hill  1969;  Odum  and  Johannes 
1975;  Patterson-Zucca  1978;  Lugo  1981). 
In  south  Florida,  E.  Heald  (pers.  comm.) 
has  observed  that  permanent  impoundment  by 
diking  which  prevents  any  tidal  exchange 
and  raises  water  levels  significantly 
during  the  wet  season  will  kill  all  adult 
red  and  black  mangrove  trees.  If  condi- 
tions behind  the  dike  remain  relatively 
dry,  the  mangroves  may  survive  for  many 
years  until  replaced  by  terrestrial  vege- 
tation. 


Mangroves  are  unusually  susceptible 
to  herbicides  (Walsh  et  al.  1973).  At 
least  250,000  acres  (100,000  ha)  of  man- 
grove forests  were  defoliated  and  killed 
in  South  Viet  Nam  by  the  U.S.  military. 
This  widespread  destruction  has  been  docu- 
mented by  Tschirley  (1969),  Orians  and 
Pfeiffer  (1970),  Westing  (1971),  and  a 
committee  of  the  U.S.  Academy  of  Sciences 
(Odum  et  al.  1974).  In  many  cases  these 
forests  were  slow  to  regenerate;  observa- 
tions by  de  Sylva  and  Michel  (1974)  indi- 
cated higher  rates  of  siltation,  greater 
water  turbidity,  and  possibly  lower  dis- 
solved oxygen  concentrations  in  swamps 
which  sustained  the  most  damage.  Teas  and 
Kelly  (1975)  reported  that  in  Florida  the 
black  mangrove  is  somewhat  resistant  to 
most  herbicides  but  the  red  mangrove  is 
extremely  sensitive  to  herbicide  damage. 
He  hypothesized  that  the  vulnerability  of 
the  red  mangrove  is  related  to  the  small 
reserves  of  viable  leaf  buds  in  this  tree. 
Following  his  reasoning,  the  stress  of  a 
single  defoliation  is  sufficient  to  kill 
the   entire   tree. 


Although  mangroves  commonly  occur  in 
areas  of  rapid  sedimentation,  they  cannot 
survive  heavy  loads  of  fine,  floculent 
materials  which  coat  the  prop  roots.  The 
instances  of  mangrove  death  from  these 
substances  have  been  briefly  reviewed  by 
Odum  and  Johannes  (1975).  Mangrove  deaths 
from  fine  muds  and  marl,  ground  bauxite 
and  other  ore  wastes,  sugar  cane  wastes, 
pulp  mill  effluent,  sodium  hydroxide 
wastes  from  bauxite  processing,  and  from 
intrusion  of  large  quantities  of  beach 
sand  have  been  documented  from  various 
areas  of  the  world. 

12.3     EFFECTS   OF   OIL   SPILLS  ON   MANGROVES 

There  is  little  doubt  that  petroleum 
and  petroleum  byproducts  can  be  extremely 
harmful  to  mangroves.  Damage  from  oil 
spills  has  been  reviewed  by  Odum  and 
Johannes  (1975),  Carlberg  (1980),  Ray  (in 
press),  and  de  la  Cruz  (in  press,  b). 
Over  100  references  detailing  the  effects 
of  oil  spills  on  mangroves  and  mangrove- 
associated  biota  are  included  in  these 
reviews. 

Petroleum  and  its  byproducts  injure 
and  kill  mangroves  in  a  variety  of  ways. 
Crude  oil  coats  roots,  rhizomes,  and  pneu- 
matophores and  disrupts  oxygen  transport 
to  underground  roots  (Baker  1971). 
Various  reports  suggest  that  the  critical 
concentration  for  crude  oil  spills  which 
may  cause  extensive  damage  is  between  100 
and  200  ml/m  of  swamp  surface  (Odum  and 
Johannes  1975).  Petroleum  is  readily 
absorbed  by  lipophylic  substances  on  sur- 
faces of  mangroves.  This  leads  to  severe 
metabolic  alterations  such  as  displacement 
of  fatty  molecules  by  oil  hydrocarbons 
leading  to  destruction  of  cellular  permea- 
bility and/or  dissolution  of  hydrocarbons 
in  lipid  components  of  chloroplasts  (Baker 
1971). 

As  with  other  intertidal  communities, 
many  of  the  invertebrates,  fishes,  and 
plants  associated  with  the  mangrove  com- 
munity are  highly  susceptible  to  petroleum 
products.  Widespread  destruction  of 
organisms  such  as  attached  algae,  oysters, 
tunicates,  crabs,  and  gobies  have  been 
reported  in  the  literature   (reviewed  by  de 


80 


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


12.4     MAN-INDUCED   MODIFICATIONS 


Damage  from  oil  spills  follows  a 
predictable  pattern  (Table  7)  which  may 
require  years  to  complete.  It  is  impor- 
tant to  recognize  that  many  of  the  most 
severe  responses,  including  tree  death, 
may  not  appear  for  months  or  even  years 
after  the   spill. 

In  Florida,  Chan  (1977)  reported  that 
red  mangrove  seedlings  and  black  mangrove 
pneumatophores  were  particularly  sensitive 
to  an  oil  spill  which  occurred  in  the 
Florida  Keys.  Lewis  (1979a,  1980b)  has 
followed  the  long-term  effects  of  a  spill 
of  150,000  liters  (39,000  gal)  of  bunker  C 
and  diesel  oil  in  Tampa  Bay.  He  observed 
short-term  (72-hour)  mortality  of  inverte- 
brates such  as  the  gastropod  Mel ongena 
corona  and  the  polychaete  Laeonerei  s 
cul veri.  Mortality  of  all  three  species 
of  mangroves  began  after  three  weeks  and 
continued  for  more  than  a  year.  Sub- 
lethal damage  included  partial  defoliation 
of  all  species  and  necrosis  of  black 
mangrove  pneumatophores;  death  depended 
upon  the  percentage  of  pneumatophores 
affected. 

In  addition  to  the  damage  from  oil 
spills,  there  are  many  adverse  impacts  on 
mangrove  forests  from  the  process  of  oil 
exploration  and  drilling  (Table  8).  This 
type  of  damage  can  often  be  reduced 
through  careful  management  and  monitoring 
of  drilling    sites. 

Although  little  is  known  concerning 
ways  to  prevent  damage  to  mangroves  once  a 
spill  has  occurred,  protection  of  aerial 
roots  seems  essential.  Prop  roots  and 
pneumatophores  must  be  cleaned  with  com- 
pounds which  will  not  damage  the  plant 
tissues.  Dispersants  commonly  used  to 
combat  oil  spills  are,  in  general,  toxic 
to  vascular  plants  (Baker  1971).  If  pos- 
sible, oil  laden  spray  should  not  be 
allowed  to  reach  leaf  surfaces.  Damage 
during  clean-up  (e.g.,  trampling,  compac- 
tion, bulldozing)  may  be  more  destructive 
than  the  untreated  effects  of  the  oil 
spill    (de   la   Cruz   in   press,  b). 


In  south  Florida,  man  has  been  re- 
sponsible for  modifications  which,  while 
not  killing  mangroves  outright,  have  al- 
tered components  of  the  mangrove  ecosys- 
tem. One  of  the  most  widespread  changes 
involves  the  alteration  of  freshwater 
runoff.  Much  of  the  freshwater  runoff  of 
the  Florida  Everglades  has  been  diverted 
elsewhere  with  the  result  that  salinities 
in  the  Everglades  estuary  are  generally 
higher  than  at  the  turn  of  the  century. 
Teas  (1977)  points  out  that  drainage  in 
the  Miami  area  has  lowered  the  water  table 
as  much  as  2  m  (6  ft). 

Interference  with  freshwater  inflow 
has  extensive  effects  on  estuaries  (Odum 
1970).  Florida  estuaries  are  no  excep- 
tion; the  effects  on  fish  and  invertebrate 
species  along  the  edge  of  Biscayne  and 
Florida  Bays  have  been  striking.  The 
mismanagement  of  freshwater  and  its 
effects  on  aquatic  organisms  have  been 
discussed  by  Tabb  (1963);  Idyll  (1965a,b); 
Tabb  and  Yokel  (1968)  and  Idyll  et  al. 
(1968).  In  addition,  Estevez  and  Simon 
(1975)  have  hypothesized  that  the  impact 
of  the  boring  isopod,  Sphaeroma  terebrans, 
may  be  more  severe  when  freshwater  flows 
from  the  Everglades  are  altered. 

One  generally  unrecognized  side 
effect  of  lowered  freshwater  flow  and  salt 
water  intrusion  has  been  the  inland  expan- 
sion of  mangrove  forests  in  many  areas  of 
south  Florida.  There  is  documented  evi- 
dence that  the  mangrove  borders  of 
Biscayne  Bay  and  much  of  the  Everglades 
estuary  have  expanded  inland  during  the 
past  30  to  40  years  (Reark  1975;  Teas 
1979;  Ball   1980). 

Sections  of  many  mangrove  forests  in 
south  Florida  have  been  replaced  by  filled 
residential  lots  and  navigation  canals. 
Although  these  canal  systems  have  not  been 
studied  extensively,  there  is  some  evi- 
dence, mostly  unpublished,  that  canals  are 
not  as  productive  in  terms  of  fishes  and 
invertebrates  as  the  natural  mangrove- 
lined    waterways    which   they   replaced. 


81 


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


Stage 


Observed  impact 


Acute 


0  to  15  days 


15  to  30  days 


Deaths  of  birds,  turtles,  fishes,  and 
invertebrates 

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


Chronic 


30  days  to  1  year 


1  year  to  5  years 


1  year  to  1 0  years  ( ? ) 


10  to  50  years  (?) 


Defoliation  and  death  of  medium-sized 
mangroves  (1-3  m),  tissue  damage  to 
aerial  roots 

Death  of  large  mangroves  (greater  than 
3  m),  loss  of  oiled  aerial  roots,  and 
regrowth  of  new  roots  (often  deformed) 

Recol oni zati on  of  oil-damaged  areas  by 
new  seedlings 

Reduction  in  litter  fall,  reduced  re- 
production, and  reduced  survival  of 
seedl 1 ngs 

Death  or  reduced  growth  of  young  trees 
colonizing  spill  site  (?) 

Increased  insect  damage  (?) 

Complete  recovery 


82 


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83 


Weinstein  et  al.  (1977)  found  that  artifi- 
cial canals  had  lower  species  diversity  of 
benthic  infauna  and  trawl -captured  fishes 
and  generally  finer  sediments  than  the 
natural  communities.  Courtney  (1975) 
reported  a  number  of  mangrove-associated 
invertebrates  which  did  not  occur  in  the 
artificial   channels. 

Mosquito  production  is  a  serious 
problem  in  black  mangrove-dominated  swamps 
in  Florida  (Provost  1969).  The  salt  marsh 
mosquitos,  Aedes  taeniorhynchus  and  A. 
sol  1 icitans,  do  not  reproduce  below  the 
mean  high  tide  mark  and  for  this  reason 
are  not  a  serious  problem  in  the  inter- 
tidal  red  mangrove  swamps.  Mosquitos  lay 
their  eggs  on  the  damp  soil  of  the  irregu- 
larly flooded  black  mangrove  zone;  these 
eggs  hatch  and  develop  when  flooded  by 
spring  tides,  storm  tides  or  heavy  rains. 
As  with  the  "high  marsh"  of  temperate 
latitudes,  there  have  been  some  attempts 
to  ditch  the  black  mangrove  zone  so  that 
it  drains  rapidly  after  flooding. 
Although  properly  designed  ditching  does 
not  appear  to  be  particularly  harmful  to 
mangrove  swamps  (other  than  the  area 
destroyed  to  dig  the  ditch  and  receive  the 
spoil),  it  is  an  expensive  practice  and 
for  this  reason  is  not  widely  practiced. 
Properly  managed  diking  can  be  an  effec- 
tive mosquito  control  approach  with  mini- 
mal side  effects  to  black  mangroves 
(Provost  1969).  Generally,  ditching  or 
diking  of  the  intertidal  red  mangrove  zone 
is  a  waste  of  money. 

Mangrove  swamps  have  been  proposed  as 
possible  tertiary  treatment  areas  for 
sewage  (see  discussion  by  Odum  and 
Johannes  1975).  To  our  knowledge,  this 
alternate  use  is  not  currently  practiced 
in  south  Florida.  Until  more  experimental 
results  are  available  on  the  assimilative 
capacities  and  long-term  changes  to  be 
expected  in  mangrove  forests  receiving 
heavy  loads  of  secondary  treated  sewage, 
it  would  be  an  environmental  risk  to  use 
mangrove  forests  for  this   purpose. 

In  many  areas  of  the  world  mangrove 
swamps  have  been  converted  to  other  uses 
such  as  aquaculture  and  agriculture  (see 
de  la  Cruz,   in  press,  a).     Although  some 


of  the  most  productive  aquaculture  ponds 
in  Indonesia  and  the  Philippines  are 
located  in  former  mangrove  swamps,  there 
is  some  question  whether  the  original 
natural  system  was  not  equally  productive 
in  terms  of  fisheries  products  at  no  cost 
to  man  (Odum  1974).  Conversion  to 
aquaculture  and  agriculture  is  cursed  with 
a  variety  of  problems  including  subsequent 
land  subsidence  and  the  "cat  clay" 
problem.  The  latter  refers  to  the 
drastically  lowered  soil  pH  which  often 
occurs  after  drainage  and  has  been  traced 
to  oxidation  of  reduced  sulfur  compounds 
(Dent  1947;  Tomlinson  1957;  Hesse  1961; 
Hart  1962,  1963;  Moorman  and  Pons  1975). 
Experience  in  Africa,  Puerto  Rico,  and 
Southeast  Asia  confirms  that  mangrove 
forests  in  their  natural  state  are  more 
valuable   than   the    "reclaimed"    land. 

12.5  PROTECTIVE  MEASURES  INCLUDING 
TRANSPLANTING 

Protection  of  mangroves  includes  (1) 
prevention  of  outright  destruction  from 
dredging  and  filling;  (2)  prevention  of 
drainage,  diking  and  flooding  (except  for 
carefully  managed  mosquito  control);  (3) 
prevention  of  any  alteration  of  hydrologi- 
cal  circulation  patterns,  particularly 
i nvol ving  tidal  exchange;  (4)  prevention 
of  introduction  of  fine-grained  materials 
which  might  clog  the  aerial  roots,  such  as 
clay,  and  sugar  cane  wastes;  (5)  preven- 
tion of  oil  spills  and  herbicide  spray 
driftage;  and  (6)  prevention  of  increased 
wave  action  or  current  velocities  from 
boat   wakes,    and  sea   walls. 

Where  mangroves  have  been  destroyed, 
they  can  be  replanted  or  suitable  alter- 
nate areas  can  be  planted,  acre  for  acre, 
through  mitigation  procedures  (see  Lewis 
et  al.  1979).  An  extensive  body  of 
literature  exists  concerning  mangrove 
planting  techniques  in  Florida  (Savage 
1972;  Carlton  1974;  Pulver  1976;  Teas 
1977;  Goforth  and  Thomas  1979;  Lewis 
1979b).  Mangroves  were  initially  planted 
in  Florida  at  least  as  early  as  1917  to 
protect  the  overseas  railway  in  the 
Florida    Keys    (Teas    1977). 

