OCS Study

BOEM 2013-214

FGB NMS 2013-002

Long-Term Monitoring at the East and 

West Flower Garden Banks National

Marine Sanctuary, 2009–2010

Volume 1: Technical Report

U.S. Department of the Interior

Bureau of Ocean Energy Management

Gulf of Mexico OCS Region

National Oceanic and Atmospheric Administration

Flower Garden Banks National Marine Sanctuary
OCS Study

BOEM 2013-214

FGB NMS 2013-002

Long-Term Monitoring at the East and 

West Flower Garden Banks National

Marine Sanctuary, 2009–2010

Volume 1: Technical Report

Authors

Michelle A. Johnston

Marissa F. Nuttall

Ryan J. Eckert

John A. Embesi

Niall C. Slowey

Emma L. Hickerson

George P. Schmahl

Prepared under BOEM Interagency Agreement

M09PG00011

by

U.S. Department of Commerce

NOAA Flower Garden Banks National Marine Sanctuary

Avenue U, Bldg. 216

Galveston, TX 77551

Published by

U.S. Department of the Interior

Bureau of Ocean Energy Management

Gulf of Mexico OCS Region

New Orleans, LA

December 2013
DISCLAIMER

This report was prepared under an interagency agreement between the Bureau of Ocean Energy 

Management (BOEM) and NOAA Flower Garden Banks National Marine Sanctuary. This report 

has been technically reviewed by BOEM, and it has been approved for publication. Approval 

does not necessarily signify that the contents reflect the views and policies of BOEM, nor does 

mention of trade names or commercial products constitute endorsement or recommendation for 

use.

REPORT AVAILABILITY

To download a PDF file of this Gulf of Mexico OCS Region report, go to the U.S. Department of 

the Interior, Bureau of Ocean Energy Management, Environmental Studies Program Information 

System website and search on OCS Study BOEM 2013-214. 

This report can be viewed at select Federal Depository Libraries. It can also be obtained from 

the National Technical Information Service; the contact information is below. 

U.S. Department of Commerce

National Technical Information Service

5301 Shawnee Rd. 

Springfield, Virginia 22312 

Phone: (703) 605-6000, 1(800)553-6847 

Fax: (703) 605-6900 

Website: http://www.ntis.gov/

CITATION

Johnston, Michelle A., M.F. Nuttall, R.J. Eckert, J.A. Embesi, N.C. Slowey, E.L. Hickerson, and 

G.P. Schmahl. 2013. Long-term monitoring at the East and West Flower Garden Banks 

National Marine Sanctuary, 2009–2010, volume 1: technical report. U.S. Dept. of 

Interior, Bureau of Ocean Energy Management, Gulf of Mexico OCS Region, New 

Orleans, Louisiana. OCS Study BOEM 2013-214. 202 pp. 

ABOUT THE COVER

The cover photograph features Marissa Nuttall, a NOAA scientific diver from the Flower 

Garden Banks National Marine Sanctuary, following a compass heading while laying out a 

random transect line at the East Flower Garden Bank long-term monitoring study site. 

Photograph by John Embesi (NOAA FGBNMS). 

iii
ACKNOWLEDGMENTS

We thank the individuals, listed below, who contributed their time and expertise to make this 

long-term monitoring effort successful. We are grateful to the NOAA and BOEM staff for their 

invaluable institutional knowledge. In particular, we acknowledge James Sinclair for his ongoing 

support and enthusiasm. 

NOAA FGBNMS PERSONNEL AND ROLE

Michelle A. Johnston, Ph.D. Project Manager, Data Analysis, Report Compilation

Marissa Nuttall Cruise Logistics,Data Acquisition and Analysis

Ryan Eckert Data Acquisition and Analysis

John Embesi Field Management, Data Acquisition and Analysis

Emma L. Hickerson Sanctuary Research Coordinator, Data Acquisition

G.P. Schmahl Sanctuary Superintendent/ Data Acquisition

PROJECT DIVERS 2009–2010

Joe Bosquez Anna Floyd G.P. Schmahl

Kevin Buch Steve Gittings Ian Shirley

Jessica Coleman Tracy Hamburger James Sinclair

Kelly Drinnen Emma Hickerson Charles Vance

Ryan Eckert Carrie Jones Kate Winters

John Embesi Marissa Nuttall

REPORT CONTRIBUTERS AND REVIEWERS 

The Flower Garden Banks National Marine Sanctuary would like to acknowledge the support 

and efforts of subject area experts who assisted with this technical report. The report benefited 

significantly from the following individuals who either provided a peer review or provided 

comments, content, images, and/or data: James Sinclair (Bureau of Ocean Energy Management), 

Michelle K. Nannen (Bureau of Ocean Energy Management), Gregory Boland (Bureau of Ocean 

Energy Management), Dr. Steve Gittings (NOAA’s Office of National Marine Sanctuaries), 

Amy Wagner (Texas A&M University), Dr. Quenton Dokken (Gulf of Mexico Foundation), Dr. 

Ivor Williams (NOAA’s National Marine Fisheries Service), and William F. Precht (Dial Cordy 

& Associates, Inc). 

v 
CONTENTS

LIST OF FIGURES ....................................................................................................................... xi

LIST OF TABLES ......................................................................................................................... xi

EXECUTIVE SUMMARY ......................................................................................................... xxi

CHAPTER 1.0: INTRODUCTION ............................................................................................. 1

1.1. Coral Reef Monitoring at the Flower Garden Banks National Marine Sanctuary ............. 1

1.2. The Flower Garden Banks National Marine Sanctuary in the Gulf of Mexico ................. 3

1.2.1. Habitat Description ............................................................................................... 3

1.2.2. The East and West Flower Garden Banks ............................................................ 5

1.3. BOEM and FGBNMS Protective Measures ...................................................................... 7

1.4. Deepwater Horizon ............................................................................................................ 9

CHAPTER 2.0: STUDY SITES ................................................................................................. 11

2.1. Study Site Methodology .................................................................................................. 11

2.1.1. 100 x 100 m Study Sites ..................................................................................... 11

2.1.2. EFGB Deep Repetitive Quadrat Stations ........................................................... 19

2.1.3. Study Site Rehabilitation .................................................................................... 20

CHAPTER 3.0: Random Transects ........................................................................................... 21

3.1. Random Transect Methodological Rationale................................................................... 21

3.2. Random Transect Field and Laboratory Methods ........................................................... 21

3.3. Statistical Analyses of the Random Transect Data .......................................................... 23

3.4. Random Transect Results ................................................................................................ 24

3.5. Random Transect Discussion ........................................................................................... 35

3.5.1. EFGB Comparison from 2009 to 2010 .............................................................. 37

3.5.2. WFGB Comparison from 2009 to 2010 ............................................................. 37

3.5.3. Comparison of Random Transect Results from 1992 to 2008 ........................... 37

CHAPTER 4.0: Repetitive Quadrats ........................................................................................ 41

4.1. Repetitive Quadrat Methodological Rationale ................................................................. 41

4.2. Repetitive Quadrat Field and Laboratory Methods .......................................................... 41

4.2.1. Repetitive Quadrat 2009 Digital Photography ................................................... 42

4.2.2. Repetitive Quadrat 2010 Digital Photography ................................................... 42

4.2.3. Repetitive Quadrat Image Analysis .................................................................... 42

4.2.4. Planimetry Analysis ........................................................................................... 43

vii 
4.3. Repetitive Quadrat Results .............................................................................................. 43

4.3.1. Repetitive Quadrat Analysis ............................................................................... 43

4.3.2. Repetitive Quadrat Planimetric Analysis ........................................................... 55

4.4. Repetitive Quadrat Discussion ......................................................................................... 61

4.4.1. Study Site Repetitive Quadrats ........................................................................... 61

4.4.2. Deep Station Quadrats ........................................................................................ 63

4.4.3. Planimetry .......................................................................................................... 64

CHAPTER 5.0: Sclerochronology ............................................................................................. 67

5.1. Sclerochronology Methodological Rationale .................................................................. 67

5.1.1. Why Montastraea faveolata? .............................................................................. 68

5.2. Sclerochronology Field and Laboratory Methods ........................................................... 69

5.3. Data Presentation and Statistical Analysis of Growth Rates ........................................... 70

5.4. Sclerochronology Results ................................................................................................ 70

5.5. Sclerochronology Discussion........................................................................................... 74

CHAPTER 6.0: Lateral Growth ................................................................................................ 77

6.1. Lateral Growth Methodological Rationale ...................................................................... 77

6.2. Lateral Growth Field and Laboratory Methods ............................................................... 77

6.2.1. Image Analysis for Lateral Growth .................................................................... 78

6.3. Data Presentation and Statistical Analysis for Lateral Growth ....................................... 80

6.4. Lateral Growth Results .................................................................................................... 80

6.5. Lateral Growth Discussion .............................................................................................. 83

CHAPTER 7.0: Perimeter Videography ................................................................................... 85

7.1. Perimeter Videography Methodological Rationale ......................................................... 85

7.2. Perimeter Videography Field and Laboratory Methods .................................................. 85

7.3. Perimeter Videography Results ....................................................................................... 85

7.3.1. EFGB Perimeter Lines ....................................................................................... 86

7.3.2. EFGB 360° Panoramic Views ............................................................................ 86

7.3.3. WFGB Perimeter Lines ...................................................................................... 87

7.3.4. WFGB 360° Panoramic Views........................................................................... 88

7.4. Perimeter Videography Discussion.................................................................................. 88

CHAPTER 8.0: Qualitative Field Observations ...................................................................... 91

8.1. Qualitative Field Observations Methodological Rationale .............................................. 91

8.2. Qualitative Field Observations Methods ......................................................................... 91

8.3. Qualitative Field Observation Results ............................................................................. 91

8.3.1. Wildlife Observations ......................................................................................... 91

8.3.2. Coral Health Observations ................................................................................. 92

8.3.3. Exotic and Invasive Species ............................................................................... 97

8.4. Qualitative Field Observations Discussion ...................................................................... 98

8.4.1. Wildlife Observations ......................................................................................... 98

viii
8.4.2. Coral Health Observations ................................................................................. 99

8.4.3. Exotic and Invasive Species ............................................................................. 101

CHAPTER 9.0: Water Quality ................................................................................................. 105

9.1. Water Quality Methodological Rationale ...................................................................... 105

9.2. Water Quality Field and Laboratory Methods ............................................................... 105

9.2.1. Sea-Bird Conductivity and Temperature Recorder .......................................... 105

9.2.2. HoboTemp Thermographs ............................................................................... 107

9.2.3. YSI Probe ......................................................................................................... 107

9.2.4. Chlorophyll a and Nutrients ............................................................................. 108

9.3. Water Quality Results .................................................................................................... 109

9.3.1. Sea-Bird Temperature and Salinity .................................................................. 109

9.3.2. HoboTemp Thermograph Data ........................................................................ 113

9.3.3. YSI Vertical Profiles ........................................................................................ 114

9.3.4. Water Samples .................................................................................................. 116

9.3.5. Hydrocarbons ................................................................................................... 118

9.4. Water Quality Discussion .............................................................................................. 118

9.4.1. Water Quality Parameters ................................................................................ 118

9.4.2. Chlorophyll a and Nutrients ............................................................................. 123

9.4.3. Sea State ........................................................................................................... 123

CHAPTER 10.0: Fish Surveys ................................................................................................. 127

10.1. Fish Surveys Methodological Rationale ...................................................................... 127

10.2. Fish Surveys Field Methods......................................................................................... 127

10.3. Fish Survey Data and Statistical Analysis ................................................................... 128

10.4. Fish Surveys Results .................................................................................................... 130

10.4.1. Sighting Frequency and Occurrence .............................................................. 135

10.4.2. Family Density, Relative Abundance, and Richness ..................................... 146

10.4.3. Diveristy and Evenness of Populations .......................................................... 150

10.4.4. Trophic Group Comparisons .......................................................................... 150

10.4.5. Biomass Analysis ........................................................................................... 165

10.4.6. Diurnal Abundance Patterns ........................................................................... 178

10.4.7. Prevalent Compared with Rare Species ......................................................... 180

10.5. Fish Surveys Discussion .............................................................................................. 182

CHAPTER 11.0: Sea Urchin and Lobster Surveys ............................................................... 185

11.1. Sea Urchin and Lobster Surveys Methodological Rationale ....................................... 185

11.2. Sea Urchin and Lobster Surveys Field Methods.......................................................... 185

11.3. Sea Urchin and Lobster Surveys Results ..................................................................... 185

11.4. Sea Urchin and Lobster Surveys Discussion ............................................................... 187

CHAPTER 12.0: Recommendations ....................................................................................... 189

LITERATURE CITED ............................................................................................................ 191

ix
LIST OF FIGURES

Figure 1.2.1. .........Map of the EFGB and WFGB in relation to the Texas-Louisiana 

continental shelf and other topographic features of the 

northwestern Gulf of Mexico. ........................................................................... 4

Figure 1.2.2. .........Topographic contour map of the FGBNMS (NOAA/FGBNMS). ................... 5

Figure 1.2.3. .........Bathymetric map of the EFGB (NOAA/FGBNMS). ........................................ 6

Figure 1.2.4. .........Bathymetric map of the WFGB (NOAA/FGBNMS). ...................................... 7

Figure 1.4.1. .........Map of the location of the Deepwater Horizon site in relation to the 

FGBNMS, including the maximum extent of observed oil from the 

spill (NOAA). ................................................................................................. 10

Figure 2.1.1. .........Topographic map of the EFGB. Inset shows the locations of the 

corners of the study site (USDOI/GS 2001). .................................................. 13

Figure 2.1.2. .........Topographic map of the WFGB. Inset shows the locations of the 

corners of the study site (USDOI/GS 2001). .................................................. 14

Figure 2.1.3. .........Locations of EFGB mooring buoys (NOAA/FGBNMS). .............................. 15

Figure 2.1.4. .........Locations of WFGB mooring buoys (NOAA/FGBNMS). ............................. 16

Figure 2.1.5. .........Locations of monitoring stations at the EFGB, 2010 

(NOAA/FGBNMS). ........................................................................................ 17

Figure 2.1.6. .........Locations of monitoring stations at the WFGB, 2010 

(NOAA/FGBNMS). ........................................................................................ 18

Figure 2.1.7. .........Bathymetric map with the deep repetitive quadrat stations in 

relation to the permanent study site at the EFGB (32–40 m or 105–

131 ft), established in April 2003. .................................................................. 19

Figure 2.1.8. .........Map showing depth and relative locations of the nine deep 

repetitive quadrat stations at the EFGB. ......................................................... 20

Figure 3.2.1. .........A diver with camera mounted on aluminum t-frame taking random 

transect photographs. The line to the diver’s left is a study site 

boundary line (NOAA/FGBNMS). ................................................................. 23

Figure 3.4.1. .........Percent cover (error bars are ± SE) of four functional categories of 

sessile benthos at the FGB from 2009 and 2010............................................. 24

Figure 3.4.2. .........Percent cover (+ SE) of the dominant coral taxa at the EFGB from 

2009 and 2010. Values are calculated from the random transect 

photographs. .................................................................................................... 25

Figure 3.4.3. .........Percent cover (+ SE) of the dominant coral taxa at the WFGB from 

2009 and 2010. ................................................................................................ 26

xi
Figure 3.4.4. .........Percent cover (+ SE) of the coral species that comprise the MASC 

at the EFGB from 2009 and 2010 (note scale difference from 

Figure 3.4.5). ................................................................................................... 29

Figure 3.4.5. .........Percent cover (+ SE) of the coral species that comprises the MASC 

at the WFGB from 2009 and 2010 (note scale difference from 

Figure 3.4.4). ................................................................................................... 30

Figure 3.4.6. .........Percent of coral observed to pale and bleach (+ SE) at the FGB in 

2009 and 2010. ................................................................................................ 31

Figure 3.4.7. .........Species composition of bleached coral from percent cover random 

transect analysis at the FGB from 2009 and 2010. ......................................... 32

Figure 3.4.8. .........Species composition of paling coral from percent cover random 

transect analysis at the FGB from 2009 and 2010. ......................................... 32

Figure 3.5.1. .........Random transect mean percent coral cover at the EFGB and 

WFGB over time, showing the consistently high coral cover. ....................... 36

Figure 3.5.2. .........Percent cover of Montastraea annularis species complex,

macroalgae, and CTB from 1992 to 2010 at (A) the EFGB and (B) 

the WFGB. ...................................................................................................... 38

Figure 4.2.1. .........Repetitive quadrat station #761 at the EFGB in 2009 

(NOAA/FGBNMS). ........................................................................................ 41

Figure 4.3.1. .........Percent cover (+ SE) of four functional categories of sessile 

benthos at the FGB in 2009 and 2010. ............................................................ 44

Figure 4.3.2. .........Percent cover (+ SE) of the dominant coral taxa in repetitive 

quadrats at the EFGB in 2009 and 2010. ........................................................ 45

Figure 4.3.3. .........Percent cover (+ SE) of the dominant coral taxa in repetitive 

quadrats at the WFGB in 2009 and 2010. ....................................................... 46

Figure 4.3.4. .........Percent cover (+ SE) of the coral species in the MASC in repetitive 

quadrats at the EFGB in 2009 and 2010. ........................................................ 49

Figure 4.3.5. .........Percent cover (+ SE) of the coral species in the MASC in repetitive 

quadrats at the WFGB in 2009 and 2010. ....................................................... 49

Figure 4.3.6. .........Percent of coral observed to pale and bleach (+ SE) in repetitive 

quadrats at the FGB in 2009 and 2010. ........................................................... 50

Figure 4.3.7. .........Percent cover (+ SE) of four functional categories of sessile 

benthos at the EFGB repetitive deep stations in 2009 and 2010. ................... 51

Figure 4.3.8. .........Percent cover (+ SE) of the dominant coral taxa at repetitive deep 

stations on the EFGB in 2009 and 2010. ........................................................ 52

Figure 4.3.9. .........Percent cover (+ SE) of the coral species that comprises the MASC 

at deep repetitive stations the EFGB deep stations from 2009 and 

2010................................................................................................................. 54

xii
Figure 4.4.1. .........Repetitive photostation 761 from the EFGB in a time series from 

2006 to 2010, showing a healthy and stable coral community. (A) 

2006; (B) 2007; (C) 2008; (D) 2009; (E) 2010 (NOAA/FGBNMS). ............. 62

Figure 4.4.2. .........Deep station coral cover (+ SE) from 2003 to 2010, averaging 

above 70% at the EFGB. ................................................................................. 64

Figure 4.4.3. .........Coral colonies from repetitive quadrat station 005 at the EFGB 

outlined for planimetric analysis from 2009 (A) and 2010 (B) for a 

combined overlay (C). .................................................................................... 65

Figure 5.1.1. .........X-ray of Montastraea faveolata showing skeletal banding 

(NOAA/FGBNMS). ........................................................................................ 68

Figure 5.4.1. .........X-radiographs of Montastraea faveolata skeletal structure in the 

plane of corallite growth from four quadrants in the EFGB longterm monitoring study site. ............................................................................. 71

Figure 5.4.2. .........X-radiographs of Montastraea faveolata skeletal structure in the 

plane of corallite growth from four quadrants in the WFGB longterm monitoring study site. ............................................................................. 72

Figure 5.4.3. .........Mean annual growth rates in cores collected in 2010 at the EFGB 

and WFGB. ..................................................................................................... 74

Figure 5.5.1. .........Mean annual growth rates (+SE) based on analysis of Montastraea

faveolata 2003, 2005, 2007, and 2010 cores from the EFGB and 

WFGB. ............................................................................................................ 76

Figure 6.2.1. .........Lateral growth station #52 on a Diploria strigosa colony with 

close-up framer at the EFGB (NOAA/FGBNMS).......................................... 78

Figure 6.2.2. .........Image analysis of Diploria strigosa lateral growth at the EFGB. ................... 79

Figure 6.4.1. .........Ternary diagram showing the proportion of Diploria strigosa

marginal tissue growing, retreating, or remaining stable at each 

station photographed at the EFGB from 2009 to 2010. .................................. 82

Figure 6.4.2. .........Ternary diagram showing the proportion of Diploria strigosa 

marginal tissue growing, retreating, or remaining stable at each 

station photographed at the WFGB from 2009 to 2010. ................................. 83

Figure 7.4.1. .........A head of Porites astreoides observed to be healthy in 2009 (A), 

but bleached in 2010 (B) at the WFGB. ......................................................... 89

Figure 8.3.1. .........Manta ray M46 observed at the EFGB in 2009 with spot and 

square ventral markings (J. Wiseman and S. Bernhardt). ............................... 92

Figure 8.3.2. .........Fluorescent orange algae, Martensia pavonia, observed at the 

EFGB in 2010 (NOAA/FGBNMS). ............................................................... 93

Figure 8.3.3. .........The Acropora palmata colony at the EFGB in (A) 2005 (B) 2006 

and (C) 2010. .................................................................................................. 95

Figure 8.3.4. .........The only Acropora palmata colony at the WFGB is shown to be 

dying from disease in (A) 2008 (B) 2009 and (C) 2010. ................................ 96

xiii
Figure 8.3.5. .........Orange cup coral (Tubastraea coccinea), an invasive species from 

the Pacific that is beginning to establish itself at the FGBNMS 

(NOAA/FGBNMS). ........................................................................................ 97

Figure 8.4.1. .........FGBNMS Manta Ray sighting catalog. The manta rays in the 

catalog have been divided into categories to aid in identifying 

individuals, each by its unique ventral markings 

(NOAA/FGBNMS). ........................................................................................ 98

Figure 8.4.2. .........Coral Reef Watch bleaching alert map for the Gulf of Mexico in 

September 2010. ........................................................................................... 100

Figure 8.4.3. .........Invasive lionfish at the West Flower Garden Bank. ..................................... 103

Figure 9.2.1. .........Sea-Bird 37-SMP MicroCAT water quality instrument at the 

EFGB (NOAA/FGBNMS)............................................................................ 106

Figure 9.2.2. .........FGBNMS research divers mounting the Sea-Bird 37-SMP 

MicroCAT water quality instrument on a rack in the sand at the 

EFGB (NOAA/FGBNMS)............................................................................ 106

Figure 9.2.3. .........FGBNMS researchers conducting seawater temperature profiles 

using a YSI probe through the moon pool of the R/V Manta

(NOAA/FGBNMS). ...................................................................................... 107

Figure 9.3.1. .........Daily average sea water temperature measured near the reef cap at 

the EFGB from 2009 to 2010........................................................................ 110

Figure 9.3.2. .........Daily average sea water temperature measured near the reef cap at 

the WFGB from 2009 to 2010. ..................................................................... 111

Figure 9.3.3. .........Salinity measured near the reef cap at the EFGB and WFGB from 

2009 and 2010. .............................................................................................. 112

Figure 9.3.4. .........Daily average seawater temperature measured near the reef cap at 

the EFGB from 2009 to 2010. Data obtained from HoboTemp 

thermographs................................................................................................. 113

Figure 9.3.5. .........Daily average seawater temperature measured near the reef cap at 

the WFGB from 2009 to 2010. Data obtained from HoboTemp 

thermographs................................................................................................. 114

Figure 9.4.1. .........Comparison of Sea-Bird and Hobo daily average sea water 

temperature readings measured near the reef cap at the EFGB from 

2009 to 2010. ................................................................................................ 119

Figure 9.4.2. .........Comparison of Sea-Bird and Hobo daily average sea water 

temperature readings measured near the reef cap at the WFGB 

from 2009 to 2010. ........................................................................................ 120

Figure 9.4.3. .........Sea water temperature measured using Sea-Bird datasonde near the 

reef cap at the EFGB in 2009 and 2010, and the historical longterm average temperature from past monitoring periods. ............................. 121

xiv
Figure 9.4.4. .........Sea water temperature measured using Sea-Bird datasonde near the 

reef cap at the WFGB in 2009 and 2010, and the historical longterm average temperature from past monitoring periods. ............................. 122

Figure 9.4.5. .........Hurricane Ida (A) and Tropical Storm Bonnie (B) storm track areas 

(NOAA Hurricane Center). ........................................................................... 126

Figure 10.4.1. .......WFGB size distribution by trophic guild. (A) herbivores, (B) 

invertivores, (C) piscivores, and (D) planktivores. ....................................... 163

Figure 10.4.2. .......EFGB size distribution by trophic guild. (A) herbivores , (B) 

invertivores, (C) piscivores, and (D) planktivores. ....................................... 164

Figure 10.4.3. .......Percent composition of biomass for each trophic guild by bank and 

year. ............................................................................................................... 167

Figure 10.4.4. .......Ranked abundance curve for the prevalence of all species recorded 

during the 2009 and 2010 surveys. ............................................................... 180

Figure 11.3.1. .......Sea urchin and lobster counts conducted at the EFGB and WFGB 

in 2009 and 2010. .......................................................................................... 186

Figure 11.4.1. .......Sea urchin and lobster densities at the FGB from 2004 to 2010. ................. 187

xv
LIST OF TABLES

Table 2.1.1. Cruise Dates at the EFGB and WFGB for 2009 and 2010 ............................ 12

Table 2.1.2. GPS Coordinates for the EFGB and WFGB Study-Site Corner 

Markers .......................................................................................................... 12

Table 3.4.1. Cover of Benthic Categories in Random Transects at the EFGB 

and WFGB from 2009 and 2010 ................................................................... 27

Table 3.4.2. Results of ANOVA on Proportional Cover Estimates from 

Random Transects from 2009 and 2010 ........................................................ 34

Table 3.5.1. Percentage of Coral Cover on Living Reefs in the Western 

Atlantic and Caribbean Region ..................................................................... 35

Table 3.5.2. EFGB and WFGB Random Transect Data for Dominant Cover 

Categories ...................................................................................................... 39

Table 4.3.1. Cover of Benthic Categories in Repetitive Quadrats at the EFGB 

and WFGB in 2009 and 2010 ........................................................................ 47

Table 4.3.2. Cover of Benthic Categories in Repetitive Deep Stations at the 

EFGB from 2009 and 2010 ........................................................................... 53

Table 4.3.3. Reef-Building Coral Colonies in Photographs from Repetitive 

Quadrat Stations Selected for Planimetric Analysis from 2009 to 

2010 ............................................................................................................... 55

Table 4.3.4. Visual Assessment of Analyzed Coral Colonies Suggest Colonies 

Did Not Suffer As Much Tissue Loss as the Planimetric Analysis 

Suggested ....................................................................................................... 58

Table 5.4.1. Estimates of Annual Growth (mm/yr) of Montastraea Faveolata

from Cores Extracted at the EFGB and WFGB in 2010 ............................... 73

Table 6.4.1. Lateral Growth Stations on Diploria Strigosa and Percent of 

Margin Growing, Retreating, or Remaining Stable of Coral Tissue 

at the EFB and WFGB from 2009 to 2010 .................................................... 81

Table 7.3.1. Comparison of Observations of the Condition of Individual Coral 

Colonies at EFGB in 2009 and 2010 ............................................................. 86

Table 7.3.2. Comparison of Observations of the Condition of Individual Coral 

Colonies at WFGB from 2009 and 2010 ....................................................... 87

Table 9.2.1. Water Sampling Schedule, Depth, and Number of Samples Taken 

at the EFGB and WFGB in 2009 to 2010 ................................................... 108

Table 9.2.2. Standard USEPA Methods Used to Analyze Water Samples Taken 

at the FGB ................................................................................................... 109

Table 9.3.1. Summary of Sea-Bird Seawater Temperature Parameters from 

2009 to 2010 ................................................................................................ 110

xvii 
Table 9.3.2. Summary of Sea-Bird Salinity Parameters from 2009 to 2010 ................... 112

Table 9.3.3. Summary of HoboTemp Seawater Temperature Parameters from 

2009 to 2010 ................................................................................................ 114

Table 9.3.4. Summary of Opportunistic YSI Seawater Temperature Vertical 

Profiles from 2009 to 2010 at the EFGB and WFGB ................................. 115

Table 9.3.5. Chlorophyll a Concentrations in Water Samples Taken at the 

EFGB and WFGB from 2009 to 2010 ......................................................... 116

Table 9.3.6. Concentrations of Ammonia, Nitrate, and TKN in Water Samples 

Taken at the EFGB and WFGB from 2009 to 2010 .................................... 117

Table 9.4.1. List of Tropical Cyclones that Entered the Gulf of Mexico from 

2002 to 2010 ................................................................................................ 124

Table 10.4.1. Complete Fish Species List, Rrophic Guilds, and Counts per Year 

per Bank ...................................................................................................... 130

Table 10.4.2. Summary of Fish Surveys at the EFGB and WFGB in 2009 and 

2010 ............................................................................................................. 133

Table 10.4.3. Species and Family Richness and Fish Density for the EFGB and 

WFGB in 2009 and 2010 ............................................................................. 134

Table 10.4.4. Mean Fish Abundance for the EFGB and WFGB in 2009 and 2010 

for Dominant Species .................................................................................. 135

Table 10.4.5. Percent Sighting Frequency for Species by Year and Bank, 

Including a Percent Sighting Frequency for All Surveys 

Conducted during the 2009 to 2010 Reporting Period ................................ 135

Table 10.4.6. Species Occurrence and Percent Sighting Frequencies in 

Parentheses for all Species Recorded at the FGB in 2009 and 2010 .......... 140

Table 10.4.7. Relative Abundance by Family for the 2009 and 2010 Surveys at 

Both the EFGB and WFGB ......................................................................... 147

Table 10.4.8. Mean Fish Density (number of individuals per 100 m2), Richness 

(number of species per family), and Mean Biomass by Family at 

the EFGB and WFGB in 2009 .................................................................... 148

Table 10.4.9. Mean Fish Density (number of individuals per 100 m2), Richness 

(number of species per family), and Mean Biomass by Family at 

the EFGB and WFGB in 2010 .................................................................... 149

Table 10.4.10. Shannon-Wiener Diversity Indices for the EFGB and WFGB in 

2009 and 2010 ............................................................................................. 150

Table 10.4.11. Comparison of Mean Density (fish per 100 m2) and Species 

Richness per Diver Survey of Herbivores, Piscivores, 

Invertivores, and Planktivores in Diver Surveys between the 

EFGB and WFGB in 2009 and 2010 ........................................................... 151

xviii
Table 10.4.12. Density Comparison between Banks for Herbivorous, Piscivorous, 

Invertivorous, and Planktivorous Fishes ..................................................... 153

Table 10.4.13. Abundances between Years for Herbivores, Piscivores, 

Invertivores, and Planktivores ..................................................................... 158

Table 10.4.14. Visual fish Survey Sampling Biomass Statistics for the EFGB and 

WFGB in 2009 and 2010 ............................................................................. 165

Table 10.4.15. Total Biomass (g/m2) for Each Family ........................................................ 165

Table 10.4.16. Comparison of Biomass (g/m2) of Herbivores, Piscivores, 

Invertivores, and Planktivores between the EFGB and WFGB in 

2009 and 2010 ............................................................................................. 168

Table 10.4.17. Two-way ANOVA for Trophic Guild Biomass (g/m2), with Fixed 

Factors of Bank and Year ............................................................................ 169

Table 10.4.18. Biomass (g/m2) of Each Species by Trophic Guild with P-values 

Shown Where Significant Differences Occur ............................................. 170

Table 10.4.19. Biomass Comparisons Between Years ........................................................ 175

Table 10.4.20 Two-way ANOVA for Great Barracuda Abundance, with Fixed 

Factors of Bank and Time of Day ............................................................... 179

Table 10.4.21. Ranked Abundance of "Prevalent" Species ................................................. 181

Table 10.5.1. Percentage of Fishes Observed in the Listed Families at Reefs 

around the Gulf of Mexico and the Caribbean Region ............................... 184

Table 11.3.1. Number of Individual Sea Urchins and Lobsters Observed During 

Surveys in 2009 and 2010 ........................................................................... 186

xix
EXECUTIVE SUMMARY

In more than 20 years of being continuously monitored, the coral reefs of the Flower 

Garden Banks National Marine Sanctuary have maintained high levels of coral cover, 

suffered minimally from hurricanes, coral bleaching, and disease outbreaks, and 

supported relatively diverse and abundant fish populations as well as other vertebrate and 

invertebrate species. No significant long-term changes have been detected in coral cover or 

diversity at the Flower Garden Banks during monitoring efforts that have taken place since 

1988, and likely not since the first measurements were made in the early 1970s. During the 

2009 to 2010 monitoring period, the analysis of monitoring data indicated that the East 

and West Flower Garden Banks were robust and productive. Based on the 2009 and 2010 

data, the average coral cover on the Flower Garden Banks coral caps is nearly 57 percent. 

Located 193 and 172 km (East and West Flower Garden Banks, respectively) offshore 

from Galveston, Texas, the banks are remotely located topographic features on the outer 

continental shelf in the Gulf of Mexico that are capped with reef-building corals. Relative 

to other coral reef systems in the Southeast and Caribbean region, the Flower Garden 

Banks (FGB) have a low diversity of stony corals, yet high coral cover (typically around 

50%). The East Flower Garden Bank (EFGB) and West Flower Garden Bank (WFGB), 

along with Stetson Bank, comprise the Flower Garden Banks National Marine Sanctuary

(FGBNMS). The reefs at these banks are afforded a certain measure of natural protection 

due to their depth underwater and geographic distance from land. 

In recent years, however, warning signs of impacts from natural and anthropogenic 

sources have been documented. Increased incidence of bleaching events, invasive 

species, and perceived reduction in fish densities at the FGBNMS (though in many cases 

not measurable) are reasons for increased vigilance and perhaps concern for the future of 

the resources. The incidence and prevalence of disease and bleaching in comparison to 

other western Atlantic coral sites, however, are low. The recent invasion by lionfish in 

the Gulf of Mexico, which are native to the Indo-Pacific, raises yet another cause for 

concern for the health and future of the bank ecosystems at the FGBNMS. This 

monitoring report represents a milestone, as it documents baseline, or at least preinvasion, ecosystem conditions before the invasion of lionfish creates phase shifts on the 

reefs.

The results of the 2009 to 2010 monitoring efforts, conducted in August 2009, and

August through November 2010, illustrate the continued stability of the coral reef 

community and associated fish populations. Random transect results revealed high coral 

cover within study sites at both banks from 2009 to 2010, and coral cover was estimated 

at 53.35% ± 4.17 and 54.49% ± 3.69, respectively, at the EFGB, and 53.84% ± 3.73 in 

2009 and to 65.95% ± 2.85 in 2010 at the WFGB. These results are consistent with 

previous monitoring efforts of high coral cover above 50% at the FGB (Dokken et al. 

1999; Dokken et al. 2001; CSA 1996; Gittings et al. 1992; Aronson et al. 2005, Zimmer 

et al. 2010), highlighting the coral stability over time. 

xxi 
Based on the results of the random transect data, the Montastraea annularis species 

complex (M. annularis, M. faveolata, and M. franksi; Weil and Knowlton 1994) was the

dominant component of coral cover at both banks; M. franksi was identified as the 

dominant species within the complex. Montastraea annularis species complex cover at 

the EFGB was estimated at 29.52% ± 12.99 in 2009 and 33.89% ± 3.84 in 2010. At the 

WFGB, cover was 27.25% ± 3.32 in 2009 and 45.98% ± 3.68 in 2010. Diploria strigosa 

was the next most abundant species during this period, with 10.08% ± 2.54 at the EFGB 

in 2009 and 9.38% ± 1.72 in 2010. The WFGB estimates were 10.84% ± 2.70 and 5.39% 

± 0.94 for the two years. 

In 2009 and 2010, macroalgae were more abundant than crustose coralline algae, fine turf 

algae, and bare rock (CTB), ranging from approximately 22–33% over both banks. The 

most dominant macroalgal cover was by fleshy algae, thick turf algae, and Dictyota spp. 

An ANOVA revealed significant effects of location (bank), and, overall, macroalgal 

cover was higher at the EFGB than the WFGB. The data for the EFGB was significantly 

different from that of the WFGB (p<0.02), suggesting variations between banks. Tukey–

Kramer a posteriori comparisons also showed that macroalgal cover was significantly

higher at the EFGB than the WFGB. 

CTB was the second-ranked non-coral category of substratum cover, ranging from 

approximately 10–15% over both banks from 2009–2010. An ANOVA revealed

significant differences between years (p<0.02), and overall, CTB cover was lower at the 

EFGB in 2009, but higher than the WFGB in 2010. Tukey–Kramer a posteriori

comparisons also showed that CTB cover was significantly higher in 2009 than in 2010. 

The Shannon-Wiener diversity index, H', was calculated from the species-specific

coral cover data from each transect in 2009 and 2010. The Shannon-Wiener diversity 

values at the EFGB were H’= 0.97 in 2009 and H’=0.95 in 2010. The values at the 

WFGB were H’= 0.98 in 2009 and H’=0.88 in 2010. The low values of H' overall

reflect the low diversity, yet high dominance by the Montastraea annularis species

complex.

Sclerochronology was used to measure the accretionary growth rates of Montastraea 

faveolata. Annual growth of M. faveolata at the EFGB averaged 0.55 mm/year (sample 

range = 0.30–0.76). At the WFGB, annual growth averaged 0.69 mm/year (range = 0.33–

0.97). When compared to the past three coring events (2003, 2005, and 2007), the 2010 

growth rate data were not substantially different. However, average growth rates at the 

EFGB decreased slightly from 2003 to 2010, while growth rates from the WFGB have 

remained relatively stable.

Random photographic transects were completed within the boundaries of the designated 

100 m2 monitoring study areas on each bank. These areas were originally selected in 

1988 because they appeared to be representative of the reef caps on each bank. Even 

now, over 20 years later, no inconsistencies among the reef character outside the 

designated study areas and the study areas themselves are apparent, suggesting that little 

long-term change is a critical component of ecosystem quality. 

xxii 
Lateral growth stations were photographed in 2009 and 2010 to measure changes in

Diploria strigosa colonies. Diploria strigosa is important at the FGB because it is the 

second largest contributor to coral cover. Net growth was positive over the 2009 to 

2010 monitoring period. 

Repetitive stations were photographed in 2009 and 2010 to monitor changes in specific 

coral reef locations over time. Like other data, repetitive quadrat data showed that coral

cover was consistently high during the 2009 to 2010 monitoring period, averaging

around 72% for both banks in all years (note that these stations were not selected as 

locations that were necessarily representative of the larger reef caps, so percent cover is 

likely not representative either). Macroalgae and CTB cover showed reverse patterns 

between banks and the incidences of bleaching, paling, and fish biting were low 

(ranging from 0.00–2.77% of area assessed). There was no evidence of coral disease

in any of the repetitive quadrats analyzed in 2009 or 2010. The coral assemblages 

remained stable at both banks, with the dominant corals being the Montastraea 

annularis species complex, Diploria strigosa, Porites astreoides, and M. cavernosa. 

In the 32–40 m deep repetitive quadrats (105–131 ft) at the EFGB, coral cover was

high, averaging approximately 81.96% between 2009 and 2010. The Montastraea 

annularis species complex and M. cavernosa were the dominant species in this depth 

range. CTB averaged 7.35%, and macroalgae averaged 10.42%.

The review of the 2009 and 2010 perimeter videos suggests that, in general, the coral

condition along the perimeter lines at the EFGB and WFGB study sites was comparable 

to that observed in past perimeter videos. The coral communities displayed low

levels of stress and high coral cover. The most distressed corals were affected by

incidences of paling and bleaching at the WFGB in 2010, followed by fish biting. No 

evidence of coral disease was observed in the perimeter videos. During the 2009 to 2010 

monitoring period, three tropical storms and two hurricanes occurred in the Gulf of 

Mexico, but there were few changes in community structure attributable to them. 

In addition to the annual data collection protocols, notable biological and

oceanographic events, such as wildlife observations, Acropora discoveries, and coral

health were qualitatively assessed and documented. April 20, 2010 marked the 

beginning of the Deepwater Horizon explosion, oil spill, and subsequent response, where 

an estimated 53,000 barrels per day escaped from the well before it was capped. While 

the oil spill caused damage to wildlife and marine habitats elsewhere in the Gulf of 

Mexico, no visible oil or oil-related impacts were observed in or near the FGBNMS.

Water quality parameters including seawater temperature and salinity were recorded at

the EFGB and WFGB using Sea-Bird 37-SMP MicroCAT datasondes (high-accuracy

conductivity and temperature recorders designed for long-term oceanographic 

deployment) from 2009 to 2010. HoboTemp thermographs were attached to each of 

the Sea-Bird platforms as backup recorders of water temperature. High seawater 

temperatures were observed during the late summer months of 2010, exceeding the 30°C 

coral bleaching threshold. 

xxiii
Fish surveys were conducted using the Bohnsack and Bannerot (1986) method in 2009 

and 2010. Fish surveys showed robust fish assemblages that were dominated by 

invertivorous fish, with healthy populations of herbivores, piscivores, and planktivores. 

An average of 57 fish species were observed per bank per year. Pomacentridae, 

Labridae, and Serranidae were the dominant fish families at both banks. Invertivores 

were the dominant fish guild, with Pomacentridae (damselfish) and Labridae (parrotfish 

and wrasses) representing the largest density. The size-frequency distributions of 

invertivores were non-normally distributed, with the majority of individuals being small 

damselfish. Interannual comparisons indicated generally stable assemblages; however, 

diversity measures were significantly different at the WFGB between 2009 and 2010. 

Following the pattern of coral species present at the FGB (low diversity compared to 

Caribbean reefs, but high coral cover), the fish assemblages reflect a similar trend of low 

diversity and high abundance (Pattengill-Semmens and Gittings 2003). 

Sea urchin surveys documented very low densities of Diadema antillarum at the EFGB

in 2009 (0.25 per 100 m2) and 2010 (0.5 per 100 m2). Higher densities were 

documented at the WFGB in 2009 (13.75 per 100 m2) and 2010 (11.0 per 100 m2).

These populations have not recovered to pre-1984 levels, which were at least 140 per 100 

m2 at the EFGB and 50 per 100 m2 at the WFGB (Gittings et al. 1998). No Panulirus argus 

(Caribbean spiny lobster) or Panuliris guttatus (spotted spiney lobster) were recorded

along transects at the EFGB or WFGB.

The FGB coral reefs remain in good condition and productive in comparison to reefs 

throughout the region. This may be in part to their remote location. Continued monitoring 

will document long-term changes in condition and will be useful for management 

decisions and future research focused on the dynamics of the robust benthic communities 

and the fish populations they support. 

xxiv 
CHAPTER 1.0: INTRODUCTION

1.1. CORAL REEF MONITORING AT THE FLOWER GARDEN BANKS 

NATIONAL MARINE SANCTUARY

The biotic assemblages of the Flower Garden Banks National Marine Sanctuary 

(FGBNMS) constitute a high coral and low algal cover reef community with a robust fish 

assemblage (Gittings et al. 1992; CSA 1996; Dokken et al. 1999, 2001, 2003; PattengillSemmens and Gittings 2003; Precht et al. 2006; Zimmer et al. 2010). Although coral 

species richness is lower at the FGBNMS than on most Caribbean reefs, 31 species of 

scleractinian corals (including deep coral species) have been documented at the 

FGBNMS (Schmahl et al. 2008). No significant long-term changes have been detected in 

coral cover or diversity at the FGBNMS from 1988 to 2008 (Zimmer et al. 2010), and 

probably not since the first measurements were taken in the mid-1970s (Gittings, 1998). In 

more than 20 years of continuous monitoring, the coral reefs of the FGBNMS have 

maintained high levels of coral cover, suffered minimally from hurricanes, coral 

bleaching and disease outbreaks, and supported relatively diverse and abundant fish and 

invertebrate populations. Though the rest of the Caribbean has experienced declines in 

zooxanthellate scleractinian coral cover (Gardner et al. 2003) and subsequent increases in 

macroalgal cover, the FGBNMS remains a stable coral reef system in the western Gulf of 

Mexico. These reefs, therefore, represent a natural laboratory for understanding the factors 

influencing stability and change in reef systems. The importance of the FGBNMS as 

representative western Atlantic coral reefs has been substantially elevated by the regional 

decline of corals. Consequently, the risk of loss (or estimated loss value) would be

elevated for the FGBNMS in the event of a severe industrial accident, expansion of the 

zone of influence of the Mississippi River, or other significant change in environmental 

conditions. 

The long-term monitoring program was initiated in 1988 by the Minerals Management 

Service (now the Bureau of Ocean Energy Management [BOEM]) to insure protective 

measures regulating potential impacts of offshore oil and gas development in the area

were effective. Gittings et al. (1992) established a single, 100 x 100 m study site at both 

the East and West Flower Garden Banks (EFGB and WFGB, respectively) to monitor 

benthic community structure from 1988 to 1991 using coral cover, relative dominance, 

species diversity, evenness, accretionary and encrusting growth rates, and water quality 

parameters as potential indicators of reef health. Comparisons between their 1988–1991 

results and those of previous studies from 1978–1982 (Rezak et al. 1985) showed no 

significant differences in any of the parameters, suggesting some degree of ecological 

stability over the period examined. During this time, coral cover was approximately 50% 

and dominated by the Montastraea annularis species complex (25%) and Diploria 

strigosa (8%) (Gittings et al. 1992). Gittings et al. (1992) considered spills from oil 

tankers, discharges of mud and drill cuttings during oil and gas exploration and 

production, noise from seismic surveys, and accidents on platforms leading to spills to be 

the greatest localized threats to these reefs.

1 
No significant changes in coral community structure were reported between 1992 and 

1995 by CSA (1996). However, variation in percent cover of individual coral species was 

detected between banks and between sampling years: 1992, 1994, and 1995. Minor 

individual coral bleaching was documented in 1990, 1992, and 1994, while 1995 was the 

first main bleaching event documented at the FGB coinciding with seawater temperatures 

in excess of 30°C (Hagman and Gittings 1992; Dokken et al. 1999, 2001, 2003). 

Montastraea cavernosa and Millepora alcicornis were the species most affected by 

bleaching, but post-bleaching mortality rates were low at 0.2%–2.8% (1992–1995) and 

were patchily distributed. The small-scale spatiotemporal variation reported by CSA did 

not appear to affect long-term landscape-scale trends in coral cover or composition. 

Dokken et al. (1999, 2003) continued the monitoring effort from 1996 through 2001 and 

documented no significant changes in coral growth or condition at the 100 x 100 m study 

sites at the EFGB and WFGB. Biodiversity inventories were conducted for algae and 

mollusks: 73 species of algae were documented as well as over 230 species of mollusks 

(Dokken et al. 2001, 2003). Fish assemblages were also documented (Pattengill 1998). 

Using the Atlantic and Gulf Rapid Reef Assessment (AGRRA) protocol in 1999, 

Pattengill-Semmens and Gittings (2003) observed high coral cover of approximately 50% 

at 20–28 m (66–92 ft), dominated by large coral colonies (mean diameter 81–93 cm or 

32–37 in), with a level of partial colony mortality (recent and long-dead portions of 

colonies) of only 13%. In concordance with earlier findings, turf was the dominant 

functional group of algae, whereas macroalgae accounted for less than 10% cover 

(Pattengill-Semmens and Gittings 2003). 

Continued monitoring of the study sites in 2002 and 2003 by Precht et al. (2006) highlighted 

the long-term stability of the coral reef communities. Coral cover was around 50% at both 

banks during those years, and no significant diseases were detected. The relative dominance 

of coral species also remained consistent with past findings. Diploria strigosa margins grew 

overall from 2001 to 2002, whereas a low sample size for 2002 to 2003 (due to replacement 

of monitoring stations in 2003) prevented firm conclusions during that time period.

Repetitive quadrat data from 2002 and 2003 revealed low prevalence of paling and 

bleaching (<0.61%) and no evidence of disease. Planimetry results showed an increase in 

surface area of selected corals at both banks. Oceanic water quality conditions prevailed at 

both banks in 2002 and 2003; however YSI maintenance issues produced data gaps, 

particularly for turbidity and PAR. Fish populations continued to be robust; however, 

Diadema and Panulirus abundance remained low. 

Zimmer et al. (2010) continued the monitoring effort from 2004 through 2008 and 

demonstrated continued stability of the coral reef community and associated fish 

populations at the 100 x 100 m study sites at the EFGB and WFGB. Coral cover averaged 

over 50% at both banks and the Montastraea annularis species complex remained the 

dominant component of coral cover. 

2 
On September 23, 2005 Hurricane Rita (Category 3, Saffir-Simpson Index) passed 

approximately 93 km (58 mi) from the EFGB on its route north to the mainland of the

United States. Two months later, Precht et al. (2008) conducted a post-hurricane 

assessment and reported that approximately 10% of the coral in repetitive quadrat stations 

at the EFGB were bleached. This was the highest level of bleaching reported for the FGB

since the bleaching event of 1995; however, there was no evidence of coral disease in any 

of the repetitive quadrats analyzed from 2004 through 2008. It should be noted that is was 

known the bleaching event was ongoing prior to the hurricane, but staff were not able to 

travel offshore to conduct a bleaching survey due to the storm. After Hurricane Rita 

passed through the Gulf of Mexico, the seawater temperature at the FGB dropped 

considerably which may have helped counteract the bleaching event. 

On September 12, 2008, Hurricane Ike (Category 3, Saffir-Simpson Index) passed 

directly over the EFGB. To monitor changes in coral reef community structure due to the 

passage of Hurricanes Rita and Ike, repetitive 8 m2 quadrats and perimeter video collected 

in November 2005 and November 2008, respectively, were assessed for hurricane 

damage. The results of the post-hurricane cruise conducted in November 2005 are 

published in a separate report (Precht et al. 2008). An estimated total area of 

approximately 2.3 m2 of coral was missing from the study-site repetitive quadrat

stations between June 2007 and November 2008 at the EFGB and WFGB, most likely

due to Hurricane Ike. The greatest loss in terms of both the number of missing coral

colonies and the total loss in area of coral cover occurred at the EFGB. Hurricane

impacts (i.e., dislodged colonies of Diploria strigosa) were only observed in perimeter

video at the EFGB. No obvious hurricane impacts were observed along perimeter lines

at the WFGB. The observed hurricane impacts were likely an underestimate of the actual 

hurricane damages because 1) only a portion of the perimeter surveys were comparable 

between June 2007 and November 2008 due to the loss of some corner locations and

shifts in line placement, and 2) the 2008 perimeter video was recorded at an angle of 90o 

to the substrate due to operator error (rather than at 45o as in previous surveys), providing 

a smaller area of view and fewer coral colonies for comparison.

1.2. THE FLOWER GARDEN BANKS NATIONAL MARINE SANCTUARY IN THE 

GULF OF MEXICO

1.2.1. Habitat Description

The Flower Garden Banks are located in the northwestern Gulf of Mexico and are part of 

a discontinuous arc of reef environments along the outer continental shelf (Rezak et al. 

1985; Figure 1.2.1). These coral reef-capped banks are the largest calcareous banks in the 

northwestern Gulf of Mexico (Bright et al. 1985). They contain the northernmost coral 

reefs in North America (Bright et al. 1984). Although coral and non-coral dominated 

communities exist on neighboring banks (e.g., Sonnier Bank, Stetson Bank, McGrail 

Bank), the reefs at Cabo Rojo in Mexico are the nearest shallow-water, true coral reefs in 

the Gulf of Mexico. 

