OCS Study 

MMS 2009-032 

 

Post-Hurricane Assessment 

of Sensitive Habitats 

of the Flower Garden Banks Vicinity 

U.S. Department of the Interior

Minerals Management Service 

Gulf of Mexico OCS Region 
U.S. Department of the Interior 

Minerals Management Service 

Gulf of Mexico OCS Region 

OCS Study 

MMS 2009-032 

Post-Hurricane Assessment 

of Sensitive Habitats 

of the Flower Garden Banks Vicinity 

Authors 

Martha L. Robbart 

Richard B. Aronson 

Kenneth J.P. Deslarzes 

William F. Precht 

Leslie Duncan 

Beth Zimmer 

Todd DeMunda 

Prepared under MMS Contract 

1435-01-06-CT-39574 (M06PC00003) 

by 

PBS&J 

2001 NW 107th Avenue 

Miami, Florida 33172 

Published by 

New Orleans

July 2009 
iii 

DISCLAIMER 

This report was prepared under contract between the Minerals Management Service (MMS), the 

National Oceanic and Atmospheric Administration (NOAA), and PBS&J Ecological Services, 

Miami, Florida, USA. This report has been technically reviewed by MMS and NOAA and has 

been approved for publication. Approval does not signify that the contents necessarily reflect the 

views and policies of MMS and NOAA, nor does the mention of trade names or commercial 

products constitute endorsement or recommendation for use. It is, however, exempt from review 

and compliance with MMS editorial standards. 

REPORT AVAILABILITY 

Extra copies of this report may be obtained from the Public Information Office (Mail Stop 5034) 

at the following address:

U.S. Department of the Interior 

Minerals Management Service 

Gulf of Mexico OCS Region 

Public Information Office (MS 5034) 

1201 Elmwood Park Boulevard 

New Orleans, Louisiana 70123-2394 

Telephone: (504) 736-2519 or 

 1-800-200-GULF 

CITATION 

Suggested citation: 

Robbart, M.L., R.B. Aronson, K.J.P. Deslarzes, W.F. Precht, L. Duncan, B. Zimmer, and T. 

DeMunda. 2009. Post-hurricane assessment of sensitive habitats of the Flower Garden 

Banks vicinity. U.S. Dept. of the Interior, Minerals Management Service, Gulf of 

Mexico OCS Region, New Orleans, Louisiana. OCS Study MMS 2009-032. 160 pp. 

ABOUT THE COVER 

Montage of photographs from Sonnier, McGrail, Geyer, Bright, and East Flower Garden Banks 

created by Martha Robbart, PBS&J. 
v 

TABLE OF CONTENTS 

Page

LIST OF FIGURES.................................................................................................................... vii

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

EXECUTIVE SUMMARY ....................................................................................................... xiii

1.0 INTRODUCTION .................................................................................................................. 1

1.1. HURRICANES IN THE NORTHWESTERN GULF OF MEXICO ..............................................................1 

1.2. STUDY BANKS IN THE NORTHWESTERN GULF OF MEXICO............................................................6 

1.2.1. General Bank Information.......................................................................................................6 

1.2.2. Sonnier Bank...............................................................................................................................7 

1.2.3. McGrail Bank..............................................................................................................................7

1.2.4. Geyer Bank ..................................................................................................................................7 

1.2.5. Bright Bank................................................................................................................................12 

1.2.6. East Flower Garden Bank......................................................................................................12 

2.0 METHODS........................................................................................................................... 15

2.1. VIDEO TRANSECT METHOD .................................................................................................................15 

2.1.1. Methodological Rationale......................................................................................................15 

2.1.2. Field Methods............................................................................................................................15 

2.1.3. Laboratory Analysis................................................................................................................17 

2.2. ROVING DIVER METHOD ......................................................................................................................19 

2.2.1. Methodological Rationale......................................................................................................19 

2.2.2. Field Methods............................................................................................................................19 

2.2.3. Laboratory Analysis................................................................................................................19 

2.3. ROV TRANSECT METHOD....................................................................................................................20 

2.3.1. Methodological Rationale......................................................................................................20 

2.3.2. Field Methods............................................................................................................................20 

2.3.3. ROV Laboratory Analysis.....................................................................................................21 

2.4. TRANSECT STATISTICAL ANALYSIS METHODOLOGY....................................................................25 

2.5. REPETITIVE QUADRATS.........................................................................................................................26 

2.5.1. Methodological Rationale......................................................................................................26 

2.5.2. Field Methods............................................................................................................................26 

2.5.3. Image Analysis of Repetitive Quadrats.............................................................................26 

2.5.4. Data Presentation and Statistical Analysis of Repetitive Quadrats...........................27 

2.6. PERIMETER VIDEOGRAPHY .................................................................................................................27 

2.6.1. Methodological Rationale......................................................................................................27 

2.6.2. Field Methods............................................................................................................................27 

2.6.3. Laboratory Methods................................................................................................................27 

2.7. HYDRODYNAMIC MODEL ANALYSIS METHODS .............................................................................28 

3.0 RESULTS ............................................................................................................................. 31

3.1. TRANSECT DATA DESCRIPTIVE STATISTICS ....................................................................................31 

3.2. SONNIER BANK........................................................................................................................................32 

3.3. MCGRAIL BANK .....................................................................................................................................37 
vi 

3.4. GEYER BANK ...........................................................................................................................................45 

3.5. BRIGHT BANK..........................................................................................................................................50 

3.6. SHANNON-WIENER DIVERSITY INDEX (H’).....................................................................................53 

3.7. MULTIVARIATE STATISTICAL RESULTS ............................................................................................53 

3.7.1. ANOSIM Analysis...................................................................................................................53 

3.7.2. MDS Analysis...........................................................................................................................55 

3.8. EAST FLOWER GARDEN BANK REPETITIVE QUADRAT DATA.....................................................57 

3.8.1. Study Site Repetitive Quadrat Percent Cover Data........................................................57 

3.8.2. Study Site Repetitive Quadrat Planimetry........................................................................60 

3.8.3. Deep Station Repetitive Quadrat Percent Cover Data...................................................61 

3.9. PERIMETER VIDEOGRAPHY ..................................................................................................................62 

3.9.1. Perimeter Lines.........................................................................................................................63 

3.9.2. East Flower Garden Bank 360° Panoramic Views.........................................................63 

3.10. RESULTS OF HYDRODYNAMIC MODEL PREDICTIONS .................................................................65 

4.0 DISCUSSION........................................................................................................................ 71

4.1. HURRICANE EFFECTS.............................................................................................................................71

4.1.1. Beneficial Aspects of Hurricanes on Coral Reefs..........................................................71 

4.1.2. Hurricane Effects at Banks Not Dominated by Corals .................................................72 

4.1.3. Hydrodynamic Modeling Predictions................................................................................73 

4.2. BENTHIC CHARACTERISTICS AND HURRICANE EFFECTS AT SONNIER, MCGRAIL, GEYER 

AND BRIGHT BANKS ..............................................................................................................................74 

4.2.1. Sonnier Bank.............................................................................................................................76

4.2.2. McGrail, Geyer, Bright, and East Flower Garden Banks.............................................77 

5.0 CONCLUSION .................................................................................................................... 81

6.0 LITERATURE CITED ....................................................................................................... 83

 

APPENDICES 

Appendix 1: List of Species...........................................................................................................89 

Appendix 2: Transect Data............................................................................................................95 

Appendix 3: Repetitive Quadrat Percent Cover Data..................................................................127 

Appendix 4: Repetitive Quadrat Planimetry Data.......................................................................129 
vii 

LIST OF FIGURES 

Page

Figure 1.1.1. Track of Hurricane Rita, September 18-26, 2005, in relation to the banks 

evaluated in this study (National Hurricane Center 2007). ...................................... 2 

Figure 1.1.2. Map displaying the probability of hurricane force winds on September 22, 

2005. ......................................................................................................................... 3 

Figure 1.1.3. (A) Sediment-scouring, (B) footprint of dislodged coral colony, and (C) coral 

colony gouged by waterborne object(s).................................................................... 5 

Figure 1.2.2. Bathymetric map of Sonnier Bank, showing location of the study site (USGS 

2003)......................................................................................................................... 8 

Figure 1.2.3. Map of the peak at Sonnier Bank (22-m depth) sampled during this study 

(courtesy of Greg Boland through Gulf Reef Environmental Action Team and 

MMS)........................................................................................................................ 9 

Figure 1.2.4. Bathymetric map of McGrail Bank, showing location of the study site (USGS 

2003)....................................................................................................................... 10 

Figure 1.2.5. Bathymetric map of Geyer Bank, showing location of the study site (USGS 

2003)....................................................................................................................... 11 

Figure 1.2.6. Bathymetric map of Bright Bank, showing location of the study site (USGS 

2003)....................................................................................................................... 13 

Figure 1.2.7. Bathymetric map of the EFGB, showing location of the study site (USGS 

2003)....................................................................................................................... 14 

Figure 2.1.2. Approximation of diver tracks, showing geographic positions of video 

transects at Sonnier Bank from 22-27 m and from 30-36.5 m. .............................. 18 

Figure 2.1.3. Diagram illustrating the method used to create transects from continuous 

video footage taken along contour lines at 30- to 36.5-m depth at Sonnier, 

Geyer, and Bright Banks. ....................................................................................... 20 

Figure 2.3.2. Seabotix LBV 300S-62 ROV at Sonnier Bank on May 22, 2007........................... 21 

Figure 2.3.3. Map of ROV track at Sonnier Bank to and from the ship to a maximum depth 

of ~60-m. ................................................................................................................ 22 

Figure 2.3.4. Map of ROV track at Geyer Bank to and from the ship to a maximum depth of 

~60-m...................................................................................................................... 23 

Figure 2.3.5. Map of ROV track at McGrail Bank to and from the ship to a maximum depth 

of ~60-m. ................................................................................................................ 24 

Figure 3.1.1. Percent live cover at Sonnier, Geyer, McGrail, and Bright Banks in all depth 

ranges. .................................................................................................................... 31 

Figure 3.2.1. (A) Xestospongia muta from 1996 video (NOAA 1996) and (B) X. muta from 

2002 video (Boland 2002). ..................................................................................... 33 

Figure 3.2.2. Percent cover of benthic categories at all depth ranges at Sonnier Bank............... 34 
viii 

Figure 3.2.3. Large rubble typical of 30- to 36.5-m depth range at Sonnier Bank...................... 36 

Figure 3.3.1. Percent cover of benthic categories in both depth ranges at McGrail Bank. ......... 40 

Figure 3.3.2. Frame capture from ROV video at McGrail Bank, showing a large colony of 

the coral Stephanocoenia intersepta. ..................................................................... 42 

Figure 3.3.3. Two-dimensional MDS plot, based on square-root-transformed Bray-Curtis 

similarities, comparing multivariate cover of the four hard-coral taxa at 

McGrail Bank within the 45- to 50-m depth range. ............................................... 43 

Figure 3.3.4. Frame capture from ROV video at McGrail Bank, showing algal nodules at 

55- to 60-m depth. .................................................................................................. 43 

Figure 3.4.1. Percent cover of benthic categories in all depth ranges at Geyer Bank. ................ 45 

Figure 3.4.2. Frame capture of Sargassum spp., typical benthic cover in the 30- to 36.5-m 

depth range at Geyer Bank. .................................................................................... 46 

Figure 3.5.1. Captured frame from video recorded at Bright Bank in September 2003 at 

36-m depth. ............................................................................................................ 50 

Figure 3.5.2. Percent cover of benthic categories at Bright Bank. .............................................. 51 

Figure 3.5.3. Frame capture showing diverse benthic cover at Bright Bank in 30- to 36.5-m 

depth. ...................................................................................................................... 51 

Figure 3.7.2. Two-dimensional MDS plot, based on square-root-transformed Bray-Curtis 

similarities, comparing multivariate benthic cover data among Sonnier, 

McGrail, Geyer, and Bright Banks at the depth ranges sampled. .......................... 56 

Figure 3.7.3. Two-dimensional MDS plot based on square-root-transformed Bray-Curtis 

similarity comparing multivariate benthic cover data between Sonnier, 

McGrail, Geyer, and Bright Banks by transect. ..................................................... 57 

Figure 3.8.1. (A) Photograph of repetitive quadrat station in June 2005 at the EFGB. (B) 

Photograph of matching repetitive quadrat station in November 2005.................. 58 

Figure 3.8.2. (A) Photograph of repetitive quadrat station in June 2005 at the EFGB. (B) 

Photograph of matching repetitive quadrat station in November 2005.................. 58 

Figure 3.8.3. (A) Photograph of repetitive quadrat station in June 2005 at the EFGB. (B) 

Photograph of matching repetitive quadrat station in November 2005.................. 59 

Figure 3.8.4. Percent cover of coral-condition categories. ......................................................... 60 

Figure 3.8.5. Mean proportional change in the planar tissue area of Montastraea annularis

species complex within repetitive quadrat stations at the EFGB between June 

2005 and November 2005, and between November 2005 and June 2006. ........... 61 

Figure 3.8.6. Coral condition at the EFGB deep stations in November 2005 and June 2006. ... 63 

Figure 3.10.1. Predicted wave velocities at Sonnier Bank during passage of Hurricane Rita..... 67 

Figure 3.10.2. Predicted wave velocities during passage of Hurricane Rita at (A) McGrail 

Bank, (B) Geyer Bank, (C) Bright Bank, and (D) EFGB. .................................... 68 
ix 

Figure 4.1.1. Typical bioerosion pattern of Diploria strigosa, resulting in mushroom-shaped 

colony at the EFGB. . ............................................................................................. 72 

Figure 4.1.2. Water temperature at the Flower Garden Banks in 2005 and mean temperature 

at the EFGB from 1990 to 2005. ............................................................................ 73 

Figure 4.2.1. Location of Sonnier, McGrail, Geyer, Bright, and East Flower Garden Banks 

in the Gulf of Mexico. ........................................................................................... 75 

Figure 4.2.2. Total coral cover at each of four banks within the study, displayed from west 

to east. .................................................................................................................... 76 

Figure 4.2.3. Percent cover of functional groups from repetitive quadrat stations at the 

EFGB in June 2005, November 2005, and June 2006. .......................................... 79 

Figure 4.2.4. Percent cover of the dominant coral taxa at the EFGB repetitive quadrat 

stations in June 2005, November 2005, and June 2006. ........................................ 80 
xi 

LIST OF TABLES 

Page

Table 1.1.1. List of tropical cyclones that entered the Gulf of Mexico (GOM) in 2004 and 

2005. .......................................................................................................................... 4 

Table 1.1.2. Shortest distance between the path of Hurricane Rita and each of the five banks 

studied........................................................................................................................ 6 

Table 1.2.1. Biological zones by depth (in meters) (adapted from Rezak et al. 1985 and 

Hickerson et al. 2008)................................................................................................ 6 

Table 2.1.1. Biological zonation of benthic communities by depth at Sonnier, McGrail, 

Geyer, and Bright Banks with corresponding depths from this study (adapted 

from Rezak et al. 1985)............................................................................................ 16 

Table 2.1.2. Summary of data collected by diver and ROV during this study. ........................... 16 

Table 2.6.1. Wave conditions during Hurricane Rita as hindcast by Oceanweather Inc. 

(2006)....................................................................................................................... 29 

Table 3.1.1. Results of one-way ANOVA and Tukey HSD test performed on square root 

transformed live cover estimates from diver and ROV transects. ........................... 32 

Table 3.2.1. Percent cover of benthic categories at Sonnier Bank from 22- to 27-m depth........ 34 

Table 3.2.2. Percent cover of benthic categories at Sonnier Bank from 30- to 36.5-m depth..... 36 

Table 3.2.3. Percent cover of benthic categories at Sonnier Bank from 45- to 50-m depth........ 38 

Table 3.2.4. Percent cover of benthic categories at Sonnier Bank from 55- to 60-m depth........ 39 

Table 3.3.1. Percent cover of benthic categories at McGrail Bank from 45- to 50-m depth....... 41 

Table 3.3.2. Percent cover of benthic categories at McGrail Bank from 55- to 60-m depth....... 44 

Table 3.4.1. Percent cover of benthic categories at Geyer Bank from 30- to 36.5-m depth........ 46 

Table 3.4.2. Percent cover of benthic categories at Geyer Bank from 45- to 50-m depth........... 48 

Table 3.4.3. Percent cover of benthic categories at Geyer Bank from 55- to 60-m depth........... 49 

Table 3.5.1. Percent cover of benthic categories at Bright Bank from 30- to 36.5-m depth. ...... 52 

Table 3.6.1. Shannon-Wiener Diversities (H') at all banks and depths surveyed........................ 53 

Table 3.7.1. Results of Pairwise ANOSIM tests with site as the factor. ..................................... 54 

Table 3.7.2. Results of Pairwise ANOSIM tests with depth as the factor. .................................. 54 

Table 3.7.3. Results of Pairwise ANOSIM tests for differences between sites across all 

depths....................................................................................................................... 55 

Table 3.7.4. Results of pairwise ANOSIM tests for differences between depths across all 

sites. ......................................................................................................................... 55 
xii 

Table 3.8.1. Percent cover by coral species of isolated fish biting (IFB), concentrated fish 

biting (CFB), paling (P), and bleaching (BL) at the EFGB in November 2005 

and June 2006. ......................................................................................................... 59 

Table 3.8.2. Percent cover (±SE) of paling, bleaching, concentrated fish biting, isolated fish 

biting, and disease at the EFGB in November 2005 and June 2006........................ 60 

Table 3.8.3. Area of coral colonies missing in November 2005 photographs at the EFGB........ 62 

Table 3.9.1. Comparison of observations of the condition of individual coral colonies at the 

EFGB along perimeter lines and in 360o panoramic views between November 

2005 (n = 815 colonies) and June 2006 (n = 736 colonies)..................................... 64 

Table 3.9.2. Affected coral colonies at the northwest corner of the study site in November 

2005 and June 2006. ................................................................................................ 65 

Table 3.9.3. Affected coral colonies at the northeast corner of the study site in November 

2005 and June 2006. ................................................................................................ 66 

Table 3.9.4. Affected coral colonies at the southeast corner in November 2005 and June 

2006. ........................................................................................................................ 66 

Table 3.10.1. Maximum bottom velocities found using stream wave theory.............................. 70 

Table 4.1.3. Results of the hydrodynamic modeling effort in relation to bank depth and 

distance from the Hurricane Rita storm track.......................................................... 74 

 
xiii 

EXECUTIVE SUMMARY 

The most active hurricane season on record in the Atlantic and Gulf of Mexico occurred in 2005, 

fueled by higher than normal sea-surface temperatures (National Climatic Data Center 2005). 

Eleven tropical cyclones entered the Gulf of Mexico in 2005, including Hurricane Rita (a 

Category 5 storm on the Saffir-Simpson Scale). Hurricane Rita was a Category 3 storm when it 

passed near the shelf edge banks on September 23, 2005 and produced wind speeds of 

126.7-138.3 mph (110-120 knots). Hurricane force winds extended 86-mi (139-km) and tropical 

storm force winds extended as far as 207-mi (333-km) from the center of the storm (National 

Hurricane Center 2007). 

Several sensitive habitats within the northwestern Gulf of Mexico were close to the path of 

Hurricane Rita, including Sonnier Bank (24-km or 15-mi east), McGrail Bank (12-km or 7-mi 

west), Geyer Bank (58-km or 36-mi west), Bright Bank (75-km or 47-mi west) and the East 

Flower Garden Bank (EFGB) (93-km or 58-mi west). Hindcast hydrodynamic models estimated 

wave heights at ~66-ft (20-m) or higher on these banks. This may have exposed some bank 

caps, even at 20- to 30-m depth. The implications for benthic community structure could have 

been catastrophic, and MMS considered it essential to characterize the banks in their posthurricane state. In order to understand the potential wave and current conditions present in the 

vicinity of Sonnier, McGrail, Geyer, Bright, and East Flower Garden Banks during the passage 

of Hurricane Rita, PBS&J conducted a wave hindcast study. On the cap at Sonnier Bank (22-m 

or ~72-ft depth) and Bright Bank (33-m or ~108-ft depth), significant wave heights during the 

passage of Hurricane Rita were estimated at ~43-ft (13-m) and ~66-ft (20-m), respectively. 

Greater wave heights may have occurred at banks with caps located in deeper water, regardless 

of the bank’s distance from the storm track. Thus, McGrail and Geyer Banks, which have caps 

located at deeper depths, may theoretically have experienced larger wave heights than Bright and 

Sonnier Banks. These larger wave height conditions were hindcast based on the potential effects 

of wave focusing, shoaling, and breaking. The estimated wave height at both McGrail and Geyer 

Banks was ~85-ft (26-m). The maximum hindcast bottom velocity was 4-m/s or ~8-kn for all 

four banks. 

This study characterized and compared the benthic habitats of four banks (Sonnier, McGrail, 

Geyer, and Bright) and recorded possible hurricane damage at these banks and the EFGB. At 

Sonnier, McGrail, Geyer, and Bright Banks, videographic records were collected by SCUBA and 

ROV in April and May 2007, at four depth ranges to assess benthic cover to the lowest possible 

taxonomic level: 22-27 m, 30-36.5 m, 45-50 m, and 55-60 m (72-89 ft, 98-120 ft, 148-164 ft, and 

180-197 ft). Video transects were qualitatively assessed for evidence of hurricane damage. To 

document recovery from Hurricane Rita at the existing long-term monitoring site on the EFGB, 

repetitive quadrats and perimeter line surveys were conducted in November 2005 and compared 

to data collected subsequently in June 2006. 

Sonnier Bank, the only bank located east of the storm track, exhibited the least live cover at all 

depth ranges (~2-38%) when compared to McGrail, Geyer, and Bright Banks (~17-86%). 

Qualitative analysis of video footage collected by divers at Sonnier Bank in 1996, 2002, and 

2005 showed differences in benthic cover compared to video collected in 2007. In previous 
xiv 

years, more live cover of a mix of algae and sponges (predominantly Neofibularia sp. and Ircinia 

spp.) was obvious. Another notable difference was the apparent disappearance of Xestospongia 

muta colonies, which were present in 1996 (~50 colonies), declined to one individual in 2002, 

and then were not recorded in 2005 or 2007. The single colony recorded in 2002 exhibited 

disease-like characteristics, with discoloration and an eroded pinacoderm. 

McGrail Bank was approximately 7-mi (12-km) from the track of Hurricane Rita and has the 

deepest reef cap of any bank in this study (45-m or 148-ft depth). Live cover at McGrail Bank 

ranged from 17-38% and was dominated by macroalgae (red, green and brown), nodules of red 

coralline algae, and coral (predominately Stephanocoenia intersepta). Previous videos were not 

available for qualitative analysis. No apparent hurricane damage, in the form of overturned or 

injured corals, was observed. 

Live cover at Geyer Bank was mostly colonized by brown macroalgae (specifically Sargassum 

spp.), corals, and sponges. Brown macroalgae, the dominant component, ranged from 22-42%. 

Corals (0-10%) and sponges (1-4%) were less prominent. According to transect video, Geyer 

Bank is the only bank in this study with an established population of the invasive, nonhermatypic coral Tubastraea coccinea. It is important to note that scientists have observed one 

or two colonies of Tubastraea coccinea at Sonnier Bank (Hickerson, personal communication, 

2008); however, T. coccinea colonies were not observed in our Sonnier Bank transect video. 

Diver video collected at Geyer Bank in 2003 showed similar benthic cover to the video that was 

recorded in 2007. Again, no obvious signs of hurricane damage were observed. 

Bright Bank exhibited the highest live cover (86%) and was dominated by macroalgae (57%), 

turf algae (12%), and corals (8%). Coral species observed included Diploria strigosa, Millepora 

alcicornis, Montastraea cavernosa, and Stephanocoenia intersepta. Diver video taken in 

September 2003 from Bright Bank revealed mostly bare substrate, low macroalgal cover, and 

few large colonies of Millepora alcicornis and Diploria strigosa. Sponge species were also 

observed, including Agelas clathrodes, Ircinia spp., and Xestospongia muta. At Bright Bank no 

hurricane damage was observed. 

The Shannon Wiener Diversity Index (H') was calculated for each bank at each depth range 

using the lowest taxonomic groupings possible (species or genus). The highest diversity of these 

four banks was at Sonnier Bank from 22-27 m (H' = 2.86), largely due to the variety of sponges 

present there. Geyer and McGrail Banks exhibited their highest diversity values in the 45- to 

50-m depth range, (H' = 2.13 and H' = 2.08, respectively). The high species richness of brown 

macroalgae accounted for these high diversity values. At Bright Bank H' = 1.81 in the 30- to 

36.5-m depth range. 

Multivariate statistical analyses were performed using benthic cover data at Sonnier, McGrail, 

Geyer, and Bright Banks. Analysis of Similarity (ANOSIM) tests showed significant differences 

between banks (Global R = 0.54, P = 0.001). Within site comparisons showed less dissimilarity 

between depths. Multidimensional scaling (MDS) highlighted the dissimilarities among banks, 

with depths within sites grouping more closely. 
xv 

Repetitive quadrat stations were photographed at the EFGB in November 2005 and June 2006 to 

document recovery from Hurricane Rita. Coral cover remained consistently high from November 

2005 to June 2006, at 61.34% ± 2.75 SE and 62.87% ± 2.32 SE, respectively. Macroalgae 

increased from November 2005 to June 2006 by ~5.4%, while CTB decreased by ~6.9%. 

Approximately 1.5% of coral colonies photographed within repetitive quadrats at the EFGB were 

missing in November 2005, most likely due to the effects of Hurricane Rita. However, this did 

not notably affect estimates of coral cover. The most obvious difference in November 2005 

repetitive quadrats was the high level of bleaching: 9.74% ± 1.07 SE of assessed coral points 

were bleached. Paling and fish biting measurements were low at 1.5% or less. The June 2006 

data showed a decrease in the level of bleaching (0.62%) compared to November 2005 and a 

slight increase in the amount of fish biting (2.31%). Coral disease was not observed in the June 

2006 repetitive quadrat photographs, although the identification of disease in these photographs 

is not reliable because of the 2-m distance from the substrate (Zimmer, personal communication, 

2005). 

The perimeter lines around the EFGB study site were videotaped in November 2005 and June 

2006 to document change at known locations along the perimeter and within the study site. 

Lower levels of bleaching and paling were seen in June 2006 compared to November 2005. 

These qualitative observations corroborate the quantitative results from the repetitive quadrat 

data. 

The unique biological characteristics of the benthic communities of Sonnier, McGrail, Geyer, 

Bright, and East Flower Garden Banks highlight their intrinsic value within the northwestern 

Gulf of Mexico ecosystem. The differences in the benthic biotas strongly suggest that these 

habitats are truly sensitive, because nearest neighbors may not be the source of recruitment. 

With predicted wave velocities of 8 knots or more acting on these banks during the passage of 

Hurricane Rita, the effects on the benthic communities could have been catastrophic. Sonnier 

Bank suffered a loss of benthic cover and hardbottom associated with the hurricane, but the 

community was recovering, with algae and sponges dominating live areas by April/May of 2007. 

McGrail, Geyer, and Bright Banks did not exhibit any obvious hurricane damage in the surveys; 

given that these banks are dominated by algae and sponges, any damage to the living benthos 

may be hard to detect after the twenty months between the passage of the storm and the survey. 

McGrail Bank, with its large Stephanocoenia intersepta colonies, exhibited no apparent damage 

to corals, which may have been protected by their considerable depth (45-m or 148-ft). 
1 

1.0 INTRODUCTION 

1.1. HURRICANES IN THE NORTHWESTERN GULF OF MEXICO

During the 20th century, numerous tropical storms and hurricanes passed over sensitive marine 

habitats in the Gulf of Mexico, including the Flower Garden Banks and other mid- and outershelf banks (Scholten and Deslarzes 1998). Strong hurricanes moved through the region of the 

Flower Garden Banks in 1900, 1909, 1915, 1957, 1961 (Category 4), 1964, 1974, 1979, and 

1980 (Category 5). In the summer of 1980, Hurricane Allen, with a surge of more than 16-ft (5-

m) and wind speeds of 60-mph (96-km/hr), caused physical damage to the coral reefs of the 

Flower Garden Banks. An example of this severe damage is the documented displacement of an 

~4409-lb (2000-kg) coral head which was moved hundreds of feet across the reef cap on the East 

Flower Garden Bank (EFGB) (C.L. Combs, personal communication, 1989).

 

The 2005 Atlantic hurricane season was the most active on record, fueled by record high seasurface temperatures in the Atlantic and Gulf of Mexico (National Climatic Data Center 2005). 

Eleven tropical cyclones entered the Gulf of Mexico in 2005. One of these cyclones, Hurricane 

Rita, passed within 58-mi (93-km) of the EFGB on September 23 as a Category 3 hurricane. 

Hurricane Rita passed 7-mi (12-km) to the east of McGrail Bank (reef cap at 45-m or ~148-ft) 

and approximately 15-mi (24-km) to the west of Sonnier Bank (reef cap at 22-m or ~72-ft) 

(Figure 1.1.1). On September 23 Hurricane Rita had wind speeds of 127-138 mph (110-120 kn). 

Hurricane force winds extended approximately 86-mi (139-km) from its center (Figure 1.1.2) 

and tropical storm force winds extended as far as 207-mi (333-km) from its center (National 

Hurricane Center 2007). Other storms in 2004 and 2005 passed more than ~311-mi (500-km) 

away from the vicinity of the Flower Garden Banks (Table 1.1.1). 

Preliminary assessments of the coral reefs at the EFGB following Hurricane Rita included 

evidence of substantial mechanical impacts, fractured and displaced corals, and sedimentscoured corals bordering sand flats (Figure 1.1.3A, B), as well as corals scarred and indented by 

waterborne objects (Figure 1.1.3C) (Hickerson and Schmahl, personal communication, 2005). 

As an example of physical damage that occurred, a large coral head, measuring ~13-ft (4-m) 

across and ~7-ft (2-m) high, was dislodged from the reef cap. Approximately one meter of 

coarse sand was removed from large sand flats. Large barrel sponges (Xestospongia muta) were 

partially or fully removed from the reef or were filled with sand (Hickerson and Schmahl, 

personal communication, 2005). In a survey in November 2005, many coral colonies were 

missing from permanent quadrats at the EFGB long-term monitoring site (Precht et al. 2008a). 

Hurricane impacts were preceded and followed by other natural disturbances at the Flower 

Garden Banks including extreme warming of the water column from August through November 

2005, which was associated with severe coral bleaching (Hickerson and Schmahl, personal 

communication, 2005), and runoff transported from the Texas-Louisiana coast to the shelf edge 

in the days and weeks following Hurricanes Katrina and Rita (NOAA CoastWatch 2005). In 

addition to its direct physical impacts, the passage of Hurricane Rita lowered sea temperatures 

(by increasing vertical mixing) and likely attenuated the coral bleaching event at the Flower 

Garden Banks. Thus, despite their physical destruction, hurricanes may also exert a positive 

influence on coral reefs (Manzello et al. 2007). 
2 

Figure 1.1.1. Track of Hurricane Rita, September 18-26, 2005, in relation to the banks evaluated 

in this study (National Hurricane Center 2007). 
3 

Figure 1.1.2. Map displaying the probability of hurricane force winds on September 22, 2005. 
4 

Table 1.1.1. 

List of tropical cyclones that entered the Gulf of Mexico (GOM) in 2004 and 2005. 

The name and category of a given cyclone is listed according to its condition when it was closest 

to the outer-shelf banks. Information on the cyclones was taken from The Weather 

Underground, Inc. (2005a and 2005b). 