Both    red   and   black    mangroves    have 


84 


been  used  in  transplanting.  As  we  men- 
tioned in  section  11,  black  mangroves  seem 
to  have  certain  advantages  over  red  man- 
groves. Properly  designed  plantings  are 
usually  75%  to  90%  successful,  although 
the  larger  the  transplanted  tree,  the 
lower  its  survival  rate  (Teas  1977). 
Pruning  probably  enhances  survival  of 
trees  other  than  seedlings  (Carlton  1974). 
Important  considerations  (Lewis  1979b; 
Teas  1977)  in  transplanting  mangroves  are: 
(1)  to  plant  in  the  intertidal  zone  and 
avoid  planting  at  too  high  or  too  low  an 
elevation,  (2)  to  avoid  planting  where  the 
shoreline  energy  is  too  great,  (3)  to 
avoid  human  vandalism,  and  (4)  to  avoid 
accumulations  of  dead  sea  grass  and  other 
wrack. 

Costs  of  transplanting  have  been 
variously  estimated.  Teas  (1977)  suggests 
$462  an  acre  ($l,140/ha)  for  unrooted 
propagules  planted  3  ft  (0.9  m)  apart, 
$1,017  an  acre  ($2,500/ha)  for  established 
seedlings  planted  3  ft  (0.9  m)  apart  and 
$87,500  ($21  6,130/ha)  for  3  year-old  nur- 
sery trees  planted  4  ft  (1.2  m)  apart. 
Lewis  (1979b)  criticized  Teas'  costs  as 
unrealistically  low  and  reported  a  project 
in  Puerto  Rico  which  used  established 
seedlings  at  a  cost  of  $5,060  an  acre 
($12,500/ha);  he  did  suggest  that  this 
cost  could  be  cut  in  half  for  larger 
projects. 


12.6       ECOLOGICAL  VALUE  OF   BLACK  VS.   RED 
MANGROVES 

One  unanswered  question  of  current 
interest  in  Florida  concerns  the  ecologi- 
cal value  of  black  mangrove  forests  com- 
pared to  intertidal  red  mangrove  forests. 
In  many  respects,  this  is  identical  to  the 
"high  marsh"  versus  "low  marsh"  debate  in 
temperate  wetlands.  One  hypothetical 
argument  which  has  been  presented  fre- 
quently in  court  cases  during  the  past 
decade  suggests  that  black  mangrove 
forests  have  less  ecological  value  than 
red  mangrove  forests  to  both  man  and 
coastal  ecosystems.  This  argument  is 
based  on  an  apparent  lack  of  substantial 
particulate  detritus  export  from  black 
mangrove   forests   above  mean   high   tide  and 


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

The  counter  argument  states  that 
black  mangrove  forests  are  important  for 
the  support  of  wildlife  and  the  export  of 
substantial  quantities  of  dissolved 
organic  matter  (DOM).  Lugo  et  al.  (1980) 
provide  evidence  that  black  mangrove 
forests  do,  in  fact,  export  large  quanti- 
ties of  DOM.  They  point  out  that  (1 ) 
black  mangrove  leaves  decompose  more 
rapidly  than  red  mangrove  leaves  and  thus 
produce  relatively  more  DOM  and  (2)  abso- 
lute export  of  carbon  from  these  forests, 
on  a  statewide  scale,  is  equal  or  greater 
than   from   red  mangrove  forests. 


12.7     THE   IMPORTANCE  OF    INTER-COMMUNITY 
EXCHANGE 

From  previous  discussions  (sections  6 
and  7.5  and  Appendices  B,  C,  D  and  E)  it 
is  clear  that  many  species  of  fishes, 
invertebrates,  birds,  and  mammals  move 
between  mangrove  forest  communities  and 
other  habitats  including  sea  grass  beds, 
coral  reefs,  terrestrial  forests,  and  the 
freshwater  Everglades.  For  example,  the 
gray  snapper,  Lut janus  gri  seus,  spends 
part  of  its  juvenile  life  in  sea  grass 
beds,  moves  to  mangrove-lined  bays  and 
rivers,  and  then  migrates  to  deeper  water 
and  coral  reefs  as  an  adult  (Croaker  1962; 
Starck  and  Schroeder  1971).  The  pink 
shrimp,  Penaeus  duorarum,  spends  its  juve- 
nile life  in  mangrove-lined  bays  and 
rivers  before  moving  offshore  to  the 
Tortugas  grounds  as  an  adult.  During  its 
juvenile  period  it  appears  to  move  back 
and  forth  from  mangrove-dominated  areas  to 
sea  grass  beds.  The  spiny  lobster, 
Panulirus  argus,  as  a  juvenile  frequently 
uses  mangrove  prop  root  communities  as  a 
refuge;  when  nearing  maturity  this  species 
moves  to  deeper  water  in  sea  grass  and 
coral  reef  communities  (see  discussion 
section  6.1).  Many  of  the  mammals  (sec- 
tion 10)  and  birds  (section  9)  move  back 
and  forth  between  mangrove  communities  and 
a  variety  of  other  environments. 

These     are     only     a     few     of     many 


85 


examples.  Clearly,  mangrove  ecosystems 
are  linked  functionally  to  other  south 
Florida  ecosystems  through  physical  pro- 
cesses such  as  water  flow  and  organic 
carbon  flux.  As  a  result,  the  successful 
management  and/or  preservation  of  many 
fishes,  mammals,  birds,  reptiles,  and 
amphibians  depends  on  proper  understanding 
and  management  of  a  variety  of  ecosystems 
and  the  processes  that  link  them.  Saving 
mangrove  stands  may  do  the  gray  snapper 
little  good  if  sea  grass  beds  are 
destroyed.  Pink  shrimp  populations  will 
be  enhanced  by  the  preservation  of  sea 
grass  beds  and  mangrove-lined  waters,  but 
shrimp  catches  on  the  Tortugas  grounds 
will  decline  if  freshwater  flow  from  the 
Everglades  is  not  managed  carefully  (Idyll 
et  al.  1968).  Successful  management  of 
south  Florida  mangrove  ecosystems, 
including  their  valuable  resources,  will 
depend  on  knowledgeable  management  of  a 
number  of  other  ecosystems  and  the 
processes  which  link  them. 


12.8     MANAGEMENT  PRACTICES:      PRESERVATION 


Based  on  years 
Florida    and    based 


of  research  in  south 
on    the    information 


reviewed  for  this  publication,  we  have 
concluded  that  the  best  management  prac- 
tice for  all  types  of  Florida  mangrove 
ecosystems  is  preservation.  Central  to 
this  concept  is  the  preservation  of 
adjacent  ecosystems  that  are  linked  signi- 
ficantly by  functional  processes.  The 
continued  successful  functioning  of  the 
mangrove  belt  of  southwest  Florida  is 
highly  dependent  on  the  continual  exis- 
tence of  the  Everglades  and  Big  Cypress 
Swamp  in  an  ecologically  healthy  condi- 
tion. 

At  no  cost  to  man,  mangrove  forests 
provide  habitat  for  valuable  birds,  mam- 
mals, amphibians,  reptiles,  fishes,  and 
invertebrates  and  protect  endangered 
species,  at  least  partially  support  exten- 
sive coastal  food  webs,  provide  shoreline 
stability  and  storm  protection,  and 
generate  aesthetically  pleasing  experi- 
ences (Figure  18).  In  situations  where 
overwhelming  economic  pressures  dictate 
mangrove  destruction,  every  effort  should 
be  made  to  ameliorate  any  losses  either 
through  mitigation  or  through  modified 
development  as  described  by  Voss  (1969) 
and  Tabb  and  Heald  (1973)  in  which  canals 
and  seawalls  are  placed  as  far  to  the  rear 
of  the  swamp  as  possible. 


86 


Figure  18.  Mangrove  islands  in  Florida  Bay  near  Upper  Matecumbe  Key.  Note  the 
extensive  stands  of  seedling  red  mangroves  which  have  become  established  (1981) 
after  a  long  period  without  major  hurricanes.  Mangrove  islands  in  the  Florida 
Keys  tend  to  expand  during  storm- free  intervals. 


87 


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105 


APPENDIX  A.   Summary  of  the  site  characteristics  and  sampling 

methodology  for  fishes  in:  A-l  -  mangrove- fringed 
tidal  streams  and  rivers,  A-2  -  mangrove-lined 
estuarine  bays  and  lagoons,  and  A-3  -  mangrove- 
lined  oceanic  bays  and  lagoons . 


106 


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109 


APPENDIX  B.   Fishes  of  mangrove  areas  of  Florida  tabulated  by 
habitat  type.   Key  to  numbered  references  appears 
at  the  end  of  the  table.   Diet  items  listed  in 
order  of  decreasing  importance. 


110 


Family  and  Species 


Habitat  Type 


B 

h  a 


*J  >i  at  >, 


Reference 


Diet 


Diet 
Reference 


Comments 


Orectolobidae  - 
carpet  sharks 

Ginglymostoma  cirratum 

nurse  shark 


5,  7 


Fish,  cephalopods,  molluscs, 
shrimp,  sea  urchins 


Randall  1967 
Clark  &  von 
Schmidt  1965 
Bohlke  & 
Chaplin  1968 


Carcharhinidae  - 
requiem  sharks 

Carcharhinus  leucas 

bull  shark 


Carcharhinus  limbatus 


blacktip  shark 


Negaprion  brevirostris 
lemon  shark 


11 


4,  5,  7 


Juveniles:  fish  (Arius  felis, 
Lophoqobius ,  Mugil  cephalus , 
Brevoortia  patronus, 
Micropogon  undulatus) ,  crus- 
taceans including  penaeid 
shrimp,  blue  crabs 

Fish  (Caranx  sp.,  Centropomus 
undecimalis,  Chilomycterus 
schoepfi,  Arius  felis,  Lacto- 
phrys  trigonnus  Lagodon 
rhomboides) ,  crabs 

Young:  crustaceans,  fish 
Adults:  fish,  crustaceans 


Odum  1971 


Clark  & 
von  Schmidt 
1965 


Randall  1967 
Clark  &  von 
Schmidt  1965 


Sphyrnidae  -  hammerhead 
sharks 

Sphyrna  tiburo  - 

bonnethead 


+    +      2,  5 


Mantis  shrimp,  shrimp,  isopods,  Bohlke  s 
barnacles,  bivalve  molluscs,    Chaplin  1968 
cephalopods,  fish 


Pristidae  -  sawfishes 
Pristis  pectinata 
smalltooth  sawfish 


+      5,  15 


Fish,  benthic  crustaceans 


Bohlke  s 
Chaplin  1968 


Rhinobatidae  - 
guitarfishes 

Rhinobatos  lenti- 


guitarfish 

Torpedinidae  -  electric 
rays 

Narcine  brasiliensis • 


+        1 


+    +    1,  17,  18 


lesser  electric  ray 

Rajidae  -  skates 
Raja  texana 
roundel  skate 


Crustacea,  fish,  annelids 


Reid   1954 


Dasyatidae  -  stingrays 
Dasyatis  americana  - 
southern  stingray 


Dasyatis  sarjina 
Atlantic  stingray 


+    +    2,  4,  5,  7 


2,  8,  13, 

17 


Fishes,  sipunculid  and  poly- 
chaete  worms ,  crabs ,  bivalves 
shrimp ,  mantis  shrimp 

Benthic  invertebrates  inclu- 
ding bivalves ,  xanthid  and 
portunid  crabs ,  shrimps , 
amphipods,  annelids,  chirono- 
mid  larvae 


Randall   1967 


Darnell   1958 


Gymnura  micrura  - 
smooth  butterfly 
ray 

Urolophus  jamaicensis  ■ 
yellow  stingray 


17       Fish,  molluscs,  annelids, 
shrimp,  other  small 
crustaceans 

1       Probably  small  burrowing 
invertebrates 


Peterson  s 
Peterson  1979 


Bohlke  s 
Chaplin  1968 


This  and  all  subsequent  Odum  1971  citations  refer 


to  W.E.  Odum  1971. 
Ill 


Family  and  Species 


Habitat  Type 


*->  >   <y  >. 


Reference 


Diet 
Reference 


Comments 


Myliobatidae  -  eagle  rays 
Aetobatus  narinari  - 
Spotted  eagle  ray 

Lepisosteidae  -  gars 

Lepisosteus  platyrhincus  -  + 
Florida  gar 


2,  1, 

15 


13, 


Clams,  oysters 


Boh Ike  & 
Chaplin  1968 


Fish  (poeciliids,  cyprinodonts ,   Odum  1971 
small  centrarchids) ,  crustaceans 
(caridean  shrimp), insect  larvae 


Elopidae  -  tarpons 
El ops  saurus  - 
ladyfish 


2,  3,  7, 

8,  13,  15 


<  45  mm:  zooplankton,  chaeto- 

gnaths,  polychaete 

worms 
>  45  mm:  caridean  &  penaeid 

shrimp,  various  small 

fish 


Odum  1971 
Austin  & 
Austin   1971 


Mega lops  atlantica  - 
tarpon 


,    8,    13,    <  45  mm:  plankton  (cyclopoid 
15  copepods) 

juveniles:  fish  (Gambusia, 
Fundulus  heteroclitus ,  Mugil 
cephalus) ,  crustaceans  (ostra- 
cods,  caridean  shrimp) 
adults:  wide  variety  of  fish, 
crabs ,  shrimp ,  ctenophores , 
insects 


Odum  1971 
Austin  & 
Austin   1971 


Obligate  air 
breathe  rs .  Juv- 
eniles  inhabit 
shallow  brackish 
pools  low  in  oxygen, 
often  containing 
H,S   (Wade   1962) 


Albulidae  -  bone  fishes 
Albula  vulpes  - 
bonefish 

Anguillidae  -  eels 
Anguilla  rostrata  - 
American  eel 


Ophichthidae  -  snake 

eels 

Myrophis  punctatus 
speckled  worm  eel 


4,  5 


8,  13 


2,  3,  17, 
18 


Clams,  snails,  shrimp,  small 
fish 


50-200  mm:  amphipods,  isopods 
180-472  mm:  xanthid  crabs,, 
caridean  shrimp,  fish 
( Lophogobi us  cyprinoides) 


Polychaetes,  Branch iostoma 
caribaeum,  sand  crabs 


Bohlke  & 

Chaplin 

1968 

Odum  1971 


Springer  & 
Woodburn 
1960, 
Reid   1954 


Members  of  this 
family  burrow 
in  mud  or  sand, 
undersampled  by 
most  methods 
(Bohlke  &  Chaplin 
1968) 


Bascanichthys  scuti- 
caris  -  whip  eels 

Ophichthus  gome si  - 
shrimp  eel 

Clupeidae  -  herrings 
Brevoortia  smithi  - 
yellowfin  sardine 


3,  17 


2,  5,  17 


Brevoortia  patronus 
Gulf  menhaden 


38-48  mm:  phy toplankton ,  zoo-    Darnell 

plankton,  plant  fragments ,       1958 

detritus 

85-103  mm:  organic  matter,  silt, 

diatoms,  foraminiferans,  copepods 


112 


Family  and  Species 


Habitat  Type 


a 


Reference 


Diet 


Diet 

Reference 


Harengula  pensacolae 
scaled  sardine 


2,  3,  8,   30  mm:   planktonic  copepods, 
13      zoea,  nauplii,  larval  fish 
64-96  mm:   amphipods, 
harpactlcoid  copepods,  isopods 
mysids,  chironomid  larvae 


