Northeastern Gulf of Mexico Coastal and Marine Ecosystem Program

Community Structure and Trophic Ecology of 

Fishes on the Pinnacles Reef Tract

Biological Sciences Report

USGS BSR 2001-0008

OCS Study MMS 2002-034

U.S. Department of the Interior

U.S. Geological Survey

U.S. Department of the Interior

Minerals Management Service

Gulf of Mexico OCS Region
Cover illustration credits:

High resolution multi-beam swath bathymetry map produced by Dr. James V. 

Gardner, USGS, Menlo Park, CA.

Fish photographs by D. C. Weaver and W. F. Smith-Vaniz. 
i

Northeastern Gulf of Mexico Coastal and Marine Ecosystem Program

Community Structure and Trophic Ecology of 

Fishes on the Pinnacles Reef Tract

Douglas C. Weaver, George D. Dennis, and Kenneth J. Sulak

June 2002

Florida Caribbean Science Center

U.S. Geological Survey 

7920 NW 71st Street

Gainesville, Florida 32653-3071

U.S. Department of the Interior

U.S. Geological Survey

U.S. Department of the Interior

Minerals Management Service

Gulf of Mexico OCS Region

Biological Sciences Report

USGS BSR 2001-0008

OCS Study MMS 2002-034
ii
iii

PROJECT COOPERATION

This study was undertaken to meet information needs identified by the U.S. Geological Survey (USGS), 

Outer Continental Shelf Ecosystem Program in concert with the Minerals Management Service (MMS).

DISCLAIMER

This report was prepared by the Florida Caribbean Science Center of the USGS. This report has been 

technically reviewed by USGS and MMS, and has been approved for publication. Approval does not signify 

that the contents necessarily reflect the views and policies of the USGS, nor does mention of trade names or 

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

review and compliance with the MMS editorial standards.

REPORT AVAILABILITY

Extra copies of this report may be obtained from the MMS Public Information Unit at the following address:

U.S. Department of the Interior

Minerals Management Service 

Public Information Office (MS 5034)

1201 Elmwood Park Boulevard

New Orleans, LA 70123-2394

Telephone: (504) 736-2519 or 

1-800-200-GULF

The following datasets are available as read-only files on the Florida Caribbean Science Center Website at 

http://www.fcsc.usgs.gov/index.html:

Located in: Pinnacles Tech Report/

Pinnacles Master Station Log.xls

Pinnacles Master Specimen Log.xls

USGS Technical Report 2001-0008.doc

USGS Technical Report 2001-0008.pdf

SUGGESTED CITATION

Weaver, D. C., G. D. Dennis, and K. J. Sulak. 2001. Northeastern Gulf of Mexico Coastal and Marine 

Ecosystem Program: Community Structure and Trophic Ecology of Demersal Fishes on the 

Pinnacles Reef Tract; Final Synthesis Report. U.S. Department of the Interior, Geological Survey, 

USGS BSR-2001-0008 and Minerals Management Service Gulf of Mexico OCS Region, New 

Orleans, LA, OCS Study MMS 2002-034, 92 pp. + apps.
iv
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PRINCIPAL INVESTIGATORS AND COLLABORATORS

Dr. Kenneth J. Sulak (PrincipaI Investigator (PI), Project Lead Scientist) 1

ken_sulak@usgs.gov

Douglas C. Weaver (Co-PI; Project Research Biologist) 1,2

doug_weaver@usgs.gov

Dr. George D. Dennis (Co-PI; Project Research Biologist) 1

george_dennis@usgs.gov

Collaborators:

Dr. Steve W. Ross 3

Dr. William F. Smith-Vaniz 1

1

Florida Caribbean Science Center, U.S. Geological Survey, 7920 NW 71st 

Street, Gainesville, FL 32653-3071.

2Department of Fisheries and Aquatic Sciences, University of Florida, 7922 NW

71st Street, Gainesville, FL 32653-3071.

3North Carolina National Estuarine Research Reserve (NCNERR), Center for

Marine Science Research, 1 Marvin Moss Lane, Wilmington, NC 28403.
vi
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CONTENTS

Page

Figures ix

Tables xiii

Executive Summary xv

Chapter 1: Community Structure of Pinnacles Reef Tract Fishes

Background and Introduction 1

Objectives 6

Methods 22

Results 34

Discussion 62

Chapter 2: Trophic Ecology of Pinnacles Reef Tract Fishes

Introduction 69

Objectives 70

Methods 70

Results and Discussion 71

Acknowledgments 83

Literature Cited 85

Appendix A. Summary of ROV surveys for 1997-2001. A-1

Appendix B. A list of reef-fish species from natural reefs in the northern Gulf of 

Mexico with a comparison among the northwestern Gulf, 

Pinnacles, and West Florida shelf. B-1

Appendix C. Food habits of reef fishes of the Pinnacles Reef Tract by trophic 

guild. C-1

Appendix D. Characteristic ichthyofauna of Pinnacles Reef Tract, displaying the 

biodiversity of fishes by trophic guild. D-1
viii
ix

LIST OF FIGURES

Figure Page

1.1. Oblique view of the Alabama Alps study site and 36 Fathom Ridge area 

along the Pinnacles Reef Tract, showing the variability in outer shelf reef 

morphology (map provided by J. Gardner, USGS, Menlo Park, CA). 2

1.2. Northeastern Gulf of Mexico outer continental shelf with polygon marking 

the combined MMS MAMES and MASPTHMS study areas, enclosing the 

Pinnacles Reef Tract, including NEGOM-CMEP and USGS study sites 

(adapted from CSA and TAMU 2001). 4

1.3. Locations of NEGOM-CMEP megasites and monitoring sites (adapted 

from CSA and TAMU 2001). 5

1.4. USGS Pinnacles Reef Tract study sites in relation to NEGOM-CMEP 

megasites (adapted from CSA and TAMU 2001). 8

1.5. A. High-resolution multi-beam swath bathymetry for 40 Fathom Fishing 

Ground area with study sites. B. Acoustic backscatter for same area 

differentiating substrate type (provided by J. Gardner, USGS, Menlo Park, 

CA). 9

1.6. High-resolution multi-beam swath bathymetry for Roughtongue Reef 

study site (provided by J. Gardner, USGS, Menlo Park, CA). 10

1.7. High-resolution multi-beam swath bathymetry for Cat's Paw Reef study 

site (provided by J. Gardner, USGS, Menlo Park, CA). 12

1.8. High-resolution multi-beam swath bathymetry for Yellowtail Reef study 

site (provided by J. Gardner, USGS, Menlo Park, CA). 13

1.9. High-resolution multi-beam swath bathymetry for Near Shoreline Ridge 

area with study sites (provided by J. Gardner, USGS, Menlo Park, CA). 14

1.10. High-resolution multi-beam swath bathymetry for Double Top Reef study 

site (provided by J. Gardner, USGS, Menlo Park, CA). 15

1.11. High-resolution multi-beam swath bathymetry for 36 Fathom Ridge area 

with study sites (provided by J. Gardner, USGS, Menlo Park, CA). 17

1.12. High-resolution multi-beam swath bathymetry for Alabama Alps study site 

(provided by J. Gardner, USGS, Menlo Park, CA). 18
x

LIST OF FIGURES

(continued)

Figure Page

1.13. High-Resolution multi-beam swath bathymetry for Ludwick and Walton 

Pinnacles area with study sites (provided by J. Gardner, USGS, Menlo 

Park, CA). 19

1.14. Distribution of sampling effort at Roughtongue Reef (NEGOM-CMEP 

Site 1) for cruises 97-01 through 2000-01. 27

1.15. Sabiki jigs used to collect small reef fishes during this study, including 

Pronotogrammus martinicensis and Hemanthias vivanus. 28

1.16. A. Suction sampler device schematic. B. NURC-UNCW Phantom DS4 

ROV with suction sampler. C. A typical collection of fishes made by the 

suction sampler. 30

1.17. Still frames taken from video surveys at the Pinnacles Reef Tract. A. 

Pronotogrammus martinicensis school along crevice in reef top at 

Yellowtail Reef, 63 m. B. Close-up of mixed group of P. martinicensis

and Hemanthias vivanus along reef face. C. Mixed school of anthiines 

along reef face at CMEP-Site 5, 75 m. D. Single P. martinicensis along 

reef face at Scamp Reef, 105 m. 40

1.18. Still frames taken from video surveys at the Pinnacles Reef Tract. A. 

Mixed school of Hemanthias vivanus and Chromis enchrysura with a 

single Pristigenys alta on reef top biotope at Yellowtail Reef, 62 m. B. 

Holacanthus bermudensis and Chaetodon aya along northern edge of reef 

top biotope at Yellowtail Reef, 63 m. C-D. Reef top biotope with welldeveloped sessile invertebrate community of gorgonians and crinoids, 

Yellowtail Reef, 62 m. 41

1.19. Still frames taken from video surveys at the Pinnacles Reef Tract. A. 

Seriola dumerili schooling above reef flat top, Yellowtail Reef, 62m. B. 