Name/Category Date 

Wind 

Speed 

(mph) or 

Hurricane 

Category 

Trajectory 

Tropical Storm Bonnie 8/10/04 50 Central GOM, ~373-mi (600-km) east of 

the Flower Garden Banks 

Hurricane Charley 8/13/04 Cat 4 Passed over the Florida Straits, 

southwest Florida 

Tropical Storm Frances 9/4/04 65 Northeast GOM, >621-mi (>1000-km) 

east of the Flower Garden Banks 

Hurricane Ivan 9/15/04 Cat 4 Passed ~342-mi (550-km) east of the 

Flower Garden Banks 

Tropical Storm Matthew 10/09/04 40 Passed ~199-mi (320-km) east of the 

Flower Garden Banks 

Tropical Storm Arlene 6/11/05 70 Central GOM, ~373-mi (600-km) east of 

the Flower Garden Banks 

Tropical Storm Bret 6/25/05 40 Southwest GOM, >497-mi (>800-km) 

south of the Flower Garden Banks 

Tropical Storm Cindy 7/5/05 70 Central GOM, ~311-mi (500-km) east of 

the Flower Garden Banks 

Hurricane Dennis 7/10/05 Cat 4 Central GOM, ~497-mi (800-km) east of 

the Flower Garden Banks 

Hurricane Emily 7/19/05 Cat 1 Southwest GOM, ~373-mi (600-km) 

south of the Flower Garden Banks 

Tropical Storm Gert 7/25/05 45 Southwest GOM, ~466-mi (750-km) 

south of the Flower Garden Banks 

Tropical Storm Jose 8/23/05 50 Southwest GOM, ~497-mi (800-km) 

south of the Flower Garden Banks 

Hurricane Katrina 8/28/05 Cat 5 Central GOM, ~311-mi (500-km) east of 

the Flower Garden Banks 

Hurricane Rita 9/23/05 Cat 3 Central GOM, ~56-mi (90-km) west of 

the Flower Garden Banks 

Tropical Storm Stan 10/3/05 40 Southwest GOM, ~528-mi (850-km) 

south of the Flower Garden Banks 

Hurricane Wilma 10/24/05 Cat 3 Southeast GOM, ~559-mi (900-km) 

southeast of the Flower Garden Banks 
5 

 

(C) 

 

Figure 1.1.3. (A) Sediment-scouring, (B) footprint of dislodged coral colony, and (C) coral 

colony gouged by waterborne object(s). Photos courtesy of Emma L. Hickerson, 

Flower Garden Banks National Marine Sanctuary. 

This study serves as 1) a baseline characterization of the benthic communities at Sonnier, 

McGrail, Geyer, and Bright Banks and 2) an assessment of possible hurricane damage at 

multiple depth ranges at these banks and at the EFGB long-term monitoring site. Sonnier, 

McGrail, Geyer, and Bright Banks were all within ~47-mi (75-km) of the passage of Hurricane 

Rita (Table 1.1.2) and have not been thoroughly characterized in terms of their benthic 

composition since the early 1980s. In order to understand how wind, wave, and current 

conditions may have acted on these topographic features during the passage of Hurricane Rita, 

hydrodynamic hindcast models were run using data from a study that modeled the effects of 

Hurricanes Katrina and Rita for the Gulf of Mexico (Oceanweather Incorporated 2006). Data 

from a November 2005 survey at the EFGB long-term monitoring site are compared to data 

collected in June 2006 at the same site. These comparisons were made to document possible 

recovery 9 months after the storm. 

(A) (B) 
6 

Table 1.1.2. 

Shortest distance between the path of Hurricane Rita and each 

of the five banks studied. 

Bank Distance (km) 

McGrail 12 

Sonnier 24 

Geyer 58 

Bright 75 

East 93 

1.2. STUDY BANKS IN THE NORTHWESTERN GULF OF MEXICO

1.2.1. General Bank Information 

Banks in the Gulf of Mexico vary in terms of their geological and biological characteristics, but 

have been grouped into three main categories: mid-shelf banks, outer-shelf banks with carbonate 

reef caps, and reefs growing on relict carbonate shelf (Rezak et al. 1985). The banks in this 

study fall within the first two categories. Mid-shelf banks and outer-shelf carbonate reef caps are 

formed by salt diapir structures uplifting the seafloor into the photic zone. Mid-shelf banks, such 

as Sonnier Bank, are characterized by uncolonized open substrate, in which the bedding of 

Tertiary limestones, claystones, and siltstones is apparent (Rezak et al. 1985). Major 

components of biological communities capping mid-shelf banks include the hydrocoral 

Millepora alcicornis, the sponge Neofibularia nolitangere, and macroalgae. At deeper depths 

algal/sponge communities characterize the mid-shelf banks (Table 1.2.1). McGrail, Geyer, 

Bright, and East Flower Garden Banks are outer-shelf banks with carbonate reef caps. 

Algal/sponge communities dominate the substrate at outer-shelf banks, with the exception of the 

Flower Garden Banks which are characterized by low diversity coral assemblages (Table 1.2.1) 

(Rezak et al. 1985). The benthic communities at these banks have not been studied in detail, 

with the exception of the Flower Garden Banks. 

Table 1.2.1. 

Biological zones by depth (in meters) 

(adapted from Rezak et al. 1985 and Hickerson et al. 2008). 

“–” represents that the zone is not present 

 Coral Reef Zone Coral Community Zone Coralline Algae Zone 

Bank 

Montastraea 

habitat Madracis Stephanocoenia Millepora/Sponge Algal/Sponge 

Sonnier – – – 18-52 52-74 

McGrail – – 45-47 – 45-82 

Geyer – – – 37-52 60-98 

Bright – – 37 – 52-74 

EFGB 15-36 28-44 36-52 – 45-90 
7 

1.2.2. Sonnier Bank 

Sonnier Bank is located at 28°20’N and 92°27’W and is the only bank in this study to lie east of 

Hurricane Rita’s path (Figure 1.1.1). At Sonnier Bank, several peaks are arranged in an arcuate 

pattern and rise from a depth of ~197-ft (60-m) up to within ~66-ft (20-m) of the sea-surface 

(Figure 1.2.2). Each peak is a fault block created by the collapse of a salt diapir (Rezak et al. 

1985). There is approximately 11,000-m2

 of bank surface area between 22-30 m before the bank 

drops off to deeper depths. Some preliminary mapping and fish community studies have been 

conducted at Sonnier Bank (Boland 1999) (Figure 1.2.3). The peak visited during this study is 

steeply sloping, with its shallowest depth at ~72-ft (22-m). 

The benthic community of Sonnier Bank is similar to other mid-shelf banks, especially Stetson 

Bank (Rezak et al. 1985). Sonnier Bank is characterized by hydrocorals and sponges, with 

Millepora alcicornis, Neofibularia nolitangere, and Ircinia spp. as the dominant living benthic 

components (Rezak et al. 1985). Nine species of scleractinian corals are found at Sonnier Bank, 

including Agaricia spp., Madracis spp., Siderastrea sp., Stephanocoenia intersepta, and 

Montastraea cavernosa; however, these corals are not abundant (Rezak and Bright 1979; Irion 

and Anuskiewicz 1999). 

1.2.3. McGrail Bank 

McGrail Bank, located at 27°57’N and 92°35’W, is a deep shelf-edge carbonate bank positioned 

atop a salt diapir. McGrail Bank has the deepest peak of any bank in this study (45-m or 148-ft) 

(Figure 1.2.4). Deep-water coral reef resources are characteristic of this bank, and coral cover is 

high (up to 30%) compared to other banks in the region (Weaver et al. 2005; Schmahl and 

Hickerson 2006). Dominant corals include Stephanocoenia intersepta, Diploria strigosa, 

Montastraea cavernosa, and Agaricia spp. (Schmahl and Hickerson 2006). Rezak et al. (1985) 

documented abundant nodules of red coralline algae as well as high cover of fleshy macroalgae 

in the algal-sponge zone at depths of ~148-154 ft (45-47 m). 

1.2.4. Geyer Bank 

Geyer Bank, located at 27°51’N and 93°04’W, is a shelf-edge carbonate bank with extensive reef 

communities (Rezak and Bright 1981). Geyer Bank is the southernmost bank surveyed during 

this study. The shallowest peaks (~128-ft or 39-m), all of which are steeply sloping, are located 

in the northern area of the bank (Figure 1.2.5). Corals include Stephanocoenia intersepta, 

Agaricia spp., Madracis spp., and Millepora alcicornis (Hickerson 2004). Geyer Bank is 

characterized by an abundance of Sargassum spp. and appeared to be the only bank with an 

established population of the invasive, non-hermatypic coral Tubastraea coccinea. It is 

important to note that scientists have observed one or two colonies of Tubastraea coccinea at 

Sonnier Bank (Hickerson, personal communication, 2008); however, T. coccinea colonies were 

not observed in our Sonnier Bank transect video. Substantial populations of the echinoid 

Diadema antillarum and large oceanic fish species have also been reported (Hickerson, personal 

communication, 2007). 
8 

Figure 1.2.2. Bathymetric map of Sonnier Bank, showing location of the study site (USDO
9 

Figure 1.2.3. Map of the peak at Sonnier Bank (22-m depth) sampled during this study 

(courtesy of Greg Boland through Gulf Reef Environmental Action Team and 

MMS). 
10 

Figure 1.2.4. Bathymetric map of McGrail Bank, showing location of the study site 

Note: The high of 32-m depicted in the figure legend is due to an outliebathymetry data. The true high or top of bank is located at 45-m. 
11 

Figure 1.2.5. Bathymetric map of Geyer Bank, showing location of the study site (USDOI, GS 

2003). 
12 

1.2.5. Bright Bank 

Bright Bank is located at 27°53’N and 93°18’W (Figure 1.2.6) and is 108-ft (33-m) at its 

shallowest depth. Bright Bank is dominated by sponges, macroalgae, and several species of 

scleractinian corals. Bare siltstone mounds interrupt the flat, coral cap of Bright Bank. Corals 

include Agaricia spp., Madracis spp., Stephanocoenia intersepta, and Montastraea cavernosa 

(Hickerson 2004). Recent bathymetric work has revealed a mud volcano and hydrocarbon seeps 

at depth (Schmahl et al. 2003). For at least the past 30 years, treasure salvage activities have 

taken place at Bright Bank, resulting in the destruction of many coral colonies (Hickerson, 

personal communication, 2007). 

1.2.6. East Flower Garden Bank 

The shallowest portion of the EFGB is located at 59-ft (18-m) and consists of a well-developed 

coral reef cap located at 27°54’N and 93°36’W (Figure 1.2.7). The reef cap at the EFGB is well 

documented and characterized by a high coral cover, low-diversity scleractinian assemblage. 

Diversity is lower than Caribbean reefs but higher than other banks in the area. The EFGB 

supports a robust reef-fish population, with regular visitation by larger pelagic species. 

An annual monitoring cruise was conducted in June 2005 at the East and West Flower Garden 

Banks. A post-hurricane assessment was performed five months later in November 2005 at the 

EFGB to assess damages that may have been caused by Hurricane Rita. Data collected in June 

2005 and November 2005 at the EFGB revealed that coral cover within the established long-term 

monitoring study site remained relatively constant (62.78% ± 2.60 and 61.34% ± 2.75, 

respectively) and species relative abundance showed stability with Montastraea annularis

species complex, Diploria strigosa, and Porites spp. as the dominant species (Precht et al. 

2008a). In June 2005 macroalgae was high at 24%, while CTB was lower at 10%. After the 

hurricane, in November 2005, the inverse relationship between macroalgae and CTB was 

evident, with 24% CTB and 13% macroalgae (Precht et al. 2008a). 

The EFGB contains four biological zones: Coral Reef Zone, Coral Community Zone, Coralline 

Algae Zone, and Deep Coral Zone (Hickerson et al. 2008). The coral cap contains the Coral 

Reef Zone which contains several major habitats that are named based on their dominant coral 

species in the coral assemblage. The primary habitat in the Coral Reef Zone is the Montastraea 

habitat; however, the Zone also contains the Madracis habitat in depths from 92-144 ft (28-44 m) 

and the Stephanocoenia habitat at depths below 118-ft (36-m). The Coral Community Zone, 

which is found at depths of 59-164 ft (18-50 m), is comprised of areas that have low densities of 

hermatypic corals or are characterized by coral reef biota, such as the hydrozoan Millepora spp., 

sponges, and macroalgae. The Coralline Algae Zone occurs at depths of 148-295 ft (45-90 m) 

and is characterized by species of crustose coralline algae that actively produce carbonate 

substrate, including algal nodules. The Deep Coral Zone occurs at depths of 295-ft (90-m) and 

greater, where active photosynthesis is not possible, and is characterized by a diverse assemblage 

of organisms, including antipatharian and gorgonian corals, azooxanthellate branching corals, 

small, solitary hard corals, crinoids, bryozoans, and sponges. 
13 

Figure 1.2.6. Bathymetric map of Bright Bank, showing location of the study site (USDOI, G
14 

Figure 1.2.7. Bathymetric map of the EFGB, showing location of the study site (USDOI, 

GS 2003). 
15 

2.0 METHODS 

2.1. VIDEO TRANSECT METHOD

2.1.1. Methodological Rationale 

This study was designed to serve as (1) a quantitative, baseline evaluation of the benthic 

communities at Sonnier, McGrail, Geyer, and Bright Banks and (2) an assessment of possible 

hurricane effects at multiple depths at these banks. The benthic habitats of these banks are not 

well known and this was the first quantitative characterization of these banks at multiple depths. 

These banks are of particular interest because Hurricane Rita passed 7-mi (12-km) east of 

McGrail Bank and 15-mi (24-km) west of Sonnier Bank in September 2005. Hindcast models 

indicated that waves may have reached heights of up to 85-ft (26-m) at these banks, which may 

have exposed some bank caps, even those that are in 66-98 ft (20-30 m) under normal sea 

conditions. The implications for benthic community structure could have been catastrophic and 

the Minerals Management Service (MMS) considered it critical to assess the banks in their posthurricane state. 

The sampling protocol for this study was developed using literature references that included 

descriptions of biological communities at banks in the northwestern Gulf of Mexico (Rezak et al. 

1985; Boland 1999; Rezak and Bright 1979; Schmahl and Hickerson 2006). Rezak et al. (1985) 

included the most complete description of benthic communities or habitat types along a depth 

gradient at several banks within the Gulf of Mexico. These data were collected by diver and 

manned submersible in the late 1970s through the early 1980s and recently modified by 

Hickerson et al. (2008) (Table 1.2.1). 

In April and May 2007, videographic transects were collected by divers and a remotely operated 

vehicle (ROV) to characterize the benthic habitats of Sonnier, McGrail, Geyer, and Bright 

Banks. The sampling depth range in this study was from 72-197 ft (22-60 m). Four distinct 

depth ranges (zones) were sampled within the habitat types described by Rezak et al. (1985) 

(Table 2.1.1 and Table 2.1.2). The discontinuity between the four depth ranges was purposefully 

established to prevent overlap at the margins of each sampling zone. 

2.1.2. Field Methods 

Divers videotaped transects within no-decompression diving limits, down to ~120-ft (36.5-m). 

A diver swam slowly along each transect, videotaping at a height of 1.3-ft (40-cm) from the 

substratum, using a digital video camera in an Ikelite® underwater housing fitted with a wideangle lens and underwater video lights. A depth gauge and scaling bar projected forward from 

the video housing, ensuring that the camera remained a constant distance from the bottom. By 

holding the video camera perpendicular to the substratum and swimming slowly along the 

transect, it was possible to produce clear stop-action images for analysis (Aronson et al. 1994; 

Murdoch and Aronson 1999; Precht et al. 2006). 
16 

Table 2.1.1. 

Biological zonation of benthic communities by depth at Sonnier, McGrail, Geyer, and Bright 

Banks with corresponding depths from this study (adapted from Rezak et al. 1985). 

Bank 

Depth 

Ranges 

(m) 

this 

study 

Depth 

Zones 

(m) 

Rezak 

et al. 

1985 Sonnier McGrail Geyer Bright 

22-27 18-52 

Millepora

sponge 

30-36.5 37-52 

Millepora

sponge 

Millepora

sponge Stephanocoenia 

45-50 45-47 

Millepora

sponge Stephanocoenia

Millepora

sponge Algal sponge 

55-60 45-82 52+ Nepheloid Algal sponge 

Algal sponge to 

74 

ND 60-98 ND 

82-? 

Antipatharian Algal sponge 

74-? 

Antipatharian 

Table 2.1.2. 

Summary of data collected by diver and ROV during this study. 

Abbreviations: D = sampled by diver; R = sampled by ROV; 

NS = not sampled; “-” = zone not present. The number 

following the dash represents number of transects. Note that 

Bright Bank was sampled by divers only. The 45- to 50-m 

and 55- to 60-m depth ranges were sampled by ROV only.

Bank 

Depth Zone (m) Sonnier McGrail Geyer Bright 

22-27 D-2 - - - 

30-36.5 D-8 - D-8 D-3 

45-50 R-7 R-12 R-4 NS 

55-60 R-6 R-8 R-7 NS 

A TrackLink® 1500 High Accuracy Ultra Short Baseline acoustic positioning system (USBL) 

with two beacons was used to track divers videotaping transects at Sonnier, Geyer, and Bright 

Banks. The TrackLink system has an accuracy of 0.25 degrees. Data were converted into a GIS 

format in order to visualize where diver and ROV transects were in geographic space. 

 

22- to 27- m Transects 

At Sonnier Bank in the 22- to 27-m (72- to 89-ft) depth range, eight 10-m (33-ft) transects were 

positioned on the cap of the bank. Starting at a U-bolt attached to the seafloor (a mooring for 
17 

vessels up to approximately 108-ft or 33-m in length), a fiberglass surveyor’s tape was laid 

~656-ft (200-m) along the entire extent of the east-west axis of the cap of the bank. Transects 

were positioned at random distances along the east-west axis of the reef cap, and extended along 

random headings from the reference line. Because the transects were positioned with reference 

to the line and were, therefore, pseudoreplicates, the data from the individual transects were 

pooled, creating what was in essence one long transect for statistical analysis. One other transect 

was laid in the 22- to 27-m (72- to 89-ft) depth range, and started ~69-ft (21-m) northeast of the 

U-bolt and videotaped in a 90° direction. The TrackLink System technology was used to record 

diver locations underwater. A map was developed using the geo-referenced locations of divers 

that represents the eight transects that were later pooled and the single long transect that ran in a 

southeast-northwest direction (Figure 2.1.2). 

30- to 36.5-m Transects 

Due to the limited bottom time associated with the depths at Sonnier, Geyer, and Bright Banks, 

transect tapes were not deployed by divers. Depth contours along the reef slopes were used as a 

substitute for transect lines to guide the videographer. A diver swam along the depth contour 

holding the video camera 1.3-ft (40-cm) above the bottom, as in the transect method described 

above. Continuous video footage was collected at each bank within this depth range. A map 

was developed using the geo-referenced locations of the divers collecting video transects and 

their actual locations (e.g., Figure 2.1.2). Discrete transects were extracted from the footage, as 

described below. 

2.1.3. Laboratory Analysis 

Digital videography provides a permanent data record and is a reliable and logistically simple 

method of obtaining benthic cover data. Each video frame covered a 1.3-ft (40-cm) wide block 

of substratum. Each 10-m (33-ft) transect thus yielded ~4-m2 of benthic imagery, with each 

video frame measuring 40-cm x 27-cm, or 1080-cm2. This frame size was selected to enable 

investigators to identify corals and many other sessile invertebrates to species level at colony 

sizes down to 1.2-in (3-cm) longest dimension. Such precision is not attainable using video 

frames that record larger areas of the substratum (Aronson et al. 1994; Murdoch and Aronson 

1999). 

22- to 27-m Transects 

The 10-m (33-ft) transects recorded at Sonnier Bank in the 22- to 27-m (72- to 89-ft) depth range 

were cut into non-overlapping still images using ULead® VideoStudio® 9. Digital filters were 

applied using the ULead® software in order to enhance image quality. The original videotape of 

the transect was used to gain more detail or different perspectives on particular still images. 

After image capture and enhancement, randomly placed dots were added to each frame using 

Coral Point Count® (CPCe), for a total of 500 dots per transect. Organisms positioned beneath 

each random dot were identified to the lowest possible taxonomic level in one of the following 

categories: corals, antipatharians, sponges, turf algae (>3 mm thallus height), brown macroalgae, 

red (non-coralline) macroalgae, green macroalgae, other live organisms, coralline (red) algae, TB 

[fine turfs (<3-mm thallus height) and bare space], rubble, sand, and unknown. After each image 

was analyzed, the data were entered into project-specific Microsoft Excel spreadsheets. 
18 

Figure 2.1.2. Approximation of diver tracks, showing geographic positions of video 

Bank from 22-27 m (center of figure) and from 30-36.5 m (continuous bottom). 
19 

Quality assurance/quality control (QA/QC) for the video analysis consisted of independent 

analysis by trained observers of the same piece of captured video. There was agreement between 

pairs of observers at least 95% of the time. 

30- to 36.5-m Transects 

Videography in the 30- to 36.5-m (98- to 120-ft) depth range at Sonnier, Geyer, and Bright 

Banks produced continuous video footage for as long as a diver could remain on the bottom. 

The following method was used to create individual transects from this continuous footage. 

Not all video footage was usable for quantitative purposes, due to changes in distance from the 

bottom or selective videotaping of interesting mobile fauna by divers. First, the video footage 

was broken into still images using ULead® VideoStudio® 9. Digital filters were applied using 

the ULead® software to enhance image quality. Images that were not suitable for quantitative 

use were removed and used for qualitative analysis where possible. From the remaining still 

images, Transect 1 (T1) began at a random time near the beginning of the tape and continued for 

37 frames (~4-m2). After the 37th frame, a random number of frames (between 5 and 15) was 

skipped to select the beginning of the next transect (T2), and so on (Figure 2.1.3). 

Transects in 30- to 36.5-m (98- to 120-ft) depth range were analyzed in the same manner as the 

22- to 27-m (72- to 89-ft) transects, using the same 13 benthic categories. See Section 2.1.3 for 

data categories and QA/QC procedures. 

2.2. ROVING DIVER METHOD

2.2.1. Methodological Rationale 

The roving diver method was used to capture landscape-scale views of all banks visited. This 

video footage can be analyzed qualitatively for possible hurricane impacts, as it provides a 

broader perspective than the video transects. 

2.2.2. Field Methods 

Divers recorded video footage both during their descents and after the transects were videotaped. 

Roving diver video was recorded from 3-7 ft (1-2 m) above the substrate, looking down at a 45° 

angle. 

2.2.3. Laboratory Analysis 

The roving diver videos were qualitatively analyzed for possible hurricane impacts. Video 

footage from previous surveys (Sonnier Bank in 1996, 2002, and 2005; Geyer Bank in 2003; and 

Bright Bank in 2003) was assessed to estimate pre-hurricane conditions and benthic community 

structure. 
20 

Figure 2.1.3. Diagram illustrating the method used to create transects from continuous video 

footage taken along contour lines at 30- to 36.5-m depth at Sonnier, Geyer, and 

Bright Banks. 

2.3. ROV TRANSECT METHOD

2.3.1. Methodological Rationale 

The ROV video footage was collected at two depth ranges, 45-50 m (148-164 ft) and 55-60 m 

(180-197 ft), to characterize benthic communities and assess possible hurricane impacts. 

Additionally, it was an important goal of the study to distinguish between storm damage and 

damage due to oil and gas activities, which might be apparent at a greater depth. 

2.3.2. Field Methods 

While the vessel was moored or anchored at each bank (Sonnier, McGrail, and Geyer), ROV 

video was collected using a Seabotix LBV 300S-62

 ROV (Figure 2.3.2). The ROV was tethered 

to the vessel with a ~984-ft (300-m) umbilical cord and was controlled using a joystick located 

within the cabin of the ship. Real-time video was displayed while the ROV traversed the 
21 

benthos with time, direction, depth, and heading displayed on the video screen. The video was 

recorded onto an external hard drive and back-up copies were recorded on DVDs. 

Figure 2.3.2. Seabotix LBV 300S-62

 ROV at Sonnier Bank on May 22, 2007. 

A TrackLink® 1500 High Accuracy Ultra Short Baseline acoustic positioning system (USBL) 

with two beacons was used to track the ROV at Sonnier, McGrail, and Geyer Banks at the 45- to 

50-m (148- to 164-ft) and 55- to 60-m (180- to 197-ft) depth ranges. The TrackLink system has 

an accuracy of 0.25 degrees. The position of the Seabotix LBV 300S-62 ROV was recorded in 

real time and these data were collected and stored. Using ArcView GIS, these data were 

converted into a visualization of the ROV track (Figure 2.3.3-2.3.5). 

The ROV operations took place on May 21, 2007 at Geyer Bank between 1501 and 1721 hours 

local time and on May 22, 2007 at Sonnier Bank between 0820 and 1155 hours. ROV operations 

at McGrail Bank were conducted at night on May 22, 2007 between 0006 and 0218 hours due to 

impending bad weather. 

2.3.3. ROV Laboratory Analysis 

The ROV video data were analyzed in the same manner as the video footage collected by divers 

at 30- to 36.5-m (98- to 120-ft) depth (Section 2.1.3.). Video was converted into nonoverlapping still images. Still images that could not be used for quantitative analysis were 

removed, but were used for qualitative analysis if possible. Transects started at a random time 

(in seconds) and proceeded for 37 frames (~10-m length, or 4-m2

 area). A random number of 

frames were skipped before beginning the next 37 frame transect. See Section 2.1.3. for data 

categories and QA/QC procedures. 
22 

Figure 2.3.3. Map of ROV track at Sonnier Bank to and from the ship to a maximum depth of 
23 

Figure 2.3.4. Map of ROV track at Geyer Bank to and from the ship to a maximum depth of ~
24 

Figure 2.3.5. Map of ROV track at McGrail Bank to and from the ship to a maximum depth o
25 

2.4. TRANSECT STATISTICAL ANALYSIS METHODOLOGY

Descriptive statistics, univariate statistics, and graphs were created using Excel®. Live cover 

data was square-root transformed and analyzed using a one-way ANOVA (Analysis of 

Variance), followed by a Tukeys HSD test. Transect data, which consisted of all groups 

analyzed to the lowest taxonomic level, were entered into an Access® database to facilitate 

analysis using SAS®. Data fell within the following larger categories: corals (scleractinians and 

hydrocorals), antipatharians, sponges, turf algae (>3-mm thallus height), brown macroalgae, red 

(non-coralline) macroalgae, green macroalgae, other live organisms, coralline algae, TB [fine 

turfs (<3-mm thallus height) and bare space], rubble, sand, and unknown. In SAS® the data were 

partitioned into subsets and transposed so that they would be in the correct format for analysis 

with PRIMER® software. The data subsets were: 

1. Bright, Geyer, and Sonnier Banks at 30- to 36.5-m (98- to 120-ft) depth (SET A). 

2. Geyer, McGrail, and Sonnier Banks at 45- to 50-m (148- to 164-ft) and 55- to 60-m (180- 

to 197-ft) depth (SET B). 

3. Geyer and Sonnier at 30- to 36.5-m (98- to 120-ft), 45- to 50-m (148- to 164-ft), and 55- 

to 60-m (180- to 197-ft) depth (SET C). 

Each data subset was exported from SAS® to Excel® and then imported into a unique PRIMER®

workstation. 

The three data subsets were analyzed using PRIMER® Version 6.1.6. In each case the data were 

square-root transformed to reduce the disproportionate effects of dominant categories. The 

transformed data were then converted to a Bray-Curtis similarity matrix. 

Cluster analyses were performed on the Bray-Curtis matrices, using the group-averages 

clustering algorithm, and dendrograms were plotted. Multiple analyses were performed to 

evaluate linkages with respect to both site and depth range. 

Multidimensional scaling (MDS) analyses were performed on the Bray-Curtis matrices, allowing 

for 50 restarts and a 0.01 minimum stress level with Kruskal fit scheme 1 in PRIMER®. As with 

the cluster analyses, multiple analyses were performed to evaluate the effects of both site and 

depth range. 

One-way Analysis of Similarity (ANOSIM) analyses were performed on the Bray-Curtis 

matrices, allowing for a maximum of 999 permutations. For the subset data “SET A,” a one-way 

ANOSIM design was utilized, as the only factor in these data was site. For “SET B” and “SET 

C,” one-way analyses were performed separately for both factors: site and depth. Additionally, 

two-way crossed ANOSIM analyses were performed, with multiple transects from a given site 

and depth range serving as replicates. 
26 

2.5. REPETITIVE QUADRATS

2.5.1. Methodological Rationale 

To monitor changes in coral reef community structure as a result of Hurricane Rita, repetitive 8-m2

quadrats were photographed at the EFGB in June 2005, November 2005, and June 2006 and 

analyzed in two ways. The first analytical approach measured the percent cover of benthic 

components in November 2005 and June 2006 using random dot analysis. The second approach 

used planimetry to analyze select coral colonies within repetitive quadrats between June 2005 and 

November 2005 and between November 2005 and June 2006. Colonies of the dominant corals at 

the Flower Garden Banks were selected for analysis if their margins were visible in the 

photographs. Since the alignment of photographs is not exactly the same each year, corals closer to 

the centers of the photographs were chosen. 

2.5.2. Field Methods 

Forty repetitive quadrat stations were photographed within the 100-m x 100-m study site at the 

EFGB in November 2005. Thirty-nine stations were re-photographed in June 2006. In November 

2005, 9 deep station repetitive quadrats were photographed at the EFGB, and 7 stations were rephotographed in June 2006. Stations were photographed using a Nikonos V camera with a 15-mm 

lens. The camera was loaded with Kodak Ektachrome 200 ASA, 36-exposure slide film and 

standard settings were applied (distance = 2-m, f8). The camera was mounted in the center of a 

T-bar camera frame. Two Ikelite 75 watt-sec strobes, placed 4-ft (1.2-m) apart, were mounted on 

the ends of the T-bar and set on TTL and slave (Gittings et al. 1992). The camera was positioned in 

a north-facing direction and leveled to ensure repetitive photographs from year to year. The 

consistent orientation of the camera was achieved with a compass and a bubble level. 

2.5.3. Image Analysis of Repetitive Quadrats 

Percent cover of corals by species; the cover of coral bleaching, paling, concentrated and isolated 

fish biting, and disease; and the cover of algae were determined by overlaying 300 random dots on 

each repetitive quadrat station photograph using CPCe® point-count software with Excel®

extensions. Percent coverage was calculated from the November 2005 images and compared to the 

June 2006 images. Forty-nine images were analyzed in November 2005: 40 from within the 100-m 

x 100-m study site and 9 from the deep station repetitive quadrats. Forty-six images were analyzed 

in June 2006: 39 from within the 100-m x 100-m study site and 7 from the deep stations. 

Planimetry was used to measure changes in tissue extent for select coral colonies photographed 

within the quadrats in successive years. Four to six coral colonies were chosen within each 

repetitive quadrat. Areal measurements were taken using Sigma Scan Pro 5® planimetry software in 

each year for matched coral colonies. Each image was calibrated to an image size of 25.2-cm by 

37.9-cm. To show effects of the hurricane, 40 image pairs were analyzed from within the 100-m x 

100-m study site and 9 image pairs were compared from the deep repetitive quadrat stations 

between June 2005 and November 2005. Thirty-seven image pairs were compared from within the 

study site and 7 image pairs were analyzed from the deep stations between November 2005 and 

June 2006. 
27 

2.5.4. Data Presentation and Statistical Analysis of Repetitive Quadrats 

Mean percent coverages of benthic categories (corals, sponges, macroalgae, other live, CTB 

[crustose coralline algae, fine turfs, and bare rock], sand and rubble, and unknown) and coral 

conditions (bleaching, paling, fish biting, and disease) were calculated using random-dot analysis 

with CPCe® software. In order to obtain an estimate of percent cover for a given benthic category, 

the total number of dots within that category was divided by the total number of dots analyzed for 

all repetitive quadrat stations (minus the number of dots that fell on tape, wand, or shadow). In 

order to obtain a percent-cover value for a particular coral condition, the total number of dots 

landing on a particular condition (i.e. bleaching, fish biting, and disease) was divided by the total 

number of dots falling on living coral in all the repetitive quadrat stations. 

