Juvenile and small resident

fishes of Florida Bay, a critical 

habitat in the Everglades 

National Park, Florida

Allyn B. Powell

Gordon Thayer

Michael Lacroix

Robin Cheshire

NOAA Professional Paper NMFS 6

U.S. Department 

of Commerce

March 2007
U.S. Department

of Commerce

Carlos M. Gutierrez

Secretary

National Oceanic

and Atmospheric

Administration

Vice Admiral

Conrad C. Lautenbacher Jr.,

USN (ret.)

Under Secretary for 

Oceans and Atmosphere

National Marine 

Fisheries Service

William T. Hogarth

Assistant Administrator 

for Fisheries

The NOAA Professional Paper NMFS 

(ISSN 1931-4590) series is published 

by the Scientific Publications Office, 

National Marine Fisheries Service, 

NOAA, 7600 Sand Point Way NE, 

Seattle, WA 98115.

The Secretary of Commerce has

determined that the publication of 

this series is necessary in the transaction of the public business required by 

law of this Department. Use of funds 

for printing of this series has been approved by the Director of the Office 

of Management and Budget.

NOAA Professional 

Papers NMFS

Scientific Editor

Dr. Adam Moles

Technical Editor

Elizabeth Calvert

National Marine Fisheries Service, NOAA

17109 Point Lena Loop Road

Juneau, AK 99801-8626

Managing Editor

Shelley Arenas

National Marine Fisheries Service

Scientific Publications Office

7600 Sand Point Way NE

Seattle, WA 98115

Editorial Committee

Dr. Ann C. Matarese National Marine Fisheries Service

Dr. James W. Orr National Marine Fisheries Service

Dr. Bruce L. Wing National Marine Fisheries Service

The NOAA Professional Paper NMFS series carries peer-reviewed, lengthy original 

research reports, taxonomic keys, species synopses, flora and fauna studies, and data-intensive reports on investigations in fishery science, engineering, and economics. Copies 

of the NOAA Professional Paper NMFS series are available free in limited numbers to 

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other scientific and technical publications in the marine sciences. Professional Papers 

are published online in PDF format at http://spo.nmfs.noaa.gov

NOTICE: This series was established in 2003 to replace the NOAA Technical Report 

NMFS series. 
NOAA Professional Paper NMFS 6

Juvenile and small resident fishes 

of Florida Bay, a critical habitat 

in the Everglades National Park, Florida

Allyn B. Powell

Gordon Thayer

Michael Lacroix

Robin Cheshire

March 2007

U.S. Department of Commerce

Seattle, Washington
Suggested reference

Powell, Allyn B., Gordon Thayer, Michael Lacroix, and Robin Cheshire. 2007. 

Juvenile and small resident fishes of Florida Bay, a critical habitat in the Everglades 

National Park, Florida. NOAA Professional Paper NMFS 6, 210 p. 

Online dissemination

This report is posted online in PDF format at http://spo.nmfs.noaa.gov 

(click on Professional Papers link). 

Copyright law

Although the contents of the Professional Papers have not been copyrighted and 

may be reprinted entirely, reference to source is appreciated. 

Proprietary products

The National Marine Fisheries Service (NMFS) does not approve, recommend, 

or endorse any proprietary product or proprietary material mentioned in this 

publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales promotion which would indicate or 

imply that NMFS approves, recommends, or endorses any proprietary product 

or proprietary material mentioned herein, or which has as its purpose an intent 

to cause directly or indirectly the advertised product to be used or purchased 

because of this NMFS publication. 

ii
CONTENTS

Chapter 1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Chapter 2. Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 Field and laboratory methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Sampling strategy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Collection methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Processing of specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Data analysis and presentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Temperature and salinity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Seagrasses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Fishes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Chapter 3. Environmental variables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Temperature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Salinity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Seagrasses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Thalassia testudinum (turtle grass) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Halodule wrightii (shoalgrass) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Syringodium filiforme (manatee grass) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Chapter 4. Family Clupeidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

Brevoortia smithi (yellowfin menhaden) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

Harengula jaguana (scaled sardine) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Opisthonema oglinum (Atlantic thread herring) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

Sardinella aurita (Spanish sardine) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

Chapter 5. Family Engraulidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

Anchoa hepsetus (striped anchovy) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

A. mitchilli (bay anchovy) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

Chapter 6. Family Ariidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46

Arius felis (hardhead catfish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46

Chapter 7. Family Synodontidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

Synodus foetens (inshore lizardfish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

Chapter 8. Family Batrachoididae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

Opsanus beta (gulf toadfish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

Chapter 9. Family Hemiramphidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

Chriodorus atherinoides (hardhead halfbeak) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

Hemiramphus brasiliensis (ballyhoo) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

Hyporhamphus unifasciatus (halfbeak) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64

Chapter 10. Family Belonidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

Strongylura notata (redfin needlefish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

S. timucu (timucu) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74

iii
Chapter 11. Families Fundulidae and Cyprinodontidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76

Floridichthys carpio (goldspotted killifish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76

Lucania parva (rainwater killifish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

Chapter 12. Families Atherinidae and Atherinopsidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

Atherinomorus stipes (hardhead silverside) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

Hypoatherina harringtonensis (reef silverside) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87

Membras martinica (rough silverside) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90

Chapter 13. Family Syngnathidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93

Anarchopterus criniger (fringed pipefish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93

Syngnathus floridae (dusky pipefish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

S. scovelli (gulf pipefish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101

Hippocampus erectus (lined seahorse) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105

H. zosterae (dwarf seahorse) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109

Chapter 14. Family Lutjanidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113

Lutjanus griseus (gray snapper) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113

L. synagris (lane snapper) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117

Chapter 15. Gerreidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121

Eucinostomus gula (silver jenny) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121

Chapter 16. Family Haemulidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125

Haemulon plumieri (white grunt) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125

H. sciurus (bluestriped grunt) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129

Orthopristis chrysoptera (pigfish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133

Chapter 17. Family Sparidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

Archosargus probatocephalus (sheepshead) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

Calamus arctifrons (grass porgy) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141

Lagodon rhomboides (pinfish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145

Chapter 18. Family Sciaenidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

Bairdiella chrysoura (silver perch) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

Cynoscion nebulosus (spotted seatrout) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153

Chapter 19. Family Sphyraenidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157

Sphyraena barracuda (great barracuda) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157

Chapter 20. Family Scaridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161

Nicholsina usta (emerald parrotfish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161

Chapter 21. Family Callionymidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165

Diplogrammus pauciradiatus (spotted dragonet) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165

Chapter 22. Family Gobiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169

Gobiosoma robustum (code goby) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169

Microgobius gulosus (clown goby) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173

M. microlepis (banner goby) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177

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Chapter 23. Family Monacanthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

Monacanthus ciliatus (fringed filefish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

M. hispidus (planehead filefish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185

Chapter 24. Family Ostraciidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189

Lactophrys quadricornis (scrawled cowfish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189

Chapter 25. Family Tetraodontidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193

Sphoeroides nephelus (southern puffer) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193

Chapter 26. Family Diodontidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197

Chilomycterus schoepfi (striped burrfish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197

Chapter 27. Overview and future research needs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201

Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201

Future research needs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203

Literature cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203

Appendix A. Abundance of fishes by salinity regime . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209

Appendix B. Uncommon species—number and area collected . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210

v
Abstract—This compendium presents information on the life history, diet, 

and abundance and distribution of 46 of 

the more abundant juvenile and small 

resident fish species, and data on three 

species of seagrasses in Florida Bay, Everglades National Park. Abundance and 

distribution of fish data were derived from 

three sampling schemes: (1) an otter trawl 

in basins (1984–1985, 1994–2001), (2) a 

surface trawl in basins (1984–1985), and 

(3) a surface trawl in channels (1984–

1985). Results from surface trawling only 

included pelagic species. Collections 

made with an otter trawl in basins on a 

bi-monthly basis were emphasized. Nonparametric statistics were used to test 

spatial and temporal differences in the 

abundance of species and seagrasses. Fish 

species accounts were presented in four 

sections – Life history, Diet, Abundance 

and distribution, and Length-frequency 

distributions. Although Florida Bay is a 

subtropical estuary, the majority of fish 

species (76%) had warm-temperate affinities; i.e., only 24% were solely tropical 

species. The five most abundant species 

collected, in descending order, by (1) otter trawl in basins were: Eucinostomus gula, 

Lucania parva, Anchoa mitchilli, Lagodon 

rhomboides, and Syngnathus scovelli; (2) 

surface trawl in basins were: Hyporhamphus 

unifasciatus, Strongylura notata, Chriodorus 

atherinoides, Anchoa hepsetus, and Atherinomorus stipes; (3) surface trawl in channels were: Hypoatherina harringtonensis, A. 

stipes, A. mitchelli, H. unifasciatus, and C. 

atherinoides.

Chapter 1. Introduction

The importance of estuaries as nursery areas for fishes has been well documented, and there is ample evidence 

that many commercial and recreational fisheries target those species 

that utilize estuaries. Estuaries provide abundant food resources and 

relatively fewer predators than coastal 

waters. They provide a variety of habitats that include the water column, 

grass beds, marshes, mangroves, and 

hard/soft bottoms. Potential impacts 

on these habitats include fishing, 

development, chemical spills, and 

freshwater input and diversion. Of recent concern has been the impact of 

man-induced activities in the Kissimmee-Okeechobee-Everglades drainage basin on estuarine systems within 

Everglades National Park. The hydrology of this drainage basin has been 

altered by water management policies to reclaim wetlands and furnish 

irrigation for agriculture, and provide flood control for the sprawling 

population of the Miami area. These 

alterations of the drainage, along with 

natural causes, have had an impact on 

the downstream estuary—Florida Bay 

(Thayer and Chester, 1989; Hoss and 

Thayer, 1993; Boesch, 1996; Able and 

Fahay, 1998; Schofield, 2003).

Florida Bay is a shallow lagoon 

roughly encompassed on the north 

by the mainland of the Florida peninsula, on the east and south by the 

Florida Keys, and to the west by the 

Gulf of Mexico (Fig.1.1). The precise 

boundaries of Florida Bay vary because of the open connection of the 

Bay with the Gulf of Mexico. Here, 

we define Florida Bay as that portion 

which is situated within the boundaries of Everglades National Park, covering an area of approximately 1700 

km2 (FMRI1). An excellent summary 

of the description of the Florida Bay 

ecosystem is given by Fourqurean and 

Robblee (1999), which is a source of 

information for this introduction.

A latticework of carbonate mud 

banks and shallow basins (1–3 m 

deep) connected by channels is characteristic of the Bay. Approximately 

200 small mangrove-lined islands, or 

keys, occur in the Bay. The carbonate 

mud banks restrict tidal flow from the 

Gulf of Mexico on the west while the 

Florida Keys isolate all but the south1 FMRI. 1995. Fisheries-independent monitoring program, 1994 annual data summary 

report. Florida Marine Research Institute, 

Florida Department of Environmental Protection, St. Petersburg, FL, 451 p.

Juvenile and small resident fishes 

of Florida Bay, a critical habitat in 

the Everglades National Park, Florida

Allyn B. Powell

Gordon Thayer1

Michael Lacroix

Robin Cheshire

Center for Coastal Fisheries and Habitat Research

National Oceanic and Atmospheric Administration

101 Pivers Island Road

Beaufort, North Carolina 28516

e-mail address: Allyn.Powell@noaa.gov

1 Current address: I. M. Systems Group Incorporated

3401 Blexhill Place

Kensington, MD 20895
2 

Figure 1.1

Florida Bay and location of geographic names used in text.
3

ern portion of the Bay from tidal cycles of the Atlantic 

Ocean. Tides along the western margin of the bay range 

61 cm; whereas on the east, adjacent to the Florida 

Keys, tides range 17 cm. Within most of the central and 

northeastern portions of the Bay, tides are generally 

wind driven. Southwest winds produce relatively higher 

tides, while northeast winds, which are dominant in 

winter, produce relatively lower tides (Fourqurean and 

Robblee, 1999; FMRI1). Lack of strong tidal circulation 

in the central and northeastern portion of the bay could 

restrict the recruitment of early-stage, planktonic fish 

larvae into those areas.

Weather in Florida Bay is subtropical and is characterized by a warm, wet season from May to October 

and a cooler, dry season from November to April. The 

majority of rainfall (75%) occurs during the wet season, 

and annual evaporation and precipitation are similar. 

Freshwater run-off into Florida Bay is mainly through 

Taylor Slough. Natural freshwater flow into the Bay has 

been disrupted since the late 1800s as a result of managed agriculture, water supply, and flood control on the 

Florida mainland (Fourqurean and Robblee, 1999). 

Florida Bay is generally oligotrophic with low phytoplankton biomass throughout. Western Florida Bay has 

relatively stable marine conditions with near Redfield N:

P ratios for seston, which are good indicators of nitrogen 

or phosphate limitations to phytoplankton. The central 

portion of the bay is occasionally hyperhaline, with high 

N:P ratios for seston, and high concentrations of dissolved inorganic matter. The eastern portion of the Bay 

has highly variable salinities and N:P ratios. Phosphate 

is a limiting factor for phytoplankton productivity in this 

eastern part of the Bay (Boyer et al., 1999; Fourqurean 

and Robblee, 1999).

The organic content of the surface sediment, which 

is linked to overall hydrology, deposition, and the plant 

community, is generally low in eastern Florida Bay and 

highest in western and northwestern Florida Bay. The 

western Bay also has greater sediment depths which 

generally correspond to a high seagrass standing crop 

(Thayer and Chester, 1989).

Basins, banks, and channels in Florida Bay are dominated by the seagrass Thalassia testudinum, followed by 

Halodule wrightii, and Syringodium filiforme. The distribution and abundance of seagrasses within the Bay is not 

uniform. Generally higher seagrass standing crop, higher seagrass short-shoot densities, and higher seagrass 

species diversity are found in western Florida Bay and 

channels throughout the Bay. The highest mean densities of fishes also are found there. Eastern Florida Bay has 

a relatively low standing crop of seagrasses, and the lowest mean density of fishes (Thayer and Chester, 1989). 

In the late 1980s environmental changes occurred 

in Florida Bay. Dense stands of Thalassia experienced 

major die-offs. Turbidity increased and subsequent algal 

blooms further reduced the penetration of light into the 

water column (Fourqurean and Robblee, 1999). Prior 

to the die-off of seagrasses, three species of fish (Lucania 

parva, Lagodon rhomboides, and Eucinostomus gula) were 

dominant in Florida Bay (Thayer and Chester, 1989; 

Thayer et al., 1999). A decade later while the die-off 

was still in progress the bay anchovy (Anchoa mitchilli), 

a zooplanktivore, increased dramatically in abundance 

(Thayer et al., 1999). Although the fish community was 

still dominated by a few species (L. parva, L. rhomboides, 

and E. gula), the major presence of A. mitchilli signaled 

a shift from a fish community associated with seagasses 

(i.e., canopy dwellers), to one dominated by a zooplanktivorus species.

This habitat change occurred rapidly, and may have 

been a consequence of long-term stressors, including 

chronic hyperhaline conditions, overdevelopment of 

seagrass beds, silting of the Bay due to lack of hurricanes, and abnormally warm summer and fall temperatures (Fourqurean and Robblee, 1999). For a detailed 

account of the changes in seagrass distribution and 

abundance prior to and after the die-off of seagrasses, 

and a conceptual model of seagrass die-off in Florida 

Bay, see Hall et al. (1999), Thayer et al. (1999), and 

Zieman et al. (1999).

These environmental changes have resulted in modifications of aspects of the ecology of the Bay beyond 

seagrass and fish. Some resident populations of sponges, 

which provide nursery habitat for spiny lobster, have also experienced die-offs. Landings of pink shrimp, which 

use seagrass beds in Florida Bay as a nursery area, have 

also declined. Thus, modifications in the Bay’s ecology 

can have far reaching impacts on its fishery resources 

since various life history stages utilize Florida Bay as a 

nursery area or as a transient or permanent residence. 

This compendium of life history information for the 

fishes and invertebrates, which we collected throughout the bay between 1984–2001, will provide future 

researchers and managers with information needed to 

evaluate changes in life history structures that may occur 

as a result of natural or anthropogenic environmental 

changes.

Biological information is provided for 46 fish species. 

Included for each species is brief life history information, diet, spatial and temporal abundance and distribution data, and length-frequency distributions to infer 

spawning, recruitment, and growth. Also included are 

temporal and spatial analyses of temperature, salinity, 

and seagrass composition and abundance. 
4 

Field and laboratory methods

Sampling strategy Two methods of stratifying Florida 

Bay were employed. For the first, sampling occurred in 

basins and channels in three strata. During the periods 

May 1984–June 1985 and July 1994–January 1999, sampling during each trip was conducted at 18 basin stations equally divided among three strata – East, Central, 

and West (Fig. 2.1). Also, six random samples per trip 

were taken in channels during 1984–1985. The samples 

in channels were not stratified by strata, and sampling at 

channels was done at fixed sites because of the limited 

trawling area. Sampling locations in the basin strata 

were determined using a grid system. Each grid cell was 

approximately 1800 m on a side. Six cells were randomly 

chosen for each basin strata and samples were taken at 

approximately the center of the cell where possible. 

For the second method, we modified our sampling 

beginning in March 1999 to conform with the South 

Florida Ecosystem Restoration Prediction and Modeling 

(SFERPM) Program Management Committee’s (PMC) 

subdivision designations. Florida Bay was stratified into 

six subdivisions (Fig. 2.1). Sampling locations in the 

basin strata were determined using a grid system. Each 

grid cell was 1800 m on a side. Because the number of 

trawlable squares in each subdivision was not equal, 

the number of stations to be sampled was weighted by 

trawlable area. Thirty-six stations were randomly chosen 

(Gulf: 4; Western: 3; Central: 10; Atlantic: 7; Northern: 

3; Eastern: 9). Samples were taken at approximately 

the center of the cell where possible. For latitude and 

longitude demarcations see Fig. 2.1; for location of 

geographic locations referred to in the text, see Fig. 1.1. 

Because of the change in the stratified design in 1999, 

caution should be used when comparisons are made 

between May 1984–January 1999 and March 1999–January 2001 as more extensive sampling from 1999 through 

2001 may result in biased comparisons.

Collection methods: 1984–2001 [A summary of 

sampling in Florida Bay by month, habitat, and gear is 

presented in Table 1.] From the initial sampling trip in 

May 1984 through January 1999, a two-boat (5-m long), 

otter trawl was employed. In March 1999, the two boats 

were replaced with one 5.5 m boat for safety reasons. 

This required re-calibrating the opening of the mouth 

of the otter trawl to calculate area sampled. The otter 

trawl had a 3.4 m head rope, 3.8 m foot rope equipped 

with a 3 mm galvanized tickler chain, 6 mm mesh in the 

body, and a 3 mm tailbag. 

A surface trawl was employed only in 1984–1985. The 

surface trawl measured 6.6 m at the head rope, 6.2 m at 

the foot rope, was 0.7 m deep, had 6 mm mesh in the 

body and 3 mm mesh in the tailbag. The surface trawl, 

without doors to open the net, was pulled by two boats 

each angled about 45° away from the intended trawl path. 

The surface and otter trawls were towed at a speed 

of approximately 2.0 m–sec for 2 min, unless the net was 

clogged with detritus. A floating marker was deployed 

at the beginning and end of each tow and the distance 

Chapter 2. Methods 

Table 1

Summary of sampling in Florida Bay by month, habitat, and gear. The symbol X indicates sampling occurred. Shaded 

areas indicate annual periods of bi-monthly, otter trawl collections in basins. Periods are separated by variable shading. For 

example, the first annual period is from July 1984 through May 1985, the second from July 1994 through May 1995.

Month1 Habitat Gear

Year Jan Mar May Jun Jul Sep Nov Channels2 Basins Otter trawl Surface trawl

1984 X X X X X X X X

1985 X X X X X X X X

1994 X X X X X X

1995 X X X X X X X

1996 X X X X X X X X X X

1997 X X X X X

1998 X X X X X X X X

1999 X X X X X X X

2000 X X X X X X X X

2001 X X X

1

 Sampling did not occur in February, April, August, October, and November.

2

 Data from samples taken in channels in 1984–1985 were only used to depict the distribution and abundance of pelagic species collected by surface trawl. 

Sampling in channels with an otter trawl occurred in 1984–1985 and 1994–1996, but data were only used for length-frequency distributions.
5

Figure 2.1

Location of Florida Bay strata (top) and subdivisions (bottom).
6 

between buoys was measured with an optical range 

finder. The area covered by each trawl transect was 

calculated knowing the distance towed and the mouth 

opening of the net. Densities (number of fish 1000 m–2) 

were used as an index of abundance. 

Temperature was measured at the surface with a 

standard hand-held thermometer, and salinity measured at the surface with a refractometer.

For seagrass we measured short-shoot density, aboveground standing crop, blade length, and species 

composition. In 1984–1985 we collected three 100 cm2 

replicates of seagrass at the beginning, middle, and end 

of each trawl line, but in July 1994 through November 

1995, we used a 1 m2 quadrate subdivided into sixteen 

0.062 m2 subsections as described by Thayer et al. 

(1994). Three random samples were taken at each location when vegetation was present. If shoots exceeded 40 

per subsection, a 0.01 m2 quadrate was used. In January 

1996, a 0.01 m2 core was used to collect seagrass.

Processing of specimens Fishes and selected invertebrates were preserved in 10% formalin, transferred 

to 95% ethyl alcohol, identified to species in the 

laboratory, and then weighed. Measurements of fishes 

were in standard length (SL), except for seahorses 

(Hippocampus) where the tip of the tail (outstretched 

body) was measured to the corona (Ginsburg, 1937). 

Fish lengths reported from the literature are in notochord length (NL) for preflexion and flexion stage 

larvae, standard length (SL), total length (TL), and 

fork length (FL). 

Plants were removed at the sediment surface, placed 

in a labeled bag on ice, and returned to the laboratory 

for density and biomass measures. At the laboratory, 

individual short shoots of each seagrass species were 

sorted by species. Individual short shoots of each species 

were counted and separated from any below ground 

materials. Carbonates and epiphytes were removed 

by soaking seagrasses in 10% phosphoric acid until 

effervescence ceased, and then re-washed in seawater. 

Seagrasses were dried at 80o

 C to a constant weight and 

weighed to the nearest 0.001 g. Data were averaged at 

each site for each species.

Data analysis and presentation

Temperature and salinity Regression models (SAS Institute, 1997; PROC REG) were used to depict patterns 

of temperature and salinity in Florida Bay. We tested 

for significant differences (∝=0.05) among stratum and 

subdivision (Fig. 2.1), year, and month. Only bi-monthly 

data (January, March, May, July, September, and November) for six annual periods were included as only 

these months were consistently sampled throughout the 

period 1984–2001 (Table 1). 

Seagrasses A non-parametric Kruskal-Wallis test 

(Sokal and Rohlf, 1981) with ∝=0.05 was used to test 

differences in seagrass short-shoot densities (number 

m–2) and seagrass above-ground biomass (g dry weight 

m–2) among stratum and subdivision (Fig. 2.1), year, and 

month. As detailed above, only bi-monthly samples for 

six annual periods were included (Table 1). 

Seagrass distribution data were presented in GIS generated, six-panel figures by stratum, subdivision, and 

year for individual species (e.g., see Figs. 3.7 and 3.8). 

In addition, we included a three-panel figure depicting mean seagrass densities, and standing crop by year, 

month, stratum, and subdivision (e.g., see Figs. 3.5 and 

3.6). Seagrass composition and short-shoot density data 

from 1984–1985 and 1994–1996 were a major part of a 

data set that was used to evaluate the composition and 

density of seagrasses relative to changes in environmental conditions in Florida Bay (Thayer et al., 1999).

Fishes A species was included when the total number 

collected was ≥50. There were fourteen pelagics and 

32 non-pelagics included (Table 2). A non-parametric 

Kruskal-Wallis test (Sokal and Rohlf, 1981) with ∝=0.05 

was used to test differences in mean densities (numbers 

1000 m–2) of a given species (all non-pelagics, and those 

pelagics that numbered ≥50) among years, months, stratum, and subdivision for otter trawl bi-monthly basin collections (1984–2001; see Table 1 for the designation of 

sampling periods). For surface trawl data (1984–1985), 

the same test was used to test differences in mean densities of only pelagic species, among months, stratum, 

and basins and channels. For surface trawl collections 

all months in 1984–1985 were included. 

Mean Wilcoxon scores (rank sums) were used to estimate the frequency (i.e., the percentage of stations) a 

species occurred. Levels of species abundance were arbitrarily assigned to the following categories: very abundant, ≥10,000 total fish collected; abundant, <10,000 

and ≥1,000; common, <1,000 and ≥100; uncommon, 

<100. Fish species accounts are presented by family, with 

each family constituting a chapter. However, the families Atherinidae and Atherinopsidae, Fundulidae, and 

Cyprinodontidae were combined to avoid confusion. 

For a summary of sampling by year, month, habitat, and 

gear, see Table 1.

Fish species data are presented in four sections. The 

sections on Life history and Diet were derived from the 

literature. The section on Abundance and distribution

consists of data from bi-monthly otter trawl collections 

(1984–2001), and/or surface trawl collections of pelagic 

species in basins and channels (1984–1985). Data from 

these two gears are presented separately. In addition, 

comparisons with other studies are included here typically following our results. The section on Length-frequency distributions consists of length measurements 
7

Table 2

Summary of selected ecological characteristics of juvenile and small resident fishes in Florida Bay reported in this study. 

Transient species spend only a portion of their life history in Florida Bay and spawn outside the Bay.

 Warm Benthic/ Canopy

Family Species Tropical temperate Resident Transient demersal Pelagic dweller

Clupeidae Brevoortia smithi X X X

Harengula jaguana X X X

Opisthonema oglinum X X X X

Sardinella aurita X X X X

Engraulidae Anchoa hepsetus X X X

A. mitchilli X X X

Ariidae Arius felis X X X 

Synodontidae Synodus foetens X X X X 

Batrachoididae Opsanus beta X X X 

Hemiramphidae Chriodorus atherinoides X X (?) X

Hemiramphus brasiliensis X X X X

Hyporhamphus unifasciatus X X (?) X

Belonidae Strongylura notata X X X

S. timucu X X X X

Cyprinodontidae Floridichthys carpio X X X

Fundulidae Lucania parva X X X

Atherinidae Atherinomorus stipes X X X

Hypoatherina harringtonensis X X X

Atherinopsidae Membras martinica X X X

Syngnathidae Anarchopterus criniger X X X X

Syngnathus floridae X X X X

S. scovelli X X X X

Hippocampus erectus X X X X

H. zosterae X X X X

Lutjanidae Lutjanus griseus X X X

L. synagris X X X

Gerreidae1 Eucinostomus gula X X X

Haemulidae Haemulon plumieri X X X X

H. sciurus X X X

Orthopristis chrysoptera X X X

Sparidae Archosargus probatocephalus X X X

Calamus arctifrons X X X

Lagodon rhomboides X X X

Sciaenidae Bairdiella chrysoura X X X

Cynoscion nebulosus X X X

Sphyraenidae Sphyraena barracuda X X X X

Scaridae Nicholsina usta X X X X

Callionymidae Diplogrammus pauciradiatus X X X

Gobiidae Gobiosoma robustum X X X

Microgobius gulosus X X X

M. microlepis X X X 

Monacanthidae Monacanthus ciliatus X X X X

M. hispidus X X 

Ostraciidae Lactophrys quadricornis X X X (?) X

Tetraodontidae Sphoeroides nephelus X X X

Diodontidae Chilomycterus schoepfi X X X

1 Eucinostomus argenteus was also identified, but most were mis-identified and were most likely E. harengulus.

from all collections, except 1984–1985 when no measurements were taken. Length-frequency distributions 

were used, when possible, to estimate life stage captured, spawning times, recruitment into Florida Bay for 

those species which spawn outside the Bay, and growth. 

Additional material from the literature was added when 

appropriate.

For each fish species an un-numbered table is presented that depicts the raw numbers collected and overall rank of the species by gear and habitat. For surface 
8 

trawl collections in basins and channels, a total of 14 

pelagic species (Table 2) in each habitat were ranked. 

For otter trawl collections in basin habitats, 39 species 

were ranked. 

The graphic presentation of data was dependent on 

gear and raw numbers of fish collected. Pelagic fish 

species collected by surface trawl in 1984–1985 were 

presented in one-panel GIS generated figures depicting their distribution in basins and channels by stratum 

(e.g., Fig. 4.1). Also included were figures (bar graphs) 

depicting the mean densities (numbers 1000 m–2) by 

month and strata (e.g., Fig. 4.2). However, when a pelagic species was collected by otter trawl in basins, and 

sample size was >50 fish, GIS generated two-panel (<100 

total fish; e.g., Fig. 4.10) or six-panel (≥100 total fish) 

figures were included (e.g., Fig. 4.5). Also included for 

this gear were figures (bar graphs) depicting the mean 

densities (numbers 1000 m–2) by year, month, and 

strata/subdivision (e.g., Fig. 4.6). For all other species 

(i.e., canopy dwellers and benthics that made up the 

majority of species) that totaled >50 fish, we included 

GIS generated two-panel (<100 total fish; e.g., Fig. 6.4) 

or six-panel figures (≥100 total fish; e.g., Fig.6.1) depicting their distribution. Also included for these species 

were figures (bar graphs) depicting the mean densities 

(numbers 1000 m–2) by year, month, and strata/subdivision (e.g., Fig. 6.2).

Juvenile and small resident fish data from 1984–1985 

and 1994–1996 were a major part of a data set that was 

used in a manuscript to evaluate the composition of 

ichthyofauna relative to changes in environmental conditions in Florida Bay (Thayer et al., 1999).
9

Temperature

Mean annual water temperatures for Florida Bay ranged 

from 25.4oC in 1999–2000 to 27.2oC in 1997–1998 (Fig. 

3.1A). Mean temperatures among strata (1984–1985, 

1994–1998) were similar and ranged from 25.9oC in 

the Central stratum to 26.9oC in the West stratum. 

Mean temperatures were similar among subdivisions 

(1999–2001) and ranged from 25.2oC in the Eastern 

subdivision to 25.8oC in the Atlantic subdivision (Fig. 

3.1B). Generally, coldest temperatures occurred in 

January, followed by March; warmest temperatures 

occurred from May through September (Fig. 3.2). 

Our observations support those of Boyer et al. (1999), 

who concluded that water temperatures in Florida Bay 

showed pronounced seasonal patterns, but no interannual trends.

Salinity

Overall, mean monthly salinities were highest in July 

and May. High salinities during this wet period are a 

result of high water temperatures, high evaporation 

rates, and long water residence times (Sogard et al., 

1987; Boyer et al., 1999). Salinities were similar among 

strata (Fig. 3.3), but differed among subdivisions (Fig. 

3.4). Salinities were similar and highest in the Atlantic, 

Central, Western, and Gulf subdivisions. Moderate 

salinities were observed in the Eastern subdivision; 

whereas salinities were low and variable in the Northern 

subdivision (Fig. 3.4), the latter an area influenced by 

freshwater inputs from Taylor Slough (Fig. 1.1; Boyer 

et al., 1999).

Our findings of yearly trends in salinity differ from 

Boyer et al. (1999). From a robust data set, they found a 

38% decrease is salinity during the period 1991–1997 in 

the eastern area of the Bay, a 29% decrease in the central area, and a 15% decrease in the western area. These 

declines were linked to wetter than normal conditions 

in 1994–1996. Although we did not find any significant 

differences among years, the pattern we observed suggested a decline in salinities from 1984–1985, while 

salinities remained relatively low during the last three 

annual sampling periods (Fig. 3.1A). 

A comprehensive study of salinity trends in Florida 

Bay is given by Orlando et al. (1997), who depicted salinity characteristics by salinity contours using 40 years 

(1955–1995) of data. They observed three persistent 

features which included a hypersaline “bull’s-eye” area 

in the north-central area of the Bay, brackish salinities 

in the northeast, and near-ocean salinities along the 

western and southern boundaries. They noted that longterm changes might include decreasing salinities in 

Chapter 3. Environmental variables

A

Year

5891- 48915991- 4991

6991

8991- 79910002- 99911002- 0002

( er ut arep me T o ) C 24.5

25.0

25.5

26.0

26.5

27.0

27.5

28.0

)t pp( yti nil aS

30.0

31.0

32.0

33.0

34.0

35.0

36.0

B

tsa E

l art ne C

tse W

nretsa E

nrehtr o N

cit nalt A

l art ne C

nretse W

fl u G

( er ut arep me T o ) C 24.5

25.0

25.5

26.0

26.5

27.0

27.5

28.0

)t pp( yti nil aS

18.0

20.0

22.0

24.0

26.0

28.0

30.0

32.0

34.0

36.0

38.0

____________ ___________________________

 Stratum Subdivision

Figure 3.1

Mean temperature and salinity by (A) year and (B) 

stratum and subdivision. Error bars are one standard 

deviation from the mean. Solid circles represent 

temperature, open circles represent salinity. For 

location of strata and subdivisions see Fig. 2.1.

the northeastern area of the Bay, while the hypersaline 

“bull’s eye” area might be intensifying.