Gonioplectrus hispanus at reef base, CMEP-Site 5, 78 m. C. Female 

Mycteroperca phenax on reef face at Scamp Reef, 105 m. D. Dominant 

male M. phenax exhibiting grey-head color phase along reef face, Scamp 

Reef, 105 m. 42

1.20. Typical ichthyofauna contributing to community structure on the 

Pinnacles Reef Tract, northeastern Gulf of Mexico, based on ROV data. 49
xi

LIST OF FIGURES

(continued)

Figure Page

1.21. Pinnacles Shallow Reef Tract fish faunal composition for reef crest biotope. 50

1.22. Pinnacles Shallow Reef Tract fish faunal composition for reef base biotope. 51

1.23. Pinnacles Shallow Reef Tract fish faunal composition for talus zone biotope. 52

1.24. Cluster of reef sites using percent similarity and unweighted pair-group 

mean average combinatorial strategy. 54

1.25. Cluster analysis of taxa using percent similarity and unweighted pairgroup mean average combinatorial strategy. 56

 1.26. Constancy and fidelity for site and taxa groups defined by cluster analysis. 57

2.1. Qualitative food web model for reef fishes of the Pinnacles Reef Tract 

constructed from stomach content analysis. 77

2.2. Quantitative food web model for reef fishes of the Pinnacles Reef Tract 

constructed from stomach content analysis. 78
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 LIST OF TABLES

Table Page

1.1. Summary of USGS Pinnacles Reef Tract study locations. 7

1.2. Summary of research activities conducted from 1997-2000. 24

1.3. Previous investigations of the ichthyofauna of the Pinnacles Reef Tract, 

comparing number of reef fishes observed by ROV in each study. 34

1.4. List of fishes documented in the Pinnacles Reef Tract from 1997 - 2000. 35

1.5. Abundance of top 30 taxa observed in ROV surveys partitioned by survey 

period (day/night) and reef tract (shallow/deep) for reef habitat. 44

1.6. Results of chi-square goodness of fit test for reef fish abundance by survey 

period (day/night) and reef tract (shallow/deep). 45

1.7. Day-night comparison of selected community parameters and top six taxa 

(abundance) paired by reef based on ROV data. 46

1.8. Biotope association of Pinnacles Reef Tract fishes based on ROV surveys 

during Cruise 97-01, combined for all study sites. 48

1.9. Summary statistics for ROV data by site group based on contributing to 

cluster analysis. 55

1.10. Abundance of top 15 predatory fishes caught by baited hook and line 

angling from 1997-2000. 59

1.11. Abundance of top 15 fish taxa caught in traps from 1997-2000. 60

1.12. Abundance of top 15 fish taxa caught in trawls in 1997. 61

2.1. Feeding guilds defined by characteristic prey type. 72

2.2. Dietary analysis of characteristic reef fishes of the Pinnacles Reef Tract by 

trophic guild. 73

2.3. Feeding guilds of deep reef fishes along the Pinnacles Reef Tract. 75
xiv
xv

EXECUTIVE SUMMARY

Topographic prominences form reef-like features along the outer continental shelf (OCS) 

edge and continental slope of the southeastern United States and Gulf of Mexico. An 

extensive set of reef features, known as the Pinnacles Reef Tract, occurs off Mississippi 

and Alabama at depths between 60 and 110 m. In this area recent research has focused 

largely on distribution of reef structures, their geology and physical characteristics, and 

processes relating to their origin. Community ecology studies have been limited in 

scope, confined mainly to distribution of sessile megafaunal and incidental observations 

of fishes. This study’s goal was to determine patterns of fish community structure and 

trophic relationships on Pinnacles reef and reef-associated biotopes. We examined 

variation in hard-bottom reef fish assemblages with reef topography and geographic 

location, to identify faunal differentiation associated with reef profile, biotopes, area, and 

depth. In addition, trophodynamic investigations were undertaken using food habits and 

food web structure of Pinnacles Reef Tract fishes. Analysis of food web structure was 

used to identify the relative importance of resident reef fishes (autochthonous) versus offreef trophic resources (allochthonous). This study was a component of the overall 

Minerals Management Service (MMS) Northeastern Gulf of Mexico Coastal and Marine 

Ecosystem Program (NEGOM-CMEP), designed to compliment other synoptic programs. 

It focused on hard-bottom features that serve as essential fish habitat for reef fishes on the 

OCS in areas of oil and gas development.

Ten sites within the combined Mississippi-Alabama Marine Ecosystems Study and 

Mississippi-Alabama Shelf Pinnacle Trend Habitat Mapping Study 

(MAMES/MASPHTHMS) area were visited on the initial cruise. An additional eight 

study sites were visited on subsequent cruises. High Resolution Multi-Beam Swath 

bathymetric maps were completed in the Pinnacles Reef Tract to provide detailed maps 

of the geomorphology of study reefs. These high-resolution images have been used to 

guide subsequent sampling. The combined sampling effort for all cruises comprised 326 

stations, apportioned into 112 angling, 63 trap, 22 bottom trawl, 58 remotely operated 

vehicle (ROV), 15 dredge/core/grab, and 37 plankton stations, with a small number of 

stations representing bottom surveys, CTD casts, light meter deployments, and 

exploratory sampling gear. All sampling combined returned over 6,000 specimens for 

food habits analyses, taxonomic verification and documentation, and subsequent life 

history analyses. Shipboard photographs documenting 113 fish species were obtained. 

ROV surveys resulted in about 85 hours of reef and reef-associated biotope observations 

and were used for analysis of resident fish fauna.

The results of this study have expanded the number of fishes documented from the 

Pinnacles Reef Tract, and is the first extensive collection of deep (greater than 60 m) reef 

fishes made in this region, and possibly worldwide. Of the 159 species documented from 

collected specimens or via videotape, 88 were identified from ROV censuses, and 70 can 

be classified as obligate reef-fishes belonging to the Caribbean reef-fish fauna. Such 

obligate reef fishes have life histories tied to three-dimensional hard-bottom habitats, and 

would not otherwise be found on the OCS open shelf. An additional 32 species are 

facultative reef associates that range broadly over OCS habitats, but occur on reef habitat, 
xvi

when available. ROV surveys revealed the dominance of planktivorous reef fishes on 

high profile reef features. Anthiine fishes (small planktivorous seabasses) numerically 

dominate the deep reef fish community, and may serve as a keystone species by foraging 

in the water column and transferring energy to the deep reef ecosystem. The diurnal 

nature of the fauna is evident in that the total number of individuals was significantly 

lower at night especially on shallow reefs. While small diurnal planktivores are 

considered to be an important link between open water plankton and the reef, nocturnal 

planktivores may play a relatively important role in the overall energetics of the reef 

community. Based on ROV analysis of fish faunal composition, species richness and 

relative abundance of reef fishes differs among biotopes (reef top, reef crest, reef face, 

reef base, talus zone, and sand flats). Cluster analysis grouped reefs by depth trend and 

relief.

At least 250 reef-associated fishes have been documented from offshore banks, reefs, and 

other hard-bottom habitats in the northern Gulf of Mexico based on previous literature. 

The suite of primary reef fishes inhabiting the Pinnacles Reef Tract represents a 

depauperate, winnowed fraction of the Caribbean reef-fish fauna. It can be hypothesized 

that the known Pinnacles reef fish diversity is diminished relative to the Caribbean due 

to: 1) distance from the source fauna, 2) depth limitation, 3) temperature limitation, 4) 

habitat limitation due to turbidity, 5) trophic limitations, 6) fishing pressure, or 7) 

incomplete sampling. Environmental conditions per se do not appear to control reef fish 

distribution in the area. Based on the analysis presented here the lack of a full range of 

reef habitats, such as coral and coralline algae dominated communities, is the greatest 

influence on taxonomic structure. In addition, heavy fishing pressure may have 

substantially altered the reef fish community. Trophic structure can assist in evaluation 

of overfishing of large predators and the associated shift in the reef fish community. The 

numerical dominance by small and medium planktivorous reef fish taxa in shelf-edge reef 

communities in the Pinnacles Reef Tract has likely been magnified by removal of 

piscivores through recent increases in commercial fishing activity in this region.

Trophodynamic investigations undertaken in the present study include feeding habits and 

food web structure of Pinnacles Reef Tract fishes. Defining trophic pathways is the key 

to developing an understanding of the ecological framework in the reef fish community. 

Analysis of food web structure also identifies relative importance of resident reef fishes 

versus off-reef trophic resources such as adjacent soft-substrate and pelagic prey in the 

diet of large predatory fishes, predator-prey relationships among fishes and invertebrates, 

and may also identify potential pathways of energy transfer from fishes to the benthic 

invertebrate assemblage through the release of fecal material.

A food web model was developed for the Pinnacles fish community, based on numerical 

contribution of prey to the diet of each predator. Over 1000 individuals were examined 

for food habits, with approximately 50% containing prey items. Fish were assigned to 

feeding guilds based on stomach contents to identify similarity in prey resources among 

species, sources of prey, and overall patterns in trophic structure. The numerically 

dominant species (greater than 80% of the individuals observed by ROV) on high profile 

reefs, roughtongue bass and red barbier, have diets dominated by calanoid copepods, 
xvii

gastropod larvae, and a variety of other pelagic mesoplankton. The generalized carnivore 

guild was represented by numerous taxa including tattler and a variety of serranids, 

scorpaenids, muraenids, and holocentrids. This guild contains the most taxa of deep reef 

fishes, although individual species are not as numerically abundant as planktivores and 

benthic carnivores in reef communities.

Anthiine fishes may serve as a keystone predator by foraging in the water column and 

transferring energy to the deep reef ecosystem. Anthiines in turn become prey for a 

variety of reef predators. Dense populations of Pinnacles fishes appear to exceed levels 

that could be supported solely by locally produced prey resources. Fishes on deep reefs 

must therefore be “subsidized” by passive transport of prey from surrounding benthic or 

pelagic habitats on water currents, or via active feeding migrations of predators to 

adjacent areas of the water column or the underwater landscape. Large predators, 

including greater amberjack and red snapper, utilize prey from pelagic, reef, and softbottom habitats and provide pathways of energy transfer.