Planimetry was calculated by taking areal measurements of dominant corals each year. The areal 

values were compared between June 2005 and November 2005, and between November 2005 and 

June 2006. Descriptive statistics were used to characterize the change in area of dominant coral 

colonies following Hurricane Rita within 8-m2

 repetitive quadrats. 

2.6. PERIMETER VIDEOGRAPHY

2.6.1. Methodological Rationale 

The perimeter lines were videotaped in November 2005 and June 2006 to document changes at 

known locations along the perimeter and within the study site. General aspects of coral 

condition were documented and compared year to year. 

2.6.2. Field Methods 

Divers videotaped two 100-m segments of the perimeter lines at the EFGB—the north and east 

lines—in November 2005 and June 2006. The videos commenced at the northwest corner and 

continued along the north line to the northeast corner, then along the east line to the southeast 

corner. The videographer maintained ~7-ft (2-m) distance above the substratum using a 

weighted line attached to the video housing. The camera was aimed down at a 45° angle to 

capture the substratum. At each corner of the study site, the videographer recorded a 360°

panoramic view of the reef. 

2.6.3. Laboratory Methods 

The video footage was reviewed to record the general condition of corals along the perimeter of 

the EFGB study site. Individual coral heads displaying disease, bleaching, paling, or tissue loss 

due to fish biting were identified and recorded. The categories were as follows: bleaching, 

paling, healthy colony, concentrated fish biting, isolated fish biting, growth infilling (tissue 

regrowth), new incidence of fish biting, surface replaced by turf algae, and unchanged from the 

previous year. Concentrated fish biting removes the coral polyps completely from an affected 

area and may be due to the activity of the parrotfish Sparisoma viride (Bruckner and Bruckner 

1998; Bruckner et al. 2000). Isolated fish biting is less dense and occurs on a smaller scale, 

typically representing the activity of damselfish within their territories. Affected coral colonies 
28 

were compared between November 2005 and June 2006 and changes in their condition were 

recorded. In addition to the comparison of coral colonies, coral species composition and fish 

counts were assessed. These analyses were qualitative and, therefore, no statistical analyses 

were performed. 

2.7. HYDRODYNAMIC MODEL ANALYSIS METHODS

In order to understand the wave and current conditions in the vicinity of Sonnier, McGrail, 

Geyer, Bright, and East Flower Garden Banks during the passage of Hurricane Rita, PBS&J 

conducted a wave study using hindcast data provided by Oceanweather, Inc. The Oceanweather, 

Inc. study was completed for MMS to provide hindcast wind and wave datasets in the Gulf of 

Mexico for Hurricanes Katrina and Rita (Table 2.6.1). This study consisted of two distinct 

analyses: (1) a numerical modeling effort to determine the magnitude of wave transformation 

from the hindcast wave location to the bank locations and (2) an analytical model to estimate the 

potential currents resulting from the transformed wave conditions. 

Numerical modeling was performed with the REF/DIF program (Kirby and Dalrymple 1994), 

which is a weakly non-linear water wave refraction and diffraction model. The model inputs are 

the hindcast wave height, period, and direction for the vicinity of the banks, as well as the water 

depth at each bank location. REF/DIF computes the predicted wave height and direction at each 

bank location, taking into account wave refraction, diffraction, and shoaling due to the sharp 

changes in water depth. The model was employed to assess the extent of the effects of bank 

bathymetry on storm waves propagating over the banks at the bank locations. The model 

showed that the banks have a sizable impact on wave conditions in their vicinity. The results 

from the modeling effort were used to alter the hindcast data. The wave heights from the 

hindcast study were increased to a point just before breaking would have occurred, which is 

approximately 60% of the water depth at each bank location. 

These new wave conditions were used as input for an analytical calculation of potential induced 

water velocities at the bank locations. This analysis utilizes Stream Function Wave Theory 

(Dean 1965), a fully non-linear formulation that is applicable for waves in deep water to near 

breaking conditions. The program (Chaplin 1999) uses transformed wave height, wave period, 

and water depth at the bank locations to estimate two-dimensional velocity throughout the water 

column induced by the storm waves. Because the hurricane waves that impacted the banks were 

quite large relative to the water depths, linear wave theory cannot be used to accurately calculate 

water motions. Therefore, a fully-non linear application is most appropriate for the calculation 

of water velocities in this case (Dean and Dalrymple 1991). The output of this analysis is 

predicted water velocity with depth, from the seafloor to the water surface, induced by the 

passage of storm waves. 

 
29 

Table 2.6.1. 

Wave conditions during Hurricane Rita as hindcast by Oceanweather Inc. (2006). 

Depth of seamount is a value obtained from bathymetry data using GIS. 

Site McGrail 

Bank 

Geyer 

Bank 

Sonnier 

Bank 

Bright 

Bank 

EFGB 

Latitude (deg N) 27.96 27.85 28.34 27.89 27.91 

Longitude (deg W) 92.6 93.06 92.46 93.31 93.6 

Hindcast model grid Basin 

(Rita) 

Basin 

(Rita) 

Basin 

(Rita) 

Basin 

(Rita) 

Basin 

(Rita) 

Hindcast node ID 51886 51308 8629 51587 51581 

Depth (m) 201 223 60 139 131 

Sig Wave Height (m) 13.6 11.7 12.6 10.9 9.85 

Assoc Period (swell, s) 14.3 15.5 14.5 15.4 15.1 

Peak Period (swell, s) 15.5 15.5 15.9 16.2 15.4 

Max surface current (m/s) 0.2 0.16 1.89 0.19 0.21 

Depth of seamount (m) 45.0 39.1 22.2 32.9 21.0 
31 

3.0 RESULTS 

3.1. TRANSECT DATA DESCRIPTIVE STATISTICS 

The transect data were collected to characterize and compare the benthic habitats of Sonnier, 

McGrail, Geyer, and Bright Banks subsequent to the passing of Hurricane Rita. Living benthic 

cover at each bank generally declined with depth (Figure 3.1.1; Appendices 1 and 2). Bright 

Bank exhibited the highest live cover of any bank (86%), while Sonnier had the lowest live cover 

(2%). Live cover data were pooled for each bank and data were square-root transformed in order 

to perform a one-way ANOVA (Analysis of Variance) (Table 3.1.1). Live cover was 

significantly different across banks (F = 16.49, df = 64, P<0.0001). Tukey’s HSD test showed 

significant differences between all pairs containing Sonnier Bank, as well as between Bright 

Bank and McGrail Bank (Table 3.1.1). 

Percent Live Cover

0

10

20

30

40

50

60

70

80

90

100

22-27 30-36.5 45-50 55-60

Depth Range (m)

Percent Cover

Sonnier Bank

Geyer Bank

McGrail Bank

Bright Bank

Figure 3.1.1. Percent live cover at Sonnier, Geyer, McGrail, and Bright Banks in all depth 

ranges. Error bars represent one standard error. 

Of the four banks assessed, Sonnier Bank was the only bank containing all four depth ranges. 

Surveys at Sonnier, Geyer, and McGrail Banks began at the reef cap and proceeded to a 

maximum depth of ~197-ft (60-m). Due to time constraints, only one depth range (30-36.5 m or 

98-120 ft) was surveyed at Bright Bank. The benthic characteristics of the EFGB have been 

thoroughly documented (long-term monitoring of the Flower Garden Banks has occurred since 

1989) and consequently, the EFGB was not included in the April and May 2007 transect data 

collection. Thus, data for the EFGB is not included in the following analyses. 
32 

Table 3.1.1. 

Results of one-way ANOVA and Tukey HSD test performed on square root transformed 

live cover estimates from diver and ROV transects. 

Asterisk (*) denotes significant difference at a 95% confidence interval. 

Square root transformed data

Bank n Mean 

Standard 

Error 

Pooled 

Standard 

Error Standard Deviation 

Sonnier Bank 23 3.5414 0.4141 0.3567 1.9861 

Geyer Bank 19 6.4916 0.3625 0.3924 1.5800 

McGrail Bank 20 5.2135 0.3491 0.3825 1.5612 

Bright Bank 3 9.2727 0.2742 0.9875 0.4749 

Source of variation 

Sum of 

Squares df 

Mean 

Square 

F 

statistic P value

Square root transformed data 144.7145 3 48.2382 16.49 <0.0001

Residual 178.4716 61 2.9258 

Total 323.1861 64 

Tukey Contrast Difference 95% Confidence Interval 

Sonnier Bank v Geyer Bank -2.9502 -4.3508183225 to -1.5495810395* 

Sonnier Bank v McGrail Bank -1.6721 -3.0534138161 to -0.2907929789* 

Sonnier Bank v Bright Bank -5.7313 -8.504656051 to -2.9580400542* 

Geyer Bank v McGrail Bank 1.2781 -0.1692634323 to 2.7254559993 

Geyer Bank v Bright Bank -2.7811 -5.5879381622 to 0.0256414189 

McGrail Bank v Bright Bank -4.0592 -6.8564494781 to -1.2620398321* 

Obvious hurricane damage, such as toppled coral heads or broken sponges, was not detected at 

these banks. Due to the dominance of algae and sponge communities, and the paucity of hard 

corals, some signs of hurricane damage typical of coral reefs (e.g., toppled coral heads) were not 

evident. Some algal taxa would be expected to grow back within six months of a hurricane, 

while sponges would have a longer recovery period. Sponges may have healed in the twenty 

months between the passage of the hurricane and this study. No overturned corals were 

documented at sites with corals (Bright and McGrail Banks), although overturned corals and 

other coral related damage were evident in videotapes and photographs from the EFGB. Percent 

live cover is given by species and depth for each bank in Appendix 1. 

3.2. SONNIER BANK

Sonnier Bank was dominated by brown macroalgae, sponges, and Millepora alcicornis in 

shallow areas, while TB (fine turfs < 3-mm and bare space) and rubble (bare or covered in fine 

turf algae) were dominant at depth. A complete list of taxa identified in the video footage is 

provided with percent cover by depth range in Appendix 1. 
33 

Video footage of Sonnier Bank from 1996 (NOAA), 2002 (MMS), and 2005 (NOAA) were 

viewed for qualitative analysis and compared to one another and the transect videos from this 

study. The video footage from 1996 and 2005 were recreational videos recorded on the reef cap 

of Sonnier Bank (courtesy of E.L. Hickerson). The video footage from 2002 was collected by 

the Gulf Reef Environmental Action Team (GREAT). The goal of the project was to establish a 

monitoring program to detect natural variation at Sonnier Bank in advance of any nearby oil and 

gas development activities (Boland, personal communication, 2008). The second, and final, 

monitoring trip occurred in May 2002, which included the videotaping of transects on the 79-ft 

(24-m) peak. 

The depths videotaped in previous years (1996, 2002, and 2005) are comparable to the 

shallowest depth range in this study, which is 22-27 m (72-89 ft). Although impossible to 

quantify, it appeared that there was more live cover of algae, sponges, and Millepora alcicornis

in earlier years. The barrel sponge Xestospongia muta was recorded in 1996 and 2002, but this 

species was absent from the 2005 videotapes (Figure 3.2.1). Xestospongia muta was not 

recorded at Sonnier Bank in 2007. Roving diver video from April-May 2007 showed areas of 

low-lying algal and sponge cover, with no evidence of overturned corals or other organisms 

removed from their growth positions. Numerous areas of rubble were observed and they were 

evident in the transect video. 

 (A) (B) 

Figure 3.2.1. (A) Xestospongia muta from 1996 video (NOAA) and (B) X. muta from 2002 

video (Boland 2002). 

22- to 27-m Depth 

Atop Sonnier Bank from 22-27 m (72-89 ft), live cover was ~38% and consisted mostly of 

brown macroalgae, Millepora alcicornis (the only coral recorded in these transects), sponges, 

and turf algae, the latter of which was unidentifiable to taxonomic group (Figure 3.1.1; Figure 

3.2.2; Table 3.2.1). Appendix 2 contains transect data for all the banks. 

30- to 36.5-m Depth 

In the 30- to 36.5-m (98- to 120-ft) depth range, live cover decreased to ~30%. The dominant 

live component of the substratum was brown macroalgae, comprising 22%. The cover of 

Millepora alcicornis declined to 1.6%. Rubble was prominent in this depth range, covering 

~40% of the substratum (Figure 3.2.2; Figure 3.2.3; Table 3.2.2). 
34 

Sonnier Bank 

0

10

20

30

40

50

60

70

80

90

100

Coral

Antipatharians

Sponge

Turf Algae

Brown Macroalgae

Red Macroalgae

Other Live

Total Coralline Algae

TB

Rubble 

Sand

Benthic Category

Percent Cover

22-27 meters

30-36.5 meters

45-50 meters

55-60 meters

Figure 3.2.2. Percent cover of benthic categories at all depth ranges at Sonnier Bank. 

Error bars represent one standard error. 

Table 3.2.1. 

Percent cover of benthic categories at Sonnier Bank from 22- to 27-m depth. 

No samples of green macroalgae or unknown categories were identified. 

SONNIER BANK 22-27 METERS 

 Mean Percent Cover Standard Error 

CORAL 

Millepora alcicornis 7.98 1.73 

Total Coral 7.98 1.73 

ANTIPATHARIANS 

Stichopathes sp. 0.20 0.20 

Total Antipatharians 0.20 0.20 

SPONGES 

Agelas clathrodes 0.61 0.09 

Ircinia spp. 0.33 0.03 

Neofibularia nolitangere 0.46 0.44 

Niphates erecta 0.05 0.05 

Spheciospongia sp. 0.25 0.25 

Unidentified Sponges 0.65 0.45 

Total Sponge 2.35 0.35 
35 

Table 3.2.1. Percent cover of benthic categories at Sonnier Bank from 

22- to 27-m depth (continued). 

SONNIER BANK 22-27 METERS 

 Mean Percent Cover Standard Error 

TURF ALGAE 

Total Turf Algae 6.34 4.24 

BROWN MACROALGAE 

Dictyota spp. 8.43 1.93 

Lobophora variegata 7.75 3.85 

Padina sp. 0.34 0.24 

Unidentified Brown 

macroalgae 2.11 2.09 

Total Brown Macroalgae 18.63 3.93 

RED MACROALGAE 

Total Red Macroalgae 1.06 1.06 

GREEN MACROALGAE 

Total Green Macroalgae 0.00 0.00 

OTHER LIVE 

Diadema antillarum 0.05 0.05 

Gastropoda 0.25 0.15 

Holothuroidea 0.05 0.05 

Total Other Live 0.35 0.15 

CORALLINE ALGAE 

Total Coralline Algae 1.41 1.19 

FINE TURF, BARE 

Rubble 19.64 0.46 

TB 41.45 5.95 

Total TB and Rubble 61.09 6.41 

SAND 

Sand 0.60 0.50 

Total Sand 0.60 0.50 

UNKNOWNS 

Total Unknown 0.00 0.00 

Total Live Cover 38.31 

TOTAL 100.0 

45- to 50-m Depth 

Live cover at 45- to 50-m (148- to 164-ft) depth was low, at 6.8 %. Live cover consisted 

primarily of turf algae, at 5% cover. Within this depth range, rubble was prevalent at 47% cover 

(Figure 3.2.2; Table 3.2.3). 

55- to 60-m Depth 

Live cover at 55- to 60-m (180- to 197-ft) depth was extremely low, at 1.8%. Sand was the 

largest space occupant at this depth, covering 87% of the substratum and rubble decreased to 

~9% cover (Figure 3.2.2; Table 3.2.4). Within this depth range, a number of fireworms were 

noted, although not at a sufficiently high density to be quantified at the sampling density 

employed (500 dots per 4-m2

). No obvious signs of hurricane damage were documented. 
36 

Figure 3.2.3. Large rubble typical of 30- to 36.5-m depth range at 

Sonnier Bank. 

Table 3.2.2. 

Percent cover of benthic categories at Sonnier Bank from 30- to 36.5-m 

depth. 

No samples of red macroalgae, green macroalgae, or unknown categories 

were identified.

SONNIER BANK 30-36.5 METERS 

 Mean Percent Cover Standard Error 

CORAL 

Millepora alcicornis 1.60 0.55 

Total Coral 1.60 0.55 

ANTIPATHARIANS 

Stichopathes sp. 0.03 0.03 

Total Antipatharians 0.03 0.03 

SPONGES 

Agelas clathrodes 0.03 0.03 

Ircinia spp. 0.28 0.13 

Neofibularia nolitangere 0.50 0.33 

Niphates erecta 0.28 0.14 

Spheciospongia sp. 0.25 0.17 

Unidentified Sponges 0.45 0.07 

Total Sponge 1.78 0.65 

TURF ALGAE 

Total Turf Algae 4.48 1.78 

BROWN MACROALGAE 

Dictyota spp. 13.48 0.99 

Lobophora variegata 1.65 0.56 
37 

Table 3.2.2. Percent cover of benthic categories at Sonnier Bank from 

30- to 36.5-m depth (continued). 

SONNIER BANK 30-36.5 METERS 

 Mean Percent Cover Standard Error 

Padina spp. 0.03 0.03 

Unidentified Brown Macroalgae 6.90 3.08 

Total Brown Macroalgae 22.1 3.80 

RED MACROALGAE 

Total Red Macroalgae 0.00 0.00 

GREEN MACROALGAE 

Total Green Macroalgae 0.00 0.00 

OTHER LIVE 

Fish 0.03 0.03 

Gastropoda 0.03 0.03 

Total Other Live 0.05 0.03 

CORALLINE ALGAE 

Total Coralline Algae 0.08 0.05 

FINE TURF, BARE 

Rubble 40.30 3.68 

TB 28.90 5.55 

Total TB and Rubble 69.20 3.61 

SAND 

Sand 0.75 0.23 

Total Sand 0.75 0.23 

UNKNOWNS 

Total Unknown 0.00 0.00 

Total Live Cover 30.06 

TOTAL 100.0 

3.3. MCGRAIL BANK

Live cover at McGrail Bank was dominated by macroalgae and coral (specifically 

Stephanocoenia intersepta) in the 45- to 50-m (148- to 164-ft) depth range and algal nodules and 

macroalgae at deeper depths (55-60 m or 180-197 ft). Rubble and sand were the major benthic 

components in both depth ranges. A complete list of taxa identified in the video footage is 

provided with percent cover by depth range in Appendix 1. 

45- to 50-m Depth 

Atop McGrail Bank at 148-ft (45-m), live cover was ~38%, consisting primarily of brown and 

green macroalgae, as well as scleractinian corals (Figure 3.1.1; Figure 3.3.1; Table 3.3.1). 

Rubble was a dominant feature at 39% cover. Differentiated from rubble were algal nodules, 

which our transect video only identified at McGrail Bank and comprised ~2% of the substratum 

cover. The mean coral cover was ~5% and consisted mostly of Stephanocoenia intersepta, at

4.8% cover (Figure 3.3.2), with Millepora alcicornis, Agaricia sp., and Montastraea cavernosa

comprising the remaining 0.7% (Appendix 2). 
38 

Table 3.2.3. 

Percent cover of benthic categories at Sonnier Bank from 45- to 50-m depth. 

No samples of antipatharians, red macroalgae, or green macroalgae were 

identified. 

SONNIER BANK 45-50 METERS 

 Mean Percent Cover Standard Error 

CORAL 

Octocorallia 0.34 0.17 

Total Coral 0.34 0.17 

ANTIPATHARIANS 

Total Antipatharians 0.00 0.00 

SPONGES 

Ircinia spp. 0.11 0.09 

Unidentified Sponges 0.43 0.13 

Total Sponge 0.54 0.12 

TURF ALGAE 

Total Turf Algae 5.03 0.53 

BROWN MACROALGAE 

Dictyota spp. 0.31 0.25 

Unidentified Brown Macroalgae 0.49 0.29 

Total Brown Macroalgae 0.80 0.34 

RED MACROALGAE 

Total Red Macroalgae 0.00 0.00 

GREEN MACROALGAE 

Total Green Macroalgae 0.00 0.00 

OTHER LIVE 

Gastropoda 0.03 0.03 

Total Other Live 0.03 0.03 

CORALLINE ALGAE 

Total Coralline Algae 0.03 0.03 

FINE TURF, BARE 

Rubble 47.00 6.41 

TB 9.77 1.83 

Total TB and Rubble 56.77 5.57 

SAND 

Sand 36.4 5.94 

Total Sand 36.40 5.94 

UNKNOWNS 

Total Unknown 0.06 0.04 

Total Live Cover 6.77 

TOTAL 100.0 
39 

Table 3.2.4. 

Percent cover of benthic categories at Sonnier Bank from 55- to 60-m depth. 

No samples of coral, brown macroalgae, red macroalgae, green 

macroalgae, or unknown categories were identified. 

SONNIER BANK 55-60 METERS 

 Mean Percent Cover Standard Error 

CORAL 

Total Coral 0.00 0.00 

ANTIPATHARIANS 

Antipatharia spp. 0.17 0.11 

Total Antipatharian 0.17 0.11 

SPONGES 

Unidentified Sponges 0.13 0.07 

Total Sponge 0.13 0.07 

TURF ALGAE 

Total Turf Algae 1.40 0.49 

BROWN MACROALGAE 

Total Brown Macroalgae 0.00 0.00 

RED MACROALGAE 

Total Red Macroalgae 0.00 0.00 

GREEN MACROALGAE 

Total Green Macroalgae 0.00 0.00 

OTHER LIVE 

Cerianthidea 0.03 0.03 

Gastropoda 0.03 0.03 

Total Other Live 0.07 0.04 

CORALLINE ALGAE 

Total Coralline Algae 0.03 0.03 

FINE TURF, BARE 

Rubble 9.37 1.39 

TB 1.40 0.38 

Total TB and Rubble 10.77 1.67 

SAND 

Fish Hole 0.13 0.10 

Sand 87.30 1.94 

Total Sand 87.43 1.98 

UNKNOWNS 

Total Unknown 0.00 0.00 

Total Live Cover 1.80 

TOTAL 100.0 
40 

McGrail Bank

0

10

20

30

40

50

60

70

80

90

100

Coral

Antipatharians

Sponge

Turf Algae

Brown Macroalgae

Green Macroalgae

Red Macroalgae

Other Live

Coralline Algae

Algal Nodules

TB

Rubble 

Sand

Benthic Category

Percent Cover

45-50 meters

55-60 meters

Figure 3.3.1. Percent cover of benthic categories in both depth ranges at McGrail Bank. 

Error bars represent one standard error. 

It should be noted that the percent cover of Stephanocoenia intersepta appeared to be much 

higher in some areas compared to others within this depth range, ranging from 0-32% within the 

transect data (Appendix 2). A multidimensional scaling (MDS) analysis was performed on the 

45- to 50-m (148- to 164-ft) transect data, based on multivariate cover of the four hard-coral taxa 

(Agaricia sp., Millepora alcicornis, Montastraea cavernosa, and Stephanocoenia intersepta.) 

The MDS plot (Figure 3.3.3) reveals that there is indeed high variability among transects in the 

composition of the coral assemblage at McGrail Bank in this depth range. The low stress level 

(0.01) of the MDS plot provides confidence in this result. No obvious signs of hurricane damage 

were documented. 

55- to 60-m Depth 

Live cover within this depth range was ~17% (Figure 3.1.1) and was comprised mostly of algal 

nodules (Figure 3.3.4) and the brown macroalga Lobophora variegata (Figure 3.3.1; Table 

3.3.2). Rubble and sand were the dominant features of McGrail Bank in the ROV transects from 

55- to 60-m (180- to 197-ft) depth (Figure 3.3.1; Table 3.3.2). No obvious signs of hurricane 

damage were documented. 
41 

Table 3.3.1. 

Percent cover of benthic categories at McGrail Bank from 45- to 50-m depth. 

No samples of turf algae were identified. 

MCGRAIL BANK 45-50 METERS 

 Mean Percent Cover Standard Error 

CORAL 

Agaricia spp. 0.08 0.04 

Millepora alcicornis 0.59 0.25 

Montastraea cavernosa 0.02 0.02 

Stephanocoenia intersepta 4.75 2.65 

Total Coral 5.43 2.84 

ANTIPATHARIANS 

Antipatharia spp. 0.52 0.50 

Total Antipatharians 0.52 0.50 

SPONGES 

Agelas clathrodes 0.19 0.08 

Ircinia spp. 0.26 0.18 

Pseudoceratina crassa 0.04 0.03 

Unidentified Sponges 1.02 0.53 

Xestospongia muta 0.37 0.21 

Total Sponge 1.88 0.85 

TURF ALGAE 

Total Turf Algae 0.00 0.00 

BROWN MACROALGAE 

Unidentified Brown Macroalgae 0.60 0.13 

Dictyota spp. 5.88 1.36 

Lobophora variegata 2.56 0.77 

Sargassum spp. 1.10 0.27 

Total Brown Macroalgae 10.15 1.40 

RED MACROALGAE 

Red Macroalgae 4.48 1.61 

Total Red Macroalgae 4.48 1.61 

GREEN MACROALGAE 

Green Macroalgae 9.50 2.43 

Total Green Macroalgae 9.50 2.43 

OTHER LIVE 

Diadema antillarum 0.29 0.16 

Fish 0.08 0.03 

Asteroidea/Ophiuroidea 0.02 0.02 

Gastropoda 0.02 0.02 

Total Other Live 0.40 0.18 

CORALLINE ALGAE 

Algal Nodules 2.44 0.50 

Crustose Coralline Algae 2.79 0.70 

Total Coralline Algae 5.22 0.58 
42 

Table 3.3.1. Percent cover of benthic categories at McGrail Bank from 

45- to 50-m depth (continued). 

MCGRAIL BANK 45-50 METERS

 Mean Percent Cover Standard Error 

FINE TURF, BARE 

Rubble 39.26 3.96 

TB 4.76 2.27 

Total TB and Rubble 44.01 2.50 

SAND 

Total Sand 18.31 1.54 

UNKNOWNS 

Total Unknown 0.09 0.04 

Total Live Cover 37.58 

TOTAL 100.0 

Figure 3.3.2. Frame capture from ROV video at 

McGrail Bank, showing a large 

colony of the coral Stephanocoenia 

intersepta. Laser lights in the center 

of the frame are 5-cm apart. 
43 

Figure 3.3.3. Two-dimensional MDS plot, based on square-root-transformed 

Bray-Curtis similarities, comparing multivariate cover of the four 

hard-coral taxa at McGrail Bank within the 45- to 50-m depth 

range. 

Figure 3.3.4. Frame capture from ROV video at 

McGrail Bank, showing algal 

nodules at 55- to 60-m depth. 

Laser lights in the center of the 

frame are 5-cm apart. 
44 

Table 3.3.2. 

Percent cover of benthic categories at McGrail Bank from 55- to 60-m depth. 

No samples of antipatharians or red macroalgae were identified. 

MCGRAIL BANK 55-60 METERS 

 Mean Percent Cover Standard Error 

CORAL 

Octocorallia 0.03 0.03 

Total Coral 0.03 0.03 

ANTIPATHARIANS 

Total Antipatharians 0.00 0.00 

SPONGES 

Unidentified Sponges 0.33 0.11 

Total Sponge 0.33 0.11 

TURF ALGAE 

Total Turf Algae 0.52 0.27 

BROWN MACROALGAE 

Unidentified Brown Macroalgae 0.24 0.08 

Dictyota spp. 0.76 0.47 

Lobophora variegata 3.55 2.37 

Total Brown Macroalgae 4.54 2.79 

RED MACROALGAE 

Total Red Macroalgae 0.00 0.00 

GREEN MACROALGAE 

Green Macroalgae 2.17 1.65 

Halimeda sp. 0.03 0.03 

Total Green Macroalgae 2.20 1.65 

OTHER LIVE 

Cerianthidea 0.19 0.10 

Fish 0.03 0.03 

Asteroidea/Ophiuroidea 0.06 0.04 

Total Other Live 0.28 0.14 

CORALLINE ALGAE 

Algal Nodules 6.51 0.46 

Crustose coralline algae 2.95 1.06 

Total Coralline Algae 9.46 1.10 

FINE TURF, BARE 

Rubble 54.99 3.38 

TB 2.06 0.76 

Total TB and Rubble 57.06 3.00 

SAND 

Total Sand 25.46 3.65 

UNKNOWNS 

Total Unknown 0.12 0.05 

Total Live Cover 17.37 

TOTAL 100.0 
45 

3.4. GEYER BANK

Geyer Bank was dominated by brown macroalgae, rubble, and TB. A complete list of taxa 

identified in the video footage is provided with percent cover by depth range in Appendix 1. 

Fish populations were the focus of diver-collected video surveys at Geyer Bank in July 2003 

(Moore 2003a), and therefore a limited qualitative comparison of the benthic communities was 

possible between 2003 and 2007. The benthic community at Geyer Bank was dominated by 

macroalgae and hydrocorals, specifically Sargassum spp. and Millepora alcicornis, respectively. 

Sponges were also observed, including Neofibularia nolitangere, Aplysina spp., Ircinia spp., and 

Xestospongia muta. Geyer Bank was the only location in this study with an observed population 

of Tubastraea coccinea. Observed T. coccinea colonies occupied all angles of the substrate, 

both vertical and horizontal surfaces. 

30- to 36.5-m Depth 

Live cover within the 30- to 36.5-m (98- to 120-ft) depth range was 60% at Geyer Bank (Figure 

3.1.1). Live cover at this depth consisted mostly of brown macroalgae, at 42% cover, with 

Sargassum being the dominant genus, at 27% (Figure 3.4.1; Figure 3.4.2; Table 3.4.1). Coral 

was present at Geyer Bank, accounting for ~10% of cover, with Millepora alcicornis as the 

dominant species (Figure 3.4.1; Table 3.4.1; Appendix 2). No obvious signs of hurricane 

damage were documented. 

Geyer Bank

0

10

20

30

40

50

60

70

80

90

100

Coral

Antipatharians

Sponge

Turf Algae

Brown Macroalgae

Green Macroalgae

Other Live

Total Coralline Algae

TB

Rubble

Sand

Benthic Category

Percent Cover

30-36.5 meters

45-50 meters

55-60 meters

Figure 3.4.1. Percent cover of benthic categories in all depth ranges at Geyer Bank. 

Error bars represent one standard error. 
46 

Figure 3.4.2. Frame capture of Sargassum spp., typical benthic 

cover in the 30- to 36.5-m depth range at Geyer 

Bank. 

Table 3.4.1. 

Percent cover of benthic categories at Geyer Bank from 30- to 36.5-m depth. 

No samples of antipatharians, red macroalgae, or green macroalgae were 

identified. 