3 
Figure 1.2.1. Map of the EFGB and WFGB in relation to the Texas-Louisiana continental 

shelf and other topographic features of the northwestern Gulf of Mexico. 

1. Stetson Bank, 2. Applebaum Bank, 3. Claypile Bank, 4. Coffee Lump Bank, 5. 

West Flower Garden Bank, 6. Horseshoe Bank, 7. East Flower Garden Bank, 8. 

MacNeil Bank, 9. 29 Fathom Bank, 10. Rankin Bank, 11. 28 Fathom Bank, 12. 

Bright Bank, 13. Geyer Bank, 14. Elvers Bank, 15. McGrail Bank, 16. Bouma Bank, 

17. Sonnier Bank, 18. Rezak Bank, 19. Sidner Bank, 20. Parker Bank, 21. 

Alderdice Bank, 22. Sweet Bank, 23. Fishnet Bank, 24. Jakkula Bank, 25. Ewing 

Bank, 26. Diaphus Bank. Red lines represent sanctuary boundaries 

(NOAA/FGBNMS).

The large-scale topographic features of the FGBNMS were created by geologic activity 

associated with salt diapirs of the Jurassic Louann Formation and consequent loading and 

uplifting of sedimentary rocks (Rezak 1981). Many such diapirs exist in the northern Gulf 

of Mexico and dozens form substantial submerged banks. The caps of some of the banks 

extend into the photic zone in clear oceanic waters, where conditions are ideal for 

colonization by species of corals, algae, invertebrates, and fish typical of coral reefs 

found in the Caribbean and western Atlantic (Figure 1.2.2). Although coral species 

richness is lower at the FGBNMS than on most Caribbean reefs, 31 species of 

scleractinian corals have been documented at the FGB (Schmahl et al. 2008) and 298 

species of tropical Atlantic fish have been reported sanctuary wide, including the 

deepwater communities (Hickerson, unpublished data). Oceanic salinity conditions 

prevail at the FGBNMS and range from 34 to 36 PSU, with water temperatures ranging 

from 18°C (in mid-February) to 30°C (in late August). Water clarity at the banks is 

excellent (commonly 30 m or more) providing ample light to photosynthesizing 

organisms. 

4 
Figure 1.2.2. Topographic contour map of the FGBNMS (NOAA/FGBNMS).

1.2.2. The East and West Flower Garden Banks

The EFGB (27o 54.5’ N, 93o 36.0’ W) is a pear-shaped dome located approximately 193 

km (120 mi) southeast of Galveston, Texas. The EFGB is 8.7 by 5.1 km (5.4 by 3.2 mi) 

in size, sloping from its shallowest point at 17 m (55 ft) to the terrigenous mud seafloor at 

a depth of 100–120 m (330–390 ft). The eastern and southern edges of the bank slope 

steeply whereas the northern and western edges descend more gently (Figure 1.2.3). The 

WFGB (27o 52.4’ N, 93o 48.8’ W) is an oblong-shaped dome located 20 km (12 mi) west 

of the EFGB and 172 km (107 mi) southeast of Galveston. It is 11.0 by 5.0 km (6.8 by 

5.0 mi) in size and larger than the EFGB (Figure 1.2.4). The two peaks that comprise the 

WFGB are aligned along an east-west axis. The WFGB study site is located on the 

eastern peak, which is 18 m (59 ft) at its shallowest. Coral species diversity at both banks 

is low, with 31 species from 18 genera represented (Schmahl et al. 2008), compared to 67 

species found on some Caribbean reefs (Goreau and Wells 1967). Shallow-water 

gorgonians and live acroporids have not been reported at the Flower Garden Banks in 

historical surveys. However, one colony of Acropora palmata was discovered in 2003 at 

the WFGB, but is in decline due to stressors such as damselfish predation, algal growth, 

etc. Another living colony of A. palmata was discovered at the EFGB, southeast of the 

study site in 2005 (Zimmer et al. 2006) and has a resident damselfish and minimal tissue 

loss. 

5 
Figure 1.2.3. Bathymetric map of the EFGB (NOAA/FGBNMS).

6 
Figure 1.2.4. Bathymetric map of the WFGB (NOAA/FGBNMS). 

Five habitat zones have been delineated at the EFGB and WFGB. This zonation scheme 

was updated by Schmahl et al. (2008) and includes the coral reef zone, the coral 

community zone, the coralline algal zones (which consist of coralline algal reefs and/or 

algal nodules), the deep coral zone, and the soft bottom zone. All monitoring at both 

banks was conducted within the coral reef zone.

1.3. BOEM AND FGBNMS PROTECTIVE MEASURES

Oil and gas activity in the vicinity of the FGBNMS has been ongoing since the 1970s. 

The former Minerals Management Service (MMS), of the U.S. Department of the Interior 

(USDOI), has regulated the development of the oil and gas industry on the Gulf of 

Mexico outer continental shelf. In 2010, MMS was reorganized and renamed the Bureau 

of Ocean Energy Management, Regulation and Enforcement (BOEMRE). In October 

2011, the agency was reorganized and the agency partnering in this monitoring effort is 

now called the Bureau of Ocean Energy Management (BOEM). 

7 
The first coral reef assessment of the FGB took place at the WFGB in 1972 (Bright and 

Pequegnat 1974). In 1973, BOEM (then the Bureau of Land Management) conducted a 

program of protective activities at the FGB coral reefs and sponsored numerous studies of 

the banks. The Topographic Features Stipulation (since 1973) was designed to protect 

sensitive, biological resources in the northwestern Gulf of Mexico, from the adverse 

effects of routine oil and gas activities (USDOI, MMS 2002) and in particular, from the 

discharge of drilling effluents. Since 1983, the stipulation has protected the biota of the 

FGB from physical damage associated with oil and gas activities including anchoring and 

rig emplacement, and potential toxic and smothering effects from drilling muds and 

cuttings discharges (USDOI, MMS 2002). The Stipulation defines a No Activity Zone 

(NAZ) around each of the banks using boundaries based on the “1⁄4, 1⁄4, 1⁄4 system”, where 

lease blocks are divided into smaller sections by successively breaking each section into 

quarters (USDOI, MMS 1998). The boundary of the NAZ overlaps the 100–120 m 

isobaths (328-394 ft) at the WFGB and the 100–130 m isobaths (328–427 ft) at the 

EFGB. No oil or gas structures, drilling rigs, pipelines, or anchoring are allowed within 

the NAZ. The Stipulation also defines a “4 Mile Zone” outside of the NAZ, within which 

operators are to shunt all drill cuttings and drilling fluids to within 10 m (33 ft) of the 

seafloor. Note that the FGB are the only topographic features with a NAZ defined by the 

1⁄4, 1⁄4, 1⁄4 system. The NAZ of all other BOEM protected banks follow defined isobaths 

surrounding the bank. 

In addition to the protections provided by the BOEM, the FGB were designated a United 

States National Marine Sanctuary in 1992 (Code of Federal Regulations, 15 CFR Part 

992, Subpart L, Section 922.120). While certain exceptions exist for oil and gas 

operations, the FGBNMS regulates, restricts and prohibits: 

(1) anchoring or mooring of all vessels within the sanctuary boundaries; 

(2) discharge of any material or matter within the sanctuary boundaries; 

(3) any alteration of the seabed within the sanctuary boundaries; 

(4) any injury or removal or attempt of injury or removal of any living or 

non-living sanctuary resource; 

(5) taking of marine mammals and sea turtles;

(6) possessing or using within the sanctuary boundaries any fishing gear 

except conventional hook and line gear; and 

(7) possessing or using explosives within the sanctuary boundaries or 

releasing electrical charges within the sanctuary boundaries. 

In July of 2001, the United States delegation to the International Maritime Organization 

(IMO), submitted a proposal to ban anchoring in FGBNMS for vessels greater than 30.48 

m (100 ft). The IMO, out of concern for impacts to corals, modified the United States’ 

proposal to prohibit all anchoring, but vessels 100 ft and under would be allowed to moor 

using existing sanctuary mooring buoys. The new international measure also ensured that 

no-anchoring zones are marked on all charts internationally. Code of Federal 

Regulations, 15 CFR Part 922.122 amended sanctuary regulations to align with IMO noanchor rule within the sanctuary. 

8 
From 1988 to 1995, the BOEM monitored the FGBNMS coral reefs to detect any 

incipient changes that may be caused by oil and gas activities, as well as by other

disturbances (Gittings et al. 1992; Gittings 1998). From 1996 until 2008, the FGBNMS 

and the BOEM partnered to continue the long-term monitoring of the FGB through a 

competitive contract. Since 2009, the FGBNMS has conducted the long-term monitoring

and the BOEM continues to support half of the monitoring effort through an interagency 

agreement contract with the FGBNMS. The decision to take on this contract in-house was 

driven mainly by the acquisition of the FGBNMS research vessel (R/V) Manta. This 

vessel was task-designed, and allows the FGBNMS to be efficient and effective in its 

research abilities. The FGBNMS has also built up a team of NOAA scientific divers and 

researchers to conduct both the field work and analysis of the monitoring data. 

1.4. DEEPWATER HORIZON

On April 20, 2010 Deepwater Horizon, a semi-submersible offshore drilling rig operating 

in lease block Mississippi Canyon 252, exploded; this resulted in the deaths of eleven 

people. More than 170 million gallons of crude oil were released into the Gulf of Mexico 

over a three-month period (NOAA 2012). Over 1 million gallons of dispersant were used, 

mostly Corexit 9500. On July 15, 2010, the leak was stopped by capping the gushing 

wellhead after it had released about 4.9 million barrels of crude oil. An estimated 53,000 

barrels per day escaped from the well before it was capped (McNutt et al. 2011). On 

September 19, 2010, the relief well process was successfully completed, and the federal 

government declared the well to be successfully plugged. The Deepwater Horizon

explosion resulted in the largest accidental marine oil spill in the history of the United 

States petroleum industry. 

NOAA acted as the lead science support agency to the United States Coast Guard during 

the spill response. The spill site was approximately 520 km (323 miles) from the EFGB

(Figure 1.4.1.), but because of the size of the spill, the potential for impact to the 

resources of the sanctuary was of great concern. As part of the National Resource 

Damage Assessment (NRDA) activities, FGBNMS personnel were assigned to the 

shallow and deep water coral response groups. In addition, semi-permeable membrane 

devices (SPMDs) were deployed at the EFGB and WFGB, as well as Stetson and Sonnier 

Banks. 

While the oil spill caused damage to wildlife and marine habitats in other areas of the 

Gulf of Mexico, no visible oil or oil-related impacts have been observed in or near the 

FGBNMS. Furthermore, no hydrocarbon signatures were found in SPMDs at the 

FGBNMS. Nevertheless, the long-term monitoring data (including previous SPMD data)

would have provided a valuable baseline from which impacts could have been detected, if 

they occurred. 

9 
Figure 1.4.1. Map of the location of the Deepwater Horizon site in relation to the 

FGBNMS, including the maximum extent of observed oil from the spill

(NOAA). 

10 
CHAPTER 2.0: STUDY SITES

2.1. STUDY SITE METHODOLOGY

The FGB are located roughly 190 km (118 mi) offshore and are submerged in water 

deeper than 18 m (59 ft). The monitoring effort was conducted from the NOAA R/V

Manta. The benthos (with an emphasis on corals and algae) was examined along 

videographic transects and stationary repetitive photoquadrats. Sclerochronology was 

used to document the accretionary growth rate of specific coral colonies, and 

photography was used at permanent stations to monitor the lateral growth of corals.

General aspects of coral condition were documented along perimeter lines at the EFGB 

and WFGB. During each annual monitoring cruise, observations of general coral reef 

health, as well as notable biological and oceanographic events were qualitatively assessed 

and documented. Water quality was assessed to characterize the reef cap and water 

column environment of the FGBNMS. Fish surveys were conducted at randomly located 

stations and sea urchin and lobster surveys were conducted along the study site perimeter 

lines. 

2.1.1. 100 x 100 m Study Sites

Data were collected within the 100 x 100 m study sites at the EFGB and WFGB in 2009 

and 2010 (Table 2.1.1). Originally established in 1988, the general locations of the study 

sites are marked by permanent mooring buoys: FGBNMS permanent mooring No. 2 at 

the EFGB (27° 53’ 35.80” N, 93° 38’ 23.90” W) and mooring No. 5 at the WFGB (27° 

52’ 50.86” N, 93° 52’ 25.34” W). Figures 2.1.1 and 2.1.2 depict the topography of the 

EFGB and WFGB, respectively, along with the locations of the 100 x 100 m study sites.

Figures 2.1.3 and 2.1.4 depict the mooring locations at EFGB and WFGB, respectively.

Headings to corners from the mooring u-bolts were used to locate the corners (Table 

2.1.2). Divers installed measuring tapes to temporarily mark the perimeters of the study 

sites and diving reels were used to mark the north/south and east/west centerlines 

(hereafter referred to as the “crosshairs”). Establishment of the perimeter and crosshairs 

divided each 100 x 100 m study site into four quadrants. The lines aided divers in 

orientation/navigation and they allowed for efficient completion of monitoring tasks. 

Each dive team was supplied with detailed underwater maps of each study site. Master 

maps were updated on the dive vessel with new data, including station numbers, 

locations, replacements, and revisions. These revisions are reflected in the current site 

maps (Figures 2.1.5 and 2.1.6). 

11 
Table 2.1.1.

Cruise Dates at the EFGB and WFGB for 2009 and 2010

EFGB WFGB

August 2009 August 2009

August 2010 August 2010

October 2010

November 2010

Table 2.1.2.

GPS Coordinates for the EFGB and WFGB Study-Site Corner Markers

EFGB WFGB

Corner North West Corner North West

NE 27o54'32.8 93o35'48.1 NE 27o52'31.8 93o48'53.6

NW 27o54'32.2 93o35'51.6 NW 27o52'31.5 93o48'56.9

SE 27o54'29.6 93o35'48.6 SE 27o52'28.7 93o48'53.2

SW 27o54'30.1 93o35'52.1 SW 27o52'28.5 93o48'56.8

Metal rods were previously installed in the reef to mark the permanent monitoring stations. 

There are two types of permanent monitoring stations within the study sites: (1) lateral 

growth stations on Diploria strigosa colonies, which are marked by two short rods per 

station; and (2) repetitive quadrats, the centers of which are marked by 0.5 m (1.6 ft) tall 

rods. Eighty repetitive quadrats and 120 lateral growth stations were maintained at the 

EFGB and WFGB.

12 
Figure 2.1.1. Topographic map of the EFGB. Inset shows the 

locations of the corners of the study site 

(USDOI/GS 2001).

13 
Figure 2.1.2. Topographic map of the WFGB. Inset shows the locations of the 

corners of the study site (USDOI/GS 2001).

14 
Figure 2.1.3. Locations of EFGB mooring buoys (NOAA/FGBNMS).

15 
Figure 2.1.4. Locations of WFGB mooring buoys (NOAA/FGBNMS).

16 
Figure 2.1.5. Locations of monitoring stations at the EFGB, 2010 (NOAA/FGBNMS). 

17 
Figure 2.1.6. Locations of monitoring stations at the WFGB, 2010 (NOAA/FGBNMS). 

18 
2.1.2. EFGB Deep Repetitive Quadrat Stations 

Nine deep repetitive quadrat stations are located outside the 100 x 100 m study site at the 

EFGB. These deep stations were established in April 2003 by BOEM and NOAA FGBNMS 

staff for comparision with the shallower repetitive photostations already in place (Precht et 

al. 2005). The stations were located east of the EFGB study site at depths between 32 m and 

40 m (105 ft and 131 ft) (Figures 2.1.7 and 2.1.8; Precht et al. 2005).

Note that the southern EFGB#2 u-bolt, formerly used as a mooring site, is no longer used; 

however, it is still marked and used as a point of reference. Contour lines at 20, 30, and 40 

m (NOAA/FGBNMS).

Figure 2.1.7. Bathymetric map with the deep repetitive quadrat stations in relation to the 

permanent study site at the EFGB (32–40 m or 105–131 ft), established in April 

2003. 

19 
Figure 2.1.8. Map showing depth and relative locations of the nine deep repetitive 

quadrat stations at the EFGB. 

2.1.3. Study Site Rehabilitation 

In August 2009 and 2010, the monitoring team conducted site rehabilitation at the EFGB 

and WFGB study site. This was necessary to replace bent or missing rods at the lateral 

growth and repetitive quadrat stations, missing station photo tags, reinstall loose rods 

with epoxy, and install missing corner markers. Photostation tags were replaced on ten

stations in 2009 and six stations in 2010. Maps depicting the locations of monitoring 

stations at the EFGB and WFGB were updated with new data, including station numbers, 

locations, replacements, and revisions for the study sites after each monitoring period. 

These revisions are reflected in the current site maps (Figures 2.1.5 and 2.1.6). 

20 
CHAPTER 3.0: RANDOM TRANSECTS

3.1. RANDOM TRANSECT METHODOLOGICAL RATIONALE

To estimate the areal coverage of benthic components such as corals, sponges, and 

macroalgae, 10 m (33 ft) fiberglass transect tapes were positioned randomly within each 

study site. Each transect originated at a random location using a randomly generated 

point on the perimeter boundary, followed by a randomly generated kick cycle distance, 

then laid out in a random direction according to a set of randomly generated numbers.

Formerly conducted using still photography, the random transect methodology was 

changed to videography by principal investigators during the previous long-term 

monitoring contract period. Initial comparisons between still and video images were 

made in 2002 and 2003. Videography was then used in 2004 through 2008. It was used

for the random transects by FGBNMS in 2009, but a decision was made to fully 

transition to digital still photography in 2010 due to the higher level resolution of the 

photographs (versus still images captured from video). 

3.2. RANDOM TRANSECT FIELD AND LABORATORY METHODS

The desired design was four transects laid randomly within each quadrant of each study 

site, for a total of 16 transects. Although only 14 transects are required by the BOEM 

long-term monitoring Scope of Work (SOW), the research team strives to complete 16 

transects every year. In some cases, however, only 14 transects were photographed. For 

each quadrant, a random number was generated between 0 and 50 to indicate the starting 

location along the first boundary line encountered (e.g., if the random number is 27 and 

the corner of the transect tape is 0, the transect is started at 27 m along the transect tape, 

and if the lowest corner of the transect tape is 50, the transect is started at 77 m along the 

transect tape). A second random number was generated between 0 and 40 to determine 

the number of fin kicks perpendicular from the boundary line into the study site. It is 

assumed that it takes 40 fin kicks to swim 50 m, assuming no opposing current. This

randomly generated number of kicks provides the start location for the first transect. A 

randomly generated number between 0 and 360 was generated for the direction of the 10

m transect from the starting point, and a measuring tape was laid down to mark the 

transect. 

The subsequent survey starting point was determined with a second set of randomly 

generated numbers. First, the number of fin kicks, between 12 and 40, was determined to 

ensure the starting point was at least 15 m away from the previous location. Second, the 

compass heading, between 1 and 360°, was established for the direction of the next 10 m 

transect. If a quadrant boundary was encountered, the line was reflected at a 90° angle 

back into the quadrant. 

In 2009, a Sony PC1000 3 CCD digital video camera with L&M Housing and HID lights

was used to collect digital videography. A diver swam slowly along each transect, 

videotaping at a height of 0.4 m (1.31 ft) off the seafloor (Aronson et al. 1994; Murdoch 

21 
and Aronson 1999). The videographer maintained an approximate 0.4 m distance above 

the benthos using a weighted line attached to the video housing. The camera was aimed 

downward at a 90° angle to record the substratum. 

The video frames covered a 40 cm (15.75 in) wide swath along each of the 10 m (33 ft) 

transects, for a total area of 4 m2 (43 ft2) per transect, or a minimum of approximately 56 m

2

(600 ft2) videotaped per study site per year. Each video frame was approximately 40 x 27 

cm (16 x 11 in) or 1080 cm2 (167 in2). Non-overlapping video frames were captured from 

each of the 16 video transects using Apple Final Cut Pro®. The original transect 

videotapes were used to gain more detail on objects or different perspectives on specific 

still images. Substrate cover was assessed from all captured images, as described below. 

In 2010, a Canon Power Shot G11 digital camera was used in an Ikelite housing with a 28 

mm equivalent wet mount lens adaptor to replace the video method. A diver mounted the 

camera on an aluminum t-frame, which allowed the camera lens to be 0.65 m above the 

substratum (Figure 3.2.1). The camera was placed at intervals marked on the tape at

55.88 cm (22 in) apart producing 19 non-overlapping images along the 10 m transect. 

Each still frame image captured an 80 x 55 cm (31.45 x 21.65 in) area. This produced a 

total photographed area of 8.36 m2

 (89.99 ft2) per transect, or a minimum of 117.04 m

2

(1259.81 ft2

) photographed per study site per year. The area captured using the digital 

still images increased from previous years to conform to image size requirements stated 

in the SOW. 

Coral cover in the still frame images was analyzed using Coral Point Count with 

Microsoft® Excel® extensions (CPCe). CPCe is Windows®-based software that 

provides a tool for the determination of coral cover using transect photographs (Kohler 

and Gill 2006). A specified number of spatially random points are distributed on a 

quadrat image, and benthic species lying under these points are identified. Microsoft® 

Excel® spreadsheets can be created automatically to further analyze the data, and users 

can create their own customized code files pertinent to the cover of coral and other 

benthic species in their region of interest. CPCe is provided by the National Coral Reef 

Institute as freeware to researchers from scientific institutions and government agencies. 

Randomly placed dots were added to each frame using CPCe, for a total of at least 500 

dots per transect (Kohler and Gill 2006). The number of images obtained from each 

transect was divided by 500, and rounded up if necessary, to obtain the number of 

random points per photo in that transect. Organisms positioned beneath each random dot 

were identified as follows: corals, sponges, and macroalgae were identified to lowest 

possible taxonomic group (macroalgae included algae longer than approximately 3 mm 

and included thick algal turfs); and crustose coralline algae, fine turfs, and bare rock were 

grouped as “CTB.” The components of the CTB group were combined for analysis not 

because they are difficult to distinguish (they are easy to tell apart), but because they are 

habitat where settlement of coral or sponges could occur. Macroalgae is not included in 

this category since settlement has already occurred. The CTB grouping was originally 

used by Aronson and Precht (2000) to characterize reefs in the Caribbean. The “other” 

22 
categories included other live components (ascidians, fish, serpulids, etc.), sand, rubble, 

and unknown. The coverages of coral bleaching, paling, concentrated and isolated fish 

biting, and disease were also determined from random transects.

After each image was analyzed, the data were entered into project-specific Microsoft® 

Excel® spreadsheets. Quality assurance/quality control (QA/QC) for the video and 

photographic methods consisted of multiple, trained scientific divers diving together on 

the study sites and identifying corals and other taxa. Statistical comparisons of 

identifications by person (no statistically significant differences) were conducted to 

confirm the same identifications in the videos and photographs to ensure that they agreed 

on species identifications within the frames.

Figure 3.2.1. A diver with camera mounted on aluminum t-frame taking random transect 

photographs. The line to the diver’s left is a study site boundary line

(NOAA/FGBNMS).

3.3. STATISTICAL ANALYSES OF THE RANDOM TRANSECT DATA

Percent coverage was calculated for each transect from the 500 analyzed points for each

of the taxa and benthic categories discussed in section 3.2. Factor plots were produced in 

CPCe to compare the average percent cover of major substrate types and coral species

between reefs and through time. Previous examination of means and variances, using 

different numbers of random dots, suggested that 500 dots per transect provided 

required accuracy and precision for estimates of the coverage of benthic components, 

23 
regardless of the transect length (Zimmer et al. 2010). Two-way Analyses of Variance

(ANOVAs) were performed to test the null hypotheses that the response variables of 

interest did not differ between banks or among years. ANOVAs were calculated for each

substratum variable with the statistical software JMP version 9.0. 

3.4. RANDOM TRANSECT RESULTS

The point counts from the random transects were grouped into four major functional 

categories (coral, sponge, macroalgae, and CTB) and expressed as percent cover ± 

standard error. In general, random transect results from 2009 and 2010 revealed high 

coral cover, followed by macroalgae cover (thick turfs and fleshy macroalgal species) 

and relatively low levels of CTB (crustose coralline algae, fine turf algae, and bare rock). 

The sponge cover was very low as not many macrosponges are found at the FGB; 

however, encrusting sponges are common but do not show up in photographs because 

they are generally found on the underside of boulders. High levels of coral cover were 

consistent with past monitoring results, however, levels of macroalgae increased from 

previous monitoring periods, while levels of CTB decreased (Figure 3.4.1). 

0

10

20

30

40

50

60

70

80

90

100

2009 EFGB 2010 EFGB 2009 WFGB 2010 WFGB

Percent Cover

Bank and Year

Percent Cover of Four Functional Catagories at the Flower 

Garden Banks, 2009 - 2010 Coral

Sponge

Macroalgae

CTB

CTB: crustose coralline algae, fine turf algae, and bare rock. The “macroalgae” category 

includes thick turfs as well as fleshy macroalgal species. Values are calculated from the 

random transect photographs.

Figure 3.4.1. Percent cover (error bars are ± SE) of four functional categories of sessile 

benthos at the FGB from 2009 and 2010. 

At the EFGB, coral cover remained stable from 2009 to 2010 (53.35% ± 4.17 to 54.49% 

± 3.69), and the sponge cover remained extremely low for both years (0.13% ± 0.05 to

0.25% ± 0.13) (Figure 3.4.2 and Table 3.4.1). Macroalgae cover (mainly fleshy algae,

24 
turfs, and Dictyota spp.) remained stable from 2009 to 2010 (31.85% ± 3.49 to 32.94% ± 

3.05), as did CTB cover (13.32% ± 1.07 to 11.15% ± 0.86). Consistent with past 

monitoring results, the Montastraea annularis species complex (MASC), containing M. 

annularis, M. faveolata, and M. franksi, continued to dominate the EFGB in 2009 

(29.52% ± 12.99) and 2010 (38.89% ± 3.84). Diploria strigosa (10.08% ± 2.54 and 

9.38% ± 1.72 in 2009 and 2010, respectively) and Porites astreoides (4.93% ± 1.29 in 

2009 and 5.11% ± 0.69 in 2010) were the next most abundant species (Figure 3.4.2 and 

Table 3.4.1). The remaining coral cover was made up of eleven species, none of 

which exceeded 4.0% individually in either 2009 or 2010 (Table 3.1.1). ShannonWeiner diversity values remained consistent at the EFGB (H’=0.97 and H’=0.95 in 2009

and 2010, respectively). 

0 5 10 15 20 25 30 35 40 45 50

MASC

Diploria strigosa

Porites astreoides

Montastraea cavernosa

Colpophyllia natans

Unidentifiable coral

Agaricia spp.

Stephanocoenia intersepta

Millepora alcicornis

Madracis spp.

Mussa angulosa

Siderastrea radians

Colpophyllia amaranthus

Siderastrea siderea

Scolymia spp.

Paracyanthus pulchellus

Oxysmilia spp.

Percent Cover

Coral Species

Percent Cover of the Coral Taxa at the East Flower Garden Bank, 

2009-2010

2009

2010

Figure 3.4.2. Percent cover (+ SE) of the dominant coral taxa at the EFGB from 2009 and 

2010. Values are calculated from the random transect photographs. 

At the WFGB, the 2009 coral cover remained consistent with previous years (53.84% ± 

3.73); however, coral cover was higher in 2010 (65.95% ± 2.85). The sponge cover 

remained extremely low for both years at the WFGB (0.10% ± 0.05 to 0.25% ± 0.10) 

(Figure 3.4.3 and Table 3.4.1). Macroalgae cover decreased from 2009 to 2010 (30.03% ± 

2.63 to 22.03% ± 2.00), while CTB cover remained relatively stable (14.93% ± 1.52 to 

10.20% ± 1.03). As with the EFGB, the Montastraea annularis species complex

continued to dominate the WFGB in 2009 (27.25% ± 3.32) and 2010 (45.98% ± 3.68). 

Diploria strigosa (10.84% ± 2.70 and 5.39% ± 0.94 in 2009 and 2010, respectively) and

Porites astreoides (3.85% ± 0.64 in 2009 and 3.20% ± 0.62 in 2010) were the next most 

abundant species (Figure 3.4.3 and Table 3.4.1). The remaining coral cover was made

up of eleven species, none of which exceeded 6.0% individually in either 2009 or 

25 
2010 (Table 3.1.1). Shannon-Weiner diversity values were higher at the WFGB in 

2009 than in 2010 (H’=0.98 and H’=0.88 in 2009 and 2010, respectively). 

0 5 10 15 20 25 30 35 40 45 50

MASC

Diploria strigosa

Porites astreoides

Montastraea cavernosa

Colpophyllia natans

Unidentifiable coral

Agaricia spp.

Stephanocoenia intersepta

Millepora alcicornis

Madracis spp.

Mussa angulosa

Siderastrea radians

Colpophyllia amaranthus

Siderastrea siderea

Scolymia spp.

Paracyanthus pulchellus

Oxysmilia spp.

Percent Cover

Coral Species

Percent Cover of the Coral Taxa at the West Flower Garden Bank, 

2009-2010

2009

2010

 Values are calculated from the random transect photographs.

Figure 3.4.3. Percent cover (+ SE) of the dominant coral taxa at the WFGB from 2009 and 

2010. 

26 
Table 3.4.1.

Cover of Benthic Categories in Random Transects at the EFGB and WFGB from 2009 and 2010

Values are expressed as percent cover ± SE.

Cover Category 2009 EFGB 2010 EFGB 2009 WFGB 2010 WFGB

Coral

Montastraea annularis

species complex 29.52 ± 12.99 33.89 ± 3.84 27.25 ± 3.32 45.98 ± 3.68

Diploria strigosa 10.08 ± 2.54 9.38 ± 1.72 10.84 ± 2.70 5.39 ± 0.94

Porites astreoides 4.93 ± 1.29 5.11 ± 0.69 3.85 ± 0.64 3.20 ± 0.62

Montastraea cavernosa 3.58 ± 1.22 3.71 ± 1.46 4.03 ± 0.97 5.09 ± 1.12

Colpophyllia natans 3.21 ± 1.00 2.45 ± 0.63 4.26 ± 1.38 1.45 ± 0.42

Unidentifiable coral 0.65 ± 0.25 0.96 ± 0.20 0.19 ± 0.07 0.62 ± 0.23

Agaricia spp. 0.34 ± 0.17 0.21 ± 0.08 0.24 ± 0.07 0.18 ± 0.08

Stephanocoenia intersepta 0.30 ± 0.13 0.27 ± 0.12 0.94 ± 0.27 2.74 ± 2.03

Millepora alcicornis 0.25 ± 0.15 0.24 ± 0.10 0.41 ± 0.16 0.73 ± 0.32

Madracis spp. 0.22 ± 0.07 0.31 ± 0.17 0.05 ± 0.04 0.06 ± 0.03

Mussa angulosa 0.10 ± 0.07 0.12 ± 0.06 0.32 ± 0.14 0.13 ± 0.07

Siderastrea radians 0.09 ± 0.06 0.00 0.00 0.00

Colpophyllia amaranthus 0.00 0.00 0.00 0.00

Siderastrea siderea 0.03 ± 0.02 0.10 ± 0.06 1.44 ± 1.33 0.33 ± 0.31

Scolymia spp. 0.00 0.00 0.02 ± 0.02 0.00

Paracyanthus pulchellus 0.00 0.00 0.00 0.00

Oxysmilia spp. 0.00 0.01 ± 0.01 0.00 0.04 ± 0.02

Total Coral 53.35 ± 4.17 54.49 ± 3.69 53.84 ± 3.73 65.95 ± 2.85

Sponge

Agelas clathrodes 0.03 ± 0.02 0.09 ± 0.06 0.06 ± 0.05 0.01 ± 0.01

Arolochroia 

(Pseudoceratina) crassa 0.01 ± 0.01 0.05 ± 0.05 0.01 ± 0.01 0.06 ± 0.06

Ectyoplasia ferox 0.00 0.03 ± 0.03 0.00 0.03 ± 0.02

Ircinia strobilina 0.03 ± 0.03 0.00 0.00 0.00

Mycale laxissima 0.00 0.00 0.00 0.01 ± 0.01

Neofibularia nolitangere 0.03 ± 0.03 0.00 0.03 ± 0.03 0.00

Unidentifiable Sponge 0.01 ± 0.01 0.07 ± 0.07 0.00 0.07 ± 0.06

Xestospongia muta 0.01 ± 0.01 0.01 ± 0.01 0.00 0.04 ± 0.03

Encrusting sponge 0.24 ± 0.11 0.28 ± 0.13 0.26 ± 0.06 0.16 ± 0.06

Spirastrelle cunctatrix 0.00 0.07 ± 0.04 0.01 ± 0.01 0.11 ± 0.06

Unidentifiable Encrusting 

Sponge 0.00 0.03 ± 0.03 0.00 0.00

Total Sponge 0.13 ± 0.05 0.25 ± 0.13 0.10 ± 0.05 0.25 ± 0.10

CTB

27 
Cover Category 2009 EFGB 2010 EFGB 2009 WFGB 2010 WFGB

Bare Substrate 6.13 ± 1.02 6.73 ± 0.77 7.34 ± 0.98 4.83 ± 0.78

Crustose Coralline Algae 2.55 ± 0.41 3.06 ± 0.38 1.46 ± 0.41 3.21 ± 0.49

Fine Turf 4.64 ± 0.42 1.36 ± 0.24 6.14 ± 0.61 2.16 ± 0.35

Total CTB 13.32 ± 1.07 11.15 ± 0.86 14.93 ± 1.52 10.20 ± 1.03

Macroalgae

Fleshy algae 13.53 ± 2.27 17.64 ± 2.18 19.74 ± 2.03 9.18 ± 1.00

Thick turf algae 11.85 ± 1.48 1.44 ± 0.35 6.65 ± 1.28 0.70 ± 0.29

Dictyota spp. 6.00 ± 1.05 9.95 ± 1.74 1.04 ± 0.25 1.62 ± 0.56

Lobophora variegata 0.48 ± 0.11 3.89 ± 0.56 2.60 ± 0.36 10.53 ± 2.08

Filamentous Algae 0.00 0.01 ± 0.01 0.00 0.00

Total Macroalgae 31.85 ± 3.49 32.94 ± 3.05 30.03 ± 2.63 22.03 ± 2.00

Other

Shadow 5.96 ± 0.65 6.66 ± 0.69 6.27 ± 1.12 7.43 ± 1.02

Sand 0.57 ± 0.33 0.21 ± 0.12 0.57 ± 0.25 0.58 ± 0.18

Substrate Rubble 0.21 ± 0.17 0.31 ± 0.20 0.12 ± 0.05 0.43 ± 0.15

Fish 0.04 ± 0.02 0.03 ± 0.02 0.01 ± 0.01 0.00

Invertebrate 0.28 ± 0.05 0.20 ± 0.06 0.11 ± 0.04 0.29 ± 0.06

Coral Condition (occurences in coral)

Bleached Coral 0.14 ± 0.10 0.02 ± 0.02 0.16 ± 0.13 6.98 ± 2.06

Paling Coral 0.76 ± 0.46 0.36 ± 0.12 0.39 ± 0.21 11.09 ± 3.33

Concentrated Fish Biting 0.00 0.05 ± 0.05 0.09 ± 0.05 0.02 ± 0.02

Isolated Fish Biting 0.03 ± 0.03 0.00 0.06 ± 0.06 0.02 ± 0.02

The combined data collected from 2009 and 2010 showed persistence of the Montastraea 

annularis species complex as the dominant coral species at the EFGB and WFGB (Table 

3.4.1). Likewise, Diploria strigosa remained the second-most prevalent coral species

during this time period. Porites astreoides and M. cavernosa were consistently the 

third and fourth most dominant corals at the EFG, while M. cavernosa was the third 

dominant and Porites astreoides was the fourth dominant coral at WFGB. The average 

coral cover between the EFGB and WFGB determined by random transect data was 

56.95% for the 2009 and 2010 reporting period. Appendix 1 in Volume II of this report 

contains the random transect data from 2009 and 2010 at the FGB. 

In past monitoring years, the Montastraea annularis species complex (M. annularis, M. 

faveolata, and M. franksi) has been difficult to differentiate using the photographic and 

videographic techniques employed in this study (primarily because the scale of the 

photograph does not allow visualization of the entire colony formation, which can be 

very helpful for identification). However, with improved digital photography resolution 

and the familiarity of the FGBNMS staff with the photostations, the components of the 

Montastraea annularis species complex were individually indentified. At the EFGB and 

WFGB, the Montastraea annularis species complex is dominated by M. franksi, followed 

28 
by M. faveolata and M. annularis. At the EFGB, M. franksi percent cover remained 

stable from 2009 to 2010 (18.50% ± 8.90 to 20.20% ± 11.00), and M. faveolata (3.80% ± 

5.42 to 3.00% ± 3.92) and M. annularis (2.84% ± 5.42 to 1.47% ± 4.34) remained stable 

as well (Figure 3.4.4). At the WFGB, M. franksi percent cover of the MASC was 

significantly higher in 2010 (21.44% ± 9.81 in 2009 and 37.54% ± 12.98 in 2010), while 

M. faveolata (1.65% ± 2.62 to 1.40% ± 4.09) and M. annularis (1.18% ± 3.88 to 0.48% ± 

1.73) remained stable (Figure 3.4.5). Approximately 5.5% of the MASC components 

were unable to be differentiated, which may be due to MASC hybridization or genotypic 

variation. 

0 5 10 15 20 25 30 35

Montastraea annularis

Unidentified MASC

Montastraea faveolata

Montastraea franksi

Percent Cover

Coral Species

Percent Cover of the Coral that Comprises MASC at the East 

Flower Garden Bank, 2009-2010

2009

2010

 Values are calculated from the random transect photographs.

Figure 3.4.4. Percent cover (+ SE) of the coral species that comprise the MASC at the EFGB 

from 2009 and 2010 (note scale difference from Figure 3.4.5). 

29 
0 10 20 30 40 50 60

Montastraea annularis

Unidentified MASC

Montastraea faveolata

Montastraea franksi

Percent Cover

Coral Species

Percent Cover of the Coral that Comprises MASC at the West 

Flower Garden Bank, 2009-2010

2009

2010

 Values are calculated from the random transect photographs.

Figure 3.4.5. Percent cover (+ SE) of the coral species that comprises the MASC at the 

WFGB from 2009 and 2010 (note scale difference from Figure 3.4.4). 

In the 2009 and 2010 random transects, the incidences of bleaching, paling, and fish 

biting were low at both banks and coral disease was absent (Table 3.4.1). The percentage 

of corals impacted by isolated/concentrated fish biting at the EFGB and WFGB was 

minimal, ranging from 0.00 to 0.09%. Less than 1% of the coral cover analyzed in the 

random transects was diseased or bleached at the EFGB in 2009 and 2010 and at the 

WFGB in 2009; however, higher incidences of bleaching (~7%) and paling (~11%) were 

observed at the WFGB in 2010 (Figure 3.4.6). 

30 
0

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

2009 EFGB 2010 EFGB 2009 WFGB 2010 WFGB

Percent of Coral

Bank and Year

Bleaching and Paling of Corals from Percent Cover Analysis 

at the Flower Garden Banks, 2009-2010 

Bleached

Paling

Values are calculated from coral cover in the random transect photographs.

Figure 3.4.6. Percent of coral observed to pale and bleach (+ SE) at the FGB in 

2009 and 2010. 

In 2009 and 2010, 58% of the bleached coral cover was in the Montastraea annularis

species complex (Figure 3.4.7). Twenty-eight percent was Montastraea cavernosa (28%). 

Of the cover that was observed to pale, 74% was in the Montastraea annularis species 

complex, followed by 14% for Montastraea cavernosa (Figure 3.4.8). The elevated 

bleaching and paling levels at the WFGB in 2010 will be further discussed in the 

Discussion section of this report. 

31 
Montastraea 

annularis 

species complex

58%

Montastraea 

cavernosa

28%

Other 

Species

14%

Species Composition from Percent Cover Analysis of Bleached 

Coral at the Flower Garden Banks, 2009-2010

Figure 3.4.7. Species composition of bleached coral from percent cover random 

transect analysis at the FGB from 2009 and 2010. 

Montastraea 

annularis 

species complex

74%

Montastraea 

cavernosa

14%

Other 

Species

12%

Species Composition from Percent Cover Analysis of Paling 

Coral, at the Flower Garden Banks, 2009-2010

Figure 3.4.8. Species composition of paling coral from percent cover random transect

analysis at the FGB from 2009 and 2010. 

The point count data that were grouped into the four major functional categories (coral, 

sponge, macroalgae, and CTB) and expressed as percent covers were analyzed by two32 
way analysis of variance (ANOVA), with Site (EFGB and WFGB) and Year (2009 and 

2010) as fixed factors. Prior to analysis, the data were tested for conformity to the 

parametric assumptions of normality and homogeneity of variances. All data were log 

transformed to assume a normal distribution. 

A two-way ANOVA on the proportional cover of all living hard corals (Scleractinia and 

Milleporina) showed no significant effect of Site or Year, and the Site x Year interaction

was also not significant (Table 3.4.2 A). A two-way ANOVA on the proportional cover

of sponges showed no significant effects of Site or Year, and the Site x Year interaction

was also not significant (Table 3.4.2 B). 

A two-way ANOVA on the proportional cover of macroalgae showed no significant

effects of Year, and the Site x Year interaction was also not significant, however, the 

effects of Site were significant (Table 3.4.2 C). Overall, macroalgal cover was higher at

the EFGB than the WFGB during the study period. Because the effect of Site was 

significant, a simple one-way ANOVA was performed on the macroalgal data for each

bank separately to examine Site variations. The data for the EFGB was significantly 

different from that of the WFGB (F=10.59, df=1, 62, P-value <0.0191). Tukey–Kramer a 

posteriori comparisons also showed that macroalgal cover was significantly higher at the 

EFGB than the WFGB. 

A two-way ANOVA on the fourth cover category, CTB ( crustose coralline algae,

fine algal turfs and bare rock), showed no significant effects of Site, and the Site x 

Year interaction was also not significant, however, the effects of Year were significant 

(Table 3.4.2 D). Overall, CTB cover was lower at the EFGB in 2009, but higher at the 

WFGB in 2010. Because the effect of Year was significant, a simple one-way ANOVA

was performed on the CTB data to examine year-to-year variations. The data for 2009 

was significantly different 2010 (F=6.14, df=1, 62, P-value <0.0177). Tukey–Kramer a 

posteriori comparisons also showed that CTB cover was significantly higher in 2009 

than in 2010. 

33 
Table 3.4.2.

Results of ANOVA on Proportional Cover Estimates from Random Transects from 2009 and 2010 

Comparisons of groups where ANOVA was performed are in bold with appropriate P-values 

where significant.

(A) Hard Corals (log transformed)

Source Sum of Squares df Mean Square F-ratio P-value

Site 0.3308 1 2.18 0.15

Year 0.3866 1 2.55 0.12

Site*Year 0.0650 1 0.26 0.43 0.52

Error 8.9455 59 0.15

Total 9.7267 62

(B) Sponges (log transformed)

Source Sum of Squares df Mean Square F-ratio P-value

Site 0.0006 1 0.0109 0.9173

Year 0.0978 1 1.7403 0.1922

Site*Year 0.0044 1 0.0346 0.0786 0.7802

Error 3.3170 59 0.0562

Total 3.4208 62

(C) Macroalgae (log transformed)

Source Sum of Squares df Mean Square F-ratio P-value

Site 0.7389 1 5.8051 0.0191

Year 0.2823 1 2.2177 0.1418

Site*Year 0.4405 1 0.4925 3.4608 0.0678

Error 7.5096 59 0.1273

Total 8.9870 62

(D) CTB (log transformed)

Source Sum of Squares df Mean Square F-ratio P-value

Site 0.0082 1 0.0618 0.8046

Year 0.7937 1 5.9532 0.0177

Site*Year 0.0681 1 0.2920 0.5107 0.4776

Error 7.8657 59 0.1333

Total 8.7418 62

Finally, the Shannon-Wiener diversity index, H', was calculated from the speciesspecific coral cover data from each transect. The three species of the Montastraea 

annularis species complex were combined for the calculation. The Shannon-Wiener

diversity index is used to give a measure of both species numbers and evenness (the 

apportionment of individuals among species), resulting in a biodiversity measure that is 

34 
useful when comparing similar habitats. The Shannon-Wiener diversity values at the 

EFGB were H’= 0.97 in 2009 and H’=0.95 in 2010. The values at the WFGB were H’= 

0.98 in 2009 and H’=0.88 in 2010. The low values of H' reflect the low diversity, yet 

high coral cover, particularly of the strong dominance of the Montastraea annularis

species complex.

In summary, combined mean coral cover averaged approximately 56% at the EFGB and

WFGB in the period 2009 through 2010. These values were consistent with 

measurements of coral cover above 50% at the FGB in previous years (Dokken et al. 

2003; Precht et al. 2006) and they are high compared to other western Atlantic reefs (e.g., 

Aronson et al. 1994; Gardner et al. 2003). Macroalgae ranked second behind the corals, 

and sponge cover was extremely low. This pattern was consistent with recent results from 

the EFGB and WFGB (Aronson et al. 2005; Precht et al. 2006; Zimmer et al. 2010) and

previous work in the Caribbean (Aronson and Precht 2000). 

3.5. RANDOM TRANSECT DISCUSSION

In a global trend of declining coral reef health and cover, the FGB continues to support 

high coral cover compared to other reefs of the western Atlantic and Caribbean region 

(Aronson et al. 1994, 2005; Gardner et al. 2003; AGRRA 2003; Pina Amargós et al. 

2008; ONMS 2011; Steneck et al. 2011) (Table 3.5.1). Gardner et al. (2003) reported

the regional decline of corals across the Caribbean basin over the last three decades,

with the average hard coral cover on reefs decreasing from approximately 50% to 10%. 

Natural and anthropogenic factors, including storm events, temperature stress, disease, 

predation, overfishing, sedimentation, eutrophication, and habitat destruction have all

played a part in the decline (Aronson and Precht 2001; Rogers and Beets 2001; Gardner

et al. 2003). 

Table 3.5.1.

Percentage of Coral Cover on Living Reefs in the Western Atlantic and Caribbean Region

Location Percent Coral Cover Source

Flower Garden Banks NMS 56 this report

Netherlands Antilles 10–47 AGRRA 2003

St. Vincent Grenadines 29–44 AGRRA 2003

Bonaire 38 Steneck et al. 2011

Turks and Caicos up to 30 AGRRA 2003

Florida Keys NMS 3–20 ONMS 2011

Cayman Islands 21 AGRRA 2003

Jardin de la Reina, Cuba 7–19 Pina Amargós et al. 2008

Akumal, Mexico 17 AGRRA 2003

Caribbean reefs that have historically displayed high coral cover are showing declines, 

mainly due to algae competition, bleaching and/or coral disease. Bonaire reported a 

decrease in coral cover from 38% to 10% in 2011 (Steneck et al. 2011), and the Florida 

Keys National Marine Sanctuary is maintaining approximately 7% coral coverage with 

35 
ranges from about 3% to 20% (ONMS 2011). The Cuban reefs within Jardin de la Reina 

range from 7% to 9% cover (Pina Amargós et al. 2008). In contrast, coral cover at the 

FGB has remained relatively stable over time (Figure 3.5.1). Univariate analysis of the 

random transect data revealed that the average coral cover at the EFGB and WFGB 

remained high from 2009 to 2010 (56.95%). Coral cover was also similar to values from

earlier studies, (Dokken et al. 1999, 2001; CSA 1996; Gittings et al. 1992), highlighting

the stability of the coral assemblage over time (Figure 3.5.1). 

Some reasons for the exceptional condition of the FGB include (1) water depth of 

the reefs, which buffers the reef cap from the effects of storm waves and variable 

seasurface temperatures; (2) the remote offshore location, which limits human access

and exposes these reefs consistently to oligotrophic, oceanic waters; (3) healthy grazer 

populations; and (4) protective federal regulations, which prevent hydrocarbon-related

effects, as well as effects from anchoring, fishing and recreational diving (Aronson et

al. 2005). The importance of the FGB, in terms of the Atlantic coral reef system as a 

whole, has been substantially elevated because of the regional decline of corals. 

Consequently, it could be argued that the estimated loss value would be higher for the 

FGB in the event of a severe industrial accident, expansion of the zone of influence

of the Mississippi River, or other significant change in environmental conditions. 

0

10

20

30

40

50

60

70

80

Percent Cover

Year

Percent Coral Cover at the Flower Garden Banks, 1978 - 2010

East Bank

West Bank

a.TAMU, b. CSA, c. TAMU-CC, d. PBSJ, e. FGBNMS

LPI Film Photography Video Digital 

Photography

No percent cover data were reported in 1993. Data for 1978 to 1982 from Gittings

et al. (1992), who reported data from Kraemer (1982); for 1988 to 1991 from

Gittings et al. (1992); for 1992 to 1995 from Continental Shelf Associates, Inc. (CSA)

(1996); for 1996 to 2001 from Dokken et al. (2003); 2002 to 2008 from PBS&J (Precht 

et al. 2006, 2008b); and FGBNMS for 2009 and 2010 (Johnston et al. 2012).

Figure 3.5.1. Random transect mean percent coral cover at the EFGB and WFGB over

time, showing the consistently high coral cover. 

36 
3.5.1. EFGB Comparison from 2009 to 2010

The random transect data for the EFGB showed similar values for coral, sponge, CTB, 

macroalgae cover and H' in both years. The three dominant coral taxa—the Montastraea 

annularis species complex, Diploria strigosa and Porites astreoides—varied little from 

2009 to 2010. The H' for the coral assemblage was low at the EFGB due to the low 

species-richness values and the dominance of a few species, namely the M. annularis

species complex and D. strigosa. The most noticeable pattern was consistent coral cover 

at the EFGB compared to the other sampling years.

3.5.2. WFGB Comparison from 2009 to 2010

The random transect data for the WFGB showed similar values for sponge, CTB, 

macroalgae cover and H' in both years. However, coral cover was significantly higher at 

the WFGB in 2010 (65.95% ± 2.85) than in 2009 (53.84% ± 3.73). Much of the variation

was likely due to the transect placement, rather than reflecting real variations. Past

studies have documented similar variations in relative abundance from year to year and

were often attributable to sampling or image analysis error (Dokken et al. 2003, 1999); 

however, photographs were analyzed multiple times in the CPCe software to ensure coral 

was identified correctly. It was determined that the randomness of the transect 

placements resulted in more areas containing high coral cover than in years past, resulting 

in an artifact from the 2010 sampling season. As with the EFGB, the three dominant 

coral taxa—the Montastraea annularis species complex, Diploria strigosa and Porites

astreoides—fluctuated to a negligible degree. Overall, the most noticeable pattern was 

higher coral cover in 2010, when compared to the other sampling years. 