The numerical dominance of planktivorous reef fish taxa in shelf-edge reef communities 

and vermilion snapper in our hook and line samples in the Pinnacles Reef Tract have 

likely been magnified by removal of piscivores through recent increases in commercial 

fishing activity in this region and historical fishing practices throughout the northwestern 

Gulf of Mexico since the mid 1800’s. While our observations of trophic structure in the 

deep reef community closely resemble those observed in the Indo-Pacific, the reef fish 

community structure may have been greatly altered by human activity through fishing 

during the past century.

Fishes and invertebrates have been characterized at the Pinnacles Reef Tract by three 

major studies and all major reef features have been precisely mapped. This area may be

the most well-known deep hard bottom area in the world. Thus we postulate that the 

majority of the reef fish species that occur at depths between 60 and 110 m are now 

documented. Yet due to the large area and diversity of habitat further scientific inquiry 

will continue to add to our knowledge of the fish fauna. Our present knowledge of the 

Pinnacles ichthyofauna can be used to characterize the fundamental taxonomic composition 

and trophic structure of deep reef-fish communities throughout southeastern United States 

and Gulf of Mexico OCS areas and therefore be useful in assessing future anthropogenic 

impacts on faunal structure.
xviii
CHAPTER 1

COMMUNITY STRUCTURE OF PINNACLES REEF TRACT FISHES

BACKGROUND AND INTRODUCTION

Topographic prominences form reef-like features along the continental shelf edge and 

continental slope of the southeastern United States and Gulf of Mexico. Such reefs have 

been known in the northern Gulf for over 70 years (Trowbridge 1930, Shepard 1937, 

Ludwick and Walton 1957, Moore and Bullis 1960). Shepard (1937) discussed the 

distribution of salt domes throughout the northwestern Gulf of Mexico based on 

fathometer surveys conducted by the U. S. Coast and Geodetic Survey in 1936, and 

Goedicke (1955) applied the term "pinnacles" to describe outer shelf features in the 

northern Gulf, including East and West Flower Garden Bank. Subsequently, Ludwick 

and Walton (1957) followed suit, using the term pinnacles to refer to steep-sided reef-like 

structures on the outer shelf in the northeastern Gulf. Historically, the term pinnacles 

included the entire class of reef structures rising from the open shelf, whether the top of 

the structure is spire-like, flat, or ragged, and regardless of the areal size or height of the 

overall structure (Fig. 1.1).

Shelf-edge prominences, including northeastern Gulf pinnacles, include structures of very 

heterogeneous origins (Gittings et al. 1992a, Gardner et al. 2001). West of the 

Mississippi, many shelf-edge banks are formed by the extrusion of salt domes (diapirs) 

from the surrounding seafloor, and are either capped with contemporary communities of 

reef-building organisms such as scleractinian corals, or are characterized by exposed 

carbonate rocks that are colonized by a diverse assemblage of sessile invertebrates 

(Rezak et al. 1985, Gittings et al. 1992a, Hardin et al. 2001). In the western Gulf off 

south Texas, shelf-edge prominences are non-diapiric drowned fossil coral-algal reefs 

(Rezak et al. 1985). East of the Mississippi, relatively few diapiric structures occur and 

are primarily located along the western portion of the Mississippi-Alabama shelf 

(Gittings et al. 1992a).

Most shelf-edge (60 to 110 m) features in this region appear to be non-diapiric, drowned, 

or fossil reefs initially formed during low sea level stands during the Pleistocene, and 

colonized by invertebrate communities [Continental Shelf Associates (CSA) 1992, Sager 

et al. 1992, Gittings et al. 1992a, Hancock 1997, Continental Shelf Associates and Texas 

A&M University (CSA & TAMU) 2001, Gardner et al. 2001]. Hard-substrate features 

supporting reef-like communities also occur on slide scar features that are exposed by 

slumping of the upper continental slope (Gardner et al. 2001) or on banks built by 

ahermatypic corals such as Lophelia pertusa (Linnaeus 1758) (Moore and Bullis 1960, 

Rogers 1999).

With few exceptions (e.g., Flower Garden Banks) outer continental shelf (OCS) reefs 

throughout the northern Gulf of Mexico lie in waters that are either too deep for sufficient 

light penetration, or seasonally too cold or too turbid to support hermatypic corals and 
2

Figure 1.1. Oblique view of the Alabama Alps study site and 36 Fathom Ridge area 

along the Pinnacles Reef Tract, showing the variability in outer shelf reef 

morphology (map provided by J. Gardner, USGS, Menlo Park, CA). 

Vertical exaggeration = 10X. Isobaths in meters.

most forms of algae (Rezak et al. 1985). In the northeastern Gulf, OCS deep reefs (>60 

m) are predominantly fossil structures that do not harbor hermatypic corals, extensive 

coralline or leafy algae communities (CSA 1992, Gittings et al. 1992a, Sager et al. 1992, 

Hancock 1997, CSA and TAMU 2001). There appears to be considerable production of 

biogenic debris by benthic organisms that colonize these reefs, as evidenced by coarse 

composition of sediment and high acoustic backscatter samples near features (Sager et al. 

1992, Sager and Schroeder 2001, Gardner et al. 2002, Gardner et al. 2001). Moreover, 

many topographic features display evidence of extensive historical subaerial erosion (i.e., 

flattening of reef tops at lower sea level stages) and submarine sculpturing by scouring 

and bioerosion (resulting in the formation of basal undercuts, hollows and caves). 

Although the major coral reef builders are absent in pinnacle megafaunal communities, 

some biological deposition continues due to growth of ahermatypic corals, bryozoans, 

calcareous worms, and other calcium depositing and cementing organisms on the surface 

of these features (Sager et al. 1992, Hancock 1997, CSA and TAMU 2001).

Reef communities of the OCS have been comprehensively studied in the northwestern 

Gulf (Bright et al. 1984, Rezak et al. 1990, Gittings et al. 1992b, Gittings and Hickerson 

1998). However, assessment of fish community structure has been primarily taxonomic 

and qualitative (Bright and Cashman 1974, Rezak et al. 1985), or semi-quantitative

(Dennis and Bright 1988a). Long-term monitoring on the Flower Garden Banks has 

concentrated on corals, with limited attention to quantitative assessment of fish 

populations (Gittings et al. 1992b).

Early investigations on the distribution of OCS reefs in the northeastern Gulf began in the 

middle of the 20th century. Relative to the technology of the time, the pioneering research of 

Ludwick and Walton (1957) was remarkable in its scope and breadth, laying the foundation 
3

for most subsequent geological and biological research on OCS deep reefs in the Gulf of 

Mexico. Using single-beam acoustic transects, Ludwick and Walton mapped the location of 

OCS pinnacles all along the northern Gulf from the Mississippi Delta to Port St. Joe, 

Florida. They produced the first bathymetric chart of the zone of pinnacles and 

reconstructed basal topography underlying the pinnacles. They measured physical and 

chemical seawater properties, mapped sediments and foraminiferal populations, and 

identified the major biological contributors to fossil reef formation from dredged reef rock. 

Ludwick and Walton also identified living sessile macrofauna and provided the first 

underwater photographs of pinnacles epifauna and bottom morphology.

Subsequent to Ludwick and Walton’s work, major study programs of the MississippiAlabama Pinnacles Reef Tract (referred to hereafter as Pinnacles Reef Tract) have 

proceeded under the auspices of the Minerals Management Service (MMS) including the 

Mississippi-Alabama Marine Ecosystems Study (MAMES) (Laswell et al. 1990, Brooks 

1991), Mississippi-Alabama Shelf Pinnacle Trend Habitat Mapping Study (MASPTHMS) 

(CSA 1992), and Northeastern Gulf of Mexico Coastal and Marine Ecosystem Program 

(NEGOM-CMEP) (CSA and TAMU 2001). These studies have conducted bathymetric and 

habitat reconnaissance, side-scan mapping, oceanographic, physical, and biological 

characterization and monitoring of pinnacle features and their hard-bottom communities. 

These investigations have provided the site locations, bathymetric charts, physical 

background, and macrofaunal framework for this study of pinnacle reef fish communities 

(Figs. 1.2, 1.3). Our community structure study complements work accomplished 

simultaneously under the NEGOM-CMEP program.

In the northeastern Gulf, recent research on OCS reefs has focused largely on distribution 

of reef structures, their geology and physical characteristics, and processes relating to 

their origin (Brooks 1991; CSA 1992, Sager et al. 1992, Sager and Schroeder 2001). 

Community ecology studies have been limited in scope, confined mainly to distribution 

of sessile megafaunal invertebrates (Gittings et al. 1992a, CSA and TAMU 2001), and 

incidental observations of fish communities (Shipp and Hopkins 1978; Gittings et al. 

1991, Thompson et al. 1999). Snyder (2001) investigated some aspects of reef fish 

community structure (taxonomic diversity, habitat association) during the NEGOMCMEP program.