Odum  1971 


Opisthonema  oglinum  - 
Atlantic  thread  herring 


2,  3,  5, 
13,  17 


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


Odum  1971 


Sardinella  anchovia 
Spanish  sardine 


17 


Engraulidae  -  anchovies 
Anchoa  cub ana  - 
Cuban  anchovy 


2,  16    Ostracods,  copepods 


Springer  & 

Woodburn 

1960 


Anchoa  hepsetus 
striped  anchovy 


2,  3,  13,   32-114  mm:   copepods,  isopods,     Springer  & 
16,  17   mysids,  caridean  shrimp,  small     Woodburn 
bivalves  1960 


Anchoa  lampro taenia 
bigeye  anchovy 

Anchoa  mitchilli  - 
bay  anchovy 


1,  2,  3,    <25  mm:   microzooplankton  Odum  1971 

5,  7,  8,   31-62  mm:   amphipods,  zooplank- 
13,  16-18  ton,  mysids,  ostracods,  plant 

detritus,  copepods,  small  molluscs, 

chironomid  larvae 


Synodontidae  - 

lizardfishes 

Synodus  foetens  - 
inshore  lizardfish 


1-3,  5,  8,  Small  fish,  crabs,  shrimp, 
17,  18   polychaete  worms 


Odum  1971 


Catostomidae  -  suckers 
Erimyzson  sucetta  " 
lake  chubsucker 


A  freshwater 
stray 


Ictaluridae  -  freshwater 
catfish 

Ictalurus  natal is  - 

yellow  bullhead 


14 


A  freshwater 
stray 


Noturus  gyrinus 
tadpole  mad torn 


14 


A  freshwater 
stray 


Arriidae  -  sea  catfishes 

Arius  felis  -  sea  catfish   + 


2,  3,  5,   100  mm:   copepods,  zooplankton 
7,  8,  13,  amphipods,  mysids,  chironomid 
17        larvae,  isopods,  small  crabs 
100-200  mm:   benthic  inverte- 
brates 

200-330  mm:   crabs,  amphipods, 
mysids,  fishes,  bark,  crayfish, 
caridean  and  penaeid  shrimp 


Odum  1971 


Bagre  marinus  - 
gafftopsail  catfish 


2,  8,  17 


262-445  mm:   blue  crabs,  small 
fishes 


Odum  1971 


Batrachoididae  -  toadfishes 
Opsanus  beta  - 
Gulf  toadfish 


1-3,  5,    18-60  mm:  amphipods,  chironomid 
7,  12,  13,  larvae,  mysids,  isopods,  few  fish 
15,  17,  18  >60  mm:   caridean  shrimp,  xanthid 

crabs,  snapping  shrimp,  mussels, 

fish,  mangrove  bark 


Odum  1971 


Salinities 
10  o/oo  > 
(Odum  1971) 


113 


Family  and  Species 


at  Type 


Htn  ma  o  <o 


Reference 


Diet 
Reference 


Comments 


Porlchthys  porosi6simus 
Atlantic  midshipman 


3,  18 


Gobiesocidae  -  clingfishes 
Gobiesox  strumosus 
skilletfish 


2,  3,  5,  8  10-32  mm:   amphipods,  isopods, 
chironomid  larvae 


Odum  1971 


Ogcocephalidae  -  batfishes 
Ogcocephalus  nasutus 
shortnose  batfish 

Ogcocephalus  radiatus 
polka-dot  batfish 


18 


Small  bivalves,  gastropods, 
polychaetes 


2,  11,  17, 
18 


Reid  1954 


Gadidae  -  codfishes 
Urophycis  floridanus 
Southern  hake 


Ophidiidae  -  cusk-eels, 
brotulas 

Gunterichthys  longipenis 

gold  brotula 

Ogilbia  cayorum 
key  brotula 

Ophidion  holbrooki 
bank  cusk-eel 


12 


17 

1,  3 

3 


Amphipods,  isopods,  mysids,  Springer  &    A  species  more 

decapod  shrimp,  polychaetes,  Woodburn      common  at  more 

insect  larvae,  fishes  (Lagodon  1960         northerly 
rhomboides,  Paralichthys  latitudes 

alblgutta) 


Exocoetidae  -  flying- 
fishes  ,  halfbeaks 

Chriodorus  atherinoides 


hardhead  halfbeak 


Hyporhamphus  unifasciatus 
halfbeak 


3,  5     juveniles:  zooplankton  including 
crab  megalops,  veligers,  cope- 
pods 

130-199  mm:  epiphytic  algae, 
detritus,  seagrass 


Carr  £ 
Adams 


1973 


Belonidae  -  needlefishes 
Strongylura  marina 
Atlantic  needlefish 


Strongylura  notata 
redfin  needlefish 


7,  15 


2,  3,  5, 
8,  13 


357-475  mm:  small  fishes,         Darnell 
insects,  shrimp,  small  amounts    1958 
of  vascular  plant  material  and 
algae 

In  grassbeds  -  Brook 

Juveniles:  polychaete  worms,      1975 
cumaceans ,  fish 
Adults:  fish,  primarily 
atherinids 


Strongylura  timucu 
timucu 


Tylosurus  crocodilus  ■ 
houndf ish 


2,  3,  11    159-378  mm:  anchovies,  shrimp 


11       250-1320  mm:  fishes,  shrimp 


Randall 

Primarily  inshore 

1967 

species, freely 

enters  fresh- 

water (Randall 

1967) 

Randall 

Open  water  and 

1967 

inshore  surface 

water  inhabitant 

{Voss  et  al . 

1969) 

114 


Habitat  Type 


Family  and  Species 


Reference 


Diet 


Diet 
Reference 


Comments 


Cyprinodontidae  -  killi- 

fishes 

Adinia  xenica  - 
diamond  killifish 


Cyprinodon  variegatus 
sheepshead  minnow 


Floridichthys  carpio 
goldspotted  killifish 


Fundulus  confluentus 
marsh  killifish 


Fundulus  chrysotus 
golden  topminnow 


Fundulus  grandis 
Gulf  killifish 


Fundulus  heteroclitus 
Mummichog 


Fundulus  seminolis 
Seminole  killifish 


Jordanella  floridae 
flagfish 


Lucania  goodei 
bluefin  killifish 


Lucania  parva 
rainwater  killifish 


Rivulus  marmoratus 
rivulus 


2,8,  13-15  Plant  detritus,  diatoms, 
amphipods ,  harpacticoid 
copepods,  insects 


2,  7,  8, 
13-15 

2,  3,  8, 
13 


Plant  detritus,  algae, 
nematodes,  small  crustaceans 


Odum  1971 


Odum  1971 


Amphipods,  ostracods,  isopods,   Odum  1971 
copepods,  chironomid  larvae, 
nematodes,  plant  detritus,  algae 


2,  8,  13-15  Caridean  shrimp,  small  fish, 

(Gambusia  aff inis) ,  amphipods , 
isopods,  adult  &  larval  insects, 
copepods,  mysids,  ostracods, 
algal  filaments 

14,  15 


Odum  1971 


Rare  in  mangrove 
zone,  headwater 
pools  only 


Odum  1971 


2,8,  13-15  Amphipods,  isopods,  xanthid 
crabs,  chironomid  larvae, 
terrestrial  insects,  snails, 
algae,  small  fish  (poeciliids) 

7       Small  crustaceans  (amphipods,    Peterson  & 
isopods,  ostracods,  tanaids,     Peterson 
copepods) ,  detritus,  polychaete   1979 
worms,  insects,  snails,  inver- 
tebrate eggs 

14-15 


13-15 


2,  8,  13-15  Small  crustaceans  (copepods, 

cladocerans,  ostracods),  insect 
larvae 


1-3,  5,  8, 
13-15,  17 


3,  8,  13,  15 


<20  mm:  planktonic  copepods 
21-37  nun:  amphipods,  mysids, 
chironomid  larvae,  ostracods, 
molluscs,  plant  detritus 


Odum  1971 


Odum  1971 


Primarily  a 
freshwater  form, 
headwater  pools 
only 

Primarily  fresh- 
water ,  common  in 
pools  in  headwater 
regions 

Headwater  pools 
and  channel 


Poeciliidae  -  livebearers 
Gambusia  affinis 
mosquitofish 


Gambusia  rhizophorae 
mangrove  gambusia 


2,  3,  7, 
13-15 


6,  9 


A  versatile  feeder:  amphipods, 
chironomid  larvae,  hydracarina, 
harpacticoid  copepods,  snails, 
ants,  adult  insects,  polychaete 
worms,  ostracods,  mosquito  pupae 
algae 


Odum  1971 


Fresh  and  brackish 
water  in  Rhizophora 
swamps ,  northern 
Cuba,  southeastern 
Florida 


115 


Family  and  Species 


Habitat  Type 


Reference 


Diet 
Reference 


Heterandria  forroosa 
least  killifish 


Poecilia  latipinna 
sailfin  molly 

Atherinidae  -  silversides 
Allanetta  harringtonensis 
reef  silver side 

Membras  martinica 
rough  silverside 


Menidia  beryl ling 
tidewater  silverside 


8,  14,  15   Chironomid  larvae,  harpacticoid  Odum  1971 
and  planktonic  copepods,  clado- 
cerans,  terrestrial  insects, 
algae,  diatoms 

5,  7,  8,    Plant  detritus,  algae,  diatoms   Odum  1971 
13-15 


39-60  mm:  copepods,  fish  larvae,  Randall 
polychaete  larvae  1967 


2,  5,  11 


2,  3,  8, 

11,  12, 

13,  17, 

18 


Small  zooplankton  crustaceans,  Peterson  & 

juvenile  &  larval  fishes,  Peterson 

insects,  detritus,  snails  1979 

Insects,  copepods,  chironomid  Odum  1971 
larvae ,  mysids ,  amphipods 


Syngnathidae  - 
pipefishes ,  seahorses 

Corythoichthys 

albirostris 

whitenose  pipefish 


Hippocampus  erectus 
lined  seahorse 


Hippocampus  zoster ae 
dwarf  seahorse 


Micrognathus  crinigerus 
fringed  pipefish 

Syngnathus  f loridae 
dusky  pipefish 

Syngnathus  louisianae 
chain  pipefish 


1,  11 


,  2,  3. 

11. 

17 

.  2,  3, 

5, 

1,    16, 

17 

18 

1,  5,  10     52-82  mm:  copepods,  micro- 
crustaceans 

1-3,  5,  11    Caridean  shrimp,  amphipods, 
tanaids,  isopods 

1-3,  11 ,     Copepods,  amphipods,  small 
16-18      shrimp 


Reid   1954 


Associated  with 
vegetated  areas  (Tabb 
&  Manning  1961) 

Intimately  associated 
with  unattached  algae 
(Tabb  &  Manning 
1961) ,  or  grassy 
areas  (Springer  & 
woodburn  1960) 


Brook  1975 


Reid   1954 


Inhabit  grassy 
flats  (Springer  & 
Woodburn   1960) 


Syngnathus  scovelli 
Gulf  pipefish 


Syngnathus  springe ri 
bull  pipefish 

Syngnathus  dunckeri 
Pugnose  pipefish 


1-3,  5,  11,   Amphipods,  isopods,  tanaids, 
16-18      copepods,  tiny  caridean 

shrimp,  gastropods  (Bittium, 
Mitrella) 

2,  17 


11 


Brook  197  5 
Springer  & 
Woodburn  1960 
Reid  1954 


Associated  with 
vegetated  areas 
(Tabb  S>  Manning 
1961) 


Syngnathus  pelagicus 
sargassum  pipefish 


Centropomidae  -  snooks 
C e n t r opojmus  parallelus 
fat  snook 


Family  as  a  whole 
shows  preference 
for  estuarine  man- 
grove habitat 
(Rivas  1962) 


Centropomus  pectinatus 
tarpon  snook 


7,  8,  13 


116 


Habitat  Type 


Family  and  Species 


u  «  Reference 


Diet 


Diet 
Reference 


Centropomus  undecimalis  + 
snook 


2,  5,  7 ,  8,   Juveniles:  caridean  shrimp,  Odum  1971 

11,  13,  14     small  cyprinodont  fishes,  Austin  & 

gobies,  mojarras  Austin 

Adults:  fish,  crabs,  penaeid  1971 

shrimp,  crayfish,  snapping 

shrimp 


By  far  most 
abundant  of 
three  species 
(Rivas   1962) 


Serranidae  -  sea  basses 
Centropristis  striata 
black  seabass 

Diplectrum  formosum 
sand  perch 

Epinephelus  itajara 
jewfish 


Epinephelus  morio 
red  grouper 


2,  3,  11, 

16-18 


2,  5,  7,  8, 
11,  13,  15 


Family  in  general  carnivorous 
on  fish,  crustaceans 

Caridean  &  penaeid  shrimp, 
copepods,  crabs,  fish 

Juveniles:  penaeid  shrimp, 
xanthid  crabs 


228-340  mm:  crustaceans, 
crabs,  fishes 


Randall 

1967 

Re  id 

1954 

Odum 

1971 

The  most  abundant 

of  the  seabasses 

in  mangrove  habitats 


Randall 
1967 


Epinephelus  striatus 
Nassau  grouper 


11     170-686  mm:  fish,  crabs, 
stomatopods ,  cephalopods , 
shrimp,  spiny  lobsters, 
gastropods,  bivalves,  isopods 


Randall 
1967 


Hypoplectrus  puella 
barred  hamlet 


11      54-98  mm:  snapping  shrimp,       Randall 
crabs,  fish,  mysids,  stomato-    1967 
pods ,  isopods 


Mycteroperca  microlepis 
gag 

Centrarchidae  -  sunfishes 
Elassoma  evergladei 
Everglades  pygmy  sunfish  + 


1,  2,  5,  11,  71-100  mm: 
17,  18    fish 


penaeid  shrimp. 