3.5.3. Comparison of Random Transect Results from 1992 to 2008

A qualitative comparison of the dominant cover components from the random transects

showed interesting results for several cover categories: Montastraea annularis species

complex, macroalgae, and CTB. In 1996 and 1997, no data were recorded for the CTB 

category (known as reef rock from previous monitoring periods). 

The Montastraea annularis species complex showed an overall increase in cover 

during the period from 1992 to 2010 at the WFGB; with the highest cover being 

recorded in 2010 and an average of approximately 32% cover. Estimates varied at the

EFGB but remained consistently at or above 30% (Figure 3.5.2). Estimates for the M.

annularis species complex were slightly lower in 1996, 1999, and 2003 at the EFGB

and in 1996, 2000, 2004, 2007, and 2009 at the WFGB. Higher MASC cover was 

observed in 1994, 2000, and 2010 at the EFGB and 1994, 2006, and 2010 at the WFGB. 

Periods of lower coral cover generally coincided with increases in the algal component 

and decreases in the CTB category. The MASC increases generally coincided with

lower macroalgal cover. The results suggest that algal overgrowth can significantly 

affect estimates of underlying benthic cover, but due to the epemeral nature of algae

blooms, does not necessarily lead to long-term reductions in those populations. 

37 
 (A)

0

10

20

30

40

50

60

Percent Cover

Year

East Flower Garden Bank Percent Cover of Montastraea annularis 

species complex, Macroalgae, and CTB from 1992-2010

MASC

Macroalgae

CTB

 (B) 

0

10

20

30

40

50

60

Percent Cover

Year

West Flower Garden Bank Percent Cover of Montastraea annularis 

species complex, Macroalgae, and CTB from 1992-2010

MASC

Macroalgae

CTB

Figure 3.5.2. Percent cover of Montastraea annularis species complex,

macroalgae, and CTB from 1992 to 2010 at (A) the EFGB and (B) 

the WFGB.

38 
Local and regional weather patterns affect benthic communities like those at the

FGB. Changes in the frequency and severity of the El Niño–Southern Oscillation

(ENSO) have been partially responsible for transitions from coral-dominated

communities to algae-dominated reef systems in the Caribbean (Glynn 1984, 1993; 

Goreau and Hayes 1994; Wilkinson and Souter 2008). In 1987, 1995, and 1998, severe 

ENSO fluctuations affected the western Atlantic, causing large-scale coral bleaching,

subsequent coral mortality, and colonization of substrate by algae (Glynn 1984; McField

1999; Aronson et al. 2000). Widespread and severe coral bleaching also occurred in the 

Caribbean in 2005, but in the absence of an El Niño event (Wilkinson and Souter

2008). T he FGB, being a system where these severe effects have not been documented, 

provides an opportunity to dissect the community dynamics of coral cover,

macroalgae, and CTB. 

Macroalgae tend to be ephemeral, with different species becoming abundant under 

certain seasonal conditions (Diaz-Pulido and Garzon-Ferreira 2002). Algae cover at the

FGB, here represented primarily within the macroalgae category, remained relatively

low until 1999, never reaching more than 6.1% at either bank. It increased dramatically

in 1999 and while fluctuating, has remained comparatively high ever since. Average 

cover has been 15% at EFGB and 13% at WFGB (Table 3.5.2 and Figure 3.5.2).

Concurrent with high algae cover, the CTB category was lower at the EFGB in 1999, 

2004, and 2010 and at the WFGB in 2000, 2004, and 2010. Overall, the most noticeable 

pattern was the inverse relationship between coral and macroalgae cover. However, this 

is a general observation, as coral does not grow and die at the same rate as algae. 

Table 3.5.2.

 

EFGB and WFGB Random Transect Data for Dominant Cover Categories 

As reported for 1992 to 1995 in Continental Shelf Associates, Inc. (CSA)

(1996); for 1996 to 2001 from Dokken et al. (2003); PBS &J from 2002 to

2008 (Precht et al. 2006, 2008b); and FGBNMS for 2009 to 2010 (this report)

EFGB Random Transect Data

Year Montastraea annularis

species complex Macroalgae CTB

1992 24.12 4.78 54.46 ± 0.00

1994 26.93 0.29 47.31 ± 0.00

1995 35.65 0.57 42.15 ± 0.00

1996 21.30 ± 14.20 6.10 ± 5.20 -

1997 21.60 ± 8.10 0.50 ± 0.60 -

1998 30.40 ± 11.10 3.20 ± 2.60 27.60 ± 5.90

1999 28.20 ± 11.70 24.70 ± 13.20 11.10 ± 8.20

2000 39.50 ± 9.60 17.30 ± 4.90 4.30 ± 1.70

2001 44.80 ± 12.90 14.90 ± 5.60 5.70 ± 3.60

2002 33.59 ± 3.86 4.06 ± 0.75 37.07 ± 2.69

39 
EFGB Random Transect Data

Year Montastraea annularis

species complex Macroalgae CTB

2003 28.47 ± 2.98 16.74 ± 2.05 28.12 ± 2.05

2004 30.14 ± 4.76 12.03 ± 2.77 20.89 ± 3.08

2005 26.80 ± 4.09 34.03 ± 2.58 11.96 ± 1.49

2006 31.45 ± 4.09 21.10 ± 2.32 23.15 ± 1.94

2007 32.44 ± 4.62 21.73 ± 2.28 24.43 ± 2.11

2008 33.58 ± 4.52 24.06 ± 2.16 17.64 ± 1.77

2009 29.52 ± 12.99 31.85 ± 3.49 13.32 ± 1.07

2010 33.89 ± 3.84 32.94 ± 3.05 11.15 ± 0.86

WFGB Random Transect Data

Year Montastraea annularis

species complex Macroalgae CTB

1992 23.02 4.45 56.56

1994 24.95 0.42 51.08

1995 31.0 2.7 45.85

1996 27.20 ± 8.30 4.50 ± 5.20 -

1997 27.70 ± 9.90 0.10 ± 0.60 -

1998 28.40 ± 11.90 2.30 ± 2.60 20.70 ± 5.90

1999 31.70 ± 8.60 18.80 ± 13.20 21.10 ± 8.20

2000 30.90 ± 11.60 22.60 ± 14.00 8.50 ± 3.70

2001 35.10 ± 12.00 25.40 ± 7.30 4.60 ± 2.90

2002 31.73 ± 3.57 19.14 ± 1.40 27.63 ± 3.14

2003 33.80 ± 4.31 8.41 ± 1.41 31.63 ± 3.04

2004 31.70 ± 2.70 14.75 ± 1.50 20.85 ± 2.11

2005 36.20 ± 3.50 18.35 ± 1.44 18.27 ± 1.67

2006 40.13 ± 3.29 12.38 ± 1.34 25.64 ± 2.06

2007 35.50 ± 3.81 17.64 ± 2.44 24.27 ± 1.89

2008 37.01 ± 4.65 12.06 ± 1.31 26.74 ± 2.41

2009 27.25 ± 3.32 30.03 ± 2.63 14.93 ± 1.52

2010 45.98 ± 32.36 22.03 ± 2.00 10.20 ± 1.03

Values listed in table are the mean percent covers for MASC coral cover, 

macroalgae, and CTB. Standard deviations are shown for 1996 to 2001 based 

on analysis by Dokken et al. (2003) and standard errors are reported from

2002 to 2010

40 
CHAPTER 4.0: REPETITIVE QUADRATS

4.1. REPETITIVE QUADRAT METHODOLOGICAL RATIONALE

Permanent quadrats, covering 5 m2 in 2009 and 8 m

2 in 2010, were photographed to 

monitor changes in the composition of benthic assemblages on the FGB. The repetitive 

quadrats were located within the EFGB and WFGB study sites, except for the deep 

stations on the EFGB. Only the central 5 m2 of the 2010 images was analyzed, as this 

area provided the clearest resolution for analysis and was comparable to the previous 

year. The photographs were analyzed in two ways. The first method measured percent 

benthic cover components in 2009 and 2010 using random-dot analysis. Second, selected 

corals within the repetitive quadrats were analyzed using planimetry to measure gain or 

loss of tissue area. 

4.2. REPETITIVE QUADRAT FIELD AND LABORATORY METHODS

In 2009, thirty-seven 5 m2 repetitive quadrats were photographed at each EFGB and WFGB. 

In 2010, thirty-four and thirty-nine 8 m2 quadrats were photographed at the EFGB and 

WFGB respectively (Figure 4.2.1). All nine EFGB deep station quadrats were 

photographed in both years.

Figure 4.2.1. Repetitive quadrat station #761 at the EFGB in 2009

(NOAA/FGBNMS).

41 
4.2.1. Repetitive Quadrat 2009 Digital Photography

In 2009, stations were photographed using a Nikon Coolpix P5000 camera in an Ikelite 

housing, with the Inon UWL-100 Type 2 wet mount wide-angle converter lens. The

camera was mounted in the center of a T-shaped camera frame, at a distance of 2 m (6.5 

ft) from the substrate. Two Ikelite substrobe DS125 strobes were mounted 1.2 m apart on 

the ends of the T-frame and set on TTL. To ensure that the same quadrats were 

photographed in the same manner each year, the frame was always oriented in a northfacing direction and kept vertical using an attached bulls-eye bubble level. This set-up 

produced images with a coverage of 251 x 194 cm.

4.2.2. Repetitive Quadrat 2010 Digital Photography

In 2010, stations were photographed in the same way, but using a Canon Power Shot 

G11 digital camera in a FIX Fish-Eye housing with a 165o dome port. Because a 

different lens was used, the distance above the bottom 1.5 m (4.9 ft) and the strobes (Inon 

Z240) were mounted 1.2 m apart. This set-up produced images with a coverage of 384 x 

239 cm.

4.2.3. Repetitive Quadrat Image Analysis

White balance and color correction were applied to each image as necessary. All images 

were corrected for barrel distortion using Abode® Photoshop® CS5’s distortion 

correction feature. The amount of distortion correction needed was determined by

analyzing photographs from pool tests with scale bars. For the 2009 images, no distortion 

correction was needed because the 2009 camera setup did not use a wide angle lens. For 

2010, a distortion correction of +217 was applied to remove the barrel distortion caused 

by the use of the 165o

 dome port. In addition, all 2010 images were cropped to match the 

2009 repetitive image area. This correction technique removes any error that could occur 

during percent cover analysis as a result from utilizing different camera setups in 2009 

and 2010. However, this does not correct for fine scale differences as seen in the 

planimetery analysis. This will be addressed in section 4.4.3. 

4.2.3.1. Percent Cover of Benthic Components

Coral cover in the images was analyzed using Coral Point Count with Excel® extensions 

(CPCe) as described in section 3.2. 

The percent cover of coral species, sponges, macroalgae, and CTB were determined by 

overlaying 100 random dots on each photograph using CPCe. This was changed from the 

previous methodology by Zimmer et al. (2010) where 100 point counts per image were 

conducted three times. Statistical results from a one-way ANOVA showed there was no 

significant difference between major cover categories in triplicate compared with single 

counts for coral species (F=0.03, df=1, P-value >0.86), sponge (F=1.0, df=1, P-value 

>0.33), macroalgae (F=0.01, df=1, P-value >0.98), and CTB (F=0.07, df=1, P-value 

>0.79). Therefore, single 100 points were used per picture for the 2009 and 2010 monitoring 

period. Mean percent cover of corals, sponges, macroalgae, CTB, along with coral 

bleaching, fish biting, and disease were calculated using 1 x 100 random-dot analysis with 

CPCe software.

42 
4.2.4. Planimetry Analysis

Planimetry was used to measure percent change in area of living tissue of selected coral 

colonies (i.e., Montastraea annularis species complex, Diploria strigosa, Colpophyllia 

natans, Montastraea cavernosa, and Porites astreoides) at repetitive quadrat stations in 

successive years. For the 2009 to 2010 interval, it was possible to compare 24 quadrats at 

the EFGB, and 37 quadrats at the WFGB. For the EFGB deep stations, all nine quadrats 

were compared for the interval. The 2009 images were scaled to match the area of the

2010 images. This was done in order to obtain a 1:1 pixel ratio between the 2009 and 

2010 photos. This was conducted using Adobe Photoshop® CS5 image size feature. 

Planimetry results were calculated by taking areal measurements of coral colonies whose

lateral margins were contained entirely within the image frame from 2009 and 2010. The 

live tissue cover of each colony was traced in Adobe Illustrator® CS2 using a Wacom®

Cintiq® 12WX Tablet. A mask was then created of the live tissue and areal measurements 

were calculated using ImageJ®. The percent change in area for each colony was 

calculated by comparing the area of 2009 to the area of 2010. The change (either 

positive=growth, or negative=retreat) in pixels was divided by the area (pixels) from 

2009 to determine proportional growth or loss of tissue area. 

4.3. REPETITIVE QUADRAT RESULTS

4.3.1. Repetitive Quadrat Analysis

4.3.1.1. Percent Cover

The point count data from the repetitive quadrats in the 100 x 100 m study sites were 

grouped into four functional categories (coral, sponge, macroalgae, and CTB) and 

expressed as percent covers, the same way the random transect point counts were

displayed. In general, repetitive quadrat results from 2009 and 2010 revealed higher coral 

cover than random transects, low levels of macroalgae (thick turfs and fleshy macroalgal 

species) and CTB (crustose coralline algae, fine turf algae, and bare rock) cover, and very

low coverage of sponges (Figure 4.3.1). The higher coral cover at repetitive stations was 

to be expected because the stations were originally selected as premium monitoring sites 

with high coral cover. They were not intended to be representative of average benthic 

cover.

43 
0

10

20

30

40

50

60

70

80

90

2009 EFGB 2010 EFGB 2009 WFGB 2010 WFGB

Percent Cover

Bank and Year

Percent Cover of Repetitive Quadrat Functional Catagories 

at the Flower Garden Banks, 2009-2010

Coral

Sponge

Macroalgae

CTB

CTB combines crustose coralline algae, fine turf algae, and bare rock.The “macroalgae” 

category includes thick turfs as well as fleshy macroalgal species. Values are calculated 

from the repetitive quadrat photographs. 

Figure 4.3.1. Percent cover (+ SE) of four functional categories of sessile benthos 

at the FGB in 2009 and 2010.

At the EFGB, the results reflected stable coral cover from 2009 to 2010 (69.27% ± 2.69 

to 72.41% ± 2.56), while the sponge cover remained extremely low for both years (0.24% 

± 0.11 to 0.19% ± 0.08) (Figure 4.3.1 and Table 4.3.1). Macroalgae cover (mainly fleshy 

algae, turfs, and Dictyota spp.) from 2009 to 2010 (19.27% ± 1.98 to 15.37% ± 1.71), and

CTB cover remained stable (10.83% ± 0.94 to 11.69% ± 1.14) as well. Consistent with 

past monitoring results, the Montastraea annularis species complex (MASC), containing 

M. annularis, M. faveolata, and M. franksi, dominated the stations at EFGB in 2009 

(46.87% ± 1.56) and 2010 (51.19% ± 1.93). Diploria strigosa (11.10% ± 1.83 and 

10.25% ± 2.17 in 2009 and 2010, respectively) and Porites astreoides (5.00% ± 0.62 in 

2009 and 1.74% ± 0.33 in 2010) were the next most abundant taxa (Figure 4.3.2 and 

Table 4.3.1). The remaining coral cover was made up of eleven species, none of 

which exceeded 6.0% in either year (Table 4.3.1). Corals that could not be 

differentiated because of camera angle or camera distortion were labeled as “unidentified 

coral.” Shannon-Weiner diversity values remained consistent at the EFGB (H’= 0.76 and 

H’= 0.73 in 2009 and 2010, respectively). 

44 
0 10 20 30 40 50 60

MASC

Diploria strigosa

Porites astreoides

Montastraea cavernosa

Unidentifiable Coral

Colpophyllia natans

Millepora alcicornis

Siderastrea siderea

Percent Cover

Coral Species

Percent Cover of Dominant Coral Taxa at the East Flower Garden Bank, 2009-2010

2009

2010

Figure 4.3.2. Percent cover (+ SE) of the dominant coral taxa in repetitive quadrats at the 

EFGB in 2009 and 2010. 

In quadrats at the WFGB, coral cover remained consistent from 2009 (72.19% ± 2.48) to 

2010 (74.19% ± 2.07). The sponge cover remained extremely low for both years at the 

WFGB (0.03% ± 0.03 to 0.11% ± 0.06) (Figure 4.3.1 and Table 4.3.1). Macroalgae

cover decreased from 2009 to 2010 (11.65% ± 1.53 to 7.46% ± 1.01), while CTB cover 

remained stable (15.41% ± 1.51 to 14.69% ± 1.21). As with the EFGB, the Montastraea 

annularis species complex dominated quadrats at the WFGB in 2009 (48.36% ± 1.85) 

and 2010 (45.69% ± 1.81). Diploria strigosa (10.24% ± 1.89 and 7.84% ± 1.23 in 2009 

and 2010, respectively) and Porites astreoides (4.29% ± 0.62 in 2009 and 2.49% ± 0.56 

in 2010) were the next most abundant species (Figure 4.3.3 and Table 4.3.1). 

Montastraea cavernosa was also abundant in 2009 (3.41% ± 0.92) and 2010 (4.47% ± 

1.16). The remaining coral cover was made up of eleven species, none of which

exceeded 10.0% in either year (Table 4.3.1). Shannon-Weiner diversity values

remained consistent at the WFGB (H’=0.73 and H’=0.70 in 2009 and 2010, 

respectively). 

45 
0 10 20 30 40 50

MASC

Diploria strigosa

Porites astreoides

Montastraea cavernosa

Unidentifiable Coral

Colpophyllia natans

Millepora alcicornis

Siderastrea siderea

Percent Cover

Coral Species

Percent Cover of Dominant Coral Taxa at the West Flower Garden Bank, 2009-2010

2009

2010

Figure 4.3.3. Percent cover (+ SE) of the dominant coral taxa in repetitive quadrats at the 

WFGB in 2009 and 2010. 

46 
Table 4.3.1.

Cover of Benthic Categories in Repetitive Quadrats at the EFGB and WFGB in 2009 and 2010 

Values are expressed as percent cover ± SE

Cover Category 2009 EFGB 2010 EFGB 2009 WFGB 2010 WFGB

Coral

Montastraea annularis

species complex (MASC) 46.87 ± 1.56 51.19 ± 1.93 48.36 ± 1.85 45.69 ± 1.81

Diploria strigosa 11.10 ± 1.83 10.25 ± 2.17 10.24 ± 1.89 7.84 ± 1.23

Porites astreoides 5.00 ± 0.62 1.74 ± 0.33 4.29 ± 0.62 2.49 ± 0.56

Montastraea cavernosa 2.43 ± 0.74 1.72 ± 0.52 3.41 ± 0.92 4.47 ± 1.16

Unidentifiable Coral 1.86 ± 0.22 5.63 ± 0.72 2.30 ± 0.38 9.56 ± 1.06

Colpophyllia natans 0.45 ± 0.17 1.10 ± 0.45 0.96 ± 0.36 1.36 ± 0.45

Millepora alcicornis 0.36 ± 0.15 0.21 ± 0.09 0.29 ± 0.12 0.43 ± 0.18

Madracis spp. 0.32 ± 0.27 0.10 ± 0.06 0.19 ± 0.09 0.06 ± 0.06

Agaricia spp. 0.26 ± 0.17 0.04 ± 0.04 0.11 ± 0.09 0.04 ± 0.04

Siderastrea siderea 0.20 ± 0.17 0.00 1.39 ± 0.87 1.30 ± 0.96

Stephanocoenia intersepta 0.15 ± 0.12 0.00 0.40 ± 0.18 0.24 ± 0.12

Mussa angulosa 0.07 ± 0.05 0.11 ± 0.06 0.18 ± 0.13 0.13 ± 0.07

Porites furcata 0.03 ± 0.03 0.00 0.00 0.00

Siderastrea spp. 0.03 ± 0.03 0.09 ± 0.07 0.08 ± 0.05 0.26 ± 0.14

Scolymia spp. 0.00 0.03 ± 0.03 0.00 0.03 ± 0.03

Total Coral 69.27 ± 2.69 72.41 ± 2.56 72.19 ± 2.48 74.19 ± 2.07

Sponge

Agelas clathrodes 0.11 ± 0.08 0.04 ± 0.04 0.00 0.11 ± 0.06

Chondrilla brown 0.06 ± 0.04 0.11 ± 0.06 0.00 0.00

Arolochroia (Pseudoceratina) 

crassa 0.04 ± 0.04 0.00 0.00 0.00

Unknown Sponge I 0.03 ± 0.03 0.04 ± 0.04 0.03 ± 0.03 0.00

Encrusting sponge 0.00 0.07 ± 0.05 0.08 ± 0.05 0.52 ± 0.18

Total Sponge 0.24 ± 0.11 0.19 ± 0.08 0.03 ± 0.03 0.11 ± 0.06

CTB

Bare Substrate 8.47 ± 0.86 4.53 ± 0.82 9.91 ± 1.14 3.24 ± 0.34

Fine Turf 2.33 ± 0.46 3.03 ± 0.62 4.29 ± 1.09 6.98 ± 0.92

Crustose Coralline Algae 0.00 4.05 ± 0.50 0.00 4.47 ± 0.70

Total CTB 10.83 ± 0.94 11.69 ± 1.14 15.41 ± 1.51 14.69 ± 1.21

Macroalgae

Fleshy algae 15.84 ± 1.64 10.39 ± 1.24 10.45 ± 1.35 5.73 ± 0.81

Dictyota spp. 1.97 ± 0.53 3.35 ± 0.76 0.40 ± 0.14 0.40 ± 0.13

Thick turf algae 1.04 ± 0.37 1.59 ± 0.41 0.08 ± 0.08 1.28 ± 0.32

Lobophora variegata 0.39 ± 0.13 0.05 ± 0.05 0.72 ± 0.23 0.05 ± 0.05

47 
Cover Category 2009 EFGB 2010 EFGB 2009 WFGB 2010 WFGB

Filamentous algae 0.03 ± 0.03 0.00 0.00 0.00

Total Macroalgae 19.27 ± 1.98 15.37 ± 1.71 11.65 ± 1.53 7.46 ± 1.01

Other

Shadow 14.63 ± 1.26 22.00 ± 1.86 15.35 ± 0.83 26.36 ± 1.61

Sand 0.14 ± 0.10 0.28 ± 0.20 0.35 ± 0.21 2.91 ± 1.65

Substrate Rubble 0.00 0.03 ± 0.03 1.21 ± 0.73 0.00

Fish 0.06 ± 0.06 0.00 0.25 ± 0.10 0.03 ± 0.03

Invertebrate 0.18 ± 0.08 0.00 0.03 ± 0.03 0.10 ± 0.08

Coral Condition (occurences in coral)

Bleached Coral 0.05 ± 0.04 0.03 ± 0.03 0.00 1.90 ± 0.38

Paling Coral 0.82 ± 0.39 0.43 ± 0.18 0.50 ± 0.18 2.77 ± 0.62

Concentrated Fish Biting 0.03 ± 0.03 0.00 0.03 ± 0.03 0.03 ± 0.03

Isolated Fish Biting 0.03 ± 0.03 0.03 ± 0.03 0.00 0.03 ± 0.03

The combined data collected in 2009 and 2010 showed persistence of the Montastraea 

annularis species complex as the dominant coral species in the repetitive quadrats at the 

EFGB and WFGB (Table 4.3.1). Diploria strigosa was the second-most prevalent coral 

species during this time period. Porites astreoides and M. cavernosa were consistently

the third and fourth most abundant corals. The average coral cover in quadrats at both 

banks was approximately 72% for the reporting period. This was much higher than the 

average random transect coral cover for both banks during the reporting period (~57%), 

but as explained above, the coral in the repetitive stations were selected partly because of

their high coral cover. Appendix 4 contains the repetitive quadrat data for 2009 and 2010

at the FGB.

In past monitoring years, the Montastraea annularis species complex (M. annularis, M. 

faveolata, and M. franksi) were not differentiated, but with higher resolution digital 

photography and the familiarity of the FGBNMS staff with the photostations, the 

components of the Montastraea annularis species complex were indentified in the 

repetitive quadrat photographs. At both banks, the Montastraea annularis species 

complex is dominated by M. franksi, followed by M. faveolata and M. annularis. At the 

EFGB, M. franksi percent cover was similar in 2009 and 2010 (35.50% ± 3.40 to 37.19% 

± 3.82), and M. faveolata (3.99% ± 1.11 to 2.81% ± 1.01) remained stable as well. M. 

annularis was not sampled in 2009, but was identified in 2010 (3.03% ± 1.42) (Figure 

4.3.4). At the WFGB, M. franksi percent cover was slightly lower in 2010 (35.21% ± 

3.62 to 31.76% ± 2.72), while M. annularis (3.44% ± 1.43 to 5.26% ± 2.00) and M. 

faveolata (0.40% ± 0.26 to 1.05% ± 0.81) remained relatively stable (Figure 4.3.5). 

Approximately 8.0% of the individual MASC components still could not be identified, 

which may be due to MASC hybridization or genotypic variation. 

In the repetitive quadrat in 2009 and 2010, the incidences of bleaching, paling, and fish 

biting were low at both banks and coral disease was absent (Table 4.3.1). The percentage 

of corals impacted by isolated/concentrated fish biting at the EFGB and WFGB was 

negligible, ranging from 0.00 to 0.03%. Less than 3% of the coral cover analyzed in the 

48 
random transect percent cover analysis exhibited disease or bleaching at the EFGB and 

WFGB in either year (Figure 4.3.6). 

0 5 10 15 20 25 30 35 40 45

Montastraea franksi

Unidentified MASC

Montastraea faveolata

Montastraea annularis

Percent Cover

Coral Species

Percent Cover of the Coral that Comprises MASC at the East 

Flower Garden Bank, 2009-2010

2009

2010

Figure 4.3.4. Percent cover (+ SE) of the coral species in the MASC in repetitive 

quadrats at the EFGB in 2009 and 2010. 

0 10 20 30 40 50

Montastraea franksi

Unidentified MASC

Montastraea faveolata

Montastraea annularis

Percent Cover

Coral Species

Percent Cover of the Coral that Comprises MASC at the West 

Flower Garden Bank, 2009-2010

2009

2010

Figure 4.3.5. Percent cover (+ SE) of the coral species in the MASC in repetitive quadrats 

at the WFGB in 2009 and 2010. 

49 
0

0.5

1

1.5

2

2.5

3

3.5

4

2009 EFGB 2010 EFGB 2009 WFGB 2010 WFGB

Percent of Coral

Bank and Year

Bleaching and Paling of Corals from Percent Cover Analysis at the 

Flower Garden Banks, 2009-2010 Bleached

Paling

Figure 4.3.6. Percent of coral observed to pale and bleach (+ SE) in repetitive quadrats 

at the FGB in 2009 and 2010. 

4.3.1.2. Deep Stations–Percent Cover

All nine EFGB deep station quadrats were photographed in both years and analyzed for

benthic cover using random dot analysis. The point count data were grouped into the 

same four functional categories (coral, sponge, macroalgae, and CTB) and expressed as 

percent covers, the same way the random transect point counts were displayed. Coral

cover was very high at the deep stations (ranging from 81–82% between 2009 and

2010), and sponge cover was extremely low (0.00–0.25%). Coral cover was followed by 

low levels of macroalgae and CTB cover (Figure 4.3.7). High levels of coral cover were 

consistent with past monitoring results. Unlike the repetitive stations in the shallower 

study sites, however, which were chosen because of the presence of high cover, these

stations were selected to show representative areas of the deep reef. Therefore, the high 

coral cover at these stations may be more indicative of average conditions at these depths 

on the bank. 

50 
CTB includes crustose coralline algae, fine turf algae, and bare rock. The “macroalgae” 

category includes thick turfs as well as fleshy macroalgal species. Values are calculated 

from the repetitive quadrat deep station photographs.

Figure 4.3.7. Percent cover (+ SE) of four functional categories of sessile benthos at the 

EFGB repetitive deep stations in 2009 and 2010. 

Coral cover remained stable from 2009 to 2010 (81.73% ± 3.90 to 82.18% ± 3.01). 

Sponge cover was not detected in 2009, and only minimally observed in 2010 (0.25% ± 

0.25) (Figure 4.3.7 and Table 4.3.2). Macroalgae cover (mainly fleshy algae, turfs, and 

Dictyota spp.) remained stable from 2009 to 2010 (10.56% ± 3.12 to 10.28% ± 2.06), as 

did CTB cover (7.72% ± 1.39 to 6.98% ± 1.53). 

Consistent with past monitoring results, the Montastraea annularis species complex 

(MASC) continued to dominate the EFGB repetitive deep stations in 2009 (51.70% ± 

4.38) and 2010 (46.70% ± 3.54). Montastraea cavernosa (15.42% ± 4.16 and 10.65% ± 

3.69 in 2009 and 2010, respectively) and Colpophyllia natans (5.10% ± 2.06 in 2009 and 

5.75% ± 3.44 in 2010) were the next most abundant species (Figure 4.3.8 and Table 

4.3.2). Montastraea cavernosa was also more dominant in the EFGB deep stations than 

on the reef cap. The remaining coral cover was made up of twelve species, none of 

which exceeded than 2% in either year (Table 4.3.2). Corals that could not be 

differentiated due to camera angle or camera distortion were label as “unidentifiable

coral.”

51 
Figure 4.3.8. Percent cover (+ SE) of the dominant coral taxa at repetitive deep stations 

on the EFGB in 2009 and 2010. 

52 
Table 4.3.2.

Cover of Benthic Categories in Repetitive Deep Stations at the EFGB from 2009 and 2010 

Values are expressed as percent cover ± SE

Cover Category 2009 EFGB Deep Station 2010 EFGB Deep Station

Coral

Montastraea annularis species 

complex (MASC) 51.70 ± 4.38 46.70 ± 3.54

Montastraea cavernosa 15.42 ± 4.16 10.65 ± 3.69

Colpophyllia natans 5.01 ± 2.06 5.75 ± 3.44

Unidentifiable Coral 2.96 ± 1.07 13.56 ± 2.22

Diploria strigosa 1.69 ± 1.18 1.75 ± 1.61

Millepora alcicornis 1.40 ± 1.14 1.07 ± 0.63

Stephanocoenia intersepta 1.32 ± 0.71 1.16 ± 0.99

Mussa angulosa 0.96 ± 0.60 0.79 ± 0.45

Agaricia spp. 0.65 ± 0.65 0.16 ± 0.16

Scolymia cubensis 0.36 ± 0.25 0.00

Madracis decactis 0.13 ± 0.13 0.00

Scolymia spp. 0.12 ± 0.12 0.00

Agaricia undata 0.00 0.16 ± 0.16

Porites astreoides 0.00 0.16 ± 0.16

Paracyanthus pulchellus 0.00 0.13 ± 0.13

Siderastrea siderea 0.00 0.12 ± 0.12

Total Coral 81.73 ± 3.90 82.18 ± 3.01

SPONGE

Ircinia felix 0.00 0.13 ± 0.13

Ircinia strobilina 0.00 0.13 ± 0.13

Total Sponge 0.00 0.25 ± 0.25

CTB

Bare Substrate 3.53 ± 0.92 3.45 ± 0.74

Fine Turf 3.18 ± 0.92 1.48 ± 0.71

Crustose Coralline Algae 0.00 2.05 ± 0.74

Total CTB 7.72 ± 1.39 6.98 ± 1.53

Macroalgae

Fleshy algae 5.47 ± 1.35 7.69 ± 1.58

Lobophora variegata 1.82 ± 0.74 2.07 ± 0.72

Think turf algae 1.78 ± 0.62 0.13 ± 0.13

Dictyota spp. 1.49 ± 0.99 0.26 ± 0.17

Peysonnelia spp. 0.00 0.13 ± 0.13

53 
Cover Category 2009 EFGB Deep Station 2010 EFGB Deep Station

Total Macroalgae 10.56 ± 3.12 10.28 ± 2.06

Other

Shadow 6.78 ± 1.90 18.56 ± 2.95

Coral Rubble 0.76 ± 0.76 0.00

Substrate Rubble 0.25 ± 0.17 0.00

Sand 0.00 0.14 ± 0.14

Coral Condition (occurences 

in coral)

Bleached Coral 0.00 0.00

Paling Coral 2.67 ± 1.46 1.89 ± 1.09

Concentrated Fish Biting 0.22 ± 0.22 0.00

Isolated Fish Biting 0.00 0.00

At the deep stations on the EFGB, the Montastraea annularis species complex is 

dominated by M. franksi, followed by M. faveolata. M. franksi dominated the MASC 

cover (45.37% ± 6.19 in 2009 and 25.32% ± 4.49 in 2010). M. faveolata was not 

detected in 2009, but was detected in 2010 (1.40% ± 1.40) (Figure 4.3.9). M. annularis

was not identified in 2009 or 2010; however, 6.33% of the individual MASC components 

were unidentifiable in 2009, and 19.98% of the components were unidentifiable in 2010.

This may also be due to MASC hybridization or genotypic variation. 

Figure 4.3.9. Percent cover (+ SE) of the coral species that comprises the MASC at 

deep repetitive stations the EFGB deep stations from 2009 and 2010. 

54 
In the 2009 and 2010 repetitive deep stations, evidence of coral disease was absent (Table 

4.3.2). The percentage of corals impacted by isolated/concentrated fish biting was 

negligible, ranging from 0.00 to 0.22%. Less than 3% of the coral cover analyzed in the 

deep stations exhibited paling or bleaching. 

4.3.2. Repetitive Quadrat Planimetric Analysis 

Some corals in repetitive quadrat photographs were analyzed using planimetry. 

Measurements of the amount of change in living area on selected coral colonies were 

conducted to document the dynamics of particular coral colonies at the EFGB (cap and 

deep stations) and WFGB. Changes were also assessed visually to either verify or refute 

planimetric measurements. Staff chose and identified coral colonies with discernable 

margins to be measured for planar areal change from 2009 to 2010. In each frame, one 

to four colonies of framework-building corals, where the margins were clearly defined, 

were chosen for analysis. Montastraea annularis species complex, the main contributor 

to coral cover at the FGB, along with Diploria strigosa and Porites asteroides colonies 

were the most common colonies selected (Table 4.3.3). 

Table 4.3.3.

Reef-Building Coral Colonies in Photographs from Repetitive Quadrat Stations Selected for 

Planimetric Analysis from 2009 to 2010 

Bank

Coral Colonies Chosen for Planimetric Analysis 

Station Number Coral 1 Coral 2 Coral 3 Coral 4

EAST BANK

001 M. cavernosa P. astreoides

003 D. strigosa P. asteroides P. astreoides M. franksi

005 C. natans M. faveolata P. astreoides P. astreoides

010 C. natans P. astreoides P. astreoides

016 M. franksi P. astreoides D. strigosa

020 P. astreoides P. astreoides P. astreoides D. strigosa

021 P. astreoides M. franksi P. astreoides M. franksi

027 D. strigosa D. strigosa C. natans

029 M. franksi P. astreoides D. strigosa

032 M. faveolata M. faveolata P. astreoides M. franksi

058 MASC M. cavernosa P. astreoides

067 D. strigosa P. astreoides D. strigosa

185 P. astreoides D. strigosa P. astreoides

715 D. strigosa D. strigosa MASC

717 D. strigosa P. astreoides

725 P. astreoides M. franksi C. natans

739 M. cavernosa M. franksi P. astreoides

751 M. franksi M. annularis P. astreoides M. annularis

55 
Bank

Coral Colonies Chosen for Planimetric Analysis 

Station Number Coral 1 Coral 2 Coral 3 Coral 4

755 P. astreoides D. strigosa P. astreoides

756 C. natans M. franksi P. astreoides

757 P. astreoides D. strigosa M. franksi

761 C. natans D. strigosa M. franksi P. astreoides

764 M. franksi P. astreoides D. strigosa P. astreoides

765 P. astreoides M. cavernosa P. astreoides

DEEP STATIONS

081 C. natans C. natans

082 M. angulosa C. natans MASC

083 M. franksi M. cavernosa M. cavernosa

084 M. cavernosa M. cavernosa M. faveolata

085 M. cavernosa M. faveolata M. cavernosa

086 M. franksi M. cavernosa

087 M. franksi MASC P. astreoides MASC

088 D. strigosa M. franksi

089 M. cavernosa M. franksi M. cavernosa

WEST BANK

018 P. astreoides P. astreoides P. astreoides P. astreoides

041 M. faveolata M. franksi P. astreoides

042 P. astreoides MASC M. cavernosa

045 M. franksi P. astreoides D. strigosa

046 M. franksi P. astreoides C. natans

049 D. strigosa M. cavernosa P. astreoides

051 P. astreoides D. strigosa D. strigosa

054 P. astreoides P. astreoides P. astreoides

061 M. annularis P. astreoides M. annularis

072 P. astreoides D. strigosa D. strigosa D. strigosa

073 D. strigosa M. franksi D. strigosa

091 D. strigosa D. strigosa D. strigosa

093 P. astreoides D. strigosa P. astreoides

125 P. astreoides C. natans P. astreoides

822 D. strigosa D. strigosa M. franksi

824 S. intersepta P. astreoides M. annularis

839 M. franksi M. franksi D. strigosa

843 P. astreoides D. strigosa D. strigosa

844 P. astreoides P. astreoides P. astreoides

847 P. astreoides P. astreoides D. strigosa

848 D. strigosa P. astreoides D. strigosa

56 
Bank

Coral Colonies Chosen for Planimetric Analysis 

Station Number Coral 1 Coral 2 Coral 3 Coral 4

857 C. natans M. franksi M. franksi

861 M. franksi P. astreoides P. astreoides

862 P. astreoides P. astreoides M. franksi

865 D. strigosa P. astreoides D. strigosa

866 S. intersepta D. strigosa P. astreoides

867 M. franksi D. strigosa

868 P. asteroides P. asteroides P. astreoides 

869 M. annularis

870 P. astreoides P. astreoides P. astreoides P. astreoides

874 M. faveolata M. annularis P. astreoides

875 P. astreoides M. franksi D. strigosa

876 P. astreoides P. astreoides

877 P. astreoides C. natans M. franksi

878 M. franksi M. franksi M. franksi

879 M. franksi D. strigosa M. franksi

NT3 D. strigosa P. astreoides P. astreoides

On average, coral colonies selected for analysis at the EFGB and WFGB were found to 

have decreased in area. However, visual photohraphics assessments of all the analyzed 

coral colonies suggested that most of the coral colonies did not suffer as much tissue loss 

as the planimetric analysis suggested (Table 4.3.4). The problems with the calculations of 

area lost or gained from 2009 to 2010 are thought to arise from using different camera 

setups in the two years and previous analog versus digital photos from previous 

monitoring periods. No statistical analyses were run on the data because of issues with 

image distortion and differing heights of the camera above the bottom derived from 

changing camera setups between years resulting in corals to appear at different angles. 

These issues are addressed further in the discussion section. 

57 
Table 4.3.4.

Visual Assessment of Analyzed Coral Colonies Suggest Colonies Did Not Suffer As Much Tissue 

Loss as the Planimetric Analysis Suggested 

Highlighted cells suggest causes or consequences of change; blue cells represent growth 

and yellow loss

Bank Station 

Number Coral 1 Coral 2 Coral 3 Coral 4

EAST BANK

001 Tissue growth in 

some bare areas

No detectable 

change

003 No detectable 

change

No detectable 

change

No detectable 

change

No detectable 

change

005

No detectable 

change

Small growth 

around bare 

areas

Loss of tissue on 

margin

Small loss of 

tissue on 

cracks

010

Loss of tissue and 

increase in orange 

sponge

No detectable 

change

No detectable 

change

016 No detectable 

change

No detectable 

change

No detectable 

change

020 No detectable 

change

No detectable 

change

No detectable 

change

No detectable 

change

021 Small tissue loss No detectable 

change

No detectable 

change

No detectable 

change

027

Small growth on 

margins, filling in 

gaps

Small loss of 

tissue on margin, 

next to M. 

franksi

No detectable 

change

029 Bare areas filling 

in with live tissue

No detectable 

change

No detectable 

change

032 Concentrated fish 

biting healed

No detectable 

change

No detectable 

change

No detectable 

change

058

No detectable 

change

Tissue loss in 

center of head, 

overgrown with 

algae

No detectable 

change

067

Small tissue loss, 

possible fish 

biting

Small growth 

around margins

No detectable 

change

185

No detectable 

change

No detectable 

change

Large tissue loss, 

overgrown with 

algae

715

No detectable 

change

Large tissue loss, 

CCA / algae 

growing on dead 

coral

No detectable 

change

717 No detectable 

change

Small tissue loss 

on margins

725 No detectable 

change

No detectable 

change

Loss of tissue

739 No detectable 

change

No detectable 

change

No detectable 

change

58 
Bank Station 

Number Coral 1 Coral 2 Coral 3 Coral 4

751

Small tissue loss No detectable 

change

No detectable 

change

Tissue 

growing over 

bare area 

755 No detectable 

change

No detectable 

change

No detectable 

change

756 No detectable 

change

No detectable 

change

Small growth on 

margin

757

Small loss of 

tissue; possible 

fish biting

Coral looks 

healthier; tissue 

growth over 

cracks; 

discoloration 

gone

Tissue growth in 

previous voids

761

Some 

discoloration on N 

margin

Small tissue loss 

in crack on S 

margin

No detectable 

change

Tissue growth 

filling in 

cracks

764

No detectable 

change

Small loss of 

tissue on margin 

near Coral 3

No detectable 

change

No detectable 

change

765 No detectable 

change

No detectable 

change

Large tissue loss

DEEP STATIONS

081 No detectable 

change

No detectable 

change

082 Small loss of 

tissue

No detectable 

change

No detectable 

change

083 No detectable 

change

No detectable 

change

Small loss of 

tissue; fish biting

084 Small tissue loss; 

fish biting

No detectable 

change

No detectable 

change

085

No detectable 

change

No detectable 

change

Small growth of 

tissue in bare 

areas

086 No detectable 

change

No detectable 

change

087

Small tissue loss, 

possible fish 

biting

No detectable 

change

No detectable 

change

No detectable 

change

088 No detectable 

change

No detectable 

change

089

Tissue loss; 

overgrown with 

algae

No detectable 

change

Tissue loss; fish 

biting

WEST BANK

018

No detectable 

change

No detectable 

change

No detectable 

change

No detectable 

change

041 No detectable 

change

No detectable 

change

No detectable 

change

042

No detectable 

change

Large tissue 

growth into 

previously bare 

areas

No detectable 

change; coral 

paling

59 
Bank Station 

Number Coral 1 Coral 2 Coral 3 Coral 4

045 No detectable 

change

Small loss of 

tissue

No detectable 

change

046 Small recovery of 

tissue

No detectable 

change

No detectable 

change

049 No detectable 

change

Large loss of 

tissue on apex

No detectable 

change

051 No detectable 

change

No detectable 

change

Small tissue 

growth

054 No detectable 

change

Small tissue 

growth

No detectable 

change

061 Small loss of 

tissue

No detectable 

change

No detectable 

change

072 No detectable 

change

No detectable 

change

No detectable 

change

No detectable 

change

073 Tissue growth 

into bare areas

Tissue loss Tissue loss

091 No detectable 

change

No detectable 

change

No detectable 

change

093 No detectable 

change

Small tissue loss No detectable 

change

125

No detectable 

change

Small loss of 

tissue on 

margins

No detectable 

change

822 No detectable 

change

No detectable 

change

No detectable 

change

824

No detectable 

change

Small loss of 

tissue

Loss of tissue; 

overgrown with 

algae

839 No detectable 

change

No detectable 

change

No detectable 

change

843 Small growth of 

tissue

No detectable 

change

No detectable 

change

844 Small loss of 

tissue on margin

No detectable 

change

No detectable 

change

847 No detectable 

change

No detectable 

change

No detectable 

change

848 No detectable 

change

No detectable 

change

No detectable 

change

857 Loss of tissue No detectable 

change

No detectable 

change

861 Small growth of 

tissue

Loss of tissue No detectable 

change

862 No detectable 

change

No detectable 

change

No detectable 

change

865 No detectable 

change

No detectable 

change

No detectable 

change

866 No detectable 

change

No detectable 

change

No detectable 

change

867 Tissue growth Small tissue loss

868 No detectable 

change

No detectable 

change

No detectable 

change

60 
Bank Station 

Number Coral 1 Coral 2 Coral 3 Coral 4

869 No detectable 

change

870 No detectable 

change

Loss of tissue No detectable 

change

No detectable 

change

874

Loss of tissue: 

fish biting

Tissue growth 

into previously 

bare areas

No detectable 

change

875 No detectable 

change

No detectable 

change

No detectable 

change

876 No detectable 

change

No detectable 

change

877

No detectable 

change

No detectable 

change; coral 

paling

No detectable 

change

878 No detectable 

change

No detectable 

change

No detectable 

change

879 Tissue growth No detectable 

change

No detectable 

change

NT3 No detectable 

change

Small loss of 

tissue

No detectable 

change

4.4. REPETITIVE QUADRAT DISCUSSION

4.4.1. Study Site Repetitive Quadrats

Repetitive quadrats were analyzed for percent cover of benthic components, including 

coral species, sponge, macroalgae, CTB, and to identify coral health indicators 

(bleaching, paling, concentrated fish biting, isolated fish biting, and disease) in 2009 

and 2010. Higher coral cover estimates (~72%) were obtained from the repetitive 

quadrats in comparison to the random transects (~57%) at both the EFGB and WFGB. 

Higher percent coral cover in repetitive quadrats relative to random transects has also 

been documented in previous reports (Dokken et al. 2003; Precht et al. 2006, 2008b; 

Zimmer et al. 2010). The most likely reason for this difference is that repetitive quadrat

stations were not installed in random locations, but were originally selected as premium 

monitoring sites with high coral cover (large coral colonies). They were not intended to 

be representative of average benthic cover.

Figure 4.4.1 shows one typical repetitive photostation (number 761) from the EFGB in a 

time series from 2006 to 2010. Like many stations, the coral community appears to be 

stable and in good health during all years. Though some colonies may appear somewhat 

paler in certain years (e.g., note large Montastraea cavernosa in upper right corner in 

2010), there is no significant tissue loss evident on any colonies. 

61 
Figure 4.4.1. Repetitive photostation 761 from the EFGB in a time series from 2006 to 2010, 

showing a healthy and stable coral community. (A) 2006; (B) 2007; (C) 2008; (D) 

2009; (E) 2010 (NOAA/FGBNMS).

Species composition at repetitive stations was similar to that in the random transects,

with the dominant corals being Montastraea annularis species complex, Diploria

strigosa, Porites astreoides, and M. cavernosa. The M. annularis species complex

had higher cover estimates in the repetitive quadrats (EFGB average from 2009 and 

2010: 49.03%; WFGB average for the same period: 47.03%) than in the random transects

(EFGB average from 2009 and 2010: 31.71%; WFGB average for the same time 

period: 36.62%). The percent cover of Porites astreoides and M. cavernosa in the 

repetitive quadrats were roughly equivalent to the percent cover in the random quadrats. 

(A) (B)

(C) (D)

(E)

62 
Coral disease was absent from analyzed quadrats at both banks in 2009 and 2010. This

may signify a decline in disease within the study areas from past monitoring efforts, 

when low levels of disease were observed (Dokken et al. 2003). Paling and bleaching

were also rare, with the highest levels seen at the WFGB in 2010 (bleached coral 1.9% 

and paling coral 2.8%). Concentrated fish biting and isolated fish biting were similarly

rare at each bank, ranging from 0.0 to 0.03% in all years. 

4.4.2. Deep Station Quadrats 

At the EFGB, nine deep stations located at 32–40 m depths (105–131 ft) were established 

in April of 2003 to monitor the deeper benthic coral community. The coral cover was

high in the deep station quadrats, averaging 82% in 2009 and 2010. This amount of coral 

cover was similar to previous monitoring periods. In 2003, the average deep station coral 

cover was estimated at 76.5%. Between 2004 and 2008, it averaged 72–86% (Figure 

4.4.2). 

The coral community observed in the deep station quadrat photographs appears to be 

both healthy and stable from 2003 to 2010, with consistent percent cover above 70%. 

Higher percent coral cover (~82%) in the deep station quadrats relative to random 

transects has also been documented in previous reports (Dokken et al. 2003; Precht et

al. 2006, 2008b; Zimmer et al. 2010). As with the shallower repetitive stations, the most

likely reason for this difference is that the deep stations were not installed in random 

locations, but were selected as premium monitoring sites with high coral cover in that 

depth range. The deep stations were dominated by Montastraea annularis species

complex. M. cavernosa was the second-most dominant coral species, unlike the

shallower study sites. Lateral growth of colonies of Montastraea annularis species

complex was variable from year to year. Low sample sizes limited statistical power, 

however, making it difficult to draw firm conclusions. 

63 
64 

Figure 4.4.2. Deep station coral cover (+ SE) from 2003 to 2010, averaging above 

70% at the EFGB. 

4.4.3. Planimetry 

Visual and planimetric analysis at repetitive stations generally revealed both small areas 

of living tissue loss and small areas of growth. Coral disease and coral bleaching were not 

apparent during the sampling period. Areas of paling were seen in a small percentage of 

corals. Areas of tissue loss were associated with fish biting and algae growth, though it is 

not clear whether algae competition caused coral mortality or was a result of bare 

substrate becoming available as a consequence of the coral tissue loss. 

Attempts to measure tissue growth and tissue loss in 2009 and 2010 were complicated by 

a change in methods employed. Visual assessments of areas with measured changes 

suggested that estimates were not accurate. The differences are thought to have arisen 

from using a different framer, lens and camera in subsequent years. In 2009, a Nikon 

CoolPix P5000 digital camera was used; in 2010, a Canon PowerShot G11 was used. The 

Canon G11 was used in conjunction with a wide-angle dome port, in order to achieve the 

targeted 8 m2

 image size. While the image area was achieved, barrel distortion was 

observed near the edges of the images. Furthermore, because the angle of acceptance of 

the lenses was different, the cameras had to be located at different heights above the 

bottom to achieve the same sample area. The distortion was corrected in Adobe 

Photoshop®; but the fact that corals were photographed from slightly different angles in 

subsequent years could not be corrected. Therefore, it was not possible to eliminate all 

the error in areal measurement of coral colonies (Figure 4.4.3). 
Note the overlay distortion and height differential (C) resulting from different camera outfits from 

each year. Camera outfit from 2010 resulted in large images, and after image correction, traced 

coral colonies appeared to have exhibited areal regression, thus skewing the analysis 

(NOAA/FGBNMS).

Figure 4.4.3. Coral colonies from repetitive quadrat station 005 at the EFGB outlined for 

planimetric analysis from 2009 (A) and 2010 (B) for a combined overlay (C). 

Because a reliable scale in each picture was not defined before photographing coral 

colonies at each repetitive station, images could not be scaled to adjust for differences in 

area photographed in the two years. Even though the T-frame pole is in each picture, the 

base of the pole itself is too small to be a reliable scale for the areal measurements. 

To eliminate difficulties in future monitoring analyses, FGBNMS staff will use the same 

camera outfit in sequential years as long as possible. The staff at the FGBNMS has also

researched a more suitable camera outfit that covers a large enough image area and has a 

lesser degree of image distortion. A fixed scale will also be added to camera T-frames in 

the future to aid in planimetric image analysis. 