The present study affords a more comprehensive evaluation of community structure, 

biotope affinities, and trophic ecology of the Pinnacles Reef Tract fish fauna. Here we 

examine variation in hard-bottom reef fish assemblages with reef topography and 

geographic location, to identify faunal differentiation associated with reef profile, 

biotopes, area, and depth. We also identify food habits and trophic structure of Pinnacles 

fishes (Chapter 2). The present study led to the initiation of important high-resolution 

multi-beam swath bathymetry of Pinnacles habitat/topography as is depicted in the study 

sites description.
Figure 1.2. Northeastern Gulf of Mexico outer continental shelf with polygon marking the combined MMS MAMES and 

MASPTHMS study areas, enclosing the Pinnacles Reef Tract, including NEGOM-CMEP and USGS study sites (adapted 

from CSA and TAMU 2001).

4
Figure 1.3. Locations of NEGOM-CMEP megasites (large numerals) and monitoring sites (small numerals) (adapted from CSA and 

TAMU 2001).

5
6

OBJECTIVES

A component of the overall NEGOM-CMEP Program, this study was designed to 

investigate aspects of fish community structure and compliment other synoptic programs. 

The study focused on hard-bottom features that serve as essential fish habitat for reef 

fishes on the OCS in areas of oil and gas development. The overall goal was to 

determine patterns of fish composition, community structure, and trophic relationships on 

Pinnacles reef and reef-associated biotopes. Project objectives were as follows:

· Using a broad range of sampling and video methods, qualitatively define taxonomic 

composition of fishes associated with steep-sided, reef-like, hard-bottom structures 

comprising the Pinnacles Reef Tract of the northeastern Gulf of Mexico.

· Quantitatively define species composition, abundance, dominance rank order, and 

species richness of fishes according to reef type, relief, and biotope. In addition, the fish 

fauna of the near reef zone and surrounding soft bottom area were also characterized.

· Define food habits and tropic structure of Pinnacles reef fishes (Chapter 2).

Study Sites

Bathymetric charts produced during the MAMES (Laswell et al. 1990, Brooks 1991), 

MASPTHMS (CSA 1992), and NEGOM-CMEP programs (CSA and TAMU 2001) were 

used to locate areas of interest during the initial USGS reconnaissance cruise in 1997. Ten 

potential sites within the combined MAMES/MASPHTHMS polygon were visited on the 

initial cruise (Table 1.1, Fig. 1.4). An additional eight study sites were visited on 

subsequent cruises. During May/June 2000, High Resolution Multi-Beam Swath (HRMBS) 

bathymetric maps were completed in the Pinnacles Reef Tract to provide detailed maps 

of the geomorphology of study reefs (Gardner et al. 2000, Gardner et al. 2001, Gardner et 

al. 2002). High-resolution images resulting from that survey are included in this report, 

and were used to guide subsequent sampling.

Many of the sites for this study were selected within NEGOM-CMEP megasites to make 

use of data collected there by other investigators (Figs. 1.3, 1.4). Study reefs were 

selected from both the shallow reef trend (65-80 m) and deep reef trend (85-110 m). 

Eight main study reefs (five shallow, three deep) selected for fish community structure 

and trophodynamics study are described below:

1) Roughtongue Reef 1- is a roughly elliptical (400 m major base diameter), high 

profile, flat-top structure with steep vertical sides (Figs. 1.5A, 1.6). The reef lies within 

NEGOM-CMEP Megasite 1 in the eastern part of the study area, in Destin Dome Lease 

Blocks 532 and 533, and corresponds to NEGOM-CMEP Site 1 (CSA and TAMU 2001) 

(Fig. 1.3). The general area containing this and the next two target reefs has historically 

 

1

 names were designated by USGS for convenient reference and are not official geographic names
Table 1.1. Summary of USGS Pinnacles Reef Tract study locations. Geographic coordinates indicate general reef location.

Reef Name Latitude Longitude Base Crest Max Relief Reef Area Corresponding

(N) DD (W) DD Depth Depth Relief Category (ha) NEGOM-CMEP Location

(m) (m) (m)

Alabama Alps 29.2518 88.3373 88 72 16 High 20.0 Megasite 5, Site 7

Cat's Paw Reef 29.4396 87.5870 78 64 14 High 23.9 Megasite 1

Corkscrew Reef 29.4426 87.5445 78 66 12 High 13.8 none

Double Top Reef 29.3920 87.9830 80 68 12 High 3.0 Megasite 3

Far Tortuga 29.5571 87.4616 70 66 4 Intermediate 74.8 none 

Ludwick and Walton Pinnacle 1 29.3268 87.7715 110 100 10 High 9.0 Megasite 4

L&W Pinnacle 2 29.3273 87.7680 110 100 10 High 7.0 Megasite 2, Site 4

L&W Pinnacle A 29.3428 87.7406 106 100 6 High 5.2 Megasite 2

L&W Pinnacle B 29.3386 87.7546 104 98 6 High 11.0 Megasite 2

L&W Pinnacle C 29.3400 87.7466 110 102 8 High 6.7 Megasite 2

L&W Pinnacle D 29.3350 87.7591 104 100 4 Intermediate 2.5 Megasite 2 

L&W Pinnacle H 29.3265 87.7625 110 104 6 High 5.3 Megasite 2

NEGOM-CMEP Site 5 29.3938 87.9810 80 68 12 High 0.7 Megasite 3, Site 5

Patch Reef Field 29.4415 87.6915 71-77 69-75 3 Low 1000.0 none

Porgy Reef 29.4371 87.6180 78 68 10 High 19.4 Megasite 1

Roughtongue Reef 29.4415 87.5785 78 64 14 High 15.4 Megasite 1, Site 1

Scamp Reef 29.3250 87.7765 110 100 10 High 4.6 Megasite 2 

Shark Reef 29.3965 88.0193 77 74 3 Low 1.2 Megasite 3

Solitary Mound 1 29.4558 87.6641 71 66 5 Intermediate 0.8 none 

Solitary Mound 2 29.4643 87.6320 72 64 8 High 12.6 none 

Triple Top Reef 29.3978 87.9915 76 68 8 High 3.6 Megasite 3

Yellowtail Reef 29.4403 87.5751 68 60 8 High 13.9 Megasite 1

7
8

Figure 1.4. USGS Pinnacles Reef Tract study sites in relation to NEGOM-CMEP megasites (open boxes) (adapted 

from CSA and TAMU 2001).
9

Figure 1.5. A. High-resolution multi-beam swath bathymetry for 40 Fathom Fishing 

Ground area with study sites. B. Acoustic backscatter for same area 

differentiating sediment type. Horizontal lines are due to sampling 

artifacts. Red indicates high backscatter and blue low backscatter 

(provided by J. Gardner, USGS, Menlo Park, CA). Isobaths in meters.
Figure 1.6. High-resolution multi-beam swath bathymetry for Roughtongue Reef study site (provided by J. Gardner, USGS, Menlo 

Park, CA). Vertical exaggeration = 9X. Transect line drawn from point A to B depicts reef in cross-section. Isobaths in 

meters.

10
11

been called the 40 Fathom Fishing Ground by fishermen, and in the MAMES program 

this area is referred to as the Eastern Delta Mounds (CSA and TAMU 2001). 

Roughtongue Reef belongs to the shallow pinnacle trend, with a base depth of 80 m (Fig. 

1.6). Extensive areas of accumulated sediments occur on the reef flat top and a diverse, 

dense invertebrate assemblage occurs throughout the interior (Hardin et al. 2001). 

Vertical rock faces, boulders, and rock outcrops occurring along the sides and 

surrounding the feature display eroded surfaces, and typically have lower invertebrate 

densities, dominated by the solitary coral Rhizopsammia manuelensis Chevalier 1966 and 

other small ahermatypic species (Hardin et al. 2001). Attached to the main reef is a 

smaller mound immediately to the south. The reef is surrounded by an extensive area of 

high acoustic backscatter resulting from coarse reef-derived sediment or hard substrates 

(Fig. 1.5B, red area). The USGS designated name refers to the common name for the 

small planktivorous serranid, Pronotogrammus martinicensis, the roughtongue bass, 

which is extremely abundant on this reef.

2) Cat’s Paw Reef is a group of six small, medium to high profile, flat-topped mounds, 

arranged in the pattern of a “cat’s paw” print, with 5-10 m relief (Figs. 1.5, 1.7). This 

cluster of mounds lies about 1000 m west of Roughtongue Reef in the 40 Fathom Fishing 

Ground, NEGOM-CMEP Megasite 1 area (Figs. 1.4, 1.5, 1.7) and is within Destin Dome 

Lease Block 532. Individual reef formations within the feature have flat top communities 

present with limited sediment cover, highly eroded and sculpted rock surfaces with 

vertical faces along edges of features. Small soft corals are abundant on horizontal 

surfaces, solitary coral colonies (including R. manuelensis), with spiral sea whips, 

antipatharians, and crinoids also common. Coarse sand substratum was present around 

reef features, and silt was not evident on sediment surfaces.

3) Yellowtail Reef, a single, elliptical (200 m base diameter), high-profile, flat-top structure, 

that reaches the shallowest crest depth (60 m) of all study sites (Table 1.1, Fig. 1.8). This 

structure also belongs to the 40 Fathom Fishing Ground group, and lies within NEGOMCMEP Megasite 1 (Fig. 1.5A) within Destin Dome Lease Block 532. It forms the 

northwestern end of a reef arc, with Cat’s Paw Reef at the center, and Roughtongue Reef 

lying at the southeastern end. Like other reef features in the group, an extensive flat top area 

is present, characterized by accumulated sediments and a dense invertebrate assemblage 

dominated by octocorals, antipatharians, sponges, and coralline algae. Rock outcrops 

characterize the northern extent of the feature, and these areas are heavily colonized by 

sessile invertebrates and coralline algae. The USGS designated name refers to the yellowtail 

reeffish, Chromis enchrysura, which is particularly abundant on this reef.