GEYER BANK 30-36.5 METERS 

 Mean Percent Cover Standard Error 

CORAL 

Millepora alcicornis 9.15 0.68 

Porites sp. 0.03 0.03 

Unidentified Scleractinia 0.18 0.08 

Stephanocoenia intersepta 0.33 0.14 

Tubastraea coccinea 0.23 0.17 

Total Coral 9.90 0.74 

ANTIPATHARIANS 

Total Antipatharian 0.00 0.00 

SPONGES 

Aplysina spp. 0.23 0.10 

Clathria spp. 0.03 0.03 

Cliona sp. 0.05 0.05 

Neofibularia nolitangere 0.10 0.10 

Unidentified Sponges 0.75 0.13 

Xestospongia muta 0.08 0.08 

Total Sponge 1.23 0.18 
47 

Table 3.4.1. Percent cover of benthic categories at Geyer Bank from 

30- to 36.5-m depth (continued). 

GEYER BANK 30-36.5 METERS 

 Mean Percent Cover Standard Error 

TURF ALGAE 

Total Turf Algae 3.18 0.81 

BROWN MACROALGAE 

Unidentified Brown Macroalgae 0.08 0.04 

Dictyota spp. 2.68 0.43 

Lobophora variegata 12.65 0.90 

Padina spp. 0.03 0.03 

Sargassum spp. 26.88 3.38 

Total Brown Macroalgae 42.30 3.90 

RED MACROALGAE 

Total Red Macroalgae 0.00 0.00 

GREEN MACROALGAE 

Total Green Macroalgae 0.00 0.00 

OTHER LIVE 

Fish 0.23 0.10 

Total Other Live 0.23 0.10 

CORALLINE ALGAE 

Total Coralline Algae 3.43 0.53 

FINE TURF, BARE 

Rubble 13.70 2.46 

TB 24.10 2.07 

Total TB and Rubble 37.80 2.91 

SAND 

Total Sand 1.83 0.54 

UNKNOWNS 

Total Unknown 0.13 0.04 

Total Live Cover 60.25 

TOTAL 100.0 

45- to 50-m Depth 

Live cover was ~30% at Geyer Bank in the 45- to 50-m (148- to 164-ft) depth range (Figure 

3.1.1). Brown macroalgae was still the dominant taxonomic group, but decreased to ~22% 

(Figure 3.4.1; Table 3.4.2). Rubble and bare space increased to 63% in this depth range. No 

obvious signs of hurricane damage were documented. 

55- to 60-m Depth 

Live cover was ~35% in the 55- to 60-m (180- to 197-ft) depth range (Figure 3.1.1). Turf algae 

and coralline algae were higher than in 45-50 m (148-164 ft), accounting for the difference in 

live cover (Figure 3.4.1; Table 3.4.3). No obvious signs of hurricane damage were documented. 
48 

Table 3.4.2. 

Percent cover of benthic categories at Geyer Bank from 45- to 50-m depth. 

No samples of red or green macroalgae were identified. 

GEYER BANK 45-50 METERS 

 Mean Percent Cover Standard Error 

CORAL 

Millepora alcicornis 2.20 0.92 

Unidentified Scleractinia 0.10 0.06 

Stephanocoenia intersepta 0.20 0.14 

Total Coral 2.50 0.77 

ANTIPATHARIANS 

Stichopathes sp. 0.10 0.10 

Total Antipatharian 0.10 0.10 

SPONGES 

Agelas clathrodes 1.30 0.62 

Unidentified Sponges 1.55 0.76 

Xestospongia muta 0.65 0.39 

Total Sponge 3.50 1.37 

TURF ALGAE 

Total Turf Algae 0.65 0.33 

BROWN MACROALGAE 

Unidentified Brown Macroalgae 8.35 1.21 

Dictyota spp. 8.30 1.01 

Lobophora variegata 0.85 0.47 

Padina spp. 0.05 0.05 

Sargassum spp. 4.90 0.83 

Total Brown Macroalgae 22.45 0.81 

RED MACROALGAE 

Total Red Macroalgae 0.00 0.00 

GREEN MACROALGAE 

Total Green Macroalgae 0.00 0.00 

OTHER LIVE 

Fish 0.35 0.17 

Total Other Live 0.35 0.17 

CORALLINE ALGAE 

Total Coralline Algae 0.80 0.73 

FINE TURF, BARE 

Rubble 32.30 2.27 

TB 31.05 2.98 

Total TB and Rubble 63.35 2.63 

SAND 

Total Sand 6.25 1.77 

UNKNOWNS 

Total Unknown 0.05 0.05 

Total Live Cover 30.35 

TOTAL 100.00 
49 

Table 3.4.3. 

Percent cover of benthic categories at Geyer Bank from 55- to 60-m depth. 

No samples of antipatharians or red macroalgae were identified. 

GEYER BANK 55-60 METERS 

 Mean Percent Cover Standard Error 

CORAL 

Millepora alcicornis 0.11 0.07 

Unidentified Scleractinia 0.03 0.03 

Stephanocoenia intersepta 0.06 0.04 

Total Coral 0.20 0.10 

ANTIPATHARIANS 

Total Antipatharian 0.00 0.00 

SPONGES 

Agelas clathrodes 0.71 0.31 

Unidentified Sponges 1.17 0.30 

Xestospongia muta 0.66 0.33 

Total Sponge 2.54 0.63 

TURF ALGAE 

Total Turf Algae 3.94 1.12 

BROWN MACROALGAE 

Unidentified Brown Macroalgae 5.97 1.90 

Dictyota spp. 15.23 4.24 

Lobophora variegata 0.14 0.07 

Sargassum spp. 0.23 0.20 

Total Brown Macroalgae 21.57 4.96 

RED MACROALGAE 

Total Red Macroalgae 0.00 0.00 

GREEN MACROALGAE 

Green Macroalgae 0.20 0.12 

Total Green Macroalgae 0.20 0.12 

OTHER LIVE 

Fish 0.37 0.23 

Gastropoda 0.03 0.03 

Total Other Live 0.40 0.23 

CORALLINE ALGAE 

Total Coralline Algae 5.74 1.39 

FINE TURF, BARE 

Rubble 43.34 7.56 

TB 11.69 1.91 

Total TB and Rubble 55.03 6.77 

SAND 

Total Sand 10.09 0.66 

UNKNOWNS 

Total Unknown 0.29 0.07 

Total Live Cover 34.60 

TOTAL 100.0 
50 

3.5. BRIGHT BANK

Bright Bank was dominated by brown and green macroalgae, turf algae, TB, corals, and coralline 

algae. A complete list of taxa identified in the video footage is provided with percent cover by 

depth range in Appendix 1. 

Diver video taken in September 2003 from Bright Bank provided qualitative information about 

the benthic community at that time (Moore 2003b). Macroalgal cover was low and Lobophora 

variegata was the only macroalgal species identifiable in 2003. The substratum videotaped was 

mostly bare (Figure 3.5.1). Large colonies of Millepora alcicornis and Diploria strigosa were 

observed. Sponge species included Agelas clathrodes, Ircinia spp., and Xestospongia muta. 

The ROV data were not collected at this bank due to weather. This limited the data collected to 

diver depths which covered only the 30- to 36.5-m (98- to 120-ft) depth range. 

Figure 3.5.1. Captured frame from video recorded at Bright Bank 

in September 2003 at 36-m depth. Poles in 

photograph are thought to be part of salvage 

operations that have occurred at Bright Bank. 

30- to 36.5-m Depth 

In this depth range, live cover at Bright Bank was ~86%, with brown, green, and turf algae as the 

dominant groups (Figure 3.1.1; Figure 3.5.2; Figure 3.5.3; Table 3.5.1). Coral cover was ~8% 

and was comprised of four species, which in order of decreasing cover were Millepora 

alcicornis, Diploria strigosa, Stephanocoenia intersepta, and Montastraea cavernosa (Table 

3.5.1; Appendix 2). No obvious signs of hurricane damage were documented. 
51 

Bright Bank 30-36.5 meters

0

10

20

30

40

50

60

70

80

90

100

Coral

Sponge

Turf Algae

Brown Macroalgae

Green Macroalgae

Red Macroalgae

Other Live

Total Coralline Algae

TB

Rubble 

Sand

Benthic Category

Percent Cover

Figure 3.5.2. Percent cover of benthic categories at Bright Bank. Error bars represent one 

standard error. 

. 

Figure 3.5.3. Frame capture showing diverse benthic cover at 

Bright Bank in 30- to 36.5-m depth. 
52 

Table 3.5.1. 

Percent cover of benthic categories at Bright Bank from 30- to 36.5-m depth. 

No samples of antipatharians were identified. 

BRIGHT BANK 30-36.5 METERS 

 Mean Percent Cover Standard Error 

CORAL 

Diploria strigosa 0.87 0.87 

Millepora alcicornis 6.93 2.38 

Montastraea cavernosa 0.07 0.07 

Stephanocoenia intersepta 0.47 0.29 

Total Coral 8.33 3.38 

ANTIPATHARIANS 

Total Antipatharians 0.00 0.00 

SPONGES 

Agelas clathrodes 0.27 0.13 

Clathria spp. 0.20 0.20 

Spheciospongia sp. 0.07 0.07 

Unidentified Sponges 0.13 0.13 

Xestospongia muta 0.33 0.33 

Total Sponge 1.00 0.81 

TURF ALGAE 

Total Turf Algae 12.13 1.33 

BROWN MACROALGAE 

Unidentified Brown Macroalgae 28.13 6.29 

Dictyota spp. 0.33 0.18 

Lobophora variegata 14.20 3.53 

Padina spp. 0.13 0.07 

Sargassum spp. 0.80 0.61 

Total Brown Macroalgae 43.60 9.62 

RED MACROALGAE 

Red Macroalgae 0.07 0.07 

Total Red Macroalgae 0.07 0.07 

GREEN MACROALGAE 

Green Macroalgae 13.13 0.82 

Total Green Macroalgae 13.13 0.82 

OTHER LIVE 

Fish 0.73 0.07 

Gastropoda 0.07 0.07 

Total Other Live 0.80 0.00 

CORALLINE ALGAE 

Total Coralline Algae 7.07 1.62 

FINE TURF, BARE 

Rubble 1.33 0.93 

TB 12.27 3.88 

Total TB and Rubble 13.60 4.77 
53 

Table 3.5.1. Percent cover of benthic categories at Bright Bank from 

30- to 36.5-m depth (continued). 

BRIGHT BANK 30-36.5 METERS 

 Mean Percent Cover Standard Error 

SAND 

Total Sand 0.20 0.20 

UNKNOWNS 

Total Unknown 0.07 0.07 

Total Live Cover 86.13 

TOTAL 100.00 

3.6. SHANNON-WIENER DIVERSITY INDEX (H') 

The Shannon-Wiener Diversity Index, H', was calculated for each bank at each depth using the 

lowest available taxonomic groupings (species or genera). The highest diversity found was at 

Sonnier Bank from 22-27 m (72-89 ft), due largely to the variety of sponges present there. 

Diversity decreased with depth at Sonnier Bank (Table 3.6.1). Both Geyer and McGrail Banks 

exhibited highest diversity values in the 45- to 50-m (148- to 164-ft) depth range. The brown 

macroalgae accounted for these high diversity values. Notably, diversity at 45-50 m (148-164 ft) 

was very low at Sonnier Bank compared to Geyer and McGrail Banks. Brown algae 

distinguishable to genus or species were virtually absent from Sonnier Bank between 45-50 m 

(148-164 ft). Turf algae, rubble, and sand were the dominant categories at Sonnier Bank from 

55-60 m (180-197 ft), accounting for the zero-value of H' in that depth range. 

Table 3.6.1. 

Shannon-Wiener Diversities (H') at all banks and depths surveyed. 

 Bank 

Depth Sonnier Geyer McGrail Bright EFGB 

22-27 2.86 – – – 2.43 

30-36.5 1.55 1.65 – 1.81 2.27 

45-50 0.23 2.13 2.08 – – 

55-60 0* 1.47 0.83 – – 

* only turf algae, rubble and sand 

3.7. MULTIVARIATE STATISTICAL RESULTS

3.7.1. ANOSIM Analysis 

All Sites and Depths (excluding 22- to 27-m dataset) 

The lowest taxonomic levels of benthic cover category data including corals, antipatharians, 

sponges, turf algae (>3-mm), brown macroalgae, green macroalgae, red (non-coralline) 

macroalgae, crustose coralline algae, “other live” (which consisted of fish, mollusks, and any 

other living organism that did not fall within one of the above categories), rubble, TB (fine turfs 

<3-mm and bare space), sand, and unknown were analyzed using multivariate techniques. 
54 

One-way Analysis of Similarity (ANOSIM) with site as factor showed that there were significant 

among-site differences (Global R = 0.54, P = 0.001). Pairwise tests showed differences between 

each possible pair of banks (Table 3.7.1). McGrail and Bright Banks were the most dissimilar 

banks (R = 0.926, P = 0.003), whereas Geyer and Sonnier Banks were the least dissimilar (R = 

0.318, P = 0.001). 

Table 3.7.1. 

Results of Pairwise ANOSIM tests with site as the factor. 

The closer R is to 1, the more dissimilarity between treatments. 

One-way ANOSIM was also conducted with depth as the factor. A significant difference was 

found among depths (Global R = 0.37, P = 0.001). Pairwise tests showed differences between 

depths, although depths were not as dissimilar as sites (Table 3.7.2). The 45- to 50-m (148- to 

164-ft) and 55- to 60-m (180- to 197-ft) depth ranges were less dissimilar than the dissimilarities 

between 30-36.5 m (98-120 ft) and 55-60 m, and between 30-36.5 m and 45-50 m (Table 3.7.2). 

In fact, adjustment of the α-value, to maintain an overall type-I error rate of 0.05, renders the 

comparison between 45-50 m and 55-60 m non-significant. 

Table 3.7.2. 

Results of Pairwise ANOSIM tests with depth as the factor. 

The closer R is to 1, the greater the dissimilarity is between treatments. 

Groups 

(m) 

RStatistic Significance 

Level 

Possible 

Permutations

Actual 

Permutations 

Number ≥

Observed 

45-50/ 

30-36.5 0.499 0.001 Very Large 999 0 

45-50/ 

55-60 0.065 0.05 Very Large 999 49 

30-36.5/ 

55-60 0.584 0.001 Very Large 999 0 

A Two-Way Crossed Analysis was conducted with site and depth as factors. The results showed 

significant differences among sites across depths (Global R = 0.9, P = 0.001), as well as 

differences among depth ranges across sites (R = 0.77, P = 0.001). 

Groups 

RStatistic Significance 

Level 

Possible 

Permutations

Actual 

Permutations 

Number ≥

Observed 

McGrail, Bright 0.926 0.003 1771 999 2 

McGrail, Geyer 0.691 0.001 Very Large 999 0 

McGrail, Sonnier 0.516 0.001 Very Large 999 0 

Bright, Geyer 0.734 0.002 1540 999 1 

Bright, Sonnier 0.71 0.002 2600 999 1 

Geyer, Sonnier 0.318 0.001 Very Large 999 0 
55 

Pairwise tests showed that Bright Bank was most different from Geyer and Sonnier Banks (Table 

3.7.3). Pairwise tests between Bright and McGrail Banks were not possible because data did not 

exist for McGrail Bank from 30-36.5 m (98-120 ft). 

Table 3.7.3. 

Results of Pairwise ANOSIM tests for differences between sites across all depths. 

The closer R is to 1, the greater the dissimilarity is between treatments. 

Groups 

RStatistic Significance 

Level 

Possible 

Permutations 

Actual 

Permutations 

Number ≥

Observed 

McGrail/Geyer 0.799 0.001 11711700 999 0 

McGrail/ Sonnier 0.866 0.001 151315164 999 0 

Bright/Geyer 1 0.006 165 165 1 

Bright/Sonnier 1 0.006 165 165 1 

Geyer/Sonnier 0.995 0.001 Very Large 199 0 

Differences between depth groups were most evident between the 30- to 36.5-m (98- to 120-ft) 

and 55- to 60-m (180- to 197-ft) depth ranges and between the 45- to 50-m (148- to 164-ft) and 

30- to 36.5-m depth ranges. The 45- to 50-m and 55- to 60-m comparison showed more 

similarity between these depth ranges with respect to benthic composition (Table 3.7.4). 

Table 3.7.4. 

Results of pairwise ANOSIM tests for differences between depths across all sites. 

The closer R is to 1, the greater the dissimilarity is between treatments. 

3.7.2. MDS Analysis 

Multidimensional scaling (MDS) highlighted the dissimilarities among banks, with depths within 

sites grouping more closely (Figure 3.7.2). The stress level was low at 0.05, indicating high 

confidence in the relationships displayed. MDS plots showed transect-level data also grouped, 

with distinct clusters being formed within depth zones (Figure 3.7.3). McGrail Bank Transect 6 

(M46) at 45-50 m (148-164 ft) lies by itself (Figure 3.7.3), due to the high percentage of 

Groups 

(m) 

RStatistic Significance 

Level 

Possible 

Permutations 

Actual 

Permutations 

Number ≥

Observed 

45-50/ 

30-36.5 0.993 0.001 3185325 999 0 

45-50/ 

55-60 0.609 0.001 Very Large 999 0 

30-36.5/ 

55-60 0.996 0.001 19324305 999 0 
56 

Stephanocoenia intersepta in that transect (31.6%). The stress level of 0.13 indicates less 

confidence in the relationships than in Figure 3.7.2. 

Figure 3.7.2. Two-dimensional MDS plot, based on square-root-transformed Bray-Curtis 

similarities, comparing multivariate benthic cover data among Sonnier, 

McGrail, Geyer, and Bright Banks at the depth ranges sampled. Data were 

pooled within depths at each site. Abbreviations: S2, Sonnier Bank 22-27 m; 

S3, Sonnier Bank 30-36.5 m; S4, Sonnier Bank 45-50 m; S5, Sonnier Bank 

55-60 m; M4, McGrail Bank 45-50 m; M5, McGrail 55-60 m; G3, Geyer Bank 

30-36.5 m; G4, Geyer Bank 45-50 m; G5, Geyer Bank 55-60 m; B3, Bright 

Bank 30-36.5 m. 
57 

Figure 3.7.3. Two-dimensional MDS plot based on square-root-transformed Bray-Curtis 

similarity comparing multivariate benthic cover data between Sonnier, 

McGrail, Geyer, and Bright Banks by transect. Abbreviations were as follows. 

The first letter stands for the bank (see caption for Figure 3.7.2), and the first 

number stands for the depth range: 2= 22-27 m, 3= 30-36.5 m, 4= 45-50 m, 5= 

55-60 m. The second number is the transect number at that depth (e.g., S21, 

Sonnier Bank between 22-27 m, transect 1). 

3.8. EAST FLOWER GARDEN BANK REPETITIVE QUADRAT DATA

3.8.1. Study Site Repetitive Quadrat Percent Cover Data 

Forty repetitive quadrats (8-m2) were photographed at the EFGB in November 2005 during the 

post-hurricane assessment effort, and 39 repetitive quadrat photographs were taken during the 

annual monitoring in June 2006. Hard coral cover within the repetitive quadrats was stable 

between November 2005 and June 2006 at 61.34% ± 2.75 SE and 62.87% ± 2.32 SE, 

respectively. Macroalgal cover increased from November 2005 to June 2006 by ~5.4%, while 

CTB decreased by~6.9%. Species abundance showed stability with Montastraea annularis

species complex, Diploria strigosa, Porites astreoides and M. cavernosa as the four dominant 

species. Appendix 3 presents repetitive quadrat percent cover categories for November 2005 and 

June 2006. 
58 

Approximately 1.5% of coral colonies photographed within repetitive quadrats at the EFGB were 

missing in November 2005, most likely due to the effects of Hurricane Rita. This did not 

noticeably affect estimates of coral cover. Diploria strigosa, Porites astreoides, and 

Montastraea spp. comprised the majority of missing coral colonies, with sizes ranging from 

76.38-6498.76-cm2 (Table 3.8.3). Examples of coral colonies that disappeared or were damaged 

between June and November 2005 are presented in Figures 3.8.1-3.8.3. 

(A) (B) 

Figure 3.8.1. (A) Photograph of repetitive quadrat station in June 2005 at the EFGB. 

(B) Photograph of matching repetitive quadrat station in November 2005. The 

yellow boxes highlight the locations of likely hurricane impacts. 

(A) (B) 

Figure 3.8.2. (A) Photograph of repetitive quadrat station in June 2005 at the EFGB. 

(B) Photograph of matching repetitive quadrat station in November 2005. The 

yellow boxes highlight the locations of likely hurricane impacts. 
59 

(A) (B) 

Figure 3.8.3. (A) Photograph of repetitive quadrat station in June 2005 at the EFGB. 

(B) Photograph of matching repetitive quadrat station in November 2005. The 

yellow boxes highlight the locations of likely hurricane impacts. 

The most noticeable difference between repetitive quadrat photographs at the EFGB in 

November 2005 and June 2006 was the level of bleaching: 9.74% ± 1.07 SE in November 2005 

compared to 0.62% ± 0.24 in June 2006. Most bleaching was observed on colonies of 

Montastraea annularis species complex, M. cavernosa, and Millepora alcicornis (Table 3.8.1). 

Measurements of paling and fish biting were low at 1.5% or less, and disease was not observed 

(Table 3.8.2). The data from June 2006 showed a decrease in the amount of bleaching and 

paling compared to November 2005 and an increase in the amount of fish biting (Figure 3.8.4). 

Coral disease was absent from the analyzed photographs (Table 3.8.2). It should be noted that 

the identification of coral disease in photographs taken at a distance of 2-m from the substrate is 

unreliable (Zimmer, personal communication, 2005). 

Table 3.8.1. 

Percent cover by coral species of isolated fish biting (IFB), concentrated fish biting (CFB), 

paling (P), and bleaching (BL) at the EFGB in November 2005 and June 2006. 

Data are from random-dot analysis of repetitive quadrats. 

EFGB November 2005 EFGB June 2006 

Coral Species IFB CFB P BL IFB CFB P BL 

Colpophyllia natans 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 

Diploria strigosa 0.000 0.000 0.126 0.173 0.000 0.000 0.000 0.096 

Millepora alcicornis 0.000 0.000 0.000 1.131 0.000 0.000 0.000 0.303 

Montastraea species complex 0.424 0.016 0.880 3.284 2.089 0.207 0.016 0.207 

Montastraea annularis 0.000 0.000 0.000 0.016 0.000 0.000 0.000 0.000 

Montastraea cavernosa 0.000 0.000 0.471 4.808 0.016 0.000 0.000 0.016 

Montastraea faveolata 0.016 0.000 0.079 0.581 0.080 0.032 0.000 0.016 

Montastraea franksi 0.408 0.016 0.801 2.687 2.009 0.175 0.016 0.191 

Porites astreoides 0.000 0.000 0.000 0.079 0.000 0.000 0.000 0.000 

Unidentifiable Coral 0.000 0.000 0.000 0.267 0.000 0.000 0.000 0.000 

Total 0.424 0.016 1.477 9.741 2.105 0.207 0.016 0.622 
60 

Table 3.8.2. 

Percent cover (±SE) of paling, bleaching, concentrated fish biting, isolated fish biting, and 

disease at the EFGB in November 2005 and June 2006. 

Data are from random-dot analysis of repetitive quadrats. 

Observation EFGB November 2005 EFGB June 2006 

Paling 1.48 ± 0.39 0.02 ± 0.01 

Bleaching 9.74 ± 1.07 0.62 ± 0.24 

Concentrated Fish Biting 0.02 ± 0.01 0.21 ± 0.07 

Isolated Fish Biting 0.42 ± 0.09 2.1 ± 0.30 

Disease 0 0 

Coral Condition at the East Flower Garden Bank 

November 2005 and June 2006

0

2

4

6

8

10

12

Paling Bleaching Concentrated Fish Biting Isolated Fish Biting Disease

Coral Condition

Percent Cover

Nov-05

Jun-06

Figure 3.8.4. Percent cover of coral-condition categories. Error bars represent one 

standard error. 

3.8.2. Study Site Repetitive Quadrat Planimetry 

Planimetry data from the repetitive quadrats showed stability in lateral growth of the 

Montastraea annularis species complex from June 2005 to November 2005 and from November 

2005 to June 2006 (Figure 3.8.5). Appendix 4 contains the planimetry data for coral colonies in 

the repetitive quadrats from June 2005-June 2006. 

Twenty-one coral heads from the 40 repetitive quadrats (8-m2) photographed were removed 

between June 2005 and November 2005. Measurements of all missing corals were made from 
61 

June 2005 photographs to obtain a total area of living coral that had been lost. An estimated total 

area of ~3-m2 of coral was removed from the repetitive quadrat stations between June and 

November 2005 (Table 3.8.3). This represents a loss of ~1.5% of coral colonies photographed 

within repetitive quadrat stations at the EFGB, most likely due to the effects of Hurricane Rita. 

Mean Proportional Change in M. annularis Species Complex

Study Site Repetitive Quadrats 

-0.15

-0.10

-0.05

0.00

0.05

0.10

0.15

0.20

Mean Proportional Change

June 2005 - November 2005

November 2005 - June 2006

Figure 3.8.5. Mean proportional change in the planar tissue area of Montastraea 

annularis species complex within repetitive quadrat stations at the 

EFGB between June 2005 and November 2005, and between 

November 2005 and June 2006. Error bars represent one standard 

error. 

3.8.3. Deep Station Repetitive Quadrat Percent Cover Data 

Nine deep stations were analyzed from November 2005 repetitive quadrat (8-m2) photographs, 

revealing that 2 coral colonies (0.5% of coral colonies photographed at the deep stations) were 

missing. Analysis of the deep quadrats revealed less bleaching than in the shallower quadrats in 

November 2005 (3.05% vs. 9.74%). In 2006, bleaching in the deep station quadrats decreased 

nearly to zero (0.24%), whereas fish biting increased during the same period (Figure 3.8.6). 
62 

Table 3.8.3. 

Area of coral colonies missing in November 2005 photographs at the EFGB. 

Asterisks (*) indicate coral colonies that were not used during the planimetry 

analysis. 

Repetitive 

Quadrat 

Station # Coral Species 

Area 

(cm2

) Area (m2) 

3 Diploria strigosa 2614.25 0.26 

4 Diploria strigosa 1290.08 0.13 

Montastraea sp. 566.48 0.06 

7 Porites astreoides 76.38 0.01 

Diploria strigosa 254.46 0.03 

Diploria strigosa * 6498.76 0.65 14b Diploria strigosa 582.75 0.06 

Madracis decactis * 1507.98 0.15 18 Porites astreoides * 461.67 0.05 

20b Montastraea sp 1335.54 0.13 

21b Diploria strigosa * 667.53 0.07 

Porites astreoides * 246.25 0.02 22 Porites astreoides * 113.94 0.01 

26 Diploria strigosa 1669.04 0.17 

Diploria strigosa 3576.90 0.36 29 Diploria strigosa 1336.90 0.13 

29b Diploria strigosa 1102.40 0.11 

Diploria strigosa 5780.06 0.58 30b Diploria strigosa 396.64 0.04 

UNK1 Diploria strigosa 1706.04 0.17 

UNK12 Montastraea 

cavernosa * 219.03 0.02 

 Total 32003.07 3.20 

3.9. PERIMETER VIDEOGRAPHY

Corals observed along perimeter lines (east and north) at the EFGB in November 2005 showed 

evidence of hurricane impacts (i.e., dislodgement, loss, or deposition of entire coral heads; 

breaking of corals; and abrasion on the reef) that were not present in the June 2005 perimeter 

video. Most distressed corals observed in June 2005 were affected by fish biting, with fewer 

incidences of paling and bleaching. Increased levels of coral stress, manifested as paling and 

bleaching, were documented in November 2005. The footage from June 2006 showed lower 

levels of paling and bleaching and an increase in fish biting compared to November 2005. No 

evidence of disease was observed in June 2005, November 2005, or June 2006 along perimeter 

lines. Despite hurricane damage and moderate levels of corals stress, the coral population at the 

EFGB is stable and healthy. 
63 

Coral Condition at East Flower Garden Bank Deep Stations

November 2005 and June 2006

0

0.5

1

1.5

2

2.5

3

3.5

4

4.5

Paling Bleaching Concentrated Fish Biting Isolated Fish Biting Disease

Observation

Percent Cover

Nov-05

Jun-06

Figure 3.8.6. Coral condition at the EFGB deep stations in November 2005 and 

June 2006. Error bars represent one standard error. 

3.9.1. Perimeter Lines 

No incidences of disease were observed at the EFGB in November 2005. Concentrated fish 

biting was documented on two colonies, isolated fish biting on five colonies, paling on three 

colonies, and bleaching on 40 colonies along the north and east perimeter lines in November 

2005. Montastraea franksi and M. cavernosa were the two coral species most affected (Table 

3.9.1). 

In June 2006 no incidences of disease were observed at the EFGB. Stresses included 

concentrated fish biting in eight colonies, isolated fish biting in four colonies, paling in five 

colonies, and bleaching in 12 colonies. Montastraea franksi and M. cavernosa again were the 

two coral species most affected (Table 3.9.1). 

3.9.2. East Flower Garden Bank 360° Panoramic Views 

At the northwest corner of the study site, 16 coral colonies exhibited bleaching in November 

2005. In June 2006, the majority of these colonies (11) appeared to have recovered from 

bleaching. The remaining colonies were still bleached (1), exhibited paling (1), or were replaced 

by turf algae (3) (Table 3.9.2). 
64 

Table 3.9.1. 

Comparison of observations of the condition of individual coral colonies at the EFGB along 

perimeter lines and in 360o panoramic views between November 2005 (n = 815 colonies) and 

June 2006 (n = 736 colonies). 

(CFB = Concentrated fish biting, IFB = Isolated fish biting, B = Bleaching, 

P = Paling, H = Healthy colony, IIFB = increased tissue loss due to isolated fish 

biting, GI = Growth infilling [tissue regrowth], U = Unchanged condition). 

Number of 

Colonies Coral Species 

EFGB 

November 2005 

EFGB 

June 2006 

1 Diploria strigosa P H 

3 Diploria strigosa B H 

2 Diploria strigosa H CFB 

1 Diploria strigosa H P 

4 Millepora alcicornis B H 

2 Millepora alcicornis B U 

1 Montastraea annularis B H 

2 Montastraea annularis H CFB 

9 Montastraea cavernosa B H 

4 Montastraea cavernosa B U 

2 Montastraea cavernosa B P 

3 Montastraea faveolata B H 

1 Montastraea faveolata H P 

1 Montastraea faveolata H CFB 

1 Montastraea faveolata P H 

2 Montastraea franksi CFB GI 

11 Montastraea franksi B H 

3 Montastraea franksi IFB IIFB 

3 Montastraea franksi H B 

1 Montastraea franksi B P 

1 Montastraea franksi P H 

3 Montastraea franksi H CFB 

1 Montastraea franksi H IFB 

2 Montastraea franksi IFB GI 

3 Porites astreoides H B 
65 

Table 3.9.2. 

Affected coral colonies at the northwest corner of the study site in November 2005 and June 

2006. 

(B = Bleaching, P = Paling, H = Healthy colony, T= turf algae, U = Unchanged 

condition) 

Northwest Corner 

Number of Colonies Coral Species EFGB 

November 2005 

EFGB 

June 2006

2 Montastraea cavernosa B H 

3 Unidentifiable B H 

3 Montastraea franksi B H 

1 Montastraea sp. B U 

1 Diploria strigosa B H 

1 Montastraea sp. B P 

3 Millepora alcicornis B T 

1 Millepora alcicornis B H 

1 Colpophyllia natans B H 

At the northeast corner, several coral colonies exhibited bleaching, whereas only one colony 

exhibited isolated fish biting in November 2005 (Table 3.9.3). The northeast corner marker was 

not in place in November 2005 or June 2006. Therefore, the “corner” shifted from year to year 

and the same coral colonies were not videotaped. Table 3.9.3, therefore, underestimates the 

numbers of affected coral colonies at the northeast corner. 