(A) (B)

(C) 

65 
CHAPTER 5.0: SCLEROCHRONOLOGY 

5.1. SCLEROCHRONOLOGY METHODOLOGICAL RATIONALE

Sclerochronology allows the determination of coral growth rates through the measurement 

of the accretionary growth bands deposited by corals in their calcium carbonate skeletons. 

One commonly measured coral growth parameter is accretionary growth. The skeletons 

of many corals, including Montastraea species, contain a consistent sequence of highand low-density bands, comparable to tree rings. As shown in Figure 5.1.1., annual 

growth is represented by each couplet of adjacent high- and low-density bands. Thus, the

rate of skeletal growth can be estimated by measuring the combined width of two 

adjacent growth bands along the length of a corallite. The skeletons of a long-lived corals 

therefore record the histories of coral growth, and it is possible to examine how current 

rates compare with those of the past. Skeletal density and mass growth are additional 

parameters that may be obtained using image analysis densitometry (e.g., Dodge and 

Kohler 1984), though these measures were not used in this study. 

Although the method of counting seasonal density bands within a coral skeleton has been 

used for some time (Knutson et al. 1972; Buddemeier et al. 1974), there still remains 

some uncertainty as to the exact cause of the density variations. In the case of 

Montastraea species living in the Gulf of Mexico and Caribbean Sea, it is generally 

believed that annual low-density bands are produced during much of the year when

favorable growth conditions exist, and annual high-density bands are produced in the 

summer when suboptimal growth conditions occur and the coral is putting more energy 

toward sexual reproduction and less into calcification. Variations in several physical 

environmental factors are known to influence coral skeletal density: (1) light (e.g., 

Macintyre and Smith 1974; Knutson et al. 1972; Wellington and Glynn 1983); (2) 

temperature (e.g., Highsmith 1979; Hudson et al. 1976); and (3) suspended sediment 

(e.g., Dodge et al. 1974; Brown and Howard 1985). Salinity and water agitation may also 

exert some control. Other factors that influence the metabolism of the coral may be 

reflected in skeletal growth, including nutrient availability and reproductive activity 

(Wellington and Glynn 1983; Szmant and Gassman 1990). The roles played by symbiotic 

zooxanthellae in influencing calcification, and endolithic algae in modifying density 

patterns, and the effects of boring organisms, are further complications. 

Lack of high-density summer band deposition, or the occurrence of high-density winter 

stress bands may correspond to times during the year when significant coral bleaching or 

other stresses exist, including cold-air outbreaks, pulses of freshwater influx from rivers, 

concentrated parrotfish biting, and damselfish territory effects (Wells 1963; Buddemeier et 

al. 1974; Dodge 1975, 1980; Hudson et al. 1976, 1989; Kaufman 1977; Highsmith 1979; 

Hudson 1981a, b; Smith et al. 1989; Leder et al. 1991; Fitt et al. 1993; Heiss and Dullo 

1995; Insalco 1996). Care must be taken to differentiate between normal, annual bands 

and other bands produced by stressful non-cyclic environmental fluctuations (Graus and 

Macintyre 1982; Leder et al.1991). 

67 
LD=low-density growth band, HD=high-density 

growth band, and SB=stress band

Figure 5.1.1. X-ray of Montastraea faveolata 

showing skeletal banding

(NOAA/FGBNMS). 

5.1.1. Why Montastraea faveolata? 

The determination of coral growth rate has been identified as one of the best

quantitative measures of assessing coral stress due to disturbance, because this parameter

integrates a variety of physiological processes (Brown and Howard 1985). It is also

widely accepted that coral growth rates may be inherently variable for a single species

within a reef zone and even within individual colonies (Buddemeier and Kinzie 1976). 

Gladfelter et al. (1978) described some species as “conservative” in their growth.

Specifically, they argued that the Montastraea annularis species complex shows 

relatively little response in growth rate to varying environmental conditions compared to 

other species. However, studies have shown significant suppression of M. faveolata

growth may occur if a coral is disturbed, for example, by short-term exposure to high 

concentrations of drilling mud (Hudson and Robbin 1980) or by changes in 

environmental parameters when a coral is transferred from an offshore location to an

inshore site (Hudson 1981b). 

SB

LD

HD

68 
A significant feature revealed by X-radiography is the presence of high-density skeletal

deposits or “stress” bands, which have been observed in sections of Montastraea 

annularis during periods of rapid chilling and mixing of shallow inshore waters

(Hudson et al. 1976; Hudson 1977, 1981a; Shinn et al. 1989) and during periods of 

increased sea surface temperatures and coral bleaching (Leder et al. 1991). 

Previous research on sclerochronology in the Montastraea annularis species complex

has been extensive (Dustan 1975; Hudson et al. 1976; Emiliani et al. 1978; Foster

1980; Hudson 1981a, 1981b; Graus and Macintyre 1982; Dodge and Lang 1983; Dodge 

and Brass 1984; Leder et al. 1991; Slowey and Crowley 1995). It has been shown that

accelerated growth in Montastraea occurs seasonally during cooler periods (Leder et

al. 1991). In Belize, Highsmith (1979) noted that, when compared with Montastraea 

cavernosa and Porites astreoides from the same locality, high- density bands of 

Montastraea annularis appeared to be deposited only for short periods of time, whereas

the low-density bands were generally produced for a greater part of the year.

5.2. SCLEROCHRONOLOGY FIELD AND LABORATORY METHODS

Random Montastraea faveolata core samples from each quadrant at the Flower Garden

Banks are usually collected every two years. Originally, the cores for the monitoring 

period were scheduled to be collected in 2009, but equipment was not available that year. 

The cores were taken at the EFGB in August of 2010 and at the WFGB in November of 

2010. At each bank, four cores were extracted from Montastraea faveolata colonies, for a 

total of eight cores. A pneumatic drill, fitted with a diamond tipped 35 mm (1.38 in) 

lapidary bit, was used to extract short cores of skeletal material from the apex of robust 

M. faveolata colonies. Corals were sampled at their apex and cores were drilled down the 

main growth axis. They were collected from the apex because growth rate is known to 

vary across the surface of an individual coral colony and it is at the apex where the 

maximum rate typically occurs. The longitudinal axis of each core was also oriented as 

closely as possible to the direction that the corallites appeared to grow. Cores were 35 

mm (1.38 in) in diameter and from 54 to 135 mm (2.13 to 5.31 in) long, spanning six to 

nineteen years of growth. Short cores spanning ten or more years of growth were 

recovered because they could be collected quickly and easily. The hole left from core 

extraction was filled with pre-drilled coral skeleton plugs and coral rubble (obtained from 

coral cores extracted during mooring drilling), and LiquidRoc 500® to prevent 

subsequent mortality and bioerosion of the sampled colony. All eight coral cores were 

transferred to the laboratory of Dr. Niall Slowey, Department of Oceanography at Texas 

A&M University, for processing and analysis.

Cores from the field were placed in containers with 95% ethanol solution and kept 

refrigerated until they were shipped in coolers to Texas A&M University for analysis. In 

the laboratory, a custom rock saw with dual diamond blades was used to cut 8-mm-thick 

slabs of coral skeletal material from along the longitudinal axis of each core. These slabs 

were washed, air dried, and then a digital X-ray system was used to collect images that 

revealed variations in the skeletal density and allowed annual couplets of low and high 

density bands, plus high-density stress bands to be identified. 

69 
Ages for the annual band couplets were assigned by counting backwards from the most 

recently deposited colony surface toward older skeletal material. Annual extension 

distances were determined by measuring the thickness of adjacent high and low density 

couplets by grayscale analysis of the digital X-radiographs using CoralXDS software 

(Helmle et al. 2002) from Nova Southeastern University. It was presumed that one year’s 

growth corresponds to the distance from the top of a high density band to the bottom of 

the adjacent low density band (i.e., the thickness of each couplet), representing the annual 

growth rate.

5.3. DATA PRESENTATION AND STATISTICAL ANALYSIS OF GROWTH RATES 

After the annual accretionary growth rates were determined for each coral core, the 

means and standard errors or standard deviations were calculated for each bank and

year. A Student’s t-test assuming equal variances compared values between the EFGB

and WFGB. 

Data are also presented in this report from prior sampling efforts. In 2005, core and X-ray 

processing was conducted at Florida International University (Miami, Florida) and Nova 

Southeastern University (Dania Beach, Florida). In 2007, core and X-ray processing was 

conducted at the Florida Keys National Marine Sanctuary–Upper Keys Office in Key 

Largo, Florida.

5.4. SCLEROCHRONOLOGY RESULTS

X-ray images of the slabs of skeletal material from each core revealed two types of 

distinct high-density bands: annual bands presumably deposited during late summer, and 

stress bands deposited during periods of otherwise rapid growth (Figures 5.4.1 and 5.4.2).

Estimates of annual growth are shown in Table 5.4.1. Standard error values were 

calculated for each bank and year when comparable measurements were available. The 

longest time periods spanned by the coral cores were from 1991 to 2010 for the EFGB 

and from 1998 to 2010 for the WFGB. The core collected from the northwest quadrant at 

EFGB was not used for extension rate comparison due to breakage that occurred around 

the midpoint of that core (i.e., the core barrel caused the two segments to grind together 

during drilling) making it impossible to assign ages to the underlying annual bands. 

The annual growth rates of Montastraea faveolata cores extracted in 2010 at the EFGB 

ranged from 3.0 mm to 7.6 mm. The mean growth for the EFGB between 1991 and 2010 

was 5.5 mm/yr (Table 5.4.1). The maximum occurred between 1996 and 1997 and the 

minimum between 1991 and 1992 (Figure 5.4.3).

The annual growth rate of Montastraea faveolata at the WFGB ranged from a 3.3 mm to 

9.7 mm between 1998 and 2010. The mean at the WFGB between 1998 and 2010 was 6.9 

mm/yr (Table 5.4.1). The maximum growth occurred between 2000 and 2001 and the 

minimum between 2009 and 2010 (Figure 5.4.3).

70 
To determine if the growth rates differed significantly between banks, a Student’s t-test 

was performed. There was a significant difference (α=0.05, P-value=0.0011) in the 

growth rate between the banks when comparable years (1998 to 2010) were analyzed.

There was also a significant difference (α=0.05, P-value=0.0010) when all years (1991 to 

2010) were combined. 

Summer high-density bands are dated. NW=northwest, 

NE=northeast, SE=southeast, SW=southwest.

Figure 5.4.1. X-radiographs of Montastraea faveolata skeletal 

structure in the plane of corallite growth from four 

quadrants in the EFGB long-term monitoring study 

site. 

(A) EFGB NW (B) EFGB NE

(C) EFGB SE (D) EFGB SW

71 
Summer high-density bands are dated. NW=northwest, 

NE=northeast, SE=southeast, SW=southwest.

Figure 5.4.2. X-radiographs of Montastraea faveolata skeletal 

structure in the plane of corallite growth from four 

quadrants in the WFGB long-term monitoring study 

site.

(A) WFGB NW (B) WFGB NE

(C) WFGB SE (D) WFGB SW

72 
Table 5.4.1.

Estimates of Annual Growth (mm/yr) of Montastraea Faveolata from Cores Extracted at the 

EFGB and WFGB in 2010

Coral cores are designated by quadrant in each study site. NW=northwest, NE=northeast, 

SE=southeast, SW=southwest, SE=Standard Error

Year East Flower Garden Bank

NW NE SE SW Mean SE

2009–2010 4.3 5.4 4.9 0.6

2008–2009 6.0 4.2 6.7 5.6 0.7

2007–2008 5.1 4.3 6.2 5.2 0.6

2006–2007 6.0 4.8 6.8 5.9 0.6

2005–2004 7.6 4.2 5.2 5.7 1.0

2004–2005 6.9 5.8 5.4 6.0 0.4

2003–2004 6.3 4.7 6.3 5.8 0.5

2002–2003 4.8 4.9 5.7 5.1 0.3

2001–2002 7.1 3.3 5.2 1.9

2000–2001 6.1 4.9 5.5 0.6

1999–2000 6.9 4.5 5.7 1.2

1998–1999 5.7 5.7

1997–1998 7.3 5.7 6.5 0.8

1996–1997 6.6 7.0 6.8 0.2

1995–1996 6.1 5.3 5.7 0.4

1994–1995 6.6 5.3 6.0 0.7

1993–1994 4.7 4.7

1992–1993 4.9 4.9

1991–1992 3.0 3.0

Mean EFGB Growth Rate (mm) 5.5 0.7

West Flower Garden Bank

Year NW NE SE SW Mean SE

2009–2010 5.1 4.7 5.5 7.3 5.7 0.6

2008–2009 5.6 7.1 3.9 7.3 6.0 0.8

2007–2008 6.7 3.3 6.1 6.9 5.8 0.8

2006–2007 6.9 5.1 7.2 8.0 6.8 0.6

2005–2004 6.5 7.2 5.5 7.8 6.8 0.5

2004–2005 7.3 5.3 7.3 9.7 7.4 0.9

2003–2004 7.1 5.8 7.0 6.6 0.4

2002–2003 7.1 4.6 7.3 6.3 0.9

2001–2002 8.0 6.1 7.5 7.2 0.6

2000–2001 8.8

10.

1 9.5 0.6

1999–2000 4.9 8.2 6.6 1.7

1998–1999 8.4 8.4

Mean WFGB Growth Rate (mm) 6.9 0.8

73 
0.00

1.00

2.00

3.00

4.00

5.00

6.00

7.00

8.00

9.00

10.00

Growth (mm)

Year

Mean Annual Extension Rate in Coral Cores from the East and West 

Flower Garden Banks, 1991-2010

EFGB

WFGB

Figure 5.4.3. Mean annual growth rates in cores collected in 2010 at the EFGB and 

WFGB. 

5.5. SCLEROCHRONOLOGY DISCUSSION

A variety of factors can affect coral growth rate including depth, salinity,

temperature, light, genetic factors, and relative position on the colony (Knutson et al.

1972; Bak 1974; Weber and White 1977; Highsmith 1979; Hudson 1981a; Hudson et al.

1989; Smith et al. 1989). 

Accretionary rates of Montastraea annularis documented over a wide geographic range 

throughout the Caribbean vary from 3.0 to 12.0 mm/yr (Weber and White 1977). 

Growth rates have been shown to vary with depth, with faster growth rates generally

occurring in shallower water (Weber and White 1977). Hudson (1981a) reported 

growth rates of M. annularis in the Florida Keys to be 6.3 mm/yr on offshore reefs and

8.2 mm/yr on mid-shelf reefs from 1928 to 1978. Hudson and Robbin (1980) and 

Deslarzes (1992) reported annual growth rates for 16 colonies of M. annularis at the 

FGB. The mean was 7.9 mm/yr from 1886 to 1907, 8.8 mm/yr from 1907 to 1957, 7.0 

mm/yr from 1957 to 1988, and 9.0 mm/yr during 1988–89 (the last year for which they 

had data). 

Dokken et al. (2001) reported a lower growth rate for 1985–1999, with an average of 6.80 

mm/yr at the EFGB and 5.13 mm/yr at the WFGB. The shorter sampling period was 

offered as a possible explanation for the observed differences. Alternative explanations 

are that the cores analyzed by Dokken et al. were not taken from the apex of the coral 

74 
head, were taken from smaller coral heads than those analyzed by Hudson and Robbin 

(1980) and Deslarzes (1992), and/or that M. franksi may have been mistakenly sampled 

instead of M. faveolata. Each possibility would result in lower estimates that are not 

readily comparable with the Hudson and Robbin (1980) and Deslarzes (1992) data. 

During the 2009–2010 sampling period, Montastraea faveolata growth ranged from 3.0

to 7.6 mm/yr at the EFGB and 3.3–9.7 mm/yr at the WFGB. These results were similar 

to the growth rates reported by Zimmer et al. (2010) for the 2004–2008 long-term 

monitoring reporting periods, but differed slightly from the growth rates reported by

Precht et al. (2006) and the past work by Dokken et al. (2003), who reported a wider

range of growth rates at the EFGB and WFGB. Growth rates for M. faveolata at the 

WFGB, and less so at the EFGB, continued to be in the middle to upper range of 

FGB growth rates as recorded by Hudson and Robbin (1980). 

When compared to the past three coring events (2003, 2005, and 2007), the 2010 data are 

not significantly different (α=0.05, P-value=0.067) with respect to mean growth rates 

(Figure 5.5.1). However, the range in annual growth decreased in 2010 for cores from the 

EFGB and for cores from the WFGB (with the exception of cores from the year 2005). 

In long-term growth studies of Montastraea faveolata corals at the Flower Garden Banks, 

Hudson and Robbin (1980) and Deslarzes (1992) observed that a late 1950s growth 

decline is a prominent feature of the records. Studies first hypothesized this decline could 

be caused by lower light levels due to bank subsidence resulting from salt dome 

dissolution (Rezak and Bright 1981), or local variations in water temperature or outflow 

from the Atchafalaya River (Dodge and Lang 1983; Szmant-Froelich 1984) with a 

positive correlation between and winter/spring temperature and growth, and a negative 

correlation between Atchafalaya River discharge and growth. The analysis by Slowey 

and Crowley (1995) supported the temperature explanation and fit it into a large scale

climatic context. They compared meteorologic and coral growth records and found that 

during the past century changes in coral growth at the FGB correspond to changes in the 

winter climate of the Gulf of Mexico/southeastern USA. Slowey and Crowley (1995) 

found that recent decadal scale variability in both coral growth and regional wintertime 

climate are closely linked to changes to the orientation of the mid-latitude atmospheric jet 

stream over North America. When the jet stream has a more north-south orientation, 

wintertime temperatures in the Gulf of Mexico/southeastern USA are colder and the 

corals exhibit slower growth and possess more winter stress bands. 

75 
0

2

4

6

8

10

12

2003 2005 2007 2010

Extension (mm)

Year

Mean Extension in Coral Cores from the East and West 

Flower Garden Banks 

EFGB

WFGB

Figure 5.5.1. Mean annual growth rates (+SE) based on analysis of Montastraea

faveolata 2003, 2005, 2007, and 2010 cores from the EFGB and WFGB. 

Variations in the orientation of the jet stream and wintertime climate are controlled by the 

Pacific North American Pattern of wintertime climate variability, with a north-south jet 

stream orientation being associated with the positive phase of the pattern (Wallace and 

Gutzler 1981). Significant interdecadal variations in the Pacific North American Pattern 

occur. Studies indicate that the pattern is among the dominant modes of extratropical 

climate variability in the Northern Hemisphere (Simmons et al. 1983; Wallace et al. 

1993). 

Hudson (1981a, 1984) observed that Montastraea faveolata corals within the Florida 

Keys National Marine Sanctuary show a decline in growth rates and deposited abundant 

winter stress bands during the late 1950’s and early 1960’s. Fluctuations in river 

discharge cannot account for changes in Florida Keys coral growth. The similarity in 

temporal variations in extension rate changes and stress banding displayed by both 

Flower Garden Banks and Florida Keys corals supports the contention that coral growth 

in both locations is most likely impacted by regional changes of wintertime climate 

(Slowey and Crowley 1995). 

In the summer of 2010, the FGB experienced an unusual warming of the water column 

and comparatively high rates of coral bleaching, which were reflected in the data at 

WFGB. As described in the random transect data, approximately 7% of the corals at the 

WFGB were bleached and 11% were paling. The M. annularis species complex was the 

species most impacted by bleaching and paling. The 2010 coral core collection occurred

during this bleaching event; the coral cores did not indicate a large drop in growth

during the 2009–2010 monitoring period. However, cores collected for the next 

monitoring period may display reduced growth rates due to the 2010 bleaching event. 

76 
CHAPTER 6.0: LATERAL GROWTH

6.1. LATERAL GROWTH METHODOLOGICAL RATIONALE

Diploria strigosa is the second largest contributor to coral cover at the FGB after the 

Montastraea annularis species complex (Bright et al. 1984; USDOI, MMS 1998; Dokken 

et al. 2003; Gittings et al. 1992; Precht et al. 2006; Precht et al. 2008b; Zimmer et al. 

2010). The lateral margins of selected D. strigosa colonies were monitored and 

photographed annually to detect any incipient changes over time and space. D. strigosa is 

more suitable than other coral species due to the conspicuous patterns and grooves that, 

when photographed, can be matched when repetitive annual photographs are overlaid. 

6.2. LATERAL GROWTH FIELD AND LABORATORY METHODS

Sixty lateral growth stations, located on the margins of Diploria strigosa colonies, were 

maintained at each bank (Figure 6.2.1). Divers were equipped with a camera, a close-up 

kit (close-up framer), and strobe. In the 2009 field season, a Nikonos V 35 mm 

underwater film camera with a Nikkor 28 mm lens was used initially at the EFGB for the 

first 35 station images. The camera aperture was set at f/22, focal distance at infinity, and 

TTL strobe control. A 13.3 x 19.7 cm image was captured. However, due to mechanical 

failure of the Nikonos camera, an alternate camera system was used for the remainder of 

the 2009 data collection. A Canon G6 digital camera with Ikelite housing and Ikelite 

digital strobe DS125 was used to capture the remaining lateral growth station images for 

2009. The camera was set at auto exposure, ISO 100, on macro and autofocus settings, 

and the strobe was set to TTL. This created an image larger than the close up kit framer, 

which was later cropped using Photoshop CS5® to the match sizes. For the 2010 field 

season, a replacement digital camera system was evaluated with the close-up kit (closeup framer), and strobe. A Canon Power Shot G11 in Fisheye Fix housing with a standard 

port was used. The camera was set at auto exposure, ISO 200, on macro and autofocus 

settings, and strobe set to TTL. The framer was placed on corner pins at each station, 

ensuring a repeated image of the station. Some stations were missing identification tags. 

Those stations that did have tags were photographed with the tag in the frame. For 

stations without tags, the current photographs were matched with past photographs using 

the ridge patterns of the Diploria strigosa colonies. 

In 2009, 41 and 40 colonies of Diploria strigosa were photographed on the EFGB and 

WFGB, respectively. In 2010, 29 and 34 lateral growth stations were photographed at the 

EFGB and WFGB, respectively. Of the photographs taken, five matching pairs were 

comparable from 2009 and 2010 for the EFGB, and nine matching pairs were comparable 

for the WFGB. Several factors contribute to the large discrepancy in the number of 

photographs collected versus the number of photographs that are useful during analysis. 

These factors include: (1) photographs are not taken in the same position or orientation 

each year due to missing bolts, photographer error, and changes in the rugosity of some 

colonies and (2) some stations no longer have margins to measure in a photograph due to 

colony growth or death. It is also important to note that more than 60 lateral growth 

77 
stations are located at each bank. In the past, stations have been abandoned and 

rehabilitated and new stations have also been installed. During some monitoring events, 

all lateral growth stations were marked for photography, both old and new, while during 

other monitoring cruises only the new stations were marked. Regardless of the method 

used to mark stations, the collection of lateral growth photographs ceased when the 

number of stations photographed totaled 60, or as close to 60 as possible given the time 

constraints at each study site. In other words, while there may have been nearly 60 stations 

photographed in two consecutive years, there were far fewer than 60 photo comparisons 

possible for the reasons listed above.

Note overgrowth of marker bolts by coral and the approach of the advancing edge of the 

coral to the station boundary, even extending beyond it in some places.

Figure 6.2.1. Lateral growth station #52 on a Diploria strigosa colony with close-up framer 

at the EFGB (NOAA/FGBNMS). 

6.2.1. Image Analysis for Lateral Growth

Images corresponding to a specific lateral growth station were compared between 

consecutive years (2009 and 2010). Lateral differences in the margins of the Diploria 

strigosa colonies were evaluated by overlaying the pairs of photographs, using Photoshop 

CS5®, and dividing the lateral growth edge of the colony vertically into areas categorized 

as “growth,” “retreat,” “stable,” and “not available” (for areas that were dark, shadowed, 

or out of focus). Using ImageJ® (a public domain, Java-based image processing program 

developed at the National Institutes of Health), the entire horizontal width of the frame 

78 
was set to 100 units, then the horizontal distances of growth, retreat, and stability were 

measured (Figure 6.2.2). These values were then combined to obtain an overall 

percentage of growth, retreat, and stability for each image. Successive photographs of a 

given colony were aligned using the colony’s ridge patterns. 

The red line is the 2009 margin and the blue line is the 2010 margin. (A) growth (red 

line) on station #12 and (B) retreat (blue line) on station #13. Sections where lines 

overlap demonstrate stasis (NOAA/FGBNMS).

Figure 6.2.2. Image analysis of Diploria strigosa lateral growth at the EFGB. 

(A)

(B)

79 
6.3. DATA PRESENTATION AND STATISTICAL ANALYSIS FOR LATERAL 

GROWTH

Proportional annual changes in the lateral margin of individual Diploria strigosa

colonies, whether positive or negative, were examined per bank per year. A percent 

change value was calculated for each comparable image, and each image was assigned an 

overall designation category of growth, retreat, or stability from 2009 to 2010. Lateral 

growth data was not collected during the 2008 annual monitoring cruise due to the 

constraints of weather; therefore, there was no comparable data from 2008 to 2009. 

Growth, retreat, and stability of colony margin tissue were calculated by subtracting the 

linear distance for each category measured for each station for the current year from the 

linear distance for each category measured during the previous year. Areas of the picture 

that were not distinguishable during analysis were not included in the total percentage. 

Percent changes were calculated by determining the linear margin gained or lost and 

dividing by the total units of the photographed colony margin and multiplying by 100. 

Due to the low sample size from 2009 through 2010, a repeated measures Analysis of 

Variance with enough power was not possible for analysis. Therefore, these data are used 

for descriptive purposes only. 

6.4. LATERAL GROWTH RESULTS

Proportion of marginal tissue growing, retreating, or remaining stable in lateral growth of 

individual colonies, whether positive or negative, were examined by site (EFGB and 

WFGB) and by year (2009 and 2010). Of the photographs taken in 2009 and 2010, only 

five were comparable for the EFGB, and nine were comparable for the WFGB. 

Throughout the 2009 to 2010 monitoring period, a large number of photographs were 

unsuitable for analysis due to a variety of reasons, including but not limited to, 

photographer error, improper orientation, missing guide bolts, and no visible margins in 

the photo due to colony growth or death. Due to the low number of comparable 

photographs, no statistics were used to analyze the lateral growth rates of change. Appendix 

3 contains the lateral growth data from 2009 and 2010 at the FGB. 

There was a high degree of variability between progression/regression percentages among 

years for both the EFGB and WFGB (Table 6.4.1). At the EFGB, there was an overall 

increase in marginal growth from 2009 to 2010. The WFGB also showed a slight increase in 

Diploria strigosa marginal growth. 

80 
Table 6.4.1.

Lateral Growth Stations on Diploria Strigosa and Percent of Margin Growing, Retreating, or 

Remaining Stable of Coral Tissue at the EFB and WFGB from 2009 to 2010

Site Percent Change from 2009 to 2010

EFGB Station Growth Retreat Stable

EFGB 10 6.73 59.88 33.39

EFGB 12 40.25 44.16 15.58

EFGB 13 91.50 8.50 0.00

EFGB 29 92.47 7.53 0.00

EFGB 34 51.56 48.44 84.66

WFGB Station Growth Retreat Stable

WFGB 68 82.43 0.00 17.57

WFGB 69 100.00 0.00 0.00

WFGB 73 59.53 12.80 30.37

WFGB 94 0.00 92.58 7.42

WFGB 95 45.37 28.97 25.28

WFGB 99 79.15 15.52 5.33

WFGB 106 24.37 63.61 12.02

WFGB 118 0.00 67.63 32.37

WFGB 121 0.00 50.30 49.70

Ternary plots were produced to compare the relative amounts of progression, regression, 

and stasis at the lateral growth stations. These compare the relative amounts of gain, loss, 

or no change without regard to change along the margins. Each station provided single 

estimates of each measure (growth, reatreat, and stability), regardless of the number of 

areas of progression or regression present at the FGB. 

At both the EFGB and WFGB, the proportions of marginal growth, from 2009 to 2010

were high. Greater than 50% of all the marginal length analyzed was advancing (Figures 

6.4.1 and 6.4.2). 

81 
82 

Proportion of D. strigosa Colony Margins Growing, Retreating, or 

Remaining Stable at the East Flower Garden Bank, 2009-2010 

Figure 6.4.1. Ternary diagram showing the proportion of Diploria strigosa marginal 

tissue growing, retreating, or remaining stable at each station 

photographed at the EFGB from 2009 to 2010. 
Proportion of D. strigosa Colony Margins Growing, Retreating, or 

Remaining Stable at the Wast Flower Garden Banks, 2009–2010 

0.1

0.1

0.1

0.2

0.2

0.2

0.3

0.3

0.3

0.4

0.4

0.4

0.5

0.5

0.5

0.6

0.6

0.6

0.7

0.7

0.7

0.8

0.8

0.8

0.9

0.9

0.9

Regression

Progression

Stasis

106

118

121

68

69

73

94

95

99

WFGB

Station

Figure 6.4.2. Ternary diagram showing the proportion of Diploria strigosa marginal 

tissue growing, retreating, or remaining stable at each station 

photographed at the WFGB from 2009 to 2010. 

6.5. LATERAL GROWTH DISCUSSION

Net lateral growth of Diploria strigosa was positive during the 2009 and 2010 study 

period; however, variability in the growth rates was high. For future monitoring efforts,

sample sizes sufficient for statistical analysis across multiple intervals (which was not 

possible in this study or 2009 and 2010 comparison) will need to be collected to ensure

that the initial area does not confound the detection and interpretation of pattern. 

Lateral growth measurements have been used for much of the monitoring history of the 

FGB and results have shown overall growth of monitored margins, with high variability

among individual colonies (Gittings et al. 1992; Dokken et al. 2001, 2003; Precht et al.

83 
2006; Zimmer et al. 2010). Lateral growth measurements do not take into account the 

fact that individual corals may grow at different rates along different margins. For 

example, corals can die or retreat much faster than they can grow or advance, since 

disease and other factors causing mortality are capable of destroying tissue at a rapid rate, 

while growth occurs at a comparatively slow and more constant rate. While some 

marginal tissue may be advancing, other tissue on the same colony may be retreating, 

affecting the overall picture of lateral change in a given colony and by extension on 

a given bank and year. Also, lateral growth measurements do not take into account the 

accretionary growth of Diploria strigosa. 

Other factors have affected the quality of lateral growth data from the long-term 

monitoring study for many years. Past researchers have encountered problems with 

locating lateral growth photostations, photographing the stations in a consistent way (e.g., 

at the same angle every time), and assessing photos that have not been taken in the proper 

orientation. Locating the photostations in the field (60 per bank) has proven to be time 

consuming due to missing pins and tags. The stations appear to have a short useful life

because the colonies can overgrow the small area within the station in a short period of 

time. With 60 stations on each bank, it has also been a laborious task to keep them 

maintained, as they have to be re-established or repaired frequently, and there is not 

enough time on the monitoring cruises to repair or find new stations since other tasks take 

a priority. As of the 2010 research season, few comparable stations remained at either 

bank. 

In 2010, many of the stations established years ago were completely overgrown by coral. 

This has resulted in the loss of many stations. Confusion by the frequent loss of marker 

pins and large changes in stations has resulted in photographs being taken in different

orientations, making analysis difficult or impossible. Analysis of the photos has been 

difficult because it is hard to obtain true repetitive photographs since the framer is placed 

directly on the colonies. Because of the direct contact, the effects of increased rugosity 

and the changing orientation of margins caused by bioerosion, and accretionary growth 

also hinders repetitive photography. It is also difficult to replace lost corner bolts at 

precise locations. Lastly, the study site maps, largely based on previous researcher’s 

maps, are continuously updated but are still not fully reliable. Due to these problems, it is 

recognized that not enough time is allotted for this effort. Improvements in methodology, 

mapping, establishing new stations, and extending cruise days to allow time for these 

tasks are necessary. 

84 
CHAPTER 7.0: PERIMETER VIDEOGRAPHY 

7.1. PERIMETER VIDEOGRAPHY METHODOLOGICAL RATIONALE

Portions of the perimeter lines were videotaped each year at the EFGB and WFGB to 

document change at known locations along the perimeter of the study sites. General 

aspects of coral condition were documented and compared year to year. 

7.2. PERIMETER VIDEOGRAPHY FIELD AND LABORATORY METHODS

Divers videotaped two 100 m segments of the perimeter lines at the EFGB (north and 

east margins) and WFGB (south and west margins) in 2009 and 2010. At the EFGB, 

divers began at the northwest corner of the 100 m x 100 m study site and videotaped the 

north line to the northeast corner, then swam the east line to the southeast corner. At the 

WFGB, divers captured footage of the south and west lines, beginning at the southeast 

corner and ending at the northwest corner. The videographer maintained an approximate 

2.0 m (6.5 ft) distance above the bottom. The camera was aimed downward at a 45° angle 

to capture the substratum. In both years, a 360° panoramic view of the reef was 

videotaped at the three corners documented during the perimeter video. In both years, a 

Sony® Handicam® DCR-TRV950 video camera in a Light and Motion® Bluefin®

housing with Light and Motion® Sunray® video lights. A red filter was used for color 

correction. 

The video footage was reviewed to record the general condition of corals along the 

perimeter of the study sites. Individual coral colonies displaying possible disease, 

bleaching, paling, and tissue loss due to fish biting were identified and recorded. 

Analysis categories were as follows: disease, bleaching, paling, concentrated fish biting,

and isolated fish biting. Concentrated fish biting (CFB) represents the concentrated biting 

that removes the coral polyps completely from an affected area and may be due to 

activity of the parrotfish Sparisoma viride (Bruckner and Bruckner 1998; Bruckner et al. 

2000). Isolated fish biting describes less dense and smaller-scale fish biting, typically 

representative of damselfish territories. Affected coral colonies were compared in 2009 

and 2010. Changes in coral colony condition were recorded. The perimeter surveys are 

intended to provide a general overview of ecosystem health. The analyses were 

qualitative; therefore, no statistical analyses were conducted on these data.

7.3. PERIMETER VIDEOGRAPHY RESULTS

The perimeter video was reviewed for a qualitative analysis of the general condition of 

corals along the perimeter of the study sites. In previous study years, fish population 

levels along the perimeter lines were also observed; however, due to the difficulty in 

identifying fish species in the videos, fish population information was collected in a more 

robust way, as described in a separate section of this report. The review of the 2009 and 

2010 perimeter videos suggests that, in general, the coral community along the 

perimeter lines at the EFGB and WFGB study sites displayed low levels of stress and

85 
high cover. The most distressed corals were affected by paling and bleaching at 

WFGB in 2010. These results were comparable to random transect and repetitive quadrat 

data, although no statistical comparisons were made. Furthermore, no evidence of disease 

was observed at either bank. 

7.3.1. EFGB Perimeter Lines 

Two incidences of bleaching were observed at the EFGB in 2009, one being a

Montastraea faveolata colony and the other a Montastraea annularis colony. In 2010,

isolated fish biting (typical of damselfish) occurred on five colonies, followed by 

algal overgrowth (one colony), paling (one colony), and bleaching (seven colonies). 

Montastraea franksi was the coral species most impacted by these stressors. A

comparison of corals affected by fish biting, disease, algal growth, paling, and

bleaching in 2009 and 2010 at the EFGB is shown in Table 7.3.1. 

Table 7.3.1.

Comparison of Observations of the Condition of Individual Coral Colonies at EFGB in 2009 and 

2010

B= Bleaching, P= Paling, A= Overgrown by algae, D= Disease, IFB= Isolated fish 

biting, CFB= Concentrated fish biting

Number of Colonies Coral Species EFGB 2009 EFGB 2010

1 Montastraea annularis B

1 Montastraea faveolata B

2009 Total Affected Colonies = 2

1 Diploria strigosa B

1 Diploria strigosa P

1 Diploria strigosa A

1 Montastraea annularis IFB

1 Montastraea cavernosa B

1 Montastraea cavernosa IFB

4 Montastraea franksi B

2 Montastraea franksi IFB

1 Porites asteroides IFB

1 Stephanocoenia intersepta B

2010 Total Affected Colonies = 14

7.3.2. EFGB 360° Panoramic Views

At the northwest corner, corals appeared to be in good condition in both 2009 and

2010. In 2009, there were two Montastraea spp. Colonies that where paling and tissue 

loss occurred due to concentrated fish biting from numerous fish present. In 2010, it 

86 
appeared these two colonies had recovered and were no longer paling. At the northeast 

corner, the corals appeared to be in good condition, with little evidence of bleaching 

stress or fish biting. Large fish populations were noted, including ocean trigger fish in 

2009. 

The southeast corner appeared to be in good health. There was little evidence of paling 

and bleaching in 2009, and one coral head displaying fish biting. The fish populations 

were similar between years and included Creole wrasse, Creolefish, and damselfish. 

7.3.3. WFGB Perimeter Lines

In 2009, concentrated fish biting occurred on three colonies, isolated fish biting on one, 

and paling on three. Montastraea faveolata was the coral most impacted by these 

stressors. In 2010 at the WFGB, coral stress included bleaching (98 colonies), paling

(27 colonies), concentrated fish biting (seven colonies), algal over growth (one colony), 

and disease (one colony). Montastraea cavernosa and Millepora alcicornis were the 

most impacted coral species (Table 7.3.2). 

Table 7.3.2.

Comparison of Observations of the Condition of Individual Coral Colonies at WFGB from 2009 

and 2010

B= Bleaching, P= Paling, A= Overgrown with algae, D= Disease, IFB= Isolated fish biting, CFB= 

Concentrated fish biting

Number of Colonies Coral Species WFGB 2009 WFGB 2010

1 Millepora alcicornis P

1 Montastraea annularis IFB

1 Montastraea faveolata P

3 Montastraea faveolata CFB

1 Stephanocoenia intersepta P

2009 Total Affected Colonies = 7

1 Colpophyllia natans B

1 Colpophyllia natans P

4 Diploria strigosa P

2 MASC B

1 MASC CFB

32 Millepora alcicornis B

46 Montastraea cavernosa B

1 Montastraea cavernosa CFB, A

8 Montastraea cavernosa P

1 Montastraea faveolata B

1 Montastraea faveolata CFB

1 Montastraea faveolata D

87 
Number of Colonies Coral Species WFGB 2009 WFGB 2010

1 Montastraea faveolata P

2 Montastraea franksi B

8 Montastraea franksi P

1 Porites astreoides B

13 Unidentified Coral B

4 Unidentified Coral CFB

5 Unidentified Coral P

2010 Total Affected Colonies = 133

7.3.4. WFGB 360° Panoramic Views

The southeast corner coral health was good in 2009 and 2010, with only one coral head 

affected by fish biting in 2009. In 2010, three colonies of Montastraea cavernosa and six 

colonies of Millepora alcicornis exhibited bleaching. 

At the southwest corner, only one Montastraea spp. colony appeared to be paling in 

2009. However, in 2010, the colony from the year before, six colonies of Montastraea 

cavernosa and 18 colonies of Millepora alcicornis exhibited paling or bleaching. Large 

fish populations were noted, including Ocean Triggerfish in 2009. Large barracudas were 

observed at the southwest corner in 2009. 

The northwest corner coral health was good in 2009, with only one conspicuous paling 

Millepora alcicornis coral head. A large school of Brown Chromis was observed near 

the corner. In 2010, many more coral heads were bleached (nine colonies of Montastraea 

cavernosa and ten colonies of Millepora alcicornis, including the colonly from 2009). 

7.4. PERIMETER VIDEOGRAPHY DISCUSSION

Videography of the perimeter lines and 360° panoramic views of the corner markers at the 

EFGB and WFGB provided a general overview of coral condition at the study sites from

2009 to 2010. Similar to the findings from the random transects, coral condition 

appeared to be relatively good at both banks in both years. There were no signs of coral

disease. In previous years, the most noticeable impacts to coral colonies were

concentrated and isolated fish biting; however, only a few incidences of fish biting were 

observed. In 2009 and especially in 2010, the most common impacts were coral paling 

and bleaching. For example, in Figure 7.4.1, a head of Porites astreoides is viewed as a 

healthy colony in the 2009 perimeter video, but shows as a bleached colony in the 2010 

perimeter video. 

It is important to note that a number of human errors may have influenced the qualitative 

data provided by the perimeter video and the 360° panoramic views. First, while the 

perimeter lines at both banks were generally in the same locations between years, the 

lines did shift (Figure 7.4.1). This is due to the affect of currents and surge on the flexible 

perimeter lines between the fixed corner markers, which are 100-m apart. In the future, 

88 
there are plans to install mid-transect eye bolts during study site refurbishment activities 

to assist in improving repeatability so that transect lines can be attached to eye bolts to 

avoid shifting lines with the current. 

Furthermore, there is an occasional need for reinstallation of missing corner markers, 

which usually is a slightly different position from previous locations. Shifting perimeter 

lines and corner marker positions result in a lack of overlapping video footage and fewer 

available coral comparisons. In addition, due to operator error, the 2 m height of the 

camera above the substratum and the 45o angle were not always maintained, changing the 

view of the bottom and affecting the analysis. 

Figure 7.4.1. A head of Porites astreoides observed to be 

healthy in 2009 (A), but bleached in 2010 (B) 

at the WFGB. 

(A)

(B) 

89 
CHAPTER 8.0: QUALITATIVE FIELD OBSERVATIONS

8.1. QUALITATIVE FIELD OBSERVATIONS METHODOLOGICAL RATIONALE

In addition to the annual data collection protocol, other biologically relevant information 

is documented on the reefs of the FGBNMS. During each annual monitoring cruise and 

other research cruises, observations of general coral reef health and notable biological 

and oceanographic events (e.g., spawning, animal behavior) were qualitatively assessed 

and documented. 

8.2. QUALITATIVE FIELD OBSERVATIONS METHODS

As divers traversed the EFGB and WFGB study sites during the 2009 and 2010 annual 

monitoring cruises, they noted and photographically documented any biologically 

relevant observations and/or events. 

8.3. QUALITATIVE FIELD OBSERVATION RESULTS

8.3.1. Wildlife Observations

8.3.1.1. Elasmobranch Sightings

During the 2009 EFGB annual monitoring cruise, three manta rays (Manta birostris)

were sighted during diving operations. Each manta ray has a unique set of markings and 

color patterns on its ventral side. These can be used to identify individual rays. Of the 

three mantas seen at the EFGB in 2009, two were newly recorded mantas and one was a 

resighting of manta number M46 (Manta Photo Catalog discussed in Section 8.4) (Figure 

8.3.1). One of the unidentified mantas was missing a portion of its posterior end and its tail. 

Manta rays were not sighted at the WFGB in 2009 during the monitoring period.

In 2010, two mantas were sighted at the EFGB, one measuring approximately 2.4 m (8 ft) in 

length and the second approximately 1.8 m (6 ft) in length. At the WFGB, a 12-foot manta 

was sighted in 2010. Also at the WFGB, two tiger sharks (Galeocerdo cuvier), one 

approximately 3 m (10 ft) in length and the other 2.4 m (8 ft) in length, were sighted in

2010. 

91 
Figure 8.3.1. Manta ray M46 observed at the EFGB in 2009 

with spot and square ventral markings (J. 

Wiseman and S. Bernhardt). 

8.3.1.2. Fish Sightings

At the EFGB deep stations, approximately five black grouper (Mycteroperca bonaci) and

yellowmouth grouper (Mycteroperca interstitialis) were observed by divers in 2009. In 

2010, a rare peppermint bass (Serranus sp.) was seen at the EFGB. In 2009 at the WFGB, a 

slipper lobster was observed, and a barracuda was seen at a cleaning station near a 

Siderastrea spp. coral head. At the WFGB in 2010, nesting sergeant major (Abudefduf 

saxatilis) damselfish were observed by divers. Two large black grouper were observed 

near the WFGB sand patch, and a large school of amberjack were also observed, although 

they are not frequently seen at the FGB. A large loggerhead sea turtle (Caretta caretta) 

was observed surfacing for a breath near the R/V Manta. 

8.3.2. Coral Health Observations

8.3.2.1. Qualitative Coral Health Assessments

During the 2009 and 2010 annual monitoring cruises, scientific divers made qualitative 

observations of coral colonies exhibiting signs of disease or other coral health issues. During 

the 2009 monitoring cruise at the EFGB, divers experienced 100 foot visibility and water 

92 
temperatures near 29°C (84°F). The Madracis auretenra (formerly named Madracis 

mirabilis) fields to the east of the study site were observed to be recovering from 

September 2008 Hurricane Ike impacts (Locke 2009). 

In 2010, divers experienced a thermocline at approximately 12 m (40 ft), up to 21 m (70 

ft) visibility, and a strong current due to a tropical depression that was moving into the 

Gulf of Mexico, causing conditions to change throughout the week. Visibility ranged 

from 15–21 m (50–70 ft). Divers observed false bleaching (white pigmentation that 

appears in certain corals), and an unidentified plague-like disease on multiple heads of M. 

annularis, M. franksi, and M. faveolata. The summer 2010 season was distinguished by 

increases of fluorescent orange algae growth at the EFGB and WFGB (Figure 8.3.2). The 

orange alga was very noticeable in the crevices of the reef, and samples were collected 

and sent to Dr. Suzanne Fredericq’s Seaweeds Laboratory at the University of Louisiana 

at Lafayette for identification. The alga was identified as Martensia pavonia, a delicate 

net-forming alga that is a member of the Delesseriaceae family and exhibits a bright 

orange color in its deteriorating state. Algal blooms are classic indicators of high nutrient 

levels on coral reefs, and tend to expand in the warm season and during the rainy season 

when more nutrients are flushed offshore from land.

Figure 8.3.2. Fluorescent orange algae, Martensia pavonia, 

observed at the EFGB in 2010 (NOAA/FGBNMS). 

There were no unusual observations at the WFGB in 2009, but in 2010, divers observed 

bleached corals at the WFGB, most likely due to warm seawater temperatures. The most 

93 
common bleached corals included Montastraea cavernosa, Millepora alcicornis, and 

Siderastrea siderea. 

8.3.2.2. Acropora palmata 

During the June 2005 monitoring cruise, a colony of Acropora palmata (elkhorn coral)

was discovered outside the EFGB study site, in close proximity to the southeast corner 

marker. Acropora palmata is a reef-building coral that has not been previously 

encountered at the FGB. The colony was located at a depth of 23.5 m (77.1 ft) and 

measured approximately 0.5 m (1.6 ft) in width and 1.0 m (3.3 ft) in height, with a 

maximum branch length of 30 cm (11.8 in) (Zimmer et al. 2006). As of October 2009, 

this Acropora palmata colony exhibited branch loss, which was likely due to the passage

of Hurricane Ike in September 2008. In addition, the colony also displayed tissue loss 

on its southern side due to an unidentified cause and a white band of exposed coral

skeleton on one of the branches. Algal farming ( non-destructive cropping of algae, 

which promotes exponential growth for grazing) by a Threespot Damselfish (Stegastes 

planifrons) was also evident on the colony and may have contributed to the coral tissue

loss. Despite these changes, in 2010 the colony displayed a normal yellow-brown color 

and appeared to be recovering and in good health (Figure 8.3.3). 

In July 2003, an A. palmata colony was found at a depth of 21.6 m (70.9 ft) on the 

WFGB. This colony included an encrusting basal plate and one small branch. As of May

2005, the colony measured 0.6 m (2 ft) wide by 0.5 m (1.6 ft) high with a maximum

branch length of 8.8 cm (3.5 in). In both 2009 and 2010, the colony appeared unhealthy, 

and suffering from disease. Algae covered the coral in places where it had already died

(Figure 8.3.4). 

94 
Figure 8.3.3. The Acropora palmata colony at the EFGB in (A) 2005 (B) 2006 and (C)

2010. 

Note the resident damselfish in photo (A) (NOAA/FGBNMS). 

(C)

(A) (B)

95 
Figure 8.3.4. The only Acropora palmata colony at the WFGB is 

shown to be dying from disease in (A) 2008 (B)

2009 and (C) 2010. 

(C)

(A)

(B)

(C)

96 
Algae are taking over in places where the coral has 

already died (NOAA/FGBNMS). 

8.3.3. Exotic and Invasive Species

In 2002, invasive orange cup coral (Tubastraea coccinea) was first documented at the 

EFGB. Orange cup coral is native to the Indo-Pacific and may have entered the South 

Atlantic and Caribbean by attaching to a ship’s hull, having its larvae discharged in 

ballast water, or being transported on reused structures such as drilling rigs or production 

platforms. This species is now common on oil and gas platforms in the Gulf of Mexico, 

and it is suspected that platforms played a role in the spread of this species. In 2009 and 

2010, sporadic colonies of cup coral were observed on the reefs near the EFGB and 

WFGB study sites (Figure 8.3.5). 

Figure 8.3.5. Orange cup coral (Tubastraea coccinea), an invasive species from the 

Pacific that is beginning to establish itself at the FGBNMS

(NOAA/FGBNMS). 

97 
8.4. QUALITATIVE FIELD OBSERVATIONS DISCUSSION

8.4.1. Wildlife Observations

8.4.1.1. Elasmobranch Sightings

Sanctuary staff and volunteers have been collecting photos and videos of mantas sighted 

in the sanctuary over many years. These photos are the basis of a FGBNMS Manta Ray 

Photo Catalog, identifying each of the known individuals and the additional of new 

individuals. The catalog can be viewed online and downloadable as a poster (Figure 

8.4.1). The manta rays in the catalog have been divided into categories to aid in 

identifying individuals. Each manta ray is placed in a category according to its unique 

underbelly markings (i.e., spots, squares, spots and squares, mostly black, mostly white, 

and tagged). The catalog is an ongoing project, so new mantas and additional sightings of 

current mantas are added at every opportunity. Of the six mantas observed during 2009 

and 2010, one could be positively identified as having been seen in the past. 

Figure 8.4.1. FGBNMS Manta Ray sighting catalog. The manta rays in the 

catalog have been divided into categories to aid in 

identifying individuals, each by its unique ventral markings

(NOAA/FGBNMS).

The two tiger sharks observed in November 2010 at the WFGB are among approximately 

20 species of sharks and rays have been documented at the Flower Garden and Stetson 

98 
Banks; some occurr seasonally, others year-round. During the late fall and winter months, 

frequent users of the sanctuary include tiger sharks, schooling scalloped hammerheads, 

and spotted eagle rays. 

8.4.2. Coral Health Observations

8.4.2.1. Qualitative Coral Health Assessments

In November of 2010, divers observed a higher than normal number of bleached coral 

colonies at the WFGB. The most common were Montastraea cavernosa and Millepora 

alcicornis. This was most likely a result of warm sea surface water temperatures in late 

summer of 2010, which exceeded the 30°C coral bleaching threshold for the Flower 

Gardens (Hagman and Gittings, 1992) during that season. Because these observations 

were taken after most of the data was collected during the August and October 2010 

cruises, the bleaching of Millepora alcicornis is not reflected in the bleaching results

from earlier sections of this report. 