4) Double Top Reef is a horseshoe shaped (100 m base diameter), high profile structure that 

consists of multiple flat-top mounds with steep vertical sides (Figs. 1.9, 1.10). This 

structure lies within NEGOM-CMEP Megasite 3 (Figs. 1.3, 1.4) in the central part of the 

study area, in Main Pass Lease Block 223. A secondary mound attached to the northeast 

corresponds to NEGOM-CMEP Site 5 (Fig. 1.9). In the MAMES program this general area 

is referred to as the Near Shoreline Ridge (Brooks 1991). Double Top Reef belongs to the 

shallow pinnacle trend in the northeastern Gulf, and also includes a similarly-shaped series 

of mounds in the study area referred to as Triple Top Reef, and an adjacent, low profile 
Figure 1.7. High-resolution multi-beam swath bathymetry for Cat's Paw Reef study site (provided by J. Gardner, USGS, Menlo Park, 

CA). Vertical exaggeration = 9X. Transect line drawn from point A to B depicts reef in cross-section. Isobaths in meters.

12
Figure 1.8. High-resolution multi-beam swath bathymetry for Yellowtail Reef study site (provided by J. Gardner, USGS, Menlo 

Park, CA). Vertical exaggeration = 9X. Transect line drawn from point A to B depicts reef in cross-section. Isobaths in 

meters.

13
Figure 1.9. High-resolution multi-beam swath bathymetry for Near Shoreline Ridge area with study sites (provided by J. Gardner, 

USGS, Menlo Park, CA).

14
Figure 1.10. High-resolution multi-beam swath bathymetry for Double Top Reef study site (provided by J. Gardner, USGS, Menlo Park, 

CA). Vertical exaggeration = 9X. Transect line drawn from point A to B depicts reef in cross-section. Isobaths in meters.

15
16

feature referred to as Pancake Reef that corresponds to NEGOM-CMEP Site 6 (Fig. 1.9). 

These features also have flat top communities characterized by high sediment cover and 

dense invertebrate assemblages dominated by octocorals and antipatharians, with few 

solitary corals. Vertical rock walls and overhangs are dominated by R. manuelensis and 

other solitary corals, and are highly sculpted and eroded. The USGS designated names 

refer to the topography of these reef structures. Double Top Reef and Triple Top Reef 

are not NEGOM-CMEP study sites.

5) Alabama Alps is a long, narrow, north-south aligned, high profile mound 

approximately 1000 m in length (Figs. 1.11, 1.12). This structure lies within NEGOMCMEP Megasite 5 in the western part of the study area, in Main Pass Lease Block 286, 

and corresponds in part to NEGOM-CMEP Site 7. In the MASPTHMS program this 

same area is referred to as Lagniappe Delta Shallow, and has historically been called the 

36 Fathom Ridge by fisherman. Alabama Alps forms the northwestern terminus of a 

long northwest to southeast-aligned ridge and pinnacle arc paralleling the shelf edge (Fig. 

1.1), and belongs to the shallow pinnacle trend of the northeastern Gulf. The top of this 

feature has sections of relatively flat terrain with scattered sections of sediment cover, 

particularly in the southern portion of the feature (Figs. 1.11, 1.12). Invertebrate 

assemblages on the flat sections are dominated by octocorals, antipatharians, and sponges 

(Hardin et al. 2001). The sides of the feature range from vertical walls to large attached 

monoliths where the solitary coral R. manuelensis is the dominant sessile invertebrate 

with crinoids, antipatharians, coralline algae, sponges, and other solitary corals present 

(Hardin et al. 2001). The USGS designated name refers to the precipitous terrain, 

particularly the near-vertical west-face scarp of the structure, and its position off the state 

of Alabama.

6) Ludwick and Walton Pinnacle 1 is the central member of a group of five medium to 

high-profile, spire-top, shelf-edge structures with 10 m maximum relief and a base depth 

of 110 m. This group belongs to the deep shelf-edge pinnacle trend in the northeastern 

Gulf (Fig. 1.13). These pinnacles form a short east-west aligned arc on the shelf-slope 

break, bordering the northern edge of a massive shelf-edge slump feature. Pinnacle 1 is 

one of those profiled and contoured by Ludwick and Walton (1957). It lies within the 

MAMES Western Delta Mounds area, NEGOM-CMEP Megasite 2 and within Destin 

Dome Lease Block 661. A fairly uniform coverage of coarse sand, rock fragments, shell 

fragments, and other calcareous debris surrounds the base of feature with occasional 

small rocky reef outcrops and patch reefs encrusted with R. manuelensis, octocorals, 

antipatharians, and crinoids. Emergent rocky features with vertical walls, rock ridges, 

and rock arches are distributed across the reef. Vertical rock faces have highly eroded 

surfaces, and are densely covered with R. manuelensis, with low coverage of other 

solitary corals, octocorals, sponges, and antipatharians.

7) Ludwick and Walton Pinnacle 2 is another of the deep shelf-edge pinnacle group. 

This structure, lying immediately to the east of Pinnacle 1, also was profiled and 

contoured by Ludwick and Walton (1957). It also lies within the MAMES Western Delta 

Mounds area, NEGOM-CMEP Megasite 2, and Destin Dome Lease Block 661. The 
17

Figure 1.11. High-resolution multi-beam swath bathymetry for 36 Fathom Ridge area 

with study sites (provided by J. Gardner, USGS, Menlo Park, CA). Isobaths 

in meters.
Figure 1.12. High-resolution multi-beam swath bathymetry for Alabama Alps study site (provided by J. Gardner, USGS, Menlo Park, 

CA). Vertical exaggeration = 9X. Transect line drawn from point A to B depicts reef in cross-section. Isobaths in meters.

18
Figure 1.13. High-resolution multi-beam swath bathymetry for Ludwick and Walton Pinnacles area with study sites (provided by J. 

Gardner, USGS, Menlo Park, CA). Vertical exaggeration = 10X. Distance from east to west = 6 km. Isobaths in meters.

19
20

feature is a broad mound that gradually slopes from the center, and is characterized by 

low relief hardbottom interspersed with a sediment veneer (Hardin et al. 2001). Scattered 

across the surface are individual rocky features reaching heights of 3 m. The elevated 

rocky features were colonized by dense populations of R. manuelensis, other solitary 

corals, octocorals, crinoids, and basket stars, while low relief hard bottom regions were 

characterized primarily by octocorals, antipatharians, and crinoids (Hardin et al. 2001). 

This pinnacle corresponds to NEGOM-CMEP Site 4.

8) Scamp Reef is a member of the Ludwick and Walton Pinnacles deep shelf-edge 

group with a precipitous southern reef face (Fig. 1.13). This structure, lying immediately 

to the west of Pinnacle 1, also was profiled and contoured by Ludwick and Walton 

(1957). It lies within the MAMES Western Delta Mounds area, NEGOM-CMEP 

Megasite 2 (Fig. 1.4), and within Viosca Knoll Lease Block 654. This feature has 

extensive vertical rock outcrops with profiles in excess of 5 m. Spectacular arches, 

overhangs and rugged topography occur along the southern face of the reef, with exposed 

rock colonized by R. manuelensis, antipatharians, crinoids, octocorals, and ahermatypic 

coral colonies. The USGS name Scamp Reef refers to the abundance of the scamp 

grouper, Mycteroperca phenax, that reside at this site.

9) Other Study Sites. Additional study sites were visited during single occasions on 

reconnaissance cruise 97-01 or subsequent research cruises, but were not intensively 

sampled during this project (Fig. 1.4). The majority of these sites are located in the eastern 

part of the study area, and are within or adjacent to CMEP Megasite 1.

Porgy Reef is a broad, gradually sloping mound, approximately 500 m in diameter, located 

west of Roughtongue Reef, and within the boundary of Megasite 1. Silty sand substrates 

surround the reef, and the interior of the feature is composed of sculpted rock outcrops. No 

visible sediments were accumulated on the reef interior, and a flat top invertebrate 

community was absent. Invertebrate assemblages were dominated by R. manuelensis, 

corkscrew sea whips, antipatharians, octocorals, and crinoids on exposed rock surfaces.

Shark Reef is an elongate, east to west oriented, low profile reef feature approximately 250 

m in length, located within CMEP Megasite 3, west of Double Top Reef and Triple Top 

Reef study sites. A well-developed invertebrate community was present on vertical walls 

and overhangs with small brown solitary corals and R. manuelensis abundant. Horizontal 

surfaces were heavily silted, and many sessile invertebrates appeared to be dead, with basal 

parts remaining but few living colonies visible. Pencil urchins were common on silted reef 

surfaces.

Solitary Mound 1 and Solitary Mound 2 are located to the northwest of Megasite 1. 

Solitary Mound 1 is a gently sloping mound of intermediate height less than 100 m in 

diameter. A coarse sand substratum surrounds the reef features, and rock surfaces are 

extremely eroded/sculpted. Large rocky caverns and depressions occur on the sides of the 

reef with R. manuelensis, stony corals, with numerous corkscrew sea whips and crinoids on 

the reef crest, and crinoids and soft corals common on vertical rock surfaces. Rocky 

outcrops around the base of the feature are colonized by R. manuelensis, octocorals, and 
21

long-spined urchins. Solitary Mound 2 is part of an intermittent ridge, and is an elongate 

feature approximately 800 m in length. This feature has a well-developed reef flat top with 

extensive sand and silt cover, where large red soft corals and other octocorals were 

abundant. Small rocky outcrops were present along the interior of the feature, along with 

numerous pits in the accumulated sediments.