Seagrasses

Thalassia testudinum (turtle grass) Turtle grass, a 

member of the Caribbean flora, is the dominant seagrass in Florida Bay both in spatial distribution and 

biomass (Zieman et al., 1989; Hemminga and Duarte, 

2000).

We observed a distinct temporal trend in short-shoot 

densities of turtle grass. Densities declined Bay-wide 

from 722 short-shoots m–2 in 1984–1985 to 12 shortshoots m–2

 in 1996. Some recovery in short-shoot densi-
10

Jul 1984-May 1985

14

18

22

26

30

34

Jul 1994-May 1995

er ut arep me T o

C

14

18

22

26

30

34

Jan 1996-Nov 1996

Month

Jan Mar May Jul Sep Nov

14

18

22

26

30

34

Sep 1997-Jul 1998

Mar 1999-Jan 2000

Mar 2000-Jan 2001

Month

Jan Mar May Jul Sep Nov

Figure 3.2

Mean monthly temperatures by year. Error bars are one standard deviation from the mean.

ties has occurred since 1996, but densities are well below 

1984–1985 levels (Figs. 3.5 and 3.7). No significant differences in monthly densities were observed.

Short-shoot densities of turtle grass differed among 

subdivisions, but not strata. Unexpectedly, short-shoot 

densities were relatively low in the Western subdivision 

(Fig. 3.5). Our original sampling design (Fig. 2.1) was 

based on the premise that there is a general increase in 

standing crop of turtle grass from the northeastern area 

of Florida Bay to the western part of the Bay (Thayer 

and Chester, 1989). Furthermore, short-shoot densities 

have been shown to be positively associated with standing crop in Florida Bay (Hall et al., 1999). 

Differences in standing crop mirrored our observations for turtle grass short-shoot densities (Figs. 3.6 and 

3.8 vs. 3.5 and 3.7), but standing crop of turtle grass 

was not positively associated with short-shoot densities. 

However, standing crop generally increased from east 

to west and was highest in western Florida Bay (Figs. 3.6 

and 3.8). Disparity between short-shoot densities and 

standing crop is related to spatial differences in blade 

length. Mean blade lengths (cm) of turtle grass were 
11

July 1984- May 1985

( yti nil aS o

/ oo )

15

20

25

30

35

40

45

50

East Stratum

Central Stratum

West Stratum 

July 1994-May 1995 ( yti nil aS o

/ oo )

15

20

25

30

35

40

45

50

Jan 1996-Nov 1996

( yti nil aS o

/ oo )

20

25

30

35

40

45

Sep 1997-Jul 1998

Month

Jan Mar May Jul Sep Nov

( yti nil aS o

/ oo )

20

25

30

35

40

45

Figure 3.3

Mean monthly salinities by stratum and year. Error bars 

are one standard deviation from the mean. For location 

of strata see Fig. 2.1.

Mar 2000-Jan 2001

Month

Jan Mar May Jul Sep Nov

( yti nil aS o

/ oo )

0

10

20

30

40

50

Mar 1999-Jan 2000

( yti nil aS o

/ oo )

0

10

20

30

40

50

Atlantic, Central, Gulf, and Western Subdivisions

Eastern Subdivision

Northern Subdivision

Figure 3.4

Mean monthly salinities by subdivision and year. Error 

bars are one standard deviation from the mean. For 

location of strata see Fig. 2.1. Subdivisions were pooled 

when there were no significant differences among 

them. 

32.6, 30.9, 13.5, 13.3, 11.7, and 9.9 in the Gulf, Western, 

Central, Atlantic, Northern, and Eastern subdivisions, 

respectively.

Halodule wrightii (shoalgrass) Shoalgrass, a member 

of the Caribbean flora, is dominant only in limited areas 

of Florida Bay where temperature and salinity extremes 

occur; i.e., where physical conditions rule out dense 

stands of turtle grass (Zieman et al., 1989; Hemminga 

and Duarte, 2000). 

We observed that short-shoot densities of shoalgrass 

declined from a bay-wide average of 142 short-shoots 

m–2 in 1984–1985 to 1 short-shoot m–2 in 1996. Densities 

from 1997–1998 through 2001 were similar to 1984–

1985 densities, but slightly lower (Figs. 3.5 and 3.9). 

Bay-wide standing-crop biomass declined from 4.2 

g m–2

 in 1984–1985 to 0.7 g m–2 in 1996 (Figs. 3.6 and 

3.10). Shoalgrass biomass was highest in the West stratum (3.8 g m–2

) and Western subdivision (7.3 g m–2). 

Low biomass values were observed in the East stratum 

(0.04 g m–2) and Atlantic and Eastern subdivision (0.04 g 

m–2). Shoalgrass was relatively more abundant in the 

central part of the Bay in 2001, perhaps indicating a 

recovery of seagrasses in this area (Fig. 3.9) as shoalgrass 

is an early colonizer in disturbed areas (Zieman and 

Adams, 1982). 

Syringodium filiforme (manatee grass) Manatee grass, 

a member of the Caribbean flora, is locally dominant in 

areas where lower light energy occurs; i.e., where turtle 

grass is light limited. It would be expected to occur in 

deeper areas in Florida Bay (Zieman et al., 1989; Hemminga and Duarte, 2000). 

We observed that short-shoot densities of manatee 

grass differed only spatially. This species had the most 

limited distribution of the seagrasses in Florida Bay 

(Figs. 3.11 and 3.12). Manatee grass was collected 

only in the West stratum, mainly in the Western and 
12

Figure 3.5

Mean number of seagrass short-shoots by (A) year and 

(B) stratum/subdivision. For location of strata and 

subdivisions see Fig. 2.1.

5891- 48915991- 49916991

8991- 79910002- 9991

1002- 0002

m/r eb mun( st oohs-tr ohs nae M 2

)

0

200

400

600

800

1000

1200

1400

Thalassia testudinum

Halodule wrightii

Syringodium filiforme

A

 ______________ _____________________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nr ehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

/msr eb mun( st oohs-tr ohs nae M 2)

0

200

400

600

800

Thalassia testudinum

Halodule wrightii

Syringodium filiforme

B

5891- 4891

5991- 499169918991- 7991

0002- 999110- 0002

m/t hgi e wyr d s mar g( por c gni dnats nae M 2)

0

20

40

60

80

100

120

140

160

Thalassia testudinum

Halodule wrightii

Syringodium filiforme

A

 ______________ _____________________________

 Stratum Subdivision

tsa El art ne Ctse W

nr etsa E

nr ehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

m/t hgi e wyr d s mar g( por c gni dnats nae M 2)

0

20

40

60

80 Thalassia testudinum

Halodule wrightii

Syringodium filiforme B

Figure 3.6

Mean above-ground standing crop of seagrasses by (A) 

year and (B) stratum/subdivision. For location of strata 

and subdivisions see Fig. 2.1.

Gulf subdivisions (Figs. 3.5 and 3.11). Although we 

found no significant differences in short-shoot densities among years, Thayer et al. (1999) and Hall et 

al. (1999) found significant declines of short-shoot 

densities following the seagrass die-off in Florida Bay. 

Our data for 1984–1985 versus 1994–1996 indicated a 

dramatic decline during the latter time period. Baywide densities of short-shoots declined from 248 m–2

to 1 m–2 (Fig. 3.5). 

The pattern of standing crop biomass, including 

the decline noted by Hall et al. (1999) and Thayer et 

al. (1999), was similar to that observed for short-shoot 

densities, with highest standing crop values observed in 

western Florida Bay (Figs. 3.6 and 3.12). 
13

Figure 3.7

Mean short-shoot densities (numbers m–2

) of turtle grass (Thalassia testudinum) by year, 

stratum, and subdivison. For location of strata and subdivisions see Fig. 2.1.
14

Figure 3.8

Mean above-ground biomass (grams dry weight m–2

) of turtle grass (Thalassia 

testudinum) by year, stratum, and subdivision. For location of strata and subdivisions 

see Figure 2.1.
15

Figure 3.9

Mean short-shoot densities (numbers m–2

) of shoalgrass (Halodule wrightii) by year, 

stratum, and subdivision. For location of strata and subdivisions see Fig. 2.1.
16

Figure 3.10

Mean standing-crop biomass (grams dry weight m–2

) of shoalgrass (Halodule wrightii) by 

year, stratum, and subdivision. For location of strata and subdivisions see Figure 2.1.
17

Figure 3.11

Mean short-shoot densities (numbers m–2

) of manatee grass (Syringodium filiforme) by 

year, stratum, and subdivision. For location of strata and subdivisions see Fig. 2.1.
18

Figure 3.12

Mean above-ground biomass (grams dry weight m–2

) of manatee grass (Syringodium 

filiforme) by year, stratum, and subdivison. For location of strata and subdivisions see 

Figure 2.1.
19

Brevoortia smithi (yellowfin menhaden)

Total numbers collected and rank of yellowfin menhaden by 

sampling method. There were 14 species collected by surface 

trawl in each habitat.

Gear Habitat Year Number Rank

Surface trawl Basins 1984–1985 37 13

Surface trawl Channels 1984–1985 76 8

Life history An Atlantic population of yellowfin menhaden occurs from Cape Lookout, North Carolina to Jupiter 

Inlet, Florida. A Gulf of Mexico population occurs from 

Florida Bay to Louisiana (Dahlberg, 1970). Yellowfin menhaden spawn during winter in nearshore waters as early as 

November, but most commonly from February to March. 

Eggs, larvae, and juveniles are pelagic (Hettler, 1968; Dahlberg, 1970). Size of larvae at hatching is approximately 3.0 

mm NL. Transformation to the juvenile stage begins at 14 

mm SL and is complete between 20–23 mm SL (Houde 

and Swanson, 1975). Sexual maturity is attained at approximately 220 mm FL (Pattillo et al., 1997). 

Chapter 4. Family Clupeidae

Diet Juveniles are filter-feeding planktivores that feed 

on phytoplankton, small zooplankton, and detritus 

(Dahlberg, 1970; Ahrenholz, 1991).

Abundance and distribution Yellowfin menhaden 

were commonly collected by surface trawl in Florida 

Bay, but were limited to the western portion (Fig. 4.1). 

These fish were most likely spawned in Gulf of Mexico 

waters during the winter. They were mainly collected in 

1984–1985, with highest densities from channel collections (Fig. 4.2). Significant differences in densities were 

only observed between months. They were collected 

only in spring, mainly in March (Fig. 4.2). They were 

never collected by otter trawl.

Tabb et al. (1962) stated that this species was abundant in muddy shallow waters of Whitewater and Coot 

Bays (salinities ranged from 12–38 ppt), when Coot Bay 

was connected to Florida Bay by the Buttonwood Canal. 

However, Thayer et al. (1987a) collected few yellowfin 

menhaden in Coot and southern Whitewater Bays after 

the connection to Florida Bay was blocked.

Length-frequency distributions No data were available.

Figure 4.1

Mean densities (numbers 1000 m–2

) of yellowfin menhaden (Brevoortia smithi) collected 

by surface trawl in Florida Bay basins and channels during 1984–1985.Florida Bay 

basins are divided into three strata. For designations of strata see Fig. 2.1.
20

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

A

Stratum

East Central West Channels

ytisne D

0.0

0.5

1.0

1.5

2.0

B

Figure 4.2

Mean densities (numbers 1000 m–2

) of yellowfin 

menhaden (Brevoortia smithi) collected by surface trawl 

in Florida Bay basins and channels during 1984–1985 

by (A) month and (B) stratum. Florida Bay basins are 

divided into three strata. For designations of strata see 

Fig. 2.1.
21

Harengula jaguana (scaled sardine)

Total numbers collected and rank of scaled sardine by sampling method. There were 39 species collected by otter trawl; 

14 by surface trawl in each habitat.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 123 25

Surface trawl Basins 1984–1985 27 14

Surface trawl Channels 1984–1985 102 7

Life history Scaled sardine reside in warm temperate 

and tropical coastal waters from New Jersey to Florida, 

throughout the Gulf of Mexico, and along Caribbean 

coastal waters to southern Brazil, Bermuda, Bahamas, 

and West Indies (Whitehead, 1978). They are extremely 

rare north of Cape Hatteras, North Carolina and not 

commonly collected north of Cape Canaveral, Florida 

(Shaw and Drullinger, 1990). This species spawns in 

nearshore waters of the northern Gulf of Mexico during 

March–August (Houde and Fore, 1973), but mainly in 

June and July (Shaw and Drullinger, 1990). In Florida 

waters (Tampa Bay), spawning appears to occur year 

round (Pierce et al., 2001). Collins and Finucane (1984) 

mainly collected larvae in May and August in Gulf of 

Mexico coastal waters off southwest Everglades National 

Park. They did not collect larvae in November or in 

estuarine waters. Eggs, larvae, and juveniles are pelagic. 

Size of larvae at hatching is 2.4 mm NL. Transformation to the juvenile stage is completed at 22–24 mm SL 

(Houde et al., 1974). Sexual maturity, estimated from 

specimens collected mainly in Biscayne Bay, is attained 

at 75–85 mm SL (Martinez and Houde, 1975).

Diet Quantitative food studies of fishes from the Whitewater Bay system, adjacent to Florida Bay, indicated that 

scaled sardines between 64 and 96 mm SL fed mainly 

on amphipods and mysids (Odum and Heald, 1972). In 

another Florida study, juveniles between 21 and 25 mm 

SL fed mainly on veligers, followed by copepods and 

cypris larvae. Between 26 and 30 mm SL, crab megalops 

dominated the diet with a marked decrease in the occurrence of veligers and copepods (Carr and Adams, 1973). 

Abundance and distribution Based on surface trawl 

data, there were no significant spatial or temporal differences in densities of scaled sardine, even though 

densities appeared to be higher in channels (Figs. 4.3 

and 4.4). Scaled sardine, like yellowfin menhaden, appeared to be most abundant in channel habitats.

Based on otter trawl data, only significant interannual 

differences were observed. Highest densities occurred 

in 1994–1995, lowest in 1999–2001 (Figs. 4.5 and 4.6). 

The distribution of this species in Florida Bay suggests 

recruitment into the Bay from both the Atlantic and 

Gulf of Mexico (Fig. 4.3).

Figure 4.3

Mean densities (numbers 1000 m–2

) of scaled sardine (Harengula jaguana) collected by 

surface trawl in Florida Bay basins and channels during 1984–1985. Florida Bay basins 

are divided into three strata. For designations of strata see Fig. 2.1.
22

Month

Jan Mar May June Jul Sep Nov

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

A

Stratum

East Central West Channels

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

3.0

B

Figure 4.4

Mean densities (numbers 1000 m–2

) of scaled sardine 

(Harengula jaguana) collected by surface trawl in 

Florida Bay basins and channels during 1984–1985 

by (A) month and (B) stratum. Florida Bay basins are 

divided into three strata. For designations of strata see 

Fig. 2.1.

Scaled sardine is one of the more abundant clupeids 

occurring in Florida Bay (FMRI1,2,3,4)and were extremely 

abundant during the summer and fall in Buttonwood 

Canal when it was connected to Florida Bay (Roessler, 

1970). Other studies of juvenile fishes in Florida Bay 

have shown that scaled sardine abundances declined 

markedly between 1994–1995 and 1996–1997, and 

this species was consistently more abundant in August 

(FMRI1,2,3,4).

Length-frequency distributions There appeared to be 

constant recruitment into Florida Bay from late spring 

to late fall. One size class was dominant; it appeared to 

be comprised of recently transformed juveniles (20–29 

mm SL) (Fig. 4.7). Sexually-mature scaled sardines 

(>75 mm) were rarely collected in Florida Bay. This 

species attains a maximum size of 180 mm TL (Robins 

and Ray, 1986), which is larger than the specimens we 

collected.

2

 FMRI. 1996. Fisheries-independent monitoring program, 1995 annual data summary report. Florida Marine Research Institute, Florida Department of Environmental Protection, St. Petersburg, FL, 

195 p.

3 FMRI. 1997. Fisheries-independent monitoring program, 1996 annual data summary report. Florida Marine Research Institute, Florida Department of Environmental Protection, St. Petersburg, FL, 

227 p.

4

 FMRI. 1998. Fisheries-independent monitoring program, 1997 annual data summary report. Florida Marine Research Institute, Florida Department of Environmental Protection, St. Petersburg, FL, 

236 p.
23

Figure 4.5

Mean densities (numbers 1000 m–2

) of scaled sardine (Harengula jaguana) from bi-monthly otter trawl collections in Florida 

Bay basins. Florida Bay is divided into three strata from 1984–1998 and six subdivisions from 1999–2001. For designations 

of strata and subdivisions see Fig. 2.1.
24

Year

5891- 4891

5991- 4991

6991

8991- 7991

0002- 99911002- 0002

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

1.4

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.2

0.4

0.6

0.8

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

0.6

A

B

C

Figure 4.6

Mean densities (numbers 1000 m–2

) of scaled sardine 

(Harengula jaguana) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
25

January

y

c

n

e

u

q

e

r

F

0

100

200

300

March

0

100

200

300

May

0

100

200

300

June

0

100

200

300

July

0

100

200

300

September

0

100

200

300

November

Size Class (mm)

41- 0142- 02

43- 03

44- 0445- 0546- 06

47- 07

48- 08

49- 09

401- 001

411- 011

421- 021431- 031441- 041

0

100

200

300

(1) (3) (5) (4) (1) (1) (3) (1) (1)

(1) (1) (1) (1) (1) (2) (1) (1) (2) (1) (1)

(7) (6) (6) (1) (2) (2) (1)

(9) (7) (1) (8) (8) (3) (1) (1) (1) (1) (1)

(1) (9) (1) (1)

(5) (3) (5) (3)

(1) (3) (1) (1) (4) (2) (6) (1) (1) (4) (1) (1) (1) (1) (1) (1)

Figure 4.7

Monthly length-frequency distributions of scaled sardine (Harengula jaguana) from Florida 

Bay. Values are shown for numbers <10 for clarity.
26

Opisthonema oglinum (Atlantic thread herring)

Total numbers collected and rank of Atlantic thread herring 

by sampling method. There were 39 species collected by otter 

trawl; 14 by surface trawl in each habitat.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 57 37

Surface trawl Basins 1984–1985 119 10

Surface trawl Channels 1984–1985 8 13

Life history Atlantic thread herring occur in coastal 

waters from Cape Cod (although not common north 

of North Carolina) to Florida, and throughout the Gulf 

of Mexico to Brazil, West Indies, and Bermuda (Whitehead, 1978; Robins and Ray, 1986; Smith, 1994). 

They spawn during May–June off North Carolina 

(Whitehead, 1978), and in the northern Gulf of Mexico 

their larvae are commonly collected in inner shelf waters during May–August (Shaw and Drullinger, 1990). 

Collins and Finucane (1984) mainly collected larvae in 

August in Gulf of Mexico coastal waters off southwest 

Everglades National Park. Larvae were also collected in 

November, February, and May. They were not collected 

by Collins and Finucane (1984) in estuarine waters. 

Eggs, larvae, and juveniles are pelagic. Size of larvae 

at hatching is 3.8–4.0 mm NL. Transformation to the 

Figure 4.8

Mean densities (numbers 1000 m–2

) of Atlantic thread herring (Opisthonema oglinum) 

collected by surface trawl in Florida Bay basins and channels during 1984–1985. Florida 

Bay basins are divided into three strata. For designations of strata see Fig. 2.1.

juvenile stage occurs between 17–25 mm SL (Richards 

et al., 1974). 

Diet Juvenile Atlantic thread herring, between 21 and 

24 mm SL, feed mainly on veligers, followed by detritus 

and copepods (Carr and Adams, 1973). The portion 

of the diet consisting of veligers decreases during ontogeny with concurrent increases in copepods, mysids, 

and megalops for juveniles 36–40 mm SL. Detritus is a 

significant part of the juvenile diet.

Abundance and distribution Atlantic thread herring, 

sampled by surface trawl, were collected only in the West 

stratum and only in May (Figs. 4.8 and 4.9). 

Based on otter trawl samples (Figs. 4.10 and 4.11), 

Atlantic thread herring were collected in highest densities in 1997–1998 and 2000–2001, and were absent in 

1994–1995 and 1999–2000. From 1984 through 1996 

highest densities were observed in the Central stratum, 

lowest in the West stratum. No significant differences 

in monthly densities were observed in the otter trawl 

collections in basins, even though September and November mean densities appeared to be higher than the 

other months. This species appeared to prefer basin 

habitats to channels, but compared to other pelagics 

(except clupeids), this species ranked relatively low in 

total numbers collected (see table above).

Other studies in Florida Bay report patterns of abundances similar to our observations for Atlantic thread 
27

herring; i.e, abundances declined markedly between 

1994–1995 and 1996–1997 (FMRI1,2,3,4). In those studies, 

Atlantic thread herring were most abundant in August 

(1994 and 1996), July (1995), and October (1997). 

Atlantic thread herring were collected in comparable 

numbers to scaled sardines, which is in concordance 

with other studies in Florida Bay (Sogard et al., 1989a; 

(FMRI1,2,3,4). Hence, they, along with scaled sardine, 

could be considered the two most abundant clupeids in 

Florida Bay. They are also one of the two most abundant 

clupeids in adjacent Biscayne Bay (Houde and Lovdal, 

1984). 

Length-frequency distributions Larvae or transforming juveniles recruited into Florida Bay throughout most 

of the year (Fig. 4.12). Smallest larvae and transforming 

juveniles (<25 mm SL) were collected by otter trawl in 

all months except January, March, and June. Length-frequency distributions in March indicate that recruitment 

into Florida Bay also occurred during winter. Few large 

juveniles were collected with the otter trawl. The largest 

Atlantic thread herring collected were ca. 155 mm SL, 

much smaller than the reported maximum length of 

300 mm TL (Robins and Ray, 1986).

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

A

Stratum

East Central West Channels

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

3.0

B

Figure 4.9

Mean densities (numbers 1000 m–2

) of Atlantic thread 

herring (Opisthonema oglinum) collected by surface 

trawl in Florida Bay basins and channels during 1984–

1985 by (A) month and (B) stratum. Florida Bay basins 

are divided into three strata. For designations of strata 

see Fig. 2.1.
28

Figure 4.10

Mean densities (numbers 1000 m–2

) of Atlantic thread herring (Opisthonema oglinum) 

from bi-monthly otter trawl collections in Florida Bay basins. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 1999–2001. For designations of 

strata and subdivisions see Fig. 2.1.
29

Year

5891- 48915991- 49916991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.00

0.05

0.10

0.15

0.20

0.25

0.30

Month

Jan Mar May Jul Sep Nov

ytisne D

0.00

0.05

0.10

0.15

0.20

0.25

0.30

0.35

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

A

B

C

Figure 4.11

Mean densities (numbers 1000 m–2

) of Atlantic thread 

herring (Opisthonema oglinum) from bi-monthly otter 

trawl collections in Florida Bay basins by (A) year, 

(B) month, and (C) stratum/subdivision. Florida Bay 

is divided into three strata from 1984–1998 and six 

subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
30

January

y

c

n

e

u

q

e

r

F

0

20

40

60

80

March

0

20

40

60

80

May

0

20

40

60

80

June

0

20

40

60

80

July

0

20

40

60

80

September

0

20

40

60

80

November

Size Class (mm)

91- 51

92- 52

93- 5394- 54

95- 55

96- 56

97- 57

98- 58

99- 59

901- 501911- 511

921- 521931- 531

941- 541

951- 551

0

20

40

60

80

Figure 4.12

Monthly length-frequency distributions of Atlantic thread herring (Opisthonema oglinum) 

from Florida Bay.
31

Sardinella aurita (Spanish sardine)

Total numbers collected and rank of Spanish sardine by sampling method. There were 39 species collected by otter trawl; 

14 by surface trawl in each habitat.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 84 30

Surface trawl Basins 1984–1985 163 9

Surface trawl Channels 1984–1985 1 14

Life history Spanish sardine occur in coastal waters 

from Cape Cod to Florida, throughout the Gulf of Mexico, and along the Caribbean coast to southern Brazil, 

Bermuda, Bahamas, and West Indies (Whitehead, 1978). 

In the eastern Gulf of Mexico Spanish sardines have a protracted spawning season with a peak during spring and 

summer. Along southwest Florida they spawn primarily 

in the fall and winter (Houde and Fore, 1973; Shaw and 

Drullinger, 1990). Size of larvae at hatching is approximately 2.5 mm NL. Transformation to the juvenile stage 

begins at 16 mm SL and is completed at 23 mm SL (Ditty 

et al., 1994). In western Florida waters, 50% of females 

mature at 147 mm FL (Shaw and Drullinger, 1990).

Diet Juvenile Spanish sardine feed on diatoms and 

small copepods (Wang and Qiu, 1986).

Abundance and distribution Spanish sardine were 

common in surface trawl collections (but only collected 

at three stations) and uncommon in otter trawl basin 

collections. Based on surface trawl data (Fig. 4.13), 

there were no significant spatial or temporal differences in densities; however, Spanish sardine were only 

collected in the Central and West strata (Fig. 4.14). This 

species, like the Atlantic thread herring, appeared to 

prefer basin habitats to channels. 

In otter trawl collections in basins (Fig. 4.15), Spanish sardine were only collected during 1999–2000 and 

2000–2001 (Fig. 4.16). There were no significant differences in densities among months, although we failed 

to collect any specimens during three (January, March, 

and July) of the six months sampled. Spatially, highest 

densities were observed in the Gulf subdivision, moderate in the Central and Eastern subdivisions, low in 

the Northern subdivision, and they were absent in the 

Atlantic or Western subdivisions (Fig. 4.16).

Other studies of juvenile fishes in Florida Bay and 

adjacent waters indicate that this species is rare in 

Florida Bay and was also rare in the Buttonwood Canal 

when it was connected to Florida Bay (Roessler, 1970; 

FMRI1,2,3,4). 

Length-frequency distributions Based on limited 

length-frequency data, Spanish sardine recruited mainly 

to Florida Bay during spring (Fig. 4.17); however, some 

recruitment may occur throughout the year. In May 

specimens were mainly larvae and transforming juveniles (14–24 mm SL). In June juveniles were dominant. 

No sexually mature (ca. 145 mm SL) Spanish sardine 

were collected. The largest specimens collected by otter 

trawl were ca. 55 mm SL, much smaller than the reported maximum size of 250 mm TL (Robins and Ray, 1986).

Figure 4.13

Mean densities (numbers 1000 m–2

) of Spanish sardine (Sardinella aurita) collected by 

surface trawl in Florida Bay basins and channels during 1984–1985. Florida Bay basins 

are divided into three strata. For designations of strata see Fig. 2.1.
32

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

A

Stratum

East Central West Channels

ytisne D

0.0

1.0

2.0

3.0

4.0

B

Figure 4.14

Mean densities (numbers 1000 m–2

) of Spanish sardine 

(Sardinella aurita) collected by surface trawl in Florida 

Bay basins and channels during 1984–1985 by (A) 

month and (B) stratum. Florida Bay basins are divided 

into three strata. For designations of strata see Fig. 2.1.
33

Figure 4.15

Mean densities (numbers 1000 m–2

) of Spanish sardine (Sardinella aurita) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
34

Year

5891- 48915991- 4991

6991

8991- 79910002- 99911002- 0002

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

3.0

A

B

C

Figure 4.16

Mean densities (numbers 1000 m–2

) of Spanish sardine 

(Sardinella aurita) from bi-monthly otter trawl collections 

in Florida Bay basins by (A) year, (B) month, and (C) 

stratum/subdivision. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–

2001. For designations of strata and subdivisions see 

Fig. 2.1.
35

Figure 4.17

Monthly length-frequency distributions of Spanish sardine (Sardinella aurita) from Florida 

Bay.

January

y

c

n

e

u

q

e

r

F

0

25

50

75

100

March

0

25

50

75

100

May

0

25

50

75

100

June

0

25

50

75

100

July

0

25

50

75

100

September

0

25

50

75

100

November

Size Class (mm)

01<

41- 01

91- 51

42- 02

92- 52

43- 03

93- 53

44- 04

94- 54

45- 05

0

25

50

75

100
36

Chapter 5. Family Engraulidae

Anchoa hepsetus (striped anchovy)

Total numbers collected and rank of striped anchovy by sampling method. There were 39 species collected by otter trawl; 

14 by surface trawl in each habitat.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 81 31

Surface trawl Basins 1984–1985 496 4

Surface trawl Channels 1984–1985 29 11

Life history Striped anchovy inhabit shallow coastal 

waters from Nova Scotia to Florida, throughout the 

Gulf of Mexico, and along the Caribbean coast to Uruguay, the Bahamas, and the West Indies (Whitehead, 

1978; Able and Fahay, 1998). Striped anchovy spawn in 

harbors, estuaries, sounds, and offshore waters (Whitehead, 1978). Spawning occurs from April–July in South 

Carolina waters (Whitehead, 1978); throughout the 

year, with peaks in April–June, off Campeche, Mexico 

(Valencia et al., 1998); and November through April in 

Biscayne Bay, Florida (Jones et al., 1978). Eggs, larvae, 

and juveniles are pelagic, and occupy the higher part 

of the water column relative to other zooplanktivorous 

fishes (Allen et al., 1995). Size at hatching is 3.6–4.0 mm 

TL; transformation to the juvenile stage occurs at approximately 30 mm SL (Jones et al., 1978). The minimal 

size at sexual maturity is 43 mm and 48 mm TL for males 

and females, respectively (Valencia et al., 1998). 

Diet Juvenile striped anchovy (25–50 mm SL) feed 

mainly on veliger larvae (Carr and Adams, 1973). Copepods, mysids, zooea, and fish larvae are also consumed, 

but their contribution is small compared to veligers.

Abundance and distribution This species was commonly collected with a surface trawl, especially in basin 

habitats, but only in the northern Central and West 

strata, and only in May (Figs. 5.1 and 5.2). These spatial 

and temporal differences in densities were significant. 

The striped anchovy was uncommon in otter trawl collections (Fig. 5.3), and its distribution and abundance 

were most likely underestimated as this species occurs 

mainly in surface waters (Allen et al., 1995). There were 

significant differences in otter trawl densities among 

strata and subdivisions, and year, but not among months 

(Fig. 5.4). Highest densities were observed in the 

West stratum, the lowest in the East stratum; and they 

were absent in the Central stratum. From 1999–2001 

this species was only collected in the Gulf subdivision 

(Fig. 5.3).

Length-frequency distributions All life history stages 

of striped anchovy were collected in Florida Bay (Fig. 

5.5). The smallest larvae were collected in May, but 

larvae and transforming juveniles(<30 mm SL) were 

collected in all months sampled. Sexually mature 

striped anchovy (30–35 mm SL) were also collected in 

all months. 

Figure 5.1

Mean densities (numbers 1000 m–2

) of striped anchovy (Anchoa hepsetus) collected by 

surface trawl in Florida Bay basins and channels during 1984–1985. Florida Bay basins 

are divided into three strata. For designations of strata see Fig. 2.1.
37

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

12.0

14.0

16.0

A

Stratum

East Central West Channels

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

B

Figure 5.2

Mean densities (numbers 1000 m–2

) of striped anchovy 

(Anchoa hepsetus) collected by surface trawl in Florida 

Bay basins and channels during 1984–1985 by (A) 

month and (B) stratum. Florida Bay basins are divided 

into three strata. For designations of strata see Fig. 2.1.
38

Figure 5.3

Mean densities (numbers 1000 m–2

) of striped anchovy (Anchoa hepsetus) from 

bi-monthly otter trawl collections in Florida Bay basins. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 1999–2001. For designations of 

strata and subdivisions see Fig. 2.1.
39

Year

5891- 48915991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

0.6

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

1.4

1.6

A

B

C

_________ ____________________

 Stratum Subdivision

Figure 5.4

Mean densities (numbers 1000 m–2

) of striped anchovy 

(Anchoa hepsetus) from bi-monthly otter trawl collections 

in Florida Bay basins by (A) year, (B) month, and (C) 

stratum/subdivision. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–

2001. For designations of strata and subdivisions see 

Fig. 2.1.
40

January

y

c

n

e

u

q

e

r

F

0

30

60

90

120

March

0

30

60

90

120

May

0

30

60

90

120

June

0

30

60

90

120

July

0

30

60

90

120

September

0

30

60

90

120

November

Size Class (mm)

01<

91- 5192- 5293- 5394- 54

95- 55

96- 56

97- 57

98- 58

99- 59

901- 501

0

30

60

90

120

Figure 5.5

Monthly length-frequency distributions of striped anchovy (Anchoa hepsetus) from Florida 

Bay.
41

Figure 5.6

Mean densities (numbers 1000 m–2

) of bay anchovy (Anchoa mitchilli) collected by 

surface trawl in Florida Bay basins and channels during 1984–1985.Florida Bay basins 

are divided into three strata. For designations of strata see Fig. 2.1.