In September of 2010, NOAA's Coral Reef Watch Program's satellite data provided nearreal time data on the reef environmental conditions in the Gulf of Mexico to quickly 

identify areas at risk for coral bleaching. The Coral Reef Watch uses remote sensing and 

in situ tools for near-real-time and long term monitoring, modeling and reporting of 

physical environmental conditions of coral reef ecosystems. Areas near and inside the 

FGBNMS boundaries were coded at bleaching alert “Watch” levels and “Alert Level 2” 

in the fall of 2010 (Figure 8.4.2). 

Coral Reef Watch is part of the NOAA Coral Reef Conservation Program (CRCP) and 

the National Environmental Satellite Data and Information Service (NESDIS). The 

satellite product suite is a key aspect to NOAA's monitoring system for coral reef 

ecosystems, the Coral Reef Ecosystem Integrated Observing System (CREIOS). 

Continuous monitoring of sea surface temperature at global scales provides researchers 

and managers with tools to understand and better manage the complex interactions 

leading to coral bleaching. When bleaching conditions occur, these tools can be used to 

trigger bleaching response plans and support appropriate management decisions. 

It should be noted that monitoring at the EFGB was conducted in August 2010, before

bleaching and before signs were observed. However, monitoring at the WFGB occurred 

in the late fall when bleaching was at its peak and significantly visible. Based on 

qualitative surveys, the EFGB and WFGB response to bleaching are similar, so this could 

be considered a temporal comparison, and signs of coral bleaching stress, or coral 

recovering from bleaching stress, may be observed at the EFGB as well. 

99 
Areas near and inside the FGBNMS boundaries were coded at bleaching alert 

“Watch” levels and “Alert Level 2” in the fall of 2010 (NOAA/CRCP). 

Figure 8.4.2. Coral Reef Watch bleaching alert map for the Gulf of Mexico in 

September 2010. 

In 2010, divers also observed the fluorescent orange algae, Martensia pavonia, at the 

EFGB and WFGB. Algal growth often indicates excessive levels of nutrients moving from 

the coastal zone into offshore habitats, stimulating production. Coral reef ecosystems are 

sensitive to high concentrations of nitrogen and phosphorus, which promote algal 

overgrowth. Besides algal cover, reefs with high levels of nutrients in the water may be 

more susceptible to coral diseases. Along developed coastlines excess nutrients come from 

human sources, such as human sewage, livestock manures, and agricultural runoff. Nutrients 

carried offshore act as fertilizers for marine plants growth. Algae can be considered an early 

warning of changes in water quality and potential impacts to coral health. 

8.4.2.2. Acropora palmata 

Historically, Acropora palmata (elkhorn coral) and A. cervicornis (staghorn coral) were

two of the most important reef-building coral species in the Caribbean (Bruckner 2002; 

Precht and Aronson 2006). On May 9, 2006, A. palmata was officially placed on the 

Endangered and Threatened Species List (71 FR 26852) as a threatened species. 

Populations of these acroporid species in the Caribbean were decimated in the 1970s and 

1980s by white band disease, with few apparent signs of recovery (Aronson and Precht 

2001b). Researchers estimate that the population of A. palmata in the Caribbean is less 

than 5% of their historical abundance (before the 1970s decline; Bruckner 2002). Threats 

to Acropora spp. include disease, coral bleaching, predation, storm damage, and human 

activities. 

100 
The Acropora palmata colony discovered on the EFGB during the June 2005 annual 

monitoring cruise represents the deepest report of A. palmata from the Caribbean and 

western Atlantic regions, as well as the first record of Acropora spp. anywhere in the 

northern Gulf of Mexico (Zimmer et al. 2006). A. palmata is typically considered a 

shallow-water species, primarily occupying depths of less than 5 m (Lighty et al. 1982). 

The virtual absence of this species from the reefs of the FGB has been ascribed to cold 

winter water temperatures, the substantial depths of the reef caps at the EFGB and 

WFGB (18 m minimum for both banks), and the remoteness of the FGB from potential 

sources of A. palmata larvae (e.g., the Florida Keys and Mexican reefs of the southern 

Gulf of Mexico; Bright et al. 1984, Schmahl et al. 2008). The A. palmata colony branch

loss is most likely due to the passage of Hurricane Ike in 2008, and tissue loss on the 

colony may be due to algal farming by damselfish. The FGBNMS researchers continue 

to monitor the status of the two A. palmata colonies, and follow-up studies are proposed 

to document and explain the turn-on and turn-off mechanisms for Acropora reef 

development on these isolated reef complexes (Schmahl et al. 2008). 

8.4.3. Exotic and Invasive Species

Invasive species have recently become a concern for marine resource managers because 

of the inherent potential negative impacts that they can cause. Tubastraea coccinea, 

orange cup coral, was first documented within the FGBNMS in 2002 (Fenner and Banks 

2004) and has further established itself on nearby oil and gas infrastructure, as well as in 

discrete locations within the sanctuary. Tubastraea coccinea began to colonize Geyer 

Bank, which is located 52 km (32 mi) east of the EFGB, and approximately 50 colonies 

of T. coccinea were removed by sanctuary divers in 2004. Since that time, T. coccinea

has become well established at Geyer Bank, and has also been documented near both the 

EFGB and WFGB study sites. 

T. coccinea is a zooxanthellate scleractinian coral that is an exotic, invasive species 

within the Caribbean and western Atlantic (Fenner 1999, 2001; Fenner and Banks 2004). 

Native to the tropical Indo-Pacific and the eastern Atlantic (Cairns 2000), T. coccinea

was first reported in the Caribbean in 1943 (Fenner and Banks 2004). No fossil evidence 

of this species has been found within the Caribbean (Cairns 1999). Tubastraea coccinea

is typically located on the undersides of rocks or massive corals, in caves, and on rock 

walls (Glynn et al. 2008). It appears to compete particularly well on artificial substrates, 

which may account for its widespread dispersal. It is a hermaphroditic brooding coral that 

releases planula larvae year round (Cairns 2000; Glynn et al. 2008) and has a mean 

growth rate of approximately 3 cm2/year (Vermeij 2006). This species reaches 

reproductive maturity at a small size (from as small as 2–10 polyps; Glynn et al. 2008) 

and at an early age (reproductively viable at approximately 1.5 years; Vermeij 2006). T.

coccinea has the ability to compete effectively by forming thin tissue outgrowths 

(“runners”) that extend over the substrate until suitable substrate is encountered, at which 

time a new polyp forms (Vermeij 2005). These competitive mechanisms may put native 

benthos at risk. 

From its native Indo-Pacific range, Tubastraea coccinea has now been introduced to the 

B

101 
waters off Asia, Africa, Australia, North America, Central America, and South America 

(IUCN 2007; Ferreira 2003; Fenner and Banks 2004; Glynn et al. 2008). This species was 

first observed in the Caribbean in 1943 by Vaughn and Wells, and has since spread 

throughout the Caribbean and Bahamas (Glynn et al. 2008), Gulf of Mexico (Fenner and 

Banks 2004), and Brazil (Figueira de Paula and Creed 2004). Possible mechanisms of 

introduction to these regions include boat/ship hulls, ballast water, transport of marine 

structures/machinery (e.g., oil platforms; Ferriera 2003; Fenner and Banks 2004). T. 

coccinea has colonized many of the oil and gas platforms in the northwestern Gulf of 

Mexico (Sammarco et al. 2006). Based on the proximity of the many T. coccineacolonized platforms to the reefs of the FGB, along with this invasive species’ effective 

dispersal capacity, the FGB is potentially at risk for further invasion by T. coccinea

(Sammarco et al. 2006). 

It is recommended that the FGBNMS continue to monitor for Tubastraea coccinea, 

Thecacera pacifica (a sea slug documented at Stetson Bank in 2006), and other exotic 

species. Exotic species have the potential to harm native species via competition for 

space or resources, or by harboring pathogens or parasites. If a T. coccinea invasion 

becomes at problem at the EFGB and/or WFGB, an opportunistic removal program 

should be initiated. In Brazil, the Universidade do Estado do Rio de Janeiro (UERJ) has 

instituted a removal/eradication program titled “Projecto Coral-Sol” to eliminate the 

potential threat of T. coccinea in that region. 

Over the last five years, one of the world’s most popular ornamental aquarium fish, the 

native Indo-Pacific lionfish (Pterios volitans/miles), has invaded much of the southeast 

Atlantic and Caribbean region, beginning with the escape of a few individuals from 

aquaria in Florida about a decade ago (Morris and Whitfield 2009) (Figure 8.4.3). The 

proliferation of this species has occurred quickly due to early sexual maturation, high 

fecundity, ability to invade many habitats, and effectiveness in competing for food (they 

are voracious predators that have no known predators in the region) (Mumby et al. 2011). 

Lionfish subject small-bodied and juvenile reef fish to greatly elevated predation, and 

coral reefs are at risk of phase shifts mediated by secondary effects of changes in fish 

trophic structure (e.g. increasing algal cover caused by the loss of herbivores). The longterm consequences on reef biodiversity and function are not yet clear, but are a matter of 

grave concern to resource managers. 

Lionfish were first captured in the southern Gulf of Mexico off the northern Yucatan 

Peninsula in December 2009. In September 2010, two lionfish were sighted at Sonnier 

Bank by Texas A&M University Galveston researchers. These were the first confirmed 

sightings of lionfish at the natural banks in the northwestern Gulf of Mexico, about 60 

miles east of East Flower Garden Bank. In July 2011, the first lionfish to invade 

sanctuary waters was observed by divers. The FGBNMS will continue to monitor the 

sanctuary for this invasive species. The FGBNMS staff is currently in the process of 

working with other staff at sanctuary sites to develop Lionfish Research and Management 

plans. While there is little scientific evidence to support using biocontrol as a natural 

means to slow the invasion of the lionfish, the EFGB and WFGB provides a unique 

opportunity to conduct an experiment to measure the effectiveness of natural predation, 

102 
and the influence of fishing on this natural control mechanism. The managers of the 

FGBNMS are currently in the process of investigating the use of an experimental 

research area to assess these questions. 

Figure 8.4.3. Invasive lionfish at the West Flower Garden Bank. 

Lionfish are native to the Indo-Pacific region and invaded the Gulf of Mexico in 2009 

(NOAA/FGBNMS).

103 
CHAPTER 9.0: WATER QUALITY

9.1. WATER QUALITY METHODOLOGICAL RATIONALE

During the reporting period, Sea-Bird and HoboTemp dataloggers deployed at the EFGB 

and WFGB recorded variations of temperature and salinity. Temperature and salinity 

depth profiles were also collected opportunistically throughout the field season, using an

YSI probe. Water samples collected quarterly from the sea surface to the reef cap at the 

EFGB and WFGB were analyzed for chlorophyll a and nutrients (ammonia, nitrate, and 

nitrite, soluble reactive phosphorous and total Kjeldahl nitrogen [TKN]). 

Hereafter, “water quality” will refer to the physical (temperature) and biological 

(chlorophyll a [chl a]), and chemical (salinity and nutrients) characteristics of the 

seawater overlying the FGB. This report presents the results of the water quality 

monitoring at the EFGB and WFGB conducted from July 2009 to November 2010, 

although temperature and salinity profiles were collected from January 2009 to December 

2010. 

9.2. WATER QUALITY FIELD AND LABORATORY METHODS

9.2.1. Sea-Bird Conductivity and Temperature Recorder

The Sea-Bird Electronics, Inc. (SBE) 37-SMP MicroCAT with RS-232 serial interface is 

a high-accuracy conductivity and temperature recorder designed for long-term 

oceanographic deployment (www.seabird.com). The MicroCATs used at the FGBNMS 

included pressure and sound velocity sensors. The primary role of the MicroCAT in this 

study was to accurately record temperature and salinity. The specifications (and typical 

stability) of the MicroCAT indicate an initial accuracy of 0.003 mS/cm (conductivity) 

and 0.002°C (temperature). The resolution of the instrument is 0.0001 mS/cm and 

0.0001°C (Figure 9.2.1). 

One Sea-Bird datasonde was deployed at the EFGB (23 m or 75 ft water depth) near buoy 

number five and one at the WFGB (27 m or 88.5 ft water depth) near buoy number two. 

Sand flats were used as deployment locations to accommodate the secure attachment of 

the datasondes to galvanized train wheels (Figure 9.2.2). Water quality data were 

recorded every 30 min. These instruments are returned to Sea-Bird Electronics, Inc., in 

Bellevue, Washington for annual calibration and maintenance.

9.2.1.1. Specific Conductance

Datasondes used a cell with four nickel electrodes to measure solution-conductance. 

Two of the electrodes were current driven, and two were used to measure the drop in 

voltage. Differences were converted into a specific conductance value and reported in 

milli-Siemens (milliohms). Salinity was later derived from the conductivity and 

105 
temperature readings according to accepted algorithms and reported as practical salinity 

units (PSU).

Figure 9.2.1. Sea-Bird 37-SMP MicroCAT water quality instrument at the 

EFGB (NOAA/FGBNMS).

Figure 9.2.2. FGBNMS research divers mounting the Sea-Bird 37-SMP 

MicroCAT water quality instrument on a rack in the sand at the 

EFGB (NOAA/FGBNMS).

106 
9.2.1.2. Temperature

The datasondes used a thermistor of sintered metallic oxide that changed predictably in 

resistance with variation in temperature. The algorithm for conversion of resistance to 

temperature was built into the datasonde software, and accurate temperature readings in 

degrees Celsius (°C), Kelvin (°K), or Fahrenheit (°F) were provided automatically. No 

user calibration or maintenance of the temperature sensor was necessary.

9.2.2. HoboTemp Thermographs

One HOBO Pro v2 Water Temperature Data Logger (HoboTemp) was attached to each 

of the Sea-Bird instruments as backup recorders of water temperature. HoboTemp 

recorders have an accuracy of ± 0.2°C and resolution is 0.02°C at 25°C. They are 

designed with a durable streamlined case for extended deployment in fresh or salt water, 

and equipped with an Optic USB interface for data offload in the field. The data loggers 

were deployed in a water depth of 23 m (75 ft) at the EFGB and in at 27 m (88.5 ft) at the 

WFGB and recorded every 30 min.

9.2.3. YSI Probe

During each cruise, opportunistic temperature profiles were measured by researchers 

using an YSI water quality sensor deployed by hand (Figure 9.2.3). Temperature and 

salinity were recorded at 20 m, 15 m, 10 m, 5 m, 4 m, 3 m, 2 m, 1 m, and the surface.

These data complement the data collected by the stationary water quality instruments on 

the sea floor at each bank by providing additional information about the conditions

throughout the water column. 

Figure 9.2.3. FGBNMS researchers conducting seawater 

temperature profiles using a YSI probe 

through the moon pool of the R/V Manta

(NOAA/FGBNMS).

107 
9.2.4. Chlorophyll a and Nutrients

Surface (<1 m), midwater (10 m), and near bottom (20 m) water samples were acquired 

at six different times on the EFGB and WFGB between July 2009 and November 2010 

(Table 9.2.1). During each sampling event, water was collected twice at each depth using 

a vertical 10 liter sampling bottle (Niskin®). Water samples were immediately transferred 

into pre-cleaned polyethylene and glass containers (tested monthly using nanopure water) 

provided by an independent, U.S. Environmental Protection Agency (USEPA) certified 

analytical laboratory (Anacon, Inc. in Houston, TX). Water samples were analyzed for 

chl a, ammonia, nitrate, nitrite, TKN, and soluble reactive phosphorous. Water samples 

for chl a analyses were collected in 1000 ml glass containers with no preservatives. 

Samples for reactive soluble phosphorous were placed in 250 ml bottles with no 

preservatives. Ammonia, nitrate, nitrite, and TKN samples were collected in 1000 ml 

bottles with sulphuric acid (H2SO4) as a preservative. One blind duplicate water sample 

was taken at one of the sampling depths on one of the banks for each sampling period. 

Within minutes of sampling, labeled sample containers were stored on ice at 4°C and a 

chain of custody was initiated. Once back onshore, the samples were sent to Anacon, Inc.

for analysis using standard USEPA methods (Table 9.2.2) to assess concentrations of chl 

a and nutrients (ammonia, nitrate and nitrite, TKN, soluble reactive phosphorous). 

Table 9.2.1. 

Water Sampling Schedule, Depth, and Number of Samples Taken at the EFGB and WFGB in

2009 to 2010 

EFGB WFGB

Sampling 

Date Depth Samples Sampling Date Depth Samples

7/7/2009 1, 10, 20 m 6 7/8/2009 1, 10, 20 m 6

11/24/2009 1, 10, 20 m 6 11/24/2009 1, 10, 20 m 6

3/14/2010 1, 10, 20 m 6 3/14/2010 1, 10, 20 m 6

5/27/2010 1, 10, 20 m 6 5/27/2010 1, 10, 20 m 6

9/11/2010 1, 10, 20 m 6 9/13/2010 1, 10, 20 m 6

11/10/2010 1, 10, 20 m 6 11/9/2010 1, 10, 20 m 6

108 
Table 9.2.2. 

Standard USEPA Methods Used to Analyze Water Samples Taken at the FGB 

mg/m3 = milligrams per cubic meter; mg/l = milligrams per liter

Parameter Method Detection Limit

Chlorophyll a 10200HPLC 1-mg/m3

Ammonia E350.3 0.03–mg/l

Nitrate E350.3 0.15–mg/l

Nitrite E353.2 0.15–mg/l

Soluble reactive phosphorous 300.0 0.40–mg/l

Soluble reactive phosphorous SM-4500-P 0.01–mg/l

Total Kjeldahl nitrogen (TKN) E351.3 0.10–mg/l

Total Kjeldahl nitrogen (TKN) E351.3 0.55–mg/l

Total Petroleum Hydrocarbons THP-GRO - 1005 < 5

In May 2010, hydrocarbons were added to the suite of analysis parameters in response to 

the Deepwater Horizon oil spill. This measure was only added to this single sampling 

event as part of the oil spill response. 

9.3. WATER QUALITY RESULTS

9.3.1. Sea-Bird Temperature and Salinity

9.3.1.1. Temperature

Seawater temperature was simultaneously measured on the reef caps of the EFGB (23 m 

or 75 ft water depth) and WFGB (27 m or 88.5 ft water depth) using a Sea-Bird 37-SMP 

MicroCAT datasonde from January 2009 to December 2010. The seawater temperature 

records were complete, with the exception of a gap from 01/01/2009 to 01/21/2009 at the 

EFGB due to a sensor malfunction that was reset on 01/22/2009. The temperature records 

include the winter minimum and summer maximum, and daily average temperatures were 

calculated. Appendix 5 contains the water quality data from 2009 and 2010 at the FGB.

At the EFGB, temperature ranged from a minimum of 20.41°C to a maximum of 29.53°C 

in 2009. The annual mean temperature was 25.11°C. In 2010, the temperature ranged 

from 17.40°C to 30.69°C. The 2010 mean temperature was 24.46°C (Table 9.3.1). 

Several thermal anomalies were recorded at the EFGB. There were episodes of low 

temperature in summer (June–September) of 2009 ranging from 25.45–27.54°C on the 

reef cap of the EFGB , as well as a sudden decrease in temperature (to 26.21°C) in late 

July 2010 at the EFGB. Late spring (March–May) temperatures in 2010 were also 

unusually low (17.40–21.40°C) at the EFGB. The temperature over the reef cap was high 

(29.26–30.69°C) during the summer and fall (July–November) of 2010. While the 

maximim temperature at the EFGB was less than 30°C in 2009, the maximum in 2010 

exceeded 30°C. This is considered the coral bleaching threshold for the Flower Garden 

Banks (Hagman and Gittings 1992) (Figure 9.3.1). 

109 
Table 9.3.1.

Summary of Sea-Bird Seawater Temperature Parameters from 2009 to 2010 

Measurement

2009 2010

EFGB WFGB EFGB WFGB

Annual Mean 

Temperature (°C) 25.11 24.95 24.46 23.96

Annual Minimum 

Temperature (°C) 20.41 19.76 17.40 17.63

Annual Maximum 

Temperature (°C) 29.53 29.50 30.69 30.64

16

18

20

22

24

26

28

30

32

1-Jan

10-Jan19-Jan28-Jan

6-Feb

15-Feb24-Feb

4-Mar

13-Mar22-Mar31-Mar

9-Apr

18-Apr27-Apr

6-May

15-May24-May

2-Jun

11-Jun20-Jun29-Jun

8-Jul

17-Jul26-Jul

4-Aug

13-Aug22-Aug31-Aug

9-Sep

18-Sep27-Sep

6-Oct

15-Oct24-Oct

2-Nov

11-Nov20-Nov29-Nov

8-Dec

17-Dec26-Dec

Temperature (C)

Date

Sea-Bird Annual Temperature Variation at the East Flower 

Garden Bank, 2009-2010

2009

2010

Data obtained from Sea-Bird datasonde.

Figure 9.3.1. Daily average sea water temperature measured near the reef cap at the 

EFGB from 2009 to 2010. 

110 
16

18

20

22

24

26

28

30

32

1-Jan9-Jan

17-Jan25-Jan

2-Feb

10-Feb18-Feb26-Feb

5-Mar

13-Mar21-Mar29-Mar

6-Apr

14-Apr22-Apr

30-Apr

8-May

16-May24-May

1-Jun9-Jun

17-Jun25-Jun

3-Jul

11-Jul19-Jul27-Jul

4-Aug

12-Aug20-Aug28-Aug

5-Sep

13-Sep21-Sep29-Sep

7-Oct

15-Oct23-Oct31-Oct

8-Nov

16-Nov24-Nov

2-Dec

10-Dec18-Dec26-Dec

Temperature (C)

Date

Sea-Bird Annual Temperature Variation at the West Flower 

Garden Bank, 2009-2010

2009

2010

Data obtained from Sea-Bird datasonde. 

Figure 9.3.2. Daily average sea water temperature measured near the reef cap at the 

WFGB from 2009 to 2010. 

At the WFGB, the annual daily mean temperature ranged from a minimum of 19.76°C to 

a maximum of 29.50°C in 2009. The annual mean temperature was 24.95°C. In 2010, 

temperature ranged from 17.63°C to 30.64°C. The annual mean temperature was 23.96°C 

(Table 9.3.1). 

Several thermal anomalies were also recorded at the WFGB. There were episodes of low 

summer (July–September) temperature in 2009 (25.79–26.29°C), and a sudden decrease 

in temperature in June (21.85°C) and late July (24.83°C). Late spring (March–May) 

temperatures were also abnormally low (17.63–19.56°C) in 2010. The temperature over 

the reef cap was high (29.76–30.64°C) during the summer and fall (July–November) of 

2010. Like the EFGB, the highest temperature in 2010 at the WFGB exceeded the 30°C 

coral bleaching threshold (Figure 9.3.2). 

9.3.1.2. Salinity

Salinity records at the WFGB were complete between January 2009 and December 2010. 

At the EFGB, there were data gaps from 01/01/2009 to 01/21/2009 and from 06/30/2010 

to 12/4/2010, both due to a sensor malfunctions Salinity ranged from approximately 31 to 

38 PSU at the EFGB and WFGB from 2009 to 2010. 

At the EFGB, the annual daily mean salinity ranged from 34.74 PSU to 36.99 PSU in 

2009. The annual mean salinity was 36.53 PSU. In 2010, salinity ranged from 34.51 PSU 

to 37.75 PSU. The annual mean salinity was 36.56 PSU (Table 9.3.2). At the WFGB, the 

111 
annual range was 30.97 to 36.68 PSU in 2009. The annual mean salinity was 35.18 PSU. 

In 2010, salinity ranged from 34.12 to 38.17 PSU. The annual mean was 36.37 PSU 

(Table 9.3.2). 

Table 9.3.2.

Summary of Sea-Bird Salinity Parameters from 2009 to 2010 

Practical salinity unit (PSU)

Measurement

2009 2010

EFGB WFGB EFGB WFGB

Annual Mean PSU 36.53 35.18 36.56 36.37

Annual Minimum PSU 34.74 30.97 34.51 34.12

Annual Maximum PSU 36.99 36.68 37.75 38.17

The WFGB experienced a period of low salinity from May to late August of 2009, even 

dropping below 31 PSU at one point. This was followed by salinities around 36 PSU in 

late August. The WFGB also experienced a smaller drop in salinity from September to 

November 2010. For the remainder of the 2009–2010 monitoring period, salinity was 

stable at approximately 36 PSU (Figure 9.3.3). 

28.000

30.000

32.000

34.000

36.000

38.000

40.000

1-Jan-09

1-Feb-09

1-Mar-09

1-Apr-09

1-May-09

1-Jun-09

1-Jul-09

1-Aug-09

1-Sep-09

1-Oct-09

1-Nov-09

1-Dec-09

1-Jan-10

1-Feb-10

1-Mar-10

1-Apr-10

1-May-10

1-Jun-10

1-Jul-10

1-Aug-10

1-Sep-10

1-Oct-10

1-Nov-10

1-Dec-10

Practical salinity unit (PSU)

Date

Sea-Bird Annual Salinity Variation at the East and West Flower 

Garden Banks, 2009-2010 

EFGB

WFGB

Data obtained from Sea-bird datasonde.

Figure 9.3.3. Salinity measured near the reef cap at the EFGB and WFGB from 2009 

and 2010. 

112 
9.3.2. HoboTemp Thermograph Data

HoboTemp thermographs were attached to each of the Sea-Bird instruments at both 

banks from January 2009 through December 2010. The HoboTemp record provided 

similar data to that captured by the Sea-Bird datasondes: low average temperatures in the 

summer of 2009, an unusually cool spring in 2010, and high summer temperatures in 

2010 (Figures 9.3.4. and 9.3.5). 

At the EFGB, the HoboTemp temperature ranged from 20.35°C to 29.48°C in 2009. The 

annual mean was 25.07°C. In 2010, the annual mean ranged from 17.33°C to 30.08°C. 

The annual mean was 24.16°C (Table 9.3.3). At the WFGB, it ranged from 19.56°C to 

29.67°C in 2009, with a mean of 24.91°C. In 2010, the range was 17.63°C to 30.77°C. 

The mean was 24.61°C (Table 9.3.3). These were not significantly different from the 

Sea-Bird temperature measurements. 

16

18

20

22

24

26

28

30

32

1-Jan 1-Mar 1-May 1-Jul 1-Sep 1-Nov

Temperature (C)

Date

Hobo Annual Temperature Variation at the East Flower 

Garden Bank, 2009-2010

2009

2010

Figure 9.3.4. Daily average seawater temperature measured near the reef cap at the 

EFGB from 2009 to 2010. Data obtained from HoboTemp 

thermographs. 

 

113 
16

18

20

22

24

26

28

30

32

1-Jan 1-Mar 1-May 1-Jul 1-Sep 1-Nov

Temperature (C)

Date

HoboAnnual Temperature Variation at the West Flower 

Garden Bank, 2009-2010

2009

2010

Figure 9.3.5. Daily average seawater temperature measured near the reef cap at the 

WFGB from 2009 to 2010. Data obtained from HoboTemp 

thermographs. 

Table 9.3.3.

Summary of HoboTemp Seawater Temperature Parameters from 2009 to 2010

Measurement

2009 2010

EFGB WFGB EFGB WFGB

Annual Mean 

Temperature (°C) 25.07 29.67 24.16 24.61

Annual Minimum 

Temperature (°C) 20.35 19.56 17.33 17.63

Annual Maximum 

Temperature (°C) 29.48 24.91 30.08 30.77

9.3.3. YSI Vertical Profiles

During each cruise, opportunistic seawater temperature profiles were measured by 

researchers using an YSI probe deployed by hand. Temperature was recorded at 20 m, 15 

m, 10 m, 5 m, 4 m, 3 m, 2 m, 1 m increments, and at the water surface (Table 9.3.4). 

These data complement the data collected by the stationary water quality instruments on 

the sea floor at each bank by giving additional information, if needed, about the 

conditions at the surface and throughout the water column. YSI profiles corroborated the 

high water temperatures in August 2010. They also indicate the lack of a pronounced 

thermocline between the surface and 20 m during any times of the year. 

114 
Table 9.3.4.

Summary of Opportunistic YSI Seawater Temperature Vertical Profiles from 2009 to 2010 at the 

EFGB and WFGB 

NA=Not available

Temperature (°C)

Depth (m)

Bank Buoy Date Time 0.0 1.0 2.0 3.0 4.0 5.0 10.0 15.0 20.0

EFGB 7 1/21/2009 20:15 21.1 21.1 21.1 21.1 21.1 21.1 21.1 21.1 21.1

EFGB 7 1/22/2009 11:05 21.1 21.1 21.1 21.1 21.1 21.1 21.1 21.1 21.1

EFGB 1 3/14/2010 11:45 18.8 18.8 18.8 18.8 18.8 18.8 18.7 18.7 18.7

EFGB 1 5/27/2010 12:21 26.9 26.9 26.9 26.9 26.9 27 26.8 25.8 25.1

EFGB 1 6/5/2010 NA 28.1 28.0 28.0 28.0 27.9 27.8 27.5 27.4 27.6

EFGB 4 7/30/2010 17:55 30.8 30.7 30.7 30.6 30.4 30.4 30.1 29.7 29.4

EFGB 4 8/31/2010 23:03 30.2 30.3 30.3 30.3 30.3 30.3 30.3 30.3 30.3

EFGB 2 8/31/2010 NA 30.2 30.2 30.2 30.2 30.2 30.2 30.2 30.2 30.2

EFGB 5 9/11/2010 7:09 29.6 29.5 29.6 29.6 29.7 29.7 29.9 29.8 29.8

EFGB 5 9/11/2010 18:35 30.7 30.5 30.2 30.0 30.0 30.0 30.0 29.9 NA

EFGB 5 9/12/2010 6:44 30.2 30.2 30.2 30.2 30.2 30.1 24.9 29.8 NA

EFGB 4 11/10/2010 15:16 25.2 25.2 25.2 25.2 25.2 25.2 25.3 25.3 NA

EFGB 4 12/4/2010 16:25 23.4 23.4 23.4 23.4 23.4 23.4 23.4 23.5 23.5

WFGB 2 1/22/2009 NA 21.1 21.1 21.1 21.1 21.1 21.1 21.1 21.1 21.1

WFGB 4 3/14/2010 13:53 18.7 18.7 18.7 18.7 18.6 18.6 18.6 18.5 18.4

WFGB 1 5/27/2010 14:45 27 27 26.9 26.9 26.9 26.9 26.7 26.6 25

WFGB 1 6/5/2010 NA 27.5 27.5 27.4 27.3 27.3 27.3 27.2 27.1 26.8

WFGB 1 7/30/2010 19:45 30.7 30.6 30.6 30.6 30.6 30.7 29.6 29.7 28.2

WFGB 1 7/31/2010 7:09 30.1 30.1 30.1 30.1 30.1 30.1 30 29.5 29.7

WFGB 2 8/30/2010 NA 30.5 30.5 30.5 30.4 30.4 30.3 30.2 30.2 30.2

WFGB 1 9/1/2010 9:54 30.2 30.2 30.2 30.2 30.2 30.2 30.2 30.2 30.3

WFGB NA 9/13/2010 6:59 30.1 30.1 30.1 30.1 30.1 30.1 30.1 30 29.9

WFGB NA 9/14/2010 11:59 30.1 30.1 30.0 30.0 30.0 30.0 29.9 29.8 29.6

WFGB 5 11/9/2010 9:55 25.2 25.2 25.2 25.2 25.2 25.2 25.2 25.3 25.3

WFGB 5 10/10/2010 9:03 25.3 25.3 25.3 25.3 25.3 25.3 25.3 25.3 25.4

115 
9.3.4. Water Samples

9.3.4.1 Chlorophyll a and Nutrients

Surface (<1 m), midwater (10 m), and near bottom (20 m) water samples were acquired 

at 6 different times on both the EFGB and WFGB for the 2009 and 2010 monitoring 

period. Water samples were analyzed for chlorophyll a (chl a) and nutrients (ammonia, 

nitrate, nitrite, soluble reactive phosphorous, and total Kjeldahl nitrogen [TKN]). 

Approximately 15% of the water samples collected contained concentrations of chl a that 

were above detectable limits (>1 mg/m3). Most were those collected in May 2010, 

specifically those from midwater and at the reef cap at the EFGB, and throughout the 

water column at the WFGB. The highest concentration detected was 2.14 mg/m3 of chl a

(Table 9.3.5). 

Table 9.3.5.

Chlorophyll a Concentrations in Water Samples Taken at the EFGB and WFGB from 2009 to 

2010

ND=Not detected at reporting limit

Chlorophyll a 

(Detection limit:

1 mg/m3)

2009 2010

7/7/2009 

– 7/8/2009 11/24/09 03/14/10 05/27/10 9/11/2010 –

9/13/2010

11/09/2010 

–

11/10/2010

EFGB Surface A ND ND ND ND ND ND

EFGB Surface B ND ND ND ND ND ND

EFGB Midwater A ND 2.14 ND 1.34 ND ND

EFGB Midwater B ND ND ND 1.07 ND ND

EFGB Reef Cap A ND ND ND 2.14 ND ND

EFGB Reef Cap B ND ND ND 1.87 ND ND

WFGB Surface A ND ND ND 1.34 ND ND

WFGB Surface B ND ND ND 1.07 ND ND

WFGB Midwater A ND ND ND 1.07 ND ND

WFGB Midwater B 2.14 ND ND 1.6 ND ND

WFGB Reef Cap A ND ND ND 1.07 ND ND

WFGB Reef Cap B ND ND ND ND ND ND

Eighteen percent of water samples contained concentrations of ammonia above detectable 

limits. Ammonia was detected in all samples taken from 07/07 to 08/2009 at the EFGB 

and WFGB and ranged from 0.05 to 0.09 mg/l (Table 9.3.6). The modal value for 

116 
ammonia levels was 0.03 mg/l. Samples containing the greatest amount of ammonia were 

obtained at the WFGB on 07/08/2009. 

In approximately 87% of water samples, the nitrate concentrations were below detection 

limits (0.15 mg/l). The only samples that contained detectable levels were those collected 

at both banks in November 2009 and the WFGB in September 2010 (Table 9.3.6). Nitrite 

was not detected in any of the tested samples.

TKN was detectable in all water samples collected at the EFGB and WFGB in 2009, but 

only sporadically after that. Concentrations ranged from 0.84 to 13.9 mg/l (Table 9.3.6). 

Approximately 10% of the samples from 2010 contained TKN concentrations above 

detectable limits, ranging from 0.10 to 1.34 mg/l. Soluble reactive phosphorous was not 

detected in any of the tested samples.

Table 9.3.6.

Concentrations of Ammonia, Nitrate, and TKN in Water Samples Taken at the EFGB and WFGB 

from 2009 to 2010 

Nitrite and soluble reactive phosphorous were not detected in any samples. ND=Not detected at 

reporting limit. 

 2009 2010

Ammonia

(Detection limit:

0.03 mg/l)

7/7/2009 

– 7/8/2009 11/24/09 03/14/10 05/27/10 9/11/2010 –

9/13/2010

11/09/2010 

–

11/10/2010

EFGB Surface A 0.06 ND ND ND ND ND

EFGB Surface B 0.06 ND ND ND ND ND

EFGB Midwater A 0.08 ND ND ND ND ND

EFGB Midwater B 0.07 ND ND ND ND ND

EFGB Reef Cap A 0.07 ND ND ND ND ND

EFGB Reef Cap B 0.06 ND ND ND ND ND

WFGB Surface A 0.06 ND ND ND ND ND

WFGB Surface B 0.08 ND ND ND ND ND

WFGB Midwater A 0.05 ND ND ND ND ND

WFGB Midwater B 0.09 ND ND 0.1 ND ND

WFGB Reef Cap A 0.06 ND ND ND ND ND

WFGB Reef Cap B 0.06 ND ND ND ND ND

Nitrate 

(Detection limit: 

0.15 mg/l)

7/7/2009 

– 7/8/2009 11/24/09 03/14/10 05/27/10 9/11/2010 –

9/13/2010

11/09/2010 

–

11/10/2010

EFGB Surface A ND 0.17 ND ND ND ND

EFGB Surface B ND ND ND ND ND ND

EFGB Midwater A ND 0.18 ND ND ND ND

EFGB Midwater B ND ND ND ND ND ND

EFGB Reef Cap A ND ND ND ND ND ND

117 
 2009 2010

Ammonia

(Detection limit:

0.03 mg/l)

7/7/2009 

– 7/8/2009 11/24/09 03/14/10 05/27/10 9/11/2010 –

9/13/2010

11/09/2010 

–

11/10/2010

EFGB Reef Cap B ND 0.18 ND ND ND ND

WFGB Surface A ND 0.2 ND ND 1.34 ND

WFGB Surface B ND ND ND ND ND ND

WFGB Midwater A ND 0.16 ND ND ND ND

WFGB Midwater B ND 0.27 ND ND 0.88 ND

WFGB Reef Cap A ND ND ND ND ND ND

WFGB Reef Cap B ND ND ND ND 2.52 ND

TKN 

(Detection limit:

0.10 mg/l)

7/7/2009 

– 7/8/2009 11/24/09 03/14/10 05/27/10 9/11/2010 –

9/13/2010

11/09/2010 

–

11/10/2010

EFGB Surface A 0.98 4.70 ND ND ND ND

EFGB Surface B 1.40 13.90 ND 1.12 ND ND

EFGB Midwater A 1.96 10.60 ND ND ND ND

EFGB Midwater B 1.26 2.40 ND ND ND ND

EFGB Reef Cap A 1.96 1.60 0.7 ND ND ND

EFGB Reef Cap B 1.26 3.70 ND ND ND ND

WFGB Surface A 1.40 5.80 ND ND 1.34 1.31

WFGB Surface B 1.40 1.90 ND ND ND ND

WFGB Midwater A 0.84 2.10 ND ND ND ND

WFGB Midwater B 1.12 10.90 ND 0.1 0.88 ND

WFGB Reef Cap A 1.54 4.4 ND ND ND ND

WFGB Reef Cap B 1.26 1.0 ND ND 2.52 ND

9.3.5. Hydrocarbons

In response to the Deepwater Horizon oil spill, total petroleum hydrocarbons (TPH) were 

added to the suite of analysis parameters for the May 2010 water quality sampling at the 

EFGB and WFGB. This measure was only added to this one sampling event as part of the 

oil spill response. Water collected at the surface, midwater, and at the reef cap were 

analyzed for hydrocarbons. Total petroleum hydrocarbon concentrations were 

undetectable (>5 mg/L) in all water samples at both the EFGB and WFGB. 

9.4. WATER QUALITY DISCUSSION

9.4.1. Water Quality Parameters

Water quality parameters investigated at the EFGB and WFGB from January 2009 

through December 2010 were temperature, salinity, chl a, and nutrients (ammonia, 

nitrate, nitrite, soluble reactive phosphorous, and TKN). The accuracy of the temperature 

118 
and salinity observations reported depended largely on the performance of the sensors 

during long-term oceanic deployments. 

9.4.1.1. Sea-Bird versus Hobo Temperature Records

HoboTemp thermographs were attached to each of the Sea-Bird 37-SMP MicroCAT 

datasondes as backup records of seawater temperature, and data were acquired at the both 

banks from January 2009 through December 2010. Concurrent Sea-Bird and Hobo data 

could therefore be compared to evaluate accuracy. Overall, both methods provided 

reliable temperature records. Statistical comparisons were conducted to compare daily 

averages from 2009 to 2010 using a two-sample t-test. No significant differences (Pvalue >0.05) were found. However, one or both of the sensors at the EFGB apparently 

began to provide inaccurate readings during July of 2010 (see July–December data in 

Figure 9.4.1). The WFGB Sea-Bird and Hobo temperature data had only slight 

differences at the WFGB in late July of 2010 (Figure 9.4.2). Both the Sea-Bird and Hobo 

Temp thermographs will continue to be used at the FGB to enhance long-term monitoring 

of temperature on the reef cap, as well as provide reliable backup records if one sensor 

fails.

16

18

20

22

24

26

28

30

32

1-Jan 1-Feb 1-Mar 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 1-Oct 1-Nov 1-Dec

Temperature (C)

Date

Comparison of Sea-Bird and Hobo Annual Temperature Variation Readings 

at the East Flower Garden Bank, 2009-2010

2009 Sea-Bird

2009 Hobo

2010 Sea-Bird

2010 Hobo

Figure 9.4.1. Comparison of Sea-Bird and Hobo daily average sea water temperature 

readings measured near the reef cap at the EFGB from 2009 to 2010. 

119 
16

18

20

22

24

26

28

30

32

1-Jan 1-Feb 1-Mar 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 1-Oct 1-Nov 1-Dec

Temperature (C)

Date

Comparison of Sea-Bird and Hobo Annual Temperature Variation Readings 

at the West Flower Garden Bank, 2009-2010

2009 Sea-Bird

2009 Hobo

2010 Sea-Bird

2010 Hobo

Figure 9.4.2. Comparison of Sea-Bird and Hobo daily average sea water temperature 

readings measured near the reef cap at the WFGB from 2009 to 2010. 

9.4.1.2. Temperature

Historical long-term averages derived from past monitoring periods show that the 

temperature minimum on the reef cap typically occurs from January to mid-March and

the temperature maximum from mid-August through mid-September (Figures 4.7.3 and 

4.7.4). The temperature on the EFGB reef cap was typically slightly warmer than that on 

the WFGB, especially during the summer months. The may be due to the WFGB 

proximity to the shelf edge is more likely to subject the reef cap to cooler water. The 

EFGB is slightly farther up on the continental shelf, perhaps subjecting it more regularly 

warmer water masses. The average temperature range on the EFGB reef cap from 1990 to 

2010 was 19.8 to 29.7°C. The average temperature range on the WFGB reef cap from 

1990 to 2010 was 19.5 to 29.3°C. 

Seasonal thermal changes over the reef caps of the EFGB and WFGB are very

apparent in the Figures 9.4.3 and 9.4.4. From a winter minimum, the temperature

gradually rises through the end of March and reaches a maximum during mid-August

through mid-September. The temperature decreases gradually starting around October

and reaches an annual minimum by mid-February through mid-March. During the 2009 

to 2010 monitoring period, there were several thermal anomalies (both positive and

negative), particularly in 2010. The most notable was the extreme warming that the 

western Atlantic region experienced during the summer of 2010. The prolonged sea

surface thermal anomaly began in the summer and continued well into the fall, causing

coral bleaching throughout the region. 

120 
Several thermal anomalies were recorded in 2009 and 2010 at the EFGB. When 

compared to historical means, the most dramatic was the very cold winter and spring 

temperatures experienced in March and April 2010. But there were also low summer 

temperatures in 2009, and a sudden decrease in temperature in late July 2010. The 

temperature over the reef cap was above average for several months during the summer 

and fall of 2010 (Figure 9.4.3). For a period of 27 days, temperature in summer 2010 

exceeded the coral bleaching threshold of 30°C. During that time, temperature exceeded 

30°C for 27 days. However, the monitoring cruise was completed that year before any 

visible bleaching. 

Several thermal anomalies were also recorded at the WFGB. Below average summer 

temperatures occurred in 2009, and like in the EFGB, a sudden decrease in temperature 

occurred in late July 2010 at the WFGB. Winter and spring temperatures were also 

abnormally low at the WFGB in 2010. The temperature over the reef cap was high during 

the summer and fall of 2010 compared with the long term average (Figure 9.4.4). The 

summer temperature in 2010 exceeded the coral bleaching threshold of 30°C at the 

WFGB, as well. During that time, temperature exceeded 30°C for 24 consecutive days. 

Coral bleaching was observed during the 2010 cruises in October and November at the 

WFGB and is described in sections 3.4 and 4.3 of this report. 

16

18

20

22

24

26

28

30

32

1-Jan 1-Mar 1-May 1-Jul 1-Sep 1-Nov

Temperature (C)

Date

Sea-Bird Annual Temperature Variation at the East Flower 

Garden Bank, 2009-2010

1990-2008 Mean Temp

2009

2010

Figure 9.4.3. Sea water temperature measured using Sea-Bird datasonde near 

the reef cap at the EFGB in 2009 and 2010, and the historical longterm average temperature from past monitoring periods. 

 

121 
16

18

20

22

24

26

28

30

32

1-Jan 1-Mar 1-May 1-Jul 1-Sep 1-Nov

Temperature (C)

Date

Sea-Bird Annual Temperature Variation at the West Flower 

Garden Bank, 2009-2010

1990-2008 Mean Temp

2009

2010

Figure 9.4.4. Sea water temperature measured using Sea-Bird datasonde near the 

reef cap at the WFGB in 2009 and 2010, and the historical long-term 

average temperature from past monitoring periods. 

9.4.1.3. Salinity

Accurate salinity data were obtained from the Sea-Bird MicroCAT for January 2009 to 

December 2010, ranging from approximately 31 to 38 PSU at the EFGB and WFGB with 

a mean of approximately 36 PSU. The data indicated lower salinity in summer months, a 

pattern consistent on both banks. During 2009, there were two annual events of low 

salinity on the reef caps of the FGB: one in May/June at the EFGB and another, more 

pronounced event from May through August at the WFGB. During 2010, there were

also two annual events of low salinity: one from May through August at the EFGB and 

another, more pronounced event from August through November at the WFGB. Although 

the data collected appear to be within the accepted limits of salinity for coral reefs located 

in the Western Atlantic (31–38 PSU; Coles and Jokiel 1992), the WFGB salinity data 

included one value below 31 PSU in June 2009. 

Future salinity data collected by the Sea-Bird MicroCAT conductivity recorder

should elucidate the occurrence and intensity of low salinity events on the reef cap of

the FGB. For now, independent measurements of salinity at and near the FGB point 

to the occurrence of substantial changes of salinity. The most probable source of low 

salinity water at the FGB is a nearshore river-seawater mix that reaches the outer 

continental shelf, eminating principally from the Mississippi and Atchafalaya River

watersheds, and subjecting the FGB occasionally to nearshore processes and to regional 

river runoff. 

122 
9.4.2. Chlorophyll a and Nutrients

Chl a concentrations in 2009 and 2010 revealed that the water column overlying the FGB 

reef caps could occasionally contain as much as 2.14 mg/m3. Not all water samples 

contained detectable levels of chl a (>1 mg/m3), and concentrations at the shelf edge in 

the northwestern Gulf of Mexico typically range from 0.1–0.3 mg/m3 (Nowlin et al.

1998). The highest values for surface chl a are typically expected in the summer (July–

August; Nowlin et al. 1998). The relatively high values of chl a (by FGB standards)

observed on 05/27/2010 may be indicative of an algal bloom, and coincided with a 

period with reduced salinity, suggesting the influence of nearshore water. There were

unfortunately no oceanographic satellite data available through NOAA’s CoastWatch 

Program to examine the occurrence of an algal bloom at the FGB in May 2009. The use 

of CoastWatch to monitor changes in chl a in the area of the FGB is certainly more 

useful than spot checks alone. The spot checks conducted here were valuable for ground 

truthing purposes and to examine vertical profiles over the reef cap. 

No definitive trends could be determined from the nutrient data. Ammonia values

were typically less than 1 mg/l from the sea surface to the reef cap, with the exception of 

samples taken on 07/07–08/2009 at the EFGB and WFGB. Nitrate levels were typically

very low (less than 0.15 mg/l) even when relatively high chlorophyll a was measured. 

TKN (organic nitrogen and ammonia) was detected in all water samples collected at the 

banks in 2009, with concentrations ranging from 0.84–13.9 mg/l (above the 0.10 mg/l

detection limit). Only about 10% of the samples from 2010 contained TKN 

concentrations above detectable limits. Soluble reactive phosphorous was not detected. 

After the Deepwater Horizon oil spill in 2010, TPH concentrations were undetectable in 

all water samples collected at the FGB, a result that is consistent with the measured 

trajectories of oil from the incident. 

The data gathered here could not elucidate annual patterns of nutrient concentrations 

at the FGB. Nowlin et al. (1998) showed that shelf edge waters in the northwestern

Gulf of Mexico are typically stripped of nutrients. When present, the probable sources 

of nutrients in the water column at the FGB are nearshore waters (Nowlin et al. 1998), 

sediments (Entsch et al. 1983), or benthic and planktonic organisms (D’Elia and Wiebe 

1990). More frequent sampling is required to understand nutrient dynamics over the reef

caps. 

9.4.3. Sea State

During the course of the 2009 to 2010 monitoring period, three tropical storms and two 

hurricanes occurred in the Gulf of Mexico. In the past, up to 11 tropical cyclones have 

affected the Gulf of Mexico in one year (Precht et al. 2006). In 2008, Hurricane Ike 

crossed over the EFGB (eventually made landfall in Galveston, Texas), damaging coral 

in the long-term monitoring study site. However, the 2009 and 2010 storms were located 

in other areas of the Gulf of Mexico and did not impact the FGBNMS. Hurricane Ida in 

2009 and Tropical Storm Bonnie in 2010 were the two cyclones that came closest to the 

FGB during the monitoring period (Figure 9.4.5). Table 9.4.1 lists all of the tropical 

cyclones that have occurred in the Gulf of Mexico since 2002. 

123 
Table 9.4.1.

List of Tropical Cyclones that Entered the Gulf of Mexico from 2002 to 2010 

Source data: NOAA, National Hurricane Center (2010). Cat=category.