Reid   1954 


Family  is  primarily 
freshwater ,  fish 
occasionally  enter 
headwater  area 
of  mangrove- 
fringed  stream 


Lepomis  auritus 
redbreast  sunfish 

Lepomis  gulosus 
warmouth 


Shrimp  (Palaemonetes),  fish 
( Gob io soma  bosci ,  Lepomis 
macrochirus) ,  detritus, 
Vallisneria,  amphipods ,  xan- 
thid crabs,  blue  crabs 


Desselle  et 
al.  1978 


Diet  from  Lake 
Pontchartrain 
salinities  1.6- 
4 . 1  o/oo 


Lepomis  macrochirus 
bluegill 


2,  15      Amphipods,  blue  crab  (Cal- 
linectes  sapidus) ,  xanthid 
crabs,  detritus,  Vallisneria, 
clams  (Rangia  cuneata) , 
sponge  (Ephydatia  fluviatilis) , 
barnacles,  insect  larvae 


Desselle  et 
al.  1978 


Diet   from  Lake 
Pontchartrain 
salinities  1.6- 
4.1  o/oo 


Lepomis  microlophus 
redear  sunfish 


2,  13-15     Chironomid  larvae,  amphipods, 
xanthid  crabs,  clam  (Rangia 
cuneata) ,  sponge  (Ephydatia 
fluviatilis) ,  detritus 


Desselle  et 
al.  1978 


Diet  from  Lake 
Pontchartrain 
salinities  1.6- 
4 . 1  o/oo 


Lepomis  punctatus 
spotted  sunfish 


8,  14,  15    Cladocerans,  small  crabs,       odum  1971 
mysids ,  chironomids ,  amphipods , 
insects,  molluscs,  isopods, 
fish,  algae 


Salinities  <  15  o/oo 
(Odum  1971) 


117 


Family  and  Species 


Habitat  Type 


■32 


3 

3  4 
to 


o   «  Reference 


Diet 


Diet 
Reference 


Mlcropterus  salmoldes 
largemouth  bass 


Apogonldae  -  cardlnalflshes 
Astrapogon  alutus 
bronze  cardinalf ish 


13-15 


1.  3 


Caridean  shrimp,  small  blue 
crabs,  crayfish,  xanthld  crabs, 
25  species  of  fish,  Vallisnerla, 
Cladophora 


Darnell  1958 


Astrapogon  stellatus 
conchfish 


Pomatomldae  -  blueflshes 
Pomatomus  saltatrlx 
bluefish 


11      Young:  mainly  fishes  (anchovies, 
sllversldes,  killifishes,  men- 
haden, shad,  spotted  seatrout), 
shrimp,  crabs,  other  small 
crustaceans,  annelids,  snails 


Peterson  & 
Peterson  1979 


Rachycentrldae  -  cobias 
Rachycentron  canadum 
cobia 


+  5,  7,  11   Fish,  crabs 


Randall  1967 


Echeneldae  -  remoras 
Echenels  neucratoides 
vhltefln  sharksucker 


2,  11    Fish,  isopods,  other  Crustacea    Randall  1967 


Members  of  this 
family  attach  to 
sharks  and  large 
bony  fishes 
(Randall  1967) 


Remora  remora 


58-175  mm:  copepods,  Isopods, 
vertebrate  muscle  tissue,  crab 
larvae,  fish  remains,  crusta- 
ceans, amphipods 


Randall  1967 


Carangidae  -  jacks,  pompanos 
Caranx  crysos  -  blue      + 
runner 


+  2,  4,  5, 

7.  11 


Family  of  swift- 
svlmmlng,  carniv- 
orous fishes, 
often  running  in 
schools,  wide- 
ranging  (Randall 
1967) 


Caranx  hippos 
crevalle  Jack 

Caranx  ruber 
bar  jack 

Chloroscombrus  chrysurus 
Atlantic  bumper 

Ollgoplltes  8aurua 
leatherjacket 


Trachlnotus  carolinus 
Florida  pompano 


Trachlnotus  falcatus 
permit 


+  2,  5,  7,   Fishes,  crustaceans 
8,  11,  13 

+    4,  11    160-547  mm:  fish,  shrimp,  myslds, 
stomatopods,  gastropods 

+  2,  11,  17, 
18 

+  2,  3,  5,   Snapping  shrimp,  penaeld  shrimp, 
8,  11,  13  larval  anchovies,  lady fish, 
harpactlcoid  copepods 

+     11      sardines  (Harengula  sp.) , 
mole  crabs  (Hlppa  sp . ) , 
bivalves  (Donax  sp.) 

+    7,  11    15-70  mm:  myslds,  shrimp, 

anchovies,  sllversldes,  crabs, 
snails 


Odum  1971 


Randall  1967 


Tabb  & 
Manning  1961 


Springer  & 

Woodburn 

1960 


Common  over  mud 
bottoms  (Randall 
1967) 


Carr  6  Adams  More  apt  to  occur 
1973         over  sandy  bottoms 
than  T.  carolinas 
(Randall  1967) 


Selene  vomer 
lookdown 


+    +2,3,  7,   Young:  shrimp  and  other 

11      crustaceans,  small  molluscs 


Peterson  6 
Peterson  1979 


118 


Habitat  Type 

Reference 

Diet 

Diet 
Reference 

Family  and  Species 

■o  u 

■H  U 

01 

q 

M 

3 

U  03 

u 

<H 

d 

u  (0 

O  CQ 

Comments 

Hemlcaranx 
amblyrhynchus  - 
bluntnose  Jack 

Caranx  latus 
horse-eye  Jack 


Lutjanidae  -  snappers 
Lut Janus  ana lis 
mutton  snapper 


Lut J anus  apod us 
schoolmaster 


Lut Janus  grlseus 
gray  snapper 


+        + 


+    +    + 


17 


12 


1,  4.  11 


Predaceous  on  other  fishes 


204-620  mm:  crabs,  fish, 
gastropods,  octopods,  hermit 
crabs,  penaeid  shrimp,  spiny 
lobster,  stomatopods 


Darnell  1958 


Randall  1967 


1,  4,  5,   Crustaceans  (shrimp,  snapping 
7,  11     shrimp,  blue  crabs,  xanthid 
crabs,  grapsid  crabs),  fish 


Nugent  1970 


1-5,  7,    <50  mm:  reside  in  grassbeds       Odum  1971 

8,  11-13,   feeding  on  small  crustaceans, 

15-18      insect  larvae 

95-254  mm:  reside  in  mangrove 
creeks  feeding  on  crustaceans 
(snapping  shrimp,  xanthid  crabs, 
penaeid  shrimp,  crayfish,  caridean 
shrimp),  fish  including  gobies, 
anchovies,  poeciliids,  eels, 
killiflshes 


Considered  by 
Gunter  (1956)  to 
be  euryhaline 


Commonly  found 
over  sand,  sea- 
grass,  rubble, 
coral  reefs 
(Randall  1967) 


By  far  the  most 
abundant  snapper 
in  mangrove 
habitats 


Lut Janus  Jocu 
dog  snapper 


Lutjanus  synagris 
lane  snapper 


Gerreidae  -  moj arras 

Diapterus  olisthostomus 
Irish  pompano 


Diapterus  plumleri       +    +    + 
striped  mojarra 


Eucinostomus  argenteus   +    +    + 
spotfin  mojarra 


Eucinostomus  gula 
silver  Jenny 


Eucinostomus  lefroyi 
mottled  mojarra 

Gerres  clnereus 
yellowfin  mojarra 


Pomadasyidae  -  grunts 


+    +    + 


1  190-630  mm:  fish,  crabs, 
octopods,  spiny  lobster, 
gastropods 

1,  2,  3,    snapping  shrimp,  crabs, 
5,  7,  11,   anchovies,  annelids,  molluscs 
16-18 


2      110-116  mm:  green  algae 
(Enteromorpha  flexuosa, 
Cladophora) ,  Ruppia  maritima, 
blue-green  algae  (Lyngba 
majuscula) 

2,  7,  8,    36-172  mm:  mysids,  amphipods, 
11-13,     harpacticoid  copepods, 
15,  18     chironomid  larvae,  ostracods, 
bivalves,  plant  detritus 

1-5,  7,    19-63  mm:  amphipods, 
8,  11-13,   chironomids,  harpacticoid 
16-18      copepods,  ostracods,  mysids, 
molluscs,  plant  detritus 


1-3,  5, 
7,  8, 
11-13, 
16-18 

10 


2,  7,  11 


19-70  mm:  amphipods,  chironomid 
larvae,  harpacticoid  copepods, 
molluscs,  mysids,  ostracods, 
plant  detritus 


Crabs,  bivalves,  gastropods, 
polychaete  worms,  shrimp, 
ostracods 

Family  carnivorous  though  rarely 
piscivorous 


Randall  1967 


Stark  & 

Schroeder 

1970 


Austin  & 
Austin  1971 


Odum  1971 


Odum  1971 


Odum  1971 


Known  from  brackish 
water  to  depths  of 
220  fathoms 
(Randall  1967) 


A  permanent 
resident  (Odum 
1971) 


Randall  1967, 
Austin  & 
Austin  1971 

Randall  1967 


Most  shelter  on 
coral  reef  by 
day,  feed  on 
grassy  flats  by 
night  (Randall 
1967) 


119 


Habitat  Type 


Family  and  Species 


o  co  Reference 


Diet 


Diet 
Reference 


Comments 


Amsotremus  virginicus 
porkfish 


Haemulon  aurolineatum 


11     112-264  mm:  brittle  stars,  crabs,  Randall 
shrimp,  polychaetes ,  isopods ,     1967 
bivalves,  stomatopods,  gastropods 

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


Haemulon  carbonarium 
Caesar  grunt 


Haemulon  flavolineatum 


French  grunt 


156-273  mm:  crabs,  gastropods,    Randall 
sea  urchins,  chitons,  poly-       1967 
chaetes,  brittle  stars,  sipun- 
culid  worms,  shrimp 

113-228  mm:  polychaetes,  crabs,    Randall 
sipunculid  worms,  chitons,        1967 
holothurians,  isopods,  shrimp, 
bivalves 


Haemulon  parrai 
sailor's  choice 


1,  7     Benthic  invertebrates  including 
shrimp ,  crabs ,  amphipods ,  gas- 
tropods ,  polychaete  worms, 
bivalves 


Randall 
1967 


Haemulon  plumieri 
white  grunt 


Haemulon  album 


1,  2,   11,  18 


margate 


Haemulon  sciurus 
bluestriped  grunt 


Orthopristis  chrysop- 

tera 

pigfish 


Spaxidae  -  porgies 

Archosargus  pr oba toe eph a - 

lus 

sheepshead 


Archosargus  rhomboidalis 
sea  bream 


Calamus  arctifrons 


grass  porgy 


Calamus  calamus 
saucereye  porgy 


I,    3,  5, 
7,  11 


1-3,  5, 
11,  16-18 


130-279  mm:  crabs,  polychaete     Randall 
worms,  sea  urchins,  sipunculid    1967 
worms,  gastropods,  shrimp,  brittle  Reid 
stars;  juveniles:  copepods,  mysids  1954 

Benthic  invertebrates  including   Randall 
crabs,  shrimp,  polychaete  worms,   1967 
amphipods ,  copepods ,  snails , 
bivalves 


Benthic  invertebrates  including 
crustaceans ,  molluscs ,  annelid 
worms 

Juveniles:  16-30  mm:  plankton 
including  copepods,  mysids, 
postlarval  shrimp 

>30  mm:  polychaetes,  shrimp, 
amphipods 


2,  3,  5,  7,  <40  mm:  in  grassbeds  -  copepods , 
8,  11-13 ,  amphipods ,  chironomid  larvae , 
17-18         mysids ,  algae,  molluscs 

>40  mm:  in  mangrove  creeks  - 
mussels,  false  mussels,  crabs, 
snapping  shrimp,  crayfish, 
hydrazoans ,  algae ,  plant 
detritus 
32-85  mm:  in  Puerto  Rico  man- 
groves -  100%  blue-green 
algae  (Lyngbya  mojuscula) 

5,  11     105-220  mm:  seagrasses  Cymodocea   Randall 
&  Thalassia,  algae, crabs,  gas-    1967 
tropods,  invertebrate  eggs, 
bivalves 


Randall 

1967 

Carr  & 

Strong  preference 

Adams 

for  vegetated  sub 

1973 

strate  in  bay 

areas  (Weinstein 

et  al.  1977) 

Odum  1971 

Austin  & 

Austin 

1971 

Usually  seen  in 
mangrove  sloughs, 
rare  on  reefs 
(Randall   1967) 


Copepods ,  amphipods ,  mysids ,      Reid 
shrimp,  bivalves,  gastropods 
(Hitrella,  Bittium) ,  polychaetes 


1954   Associated  with 

grassy  flats  (Tabb  s. 
Manning   1961) 


190-250  mm:  polychaetes,  brittle   Randall 
stars,  bivalves,  hermit  crabs,     1967 
sea  urchins,  gastropods,  chitons 


120 


Habitat  Type 


Family  and  Species 


01 

a 

■H 

«  OJ 

3 

TT   U 

i-(  U 

on  a 

H  W 

Diet 


Diet 
Reference 


Lagodon  rhomboides 
pinfish 


1-3,  5,  7,  B,  In  mangrove  creek  -  scorched 
11,  12,  16-   mussel,  mysids,  amphipods, 
18       false  mussel 

In  Whitewater  Bay  -  100%  plant 

material 


Odum  1971  Strong  preference 
Reid  1954  for  vegetated  sub- 
strate in  bay  areas 
(Weinstein  et  al. 
1977) 


Sciaenidae  -  drums 
Balrdiella  batabana 
blue  croaker 

Balrdiella  chrysura 
silver  perch 


Cynoscion  arenarlus 
sand  seatrout 


Cynoscion  nebulosus 
spotted  seatrout 


3,  11 


Lelostomus  xanthurus 
spot 


Menticirrhus  americanus 
Southern  kingfish 


Menticirrhus  littoralis 
Gulf  kingfish 

Micropogon  undulatus 
Atlantic  croaker 


Pogonlas  cromis 
black  drum 


Sciaenops  ocellata 
red  drum 


+    1-3,  8,  Il- 
ls, 16-18 

Larvae:  copepods,  larval  fish 
(Henidia  beryllina) 

Odum  1971 

127-181  mm:  fish  (Anchoa 
mitchilli) ,  mysids 

2,  12,  17, 

18 

Mostly  fish,  caridean  shrimp, 
mysids,  amphipods,  crab  zoea 

Springer  & 

Woodburn 

1960 

+   1-3,  5,  7, 
8,  11-13, 
15,  17,  18 

<50  mm:  copepods,  planktonic 
Crustacea 

50-275  mm:  fish  (Mugil  cephalus, 
Lagodon  rhomboides,  Eucino- 
stomus  gula,  E.  argenteus, 
Cyprinodon  varieqatus, 
Gobiosoma  robustum,  Anchoa 
mitchilli) 

Odum  1971 

2,  7,  12, 
17-18 

<40  mm:  planktonic  organisms 
>40  mm:  filamentous  algae, 
desmids,  forams,  amphipods. 