At the southeast corner, several coral colonies that exhibited bleaching in November 2005 

appeared healthy in June 2006. A single, unidentifiable colony appeared healthy in November 

2005 but exhibited paling in June 2006. Similar to the northeast corner, the southeast corner 

marker was missing in November 2005 and June 2006. Therefore the “corner” shifted from year 

to year and the same coral colonies were not videotaped. Table 3.9.4, therefore, underestimated 

the numbers of affected coral colonies at the southeast corner. 

3.10. RESULTS OF HYDRODYNAMIC MODEL PREDICTIONS 

The REF/DIF model was run for Bright Bank, 47-mi (75-km) away from the track of Hurricane 

Rita to determine the potential effects of the bank on waves generated by the storm. Results of 

this modeling exercise showed that wave heights increased dramatically in the vicinity of the 

banks due to the large difference in water depth between the seabed and the summit of the bank. 

The model further showed that wave height, the distance from trough to crest, would have been 

highest directly over the peak of the bank. Per wave theory, the modeled bottom velocity is 

higher at the summit of the bank and decreases along the bank slope as depth increases. 

As discussed in Section 2.7, the guidance provided by the modeling effort suggests that wave 

heights are potentially increased to just below the wave-breaking point in the vicinity of the 

banks. Therefore, the wave heights from the hindcast study were increased to just below 
66 

breaking (approximately 60% of the water depth at each bank location) in order to estimate the 

water velocities induced by the waves on the banks. These transformed waves were then input 

into the Stream Function Wave Theory program, discussed in Section 2.7, to estimate water 

velocity. 

At Sonnier Bank, the transformed significant wave height during the passage of Hurricane Rita 

was estimated as 43-ft (13-m) (Figure 3.10.1) and the maximum bottom velocity predicted was 

4-m/s or ~8 knots (Table 3.10.1). Higher waves and water velocities may have occurred at 

deeper banks (Figure 3.10.2 A-D, Table 3.10.1). 

Table 3.9.3. 

Affected coral colonies at the northeast corner of the study site in November 2005 

and June 2006. 

(B = Bleaching, P = Paling, H = Healthy colony, IFB = isolated fish biting, GI = 

Growth infilling [tissue regrowth]). 

Northeast Corner 

Number of Colonies Coral Species EFGB 

November 2005 

EFGB 

June 2006

1 Montastraea faveolata IFB GI 

2 Diploria strigosa B H 

1 Millepora alcicornis B H 

1 Montastraea cavernosa B H 

2 M. cavernosa B P 

1 M. faveolata B H 

1 Montastraea spp. B H 

Table 3.9.4. 

Affected coral colonies at the southeast corner in November 2005 and June 2006. 

(B = Bleaching, P = Paling, H = Healthy colony). 

Southeast Corner 

Number of 

Colonies Coral Species EFGB November 2005

EFGB 

June 2006

2 Montastraea franksi B H 

1 Unidentifiable B H 

1 Unidentifiable H P 
67 

Figure 3.10.1. Predicted wave velocities at Sonnier Bank during passage of 

Hurricane Rita. The x-axis represents depth and the z-axis 

represents height above and below sea level. The maximum 

wave height (H) predicted by the hindcast is 60% of the depth 

(h) of the top of bank. 
68 

(A) McGrail Bank 

(B) Geyer Bank 

Figure 3.10.2. Predicted wave velocities during passage of 

Hurricane Rita at (A) McGrail Bank, (B) Geyer 

Bank, (C) Bright Bank, and (D) EFGB. The 

x-axis represents depth and the z-axis represents 

height above and below sea level. The maximum 

wave height (H) predicted by the hindcast is 60% 

of the depth (h) of the top of bank. 
69 

(C) Bright Bank 

(D) EFGB 

Figure 3.10.2. Predicted wave velocities during passage of 

Hurricane Rita at (A) McGrail Bank, (B) Geyer 

Bank, (C) Bright Bank, and (D) EFGB. The 

x-axis represents depth and the z-axis represents 

height above and below sea level. The 

maximum wave height (H) predicted by the 

hindcast is 60% of the depth (h) of the top of 

bank (continued). 
70 

Table 3.10.1. 

Maximum bottom velocities found using stream wave theory. 

The maximum wave height (H) predicted by the hindcast is 60% of the 

depth (h) of the top of bank. 

Bank 

Maximum Bottom 

Velocity 

(m/s, H from hindcast) 

Maximum Bottom 

Velocity 

(m/s, H = 0.6h) 

Sonnier Bank 3.5 4 

McGrail Bank 2.5 4 

Geyer Bank 2 4 

Bright Bank 2.5 4.5 

EFGB 3.17 3.77 
71 

4.0 DISCUSSION 

4.1. HURRICANE EFFECTS 

The physical effects of hurricanes on shallow coral reefs have been well documented (Woodley 

et al. 1981; Rogers et al. 1991; Hughes and Connell 1999; Steneck and Lang 2003). Physical 

damage to shallow coral reefs can include breaking, smashing, tumbling, and scouring; the 

destruction of corals and other benthic organisms; and the removal of soft corals and algae 

(Woodley et al. 1981). Branching and plating corals are in particular jeopardy because of their 

delicate structure; however, many branching corals capitalize on disturbance for dispersal and 

asexual reproduction (Woodley et al. 1981; Highsmith 1982). In this way, hurricanes open large 

areas of substratum. These areas are quickly colonized by fast-growing filamentous algae and 

macroalgae to the exclusion of corals, which grow at a much slower rate (Woodley et al. 1981; 

Gardner et al. 2003). Subsequent to hurricane impacts, coral reefs have a greater rate of loss of 

coral cover than reefs that have not been impacted by hurricanes (Gardner et al. 2005). 

The effects of hurricanes on deep coral reefs (18-40 m or 59-131 ft), such as the East and West 

Flower Garden Banks, which are dominated by large colonies of massive corals and few 

branching corals, have not been well documented; although, hurricane impacts at the EFGB in 

November 2005 are described by Precht et al. (2008a) and were anecdotally described in 1980 

after Hurricane Allen (Combs, personal communication, 1989). Hurricane impacts observed at 

the EFGB in October 2005 included substantial mechanical damage, fractured and displaced 

corals, sediment-scoured corals, and corals scarred and dented by waterborne objects (Hickerson 

and Schmahl, personal communication, 2005). 

Although the water depth of the banks provides some protection, bioeroded coral colonies 

remain susceptible to breakage and dispersal from the waves and high water velocities associated 

with hurricanes. At the EFGB, repetitive quadrat data collected in November 2005 showed that 

62% of missing coral colonies were Diploria strigosa. D. strigosa colonies at the East and West 

Flower Garden Banks experience intense bioerosion, forming mushroom-shaped colonies 

(Figure 4.1.1). More research is needed to confirm this qualitative assessment of the 

susceptibility of D. strigosa to hurricane damage at the Flower Garden Banks. 

4.1.1. Beneficial Aspects of Hurricanes on Coral Reefs 

In contrast to the physically destructive aspects of hurricanes, cooling water temperatures 

associated with the passage of hurricanes has been shown to relieve coral bleaching and thereby 

benefit coral reefs (Manzello et al. 2007). During the summer of 2005, sea-surface temperatures 

were elevated (above average) in the Florida Reef Tract and the United States Virgin Islands 

(U.S.V.I.) (Manzello et al. 2007). Bleaching levels were similar on the Florida Reef and the 

U.S.V.I. through the end of September 2005. In October 2005, bleaching began to decline on the 

Florida Reef Tract due to Hurricane Rita and the cool water associated with the passage of that 

storm. Further cooling occurred on the Florida Reef Tract with the passage of Hurricane Wilma, 

which accelerated the recovery of bleached corals within ~2 weeks of its passage (Manzello et al. 
72 

2007). In contrast to the Florida Reef Tract, no hurricanes passed near the U.S.V.I. during the 

2005 hurricane season and bleaching continued unabated. Bleaching in the U.S.V.I. corals lasted 

well into 2006 (Manzello et al. 2007). 

Figure 4.1.1. Typical bioerosion pattern of Diploria strigosa, resulting in mushroom-shaped 

colony at the EFGB. Image courtesy of PBS&J. 

Hurricane Rita brought cooler water temperatures to the EFGB in late September 2005 (Figure 

4.1.2.). Sea temperatures had been near or above 30° C for approximately two months—from 

July 2005 until September 23 at 1933 hours local time - when temperature decreased abruptly by 

1.5° C. However, Figure 4.1.2 does show that the temperature does climb back up and remains 

above the mean from previous years for several months. Due to the elevated temperatures, coral 

bleaching occurred at the Flower Garden Banks during the summer and fall of 2005. Bleaching 

surveys completed in October 2005 by FGBNMS showed 48% of corals surveyed were bleached 

or partially bleached (Hickerson, personal communication, 2005). Repetitive quadrat data 

collected at the EFGB in November 2005 showed that the percent cover of bleached coral was 

9.74% ±1.07 SE. This is the highest level of bleaching recorded since the beginning of the longterm monitoring program (Gittings et al. 1992; Continental Shelf Associates Inc. 1996; Dokken 

et al. 1999; Dokken et al. 2001; Precht et al. 2006). Bleached corals at the EFGB had recovered 

by June 2006. 

4.1.2. Hurricane Effects at Banks Not Dominated by Corals 

Little is known about the effects of hurricanes on benthic communities not dominated by corals, 

such as those at Sonnier, McGrail, Geyer, and Bright Banks. This study suggests that hurricane 

impacts to algal and sponge dominated banks may be harder to detect more than a year after the 

hurricane has passed (20 months in this case). Since the passage of Hurricane Rita, algal 

communities have had time to re-grow and sponges may have healed/regenerated. Furthermore, 

corals, sponges, and other organisms that may have been removed from these banks during the 

hurricane were likely fractured and dispersed by the predicted water velocity of ~4-m/s or 8 

knots on the reef cap, leaving little or no detectable remnants 20 months after the storm. Vast 

areas of rubble, documented at Sonnier Bank and at deeper depths on McGrail and Geyer Banks 
73 

may have represented hurricane effects; however, this may not be the case as normal reef 

processes also create rubble zones on banks in the northwestern Gulf of Mexico (Rezak et al. 

1985). 

18.0

20.0

22.0

24.0

26.0

28.0

30.0

32.0

11-Mar26-Mar10-Apr25-Apr

10-May25-May

9-Jun

24-Jun

9-Jul

24-Jul

8-Aug

23-Aug

7-Sep

22-Sep

7-Oct

22-Oct

6-Nov

21-Nov

Date

Temperature (°C)

EFGB YSI 2005 EFGB Hobo 2005 Mean EFGB 1990-2005 WFGB Hobo 2005

18.0

20.0

22.0

24.0

26.0

28.0

30.0

32.0

11-Mar26-Mar10-Apr25-Apr

10-May25-May

9-Jun

24-Jun

9-Jul

24-Jul

8-Aug

23-Aug

7-Sep

22-Sep

7-Oct

22-Oct

6-Nov

21-Nov

Date

Temperature (°C)

EFGB YSI 2005 EFGB Hobo 2005 Mean EFGB 1990-2005 WFGB Hobo 2005

Figure 4.1.2. Water temperature at the Flower Garden Banks in 2005 and mean temperature at 

the EFGB from 1990 to 2005. Water temperature was measured using YSI 

6600 Series datasondes (YSI) and HoboTemp recorders (Hobo). A vertical line 

drawn through the temperature profiles shows the timing of the passage of 

Hurricane Rita near the Flower Garden Banks (September 23, 2005). The 

1990-1997 temperature data are from Gittings et al. 1992, Continental Shelf 

Associates 1996, and Dokken et al. 1999. The 1998-2005 data are from 

Dokken et al. 2003, Precht et al. 2006, and Precht et al. 2008b. Abbreviation: 

WFGB-West Flower Garden Bank. 

4.1.3. Hydrodynamic Modeling Predictions 

A wave study was conducted to better understand the wave and current conditions in the vicinity 

of Sonnier, McGrail, Geyer, Bright, and East Flower Garden Banks during the passage of 

Hurricane Rita. A numerical modeling effort (REF/DIF model) was employed to assess the 

effects of generalized bank bathymetry on storm waves propagating over the banks. It is 

important to note that this model does not take into account any bank-specific geometry; rather, 

the model utilizes a simplified bank shape. In addition, wave kinematics were calculated using 

stream function wave theory to more accurately estimate the bottom current velocity on the 

banks. This analysis provided a predicted water velocity of ~4-m/s (~8 knots) at all banks 

included in this study. 
74 

Table 4.1.3. 

Results of the hydrodynamic modeling effort in relation to bank depth and distance from the 

Hurricane Rita storm track.

 

The REF/DIF model wave height estimates (Table 4.1.3) represent the largest theoretical wave 

heights that may have occurred at each bank cap. Based on these results, larger wave heights 

may have occurred at banks with caps located in deeper water (i.e., McGrail and Geyer Banks). 

However, it is interesting to note that the analysis of transect video in this study did not show the 

greatest damages at McGrail and Geyer Banks, as might be expected by the results of the wave 

model. Furthermore, the shallowest banks included in this study (i.e., EFGB at 18-m and 

Sonnier Bank at 22-m depth) experienced the most observable damage, with qualitative 

observations of significant mechanical impacts at EFGB during post-hurricane assessments and 

the appearance of bare substrate and rubble fields in transect video from Sonnier Bank in 2007. 

While one might assume that the greatest damage would likely occur at banks located closest to 

the storm track, the results of this study do not support this assumption. While McGrail Bank is 

located closest to the storm track (12-km or 7-mi), no hurricane damages were observed at this 

bank. Also, the significant damages were noted at the EFGB, which is the bank in this study 

located furthest from the storm track (93-km or 58-mi). Significant damages were also detected 

at Sonnier Bank, which is the only bank in this study located east of the Hurricane Rita storm 

track. It is common knowledge that the right side of a hurricane (relative to its direction of 

travel) is the most powerful portion of the storm in terms of wind speed and storm surge 

(USDOC, NOAA 1999). This may be a contributing reason for the damages observed at Sonnier 

Bank. 

4.2. BENTHIC CHARACTERISTICS AND HURRICANE EFFECTS AT SONNIER, MCGRAIL, GEYER, AND 

BRIGHT BANKS

Sonnier, McGrail, Geyer, and Bright Banks are characterized by unique biological communities, 

despite their geographic and oceanographic proximity (Figure 4.2.1). This study found that for 

these banks, within bank variability was lower than among bank variability. Variability among 

transects within a depth range was lower than the variability among depth ranges, which was less 

than variability among banks, as shown by the MDS plots in Section 3.7. These results suggest 

that the variability of the benthic communities is scale-dependent. Variability of coral reef 

Bank Bank Depth (m) Distance from 

Hurricane Rita 

storm track 

(km) 

Maximum 

Bottom Velocity 

(m/s) 

Maximum 

Wave Height 

at Top of Bank 

(m) 

McGrail 45 12 4 26 

Sonnier 22 24 4 13 

Geyer 39 58 4 26 

Bright 33 75 4.5 20 

East Flower 

Garden Bank 18 93 3.77 13 
75 

assemblages has been shown to be scale-dependent by other researchers (O'Neill et al. 1986; 

Hatcher et al. 1987; Wiens 1989; Murdoch and Aronson 1999). The dissimilarity between 

Sonnier, McGrail, Geyer, and Bright Banks is likely related to the differing influences of 

oceanographic currents delivering propagules of benthic organisms (Lugo-Fernandez et al. 

2001). 

Figure 4.2.1. Location of Sonnier, McGrail, Geyer, Bright, and East Flower Garden 

Banks in the Gulf of Mexico. Black line shows the track of Hurricane 

Rita. 

Observed and simulated drift studies conducted in 1993, 1997, and 1998 suggested that the 

distribution of coral larvae is driven by five modes in the Flower Garden Banks region (LugoFernandez et al. 2001). Drifters released from the EFGB contacted 10 shallow topographic 

banks that are suitable for coral recruitment, based on their minimum depths being <180-ft 

(<55-m). The study showed that banks closest to the EFGB should have the highest coral cover 

because those close banks, which included McGrail, Geyer, Bright, and Sonnier Banks, were 

more likely to be contacted by drifters (simulated larvae) from the EFGB. The results of our 

study show that coral cover was indeed highest at Bright Bank (located ~25-km or ~16-mi east 

of the EFGB) and decreased in an eastward direction from Geyer (49.3-km or 31-mi), to McGrail 

(94.4-km or 59-mi), to Sonnier Bank (116.7-km or 73-mi) (Figure 4.2.2). Thus, our initial 

sampling of these banks supports the conclusions of Lugo-Fernandez et al. (2001). An additional 

bank that would be interesting to characterize from this standpoint is Rankin -1, which is 7-mi 

(11-km) from the EFGB and 171-ft (52-m) deep at its highest point. According to LugoFernandez et al. (2001), and if the results of this study are correct, Rankin-1 should have higher 

coral cover than Bright Bank. 
76 

Percent Coral Cover for All Depth Ranges

0

2

4

6

8

10

12

14

Bright Bank Geyer Bank McGrail Bank Sonnier Bank

Bank

Percent Cover

Figure 4.2.2. Total coral cover at each of four banks within the study, displayed from west to 

east. Error bars represent one standard error. 

4.2.1. Sonnier Bank 

As the shallowest bank located on the east side of the hurricane track, with hindcast water 

velocities up to 4-m/s (8-kn), and estimated waves up to 43-ft (13-m), Sonnier Bank likely 

sustained hurricane damage in late September 2005. Benthic survey results from 2007 showed 

that Sonnier Bank had lower live cover than any of the other banks in this study (Figure 3.1.1). 

At the shallowest depth (72-ft or 22-m) on Sonnier Bank, divers with previous experience at 

Sonnier Bank noted that the bank surface appeared very different from previous visits. Areas of 

bare bedrock appeared in places where live cover had been before (Hickerson, personal 

communication, 2007). Although they apparently were strongly affected by Hurricane Rita, 

algal and sponge populations at Sonnier Bank remain and if left undisturbed should recover to 

pre-hurricane conditions. 

Qualitative analysis of diver video from the reef cap (22-27 m or 72-89 ft) at Sonnier Bank from 

1996, 2002, and 2005 showed some differences in benthic cover compared to video taken in the 

same depth range in 2007. In 1996, video footage showed areas of Sonnier Bank covered with 

attached flora and fauna, mostly of algae and sponges. Common sponges were individuals of the 

genera Neofibularia and Ircinia. Notably, large Xestospongia muta colonies inhabited high 

points along the bank’s ridge (Figure 3.2.1.A). An estimated 50 Xestospongia colonies were 

recorded in video footage taken in 1996. In 2002, the substratum was covered in a mix of algae 

and sponges. Dominant genera were again Neofibularia and Ircinia. Only one colony of 

Xestospongia was captured on video in 2002 and it appeared to be in poor condition, with 
77 

discoloration and a loss of rugosity on its pinacoderm (Figure 3.2.1.B). No Xestospongia

colonies were present in the video footage from 2005 or 2007. 

A decline in sponge diversity and abundance has been recorded at different locations around the 

world in the last 30 years (Wulff 2006). Sponge disease has been implicated in at least some of 

these rapidly declining sponge populations (Webster 2007). Disease has been disproportionally 

reported in large sponge species, such as Xestospongia muta (Wulff 2006). This may be due to 

the fact that these large sponges take weeks to months to die from disease, and are therefore 

more easily documented (Wulff 2006). From 1996-2006, disease was reported across the 

Caribbean in X. muta in Belize (Paz 1997), Curacao (Nagelkerken et al. 2000), Mexico (Gammill 

and Fenner 2005), and the Florida Keys (Cowart et al. 2006). With the photographic record of 

an intact but deteriorating X. muta in 2002 (Figure 3.2.1.B) and sponge disease documented 

across the Caribbean during this time, it may be that a water-borne pathogen was responsible for 

the demise of X. muta populations at Sonnier Bank prior to 2002. 

4.2.2. McGrail, Geyer, Bright, and East Flower Garden Banks 

As shown in MDS plots, Sonnier, McGrail, Geyer, and Bright Banks supported a unique benthic 

assemblage of algae, sponges, and corals. The EFGB, which was approximately 58-mi (93-km) 

from the path of Hurricane Rita, sustained measurable damage including: toppled, broken and 

shattered corals; scouring; shifting of sand flats; and gouging of coral heads by waterborne 

projectiles. All of these damages were likely related to water movement associated with the 

Category 3 storm. With McGrail, Bright, and Geyer Banks located in between the EFGB and 

Sonnier Bank, similar wind and wave activity was hindcast to have acted upon these banks, 

dependent partially upon their depth (Section 3.10). 

The 45- to 50-m (148- to 164-ft) depth range at McGrail Bank contains large colonies of 

Stephanocoenia intersepta. The mean coral cover in this depth range was ~5% and consisted 

mostly of S. intersepta (4.8%). However, the percent cover of S. intersepta ranged from 0-32% 

within the 45- to 50-m transect data. An MDS analysis was performed on the 45- to 50-m 

transect data, based on multivariate cover of the four hard coral taxa. The MDS analysis 

revealed that there is a high variability among transects in the composition of the coral 

assemblage at McGrail Bank in this depth range (Figure 3.3.3). This suggests that the coral 

cover at McGrail Bank in the 45- to 50-m depth range is patchy in nature. 

Vast areas of algal nodules at depth are present at McGrail Bank. The depth of this bank, 148-ft 

(45-m), may have protected the scleractinians and algal nodules from hurricane damage, even 

though the hurricane essentially passed over the bank (12-km or 7-mi from the centerline of the 

storm track). Wave heights of up to 85-ft (26-m) were modeled to have passed over McGrail 

Bank (Section 3.10). Undoubtedly, the pressure and velocity of water during the storm affected 

the top of the bank, even at 148-ft (45-m). 

The absence of baseline (pre-hurricane) data with which to compare the observations and data 

gathered at McGrail Bank during this study prevents statistical comparisons and limits our ability 

to draw inferences from the data. Because ROV footage was taken at night, landscape-scale 

views were not possible to discern. It may be that hurricane damage was not detected for this 
78 

reason. The colonies that were observed appeared undamaged; however, the high variability of 

coral cover observed in the 12 transects in the 45- to 50-m (148- to 164-ft) depth range could 

have been a consequence of the hurricane. On the other hand, that high variability could reflect 

the patchiness of the benthic biota caused by variability of coral recruitment or post-settlement 

processes, or previous perturbations. 

Qualitative video from 2003 was analyzed for pre-hurricane conditions at Geyer and Bright 

Banks. The subject of both videos was the fish populations at these banks; therefore, a thorough 

characterization of the benthic community structure was not possible. Video from Bright Bank 

showed largely bare, uncolonized areas, with Lobophora variegata being the only discernible 

macroalgal genus. Underwater blasting at the site of a purported shipwreck was to have taken 

place at Bright Bank sporadically over the past 30 years (Hickerson, personal communication, 

2007). Video from Geyer Bank shot in 2003 showed coverage of the benthos by brown 

macroalgae, including Sargassum spp., which was the dominant macroalgae in 2007. From 

these pre-hurricane videos we can see: (1) that the unique biological characteristics were not 

established after the hurricane; and (2) that as of April-May of 2007 the benthic assemblages had 

recovered to some extent. The types of hurricane impacts observed at the EFGB were not 

observed at these other banks because they are not dominated by scleractinian corals. 

Qualitative observations made one month after the storm at the EFGB indicated the occurrence 

of substantial mechanical damage: fractured and displaced corals, sediment-scoured corals, and 

corals scarred by waterborne objects (Hickerson and Schmahl, personal communication, 2005). 

Data collected in November 2005 revealed that approximately 1.5% of coral colonies 

photographed within repetitive quadrats at the EFGB were missing, most likely due to the effects 

of Hurricane Rita. However, this did not noticeably affect estimates of coral cover. 

Data collected in June 2005, November 2005, and June 2006 at the EFGB revealed that coral 

cover within the long-term monitoring site remained relatively constant (62.78% ± 2.60, 61.34% 

± 2.75, 62.87% ± 2.32, respectively) and species relative abundance showed stability with 

Montastraea annularis species complex, Diploria strigosa, and Porites spp. as the dominant 

species (Figures 4.2.3 and 4.2.4; Precht et al. 2008a). In June 2005 macroalgae was high at 24%, 

while CTB was lower at 10%. After the hurricane, in November 2005, the inverse relationship 

between macroalgae and CTB was evident, with 24% CTB and 13% macroalgae (Precht et al. 

2008a). Macroalgae cover is seasonally influenced and with the passage of Hurricane Rita it is 

likely that macroalgae was removed from the substratum. Between November 2005 and June 

2006, macroalgae increased by ~5.4%, while CTB decreased by ~6.9%. 
79 

East Flower Garden Bank Study Site Repetitive Quadrat Functional Groups 

June 2005, November 2005, and June 2006

0

10

20

30

40

50

60

70

Total Coral Sponge Macroalgae CTB

Benthic Category

Percent Cover

Jun-05

Nov-05

Jun-06

Figure 4.2.3. Percent cover of functional groups from repetitive quadrat stations at the 

EFGB in June 2005, November 2005, and June 2006. Error bars represent 

one standard error. 
80 

Percent Cover of Dominant Coral Taxa at East Flower Garden Bank 

Study Site Repetitive Quadrat Stations

June 2005, November 2005, and June 2006

0

5

10

15

20

25

30

35

40

45

50

Colpophyllia

natans

Diploria

strigosa

Madracis spp. Millepora

alcicornis

Montastraea

annularis

complex

Montastraea

cavernosa

Porites

astreoides

Coral Species

Percent Cover

Jun-05

Nov-05

Jun-06

Figure 4.2.4. Percent cover of the dominant coral taxa at the EFGB repetitive quadrat 

stations in June 2005, November 2005, and June 2006. Error bars represent 

one standard error. 
81 

5.0 CONCLUSION 

The unique biological characteristics of the remote benthic communities at Sonnier, McGrail, 

Geyer, Bright, and East Flower Garden Banks highlight their intrinsic value within the 

northwestern Gulf of Mexico ecosystem. With predicted wave velocities of 8 knots acting on 

these banks during the passage of Hurricane Rita, the damage could have been catastrophic. The 

apparent recovery of corals from bleaching at the EFGB associated with the 2005 warming event 

was documented in the June 2006 photographic data. The dataset generated for Sonnier, 

McGrail, Geyer, and Bright Banks suggests that Sonnier Bank suffered a loss of benthic cover 

but is recovering, with algae and sponges continuing to dominate. It is, however, unknown what 

impacts can be attributed to prior disturbances, or the passage of the hurricane. No obvious 

hurricane damage was apparent at McGrail, Geyer, and Bright Banks and because these three 

banks are dominated by algae and sponges, any recovery from damage might have been 

completed in the 20 months following the storm. McGrail Bank, with its large Stephanocoenia 

intersepta colonies, had no apparent coral damage and may have been protected by its depth 

(148-ft or 45-m); however, the absence of baseline (pre-hurricane) data limits the conclusions 

that can be drawn from the data collected during this study. 

This dataset serves as a baseline survey across multiple depth ranges and can be used in the 

future to compare with follow up studies as well as studies at other locations throughout the 

region. Although some aspects of the dispersal of coral larvae have been explained (LugoFernandez et al. 2001), the dispersal of other benthic organisms in the northwestern Gulf of 

Mexico is not well understood. Study of the colonization of these banks by algae and sponges is 

one area that should be considered for future research. Resource managers should view the 

present study as a characterization of these banks, which have proven to be biologically unique 

from one another. Increased protection, combined with a long term monitoring of Sonnier Bank, 

would benefit documentation of recovery at a mid-shelf bank. Increased protection to Bright 

Bank would halt treasure salvage activities and stop destruction of the resource. Increased 

protection for all banks of interest could decrease the pressure on the resources and increase the 

resiliency of the banks to recover from natural events. 

 

Recommendations 

• Conduct additional dispersal or drift studies (especially a study of genetic connections) to 

understand the oceanographic dynamics that supply the banks with their unique non-coral 

benthic components. 

• Study fish populations of these banks, which appeared from qualitative, visual 

observations to be robust, but which were not within the scope of this study. 

Understanding the connectivity of fish populations would be particularly interesting and 

important. 

• Detailed surveys should be conducted at McGrail Bank, including sclerochronology of 

the Stephanocoenia intersepta colonies. 

• A GIS mapping initiative is recommended. Video transects shot by divers and ROV 

should be used to create a GIS benthic cover/habitat database for these banks and other 

areas. 
83 

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Appendix 1: List of Species

SONNIER BANK

Percent 

Cover

Standard 

Error

Percent 

Cover

Standard 

Error

Antipatharians

Antipatharian spp. 0.00 0.00 0.00 0.00

Stichopathes sp. 0.20 0.20 0.03 0.03

Brown Macroalgae

Dictyota spp. 8.43 1.93 13.48 0.99

Lobophora variegata 7.75 3.85 1.65 0.56

Padina spp. 0.34 0.24 0.03 0.03

Sargassum spp.* 0.00 0.00 0.00 0.00

Unidentified Brown Macroalgae 2.11 2.09 6.90 3.08

Corals

Millepora alcicornis 7.98 1.73 1.60 0.55

Octocorallia 0.00 0.00 0.00 0.00

Siderastrea sp.* 0.00 0.00 0.00 0.00

Stephanocoenia intersepta* 0.00 0.00 0.00 0.00

Coralline Algae

Coralline Algae 1.41 1.19 0.08 0.05

Other Live

Cerianthidea 0.00 0.00 0.00 0.00

Diadema antillarum 0.05 0.05 0.00 0.00

Fish 0.00 0.00 0.03 0.03

Hermodice sp.* 0.00 0.00 0.00 0.00

Gastropoda 0.25 0.15 0.03 0.03

Holothuroidea 0.05 0.05 0.00 0.00

 Actinopygia agassizii

Spirobranchus giganteus* 0.00 0.00 0.00 0.00

Red Macroalgae

Red Macroalgae 1.06 1.06 0.00 0.00

Sponges

Agelas clathrodes 0.61 0.09 0.03 0.03

Ircinia spp. 0.33 0.03 0.28 0.13

 Ircinia strobilina

Neofibularia nolitangere 0.46 0.44 0.50 0.33

Niphates erecta 0.05 0.05 0.28 0.14

Spheciospongia sp. 0.25 0.25 0.25 0.17

Unidentified Sponges 0.65 0.45 0.45 0.07

Turf Algae

Turf Algae 6.34 4.24 4.48 1.78

Total Live Cover 38.32 30.10

22-27 m 30-36.5 m

* Species observed but not sampled. 89
Appendix 1: List of Species

SONNIER BANK

Antipatharians

Antipatharian spp.

Stichopathes sp.

Brown Macroalgae

Dictyota spp.

Lobophora variegata

Padina spp.

Sargassum spp.*

Unidentified Brown Macroalgae

Corals

Millepora alcicornis

Octocorallia

Siderastrea sp.*

Stephanocoenia intersepta*

Coralline Algae

Coralline Algae

Other Live

Cerianthidea

Diadema antillarum

Fish

Hermodice sp.*

Gastropoda

Holothuroidea

 Actinopygia agassizii

Spirobranchus giganteus*

Red Macroalgae

Red Macroalgae

Sponges

Agelas clathrodes

Ircinia spp.

 Ircinia strobilina

Neofibularia nolitangere

Niphates erecta

Spheciospongia sp.