Patch Reef Field is located to the east of Megasite 1 and is described in detail by Gittings et 

al. (1991). It is an area with numerous, low profile hard bottom outcroppings. Reef features 

in this area are only 2 to 3 m in maximum profile. Sessile invertebrates are relatively 

common on rock surfaces. At higher elevations, the reef community resembles larger 

features, with abundant solitary corals, octocorals, antipatharians, and sponges. Eroded, 

sculpted reef faces are encrusted with R. manuelensis and other solitary corals on overhangs, 

and corkscrew sea whips and antipatharians are common along the top and sides of rock 

outcroppings. As noted by Gittings et al. (1991), a thin layer of silt was present on reef 

surfaces, and many areas of the reef base, reef face, and small patch reefs were heavily 

silted. Flocculent silt was also present in the near-bottom water column. Most of the 

footage on this dive was over open sand bottom, with many small individual patch reefs 

encountered during the dive, as also observed by Gittings et al. (1991).

Far Tortuga Reef is the site farthest east in this study and is a broad, gradually sloping reef 

of intermediate profile, approximately 800 m in diameter. Scattered rock outcrops on the 

feature have typical invertebrate assemblages, with small octocorals and antipatharians 

abundant on rock surfaces, and R. manuelensis, crinoids and sponges also common. The 

reef feature was composed of relatively small, low profile outcrops interspersed with sandy 

patches, with a silt veneer over coarse sand around the reef outcrops. High profile 

formations or a flat-top community were not encountered.

Corkscrew Reef is a series of three high profile, gradually sloping mounds, each 

approximately 300 m in diameter that are located directly to the east of Megasite 1 (Table

1.1). This site is characterized by coarse sand with low silt surrounding the reef, and coarse 

sand on the reef. Reef surfaces were highly eroded, sculpted rock faces with R. manuelensis 

and large solitary corals on surfaces. Patchy rock outcrops with low invertebrate cover and 

abundant corkscrew sea whips and antipatharians on the top of reef features. No welldeveloped flat top community was visible at this site, with the rocky crests of reef features 

harbored dense assemblages of corkscrew sea whips and other antipatharians.

Ludwick and Walton Pinnacles A-D, H includes a number of additional sites distributed 

throughout the Ludwick and Walton deep shelf-edge pinnacles tract, ranging from 200 to 

1000 m in diameter (Fig. 1.13, Table 1.1). Reefs along the Ludwick and Walton Pinnacles 

tract are surrounded by sand and silt substrate at the base of the feature, with elevated highly 

sculpted and eroded rock surfaces and extensive caves and depressions. Dominant 

invertebrates on the deep pinnacles are R. manuelensis, other solitary corals, octocorals, 

antipatharians, and crinoids. Invertebrate density is highest on vertical surfaces and rocky 

crests. Flat top reef communities with accumulated sediments and dense invertebrate 

assemblages characteristic of high profile shallow reefs were not observed at any of these 

sites.
22

METHODS

Cruise Activities

The initial reconnaissance and sampling cruise USGS 97-01 was conducted on the R/V 

SUNCOASTER 4-13 August 1997 (92 stations). The remaining six USGS-funded cruises 

(234 stations) were completed on the R/V TOMMY MUNRO (Table 1.2). The combined 

sampling effort for all cruises comprised 326 stations, apportioned into 112 angling, 63 trap, 

22 bottom trawl, 58 remotely operated vehicle (ROV), 15 dredge/core/grab, and 37 plankton 

stations, with a small number of stations representing bottom surveys, CTD casts, light 

meter deployments, and exploratory sampling gear (Table 1.2). All sampling combined 

returned over 6,000 specimens for food habits analyses, taxonomic verification and 

documentation, and subsequent life history analyses. Shipboard photographs documenting 

113 species were obtained. ROV tapes resulted in about 85 hours of observation of reef and 

reef-associated biotopes and the resident fish fauna for analysis.

Sampling was conducted during all seven research cruises spanning the time interval of 

1997-2000 (Table 1.2). Positions for all sampling sites were determined by Differential 

Global Positioning System (DGPS) and charts were provided by CSA-TAMU collaborators. 

Sampling conducted during the 2000-01 cruise was further georeferenced using digital maps 

of HRMBS topography and vessel position tracked in ArcView software to display station 

location (Fig. 1.14). Operations were conducted continuously 24 hours per day.

Hook and Line Angling

During all cruises, demersal and pelagic fishes from hard-bottom reefs were obtained 

using hook and line angling with bait and large hooks (for macro-carnivores) and angling 

with unbaited, multi-hook “Sabiki” jigs (for planktivores and micro-carnivores) (Fig. 

1.15). Sabiki jigs are constructed of a variety of small hook sizes (5 to 20 mm in length) 

with added strips of iridescent film to imitate shrimps or other small planktonic 

organisms, and each pre-packaged unit contains from 4 to 7 hooks. Sabiki jigs were 

deployed using 142 to 340 g lead sinkers, depending upon water depth and currents, and 

were successfully used to collect fishes to 110 m depth. Fishes were primarily collected 

during daylight hours, as the majority of individuals captured using this technique were 

visual planktivores.

Traps

A variety of sizes and mesh traps including large (1.8 m X 1.5 m X 0.6 m) chevron traps 

with 25 mm X 12 mm mesh to box traps (1.2 m X 1.2 m X 0.6 m) of 25 mm hexagonal 

wire were used to obtain fish specimens to confirm visual identification and sample 

cryptic taxa not easily observed by the ROV. Some traps were equipped with 3 or 6 mm 

Vexar mesh liner to retain small specimens. Traps were deployed in trap lines made up 

of two to four traps. Also attached to these lines were smaller collection devices such as 

minnow traps, PVC bundles, and bottles in an attempt to attract and capture small shelter 

seeking fishes.
23

Trawls

A 4.9 m semi-balloon trawl with a 3.8 cm mesh body and a 0.6 cm mesh liner was used 

to collect bottom fishes near reefs at 13 stations and a 7.6 m semi-balloon trawl with a 3.8 

cm mesh body and a 0.6 cm mesh liner at five stations. Both trawls were equipped with a 

“Texas” roller rig made up of 7.6 cm rubber disks on a chain in front of the head rope to 

reduce hangs on reef structures.

Specimen Disposition

In-situ collections provided voucher specimens to verify and document species 

identifications, and material for food habits studies. On board the research vessel, 

individual voucher fish specimens were carefully preserved, pinned out to display 

coloration and diagnostic features, and photographed using digital and 35 mm cameras on 

a water bath light table (Randall 1961). Photographs were taken by D. Weaver, W. 

Smith-Vaniz, and J. Caruso. Specimens were preserved in 10% buffered formalin or 

frozen in the field, and later transferred to 70% ethanol in the laboratory. Selected 

voucher specimens were catalogued in the National Museum of Natural History (NMNH) 

and Florida Museum of Natural History (FMNH) ichthyology collections. The remaining 

voucher material, metadata, and voucher photographs are maintained at the Florida 

Caribbean Science Center (FCSC).

Other gears

Additional shipboard activities included plankton sampling to document pelagic prey 

availability to the deep reef fish community, and collection of physicochemical 

parameters of the water column. These data will be reported elsewhere.

ROV Methodology

Collection of specimens was complemented by use of ROVs to conduct habitat 

reconnaissance and document faunal composition via color video camera. All dives were 

recorded to videotape for subsequent laboratory analyses. A Phantom DS4 ROV, 

equipped with a color video camera, was the primary system used for in-situ observations. 

Three different Phantom ROVs were employed during the seven Pinnacles research cruises. 

United States Navy (97-01) and NMFS (99-03) ROVs were operated by NMFS Mississippi 

Laboratories during the first three years of the program, and the NOAA/NURP Phantom 

DS4 was operated by NURC-UNCW on the remaining cruises. Due to technical difficulties 

and image quality, only ROV data from cruises 97-01, 99-03, and 2000-01 were used for 

quantitative purposes (Appendix A). ROV observations were used to define and compare 

fish faunal composition at each main target study reef by depth, reef height (high, medium 

and low profile), and biotope.
Table 1.2. Summary of research activities conducted from 1997-2000. Number of stations listed by study site and sampling gear.

24

 Main Study Sites Other Study Sites

 Cruise No. 

 Alabama Alps

 Cat's Paw Reef

 Double Top Reef

 Roughtongue Reef

 Triple Top Reef

 Yellowtail Reef

Patch Reef Field Corkscrew Reef

 CMEP- Site 5

 Far Tortuga Reef

 L & W Pinnacle 1 L & W Pinnacle 2

 L & W Pinnacles A-H

 Scamp Reef

 Porgy Reef

 Shark Reef

 Solitary Mound 1  Solitary Mound 2 

Date Sampling Gear

ROV Day 1 2 4 1 5 1 1 1 2 1

ROV Night 1 1 1 1

Angling AM 2 4

Angling PM 2 3 1 1 1 1 2 1

97-01 Angling Night 2 1 1 1 1 1

4-13 Aug 1997 Plankton

Traps 2 1 1 1 1

Trawls 2 1 1 1 2

CTD 1 2 1 1 1

ROV Day

ROV Night

Angling AM 1 4 1

Angling PM 1 1 2

98-02 Angling Night 1 1

8-10 Oct 1998 Plankton

Traps 3 3

Trawl

CTD 

ROV Day

ROV Night

Angling AM 3 2 2

Angling PM 1 1 1 1

 99-01 Angling Night 1

18-23 Feb 1999 Plankton 5 1 1

Traps 4 1 1

Trawl

CTD 
Table 1.2. (continued).