Springer  & 

Woodburn 

1960 

2,  11-12, 
17-18 


11,  12 


+    2,  7,  11, 
12,  15 


2,  3j  5,  8, 

11-13,  15, 

17 


Equetus  acuminatus 
high-hat 


mysids,  copepods,  ostracods, 
isopods ,  chaetognaths ,  bi- 
valves, snails,  polychaete 
worms 

Fish,  benthic  crustaceans 


Polychaetes ,  bivalves  (Dona*) , 
sand  crab  Emerita),  razor  clams 

Juveniles:  copepods,  mysids, 
caridean  shrimp,  polychaete 
worms,  insect  larvae,  iso- 
pods, small  bivalves 

<100  mm:  molluscs,  xanthid 

crabs 
>100  mm:  bivalves,  amphipods, 

blue  crabs,  penaeid  shrimp, 

caridean  shrimp 

<10  mm:  planktonic  organisms 

(copepods,  crab  zoea,  larval 

fish) 
34-42  mm:  mysids,  amphipods, 

caridean  shrimp 
>50  mm:  xanthid  &  portunid 

crabs,  penaeid  shrimp, 

small  fish 
308-403  mm:  xanthid  crabs 

68-152  mm:  shrimp  S  shrimp 
larvae,  isopods,  stoma topod 
larvae ,  copepods ,  amphipods 


Springer  & 

Woodburn 

1960 

Springer  &   Most  common  off  sandy 
Woodburn    beaches  (Springer  & 
1960       woodburn  1960) 

Springer  & 

Woodburn 

1960 


Darnell 
1958 


Odum  1971 


Randall 
1967 


Characteristic  of 
coral  reefs 
(Randall   1967) 


121 


Habitat  Type 


Family  and  Species 


a 

•S 

H 

a 

3 

a 

■d 

u 

*-> 

CO    « 

H 

■s> 

u  oa 

Diet 
Reference 


Ephippidae  -  spadefishes 
Chaetodipterus  faber 
■Atlantic  spadef ish 


Pomacentridae  - 
damsel fishes 

Abudefduf  saxatilis 


sergeant  major 


2,  3,  5, 

11,  16-18 


Worms,  crustaceans,  debris       Darnell 

1961 


101-135  mm:  copepods,  algae,     Randall 
fish  eggs,  fish,  shrimp  larvae,   1967 
polychaetes 


Juveniles  (7-12  mm) 
inhabit  very  shallow 
nearshore  sandy 
beaches.   Bear  a 
deceptive  resemblance 
to  infertile  red 
mangrove  seed  pods 
(Breder  1946) 

Characteristic  family 
of  coral  reefs  (Ran- 
dall 1967) 

A  habitat  generalist: 
reefs ,  grassbeds , 
rock  piles,  wharfs 
(Bohlke  &  Chaplin 
1968) 


Labridae  -  wrasses 

Halichoeres  bivittatus 
slippery  dick 


67-153  mm:  crabs,  sea  urchins,   Randall 
polychaetes ,  gastropods ,  brittle  1967 
stars,  bivalves ,  shrimp,  fish, 
hermit  crabs 


Shallow  water  patch 
reefs,  sand  bottoms, 
gr a s sbeds  ( Randa 1 1 
1967) 


Scaridae  -  parrotfishes 
Nicholsina  usta 
emerald  parrotfish 


+   1,  2,  11,  18 


Family  herbivorous,  feeding 
primarily  on  algae  growing 


on  hard  substrates, 
on  seagrasses 


secondarily 


Randal]      Family  characteris- 
1967         tic  of  coral  reefs, 
ranging  into  grass- 
beds 


Scarus  coeruleus 
blue  parrotfish 


Scarus  croicensis 


striped  parrotfish 

Spar i soma  chrysopterum 
redtail  parrotfish 

Sparisoma  rubripinne 
redfin  parrotfish 

Sparisoma  viride 
stoplight  parrotfish 

Mugilidae  -  mullets 
Mugil  cephalus 
striped  mullet 

Mugil  curema 
white  mullet 


11 


+    2,  3,  5,  7, 

11-13,  15 


+    2,  5,  7, 
11-12 


Inorganic  sediments,  fine 
detritus ,  micro-algae 

25-73  mm:  plant  detritus,  blue- 
green  algae  (Lyngbya  majuscula) 


Odum  1971 


Austin  & 

Austin 

1971 


Requires  near  marine 
salinities  {Tabb  & 
Manning   1961) 


Mugil  trichodon 
fantail  mullet 

Sphyraenidae  -  barracudas 
Sphyraena  barracuda 
great  barracuda 

Opistognathidae  -  jawfishes 
Opistognathus  maxillosus 
mottled  jawfish 


+    2,  7,  11,  12 


1-5,  7,  8, 
11,  13 


135-369  mm:  fish  (Eucinostomus 
quia,  Menidia  beryllina,  Archo- 
sargus  probatocephalus) 

53-110  mm:  shrimp,  isopods, 
fishes,  polychaetes,  mysids, 
copepods 


Odum  1971 


Randa 1 1 
1967 


Salinities  >10  o/oo 
(Odum  1971) 


Family  lives  in 
burrows  in  sediment, 
often  in  vicinity 
of  reefs  (Randall 
1967) 


122 


Family  and  Species 


Habitat  Type 


0} 

3 

1 

0) 

3 

M 

*->    >> 

u 

<n    cu 

03 

Cx3     32 

o  «  Reference 


Diet 
Reference 


Clinidae  -  clinids 
Chaenopsis  ocellata 
bluethroat  pikeblenny 


Family  appears  to  be  carnivorous  Randall 
on  benthic  invertebrates         1967 


Inshore  on  rock, 
coral  or  rubble 
substrates  (Ran- 
dall  1967) 


Paraclinus  marmoratus 
marbled  blenny 

Paraclinus  fasciatus 
banded  blenny 

Stathmonotus  hemphilli 
blackbelly  blenny 


+    1,  5,  11 


Blenniidae  -  combtooth 
blennies 

Chasmodes  saburrae 

Florida  blenny 


Blennius  marmoreus 


seaweed  blenny 


1-3,  11,  17,   21-25  mm:  amphipods 
18       25-60  mm:  amphipods, 
polychaetes ,  snails 


detritus, 


Algae,  organic  detritus, 
brittle  stars,  polychaetes, 
hydro ids 


Carr  & 

Adams 

1973 

Randall 
1967 


Common  brackish 
water  blenny  (Tabb 
&  Manning  1961) 


Blennius  nicholsi 
highfin  blenny 

Callionymidae  -  dragonets 
Callionymus  pauciradiatus 
spotted  dragonet 


Eleotridae  -  sleepers 
Dormitator  maculatus 
fat  sleeper 

Gobiidae  -  gobies 

Bathygobius  sopor a tor 
frillfin  goby 

Gobionellus  hastatus 
sharptail  goby 


13,  15 


2,  3,  8,11,  Caridean  shrimp,  chironomid 
17       larvae,  amphipods 


Filamentous  algae  (Entero- 
morpha) ,  ostracods,  copepods, 
insect  larvae 


Odum  1971 


Springer  & 

Woodburn 

1960 


Freshwater  and 
low  salinity  areas 
(Darnell   1961) 


Gobionellus  shufeldti 
freshwater  goby 

Gobionellus  smaragdus 
emerald  goby 


2,  17,  18 


3,  8,  10,  11, 
15 


Gobiosoma  bosci 
naked  goby 


2,  12     Small  crustaceans  including      Petersons 
amphipods,  annelids,  fish,       Peterson 
fish  eggs  1979 


Gobiosoma  longipala 
twoscale  goby 


Gobiosoma  macrodon 
tiger  goby 


Gobiosoma  robustum 
code  goby 

Lophogobius  cyprinoides 
crested  goby 


1-3,  5,  8,    Amphipods,  mysids,  chironomid 
11,  16-18     larvae 

1-3,  7,  8,    A  versatile  feeder:  amphipods, 
13       mangrove  detritus,  filamentous 
algae,  mysids,  caridean  & 
penaeid  shrimp,  polychaete 
worms ,  ostracods ,  bivalves , 
chironomid  larvae,  harpacticoid 
copepods,  isopods,  xanthid 
crabs ,  snails 


Odum  1971 


Odum,  1971 


123 


Family  and  Species 


Habitat 

rype 

a 

u 

a 

h 

•H 

■ 

a 

(Q    (U 

9 

1 

4J     > 

41     >» 

■H    4J 

a>   Q 

O    Q 

H  v. 

W   03 

O   03 

Diet 
Reference 


Comments 


Microgobius  gulosus 
clown  goby 


Microgobius  microlepis 
banner  goby 

Microgobius  thalassinus 
green  goby 


2,5,8,  11-  Amphipods,  copepods,  chironomid  Odum  1971 
13,  15,  17,    larvae 


18 

5 


3,  12 


Planktonic  organisms 


Bird song 
1981 


Small  crustaceans  including      Peterson  & 
amphipods,  other  invertebrates   Peterson 

1979 


Scombridae  -  mackerels, 
tunas 

Scomberomorus  maculatus 

Spanish  mackerel 


2,  11,  12,    Adults  feeding  on  penaeid        Tahb  & 
15       shrimp  migrating  from  tidal      Manning 
stream  1961 


Scomberomorus  cavalla 
king  mackerel 


11 


350-1022 


fish 


Randall 
1967 


Scorpaenidae  -  scorpion- 
fishes 

Scorpaena  brasiliensis 

barbfish 

Scorpaena  grandicornis 
plumed  scorpionfish 


11     Shrimp,  other  crustaceans, 
fish 

1       37-102  ram:  shrimp,  fish, 
unidentified  crustaceans 


1967 

Randall 

Most  often   found 

1967 

in  seagrass 

Triglidae  -  searobins 
Prionotus  salmonicolor 
blackwing  searobin 

Prionotus  scitulus 
leopard  searobin 

Prionotus  tribulus 
blghead  searobin 


+   1-3,  11,     Small  molluscs,  shrimp,  crabs 
16-18       fish,  small  crustaceans 
(ostracods ,  cumaceans) 

+   1-3,  11-13,   Shrimp,  crabs,  fishes,  amphi- 
17 i  18     pods,  copepods,  annelids, 
bivalves,  sea  urchins 


Peterson  & 
Peterson  1979 


Peterson  & 
Peterson  1979 


Bothidae  -  lefteye 
flounders 

Bo thus  ocellatus 

eyed  flounder 

Citharichthys  macrops 
spotted  whiff 

Citharichthys 

spllopterus 
bay  whiff 


i,  n 


1,  17,  18 


68-130  ram:  fish,  crabs,  shrimp, 
amphipods 


Mainly  myslds,  also  shrimp, 
crabs,  copepods,  amphipods, 
fishes,  annelids 


Randall  1967 


Peterson  &   Recorded  from 
Peterson     salinity  range 
1979        2.5-36.7  o/oo 
(Darnell  1961) 


Etropus  crossotus 
fringed  flounder 


Farallchthys  albigutta 
Gulf  flounder 


Parallchthys  lethostlgma 
Southern  flounder 


3,  11,  16 


1-3,  7,  11, 

12,  17,  18 


Calanoid  copepods,  cumaceans,  Peterson  & 

amphipods,  myslds,  shrimp,  Peterson 

crabs,  isopods,  annelids,  1979 
molluscs,  fishes 

<A5  mm:  small  crustaceans.  Springer  & 

including  amphipods,  small  Woodburn 

fish  I960-,  Reid 

>45  mm:  fish  (pigflsh,  plnfish,  1954 
lizardflsh,  bay  anchovy, 
labrids),  crustaceans 

Mainly  fishes  (mullet,  menha-  Peterson  & 

den,  shad,  anchovies,  plnfish,  Peterson 

mojarras,  croakers),  crabs,  1979 
mysids,  molluscs,  penaeid 
shrimp,  amphipods 


124 


Family  and  Species 


Habitat  Type 


Reference 


Diet 


Diet 
Reference 


Comments 


Syacium  papillosum 
dusky  flounder 


Soleidae  -  soles 
Achirus  lineatus 
lined  sole 


1-3,  5,  8, 

11-13,  17- 

18 


32-74  mm:  chironomid  larvae, 
polychaete  worms,  foraminiferans 


Odum  1971 


Trinectes  inscriptus 
scrawled  sole 


Trinectes  maculatus 
hogchoker 


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


14-110  mm:  amphipods ,  mysids 


Odum  1971 


Cynoglossidae  -  tongue- 
fishes 

Symphurus  plagiusa 
blackcheek  tonguefish 


Balistidae  -  triggerfishes 
&  filefishes 

Aluterus  schoepfi 

orange  filefish 


Balistes  vetula 


queen  triggerfish 


Monacanthus  ciliatus 
fringed  filefish 


Monacanthus  hispidus 
planehead  filefish 


Balistes  capriscus 
gray  triggerfish 


1,  3,  11, 
12,  16-18 


1,  11 


35-102  mm:  polychaete  worms, 
ostracods,  portunid  crabs, 
Ruppia  and  Halodule  plant 
tips 


Seagrasses,  algae,  hermit 
crabs,  gastropods 


11       130-480  mm:  sea  urchins,  crabs, 
bivalves,  brittle  stars,  poly- 
chaetes,  hermit  crabs,  gastro- 
pods, algae 

1,  11,  17  47-97  mm:  Algae,  organic  detri- 
tus ,  seagrass ,  copepods,  shrimp 
&  shrimp  larvae,  amphipods, 
tanaids,  polychaetes,  molluscs 

1-3,  11,    Detritus,  bryozoans,  annelids, 
16-18      harpacticoid  copepods ,  amphi- 
pods, hermit  crabs,  molluscs, 
algae,  sea  urchins 


Austin 
Austin 
1971 


Randall 
1967 


Randall 
1967 


Randall, 
1967 

Springer  & 
Woodburn 
1960 

Peterson  & 

Peterson 

1979 


Associated  with 
grassbeds ,  sponge/sea 
fan  habitats  (Ran- 
dall  1967,  Voss 
et  al.   1969) 

Solitary  reef  fish 
ranging  into  grass- 
beds 


Closely  associated 
with  vegetated  areas 
(Tabb  &  Manning 
1961) 

Associated  with 
vegetated  areas  {Tabb 
&  Manning  1961) 


Ostraciidae  -  boxfishes 
Lactophrys  quadracornis 
scrawled  cowfish 


Lactophrys  trigonus 
trunkfish 


Lactophrys  triqueter 
smooth  trunkfish 


Tetraodontidae  -  puffers 
Sphoeroides  nephelus 
southern  puffer 


I,  2,  5,  7, 

II,  16-18 


1,  4,  11 


1-3,  5,  11, 
16-18 


Vegetation ,  algae ,  bivalves 


109-395  mm:  crabs,  bivalves, 
polychaetes,  sea  urchins,  algae, 
seagrass,  gastropods,  amphipods 

93-250  mm:  polychaetes,  sipun- 
culid  worms,  crabs,  shrimp, 
gastropods,  hermit  crabs,  sea 
urchins,  bivalves 


Juveniles :  detritus ,  fecal 
pellets,  zooplankton,  poly- 
chaetes, gastropods,  crabs, 
shrimp 
Adults :  small  crabs ,  bivalves 


Reid  1954  Young  mimic  sea- 
grass blades 
(Bohlke  &  Chaplin 
1968) 


Randall 

Primarily  a  resident 

1967 

of  seagrass  (Randall 

1967) 

Randall 

Primarily  a  reef 

1967 

species  (Randall 

1967) 

Carr  & 

Adams 

1973 


125 


Habitat  Type 


Family  and  Species 


M  3,      Reference 


Diet 


Diet 
Reference 


Comments 


Sphoeroides  spengleri 
bandtall  puffer 


Sphoeroides  testudineus  + 
checkered  puffer 

Diodontidae  -  porcupine- 
fishes 

Chilomycterus  antenna tus 

bridled  burrfish 


Chilomycterus  ant 11 la rum 
web  burrfish 


1»  7,  11   Crabs,  bivalves,  snails, 
polychaetes,  amphipods, 
shrimp 


1,  7     85-92  mm:  portunid  megalops 
larvae,  gastropods 


11      Gastropods,  hermit  crabs, 
isopods,  crabs,  shrimp 


Randall 

Inhabits  sea- 

1967 

grass,  reef, 

rubble,  man- 

groves (Randall 

1967;  Voss  et  al 

1969) 

Austin  & 

Austin  1971 

Randall 

Reefs  and  grass 

1967 

beds  (Voss 

et  al.  1969) 

Chilomycterus  schoepf 1 
striped  burrfish 


1-3,  5,    Gastropods,  barnacles,  crabs, 
11,  16-18  amphipods 


Springer  & 

Uoodburn 

1960 


Associated  with 

grassbeds  (Voss 

et  al.  1969) 

Salinities 

>25  o/oo  (Springer 

&  Woodburn  1960) 


Reference  Numbers  Key 


1.  Bader  &  Roessler  1971 

2.  Carter  et  al.  1973 

3.  Clark  1970 

4.  Holm  1977 

5.  Hudson  et  al.  1970 

6.  Rush  Ian  &  Lodge  1974 

7.  Nugent  1970 

8.  Odum  1971 

9.  Rivas  1969 


10.  Seaman  et  al.  1973 

11.  Schmidt  1979 

12.  Springer  6,   Woodburn  1960 

13.  Tabb  1966 

14.  Tabb,  Dubrow  &  Manning  1962 

15.  Tabb  &  Manning  1961 

16.  Weinstein  et  al.  1977 

17.  Yokel  1975a 

18.  Yokel  1975b 


126 


APPENDIX  C.  Amphibians  and  reptiles  recorded  from  south  Florida  mangrove 
swamps . 