Unidentified Sponges

Turf Algae

Turf Algae

Total Live Cover

Percent 

Cover

Standard 

Error

Percent 

Cover

Standard 

Error

0.00 0.00 0.17 0.11

0.00 0.00 0.00 0.00

0.31 0.25 0.00 0.00

0.00 0.00 0.00 0.00

0.00 0.00 0.00 0.00

0.00 0.00 0.00 0.00

0.49 0.29 0.00 0.00

0.00 0.00 0.00 0.00

0.34 0.17 0.00 0.00

0.00 0.00 0.00 0.00

0.00 0.00 0.00 0.00

0.03 0.03 0.03 0.03

0.00 0.00 0.03 0.03

0.00 0.00 0.00 0.00

0.00 0.00 0.00 0.00

0.00 0.00 0.00 0.00

0.03 0.03 0.03 0.03

0.00 0.00 0.00 0.00

0.00 0.00 0.00 0.00

0.00 0.00 0.00 0.00

0.00 0.00 0.00 0.00

0.11 0.09 0.00 0.00

0.00 0.00 0.00 0.00

0.00 0.00 0.00 0.00

0.00 0.00 0.00 0.00

0.43 0.13 0.13 0.07

5.03 0.53 1.40 0.49

6.77 1.79

45-50 m 55-60 m

* Species observed but not sampled. 90
Appendix 1: List of Species

MCGRAIL BANK

Percent Cover Standard Error Percent Cover Standard Error

Antipatharian spp. 0.52 0.50 0.00 0.00

Dictyota spp. 5.88 1.36 0.76 0.47

Lobophora sp. 2.56 0.77 3.55 2.37

Sargassum spp. 1.10 0.27 0.00 0.00

Unidentified Brown Macroalgae 0.60 0.13 0.24 0.08

Agaricia spp. 0.08 0.04 0.00 0.00

Millepora alcicornis 0.59 0.25 0.00 0.00

Montastraea cavernosa 0.02 0.02 0.00 0.00

Siderastrea sp.* 0.00 0.00 0.00 0.00

Octocorallia 0.00 0.00 0.03 0.03

Stephanocoenia intersepta 4.75 2.65 0.00 0.00

Algal Nodules 2.44 0.50 6.51 0.46

Coralline Algae 2.79 0.70 2.95 1.06

Green Macroalgae 9.50 2.43 2.17 1.65

Halimeda sp. 0.00 0.00 0.03 0.03

Asteroidea/Ophiuroidea 0.02 0.02 0.06 0.04

Cerianthidea 0.00 0.00 0.19 0.10

Diadema antillarum 0.29 0.16 0.00 0.00

Fish 0.08 0.03 0.03 0.03

Gastropoda 0.02 0.02 0.00 0.00

Holothuroidea* 0.00 0.00 0.00 0.00

Red Macroalgae 4.48 1.61 0.00 0.00

Agelas clathrodes 0.19 0.08 0.00 0.00

Geodia spp.* 0.00 0.00 0.00 0.00

Ircinia spp. 0.26 0.18 0.00 0.00

Pseudoceratina crassa 0.04 0.03 0.00 0.00

Unidentified Sponges 1.02 0.53 0.33 0.11

Xestospongia muta 0.37 0.21 0.00 0.00

Turf Algae 0.00 0.00 0.52 0.27

Total Live Cover 37.60 17.37

45-50 m 55-60 m

Sponges

Other Live

Green Macroalgae

Corals

Brown Macroalgae

Antipatharians

Red Macroalgae

Coralline Algae

Turf Algae

* Species observed but not sampled. 91
Appendix 1: List of Species

GEYER BANK

Percent Cover Standard Error Percent Cover Standard Error

Antipatharians

Antipatharian spp.* 0.00 0.00 0.00 0.00

Stichopathes sp. 0.00 0.00 0.10 0.10

Brown Macroalgae

Dictyota spp. 2.68 0.43 8.30 1.01

Lobophora variegata 12.65 0.90 0.85 0.47

Padina spp. 0.03 0.03 0.05 0.05

Sargassum spp. 26.88 3.38 4.90 0.83

Unidentified Brown Macroalgae 0.08 0.04 8.35 1.21

Corals

Agaricia spp.* 0.00 0.00 0.00 0.00

Millepora alcicornis 9.15 0.68 2.20 0.92

Porites spp. 0.03 0.03 0.00 0.00

Stephanocoenia intersepta 0.33 0.14 0.20 0.14

Tubastraea coccinea 0.23 0.17 0.00 0.00

Unidentifed Scleractinia 0.18 0.08 0.10 0.06

Coralline Algae

Coralline Algae 3.43 0.53 0.80 0.73

Green Macroalgae

Green Macroalgae 0.00 0.00 0.00 0.00

Other Live

Fish 0.23 0.10 0.35 0.17

 Gymnothorax moringa

 Muraena retifera

Gastropoda 0.00 0.00 0.00 0.00

Spirobranchus giganteus* 0.00 0.00 0.00 0.00

Sponges

Agelas clathrodes 0.00 0.00 1.30 0.62

Aplysina spp. 0.23 0.10 0.00 0.00

 Aplysina fistularis

Axinellid sponge* 0.00 0.00 0.00 0.00

Clathria spp. 0.03 0.03 0.00 0.00

Cliona sp. 0.05 0.05 0.00 0.00

Neofibularia nolitangere 0.10 0.10 0.00 0.00

Niphates erecta* 0.00 0.00 0.00 0.00

Unidentifed Sponges 0.75 0.13 1.55 0.76

Xestospongia muta 0.08 0.08 0.65 0.39

Turf Algae

Turf Algae 3.18 0.81 0.65 0.33

Total Live Cover 60.25 30.35

30-36.5 m 45-50 m

* Species observed but not sampled. 92
Appendix 1: List of Species

GEYER BANK

Antipatharians

Antipatharian spp.*

Stichopathes sp.

Brown Macroalgae

Dictyota spp.

Lobophora variegata

Padina spp.

Sargassum spp.

Unidentified Brown Macroalgae

Corals

Agaricia spp.*

Millepora alcicornis

Porites spp.

Stephanocoenia intersepta

Tubastraea coccinea

Unidentifed Scleractinia

Coralline Algae

Coralline Algae

Green Macroalgae

Green Macroalgae

Other Live

Fish

 Gymnothorax moringa

 Muraena retifera

Gastropoda

Spirobranchus giganteus*

Sponges

Agelas clathrodes

Aplysina spp.

 Aplysina fistularis

Axinellid sponge*

Clathria spp.

Cliona sp.

Neofibularia nolitangere

Niphates erecta*

Unidentifed Sponges

Xestospongia muta

Turf Algae

Turf Algae

Total Live Cover

Percent Cover Standard Error

0.00 0.00

0.00 0.00

15.23 4.24

0.14 0.07

0.00 0.00

0.23 0.20

5.97 1.90

0.00 0.00

0.11 0.07

0.00 0.00

0.06 0.04

0.00 0.00

0.03 0.03

5.74 1.39

0.20 0.12

0.37 0.23

0.03 0.03

0.00 0.00

0.71 0.31

0.00 0.00

0.00 0.00

0.00 0.00

0.00 0.00

0.00 0.00

0.00 0.00

1.17 0.30

0.66 0.33

3.94 1.12

34.59

55-60 m

* Species observed but not sampled. 93
Appendix 1: List of Species

BRIGHT BANK

Percent Cover Standard Error

Stichopathes sp.* 0.00 0.00

Dictyota spp. 0.33 0.18

Lobophora sp. 14.20 3.53

Padina spp. 0.13 0.07

Sargassum spp. 0.80 0.61

Unidentified Brown Macroalgae 28.13 6.29

Diploria strigosa 0.87 0.87

Millepora alcicornis 6.93 2.38

Montastraea cavernosa 0.07 0.07

Stephanocoenia intersepta 0.47 0.29

Coralline Algae 7.07 1.62

Green Macroalgae 13.13 0.82

 Anadyomene sp.

 Ulva sp.

Fish 0.73 0.07

Gastropoda 0.07 0.07

Spirobranchus giganteus* 0.00 0.00

Red Macroalgae 0.07 0.07

Agelas clathrodes 0.27 0.13

Aplysina spp.* 0.00 0.00

Clathria spp. 0.20 0.20

Ircinia spp.* 0.00 0.00

Pseudoceratina crassa* 0.00 0.00

Spheciospongia sp. 0.07 0.07

Unidentified Sponges 0.13 0.13

Xestospongia muta 0.33 0.33

Turf Algae 12.13 1.33

Total Live Cover 86.13

Turf Algae

Red Macroalgae

Coralline Algae

30-36.5 m

Sponges

Other Live

Green Macroalgae

Corals

Brown Macroalgae

Antipatharians

* Species observed but not sampled. 94
Appendix 2: Transect Data

Sonnier Bank 22-27 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Mean

Standard 

Deviation

Standard 

Error

Millepora alcicornis 6.25 9.70 7.98 2.44 1.73

Total Coral 6.25 9.70 7.98 2.44 1.73Stichopathes sp. 0.00 0.40 0.20 0.28 0.20Total Antipatharians 0.00 0.40 0.20 0.28 0.20Agelas clathrodes 0.53 0.70 0.61 0.12 0.09Ircinia spp. 0.35 0.30 0.33 0.04 0.03Neofibularia nolitangere 0.03 0.90 0.46 0.62 0.44Niphates erecta 0.00 0.10 0.05 0.07 0.05Spheciospongia sp. 0.00 0.50 0.25 0.35 0.25Unidentified Sponges 1.10 0.20 0.65 0.64 0.45Total Sponge 2.00 2.70 2.35 0.49 0.35Total Turf Algae 10.58 2.10 6.34 5.99 4.24Dictyota sp. 10.35 6.50 8.43 2.72 1.93Unidentified Brown macroalgae 0.03 4.20 2.11 2.95 2.09Lobophora variegata 11.60 3.90 7.75 5.44 3.85Padina sp. 0.58 0.10 0.34 0.34 0.24Total Brown Macroalgae 22.55 14.70 18.63 5.55 3.93Total Red Macroalgae 2.13 0.00 1.06 1.50 1.06Diadema antillarum 0.00 0.10 0.05 0.07 0.05Holothuroidea 0.10 0.00 0.05 0.07 0.05Gastropoda 0.40 0.10 0.25 0.21 0.15Total Other Live 0.50 0.20 0.35 0.21 0.15ANTIPATHARIANSSPONGESCORALRED MACROALGAEOTHER LIVETURF ALGAEBROWN MACROALGAE

95
Appendix 2: Transect Data

Sonnier Bank 22-27 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Mean

Standard 

Deviation

Standard 

Error

Total Coralline Algae 0.23 2.60 1.41 1.68 1.19

TB 35.50 47.40 41.45 8.41 5.95Rubble 19.18 20.10 19.64 0.65 0.46Total TB and Rubble 54.68 67.50 61.09 9.07 6.41Total Sand 1.10 0.10 0.60 0.71 0.50Total Live Cover 44.23 32.40 38.31TOTAL 100 100 100SANDFINE TURF, BARECORALLINE ALGAE

96
Appendix 2: Transect Data

Sonnier Bank 30-36.5 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect 5 CORALMillepora alcicornis 1.40 0.60 0.00 3.20 0.60 Total Coral 1.40 0.60 0.00 3.20 0.60 ANTIPATHARIANSStichopathes sp. 0.00 0.00 0.00 0.00 0.00 Total Antipatharians 0.00 0.00 0.00 0.00 0.00 SPONGESAgelas clathrodes 0.20 0.00 0.00 0.00 0.00 Ircinia spp. 0.20 0.00 0.60 1.00 0.00 Neofibularia nolitangere 0.00 0.00 1.00 2.60 0.00 Niphates erecta 0.20 0.00 0.00 0.80 0.00 Spheciospongia sp. 0.00 0.00 0.00 1.40 0.40 Unidentified Sponges 0.40 0.20 0.60 0.20 0.40 Total Sponge 1.00 0.20 2.20 6.00 0.80 TURF ALGAETotal Turf Algae 16.40 2.80 0.20 2.20 3.00 BROWN MACROALGAEUnidentified Brown Macroalgae 0.20 0.00 21.60 19.20 7.60 Dictyota spp. 9.60 14.00 15.80 18.20 12.00 Lobophora variegata 0.20 0.60 1.60 0.20 0.40 Padina spp. 0.00 0.00 0.00 0.00 0.20 Total Brown Macroalgae 10.00 14.60 39.00 37.60 20.20 OTHER LIVEFish 0.00 0.20 0.00 0.00 0.00 Gastropoda 0.00 0.00 0.00 0.00 0.00 Total Other Live 0.00 0.20 0.00 0.00 0.00 

97
Appendix 2: Transect Data

Sonnier Bank 30-36.5 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect 5 CORALLINE ALGAETotal Coralline Algae 0.40 0.00 0.00 0.20 0.00 0FINE TURF, BARETB 27.00 57.20 11.40 12.60 19.20 3Rubble 42.20 24.40 47.00 37.80 55.80 2Total TB and Rubble 69.20 81.60 58.20 50.40 75.00 6SANDTotal Sand 1.60 0.00 0.40 0.40 0.40 1Total Live Cover 29.20 18.40 41.40 49.20 24.60 2TOTAL 100 100 100 100 100 1

98
Appendix 2: Transect Data

Sonnier Bank 30-36.5 meters

MAJOR CATEGORY (% of transect) Transect 7 Transect 8 Mean

Standard 

Deviation

Stan

Er

CORALMillepora alcicornis 0.00 3.60 1.60 1.56 0.55Total Coral 0.00 3.60 1.60 1.56 0.55ANTIPATHARIANSStichopathes sp. 0.00 0.20 0.03 0.07 0.03Total Antipatharians 0.00 0.20 0.03 0.07 0.03SPONGESAgelas clathrodes 0.00 0.00 0.03 0.07 0.03Ircinia spp. 0.20 0.00 0.28 0.35 0.13Neofibularia nolitangere 0.40 0.00 0.50 0.92 0.33Niphates erecta 0.20 1.00 0.28 0.40 0.14Spheciospongia sp. 0.00 0.20 0.25 0.49 0.17Unidentified Sponges 0.60 0.80 0.45 0.21 0.07Total Sponge 1.40 2.00 1.78 1.84 0.65TURF ALGAETotal Turf Algae 5.40 2.60 4.48 5.02 1.78BROWN MACROALGAEUnidentified Brown Macroalgae 1.40 1.20 6.90 8.71 3.08Dictyota spp. 13.60 10.40 13.48 2.81 0.99Lobophora variegata 3.40 2.60 1.65 1.58 0.56Padina spp. 0.00 0.00 0.03 0.07 0.03Total Brown Macroalgae 18.40 14.20 22.06 10.75 3.80OTHER LIVEFish 0.00 0.00 0.03 0.07 0.03Gastropoda 0.20 0.00 0.03 0.07 0.03Total Other Live 0.20 0.00 0.05 0.09 0.03

99
Appendix 2: Transect Data 

Sonnier Bank 30-36.5 meters

MAJOR CATEGORY (% of transect) Transect 7 Transect 8 Mean

Standard 

Deviation

Stan

Er

CORALLINE ALGAETotal Coralline Algae 0.00 0.00 0.08 0.15 0.05

FINE TURF, BARETB 24.80 39.60 28.90 15.66 5.54Rubble 49.40 36.00 40.30 10.42 3.68Total TB and Rubble 74.20 75.60 69.20 10.20 3.61SANDTotal Sand 0.40 1.80 0.75 0.65 0.23Total Live Cover 25.40 22.60 30.05TOTAL 100 100 100

100
Appendix 2: Transect Data

Sonnier Bank 45-50 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect 5 TraOctocorallia 0.00 0.00 0.80 1.00 0.60 0.00 Total Coral 0.00 0.00 0.80 1.00 0.60 0.00 Ircinia spp. 0.00 0.00 0.20 0.00 0.00 0.60 Unidentified Sponges 0.60 0.60 0.20 0.00 1.00 0.20 Total Sponge 0.60 0.60 0.40 0.00 1.00 0.80 Total Turf Algae 4.60 2.60 6.80 4.40 5.80 4.80 Unidentified Brown Macroalgae 1.40 1.80 0.20 0.00 0.00 0.00 Dictyota spp. 0.00 0.00 1.80 0.20 0.20 0.00 Total Brown Macroalgae 1.40 1.80 2.00 0.20 0.20 0.00 Gastropoda 0.00 0.00 0.00 0.00 0.00 0.00 Total Other Live 0.00 0.00 0.00 0.00 0.00 0.00 Total Coralline Algae 0.00 0.00 0.00 0.20 0.00 0.00 TB 5.00 7.00 12.80 8.20 6.20 10.2Rubble 44.40 42.00 52.20 52.20 78.80 33.4Total TB and Rubble 49.40 49.00 65.00 60.40 85.00 43.6Total Sand 44.00 46.00 25.00 33.80 7.20 50.6Total Unknown 0.00 0.00 0.00 0.00 0.20 0.20 Total Live Cover 6.60 5.00 10.00 5.80 7.60 5.60 TOTAL 100 100 100 100 100 100 OTHER LIVEBROWN MACROALGAETURF ALGAESPONGESUNKNOWNSSANDFINE TURF, BARECORALLINE ALGAECORAL

101
Appendix 2: Transect Data

Sonnier Bank 45-50 meters

MAJOR CATEGORY (% of transect) Mean Standard Deviation Standard ECORALOctocorallia 0.34 0.44 0.17Total Coral 0.34 0.44 0.17SPONGESIrcinia spp. 0.11 0.23 0.09Unidentified Sponges 0.43 0.34 0.13Total Sponge 0.54 0.32 0.12TURF ALGAETotal Turf Algae 5.03 1.39 0.53BROWN MACROALGAEUnidentified Brown Macroalgae 0.49 0.77 0.29Dictyota spp. 0.31 0.66 0.25Total Brown Macroalgae 0.80 0.89 0.34OTHER LIVEGastropoda 0.03 0.08 0.03Total Other Live 0.03 0.08 0.03CORALLINE ALGAETotal Coralline Algae 0.03 0.08 0.03FINE TURF, BARETB 9.77 4.83 1.83Rubble 47.00 16.95 6.41Total TB and Rubble 56.77 14.75 5.57SANDTotal Sand 36.40 15.71 5.94UNKNOWNSTotal Unknown 0.06 0.10 0.04Total Live Cover 6.77TOTAL 100

102
Appendix 2: Transect Data

Sonnier Bank 55-60 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect ANTIPATHARIANSAntipatharia spp. 0.00 0.00 0.00 0.40 0.60 Total Antipatharian 0.00 0.00 0.00 0.40 0.60 SPONGESUnidentified Sponges 0.20 0.00 0.00 0.20 0.40 Total Sponge 0.20 0.00 0.00 0.20 0.40 TURF ALGAETotal Turf Algae 3.80 1.00 0.60 0.60 1.20 OTHER LIVECerianthidea 0.20 0.00 0.00 0.00 0.00 Gastropoda 0.00 0.00 0.20 0.00 0.00 Total Other Live 0.20 0.00 0.20 0.00 0.00 CORALLINE ALGAECrustose Coralline Algae 0.20 0.00 0.00 0.00 0.00 Total Coralline Algae 0.20 0.00 0.00 0.00 0.00 FINE TURF, BARETB 2.60 2.20 1.40 1.00 1.20 Rubble 13.80 12.20 10.40 6.40 5.00 Total TB and Rubble 16.40 14.40 11.80 7.40 6.20 SANDFish Hole 0.00 0.00 0.20 0.60 0.00 Sand 79.20 84.60 87.20 90.80 91.60 Total Sand 79.20 84.60 87.40 91.40 91.60 Total Live Cover 4.40 1.00 0.80 1.20 2.20 TOTAL 100 100 100 100 100 

103
Appendix 2: Transect Data

Sonnier Bank 55-60 meters

MAJOR CATEGORY (% of transect) Mean Standard Deviation Standard ErrANTIPATHARIANSAntipatharia spp. 0.17 0.27 0.11Total Antipatharian 0.17 0.27 0.11SPONGESUnidentified Sponges 0.13 0.16 0.07Total Sponge 0.13 0.16 0.07TURF ALGAETotal Turf Algae 1.40 1.21 0.49OTHER LIVECerianthidea 0.03 0.08 0.03Gastropoda 0.03 0.08 0.03Total Other Live 0.07 0.10 0.04CORALLINE ALGAECrustose Coralline Algae 0.03 0.08 0.03Total Coralline Algae 0.03 0.08 0.03FINE TURF, BARETB 1.40 0.92 0.38Rubble 9.37 3.39 1.39Total TB and Rubble 10.77 4.09 1.67SANDFish Hole 0.13 0.24 0.10Sand 87.30 4.76 1.94Total Sand 87.43 4.85 1.98Total Live Cover 1.80TOTAL 100

104
Appendix 2: Transect Data

McGrail Bank 45-50 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect 5 TrCORALAgaricia sp. 0.00 0.00 0.00 0.00 0.00 0.46 Millepora alcicornis 0.00 0.45 1.11 1.11 0.00 1.84 Montastraea cavernosa 0.00 0.00 0.22 0.00 0.00 0.00 Stephanocoenia intersepta 0.00 0.22 12.03 6.22 2.32 31.5Total Coral 0.00 0.67 13.36 7.33 2.32 33.8ANTIPATHARIANSAntipatharia spp. 0.00 0.00 0.00 0.00 0.00 0.00 Total Antipathrians 0.00 0.00 0.00 0.00 0.00 0.00 SPONGESAgelas clathrodes 0.00 0.00 0.45 0.00 0.21 0.92 Ircinia spp. 0.00 0.00 0.00 1.33 0.00 1.84 Pseudoceratina crassa 0.00 0.00 0.00 0.22 0.00 0.00 Unidentified Sponges 0.00 0.00 0.67 0.22 0.21 5.76 Xestospongia muta 0.00 0.00 0.67 0.00 1.90 1.84 Total Sponge 0.00 0.00 1.78 1.78 2.32 10.3BROWN MACROALGAEUnidentified Brown Macroalgae 0.68 0.67 0.67 0.67 0.21 0.00 Dictyota spp. 7.94 4.24 7.80 16.44 12.66 2.07 Lobophora variegata 3.40 0.89 1.56 0.44 0.00 0.00 Sargassum spp. 0.00 0.00 0.67 2.44 1.48 0.23 Total Brown Macroalgae 12.02 5.80 10.69 20.00 14.35 2.30 GREEN MACROALGAEGreen Macroalgae 5.90 11.61 18.04 12.89 31.22 1.84 Total Green Macroalgae 5.90 11.61 18.04 12.89 31.22 1.84 RED MACROALGAERed Macroalgae 0.91 3.35 0.00 0.00 0.00 0.00 Total Red Macroalgae 0.91 3.35 0.00 0.00 0.00 0.00 

105
Appendix 2: Transect Data

McGrail Bank 45-50 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect 5 TraOTHER LIVEDiadema antillarum 0.00 0.00 0.67 0.44 0.00 1.84 Fish 0.00 0.22 0.00 0.22 0.00 0.00 Asteroidea/Ophiuroidea 0.00 0.00 0.00 0.22 0.00 0.00 Gastropoda 0.00 0.00 0.00 0.00 0.00 0.23 Total Other Live 0.00 0.22 0.67 0.89 0.00 2.07 CORALLINE ALGAECrustose Coralline Algae 1.59 0.89 2.00 1.78 3.38 8.99 Algal Nodules 5.44 4.46 2.45 2.67 1.05 0.00 Total Coralline Algae 7.03 5.36 4.45 4.44 4.43 8.99 FINE TURF, BARETB 1.13 0.22 2.67 2.22 1.48 28.80 Rubble 47.39 53.57 30.07 34.44 27.64 4.38 Total TB and Rubble 48.53 53.79 32.74 36.67 29.11 33.18 SANDTotal Sand 25.17 19.20 18.04 15.78 16.24 7.37 UNKNOWNSTotal Unknown 0.45 0.00 0.22 0.22 0.00 0.00 Total Live Cover 25.85 27.01 49.00 47.33 54.64 59.45 TOTAL 100 100 100 100 100 100 

106
Appendix 2: Transect Data

McGrail Bank 45-50 meters

MAJOR CATEGORY (% of transect) Transect 8 Transect 9 Transect 10 Transect 11 TrCORALAgaricia sp. 0.23 0.00 0.00 0.22 0.00Millepora alcicornis 2.56 0.00 0.00 0.00 0.00Montastraea cavernosa 0.00 0.00 0.00 0.00 0.00Stephanocoenia intersepta 2.56 0.00 0.00 0.00 0.00Total Coral 5.36 0.00 0.00 0.22 0.00ANTIPATHARIANSAntipatharia spp. 0.00 0.00 0.00 0.22 0.00Total Antipathrians 0.00 0.00 0.00 0.22 0.00SPONGESAgelas clathrodes 0.00 0.22 0.00 0.00 0.00Ircinia spp. 0.00 0.00 0.00 0.00 0.00Pseudoceratina crassa 0.23 0.00 0.00 0.00 0.00Unidentified Sponges 3.73 0.00 0.00 0.00 0.23Xestospongia muta 0.00 0.00 0.00 0.00 0.00Total Sponge 3.96 0.22 0.00 0.00 0.23BROWN MACROALGAEUnidentified Brown Macroalgae 0.23 0.22 0.22 0.88 1.15Dictyota spp. 2.80 2.69 2.21 0.44 5.29Lobophora variegata 2.33 4.93 3.10 8.13 5.98Sargassum spp. 1.40 1.12 0.00 1.54 2.53Total Brown Macroalgae 6.76 8.97 5.53 10.99 14.9GREEN MACROALGAEGreen Macroalgae 6.53 3.14 1.77 3.96 7.36Total Green Macroalgae 6.53 3.14 1.77 3.96 7.36RED MACROALGAERed Macroalgae 4.90 5.38 15.27 12.97 10.8Total Red Macroalgae 4.90 5.38 15.27 12.97 10.8

107
Appendix 2: Transect Data

McGrail Bank 45-50 meters

MAJOR CATEGORY (% of transect) Transect 8 Transect 9 Transect 10 Transect 11 TrOTHER LIVEDiadema antillarum 0.47 0.00 0.00 0.00 0.00Fish 0.23 0.00 0.00 0.00 0.00Asteroidea/Ophiuroidea 0.00 0.00 0.00 0.00 0.00Gastropoda 0.00 0.00 0.00 0.00 0.00Total Other Live 0.70 0.00 0.00 0.00 0.00CORALLINE ALGAECrustose Coralline Algae 5.59 1.57 0.88 2.86 3.45Algal Nodules 0.47 1.35 1.99 1.76 4.83Total Coralline Algae 6.06 2.91 2.88 4.62 8.28FINE TURF, BARETB 8.86 2.91 2.21 2.64 2.99Rubble 38.23 49.78 50.44 44.62 43.6Total TB and Rubble 47.09 52.69 52.65 47.25 46.6SANDTotal Sand 18.65 26.68 21.90 19.78 11.4UNKNOWNSTotal Unknown 0.00 0.00 0.00 0.00 0.23Total Live Cover 34.27 20.63 25.44 32.97 41.6TOTAL 100 100 100 100 100

108
Appendix 2: Transect Data

McGrail Bank 45-50 meters

MAJOR CATEGORY (% of transect) Mean Standard Deviation Standard ErrorCORALAgaricia sp. 0.08 0.15 0.04Millepora alcicornis 0.59 0.88 0.25Montastraea cavernosa 0.02 0.06 0.02Stephanocoenia intersepta 4.75 9.17 2.65Total Coral 5.43 9.84 2.84ANTIPATHARIANSAntipatharia spp. 0.52 1.73 0.50Total Antipathrians 0.52 1.73 0.50SPONGESAgelas clathrodes 0.19 0.29 0.08Ircinia spp. 0.26 0.63 0.18Pseudoceratina crassa 0.04 0.09 0.03Unidentified Sponges 1.02 1.84 0.53Xestospongia muta 0.37 0.73 0.21Total Sponge 1.88 2.96 0.85BROWN MACROALGAEUnidentified Brown Macroalgae 0.60 0.46 0.13Dictyota spp. 5.88 4.73 1.36Lobophora variegata 2.56 2.65 0.77Sargassum spp. 1.10 0.92 0.27Total Brown Macroalgae 10.15 4.84 1.40GREEN MACROALGAEGreen Macroalgae 9.50 8.40 2.43Total Green Macroalgae 9.50 8.40 2.43RED MACROALGAERed Macroalgae 4.48 5.57 1.61Total Red Macroalgae 4.48 5.57 1.61

109
Appendix 2: Transect Data

McGrail Bank 45-50 meters

MAJOR CATEGORY (% of transect) Mean Standard Deviation Standard ErrorOTHER LIVEDiadema antillarum 0.29 0.55 0.16Fish 0.08 0.11 0.03Asteroidea/Ophiuroidea 0.02 0.06 0.02Gastropoda 0.02 0.07 0.02Total Other Live 0.40 0.62 0.18CORALLINE ALGAECrustose Coralline Algae 2.79 2.42 0.70Algal Nodules 2.44 1.72 0.50Total Coralline Algae 5.22 2.01 0.58FINE TURF, BARETB 4.76 7.88 2.27Rubble 39.26 13.73 3.96Total TB and Rubble 44.01 8.67 2.50SANDTotal Sand 18.31 5.32 1.54UNKNOWNSTotal Unknown 0.09 0.15 0.04Total Live Cover 37.58TOTAL 100

110
Appendix 2: Transect Data

McGrail Bank 55-60 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect 5 TrCORALOctocorallia 0.00 0.00 0.00 0.00 0.00 0.00 Total Coral 0.00 0.00 0.00 0.00 0.00 0.00 SPONGESUnidentified Sponges 0.00 0.22 0.00 0.00 0.49 0.7Total Sponge 0.00 0.22 0.00 0.00 0.49 0.7TURF ALGAETotal Turf Algae 0.00 0.22 0.24 0.00 0.25 0.00 BROWN MACROALGAEUnidentified Brown Macroalgae 0.45 0.00 0.48 0.00 0.25 0.4Dictyota spp. 0.45 0.00 0.00 0.00 0.00 0.00 Lobophora variegata 4.92 0.00 0.00 0.00 0.00 0.00 Total Brown Macroalgae 5.82 0.00 0.48 0.00 0.25 0.48 GREEN MACROALGAEGreen Macroalgae 13.65 1.55 0.00 0.00 0.25 0.4Halimeda sp. 0.00 0.00 0.24 0.00 0.00 0.00 Total Green Macroalgae 13.65 1.55 0.24 0.00 0.25 0.48 OTHER LIVECerianthidea 0.00 0.00 0.00 0.75 0.00 0.24 Fish 0.00 0.00 0.00 0.00 0.00 0.00 Asteroidea/Ophiuroidea 0.00 0.00 0.00 0.25 0.00 0.00 Total Other Live 0.00 0.00 0.00 1.00 0.00 0.24 CORALLINE ALGAECrustose coralline algae 4.03 0.00 0.96 1.25 1.72 0.9Algal Nodules 6.49 4.42 7.69 7.77 8.09 5.4Total Coralline Algae 10.51 4.42 8.65 9.02 9.80 6.43 

111
Appendix 2: Transect Data

McGrail Bank 55-60 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect 5 TraFINE TURF, BARETB 1.12 1.55 1.20 1.25 0.49 1.19 Rubble 46.09 50.22 51.92 60.40 69.61 66.90 Total TB and Rubble 47.20 51.77 53.13 61.65 70.10 68.10 SANDTotal Sand 22.82 41.59 37.02 28.32 18.87 23.57 UNKNOWNSTotal Unknown 0.00 0.22 0.24 0.00 0.00 0.00 Total Live Cover 29.98 6.42 9.62 10.03 11.03 8.33 TOTAL 100 100 100 100 100 100 