Anchoa mitchilli (bay anchovy)

Total numbers collected and rank of bay anchovy by sampling 

method. There were 39 species collected by otter trawl; 14 by 

surface trawl in each habitat.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 9284 3

Surface trawl Basins 1984–1985 181 8

Surface trawl Channels 1984–1985 409 3

Life history The bay anchovy ranges from the Gulf of 

Maine to Florida, throughout the Gulf of Mexico, and 

along the Caribbean coast to Belize. It is rare in the 

Yucatan, Gulf of Maine, and Florida Keys, and absent 

from the West Indies (Robins and Ray, 1986; Leak and 

Houde, 1987; Pattillo et al., 1997). This species most 

likely constitutes the greatest biomass of ichthyofauna 

in the southeast and U. S. Gulf of Mexico (Pattillo et 

al., 1997). In adjacent Biscayne Bay, bay anchovy eggs 

occurred throughout the year, but were most abundant 

in summer, and rare to absent during winter (Jones et 

al., 1978; Houde and Lovdal, 1984). Whereas, in Tampa 

Bay, Florida, spawning occurs when water temperatures 

reach 20°C and ceases by November (Pattillo et al., 

1997). Eggs, larvae, and juveniles are pelagic, but bay anchovy dominate the lower portion of the water column 

relative to other zooplanktivorous fishes (Allen et al., 

1995). Size at hatching is 1.8–2.7 mm TL; transformation to the juvenile stage begins at 16 mm SL and is complete by 22 mm SL. Males and females mature at lengths 

of approximately 40 mm SL (Zastrow et al., 1991). 

Diet In the Whitewater Bay system, larval and juvenile 

bay anchovy <25 mm SL fed mainly on microzooplankton (copepods, copepodites, and naupli) by size-selective capture of individual organisms (Odum and Heald, 

1972). 

Abundance and distribution Bay anchovy were common in surface trawl collections, and densities differed 

among strata and months (Figs. 5.6 and 5.7). Highest 

densities were observed in May, the lowest in January 

and July. Densities were highest in the West stratum. 

Unlike the striped anchovy, they appeared to prefer 

channel habitats to basins. 

Bay anchovy were abundant in otter trawl collections. 

Considerable interannual variability was observed in 

the abundance of bay anchovy (Figs. 5.8 and 5.9). 

Relatively low densities were observed in 1984–1985 and 

2000–2001, and to a lesser degree in 1998. Relatively 

high densities were observed in 1994–1995 and 1996. 

Significant spatial and temporal differences in densities 

were observed (Fig. 5.9). Highest densities were observed in November and lowest in July. From 1984–1998, 

bay anchovy were collected at highest densities in the 

Central stratum with lower values in the East and West 
42

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

12.0

14.0

A

Stratum

East Central West Channels

ytisne D

0.0

2.0

4.0

6.0

8.0 B

Figure 5.7

Mean densities (numbers 1000 m–2

) of bay anchovy 

(Anchoa mitchilli) collected by surface trawl in Florida 

Bay basins and channels during 1984–1985 by (A) 

month and (B) stratum. Florida Bay basins are divided 

into three strata. For designations of strata see Fig. 2.1.

strata; from 1999–2001, highest densities occurred in 

the Northern, Central, and Gulf subdivisions. They were 

not collected in the Atlantic subdivision. 

Because this species occurs in the lower water column (Allen et al., 1995), otter trawl densities probably 

are good estimators of its distribution and abundance. 

Highest values we observed followed seagrass dieoff, persistent turbidity, and phytoplankton blooms 

(Fourqurean and Robblee, 1999), which might be due 

to their attraction to unvegetated bottoms and high 

turbidity (Pattillo et al., 1997).

Contrary to our findings, Sogard et al. (1987) reported significant differences in densities of bay anchovy 

among sites, with highest densities occurring at Eagle 

Key (Fig. 1.1).

Length-frequency distributions Based on length-frequency distributions, spawning occurred throughout 

the year in Florida Bay (Fig. 5.10). Otter trawl collections in Florida Bay were dominated by transforming 

juveniles (16–22 mm SL) and juveniles (22–40 mm 

SL). Growth of bay anchovy was difficult to discern 

from length-frequency distributions because of their 

protracted spawning period. In Biscayne Bay, bay anchovy (<14 mm SL) grow approximately 0.50 mm per 

day (Leak and Houde, 1987). Anchovies collected near 

Beaufort, North Carolina, grew exponentially (approximately 4% per day) during their first 49 days (Fives et 

al., 1986). Fish spawned in spring could grow to maturity 

and spawn during their first summer.
43

Figure 5.8

Mean densities (numbers 1000 m–2

) of bay anchovy (Anchoa mitchilli) from bi-monthly otter 

trawl collections in Florida Bay basins. Florida Bay is divided into three strata from 1984–1998 

and six subdivisions from 1999–2001. For designations of strata and subdivisions see Fig. 2.1.
44

Year

5891- 4891

5991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

10.0

20.0

30.0

40.0

50.0

60.0

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

10.0

20.0

30.0

40.0

50.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

10.0

20.0

30.0

40.0

50.0

A

B

C

Figure 5.9

Mean densities (numbers 1000 m–2

) of bay anchovy 

(Anchoa mitchilli) from bi-monthly otter trawl collections 

in Florida Bay basins by (A) year, (B) month, and (C) 

stratum/subdivision. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–

2001. For designations of strata and subdivisions see 

Fig. 2.1.
45

Figure 5.10

Monthly length-frequency distributions of bay anchovy (Anchoa mitchilli) from Florida Bay. 

Values are shown for numbers <100 for clarity.

January

y

c

n

e

u

q

er F

0

1000

2000

3000

4000

March

0

1000

2000

3000

4000

May

0

1000

2000

3000

4000

June

0

1000

2000

3000

4000

July

0

1000

2000

3000

4000

September

0

1000

2000

3000

4000

November

Size Class (mm)

01<

91- 51

92- 5293- 53

94- 54

95- 55

96- 56

0

1000

2000

3000

4000

(12) (36) (7)

(79) (37) (2)

(39) (83) (31) (8) (2)

(32) (8)

(20) (59) (20) (3) (2)

(8) (73) (4)

(31) (33) (8) (2)
46

Chapter 6. Family Ariidae

Arius felis (hardhead catfish)

Total number collected and rank of hardhead catfish by sampling method. There were 39 species collected.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 120 26

Life history The hardhead catfish ranges from Cape 

Cod, Massachusetts, to Florida and throughout the Gulf 

of Mexico to the Yucatan Peninsula, but is not common 

north of Virginia. It is generally found in turbid waters 

over muddy bottoms in brackish and marine environments, occasionally entering fresh water (Breder, 1948; 

Taylor and Menezes, 1978). It is a ubiquitous species in 

shallow brackish and marine waters of the northern Gulf 

of Mexico, and juveniles are absent from seagrass beds. 

In the Gulf of Mexico, breeding occurs from May to 

September in shallow waters of secondary and primary 

bays, and inlets (Pattillo et al., 1997). Eggs are brooded 

in the mouths of males. Size at hatching is 29–45 mm 

TL, and juveniles are released from the males at 31–62 

mm TL (Pattillo et al., 1997). 

Diet Juvenile hardhead catfish are nocturnal bottom 

feeders with copepods being the most important food 

source for juveniles <100 mm TL; for individuals >100 

and <200 mm TL, benthic macro-invertebrates are important food sources (Darnell, 1958). 

Abundance and distribution Hardhead catfish were 

common in otter trawl collections. Significant differences in hardhead catfish densities were observed spatially, and among years, but not months. Densities were 

highest in western Florida Bay (Figs. 6.1 and 6.2). They 

were absent in the Atlantic and Eastern subdivision and 

rarely collected in the East stratum and Central and 

Northern subdivisions. There was a decline in densities from 1984–1985, increases in 1996 and 1997–1998, 

then decreases to lowest densities in 1999–2000 and 

2000–2001 (Fig. 6.2).

Our findings in western Florida are in concordance 

with those of Sogard et al. (1989a), while the decline 

in densities from 1984–1985 mirrored the die-off of 

seagrasses (Fourqurean and Robblee, 1999).

Length-frequency distributions Spawning occurred 

in summer and fall, based on the presence of specimens 

<50 mm (Fig. 6.3). Although spawning was contracted, 

we could not discern any patterns of growth by following modes, nor could we identify size classes. There 

was a wide range of lengths from July through January. 

This species attains a maximum length of 600 mm TL 

(Robins and Ray, 1986).
47

Figure 6.1

Mean densities (numbers 1000 m–2

) of hardhead catfish (Arius felis) from bi-monthly 

otter trawl collections in Florida Bay basins. Florida Bay is divided into three strata 

from 1984–1998 and six subdivisions from 1999–2001. For designations of strata and 

subdivisions see Fig. 2.1.
48

Year

5891- 48915991- 49916991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

0.1

0.2

0.3

0.4

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

0.2

0.4

0.6

0.8

A

B

C

__________

 Stratum

______________________

 Subdivision

Figure 6.2

Mean densities (numbers 1000 m–2

) of hardhead catfish 

(Arius felis) from bi-monthly otter trawl collections in 

Florida Bay basins by (A) year, (B) month, and (C) 

stratum/subdivision. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–

2001. For designations of strata and subdivisions see 

Fig. 2.1.
49

Figure 6.3

Monthly length-frequency distributions of hardhead catfish (Arius felis) from Florida Bay.

January

y

c

n

e

u

q

e

r

F

0

10

20

30

March

0

10

20

30

May

0

10

20

30

June

0

10

20

30

July

0

10

20

30

September

0

10

20

30

November

Size Class (mm)

94- 0496- 06

98- 08

901- 001921- 021

941- 041961- 061981- 081

902- 002922- 022

942- 042

962- 062

982- 082

903- 003

923- 023

943- 043

0

10

20

30
50

tidewater silversides are most frequently eaten, but gobies, killifishes, silver perch, pipefish, spotted seatrout, 

and puffers are also consumed.

Abundance and distribution Inshore lizardfish were 

common in otter trawl collections. There were no significant differences in densities of inshore lizardfish, 

between strata, but there were between subdivisions. 

They were more common in the Western, Central, and 

Atlantic subdivisions, and rarely collected in the Northern and Eastern subdivisions; the latter where salinities 

are relatively lower (Figs. 3.1, 7.1, and 7.2). Significant 

differences in densities between months and years were 

observed (Fig. 7.2). However, no distinct monthly patterns could be discerned. They were more abundant in 

the spring and late fall and least abundant in early fall. 

This species was more frequently encountered in 1997–

1998 and 1999–2000; it was less frequently encountered 

in 1994–1995 (Fig. 7.1).

The abundance of inshore lizardfish was negatively 

correlated with its reported preferred food. This species 

was more frequently encountered when its preferred 

food source, bay anchovy (Carr and Adams, 1973), was 

rarely encountered and less frequently encountered 

when its food source was most abundant (Figs. 5.9 and 

7.1). 

Length-frequency distributions Inshore lizardfish 

recruit into Florida Bay throughout the year (Fig. 7.3). 

A wide size range of juvenile sizes was collected. This 

species attains a maximum length of 450 mm, but we 

rarely collected specimens > ca. 230 mm SL, which 

could be either due to gear avoidance or the species 

leaving Florida Bay at this size.

Chapter 7. Family Synodontidae

Synodus foetens (inshore lizardfish)

Total number collected and rank of inshore lizardfish by sampling method. There were 39 species collected.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 262 18

Life history Inshore lizardfish range from Massachusetts to Brazil including the northern Gulf of Mexico. 

It is rare north of South Carolina, and is not found 

north of Cape Hatteras during winter. It occurs most 

frequently on mud bottoms in shallow waters, but has 

been taken at depths of 200 m (Breder, 1948; Anderson et al., 1966; Robins and Ray, 1986; Able and Fahay, 

1998). In New Jersey estuarine waters, inshore lizardfish 

are most abundant in macroalgae beds over sandy substrates, where they burrow into the substrate, but are 

also commonly collected over silty bottoms (Able and 

Fahay, 1998). Inshore lizardfish spawn in continental 

shelf waters throughout the year, possibly more intense 

in spring (Jones et al., 1978). Eggs, larvae, and prejuveniles are pelagic. Larvae hatch at approximately 2.5 

mm NL, transformation to the prejuvenile stage occurs 

at approximately 16 mm TL, and transformation to the 

juvenile stage occurs at 30–44 mm TL. Juveniles begin 

to settle to the bottom at approximately 30–40 mm TL 

(Jones et al., 1978; Able and Fahay, 1998). Size at sexual 

maturity is not known.

Diet Juvenile inshore lizardfish (41–140 mm SL) feed 

almost exclusively on fish and apparently are extremely 

opportunistic (Carr and Adams, 1973). Anchovies and 
51

Figure 7.1

Mean densities (numbers 1000 m–2

) of inshore lizardfish (Synodus foetens) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
52

Year

5891- 4891

5991- 4991

6991

8991- 79910002- 9991

1002- 0002

ytisne D

0.0

0.2

0.4

0.6

0.8

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.2

0.4

0.6

0.8

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

1.0

2.0

3.0

4.0

A

B

C

Figure 7.2

Mean densities (numbers 1000 m–2

) of inshore lizardfish 

(Synodus foetens) from bi-monthly otter trawl collections 

in Florida Bay basins by (A) year, (B) month, and (C) 

stratum/subdivision. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–

2001. For designations of strata and subdivisions see 

Fig. 2.1.
53

Figure 7.3

Monthly length-frequency distributions of inshore lizardfish (Synodus foetens) from Florida 

Bay.

January

y

c

n

e

u

q

e

r

F

0

10

20

30

40

March

0

10

20

30

40

May

0

10

20

30

40

June

0

10

20

30

40

July

0

10

20

30

40

September

0

10

20

30

40

November

Size Class (mm)

92- 02

94- 0496- 0698- 08901- 001

921- 021941- 041

961- 061

981- 081

902- 002

922- 022

942- 042962- 062982- 082903- 003

923- 023

943- 043

963- 063

0

10

20

30

40
54

est densities were observed in May and July, lowest in 

September and January.

Gulf toadfish appear to have an affinity for the seagrass Thalassia testudinum, which is more dense in western and central Florida Bay (Fig. 3.6). Sheridan et al. 

(1997) reported that highest densities of gulf toadfish 

in Florida Bay were consistently higher in T. testudinum

grass beds than in either Halodule wrightii grass beds, 

macroalgae, or mud patches, while Serafy et al. (1997a) 

concludes that shallow Thalassia beds, where high nocturnal dissolved oxygen levels occur, might be important 

habitat features for this species. However, in a study in 

the Bay after the die-off of seagrasses in the late 1980s, 

declines in standing crops of seagrass were correlated 

with increases in the abundance of gulf toadfish (Matheson et al., 1999). Contrary to these latter findings, we 

observed significantly higher densities in 1984–1985 

(pre-seagrass die-off) than 1994–1995 (post-seagrass 

die-off) in basin habitats.

Serafy et al. (1997a) observed seasonal differences in 

densities of gulf toadfish at some sites in adjacent Biscayne Bay. These differences were explained by spawning behavior; i.e., concentration of spawning adults in 

shallow T. testudinum habitat during winter. Based on 

their speculation, our lowest concentrations in January 

could be indicative of movement from deeper basins to 

shallow banks, the latter area not sampled by us. Our 

highest concentrations in May and July are more difficult to explain.

Length-frequency distributions The distribution 

of small juveniles indicated that spawning occurred 

throughout the year in Florida Bay (Fig. 8.3). This is 

contrary to the spawning season reported by Breder 

(1941) for western Florida (February–March). Juveniles 

ranged from 20 mm SL to 190 mm SL, which was considerably different than gulf toadfish in Biscayne Bay 

(28–268 mm TL) (Serafy et al., 1997a). This difference 

could be attributed to differences in gear and mesh size. 

Chapter 8. Family Batrachoididae

Opsanus beta (gulf toadfish)

Total number collected and rank of gulf toadfish by sampling 

method. There were 39 species collected.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 649 13

Life history Gulf toadfish range from Cape Canaveral, 

Florida, Little Bahama Bank, and throughout the Gulf 

of Mexico to Campeche (Robins and Ray, 1986). In 

general, it is common in seagrass beds and rocky cuts in 

estuaries, and in shallow coastal waters (Robins and Ray, 

1986). Gulf toadfish spawn in western Florida from February to March at temperatures 15–22°C (Breder, 1941). 

Eggs are demersal and attached in empty shells and 

sponge cavities. Eggs are guarded by the male. Although 

little information is available for gulf toadfish, life histories are probably similar to its temperate congener, O. 

tau (oyster toadfish). Oyster toadfish have no definable 

larvae as the juvenile stage is attained at yolk absorption. 

Juveniles become free swimming at 16–18 mm TL. Gulf 

toadfish are sexually mature at approximately 95 mm SL 

and attain a length of approximately 65 mm SL after a 

year of growth (Breder, 1941).

Diet We are unaware of any information on diet.

Abundance and distribution Gulf toadfish were common in otter trawl collections in basins. There were significant differences in densities of gulf toadfish between 

strata and subdivisions (Figs. 8.1 and 8.2). Densities 

were highest in Central and West strata and Central 

and Western subdivisions. Considerable densities were 

also observed in the Eastern subdivision (northern and 

eastern stations). We observed significant intra-annual 

differences in densities of gulf toadfish (Fig. 8.2). High-
55

Figure 8.1

Mean densities (numbers 1000 m–2

) of gulf toadfish (Opsanus beta) from bi-monthly 

otter trawl collections in Florida Bay basins. Florida Bay is divided into three strata 

from 1984–1998 and six subdivisions from 1999–2001. For designations of strata and 

subdivisions see Fig. 2.1.
56

Figure 8.2

Mean densities (numbers 1000 m–2

) of gulf toadfish 

(Opsanus beta) from bi-monthly otter trawl collections 

in Florida Bay basins by (A) year, (B) month, and (C) 

stratum/subdivision. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–

2001. For designations of strata and subdivisions see 

Fig. 2.1.

Year

5891- 48915991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

1.4

1.6

1.8

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

1.4

1.6

1.8

2.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

3.0

A

B

C
57

Figure 8.3

Monthly length-frequency distributions of gulf toadfish (Opsanus beta) from Florida Bay. 

Values are shown for numbers <5 for clarity.

January

y

c

n

e

u

q

e

r

F

0

50

100

150

200

March

0

50

100

150

200

May

0

50

100

150

200

June

0

50

100

150

200

July

0

50

100

150

200

September

0

50

100

150

200

November

Size Class (mm)

02<

93- 03

95- 0597- 0799- 09

911- 011

931- 031951- 051

971- 071

991- 091

912- 012932- 032

0

50

100

150

200

(4) (4) (4) (2) (2) (2) (1)

(1) (1) (1) (1) (1)

(4) ( (2) 2)

(4) (1) (1) (2)

(3) (4) (1) (1) (1) (2) (1) (1) (1)

(3) (3) (2) (4) (3)

(3) (2) (3) (2) (1)
58

Chapter 9. Family Hemiramphidae

Chriodorus atherinoides (hardhead halfbeak)

Total numbers collected and rank of hardhead halfbeak by 

sampling method. There were a total of 14 species collected 

by surface trawl in each habitat.

Gear Habitat Year Number Rank

Surface trawl Basins 1984–1985 561 3

Surface trawl Channels 1984–1985 150 5

Life history Hardhead halfbeaks range from the Bahamas, southeast Florida, and northern Gulf of Mexico 

to Cuba and Yucatan. They are a pelagic, schooling fish 

that inhabits bays and waterways and is typically associated with seagrass beds (Böhkle and Chaplin, 1968; 

Robins and Ray, 1986). Very little is known about life 

history characteristics of this species. Breder (1934) 

collected hardhead halfbeaks ranging 77–105 mm with 

ripening ovaries in January. 

Diet Breder (1934) noted that this species feeds in the 

alga Batophora, which is populated by diatoms and other 

small organisms.

Abundance and distribution Hardhead halfbeaks 

were common in surface trawl collections in channels 

and basins in 1984–1985, and exhibited significant temporal and spatial differences. They were more abundant 

in the East and Central strata and in channels (Figs. 

9.1 and 9.2). Hardhead halfbeak were most abundant 

in June and July, and less abundant from September 

through March (Fig. 9.2). Hardhead halfbeak are 

considered to be the numerically dominant species in 

eastern Florida Bay (Schmidt, 1979). 

Length-frequency distributions We observed two size 

classes of hardhead halfbeaks in Florida Bay that could 

represent juveniles and adults (Fig. 9.3). Although there 

is limited data from September through March, this species could spawn in all seasons. Based on May and June 

length-frequency distributions, fish were estimated to 

grow approximately 80 mm SL per year. 

Figure 9.1

Mean densities (numbers 1000 m–2

) of hardhead halfbeak (Chriodorus atherinoides) 

collected by surface trawl in Florida Bay basins and channels during 1984–1985. Florida 

Bay basins are divided into three strata. For designations of strata see Fig. 2.1.
59

Figure 9.2

Mean densities (numbers 1000 m–2

) of hardhead 

halfbeak (Chriodorus atherinoides) collected by surface 

trawl in Florida Bay basins and channels during 1984–

1985 by (A) month and (B) stratum. Florida Bay basins 

are divided into three strata. For designations of strata 

see Fig. 2.1.

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

12.0

A

Stratum

East Central West Channels

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

6.0 B
60

January

y

c

n

e

u

q

e

r

F

0

25

50

75

March

0

25

50

75

May

0

25

50

75

June

0

25

50

75

July

0

25

50

75

September

0

25

50

75

November

Size Class (mm)

92- 0294- 0496- 06

98- 08

901- 001921- 021941- 041

961- 061

981- 081

0

25

50

75

Figure 9.3

Monthly length-frequency distributions of hardhead halfbeak (Chriodorus atherinoides) from 

Florida Bay.
61

Figure 9.4

Mean densities (numbers 1000 m–2

) of ballyhoo (Hemiramphus brasiliensis) collected by 

surface trawl in Florida Bay basins and channels during 1984–1985. Florida Bay basins 

are divided into three strata. For designations of strata see Fig. 2.1.

Hemiramphus brasiliensis (ballyhoo) 

Total numbers collected and rank of ballyhoo by sampling 

method. There were a total of 14 species collected by surface 

trawl in each habitat.

Gear Habitat Year Number Rank

Surface trawl Basins 1984–1985 62 12

Surface trawl Channels 1984–1985 40 10

Life history The ballyhoo occurs on both sides of the 

Atlantic Ocean. In the western Atlantic it ranges from 

Woods Hole, Massachusetts to Rio de Janeiro, Brazil, 

and throughout the Gulf of Mexico, Caribbean Sea, and 

the Bahamas. It is a straggler north of Florida (Breder, 

1948; Collette, 1978a; Hardy, 1978). They are inshore, 

pelagic fish occurring in large schools (Collette, 1978a) 

and are an important bait fishery in Florida Bay. Florida 

Bay has been identified as a spawning area for ballyhoo 

(McBride et al., 2003). Spawning occurs from March 

through August in Florida waters and in Florida Bay in 

association with mud banks and basins (Berkeley and 

Houde, 1978; McBride et al., 2003). Eggs are demersal and have long filaments attached to the chorion 

(Berkeley and Houde, 1978). Larvae are 5–7 mm SL at 

hatching and are well developed (Berkeley and Houde, 

1978). The smallest juvenile that has been described was 

35 mm (Hardy, 1978). Females attain sexual maturity in 

their first year (Berkeley and Houde, 1978). Ballyhoo 

attain a maximum size of 400 mm (Robins and Ray, 

1986).

Diet Ballyhoo (<180 mm FL) consume mainly decapods and copepods, and with increasing growth there 

is a change in food habits toward increases in seagrasses 

(Berkeley and Houde, 1978).

Abundance and distribution Ballyhoo were commonly collected by surface trawl in basins and channels, and never collected by otter trawl. Densities were 

different between strata, but not months (Figs. 9.4 and 

9.5). Based on data in 1984–1985, ballyhoo occurred in 

all strata, but were more abundant in the West stratum. 

This species is rarely collected on Florida Bay mud 

banks (Sogard et al., 1989a).

Length-frequency distributions Ballyhoo spawn in 

Florida Bay during late spring and early summer (Fig. 

9.6). Because of limited data, we were unable to discern 

age or size classes, or estimate growth.
62

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

1.4 A

Stratum

East Central West Channels

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0 B

Figure 9.5

Mean densities (numbers 1000 m–2

) of ballyhoo 

(Hemiramphus brasiliensis) collected by surface trawl 

in Florida Bay basins and channels during 1984–1985 

by (A) month and (B) stratum. Florida Bay basins are 

divided into three strata. For designations of strata see 

Fig. 2.1.
63

January

y

c

n

e

u

q

e

r

F

0

5

10

15

March

0

5

10

15

May

0

5

10

15

June

0

5

10

15

July

0

5

10

15

September

0

5

10

15

November

Size Class (mm)

93- 0395- 05

97- 0799- 09

911- 011

931- 031

951- 051

971- 071

991- 091

912- 012

932- 032952- 052

972- 072992- 092

0

5

10

15

Figure 9.6

Monthly length-frequency distributions of ballyhoo (Hemiramphus brasiliensis) from Florida 

Bay.
64

at sexual maturity (female) is approximately 150 mm 

(Hardy, 1978).

Diet We are unaware of any information on diet.

Abundance and distribution Halfbeak were abundant 

in surface trawl collections in 1984–1985 and common 

in otter trawl collections. It was the most abundant 

pelagic species collected by surface trawl in basins in 

1984–1985. We observed significant differences in 

halfbeak densities between strata and months for the 

surface trawl data (Figs. 9.7 and 9.8). Highest densities 

were observed in the West stratum, lowest in the Central 

stratum. Temporally, we observed highest densities in 

September (Fig. 9.8). 

Although we collected 119 halfbeak in otter trawl collections in basins (Fig. 9.9 and 9.10), they were mainly 

collected at one station in 1984–1985. There were no 

significant spatial or temporal differences, which is to be 

expected given the high number of zero catches.

Length-frequency distributions Spawning and/or 

recruitment occurred in spring and summer (Fig. 9.11). 

Sexually mature fish (approximately 150 mm SL) were 

present during all months. Multiple cohorts, and possibly two age groups, were present from at least May 

through November.

Figure 9.7

Mean densities (numbers 1000 m–2

) of halfbeak (Hyporhamphus unifasciatus) collected 

by surface trawl in Florida Bay basins and channels during 1984–1985. Florida Bay 

basins are divided into three strata. For designations of strata see Fig. 2.1.

Hyporhamphus unifasciatus (halfbeak)

Total numbers collected and rank of halfbeak by sampling 

method. There were a total of 39 species collected by otter 

trawl; 14 by surface trawl in each habitat.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 119 27

Surface trawl Basins 1984–1985 1249 1

Surface trawl Channels 1984–1985 276 4

Life history Halfbeak occur in Bermuda, peninsular 

Florida, the West Indies, and from Campeche Bay 

southward to Uruguay. It is a pelagic, inshore schooling species that frequently enters estuaries (Collette, 

1978a). Little information on its life history is known. 

Hardy (1978) summarized life history information 

on the halfbeak, but because of taxonomic confusion 

between H. meeki and H. unifasciatus (Banford and Collette, 1993), Hardy’s summary pertains mainly to H. 

meeki. Halfbeak spawn in March in the West Indies. Eggs 

have filaments attached to the chorion and have been 

found on Thalassia in shallow water in Puerto Rico. Size 
65

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

5.0

10.0

15.0

20.0

25.0

30.0

35.0

40.0

A

Stratum

East Central West Channels

ytisne D

0.0

5.0

10.0

15.0

20.0

25.0

30.0

B

Figure 9.8

Mean densities (numbers 1000 m–2

) of halfbeak 

(Hyporhamphus unifasciatus) collected by surface trawl 

in Florida Bay basins and channels during 1984–1985 

by (A) month and (B) stratum. Florida Bay basins are 

divided into three strata. For designations of strata see 

Fig. 2.1.
66

Figure 9.9

Mean densities (numbers 1000 m–2

) of halfbeak (Hyporhamphus unifasciatus) from bimonthly otter trawl collections in Florida Bay basins. Specimens were only collected in 

1984–1985 and 1997–1998.Florida Bay is divided into three strata in 1984–1985. For 

designations of strata see Fig. 2.1.
67

Figure 9.10

Mean densities (numbers 1000 m–2

) of halfbeak (Hyporhamphus unifasciatus) from bi-monthly otter trawl collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum. Specimens were only collected in 

1984–1985. Florida Bay is divided into three strata in 

1984–1985. For designations of strata see Fig. 2.1.

Year

5891- 4891

5991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

0.4

0.8

1.2

1.6

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.4

0.8

1.2

1.6

 Stratum 

tsa E

l art ne C

tse W

ytisne D

0.0

0.4

0.8

1.2

A

B

C
68

Figure 9.11

Monthly length-frequency distributions of halfbeak (Hyporhamphus unifasciatus) from Florida 

Bay.

January

y

c

n

e

u

q

e

r

F

0

40

80

120

160

March

0

40

80

120

160

May

0

40

80

120

160

June

0

40

80

120

160

July

0

40

80

120

160

September

0

40

80

120

160

November

Size Class (mm)

02<93- 03

95- 05

97- 0799- 09

911- 011931- 031951- 051

971- 071991- 091912- 012932- 032

0

40

80

120

160
69

Abundance and distribution Redfin needlefish were 

commonly collected by surface trawl particularly in basins in 1984–1985. They were collected at many stations 

in all strata, but densities were significantly higher in the 

Central stratum (Figs. 10.1 and 10.2). The distribution 

of densities among months was clearly uni-modal with 

significantly highest densities occurring in July (Fig. 

10.2), but they occurred at a greater percentage of stations in September and May (Fig. 10.1). 

Redfin needlefish were uncommon in otter trawl collections (Fig.10.3). We observed significant differences 

in densities between strata, subdivisions, and months, 

but not years (Fig. 10.4). Contrary to the surface trawl 

data, densities were highest in the East stratum. High 

densities were observed in the Northern and Eastern 

subdivisions, and in July. 

Sogard et al. (1989a) reported gillnet catches of redfin needlefish at three of their four sites in Florida Bay. 

They were Cowpen Keys, Buchanan Key, and Oyster 

Keys. Lower catch-per-unit-effort was observed at Eagle 

Key (Fig. 1.1).

Length-frequency distributions Recruitment of redfin needlefish was protracted, and spawning most 

likely occurs in all seasons, although minimally in winter 

(Fig. 10.5).

Figure 10.1

Mean densities (numbers 1000 m–2

) of redfin needlefish (Strongylura notata) collected 

by surface trawl in Florida Bay basins and channels during 1984–1985. Florida Bay 

basins are divided into three strata. For designations of strata see Fig. 2.1.

Chapter 10. Family Belonidae

Strongylura notata (redfin needlefish)

Total numbers collected and rank of redfin needlefish by 

sampling method. There were a total of 39 species collected 

by otter trawl; 14 by surface trawl in each habitat.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 77 33

Surface trawl Basins 1984–1985 595 2

Surface trawl Channels 1984–1985 111 6

Life history Redfin needlefish range from Bermuda, 

Florida, Bahamas to the Lesser Antilles, and to Central 

America. Its preferred habitat is coastal waters, mainly 

bays and inlets (Robins and Ray, 1986). Very little information on the life history of this species is available. 

Redfin needlefish spawn in red mangrove (Rhizophora 

mangel) aerial root tangles. Eggs are assumed to be demersal and have external adhesive filaments attached to 

the chorion. Size at hatching is approximately 12 mm 

SL (Breder, 1959). 