Name Date

Wind

Speed 

(mph) or 

Category

Location and Distance

Tropical Storm Bertha 8/4/02 40 Northeast GOMEX, ~ 250 km north of the FGB

Tropical Storm Edouard 9/1/02 65 Northeast GOMEX, ~500 km east of the FGB

Tropical Storm Fay 9/7/02 60 Northwest GOMEX, ~100 km west of the FGB

Tropical Storm Hanna 9/12/02 50 Passed more than 400 km east of the FGB

Tropical Storm Bill 6/29/03 60 Passed within 200 km east of the FGB

Tropical Storm Grace 8/30/03 40 Passed within 100 km of the FGB

Tropical Storm Henri 9/3/03 50 Passed 700 km to the east of the FGB

Tropical Storm Larry 10/2/03 60 Took place in the southern GOMEX

Tropical Storm Bonnie 08/10/04 50

Central GOMEX, ~401 km (249 mi) east of the 

EFGB

Hurricane Charley 08/13/04 Cat 4

Florida Straits, ~1132 km (703 mi) from the 

EFGB

Tropical Storm Frances 09/04/04 65

Northeast GOMEX, ~924 km (574 mi) east of 

the EFGB

Hurricane Ivan 09/15/04 Cat 4

FGB Eastern GOMEX, ~168 km (104 mi) east 

of the EFGB

Tropical Storm Matthew 10/09/04 40

Northwest GOMEX, ~191 km (119 mi) east of 

the EFGB

Tropical Storm Arlene 06/11/05 69

GB Central GOMEX, ~635 km (395 mi) east of 

the EFGB

Tropical Storm Bret 06/25/05 40

Southwest GOMEX, ~805 km (500 mi) 

southwest of the WFGB

Tropical Storm Cindy 07/05/05 70

Central GOMEX, ~307 km (191 mi) east of the 

EFGB

Hurricane Dennis 07/10/05 Cat 4

Central GOMEX, ~686 km (426 mi) east of the 

EFGB

Hurricane Emily 07/19/05 Cat 1

Southwest GOMEX, ~437 km (272 mi) south 

of the WFGB

Tropical Storm Gert 07/25/05 45

Southwest GOMEX, ~924 km (574 mi) south 

of the WFGB

Tropical Storm Jose 08/23/05 50

Central GOMEX, ~394 km (245 mi) east of the 

EFGB

Hurricane Katrina 08/28/05 Cat 5

Central GOMEX, ~394 km (245 mi) east of the 

EFGB

Hurricane Rita 09/23/05 Cat 3

Central GOMEX, ~93 km (58 mi) east of the 

EFGB

Tropical Storm Stan 10/3/2005 40

Southwest GOMEX, ~862 km (535 mi) south 

of the EFGB

Hurricane Wilma 10/24/05 Cat 3

Southeast GOMEX, ~965 km (600 mi) 

southeast of the EFGB

Tropical Storm Alberto 06/12/06 69

Eastern GOMEX, ~659 km (409 mi) southeast 

of the EFGB

Tropical Storm Barry 06/02/07 58 Southeast GOMEX,~926 km (576 mi) 

124 
Name Date

Wind

Speed 

(mph) or 

Category

Location and Distance

southeast of the EFGB

Tropical Storm Erin 08/15/07 40

Western GOMEX, ~216 km (134 mi) east of 

the WFG

Hurricane Humberto 09/13/07 Cat 1

Northwest GOMEX, ~123 km (76 mi) west of 

the WFGB

Hurricane Lorenzo 09/25/07 Cat 1

Southwest GOMEX, 680 km (423 mi) 

southwest of the WFGB

Hurricane Dolly 07/22/08 Cat 1

South central GOMEX, ~360 km (224 mi) 

southeast of the WFGB

Tropical Storm Edouard 08/05/08 63

Central GOMEX, ~139 km (86 mi) northeast of 

the EFGB

Tropical Storm Fay 08/23/08 45

Eastern GOMEX, ~512 km (318 mi) east of the 

EFGB

Hurricane Gustav 09/01/08 Cat 3

East central GOMEX, ~300 km (186 mi) east 

of the EFGB

Hurricane Ike 09/13/08 Cat 2

Central GOMEX, ~0.7 km (0.4 mi) from the 

EFGB

Tropical Storm Marco 10/07/08 63

Southwest GOMEX, ~916 km (569 mi) south 

of the WFGB

Tropical Storm 

Claudette 8/16/2009 60

Eastern GOMEX, ~1,207 km (750 mi) 

southeast of the EFGB

Hurricane Ida

11/11/200

9 Cat 2

Central GOMEX, ~450 km (280 mi) northeast 

of the EFGB

Hurricane Alex 6/30/2010 Cat 2

Southwest GOMEX, ~482 km (300 mi) 

southwest of the WFGB

Tropical Storm Bonnie 7/24/2010 45

Eastern GOMEX, ~402 km (250 mi) southeast 

of the EFGB

Tropical Storm Hermine 9/5/2010 70

Southwest GOMEX, ~467 km (290 mi) 

southwest of the WFGB

125 
Figure 9.4.5. Hurricane Ida (A) and Tropical Storm Bonnie (B) 

storm track areas (NOAA Hurricane Center).

 (A)

 (B)

126 
CHAPTER 10.0: FISH SURVEYS

10.1. FISH SURVEYS METHODOLOGICAL RATIONALE

Surveys of fish assemblages have been conducted at the FGBNMS since the early 1980s; 

however, these surveys have not generally been part of the long-term monitoring study

(Boland et al. 1983; Rezak et al. 1985; Dennis and Bright 1988; Pattengill 1998). Fish 

surveys were officially added to the long-term monitoring protocol in 2002.The fish 

assemblages of the coral reef zone at the EFGB and WFGB are primarily composed of 

Caribbean reef species; however, the total number of species is lower in comparison. 

Certain families such as the snappers (Lutjanidae) and grunts (Haemulidae) are 

underrepresented at the FGB mainly due to lack of diverse and nearby seagrass and 

mangrove habitats (Jones and Clark 1981; Lukens 1981; Rezak et al. 1985; Mumby et al. 

2004). The influence of nearby offshore gas and petroleum production platforms on fish 

assemblages at the FGBNMS has been under continuous investigation (Rooker et al. 1997).

Therefore, continued monitoring of the FGBNMS is vital to increasing our understanding of 

this unique habitat in light of the ongoing, as well as the changing, natural and 

anthropogenic pressures on fish populations. 

10.2. FISH SURVEYS FIELD METHODS

Twenty-four stationary visual fish surveys were conducted at each bank in 2009. In 2010, 

25 surveys were conducted on the reef cap at the EFGB, and 26 were conducted at the 

WFGB (a minimum of six in each quadrant of the study sites). Fishes were visually 

assessed using SCUBA and a stationary visual census technique (Bohnsack and Bannerot 

1986). Observations of fishes were restricted to an imaginary cylinder with a radius 7.5 

m (24.6 ft) from the diver, extending to the surface. All fish species observed within the 

first five minutes of the survey were recorded as the diver slowly rotated in place. 

Immediately following this five-minute observation period, one rotation was conducted 

for each species noted in the original five-minute period to record abundance (number of 

individuals per species) and total length (within eight categories). Transitory or schooling 

species were counted and measured at the time the individuals moved through the 

cylinder during the initial five-minute period. After the initial five-minute period, no 

additional species were added. Each survey required 10 to 15 minutes. 

For each quadrant, a random number was generated between 0 and 50 to indicate the 

starting location along the first boundary line encountered (e.g., if the random number is 

27 and the corner of the transect tape is 0, the transect is started at 27 m along the transect 

tape, and if the lowest corner of the transect tape is 50, the transect is started at 77 m 

along the transect tape). A second random number was generated between 0 and 40 to 

determine the number of fin kicks perpendicular from the boundary line into the study 

site. It is assumed that it takes 40 fin kicks to swim 50 m if there is not a strong current. 

Those criteria provided the location for the first fish count. Subsequent survey starting 

points were determined with a second set of randomly generated numbers with the first 

number providing a heading, between 0o and 360o

, and the second providing the number 

127 
of fin kicks, between 12 and 40, to ensure the starting point was at least 15 m away from 

the previous location. A third number was generated to provide a random heading, 

between 0o

 and 360o, in which to lay the tape marking the 7.5 m radius of the survey. 

Fish survey dives began in the early morning (after 0700), and were repeated by two to 

three divers throughout the day until dusk. Survey locations were spread randomly within 

the 100 x 100 m study site. Survey depths ranged from 20–24 m (65–80 ft). 

10.3. FISH SURVEY DATA AND STATISTICAL ANALYSIS

Fish densities are expressed as the number of fish per 100 m2. For each bank and year, 

densities were calculated by dividing the mean number of individuals per survey by the 

horizontal area of the survey cylinder (176.7 m2). This value was then multiplied by

100 to provide fish densities per 100 m2.

Percent sighting frequency for each species is expressed as the percentage of the total 

number of times the species was recorded out of the total number of surveys for the 

site (bank and year). Species richness is the total number of species for each site (bank

and year). Relative abundance is expressed as the number of individuals of one species 

divided by the total number of all species observed. 

Size-frequency distributions for four trophic guilds (herbivores, piscivores, invertivores, 

and planktivores) were calculated for each bank and year by dividing the number of 

herbivores, piscivores, invertivores, and planktivores in each size class by the total

number observed in each trophic guild. Size classes included <5 cm, 5–10 cm, 10–15 

cm, 15–20 cm, 20–25 cm, 25–30 cm, 30–35 cm and >35 cm. The herbivore guild is 

comprised of Acanthuridae, Blennidae, Kyphosidae, Scaridae, and members of the 

Balistidae (Black Durgon), Gobiidae (Goldspot Goby), Pomacanthidae (Queen 

Angelfish), and Pomacentridae (Cocoa, Bi-Color, Yellowtail, and Dusky Damselfish) 

families. The piscivore guild is comprised of Aulostomidae, Carangidae, Inermiidae, 

Muraenidae, Sphyraenidae, and members of the Lutjanidae (Dog Snapper), and 

Serranidae (Yellowmouth, Tiger, Black, and Yellowfin Groupers, Graysby, and Scamp) 

families. The invertivore guild is comprised of Apogonidae, Chaetodontidae, Cirrhitidae, 

Dasyatidae, Diodontidae, Holocentridae, Monacanthidae, Mullidae, Muraenidae, 

Ostraciidae, Tetraodontidae, and members of the Balistidae (Ocean and Queen 

Triggerfish), Gobiidae (Neon Goby), Labridae (Spanish and Spotfin Hogfish, Clown, 

Yellowhead, Slippery Dick, and Bluehead Wrasse, and Pudding Wife), Lutjanidae (Grey 

Snapper), and Pomacanthidae (Rock Beauty and French Angelfish), Pomacentridae

(Brown Chromis, Sergeant Major, and Three-Spot Damselfish), and Serranidae (Rock 

and Red Hinds, and Coney) families. The planktivore guild is comprised of Echeneidae, 

and members of the Labridae (Creole Wrasse), Pomacentridae (Blue Chromis, Purple 

Reeffish, and Sunshinefish), and Serranidae (Creolefish) families.

128 
Diversity was calculated using the Shannon-Wiener diversity index, H':

 k

H '= −∑ pi log pi

i=1 

where k is the number of species present and pi is the relative abundance of each

species, calculated as the proportion of individuals of a given species to the total

number of individuals observed. Species evenness (J') was determined for each site

and year using the following calculation: 

J '= H '

 H 'max

where H'max was the maximum possible diversity (H'max = log k).

To allow the valid application of parametric analyses of variance, fish abundances were 

log10+1 transformed to make them normal, homoscedastic, and additive (Zar 1984; 

Aronson et al. 1994; Edmunds and Carpenter 2001). Two-sample t-tests (two-tailed) 

were used to compare densities and species-richness values by bank and year. In 

addition, observations of manta rays were removed from all statistical and biomass 

analyses due to their rare nature.

Percent sighting frequency for each species was expressed as the percentage of the 

total number of times the species was recorded out of the total number of surveys 

for the site (bank). Species richness is the number of species recorded at each site

(bank). Fish abundances were log10+1 transformed as mentioned above and twosample t-tests (two-tailed) were used to compare the fish abundances and species

richness by bank. Size frequency distributions for the herbivores, piscivores, 

invertivores, and planktivores were as described in the previous section. 

Fish biomass, an important aspect of coral reef ecology, was computed by converting 

length data to weights using the allometric length-weight conversion formula:

W = α*Lβ

where W = individual weight (grams), L = length of fish (cm), and α and β are constants 

for each species generated from the regression of its length and weight, derived from

FishBase (2009) and Bohnsack and Harper (1988). Because lengths for every

individual fish were not recorded, mean lengths for each species size categories were

used. A species-biomass per unit area estimate (g/m2) was calculated by dividing the 

mean biomass for a species across all surveys by the area of a diver survey (176.7 m2). 

Coupling both biomass and abundance was useful in assessing the fish communities at 

the EFGB and WFGB.

129 
All comparisons of density or biomass were conducted using parametric models on 

log(n+1) transformed means (either ANOVAs or two-tailed t-tests, unless otherwise 

stated). Species prevalence was determined by calculating the presence/absence of 

each species in each survey and summing across surveys. Tests of differences in

prevalence of species between banks were performed using proportions tests based on 

binomial distributions. All tests were conducted using JMP9® statistical software 

package. 

The NOAA National Centers for Coastal Ocean Science (NCCOS) BioGeography 

Branch conducted annual fish and benthic community structure surveys at EFGB and 

WFGB in 2006, 2007, 2009, and 2010. These surveys are conducted randomly on the 

entire coral cap in high relief and low relief habitats, and couple fish abundance data with 

benthic communities. Fish species abundance, size and distribution were characterized 

using 75 belt transect surveys, showing that density and biomass and species richness 

were higher at WFGB than EFGB. Additional information can be found at the NCCOS 

website (http://ccma.nos.noaa.gov/ecosystems/sanctuaries/ fgb_nms). 

10.4. FISH SURVEYS RESULTS

Fish surveys were conducted at both banks during 2009 and 2010. In 2009, fish surveys 

were conducted in August at both the EFGB and WFGB. In 2010, fish surveys were 

conducted in August at the EFGB. For the WFGB, several surveys were completed in 

August, but due to inclement weather, the remaining surveys were collected in October. 

Table 10.4.1 shows a complete list of species and their abundance per year per bank. 

Table 10.4.1. 

Complete Fish Species List, Rrophic Guilds, and Counts per Year per Bank

 

P=Piscivore, I=Invertivore, PL=Planktivore, and H=Herbivore

EFGB WFGB

Species Name Trophic 

Guild 2009 2010 2009 2010 

Lutjanidae: Lutjanus jocu (Snapper, Dog) P 8 0 0 5

Pomacentridae: Chromis multilineata (Chromis, Brown) I 2532 1451 1815

151

2

Serranidae: Paranthias furcifer (Creolefish) PL 923 760 570

117

7

Labridae: Thalassoma lucasanum (Wrasse, Bluehead) I 748 283 566 283

Pomacentridae: Stegastes planifrons (Damselfish, Threespot) I 141 71 157 55

Pomacentridae: Stegastes variabilis (Damselfish, Cocoa) H 29 69 40 83

Serranidae: Mycteroperca interstitialis (Grouper, 

Yellowmouth) P 12 6 4 6

Kyphosidae: Kyphosus incisor / Kyphosus sectatrix 

(Chub) H 354 64 18 49

Scaridae: Scarus vetula (Parrotfish, Queen) H 54 71 56 68

Cirrhitidae: Amblycirrhitus pinos (Hawkfish, 

Redspotted) I 3 1 3 3

130 
EFGB WFGB

Species Name Trophic 

Guild 2009 2010 2009 2010 

Pomacentridae: Stegastes partitus (Damselfish, Bi-color) H 364 132 235 82

Acanthuridae: Acanthurus coeruleus (Tang, Blue) H 120 83 140 64

Atherinopsidae: Menidiinae sp. (Silversides) H 910 0 0 0

Pomacentridae: Chromis cyanea (Chromis, Blue) PL 525 40 449 134

Tetraodontidae: Canthigaster rostrata (Puffer, 

Sharpnose) I 106 79 125 91

Labridae: Clepticus parrae (Wrasse, Creole) PL 1710 291 542 361

Acanthuridae: Acanthurus chirurgus (Doctorfish) H 11 3 11 3

Pomacanthidae: Holacanthus ciliaris (Angelfish, Queen) H 3 0 5 2

Labridae: Bodianus rufus (Hogfish, Spanish) I 48 63 85 90

Blenniidae: Ophioblennius macclurei (Blenny, Redlip) H 27 0 39 4

Chaetodontidae: Chaetodon ocellatus (Butterflyfish, 

Spotfin) I 9 0 8 8

Pomacentridae: Chromis scotti (Reeffish, Purple) PL 139 23 240 81

Scaridae: Sparisoma aurofrenatum (Parrotfish, Redband) H 53 25 10 13

Serranidae: Mycteroperca tigris (Grouper, Tiger) P 4 7 0 11

Pomacentridae: Microspathodon chrysurus (Damselfish, 

Yellowtail) H 16 19 33 7

Pomacanthidae: Holacanthus tricolor (Angelfish, Rock 

Beauty) I 19 5 11 1

Chaetodontidae: Chaetodon sedentarius (Butterflyfish,

Reef) I 17 29 23 36

Monacanthidae: Cantherhines pullus (Filefish, Orange 

Spotted) I 6 1 4 0

Pomacentridae: Chromis insolata (Sunshinefish) PL 104 7 42 15

Scaridae: Sparisoma viride (Parrotfish, Stoplight) H 23 38 51 30

Carangidae: Caranx latus (Jack, Horse-eye) P 83 0 17 5

Carangidae: Caranx lugubris (Jack, Black) P 5 5 10 0

Balistidae: Canthidermis sufflamen (Triggerfish, Ocean) I 4 2 3 0

Serranidae: Cephalopholis cruentata (Graysby) P 12 17 15 19

Ostraciidae: Lactophrys triqueter (Trunkfish, Smooth) I 5 10 9 5

Labridae: Halichoeres maculipinna (Wrasse, Clown) I 14 0 49 3

Acanthuridae: Acanthurus bahianus (Surgeonfish, 

Ocean) H 9 24 6 15

Balistidae: Melichthys niger (Durgon, Black) H 12 23 39 30

Pomacentridae: Abudefduf saxatilis (Sergeant Major) I 9 5 27 15

Gobiidae: Elacatinus oceanops (Goby, Neon) I 3 11 2 2

Sphyraenidae: Sphyraena barracuda (Barracuda, Great) P 6 47 18 75

Mobulidae: Manta birostris (Ray, Manta) PL 1 1 0 0

Serranidae: Mycteroperca bonaci (Grouper, Black) P 1 0 0 1

131 
EFGB WFGB

Species Name Trophic 

Guild 2009 2010 2009 2010 

Pomacanthidae: Pomacanthus paru (Angelfish, French) I 8 7 1 1

Scaridae: Scarus taeniopterus (Parrotfish, Princess) H 5 34 1 17

Serranidae: Epinephelus adscensionis (Hind, Rock) I 2 2 0 0

Serranidae: Epinephelus guttatus (Hind, Red) I 1 0 0 0

Serranidae: Mycteroperca phenax (Scamp) P 2 1 1 0

Labridae: Halichoeres garnoti (Wrasse, Yellowhead) I 8 4 18 19

Mullidae: Pseudupeneus maculatus (Goatfish, Spotted) I 2 0 0 0

Inermiidae: Emmelichthyops atlanticus (Bonnetmouth) P 2510 0 12 0

Ostraciidae: Acanthostracion polygonius (Cowfish, 

Honeycomb) I 1 1 4 0

Echeneidae: Echeneidae sp. (Remora) PL 2 0 0 0

Chaetodontidae: Prognathodes aculeatus (Butterflyfish, 

Longsnout) I 2 4 11 7

Pomacentridae: Stegastes adustus (Damselfish, Dusky) H 5 7 16 4

Scaridae: Sparisoma atomarium (Parrotfish, 

Greenblotch) H 2 0 2 0

Carangidae: Carangoides ruber (Jack, Bar) P 41 16 6 31

Labridae: Bodianus pulchellus (Hogfish, Spotfin) I 1 0 0 0

Ostraciidae: Lactophrys bicaudalis (Trunkfish, Spotted) I 1 0 3 0

Carangidae : Caranx crysos (Blue runner) P 4 0 2 0

Muraenidae: Gymnothorax meleagris (Moray, Spotted) P 1 0 1 1

Lutjanidae: Lutjanus agennes (Snapper, Grey) I 0 5 11 5

Apogonidae: Apogon sp. (Cardinal fish sp.) I 0 0 1 0

Aulostomidae: Aulostomus maculatus (Trumpetfish) P 0 0 2 0

Holocentridae: Holocentrus rufus (Squirrelfish, 

Longspine) I 0 1 4 3

Carangidae: Elagatis bipinnulata (Rainbow runner) P 0 0 100 0

Diodontidae: Diodon holocanthus (Ballonfish) I 0 0 1 1

Monacanthidae: Cantherhines macrocerus (Filefish, 

Whitespotted) I 0 1 2 0

Mullidae: Mulloidichthys martinicus (Goatfish, Yellow) I 0 3 9 0

Gobiidae: Gnatholepis thompsoni (Goby, Goldspot) H 0 0 1 0

Muraenidae: Gymnothorax miliaris (Moray, Goldentail) I 0 0 1 0

Dasyatidae: Dasyatis americana (Stingray, Southern) I 0 0 1 0

Labridae: Halichoeres radiatus (Pudding wife) I 0 2 2 0

Labridae: Halichoeres bivittatus (Wrasse, Slippery Dick) I 0 0 0 11

Scaridae: Scarus iseri (Parrotfish, Striped) H 0 5 0 2

Tetraodontidae: Sphoeroides spengleri (Puffer, Bandtail) I 0 0 0 0

Chaetodontidae: Chaetodon striatus (Butterflyfish, 

Banded) I 0 1 0 1

132 
EFGB WFGB

Species Name Trophic 

Guild 2009 2010 2009 2010 

Balistidae: Balistes vetula (Triggerfish, Queen) I 0 2 0 0

Serranidae: Liopropoma rubre (Bass, Peppermint) I 0 1 0 0

Lutjanidae: Lutjanus argentiventris (Snapper, 

Yellowtail) PL 0 0 0 0

Holocentridae: Holocentrus adscensionis (Squirrelfish) I 0 0 0 0

Serranidae: Cephalopholis fulva (Grouper, Coney) I 0 0 0 0

Serranidae: Mycteroperca venenosa (Grouper, 

Yellowfin) P 0 0 0 0

During the 2009 and 2010 monitoring period, 99 Bohnsack and Bannerot visual fish 

census surveys were conducted at each bank. The highest number conducted during a 

single sampling effort was 26, at the WFGB in 2010. The smallest number was 24, at 

both banks in 2009. Each survey represents one sample. The long-term monitoring at the 

EFGB and WFGB SOW requires a minimum of 24 samples to be collected at each Bank 

per year. Each sample covered 176.7 m2

, resulting in an coverage of just under 44% of 

the 100 m x100 m study sites during 2009 and 2010 (Table 10.4.2). 

Table 10.4.2. 

Summary of Fish Surveys at the EFGB and WFGB in 2009 and 2010

EFGB 

2009

EFGB 

2010

WFGB 

2009

WFGB 

2010

Number Samples (n) 24 25 24 26

% area of study site sampled 42 44 42 46

Fish cylinder samples area 

(m2) 176.7 176.7 176.7 176.7

Area sampled (m2) 4240.8 4417.5 4240.8 4594.2

Total Fish Abundance 12188 3863 5690 4547

An average of 11.71 ± 1.78 fish families per survey (sample) were recorded at the WFGB 

in 2009, while an average of 9.42 ± 1.86 per survey were recorded in 2010 (See Table 

10.4.3). A mean of 23 ± 3.56 fish families were observed per year and bank. 

Mean species richness on the banks (number of species recorded) was 57 ± 5.89 based on 

samples collected in 2009 and 2010. A total of 78 species were recorded in all surveys 

combined. The highest species richness per survey was recorded at the WFGB in 2009 

(22.08 ± 2.00), and the lowest was at the WFGB in 2010 (17.42 ± 4.04) (See Table

10.4.3). When species richness was compared using a two-way ANOVA, a significant 

difference was observed between years (α=0.005, P-value<0.0001), but not between 

banks. However, a significant interaction exists between the two factors (α=0.005, Pvalue=0.0209). Analysis with Students t-test showed a significant difference in species 

richness between the EFGB and WFGB in 2009 (α=0.005, P-value=0.0088), but not in 

133 
2010. A significant difference was also observed between 2009 and 2010 at both the 

EFGB (α=0.005, P-value=0.0425) and WFGB (α=0.005, P-value= <.0001). 

The highest average fish density (number per 100 m

2

) was observed at the EFGB in 2009 

(490.79 ± 397.02), and the lowest average was at the EFGB in 2010 (154.52 ± 54.68) 

(Table 10.4.3). The high fish density value at EFGB in 2009 was primarily due to an 

abundance of Bonnetmouth (Emmelichthyops atlanticus), Silversides (Menidiinae sp.), 

Creole Wrasse (Clepticus parrae), and Brown Chromis (Chromis multilineata) (mean 

densities of 59.19, 21.46, 40.32, and 59.71 per 100 m

2 respectively) (Table 10.4.3). 

Variations in species abundance between years and banks are predominantly attributed to 

the presence or absence of dense schooling fish species, such as Bonnetmouth 

(Emmelichthyops atlanticus) and Silversides (Menidiinae sp.), that typically consist of 

100’s of individuals. When a two-way ANOVA is run of the log (n+1) transformed fish 

density data, a significant difference is seen between years (α=0.005, P-value<0.0001), 

and between banks (α=0.005, P-value=0.0294) and a significant interaction exists 

between the two factors (α=0.005, P-value=0.0.0147). In a Students t-test, a significant 

difference in fish density was observed between the EFGB and WFGB in 2009 (α=0.005, 

P-value=0.0047), but not in 2010. Both the EFGB and WFGB showed a significant 

difference in fish densities between 2009 and 2010 (α=0.005, P-value<0.0001 and Pvalue=0.0088 respectively). 

Table 10.4.3. 

Species and Family Richness and Fish Density for the EFGB and WFGB in 2009 and 2010 

Values are expressed as richness or abundance ± SD

EFGB 2009 EFGB 2010 WFGB 2009 WFGB 2010

Species Richness 61 53 63 51

Family Richness 25 20 27 20

Total Fish Abundance 12188 3863 5690 4547

Mean Abundance/Survey

490.79 

(±397.02 )

154.52 

(±54.68 )

237.08 

(±84.47)

174.88 

(±101.81)

Mean Abundance/100m2

(Density) 275.9 87.4 134.2 99.0

Mean Species 

Richness/Survey 19.96 (±3.34) 18.24 (±2.47) 22.08 (±2.00) 17.42 (±4.04)

Mean Species Richness/m2 0.11 0.10 0.12 0.10

Mean Family Richness 10.46 (±1.77) 9.92 (±1.47) 11.71 (±1.78) 9.42 (±1.86)

Mean Family Richness/m2 0.06 0.06 0.07 0.05

Fish abundance varied between years and banks. Brown Chromis (Chromis multilineata) 

were the most abundant fish at both banks for both years. Other high-ranked species 

included Bonnetmouth (Emmelichthyops atlanticus), Creole Wrasse (Clepticus parrae), 

and Silversides (Menidiinae sp.) (Table 10.4.4).

134 
Table 10.4.4. 

Mean Fish Abundance for the EFGB and WFGB in 2009 and 2010 for Dominant Species

Mean Abundance/100 m2 (Density)

Species EFGB 2009 EFGB 2010 WFGB 2009 WFGB 2010

Bonnetmouth 59.19 0.00 0.28 0.00

Silversides 21.46 0.00 0.00 0.00

Creole Wrasse 40.32 6.59 12.78 7.86

Brown Chromis 59.71 32.85 42.80 32.91

10.4.1. Sighting Frequency and Occurrence

For the various trophic guilds at the FGB (Piscivore, Invertivore, Plankivore, and 

Herbivore), the percent sighting frequency (the percentage of times in which the species 

was observed) varied between year and Bank. The most frequently sighted species was 

Creolefish (Paranthias furcifer), sighted in 95% of surveys from 2009 to 2010 at both 

banks. Other top-ranked species included Bluehead Wrasse (Thalassoma lucasanum), Bicolor Damselfish (Stegastes partitus), Blue Chromis (Chromis cyanea), Spanish Hogfish 

(Bodianus rufus), Creole Wrasse (Clepticus parrae), Brown Chromis (Chromis 

multilineata), Blue Tang (Acanthurus coeruleus), and Sharpnose Puffer (Canthigaster 

rostrata) with sighting frequencies ranging from 74% to 94% (Table 10.4.5) 

Table 10.4.5. 

Percent Sighting Frequency for Species by Year and Bank, Including a Percent Sighting 

Frequency for All Surveys Conducted during the 2009 to 2010 Reporting Period

P=Piscivore, I=Invertivore, PL=Plankivore, and H=Herbivore

Species Trophic 

Guild

EFGB WFGB % of 

Total 

Surveys 

Sighted 2009 2010 2009 2010

Lutjanidae: Lutjanus jocu (Snapper, 

Dog) P 20.8 0.0 0.0 15.4 9.1

Pomacentridae: Chromis multilineata

(Chromis, Brown) I 91.7 92.0 87.5 88.5 89.9

Serranidae: Paranthias furcifer

(Creolefish) PL 95.8 96.0 100.0 88.5 94.9

Labridae: Thalassoma lucasanum

(Wrasse, Bluehead) I 95.8 96.0 95.8 88.5 93.9

Pomacentridae: Stegastes planifrons

(Damselfish, Three-spot) I 70.8 60.0 87.5 50.0 66.7

Pomacentridae: Stegastes variabilis

(Damselfish, Cocoa) H 29.2 60.0 45.8 69.2 51.5

135 
Species Trophic 

Guild

EFGB WFGB % of 

Total 

Surveys 

Sighted 2009 2010 2009 2010

Serranidae: Mycteroperca 

interstitialis (Grouper, Yellowmouth) P 37.5 24.0 16.7 19.2 24.2

Kyphosidae: Kyphosus incisor / 

Kyphosus sectatrix (Chub) H 70.8 48.0 41.7 42.3 50.5

Scaridae: Scarus vetula (Parrotfish, 

Queen) H 70.8 72.0 83.3 76.9 75.8

Cirrhitidae: Amblycirrhitus pinos

(Hawkfish, Redspotted) I 12.5 4.0 12.5 11.5 10.1

Pomacentridae: Stegastes partitus

(Damselfish, Bi-color) H 95.8 96.0 95.8 73.1 89.9

Acanthuridae: Acanthurus coeruleus

(Tang, Blue) H 87.5 92.0 87.5 92.3 89.9

Atherinopsidae: Menidiinae sp. 

(Silversides) H 8.3 0.0 0.0 0.0 2.0

Pomacentridae: Chromis cyanea

(Chromis, Blue) PL 95.8 60.0 91.7 69.2 78.8

Tetraodontidae: Canthigaster 

rostrata (Puffer, Sharpnose) I 87.5 96.0 87.5 80.8 87.9

Labridae: Clepticus parrae (Wrasse, 

Creole) PL 95.8 76.0 87.5 38.5 73.7

Acanthuridae: Acanthurus chirurgus

(Doctorfish) H 25.0 8.0 12.5 7.7 13.1

Pomacanthidae: Holacanthus ciliaris 

(Angelfish, Queen) H 12.5 0.0 16.7 3.8 8.1

Labridae: Bodianus rufus (Hogfish, 

Spanish) I 70.8 80.0 91.7 96.2 84.8

Blenniidae: Ophioblennius macclurei

(Blenny, Redlip) H 37.5 0.0 54.2 11.5 25.3

Chaetodontidae: Chaetodon ocellatus

(Butterflyfish, Spotfin) I 20.8 0.0 16.7 19.2 14.1

Pomacentridae: Chromis scotti

(Reeffish, Purple) PL 41.7 16.0 70.8 61.5 47.5

Scaridae: Sparisoma aurofrenatum 

(Parrotfish, Redband) H 45.8 44.0 25.0 23.1 34.3

Serranidae: Mycteroperca tigris 

(Grouper, Tiger) P 16.7 24.0 0.0 30.8 18.2

Pomacentridae: Microspathodon 

chrysurus (Damselfish, Yellowtail) H 50.0 36.0 62.5 19.2 41.4

Pomacanthidae: Holacanthus tricolor

(Angelfish, Rock Beauty) I 41.7 20.0 37.5 3.8 25.3

Chaetodontidae: Chaetodon 

sedentarius (Butterflyfish, Reef) I 41.7 48.0 54.2 57.7 50.5

Monacanthidae: Cantherhines pullus

(Filefish, Orange Spotted) I 20.8 4.0 16.7 0.0 10.1

Pomacentridae: Chromis insolata

(Sunshinefish) PL 41.7 16.0 54.2 19.2 32.3

Scaridae: Sparisoma viride

(Parrotfish, Stoplight) H 50.0 68.0 79.2 38.5 58.6

Carangidae: Caranx latus (Jack, 

Horse-eye) P 29.2 0.0 16.7 11.5 14.1

136 
Species Trophic 

Guild

EFGB WFGB % of 

Total 

Surveys 

Sighted 2009 2010 2009 2010

Carangidae: Caranx lugubris (Jack, 

Black) P 16.7 8.0 29.2 0.0 13.1

Balistidae: Canthidermis sufflamen

(Triggerfish, Ocean) I 16.7 4.0 8.3 0.0 7.1

Serranidae: Cephalopholis cruentata

(Graysby) P 41.7 48.0 45.8 50.0 46.5

Ostraciidae: Lactophrys triqueter 

(Trunkfish, Smooth) I 16.7 36.0 29.2 15.4 24.2

Labridae: Halichoeres maculipinna

(Wrasse, Clown) I 20.8 0.0 33.3 7.7 15.2

Acanthuridae: Acanthurus bahianus

(Surgeonfish, Ocean) H 25.0 40.0 16.7 26.9 27.3

Balistidae: Melichthys niger

(Durgon, Black) H 29.2 60.0 70.8 53.8 53.5

Pomacentridae: Abudefduf saxatilis

(Sergeant Major) I 16.7 8.0 33.3 26.9 21.2

Gobiidae: Elacatinus oceanops

(Goby, Neon) I 8.3 24.0 8.3 3.8 11.1

Sphyraenidae: Sphyraena barracuda 

(Barracuda, Great) P 25.0 68.0 66.7 84.6 61.6

Mobulidae: Manta birostris (Ray, 

Manta) PL 8.3 4.0 0.0 0.0 3.0

Serranidae: Mycteroperca bonaci 

(Grouper, Black) P 4.2 0.0 0.0 3.8 2.0

Pomacanthidae: Pomacanthus paru

(Angelfish, French) I 16.7 16.0 4.2 3.8 10.1

Scaridae: Scarus taeniopterus 

(Parrotfish, Princess) H 12.5 40.0 4.2 23.1 20.2

Serranidae: Epinephelus adscensionis

(Hind, Rock) I 8.3 8.0 0.0 0.0 4.0

Serranidae: Epinephelus guttatus 

(Hind, Red) I 4.2 0.0 0.0 0.0 1.0

Serranidae: Mycteroperca phenax

(Scamp) P 8.3 4.0 4.2 0.0 4.0

Labridae: Halichoeres garnoti

(Wrasse, Yellowhead) I 12.5 8.0 29.2 30.8 20.2

Mullidae: Pseudupeneus maculatus

(Goatfish, Spotted) I 4.2 0.0 0.0 0.0 1.0

Inermiidae: Emmelichthyops 

atlanticus (Bonnetmouth) P 33.3 0.0 4.2 0.0 9.1

Ostraciidae: Acanthostracion 

polygonius (Cowfish, Honeycomb) I 4.2 4.0 16.7 0.0 6.1

Echeneidae: Echeneidae sp. 

(Remora) PL 4.2 0.0 0.0 0.0 1.0

Chaetodontidae: Prognathodes 

aculeatus (Butterflyfish, Longsnout) I 8.3 12.0 25.0 23.1 17.2

Pomacentridae: Stegastes adustus

(Damselfish, Dusky) H 8.3 12.0 33.3 7.7 15.2

Scaridae: Sparisoma atomarium

(Parrotfish, Greenblotch) H 4.2 0.0 4.2 0.0 2.0

137 
Species Trophic 

Guild

EFGB WFGB % of 

Total 

Surveys 

Sighted 2009 2010 2009 2010

Carangidae: Carangoides ruber 

(Jack, Bar) P 12.5 24.0 8.3 26.9 18.2

Labridae: Bodianus pulchellus

(Hogfish, Spotfin) I 4.2 0.0 0.0 0.0 1.0

Ostraciidae: Lactophrys bicaudalis 

(Trunkfish, Spotted) I 4.2 0.0 8.3 0.0 3.0

Carangidae: Caranx crysos (Blue 

runner) P 4.2 0.0 4.2 0.0 2.0

Muraenidae: Gymnothorax meleagris 

(Moray, Spotted) P 4.2 0.0 4.2 3.8 3.0

Lutjanidae: Lutjanus agennes

(Snapper, Grey) I 0.0 16.0 16.7 11.5 11.1

Apogonidae: Apogon sp. (Cardinal 

fish sp.) I 0.0 0.0 4.2 0.0 1.0

Aulostomidae: Aulostomus 

maculatus (Trumpetfish) P 0.0 0.0 4.2 0.0 1.0

Holocentridae: Holocentrus rufus 

(Squirrelfish, Longspine) I 0.0 4.0 16.7 7.7 7.1

Carangidae: Elagatis bipinnulata 

(Rainbow runner) P 0.0 0.0 4.2 0.0 1.0

Diodontidae: Diodon holocanthus

(Ballonfish) I 0.0 0.0 4.2 3.8 2.0

Monacanthidae: Cantherhines 

macrocerus (Filefish, Whitespotted) I 0.0 4.0 4.2 0.0 2.0

Mullidae: Mulloidichthys martinicus 

(Goatfish, Yellow) I 0.0 8.0 12.5 0.0 5.1

Gobiidae: Gnatholepis thompsoni 

(Goby, Goldspot) H 0.0 0.0 4.2 0.0 1.0

Muraenidae: Gymnothorax miliaris

(Moray, Goldentail) I 0.0 0.0 4.2 0.0 1.0

Dasyatidae: Dasyatis americana

(Stingray, Southern) I 0.0 0.0 4.2 0.0 1.0

Labridae: Halichoeres radiatus 

(Pudding wife) I 0.0 8.0 8.3 0.0 4.0

Labridae: Halichoeres bivittatus

(Wrasse, Slippery Dick) I 0.0 0.0 0.0 11.5 3.0

Scaridae: Scarus iseri (Parrotfish,

Striped) H 0.0 8.0 0.0 3.8 3.0

Tetraodontidae: Sphoeroides 

spengleri (Puffer, Bandtail) I 0.0 4.0 0.0 3.8 2.0

Chaetodontidae: Chaetodon striatus

(Butterflyfish, Banded) I 0.0 4.0 0.0 0.0 1.0

Balistidae: Balistes vetula

(Triggerfish, Queen) I 0.0 4.0 0.0 0.0 1.0

Serranidae: Liopropoma rubre (Bass, 

Peppermint) I 0.0 1.0 0.0 0.0 1.0

Lutjanidae: Lutjanus argentiventris

(Snapper, Yellowtail) PL 0.0 0.0 0.0 0.0 0.0

Holocentridae: Holocentrus 

adscensionis (Squirrelfish) I 0.0 0.0 0.0 0.0 0.0

138 
Species Trophic 

Guild

EFGB WFGB % of 

Total 

Surveys 

Sighted 2009 2010 2009 2010

Serranidae: Cephalopholis fulva

(Grouper, Coney) I 0.0 0.0 0.0 0.0 0.0

Serranidae: Mycteroperca venenosa

(Grouper, Yellowfin) P 0.0 0.0 0.0 0.0 0.0

Species occurrence in samples was compared between banks in both 2009 and 2010 for 

all observed species using a Student’s t-test (Table 10.4.6). In 2009, of 73 species 

observed, a significant difference was observed between the banks for 11 species. This 

included Dog Snapper (Lutjanus jocu, α=0.005, P-value=0.0218), Chub (Kyphosus 

incisor/Kyphosus sectatrix, α=0.005, P-value=0.0426), Purple Reeffish (Chromis scotti, 

α=0.005, P-value=0.0426), Tiger Grouper (Mycteroperca tigris, α=0.005, Pvalue=0.0428), Stoplight Parrotfish (Sparisoma viride, α=0.005, P-value=0.0353), Black 

Durgon (Melichthys niger, α=0.005, P-value=0.0032), Great Barracuda (Sphyraena 

barracuda, α=0.005, P-value=0.0031), Bonnetmouth (Emmelichthyops atlanticus, 

α=0.005, P-value=0.0103), Dusky Damselfish (Stegastes adustus, α=0.005, Pvalue=0.0346), Grey Snapper (Lutjanus agennes, α=0.005, P-value=0.0428), and 

Longspine Squirrelfish (Holocentrus rufus, α=0.005, P-value=0.0428). In 2010, of 63 

species observed, a significant difference was observed between the banks for 8 species. 

This included Dog Snapper (Lutjanus jocu, α=0.005, P-value=0.043), Bi-Color 

Damselfish (Stegastes partitus, α=0.005, P-value=0.0243), Creole Wrasse (Clepticus 

parrae, α=0.005, P-value=0.006), Spotfin Butterflyfish (Chaetodon ocellatus, α=0.005, 

P-value=0.0221), Purple Reeffish (Chromis scotti, α=0.005, P-value=0.0006), Stoplight 

Parrotfish (Sparisoma viride, α=0.005, P-value=0.0349), Neon Goby (Elacatinus 

oceanops, α=0.005, P-value=0.042), and Yellowhead Wrasse (Halichoeres garnoti, 

α=0.005, P-value=0.0406).

139 
Table 10.4.6. 

Species Occurrence and Percent Sighting Frequencies in Parentheses for all Species Recorded 

at the FGB in 2009 and 2010 

Numbers in "Both Banks", "EFGB", and "WFGB" represent the number of surveys in which the 

species was observed with percent sighting frequency in parentheses. "P-value" represents the 

probability that species occurrence is randomly distributed between Banks (Student's t-test). 

Statistically significant comparisons are shown in bold. NS= Not significant.

Species

2009 2010

Both 

Banks EFGB WFGB P-value Both Banks EFGB WFGB P-value

Lutjanidae: 

Lutjanus jocu 

(Snapper, Dog) 5 (10) 5 (21) - 0.0218 4 (8) - 4 (15) 0.043

Pomacentridae: 

Chromis 

multilineata 

(Chromis, Brown) 43 (88) 22 (88) 21 (88) NS 46 (90) 23 (92) 23 (88) NS

Serranidae: 

Paranthias furcifer

(Creolefish) 47 (96) 23 (92) 24 (100) NS 47 (92) 24 (96) 23 (88) NS

Labridae: 

Thalassoma 

lucasanum

(Wrasse, Bluehead) 46 (94) 23 (92) 23 (96) NS 47 (92) 24 (96) 23 (88) NS

Pomacentridae: 

Stegastes 

planifrons 

(Damselfish, 

Three-spot) 38 (78) 17 (68) 21 (88) NS 28 (55) 15 (60) 13 (50) NS

Pomacentridae: 

Stegastes variabilis 

(Damselfish, 

Cocoa) 18 (37) 7 (28) 11 (46) NS 33 (65) 15 (60) 18 (69) NS

Serranidae: 

Mycteroperca 

interstitialis 

(Grouper, 

Yellowmouth) 13 (27) 9 (36) 4 (17) NS 11 (22) 6 (24) 5 (19) NS

Kyphosidae: 

Kyphosus incisor / 

Kyphosus sectatrix 

(Chub) 27 (55) 17 (68) 10 (42) 0.0426 23 (45) 12 (48) 11 (42) NS

Scaridae: Scarus 

vetula (Parrotfish, 

Queen) 37 (76) 17 (68) 20 (83) NS 38 (75) 18 (72) 20 (77) NS

Cirrhitidae: 

Amblycirrhitus 

pinos (Hawkfish, 

Redspotted) 6 (12) 3 (12) 3 (13) NS 4 (8) 1 (4) 3 (12) NS

Pomacentridae: 46 (94) 23 (92) 23 (96) NS 43 (84) 24 (96) 19 (73) 0.0243

140 
Species

2009 2010

Both 

Banks EFGB WFGB P-value Both Banks EFGB WFGB P-value

Stegastes partitus 

(Damselfish, Bicolor)

Acanthuridae: 

Acanthurus 

coeruleus (Tang, 

Blue) 42 (86) 21 (84) 21 (88) NS 47 (92) 23 (92) 24 (92) NS

Atherinopsidae: 

Menidiinae sp. 

(Silversides) 2 (4) 2 (8) - NS - - - -

Pomacentridae: 

Chromis cyanea 

(Chromis, Blue) 45 (92) 23 (92) 22 (92) NS 33 (65) 15 (60) 18 (69) NS

Tetraodontidae: 

Canthigaster 

rostrata (Puffer, 

Sharpnose) 42 (86) 21 (84) 21 (88) NS 45 (88) 24 (96) 21 (81) NS

Labridae: Clepticus 

parrae (Wrasse, 

Creole) 44 (90) 23 (92) 21 (88) NS 29 (57) 19 (76) 10 (38) 0.006

Acanthuridae: 

Acanthurus 

chirurgus 

(Doctorfish) 9 (18) 6 (24) 3 (13) NS 4 (8) 2 (8) 2 (8) NS

Pomacanthidae: 

Holacanthus 

ciliaris (Angelfish, 

Queen) 7 (14) 3 (12) 4 (17) NS 1 (2) - 1 (4) NS

Labridae: Bodianus 

rufus (Hogfish, 

Spanish) 39 (80) 17 (68) 22 (92) NS 45 (88) 20 (80) 25 (96) NS

Blenniidae: 

Ophioblennius 

macclurei (Blenny, 

Redlip) 22 (45) 9 (36) 13 (54) NS 3 (6) - 3 (12) NS

Chaetodontidae: 

Chaetodon 

ocellatus 

(Butterflyfish, 

Spotfin) 9 (18) 5 (20) 4 (17) NS 5 (10) - 5 (19) 0.0221

Pomacentridae: 

Chromis scotti 

(Reeffish, Purple) 27 (55) 10 (40) 17 (71) 0.0426 20 (39) 4 (16) 16 (62) 0.0006

Scaridae: 

Sparisoma 

aurofrenatum

(Parrotfish, 

Redband) 17 (35) 11 (44) 6 (25) NS 17 (33) 11 (44) 6 (23) NS

Serranidae: 

Mycteroperca tigris 

(Grouper, Tiger) 4 (8) 4 (16) - 0.0428 14 (27) 6 (24) 8 (31) NS

141 
Species

2009 2010

Both 

Banks EFGB WFGB P-value Both Banks EFGB WFGB P-value

Pomacentridae: 

Microspathodon 

chrysurus 

(Damselfish, 

Yellowtail) 27 (55) 12 (48) 15 (63) NS 14 (27) 9 (36) 5 (19) NS

Pomacanthidae: 

Holacanthus 

tricolor (Angelfish, 

Rock Beauty) 19 (39) 10 (40) 9 (38) NS 6 (12) 5 (20) 1 (4) NS

Chaetodontidae: 

Chaetodon 

sedentarius 

(Butterflyfish, 

Reef) 23 (47) 10 (40) 13 (54) NS 27 (53) 12 (48) 15 (58) NS

Monacanthidae: 

Cantherhines 

pullus (Filefish, 

Orange Spotted) 9 (18) 5 (20) 4 (17) NS 1 (2) 1 (4) - NS

Pomacentridae: 

Chromis insolata 

(Sunshinefish) 23 (47) 10 (40) 13 (54) NS 9 (18) 4 (16) 5 (19) NS

Scaridae: 

Sparisoma viride

(Parrotfish, 

Stoplight) 31 (63) 12 (48) 19 (79) 0.0353 27 (53) 17 (68) 10 (38) 0.0349

Carangidae: 

Caranx latus (Jack, 

Horse-eye) 11 (22) 7 (28) 4 (17) NS 3 (6) - 3 (12) NS

Carangidae: 

Caranx lugubris 

(Jack, Black) 11 (22) 4 (16) 7 (29) NS 2 (4) 2 (8) - NS

Balistidae: 

Canthidermis 

sufflamen

(Triggerfish, 

Ocean) 6 (12) 4 (16) 2 (8) NS 1 (2) 1 (4) - NS

Serranidae: 

Cephalopholis 

cruentata

(Graysby) 21 (43) 10 (40) 11 (46) NS 25 (49) 12 (48) 13 (50) NS

Ostraciidae: 

Lactophrys 

triqueter 

(Trunkfish, 

Smooth) 11 (22) 4 (16) 7 (29) NS 13 (25) 9 (36) 4 (15) NS

Labridae: 

Halichoeres 

maculipinna

(Wrasse, Clown) 13 (27) 5 (20) 8 (33) NS 2 (4) - 2 (8) NS

Acanthuridae: 

Acanthurus 

bahianus 10 (20) 6 (24) 4 (17) NS 17 (33) 10 (40) 7 (27) NS

142 
Species

2009 2010

Both 

Banks EFGB WFGB P-value Both Banks EFGB WFGB P-value

(Surgeonfish, 

Ocean)

Balistidae: 

Melichthys niger 

(Durgon, Black) 24 (49) 7 (28) 17 (71) 0.0032 29 (57) 15 (60) 14 (54) NS

Pomacentridae: 

Abudefduf saxatilis 

(Sergeant Major) 12 (24) 4 (16) 8 (33) NS 9 (18) 2 (8) 7 (27) NS

Gobiidae: 

Elacatinus 

oceanops (Goby, 

Neon) 4 (8) 2 (8) 2 (8) NS 7 (14) 6 (24) 1 (4) 0.042

Sphyraenidae: 

Sphyraena 

barracuda 

(Barracuda, Great) 22 (45) 6 (24) 16 (67) 0.0031 39 (76) 17 (68) 22 (85) NS

Mobulidae: Manta 

birostris (Ray, 

Manta) 2 (4) 2 (8) - NS 1 (2) 1 (4) - NS

Serranidae: 

Mycteroperca 

bonaci (Grouper, 

Black) 1 (2) 1 (4) - NS 1 (2) - 1 (4) NS

Pomacanthidae: 

Pomacanthus paru 

(Angelfish, French) 5 (10) 4 (16) 1 (4) NS 5 (10) 4 (16) 1 (4) NS

Scaridae: Scarus 

taeniopterus 

(Parrotfish, 

Princess) 4 (8) 3 (12) 1 (4) NS 16 (31) 10 (40) 6 (23) NS

Serranidae: 

Epinephelus 

adscensionis (Hind, 

Rock) 2 (4) 2 (8) - NS 2 (4) 2 (8) - NS

Serranidae: 

Epinephelus 

guttatus (Hind, 

Red) 1 (2) 1 (4) - NS - - - -

Serranidae: 

Mycteroperca 

phenax (Scamp) 3 (6) 2 (8) 1 (4) NS 1 (2) 1 (4) - NS

Labridae: 

Halichoeres 

garnoti (Wrasse, 

Yellowhead) 10 (20) 3 (12) 7 (29) NS 10 (20) 2 (8) 8 (31) 0.0406

Mullidae: 

Pseudupeneus 

maculatus

(Goatfish, Spotted) 1 (2) 1 (4) - NS - - - -

Inermiidae: 

Emmelichthyops 

atlanticus 9 (18) 8 (32) 1 (4) 0.0103 - - - -

143 
Species

2009 2010

Both 

Banks EFGB WFGB P-value Both Banks EFGB WFGB P-value

(Bonnetmouth)

Ostraciidae: 

Acanthostracion 

polygonius

(Cowfish, 

Honeycomb) 5 (10) 1 (4) 4 (17) NS 1 (2) 1 (4) - NS

Echeneidae: 

Echeneidae sp. 

(Remora) 1 (2) 1 (4) - NS - - - -

Chaetodontidae: 

Prognathodes 

aculeatus 

(Butterflyfish, 

Longsnout) 8 (16) 2 (8) 6 (25) NS 9 (18) 3 (12) 6 (23) NS

Pomacentridae: 

Stegastes adustus 

(Damselfish, 

Dusky) 10 (20) 2 (8) 8 (33) 0.0346 5 (10) 3 (12) 2 (8) NS

Scaridae: 

Sparisoma 

atomarium

(Parrotfish, 

Greenblotch) 2 (4) 1 (4) 1 (4) NS - - - -

Carangidae: 

Carangoides ruber

(Jack, Bar) 5 (10) 3 (12) 2 (8) NS 13 (25) 6 (24) 7 (27) NS

Labridae: Bodianus 

pulchellus

(Hogfish, Spotfin) 1 (2) 1 (4) - NS - - - -

Ostraciidae: 

Lactophrys 

bicaudalis

(Trunkfish, 

Spotted) 3 (6) 1 (4) 2 (8) NS - - - -

Carangidae: 

Caranx crysos

(Blue runner) 2 (4) 1 (4) 1 (4) NS - - - -

Muraenidae: 

Gymnothorax 

meleagris (Moray, 

Spotted) 2 (4) 1 (4) 1 (4) NS 1 (2) - 1 (4) NS

Lutjanidae: 

Lutjanus agennes

(Snapper, Grey) 4 (8) - 4 (17) 0.0428 7 (14) 4 (16) 3 (12) NS

Apogonidae: 

Apogon sp. 