127 


AMPHIBIANS  AND  REPTILES  OF  FLORIDA'S  MANGROVES 


Species 


Status 


Food  Habits 


Mud  Turtle 

(Kinosternon  subrubrum) 


Abundant 


Insects,  crustaceans, 
mollusks 


Striped  Mud  Turtle 
(Kinosternon  bauri ) 


Common 


Algae,  snails,  dead 
fish 


Ornate 

Diamondback  Terrapin 
(Malaclemys  terrapin 
macrospilota  and 
M.t_.  rhizophorarum) 

Florida  Red-bellied  Turtle 
(Chrysemys  nelsoni) 

Chicken  Turtle 

(Deirochelys  reticularia) 

Green  Turtle 

(Chelonia  mydas) 

Hawksbill 

(Eretmochelys   imbricata) 

Loggerhead 

(Caretta  caretta) 

Atlantic  Ridley 

(Lepidochelys  kempii) 

Florida  Softshell 
(Trionyx  ferox) 

Green  Anole 

(Anolis  carolinensis) 

Cuban  Brown  Anole 
(Anolis  sagrei) 

Bahaman  Bank  Anole 
(Anolis  distichus) 

Green  Water  Snake 
(Nerodia  cyclopion) 

Mangrove  Water  Snake 
(Nerodia  fasciata 
compressicauda) 


Uncommon 


Littorina,  Melampus,  Uca . 
Anomalocardia 


Rare  -  Uncommon     Sagittaria,  Lemna,  Naias 


Uncommon 


Uncommon 


Rare 


Common 


Uncommon 


Common 


Common 


Common 


Uncommon 


Common 


Common 


Crayfish,  insects,  Nuphar 


Mangrove  roots  and  leaves , 
seagrasses 

Rhizophora:   fruits,  leaves 
wood ,  bark 

Crabs,  jellyfish,  tuni- 
cates 

Snails,  crabs,  clams 


Snails,  crayfish,  mussels, 
frogs,  fish,  waterfowl 

Insects 


Insects 


Insects 


Fish 


Fish,  invertebrates 


128 


AMPHIBIANS  AND  REPTILES  OF  FLORIDA'S  MANGROVES  (concluded) 


Species 


Status 


Food  Habits 


Striped  Swamp  Snake  Uncommon 

(Liodytes  alleni) 

Eastern  Indigo  Snake  Uncommon 

(Drymarchon  corais) 

Rat  Snake  Uncommon 

(Elaphe  obsoleta) 

Eastern  Cottonmouth  Uncommon 

(Agkistrodon  piscivorus) 

American  Alligator  Common 

(Alligator  mississippiensis) 

American  Crocodile  Rare 

(Crocodylus  acutus) 

Giant  Toad  Common 

(Bufo  marlnus) 


Crayfish,  sirens,  frogs 


Small  mammals,  birds, 
frogs 

Small  mammals ,  birds 


Fish,  frogs,  snakes, 
birds ,  small  mammals 

Fish,  waterbirds 


Fish,  waterbirds 


Invertebrates 


Squirrel  Treefrog 
(Hyla  squirella) 

Cuban  Treefrog 

(Hyla  septentrionalis) 


Abundant 


Common 


Insects 


Insects,  frogs,  toads, 
lizards 


References : 


Carr  and  Goin  1955;  Ernst  &  Barbour   1972; 
Mahmuud  1965;  L.  Narcisse,  R.N.  "Ding"  Darling 
Fed.  Wildlife  Refuge,  Sanibell  Is.,  Fla. ; 
personal  communication  (1981)  . 


129 


APPENDIX  D.  Avifauna  of  south  Florida  mangrove  swamps. 


130 


WADING  BIRDS 


Common  Name 
(Latin  name) 


Season  of 

Occurrence3    Nestinya    Food  Habits 


References 


Great  Egret 

(Casmerodius  albus) 


Howell,  1932 

Kushlan  &  White  1977a 


Snowy  Egret 

(Egretta  thula) 


Fish 


Howell   1932 

Kushlan  6  White  1977a 

Ffrench  1966 


Cattle  Egret 

(Bubulcus  ibis) 


Fish 


Howell   1932 

Kushlan  &  White  1977a 


Great  White  Heron 
(Ardea  herodias 
occidentalis) 


Fish 


Howell   1932 

Kushlan  &  White   1977a 


Great  Blue  Heron 
(Ardea  herodias) 


Howell  1932 

Kushlan  &  White   1977a 


Reddish  Egret 
(Dichromanassa 
ruf escens) 


Howell  1932 

Kushlan  &  White  1977a 


Louisiana  Heron         Common 
(Hydranassa  tricolor) 


Kushlan  s.   White  1977a 
Maxwell  &   Kale  1977 
Girard  &  Taylor  1979 


Little  Blue  Heron 
(Florida  caerulea) 


Kushlan  &  White  1977a 
Maxwell  &   Kale  1977 
Girard  &  Taylor  1979 


Green  Heron  Common 

(Butorides  striatus) 


Fish 


Robertson  &  Kushlan  1974 
Maxwell  &   Kale,  1977 
Girard  &  Taylor  1979 


Black-crowned  Night 
Heron 

(Nycticorax 
nycticorax) 


Fish ,  crustaceans , 
frogs,  mice 


Ffrench  1966 
Maxwell  &  Kale  1977 
Girard  &   Taylor  1979 


Yellow-crowned  Night 
Heron 

(Nyctanassa  violacea) 


Fish,  crayfish, 
crabs 


Ffrench  1966 

Girard  &  Taylor  1979 


Least  Bittern 

(Ixobrychus  exilis) 


Fish 


Ffrench  1966 


American  Bittern 
(Botaurus 
lentiginosus) 


W,T 


Crayfish,  frogs, 
small  fishes 


Narcisse ,  pers .  comm . 


Wood  Stork 

(Mycteria  americana) 


Common 
(locally 
abundant) 


Kahl   1964 

Ogden  et  al.   1976 

Kushlan  1979 


Glossy  Ibis 

(Plegadis  falci- 
nellus 


Uncommon 


Fish 


Bacon  1970 
Howell  1932 


White  Ibis 

(Eudocimus  albus) 


Fish,  crabs  (Uca) 


Kushlan  1979 

Kushlan  &  Kushlan  1975 

Girard  &  Taylor   1979 


Roseate  Spoonbill 
(Ajaia  ajaja) 

Sandhill  crane 

(Grus  canadensis) 


Rare  to 
Uncommon 


Shrimp,  fish, 
aquatic  vegetation 

Roots,  rhizomes  of 
Cyperus  &  Sagit- 

taria 


Kushlan  &  White  1977a 
Howell   1932 

Ogden  1969 
Howell   1932 


Limpkin 

(Aramus  guar a una) 


Snails  (Pomacea) 


Howell   1932 
Bacon  1970 


131 


PROBING  SHORE  BIRDS 


Common  Name 
(Latin  name) 


Abundance 


Season  of 
Occurrence3 


Nesting* 


Food  Habits 


References 


King  Rail 

(Rail us  elgans) 


Clapper  Rail 

(Rallus  longiro- 
stris) 

Virginia  Rail 

(Rallus  limicola) 

Sora 

(Porzana  Carolina) 


Black  Rail 
(Laterallus 
jamaicensis) 

Semipalmated  Plover 
(Charadrius  semi  - 
palmatus) 

Wilson's  Plover 

(Charadrius  wilsonia) 

Black-bellied  Plover 
(Pluvialis 
squatarola) 

Ruddy  Turnstone 

(Arenaria  lnterpres) 

Common  Snipe 

(Capella  gallinago) 


Common 


Uncommon- 
common 


Rare 


Uncommon  to 

locally 

abundant 


Rare 


Locally 
common 


Locally 
common 


Yr 


W,T 


Uncommon 


W,T 


Beetles,  grass- 
hoppers, aquatic 
bugs 

Crabs,  shrimp 


Beetles,  snails, 
spiders 

Insects,  seeds  of 
emergent  aquatic 
plants 

Beetles,  snails 


Crustaceans, 
mollusks 


Crabs,  shrimp, 
crayfish 

Crabs ,  mollusks 


Insects,  crus- 
taceans ,  mollusks 

Mollusks,  insects, 
worms 


Narcise,  pers.  comm. 
Martin  et  al.  1951 


Howell  1932 
Ffrench  1966 
Bacon  1970 

Marcisse,  pers.  coram. 
Martin  et  al.  1951 

Howell  1932 
Bacon  1970 


Narcisse,  pers-  coram. 


Ffrench  1966 

Bacon  1970 

Baker  &  Baker  1973 

Howell  1932 
Bacon  1970 

Howell  1932 
Bacon  1970 
Ffrench  1966 

Ogden  1969 
Howell  1932 

Howell  1932 
Bacon  1970 


Long-billed  Curlew 

(Numenlus  americanus) 

Whimbrel 

(Numenius  phaeopus) 


Spotted  Sandpiper 
(Ac tit is  macularia) 


Solitary  Sandpiper 
(Tringa  solitaria) 

Willet 

(Catoptrophorus 
semipalmatus) 

Greater  Yellowlegs 
(Tringa 
melanoleucas) 

Lesser  Yellowlegs 
(Tringa   flavipes) 


Rare-uncommon    WtT 


Abundant 


Common 


Common 


Red  Knot 

(Calidrls  canutus) 

Dunlin 

(Calidris  alplna) 


White-rumped  Sandpiper   Rare 
(Calidris  fusclcollls) 


Common 

Uncommon 
Common 


W,T 


W,T 


Yr 


W,T 


W,T 


W,T 


Crustaceans, 
insects 

Mollusks,  crus- 
taceans ,  worms , 
insects 

Mollusks,  crus- 
taceans 


Ogden  1969 


Ogden  1968 
Howell  1932 


Ffrench  1966 
Bacon  1970 
Russel  1980 


Crustaceans,  aquatic  Howell  1932 
insects,  small  frogs  Bacon  1970 

Crabs,  crayfishes,    Howell  1932 
killifishes  Bacon  1970 


Fishes,  crabs , 
crustaceans 


Snails,  mollusks, 
crabs 


Marine  worms , 
crustaceans 

Marine  worms, 
mollusks 


Howell  1932 
Ffrench  1966 
Bacon  1970 

Ffrench  1966 

Bacon  1970 

Baker  &  Baker  1973 

Howell  1932 
Ogden  1964 

Ogden  1964 

Baker  &  Baker  1973 


Chironomids,  snails  Howell  1932 
Bacon  1970 


132 


PROBING   SHOREBIRDS    (concluded) 


Common  Name 
(Latin  name) 

Abundance 

Season  of 
Occurrence3 

Least  Sandpiper 

(Calidris  minutilla) 

Common 

W,T 

Short-billed  Dowitcher 
(Limnodromus  griseus) 

Common 

W,T 

Stilt  Sandpiper 
(Micropalama 
himantopus) 

Rare-uncommon 

W,T 

Semipalmated  Sandpiper 
(Calidris  pusilla) 

Common- 
abundant 

W,T 

Western  Sandpiper 
(Calidris  mauri) 

Common- 
abundant 

W,T 

Marbled  Godwit 
(Limosa  fedoa) 

Rare-common 

W 

Nesting3 


Food  Habits 


References 


American  Avocet 
(Recurvlrostra 
americana) 


Uncommon 


W,T 


Pupae  of  beetles 
and  flies 

Mollusks, 
crustaceans 

Chironomids 


Mollusks,  insects 


Chironomids 


Crustaceans, 
mollusks,  seeds  of 
emergent  aquatic 
plants 

Marine  worms, 
aquatic  insects 


Bacon  1970 

Baker  &  Baker  1973 

Bacon  1970 

Baker  6.  Baker  1973 

Howell  1932 
Bacon  1970 


Bacon  1970 

Baker  &  Baker  1973 

Howell  1932 
Bacon  1970 

Howell  1932 


Ogden  1969 


Black-necked  Stilt       Common 
(Himantopus  mexicanus) 


Aquatic  beetles 


Howell  1932 
Bacon  1970 


133 


SURFACE  AND  DIVING  BIRDS 


Common  Name 

Season  of 

(Latin  name) 

Abundance 

Occurrence' 

Common  Loon 

Occasional 

w 

(Gavia  immer) 

Horned  Grebe 

Uncommon 

w 

(Podiceps  auritus) 

Pied-billed  Grebe 

Uncommon- 

Yr 

(Podilymb'us 

common 

podiceps) 

White  Pelican 

Rare 

S 

(Pelecanus 

Common 

w 

ery throrhy nchos ) 

Brown  Pelican 

Common 

Yr 

(Pelecanus 

occidentalia) 

Double-crested 

Common 

Yr 

cormorant 

(Phalacrocorax 

auritus) 

Anhinga 

Common 

Yr 

(Anhinga  anhinga) 

Fulvous  Whistling  Duck 

Uncommon 

w 

(Dendrocygna 

bicolor) 

Mallard 

Uncommon 

W,T 

(Anas  platyrhynchos) 

Black  Duck 

Rare 

W,T 

(Anas  rubripes) 

Mottled  Duck 

Uncommon 

Yr 

(Anas  fulvigula) 

Gad wall 

Uncommon 

W,T 

(Anas  strepera) 

Pintail 

Abundant 

W,T 

(Anas  acuta) 

Green-winged  Teal 

Abundant 

W,T 

(Anas  crecca  carollnensis) 

Blue-winged  Teal 

Abundant 

Yr 

(Anas  discors) 

Common 

American  Wigeon 

W,T 

(Anas  americana) 

Northern  Shoveler 

Common 

W,T 

(Anas  clypeata) 

Wood  Duck 

Rare 

W 

(Aix  sponsa) 

Redhead 

Rare 

W 

(Aythya  americana) 

Nesting' 


Food  Habits 


References 


Fish,  crabs,  mollusks   Narcisse,  pers.  co 


Fish,  aquatic  insects,   Ogden  1969 
mollusks 


Crayfish,  fish, 
mollusks 


Widgeon  grass 


Mollusks,  crusta- 
ceans, widgeon  grass 

Polygonum,  snails, 
Ruppia 

Ruppia ,  Zostera, 
mollusks 

Saggitaria,  mollusks, 
Cyperus 

Ruppia ,  Zostera.- 
aquatic  insects 

Cyperus ,  snails, 
insects,  crustaceans 

Ruppia ,  Zostera, 
mollusks 

mollusks,  aquatic 
insects,  Ruppia, 
Zostera 

Nuts,  seeds 


Narcisse,  pare,  coram. 