112
Appendix 2: Transect Data

McGrail Bank 55-60 meters

MAJOR CATEGORY (% of transect) Transect 8 Mean

Standard 

Deviation

Standard 

Error

CORALOctocorallia 0.00 0.03 0.09 0.03

Total Coral 0.00 0.03 0.09 0.03SPONGESUnidentified Sponges 0.48 0.33 0.32 0.11Total Sponge 0.48 0.33 0.32 0.11TURF ALGAETotal Turf Algae 1.67 0.52 0.75 0.27BROWN MACROALGAEUnidentified Brown Macroalgae 0.00 0.24 0.22 0.08Dictyota spp. 3.59 0.76 1.34 0.47Lobophora variegata 19.38 3.55 6.72 2.37Total Brown Macroalgae 22.97 4.54 7.90 2.79GREEN MACROALGAEGreen Macroalgae 1.20 2.17 4.67 1.65Halimeda sp. 0.00 0.03 0.08 0.03Total Green Macroalgae 1.20 2.20 4.66 1.65OTHER LIVECerianthidea 0.00 0.19 0.29 0.10Fish 0.24 0.03 0.08 0.03Asteroidea/Ophiuroidea 0.00 0.06 0.12 0.04Total Other Live 0.24 0.28 0.39 0.14CORALLINE ALGAECrustose coralline algae 8.37 2.95 3.00 1.06Algal Nodules 5.50 6.51 1.30 0.46Total Coralline Algae 13.88 9.46 3.12 1.10

113
Appendix 2: Transect Data

McGrail Bank 55-60 meters

MAJOR CATEGORY (% of transect) Transect 8 Mean

Standard 

Deviation

Standard 

Error

FINE TURF, BARETB 7.18 2.06 2.14 0.76

Rubble 43.54 54.99 9.57 3.38Total TB and Rubble 50.72 57.06 8.49 3.00SANDTotal Sand 8.61 25.46 10.33 3.65UNKNOWNSTotal Unknown 0.24 0.12 0.13 0.05Total Live Cover 40.43 17.37TOTAL 100 100

114
Appendix 2: Transect Data

Geyer Bank 30-36.5 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect 5 TrCORALMillepora alcicornis 7.80 8.80 8.60 8.40 10.00 7.00 Porites sp. 0.00 0.00 0.00 0.20 0.00 0.00 Unidentified Scleractinia 0.00 0.00 0.20 0.00 0.00 0.40 Stephanocoenia intersepta 0.80 0.40 0.40 0.00 0.00 0.00 Tubastraea coccinea 1.40 0.00 0.20 0.00 0.20 0.00 Total Coral 10.00 9.20 9.40 8.60 10.20 7.40 SPONGESAplysina spp. 0.40 0.60 0.00 0.00 0.00 0.20 Clathria spp. 0.00 0.00 0.00 0.00 0.00 0.00 Cliona sp. 0.00 0.00 0.40 0.00 0.00 0.00 Neofibularia nolitangere 0.00 0.00 0.80 0.00 0.00 0.00 Unidentified Sponges 1.20 1.20 0.40 0.20 0.80 0.60 Xestospongia muta 0.00 0.00 0.00 0.00 0.00 0.60 Total Sponge 1.60 1.80 1.60 0.20 0.80 1.40 TURF ALGAETotal Turf Algae 7.80 1.80 2.00 2.60 2.60 0.40 BROWN MACROALGAEUnidentified Brown Macroalgae 0.20 0.00 0.00 0.00 0.20 0.20 Dictyota spp. 3.60 3.00 1.00 3.60 1.60 2.80 Lobophora variegata 10.60 12.60 9.80 12.60 12.80 12.2Padina spp. 0.00 0.00 0.00 0.20 0.00 0.00 Sargassum spp. 7.00 25.60 30.00 32.40 27.00 38.4Total Brown Macroalgae 21.40 41.20 40.80 48.80 41.60 53.6OTHER LIVEFish 0.40 0.80 0.00 0.20 0.20 0.20 Total Other Live 0.40 0.80 0.00 0.20 0.20 0.20 

115
Appendix 2: Transect Data

Geyer Bank 30-36.5 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect 5 TrCORALLINE ALGAETotal Coralline Algae 5.00 1.40 2.20 2.00 3.60 3.60 FINE TURF, BARETB 25.60 32.80 31.40 21.20 19.20 19.2Rubble 27.20 8.80 7.80 13.20 20.40 13.8Total TB and Rubble 52.80 41.60 39.20 34.40 39.60 33.0SANDTotal Sand 0.80 2.00 4.80 3.00 1.40 0.40 UNKNOWNSTotal Unknown 0.20 0.20 0.00 0.20 0.00 0.00 Total Live Cover 46.20 56.20 56.00 62.40 59.00 66.6TOTAL 100 100 100 100 100 100 

116
Appendix 2: Transect Data

Geyer Bank 30-36.5 meters

MAJOR CATEGORY (% of transect) Transect 8 Mean Standard Deviation Standard CORALMillepora alcicornis 9.20 9.15 1.94 0.68Porites sp. 0.00 0.03 0.07 0.03Unidentified Scleractinia 0.60 0.18 0.23 0.08Stephanocoenia intersepta 0.00 0.33 0.40 0.14Tubastraea coccinea 0.00 0.23 0.48 0.17Total Coral 9.80 9.90 2.10 0.74SPONGESAplysina spp. 0.00 0.23 0.27 0.10Clathria spp. 0.20 0.03 0.07 0.03Cliona sp. 0.00 0.05 0.14 0.05Neofibularia nolitangere 0.00 0.10 0.28 0.10Unidentified Sponges 1.00 0.75 0.37 0.13Xestospongia muta 0.00 0.08 0.21 0.08Total Sponge 1.20 1.23 0.52 0.18TURF ALGAETotal Turf Algae 5.00 3.18 2.28 0.81BROWN MACROALGAEUnidentified Brown Macroalgae 0.00 0.08 0.10 0.04Dictyota spp. 4.40 2.68 1.22 0.43Lobophora variegata 18.40 12.65 2.56 0.90Padina spp. 0.00 0.03 0.07 0.03Sargassum spp. 33.20 26.88 9.56 3.38Total Brown Macroalgae 56.00 42.30 11.03 3.90OTHER LIVEFish 0.00 0.23 0.27 0.10Total Other Live 0.00 0.23 0.27 0.10

117
Appendix 2: Transect Data

Geyer Bank 30-36.5 meters

MAJOR CATEGORY (% of transect) Transect 8 Mean Standard Deviation Standard CORALLINE ALGAETotal Coralline Algae 3.80 3.43 1.51 0.53FINE TURF, BARETB 17.20 24.10 5.85 2.07Rubble 6.60 13.70 6.97 2.46Total TB and Rubble 23.80 37.80 8.24 2.91SANDTotal Sand 0.20 1.83 1.52 0.54UNKNOWNSTotal Unknown 0.20 0.13 0.10 0.04Total Live Cover 75.80 60.25TOTAL 100 100

118
Appendix 2: Transect Data

Geyer Bank 45-50 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Mean

Stan

Dev

CORALUnidentified Scleractinia 0.20 0.00 0.00 0.20 0.10 0.12 Millepora alcicornis 0.00 4.00 3.40 1.40 2.20 1.84 Stephanocoenia intersepta 0.60 0.00 0.20 0.00 0.20 0.28 Total Coral 0.80 4.00 3.60 1.60 2.50 1.54 ANTIPATHARIANSStichopathes sp. 0.00 0.40 0.00 0.00 0.10 0.20 Total Antipatharians 0.00 0.40 0.00 0.00 0.10 0.20 SPONGESAgelas clathrodes 0.00 1.80 2.80 0.60 1.30 1.25 Unidentified Sponges 0.60 1.80 3.60 0.20 1.55 1.53 Xestospongia muta 0.00 1.60 0.00 1.00 0.65 0.79 Total Sponge 0.60 5.20 6.40 1.80 3.50 2.74 TURF ALGAETotal Turf Algae 0.20 1.40 1.00 0.00 0.65 0.66 BROWN MACROALGAEUnidentified Brown macroalgae 6.40 11.80 7.00 8.20 8.35 2.42 Dictyota spp. 6.20 7.20 10.80 9.00 8.30 2.03 Lobophora variegata 0.60 0.00 2.20 0.60 0.85 0.94 Padina spp. 0.20 0.00 0.00 0.00 0.05 0.10 Sargassum spp. 7.20 5.00 3.60 3.80 4.90 1.65 Total Brown Macroalgae 20.60 24.00 23.60 21.60 22.45 1.62 OTHER LIVEFish 0.20 0.80 0.40 0.00 0.35 0.34 Total Other Live 0.20 0.80 0.40 0.00 0.35 0.34 CORALLINE ALGAETotal Coralline Algae 0.00 3.00 0.20 0.00 0.80 1.47 

119
Appendix 2: Transect Data

Geyer Bank 45-50 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Mean

Sta

Dev

FINE TURF, BARETB 31.60 22.80 37.00 32.80 31.05 5.97 Rubble 36.80 33.40 26.00 33.00 32.30 4.53 Total TB and Rubble 68.40 56.20 63.00 65.80 63.35 5.25 SANDTotal Sand 9.00 5.00 1.80 9.20 6.25 3.54 UNKNOWNSTotal Unknown 0.20 0.00 0.00 0.00 0.05 0.10 Total Live Cover 22.40 38.80 35.20 25.00 30.35TOTAL 100 100 100 100 100

120
Appendix 2: Transect Data

Geyer Bank 55-60 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect 5 TrCORALMillepora alcicornis 0.00 0.40 0.00 0.40 0.00 0.00 Unidentified Scleractinia 0.20 0.00 0.00 0.00 0.00 0.00 Stephanocoenia intersepta 0.20 0.00 0.00 0.20 0.00 0.00 Total Coral 0.40 0.40 0.00 0.60 0.00 0.00 SPONGESAgelas clathrodes 0.00 0.20 1.00 0.40 2.40 0.40 Unidentified Sponges 0.00 1.40 0.40 1.60 1.20 1.20 Xestospongia muta 0.00 0.00 2.00 1.00 1.60 0.00 Total Sponge 0.00 1.60 3.40 3.00 5.20 1.60 TURF ALGAETotal Turf Algae 0.60 2.00 2.20 3.80 9.00 6.80 BROWN MACROALGAEUnidentified Brown Macroalgae 1.20 2.60 15.60 8.20 1.60 5.80 Dictyota spp. 1.40 4.20 12.00 14.00 17.40 34.2Lobophora variegata 0.40 0.00 0.00 0.40 0.00 0.00 Sargassum spp. 0.00 0.00 0.20 1.40 0.00 0.00 Total Brown Macroalgae 3.00 6.80 27.80 24.00 19.00 40.0GREEN MACROALGAEGreen Macroalgae 0.80 0.20 0.00 0.00 0.40 0.00 Total Green Macroalgae 0.80 0.20 0.00 0.00 0.40 0.00 OTHER LIVEFish 0.00 0.00 0.00 0.00 0.20 1.60 Gastropoda 0.20 0.00 0.00 0.00 0.00 0.00 Total Other Live 0.20 0.00 0.00 0.00 0.20 1.60 CORALLINE ALGAETotal Coralline Algae 0.80 5.00 7.20 6.60 10.40 9.00 

121
Appendix 2: Transect Data

Geyer Bank 55-60 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Transect 4 Transect 5 TraFINE TURF, BARETB 7.00 11.00 22.60 9.60 9.20 12.00 Rubble 75.20 63.80 28.00 44.20 34.20 18.20 Total TB and Rubble 82.20 74.80 50.60 53.80 43.40 30.20 SANDTotal Sand 11.80 9.00 8.40 7.80 12.40 10.60 UNKNOWNSTotal Unknown 0.20 0.20 0.40 0.40 0.00 0.20 Total Live Cover 5.80 16.00 40.60 38.00 44.20 59.00 TOTAL 100 100 100 100 100 100 

122
Appendix 2: Transect Data

Geyer Bank 55-60 meters

MAJOR CATEGORY (% of transect) Mean Standard Deviation Standard ErrorCORALMillepora alcicornis 0.11 0.20 0.07Unidentified Scleractinia 0.03 0.08 0.03Stephanocoenia intersepta 0.06 0.10 0.04Total Coral 0.20 0.26 0.10SPONGESAgelas clathrodes 0.71 0.81 0.31Unidentified Sponges 1.17 0.79 0.30Xestospongia muta 0.66 0.87 0.33Total Sponge 2.54 1.66 0.63TURF ALGAETotal Turf Algae 3.94 2.95 1.12BROWN MACROALGAEUnidentified Brown Macroalgae 5.97 5.03 1.90Dictyota spp. 15.23 11.23 4.24Lobophora variegata 0.14 0.19 0.07Sargassum spp. 0.23 0.52 0.20Total Brown Macroalgae 21.57 13.11 4.96GREEN MACROALGAEGreen Macroalgae 0.20 0.31 0.12Total Green Macroalgae 0.20 0.31 0.12OTHER LIVEFish 0.37 0.62 0.23Gastropoda 0.03 0.08 0.03Total Other Live 0.40 0.60 0.23CORALLINE ALGAETotal Coralline Algae 5.74 3.67 1.39

123
Appendix 2: Transect Data

Geyer Bank 55-60 meters

MAJOR CATEGORY (% of transect) Mean Standard Deviation Standard ErrorFINE TURF, BARETB 11.69 5.06 1.91Rubble 43.34 19.99 7.56Total TB and Rubble 55.03 17.91 6.77SANDTotal Sand 10.09 1.74 0.66UNKNOWNSTotal Unknown 0.29 0.20 0.07Total Live Cover 34.60TOTAL 100

124
Appendix 2: Transect Data

Bright Bank 30-36.5 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Mean Standard Deviation CORALDiploria strigosa 0.00 2.60 0.00 0.87 1.50 Millepora alcicornis 7.40 10.80 2.60 6.93 4.12 Montastraea cavernosa 0.00 0.20 0.00 0.07 0.12 Stephanocoenia intersepta 0.00 1.00 0.40 0.47 0.50 Total Coral 7.40 14.60 3.00 8.33 5.86 SPONGESAgelas clathrodes 0.40 0.40 0.00 0.27 0.23 Clathria spp. 0.00 0.60 0.00 0.20 0.35 Spheciospongia sp. 0.00 0.20 0.00 0.07 0.12 Unidentified Sponges 0.00 0.40 0.00 0.13 0.23 Xestospongia muta 0.00 1.00 0.00 0.33 0.58 Total Sponge 0.40 2.60 0.00 1.00 1.40 TURF ALGAETotal Turf Algae 10.80 14.80 10.80 12.13 2.31 BROWN MACROALGAEUnidentified Brown Macroalgae 25.80 18.60 40.00 28.13 10.89 Dictyota spp. 0.40 0.60 0.00 0.33 0.31 Lobophora variegata 8.40 13.60 20.60 14.20 6.12 Padina spp. 0.20 0.00 0.20 0.13 0.12 Sargassum spp. 0.40 0.00 2.00 0.80 1.06 Total Brown Macroalgae 35.20 32.80 62.80 43.60 16.67 GREEN MACROALGAEGreen Macroalgae 14.40 13.40 11.60 13.13 1.42 Total Green Macroalgae 14.40 13.40 11.60 13.13 1.42 RED MACROALGAERed Macroalgae 0.20 0.00 0.00 0.07 0.12 Total Red Macroalgae 0.20 0.00 0.00 0.07 0.12 

125
Appendix 2: Transect Data 

Bright Bank 30-36.5 meters

MAJOR CATEGORY (% of transect) Transect 1 Transect 2 Transect 3 Mean Standard DeviatiOTHER LIVEFish 0.80 0.80 0.60 0.73 0.12 Gastropoda 0.00 0.00 0.20 0.07 0.12 Total Other Live 0.80 0.80 0.80 0.80 0.00 CORALLINE ALGAETotal Coralline Algae 7.20 9.80 4.20 7.07 2.80 FINE TURF, BARETB 19.60 10.80 6.40 12.27 6.72 Rubble 3.20 0.40 0.40 1.33 1.62 Total TB and Rubble 22.80 11.20 6.80 13.60 8.27 SANDTotal Sand 0.60 0.00 0.00 0.20 0.35 UNKNOWNSTotal Unknown 0.20 0.00 0.00 0.07 0.12 Total Live Cover 76.40 88.80 93.20 86.13TOTAL 100 100 100 100

126
Appendix 3: Repetitive Quadrat Percent Cover Data

Study Site Repetitive Quadrat Stations

MEAN

STANDARD 

ERROR MEAN

STANDARD 

ERROR

Agaricia agaricites 0.00 0.00 0.02 0.01

Colpophyllia natans 0.59 0.27 0.96 0.50

Diploria strigosa 9.02 1.55 10.11 1.49

Madracis spp. 0.40 0.15 0.44 0.14

Millepora alcicornis 0.68 0.16 0.51 0.17

Montastraea cavernosa 3.85 0.88 4.14 0.93

Montastraea annularis spp. complex 40.34 2.91 40.20 2.77

Mussa angulosa 0.02 0.01 0.03 0.03

Porites astreoides 4.06 0.51 5.88 0.64

Siderastrea siderea 0.09 0.08 0.11 0.10

Stephanocoenia intersepta 0.24 0.11 0.13 0.05

Unidentified Scleractinia 2.06 0.33 0.34 0.09

TOTAL CORAL 61.34 2.75 62.87 2.32

Agelas clathrodes 0.17 0.08 0.13 0.09

Pseudoceratina crassa 0.07 0.03 0.00 0.00

Unidentified Sponges 0.19 0.06 0.05 0.03

Xestospongia muta 0.06 0.04 0.08 0.08

TOTAL SPONGE 0.49 0.16 0.26 0.13

Dictyota spp. 0.42 0.19 4.18 0.75

General Macroalgae 1.40 0.35 0.00 0.00

Lobophora variegata 0.00 0.00 14.12 1.25

Turf Algae 11.22 0.82 0.15 0.07

TOTAL MACROALGAE 13.04 1.02 18.45 1.36

Other 0.12 0.05 0.45 0.09

Serpulidae 0.19 0.05 0.16 0.04

TOTAL OTHER LIVE 0.31 0.06 0.61 0.10

Coralline Algae 3.29 0.57 10.50 0.94

CTB 20.53 1.65 6.42 0.88

TOTAL CTB 23.81 1.73 16.91 1.24

TOTAL SAND AND RUBBLE 0.79 0.44 0.74 0.36

TOTAL UNKNOWN 0.21 0.06 0.15 0.05

TOTAL COVER 100.00 100.00

SPONGE

CORAL

EAST BANK 

NOVEMBER 2005

EAST BANK JUNE 

2006

MACROALGAE

UNKNOWNS

SAND AND RUBBLE

CRUSTOSE CORALLINE, TURF, BARE

OTHER LIVE

127
Appendix 3: Repetitive Quadrat Percent Cover Data

Deep Repetitive Quadrat Stations

MEAN

STANDARD 

ERROR MEAN

STANDARD 

ERROR

Colpophyllia natans 8.86 3.16 3.57 1.95

Diploria strigosa 2.91 1.15 3.17 2.35

Madracis spp. 0.24 0.17 0.00 0.00

Millepora alcicornis 0.68 0.46 0.68 0.31

Montastraea cavernosa 15.84 3.02 17.74 3.11

Montastraea annularis spp. complex 42.02 4.81 41.21 4.40

Mussa angulosa 2.71 1.38 3.63 1.78

Porites astreoides 0.00 0.00 0.28 0.29

Scolymia cubensis 0.00 0.00 0.11 0.13

Stephanocoenia intersepta 0.00 0.00 0.85 0.48

Unidentified Scleractinia 1.24 1.17 0.74 0.39

TOTAL CORAL 74.50 4.44 72.00 4.60

Agelas clathrodes 0.00 0.00 0.74 0.36

Unidentified Sponges 0.28 0.29 0.28 0.16

TOTAL SPONGE 0.28 0.29 1.02 0.46

Dictyota spp. 0.00 0.00 3.68 0.86

General Macroalgae 0.36 0.23 0.00 0.00

Lobophora variegata 0.00 0.00 3.63 1.11

Turf Algae 8.98 4.18 0.45 0.46

TOTAL MACROALGAE 9.34 4.38 7.77 2.07

Other 0.04 0.05 0.17 0.09

TOTAL OTHER LIVE 0.04 0.05 0.17 0.09

Coralline Algae 4.51 2.23 13.83 2.53

CTB 7.74 2.50 1.93 0.97

TOTAL CTB 12.25 2.91 15.76 2.75

TOTAL SAND AND RUBBLE 2.11 2.22 3.00 1.58

TOTAL UNKNOWN 1.48 1.46 0.28 0.20

TOTAL COVER 100.00 100.00

SAND AND RUBBLE

UNKNOWNS

SPONGE

MACROALGAE

OTHER LIVE

CRUSTOSE CORALLINE, TURF, BARE

EAST BANK 

NOVEMBER 2005

EAST BANK JUNE 

2006

CORAL

128
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrat Stations

June 2005 - November 2005

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral head 

in June 

2005 (cm2)

Area of 

coral 

head in 

Nov 2005 

(cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Montastraea sp. 17.73 17.28 -0.44 -0.05 -2.51 

2 Diploria strigosa 4.83 4.94 0.11 0.01 2.24 

3 Montastraea sp. 6.71 4.89 -1.82 -0.19 -27.17 

4 Montastraea sp. 9.67 7.39 -2.28 -0.24 -23.62 

5 Montastraea sp. 6.47 5.69 -0.78 -0.08 -12.08 

1 Montastraea sp. 35.58 37.65 2.08 0.22 5.84 

2 Montastraea sp. 19.30 21.08 1.79 0.19 9.25 

3 Montastraea sp. 10.62 11.60 0.98 0.10 9.22 

4 Montastraea sp. 12.99 9.35 -3.64 -0.38 -28.02 

1 Diploria strigosa 86.86 67.62 -19.24 -2.01 -22.15 

2 Montastraea sp. 27.46 17.75 -9.71 -1.02 -35.37 

3 Diploria strigosa 11.36 11.83 0.47 0.05 4.18 

4 Montastraea sp. 44.88 37.66 -7.22 -0.76 -16.09 

1 Diploria strigosa 14.22 16.28 2.05 0.21 14.43 

2 Montastraea sp. 46.66 59.23 12.57 1.32 26.94 

3 Diploria strigosa 7.74 10.49 2.75 0.29 35.60 

4 Diploria strigosa 32.43 0.00 -32.43 -3.40 -100.00 

5 Montastraea sp. 33.67 29.47 -4.21 -0.44 -12.49 

1 Montastraea sp. 29.65 30.97 1.32 0.14 4.46 2 Montastraea sp. 18.85 20.21 1.36 0.14 7.20 3 Montastraea sp. 53.09 57.01 3.92 0.41 7.38 

4 Diploria strigosa 11.62 13.52 1.91 0.20 16.40 

5 Montastraea sp. 236.46 253.49 17.04 1.78 7.21 

2 1.20 0.133 -19.26 -2.02b

2.67

14

-5.22 -0.5-35.70 -3.725.54

129
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrat Stations

June 2005 - November 2005

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral head 

in June 

2005 (cm2)

Area of 

coral 

head in 

Nov 2005 

(cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Montastraea sp. 15.88 15.02 -0.86 -0.09 -5.44 

2 Montastraea sp. 24.71 9.27 -15.44 -1.62 -62.48 

3 Diploria strigosa 16.00 0.00 -16.00 -1.68 -100.00 

4 Montastraea sp. 17.86 8.18 -9.67 -1.01 -54.18 

5 Montastraea sp. 12.20 8.08 -4.12 -0.43 -33.78 

1 Montastraea sp. 19.55 24.35 4.81 0.50 24.58 

2 Montastraea sp. 29.20 36.77 7.57 0.79 25.94 

3 Montastraea sp. 11.95 12.90 0.95 0.10 7.93 

4 Montastraea sp. 47.37 46.25 -1.11 -0.12 -2.35 

5 Montastraea sp. 5.95 6.19 0.24 0.03 4.06 

1 Montastraea sp. 27.32 48.58 21.26 2.23 77.83 

2 Montastraea sp. 7.03 0.00 -7.03 -0.74 -100.00 3 Porites astreoides 0.95 0.00 -0.95 -0.10 -100.00 

4 Diploria strigosa 3.16 0.00 -3.16 -0.33 -100.00 5 Diploria strigosa 4.32 4.63 0.31 0.03 7.17 

1 Montastraea sp. 21.76 49.25 27.49 0.29 126.32 

2 Montastraea sp. 27.54 30.70 3.15 0.03 11.45 3 Montastraea sp. 67.66 76.42 8.77 0.09 12.96 4 Montastraea sp. 34.14 41.86 7.73 0.08 22.63 

5 Montastraea sp. 137.73 124.80 -12.93 -0.14 -9.39 

1 Diploria strigosa 17.67 13.12 -4.55 -0.48 -25.76 

2 Diploria strigosa 11.21 16.93 5.73 0.60 51.13 

3 Montastraea sp. 110.87 79.94 -30.93 -3.24 -27.90 4 Diploria strigosa 19.52 17.19 -2.33 -0.24 -11.95 

10 -3.3

12.45 1.30-32.08

4b -46.1059

-4.8

7 10.44 1.09

34.21 0.36

130
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrat Stations

June 2005 - November 2005

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral head 

in June 

2005 (cm2)

Area of 

coral 

head in 

Nov 2005 

(cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Diploria strigosa 11.55 7.86 -3.69 -0.39 -31.98 

2 Montastraea sp. 107.26 99.41 -7.85 -0.82 -7.32 

3 Montastraea sp. 52.30 50.28 -2.02 -0.21 -3.86 

4 Montastraea sp. 45.93 32.78 -13.16 -1.38 -28.64 

1 Porites astreoides 4.36 4.53 0.17 0.02 4.00 

2 Porites astreoides 3.57 3.91 0.34 0.04 9.59 

3 Diploria strigosa 5.27 6.07 0.80 0.08 15.14 

4 Diploria strigosa 3.41 4.12 0.71 0.07 20.98 

5 Diploria strigosa 15.87 13.88 -1.99 -0.21 -12.55 

1 Diploria strigosa 46.10 47.49 1.39 0.15 3.02 

2 Montastraea sp. 2.75 3.45 0.70 0.07 25.40 

3 Diploria strigosa 84.50 82.50 -2.00 -0.21 -2.37 

4 Montastraea cavernosa 28.74 1.16 -27.58 -2.89 -95.97 

1 Montastraea sp. 13.40 20.64 7.24 0.76 54.06 2 Montastraea sp. 24.30 18.72 -5.57 -0.58 -22.94 3 Diploria strigosa 2.37 2.30 -0.07 -0.01 -2.75 

4 Montastraea sp. 22.62 25.11 2.49 0.26 10.99 5 Montastraea cavernosa 17.73 0.59 -17.14 -1.79 -96.68 

1 Diploria strigosa 8.58 9.74 1.16 0.12 13.57 

2 Montastraea sp. 1.81 2.83 -5.75 -0.60 -316.77 

3 Montastraea sp. 14.02 19.59 17.77 1.86 126.82 

4 Diploria strigosa 8.63 6.72 -7.30 -0.76 -84.57 

5 Diploria strigosa 7.23 0.00 -7.23 -0.76 -100.00 

1211 -2.8

-1.3-0.10.00-2.8

-13.05

14b -1.34

0.04

1412b -27.49

-26.72

131
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrat Stations

June 2005 - November 2005

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral head 

in June 

2005 (cm2)

Area of 

coral 

head in 

Nov 2005 

(cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Montastraea sp. 22.75 32.33 9.58 1.00 42.08 

2 Montastraea sp. 12.35 11.08 -1.27 -0.13 -10.31 

3 Montastraea sp. 19.61 18.00 -1.61 -0.17 -8.20 

4 Montastraea sp. 5.83 5.86 0.03 0.00 0.52 

1 Montastraea sp. 32.86 33.67 0.81 0.08 2.45 

2 Diploria strigosa 20.86 20.61 -0.25 -0.03 -1.18 3 Diploria strigosa 30.35 34.95 4.60 0.48 15.17 

4 Diploria strigosa 66.91 77.25 10.34 1.08 15.46 

5 Montastraea sp. 13.85 13.19 -0.67 -0.07 -4.80 

1 Montastraea sp. 69.60 59.14 -10.46 -1.10 -15.03 

2 Montastraea sp. 185.00 192.76 7.76 0.81 4.19 

3 Diploria strigosa 14.85 14.83 -0.02 0.00 -0.13 

4 Diploria strigosa 20.88 18.18 -2.71 -0.28 -12.96 

5 Montastraea sp. 67.54 64.66 -2.88 -0.30 -4.26 

1 Montastraea sp. 4.51 5.01 0.49 0.05 10.92 

2 Diploria strigosa 1.07 1.28 0.21 0.02 19.42 

3 Montastraea sp. 5.13 4.65 -0.47 -0.05 -9.22 

4 Diploria strigosa 5.46 5.41 -0.05 0.00 -0.84 

1 Montastraea sp. 89.40 85.19 -4.21 -0.44 -4.71 

2 Montastraea cavernosa 43.21 0.07 -43.14 -4.52 -99.84 

3 Diploria strigosa 22.50 23.91 1.41 0.15 6.29 

4 Montastraea cavernosa 25.52 0.05 -25.47 -2.67 -99.81 

0.18 0.02

20 -71.41 -7.415 6.72 0.7018b

-8.31 -0.8

17 14.84 1.5518

132
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrat Stations

June 2005 - November 2005

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral head 

in June 

2005 (cm2)

Area of 

coral 

head in 

Nov 2005 

(cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

c

Q

1 Montastraea sp. 2.47 2.67 0.20 0.02 8.09 2 Diploria strigosa 3.33 3.36 0.03 0.00 0.93 3 Diploria strigosa 4.29 3.80 -0.50 -0.05 -11.55 4 Montastraea sp. 9.39 9.59 0.20 0.02 2.17 5 Montastraea sp. 12.90 0.00 -12.90 -1.35 -100.03 

1 Montastraea sp. 22.38 17.82 -4.57 -0.48 -20.40 

2 Montastraea sp. 15.50 6.09 -9.41 -0.99 -60.71 3 Montastraea sp. 22.26 23.81 1.55 0.16 6.98 4 Montastraea cavernosa 52.87 59.47 -52.51 -5.50 -99.31 5 Montastraea sp. 53.20 68.43 15.23 1.59 28.63 1 Montastraea sp. 83.30 44.85 -38.45 -4.03 -46.16 2 Montastraea cavernosa 13.55 12.17 -1.39 -0.15 -10.25 3 Diploria strigosa 46.30 54.58 8.29 0.87 17.90 4 Diploria strigosa 46.36 49.96 3.61 0.38 7.78 5 Diploria strigosa 21.78 18.12 -3.66 -0.38 -16.81 

1 Montastraea sp. 10.62 20.51 9.89 1.04 93.20 2 Diploria strigosa 22.98 21.50 -1.47 -0.15 -6.42 3 Colpophyllia natans 11.47 0.06 -11.41 -1.20 -99.47 4 Diploria strigosa 34.89 31.59 -3.30 -0.35 -9.45 5 Montastraea sp. 53.94 33.97 -19.97 -2.09 -37.02 

1 Montastraea sp. 28.48 22.35 -6.13 -0.64 -21.53 2 Montastraea sp. 64.89 48.27 -16.62 -1.74 -25.61 3 Montastraea sp. 21.68 18.61 -3.08 -0.32 -14.18 4 Diploria strigosa 7.28 8.67 1.39 0.15 19.10 5 Diploria strigosa 20.11 17.44 -2.68 -0.28 -13.31 