(Cardinal fish sp.) 1 (2) - 1 (4) NS - - - -

Aulostomidae: 

Aulostomus 

maculatus 

(Trumpetfish) 1 (2) - 1 (4) NS - - - -

144 
Species

2009 2010

Both 

Banks EFGB WFGB P-value Both Banks EFGB WFGB P-value

Holocentridae: 

Holocentrus rufus 

(Squirrelfish, 

Longspine) 4 (8) - 4 (17) 0.0428 3 (6) 1 (4) 2 (8) NS

Carangidae: 

Elagatis 

bipinnulata 

(Rainbow runner) 1 (2) - 1 (4) NS - - - -

Diodontidae: 

Diodon 

holocanthus 

(Ballonfish) 1 (2) - 1 (4) NS 1 (2) - 1 (4) NS

Monacanthidae: 

Cantherhines 

macrocerus

(Filefish, 

Whitespotted) 1 (2) - 1 (4) NS 1 (2) 1 (4) - NS

Mullidae: 

Mulloidichthys 

martinicus

(Goatfish, Yellow) 3 (6) - 3 (13) NS 2 (4) 2 (8) - NS

Gobiidae: 

Gnatholepis 

thompsoni (Goby, 

Goldspot) 1 (2) - 1 (4) NS - - - -

Muraenidae: 

Gymnothorax 

miliaris (Moray, 

Goldentail) 1 (2) - 1 (4) NS - - - -

Dasyatidae: 

Dasyatis americana

(Stingray, 

Southern) 1 (2) - 1 (4) NS - - - -

Labridae: 

Halichoeres 

radiatus (Pudding 

wife) 2 (4) - 2 (8) NS 2 (4) 2 (8) - NS

Labridae: 

Halichoeres 

bivittatus (Wrasse, 

Slippery Dick) - - - - 3 (6) - 3 (12) NS

Scaridae: Scarus 

iseri (Parrotfish, 

Striped) - - - - 3 (6) 2 (8) 1 (4) NS

Chaetodontidae: 

Chaetodon striatus 

(Butterflyfish, 

Banded) - - - - 2 (4) 1 (4) 1 (4) NS

Balistidae: Balistes 

vetula (Triggerfish, 

Queen) - - - - 1 (2) 1 (4) - NS

Serranidae: - - - - 1 (2) 1 (4) - NS

145 
Species

2009 2010

Both 

Banks EFGB WFGB P-value Both Banks EFGB WFGB P-value

Liopropoma rubre 

(Bass, Peppermint)

10.4.2. Family Density, Relative Abundance, and Richness 

The families with the highest total relative abundance included the damselfishes 

(Pomacentridae), wrasses (Labridae), and groupers and seabasses (Serranidae) (Table 

10.4.7). In 2009, damselfishes were recorded at a maximum density of 91.11 per 100 m2 

at the EFGB, while the minimum density of 41.29 per 100 m2 was recorded at the EFGB 

in 2010. A maximum density for the wrasse family of 59.63 per 100 m2 was recoded at 

the EFGB in 2009, while a minimum of 14.56 per 100 m2 was recorded at the EFGB in 

2010. The grouper and seabass family observed a maximum density of 26.42 per 100 m2 

in 2010 at the WFGB, and a minimum density of 13.91 per 100 m2 in 2009 at the 

WFGB. A maximum density of 3.92 per 100 m2 was recorded for the parrotfish family at 

the EFGB in 2010, while a minimum density of 2.83 per 100 m2 was recorded at the 

WFGB in both 2009 and 2010 (Table 10.4.8, Table 10.4.9, and Table 10.4.10). 

Families with the highest species richness included the damselfishes (Pomacentridae), 

groupers and seabasses (Serranidae), wrasses (Labridae), and parrotfishes (Scaridae). 

Between banks and year, the damselfishes (Pomacentridae) were represented by 10 

species. Groupers and seabasses were represented by a maximum of 8 species at the 

EFGB in 2009, and a minimum of 4 species at the WFGB in 2009. Wrasses were 

represented on average by 6 species, except for a minimum of 5 species at the EFGB in 

2010. The parrotfishes were represented by 5 species between years and banks (Table 

10.4.8 and Table 10.4.9). 

146 
Table 10.4.7. 

Relative Abundance by Family for the 2009 and 2010 Surveys at Both the EFGB and WFGB

Family Relative Abundance

Pomacentridae 41.5

Labridae 20.1

Serranidae 13.7

Inermiidae 9.7

Atherinopsidae 3.5

Scaridae 2.2

Acanthuridae 1.9

Kyphosidae 1.9

Tetraodontidae 1.5

Carangidae 1.3

Chaetodontidae 0.6

Sphyraenidae 0.6

Balistidae 0.4

Blenniidae 0.3

Pomacanthidae 0.2

Ostraciidae 0.2

Lutjanidae 0.1

Gobiidae 0.1

Monacanthidae 0.1

Mullidae 0.1

Cirrhitidae <0.1

Holocentridae <0.1

Muraenidae <0.1

Mobulidae <0.1

Aulostomidae <0.1

Diodontidae <0.1

Echeneidae <0.1

Apogonidae <0.1

Dasyatidae <0.1

147 
Table 10.4.8. 

Mean Fish Density (number of individuals per 100 m2), Richness (number of species per family), 

and Mean Biomass by Family at the EFGB and WFGB in 2009

Family

Density (fishes/100 m2) Species Richness Mean Biomass 

(g/m2)

EFGB WFGB EFGB WFGB EFGB WFGB

Acanthuridae 3.30 3.70 3 3 4.11 5.78

Atherinopsidae 21.46 0.00 1 0 0.59 0.00

Apogonidae 0.00 0.02 0 1 0.00 0.00

Aulostomidae 0.00 0.05 0 1 0.00 0.01

Balistidae 0.38 0.99 2 2 2.36 6.92

Blenniidae 0.64 0.92 1 1 0.01 0.02

Carangidae 3.14 3.18 3 5 15.84 63.97

Chaetodontidae 0.66 0.99 3 3 0.37 0.52

Cirrhitidae 0.07 0.07 1 1 0.00 0.00

Dasyatidae 0.00 0.02 0 1 0.00 1.81

Diodontidae 0.00 0.02 0 1 0.00 0.06

Echeneidae 0.05 0.00 1 0 0.03 0.00

Gobiidae 0.07 0.07 1 2 0.00 0.00

Holocentridae 0.00 0.09 0 1 0.00 0.16

Inermiidae 59.19 0.28 1 1 1.85 0.00

Kyphosidae 8.35 0.42 1 1 71.08 2.88

Labridae 59.63 29.76 6 6 7.46 10.78

Lutjanidae 0.19 0.26 1 1 1.27 0.94

Monacanthidae 0.14 0.14 1 2 0.08 0.34

Mullidae 0.05 0.21 1 1 0.05 0.45

Muraenidae 0.02 0.05 1 2 1.12 1.12

Ostraciidae 0.17 0.38 3 3 0.18 0.80

Pomacanthidae 0.71 0.40 3 3 2.73 1.86

Pomacentridae 91.11 72.01 10 10 5.60 4.07

Scaridae 3.23 2.83 5 5 6.38 10.01

Serranidae 22.57 13.91 8 4 38.51 18.80

Sphyraenidae 0.14 0.42 1 1 1.79 3.56

Tetraodontidae 2.50 2.95 1 1 0.08 0.09

148 
Table 10.4.9. 

Mean Fish Density (number of individuals per 100 m2), Richness (number of species per family), 

and Mean Biomass by Family at the EFGB and WFGB in 2010

Family Density (fishes/100 m2) Species Richness Mean Biomass 

(g/m2)

EFGB WFGB EFGB WFGB EFGB WFGB

Acanthuridae 2.49 1.78 3 3 3.55 3.59

Atherinopsidae 0.00 0.00 0 0 0.00 0.00

Apogonidae 0.00 0.00 0 0 0.00 0.00

Aulostomidae 0.00 0.00 0 0 0.00 0.00

Balistidae 0.61 0.65 3 1 3.96 3.78

Blenniidae 0.00 0.09 0 1 0.00 0.00

Carangidae 0.48 0.78 2 2 2.31 3.70

Chaetodontidae 0.77 1.13 3 4 0.23 0.38

Cirrhitidae 0.02 0.07 1 1 0.00 0.00

Dasyatidae 0.00 0.00 0 0 0.00 0.00

Diodontidae 0.00 0.02 0 1 0.00 0.05

Echeneidae 0.00 0.00 0 0 0.00 0.00

Gobiidae 0.25 0.04 1 1 0.01 0.00

Holocentridae 0.02 0.07 1 1 0.05 0.06

Inermiidae 0.00 0.00 0 0 0.00 0.00

Kyphosidae 1.45 1.07 1 1 10.07 3.72

Labridae 14.56 16.69 5 6 1.92 1.51

Lutjanidae 0.11 0.22 1 2 0.48 3.21

Monacanthidae 0.05 0.00 2 0 0.24 0.00

Mullidae 0.07 0.00 1 0 0.08 0.00

Muraenidae 0.00 0.02 0 1 0.00 0.02

Ostraciidae 0.25 0.11 2 1 0.32 0.03

Pomacanthidae 0.27 0.09 2 3 2.13 0.71

Pomacentridae 41.29 43.27 10 10 2.79 2.31

Scaridae 3.92 2.83 5 5 13.19 7.98

Serranidae 17.97 26.42 7 5 25.38 38.34

Sphyraenidae 1.06 1.63 1 1 6.14 17.68

Tetraodontidae 1.79 1.98 1 1 0.16 0.07

149 
10.4.3. Diveristy and Evenness of Populations

Diversity and evenness was measured between banks and years by calculating ShannonWiener diversity indices (See Table 10.4.10). Diversity varied between banks and years,

with the greatest diversity occurring at the WFGB in 2009. The lowest diversity was 

calculated for the WFGB in 2010. Evenness between banks and years remained relatively 

stable, with the least variation in communities occurring at the WFGB in 2009. 

Table 10.4.10. 

Shannon-Wiener Diversity Indices for the EFGB and WFGB in 2009 and 2010

Index

2009 2010

EFGB WFGB EFGB WFGB

Samples (n) 24 24 25 26

Diversity (H') 1.05 1.12 1.01 0.99

Diversity 

(H'max) 1.78 1.80 1.72 1.71

Evenness (J') 0.59 0.62 0.59 0.58

When compared between banks, the WFGB had a significantly higher difference in 

diversity and evenness than was observed at the EFGB in 2009 (α=0.005, Pvalue=0.0028 and 0.0367 respectively). No significant difference in diversity or evenness 

was observed between the EFGB and WFGB in 2010. When diversity and evenness was 

compared between years at each bank, no significant difference was observed between 

years except for diversity at the WFGB between 2009 and 2010 (α=0.005, Pvalue=0.0050), suggesting a significantly greater species diversity at the WFGB in 2009 

than 2010. 

10.4.4. Trophic Group Comparisons

Species were grouped by trophic guild into four major categories: Herbivores, Piscivores, 

Invertivores, and Planktivores. As defined by NOAA’s Center for Coastal Monitoring 

and Assessment (CCMA) BioGeography Branch fish-trophic level database, the 

herbivore guild is comprised of Acanthuridae, Blennidae, Kyphosidae, Scaridae, and 

members of the Balistidae (Black Durgon), Gobiidae (Goldspot Goby), Pomacanthidae 

(Queen Angelfish), and Pomacentridae (Cocoa, Bi-Color, Yellowtail, and Dusky 

Damselfish) families. The piscivore guild is comprised of Aulostomidae, Carangidae, 

Inermiidae, Muraenidae, Sphyraenidae, and members of the Lutjanidae (Dog Snapper), 

and Serranidae (Yellowmouth, Tiger, Black, and Yellowfin Groupers, Graysby, and 

Scamp) families. The invertivore guild is comprised of Apogonidae, Chaetodontidae, 

Cirrhitidae, Dasyatidae, Diodontidae, Holocentridae, Monacanthidae, Mullidae, 

Muraenidae, Ostraciidae, Tetraodontidae, and members of the Balistidae (Ocean and 

Queen Triggerfish), Gobiidae (Neon Goby), Labridae (Spanish and Spotfin Hogfish, 

Clown, Yellowhead, Slippery Dick, and Bluehead Wrasse, and Puddingwife), Lutjanidae 

(Grey Snapper), and Pomacanthidae (Rock Beauty and French Angelfish), Pomacentridae 

(Brown Chromis, Sergeant Major, and Three-Spot Damselfish), Serranidae (Rock and 

150 
Red Hinds, and Coney) families. The planktivore guild is comprised of Echeneidae, and 

members of the Labridae (Creole Wrasse), Pomacentridae (Blue Chromis, Purple 

Reeffish, and Sunshinefish), and Serranidae (Creolefish) families. The mean density of 

herbivores ranged from 10 per 100 m2 at the WFGB in 2010 to 47 per 100 m2 at the 

EFGB in 2009. The mean density of piscivores ranged from 3 per 100 m2 at the WFGB in 

2010 to 63 per 100 m2 at the EFGB in 2009. The invertivore trophic guild mean densities 

ranged from 2 per 100 m

2 at the EFGB in 2010 to 87 per 100 m2 at the EFGB in 2009. 

The mean density of planktivores ranged from 25 per 100 m2 at the EFGB in 2010 to 80 

per 100 m

2 at the EFGB in 2009 (Table 10.4.11). In both 2009 and 2010, the EFGB had a 

significantly higher difference in herbivore density than WFGB, P-value=0.0131 and 

0.0244 respectively (α=0.005). The EFGB was also observed to be significantly higher in 

the piscivore density than the WFGB in 2009 (α=0.005, P-value=0.006), and significantly 

higher in the planktivore density than the WFGB in 2009 (α=0.005, P-value=0.0494). In 

yearly comparisons, a significant difference was observed between the densities of all 

trophic guilds at the EFGB between 2009 and 2010. Herbivore, piscivore, invertivore, 

and planktivore densities were all significantly different between 2009 and 2010, Pvalues=0.0013, 0.0036, 0.0021, and <0.0001 respectively (α=0.005). However, the 

WFGB only showed a significant difference between the densities of herbivores 

(α=0.005, P-value=0.0003) and invertivores (α=0.005, P-value=0.0193) between 2009 

and 2010. 

Table 10.4.11. 

Comparison of Mean Density (fish per 100 m2) and Species Richness per Diver Survey of 

Herbivores, Piscivores, Invertivores, and Planktivores in Diver Surveys between the EFGB and 

WFGB in 2009 and 2010

NS= Not significant

Category

2009 2010

EFGB WFGB Pvalue Significance EFGB WFGB Pvalue Significance

Herbivore 

Density 47 16 0.0131 S 14 10 0.0244 S

Piscivore 

Density 63 4 0.006 S 46 3 0.0702 NS

Invertivore 

Density 87 70 0.4316 NS 2 47 0.9135 NS

Planktivore 

Density 80 43 0.0494 S 25 38 0.5237 NS

Herbivore 

Richness 16 16 0.1167 NS 14 15 0.0138 S

Piscivore 

Richness 13 12 0.246 NS 7 9 0.0955 NS

Invertivore 

Richness 25 30 <.0001 S 25 22 0.7303 NS

Planktivore 

Richness 6 5 0.2286 NS 5 7 0.6425 NS

Species richness was recorded per trophic guild by year and bank. Herbivore richness 

ranged from 14 species at the EFGB in 2010 to 16 species at both the EFGB and WFGB 

151 
in 2009. Piscivore richness ranged from 7 species at the EFGB in 2010 to 13 species at 

the EFGB in 2009. Invertivore richness was recorded at a minimum of 22 species at the 

WFGB in 2010 and a maximum of 25 species at the EFGB in 2009. Planktivore richness 

ranged from a low of 5 species at the WFGB in 2009 and EFGB in 2010, to a high of 7 

species at the WFGB in 2010. A statistically significant difference was observed between 

the herbivore richness at the EFGB and WFGB in 2010 (α=0.005, P-value=0.0138) and 

between the invertivore richness at the EFGB and WFGB in 2009 (α=0.005, Pvalue<0.0001) (Table 10.4.11).

Species were grouped into trophic guild, and densities were compared between banks for 

each year (Table 10.4.12). The herbivore guild in 2009 showed significant differences in 

the density of Chub (Kyphosus incisor/Kyphosus sectatrix, α=0.005, P-value=0.0044), 

Redband Parrotfish (Sparisoma aurofrenatum, α=0.005, P-value=0.0169), Stoplight 

Parrotfish (Sparisoma viride, α=0.005, P-value=0.0076), and Black Durgon (Melichthys 

niger, α=0.005, P-value=0.0081). In 2010, the herbivore guild showed significant 

differences in the density of Bi-Color Damselfish (Stegastes partitus, α=0.005, Pvalue=0.007), while all other species showed no significant difference. The piscivore 

guild in 2009 showed significant differences in the density of Dog Snapper (Lutjanus 

jocu, α=0.005, P-value=0.0262), Tiger Grouper (Mycteroperca tigris, α=0.005, Pvalue=0.0428), Great Barracuda (Sphyraena barracuda, α=0.005, P-value=0.0021), and 

Bonnetmouth (Emmelichthyops atlanticus, α=0.005, P-value=0.0044). In 2010, no 

significant difference was observed in the densities of any piscivore species. The 

invertivore guild in 2009 showed significant differences in the densities of Spanish 

Hogfish (Bodianus rufus, α=0.005, P-value=0.0162) and Longspine Squirrelfish 

(Holocentrus rufu, α=0.005, P-value=0.0428). In 2010, this guild showed only a 

significant difference in the density of Spotfin Butterflyfish (Chaetodon ocellatus, 

α=0.005, P-value=0.0363). The planktivore guild showed significant differences in the 

densities of Creole Wrasse (Clepticus parrae, α=0.005, P-value=0.0424) and Purple 

Reeffish (Chromis scotti, α=0.005, P-value=0.0342). In 2010, the planktivore guild 

showed significant differences in the densities of Blue Chromis (Chromis cyanea, 

α=0.005, P-value=0.0416) and Purple Reeffish (Chromis scotti, α=0.005, Pvalue=0.0047). 

152 
Table 10.4.12. 

Density Comparison between Banks for Herbivorous, Piscivorous, Invertivorous, and 

Planktivorous Fishes 

Significant differences show P-values in bold. NS = Not significant.

Trophic 

Guild Species

2009 2010

Number/100 m2 Number/100 m2

EFGB WFGB Pvalue EFGB WFGB Pvalue

Herbivores

All Herbivores 47 16 13.5 10.3

Pomacentridae: Stegastes 

variabilis (Damselfish, 

Cocoa) 0.7 0.9 NS 1.6 1.8 NS

Kyphosidae: Kyphosus 

incisor / Kyphosus 

sectatrix (Chub) 8.3 0.4 0.0044 1.4 1.1 NS

Scaridae: Scarus vetula 

(Parrotfish, Queen) 1.3 1.3 NS 1.6 1.5 NS

Pomacentridae: Stegastes 

partitus (Damselfish, Bicolor) 8.6 5.5 NS 3.0 1.8 0.007

Acanthuridae: Acanthurus 

coeruleus (Tang, Blue) 2.8 3.3 NS 1.9 1.4 NS

Atherinopsidae: 

Menidiinae sp. 

(Silversides) 21.5 0.0 NS 0.0 0.0 NS

Acanthuridae: Acanthurus 

chirurgus (Doctorfish) 0.3 0.3 NS 0.1 0.1 NS

Pomacanthidae: 

Holacanthus ciliaris 

(Angelfish, Queen) 0.1 0.1 NS 0.0 0.0 NS

Blenniidae: Ophioblennius 

macclurei (Blenny, Redlip) 0.6 0.9 NS 0.0 0.1 NS

Scaridae: Sparisoma 

aurofrenatum (Parrotfish, 

Redband) 1.2 0.2 0.0169 0.6 0.3 NS

Pomacentridae: 

Microspathodon chrysurus 

(Damselfish, Yellowtail) 0.4 0.8 NS 0.4 0.2 NS

Scaridae: Sparisoma viride

(Parrotfish, Stoplight) 0.5 1.2 0.0076 0.9 0.7 NS

Acanthuridae: Acanthurus 

bahianus (Surgeonfish, 

Ocean) 0.2 0.1 NS 0.5 0.3 NS

Balistidae: Melichthys 

niger (Durgon, Black) 0.3 0.9 0.0081 0.5 0.7 NS

Scaridae: Scarus 

taeniopterus (Parrotfish, 

Princess) 0.1 0.0 NS 0.8 0.4 NS

Pomacentridae: Stegastes 

adustus (Damselfish, 

Dusky) 0.1 0.4 NS 0.2 0.1 NS

153 
Trophic 

Guild Species

2009 2010

Number/100 m2 Number/100 m2

EFGB WFGB Pvalue EFGB WFGB Pvalue

Scaridae: Sparisoma 

atomarium (Parrotfish, 

Greenblotch) 0.0 0.0 NS 0.0 0.0 NS

Gobiidae: Gnatholepis 

thompsoni (Goby, 

Goldspot) 0.0 0.0 NS 0.0 0.0 NS

Scaridae: Scarus iseri

(Parrotfish, Striped) 0.0 0.0 NS 0.1 0.0 NS

Piscivores

All Piscivores 63.4 4.4 46.3 3.4

Lutjanidae: Lutjanus jocu

(Snapper, Dog) 0.2 0.0 0.0262 0.0 0.1 NS

Serranidae: Mycteroperca 

interstitialis (Grouper, 

Yellowmouth) 0.3 0.1 NS 0.1 0.1 NS

Serranidae: Mycteroperca 

tigris (Grouper, Tiger) 0.1 0.0 0.0428 0.2 0.2 NS

Carangidae: Caranx latus 

(Jack, Horse-eye) 2.0 0.4 NS 0.0 0.1 NS

Carangidae: Caranx 

lugubris (Jack, Black) 0.1 0.2 NS 0.1 0.0 NS

Serranidae: Cephalopholis 

cruentata (Graysby) 0.3 0.4 NS 0.4 0.4 NS

Sphyraenidae: Sphyraena 

barracuda (Barracuda, 

Great) 0.1 0.4 0.0021 1.1 1.6 NS

Serranidae: Mycteroperca 

bonaci (Grouper, Black) 0.0 0.0 NS 0.0 0.0 NS

Serranidae: Mycteroperca 

phenax (Scamp) 0.0 0.0 NS 0.0 0.0 NS

Inermiidae: 

Emmelichthyops atlanticus

(Bonnetmouth) 59.2 0.3 0.0044 0.0 0.0 NS

Carangidae: Carangoides 

ruber (Jack, Bar) 1.0 0.1 NS 0.4 0.7 NS

Carangidae: Caranx crysos 

(Blue runner) 0.1 0.0 NS 0.0 0.0 NS

Muraenidae: Gymnothorax 

meleagris (Moray, 

Spotted) 0.0 0.0 NS 0.0 0.0 NS

Aulostomidae: Aulostomus 

maculatus (Trumpetfish) 0.0 0.0 NS 0.0 0.0 NS

Carangidae: Elagatis 

bipinnulata (Rainbow 

runner) 0.0 2.4 NS 0.0 0.0 NS

Serranidae: mycteroperca 

venenosa (Grouper, 

Yellowfin) 0.0 0.0 NS 0.0 0.0 NS

Invertivores

All Invertivores 87.1 69.8 2.2 46.9

Pomacentridae: Chromis 

multilineata (Chromis, 59.7 42.8 NS 32.8 32.9 NS

154 
Trophic 

Guild Species

2009 2010

Number/100 m2 Number/100 m2

EFGB WFGB Pvalue EFGB WFGB Pvalue

Brown)

Labridae: Thalassoma 

lucasanum (Wrasse, 

Bluehead) 17.6 13.3 NS 6.4 6.2 NS

Pomacentridae: Stegastes 

planifrons (Damselfish, 

Three-spot) 3.3 3.7 NS 1.6 1.2 NS

Cirrhitidae: Amblycirrhitus 

pinos (Hawkfish, 

Redspotted) 0.1 0.1 NS 0.0 0.1 NS

Tetraodontidae: 

Canthigaster rostrata

(Puffer, Sharpnose) 2.5 2.9 NS 1.8 2.0 NS

Labridae: Bodianus rufus

(Hogfish, Spanish) 1.1 2.0 0.0162 1.4 2.0 NS

Chaetodontidae: 

Chaetodon ocellatus 

(Butterflyfish, Spotfin) 0.2 0.2 NS 0.0 0.2 0.0265

Pomacanthidae: 

Holacanthus tricolor 

(Angelfish, Rock Beauty) 0.4 0.3 NS 0.1 0.0 NS

Chaetodontidae: 

Chaetodon sedentarius 

(Butterflyfish, Reef) 0.4 0.5 NS 0.7 0.8 NS

Monacanthidae: 

Cantherhines pullus

(Filefish, Orange Spotted) 0.1 0.1 NS 0.0 0.0 NS

Balistidae: Canthidermis 

sufflamen (Triggerfish, 

Ocean) 0.1 0.1 NS 0.0 0.0 NS

Ostraciidae: Lactophrys 

triqueter (Trunkfish, 

Smooth) 0.1 0.2 NS 0.2 0.1 NS

Labridae: Halichoeres 

maculipinna (Wrasse, 

Clown) 0.3 1.2 NS 0.0 0.1 NS

Pomacentridae: Abudefduf 

saxatilis (Sergeant Major) 0.2 0.6 NS 0.1 0.3 NS

Gobiidae: Elacatinus 

oceanops (Goby, Neon) 0.1 0.0 NS 0.2 0.0 NS

Pomacanthidae: 

Pomacanthus paru

(Angelfish, French) 0.2 0.0 NS 0.2 0.0 NS

Serranidae: Epinephelus 

adscensionis (Hind, Rock) 0.0 0.0 NS 0.0 0.0 NS

Serranidae: Epinephelus 

guttatus (Hind, Red) 0.0 0.0 NS 0.0 0.0 NS

Labridae: Halichoeres 

garnoti (Wrasse, 

Yellowhead) 0.2 0.4 NS 0.1 0.4 NS

155 
Trophic 

Guild Species

2009 2010

Number/100 m2 Number/100 m2

EFGB WFGB Pvalue EFGB WFGB Pvalue

Mullidae: Pseudupeneus 

maculatus (Goatfish, 

Spotted) 0.0 0.0 NS 0.0 0.0 NS

Ostraciidae: 

Acanthostracion 

polygonius (Cowfish, 

Honeycomb) 0.0 0.1 NS 0.0 0.0 NS

Chaetodontidae: 

Prognathodes aculeatus 

(Butterflyfish, Longsnout) 0.0 0.3 NS 0.1 0.2 NS

Labridae: Bodianus 

pulchellus (Hogfish, 

Spotfin) 0.0 0.0 NS 0.0 0.0 NS

Ostraciidae: Lactophrys 

bicaudalis (Trunkfish, 

Spotted) 0.0 0.1 NS 0.0 0.0 NS

Lutjanidae: Lutjanus 

agennes (Snapper, Grey) 0.0 0.3 NS 0.1 0.1 NS

Apogonidae: Apogon sp. 

(Cardinal fish sp.) 0.0 0.0 NS 0.0 0.0 NS

Holocentridae: 

Holocentrus rufus

(Squirrelfish, Longspine) 0.0 0.1 0.0428 0.0 0.1 NS

Diodontidae: Diodon 

holocanthus (Ballonfish) 0.0 0.0 NS 0.0 0.0 NS

Monacanthidae: 

Cantherhines macrocerus 

(Filefish, Whitespotted) 0.0 0.0 NS 0.0 0.0 NS

Mullidae: Mulloidichthys 

martinicus (Goatfish, 

Yellow) 0.0 0.2 NS 0.1 0.0 NS

Muraenidae: Gymnothorax 

miliaris (Moray, 

Goldentail) 0.0 0.0 NS 0.0 0.0 NS

Dasyatidae: Dasyatis 

americana (Stingray, 

Southern) 0.0 0.0 NS 0.0 0.0 NS

Labridae: Halichoeres 

radiatus (Pudding wife) 0.0 0.0 NS 0.0 0.0 NS

Labridae: Halichoeres 

bivittatus (Wrasse, 

Slippery Dick) 0.0 0.0 NS 0.0 0.2 NS

Tetraodontidae: 

Sphoeroides spengleri 

(Puffer, Bandtail) 0.0 0.0 NS 0.0 0.0 NS

Chaetodontidae: 

Chaetodon striatus 

(Butterflyfish, Banded) 0.0 0.0 NS 0.0 0.0 NS

Balistidae: Balistes vetula

(Triggerfish, Queen) 0.0 0.0 NS 0.0 0.0 NS

156 
Trophic 

Guild Species

2009 2010

Number/100 m2 Number/100 m2

EFGB WFGB Pvalue EFGB WFGB Pvalue

Serranidae: Liopropoma 

rubre (Bass, Peppermint) 0.0 0.0 NS 0.0 0.0 NS

Holocentridae: holocentrus 

adscensionis (Squirrelfish) 0.0 0.0 NS 0.0 0.0 NS

Serranidae: Cephalopholis 

fulva (Grouper, Coney) 0.0 0.0 NS 0.0 0.0 NS

Planktivores

All Planktivores 80.2 43.5 25.0 38.5

Serranidae: Paranthias 

furcifer (Creolefish) 21.8 13.4 NS 17.2 25.6 NS

Pomacentridae: Chromis 

cyanea (Chromis, Blue) 12.4 10.6 NS 0.9 2.9 0.0416

Labridae: Clepticus parrae

(Wrasse, Creole) 40.3 12.8 0.0424 6.6 7.9 NS

Pomacentridae: Chromis 

scotti (Reeffish, Purple) 3.3 5.7 0.0342 0.5 1.8 0.0047

Pomacentridae: Chromis 

insolata (Sunshinefish) 2.5 1.0 NS 0.2 0.3 NS

Echeneidae: Echeneidae

sp. (Remora) 0.0 0.0 NS 0.0 0.0 NS

Lutjanidae: Lutjanus 

argentiventris (Snapper, 

Yellowtail) 0.0 0.0 NS 0.0 0.0 NS

When species densities were compared at each bank between years (Table 10.4.13), a 

significant difference was observed between 16 species at the EFGB between 2009 and 

2010. This included the herbivorious Cocoa Damselfish (Stegastes variabilis, α=0.005, Pvalue=0.0425), Bi-Color Damselfish (Stegastes partitus, α=0.005, P-value<0.0001), 

Redlip Blenny (Ophioblennius macclurei, α=0.005, P-value=0.0025), and Princess 

Parrotfish (Scarus taeniopterus, α=0.005, P-value=0.0097), the piscivorous Dog Snapper 

(Lutjanus jocu, α=0.005, P-value=0.0262), Horse-Eye Jack (Caranx latus, α=0.005, Pvalue=0.015), Great Barracuda (Sphyraena barracuda, α=0.005, P-value=0.0002), and 

Bonnetmouth (Emmelichthyops atlanticus, α=0.005, P-value=0.0027), the invertivorous 

Bluehead Wrasse (Thalassoma lucasanum, α=0.005, P-value=0.0015), Spotfin 

Butterflyfish (Chaetodon ocellatus, α=0.005, P-value=0.0242), Rock Beauty 

(Holacanthus tricolor, α=0.005, P-value=0.0305), and Clown Wrasse (Halichoeres 

maculipinna, α=0.005, P-value=0.0241), and the planktivorous Blue Chromis (Chromis 

cyanea, α=0.005, P-value<0.0001), Creole Wrasse (Clepticus parrae, α=0.005, Pvalue=0.0002), Purple Reeffish (Chromis scotti, α=0.005, P-value=0.0238), and 

Sunshinefish (Chromis insolata, α=0.005, P-value=0.008). 

At the WFGB, a significant difference in density was observed between 18 species 

between 2009 and 2010. This included the herbivorous Bi-Color Damselfish (Stegastes 

partitus, α=0.005, P-value<0.0001), Redlip Blenny (Ophioblennius macclurei, α=0.005, 

P-value=0.001), Yellowtail Damselfish (Microspathodon chrysurus, α=0.005, Pvalue=0.0009), Stoplight Parrotfish (Sparisoma viride, α=0.005, P-value=0.0161), and 

157 
Princess Parrotfish (Scarus taeniopterus, α=0.005, P-value=0.0256), the piscivorous 

Tiger Grouper (Mycteroperca tigris, α=0.005, P-value=0.0051), Black Jack (Caranx 

lugubris, α=0.005, P-value=0.0094), and Great Barracuda (Sphyraena barracuda, 

α=0.005, P-value<0.0001), the invertivorous Bluehead Wrasse (Thalassoma lucasanum, 

α=0.005, P-value=0.0044), Three-Spot Damselfish (Stegastes planifrons, α=0.005, Pvalue=0.0004), Rock Beauty (Holacanthus tricolor, α=0.005, P-value=0.0047), OrangeSpotted Filefish (Cantherhines pullus, α=0.005, P-value=0.0482), Clown Wrasse 

(Clepticus parrae, α=0.005, P-value=0.0102), and Honeycomb Cowfish 

(Acanthostracion polygonius, α=0.005, P-value=0.0482), and the planktivorous Blue 

Chromis (Chromis cyanea, α=0.005, P-value=0.0023), Creole Wrasse (Clepticus parrae, 

α=0.005, P-value=0.0008), Purple Reeffish (Chromis scotti, α=0.005, P-value=0.007), 

and Sunshinefish (Chromis insolata, α=0.005, P-value=0.0141) (Table 10.4.13) 

Table 10.4.13. 

Abundances between Years for Herbivores, Piscivores, Invertivores, and Planktivores 

Comparisons of groups where student's t-tests were performed are in bold with appropriate Pvalues where significant. NS = Not significant

Trophic 

Guild Species

EFGB WFGB

Number/100 m2 Number/100 m2

2009 2010 P-value 2009 2010 P-value

Herbivores

All Herbivores 47 13.5 16 10.3

Pomacentridae: Stegastes 

variabilis (Damselfish, 

Cocoa) 0.7 1.6 0.0425 0.9 1.8 NS

Kyphosidae: Kyphosus 

incisor / Kyphosus 

sectatrix (Chub) 8.3 1.4 NS 0.4 1.1 NS

Scaridae: Scarus vetula 

(Parrotfish, Queen) 1.3 1.6 NS 1.3 1.5 NS

Pomacentridae: Stegastes 

partitus (Damselfish, Bicolor) 8.6 3.0 <0.0001 5.5 1.8 <0.0001

Acanthuridae: Acanthurus 

coeruleus (Tang, Blue) 2.8 1.9 NS 3.3 1.4 NS

Atherinopsidae: 

Menidiinae sp. 

(Silversides) 21.5 0.0 NS 0.0 0.0 NS

Acanthuridae: Acanthurus 

chirurgus (Doctorfish) 0.3 0.1 NS 0.3 0.1 NS

Pomacanthidae: 

Holacanthus ciliaris 

(Angelfish, Queen) 0.1 0.0 NS 0.1 0.0 NS

Blenniidae: Ophioblennius 

macclurei (Blenny, 

Redlip) 0.6 0.0 0.0025 0.9 0.1 0.001

Scaridae: Sparisoma 

aurofrenatum (Parrotfish, 

Redband) 1.2 0.6 NS 0.2 0.3 NS

Pomacentridae: 0.4 0.4 NS 0.8 0.2 0.0009

158 
Trophic 

Guild Species

EFGB WFGB

Number/100 m2 Number/100 m2

2009 2010 P-value 2009 2010 P-value

Microspathodon chrysurus 

(Damselfish, Yellowtail)

Scaridae: Sparisoma viride

(Parrotfish, Stoplight) 0.5 0.9 NS 1.2 0.7 0.0161

Acanthuridae: Acanthurus 

bahianus (Surgeonfish, 

Ocean) 0.2 0.5 NS 0.1 0.3 NS

Balistidae: Melichthys 

niger (Durgon, Black) 0.3 0.5 NS 0.9 0.7 NS

Scaridae: Scarus 

taeniopterus (Parrotfish, 

Princess) 0.1 0.8 0.0097 0.0 0.4 0.0256

Pomacentridae: Stegastes 

adustus (Damselfish, 

Dusky) 0.1 0.2 NS 0.4 0.1 NS

Scaridae: Sparisoma 

atomarium (Parrotfish, 

Greenblotch) 0.0 0.0 NS 0.0 0.0 NS

Gobiidae: Gnatholepis 

thompsoni (Goby, 

Goldspot) 0.0 0.0 NS 0.0 0.0 NS

Scaridae: Scarus iseri

(Parrotfish, Striped) 0.0 0.1 NS 0.0 0.0 NS

Piscivores

All Piscivores 63.4 46.3 4.4 3.4

Lutjanidae: Lutjanus jocu

(Snapper, Dog) 0.2 0.0 0.0262 0.0 0.1 NS

Serranidae: Mycteroperca 

interstitialis (Grouper, 

Yellowmouth) 0.3 0.1 NS 0.1 0.1 NS

Serranidae: Mycteroperca 

tigris (Grouper, Tiger) 0.1 0.2 NS 0.0 0.2 0.0051

Carangidae: Caranx latus 

(Jack, Horse-eye) 2.0 0.0 0.015 0.4 0.1 NS

Carangidae: Caranx 

lugubris (Jack, Black) 0.1 0.1 NS 0.2 0.0 0.0094

Serranidae: Cephalopholis 

cruentata (Graysby) 0.3 0.4 NS 0.4 0.4 NS

Sphyraenidae: Sphyraena 

barracuda (Barracuda, 

Great) 0.1 1.1 0.0002 0.4 1.6 <0.0001

Serranidae: Mycteroperca 

bonaci (Grouper, Black) 0.0 0.0 NS 0.0 0.0 NS

Serranidae: Mycteroperca 

phenax (Scamp) 0.0 0.0 NS 0.0 0.0 NS

Inermiidae: 

Emmelichthyops atlanticus

(Bonnetmouth) 59.2 0.0 0.0027 0.3 0.0 NS

Carangidae: Carangoides 

ruber (Jack, Bar) 1.0 0.4 NS 0.1 0.7 NS

Carangidae: Caranx crysos 

(Blue runner) 0.1 0.0 NS 0.0 0.0 NS

159 
Trophic 

Guild Species

EFGB WFGB

Number/100 m2 Number/100 m2

2009 2010 P-value 2009 2010 P-value

Muraenidae: Gymnothorax 

meleagris (Moray, 

Spotted) 0.0 0.0 NS 0.0 0.0 NS

Aulostomidae: Aulostomus 

maculatus (Trumpetfish) 0.0 0.0 NS 0.0 0.0 NS

Carangidae: Elagatis 

bipinnulata (Rainbow 

runner) 0.0 0.0 NS 2.4 0.0 NS

Serranidae: mycteroperca 

venenosa (Grouper, 

Yellowfin) 0.0 0.0 NS 0.0 0.0 NS

Invertivores

All Invertivores 87.1 2.2 69.8 46.9

Pomacentridae: Chromis 

multilineata (Chromis, 

Brown) 59.7 32.8 NS 42.8 32.9 NS

Labridae: Thalassoma 

lucasanum (Wrasse, 

Bluehead) 17.6 6.4 0.0015 13.3 6.2 0.0044

Pomacentridae: Stegastes 

planifrons (Damselfish, 

Three-spot) 3.3 1.6 NS 3.7 1.2 0.0004

Cirrhitidae: Amblycirrhitus 

pinos (Hawkfish, 

Redspotted) 0.1 0.0 NS 0.1 0.1 NS

Tetraodontidae: 

Canthigaster rostrata

(Puffer, Sharpnose) 2.5 1.8 NS 2.9 2.0 NS

Labridae: Bodianus rufus

(Hogfish, Spanish) 1.1 1.4 NS 2.0 2.0 NS

Chaetodontidae: 

Chaetodon ocellatus 

(Butterflyfish, Spotfin) 0.2 0.0 0.0242 0.2 0.2 NS

Pomacanthidae: 

Holacanthus tricolor 

(Angelfish, Rock Beauty) 0.4 0.1 0.0305 0.3 0.0 0.0047

Chaetodontidae: 

Chaetodon sedentarius 

(Butterflyfish, Reef) 0.4 0.7 NS 0.5 0.8 NS

Monacanthidae: 

Cantherhines pullus

(Filefish, Orange Spotted) 0.1 0.0 NS 0.1 0.0 0.0428

Balistidae: Canthidermis 

sufflamen (Triggerfish, 

Ocean) 0.1 0.0 NS 0.1 0.0 NS

Ostraciidae: Lactophrys 

triqueter (Trunkfish, 

Smooth) 0.1 0.2 NS 0.2 0.1 NS

Labridae: Halichoeres 

maculipinna (Wrasse, 

Clown) 0.3 0.0 0.0241 1.2 0.1 0.0102

160 
Trophic 

Guild Species

EFGB WFGB

Number/100 m2 Number/100 m2

2009 2010 P-value 2009 2010 P-value

Pomacentridae: Abudefduf 

saxatilis (Sergeant Major) 0.2 0.1 NS 0.6 0.3 NS

Gobiidae: Elacatinus 

oceanops (Goby, Neon) 0.1 0.2 NS 0.0 0.0 NS

Pomacanthidae: 

Pomacanthus paru

(Angelfish, French) 0.2 0.2 NS 0.0 0.0 NS

Serranidae: Epinephelus 

adscensionis (Hind, Rock) 0.0 0.0 NS 0.0 0.0 NS

Serranidae: Epinephelus 

guttatus (Hind, Red) 0.0 0.0 NS 0.0 0.0 NS

Labridae: Halichoeres 

garnoti (Wrasse, 

Yellowhead) 0.2 0.1 NS 0.4 0.4 NS

Mullidae: Pseudupeneus 

maculatus (Goatfish, 

Spotted) 0.0 0.0 NS 0.0 0.0 NS

Ostraciidae: 

Acanthostracion 

polygonius (Cowfish, 

Honeycomb) 0.0 0.0 NS 0.1 0.0 0.0482

Chaetodontidae: 

Prognathodes aculeatus 

(Butterflyfish, Longsnout) 0.0 0.1 NS 0.3 0.2 NS

Labridae: Bodianus 

pulchellus (Hogfish, 

Spotfin) 0.0 0.0 NS 0.0 0.0 NS

Ostraciidae: Lactophrys 

bicaudalis (Trunkfish, 

Spotted) 0.0 0.0 NS 0.1 0.0 NS

Lutjanidae: Lutjanus 

agennes (Snapper, Grey) 0.0 0.1 NS 0.3 0.1 NS

Apogonidae: Apogon sp. 

(Cardinal fish sp.) 0.0 0.0 NS 0.0 0.0 NS

Holocentridae: 

Holocentrus rufus

(Squirrelfish, Longspine) 0.0 0.0 NS 0.1 0.1 NS

Diodontidae: Diodon 

holocanthus (Ballonfish) 0.0 0.0 NS 0.0 0.0 NS

Monacanthidae: 

Cantherhines macrocerus 

(Filefish, Whitespotted) 0.0 0.0 NS 0.0 0.0 NS

Mullidae: Mulloidichthys 

martinicus (Goatfish, 

Yellow) 0.0 0.1 NS 0.2 0.0 NS

Muraenidae: Gymnothorax 

miliaris (Moray, 

Goldentail) 0.0 0.0 NS 0.0 0.0 NS

Dasyatidae: Dasyatis 

americana (Stingray, 

Southern) 0.0 0.0 NS 0.0 0.0 NS

161 
Trophic 

Guild Species

EFGB WFGB

Number/100 m2 Number/100 m2

2009 2010 P-value 2009 2010 P-value

Labridae: Halichoeres 

radiatus (Pudding wife) 0.0 0.0 NS 0.0 0.0 NS

Labridae: Halichoeres 

bivittatus (Wrasse, 

Slippery Dick) 0.0 0.0 NS 0.0 0.2 NS

Tetraodontidae: 

Sphoeroides spengleri 

(Puffer, Bandtail) 0.0 0.0 NS 0.0 0.0 NS

Chaetodontidae: 

Chaetodon striatus 

(Butterflyfish, Banded) 0.0 0.0 NS 0.0 0.0 NS

Balistidae: Balistes vetula

(Triggerfish, Queen) 0.0 0.0 NS 0.0 0.0 NS

Serranidae: Liopropoma 

rubre (Bass, Peppermint) 0.0 0.0 NS 0.0 0.0 NS

Holocentridae: 

holocentrus adscensionis 

(Squirrelfish) 0.0 0.0 NS 0.0 0.0 NS

Serranidae: Cephalopholis 

fulva (Grouper, Coney) 0.0 0.0 NS 0.0 0.0 NS

Planktivores

All Planktivores 80.2 25.0 43.5 38.5

Serranidae: Paranthias 

furcifer (Creolefish) 21.8 17.2 NS 13.4 25.6 NS

Pomacentridae: Chromis 

cyanea (Chromis, Blue) 12.4 0.9 <0.0001 10.6 2.9 0.0023

Labridae: Clepticus parrae

(Wrasse, Creole) 40.3 6.6 0.0002 12.8 7.9 0.0008

Pomacentridae: Chromis 

scotti (Reeffish, Purple) 3.3 0.5 0.0238 5.7 1.8 0.007

Pomacentridae: Chromis 

insolata (Sunshinefish) 2.5 0.2 0.008 1.0 0.3 0.0141

Echeneidae: Echeneidae

sp. (Remora) 0.0 0.0 NS 0.0 0.0 NS

Lutjanidae: Lutjanus 

argentiventris (Snapper, 

Yellowtail) 0.0 0.0 NS 0.0 0.0 NS

162 
Size-frequency distributions were graphed for each trophic guild by bank. Herbivores, 

invertivores, and planktivores were dominated by smaller individuals, whereas piscivores 

were substantially larger individuals (See Figure 10.4.1 and Figure 10.4.2).

(A) (B)

(C) (D)

Blue columns are 2009 data; red columns are 2010 data.

Figure 10.4.1. WFGB size distribution by trophic guild. (A) herbivores, (B) invertivores, (C) 

piscivores, and (D) planktivores. 

163 
(A) (B)

(C) (D) 

Blue columns are 2009 data; red columns are 2010 data.

Figure 10.4.2. EFGB size distribution by trophic guild. (A) herbivores , (B) invertivores, (C) 

piscivores, and (D) planktivores. 

164 
10.4.5. Biomass Analysis

Mean biomass for the 2009 and 2010 surveys was calculated to be 113.28 g per 1 m2

(±195.68 SD). The mean biomass ranged from 73 g/m2 (EFGB in 2010) to 162.52 g/m2

(EFGB in 2009) (Table 10.4.14). No significant difference was observed in biomass 

between bank or year when evaluated with a Student’s t-test. 

Table 10.4.14. 

Visual fish Survey Sampling Biomass Statistics for the EFGB and WFGB in 2009 and 2010

Category

2009 2010

EFGB WFGB EFGB WFGB

Total Biomass in All Surveys (g/m2) 689199 572273 322479 400455

Mean Biomass (g/m2) 162.52 134.94 73.00 87.17

When grouped by family, chubs (Kyphosidae) contributed the most biomass to the EFGB 

surveys in 2009 comprising 44.2% of the total biomass. This was primarily due to a large 

school of 200 chub recorded during one survey. Similar to the exclusion of rare manta ray 

sightings, it is reasonable to exclude other rare occurrences that strongly skew results.

When this record was removed, chub fell to 21.2% of the total biomass, and the family 

contributing most to the 2009 EFGB surveys was the groupers and seabass (Serranidae) 

at 33.6%. At the WFGB in 2009, the jack family (Carangidae) contributed 47.3% of the 

biomass. This again was dominantly due to one large school (100 rainbow runner) 

recorded during one survey. When this record was removed, the jack family (Carangidae) 

biomass contribution fell to 5.9%, and the grouper and seabass family (Serranidae) 

became the dominant contributor, with 24.9% of biomass. The data from both the EFGB 

and WFGB in 2010 was comparatively free of anomalous data, and the groupers and 

seabass (Serranidae) contributed most to biomass, at 34.8% (EFGB) and 44.0% (WFGB) 

(Table 10.4.15). 

Table 10.4.15. 

Total Biomass (g/m2) for Each Family 

In parentheses are the percent contributions of each family to the total biomass of the 

bank per year. *The EFGB 2009 data shows biomass with the removal of one record of a 

large school of chub (Kyphosidae). **WFGB 2009 data shows biomass with the removal 

of one record of a large school of rainbow runners from the jack family (Carangidae).

Family

Total Biomass (g/m2) 

2009 2010

EFGB* WFGB** EFGB WFGB

Acanthuridae 98.64 (3.6) 138.68 (7.7) 88.82 (4.9) 93.36 (4.1)

Atherinopsidae -- -- -- --

Apogonidae 14.28 (0.5) 0 (0) -- --

165 
Family

Total Biomass (g/m2) 

2009 2010

EFGB* WFGB** EFGB WFGB

Aulostomidae -- 0.34 (0) -- --

Balistidae 56.57 (2.1) 166.08 (9.2) 98.96 (5.4) 98.4 (4.3)

Blenniidae 0.23 (0) 0.47 (0) -- 0.07 (0)

Carangidae 380.16 (13.8) 106.02 (5.9) 57.79 (3.2) 96.09 (4.2)

Chaetodontidae 8.78 (0.3) 12.45 (0.7) 5.72 (0.3) 9.93 (0.4)

Cirrhitidae 0.02 (0) 0.03 (0) 0 (0) 0.04 (0)

Dasyatidae -- 43.47 (2.4) -- --

Diodontidae -- 1.41 (0.1) -- 1.41 (0.1)

Echeneidae 0.84 (0) -- -- --

Gobiidae 0 (0) 0 (0) 0.14 (0) 0 (0)

Holocentridae -- 3.76 (0.2) 1.16 (0.1) 1.55 (0.1)

Inermiidae 44.28 (1.6) 0.01 (0) -- --

Kyphosidae 583.82 (21.2) 69.12 (3.8) 251.78 (13.8) 96.7 (4.3)

Labridae 179.14 (6.5) 258.75 (14.3) 48.11 (2.6) 39.26 (1.7)

Lutjanidae 30.45 (1.1) 22.65 (1.3) 12.12 (0.7) 83.58 (3.7)

Monacanthidae 1.84 (0.1) 8.2 (0.5) 5.9 (0.3) --

Mullidae 1.23 (0) 10.72 (0.6) 1.88 (0.1) --

Muraenidae 26.83 (1) 26.88 (1.5) -- 0.43 (0)

Ostraciidae 4.21 (0.2) 19.11 (1.1) 7.92 (0.4) 0.9 (0)

Pomacanthidae 65.63 (2.4) 44.69 (2.5) 53.3 (2.9) 18.44 (0.8)

Pomacentridae 134.44 (4.9) 97.68 (5.4) 69.66 (3.8) 60.17 (2.7)

Scaridae 153.22 (5.6) 240.35 (13.3) 329.7 (18.1) 207.45 (9.2)

Serranidae 924.2 (33.6) 451.17 (24.9) 634.53 (34.8) 996.86 (44.0)

Sphyraenidae 42.99 (1.6) 85.36 (4.7) 153.56 (8.4) 459.71 (20.3)

Tetraodontidae 1.95 (0.1) 2.11 (0.1) 3.94 (0.2) 1.94 (0.1)

Although biomass was dominated by the grouper and seabass family (Serranidae) at both 

banks in 2009 and 2010 (when observations of large schools or large rays are removed), a 

single species, Creolefish (Paranthias furcifer), contributes 92.06% ± 3.65 of the biomass 

for the family, ranging from 86.84% at the WFGB in 2010 to 95.18% at the EFGB in 

2010. 