Narcisse,  pers.  conm. 


Ffrench  1966 
Bacon  1970 


Kushlan  &  White  1977a 


Ffrench   1966 


Ogden  1969 
Smith,  pers.  obs . 


Ogden   1969 

Kushlan  et  al.,  in  prep. 

Ogden  1969 


LaHunt  &  Cornwell   1970 
Kushlan  et  al.,  in  prep. 

Ogden   1969 


Narcisse,  pers.  comm. 
Kushlan  et  al.,in  prep. 


Narcisse,  pers.  comm. 

Kushlan  et  al.,  in  prep. 

Narcisse,  pers.  coram. 
Ffrench   1966 

Narcisse,  pers.  comm. 

Kushlan  et  al.  ,  in  prep. 

Narcisse,  pers.  comm. 


Ogden   1959 


Snails,  clams,  aquatic  Ogden  1969 

insects,  Ruppia,  Zos- 

tera 


Ring-necked  Duck  Abundant 

(Aythya   collar is) 


Polygonum,    Ruppia , 
crayfish,    snails 


Ogden  1969 

Kushlan  et  al.,  in  prep. 


Canvasback  Uncommon 

(Aythya  valiaineria) 


Vallisneria,  Ruppia , 
Zostera 


Ogden   1969 

Kushlan  et  al.,  in  prep. 


134 


SURFACE  AND  DIVING  BIRDS    (concluded) 


Common  Name 
(Latin  name) 


Abundance 


Season  of 
Occurrence 


Nesting ' 


Food  Habits 


References 


Lesser  Scaup  Common- 

(Aythya  affinls)       abundant 


Mollusks,  Ruppia 


Narcisse,  pers.  coram. 

Ogden  1969 

Kushlan  et  al.,  In  prep. 


Bufflehead 

(Bucephala  albeola) 


Rare 


Ruddy  Duck  Common  W 

(Oxyura  jamaicensis) 


Gastropods,  crabs,   Ogden  1969 

crustaceans         Kushlan  et  al.,  in  prep. 


Potamogeton,  Najas,   Ogden  1969 
Zostera,   Ruppia,    Kushlan  et  al. 
mollusks 


in  prep. 


Hooded  Merganser 
(Lophodytes 
cucullatus) 

Red-breasted  Merganser 
(Mergus  serrator ) 

Purple  Gallinule 
(Porphyrula 
martlnica) 

Common  Gallinule 

(Gallinula  chloropus) 

American  Coot 

(Fulica  americana) 


Rare-uncommon 


Common 


Rare 


Common 


Abundant 


W.T 


Yr 


Yr 


W,T 


Fish 


Fish 


Ogden  1969 


Narcisse,  pers.  coram. 


Aquatic  insects,     Narcisse,  pers.  coram, 
mollusks,  Ffrench  1966 

Eleocharis,  Paspalum 


Seeds,  aquatic 
insects 

Ruppia,  Na j as , 
Potamogeton, 
aquatic  insects 


Narcisse,  pers.  coram. 
Ffrench  1966 

Narcisse,  pers.  coram. 


135 


AERIALLY   SEARCHING 


Common  Name 
(Latin  name) 


Abundance 


Season  of 
Occurrence 


Nesting"    Food  Habits 


References 


Herring  Gull 

(Larus  argentatus) 

Ring-billed  Gull 

(Larus  delawarensis) 


Common 


Fish,  mollusks, 
crustaceans 

Fish,  insects, 
mollusks 


Narcisse,  pers.  comm. 
Ogden  1969 

Narcisse,  pers.  comm. 
Ogden  1969 


Laughing  Gull  Common 

(Larus  atricilla) 

Bonaparte's  Gull       Uncommon 
(Larus  Philadelphia) 

Gull-billed  Tern        Uncommon 
(Gelochelidon 
nilotica) 

Forster's  Tern         Uncommon- 
( Sterna  fosteri)      common 


Fish,  shrimp,  crabs 


Fish,  insects 


Narcisse,  pers.  comm. 
Ogden   1969 

Ogden   1969 


Mayflies,  dragonflies   Ogden   1969 


Fish 


Narcisse,  pers.  comm. 
Ogden   1969 


Common  Tern 

(Sterna  hirundo) 


Fish 


Ogden   1969 


Least  Tern 

(Sterna  albifrons) 


Fish 


Narcisse,  pers.  comm. 
Ogden  1969 


Royal  Tern 

(Thalasseus  maxima) 


Common 


W,T 


Fish 


Ogden   1969 


Sandwich  Tern 
(Sterna  sand- 
vicensis) 


Uncommon 


Yr 


Fish 


Narcisse,  pers.  comm. 
Ogden   1969 


Caspian  Tern 

(Sterna  caspia) 


Uncommon 


Fish 


Ogden   1969 


Black  Skimmer  Common 

(Rynchops  nigra) 

Belted  Kingfisher       Common 
(Megaceryle  alcyon) 

Fish  Crow  Common 

(Corvus  ossifragus) 


Yr 


Fish 


Fish 


Fish 


Ogden  1969 


Narcisse,  pers.  comm. 


Narcisse,  pers.  comm. 


136 


BIRDS  OF  PREY 


Common  Name 
(Latin  name) 


Season  of 
Abundance     Occurrence*    Nesting3    Food  Habits 


References 


Magnificent  Frigate-    Common  S 
bird  Uncommon  W 

(Fregata  magnificens) 


Turkey  Vulture 
(Cathartes  aura) 


Common 


Fish 


Narcisse, 


pera.  com. 


Smith,  pers.  obs. 


Narcisse,  pers.  comm. 
Orians  1969 


Black  Vulture  Common 

(Coragyps  atratus) 

Swallow-tailed  Kite     Common 
(Elanoides  forf ica- 
tus) 

Sharp-shinned  Hawk      Uncommon 
(Accipiter  striatus) 

Cooper's  Hawk  Uncommon 

(Accipiter  cooper ii) 

Red-tailed  Hawk        Uncommon 
(Buteo  jamaicensis) 

Red-shouldered  Hawk     Common 
(Buteo  lineatus) 


Broad-winged  Hawk 
(Buteo  platypterus) 

Swainson's  Hawk 
(Buteo  swainsoni) 

Short-tailed  Hawk 
(Buteo  brachyurus) 

Bald  Eagle 
(Haliaeetus 
leucocephalus) 

Marsh  Hawk 

(Circus  cyaneus) 

Osprey 

(Pandion  haliaetus) 

Peregrine  Falcon 
(Falco  peregrinus) 


Merlin 

(Falco  columbarius) 

American  Kestrel       Common 
(Falco  sparverius) 

Barn  Owl  Uncommo 

(Tyto  alba) 

Great  Horned  Owl       Uncommo 
(Bubo  virginianus) 

Barred  Owl  Uncommo 

(Strix  varla) 


Rare- locally 
common  (Fla. 
Bay) 

Uncommon 


Very  rare- 
locally  common 
(Fla.  Bay) 

Uncommon 


Carrion 


Snakes,  lizards, 
frogs 


Robertson  &  Kushlan 

1974 
Orians  1969 

Howell  1932 
Snyder   1974 


Smaller  passerines  Howell  1932 
Larger  passerines  Howell  1932 
Small  mammals,  birds    Howell  1932 


Snakes,  frogs, 
lizards.  Insects 


Insects,  small 
mammals 


Howel]   1932 
Robertson  &  Kushlan. 
1974 

Howell   1932 


Small  mammals,  grass-   Howell  1932 
hoppers 


Small  birds 


Fishes 


Howell  1932 


Howell  1932 


Small  mammals,  shore-   Howell  1932 
birds 


Fishes 


Howell  1932 


Waterfowl,  shorebirds   Nisbet  1968 
Ogden  1969 
Howell  1932 

Small  birds,  shore-     Howell  1932 
birds 


Insects 


Small  mammals 


Waterfowl ,  small 
mammals 


Hovel)   1932 


Howell  1932 


Howell  1932 


Y      Small  mammals,  frogs,   Howell  1932 
snakes 


137 


ARBOREAL  BIRDS 


Common  Name 
(Latin  name) 


Mourning  Dove 

(Zenaidura  macroura) 

White-crowned  Pigeon 
(Columba 
leucocephala) 

Mangrove  Cuckoo 
(Coccyzus  minor) 


Abundance 


Season  of 
Occurrence3 


Nesting3    Food  Habits 


Uncommon 


Yr 


Seeds 


Berries,  seeds 
fruits 


Caterpillars, 
man t ids 


References 


Emlen  1977 


Howell  1932 

Robertson  &  Kushlan  1974 


Howell  1932 
Ffrench  1966 

Robertson  &  Kushlan  1974 
Martin  et  al.  1951 


Yellow-billed  Cuckoo     Common 
(Coccyzus  americanus) 


Caterpillars, 
beetles 


Howell  1932 
Ffrench  1966 
Martin  et  al.  1951 


Smooth-billed  Anl       Rare  Yr 

(Crotophaga  anl) 

Chuck-will 's-widow      Uncommon        Yr 
(Caprimulgus 
carolinensis) 

Common  Flicker  Uncommon        Yr 

(Colaptes  auratus) 

Pileated  Woodpecker      Uncommon        Yr 
(Dryocopus  pileatus) 


Mosqultos,  moths 


Ants,  beetles, 
fruits  in  winter 

Beetles,  berries, 
fruits 


Howell  1932 
Ffrench  1966 

Martin  et  al.  1951 
Narcisse,  pers.  comm. 


Narcisse,  pers.  coram. 
Martin  et  al.  1951 

Howell  1932 
Robertson  1955 
Robertson  4  Kushlan  1974 


Red-bellied  Woodpecker   Common 
(Melanerpes  carolinus) 


Beetles,  ants, 

grasshoppers, 

crickets 


Narcisse,  pers.  coram. 
Martin  et  al.  1951 


Red-headed  Woodpecker 
(Melanerpes 
erythrocephalus) 

Yellow-bellied 
Sapsucker 

(Sphyraplcus  varius) 

Hairy  Woodpecker 
(Picoides 
vlllosus) 

Eastern  Kingbird 

(Tyrannus  tyrannus) 

Gray  Kingbird 
(Tyrannus 
dominicensis) 

Western  Kingbird 

(Tyrannus  vertlcalus) 

Great  Crested 
Flycatcher 

(Mylarchus  crlnitus) 

Acadian  Flycatcher 

(Empldonax  virescens) 

Eastern  Phoebe 

(Sayornis  phoebe) 

Eastern  Wood  Pewee 
(Contopus  vlrens) 


Uncommon 


Common 


Uncommon 
(common  S) 


Rare 


Common 


Rare-uncommon 


Yr 


W,T 


S,T 


W,T 


Beetles,  ants, 

grasshoppers, 

caterpillars 

Beetles,  ants-, 
caterpillars 


Insects,  beetle 
larvae 


Ants,  wasps, 
grasshoppers 

Bees,  wasps, 
beetles,  dragon 


Bees,  wasps, 
grasshoppers 

Insects,  berries 


Narcisse,  pers.  comm. 
Martin  et  al.  1951 


Narcisse,  pers.  coram. 
Martin  et  al.  1951 


Emlen  1977 


Narcisse,  pers.  comm. 
Martin  et  al.  1951 

Howell  1932 

Robertson  &  Kushlan  1974 


Narcisse,  pers.  comm. 
Martin  et  al.  1951 


Howell  1932 
Robertson  1955 


Small  flying  insects  Morton  1980 


Bees,  wasps,  ants    Narcisse,  pers.  comm. 
Martin  et  al.  1951 

Bees,  wasps,  ants,   Narcisse,  pers.  comm. 
moths  Howell  1932 


138 


ARBOREAL  BIRDS  (continued) 


Common   *<ame 
(Latin  name) 


Season  of 
Occurrence* 


Nesting 


Food    Habits 


References 


Barn  Swallow 

(Hirundo  rustica) 

Blue  Jay 

(Cyanocitta  crlstata) 

Tufted  titmouse 
(Parus  bicolor) 

Carolina  Wren 
(Thryothorus 
ludovicianus) 

Mockingbird 

(Mimus  polyglottos) 

Catbird 

(Dumetella  caro- 
linensis) 


Locally  common 


Very  rare- 
rare 


W,T 


Grasshoppers,  cater- 
pillars, beetles 

Caterpillars ,   wasps , 
bees 

Ants,  flies,  milli- 
peds 


Fruits ,  berries 


Fruits,  insects 


Howell  1932 
Bacon  1970 

Narcisse,  pers.  comm. 
Martin  et  al.   1951 

Howell  1932 

Robertson  &  Kushlan  1974 

Narcisse ,  pers .  comm . 
Martin  et  al.   1951 


Robertson  1955 


Narcisse ,  pers .  comm . 
Martin  et  al.   1951 


Brown  Thrasher 

(Toxo stoma  rufum) 


Narcisse,  pers.  comm. 
Martin  et  al.   1951 


American  Robin         Abundan 
(Turdus  migratorius) 

Blue-gray  Gnatcatcher   Uncommo 
(Polioptila  caerulea) 

Ruby-crowned  Kinglet    Uncommo 
(Regulus  calendula) 

White-eyed  Vireo        Uncommo 
(Vireo  griseus) 

Black-whiskered  vireo   Uncommo 
(vireo  altiloquus) 

Red-eyed  Vireo         Uncommo 
(Vireo  olivaceus) 

Yellow-throated  Vireo   Uncommo 
(Vireo  f lavifrons) 

Black-and-white         Fairly 
Warbler  common 

(Mniotilta  varia) 

Worm-eating  Warbler     Uncommo 
(Helmitheros  vermi- 
vorus) 


W,T 


Worms ,  berries, 
insects 

Insects ,  especially 
Hymenopterans 

Wasps,  ants 
Butterflies,  moths 

Spiders ,  caterpillars 
Caterpillars ,  beetles 
Butterflies ,  moths , 
Wood  boring  insects 

Caterpillars,  spiders 


Narcisse,  pers.  comm. 
Martin  et  al. .  1951 


Narcisse ,  pers .  comm. 
Howell  1932 


Narcisse,  pers.  comm. 
Howell   1932 


Robertson   1955 


Howell   1932 

Robertson  &  Kushlan,  1974 


Narcisse,  pers .  comm. 
Howell.  1932 


Morton   1980 


Lack  and  Lack   1972 
Keast   1980 
Ogden   1969 

Ogden   1969 

Kushlan ,  pers .  comm . 