20b -12.96 -1

-26.26 -2-27.11 -2-49.71 -5-31.61 -3

212221b23

133
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrat Stations

June 2005 - November 2005

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral head 

in June 

2005 (cm2)

Area of 

coral 

head in 

Nov 2005 

(cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Montastraea sp. 23.02 16.58 -6.44 -0.67 -27.97 2 Montastraea cavernosa 20.62 0.01 -20.60 -2.16 -99.93 3 Montastraea sp. 48.33 43.81 -4.52 -0.47 -9.34 4 Diploria strigosa 9.37 11.01 1.64 0.17 17.53 5 Montastraea sp. 62.26 84.79 22.53 2.36 

1 Montastraea sp. 1.49 1.60 0.11 0.01 7.14 

2 Montastraea sp. 7.82 9.66 1.84 0.19 23.49 3 Montastraea sp. 1.91 2.40 0.49 0.05 25.64 4 Montastraea sp. 3.69 4.05 0.36 0.04 9.78 

1 Diploria strigosa 8.83 10.48 1.65 0.17 18.71 2 Diploria strigosa 19.70 21.73 2.03 0.21 10.31 3 Diploria strigosa 7.94 7.35 -0.59 -0.06 -7.49 4 Diploria strigosa 20.70 0.00 -20.70 -2.17 -100.00 

5 Diploria strigosa 15.04 5.37 -9.67 -1.01 -64.29 

1 Montastraea sp. 12.52 11.14 -1.38 -0.14 -11.03 2 Montastraea sp. 8.26 8.51 0.25 0.03 3.03 3 Diploria strigosa 4.42 3.97 -0.45 -0.05 -10.23 4 Diploria strigosa 20.25 18.78 -1.47 -0.15 -7.28 5 Diploria strigosa 10.84 10.41 -0.43 -0.04 -3.93 1 Montastraea sp. 4.60 5.11 -3.72 -0.39 -80.85 

2 Montastraea sp. 16.68 22.18 2.48 0.26 14.90 3 Montastraea sp. 11.01 12.17 4.23 0.44 38.38 4 Diploria strigosa 15.16 18.64 -2.06 -0.22 -13.60 5 Montastraea sp. 241.20 184.83 169.79 17.78 70.39 

-3.48 -0.3-45.73 -4.7

24

-27.29 -2.8

25 2.79 0.29

-7.39 -0.7

2626b27

134
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrat Stations

June 2005 - November 2005

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral head 

in June 

2005 (cm2)

Area of 

coral 

head in 

Nov 2005 

(cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Diploria strigosa 21.76 27.56 5.80 0.61 26.67 

2 Diploria strigosa 26.91 30.70 3.79 0.40 14.10 

3 Diploria strigosa 97.97 92.35 -5.62 -0.59 -5.73 4 Diploria strigosa 44.37 0.00 -44.37 -4.65 -100.00 

5 Diploria strigosa 16.58 0.00 -16.58 -1.74 -100.00 

1 Diploria strigosa 11.23 10.66 -0.57 -0.06 -5.09 2 Montastraea sp. 10.26 10.93 0.67 0.07 6.57 3 Montastraea cavernosa 71.95 0.61 -71.34 -7.47 -99.16 

4 Diploria strigosa 13.67 0.00 -13.67 -1.43 -100.00 

1 Montastraea cavernosa 53.02 1.26 -51.76 -5.42 -97.62 

2 Colpophyllia natans 56.14 56.05 -0.09 -0.01 -0.16 

3 Montastraea cavernosa 82.55 0.48 -82.06 -8.59 -99.42 

4 Montastraea cavernosa 32.05 3.02 -29.03 -3.04 -90.57 

5 Diploria strigosa 49.65 27.35 -22.31 -2.34 -44.92 

1 Montastraea sp. 16.46 8.48 -7.98 -0.84 -48.46 

2 Diploria strigosa 71.70 0.00 -71.70 -7.51 -100.00 

3 Diploria strigosa 4.92 0.00 -4.92 -0.52 -100.00 

4 Diploria strigosa 7.62 7.58 -0.03 0.00 -0.42 

5 Diploria strigosa 9.59 10.63 1.04 0.11 10.84 

1 Montastraea sp. 23.67 26.94 3.27 0.34 13.83 2 Montastraea sp. 1.96 2.22 0.26 0.03 13.11 

3 Montastraea sp. 5.46 6.60 1.14 0.12 20.91 

4 Montastraea sp. 6.98 7.68 0.71 0.07 10.15 

5 Montastraea sp. 25.58 23.76 -1.83 -0.19 -7.14 

-83.58 -8.7

31 3.55 0.37

-56.97 -5.9-185.24 -19.-84.92 -8.8

293030b29b

135
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrat Stations

June 2005 - November 2005

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)**unknown repetitive quadrat numberNote: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral head 

in June 

2005 (cm2)

Area of 

coral 

head in 

Nov 2005 

(cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

Tot

chan

A

wit

Qua

1 Montastraea sp. 10.58 10.38 -0.20 -0.02 -1.88 2 Montastraea sp. 15.47 11.89 -3.59 -0.38 -23.18 

3 Montastraea sp. 55.16 55.49 0.33 0.03 0.60 4 Montastraea cavernosa 21.42 0.40 -21.03 -2.20 -98.14 

1 Montastraea sp. 121.54 48.42 -73.12 -7.66 -60.16 

2 Montastraea sp. 78.55 0.35 -78.20 -8.19 -99.55 

3 Montastraea sp. 47.03 2.31 -44.73 -4.68 -95.10 

4 Montastraea sp. 15.12 0.19 -14.93 -1.56 -98.77 

1 Diploria strigosa 74.52 62.14 -12.38 -1.30 -16.61 

2 Diploria strigosa 15.18 15.00 -0.18 -0.02 -1.18 3 Colpophyllia natans 78.89 91.83 12.94 1.36 16.40 4 Diploria strigosa 10.33 11.26 0.93 0.10 9.04 5 Diploria strigosa 26.31 30.66 4.35 0.46 16.53 6 Diploria strigosa 41.88 43.17 1.30 0.14 3.09 

1 Montastraea sp. 7.95 8.34 0.40 0.04 4.97 

2 Diploria strigosa 19.15 22.04 2.89 0.30 15.07 3 Montastraea sp. 26.47 26.14 -0.33 -0.03 -1.25 4 Diploria strigosa 21.16 0.00 -21.16 -2.22 -100.00 

5 Montastraea sp. 14.68 12.91 -1.76 -0.18 -12.01 

1 Colpophyllia natans 11.94 11.71 -0.23 -0.02 -1.91 

2 Diploria strigosa 15.52 26.76 11.23 1.18 72.38 3 Montastraea sp. 25.25 16.12 -9.13 -0.96 -36.17 

4 Diploria strigosa 20.26 22.01 1.75 0.18 8.63 5 Montastraea sp. 26.49 27.47 0.99 0.10 3.72 

UNK2**

-24.48 -2.56

36 -210.98

4.61 0.48

-22.0

UNK1** -19.97 -2.093833

6.96 0.73

136
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrats

November 2005 - June 2006

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)** Measurements from June 2005 (rather than November 2005)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral 

head in 

Nov 2005 

(cm2)

Area of 

coral 

head in 

June 2006 

(cm2) 

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Montastraea sp. 17.28 20.39 3.11 0.33 18.00 

2 Diploria strigosa 4.94 5.68 0.74 0.08 14.90 

3 Montastraea sp. 4.89 7.05 2.16 0.23 44.28 

4 Montastraea sp. 7.39 8.36 0.97 0.10 13.13 

5 Montastraea sp. 5.69 7.13 1.45 0.15 25.42 

1 Montastraea sp. 37.65 43.71 6.05 0.63 16.08 

2 Montastraea sp. 21.08 19.19 -1.90 -0.20 -8.99 3 Montastraea sp. 11.60 11.98 0.38 0.04 3.30 4 Montastraea sp. 9.35 12.26 2.91 0.30 31.11 

1 Diploria strigosa 67.62 85.13 17.51 1.83 25.90 2 Montastraea sp. 17.75 34.76 17.02 1.78 95.87 

3 Diploria strigosa 11.83 11.28 -0.55 -0.06 -4.64 

4 Montastraea sp. 37.66 42.12 4.47 0.47 11.86 

1 Diploria strigosa 16.28 15.02 -1.26 -0.13 -7.73 

2 Montastraea sp. 59.23 50.24 -8.99 -0.94 -15.19 

3 Diploria strigosa 10.49 10.64 0.15 0.02 1.43 

4 Diploria strigosa 0.00 0.00 0.00 0.00 0.00 

5 Montastraea sp. 29.47 33.07 3.60 0.38 12.21 

1 Montastraea sp. 30.97 34.47 3.49 0.37 11.28 2 Montastraea sp. 20.21 21.50 1.30 0.14 6.42 

3 Montastraea sp. 57.01 60.13 3.12 0.33 5.48 4 Diploria strigosa 13.52 12.45 -1.07 -0.11 -7.91 

5 Montastraea sp. 253.49 248.29 -5.21 -0.55 -2.05 

43 -6.50 -0.682b2** 7.45

1.64 0.17

1 8.43 0.88

38.45 4.03

0.78

137
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrats

November 2005 - June 2006

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral 

head in 

Nov 2005 

(cm2)

Area of 

coral 

head in 

June 2006 

(cm2) 

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Montastraea sp. 24.35 26.20 1.84 0.19 7.57 2 Montastraea sp. 36.77 42.38 5.61 0.59 15.26 3 Montastraea sp. 12.90 12.53 -0.37 -0.04 -2.84 4 Montastraea sp. 46.25 44.78 -1.47 -0.15 -3.19 

5 Montastraea sp. 6.19 5.67 -0.52 -0.05 -8.44 

1 Montastraea sp. 48.58 44.76 -3.82 -0.40 -7.86 

2 Montastraea sp. 0.00 0.00 0.00 0.00 0.00 3 Porites astreoides 0.00 0.00 0.00 0.00 0.00 4 Diploria strigosa 0.00 0.00 0.00 0.00 0.00 5 Diploria strigosa 4.63 4.17 -0.46 -0.05 -9.85 

1 Diploria strigosa 13.12 12.46 -0.66 -0.07 -5.04 2 Diploria strigosa 16.93 16.75 -0.18 -0.02 -1.07 3 Montastraea sp. 79.94 119.15 39.21 4.10 49.04 4 Diploria strigosa 17.19 16.69 -0.51 -0.05 -2.94 

1 Diploria strigosa 7.86 10.71 2.85 0.30 36.28 2 Montastraea sp. 99.41 102.93 3.52 0.37 3.54 3 Montastraea sp. 50.28 54.50 4.22 0.44 8.39 4 Montastraea sp. 32.78 44.98 12.21 1.28 37.25 

1 Porites astreoides 4.53 5.08 0.55 0.06 12.04 2 Porites astreoides 3.91 4.38 0.47 0.05 12.02 3 Diploria strigosa 6.07 6.52 0.45 0.05 7.35 

4 Diploria strigosa 4.12 4.52 0.40 0.04 9.59 5 Diploria strigosa 13.88 14.67 0.79 0.08 5.70 

1051112 0.28

5.09 0.53

7 -4.27 -0.4

2.6537.86 3.9622.80 2.39

138
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrats

November 2005 - June 2006

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)** Measurements from June 2005 (rather than November 2005)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral 

head in 

Nov 2005 

(cm2)

Area of 

coral 

head in 

June 2006 

(cm2) 

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Diploria strigosa 47.49 50.65 3.16 0.33 6.66 

2 Montastraea sp. 3.45 2.79 -0.66 -0.07 -19.01 3 Diploria strigosa 82.50 83.03 0.53 0.06 0.64 4 Montastraea cavernosa 21.78 21.05 -0.74 -0.08 -3.39 

1 Diploria strigosa 9.74 10.40 0.66 0.07 6.77 2 Montastraea sp. 2.83 2.05 -0.78 -0.08 -27.40 

3 Montastraea sp. 19.59 24.57 4.98 0.52 25.42 

4 Diploria strigosa 6.72 8.51 1.79 0.19 26.66 

5 Diploria strigosa 0.00 0.00 0.00 0.00 0.00 

1 Montastraea sp. 32.33 34.15 1.82 0.19 5.64 

2 Montastraea sp. 11.08 14.90 3.82 0.40 34.46 3 Montastraea sp. 18.00 19.55 1.55 0.16 8.61 

4 Montastraea sp. 5.86 6.81 0.95 0.10 16.21 

1 Montastraea sp. 33.67 32.59 -1.07 -0.11 -3.18 

2 Diploria strigosa 20.61 18.25 -2.37 -0.25 -11.48 

3 Diploria strigosa 34.95 29.19 -5.76 -0.60 -16.49 

4 Diploria strigosa 77.25 69.69 -7.56 -0.79 -9.78 

5 Montastraea sp. 13.19 4.54 -8.65 -0.91 -65.59 

1 Montastraea sp. 12.42 13.74 1.32 0.14 10.58 2 Diploria strigosa 6.24 7.29 1.05 0.11 16.90 

3 Montastraea sp. 41.55 30.34 -11.21 -1.17 -26.98 4 Diploria strigosa 48.76 48.81 0.06 0.01 0.12 

5 Diploria strigosa 94.53 107.67 13.13 1.38 13.89 

15 8.14 0.8517 -25.41 -2.661417b**

6.65 0.704.35 0.46

12b 2.30 0.24

139
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrats

November 2005 - June 2006

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral 

head in 

Nov 2005 

(cm2)

Area of 

coral 

head in 

June 2006 

(cm2) 

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Montastraea sp. 59.14 68.22 9.09 0.95 15.37 

2 Montastraea sp. 192.76 198.23 5.47 0.57 2.84 

3 Diploria strigosa 14.83 16.26 1.43 0.15 9.63 

4 Diploria strigosa 18.18 18.36 0.19 0.02 1.02 

5 Montastraea sp. 64.66 73.54 8.88 0.93 13.73 

1 Montastraea sp. 5.01 4.46 -0.54 -0.06 -10.86 

2 Diploria strigosa 1.28 1.20 -0.09 -0.01 -6.78 

3 Montastraea sp. 4.65 8.52 3.87 0.41 83.12 

4 Diploria strigosa 5.41 5.46 0.04 0.00 0.82 

1 Montastraea sp. 85.19 90.82 5.63 0.59 6.61 

2 Montastraea cavernosa 33.48 42.96 9.48 0.99 28.32 

3 Diploria strigosa 23.91 24.73 0.82 0.09 3.44 

4 Montastraea cavernosa 28.18 30.03 1.85 0.19 6.56 

1 Diploria strigosa 2.67 2.59 -0.08 -0.01 -3.09 

2 Diploria strigosa 3.36 3.89 0.53 0.06 15.78 

3 Diploria strigosa 3.80 4.23 0.43 0.05 11.41 

4 Montastraea sp. 9.59 9.55 -0.04 0.00 -0.41 

5 Montastraea sp. 0.00 0.00 0.00 0.00 0.00 

1 Montastraea sp. 17.82 24.32 6.51 0.68 36.51 

2 Montastraea sp. 6.09 4.15 -1.95 -0.20 -31.95 

3 Montastraea sp. 23.81 25.14 1.32 0.14 5.56 

4 Montastraea cavernosa 59.47 58.16 -1.31 -0.14 -2.21 5 Montastraea sp. 68.43 60.07 -8.36 -0.88 -12.22 

20b 0.84 0.0920 17.78 1.8618b 3.28 0.34

2.62

-3.7925.05

2118

-0.4

140
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrats

November 2005 - June 2006

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral 

head in 

Nov 2005 

(cm2)

Area of 

coral 

head in 

June 2006 

(cm2) 

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Montastraea sp. 44.85 45.85 1.00 0.11 2.24 

2 Montastraea cavernosa 12.17 14.05 1.89 0.20 15.50 3 Diploria strigosa 54.58 46.21 -8.37 -0.88 -15.33 4 Diploria strigosa 49.96 50.47 0.51 0.05 1.02 

5 Diploria strigosa 18.12 21.51 3.39 0.36 18.71 

1 Montastraea sp. 20.51 18.16 -2.35 -0.25 -11.45 

2 Diploria strigosa 21.50 21.09 -0.42 -0.04 -1.93 3 Colpophyllia natans 10.86 12.91 2.05 0.21 18.91 4 Diploria strigosa 31.59 32.22 0.63 0.07 1.99 5 Montastraea sp. 33.97 46.81 12.83 1.34 37.78 

1 Montastraea sp. 22.35 27.45 5.11 0.53 22.85 2 Montastraea sp. 48.27 63.24 14.97 1.57 31.01 3 Montastraea sp. 18.61 27.71 9.10 0.95 48.90 4 Diploria strigosa 8.67 8.00 -0.67 -0.07 -7.74 5 Diploria strigosa 17.44 28.29 10.86 1.14 62.26 

1 Montastraea sp. 16.58 24.69 8.10 0.85 48.87 

2 Montastraea cavernosa 16.02 18.71 2.70 0.28 16.83 

3 Montastraea sp. 43.81 45.04 1.23 0.13 2.82 

4 Diploria strigosa 11.01 11.12 0.12 0.01 1.05 

5 Montastraea sp. 84.79 79.03 -5.76 -0.60 -6.79 

1 Montastraea sp. 1.60 1.66 0.06 0.01 3.64 

2 Montastraea sp. 9.66 10.22 0.56 0.06 5.79 3 Montastraea sp. 2.40 0.00 -2.40 -0.25 -100.00 

4 Montastraea sp. 4.05 3.84 -0.21 -0.02 -5.14 

24

12.75 1.34

2523 39.36 4.12

6.39-1.99 -0.2

0.67

2221b -1.58 -0.17

141
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrats

November 2005 - June 2006

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral 

head in 

Nov 2005 

(cm2)

Area of 

coral 

head in 

June 2006 

(cm2) 

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Diploria strigosa 10.48 11.73 1.25 0.13 11.92 2 Diploria strigosa 21.73 24.51 2.78 0.29 12.78 3 Diploria strigosa 7.35 8.11 0.77 0.08 10.43 4 Diploria strigosa 0.00 0.00 0.00 0.00 0.00 5 Diploria strigosa 5.37 6.18 0.81 0.08 15.07 1 Montastraea sp. 11.14 4.26 -6.88 -0.72 -61.74 2 Montastraea sp. 8.51 8.51 0.00 0.00 0.03 3 Diploria strigosa 3.97 4.73 0.76 0.08 19.29 4 Diploria strigosa 18.78 20.98 2.20 0.23 11.74 5 Diploria strigosa 10.41 11.76 1.35 0.14 12.93 

1 Diploria strigosa 27.56 13.24 -14.32 -1.50 -51.95 2 Diploria strigosa 30.70 31.90 1.20 0.13 3.90 3 Diploria strigosa 92.35 113.21 20.86 2.18 22.58 4 Diploria strigosa 0.00 0.00 0.00 0.00 0.00 5 Diploria strigosa 0.00 0.00 0.00 0.00 0.00 1 Diploria strigosa 10.66 10.33 -0.34 -0.04 -3.14 2 Montastraea sp. 10.93 10.49 -0.44 -0.05 -4.07 3 Montastraea cavernosa 66.97 76.30 9.34 0.98 13.95 4 Diploria strigosa 0.00 0.00 0.00 0.00 0.00 1 Montastraea cavernosa 35.61 12.57 -23.04 -2.41 -64.69 2 Colpophyllia natans 56.05 71.58 15.53 1.63 27.71 3 Montastraea cavernosa 55.00 49.09 -5.91 -0.62 -10.75 4 Montastraea cavernosa 36.50 46.98 10.48 1.10 28.72 5 Diploria strigosa 27.35 12.27 -15.08 -1.58 -55.13 

-2.56 -0.25.60 0.59

0.90

7.74 0.81-18.01 -1.88.56

29302626b29b

142
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrats

November 2005 - June 2006

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral 

head in 

Nov 2005 

(cm2)

Area of 

coral 

head in 

June 2006 

(cm2) 

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Montastraea sp. 15.59 14.98 -0.61 -0.06 -3.90 2 Diploria strigosa 0.00 0.00 0.00 0.00 0.00 

3 Diploria strigosa 0.00 0.00 0.00 0.00 0.00 

4 Diploria strigosa 7.58 8.84 1.25 0.13 16.52 

5 Diploria strigosa 10.63 11.53 0.90 0.09 8.44 

1 Montastraea sp. 26.94 23.21 -3.72 -0.39 -13.83 

2 Montastraea sp. 2.22 2.18 -0.04 0.00 -1.94 3 Montastraea sp. 6.60 6.86 0.26 0.03 3.98 

4 Montastraea sp. 7.68 8.38 0.70 0.07 9.11 

5 Montastraea sp. 23.76 28.69 4.93 0.52 20.76 

1 Montastraea sp. 10.38 14.03 3.65 0.38 35.15 

2 Montastraea sp. 11.89 13.55 1.66 0.17 13.98 

3 Montastraea sp. 55.49 49.56 -5.93 -0.62 -10.68 

4 Montastraea cavernosa 35.41 34.54 -0.86 -0.09 -2.44 1 Montastraea sp. 48.42 94.34 45.92 4.81 94.85 

2 Montastraea sp. 0.35 0.85 0.50 0.05 141.24 3 Montastraea sp. 2.31 0.00 -2.31 -0.24 -100.00 

4 Montastraea sp. 0.19 0.00 -0.19 -0.02 -100.00 

1 Diploria strigosa 62.14 72.52 10.38 1.09 16.70 

2 Diploria strigosa 15.00 16.67 1.67 0.18 11.16 

3 Colpophyllia natans 91.83 113.04 21.21 2.22 23.09 

4 Diploria strigosa 11.26 12.78 1.52 0.16 13.51 

5 Diploria strigosa 30.66 35.59 4.93 0.52 16.07 

6 Diploria strigosa 43.17 50.65 7.48 0.78 17.32 

-1.48 -0.1

36 43.93

1.54 0.16

4.60

38 47.18 4.9430b3331 2.13 0.22

143
Appendix 4: Repetitive Quadrat Planimetry Data

Study Site Repetitive Quadrats

November 2005 - June 2006

* The RQS # is an identifier used during planimetry analysis (not necessarily corresponding to the station number)*** unknown repetitive quadrat numberNote: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

RQS #* Coral Head # Species

Area of 

coral 

head in 

Nov 2005 

(cm2)

Area of 

coral 

head in 

June 2006 

(cm2) 

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

To

cha

A

w

Qu

1 Montastraea sp. 8.34 8.97 0.63 0.07 7.51 2 Diploria strigosa 22.04 20.57 -1.47 -0.15 -6.66 3 Montastraea sp. 26.14 22.47 -3.67 -0.38 -14.05 4 Diploria strigosa 0.00 0.00 0.00 0.00 0.00 5 Montastraea sp. 12.91 14.11 1.20 0.13 9.29 

1 Colpophyllia natans 11.71 11.17 -0.54 -0.06 -4.58 

2 Diploria strigosa 26.76 17.47 -9.29 -0.97 -34.71 

3 Montastraea sp. 16.12 23.21 7.09 0.74 43.97 

4 Diploria strigosa 22.01 21.92 -0.09 -0.01 -0.41 

5 Montastraea sp. 27.47 31.00 3.52 0.37 12.83 

UNK2*** 0.70 0.07UNK1*** -3.31 -0.3

144
Appendix 4: Repetitive Quadrat Planimetry Data

Deep Repetitive Quadrat Stations

June 2005 - November 2005

Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

Station # Coral Head # Species

Area of coral 

head in June 

2005 (cm2)

Area of coral 

head in Nov 

2005 (cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

c

1 Colpophyllia natans 89.49 86.65 -2.84 -0.30 -3.17 

2 Colpophyllia natans 4.11 5.06 0.95 0.10 22.99 

3 Diploria strigosa 68.85 55.06 -13.78 -1.44 -20.02 

4 Montastraea sp. 62.28 76.90 14.61 1.53 23.46 

1 Montastraea cavernosa 2.50 3.39 0.90 0.09 35.89 

2 Montastraea cavernosa 16.90 24.07 7.16 0.75 42.38 

3 Colpophyllia natans 15.51 15.24 -0.27 -0.03 -1.75 

4 Montastraea sp. 29.71 29.72 0.01 0.00 0.04 

1 Montastraea sp. 13.36 14.08 0.71 0.07 5.34 2 Montastraea cavernosa 5.33 4.98 -0.35 -0.04 -6.55 

3 Montastraea sp. 25.10 29.15 4.05 0.42 16.13 4 Montastraea cavernosa 23.45 28.50 5.05 0.53 21.53 

5 Montastraea sp. 23.00 26.42 3.41 0.36 14.83 

1 Montastraea cavernosa 39.48 32.72 -6.76 -0.71 -17.12 

2 Montastraea cavernosa 19.47 18.64 -0.82 -0.09 -4.23 

3 Colpophyllia natans 11.22 11.43 0.21 0.02 1.91 4 Montastraea sp. 22.78 25.74 2.96 0.31 13.01 5 Montastraea cavernosa 6.03 7.51 1.48 0.16 24.64 

1 Montastraea sp. 1.97 2.23 0.26 0.03 13.44 

2 Montastraea cavernosa 0.63 1.09 0.46 0.05 72.46 

3 Montastraea cavernosa 39.87 37.30 -2.58 -0.27 -6.46 

4 Montastraea sp. 13.55 16.34 2.78 0.29 20.53 5 Montastraea cavernosa 17.00 16.19 -0.81 -0.08 -4.75 

81 -1.07 -82 7.80 083 12.87 184 -2.92 -85 0.12 0

145
Appendix 4: Repetitive Quadrat Planimetry Data

Deep Repetitive Quadrat Stations

June 2005 - November 2005

Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

Station # Coral Head # Species

Area of coral 

head in June 

2005 (cm2)

Area of coral 

head in Nov 

2005 (cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

1 Colpophyllia natans 159.85 183.68 23.83 2.50 14.91 

2 Montastraea cavernosa 49.02 68.44 19.42 2.03 39.62 

3 Montastraea cavernosa 25.29 27.52 2.22 0.23 8.79 

4 Montastraea cavernosa 43.83 29.35 -14.48 -1.52 -33.03 

1 Montastraea sp. 30.63 26.62 -4.02 -0.42 -13.12 

2 Montastraea sp. 11.30 10.10 -1.20 -0.13 -10.66 

3 Montastraea sp. 34.94 38.58 3.64 0.38 10.41 

4 Montastraea sp. 22.44 22.68 0.25 0.03 1.11 

5 Diploria strigosa 1.74 1.57 -0.17 -0.02 -9.58 

1 Montastraea sp. 15.48 15.72 0.24 0.03 1.56 2 Montastraea cavernosa 31.38 31.39 0.01 0.00 0.04 

3 Montastraea sp. 12.74 15.60 2.86 0.30 22.44 

4 Colpophyllia natans 7.09 5.79 -1.30 -0.14 -18.32 

5 Montastraea sp. 30.43 32.97 2.54 0.27 8.34 

1 Montastraea sp. 11.11 10.62 -0.48 -0.05 -4.36 

2 Montastraea cavernosa 4.26 3.66 -0.60 -0.06 -14.18 

3 Montastraea cavernosa 42.11 41.38 -0.73 -0.08 -1.73 

4 Montastraea sp. 4.87 3.66 -1.21 -0.13 -24.82 

5 Montastraea cavernosa 10.20 13.27 3.08 0.32 30.16 

89 0.05 8788 4.35 86 31.00 

-1.50

146
Appendix 4: Repetitive Quadrat Planimetry Data

Deep Repetitive Quadrat Stations

November 2005 - June 2006

Note: Montastraea sp. refers to colonies belonging to the Montastraea annularis species complex

Station # Coral Head # Species

Area of 

coral head 

in 

November 2005 (cm2)

Area of 

coral head 

in June 

2006 (cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

1 Montastraea cavernosa 3.39 2.85 -0.54 -0.06 -15.93 2 Montastraea cavernosa 24.07 12.42 -11.65 -1.22 -48.40 

3 Colpophyllia natans 15.24 11.88 -3.36 -0.35 -22.06 4 Montastraea sp. 29.72 30.92 1.20 0.13 4.04 

1 Montastraea sp. 14.08 11.04 -3.03 -0.32 -21.54 

2 Montastraea cavernosa 4.98 3.85 -1.13 -0.12 -22.64 3 Montastraea sp. 29.15 22.40 -6.75 -0.71 -23.15 

4 Montastraea cavernosa 28.50 23.09 -5.41 -0.57 -19.00 

5 Montastraea sp. 26.42 24.42 -1.99 -0.21 -7.54 

1 Montastraea cavernosa 32.72 38.51 5.79 0.61 17.69 

2 Montastraea cavernosa 18.64 21.64 3.00 0.31 16.09 

3 Colpophyllia natans 11.43 11.48 0.05 0.01 0.43 

4 Montastraea sp. 25.74 23.56 -2.19 -0.23 -8.49 

5 Montastraea cavernosa 7.51 6.23 -1.28 -0.13 -16.99 1 Montastraea sp. 2.23 1.85 -0.39 -0.04 -17.33 

2 Montastraea cavernosa 1.09 0.83 -0.26 -0.03 -24.17 3 Montastraea cavernosa 37.30 39.51 2.21 0.23 5.93 

4 Montastraea sp. 16.34 12.90 -3.44 -0.36 -21.05 

5 Montastraea cavernosa 16.19 20.81 4.62 0.48 28.54 

1 Colpophyllia natans 183.68 143.07 -40.61 -4.25 -22.11 

2 Montastraea cavernosa 68.44 70.29 1.85 0.19 2.70 

3 Montastraea cavernosa 27.52 24.07 -3.45 -0.36 -12.53 

4 Montastraea cavernosa 29.35 36.93 7.58 0.79 25.81 

-14.35

8482858683

-34.632.745.37-18.31

0---0

147
Appendix 4: Repetitive Quadrat Planimetry Data

Deep Repetitive Quadrat Stations

November 2005 - June 2006

Note: Montastraea sp . refers to colonies belonging to the Montastraea annularis species complex

Station # Coral Head # Species

Area of 

coral head 

in 

November 2005 (cm2)

Area of 

coral head 

in June 

2006 (cm2)

Change in 

Area (cm2)

% Change 

in Area 

% change 

in coral 

colony 

area

Total 

change in 

Area 

within 

Quadrat 

(cm2)

1 Montastraea sp. 26.62 32.42 5.81 0.61 21.82 2 Montastraea sp. 10.10 11.87 1.78 0.19 17.61 

3 Montastraea sp. 38.58 38.40 -0.18 -0.02 -0.47 

4 Montastraea sp. 22.68 24.72 2.03 0.21 8.97 

5 Diploria strigosa 1.57 1.76 0.18 0.02 11.67 

1 Montastraea sp. 10.62 11.69 1.07 0.11 10.03 

2 Montastraea cavernosa 3.66 5.44 1.79 0.19 48.83 

3 Montastraea cavernosa 41.38 42.05 0.67 0.07 1.61 

4 Montastraea sp. 3.66 4.81 1.14 0.12 31.25 

5 Montastraea cavernosa 13.27 12.20 -1.07 -0.11 -8.05 

8987

3.599.62

01

148
The Department of the Interior Mission

As the Nation's principal conservation agency, the Department of the Interior has responsibility 

for most of our nationally owned public lands and natural resources. This includes fostering 

sound use of our land and water resources; protecting our fish, wildlife, and biological diversity; 

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development and environmental protection.