Diet Reid (1954) reported that individuals ranging 

from 75–125 mm contained various species of shrimp 

in their diets. 
70

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

2.0

4.0

6.0

8.0

A

Stratum

East Central West Channels

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

B

Figure 10.2

Mean densities (numbers 1000 m–2

) of redfin needlefish 

(Strongylura notata) collected by surface trawl in Florida 

Bay basins and channels during 1984–1985 by (A) 

month and (B) stratum. Florida Bay basins are divided 

into three strata. For designations of strata see Fig. 2.1.
71

Figure 10.3

Mean densities (numbers 1000 m–2

) of redfin needlefish (Strongylura notata) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
72

Year

5891- 48915991- 4991

6991

8991- 7991

0002- 99911002- 0002

ytisne D

0.0

0.1

0.2

0.3

0.4

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

A

B

C

Figure 10.4

Mean densities (numbers 1000 m–2

) of redfin 

needlefish (Strongylura notata) from bi-monthly otter 

trawl collections in Florida Bay basins by (A) year, 

(B) month, and (C) stratum/subdivision. Florida Bay 

is divided into three strata from 1984–1998 and six 

subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
73

January

y

c

n

e

u

q

e

r

F

0

10

20

30

40

March

0

10

20

30

40

May

0

10

20

30

40

June

0

10

20

30

40

July

0

10

20

30

40

September

0

10

20

30

40

November

Size Class (mm)

02<

93- 03

95- 0597- 0799- 09

911- 011

931- 031

951- 051

971- 071991- 091

912- 012932- 032952- 052

972- 072

992- 092

913- 013

933- 033

953- 053

973- 073

993- 093

0

10

20

30

40

Figure 10.5

Monthly length-frequency distributions of redfin needlefish (Strongylura notata) from Florida 

Bay.
74

S. timucu (timucu)

Total numbers collected and rank of timucu by sampling 

method. There were a total of 14 species collected by surface 

trawl in each habitat.

Gear Habitat Year Number Rank

Surface trawl Basins 1984–1985 112 11

Surface trawl Channels 1984–1985 15 12

Life history Timucu range from Jupiter Inlet on the 

east coast of Florida to Pensacola on the west coast of 

Florida, the Bahamas, the Greater and Lesser Antilles, 

and from Yucatan, Mexico to Brazil (Collette, 1978b). 

It is an inshore species that frequents most of the same 

habitats as the redfin needlefish. Life history information on this species is lacking. 

Diet Carr and Adams (1973) reported on the diets of 

Atlantic needlefish (S. marina) in a west coast Florida estuary, but were unaware at the time of their analysis that 

timucu was a valid species that occurred sympatrically 

with Atlantic needlefish in the eastern Gulf of Mexico. 

Hence, some of the smaller fish they analyzed could 

have been timucu. They observed a diet that progresses 

from small crustaceans to fish during ontogeny. Small 

juveniles (35–50 mm SL) fed on small crustaceans, especially amphipods, mysids, and very small shrimp. At 

approximately 50 mm SL and greater, they fed almost 

entirely on fish, especially anchovies and atherinids. 

Abundance and distribution Timucu has a more 

restricted distribution in Florida Bay than its congener 

redfin needlefish (Figs. 10.1 and 10.6). Timucu, collected mainly by surface trawl in basins in 1984–1985, 

was rarely collected in the East and Central strata. High 

densities were collected in the West stratum (Figs. 10.6 

and 10.7). Significant differences in densities occurred 

between months. Timucu were most abundant in March 

through May and, like their congener, least abundant in 

January (Fig. 10.7).

Based on the study of timucu occurrence on Florida 

Bay mud banks (Sogard et al., 1989a) and this study, the 

species occurs mainly in western Florida Bay.

Length-frequency distributions No measurements 

were taken in 1984–1985. 

Figure 10.6

Mean densities (numbers 1000 m–2

) of timucu (Strongylura timucu) collected by surface 

trawl in Florida Bay basins and channels during 1984–1985. Florida Bay basins are 

divided into three strata. For designations of strata see Fig. 2.1.
75

Figure 10.7

Mean densities (numbers 1000 m–2

) of timucu 

(Strongylura timucu) collected by surface trawl in Florida 

Bay basins and channels during 1984–1985 by (A) 

month and (B) stratum. Florida Bay basins are divided 

into three strata. For designations of strata see Fig. 2.1.

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

0.5

1.0

1.5

2.0

A

Stratum

East Central West Channels

ytisne D

0.0

0.5

1.0

1.5

2.0

B
76

Chapter 11. Families Fundulidae and Cyprinodontidae

Floridichthys carpio (goldspotted killifish)

Total numbers collected and rank of goldspotted killifish by 

sampling method. There were a total of 39 species collected 

by otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 1652 8

Life history The goldspotted killifish, a cyprinodont, 

ranges from southeastern Florida throughout the Gulf 

of Mexico. It is a small resident forage fish that occupies 

marine habitats (but enters brackish waters) on tidal 

flats and creeks (Robins and Ray, 1986). The ecological 

classification of this species is problematic (Matheson 

et al., 1999). It does not fit the definition of a benthic 

species, but it spends most of its time near the bottom. It 

is associated with seagrass beds, but is most abundant in 

sparse beds (Sogard et al., 1989a). On the other hand, 

Thayer et al. (1987b) found that in Florida Bay and 

adjacent estuarine waters this species prefers the mangrove root habitat rather than adjacent seagrass beds. 

Goldspotted killifish are annuals, with sexual maturity 

attained during the first year of life. They are benthic 

spawners; eggs are laid one at a time and are attached to 

the substrate. Multiple spawning occurs in this species 

(Nordlie, 2000).

Diet The diet of goldspotted killifish in northeastern 

Florida Bay has been related to habitat quality. Ley et 

al. (1994) observed a systematic variation of diet along 

an estuarine gradient. Goldspotted killifish ingested low 

quality food (algae) in areas where salinity was highly 

variable. In areas with less variability in salinity, high 

quality food (benthic invertebrates) was ingested. Ley et 

al. proposed that highly variable amounts of submerged 

aquatic vegetation in low quality food areas reduce 

abundances of benthic invertebrates.

Various studies report differences in diets of goldspotted killifish that can be explained by seasonal and 

ontogenetic differences. Motta et al. (1995) examined 

fish 46–87 mm SL from Tampa Bay, Florida, and found 

a diverse diet consisting of copepods, ostracods, eggs, 

gastropods, and amphipods. In the Whitewater Bay 

system, this species (14–59 mm SL) also consumed an 

extremely diverse group of organisms, with detritus and 

small amphipods being the dominant items (Odum and 

Heald, 1972). 

Abundance and distribution Goldspotted killifish 

were abundant in otter trawl collections. They were collected in higher densities in the Central stratum and 

Northern subdivision, and were not collected in the 

Gulf subdivision (Figs. 11.1 and 11.2).

There was a decrease in abundance of this species 

following environmental changes in the late 1980s 

(Figs. 11.1 and 11.2). Mean annual densities declined 

considerably from 1984–1985 to 1996. They were not 

collected in the Central stratum in 1996. In 1997–1998 

mean yearly densities were similar to those observed in 

1994–1995 (Figs. 11.1 and 11.2). Highest mean annual 

densities were observed in 1999–2001, but this could be 

a result of an increase in sampling effort in the Eastern 

and Northern subdivisions. 

Matheson et al. (1999) also observed a decrease in 

abundance of this species between the 1980s and 1990s 

on mud banks in Florida Bay. During the 1980s, Sogard 

et al. (1987) reported this species abundant at all their 

sampling sites on mud banks in Florida Bay. Similarly, 

Schmidt (1979) collected goldspotted killifish throughout Florida Bay, but it was most abundant in eastern 

Florida Bay. On the Atlantic Ocean side of the Florida 

Keys adjacent to Florida Bay, Springer and McErlean 

(1962) collected this species only in beds of Halodule 

wrightii in soft sediments. They suggested that goldspotted killifish burrow when seagrass is exposed. 

Length-frequency distributions Goldspotted killifish 

spawned throughout the year in Florida Bay (Fig. 11.3). 

The largest specimens were collected in June and averaged 55–59 mm SL. In adjacent waters to Florida Bay on 

the Atlantic Ocean side, Springer and McErlean (1962) 

reported a maximum length of 49 mm SL, which is in 

concordance with our findings. They only collected 

juveniles (ca. 15 mm SL) of this species in June, July, 

and September, whereas we collected juveniles during 

all months in which we sampled (Fig. 11.3). The maximum reported size of this species is 65 mm TL (Robins 

and Ray, 1986). 
77

Figure 11.1

Mean densities (numbers 1000 m–2

) of goldspotted killifish (Floridichthys carpio) from 

bi-monthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
78

Year

5891- 4891

5991- 4991

6991

8991- 79910002- 99911002- 0002

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

5.0

10.0

15.0

20.0

25.0

30.0

35.0

A

B

C

Figure 11.2

Mean densities (numbers 1000 m–2

) of goldspotted 

killifish (Floridichthys carpio) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
79

January

y

c

n

e

u

q

e

r

F

0

200

400

600

Col 2 

March

0

200

400

600

May

0

200

400

600

June

0

200

400

600

July

0

200

400

600

September

0

200

400

600

November

Size Class (mm)

01<

91- 5192- 5293- 5394- 54

95- 55

96- 56

0

200

400

600

(31) (3) (3)

(5) (49) (14) (9) (2)

(7) (31) (13) (11) (5)

(11) (36) (15) (12) (7) (4) (4)

(10) (19) (9) (4) (2) (1)

(8) (24) (3) (4) (1)

(35) (28) (9) (3)

Figure 11.3

Monthly length-frequency distributions of goldspotted killifish (Floridichthys carpio) from 

Florida Bay. Values are shown for numbers <50 for clarity.
80

Lucania parva (rainwater killifish)

Abundance (numbers) and rank of rainwater killifish by 

sampling method. There were a total of 39 species collected 

by otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 11,132 2

Life history The rainwater killifish, a fundulid, occurs 

from Massachusetts and the northern Gulf of Mexico to 

the Florida Keys and northeast Mexico (Robins and Ray, 

1986). It is a euryhaline species that is associated with 

submerged aquatic vegetation (Jordan, 2002).

Rainwater killifish hatch at 4.0 to 5.5 mm TL and 

reach a maximum size at 43 mm TL (Able and Fahay, 

1998). The smallest juvenile described is 20 mm (Hardy, 

1978). This species is sexually mature at approximately 

23 mm SL and at ages of less than 2 months (Matheson 

et al., 1999). 

Diet In the Whitewater Bay system adjacent to Florida 

Bay, fish less than 20 mm SL feed exclusively on planktonic copepods, whereas the diet of fish 21 to 37 mm SL 

from the same area was more diverse. These larger fish 

have a diverse diet, and mainly consume amphipods and 

mysids (Odum and Heald, 1972).

Abundance and distribution Rainwater killifish were 

very abundant in otter trawl collections (Fig. 11.4). 

There were significant differences in densities among 

years, months, strata, and subdivisions. Mean annual 

densities followed a pattern similar to goldspotted 

killifish (Figs. 11.2 and 11.5). There was a decline 

from 1984–1985 to 1996, following the seagrass die-off 

and accompanying environmental changes. Densities 

increased from 1997 to 2000–2002, when densities 

mirrored those of 1984–1985. Rainwater killifish were 

collected at highest mean monthly densities during the 

middle of the wet season; i.e., September (Fig. 11.5).

Spatially, rainwater killifish were collected at highest 

densities in the West strata and Western subdivision, but 

occurred at more stations in the Northern subdivision 

(Figs. 11.4 and 11.5). They were relatively rare in the 

Gulf subdivision and absent in 1984–1985 in the East 

stratum.

Over mud banks of Florida Bay, this species was abundant at Cowpen Keys (Fig. 1.1) (Sogard et al., 1987) 

located in our Atlantic subdivision where we observed 

only moderate densities (Figs. 11.4 and 11.5). Schmidt 

(1979) found this species to be most abundant in eastern Florida Bay.

Length-frequency distributions Based on the presence of small larvae, spawning occurred throughout the 

year (Fig. 11.6). Because of multiple cohorts, a progression of size classes could not be used to estimate growth. 

Able and Fahay (1998) estimated summer growth to be 

0.3 mm per day in the Middle Atlantic Bight, with little 

growth during winter. Length at the end of the first year 

is 30 to 40 mm TL (ca. 20–29 mm SL, based on measurements of Able and Fahay’s rainwater killifish illustration). The reported maximum size of fish in Middle 

Atlantic Bight (43 mm TL; ca. 33 mm SL) is similar to 

the maximum size we observed.
81

Figure 11.4

Mean densities (numbers 1000 m–2

) of rainwater killifish (Lucania parva) from 

bi-monthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
82

Year

5891- 4891

5991- 4991

6991

8991- 7991

0002- 99911002- 0002

ytisne D

0.0

10.0

20.0

30.0

40.0

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

10.0

20.0

30.0

40.0

50.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

40.0

80.0

120.0

160.0

A

B

C

Figure 11.5

Mean densities (numbers 1000 m–2

) of rainwater killifish 

(Lucania parva) from bi-monthly otter trawl collections 

in Florida Bay basins by (A) year, (B) month, and (C) 

stratum/subdivision. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–

2001. For designations of strata and subdivisions see 

Fig. 2.1.
83

Figure 11.6

Monthly length-frequency distributions of rainwater killifish (Lucania parva) from Florida 

Bay. Values are shown for numbers <100 for clarity.

January

y

c

n

e

u

q

e

r

F

0

2500

5000

March

0

2500

5000

May

0

2500

5000

June

0

2500

5000

July

0

2500

5000

September

0

2500

5000

November

Size Class (mm)

01<

91- 51

92- 52

93- 53

0

2500

5000

(11) (48)

(2) (35) (94) (4)

(45) (6)

(41)

(14) (24) (3)

(9) (75) (5)

(8) (3)
84

Chapter 12. Families Atherinidae and Atherinopsidae

Atherinomorus stipes (hardhead silverside)

Total numbers collected and rank of hardhead silversides by 

sampling method. There were a total of 14 species collected by 

surface trawl in each habitat.

Gear Habitat Year Number Rank

Surface trawl Basins 1984–1985 457 5

Surface trawl Channels 1984–1985 475 2

Life history The hardhead silverside, an atherinid and 

a coastal pelagic species, ranges from south Florida, 

the Bahamas, and the Yucatan to Brazil (Robins and 

Ray, 1986). Despite its abundance within its range, very 

little life history information is available. It spawns during June in south Florida. Like other atherinids, the 

eggs probably have filaments attached that entangle on 

submerged aquatic vegetation or other objects (Breder 

and Rosen, 1966).

Diet Hardhead silversides are zooplanktivores that 

primarily feed on shrimp larvae and copepods (Randall, 

1967).

Abundance and distribution Hardhead silversides 

were commonly collected with a surface trawl in 

basins and channels in 1984–1985. They exhibited 

higher densities in the Central stratum and least 

in the East stratum (Figs.12.1 and 12.2). There 

were no significant differences in densities among 

months. 

This species is strongly associated with mangrove prop 

root habitats in Whitewater Bay, Coot Bay, and Florida 

Bay (Thayer et al., 1987b), and mesohaline waters in 

northern Florida Bay and adjacent estuarine waters 

(Powell et al., 2002).

Length-frequency distributions Small individuals 

(15–19 mm SL), indicative of recent spawning, were 

collected in all seasons, but were absent in January, 

March, and July (Fig. 12.3). However, the occurrence 

of fish 20–24 mm SL indicated spawning occurred 

throughout most of the year. Hardhead silversides attain a maximum length of approximately 125 mm SL 

(Breder, 1948), a size much larger than we collected. 

This species could leave basin and channel habitats and 

recruit into other habitats not sampled by us at approximately 60–64 mm SL. 

Figure 12.1

Mean densities (numbers 1000 m–2

) of hardhead silversides (Atherinomorus stipes) 

collected by surface trawl in Florida Bay basins and channels during 1984–1985. Florida 

Bay basins are divided into three strata. For designations of strata see Fig. 2.1.
85

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

12.0

14.0

16.0

A

Stratum

East Central West Channels

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

B

Figure 12.2

Mean densities (numbers 1000 m–2

) of hardhead 

silversides (Atherinomorus stipes) collected by surface 

trawl in Florida Bay basins and channels during 1984–

1985 by (A) month and (B) stratum. Florida Bay basins 

are divided into three strata. For designations of strata 

see Fig. 2.1.
86

January

y

c

n

e

u

q

er F

0

250

500

750

1000

March

0

250

500

750

1000

May

0

250

500

750

1000

June

0

250

500

750

1000

July

0

250

500

750

1000

September

0

250

500

750

1000

November

Size Class (mm)

91- 51

92- 5293- 5394- 54

95- 55

96- 56

0

250

500

750

1000

Figure 12.3

Monthly length-frequency distributions of hardhead silversides (Atherinomorus stipes) from 

Florida Bay.
87

Figure 12.4

Mean densities (numbers 1000 m–2

) of reef silversides (Hypoatherina harringtonensis) 

collected by surface trawl in Florida Bay basins and channels during 1984–1985. Florida 

Bay basins are divided into three strata. For designations of strata see Fig. 2.1.

Hypoatherina harringtonensis (reef silverside)

Total numbers collected and rank of reef silverside by sampling 

method. There were a total of 14 species collected by surface 

trawl in each habitat.

Gear Habitat Year Number Rank

Surface trawl Basins 1984–1985 366 7

Surface trawl Channels 1984–1985 844 1

Life history The reef silverside, an atherinid and a 

coastal and offshore pelagic zooplanktivore, ranges 

from Bermuda, south Florida, the Bahamas, and the 

Yucatan to northern South America (Robins and Ray, 

1986). Very little life history information is available for 

this species. Like other atherinids, the eggs probably 

have filaments attached that entangle on submerged 

aquatic vegetation or other objects (Breder and Rosen, 

1966). 

Diet Reef silverside feeds predominantly on copepods 

(Randall, 1967).

Abundance and distribution Reef silverside were 

abundant in surface trawl collections from basins and 

channels. They were collected in significantly higher 

densities in the East stratum and lower densities in the 

West stratum. They were primarily collected at channel 

stations (Figs. 12.4 and 12.5). The distribution of this 

species was dissimilar to the hardhead silverside (Figs. 

12.1 and 12.2), but like the hardhead silversides, it was 

abundant in surface trawl collections in channels.

This species was rarely collected in the mangrove prop 

habitat in Coot Bay, Whitewater Bay, and Florida Bay 

(Thayer et al., 1987b), and was not collected in northern Florida Bay and adjacent estuarine waters (Powell 

et al., 2002).

Length-frequency distributions Small reef silversides 

(10–14 mm SL), indicative of recent spawning, were 

collected in May, and spawning probably occurred in 

August based on the presence of 15–19 mm SL fish in 

September (Fig. 12.6). Growth could not be estimated 

and size classes were difficult to discern, although there 

appeared to be multiple size classes in May, June, and 

September. Based on the maximum reported size of approximately 75 mm SL (Breder, 1948) larvae, juveniles 

and adults were probably collected.
88

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

5.0

10.0

15.0

20.0

A

Stratum

East Central West Channels

ytisne D

0.0

5.0

10.0

15.0

20.0

B

Figure 12.5

Mean densities (numbers 1000 m–2

) of reef silversides 

(Hypoatherina harringtonensis) collected by surface trawl 

in Florida Bay basins and channels during 1984–1985 

by (A) month and (B) stratum. Florida Bay basins are 

divided into three strata. For designations of strata see 

Fig. 2.1.
89

January

y

c

n

e

u

q

e

r

F

0

50

100

150

200

March

0

50

100

150

200

May

0

50

100

150

200

June

0

50

100

150

200

July

0

50

100

150

200

September

0

50

100

150

200

November

Size Class (mm)

41- 01

42- 02

43- 03

44- 0445- 0546- 06

0

50

100

150

200

(4) (7)

(7)

(2) (5)

(1) (1) (1)

(2) (1)

Figure 12.6

Monthly length-frequency distributions of reef silversides (Hypoatherina harringtonensis) from 

Florida Bay. Values are shown for numbers <10 for clarity.
90

Membras martinica (rough silverside)

Total numbers collected and rank of rough silverside by sampling method. There were a total of 14 species collected by 

surface trawl in each habitat.

Gear Habitat Year Number Rank

Surface trawl Basins 1984–1985 408 6

Surface trawl Channels 1984–1985 63 9

Life history The rough silverside, an atherinopsid 

and a pelagic zooplanktivore, ranges from New York to 

Florida and the northern Gulf of Mexico and northern 

Mexico. It is most abundant in the southern part of its 

range (Robins and Ray, 1986). This species inhabits 

shallow seagrass flats and is usually found along exposed 

shorelines and beaches over firm substrate (Murdy et 

al., 1997). In Chesapeake Bay, spawning is protracted 

and occurs from May to early August. Sexual maturity 

is attained at approximately 50 mm TL. Eggs have filaments that attach to vegetation on sand flats. The juvenile stage is attained at approximately 15 mm SL (Kuntz, 

1916; Martin and Drewry, 1978; Murdy et al., 1997).

Diet This species feeds on copepods and other planktonic crustaceans (Murdy et al., 1997).

Abundance and distribution Rough silversides were 

commonly collected by surface trawl in basins and 

channels in 1984–1985. Although densities appeared 

greater in the West strata and in June, there were no 

significant differences in densities among strata and 

months (Figs. 12.7 and 12.8), although they were 

collected at a greater percentage of sites in the East 

stratum. This species appeared to slightly prefer basin 

habitats to channels.

In Whitewater Bay, Coot Bay, and Florida Bay, this 

species is strongly associated with red mangrove prop 

roots, but absent in adjacent seagrass beds (Thayer et 

al., 1987b). It has not been collected in northern Florida 

Bay and adjacent waters (Powell et al., 2002).

Length-frequency distributions Based on the presence of small rough silverside (15–24 mm SL), spawning 

occurred at least from April through August (Fig. 12.9). 

Juveniles dominated collections in May and September, 

while adults were more common in June. In Gulf of 

Mexico waters, juvenile rough silversides grew 9 mm per 

month (Martin and Drewry, 1978).

Figure 12.7

Mean densities (numbers 1000 m–2

) of rough silversides (Membras martinica) collected 

by surface trawl in Florida Bay basins and channels during 1984–1985. Florida Bay 

basins are divided into three strata. For designations of strata see Fig. 2.1.
91

Month

Jan Mar May Jun Jul Sep Nov

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

6.0

7.0

8.0

A

Stratum

East Central West Channels

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

6.0

B

Figure 12.8

Mean densities (numbers 1000 m–2

) of rough silversides 

(Membras martinica) collected by surface trawl in Florida 

Bay basins and channels during 1984–1985 by (A) 

month and (B) stratum. Florida Bay basins are divided 

into three strata. For designations of strata see Fig. 2.1.
92

January

y

c

n

e

u

q

e

r

F

0

25

50

75

100

125

March

0

25

50

75

100

125

May

0

25

50

75

100

125

June

0

25

50

75

100

125

July

0

25

50

75

100

125

September

0

25

50

75

100

125

November

Size Class (mm)

91- 51

92- 52

93- 5394- 54

95- 5596- 5697- 5798- 5899- 59

0

25

50

75

100

125

Figure 12.9

Monthly length-frequency distributions of rough silversides (Membras martinica) from Florida 

Bay.
93

Chapter 13. Family Syngnathidae

Anarchopterus criniger (fringed pipefish)

Total numbers collected and rank of fringed pipefish by sampling method. There were a total of 39 species collected by 

otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 459 15

Life history Fringed pipefish range from North Carolina, the Bahamas, and northern Gulf of Mexico to 

Brazil (Robins and Ray, 1986). Like all syngnathids, 

females deposit eggs in the marsupium (brood pouch) 

of the male where eggs are then incubated. Young exit 

through a lengthwise seam in the marsupium (Robins and Ray, 1986; Murdy et al., 1997). In Cedar Key, 

Florida, Reid (1954) observed breeding males in May, 

August, and September.

The fringed pipefish is listed as vulnerable under 

the Distinct Population Segments concept defined as 

“populations that are markedly separated from other 

populations of the same taxon as a consequence of 

physical, ecological, or behavioral factors to which it 

belongs” (Musick et al., 2000). The risk factors are: it 

is rare, but locally common in Florida Bay grassbeds, 

which have undergone seagrass losses (Schmidt, 1979; 

Fourqurean and Robblee, 1999).

Diet The diet of fringed pipefish consists of copepods 

and other microcrustaceans (Reid, 1954), which are 

sucked into an elongated tube-like snout with a small 

trap door at the tip, which works like a “slurp gun” 

(Robins and Ray, 1986). 

Abundance and distribution Fringed pipefish were 

common in otter trawl collections in basins. There were 

no significant differences in densities of fringed pipefish between strata, but there were differences between 

subdivisions. This species occurred at greatest densities 

in the Western subdivision; at moderate densities in the 

Eastern, Atlantic, and Central subdivisions; and at least 

densities in the Northern and Gulf subdivisions (Figs. 

13.1 and 13.2). Among areas, it occurred at a greater 

frequency of stations in the Western and Eastern subdivisions. Temporally, there were significant inter-annual 

differences in densities. This species was least common 

in 1984–1985 prior to the seagrass die-off in western 

Florida Bay (where this species was most abundant 

in this study), and more common after the die-off of 

seagrasses in 1994–1995 (Figs. 13.1 and 13.2). Highest 

densities occurred in 1999–2000 and 2000–2001. These 

most recent high densities could be related to an increase in spatial sampling (Fig. 13.1). On Florida Bay 

mud banks, this species was most common at Buchanan 

Keys (Fig. 1.1) (Sogard et al., 1987).

Length-frequency distributions Based on lengthfrequency distributions, spawning was protracted in 

Florida Bay and occurred throughout all seasons (Fig. 

13.3). Because of multiple cohorts, growth of fringed 

pipefish could not be estimated by following progressive size classes. Smallest pipefish (<20 mm SL) were 

collected in two of seven months that we sampled. The 

largest adults were 90–99 mm SL. 
94

Figure 13.1

Mean densities (numbers 1000 m–2

) of fringed pipefish (Anarchopterus criniger) from 

bi-monthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
95

Year

5891- 48915991- 4991

69918991- 7991

0002- 99911002- 0002

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

1.4

1.6

1.8

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

3.0

3.5

A

B

C

Figure 13.2

Mean densities (numbers 1000 m–2

) of fringed pipefish 

(Anarchopterus criniger) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
96

January

y

c

n

e

u

q

e

r

F

0

25

50

75

March

0

25

50

75

May

0

25

50

75

June

0

25

50

75

July

0

25

50

75

September

0

25

50

75

November

Size Class (mm)

02<

93- 03

95- 05

97- 07

99- 09

0

25

50

75

Figure 13.3

Monthly length-frequency distributions of fringed pipefish (Anarchopterus criniger) from 

Florida Bay.
97

Syngnathus floridae (dusky pipefish)

Total numbers collected and rank of dusky pipefish by sampling method. There were a total of 39 species collected by 

otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 1683 6

Life history Dusky pipefish range from Chesapeake 

Bay, Bermuda, Bahamas, and northern Gulf of Mexico 

to Panama. They are common in seagrass beds (Robins 

and Ray, 1986). A description of their reproductive 

habits is given above for the fringed pipefish. Minimum 

size at sexual maturity for males is 116 mm SL; females, 

137 mm SL (Reid, 1954). Spawning occurs throughout 

the year in Cedar Key, Florida, but spawning is minimal 

from November through January (Reid, 1954). 

Diet The diet of the dusky pipefish consists mainly 

of very small crustaceans (Murdy et al., 1997). Near 

Cedar Key, Florida, Reid (1954) reported copepods as a 

dominant food item, followed by amphipods and small 

shrimp. A description of their feeding behavior is given 

above for the fringed pipefish.

Abundance and distribution Dusky pipefish were 

abundant in otter trawl collections in basins. Temporally, there were significant differences in densities for 

dusky pipefish between years, but not months (Figs. 

13.4 and 13.5). In concordance with the seagrass die-off, 

there was a decrease in mean densities from 1984–1985 

to 1996, then a progressive increase until 2000–2001 

when mean densities were at their maximum. They 

were, however, collected at a greater frequency of sites 

in 1984–1985 (Fig. 13.4). Spatially, there were significant differences in mean densities between strata and 

subdivision. Relatively high densities were collected in 

the West stratum compared to the East and Central 

strata (Figs. 13.4 and 13.5). Relatively high densities 

were observed in the Western subdivision providing 

further evidence that this species is associated with lush 

seagrass beds (Thayer and Chester, 1989). This species, 

along with the gulf pipefish, are the two most abundant 

pipefishes in Florida Bay (this study; Schmidt, 1979; 

FMRI1,2,3,4).

Length-frequency distributions Based on length-frequency distributions, spawning occurred throughout 

the year in Florida Bay (Fig. 13.6). Two size classes could 

be identified from November length-frequency distributions, with modes at 90–99 and 180–189 mm SL that 

could represent juveniles and adults. 
98

Figure 13.4

Mean densities (numbers 1000 m–2

) of dusky pipefish (Syngnathus floridae) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
99

Year

5891- 4891

5991- 49916991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

6.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

10.0

20.0

30.0

40.0

A

B

C

Figure 13.5

Mean densities (numbers 1000 m–2

) of dusky pipefish 

(Syngnathus floridae) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
100

January

y

c

n

e

u

q

e

r

F

0

25

50

75

100

125

March

0

25

50

75

100

125

May

0

25

50

75

100

125

June

0

25

50

75

100

125

July

0

25

50

75

100

125

September

0

25

50

75

100

125

November

Size Class (mm)

92- 02

94- 04

96- 06

98- 08901- 001921- 021941- 041

961- 061

981- 081902- 002

922- 022

942- 042

962- 062

0

25

50

75

100

125

Figure 13.6

Monthly length-frequency distributions of dusky pipefish (Syngnathus floridae) from Florida 

Bay.
101

S. scovelli (gulf pipefish)

Total numbers collected and rank of gulf pipefish by sampling 

method. There were a total of 39 species collected by otter 

trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 2426 6

Life history Gulf pipefish range from northern Georgia and the northern Gulf of Mexico to northern South 

America (Targett, 1984; Robins and Ray, 1986). A description of their reproductive habits is given above for the 

fringed pipefish. Spawning occurs throughout the year 

in waters near Cedar Key, Florida (Reid, 1954). Males attain sexual maturity at 85 mm SL, females at 99 mm SL. 

Young leave the brood pouch at 15 mm SL (Reid, 1954).

Diet Copepods are the dominant item in the diet of 

gulf pipefish in waters near Cedar Key, Florida. Amphipods and small shrimp are also eaten, and occasionally 

gastropods (Reid, 1984). In Tampa Bay, Florida, this 

species feeds mainly on cypris larvae and amphipods, 

a diet that overlaps with the zooplanktivore, the scaled 

sardine, Harengula jaguana (Motta et al., 1995). A description of their feeding behavior is given above for 

the fringed pipefish.

Abundance and distribution Gulf pipefish were 

abundant in otter trawl collections. There were 

significant temporal and spatial differences in gulf 

pipefish densities. Densities were relatively low from 

1984–1985 through 1996 and relatively high from 

1997–1998 through 2000–2001 (Figs. 13.7 and 13.8), 

and thus were not necessarily correlated with seagrass 

die-off, which occurred in the late 1980s. Monthly 

densities were highest in July and September, and 

relatively low but approximately equal in density from 

November through May (Figs. 13.7 and 13.8). Spatially, from 1984–1985 through 1998, densities were 

highest in the West stratum and lowest in the East stratum (Fig. 13.8). From 1999–2000 through 2000–2001, 

densities were highest in the Western and Northern 

subdivisions and relatively low in the Atlantic, Gulf, 

and Eastern subdivisions. Gulf pipefish occurred at a 

greater percentage of stations in the Northern subdivision, least in the Gulf subdivision. This species, along 

with the dusky pipefish, are the two most abundant 

pipefishes in Florida Bay (this study; Schmidt, 1979; 

FMRI1,2,3,4).