25

 Main Study Sites Other Study Sites

 Cruise No. 

 Alabama Alps

 Cat's Paw Reef

 Double Top Reef

 Roughtongue Reef

 Triple Top Reef

 Yellowtail Reef

 Patch Reef Field

 Corkscrew Reef

 CMEP- Site 5

 Far Tortuga Reef

 L & W Pinnacle 1 L & W Pinnacle 2

 L & W Pinnacles A-H

 Scamp Reef

 Porgy Reef

 Shark Reef

 Solitary Mound 1  Solitary Mound 2 

Date Sampling Gear

ROV Day 1 1

ROV Night

Angling AM 2 2 2 2 2

99-02 Angling PM 1 3 2 1 1 1 1

30 May- Angling Night 1 1 1

4 June 1999 Plankton 2 4 2 2 4 1

Traps 4 4 6 4 1

Trawl

CTD 

ROV Day 2 1 2 2

ROV Night 1 1

Angling AM 1 1

Angling PM 3

99-03 Angling Night 1

20-22 Aug 1999 Plankton 2 1

Traps 1

Trawl 1

CTD 1 1 1 1
Table 1.2. (continued).

26

26

 Main Study Sites Other Study Sites

 Cruise No. 

 Alabama Alps

 Cat's Paw Reef

 Double Top Reef

 Roughtongue Reef

 Triple Top Reef

 Yellowtail Reef

 Patch Reef Field

 Corkscrew Reef

 CMEP- Site 5

 Far Tortuga Reef

 L & W Pinnacle 1 L & W Pinnacle 2

 L & W Pinnacles A-H

 Scamp Reef

 Porgy Reef

 Shark Reef

 Solitary Mound 1  Solitary Mound 2 

Date Sampling Gear

ROV Day23111

ROV Night111

Angling AM13311

Angling PM21311

2000-01 Angling Night11

5-10 Mar 2000 Plankton211

Traps62715

Trawl4

CTD 

ROV Day12

ROV Night

Angling AM11

Angling PM1

2000-02 Angling Night

5-6 Sept 2000 Plankton11

Traps

Trawl

CTD 
27

Figure 1.14. Distribution of sampling effort at Roughtongue Reef (NEGOM-CMEP 

Site 1) for cruises 97-01 through 2000-01. Each symbol represents the 

position of the research vessel during sampling gear deployment. Gray 

circles-angling, black triangles-traps, open squares-ROV dives.
Figure 1.15. Sabiki jigs used to collect small reef fishes during this study, including Pronotogrammus martinicensis (top and 

middle) and Hemanthias vivanus (bottom).

28
Transecting methodology were drawn from those previously employed in ROV/ 

submersible/camera sled studies that have shown that videotape analysis of large

mobile demersal fishes requires a moving image and a low oblique perspective to capture 

diagnostic features of morphology and coloration (Grassle et al. 1975, Cohen and Pawson 

1977, Uzmann et al. 1977, Rice et al. 1979, Parker and Ross 1986, Hecker 1987, Butler et 

al. 1991, Sulak and Ross 1993, 1996, Adams et al. 1995, Felley and Vecchione 1995). 

ROV transects were conducted at a constant speed of 0.10-0.15 m/sec across the bottom, 

with altitude of the ROV held to a maximum of 1.0 m (distance between sea floor and 

video camera lens, varying somewhat according to terrain). Horizontal swath width was 

resolved as the lower border of the video frame with the video camera zoom set to full 

wide-angle, and pan and tilt angles set to about 25° and 20°, respectively. Video was 

recorded on board the ship on S-VHS or Hi-8 videotape with audio annotation.

In situ collections of small reef fishes were accomplished using a suction sampler fitted to 

the ROV (Fig. 1.16) that dispensed a suspended rotenone solution (co-designed by D. 

Weaver and L. Horn, UNC-Wilmington). Powdered rotenone (Prentiss, Inc.) was 

suspended in 3 L of concentrated Ivory Liquid dishwashing detergent, and mixed with 8 

L of seawater. Approximately 3 kg of standard table salt and 10 kg of granular sugar was 

added to increase the specific gravity of the solution to help maintain contact with the 

bottom. The suspended rotenone was passed through a standard kitchen strainer to 

remove large clumps, and loaded into the dual reservoir chambers of the sampler. A 

standard marine bilge pump, encased in an aluminum housing and powered from the 

surface, was used to dispense the rotenone solution and collect fishes at select reef sites 

during ROV dives. Fishes were retained in a 5 mm mesh liner fitted inside the first PVC 

chamber (Fig. 1.16).

Videotape Analysis

Videotapes were transferred to S-VHS format and analyzed on a commercial quality S-VHS 

VCR with variable speed editor/jogger control (slow-motion, frame by frame advance) to 

facilitate species identification at slow speed advance and enumerate individuals. Videotape 

analysis was used to quantify fish abundance by taxa and biotope. Incidental observations 

of pelagic fishes also were recorded and individuals identified. In addition to moving 

videotape transects, the zoom function of the NURC-UNCW ROV video camera was 

periodically used to capture close-up images of individuals to facilitate identification during 

cruises 2000-01 and 2000-02. These still video segments were used to verify species 

identifications made during transect surveys. All fish species were identified to the lowest 

possible taxonomic unit, with confirmation from close-up video images and by reference to 

voucher specimens collected by angling, trapping, or ROV suction sampler.

Habitat Parameters

In addition to defining fish community structure, videotape screening was used to make 

qualitative notes on the physical habitat and megafaunal invertebrates associated with 

particular biotopes and fish assemblages. Where appropriate (e.g., high profile, flat 
30

Figure 1.16. A. Suction sampler device schematic. B. NURC-UNCW Phantom DS4 

ROV with suction sampler. C. A typical collection of fishes made by the 

suction sampler.

 A 

B Photograph by Tom Potts, NURC C
31

topped features) fish observed were assigned to one of six biotopes: reef top, reef crest, 

reef base, reef talus (circum-reef sediment apron), and soft bottom.

Reef Top is referred to as flat reef tops or reef flats by Gittings et al. (1992a) and 

describes the biotope that develops on large, flat-topped mounds such as Roughtongue 

Reef, Yellowtail Reef, and CMEP-Site 5. This biotope is identified by extensive pockets 

of accumulated sediments and rocky debris, interspersed with low profile rock outcrops, 

ridges, and crevices. Invertebrate assemblages on the reef flat top community exhibit 

high density and species richness, and are dominated by erect sponges, octocorals 

(particularly sea fans, including Nicella sp.), comatulid crinoids, antipatharians, 

gorgonocephalid basket stars, bryozoans, and coralline algae (Gittings et al. 1992a, 

Hardin et al. 2001). Our reef top category corresponds to top interior locations of Hardin 

et al. (2001) and continuous hard bottom category of Snyder (2001).

Reef Crest biotopes were defined as the interface between the near vertical, rocky reef 

face and the sediment covered flat top characterized by dense assemblages of 

invertebrates. Reef crest biotopes typically were characterized by extensive rocky 

outcrops with small areas of sediment cover and low invertebrate densities. We 

distinguished the reef crest ecotone from the adjacent flat reef top and vertical reef face 

biotopes to identify the possible influence of position in the leading edge of water 

currents on the reef fish community. Our reef crest category corresponds to the top edge 

location listed by Hardin et al. (2001).

Reef Face as referred to by Gittings et al. (1992a), are rugged, often near-vertical rocky 

surfaces that characterize the steep-sided edges of large flat-topped features and the steep 

walls of narrow, spire shaped deep water pinnacles. Reef face communities are 

characterized by a lower overall density of epifauna than reef tops, with an abundance of 

ahermatypic corals, including R. manuelensis, Madrepora sp., and Madracis/Oculina sp., 

comatulid crinoids, a variety of octocoral fans, antipatharians including the spiral whip 

Stichopathes lutkeni Brooks, 1889, coralline algae (to about 75 m), and sea urchins 

(Gittings et al. 1992a, Hardin et al. 2001). This biotope is described as reef side by 

Hardin et al. (2001) and Snyder (2001).

Reef Base was defined as the ecotone between the steep reef face and the talus zone, 

where the rugged rocky face of the reef was often undercut and formed small caves and 

overhangs. This region contained both rocky vertical faces with abundant solitary corals 

and the coarse sediments and rocky debris/talus resulting from bioerosion occurring on 

the reef above. This biotope is also identified by Hardin et al. (2001) and Snyder (2001).

Reef Talus (circum-reef sediment apron) is listed as debris fields or rubble flats by 

Gittings et al. (1991, 1992a). These are flat areas of reef debris and coarse carbonate 

sediments occurring on the flanks of large, high-relief mounds (Gittings et al. 1992a, 

Sager and Schroeder 2001). Coarse sediments and debris are produced by shell material 

resulting from bioerosion, and rock fragments from the main reef. Small rocky outcrops 

in this biotope are often encrusted with solitary corals, small octocoral fans, and 

antipatharians (Gittings et al. 1992a, Hardin et al. 2001). The reef talus zone is 
32

identifiable on HRMBS maps as the area of highest acoustic backscatter (red) to the south 

and west of hard bottom features (Gardner et al. 2000) (Fig. 1.5B).