Prothonotary  Warbler 
(Protonotaria  citrea) 


Ffrench.  1966 
Russel   1980 


Yel low-throated 
Warbler 

(Pendroica  dominica) 


Beetles,  moths, 
spiders 


Morton   1980 


Yellow  Warbler 

(Pendroica  petechia) 


Y e 1 1 ow- rumpe d 

Warbler  Abundant 

(Pendroica  corona ta) 

Prairie  Warbler         Uncommon 
(Pendroica  discolor) 

Palm  Warbler  Abundant 

(Pendroica  palmar urn) 


yr 


W,T 


Pipterans ,  bayberries 
Moths,  beetles,  flies 
Insects 


Haverschmidt  1965 

Ffrench  1966 

Orians  i969 

Terborgh  &  Faaborg  1980 

Narcisse,  pers .  comm. 


Lack  &  Lack  1972 
Robertson  &  Kushlan   1974 


Lack  &  Lack   1972 
Emlen,  1977 


139 


ARBOREAL  BIRDS  (continued) 


Common  Name 
(Latin  name) 


Abundance 


Season  of 
Occurrence  a 


Nesting*    Food  Habits 


References 


Blackpoll  Warbler       Uncommon 
(Dendroica  striata) 


Ffrench   1966 


Bay-breasted  Warbler    Rare 
(Dendroica  castanea) 


Morton   1980 


Black-throated  Green    Uncommon 
Warbler 

(Dendroica  virens) 


Aphids ,  leaf-rollers 
and  other  insects 


Ogden   1969 

Kushlan ,  pers .  comm . 


Chestnut-sided  Warbler  Rare 
(Dendroica  pensyl- 
yanica) 

Cape  May  Warbler        Uncommon 
(Dendroica  tigrina)    Common 

Black-throated  Gray     Rare 
Warbler 

(Dendroica  nigrescens) 

Black-throated  Blue     Uncommon 
Warbler  Common 

(Dendroica  caeru- 
lescens) 


Beetles,  flies,  ants 


Morton   1980 


Ogden   1969 


Ogden   1969 
Kushlan,  pers.  comm. 
Hutto  1980 

Kushlan,  pers.  comm. 
Ogden   1969 


Northern  Waterthrush    Abundant 
(Seiurus  novebora-    Rare 
censis) 


Schwartz   1964 
Ffrench   1966 
Bacon  1970 
Russell  1980 


Yellowthroat  Common 

(Geothlypus  trichas) 


Grasshoppers ,  crickets ,  Narcisse,  pers .  comm. 
ants,  wasps  Howell   1932 

Lack  &  Lack  1972 


American  Redstart      Common 
(Setophaga  ruticilla) 


Caterpillars 


Bennett  1980 
Ffrench  1966 
Bacon   1970 


Tennessee  Warbler       Uncommon 
(Vermivora  peregrina) 

Nasheville  Warbler      Rare 
(Vermivora  ruf i- 
capllla) 

Orange-crowned  Warbler  Common 
(Vermivora  celata) 

Golden-winged  Warbler   Rare 
(Vermivora  chrysop- 
tera) 


Northern  Parula 

(Parula  americana) 

Ovenbird 

(Seiurus  aurocapll- 
lus) 

Kentucky  Warbler 

(Oporornis  formosus) 

Mourning  Warbler 

(Oporornis  Philadel- 
phia) 

Yellow-breasted  Chat 
(Icteria  virens) 


Rar e - un co mmo n 


Hymenoptera 


Morton  1980 


Hutto  1980 


Hutto   1980 


Morton  1980 


Lack  and  Lack   1972 


Beetles,  crickets,      Lack  and  Lack  1972 
grasshoppers 


Beetles,  caterpillars,   Morton  1980 
ants 


Hymenoptera 


Morton   1980 


Hutto.  1980 


140 


ARBOREAL  BIRDS  (concluded) 


Common  Nnmc 
(Latin  name) 


Abundance 


Season  of 
Occurrence3 


Wilson's  Warbler 
(Wilsonia  pusilla) 


Rar  e - uncommon 


Red-winged  Blackbird    Common 
(Agelaius  phoeniceus) 

Boat-tailed  Grackle     Uncommo 
(Quiscalus  major) 

Common  Grackle  Uncommo 

(Quiscalus  guiscula) 

Card  inal  Common 

(Cardinalis 
cardinalis) 

Orchard  Oriole  Rare 

(Icterus  spurius) 

Indigo  Bunting  Uncommo 

(Passerina  cyanea) 


Summer  Tanager 
(Piranga  rubra) 

Dickcissel 

(Spiza  americana) 

Rufous-sided  Townee 
(Pipllo  erythroph- 
thalmus ) 

Swamp  Sparrow 

(Melospiza  georgiana) 


Uncommon 


Uncommon 


Yr 


W,T 


Yr 


W,T 


Nesting3    Food  Habits 


Seeds,  insects 


Crayfish,  crabs, 
shrimp 

Insects,  cater- 
pillars 

Insects,  seeds 


References 


Hutto   1980 

Ramos  and  Warner   1980 

Howel]   1932 
Robertson   1955 

Robertson   1955 
Girard  &  Taylor   1979 

Howell   1932 
Robertson   1955 

Robertson   1955 


Grasshoppers,  beetles   Morton   1980 


Grasshoppers ,  cater- 
pillars 

Hymenoptera 


Narcisse,  pers.  comm. 
Howell   1932 

Morton  1980 


Caterpillars,  beetles   Bacon  1970 

Martin  et  al.   1951 


Caterpillars,  bay- 
berries,  fruits 


Ants,  flies,  seeds 


Narcisse,  pers.  comm. 
Howell   1932 


Narcisse,  pers.  comm. 
Howell   1932 


Tr  =  year  round  resident 
S  =  summer  resident 
W  =  winter  resident 

T  =  transient,  present  only  during  spring  and  fall  migration 
Y  =  species  breeds  in  mangroves 

bL.  Narcisse,  R.N.  "Ding"  Darling  Fed.  Wildlife  Refuge,  Sanibel  Island,  Fla.  (1981). 
J. A.  Kushlan,  So.  Fla.  Res.  Ctr. ,  Everglades  Natl.  Park,  Homestead,  Fla. 


141 


APPENDIX  E.   Mammals  of  south  Florida  mangrove  swamps. 


142 


MAMMALS  OF  FLORIDA  MANGROVES 


Species 


Status 


Food  Habits 


Virginia  Opossum 

(Didelphis  virginiana) 


Abundant 


Fruits,  berries,  insects, 
frogs,  snakes,  small 
birds  and  mammals 


Short-tailed  Shrew 
(Blarina  brevicauda) 


Uncommon 


Insects 


Marsh  Rabbit 

(Sylvilagus  palustris) 

Gray  Squirrel 

(Sciurus  carolinensis) 


Abundant 


Occasional 


Emergent  aquatics 


Fruits,  berries,  mast, 
seeds 


Fox  Squirrel 

(Sciurus  niger) 

Marsh  Rice  Rat 

(Oryzomys  palustris) 

Cudjoe  Key  Rice  Rat 
(Oryzomys  argentatus) 


Rare 


Uncommon 


Rare 


Cotton  Rat  Abundant 

(Sigmodon  hispidus) 

Gray  Fox  Uncommon 

(Urocyon  ciner eoar genteus ) 

Black  Bear  Rare 

(Ursus  americanus) 


Fruits,  berries,  mast 


Seeds  of  emergent  plants, 
insects,  crabs 

Seeds,  insects,  crabs 


Sedges,  grasses,  cray- 
fish, crabs,  insects 

Small  mammals,  birds 


Fruits,  berries,  fish, 
mice 


Raccoon 

(Procyon  lotor) 

Mink 

(Mustela  vison) 


Abundant 


Rare 


Crayfish,  frogs,  fish 


Small  mammals,  fish, 
frogs ,  snakes ,  aquatic 
insects 


Striped  Skunk 

(Mephitis  mephitis) 


River  Otter 

(Lutra   canadensis) 


Common 


Uncommon 


Bird  eggs  and  young 
frogs,  mice,  larger 
invertebrates 

Crayfish,  fish,  mussels 


143 


MAMMALS  OF  FLORIDA  MANGROVES  (concluded) 


Species 


Status 


Food  Habits 


Panther 

(Felis  concolor) 


Very  rare 


Deer,  rabbits,  mice, 
birds 


Bobcat 

(Felis  rufus) 


Common 


Rabbits,  squirrels, 
birds 


White-tailed  Deer 

(Odocoileus  virglnianus) 


Common 


Emergent  aquatics,  nuts, 
acorns,  occasionally 
mangrove  leaves 


Key  Deer 

(O.v.  clavium) 


Common  on  cer-     Emergent  aquatics  and 
tain  Florida  Keys  other  vegetation 
(no  longer  on 
mainland) 


Black  Rat 

(Rattus  rattus) 


Common 


Bottle-nosed  Dolphin 
(Tursiops  truncatus) 

West  Indian  Manatee 
(Trichechus  manatus) 


Uncommon 


Uncommon 


Fish 


Submerged  aquatics, 
Zostera,  Ruppia ,  Halodule, 
Syringodium,  Cymodocea  , 
Thalassia 


References:   Layne   1974;  Hamilton  and  Whittaker   1979; 

L.  Narcisse,  R.N.  "Ding"  Darling  Fed.  Wildlife 
Refuge,  Sanibel  Island,  Fla.;  personal  commu- 
nication. 


144 


50272  -101 

REPORT   DOCUMENTATION 
PAGE 


l._REPORT   NO. 

FWS/OBS-81/24 


3.    Recipient's  Accession   No. 


4.  Title  and  Subtitle 

THE  ECOLOGY  OF  THE  MANGROVES  OF  SOUTH  FLORIDA: 
PROFILE 


A  COMMUNITY 


5.    Report   Date 

January  1982 


7.  Author(s) 

William  E.  Odum,  Carole  C.  Mclvor,  Thomas  J.  Smith,  III 


8.    Performing  Organization  Rept.  No. 


s.  Address  of  Authors 

Department  of  Environmental  Sciences 
University  of  Virginia 
Charlottesville,  Virginia  22901 


10.  Proiect/Task/Work  Unit  No. 

11.  Contract(C)  or  Grant(G)  No. 
(C) 

(G) 


12.  Sponsoring  Organization  Name  and  Address 

Office  of  Biological  Services 
Fish  and  Wildlife  Service 
U.S.  Department  of  the  Interior 
Washington,  D.C.  20240 


New  Orleans  OCS  Office 
Bureau  of  Land  Management 
U.S.  Department  of  the  Interior 
New  Orleans,  Louisiana  70130 


13.  Type  of  Report  &  Period  Covered 


15.  Supplementary  Notes 


16.   Abstract  (Limit:  200  words) 

A  detailed  description  is  given  of  the  community  structure  and  ecosystem  processes 
of  the  mangrove  forests  of  south  Florida.  This  description  is  based  upon  a  compilation  of 
data  and  hypotheses  from  published  and  unpublished  sources. 

Information  covered  ranges  from  details  of  mangrove  distribution,  primary  production, 
and  diseases  to  aspects  of  reproduction,  biomass  partitioning,  and  adaptations  to  stress. 
Mangrove  ecosystems  are  considered  in  terms  of  zonation,  succession,  litter  fall  and 
decomposition,  carbon  export,  and  energy  flow. 

Most  of  the  components  of  mangrove  communities  are  cataloged  and  discussed;  these 
include  microorganisms,  plants  other  than  mangroves,  invertebrates,  fishes,  reptiles, 
amphibians,  birds,  and  mammals. 

Finally,  two  sections  summarize  the  value  of  mangrove  ecosystems  to  man  and  present 
ways  to  manage  this  type  of  habitat.  It  is  concluded  that  mangrove  forests,  which  cover 
between  430,000  and  500,000  acres  (174,000  -  202,000  ha)  in  Florida,  are  a  resource  of 
great  value  and  should  be  protected  and  preserved  wherever  possible. 


17.  Document  Analysis     a.   Descriptors 

Ecology,  value,  management,  fauna 

b.    Identifiers/Open  Ended   Terms 

Mangroves,  halophytes,  coastal  wetlands,  ecosystem,  South  Florida 


c.  COSATI  Field/Group 


18.  Availability  Statement 

Unlimited 


19.  Security  Class  (This  Report) 

Unr.lassifipH 


20.  Security  Class  (This  Page) 


21.  No.  of  Pages 

154 


22.    Price 


(See  ANSI-Z39.18) 


See  Instructions  on   Reverse 


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(Formerly  NTIS-35) 
Department  of  Commerce 


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©■© 


Headquarters  -  Office  of  Biological 
Services,  Washington,  D.C. 

National  Coastal  Ecosystems  Team, 

Slidell.  La. 

Regional  Offices 


U.S.  FISH  AND  WILDLIFE  SERVICE 
REGIONAL  OFFICES 


REGION   1 

Regional  Director 

U.S.  Fish  and  Wildlife  Service 

Lloyd  Five  Hundred  Building.  Suite  1692 

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REGION  2 

Regional  Director 

U.S.  Fish  and  Wildlife  Service 

P.O.Box  1306 

Albuquerque.  New  Mexico  87 1 03 

REGION  3 

Regional  Director 
U.S.  Fish  and  Wildlife  Service 
Federal  Building,  Foit  Snelling 
Twin  Cities.  Minnesota  55111 


REGION  4 

Regional  Director 
U.S.  Fish  and  Wildlife  Service 
Richard  B.  Russell  Building 
75  Spring  Street,  S.W. 
Atlanta,  Georgia  30303 

REGION  5 

Regional  Director 

U.S.  Fish  and  Wildlife  Service 

One  Gateway  Center 

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REGION  6 

Regional  Director 

U.S.  Fish  and  Wildlife  Service 

P.O.  Box  25486 

Denver  Federal  Center 

Denver,  Colorado  80225 


REGION   7 

Regional  Directoi 
U.S.  Fish  and  Wildlife  Service 
101  1  E.Tudor  Road 
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DEPARTMENT  OF  THE  INTERIOR 

U.S.  FISH  AND  WILDLIFE  SERVIDE 


As  the  Nation's  principal  conservation  agency,  the  Department  of  the  Interior  has  respon- 
sibility for  most  of  our  nationally  owned  public  lands  and  natural  resources.  This  includes 
fostering  the  wisest  use  of  our  land  and  water  resources,  protecting  our  fish  and  wildlife, 
preserving  the-environmental  and  cultural  values  of  our  national  parks  and  historical  places, 
and  providing  for  the  enjoyment  of  life  through  outdoor  recreation.  The  Department  as- 
sesses our  energy  and  mineral  resources  and  works  to  assure  that  their  development  is  in 
the  best  interests  of  all  our  people.  The  Department  also  has  a  major  responsibility  for 
American  Indian  reservation  communities  and  for  people  who  live  in  island  territories  under 
U.S.  administration.