Length-frequency distributions Length-frequency 

distributions were unimodal and similar (Fig. 13.9), 

suggesting that spawning occurred throughout the 

year. Both juvenile and adult gulf pipefish were 

collected.
102

Figure 13.7

Mean densities (numbers 1000 m–2

) of gulf pipefish (Syngnathus scovelli) from 

bi-monthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
103

Year

5891- 4891

5991- 49916991

8991- 79910002- 99911002- 0002

ytisne D

0.0

2.0

4.0

6.0

8.0

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

5.0

10.0

15.0

20.0

25.0

30.0

A

B

C

Figure 13.8

Mean densities (numbers 1000 m–2

) of gulf pipefish 

(Syngnathus scovelli) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
104

January

y

c

n

e

u

q

e

r

F

0

100

200

300

March

0

100

200

300

May

0

100

200

300

June

0

100

200

300

July

0

100

200

300

September

0

100

200

300

November

Size Class (mm)

02<

93- 03

95- 05

97- 07

99- 09

911- 011

931- 031951- 051

971- 071

0

100

200

300

(5) (5) (3) (1) (1) (1)

(1) (2)

(1) (8) (3) (2)

(9)

(2) (1) (5) (1) (1)

(1) (9) (1) (1)

(4) (1) (6) (6) (1)

Figure 13.9

Monthly length-frequency distributions of gulf pipefish (Syngnathus scovelli) from Florida Bay. 

Values are shown for numbers <10 for clarity.
105

Hippocampus erectus (lined seahorse)

Total numbers collected and rank of lined seahorse by sampling method. There were a total of 39 species collected by 

otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 78 32

Life history Lined seahorse range from Nova Scotia 

to Uruguay or Argentina, northern Gulf of Mexico, 

Bermuda, and the Caribbean Sea. They are rare north 

of Cape Cod, Massachusetts (Breder, 1948; Robins and 

Ray, 1986; Murdy et al., 1997). A description of their reproductive habits is given above for the fringed pipefish. 

The size of expulsion from the male brood pouch is approximately 7 mm TL (Breder, 1948). Lined seahorses 

are monogamous. Males mature at 80 mm TL, females 

at 60 mm TL (Teixeira and Musick, 2001). This species 

has been presumed to spawn throughout the year in 

Florida waters, although males with distended brood 

pouches were only collected in December (Reid, 1954). 

Diet Lined seahorse are voracious ambush predators 

and feed mainly on amphipods (Murdy et al., 1997; 

Teixeira and Musick, 2001). Shrimp and microcrustaceans also occur in their diets (Reid, 1954). A description of their feeding behavior is given above for the 

fringed pipefish.

Abundance and distribution The lined seahorse was 

uncommon in otter trawl collections in Florida Bay basins 

(Figs. 13.10 and 13.11). Although densities appeared low 

in 1996, there were no significant differences in densities among years, but differences were observed among 

months (Fig. 13.11). This species was most abundant 

in July. Spatially, differences in densities were observed 

among both strata and subdivisions (Figs. 13.10 and 

13.11). This species is relatively more common in western Florida Bay, but rarely collected in the Gulf subdivision, and never collected in the Northern subdivision. 

The high observed densities in July coincide with the 

first part of the wet season, when low water levels, high temperatures, and increasing salinities have been observed 

(Sogard et al., 1987). Lined seahorses do not appear to 

utilize mud banks in Florida Bay (Sogard et al., 1987).

Length-frequency distributions Based on our lengthfrequency distributions, small fish in March were probably spawned in winter (Fig. 13.12). In addition, there is 

evidence for spawning in May and September, but based 

on a limited number of specimens we were unable to 

demonstrate that spawning occurs throughout the year. 

No discrete modal groups were observed, which is probably a result of small sample size. Juveniles recently released (approximately 7 mm TL) from the male brood 

patch and small juveniles were not encountered (≤20–

24 mm TL). Recently released lined seahorses were 

probably extruded through the net, but we are puzzled 

by the lack of specimens between 20–29 mm TL. A wide 

range of sizes was collected in most months (Fig. 13.12).
106

Figure 13.10

Mean densities (numbers 1000 m–2

) of lined seahorse (Hippocampus erectus) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
107

Year

5891- 48915991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.00

0.05

0.10

0.15

0.20

0.25

Month

Jan Mar May Jul Sep Nov

ytisne D

0.00

0.10

0.20

0.30

0.40

0.50

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.00

0.20

0.40

0.60

0.80

A

B

C

Figure 13.11

Mean densities (numbers 1000 m–2

) of lined seahorse 

(Hippocampus erectus) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) 

month, and (C) stratum/subdivision. Florida Bay 

is divided into three strata from 1984–1998 and six 

subdivisions from 1999–2001. For designations of 

strata and subdivisions see Fig. 2.1.
108

January

y

c

n

e

u

q

e

r

F

0

2

4

6

March

0

2

4

6

May

0

2

4

6

June

0

2

4

6

July

0

2

4

6

September

0

2

4

6

November

Size Class (mm)

42- 0243- 03

44- 04

45- 0546- 0647- 0748- 08

49- 09401- 001

411- 011421- 021

431- 031441- 041451- 051

0

2

4

6

Figure 13.12

Monthly length-frequency distributions of lined seahorse (Hippocampus erectus) from Florida 

Bay.
109

H. zosterae (dwarf seahorse)

Total numbers collected and rank of dwarf seahorse by sampling method. There were a total of 39 species collected by 

otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 1529 9

Life history In Florida, dwarf seahorses range from 

Biscayne Bay to Pensacola, and also occur in the Bahamas and Cuba (Böhlke and Chaplin, 1968) and in the 

entire Gulf of Mexico (Robins and Ray, 1986). At Cedar 

Key, Florida, breeding begins in early or mid-February 

and terminates by late October or early November. The 

breeding season has been correlated with day length. 

Males mature at approximately 25 mm at an age of 

approximately three months. A description of their reproductive habits is given above for the fringed pipefish. 

Recently released young expelled from the male brood 

pouch average 7–9 mm. At least three broods are produced per year (Strawn, 1958).

The dwarf pipefish is listed as vulnerable under 

the Distinct Population Segments concept defined as 

“populations that are markedly separated from other 

populations of the same taxon as a consequence of 

physical, ecological, or behavioral factors to which it 

belongs” (Musick et al., 2000).

Diet We are unaware of any information on the diet 

of this species.

Abundance and distribution The dwarf seahorse was 

abundant in otter trawl collections in basins in Florida 

Bay. Spatially, there were significant differences in 

densities among strata and subdivision (Figs. 13.13 and 

13.14). This species was collected at highest densities 

in the West stratum, but occurred at more sites in the 

Central stratum. Relatively higher densities occurred 

in the Western subdivision, while relatively low densities were observed in the Atlantic, Gulf, and Northern 

subdivisions. Temporally, there were significant differences among months and years (Figs. 13.13 and 13.14). 

We believe these yearly differences are related to the 

change in the sampling design that occurred in 1999. 

Densities were very similar from 1984 through 1998 

when strata were employed and very similar from 1999 

through 2001 when subdivisions were employed. Dwarf 

seahorse were collected at highest densities in July and 

September, which coincides with the wet season.

Similar to our findings, dwarf seahorse have been 

collected at highest densities over Florida Bay mud 

banks in western Florida Bay (Oyster Key), and lowest 

at Cowpen Keys, located in our Atlantic subdivision (Fig. 

1.1; Sogard et al., 1987). Dwarf pipefish have a greater 

affinity for seagrass than mangrove habitats (Thayer 

et al., 1987b) and appear to have a preference for the 

seagrasses Thalassia testudinum and Halodule wrightii over 

algae and mud substrate (Sheridan et al., 1997).

Length-frequency distributions Dwarf seahorse 

length-frequency distribution patterns were similar to 

that observed for gulf pipefish (Figs. 13.9 and 13.15), 

and much smaller fish were collected compared to 

the lined seahorse (Fig. 13.12). Length-frequency 

distributions were uni-modal. Monthly modes were approximately similar, suggesting that spawning occurred 

throughout the year. Growth could not be confirmed 

from distributions, but in the laboratory fish grew from 

8 mm to 18 mm in 17 days (Strawn, 1958).
110

Figure 13.13

Mean densities (numbers 1000 m–2

) of dwarf seahorse (Hippocampus zosterae) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
111

Year

5891- 4891

5991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

6.0

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

6.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

12.0

14.0

16.0

A

B

C

Figure 13.14

Mean densities (numbers 1000 m–2

) of dwarf seahorse 

(Hippocampus zosterae) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
112

Figure 13.15

Monthly length-frequency distributions of dwarf seahorse (Hippocampus zosterae) from Florida 

Bay. Values are shown for numbers <10 for clarity.

January

y

c

n

e

u

q

e

r

F

0

100

200

300

March

0

100

200

300

May

0

100

200

300

June

0

100

200

300

July

0

100

200

300

September

0

100

200

300

November

Size Class (mm)

01<

91- 5192- 52

93- 53

94- 5495- 55

0

100

200

300

(1) (3) (3)

(3) (5)

(6) (5) (2) (1)

(2) (4)

(1) (4) (4) (2)

(6) (4)

(4) (3)
113

Chapter 14. Family Lutjanidae

Lutjanus griseus (gray snapper)

Total numbers collected and rank of gray snapper by sampling 

method. There were a total of 39 species collected by otter 

trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 132 24

Life history The gray snapper, a shallow-water tropical 

species, ranges in the western Atlantic from Florida to 

Brazil including the Gulf of Mexico, Bermuda, the Bahamas, and West Indies. Juveniles occur as far north as 

Massachusetts. Transforming juveniles are commonly 

collected in North Carolina inlets, but the fate of these 

juveniles is unknown (Vergara, 1978a; Burton, 2001; 

Tzeng et al., 2003). Gray snapper spawn mainly from 

June through September in waters adjacent to Florida 

Bay with peaks in June and July (Starck and Schroeder, 1970; Rutherford et al., 1983, 1989). However, in 

other southwestern Florida waters they mainly spawn 

in winter and summer (Allman and Grimes, 2002). 

Males mature at approximately 185 mm SL, females 

at 200 mm SL (Starck and Schroeder, 1970). Eggs and 

larvae are pelagic (Thresher, 1991). Recently hatched 

larvae are approximately 2.5 mm NL and settlement 

occurs at 10–15 mm SL (Richards et al., 1994; Tseng et 

al., 2003), although Lindeman (1997) reported larger 

sizes (15–19 mm SL) at settlement. The average duration of the pre-settlement stage is 25–27-d (Allman and 

Grimes, 2002; Tzeng et al., 2003) and 33-d (Lindeman, 

1997).

Diet Gray snapper from Florida Bay, <250 mm SL, 

fed almost entirely on crustaceans and fishes (Hettler, 

1989). Copepods, amphipods, and mysids were important dietary items for fish <50 mm SL. Penaeid shrimp, 

the overall dominant prey item, and caridean shrimp 

increased in frequency of occurrence with increasing 

fish size. The fish component was dominated by rainwater killifish.

Abundance and distribution Gray snapper were common in otter trawl collections in basins. There were 

no significant differences in densities of gray snapper 

among years, but there were among months (Figs. 14.1 

and 14.2). Highest densities were observed in spring 

and fall (Fig. 14.2); however in March gray snapper were 

collected at fewer stations. Spatially, there were differences in densities among strata and subdivisions. Gray 

snapper were collected at highest densities in western 

Florida Bay and the Atlantic subdivision (Figs. 14.1 and 

14.2), suggesting they recruit into Florida Bay from the 

Florida Keys side as well as from the southwest Florida 

shelf.

Habitat for juvenile gray snapper is stage specific. 

Studies in Biscayne Bay (Lindeman et al., 1998) and 

the Florida Keys (Starck and Schroeder, 1970) indicate 

recently settled fish prefer seagrass beds, while older 

juveniles (approximately 90 mm SL and greater) utilize seagrass beds, mangrove prop and drop roots, and 

edges of channels. Fish >200 mm SL move offshore to 

reefs and wrecks. In the Atlantic subdivision, Chester 

and Thayer (1990) suggested that gray snapper recruit 

mainly to channels where seagrass communities are 

more diverse and luxuriant than those at adjacent basins 

of the Atlantic subdivision. Older juveniles apparently 

prefer mangrove habitats to adjacent seagrass sites in 

Florida Bay and adjacent waters (Thayer et al., 1987b), 

and in a Caribbean study, juvenile gray snapper (>50 

mm TL) also preferred mangrove habitats over seagrass 

beds (Cocheret de la Morinière et al., 2002).

Length-frequency distributions We collected juvenile 

gray snapper throughout the year, but the smallest fish 

were collected in fall and winter (Fig. 14.3). Starck and 

Schroeder (1970) reported that 10–20 mm SL fish are 

abundant in July and August in the Florida Keys and suggested that spawning peaks in June and July. We collected that size class in September and November, but many 

of the smallest individuals probably escaped through 

our gear. This suggested a somewhat later spawning date 

for gray snapper that recruit into Florida Bay.

Based on an age and growth study of adult gray snapper from south Florida (Burton, 2001), our collections 

consisted mainly of young-of-year, yearlings, and twoyear olds. The yearlings and two-year olds were most 

apparent in June and July, while young-of-year and yearlings were most apparent in September and November 

(Fig. 14.3). Juvenile gray snapper (32–110 mm SL) have 

been reported to grow between 0.60 and 1.02 mm d–1

(Allman and Grimes, 2002).
114

Figure 14.1

Mean densities (numbers 1000 m–2

) of gray snapper (Lutjanus griseus) from bi-monthly 

otter trawl collections in Florida Bay basins. Florida Bay is divided into three strata 

from 1984–1998 and six subdivisions from 1999–2001. For designations of strata and 

subdivisions see Fig. 2.1.
115

Figure 14.2

Mean densities (numbers 1000 m–2

) of gray snapper 

(Lutjanus griseus) from bi-monthly otter trawl collections 

in Florida Bay basins by (A) year, (B) month, and (C) 

stratum/subdivision. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–

2001. For designations of strata and subdivisions see 

Fig. 2.1.

Year

5891- 48915991- 4991

6991

8991- 79910002- 99911002- 0002

ytisne D

0.00

0.10

0.20

0.30

0.40

0.50

Month

Jan Mar May Jul Sep Nov

ytisne D

0.00

0.10

0.20

0.30

0.40

0.50

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.00

0.20

0.40

0.60

0.80

1.00

A

B

C
116

January

y

c

n

e

u

q

e

r

F

0

5

10

15

20

March

0

5

10

15

20

May

0

5

10

15

20

June

0

5

10

15

20

July

0

5

10

15

20

September

0

5

10

15

20

November

Size Class (mm)

02<

93- 03

95- 0597- 07

99- 09

911- 011

931- 031

951- 051971- 071

991- 091

912- 012

932- 032

952- 052

972- 072992- 092

0

5

10

15

20

Figure 14.3

Monthly length-frequency distributions of gray snapper (Lutjanus griseus) from Florida Bay.
117

L. synagris (lane snapper)

Total numbers collected and rank of lane snapper by sampling 

method. There were a total of 39 species collected by otter 

trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 630 13

Life history Lane snapper range from North Carolina 

around the Florida peninsula, throughout the Gulf of 

Mexico, southward to southeastern Brazil, the Bahamas, 

Bermuda, and the West Indies. Adults are most common around the Antilles, on the Campeche Bank, off 

Panama, and the northern coast of South America (Vergara, 1978a). Lane snapper have a protracted spawning 

period. Off east Florida, lane snapper spawn from April 

to July (Herrema et al., 1985), and off Key West, Florida 

from June to September with peaks in July and August 

(Domeier et al., 1996). The minimum size at sexual 

maturity of fish from Key West is 182 mm SL and 198 

mm SL for males and females, respectively (Domeier et 

al., 1996). Eggs and larvae are pelagic (Thresher, 1991). 

Recently-hatched larvae are approximately 2.4 mm NL 

and settlement occurs at 10–15 mm SL (Richards et al., 

1994; Lindeman, 1997).

Diet Juvenile lane snapper in the northern Gulf of 

Mexico mainly feed, in descending order, on the shrimp, 

Latreutes parvlus, and fish (Franks and VanderKooy, 

2000).

Abundance and distribution Lane snapper were common in otter trawl collections in basins and are the most 

abundant snapper in the Bay. Temporally, there were 

significant differences in lane snapper among years 

and months. Relatively high densities were collected in 

1999–2000 and in fall and winter (Figs. 14.4 and 14.5). 

Low densities were observed in 1996 and in July. Spatially, there were significant differences in densities among 

strata and subdivisions. This species is most abundant in 

western Florida Bay. It was not collected in the Northern 

subdivision, and its presence in the Atlantic subdivision 

suggests, like gray snapper, it recruits into Florida Bay 

from the Florida Keys as well as from the southwest 

Florida shelf (Figs. 14.4 and 14.5).

Like gray snapper, habitat for lane snapper is stage 

specific. Early juveniles prefer mangrove prop root 

canopies, but recently settled snapper are rare there 

(Lindeman et al., 1998). On the other hand, in Florida 

Bay and adjacent waters, unlike gray snapper, lane snapper do not use mangrove prop root habitats, but prefer 

adjacent seagrass beds (Thayer et al., 1987b). 

Length-frequency distributions Based on lengthfrequency distributions, lane snapper appeared to 

recruit into Florida Bay in summer, fall, and possibly 

winter (Fig. 14.6). Snapper <30 mm SL collected in July, 

September, and November indicated summer and fall 

recruitment. The presence of small juveniles in January 

and March indicated winter spawning. 

Unpublished data5 on juvenile lane snapper (approximately 25–125 mm SL) collected in Florida Bay 

indicated that growth rates ranged from 0.6 mm day–1

to 0.7 mm day–1. Back-calculated lengths of lane snapper 

from east Florida waters at age-1 and age-2 were 135 mm 

TL and 196 mm TL, respectively (Manooch and Mason, 

1984). Based on our length-frequency distributions, we 

collected mainly young-of-the year (Fig. 14.6). This species appears to leave the Bay at a much smaller size than 

the gray snapper (Fig. 14.3).

5 Settle, L. Unpublished data (March 2003). NOAA, Center for 

Coastal Fisheries and Habitat Research, 101 Pivers Island Road, 

Beaufort, NC 28516.
118

Figure 14.4

Mean densities (numbers 1000 m–2

) of lane snapper (Lutjanus synagris) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
119

Year

5891- 4891

5991- 49916991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

3.0

3.5

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nr ehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

A

B

C

Figure 14.5

Mean densities (numbers 1000 m–2

) of lane snapper 

(Lutjanus synagris) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
120

Figure 14.6

Monthly length-frequency distributions of lane snapper (Lutjanus synagris) from Florida 

Bay.

January

y

c

n

e

u

q

e

r

F

0

25

50

75

100

125

March

0

25

50

75

100

125

May

0

25

50

75

100

125

June

0

25

50

75

100

125

July

0

25

50

75

100

125

September

0

25

50

75

100

125

November

Size Class (mm)

02<

93- 03

95- 05

97- 0799- 09911- 011931- 031

951- 051

971- 071

0

25

50

75

100

125
121

Chapter 15. Family Gerreidae

Eucinostomus gula (silver jenny) 

Total numbers collected and rank of silver jenny by sampling 

method. There were a total of 39 species collected by otter 

trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 13,347 1

Life history Silver jenny range from Massachusetts to 

Argentina, including the northern Gulf of Mexico, Bahamas, and Bermuda, but occur only as stragglers north 

of Beaufort, North Carolina (Böhkle and Chaplin, 1968; 

Johnson, 1978; Robins and Ray, 1986). There is very 

little information on the reproductive habits of this species. Silver jenny have been reported to spawn offshore 

during late winter and spring. Larvae have been collected in Texas during June–October but we question if 

this data include more than one species of Eucinostomus

(Johnson, 1978). In the Whitewater Bay–Shark River 

estuary, Schmidt (1989) observed maturing female and 

male specimens as small as 60 and 73 mm SL.

Diet Juveniles from northwest Florida waters show 

a clear transition from a zooplanktivore (11–15 mm 

SL) feeding exclusively on copepods, to a benthic carnivore (40–60 mm SL) feeding almost exclusively on 

polychaetes (Carr and Adams, 1973). In the Whitewater 

Bay system (North River), Odum and Heald (1972) 

reported that harpacticoid copepods were also an important food item, but polychaetes were insignificant 

in the diets of silver jenny and appeared to be replaced 

by amphipods. This suggested that polychaetes are rare 

in the sediments of this habitat. On the other hand, in 

Note: We identified two species of Eucinostomus: E. 

argenteus and E. gula. The majority of E. argenteus were 

most likely E. harengulus. Because of this conflict we did 

not include these taxa. In addition, because of the morphological similarities between young (Matheson and 

Gilmore, 1995), we only felt confident in identifying E. 

gula that were ≥30 mm SL.

the Whitewater Bay–Shark River estuary in close proximity to Odum and Heald’s site, Schmidt (1989) found 

nereid polychaetes to be the most important food item 

consumed by silver jenny, followed by crustaceans (isopods, amphipods, and tanaids). In southeast Florida 

waters, Brook (1977) also found that the incidence of 

copepods decreased with increasing fish size. Brook 

concluded that since the copepods were almost exclusively harpacticoid, silver jenny feed among grass blades 

and on the bottom.

Abundance and distribution Silver jenny were the 

most abundant species collected by otter trawl in basins 

(Figs. 15.1 and 15.2). Differences in mean densities 

were observed among years and months. This species 

was most abundant in 1984–1985, with lowest values observed in 1994–1995, following the die-off of seagrasses 

(Figs. 15.1 and 15.2). From 1994 to 2000, there was a 

progressive increase in mean densities with a slight decrease in 2000–2001. Seasonally, highest densities were 

observed during the wet season months (Fig. 15.2).

Silver jenny could be considered a ubiquitous species 

in Florida Bay, although they were collected at highest 

mean densities in the West stratum and Western subdivision (Figs. 15.1 and 15.2), where standing crop of 

seagrasses and species diversity is highest (Zieman and 

Fourqurean, 1985; Thayer and Chester, 1989). In the 

Indian River Lagoon, Florida, silver jenny were commonly collected in open waters, often near seagrass 

beds (Matheson and Gilmore, 1995). They also have 

an affinity for mangrove prop root habitat (Thayer et 

al., 1987b). 

Length-frequency distributions Silver jenny recruited 

into Florida Bay throughout the year (Fig. 15.3). Peak 

recruitment occurred in fall, but could have also occurred in August, a month when we did not sample. The 

range of sizes was truncated in June then increased to 

a maximum in January (Fig. 15.3). This could indicate 

a movement of larger fish out of the Bay after January 

until June; from July through January some larger fish 

remain in the Bay.
122

Figure 15.1

Mean densities (numbers 1000 m–2

) of silver jenny (Eucinostomus gula) from bi-monthly 

otter trawl collections in Florida Bay basins. Florida Bay is divided into three strata 

from 1984–1998 and six subdivisions from 1999–2001. For designations of strata and 

subdivisions see Fig. 2.1.
123

Figure 15.2

Mean densities (numbers 1000 m–2

) of silver jenny 

(Eucinostomus gula) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.

Year

5891- 4891

5991- 49916991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

10.0

20.0

30.0

40.0

50.0

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

10.0

20.0

30.0

40.0

50.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

20.0

40.0

60.0

80.0

100.0

A

B

C
124

Figure 15.3

Monthly length-frequency distributions of silver jenny (Eucinostomus gula) from Florida Bay. 

Values are shown for numbers <100 for clarity.

January

y

c

n

e

u

q

e

r

F

0

500

1000

1500

2000

March

0

500

1000

1500

2000

May

0

500

1000

1500

2000

June

0

500

1000

1500

2000

July

0

500

1000

1500

2000

September

0

500

1000

1500

2000

November

Size Class (mm)

93- 0395- 05

97- 0799- 09

911- 011

931- 031

0

500

1000

1500

2000

(65) (68) (18) (3) (2) (1)

(85) (32) (13) (8) (1)

(98) (51) (14) (4)

(45) (20) (1)

(42) (8)

(1) (1)

(75) (38) (10) (1)

(73) (17) (4) (1)

(1) (1)
125

Chapter 16. Family Haemulidae

Haemulon plumieri (white grunt)

Total numbers collected and rank of white grunt by sampling 

method. There were a total of 39 species collected by otter 

trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 1606 8

Life history White grunt range from the Cape Hatteras, North Carolina, to southern Florida, the Bahamas, and Bermuda throughout the West Indies and 

Caribbean Sea, to Brazil (Breder, 1948; Courtenay and 

Sahlman, 1978). They spawn year round in the Caribbean, with peaks in March or April and minimal spawning in June–July (Munro et al., 1973; Gaut and Munro, 

1983). Off east Florida, white grunt spawn in May–June 

(Herrema et al., 1985). Eggs and larvae are pelagic. 

White grunt are 2.7 mm NL at hatching. The flexion 

stage begins at 6.0 mm NL, postflexion at 7.0 mm SL 

(Saksena and Richards, 1975). Based on general data 

on western Atlantic haemulids, white grunt settle out 

of the plankton at approximately 6 mm SL (Lindeman 

et al., 1998). There are at least two demersal life stage 

terminologies described for white grunt. Helfman et al. 

(1982) depicts fish 8–12 mm SL as small juveniles, 12–24 

mm SL as medium juveniles, 24–32 mm SL as transitional juveniles, and 32–96 as large juveniles. Appeldoorn et 

al. (1997) depicts fish 9–37 mm SL as stage 0 juveniles, 

37–65 mm SL as stage 1 juveniles, 65–120 mm SL as 

stage 2 juveniles, and 120–155 as subadults. 

Diet Carr and Adams (1973) reported on the diets 

of juveniles between 21 and 40 mm SL. The smallest 

juveniles (21–25 mm SL) feed mainly on copepods. 

Juveniles between 26–35 mm SL feed mainly on copepods, but mysids and postlarval shrimp are also consumed. Mysids and postlarval shrimp are the dominant 

item (60% of stomach contents) of the larger juveniles 

(36–40 mm SL), but copepods are also consumed. The 

presence of sand grains in the diet of all the size ranges 

indicates that juveniles feed on or near the bottom.

Abundance and distribution White grunt were abundant in otter trawl collections in basins (Figs. 16.1 and 

16.2) and were the most abundant Haemulon spp. collected. There were no significant differences in densities among months or years. There were significant 

differences in densities among strata and subdivisions. 

White grunt were most abundant in western Florida Bay, 

and absent in the north-central and eastern areas of the 

Bay. Based on the distribution of juveniles, it appeared 

that white grunt recruit into Florida Bay mainly from 

Gulf of Mexico waters, however some recruitment appeared to occur from the Atlantic (Fig. 16.1). 

White grunt were the most common grunt collected 

in grassy shore habitat in the Atlantic side of the Florida 

Keys adjacent to Florida Bay (Springer and McErlean, 

1962). In Biscayne Bay, Florida, recently settled white 

grunt utilize a variety of habitats including attached 

seagrasses and macroalgae, detached macroalgae, invertebrate assemblages (mixture of gorgonians, sponges, 

and corals), and sedimentary habitats. Juveniles 15–50 

mm SL use habitats similar to recently settled fish, but 

also use exposed hard bottom and red mangrove prop 

roots. Juveniles >50 mm SL use red mangrove prop root 

habitats and invertebrate assemblages and are absent 

from attached seagrass beds or macroalgae (Lindeman, 

1997). To the contrary, white grunt were not collected 

in mangrove prop roots in Florida Bay (Thayer et al., 

1987b).

Length-frequency distributions Following Appeldoorn et al. (1997), our collections mainly were comprised of stage 0, stage 1, and stage 2 juveniles. Subadults 

and adults were rarely collected (Fig. 16.3). Recruitment 

of white grunt into Florida Bay occurred throughout the 

sampling period, indicating that spawning in adjacent 

coastal waters occurred throughout the year. Lindeman 

(1997) reported that juvenile white grunt (17–38 mm 

SL) grow 0.38 mm day–1.
126

Figure 16.1

Mean densities (numbers 1000 m–2

) of white grunt (Haemulon plumieri) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
127

Year

5891- 48915991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.00

1.00

2.00

3.00

4.00

5.00

6.00

7.00

Month

Jan Mar May Jul Sep Nov

ytisne D

0.00

1.00

2.00

3.00

4.00

5.00

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nr ehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.00

5.00

10.00

15.00

20.00

25.00

A

B

C

Figure 16.2

Mean densities (numbers 1000 m–2

) of white grunt 

(Haemulon plumieri) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
128

January

y

c

n

e

u

q

e

r

F

0

50

100

150

200

March

0

50

100

150

200

May

0

50

100

150

200

June

0

50

100

150

200

July

0

50

100

150

200

September

0

50

100

150

200

November

Size Class (mm)

02<

93- 03

95- 05

97- 0799- 09

911- 011

931- 031

951- 051

971- 071

0

50

100

150

200

(7) (2)

(1) (8) (1) (1)

(6) (9) (3) (2)

(5) (8) (5)

(5) (3)

(2) (5) (3) (3) (3) (1)

(4) (3) (1) (3)

Figure 16.3

Monthly length-frequency distributions of white grunt (Haemulon plumieri) from Florida Bay. 

Values are shown for numbers <10 for clarity.
129

H. sciurus (bluestriped grunt) 

Total numbers collected and rank of bluestriped grunt by 

sampling method. There were a total of 39 species collected 

by otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 69 35

Life history Bluestriped grunt range from south 

Florida, the Bahamas, the West Indies, Bermuda, lower 

Gulf of Mexico, and the Caribbean Sea south to Brazil 

(Courtenay and Sahlman, 1978). In the Caribbean, 

they spawn mainly from November through April with 

a maximum in January–February. Spawning is minimal 

from May through October. Females mature at approximately 220 mm FL (Gaut and Munro, 1983). The eggs 

and larvae of bluestriped grunt are pelagic (Thresher, 

1991). Based on general data on western Atlantic haemulids, the postflexion stage occurs from ca. 6–10 mm 

SL, transforming juvenile stage ca. 10–15 mm SL, early 

juvenile stage ca. 15–20 mm SL to ca. 50 mm SL, and 

late juvenile stage from ca. 50 mm SL to 110–170 mm 

SL (Lindeman, 1997).

Diet The diet of bluestriped grunt most likely follows 

that of other grunt species as described by Lindeman 

(1997). Newly settled fish (ca. 7–20 mm SL) are diurnal 

plankton feeders; early juveniles (ca. 20–50 mm SL) 

feed on both plankton and benthic invertebrates and 

might be nocturnal feeders. Juveniles (ca. 50–100 mm 

SL) are nocturnal benthic macro-invertebrate feeders. 

Abundance and distribution The bluestriped grunt 

was uncommon in otter trawl collections in basins. Significant differences in densities were observed among 

years, but not months. Highest densities occurred, in 

descending order, in 1999–2000, 1984–1985, and 1997–

1998 (Figs. 16.4 and 16.5). They occurred at a greater 

percentage of stations in 1997–1998. We observed no 

significant differences in densities among strata. There 

were, however, differences in densities among subdivisions. Highest densities occurred in the Atlantic, Gulf, 

and Western subdivisions, suggesting that recruitment 

into Florida Bay occurred from both the Atlantic and 

Gulf of Mexico (Figs. 16.4 and 16.5).

In Biscayne Bay, newly settled early juveniles and juveniles were observed in habitats similar to white grunt 

(see above). In the Caribbean and Belize, bluestriped 

grunt juveniles prefer mangroves (Sedberry and Carter, 

1993; Cocheret de la Morinière, 2002) or seagrass beds 

(Nagelkerken et al., 2000). In Belize, bluestriped grunt 

appear to increase in abundance in mangrove creeks 

during January–June and decrease during July–December. Among mangrove, seagrass, and reef sites in the 

Caribbean, the mean size of individuals differs among 

habitats. Smallest average size fish were found in seagrass 

habitat, largest on reefs. Given the choice, bluestriped 

grunt prefer mangroves during the day as resting sites 

and seagrasses at night where they forage (Cocheret de 

la Morinière, 2002). They are common in grassy shore 

habitat in the Atlantic side of the Florida Keys adjacent 

to Florida Bay (Springer and McErlean, 1962).

Length-frequency distributions Based on Lindeman’s 

(1997) stage terminology (see above), all juvenile stages 

were present, but adults and sexually mature females 

(ca. 220 mm FL) were absent or rare (Fig.16.6). Recruitment into Florida Bay occurred throughout the 

year, indicating either that spawning occurred locally 

throughout the year or bluestriped grunt cohorts were 

from various sources with different spawning times. 