Biomass was grouped by trophic guild for each year and bank. The contribution of each 

trophic guild to total biomass varied between year and bank, with herbivores comprising 

the greatest biomass at the EFGB in both 2009 and 2010 (Figure 10.4.3). A significantly 

higher difference was observed between the piscivores (α=0.005, P-value=0.0074) at the 

WFGB and a significantly higher difference was observed for the invertivores (α=0.005, 

P-value=0.0401) at the EFGB in 2010. No significant difference observed in trophic 

guilds between 2009 and 2010 at each bank, except the invertivore guild at the WFGB 

showed a significantly lower difference in biomass in 2010 (Table 10.4.16). 

166 
The interaction of bank and year on trophic guild biomass was analyzed using a two-way 

ANOVA (Table 10.4.17). The herbivore guild showed no significant difference between 

bank or year, and no significant interaction. The piscivore guild showed no significant 

difference between bank or year, but a significant interaction existed (α=0.005, Pvalue=0.0288). The invertivore guild showed no significant difference in bank, but a 

significant difference between years (α=0.005, P-value=0.0032), with a significiant 

interaction (α=0.005, P-value=0.0151). The planktivore guild showed no significant 

difference between bank, but there was a significant difference between years (α=0.005, 

P-value=0.0231), with no significant interaction. 

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

EFGB WFGB EFGB WFGB

2009 2010

Percent Composition (%)

Bank and Year

Percent Composition of Fish Biomass by Trophic Guild at the 

Flower Garden Banks, 2009-2010

Planktivore Biomass

Invertivore Biomass

Piscivore Biomass

Herbivore Biomass

Figure 10.4.3. Percent composition of biomass for each trophic guild by bank and year.

167 
Table 10.4.16. 

Comparison of Biomass (g/m2) of Herbivores, Piscivores, Invertivores, and Planktivores between 

the EFGB and WFGB in 2009 and 2010 

S = Significant, NS = Not significant

Category

2009 2010

EFGB WFGB P-value Significance EFGB WFGB P-value Significance

Herbivore 

Biomass 84.65 26.15 0.41 NS 30.51 19.60 0.095 NS

Piscivore 

Biomass 24.98 69.56 0.51 NS 9.96 28.83 0.0074 S

Invertivore 

Biomass 9.27 11.12 0.17 NS 7.12 4.49 0.0401 S

Planktivore 

Biomass 42.97 28.12 0.36 NS 25.40 34.25 0.80 NS

168 
Table 10.4.17. 

Two-way ANOVA for Trophic Guild Biomass (g/m2), with Fixed Factors of Bank and Year 

DF=Degrees of freedom, SS=Sum of squares, MS=Mean of squares, 

Sig=Significance; NS= Not significant, and S= Significant

Trophic 

Guild

Variance 

Source DF SS MS Sig.

Pvalue

Herbivore

Model 3 0.688766 0.229589

Bank 1

0.3784985

2 NS 0.0997

Year 1

0.2872255

2 NS 0.1508

Bank x Year 1

0.0111036

3 NS 0.7765

Error 95 13.010032 0.136948

C. Total 98 13.698798

Piscivore

Model 3 2.376345 0.792115

Bank 1 0.4843872 NS 0.2488

Year 1 0.0509231 NS 0.7076

Bank x Year 1 1.7737354 S 0.0288

Error 95 34.177862 0.359767

C. Total 98 36.554207

Invertivore

Model 3 1.767507 0.589169

Bank 1 0.0311178 NS 0.5993

Year 1 1.0212366 S 0.0032

Bank x Year 1 0.6855067 S 0.0151

Error 95 10.638045 0.111979

C. Total 98 12.405553

Planktivore

Model 3 1.954696 0.651565

Bank 1 0.1831716 NS 0.4539

Year 1 1.7283919 S 0.0231

Bank x Year 1 0.0421571 NS 0.7191

Error 95 30.770688 0.323902

C. Total 98 32.725385

Biomass of each species was compared between banks for 2009 and 2010 (Table 

10.4.18). Significant differences were only observed in species from the herbivorous and 

piscivorous fish guilds in 2009, whereas all trophic guilds possessed species with 

significant differences in biomass between banks in 2010. In 2009, the significantly 

different herbivorous fish biomasses include Chub (Kyphosus incisor / Kyphosus 

sectatrix, α=0.005, P-value=0.0043) and Black Durgon (Melichthys niger, α=0.005, Pvalue=0.0186), and the piscivorous fishes included Dog Snapper (Lutjanus jocu, 

α=0.005, P-value=0.0239), Great Barracuda (Sphyraena barracuda, α=0.005, P169 
value=0.0386), and Bonnetmouth (Emmelichthyops atlanticus, α=0.005, Pvalue=0.0489). In 2010, the significantly different herbivorous fish biomasses include BiColor Damselfish (Stegastes partitus, α=0.005, P-value=0.0216), Stoplight Parrotfish 

(Sparisoma viride, α=0.005, P-value=0.0231), the piscivorous Great Barracuda 

(Sphyraena barracuda, α=0.005, P-value=0.0207), the invertivorous Spotfin Butterflyfish 

(Chaetodon ocellatus, α=0.005, P-value=0.0363), and the Creole Wrasse (Clepticus 

parrae, α=0.005, P-value=0.0431). 

Table 10.4.18. 

Biomass (g/m2) of Each Species by Trophic Guild with P-values Shown Where Significant 

Differences Occur

NS = Not significant

Trophic 

Guild Species

2009 2010

Biomass g/m2 Biomass g/m2

EFGB WFGB P-value EFGB WFGB P-value

Herbivores

All Herbivores 84.65 26.15 30.51 19.60

Pomacentridae: Stegastes 

variabilis (Damselfish, 

Cocoa) 0.04 0.06 NS 0.10 0.17 NS

Kyphosidae: Kyphosus incisor 

/ Kyphosus sectatrix (Chub) 71.08 2.88 0.0043 10.07 3.72 NS

Scaridae: Scarus vetula 

(Parrotfish, Queen) 3.10 4.61 NS 4.39 3.95 NS

Pomacentridae: Stegastes 

partitus (Damselfish, Bicolor) 0.55 0.27 NS 0.20 0.08 0.0216

Acanthuridae: Acanthurus 

coeruleus (Tang, Blue) 3.79 5.34 NS 3.00 2.97 NS

Atherinopsidae: 

Menidiinae sp. (Silversides) 0.59 0.00 NS 0.00 0.00 NS

Acanthuridae: Acanthurus 

chirurgus (Doctorfish) 0.15 0.29 NS 0.03 0.01 NS

Pomacanthidae: Holacanthus 

ciliaris (Angelfish, Queen) 0.30 0.98 NS 0.00 0.44 NS

Blenniidae: Ophioblennius 

macclurei (Blenny, Redlip) 0.01 0.02 NS 0.00 0.00 NS

Scaridae: Sparisoma 

aurofrenatum (Parrotfish, 

Redband) 0.18 0.36 NS 0.48 0.34 NS

Pomacentridae:

Microspathodon chrysurus 

(Damselfish, Yellowtail) 0.24 0.50 NS 0.27 0.27 NS

Scaridae: Sparisoma viride

(Parrotfish, Stoplight) 2.99 4.96 NS 6.12 2.88 0.0231

Acanthuridae: Acanthurus 

bahianus (Surgeonfish, 

Ocean) 0.18 0.14 NS 0.53 0.62 NS

Balistidae: Melichthys niger

(Durgon, Black) 1.64 6.60 0.0186 3.12 3.78 NS

Scaridae: Scarus taeniopterus 0.11 0.09 NS 2.12 0.80 NS

170 
Trophic 

Guild Species

2009 2010

Biomass g/m2 Biomass g/m2

EFGB WFGB P-value EFGB WFGB P-value

(Parrotfish, Princess)

Pomacentridae: Stegastes 

adustus (Damselfish, Dusky) 0.00 0.02 NS 0.01 0.01 NS

Scaridae: Sparisoma 

atomarium (Parrotfish, 

Greenblotch) 0.00 0.00 NS 0.00 0.00 NS

Gobiidae: Gnatholepis 

thompsoni (Goby, Goldspot) 0.00 0.00 NS 0.00 0.00 NS

Scaridae: Scarus iseri

(Parrotfish, Striped) 0.00 0.00 NS 0.08 0.00 NS

Piscivores

All Piscivores 24.98 69.56 9.96 28.83

Lutjanidae: Lutjanus jocu

(Snapper, Dog) 1.27 0.00 0.0239 0.00 2.39 NS

Serranidae: Mycteroperca 

interstitialis (Grouper, 

Yellowmouth) 0.85 0.24 NS 0.67 0.23 NS

Serranidae: Mycteroperca 

tigris (Grouper, Tiger) 1.44 0.00 NS 0.31 4.10 NS

Carangidae: Caranx latus 

(Jack, Horse-eye) 14.63 3.24 NS 0.00 3.64 NS

Carangidae: Caranx lugubris

(Jack, Black) 0.69 1.07 NS 2.16 0.00 NS

Serranidae: Cephalopholis 

cruentata (Graysby) 0.24 0.53 NS 0.50 0.59 NS

Sphyraenidae: Sphyraena 

barracuda (Barracuda, Great) 1.79 3.56 0.0386 6.14 17.68 0.0207

Serranidae: Mycteroperca 

bonaci (Grouper, Black) 0.14 0.00 NS 0.00 0.13 NS

Serranidae: Mycteroperca 

phenax (Scamp) 0.07 0.14 NS 0.02 0.00 NS

Inermiidae: Emmelichthyops 

atlanticus (Bonnetmouth) 1.85 0.00 0.0489 0.00 0.00 NS

Carangidae: Carangoides 

ruber (Jack, Bar) 0.52 0.10 NS 0.15 0.06 NS

Carangidae: Caranx crysos 

(Blue runner) 0.37 0.35 NS 0.00 0.00 NS

Muraenidae: Gymnothorax 

meleagris (Moray, Spotted) 1.12 1.12 NS 0.00 0.02 NS

Aulostomidae: Aulostomus 

maculatus (Trumpetfish) 0.00 0.01 NS 0.00 0.00 NS

Carangidae: Elagatis 

bipinnulata (Rainbow runner) 0.00 59.20 NS 0.00 0.00 NS

Serranidae: Mycteroperca 

venenosa (Grouper, 

Yellowfin) 0.00 0.00 NS 0.00 0.00 NS

Invertivores

All Invertivores 9.27 11.12 7.12 4.49

Pomacentridae: Chromis 

multilineata (Chromis, 

Brown) 4.04 1.63 NS 1.65 1.23 NS

171 
Trophic 

Guild Species

2009 2010

Biomass g/m2 Biomass g/m2

EFGB WFGB P-value EFGB WFGB P-value

Labridae: Thalassoma 

lucasanum (Wrasse, 

Bluehead) 0.25 0.36 NS 0.21 0.13 NS

Pomacentridae: Stegastes 

planifrons (Damselfish, 

Three-spot) 0.18 0.49 NS 0.20 0.25 NS

Cirrhitidae: Amblycirrhitus 

pinos (Hawkfish, Redspotted) 0.00 0.00 NS 0.00 0.00 NS

Tetraodontidae: Canthigaster 

rostrata (Puffer, Sharpnose) 0.08 0.09 NS 0.16 0.07 NS

Labridae: Bodianus rufus

(Hogfish, Spanish) 0.28 0.61 NS 0.41 0.64 NS

Chaetodontidae: Chaetodon 

ocellatus (Butterflyfish, 

Spotfin) 0.20 0.11 NS 0.00 0.08 0.0363

Pomacanthidae: Holacanthus 

tricolor (Angelfish, Rock 

Beauty) 0.37 0.56 NS 0.09 0.03 NS

Chaetodontidae: Chaetodon 

sedentarius (Butterflyfish, 

Reef) 0.16 0.32 NS 0.20 0.28 NS

Monacanthidae: Cantherhines 

pullus (Filefish, Orange 

Spotted) 0.08 0.07 NS 0.01 0.00 NS

Balistidae: Canthidermis 

sufflamen (Triggerfish, 

Ocean) 0.72 0.32 NS 0.20 0.00 NS

Ostraciidae: Lactophrys 

triqueter (Trunkfish, Smooth) 0.08 0.21 NS 0.13 0.03 NS

Labridae: Halichoeres 

maculipinna (Wrasse, Clown) 0.05 0.02 NS 0.00 0.00 NS

Pomacentridae: Abudefduf 

saxatilis (Sergeant Major) 0.09 0.58 NS 0.03 0.07 NS

Gobiidae: Elacatinus 

oceanops (Goby, Neon) 0.00 0.00 NS 0.01 0.00 NS

Pomacanthidae: Pomacanthus 

paru (Angelfish, French) 2.06 0.32 NS 2.04 0.24 NS

Serranidae: Epinephelus 

adscensionis (Hind, Rock) 0.15 0.00 NS 0.08 0.00 NS

Serranidae: Epinephelus 

guttatus (Hind, Red) 0.01 0.00 NS 0.00 0.00 NS

Labridae: Halichoeres garnoti

(Wrasse, Yellowhead) 0.02 0.08 NS 0.01 0.04 NS

Mullidae: Pseudupeneus 

maculatus (Goatfish, Spotted) 0.05 0.00 NS 0.00 0.00 NS

Ostraciidae: Acanthostracion 

polygonius (Cowfish, 

Honeycomb) 0.09 0.54 NS 0.19 0.00 NS

Chaetodontidae: 

Prognathodes aculeatus 

(Butterflyfish, Longsnout) 0.01 0.09 NS 0.02 0.01 NS

172 
Trophic 

Guild Species

2009 2010

Biomass g/m2 Biomass g/m2

EFGB WFGB P-value EFGB WFGB P-value

Labridae: Bodianus pulchellus 

(Hogfish, Spotfin) 0.00 0.00 NS 0.00 0.00 NS

Ostraciidae: Lactophrys 

bicaudalis (Trunkfish, 

Spotted) 0.00 0.05 NS 0.00 0.00 NS

Lutjanidae: Lutjanus agennes 

(Snapper, Grey) 0.00 0.94 NS 0.48 0.82 NS

Apogonidae: Apogon sp. 

(Cardinal fish sp.) 0.00 0.00 NS 0.00 0.00 NS

Holocentridae: Holocentrus 

rufus (Squirrelfish, 

Longspine) 0.00 0.16 NS 0.05 0.06 NS

Diodontidae: Diodon 

holocanthus (Ballonfish) 0.00 0.06 NS 0.00 0.05 NS

Monacanthidae: Cantherhines 

macrocerus (Filefish, 

Whitespotted) 0.00 0.27 NS 0.23 0.00 NS

Mullidae: Mulloidichthys 

martinicus (Goatfish, Yellow) 0.00 0.45 NS 0.08 0.00 NS

Muraenidae: Gymnothorax 

miliaris (Moray, Goldentail) 0.00 0.00 NS 0.00 0.00 NS

Dasyatidae: Dasyatis 

americana (Stingray, 

Southern) 0.00 1.81 NS 0.00 0.00 NS

Labridae: Halichoeres 

radiatus (Pudding wife) 0.00 0.01 NS 0.01 0.00 NS

Labridae: Halichoeres 

bivittatus (Wrasse, Slippery 

Dick) 0.00 0.00 NS 0.00 0.00 NS

Tetraodontidae: Sphoeroides 

spengleri (Puffer, Bandtail) 0.00 0.00 NS 0.00 0.00 NS

Chaetodontidae: Chaetodon 

striatus (Butterflyfish, 

Banded) 0.00 0.00 NS 0.01 0.01 NS

Balistidae: Balistes vetula

(Triggerfish, Queen) 0.00 0.00 NS 0.63 0.00 NS

Serranidae: Liopropoma rubre

(Bass, Peppermint) 0.00 0.00 NS 0.00 0.00 NS

Holocentridae: holocentrus 

adscensionis (Squirrelfish) 0.00 0.00 NS 0.00 0.00 NS

Serranidae: Cephalopholis 

fulva (Grouper, Coney) 0.00 0.00 NS 0.00 0.00 NS

Planktivores

All Planktivores 42.97 28.12 25.40 34.25

Serranidae: Paranthias 

furcifer (Creolefish) 35.60 17.89 NS 23.80 33.29 NS

Pomacentridae: Chromis 

cyanea (Chromis, Blue) 0.44 0.47 NS 0.30 0.16 NS

Labridae: Clepticus parrae

(Wrasse, Creole) 6.87 9.71 NS 1.28 0.71 0.0431

Pomacentridae: Chromis 0.03 0.03 NS 0.02 0.07 NS

173 
Trophic 

Guild Species

2009 2010

Biomass g/m2 Biomass g/m2

EFGB WFGB P-value EFGB WFGB P-value

scotti (Reeffish, Purple)

Pomacentridae: Chromis 

insolata (Sunshinefish) 0.01 0.01 NS 0.01 0.01 NS

Echeneidae: Echeneidae sp. 

(Remora) 0.03 0.00 NS 0.00 0.00 NS

Lutjanidae: Lutjanus 

argentiventris (Snapper, 

Yellowtail) 0.00 0.00 NS 0.00 0.00 NS

When banks were compared between years (Table 10.4.19), significant differences were

for five herbivores, Cocoa Damselfish (Stegastes variabilis, α=0.005, P-value=0.0387), 

Chub (Kyphosus incisor / Kyphosus sectatrix, α=0.005, P-value=0.0315), Bi-Color 

Damselfish (Stegastes partitus, α=0.005, P-value=0.041), Redlip Blenny (Ophioblennius 

macclurei, α=0.005, P-value=0.0467), and Princess Parrotfish (Scarus taeniopterus, 

α=0.005, P-value=0.0088), and four piscivores Dog Snapper (Lutjanus jocu, α=0.005, Pvalue=0.0239), Horse-Eye Jack (Caranx latus, α=0.005, P-value=0.0086), Great 

Barracuda (Sphyraena barracuda, α=0.005, P-value=0.0029), and Bonnetmouth 

(Emmelichthyops atlanticus, α=0.005, P-value=0.0486). At the WFGB, significant 

difference in biomass between 2009 and 2010 were found for five herbivores, Cocoa 

Damselfish (Stegastes variabilis, α=0.005, P-value=0.0247), Bi-Color Damselfish 

(Stegastes partitus, α=0.005, P-value=0.0008), Redlip Blenny (Ophioblennius macclurei, 

α=0.005, P-value=0.0153), Stoplight Parrotfish (Sparisoma viride, α=0.005, Pvalue=0.0234), and Princess Parrotfish (Scarus taeniopterus, α=0.005, P-value=0.0463), 

three piscivoresTiger Grouper (Mycteroperca tigris, α=0.005, P-value=0.0129), Black 

Jack (Caranx lugubris, α=0.005, P-value=0.006), and Great Barracuda (Sphyraena 

barracuda, α=0.005, P-value=0.0007), two invertivores, Rock Beauty (Holacanthus 

tricolor, α=0.005, P-value=0.0137) and Honeycomb Cowfish (Acanthostracion 

polygonius, α=0.005, P-value=0.0471), and two planktivoresBlue Chromis (Chromis 

cyanea, α=0.005, P-value=0.0308) and Creole Wrasse (Clepticus parrae, α=0.005, Pvalue=0.0003). 

174 
Table 10.4.19. 

Biomass Comparisons Between Years

Significant differences are shown in bold; NS = Not significant

Trophic 

Guild Species

EFGB WFGB

Biomass g/m2 Biomass g/m2

2009 2010 P-value 2009 2010 P-value

Herbivores

All Herbivores 84.65 30.51 26.15 19.60

Pomacentridae: Stegastes variabilis

(Damselfish, Cocoa) 0.04 0.10 0.0387 0.06 0.17 0.0247

Kyphosidae: Kyphosus incisor / 

Kyphosus sectatrix (Chub) 71.08 10.07 0.0315 2.88 3.72 NS

Scaridae: Scarus vetula (Parrotfish, 

Queen) 3.10 4.39 NS 4.61 3.95 NS

Pomacentridae: Stegastes partitus 

(Damselfish, Bi-color) 0.55 0.20 0.041 0.27 0.08 0.0008

Acanthuridae: Acanthurus coeruleus 

(Tang, Blue) 3.79 3.00 NS 5.34 2.97 NS

Atherinopsidae: Menidiinae sp. 

(Silversides) 0.59 0.00 NS 0.00 0.00 NS

Acanthuridae: Acanthurus chirurgus

(Doctorfish) 0.15 0.03 NS 0.29 0.01 NS

Pomacanthidae: Holacanthus ciliaris 

(Angelfish, Queen) 0.30 0.00 NS 0.98 0.44 NS

Blenniidae: Ophioblennius macclurei

(Blenny, Redlip) 0.01 0.00 0.0467 0.02 0.00 0.0153

Scaridae: Sparisoma aurofrenatum

(Parrotfish, Redband) 0.18 0.48 NS 0.36 0.34 NS

Pomacentridae: Microspathodon 

chrysurus (Damselfish, Yellowtail) 0.24 0.27 NS 0.50 0.27 NS

Scaridae: Sparisoma viride

(Parrotfish, Stoplight) 2.99 6.12 NS 4.96 2.88 0.0234

Acanthuridae: Acanthurus bahianus 

(Surgeonfish, Ocean) 0.18 0.53 NS 0.14 0.62 NS

Balistidae: Melichthys niger (Durgon, 

Black) 1.64 3.12 NS 6.60 3.78 NS

Scaridae: Scarus taeniopterus

(Parrotfish, Princess) 0.11 2.12 0.0088 0.09 0.80 0.0463

Pomacentridae: Stegastes adustus

(Damselfish, Dusky) 0.00 0.01 NS 0.02 0.01 NS

Scaridae: Sparisoma atomarium

(Parrotfish, Greenblotch) 0.00 0.00 NS 0.00 0.00 NS

Gobiidae: Gnatholepis thompsoni

(Goby, Goldspot) 0.00 0.00 NS 0.00 0.00 NS

Scaridae: Scarus iseri (Parrotfish, 

Striped) 0.00 0.08 NS 0.00 0.00 NS

Piscivores

All Piscivores 24.98 9.96 69.56 28.83

Lutjanidae: Lutjanus jocu (Snapper, 

Dog) 1.27 0.00 0.0239 0.00 2.39 NS

Serranidae: Mycteroperca 

interstitialis (Grouper, Yellowmouth) 0.85 0.67 NS 0.24 0.23 NS

175 
Trophic 

Guild Species

EFGB WFGB

Biomass g/m2 Biomass g/m2

2009 2010 P-value 2009 2010 P-value

Serranidae: Mycteroperca tigris

(Grouper, Tiger) 1.44 0.31 NS 0.00 4.10 0.0129

Carangidae: Caranx latus (Jack, 

Horse-eye) 14.63 0.00 0.0086 3.24 3.64 NS

Carangidae: Caranx lugubris (Jack, 

Black) 0.69 2.16 NS 1.07 0.00 0.006

Serranidae: Cephalopholis cruentata 

(Graysby) 0.24 0.50 NS 0.53 0.59 NS

Sphyraenidae: Sphyraena barracuda

(Barracuda, Great) 1.79 6.14 0.0029 3.56 17.68 0.0007

Serranidae: Mycteroperca bonaci 

(Grouper, Black) 0.14 0.00 NS 0.00 0.13 NS

Serranidae: Mycteroperca phenax

(Scamp) 0.07 0.02 NS 0.14 0.00 NS

Inermiidae: Emmelichthyops 

atlanticus (Bonnetmouth) 1.85 0.00 0.0486 0.00 0.00 NS

Carangidae: Carangoides ruber (Jack, 

Bar) 0.52 0.15 NS 0.10 0.06 NS

Carangidae: Caranx crysos (Blue 

runner) 0.37 0.00 NS 0.35 0.00 NS

Muraenidae: Gymnothorax meleagris 

(Moray, Spotted) 1.12 0.00 NS 1.12 0.02 NS

Aulostomidae: Aulostomus maculatus 

(Trumpetfish) 0.00 0.00 NS 0.01 0.00 NS

Carangidae: Elagatis bipinnulata 

(Rainbow runner) 0.00 0.00 NS 59.20 0.00 NS

Serranidae: Mycteroperca venenosa 

(Grouper, Yellowfin) 0.00 0.00 NS 0.00 0.00 NS

Invertivores

All Invertivores 9.27 7.12 11.12 4.49

Pomacentridae: Chromis multilineata 

(Chromis, Brown) 4.04 1.65 NS 1.63 1.23 NS

Labridae: Thalassoma lucasanum 

(Wrasse, Bluehead) 0.25 0.21 NS 0.36 0.13 NS

Pomacentridae: Stegastes planifrons 

(Damselfish, Three-spot) 0.18 0.20 NS 0.49 0.25 NS

Cirrhitidae: Amblycirrhitus pinos

(Hawkfish, Redspotted) 0.00 0.00 NS 0.00 0.00 NS

Tetraodontidae: Canthigaster rostrata

(Puffer, Sharpnose) 0.08 0.16 NS 0.09 0.07 NS

Labridae: Bodianus rufus (Hogfish, 

Spanish) 0.28 0.41 NS 0.61 0.64 NS

Chaetodontidae: Chaetodon ocellatus 

(Butterflyfish, Spotfin) 0.20 0.00 NS 0.11 0.08 NS

Pomacanthidae: Holacanthus tricolor 

(Angelfish, Rock Beauty) 0.37 0.09 NS 0.56 0.03 0.0137

Chaetodontidae: Chaetodon 

sedentarius (Butterflyfish, Reef) 0.16 0.20 NS 0.32 0.28 NS

Monacanthidae: Cantherhines pullus

(Filefish, Orange Spotted) 0.08 0.01 NS 0.07 0.00 NS

Balistidae: Canthidermis sufflamen 0.72 0.20 NS 0.32 0.00 NS

176 
Trophic 

Guild Species

EFGB WFGB

Biomass g/m2 Biomass g/m2

2009 2010 P-value 2009 2010 P-value

(Triggerfish, Ocean)

Ostraciidae: Lactophrys triqueter 

(Trunkfish, Smooth) 0.08 0.13 NS 0.21 0.03 NS

Labridae: Halichoeres maculipinna

(Wrasse, Clown) 0.05 0.00 NS 0.02 0.00 NS

Pomacentridae: Abudefduf saxatilis

(Sergeant Major) 0.09 0.03 NS 0.58 0.07 NS

Gobiidae: Elacatinus oceanops 

(Goby, Neon) 0.00 0.01 NS 0.00 0.00 NS

Pomacanthidae: Pomacanthus paru

(Angelfish, French) 2.06 2.04 NS 0.32 0.24 NS

Serranidae: Epinephelus adscensionis 

(Hind, Rock) 0.15 0.08 NS 0.00 0.00 NS

Serranidae: Epinephelus guttatus

(Hind, Red) 0.01 0.00 NS 0.00 0.00 NS

Labridae: Halichoeres garnoti

(Wrasse, Yellowhead) 0.02 0.01 NS 0.08 0.04 NS

Mullidae: Pseudupeneus maculatus 

(Goatfish, Spotted) 0.05 0.00 NS 0.00 0.00 NS

Ostraciidae: Acanthostracion 

polygonius (Cowfish, Honeycomb) 0.09 0.19 NS 0.54 0.00 0.0471

Chaetodontidae: Prognathodes 

aculeatus (Butterflyfish, Longsnout) 0.01 0.02 NS 0.09 0.01 NS

Labridae: Bodianus pulchellus 

(Hogfish, Spotfin) 0.00 0.00 NS 0.00 0.00 NS

Ostraciidae: Lactophrys bicaudalis

(Trunkfish, Spotted) 0.00 0.00 NS 0.05 0.00 NS

Lutjanidae: Lutjanus agennes 

(Snapper, Grey) 0.00 0.48 NS 0.94 0.82 NS

Apogonidae: Apogon sp. (Cardinal 

fish sp.) 0.00 0.00 NS 0.00 0.00 NS

Holocentridae: Holocentrus rufus

(Squirrelfish, Longspine) 0.00 0.05 NS 0.16 0.06 NS

Diodontidae: Diodon holocanthus 

(Ballonfish) 0.00 0.00 NS 0.06 0.05 NS

Monacanthidae: Cantherhines 

macrocerus (Filefish, Whitespotted) 0.00 0.23 NS 0.27 0.00 NS

Mullidae: Mulloidichthys martinicus

(Goatfish, Yellow) 0.00 0.08 NS 0.45 0.00 NS

Muraenidae: Gymnothorax miliaris

(Moray, Goldentail) 0.00 0.00 NS 0.00 0.00 NS

Dasyatidae: Dasyatis americana

(Stingray, Southern) 0.00 0.00 NS 1.81 0.00 NS

Labridae: Halichoeres radiatus 

(Pudding wife) 0.00 0.01 NS 0.01 0.00 NS

Labridae: Halichoeres bivittatus 

(Wrasse, Slippery Dick) 0.00 0.00 NS 0.00 0.00 NS

Tetraodontidae: Sphoeroides 

spengleri (Puffer, Bandtail) 0.00 0.00 NS 0.00 0.00 NS

Chaetodontidae: Chaetodon striatus 0.00 0.01 NS 0.00 0.01 NS

177 
Trophic 

Guild Species

EFGB WFGB

Biomass g/m2 Biomass g/m2

2009 2010 P-value 2009 2010 P-value

(Butterflyfish, Banded)

Balistidae: Balistes vetula

(Triggerfish, Queen) 0.00 0.63 NS 0.00 0.00 NS

Serranidae: Liopropoma rubre (Bass, 

Peppermint) 0.00 0.00 NS 0.00 0.00 NS

Holocentridae: holocentrus 

adscensionis (Squirrelfish) 0.00 0.00 NS 0.00 0.00 NS

Serranidae: Cephalopholis fulva

(Grouper, Coney) 0.00 0.00 NS 0.00 0.00 NS

Planktivores

All Planktivores 42.97 25.40 28.12 34.25

Serranidae: Paranthias furcifer

(Creolefish) 35.60 23.80 NS 17.89 33.29 NS

Pomacentridae: Chromis cyanea

(Chromis, Blue) 0.44 0.30 NS 0.47 0.16 0.0308

Labridae: Clepticus parrae (Wrasse, 

Creole) 6.87 1.28 NS 9.71 0.71 0.0003

Pomacentridae: Chromis scotti

(Reeffish, Purple) 0.03 0.02 NS 0.03 0.07 NS

Pomacentridae: Chromis insolata

(Sunshinefish) 0.01 0.01 NS 0.01 0.01 NS

Echeneidae: Echeneidae sp. (Remora) 0.03 0.00 NS 0.00 0.00 NS

Lutjanidae: Lutjanus argentiventris

(Snapper, Yellowtail) 0.00 0.00 NS 0.00 0.00 NS

10.4.6. Diurnal Abundance Patterns

Previous studies have observed differences in the behaviors of several species of fish in 

the morning versus the afternoon (Zimmer et al. 2010). Three species of interest included 

Great Barracuda (Sphyraena barracuda), Creole Wrasse (Clepticus parrae), and 

Creolefish (Paranthias furcifer) as they may exhibit differences in behaviors throughout 

the day, resulting in disparate data on morning dives compared to afternoon dives. 

Yearly comparisons were made between the abundance of these species between the time 

of day (morning or afternoon) and bank. At the EFGB in 2009, 11 surveys were 

conducted in the morning (Dawn to 12:00 PM) and 13 surveys were conducted in the 

afternoon (12:00 PM to Dusk). At the WFGB in 2009, 6 surveys were conducted in the 

morning and 18 in the afternoon. At the EFGB in 2010, 8 surveys were conducted in the 

morning and 17 in the afternoon. At the WFGB in 2010, 19 surveys were conducted in 

the morning and 8 in the afternoon. No significant differences were found between time 

of surveys for the three species, though a difference was found for Great Barracuda 

(Sphyraena barracuda) between banks in 2009 (Table 10.4.20). 

178 
Table 10.4.20

Two-way ANOVA for Great Barracuda Abundance, with Fixed Factors of Bank and Time of Day

DF=Degrees of freedom, SS=Sum of squares, MS=Mean of squares, Sig=Significant, 

P=Probability, NS=Not significant, and S=Significant

Var.

Source

2009 2010

DF SS MS Sig. Pvalue DF SS MS Sig. Pvalue

Model 3 0.269392 0.089797 3 0.1230975 0.041033

Bank 1 0.161878 S 0.0101 1 0.12219133 NS 0.1692

Time 

of Day 1 0.000642 NS 0.8663 1 0.02584458 NS 0.5239

Bank x 

Time 1 0.031294 NS 0.2434 1 0.0001781 NS 0.9577

Error 44 0.98477 0.022381 48 3.0099847 0.062708

C. 

Total 47 1.254162 51 3.1330823

 

179 
10.4.7. Prevalent Compared with Rare Species

A rank abundance was used to determine and remove “rare” species. Species were 

considered “rare” if they were recorded in less than 20% of samples. “Prevalent” species

occurred in ≥20% of samples (Figure 10.4.4). A total of 30 species was considered 

“prevalent” using this method (Table 10.4.21). 

0

10

20

30

40

50

60

70

80

90

100

0 10 20 30 40 50 60 70

Prevalence (%)

Species Rank

Ranked abundance curve for the prevalence of all species recorded at the 

Flower Garden Banks, 2009-2010

The dashed line represents the 20% sighting frequency threshold in determining 

"prevalent" compared to "rare" species.

Figure 10.4.4. Ranked abundance curve for the prevalence of all species recorded 

during the 2009 and 2010 surveys. 

180 
Table 10.4.21. 

Ranked Abundance of "Prevalent" Species

Rank Species Trophic 

Guild

Percent

with rare 

removed

1 Serranidae: Paranthias furcifer (Creolefish) PL 95

2 Labridae: Thalassoma lucasanum (Wrasse, Bluehead) I 94

3 Pomacentridae: Chromis multilineata (Chromis, Brown) I 90

4 Pomacentridae: Stegastes partitus (Damselfish, Bi-color) H 90

5 Acanthuridae: Acanthurus coeruleus (Tang, Blue) H 90

6 Tetraodontidae: Canthigaster rostrata (Puffer, Sharpnose) I 88

7 Labridae: Bodianus rufus (Hogfish, Spanish) I 85

8 Pomacentridae: Chromis cyanea (Chromis, Blue) PL 79

9 Scaridae: Scarus vetula (Parrotfish, Queen) H 76

10 Labridae: Clepticus parrae (Wrasse, Creole) PL 74

11 Pomacentridae: Stegastes planifrons (Damselfish, Three-spot) I 67

12 Sphyraenidae: Sphyraena barracuda (Barracuda, Great) P 62

13 Scaridae: Sparisoma viride (Parrotfish, Stoplight) H 59

14 Balistidae: Melichthys niger (Durgon, Black) H 54

15 Pomacentridae: Stegastes variabilis (Damselfish, Cocoa) H 52

16 Kyphosidae: Kyphosus incisor / Kyphosus sectatrix (Chub) H 51

17 Chaetodontidae: Chaetodon sedentarius (Butterflyfish, Reef) I 51

18 Pomacentridae: Chromis scotti (Reeffish, Purple) PL 47

19 Serranidae: Cephalopholis cruentata (Graysby) P 46

20

Pomacentridae: Microspathodon chrysurus (Damselfish, 

Yellowtail) H 41

21 Scaridae: Sparisoma aurofrenatum (Parrotfish, Redband) H 34

22 Pomacentridae: Chromis insolata (Sunshinefish) PL 32

23 Acanthuridae: Acanthurus bahianus (Surgeonfish, Ocean) H 27

24 Blenniidae: Ophioblennius macclurei (Blenny, Redlip) H 25

25

Pomacanthidae: Holacanthus tricolor (Angelfish, Rock 

Beauty) I 25

26

Serranidae: Mycteroperca interstitialis (Grouper, 

Yellowmouth) P 24

27 Ostraciidae: Lactophrys triqueter (Trunkfish, Smooth) I 24

28 Pomacentridae: Abudefduf saxatilis (Sergeant Major) I 21

29 Scaridae: Scarus taeniopterus (Parrotfish, Princess) H 20

30 Labridae: Halichoeres garnoti (Wrasse, Yellowhead) I 20

181 
10.5. FISH SURVEYS DISCUSSION

Perhaps two of the most important factors that shape the fish assemblages at the FGB are: 

(1) they occur near the northern latitudinal limit of coral reefs and are remote from other 

tropical reefs and (2) they occur in close proximity to offshore hydrocarbon production 

platforms. As a remote outpost of Caribbean coral reef biota, the fish assemblage has

been reported to be low in diversity yet high in biomass (Pattengill-Semmens and

Gittings 2003; Zimmer et al. 2010). They also differ from fish assemblages in other reef

and hard bottom systems in the Gulf of Mexico and Caribbean in that the have limited

representation by lutjanids and haemulids (Rooker et al. 1997). The large number of oil 

and gas production platforms in the Gulf of Mexico, as well as the addition of mooring 

buoys at the banks in 1990, may have promoted the dispersal of additional fish species

and allowed some to reach the FGB and establish themselves (Boland et al. 1983; 

Rooker et al. 1997; Gittings 1998). 

Fishing pressure, water quality (including temperature and planktonic composition), 

and current flow patterns also affect fish assemblages at the FGB to varying degrees. 

Since the late 1800’s, fishermen have conducted long-line fishing around the Flower 

Gardens (Scarborough-Bull 1988). Since 1992, commercial fishing with bottom longlines, traps, nets, and bottom trawls are prohibited within the sanctuary’s boundaries.

Although hand-line and hook and line fishing, including bandit reels (powered reels), are 

allowed within the boundaries, the distance from shore does reduce fishing pressure to 

some extent, therefore providing some protection to fish populations. 

The stationary fish surveys conducted in 2009 and 2010 revealed a thriving reef fish 

assemblage, as observed in previous annual monitoring surveys (Precht et al. 2006; 

Zimmer et al. 2010) and in other reef-fish surveys conducted at the FGB (Rooker et al. 

1997). But numerous other approaches have been used over the years to collect fish data 

from the Flower Garden Banks. 

 

Extensive surveys conducted in 1980–1982 by Boland et al. (1983) used, among 

other things, remote video to collect data on fish assemblages. A decade later, when the 

Reef Environmental Education Foundation began surveys at the Flower Garden Banks, 

the roving diver technique (RDT) was used (Pattengill-Semmens and Gittings 2003). 

Rooker et al. (1997) and Pattengill et al. (1998) were the first to use the stationary visual 

census technique of Bohnsack and Bannerot (1986) at the Flower Gardens. 

Each of the techniques employed have strengths and weaknesses. Comparisons of visual

census techniques (e.g., Bortone et al. 1989) have revealed, for example, that the RDT

produces good data on diversity and sighting frequency, but is not as well suited as

stationary surveys for recording fish counts, sizes, or densities. 

The FGB fish populations on the reef cap were dominated by the families

Pomacentridae, Labridae, Serranidae, and Scaridae. This was also found in previous 

monitoring (Precht et al. 2006; Zimmer et al. 2010). The pomacentrids were the most 

182 
diverse, with 10 species. The labrids and serranids were represented by a moderate

number of species. Creolefish (Paranthias furcifer) were one of the most abundant 

species on the reefs, along with large schools of creole wrasse (Clepticus parrae). 

Similar to other Caribbean coral reef communities, blue and brown chromis (Chromis 

cyanea and C. multilineata) were commonly seen in schools above coral formations. 

Also as expected in Caribbean communities, groups of female or male bluehead wrasse 

(Thalassoma bifasciatum) in intermediate or juvenile phases were regularly seen on 

diver surveys in low areas between and just above coral formations, with one or two

males in close association. Although lower in number because of their solitary nature, 

Caribbean sharp-nose puffer (Canthigaster rostrata) actively explored crevices and low 

areas. The redlip blenny, (Ophioblennius atlanticus), was commonly seen perched on 

the face of coral formations. 

Some species were likely underestimated because of the techniques used. Many blenny 

and goby species, for example, are small and cryptic. The stationary counting

technique used here would underestimate their abundance and perhaps the number of

species. Furthermore, surveys also intentionally excluded sand-covered bottom areas, so 

diver surveys were less likely to have recorded species associated with that habitat.

While invertivores were the dominant fish guild at the Flower Garden Banks, a healthy 

assemblage of herbivorous fishes was recorded. Herbivore populations appear to have 

responded to the drastic decline in Diadema antillarum at the FGB, which occurred in 

1983 and 1984 (Gittings et al. 1992). As a group, acanthurids, scarids, and 

Microspathodon chrysurus were relatively high in density at both banks in 2009 and 

2010. While the FGB has lower species richness (including fewer scarid species) and a 

lower overall abundance of herbivorous fishes than Caribbean reefs (Rezak et al. 1985; 

Dennis and Bright 1988), the percentages of acanthurids and scarids is similar to

deep/fore reefs of far western Cuba and Akumal, Yucatan, Mexico (Table 10.5.1.; Claro 

and Cantelar Ramos 2003; Steneck and Lang 2003). The low algal cover reported here, 

despite the lack of an actively grazing Diadema antillarum population, may be the 

result of this apparently adaptive community of herbivores. Finally, algal farming

pomacentrids were abundant as well. There were more of these gardeners recorded in 

2009 than 2010 at the FGB. 

While the species richness of reef-associated carnivores (certain serranids and lutjanids) 

was high, density values were somewhat low. The density of reef associated carnivores 

appears depressed in relation to previous studies. It is not possible, at this point, to 

determine whether this is due to recent fishing pressure (which is allowed within the 

FGBNMS by hook and line only). 

183 
Table 10.5.1.

Percentage of Fishes Observed in the Listed Families at Reefs around the Gulf of Mexico and the

Caribbean Region

Combined herbivore percentages shown at bottom 

Family

Maria La Gorda,

Cuba

Akumal, Yucatan,

Mexico

FGBNMS, USA

Acanthuridae 17% 22% 15%

Balistidae 37% 0% 9%

Chaetodontidae 7% 5% 9%

Lutjanidae 5% 7% 1%

Pomacanthidae 4% 2% 4%

Scaridae 25% 58% 24%

Serranidae 6% 6% 8%

Acanthuridae and

Scaridae 42% 80% 39%

(Claro and Cantelar Ramos 2003; Steneck and Lang 2003; Precht et al. 2006)

184 
CHAPTER 11.0: SEA URCHIN AND LOBSTER SURVEYS

11.1. SEA URCHIN AND LOBSTER SURVEYS METHODOLOGICAL RATIONALE

The long-spined sea urchin, Diadema antillarum, was an important herbivore on coral reefs 

throughout the Caribbean until 1983 and 1984. At that time, an unknown pathogen 

decimated populations throughout the region, including the FGBNMS. Since then, patchy 

but limited recovery has been documented in the region (Edmunds and Carpenter 2001).

Diadema antillarum populations at the FGBNMS pre-1984 exceeded 1 individual/m2

(hindcast surveys were made from archived transect photos taken during daytime hours, 

which would underestimate densities; Gittings et al. 1992; Aronson et al. 2005). 

Lobsters are commercially important species throughout much of the Caribbean and Gulf of 

Mexico; however, population dynamics of Caribbean spiny lobster (Panulirus argus) in the 

FGBNMS are not well understood.

11.2. SEA URCHIN AND LOBSTER SURVEYS FIELD METHODS 

Due to the nocturnal nature of these species, visual surveys were conducted at night, a 

minimum of 1.5 hours after sunset. In 2009 and 2010, surveys for Diadema antillarum

(long-spined sea urchin), Panulirus argus (Caribbean spiny lobster) and P. guttatus

(spotted spiny lobster) were conducted along the northern and eastern perimeter lines at 

the EFGB and along the southern and western boundaries at the WFGB. Two belt 

transects 2 m wide and 100 m long were surveyed by diver teams on each bank, thus

totaling 400 m

2

per bank each year. Surveys began with the northeast corner at the EFGB 

study site and the southeast corner at the WFGB study site. All observed species of sea 

urchin and lobster were recorded.

Due to low sample abundance, only qualitative analyses were possible for the lobster and 

sea urchin surveys. 

11.3. SEA URCHIN AND LOBSTER SURVEYS RESULTS

The number of individuals recorded during each survey in 2009 and 2010 are listed in 

Table 11.3.1. No lobsters were observed on surveys at either bank in 2009 or 2010. At the 

EFGB in 2009, one Diadema antillarum (0.25 per 100 m2) was documented, and in 2010,

two individuals (0.5 per 100 m2) were documented. This correlates to very low densities

of Diadema antillarum at the EFGB in 2009 (0.25 individuals/100 m2) and 2010 (0.5 

individuals/100 m2). 

Considerably more Diadema antillarum were found at the WFGB during this monitoring 

period. In 2009, 55 Diadema antillarum (13.75 per 100 m2) were documented. In 2010, 

44 (11.0 per 100 m2) were found. This correlates to higher densities at the WFGB in 

2009 (13.75 individuals/100 m2) and 2010 (11.0 individuals/100 m2) (Figure 11.3.1). 

185 
 

Table 11.3.1.

Number of Individual Sea Urchins and Lobsters Observed During Surveys in 2009 and 2010

No. of Individuals 

Observed

Sea 

Urchins Lobsters

Diadema 

antillarum

Panulirus 

argus

Panulirus 

guttatus

EFGB 2009 1 0 0

EFGB 2010 2 0 0

WFGB 2009 55 0 0

WFGB 2010 44 0 0

Figure 11.3.1. Sea urchin and lobster counts conducted at the EFGB and WFGB in 2009 

and 2010.

186 
11.4. SEA URCHIN AND LOBSTER SURVEYS DISCUSSION

Since 2004, lobster counts on surveys have ranged from zero to two. They are, however, 

occasionally encountered by divers at other times, so they do occur on the banks in low 

abundance (Figure 11.4.1). 

After the mass die off in 1983, D. antillarum populations have not recovered to pre-1983 

levels, which were at least 140 individuals/100 m2 at the EFGB and 50 individuals/100 m2 

at the WFGB (Gittings et al. 1998). Post-1984 sea urchin densities dropped to near zero

(Gittings and Bright 1987). Sea urchin populations at the EFGB remained low during this 

monitoring period and were similar to those reported in previous studies (Zimmer et al. 

2010). Populations at the WFGB have been consistently higher than the EFGB. Both

2009 and 2010 had abundances among the higest recorded at the WFGB since the die off

(Figure 11.4.1). The previous fluctuations in annual density estimates suggest caution in 

declaring a recovering sea urchin population on the bank; continued monitoring will be 

required to track and compare temporal changes at both banks. 

Data for 2004 to 2008 from PBS&J (Precht et al. 2006, 2008b); and FGBNMS for 2009 

and 2010 (Johnston et al. 2012).

Figure 11.4.1. Sea urchin and lobster densities at the FGB from 2004 to 2010. 

187 
CHAPTER 12.0: RECOMMENDATIONS

Throughout this report, a number of deficiencies in existing protocols were mentioned, 

and numerous adjustments would improve the efficiency and quality of data collection 

efforts. The following are recommendations for improving the monitoring protocols and 

increasing the scientific value of the monitoring program. 

Operational Changes

Below are recommendations that should be considered as immediate improvements to the 

monitoring program. Most are recommended because they have been identified as 

deficiencies of the current protocols. 

• Install HOBO data loggers at multiple depths.

• Install mid-transect eye bolts along the study site perimeter to improve perimeter 

video repeatability so that lines will not shift with the current. 

• Upload data and service/exchange the Sea-Bird water quality monitoring 

equipment more frequently (5–8 times per year) to obtain more consistent and 

accurate results for water quality parameters.

• Improve lateral station methodology and mapping to remove comparison 

inconsistencies from year to year. Establish new stations and extend cruise days to 

allow time for this task. 

• Add scale bar to T-frame to more accurately measure year-to-year change in 

colony size. 

• Standardize camera, lens, and pole length on T-frame so that photos are always 

take from the same height above the reef at every station.

• Monitor previously-identified exotic/invasive species on the reefs of the FGB,

including Tubastraea coccinea, and Thecacera pacifica. Monitor for newly

arriving exotic/invasive species, such as Pterois volitans/miles. This will help 

support management decisions regarding removal programs for exotic or invasive 

species.

• Obtain additional measurements of the coral heads sampled for sclerochronology,

such as width and height, as well as depth of samples.

• Use multiple fish survey techniques, including belt transect, roving methods, and

stationary surveys to obtain more comprehensive data, particularly improving data 

on richness, cryptic species, and fish biomass. 

189 
• Improve methods to detect new species records and range extensions to the banks 

(e.g., Acropora palmata). Obtain estimate of tissue loss or growth of Acropora 

palmata for basic condition analyses. 

• Future monitoring efforts should include a review of fish biomass and trophic 

structure at the FGB from 1999 (or earlier if data available) to the present. This

will prove to be a useful resource status evaluation tool and help support 

management decisions. 

• Increase the number of lobster and sea urchin surveys instead of the same single 

transect every year (perhaps conduct them concurrently while doing random 

transects photos) and note all lobster and urchin species observed during all 

diving operations. 

Experimental Evaluations

The following recommendations may not be critical immediate needs, but represent areas 

that should be investigated as potential improvements to the monitoring program.

• Install deep stations at the WFGB to compare to deep station coral cover on the 

EFGB.

• Add survey for coral size class distribution and for coral recruits.

• Additional species other than barracuda and creole wrasse may exhibit 

differences in abundance throughout the day, resulting in disparate data on 

morning dives compared to afternoon dives. Diver surveys should be evenly

distributed through out daylight hours over a period of two days. More 

importantly, sampling should be done at the same times throughout the sampling 

effort. This will allow for the inclusion of changing fish behavior and to test

differences between the different sampling times. 

• During sea urchin surveys, note the color of the sea urchin spines (e.g., black or 

white) as part of data collection protocol in order to evaluate anecdotal reports of 

greater survival among the white-spined morphotype. 

 

Additional Recommendations 

• Continue and increase the number of presentations and peer-reviewed publications 

resulting from this work. 

190 
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The Department of the Interior Mission

As the Nation’s principal conservation agency, the Department of the Interior has 

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major responsibility for American Indian reservation communities and for people 

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The Bureau of Ocean Energy Management (BOEM) works to manage the 

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