Sedberry and Carter (1993) observed that the maximum mean size of juvenile fish from Belize occurred in 

September and that length-frequency distributions were 

bimodal with a strong mode of smaller fish during winter and early spring. Because of constant recruitment, 

our modes were weak (Fig. 16.6).
130

Figure 16.4

Mean densities (numbers 1000 m–2

) of bluestriped grunt (Haemulon sciurus) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
131

Year

5891- 48915991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.00

0.05

0.10

0.15

0.20

0.25

Month

Jan Mar May Jul Sep Nov

ytisne D

0.00

0.05

0.10

0.15

0.20

0.25

0.30

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.00

0.10

0.20

0.30

0.40

0.50

A

B

C

Figure 16.5

Mean densities (numbers 1000 m–2

) of bluestriped 

grunt (Haemulon sciurus) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
132

January

y

c

n

e

u

q

e

r

F

0

10

20

30

40

March

0

10

20

30

40

May

0

10

20

30

40

June

0

10

20

30

40

July

0

10

20

30

40

September

0

10

20

30

40

November

Size Class (mm)

02<

93- 03

95- 05

97- 07

99- 09

911- 011

931- 031951- 051

971- 071

991- 091912- 012

0

10

20

30

40

Figure 16.6

Monthly length-frequency distributions of bluestriped grunt (Haemulon sciurus) from Florida 

Bay.
133

Orthopristis chrysoptera (pigfish)

Total numbers collected and rank of pigfish by sampling method. There were a total of 39 species collected by otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 1246 11

Life history Pigfish range along the Atlantic coast from 

New Jersey (uncommon north of Virginia) to the tip of 

Florida, throughout the Gulf of Mexico to Yucatan, and 

in Bermuda. Adults inhabit coastal waters and occur 

over mud and sand substrates (Courtenay and Sahlman, 

1978; Johnson, 1978). In Florida waters, pigfish spawn 

during spring, but studies in the Gulf of Mexico report 

spawning in late winter (Sutter and McIlwain, 1987). 

Pigfish eggs are pelagic, and larvae are approximately 

1.4 mm NL at hatching. The juvenile stage is attained 

at 20 mm SL and sexual maturity is usually attained at 

200–215 mm TL. Pigfish reach a maximum length of 

460 mm SL (Hildebrand and Cable, 1930; Johnson, 

1978; Sutter and McIlwain, 1987). 

Diet Carr and Adams (1973) examined the diets of 

pigfish between 16 and 80 mm SL that were collected 

near Crystal River, Florida. They found that juvenile 

pigfish pass through two distinct feeding stages with a 

gradual transition from planktivory to carnivory. Juveniles 16–30 mm SL are planktivorous, mainly feeding 

on copepods, mysids, and postlarval shrimp. The smallest fish (16–20 mm SL) feed mainly on copepods and 

with subsequent growth, mysids and shrimp dominated 

their diet. At approximately 26 mm SL, polychaetes appear in their diets. Transition is complete at 41–45 mm 

SL, at which time pigfish mainly consume polychaetes, 

mysids, and small quantities of amphipods. With increasing size, polychaetes decline in importance and 

caridean and penaeid shrimp became dominant food 

items. 

In the Whitewater Bay–Shark River estuary, pigfish 

(20–39 mm and 40–89 mm SL) consume crustaceans 

(mainly gammaridean amphipods and isopods) more 

frequently than polychaetes and mollusks. However, 

larger fish feed on a greater proportion of polychaetes 

and mollusks (Schmidt, 1993). In the northern Gulf of 

Mexico, Howe (2001) noted two distinct size-specific 

diets. Small pigfish (108–150 mm SL) mainly consume, 

in descending order, polychaetes, penaeid shrimp, 

and sea anemones; while larger fish (151–198 mm SL) 

mainly consume, in descending order, penaeid shrimp, 

polychaetes, and fishes. 

Abundance and distribution Pigfish were abundant 

in otter trawl collections in basins, and were the most 

abundant grunt collected in Florida Bay. Significant 

differences in densities were observed among months 

and years, although differences among years could be 

biased by the increase in spatial sampling in the Northern and Eastern subdivisions during 1999–2001 (Fig. 

16.7). Monthly, highest densities occurred in spring, 

lowest in summer and fall (Fig. 16.8). There were significant differences in densities of pigfish among strata 

and subdivisions. This species mainly occurs in western 

Florida Bay (Figs. 16.7 and 16.8).

At Cedar Key, Florida, pigfish apparently occupy different habitats seasonally (Reid, 1954). Reid collected 

the majority of pigfish from May through November. 

During spring and early summer, juveniles occupy shallow flats with considerable seagrass, and as summer 

and early fall progress, juveniles (75–115 mm SL) are 

mainly in channels and deep flats with little vegetation. 

Unlike our findings, Tabb and Manning (1961) collected juveniles from July through October in western 

Florida Bay.

Apparently, pigfish spawn in Gulf of Mexico waters 

and recruit into the Bay from the Gulf. Pigfish do not 

appear to utilize mangrove prop root habitats in Florida 

Bay and adjacent waters (Thayer et al., 1987b), and have 

not been collected in a grassy shore habitat in the Atlantic side of the Florida Keys (Springer and McErlean, 

1962), further indicating this species recruits into the 

Bay from the Gulf of Mexico. 

Length-frequency distributions Pigfish recruited 

into Florida Bay at a size <25 mm SL during winter 

and early spring (Fig. 16.9). It appeared that two age 

classes (young-of-the-year and yearlings) were collected 

at least from January through July. Sexually mature fish 

(>200 mm SL) were rarely encountered. Near the end 

of the growing season (November) young-of the-year 

fish ranged from 70 to 190 mm SL with a slight mode 

at 120–129 mm SL. 
134

Figure 16.7

Mean densities (numbers 1000 m–2

) of pigfish (Orthopristis chrysoptera) from bi-monthly 

otter trawl collections in Florida Bay basins. Florida Bay is divided into three strata 

from 1984–1998 and six subdivisions from 1999–2001. For designations of strata and 

subdivisions see Fig. 2.1.
135

Year

5891- 4891

5991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

3.0

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

6.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nr ehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

5.0

10.0

15.0

20.0

25.0

A

B

C

Figure 16.8

Mean densities (numbers 1000 m–2

) of pigfish (Orthopristis chrysoptera) from bi-monthly otter trawl collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
136

Figure 16.9

Monthly length-frequency distributions of pigfish (Orthopristis chrysoptera) from Florida Bay.

January

y

c

n

e

u

q

e

r

F

0

100

200

March

0

100

200

May

0

100

200

June

0

100

200

July

0

100

200

September

0

100

200

November

Size Class (mm)

02<93- 03

95- 05

97- 07

99- 09

911- 011931- 031951- 051971- 071991- 091

912- 012

932- 032

0

100

200

(7) (8) (9) (5) (4) (1) (1)

(8) (8)

(2) (9)

(6) (1) (5) (1) (1)

(2) (2)

(2) (2) (2) (3) (9) (3)

(8) (8) (3)

(8) (5) (5) (8) (2) (1) (1)

(2) (1) (4) (1)
137

Chapter 17. Family Sparidae

Archosargus probatocephalus (sheepshead)

Total numbers collected and rank of sheepshead by sampling 

method. There were a total of 39 species collected by otter 

trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 89 28

Life history Sheepshead range from Nova Scotia to 

Florida, throughout the Gulf of Mexico and Caribbean 

Sea, to Brazil. They are absent from the West Indies 

and Bermuda. Late larvae and juveniles (ca. <50 mm) 

inhabit seagrass beds; larger juveniles and adults are 

associated with rocks, pilings, wrecks, and bulkheads 

(Johnson, 1978; Randall and Vergara, 1978; Robins 

and Ray, 1986; Able and Fahay, 1998). Spawning occurs 

offshore during February through April in Louisiana 

and east Florida (Johnson, 1978; Render and Wilson, 

1992). Sexual maturity is first attained at age-2 for both 

sexes (ca. 300 mm FL). Eggs are pelagic and sheepshead 

are 1.6–1.7 mm NL at hatching. Settlement occurs at 

ca. 8 mm SL and the mean size attained in the juvenile 

period is 19 mm SL (Parsons and Peters, 1989; Beckman et al., 1991; Render and Wilson, 1992; Tucker and 

Alshuth, 1997).

Diet Sheepshead undergo ontogenetic shifts in diet 

from a juvenile diet composed of soft-bodied forms 

(e.g., copepods, amphipods, and mysids) that are associated with seagrasses to a diet composed of hard 

prey items (e.g., crabs, barnacles, oysters, and clams). 

Their consumption of hard prey items is linked to the 

development (>60 mm SL) of an apparatus termed oral 

jaw crushers. At approximately 60 mm SL, sheepshead 

leave grassbeds and occupy adult habitats (described 

above) when barnacles become an important dietary 

item. Ontogenetic shifts in their diet are exemplified 

by their consumption of crabs. At 30–90 mm SL, sheepshead consume crab zoea; at 91–180 mm SL, small stone 

crabs; and at >80 mm SL, large stone crabs. Similarly, 

smaller fish consume small, soft-shelled bivalves, while 

more robust bivalves increasingly are consumed during ontogeny (Johnson, 1978; Hernandez and Motta, 

1997).

In the Whitewater Bay system (North River), Odum 

and Heald (1972) observed that newly recruited sheepshead feed in seagrass beds, first restricted to copepods 

then to amphipods, chironomods, and mysids. At approximately 35–40 mm SL they move into the hard 

substrate of the North River and move upstream where 

their diet consists of encrusting forms. 

Abundance and distribution Sheepshead were uncommon in otter trawl collections in basins, indicating 

that Florida Bay is not an important nursery area for 

this species. Significant differences in densities were 

observed among years, but not months. Highest densities were observed in 1996, and they were absent from 

collections in three annual periods (Figs. 17.1 and 

17.2). Sheepshead were mainly collected in western 

Florida Bay, indicating that recruitment was from the 

Gulf of Mexico. 

Sheepshead have not been collected in grassy shore 

habitat in the Atlantic side of the Florida Keys adjacent 

to Florida Bay (Springer and McErlean, 1962), further 

indicating this species recruits into the Bay from the 

Gulf of Mexico. In Tampa Bay, sheepshead are mainly 

associated with the macroalgae, Diplanthera. They move 

off the grass beds in August (mean size = 41 mm SL) and 

enter the adult habitat (Springer and Woodburn, 1960).

In the Whitewater Bay system (North River), Odum 

and Heald (1972) found sheepshead to be the second 

most abundant gamefish species. They spend their first 

months in Whitewater Bay seagrass beds. At about 35–40 

mm SL they begin to move into the mouth of the North 

River. The influx of these small fish begins at the mouth 

of the river in June and lasts until late fall. Movement 

from the Bay to the river system appears to be related 

to diet (see above). 

Length-frequency distributions Sheepshead recruit 

into Florida Bay in spring and early summer (Fig. 17.3). 

Based on modes of young-of-the-year from March to 

November, fish grew 40 mm in SL in six months, a relatively slow rate of growth. Annuli form on sheepshead 

sagittae during April–May (Beckman et al., 1991), and 

based on that study, we ascribed a mode observed of 

70–79 mm SL in March as one growth year. Johnson 

(1978) summarized growth rates of sheepshead from 

Beaufort, North Carolina, and Tampa Bay, Florida, 

and also concluded that this species is slow growing. 

Sheepshead from Tampa Bay, which averaged 21 mm 

SL in June, 29 mm SL in July, and 42 mm SL in August 

(Springer and Woodburn, 1960), were slightly smaller 

than our specimens (Fig. 17.3). The progressive decline 

in our modal frequencies from May (mode = 20–29 mm 

SL) to November (mode = 60–69 mm SL) could indicate a movement of juveniles into their adult habitat as 

reported by Springer and Woodburn (1960) in Tampa 

Bay (see above), although this is confounded by natural 

mortality.
138

Figure 17.1

Mean densities (numbers 1000 m–2

) of sheepshead (Archosargus probatocephalus) from 

bi-monthly otter trawl collections in Florida Bay basins. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 1999–2001. For designations of 

strata and subdivisions see Fig. 2.1.
139

Year

5891- 48915991- 499169918991- 79910002- 99911002- 0002

ytisne D

0.00

0.20

0.40

0.60

0.80

Month

Jan Mar May Jul Sep Nov

ytisne D

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.00

0.20

0.40

0.60

0.80

A

B

C

Figure 17.2

Mean densities (numbers 1000 m–2

) of sheepshead 

(Archosargus probatocephalus) from bi-monthly otter 

trawl collections in Florida Bay basins by (A) year, 

(B) month, and (C) stratum/subdivision. Florida Bay 

is divided into three strata from 1984–1998 and six 

subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
140

January

y

c

n

e

u

q

e

r

F

0

20

40

60

March

0

20

40

60

May

0

20

40

60

June

0

20

40

60

July

0

20

40

60

September

0

20

40

60

November

Size Class (mm)

02<

93- 0395- 0597- 07

99- 09

911- 011931- 031951- 051971- 071991- 091

912- 012

932- 032

0

20

40

60

Figure 17.3

Monthly length-frequency distributions of sheepshead (Archosargus probatocephalus) from 

Florida Bay.
141

Calamus arctifrons (grass porgy)

Total numbers collected and rank of grass porgy by sampling 

method. There were a total of 39 species collected by otter 

trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 71 34

Life history Grass porgy range primarily along the 

west coast of Florida in shallow water, but occasionally 

stray to Louisiana and Texas. The adults prefer shallow 

seagrass beds and other vegetated bottoms, and are 

also found over hard bottoms. Grass porgy are most 

likely hermaphroditic. Spawning apparently occurs in 

offshore waters during winter and spring (Darcy, 1986). 

Based on the presence of small fish collected at Cedar 

Key, Florida, Reid (1954) suggested that spawning occurs during early spring. Size and age at maturity are 

unknown. They attain a maximum length of 220 mm 

(type of measurement unspecified) (Darcy, 1986).

Diet Grass porgy ranging from 19 to 140 mm SL were 

found to mainly feed on crustaceans (copepods, amphipods, mysids, and shrimps), occasionally on pelycepod 

and gastropod mollusks, and rarely on polychaetes 

(Reid, 1954).

Abundance and distribution Grass porgy were uncommon in otter trawl collections in basins (Figs. 17.4 and 

17.5), indicating that Florida Bay is a marginal nursery 

area for this species. We observed significant differences 

in densities among years, but not months. Densities 

were highest in 2001 and lowest in 1994–1995 (Figs. 17.4 

and 17.5) following the die-off of seagrass. However, 

grass porgy were collected at a greater percentage of 

stations in 1997–1998 and 1984–1985 (Fig. 17.4). Grass 

porgy mainly occurred in western Florida Bay, although 

few fish were collected in the Atlantic subdivision, indicating that recruitment into Florida Bay is mainly from 

the Gulf of Mexico, although recruitment through the 

Florida Keys is possible (Figs. 17.4 and 17.5). 

Grass porgy are restricted to shallow water, probably 

because of their affinity for benthic vegetation (Darcy, 

1986), but they have not been collected on mud banks 

in Florida Bay (Sogard et al., 1987, 1989a; Sheridan et 

al., 1997; Matheson et al., 1999). They apparently do not 

use mangrove prop root habitats (Thayer et al., 1987b). 

They were rarely collected in grassy shore habitat in the 

Atlantic side of the Florida Keys adjacent to Florida Bay 

(Springer and McErlean, 1962), which could be related 

to minimal recruitment to the Bay from the Keys. Although uncommon in Florida Bay, this species has been 

reported to be abundant on the west Florida shelf with 

highest abundances recorded in shallow (9–18 m) water 

associated with hard bottom (Darcy, 1986). The lack of 

hard bottom habitat in western Florida Bay could limit 

their use of Florida Bay. 

In western Florida Bay, Tabb and Manning (1961) 

reported that grass porgy were abundant during 1957–

1959, but in future years they were uncommon. They 

attributed their decline to an increase in turbid waters 

from Joe Kemp Key to East Cape Sable. If they are sensitive to turbid waters, this could be a potential indicator 

species for Florida Bay water quality. Their sensitivity to 

turbid waters could possibly be a factor that explains 

their absence from turbid northwestern Gulf of Mexico 

waters (Darcy, 1986).

Length-frequency distributions Based on the collection of small fish, recruitment into Florida Bay occurred 

from February through May (Fig. 17.6). The growth rate 

of grass porgy is unknown (Darcy, 1986) and, based on 

limited data, we estimated grass porgy to attain a modal 

length of 90–99 mm SL in March after a year of growth 

(Fig. 17.6). Based on July through November modes, 

grass porgy grew approximately 0.3 mm SL day–1 (10 mm 

SL per month), which would relate to 120 mm SL at age-1.
142

Figure 17.4

Mean densities (numbers 1000 m–2

) of grass porgy (Calamus arctifrons) from bi-monthly 

otter trawl collections in Florida Bay basins. Florida Bay is divided into three strata from 

1984–1998 and six subdivisions from 1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
143

Year

5891- 4891

5991- 49916991

8991- 7991

0002- 99911002- 0002

ytisne D

0.00

0.10

0.20

0.30

0.40

Month

Jan Mar May Jul Sep Nov

ytisne D

0.00

0.04

0.08

0.12

0.16

0.20

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.00

0.40

0.80

1.20

1.60

A

B

C

Figure 17.5

Mean densities (numbers 1000 m–2

) of grass porgy 

(Calamus arctifrons) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
144

January

y

c

n

e

u

q

e

r

F

0

5

10

15

March

0

5

10

15

May

0

5

10

15

June

0

5

10

15

July

0

5

10

15

September

0

5

10

15

November

Size Class (mm)

02<

93- 03

95- 0597- 0799- 09911- 011931- 031951- 051971- 071

991- 091

0

5

10

15

Figure 17.6

Monthly length-frequency distributions of grass porgy (Calamus arctifrons) from Florida Bay.
145

Lagodon rhomboides (pinfish)

Total numbers collected and rank of pinfish by sampling method. There were a total of 39 species collected by otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 7626 4

Life history Pinfish range from Cape Cod southward 

along the Atlantic coast to Florida, around the tip of 

Florida, throughout the Gulf of Mexico to Yucatan, 

Mexico, along northern Cuba, and Bermuda. It is not 

abundant north of Delaware (Breder, 1948; Caldwell, 

1957; Randall and Vergara, 1978). Caldwell (1957) 

noted that “ . . . pinfish is amazingly ubiquitous in its 

ecological distribution. It is found associated with wide 

ranges of temperature, salinity, depth, bottom type, 

current, and other ecological conditions, and this tolerance of the species to wide ranges of physical factors 

permits it to exist in many types of habitats.” Pinfish 

spawn in offshore waters, and numerous studies in the 

Gulf of Mexico (summarized by Darcy, 1985) suggest 

that spawning occurs during fall, winter, and spring. 

Collins and Finucane (1984) conducted an ichthyoplankton survey of inshore Gulf of Mexico waters (≥10 

m water depth) adjacent to the Florida Everglades and 

did not collect any pinfish larvae. However, Houde et 

al.7 sampled offshore of Collins and Finucane (1984) 

and collected pinfish larvae, mainly, during winter. A 

great percentage (>60%) of larvae were taken in waters 

<30 m, but larvae were also collected in waters of 7–64 

m depths. Pinfish eggs are pelagic, larvae hatch at 2.3 

mm TL (Darcy, 1985), and the juvenile stage is attained 

at 13 mm SL (Zieske, 1989). Ingress into estuaries occurs at 10–20 mm SL (Tabb and Manning, 1961; Darcy, 

1985). Females mature at approximately 80–100 mm SL 

(Hansen, 1970; Schmidt, 1993). 

Diet Pinfish are visual feeders and generally are planktivores as small juveniles, then shift to larger benthic 

organisms and algae during ontogeny (Darcy, 1985). 

Differences in seagrass composition appear to influence the diet of pinfish (Carr and Adams, 1973; Stoner, 

1980). Carr and Adams (1973) study site was dominated by Ruppia maritima and Halodule wrightii; whereas, 

Stoner’s sites were dominated by Thalassia testudinum. In 

Florida Bay the distribution of pinfish (Fig. 17.7) mainly 

coincides with T. testudinum (Thayer and Chester, 1989); 

hence, we expect the diet of pinfish in Florida Bay to 

generally resemble the diet of pinfish from Apalachee 

Bay. Where H. wrightii dominates (e.g., western and 

northeastern Florida Bay, Figs. 3.6 and 3.7), the diet 

might resemble that of pinfish from Crystal River.

Abundance and distribution Pinfish were abundant 

in otter trawl collections in basins (Figs. 17.7 and 17.8) 

indicating that Florida Bay is a nursery area for this 

forage species. We observed significant differences in 

densities among years, but not months, even though 

densities in May appeared to be relatively high. Densities were relatively low following the die-off of seagrass 

(1994–1996), highest in 1997–1998, but then depressed 

in 1999–2000 (Figs. 17.7 and 17.8). Seagrass die-off 

could have influenced the density of pinfish, but this 

species spawns in offshore waters where biological and 

physical conditions could confound our understanding 

of the effect of seagrass die-off on pinfish populations.

At Cedar Key, Florida, pinfish occurred throughout 

the year, but numbers diminish during winter (Reid, 

1954). In western Florida Bay, pinfish have been reported to occur in shallow water during warmer months 

and move to deeper waters during winter (Tabb and 

Manning, 1961). In west coast Florida waters, Nelson 

(1998) mainly collected young-of-the-year pinfish in 

water depths <3.5 m, and concluded their distribution 

is depth limited due to their dependence on seagrass 

for cover and food. This species, which is considered to 

be ubiquitous and tolerant of a wide range of physical 

and biological conditions (Caldwell, 1957), was rarely 

collected in the Eastern subdivision and absent from the 

Northern subdivision. Recruitment constraints could 

limit their distribution in these areas. 

Length-frequency distributions Based on the appearance of fish <20 mm SL, pinfish recruited into 

Florida Bay during winter and spring (Fig. 17.9). Our 

length-frequency patterns resemble that for Cedar Key, 

Florida (Reid, 1954), in that collections are dominated 

by young-of-the-year fish. However, in November, the 

mode for young-of-the-year pinfish in Florida Bay is 

80–99 mm SL; in Cedar Key the mode is at 60–70 mm 

SL, suggesting faster growth of Florida Bay fish. Nelson 

(1998) calculated monthly instantaneous growth rates 

(Ricker, 1975) for pinfish from April through July for 

three west coast Florida Bays. The average monthly 

instantaneous growth rate was 0.18 (20% monthly 

increase in body length), which Nelson (1998) considered to be fast growth. We calculated instantaneous 

growth rates from modal lengths from March through 

November (Fig. 17.9) and observed a similar rate 

(0.17), suggesting that environmental (e.g., temperature, salinity) and biological (e.g., prey abundance and 

quality) conditions in these Florida waters are suitable 

for pinfish habitat.

7 Houde, E. D, J. C. Leak, C. E. Dowd, S. A. Berkeley, and W. J. Richards. 1979. Ichthyoplankton abundance and diversity in the eastern 

Gulf of Mexico. Report to the Bureau of Land Management under 

contract no. AA550-CT7-28, June 1979, 546 p.
146

Figure 17.7

Mean densities (numbers 1000 m–2

) of pinfish (Lagodon rhomboides) from bi-monthly 

otter trawl collections in Florida Bay basins. Florida Bay is divided into three strata 

from 1984–1998 and six subdivisions from 1999–2001. For designations of strata and 

subdivisions see Fig. 2.1.
147

Year

5891- 4891

5991- 4991

6991

8991- 79910002- 9991

1002- 0002

ytisne D

0

5

10

15

20

25

30

Month

Jan Mar May Jul Sep Nov

ytisne D

0

10

20

30

40

50

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0

20

40

60

80

100

120

A

B

C

Figure 17.8

Mean densities (numbers 1000 m–2

) of pinfish (Lagodon 

rhomboides) from bi-monthly otter trawl collections in 

Florida Bay basins by (A) year, (B) month, and (C) 

stratum/subdivision. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–

2001. For designations of strata and subdivisions see 

Fig. 2.1.
148

Figure 17.9

Monthly length-frequency distributions of pinfish (Lagodon rhomboides) from Florida Bay. 

Values are shown for numbers <25 for clarity.

January

y

c

n

e

u

q

e

r

F

0

500

1000

1500

2000

March

0

500

1000

1500

2000

May

0

500

1000

1500

2000

June

0

500

1000

1500

2000

July

0

500

1000

1500

2000

September

0

500

1000

1500

2000

November

Size Class (mm)

02<

93- 03

95- 05

97- 0799- 09911- 011931- 031951- 051

0

500

1000

1500

2000

(11) (9) (10) (12) (5)

(2) (18) (5)

(14) (4)

(21) (16) (5)

(11) (8) (1)

(1) (12) (1)

(15) (6) (3)
149

Chapter 18. Family Sciaenidae

Bairdiella chrysoura (silver perch)

Total numbers collected and rank of silver perch by sampling 

method. There were a total of 39 species collected by otter 

trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 1296 10

Life history Silver perch range along the Atlantic coast 

from Connecticut to Florida, around the tip of Florida, 

and throughout the Gulf of Mexico to Rio Grande 

(Chao, 1978). In the northern part of their range they 

appear in bays only in summer (Breder, 1948). Larvae 

have been collected in all months sampled (February, 

May, August, and September) in Gulf of Mexico coastal 

and estuarine waters adjacent to Everglades National 

Park (Collins and Finucane, 1984). Running ripe male 

and female fish have been observed in Conchie Channel 

in northwest Florida Bay (Tabb and Manning, 1961), 

but larvae have not been collected in the Bay (Powell, 

2002). Eggs are pelagic and hatching occurs at 1.5–1.8 

mm NL. Transformation to the juvenile stage occurs at 

10–12 mm SL (Ditty and Shaw, 1994). 

Diet Carr and Adams (1973) reported on the diets 

of juvenile silver perch (96–160 mm SL) collected in 

northwestern Florida. Smallest juveniles (6–15 mm SL) 

consume copepods exclusively. For fish 16–40 mm SL, 

consumption of copepods decreases and mysids and 

very small shrimp increases. For fish greater than 40 

mm SL, small caridean shrimp dominates the diet and 

small fish, amphipods, and polychaetes are occasionally 

significant dietary items in larger fishes. In the Whitewater Bay–Shark River estuary, small silver perch (≤50 

mm SL) consume mainly crustaceans (in descending 

order, isopods, amphipods, and non-penaeid shrimp). 

Intermediate size fish (51–92 mm SL) consume mainly 

non-penaeid shrimp. Largest silver perch (≥93 mm SL) 

have a diverse diet that is dominated by crustaceans 

(in descending order, non-penaeid shrimp, penaeids, 

amphipods, and isopods) (Schmidt, 1993). 

Abundance and distribution Silver perch were abundant in otter trawl collections in basins (Figs. 18.1 and 

18.2), indicating that Florida Bay basins are a valuable 

nursery area for this species. We observed significant 

differences in mean densities among years, but not 

months. This species was most abundant in 1996 following a decline in densities from the previous year. Silver 

perch were distributed mainly in western and northcentral Florida Bay. In the Western subdivision, silver 

perch were most abundant and occurred at a majority 

of stations (Figs. 18.1 and 18.2). 

This species is uncommon in adjacent Biscayne Bay, 

which is influenced by Straits of Florida waters (Serafy 

et al., 1997b), and does not appear to inhabit mangrove 

prop root habitats in Florida Bay and adjacent waters 

(Thayer et al., 1987b). It is common on western Florida 

Bay mud banks where it is nocturnally active (Sogard et 

al., 1989a; 1989b).

Length-frequency distributions Silver perch recruited in Florida Bay throughout the year (Fig. 18.3), 

indicating protracted spawning. Recruitment appeared 

to be most intense in April (based on the May distribution), May, and June; hence, recruits during these three 

months supplied the greatest proportion to the age-0 

age class. Based on modes from June through November, silver perch grew approximately 16 mm SL per 

month. Based on the slight mode in March, fish attain 

an average length of 110–119 mm SL after one year’s 

growth. Silver perch from Florida Bay are apparently 

slower growing than fish from temperate estuaries. Fish 

from Delaware Bay grew 34 mm per month from June 

through September and attained a size of 130 mm at 

age-1 (Johnson, 1978). 
150

Figure 18.1

Mean densities (numbers 1000 m–2

) of silver perch (Bairdiella chrysoura) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
151

Year

5891- 4891

5991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

6.0

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

5.0

10.0

15.0

20.0

A

B

C

Figure 18.2

Mean densities (numbers 1000 m–2

) of silver perch 

(Bairdiella chrysoura) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
152

January

y

c

n

e

u

q

e

r

F

0

100

200

300

400

March

0

100

200

300

400

May

0

100

200

300

400

June

0

100

200

300

400

July

0

100

200

300

400

September

0

100

200

300

400

November

Size Class (mm)

02<

93- 03

95- 0597- 07

99- 09

911- 011

931- 031

951- 051

971- 071

0

100

200

300

400

(2) (7) (7) (2) (1)

(6) (4) (6) (7) (8) (2) (7)

(1) (4) (4) (9) (6) (1)

(2) (8) (8) (2) (2)

(8) (1) (4) (1) (3) (3) (4)

(6) (7) (1)

(8) (4) (1)

Figure 18.3

Monthly length-frequency distributions of silver perch (Bairdiella chrysoura) from Florida Bay. 

Values are shown for numbers <10 for clarity.
153

Cynoscion nebulosus (spotted seatrout)

Total numbers collected and rank of spotted seatrout by sampling method. There were a total of 39 species collected by 

otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 215 19

Life history Spotted seatrout range from New York to 

the tip of Florida and throughout the Gulf of Mexico 

to Laguna Madre, Mexico. They are uncommon north 

of Delaware. They are generally found in estuaries 

and shallow coastal waters over sand bottoms and in 

seagrass beds. They spend their entire life history in 

Florida Bay (Breder, 1948; Chao, 1978; Rutherford et 

al., 1989). They have a protracted spawning season, 

are multiple spawners, and reach sexual maturity at 

an early age (Brown-Peterson and Warren, 2001). 

Hatchdate distributions calculated for spotted seatrout in Florida Bay (Powell et al., 2004), along with 

early stage larval collections (Powell, 2003), indicate 

spotted seatrout spawn between March and October 

with the majority of spawning occurring between 27° 

and 35°C , with very little spawning between 20° and 

26°C. However, Stewart (1961) observed that spotted 

seatrout in Florida Bay spawn throughout the year 

with peaks in spring and fall. Another larval fish study 

in Florida Bay indicated that some spawning occurs 

as early as February and continues into December 

(Rutherford et al., 1989). 

Peak spawning activity of spotted seatrout is highly 

variable (McMichael and Peters, 1989; Brown-Peterson 

and Warren, 2001). Spawning peaks, based on larval 

collections in Florida Bay, occur in June, August, and 

September (Powell, 2003), and early May, late June, and 

late August/early September, based on 1995 hatchdate 

distributions (Powell et al., 2004). McMichael and Peters (1989) observed two spawning peaks – spring and 

summer in Florida waters. Tucker and Faulkner (1987) 

observed that older fish participate in two peak spawning periods, and a portion of the larger spring-spawned 

fish (age-1+) enter the spawning population for their 

first time during their second summer, augmenting the 

summer spawn. 

Eggs of spotted seatrout are pelagic and larvae hatch 

at 1.3–1.6 mm NL. Settlement occurs between 4–8 mm 

SL and transformation to the juvenile stage occurs 

at 10–12 m SL (Ditty and Shaw, 1994; Rooker et al., 

1998).

Diet Hettler (1989) reported on the food habits of 

juvenile spotted seatrout from Florida Bay, mainly 

from seagrass beds. Smallest juveniles (<30 mm SL) 

collected in seagrass beds feed mainly on amphipods 

(Gammarus spp.), mysids, and carideans (Periclimenes sp. 

and Tozeuma carolinensis). Pink shrimp (Farfantepenaeus 

duorarum) dominate the diet of fish >30 mm SL. At 50 

mm SL there is an increase of fish in the diet, mainly 

the rainwater killifish (Lucania parva). Hettler (1989) 

also observed seasonal shifts in diet that might coincide 

with the abundance of prey, particularly the seasonal 

abundance of pink shrimp.

Abundance and distribution Spotted seatrout were 

common in otter trawl basin collections (Figs. 18.4 

and 18.5). Significant differences in mean densities 

was not observed among years or months, although 

spotted seatrout were collected at a greater percentage 

of stations in 1996. There were significant differences 

among strata and subdivisions. Spotted seatrout were 

most common in western Florida Bay, but there is an 

indication that their distribution has extended into the 

northern central portion of the Bay where hyperhaline 

conditions can occur. 

Thayer et al. (1999), in a comparison between preand post-seagrass die-off (1984–1995 vs. 1994–1996), 

also observed the extension of spotted seatrout into 

the north central Bay in their 1994–1996 sampling. 

This extension could be explained by the absence of 

hyperhaline conditions during this period (Boyer et 

al., 1999). 

Somewhat different than our results, staff at the 

Florida Marine Research Institute (FMRI), sampling 

fixed stations with trawls and seines in Florida Bay, collected a considerable number of spotted sea trout in 

1994 (n = 317) and 1995 (n = 598) relative to 1996 (n = 

130) and 1997 (n = 128). The majority of their fish were 

also collected in western and north-central Florida Bay 

(FMRI1,2,3,4).

Length-frequency distributions Length-frequency 

distributions for spotted seatrout were similar from May 

through November, which is indicative of their protracted spawning habit (Fig. 18.6). Juveniles were fully 

recruited to our gear at 20–29 mm SL, which equates 

to 35–50 days old. Growth of larvae and juveniles (<80 

mm SL) is best described by the equation loge SL = 

–1.31 + 1.2162 (loge age), and for juveniles (12–80 mm 

SL) by the equation SL = –7.50 + 0.8417 (age). Hence 

juveniles 12–80 mm SL grow 0.8 mm in SL day–1 (Powell 

et al., 2004). 
154

Figure 18.4

Mean densities (numbers 1000 m–2

) of spotted seatrout (Cynoscion nebulosus) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
155

Figure 18.5

Mean densities (numbers 1000 m–2

) of spotted seatrout 

(Cynoscion nebulosus) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.

Year

5891- 4891

5991- 4991

6991

8991- 79910002- 9991

1002- 0002

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

1.0

2.0

3.0

4.0

5.0

A

B

C
156

Figure 18.6

Monthly length-frequency distributions of spotted seatrout (Cynoscion nebulosus) from Florida 

Bay.

January

y

c

n

e

u

q

e

r

F

0

25

50

75

100

March

0

25

50

75

100

May

0

25

50

75

100

June

0

25

50

75

100

July

0

25

50

75

100

September

0

25

50

75

100

November

Size Class (mm)

02<

93- 03

95- 05

97- 07

99- 09

911- 011

931- 031

951- 051

971- 071

991- 091

912- 012

932- 032

952- 052972- 072982>

0

25

50

75

100
157

8 Settle, L. R., M. D. Greene, E. H. Laban, and M. Lacroix. Unpublished manuscript. Growth, condition, survival and spawning-date 

distribution of juvenile great barracuda, Sphryaena barracuda (Walbaum, 1792) in Florida Bay. NOAA, Center for Coastal Fisheries and 

Habitat Research, 101 Pivers Island Road, Beaufort, NC 28516.

Chapter 19. Family Sphyraenidae

Sphyraena barracuda (great barracuda)

Total numbers collected and rank of great barracuda by sampling method. There were a total of 39 species collected by 

otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 88 29

Life history Great barracuda range along the Atlantic 

coast from Massachusetts to Brazil, throughout the Gulf 

of Mexico and Caribbean Sea, West Indies, Bahamas, 

and Bermuda. It is a straggler north of South Carolina. 

Small individuals inhabit shallow waters, often forming schools, while adults are solitary and inhabit reefs 

(Breder, 1948; Vergara, 1978b). Great barracuda have 

been reported to spawn in the Miami, Florida, area 

from April through October (DeSylva, 1963). Based 

on hatchdate distributions from juveniles collected in 

Florida Bay, spawning occurs throughout the year with 

peaks in summer, but 90% of the spawning is complete 

by October. Spawning also occurs throughout the lunar 

year with peaks during the full and new months (Settle 

et al.8). Some males mature at 423 mm SL, with most 

mature at 460 mm SL (age-2). Females first mature at 

533 mm SL (age-3), but most do not mature until 606 

mm SL (age-4) (DeSylva, 1963). Eggs are pelagic, size 

at hatching is unknown, and transformation to the juvenile stage occurs at approximately 12 mm SL (Ditty 

et al., 1999). 

Diet Great barracuda are piscivorus. The diets of juveniles (64–322 mm SL) from Florida Bay mainly consisted 

of two cyprinodontids – goldspotted killifish (Floridicthys 

carpio) and rainwater killifish (Lucania parva) (Schmidt, 

1989). These two species are prevalent in seagrass habitat, which was where the great barracuda were collected. 

The diet of fish from mangrove prop root habitats might 

differ from those occupying seagrass habitats.

Fishes also dominated the diets of young (<276 mm 

SL) great barracuda collected in other Florida waters 

(off Miami), although the diet differed from fish from 

Florida Bay, with the majority of fishes being small 

pelagic schooling fishes, especially atherinids and clupeiforms. Young great barracuda collected from the 

Bahamas also show a preference for fish. Fish smaller 

than approximately 70 mm SL feed mainly on gobiids; 

whereas fish >70 mm SL feed mainly on pelagic schooling fishes. This could indicate a transition between habitats and/or prey availability. Smaller fish tend to inhabit 

detritus-rich shallow water where gobiids, cyprinodontids, and small gerreids are abundant. Older great barracuda inhabit deeper water where pelagic prey species 

(mainly belonids, carangids, mugilids, clupeiforms, and 

scombrids) are more common (DeSylva, 1963). 

Abundance and distribution Great barracuda were 

uncommon in otter trawl collections in basins (Figs. 

19.1 and 19.2). Spatially, there were no significant differences in densities. Temporally, there were no significant 

differences in mean densities among years, but there 

were among months. Densities increased steadily from 

March through November. 

Our estimates of densities in basins probably underestimated the abundance of this species in Florida Bay 

because juveniles are highly associated with mangrove 

prop root habitat (Thayer et al., 1987b; Ley and McIvor, 

2002). We believe that the low densities of fishes we 

observed in March and May can be attributed to recruitment and gear avoidance. Great barracuda recruit into 

Florida Bay at approximately 23–26 mm SL or age 30-d 

(Settle et al.8). As peak spawning occurs during summer, recruitment should begin to increase in July and, 

with the build-up of cohorts, reach maximum in fall. 

However, as the fish grow they avoid the gear, hence 

densities of juveniles would appear to decline during 

winter and spring. In addition, larger fish move into 

deeper waters and eventually move out of Florida Bay 

and on to reefs. 

Length-frequency distributions Great barracuda recruited into Florida Bay throughout the year (Fig. 19.3) 

and were first collected by our gear at approximately 

23–26 mm SL (30-d). Growth after ingress averages 2 

mm in SL d–1 for the first 60-d of residency in Florida 

Bay, then declines; fish 90-d post-ingress average 185 

mm SL, 120-d post-ingress average 210 mm SL (Settle 

et al.8). We observed two patterns of length-frequency 

distributions. From September through March a wider 

size range of fishes were collected that might be related 

to the availability of larger prey.
158

Figure 19.1

Mean densities (numbers 1000 m-2) of great barracuda (Sphyraena barracuda) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata and 

subdivisions see Fig. 2.1.
159

Year

5891- 48915991- 4991

6991

8991- 79910002- 9991

1002- 0002

ytisne D

0.00

0.05

0.10

0.15

0.20

0.25

0.30

Month

Jan Mar May Jul Sep Nov

ytisne D

0.00

0.05

0.10

0.15

0.20

0.25

0.30

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

A

B

C

Figure 19.2

Mean densities (numbers 1000 m–2

) of great barracuda 

(Sphyraena barracuda) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
160

Figure 19.3

Monthly length-frequency distributions of great barracuda (Sphyraena barracuda) from 

Florida Bay.

January

y

c

n

e

u

q

e

r

F

0

10

20

March

0

10

20

May

0

10

20

June

0

10

20

July

0

10

20

September

0

10

20

November

Size Class (mm)

92- 02

94- 0496- 0698- 08901- 001921- 021

941- 041

961- 061981- 081

902- 002

922- 022

942- 042962- 062982- 082

903- 003

0

10

20
161

Chapter 20. Family Scaridae

Nicholsina usta (emerald parrotfish)

Total numbers collected and rank of emerald parrotfish by 

sampling method. There were a total of 39 species collected 

by otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 55 39

Life history Emerald parrotfish range along the Atlantic from New Jersey to Brazil, Caribbean Sea, and 

in the West Indies only from Cuba and Hispaniola. 

It is probably uncommon north of South Carolina. 

The preferred habitat appears to be turtle grass beds 

(Breder, 1948; Randall, 1978). The eggs of other scarid 

species that have been studied are pelagic (Thresher, 

1991). 

Diet Based on the preferred habitat (Thalassia 

testudinum beds), dentition, and pharyngeal bone 

morphology, emerald parrotfish most probably feed 

on plant material. Unlike most other scarid species, 

that have fused teeth that form beak-like dental plates 

that allow them to graze on algae from corals; emerald 

parrotfish front teeth are fused basally, but the tips are 

separate and pointed (Randall, 1983). Their dentition, along with their pharyngeal bone morphology 

(pharyngeal mill), appears to be adapted to consume 

seagrasses. 

Abundance and distribution Emerald parrotfish were 

uncommon in otter trawl collections in basins (Figs. 20.1 

and 20.2) and were ranked lowest. Significant differences in mean densities were observed among years but not 

months. This species was collected in highest densities 

in recent collections, and despite being uncommon, it 

was collected in every bi-monthly sampling period (Figs. 

20.1 and 20.2). Spatially, differences among stratum and 

subdivision were observed. Emerald parrotfish were collected almost entirely in western Florida Bay. 

Tabb and Manning (1961) reported that this species 

was moderately common in western Florida Bay in clear 

waters of Thalassia flats. It was rarely collected in grassy 

shore habitat on the Atlantic side of the Florida Keys 

adjacent to Florida Bay (Springer and McErlean, 1962). 

This species has not been collected from Florida Bay 

mud banks (Sogard et al., 1987; 1989a).

Length-frequency distributions We were unable to 

discern any patterns in recruitment or growth of emerald parrotfish from length-frequency distributions (Fig. 

20.3). A relatively wide range of sizes was encountered 

during most months. Emerald parrotfish, which attain 

a maximum length of 300 mm (Robins and Ray, 1986), 

appeared to leave Florida Bay basins at >200 mm SL and 

move to deeper water.
162

Figure 20.1

Mean densities (numbers 1000 m–2

) of emerald parrotfish (Nicholsina usta) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
163

Year

5891- 48915991- 4991

6991

8991- 79910002- 9991

1002- 0002

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

0.6

Month

Jan Mar May Jul Sep Nov

ytisne D

0.00

0.05

0.10

0.15

0.20

0.25

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.00

0.20

0.40

0.60

0.80

1.00

1.20

1.40

1.60

A

B

C

Figure 20.2

Mean densities (numbers 1000 m–2

) of emerald 

parrotfish (Nicholsina usta) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
164

January

y

c

n

e

u

q

e

r

F

0

2

4

6

March

0

2

4

6

May

0

2

4

6

June

0

2

4

6

July

0

2

4

6

September

0

2

4

6

November

Size Class (mm)

02<93- 0395- 0597- 0799- 09911- 011

931- 031951- 051971- 071991- 091

0

2

4

6

Figure 20.3

Monthly length-frequency distributions of emerald parrotfish (Nicholsina usta) from Florida 

Bay.
165

Chapter 21. Family Callionymidae

Diplogrammus pauciradiatus (spotted dragonet)

Total numbers collected and rank of spotted dragonet by 

sampling method. There were a total of 39 species collected 

by otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 66 36

Life history Spotted dragonet range from North 

Carolina to Columbia and Bermuda. They inhabit seagrass beds in sub-tropical waters and the continental 

shelf (18–55 m) in temperate waters. Spawning occurs 

throughout the year in tropical waters and from spring 

through fall in temperate and sub-tropical waters (Powell and Greene, 2000). Based on larval collections, peak 

spawning in adjacent Biscayne Bay occurs in spring 

(Houde and Lovdal, 1984), whereas in Florida Bay peak 

spawning occurs from May through July and September 

(Powell, 2002). Males first mature at 12–15 mm SL, 

females at 11–12 mm SL. Eggs and larvae are pelagic. 

Spotted dragonet hatch at <1.2 mm NL and size at settlement is approximately 8 mm SL (Davis, 1966; Powell and 

Greene, 2000). This species, which is a resident species 

in Florida Bay, lives for only one year (Sogard, 1984). 

Diet Spotted dragonet in adjacent Biscayne Bay fed 

mainly on meiofauna. Harpacticoid copepods were 

the most important food item throughout ontogeny, 

followed by smaller amounts of ostracods, mites, nematodes, gastropods, and amphipods. They have small 

mouths and highly protrusible jaws that allow them to 

select small individual prey (Sogard, 1984).

Abundance and distribution Spotted dragonet were 

uncommon in otter trawl collections in basins (Figs. 

21.1 and 21.2). Temporally, there were only significant 

differences in mean densities among months. They were 

collected in greatest densities during summer and to a 

lesser degree in winter (Fig. 21.2). Spatially, there were 

differences among strata and subdivisions. Juveniles and 

adults of this species mainly were collected in the central 

and eastern section of Florida Bay (Figs. 21.1 and 21.2), 

mirroring the distribution of their larvae (Powell et al., 

1989; Powell, 2002).

The temporal differences we observed among months 

could be linked to differential spawning activity. Significant differences in larval densities among months and 

peak spawning periods have been observed (Powell, 

2002). Hence, high densities of spotted dragonet during 

certain periods could be a result of differential spawning activity.

On Florida Bay mud banks, this species was almost 

exclusively collected at Buchanan Keys (Fig. 1.1), a site 

with no freshwater input, minor salinity changes, and 

dense stands of Thalassia testudinum and green algae 

(Sogard et al., 1987). Although spotted dragonet were 

uncommon in our otter trawl collections, their larvae 

are a dominant component of the ichthyoplankton in 

Biscayne Bay and Florida Bay (Houde and Lovdal, 1984; 

Powell et al., 1989; Powell, 2002).

Length-frequency distributions Our collections 

were mainly comprised of adults (approximately >15 

mm SL), though juveniles were most likely extruded 

through our gear (Fig. 21.3). Fish >40–44 mm SL 

were rarely collected, which suggested that it attains a 

length of 40–44 mm SL after one year of growth and 

dies shortly thereafter. We observed modes at 20–

24 mm SL in four of seven months that probably 

have little biological meaning, but represent the size 

that spotted dragonet are fully recruited to the gear 

(Fig. 21.3).
166

Figure 21.1

Mean densities (numbers 1000 m–2

) of spotted dragonet (Diplogrammus pauciradiatus) 

from bi-monthly otter trawl collections in Florida Bay basins. Florida Bay is divided into 

three strata from 1984-1998 and six subdivisions from 1999–2001. For designations of 

strata and subdivisions see Fig. 2.1.
167

Year

5891- 4891

5991- 49916991

8991- 7991

0002- 99911002- 0002

ytisne D

0.00

0.05

0.10

0.15

0.20

0.25

Month

Jan Mar May Jul Sep Nov

mytisne D 2)

0.00

0.05

0.10

0.15

0.20

0.25

0.30

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.00

0.10

0.20

0.30

0.40

0.50

A

B

C

Figure 21.2

Mean densities (numbers 1000 m–2

) of spotted 

dragonet (Diplogrammus pauciradiatus) from bi-monthly 

otter trawl collections in Florida Bay basins by (A) year, 

(B) month, and (C) stratum/subdivision. Florida Bay 

is divided into three strata from 1984–1998 and six 

subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
168

Figure 21.3

Monthly length-frequency distributions of spotted dragonet (Diplogrammus pauciradiatus) 

from Florida Bay.

January

y

c

n

e

u

q

e

r

F

0

5

10

15

March

0

5

10

15

May

0

5

10

15

June

0

5

10

15

July

0

5

10

15

September

0

5

10

15

November

Size Class (mm)

01<91- 5192- 52

93- 53

94- 5495- 55

0

5

10

15
169

Chapter 22. Family Gobiidae

Gobiosoma robustum (code goby)

Total numbers collected and rank of code goby by sampling 

method. There were a total of 39 species collected by otter 

trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 313 16

Life history Code goby range from northeast Florida 

to the Florida Keys and throughout the Gulf of Mexico. 

They are generally found in seagrass beds and algae 

mats in shallow protected fully saline waters (Robins 

and Ray, 1986). Code gobies spawn throughout the year 

in Gulf of Mexico waters. Eggs are demersal and attach 

by adhesive threads to shells or sponges, and larvae are 

pelagic. The juvenile stage is reached at 6–8 mm SL and 

males attain sexual maturity at 16 mm SL, females at 

16–22 mm SL (Breder, 1942; Pattillo et al., 1997). 

Diet Seasonal variation was observed in the diets of 

fish 11–20 mm SL from Texas waters that were linked 

to prey availability (Huh and Kitting, 1985). During 

spring and fall, amphipods and polychaetes were dominant diet items, whereas during winter copepods were 

heavily preyed on. Fish >20 mm SL had a consistent 

diet throughout the year regardless of prey availability. Amphipods were the dominant prey for this size 

group (Huh and Kitting, 1985). In the Whitewater Bay 

system (North River), Odum and Heald (1972) found 

little seasonal changes in the diets of fish 15–35 mm 

SL. Fish consumed mainly amphipods with smaller 

quantities of chironomid larvae, mysids, and cladocerans. Smaller fish (7–15 mm SL) fed on harpacticoid 

copepods, juvenile mysids, cumaceans, and many pennate diatoms. 

Abundance and distribution Code goby were common in otter trawl collections in basins (Figs. 22.1 and 

22.2). There were no significant differences in mean 

densities among months or years, but there were differences in mean densities among strata and subdivisions. 

This species was most abundant in central and western 

Florida Bay (Figs. 22.1 and 22.2). 

Sogard et al. (1987) collected the greatest number 

of the code goby over a mudbank near Buchanan Keys 

(Fig. 1.1), located in our Atlantic subdivision where we 

encountered the lowest densities. Their mudbank station was characterized by lush T. testudinum stands and 

high coverage of green algae, which suggested that in 

areas where overall seagrass densities are relatively low, 

the code goby will exploit locally lush seagrass stands 

(Schofield, 2003). This species also has an affinity for 

mangrove prop root habitat (Thayer et al., 1987b). 

Code goby have been observed to occur at low densities 

when prey densities are low (Huh and Kitting, 1985).

Length-frequency distributions Length-frequency 

distributions were uni-modal and monthly distributions 

were quite similar, which suggested that recruitment occurred throughout the year (Fig. 22.3). Both juveniles 

(approximately <20 mm SL) and adults were collected, 

but we did not collect large adults that attain a size of 

75 mm TL (Robins and Ray, 1986).
170

Figure 22.1

Mean densities (numbers 1000 m–2

) of code goby (Gobiosoma robustum) from bi-monthly 

otter trawl collections in Florida Bay basins. Florida Bay is divided into three strata 

from 1984–1998 and six subdivisions from 1999–2001. For designations of strata and 

subdivisions see Fig. 2.1.
171

Year

5891- 48915991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

3.0

A

B

C

Figure 22.2

Mean densities (numbers 1000 m–2

) of code goby 

(Gobiosoma robustum) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
172

January

y

c

n

e

u

q

e

r

F

0

50

100

150

March

0

50

100

150

May

0

50

100

150

June

0

50

100

150

July

0

50

100

150

September

0

50

100

150

November

Size Class (mm)

01<91- 51

92- 5293- 5394- 54

0

50

100

150

Figure 22.3

Monthly length-frequency distributions of code goby (Gobiosoma robustum) from Florida Bay.
173

Microgobius gulosus (clown goby) 

Total numbers collected and rank of clown goby by sampling 

method. There were a total of 39 species collected by otter 

trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 307 17

Life history The clown goby ranges along the Atlantic coast from Maryland to south Florida, around the 

tip of Florida, and throughout the northern Gulf of 

Mexico to Texas, where it occurs in muddy waters in 

variable salinities (Robins and Ray, 1986). In estuaries 

near Cape Canaveral, spawning, based on gonosomatic 

indices, was reported to begin in mid-spring, decline 

and remain low through fall and winter, then increase 

in intensity in late winter (Provancha and Hall, 1991). 

Both Reid (1954) and Kilby (1955) reported year round 

spawning in northwestern Florida estuarine waters. Eggs 

are demersal (Thresher, 1991) and larvae are pelagic. 

Females appear to mature at 35 mm TL (Provancha and 

Hall, 1991). 

Diet Clown goby are opportunistic feeders. In eastern 

Florida estuaries, clown goby diets vary by location. 

Between two sites the diet is dominated either by crustaceans (tanaids, amphipods, decapods, and cumaceans) 

or annelids (Provancha and Hall, 1991). Reid (1954) 

examined clown goby diet (15–57 mm SL) from 

Cedar Key, Florida, and found they mainly consume 

copepods, mysids, and amphipods. In the Whitewater 

Bay system (North River), Odum and Heald (1972) 

reported that the diet of the clown goby is similar to 

that of the code goby. They consume mainly amphipods 

followed by hapacticoid copepods, chironomid larvae, 

and cumaceans; cladocerans, algal strands, plant detritus, and mysids make up a smaller percentage. Clown 

gobies (11–40 mm SL) from Crystal River, Florida, are 

considered to be omnivores, but all size classes consume a considerable amount of detritus. Fish in the 

size range of 11–25 mm SL consume mainly detritus 

and copepods; those 26–40 mm SL shift to a diet of 

primarily detritus, amphipods, and polychaetes. Sand 

is present in the stomachs of all size classes, indicating 

juveniles of this species feed directly on the bottom 

(Carr and Adams, 1973).

Abundance and distribution Clown goby were common in otter trawl collections (Figs. 22.4 and 22.5). 

Temporally, there were significant differences in mean 

densities among years; this was most probably influenced 

by change in sampling design, since prior to 1999 sampling rarely occurred in north-central and northeastern 

Florida Bay (Fig. 22.4). Significant differences among 

months were also observed. Highest densities were observed in July (Fig. 22.5), which could be influenced by 

recruitment patterns (Fig. 22.6). Spatially, there were 

significant differences in mean densities. This species 

was markedly more abundant in north-central and 

northeastern Florida Bay (Figs. 22.4 and 22.5). 

Provancha and Hall (1991) also observed variation 

in clown goby mean monthly densities and suggested 

that differences in densities could also be influenced 

by recruitment patterns. However, their observations of 

clown goby abundance patterns at Cape Canaveral, Florida, differed from what we observed in Florida Bay. 

Our results are in concordance with those of Sogard 

et al. (1987) who found highest densities of this species on mud banks in sparsely vegetated northeastern 

Florida Bay. Both the code goby (see above) and the 

clown goby are ecologically similar (Schofield, 2003); 

however, their relative densities are dissimilar in the 

Bay. Their distributions overlap somewhat, but the code 

goby is most prevalent in western and central Florida 

Bay where seagrass densities are relatively lush, while 

the clown goby inhabits sparsely vegetated sites. Clown 

goby prefers muddy sediments and quiet waters where 

it burrows in the bottom (Birdsong, 1981). When paired 

with the code goby in laboratory experiments, the clown 

goby is pushed out of a structured habitat that is the preferred habitat of the code goby (Schofield, 2003). Based 

on these results, Schofield (2003) concluded that when 

these species occur sympatrically, competition appears 

to modify their habitat selection. 

Length-frequency distributions The majority of clown 

goby were young-of-the-year (30–34 mm SL) based on 

estimates by Provancha and Hall (1991; 35–40 mm TL 

at end of first year), but few were ripe adults (>35 mm 

TL; ca. 30 mm SL) (Fig. 22.6). Spawning appeared to 

occur throughout the year.
174

Figure 22.4

Mean densities (numbers 1000 m–2

) of clown goby (Microgobius gulosus) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
175

Year

5891- 48915991- 49916991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

1.4

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

1.4

1.6

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

A

B

C

Figure 22.5

Mean densities (numbers 1000 m–2

) of clown goby 

(Microgobius gulosus) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
176

January

y

c

n

e

u

q

e

r

F

0

10

20

30

40

March

0

10

20

30

40

May

0

10

20

30

40

June

0

10

20

30

40

July

0

10

20

30

40

September

0

10

20

30

40

November

Size Class (mm)

01<

91- 5192- 52

93- 5394- 54

95- 55

0

10

20

30

40

Figure 22.6

Monthly length-frequency distributions of clown goby (Microgobius gulosus) from Florida 

Bay.
177

Microgobius microlepis (banner goby)

Total numbers collected and rank of banner goby by sampling 

method. There were a total of 39 species collected by otter 

trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 156 22

Life history Banner goby range from south Florida to 

Yucatan and Belize, and inhabit shallow water muddy 

bottom where they burrow, at least in the mating season 

(Longley and Hildebrand, 1940; Robins and Ray, 1986). 

Spawning in the Dry Tortugas and Florida Keys occurs 

throughout summer and might occur in burrows, but 

no eggs have been found in the burrows examined 

(Longley and Hildebrand, 1940; Birdsong, 1981). Eggs 

are demersal (Thresher, 1991) and larvae are pelagic. 

Diet Based on the examination of a few specimens, 

this species feeds on copepods and ostracods (Longley 

and Hildebrand, 1940).

Abundance and distribution Banner goby were common in otter trawl collections in basins (Figs. 22.7 and 

22.8). Mean densities differed among years, but not 

months. Significant differences among years could 

partially be explained by increased sampling in eastern Florida Bay, although densities in 2000–2001 were 

comparable to those of 1996 when sampling was not as 

extensive in eastern Florida Bay (Figs. 22.7 and 22.8). 

There were significant differences in densities among 

strata and subdivisions. Banner goby were most abundant in the Eastern subdivision, where seagrass shortshoot densities are low and sediments shallow (Thayer 

and Chester, 1989).

This species was not collected in northeastern Florida 

Bay where fluctuations in freshwater inflow cause wide 

fluctuations in salinity (Schmidt, 1979), and rarely in 

western Florida Bay (excluding the Gulf subdivision) 

where lush seagrass beds occur (Thayer and Chester, 

1989). Banner goby have not been collected on mud 

banks in Florida Bay (Sogard et al., 1987; Sheridan et 

al., 1997). This species was not collected in a grassy 

shore habitat on the Atlantic side of the Florida Keys 

adjacent to Florida Bay (Springer and McErlean, 1962) 

or in early studies in northwestern Florida Bay (Tabb 

and Manning, 1961). In the Florida Keys, abundances 

of banner gobies vary considerably among years in any 

given locality (Birdsong, 1981).

Length-frequency distributions Banner goby have a 

protracted spawning period based on length-frequency 

distributions, but spawning might be discontinuous 

throughout the year (Fig. 22.9). The absence of small 

fish (<15 mm SL) in March and June suggested that 

spawning was minimal or did not occur during certain 

months. However, the absence and relatively small numbers of fish <15 mm SL could mean that this size class 

was not vulnerable to our gear. Without growth data it 

would be difficult to link monthly cohorts with the size 

class distributions to determine the months when fish 

<15 mm SL were spawned.
178

Figure 22.7

Mean densities (numbers 1000 m–2

) of banner goby (Microgobius microlepis) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
179

Year

5891- 48915991- 4991

6991

8991- 79910002- 99911002- 0002

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

1.2

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

0.5

1.0

1.5

2.0

A

B

C

Figure 22.8

Mean densities (numbers 1000 m–2

) of banner goby 

(Microgobius microlepis) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
180

Figure 22.9

Monthly length-frequency distributions of banner goby (Microgobius microlepis) from Florida 

Bay.

January

y

c

n

e

u

q

e

r

F

0

30

60

March

0

30

60

May

0

30

60

June

0

30

60

July

0

30

60

September

0

30

60

November

Size Class (mm)

01<

91- 51

92- 52

93- 53

0

30

60
181

Chapter 23. Family Monacanthidae

Monacanthus ciliatus (fringed filefish)

Abundance (numbers) and rank of fringed filefish by sampling method. There were a total of 39 species collected by 

otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 627 14

Life history Fringed filefish range along the Atlantic 

coast from Newfoundland to Argentina and throughout the Gulf of Mexico, Caribbean, Bahamas, and Bermuda. They occur in shallow water to 50 m over sandy 

and rocky bottoms, but mostly in seagrass beds (Tyler, 

1978a). Based on the rarity of larvae of this family, 

fringed filefish probably spawn outside of Florida Bay 

(Powell et al., 1989; Powell, 2002). The eggs are demersal and termed “balistid type” in that eggs are tended 

for a short period of time, if at all, before they hatch 

rapidly into yolk-sac larvae, similar to larvae hatched 

from pelagic eggs. The larvae from “balistid type” eggs 

generally remain in the plankton longer than demersal 

spawners, which implies extensive dispersal (Thresher, 

1991).

Diet Young fish (39–55 mm SL) feed primarily on 

copepods and mollusks, but amphipods and ostracods 

are also consumed (Reid, 1954).

Abundance and distribution Fringed filefish were 

common in otter trawl collections in basins (Figs. 23.1 

and 23.2). Densities did not differ significantly among 

months or years, but differed among strata and subdivisions. Although our monthly densities were marginally 

non-significant (P = 0.07), we observed highest densities 

in September, lowest in March. There were significant 

differences in densities among strata and subdivisions. 

Fringed filefish is most abundant in western Florida Bay, 

which could be attributed to lush seagrass beds (Thayer 

and Chester, 1989) and more intense recruitment from 

Gulf of Mexico waters (Figs. 23.1 and 23.2). 

Other studies in Florida Bay indicate that this species 

is common in summer, fall, and early winter, uncommon to rare from January through April, and generally 

absent in May (Schmidt, 1979; FMRI1,2,3,4). In grassy 

shore habitats on the Atlantic side of the Florida Keys 

adjacent to Florida Bay, Springer and McErlean (1962) 

found this species to be more abundant from October 

through December and rare to absent from January 

through August. From the years 1994 through 1997, 

Florida Marine Institute staff reported a steady decline 

in abundance (FMRI1,2,3,4). similar to what we observed 

(Fig. 23.2). 

Tabb and Manning (1961) found the fringed filefish 

to be the most common filefish in western Florida Bay 

in higher salinity waters and associated with Thalassia 

testudinum. There appears to be only minor recruitment from the Florida Keys, although this species is 

common in grassy shore habitats on the Atlantic side of 

the Florida Keys adjacent to Florida Bay (Springer and 

McErlean, 1962).

Length-frequency distributions Based on specimens 

<20 mm SL, fringed filefish appeared to recruit during 

most months (Fig. 23.3). Fringed filefish reach a length 

of 200 mm TL (Robins and Ray, 1986) but our collections are comprised of much smaller fish. Although we 

are unaware of the size at sexual maturity of this species, 

Clark (1950) identified adult males at 58 mm SL and 

females at 43 mm SL. The majority of our fish could be 

juveniles and young adults, as could those collected by 

Springer and McErlean (1962) in grassy shore habitats 

on the Atlantic side of the Florida Keys adjacent to 

Florida Bay. Speculatively, Florida Bay provides nursery 

habitat for fringed filefish as well as do grassy shore 

habitats of the Florida Keys.
182

Figure 23.1

Mean densities (numbers 1000 m–2

) of fringed filefish (Monacanthus ciliatus) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
183

Year

5891- 4891

5991- 4991

6991

8991- 79910002- 9991

1002- 0002

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

3.0

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

12.0

14.0

A

B

C

Figure 23.2

Mean densities (numbers 1000 m–2

) of fringed filefish 

(Monacanthus ciliatus) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
184

January

y

c

n

e

u

q

e

r

F

0

20

40

60

80

March

0

20

40

60

80

May

0

20

40

60

80

June

0

20

40

60

80

July

0

20

40

60

80

September

0

20

40

60

80

November

Size Class (mm)

41- 0142- 02

43- 0344- 04

45- 05

46- 06

47- 07

48- 08

49- 09

0

20

40

60

80

Figure 23.3

Monthly length-frequency distributions of fringed filefish (Monacanthus ciliatus) from Florida 

Bay.
185

M. hispidus (planehead filefish)

Abundance (numbers) and rank of planehead filefish by 

sampling method. There were a total of 39 species collected 

by otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 179 21

Life history Planehead filefish range from Nova Scotia to Brazil, northern Gulf of Mexico, and Bermuda. 

They are absent in the Bahamas (Robins and Ray, 

1986). Spawning is prolonged and most likely occurs 

throughout the year south of Cape Hatteras (Fahay, 

1983). Based on the rarity of larvae of this family, 

planehead filefish probably spawn outside of Florida 

Bay (Powell et al., 1989; Powell, 2002); however, Matheson et al. (1999) list this as a resident species in Florida 

Bay. Pelagic eggs have been reported (Martin and Drewry, 1978), but are most likely the “balistid type” in that 

eggs are demersal and are tended for a short period of 

time, if at all, before they hatch rapidly into yolk-sac 

larvae, similar to larvae hatched from pelagic eggs. 

The larvae from “balistid type” eggs generally remain 

in the plankton longer than demersal spawners, which 

implies extensive dispersal (Thresher, 1991). Young 

planehead filefish larvae occur near the bottom, 

while the juvenile stage is associated with sargassum 

weed. The transition from larvae to juveniles occurs 

at approximately 8 mm SL (Fahay, 1983; Rogers et al., 

2001). Females at 81 mm SL have been observed with 

large eggs, maturing males exhibited secondary sexual 

characters at 104 mm SL (Berry and Vogele, 1961).

Diet Planehead filefish, 15–82 mm SL, collected at Cedar Key, Florida, had a diet similar to the fringed filefish 

(see above). They consumed mainly amphipods, with 

fewer amounts of pelecypod mollusks and polychaetes 

(Reid, 1954).

Abundance and distribution Planehead filefish were 

common in otter trawl collections in basins, but less 

abundant than its congener, the fringed filefish (Figs. 

23.1, 23.2, 23.4, and 23.5). Densities differed significantly among years, but not months, although monthly 

differences were marginally non-significant (P = 0.09). 

Planehead filefish were considerably more abundant 

in recent years, and more abundant in January (Fig. 

23.5). Densities differed among strata and subdivisions. 

Like the fringed filefish, the planehead filefish was most 

abundant in western Florida Bay, which could be attributed to lush seagrass beds (Thayer and Chester, 1989) 

and more intense recruitment from Gulf of Mexico 

waters (Figs. 23.1 and 23.2). 

This species might also recruit into the Atlantic 

subdivision from the Florida Keys, as it is abundant in 

grassy shore habitats on the Atlantic side of the Florida 

Keys adjacent to Florida Bay; but as in Florida Bay, not 

as abundant as the fringed filefish (Springer and McErlean, 1962).

Seasonally, Springer and McErlean (1962) observed 

the opposite. They collected relatively few fish in January compared to November, February, and March, and 

Schmidt (1979) found them to be most abundant in 

Florida Bay in summer and least in fall. 

Length-frequency distributions Planehead filefish recruited into Florida Bay throughout the year (Fig.23.6). 

Their length-frequency pattern differs from the fringed 

filefish (Fig. 23.3) in that smaller planehead filefish 

were more dominant. A relatively narrow range of sizes 

was observed in January and July, while a wider range 

was observed in fall collections that could be related 

to the intensity of recruitment (Fig. 23.6). Rogers et 

al (2001) reported that maximum growth of juveniles 

approximately 8–30 mm SL was 0.8 mm day–1. Based on 

modal sizes (Fig. 23.6) during the period July (30–34 

mm SL), September (40–44 mm SL), and November 

(55–59 mm SL), juveniles grew approximately 0.2 mm 

SL day–1. 
186

Figure 23.4

Mean densities (numbers 1000 m–2

) of planehead filefish (Monacanthus hispidus) from 

bi-monthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
187

Year

5891- 48915991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

0.2

0.4

0.6

0.8

1.0

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.2

0.4

0.6

0.8

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

2.0

4.0

6.0

8.0

10.0

12.0

14.0

A

B

C

Figure 23.5

Mean densities (numbers 1000 m–2

) of planehead 

filefish (Monacanthus hispidus) from bi-monthly otter 

trawl collections in Florida Bay basins by (A) year, 

(B) month, and (C) stratum/subdivision. Florida Bay 

is divided into three strata from 1984–1998 and six 

subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
188

Figure 23.6

Monthly length-frequency distributions of planehead filefish (Monacanthus hispidus) from 

Florida Bay.

January

y

c

n

e

u

q

e

r

F

0

10

20

30

March

0

10

20

30

May

0

10

20

30

June

0

10

20

30

July

0

10

20

30

September

0

10

20

30

November

Size Class (mm)

41- 0142- 02

43- 03

44- 04

45- 05

46- 06

47- 0748- 0849- 09

401- 001

411- 011421- 021

431- 031

0

10

20

30
189

Chapter 24. Family Ostraciidae

Lactophrys quadricornis (scrawled cowfish)

Abundance (numbers) and rank of scrawled cowfish by sampling method. There were a total of 39 species collected by 

otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 145 23

Life history Scrawled cowfish range from Massachusetts to Brazil, Bermuda, Gulf of Mexico, and the Caribbean, and generally inhabit seagrass beds. It is a straggler north of the Carolinas (Breder, 1948; Tyler, 1978b). 

In the Caribbean, scrawled cowfish spawn throughout 

the year with maximum spawning occurring in May or 

earlier (Munro et al., 1973). Spawning occurs in bays, 

nearshore, and offshore (Martin and Drewry, 1978). 

Eggs have been collected in adjacent Biscayne Bay 

(Palko and Richards, 1969), but based on ichthyoplankton surveys there is no solid evidence that this species 

spawns in Florida Bay (Powell et al., 1989; Powell, 2002). 

Larval scrawled cowfish have been collected in Gulf of 

Mexico inshore waters off southwestern Florida, but not 

in estuarine waters (Collins and Finucane, 1984). Eggs 

and larvae are pelagic. Larvae hatch at 6 mm NL, but 

shrink during the preflexion stage to at least 2.5 mm NL 

(Palko and Richards, 1969; Aboussouan and Leis, 1984). 

The juvenile stage is attained at 10 mm TL, and size at 

sexual maturity is unknown (Martin and Drewry, 1978). 

Diet Reid (1954) reported scrawled cowfish (13–65 

mm SL) consume undetermined vegetation, algae, and 

pelecypods. This species (size unknown) also feeds on 

tunicates, gorgonians, sponges, and slow-moving crustaceans (Tyler, 1978b).

Abundance and distribution Scrawled cowfish were 

common in otter trawl collections in basins (Figs. 24.1 

and 24.2). Temporally, there were significant differences in mean densities among years, but not months. 

There was a decline from 1984–1985 to 1994–1995, 

the latter date following the die-off of seagrasses and 

increased turbidity. Densities increased from 1994–1995 

through 2000–2001 (Fig. 24.2). Spatially, significant 

differences in scrawled cowfish densities were observed 

among strata and subdivisions. This species was most 

abundant in western Florida Bay (Figs. 24.1 and 24.2), 

where seagrass standing crop and diversity is greatest 

(Thayer and Chester, 1989). Moderate densities were 

observed in waters adjacent to the Keys (Atlantic subdivision) suggesting maximum recruitment from Gulf 

of Mexico waters and moderate recruitment from the 

Straits of Florida. 

Scrawled cowfish are common in western Florida Bay 

Thalassia beds, prefer clear water and moderately high 

salinities (Springer and Woodburn, 1960; Tabb and 

Manning, 1961), and could be a good indicator of seagrass health and water quality. We did not observe any 

differences in densities among months. However, other 

workers in Florida Bay reported that it was most abundant in winter when western Florida Bay water tends 

to be less turbid (Tabb and Manning, 1961), or most 

abundant in summer and fall (Schmidt, 1979). 

Although we observed moderate densities in waters 

adjacent to the Keys, and suggested moderate recruitment from this area, this species was found to be 

uncommon in the Atlantic Ocean side of the Florida 

Keys adjacent to Florida Bay (Springer and McErlean, 

1962). This species was rarely collected on Florida Bay 

mud banks (Sogard et al., 1987; 1989a), and absent in 

mangroves (Thayer et al., 1987b). 

Length-frequency distributions Recruitment into 

Florida Bay by scrawled cowfish is protracted, although 

it appeared that recruitment did not occur in late spring 

and early summer (Fig. 24.3). Based on length-frequency distributions, spawning of scrawled cowfish recruiting 

into Florida Bay appeared to be more protracted than 

in Tampa Bay where this species has been reported to 

spawn from June–September (Springer and Woodburn, 

1960). We collected a wide range of sizes that were similar to those reported from Tampa Bay (Springer and 

Woodburn, 1960), but differed from that reported in 

western Florida Bay by Tabb and Manning (1961). The 

latter investigators did not collect any scrawled cowfish 

>120 mm SL, suggesting to them that only young cowfish utilize Florida Bay and spawning does not occur in 

western Florida Bay.
190

Figure 24.1

Mean densities (numbers 1000 m–2

) of scrawled cowfish (Lachtophrys quadricornis) from 

bi-monthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
191

Year

5891- 48915991- 4991

69918991- 79910002- 9991

1002- 0002

ytisne D

0.0

0.2

0.4

0.6

0.8

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.1

0.2

0.3

0.4

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

0.5

1.0

1.5

2.0

2.5

3.0

A

B

C

Figure 24.2

Mean densities (numbers 1000 m–2

) of scrawled cowfish 

(Lachtophrys quadricornis) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
192

Figure 24.3

Monthly length-frequency distributions of scrawled cowfish (Lachtophrys quadricornis) from 

Florida Bay.

January

y

c

n

e

u

q

e

r

F

0

5

10

15

March

0

5

10

15

May

0

5

10

15

June

0

5

10

15

July

0

5

10

15

September

0

5

10

15

November

Size Class (mm)

02<

93- 0395- 05

97- 0799- 09

911- 011931- 031

951- 051

971- 071

991- 091

0

5

10

15
193

Chapter 25. Family Tetraodontidae

Sphoeroides nephelus ( southern puffer)

Abundance (numbers) and rank of southern puffer by sampling method. There were a total of 39 species collected by 

otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 55 39

Life history Southern puffer range along the Atlantic coast from northeast Florida to southern Florida, 

throughout the Caribbean to Martinique, the Bahamas, 

in the eastern Gulf of Mexico from the tip of Florida to 

western Florida, and in the Yucatan. It is rare along the 

Central American coast and is a straggler on the Atlantic 

coast to North Carolina (Shipp, 1974). At Cedar Key, 

Florida, Reid (1954) collected ripe males in October 

(189 mm SL) and March (155 mm SL). Young (approximately 15 mm SL) were collected in June and October, 

which led Reid to believe that spawning occurs in spring 

and summer. Eggs are “balistid type” in that they are 

demersal and are tended for a short period of time, if at 

all, before they hatch rapidly into yolk-sac larvae, similar 

to larvae hatched from pelagic eggs (Thresher, 1991). 

Larvae are pelagic (Leis, 1984). 

Diet Small southern puffer (6–25 mm SL) are detritivores and feed directly on the bottom (Carr and Adams, 

1973). Reid (1954) reported that diets of southern 

puffer (no sizes given) from Cedar Key, Florida, consists 

of crabs and mollusks.

Abundance and distribution Southern puffer were 

common in otter trawl collections in basins (Figs. 25.1 

and 25.2). Temporally, mean densities differed significantly among years and months. There was a marked 

decline in densities between 1984–1985 and 1994–1995, 

following the die-off of seagrass. From 1996 to 2001, densities were similar, but not as high as 1984–1985. Southern puffer appeared to be most abundant from January 

through March, which might be related to recruitment 

into the Bay (Fig. 25.2). Spatially, mean densities differed significantly among strata and subdivision. Highest densities occurred in western Florida Bay. Southern 

puffer appeared to have an unusual distribution in that 

they were also abundant in northeastern Florida Bay 

(Figs. 25.1 and 25.2). However, densities of fish in the 

northeast were biased by one collection where relatively 

high densities were observed (Fig. 25.1). 

In western Florida Bay, Tabb and Manning (1961) 

observed that larger southern puffer moved offshore, 

whereas young (20–30 mm SL) were common in tidal 

channels. Also in Florida Bay, Schmidt (1979) collected 

greater numbers in winter and spring and lesser in 

summer and fall, which is similar to our findings. This 

species was rarely collected on Florida Bay mud banks 

(Sogard et al., 1987; 1989a) and was absent in mangrove 

prop root habitats in the Bay (Thayer et al., 1987b). 

Southern puffer was uncommon in a grassy shore habitat on the Atlantic side of the Florida Keys adjacent to 

Florida Bay (Springer and McErlean, 1962).

Length-frequency distributions Small southern puffer occurred throughout the year, indicating protracted 

recruitment into the Bay (Fig. 25.3). Smaller fish, most 

likely juveniles, dominated collections. In concordance 

with Tabb and Manning (1961; see above) and based on 

January frequencies, larger fish appeared to leave the 

Bay during early winter.
194

Figure 25.1

Mean densities (numbers 1000 m–2

) of southern puffer (Sphoeroides nephalus) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
195

Year

5891- 4891

5991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.00

0.04

0.08

0.12

0.16

0.20

Month

Jan Mar May Jul Sep Nov

ytisne D

0.00

0.05

0.10

0.15

0.20

0.25

0.30

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.00

0.10

0.20

0.30

0.40

0.50

A

B

C

Figure 25.2

Mean densities (numbers 1000 m–2

) of southern puffer 

(Sphoeroides nephalus) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
196

Figure 25.3

Monthly length-frequency distributions of southern puffer (Sphoeroides nephalus) from Florida 

Bay.

January

y

c

n

e

u

q

e

r

F

0

5

10

15

March

0

5

10

15

May

0

5

10

15

June

0

5

10

15

July

0

5

10

15

September

0

5

10

15

November

Size Class (mm)

02<

93- 03

95- 05

97- 07

99- 09

911- 011931- 031951- 051

971- 071

991- 091

912- 012

0

5

10

15
197

Chapter 26. Family Diodontidae

Chilomycterus schoepfi (striped burrfish)

Abundance (numbers) and rank of striped burrfish by sampling method. There were a total of 39 species collected by 

otter trawl.

Gear Habitat Year Number Rank

Otter trawl Basins 1984–2001 185 20

Life history Striped burrfish range along the Atlantic 

coast from North Carolina (straying to Maine and Nova 

Scotia) to Brazil, the Bahamas (rare), and northern 

Gulf of Mexico, where it is common. It is absent from 

most of the West Indies. It is very common in seagrass 

beds (Robins and Ray, 1986). There is little information 

available on the reproductive habits of striped burrfish. 

Spawning might take place in spring in Gulf of Mexico 

waters off west Florida (Reid, 1954; Springer and Woodburn, 1960). Eggs and larvae are pelagic (Leis, 1984; 

Thresher, 1991). 

Diet The diet of fish 126–215 mm SL from Tampa 

Bay, Florida, consisted of large gastropods, bivalves, and 

barnacles that are crushed by bony jaw plates (Bigelow 

and Schroeder, 1953; Motta et al., 1995).

Abundance and distribution Striped burrfish were 

common in otter trawl collections in basins (Figs. 26.1 

and 26.2). Temporally, there were no significant differences in mean densities among years or months, 

although there was a steady increase in densities from 

1984–1985 to 2000–2001 (Fig. 26.2). Spatially, we observed significant differences in mean densities among 

strata and subdivisions. This species is most abundant 

in western Florida Bay and least abundant in eastern 

Florida Bay (Figs. 26.1 and 26.2).

In Cedar Key, Florida, Reid (1954) collected this 

species mainly from May through November. In 

Tampa Bay, Florida, Springer and Woodburn (1960) 

collected this species mainly in the fall and less frequently during winter. Tabb and Manning (1961) 

also found the striped burrfish to be abundant in 

western Florida Bay in seagrass beds and shallow 

clear water. This species was rarely collected in a 

grassy shore habitat on the Atlantic Ocean side of 

the Florida Keys adjacent to Florida Bay (Springer 

and McErlean, 1962), absent from the mangrove 

prop root habitat (Thayer et al., 1987b), and rare on 

Florida Bay mud banks (Sogard et al., 1987; 1989a).

Length-frequency distributions Striped burrfish 

appeared to recruit into Florida Bay throughout the 

entire year (Fig. 26.3), suggesting that this species 

spawns throughout the year. Based on distributions 

in May and June, spawning appeared to be more 

intense in spring, which is in concordance with findings of Springer and Woodburn (1960) in Tampa Bay, 

Florida.
198

Figure 26.1

Mean densities (numbers 1000 m–2

) of striped burrfish (Chilomycterus shoepfi) from bimonthly otter trawl collections in Florida Bay basins. Florida Bay is divided into three 

strata from 1984–1998 and six subdivisions from 1999–2001. For designations of strata 

and subdivisions see Fig. 2.1.
199

Year

5891- 48915991- 4991

6991

8991- 7991

0002- 9991

1002- 0002

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

0.6

Month

Jan Mar May Jul Sep Nov

ytisne D

0.0

0.1

0.2

0.3

0.4

0.5

_________ ____________________

 Stratum Subdivision

tsa E

l art ne C

tse W

nr etsa E

nrehtr o N

cit nalt A

l art ne C

nr etse W

fl u G

ytisne D

0.0

1.0

2.0

3.0

4.0

A

B

C

Figure 26.2

Mean densities (numbers 1000 m–2

) of striped burrfish 

(Chilomycterus shoepfi) from bi-monthly otter trawl 

collections in Florida Bay basins by (A) year, (B) month, 

and (C) stratum/subdivision. Florida Bay is divided into 

three strata from 1984–1998 and six subdivisions from 

1999–2001. For designations of strata and subdivisions 

see Fig. 2.1.
200

January

y

c

n

e

u

q

e

r

F

0

5

10

15

March

0

5

10

15

May

0

5

10

15

June

0

5

10

15

July

0

5

10

15

September

0

5

10

15

November

Size Class (mm)

02<

93- 03

95- 0597- 07

99- 09

911- 011

931- 031

951- 051

971- 071

991- 091

912- 012

932- 032

0

5

10

15

Figure 26.3

Monthly length-frequency distributions of striped burrfish (Chilomycterus shoepfi) from Florida 

Bay.
201

Overview

We have presented broad patterns in the abundance, 

distribution, and recruitment of 46 juvenile and small 

resident fish species from Florida Bay, along with brief 

information on life histories and diets. Approximately 

56% of the fish species had tropical affinities (Antillean fauna), whereas 76% had warm temperate affinities (Gulf of Mexico and Carolinean fauna) (Table 2). 

Twenty-four percent were solely tropical, whereas 43% 

were solely warm temperate. Tabb and Roessler (1989), 

in a review of juvenile fish studies in Florida Bay and 

adjacent waters, concluded that the fish fauna in northcentral to northwestern Florida Bay, as compared to 

Alligator Reef (adjacent to Florida Bay in the Straits 

of Florida), was more warm temperate than tropical. 

Holmquist et al. (1989a, 1989b) also noted that the 

crustacean fauna on Florida Bay mud banks appeared 

to be more temperate than tropical.

There were approximately equal numbers of resident and transient fish species collected in Florida Bay 

(Table 2). The pelagic species fell into two discrete 

resident and transient fish groups. Pelagic transient 

fishes included all the clupeid species. Pelagic resident 

fishes included the engraulids (all species), exocoetids 

and belonids, and the atherinids (all species). Demersal/benthic species represented a diverse group of 

families, equally divided among resident and transient 

species. The majority of the seagrass canopy dwellers 

were transient species and included some of the most 

abundant fishes (e.g., silver jenny, silver perch, and 

pinfish) and juveniles of an important recreational and 

commercial species (gray snapper). Although only four 

families of resident canopy dwellers were collected in 

Florida Bay (fundulids, cyprinodonts, syngnathids, and 

sciaenids), many of the species collected were abundant 

(cyprinodont: goldspotted killifish; syngnathids: dwarf 

seahorse, dusky pipefish, gulf pipefish), or very abundant (fundulid: rainwater killifish). The only sciaenid 

resident canopy-dweller collected was the spotted seatrout, an important recreational species.

The majority of species collected in basins by otter 

trawl exhibited significant spatial differences in densities (n = 46 species tested) both spatially (79%) and 

yearly (62%), but not monthly (46%). There were no 

discrete patterns that could explain the differences, but 

differences could pose questions for future research. 

Of those species that exhibited differences in densities 

among years (n = 25), 72% were resident fishes and 67% 

were transient. This provides little insight into factors 

influencing abundance. For example, dramatic unequal 

differences among years between resident and transient 

Chapter 27. Overview and future research needs

species could provide insight into whether factors inside 

or outside Florida Bay influenced abundances. There 

were also similar proportions of seagrass canopy dwellers (68%; n = 22 species), benthics (60%; n = 10 species), 

and pelagics (57%; n = 7 species from otter trawl collections in basins) that exhibited significant differences in 

densities among years. The canopy dwellers included 

almost all of the most abundant fishes (>1000); i.e, the 

transients – silver perch, silver jenny, white grunt, pinfish, and pigfish; the residents – goldspotted killifish, 

dwarf seahorse, rainwater killifish, dusky pipefish, and 

gulf pipefish; and one pelagic – bay anchovy. Following 

the seagrass die-off in the late 1980s, lowest densities of 

seagrass canopy dwellers were observed in 1994–1995 

(gulf pipefish, silver jenny, white grunt, and silver 

perch) and in 1996 (goldspotted killifish, rainwater 

killifish, dusky pipefish, and pinfish) that could be attributed to seagrass die-off. We attributed the dramatic 

increase in bay anchovy following seagrass die-off to 

increases in algae blooms and turbidity. In general, 

we observed relatively high densities of the abundant 

transients during the last sampling period (2000–2001) 

with the exception of pigfish when lowest densities for 

this species were observed. The majority of the most 

abundant species (listed above) occurred at greatest 

densities in the Western subdivision. The exceptions 

were bay anchovy, which was most abundant in the 

Central and Northern subdivisions, and gulf pipefish, 

which was equally abundant in the Western and Northern subdivisions. 

The richness of fauna in western Florida Bay has 

been well documented. Thayer and Chester (1989) 

concluded that richer and diverse fish communities are 

found in western Florida Bay where sediments are more 

organic and seagrass populations are more diverse. 

They attributed the low densities of fishes in the central 

part of the Bay to restricted circulation. Holmquist et 

al. (1989a; 1989b) also suggested that the mud bank 

network impedes circulation that acts as a barrier to 

recruitment of Gulf of Mexico and Atlantic crustaceans 

to the inner Bay. Tabb and Roessler (1989) believed that 

hypersalinity, which occurs predominantly in the central 

part of the Bay (Orlando et al., 1997), is a severe constraint on Florida Bay fish abundance and occurrence. 

In summary, frequently considered hypotheses for the 

distributional limits of species include physiological 

constraints, recruitment, competition, and predation, 

which might not be mutually exclusive (Quinn and 

Dunham, 1983). Very few studies conducted in Florida 

Bay have tested those hypotheses.

Notably absent from our study were the recreationally 

important tarpon (Megalops atlanticus), snook (Centropo-
202

mis undecimalis), and red drum (Sciaenops ocellatus). Despite their importance to the sport fishery in Florida Bay, 

the Bay is not a nursery area for these species. We are 

unaware of juvenile snook being collected in Florida Bay 

or adjacent waters (e.g., Whitewater Bay–Shark River 

estuary) in numbers that indicate nursery habitat. We 

believe that snook enter Florida Bay and adjacent waters 

as sub-adults from areas north of Florida Bay. Juveniles 

have been collected in east-central Florida (Gilmore et 

al., 1983), Tampa Bay and the Naples area (McMichael 

et al., 1989; Peters et al., 1998), and the Ten Thousand 

Islands (Carter et al., 1973).

Tarpon juveniles have been reported from shallow 

waters of the Whitewater-Coot Bay system (Tabb and 

Manning, 1961). They have been reported to inhabit 

salt marsh and mangrove ponds, tidal creeks, rivers, 

ditches, beaches, and mosquito-control impoundments 

where these habitats are shallow with no submerged vegetation and are lined by reeds or mangroves (Zale and 

Merrifield, 1989). This species was not collected during 

an extensive sampling program in quiet backwaters of 

Florida Bay and adjacent mesohaline waters (Powell et 

al., 2002).

Red drum is one of the most popular species sought 

by anglers in Florida Bay, but juveniles are rarely encountered. An extensive sampling program in northern 

Florida Bay and adjacent waters with characteristics of 

other areas where red drum occur (quiet backwaters, 

with oligohaline and mesohaline salinities) were unsuccessful in collecting any red drum juveniles (Powell 

et al., 2002). Oligohaline and meohaline waters in the 

Whitewater Bay–Shark River estuary might be a nursery 

area for this species (Odum and Heald, 1972). 

Future research needs

We recommend the following research needs that 

should compliment and supplement this study:

1. Using this study’s database from 1994, a multivariate analysis should be conducted to supplement the 

multivariate analysis performed with 1984–1985 

data (Thayer and Chester, 1989) to evaluate fishhabitat associations among years.

2. There is very little life history information (especially process-oriented) on juvenile and small resident 

fishes in Florida Bay, particularly for the abundant 

forage fishes (e.g., silver jenny, Eucinostomus gula; 

silver perch, Bairdiella chrysoura; rainwater killifish, 

Lucania parva). Yet these species are undoubtedly 

ecologically important and the resident species, 

in particular, could be good indicators of water or 

habitat quality.

3. The low densities of fishes in the interior of the Bay 

might be a function of limited recruitment resulting from the lack of circulation in the Bay. Research 

should be directed that addresses the low densities 

of fauna in the interior of the Bay, taking into account not only recruitment, but also physiological 

constraints, competition, and predation.

4. Fruitful research could be conducted by using techniques such as otolith microchemistry to determine 

the source of snook, tarpon, and red drum that inhabit Florida Bay as adults. Adult populations could 

be enhanced by determining spawning and nursery 

areas and preserving these areas.
203

Acknowledgments

Numerous individuals spent many hours discussing sampling designs, collecting samples, analyzing samples and 

data, and providing excellent advice. Particular thanks 

are expressed to Dave Colby, Peter Crumley, Monica 

Daniels, Dave Evans, Don Field, Mark Fonseca, Jim 

Fourqurean, Mike Greene, Peggy Harrigan, G. Hendrix, 

Bill Hettler, Don Hoss, Mike Johnson, Jud Kenworthy, 

Larry Krepp, Patti Marraro, Dave Meyer, William Odum, 

Dave Peters, George Powell, Keith Rittmaster, Jose 

Rivera, Roger Robbins, Mike Robblee, Ed Rutherford, 

Larry Settle, Julie Scope, Sue Sogard, DeWitt Smith, 

Vicki Thayer, Tom Schmidt, Jim Tilmant, Allison Veshlow, Harvey Walsh, Paula Whitfield, Lisa Wood, Mark 

Wuenschel, and Jay Zieman. We are also grateful to 

the following people who reviewed the manuscript and 

provided valuable comments: Dean Ahrenholz, Jon 

Hare, Joseph Smith, Patti Marraro, and an anonymous 

reviewer. Thank you all.

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209

Appendix Table A

Densities (numbers 1000 m-2) of juvenile and small resident fishes reported herein by salinity regime. There were 83 

collections in hyperhaline, 698 in euhaline, 357 in polyhaline, and 30 in mesohaline waters. Densities are derived from all 

collections from 1984-2001. 

Hyperhaline Euhaline Polyhaline Mesohaline

Family Species (≥40 ppt) (30-39 ppt) (19-29 ppt) (5-18 ppt)

Clupeidae Brevoortia smithi 0 0.22 <0.01 0

Harengula jaguana 0.07 0.22 2.38 0

Opisthonema oglinum 0.28 0.11 0.23 1.66

Sardinella aurita 0 0.5 0.05 1.62

Engraulidae Anchoa hepsetus 0 0.33 0.41 0

A. mitchilli 8.07 15.89 62.97 19.59

Ariidae Arius felis 0.21 0.23 0.48 0.22

Synodontidae Synodus foetens 0.47 0.59 0.27 0.05

Batrachoididae Opsanus beta 2.46 1.68 1.44 1.67

Hemiramphidae Chriodorus atherinoides 0.02 0.04 0.05 0.11

Hemirhamphus brasiliensis 0 0.04 0 0

Hyporhamphus unifasciatus 0 0.25 0 0

Belonidae Strongylura notata 0.18 0.19 0.13 0.91

S. timucu 0.04 0.21 0 0

Cyprinodontidae Floridichthys carpio 4.74 4.36 4.88 10.13

Fundulidae Lucania parva 44.86 37.85 15.31 22.57

Atherinidae Atherinomorus stipes 0 1.03 0.47 0

Hypoatherina harringtonensis 0 <0.01 0.02 0

Atherinopsidae Membras martinica 0 <0.01 0 0

Syngnathidae Anarchopterus criniger 0.38 1.03 0.99 2.12

Syngnathus floridae 4.11 3.72 3.59 1.39

S. scovelli 6.84 5.68 4.47 10.34

Hippocampus erectus 0.15 0.15 0.12 0

H. zosterae 0.15 3.44 0.12 0

Lutjanidae Lutjanus griseus 0.7 0.55 0.26 0.14

Lutjanus synagris 0.15 1.85 0.68 0.25

Gerreidae Eucinostomus gula 37.43 29.02 20.91 13.17

Haemulidae Haemulon plumieri 1.86 4.8 2.65 0

H. sciurus 0.28 0.3 0.31 0

Orthopristis chrysoptera 1.52 2.94 1.95 0

Sparidae Archosargus probatocephalus 0.02 0.24 0 0

Calamus arctifrons 0.06 0.21 0.06 0

Lagodon rhomboides 12.9 22.74 5.47 0

Sciaenidae Bairdiella chrysoura 2.79 3.61 1.6 0

Cynoscion nebulosus 1.18 0.61 0.28 0.12

Sphyraenidae Sphyraena barracuda 0.26 0.06 0.18 0.16

Scaridae Nicholsina usta 0.18 0.19 0.04 0

Callionymidae Diplogrammus pauciradiatus 0.16 0.2 0.16 0.17

Gobiidae Gobiosoma robustum 0.56 0.94 0.82 0.24

Microgobius gulosus 0.69 0.61 0.73 5.2

M. microlepis 0.26 0.1 0.62 0.07

Monacanthidae Monacanthus ciliatus 0.91 1.9 0.6 0

M. hispidus 0.32 0.37 0.26 0

Ostraciidae Lactophrys quadricornis 0.24 0.4 0.13 0

Tetraodontidae Sphoeroides nephalus 0.1 0.16 0.06 0

Diodontidae Chilomycterus shoepfi 0.52 0.48 0.33 0

Appendix A. Abundance of fishes by salinity regime
210

Appendix Table B

The total number of fishes and area collected of uncommon (<50 and ≥10) species in basins by otter trawl. Simple check 

mark (✓) indicates occurrence; bold (✔) relatively more abundant (arbitrarily assigned). Florida Bay was stratified into 

strata from 1984-1998, and subdivision from 1999-2001. For location of strata and subdivisions see Fig. 2.1. 

Strata1 Subdivision2

Family Species Number E C W E N A C W G

Ariidae Bagre marinus 20 ✓ ✔ ✔

Bythitidae Gunterichthys longipenis 13 ✓ ✓ ✔ ✓

Poeciliidae Poecilia latipinna 32 ✔ 

Syngnathidae Syngnathus louisianae 47 ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✔ ✓

Triglidae Prionotus scitulus 11 ✓ ✓ ✓ ✔ ✔ ✔

Serranidae Diplectrum formosum 11 ✓ ✔ ✓ ✔

Hypoplectrus unicolor 23 ✓ ✔ ✔

Lutjanidae Ocyurus chrysurus 10 ✓ ✓ ✓ ✔

Haemulidae Haemulon aurolineatum 34 ✔ ✓

Haemulon parra 20 ✓ ✓ ✔ ✓

Sparidae Archosargus rhomboidalis 22 ✓ ✓ ✔

Sciaenidae Bairdiella batabana 19 ✓ ✔ ✔

Ephippidae Chaetodipterus faber 42 ✓ ✓ ✓ ✔ ✔

Labridae Lachnolaimus maximus 12 ✓ ✓ ✓ ✔

Scaridae Sparisoma rubripinne 11 ✔ ✔

Labrisomidae Paraclinus fasciatus 40 ✓ ✓ ✓ ✓ ✔ ✔ ✓ ✓

P. marmoratus 42 ✓ ✓ ✓ ✓ ✓ ✓ ✔

Blennidae Chamodes saburrae 24 ✓ ✓ ✓ ✓ ✓ ✔

Hypsoblennius ionthas 10 ✓ ✓ ✔ ✔ ✔

Paralichthyidae Paralichthys albigutta 10 ✓ ✓ ✓ ✓ ✔ ✓

Achiridae Achirus lineatus 33 ✓ ✓ ✓ ✓ ✔ ✔ ✓

Cynoglossidae Symphurus plagiusa 24 ✓ ✓ ✓ ✓ ✓ ✔

Ostraciidae Lactophrys trigonis 12 ✓ ✓ ✓ ✓ ✔

Tetraodontidae Sphoeroides spengleri 16 ✓ ✓ ✓ ✔ ✓ ✓

1 E = East; C = Central; W = West

2

 E = Eastern; N = Northern; A = Atlantic; C = Central; W = Western; G = Gulf

Appendix B. Uncommon species—number and area collected
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