Soft Bottom/Sand Plain refers to the relatively flat substrate surrounding reef features, 

characterized by fine to coarse quartz sand without visible portions of shell hash or rock 

rubble. These areas are often flat and featureless, but occasionally are contoured by 

ripples, gentle waves, or numerous excavated burrows, pits, and mounds that add to the 

topography. Sessile invertebrates in this habitat are limited to occasional small 

octocorals or antipatharians that attach to buried rock surfaces. Non-reef associated soft 

bottom is distinguishable in acoustic backscatter maps as areas of lowest backscatter 

(Fig. 1.5B, blue).

Data Analysis

Survey Period and Reef Tract Comparisons

ROV dives were divided into day and night periods, and data from crepuscular periods 

were excluded from this analysis. A two-way chi-square goodness of fit test was used to 

compare abundance between survey period and reef tract. A sequential Bonferroni 

procedure was used to adjust significance values for multiple tests (Sokal and Rohlf, 

1995). At five reefs, Alabama Alps, Ludwig & Walton Pinnacles, Scamp, Yellowtail, and 

Roughtongue Reefs, both day and night data were obtained by ROV. Although sample 

sizes were small, a paired t-test was used to test day-night differences in total fish 

abundance, number of taxa, and top six taxa abundance (Sokal and Rohlf, 1995). All 

data were standardized by effort measured as video survey time in minutes.

Cluster Analysis

We used cluster analysis to help explore and define the structure of fish assemblages 

among reefs and biotopes. Data were summarized by reef and survey period. Data from 

Ludwick and Walton Pinnacles 1, 2, A, B and C were combined into a single reef 

category. Raw counts were standardized by effort expressed as number of fish per 

minute of video survey time for comparisons among reefs. Survey time was not recorded 

by biotope so data were double standardized for analysis. Taxa were restricted to those 

that occurred at two or more sites to reduce the potential effect of bias from rare species 

(Wolda, 1981). Two measures of similiarity, percent similarity and Morisita, were used 

for comparison as they emphasize different attributes (Bloom, 1981; Wolda, 1981). An 

agglomerative hierarchical clustering strategy was used with an unweighted pair-group 

mean average sorting algorithm to obtain grouping of reefs and taxa (Boesch, 1977). The 

COMPAH computer program was used to perform the analysis (Gallagher, 1998). 

Clusters were subjectively divided into groups to facilitate comparisons. We compared 

cluters from the two similarity measures. If a reef or taxa was found in different groups 

between clusters it was considered poorly resolved and a weak member of the group; thus 

not indicative of the group structure.
33

Percent Similarity

Morisita-Horn Similarity

Both sites (reefs) and taxa were clustered and nodal analysis was performed to enhance 

the interpretation of the data (Boesch, 1977). The relationship between reefs and taxa 

groups was measured as constancy, the faithfulness of a taxa group to a particular reef 

group and fidelity, the degree to which taxa are limited to reef groups. Constancy and 

fidelity were arbitrarily categorized from high to low to better elucidate relationships 

between reefs and taxa.

Constancy

Cij = aij

(Ni Nj)

Fidelity

Fij = (aij ? Nj)

(Nj ? aij)

where aij number of occurrences of taxa

from taxa group i in site group j.

Ni total number taxa in taxa group i.

Nj total number taxa in taxa group j.
34

RESULTS

Faunal Composition

The results of this study have expanded the number of fishes documented from the 

Pinnacles Reef Tract, and is the first extensive collection of deep reef fishes made in this 

region, and possibly worldwide (Table 1.3). Of the 159 species documented (Table 1.4) 

from collected specimens or via videotape, 88 were identified from ROV censuses, and 

65 can be classified as obligate reef-fishes belonging to the Caribbean reef-fish fauna. 

Such obligate reef fishes have life histories tied to three-dimensional, hard-bottom 

habitats, and would not otherwise be found on the OCS open shelf (Figs. 1.17-1.19). An 

additional 32 species are facultative reef associates that range broadly over OCS habitats, 

but occur on reef habitat, when available. The remaining species include primarily 

Carolinian soft-substrate species, including both coastal and shelf-dwelling fishes, along 

with a small number of pelagic and deep midwater species that were found over reef 

features with varying frequency and regularity.

Table 1.3. Previous investigations of the ichthyofauna of the Pinnacles Reef Tract, 

comparing number of reef fishes observed by ROV in each study.

Study Reef Fish Taxa Total Fish Taxa Voucher Specimens?

Darnell, 1991b 39 70 No

Snyder, 2001 49 76 No

This study 65 88 Yes
35

Table 1.4. List of fishes documented in the Pinnacles Reef Tract from 1997-2000. 

Scientific and common names follow Robins et al. (1991) except where noted 

with asterisk. Columns denoted as PRF - primary reef fishes, A - hook and 

line angling, As - Sabiki angling, Ro – ROV observed, Rc – ROV collected, 

Tp – Trap, Tl – Trawl, and P – voucher specimen photograph.

Scientific Name Common Name PRF A As Ro Rc Tp Tl P

Lamnidae - mackerel sharks

Isurus oxyrinchus shortfin mako X X

Scyliorhinidae - cat sharks

Scyliorhinus retifer chain dogfish X X X

Triakidae – houndsharks

Mustelus canis smooth dogfish X X X

Mustelus sinsusmexicanus gulf smoothhound X

Carcharhinidae - requiem sharks

Carcharhinus falciformis silky shark X

Rhizoprionodon terraenovae Atlantic sharpnose shark X

Sphyrnidae – hammerheads

Sphyrna lewini scalloped hammerhead X

Squatinidae - angel sharks

Squatina dumeril Atlantic angel shark X X

Rajidae - skates

Raja eglanteria clearnose skate X

Raja texana roundel skate X X

Muraenidae – morays

Gymnothorax hubbsi1lichen moray X X X

Gymnothorax kolpos blacktail moray X X X X X

Gymnothorax moringa spotted moray X X X

Gymnothorax nigromarginatus blackedge moray X X X

Gymnothorax saxicola honeycomb moray X X X

Muraena retifera reticulated moray X X X X

Ophichthidae - snake eels

Ophichthus gomesii shrimp eel X X

Ophichthus puncticeps palespotted eel X X X

Congridae - conger eels

Conger oceanicus conger eel X

Rhynchoconger gracilior* whiptail conger X

Nettostomatidae - duckbill eels

Hoplunnis diomedianus blacktail pike-conger X X

Clupeidae - herrings

Etrumeus teres round herring X

Argentinidae – argentines

Glossanodon pygmaeus pygmy argentine X

Aulopidae – flagfins

Aulopus filamentosus yellowfin flagfin X X

Synodontidae - lizardfishes

Saurida normani shortjaw lizardfish X X X

Synodus foetens inshore lizardfish X

Synodus intermedius sand diver X X X X

Trachinocephalus myops snakefish X X

Myctophidae - lanternfishes X

Carapidae - pearlfishes

Echiodon dawsoni chain pearlfish X

2

1

new Gulf of Mexico record

2

stomach content 

*revised nomenclature
36

Table 1.4. (continued).

Scientific Name Common Name PRF A As Ro Rc Tp Tl P

Ophidiidae - cusk-eels

Brotula barbata bearded brotula X X X X X

Lepophidium brevibarbe blackedge cusk-eel X X

Lepophidium jeannae mottled cusk-eel X X

Ophidion beani longnose cusk-eel X

Otophidium omostigmum polka-dot cusk-eel X

Moridae - codlings

Physiculus fulvus metallic codling X

Bregmacerotidae - codlets

Bregmaceros cantori pale codlet X X X

Phycidae* – phycid hakes

Urophycis floridana southern hake X X

Urophycis regia spotted hake X X

Batrachoididae – toadfishes

Opsanus pardus leopard toadfish X X X X

Porichthys plectrodon Atlantic midshipman X X

Antennariidae – frogfishes

Antennarius ocellatus ocellated frogfish X

Antennarius radiosus singlespot frogfish X X

Ogcocephalidae – batfishes

Halieutichthys aculeatus pancake batfish X X

Ogcocephalus corniger longnose batfish X X X

Ogcocephalus declivirostris slantbrow batfish X X

Holocentridae – squirrelfishes

Corniger spinosus spinycheek soldierfish X X X X

Holocentrus adscensionis squirrelfish X X X X

Ostichthys trachypoma bigeye soldierfish X X

Sargocentron bullisi* deepwater squirrelfish X X X X

Caproidae - boarfishes

Antigonia capros deepbody boarfish X X

Syngnathidae - pipefishes

Hippocampus sp. seahorse X

2

Aulostomidae – trumpetfishes

Aulostomus maculatus trumpetfish X X X

Fistulariidae – cornetfishes

Fistularia petimba red cornetfish X X X X

Scorpaenidae - scorpionfishes

Pontinus rathbuni highfin scorpionfish X X X X

Scorpaena agassizii longfin scorpionfish X X

Scorpaena dispar hunchback scorpionfish X X X X

Triglidae – searobins

Bellator militaris horned searobin X X

Prionotus paralatus Mexican searobin X X

Prionotus roseus bluespotted searobin X X

Prionotus rubio blackwing searobin X X

Prionotus stearnsi shortwing searobin X X X

PRF – primary reef fish A – Angling As – Sabiki angling Ro – ROV observed Rc –ROV collected

 Tp – Trap Tl – Trawl P – Photograph

2

stomach content

*revised nomenclature
37

Table 1.4. (continued).

Scientific Name Common Name PRF A As Ro Rc Tp Tl P

Serranidae - sea basses

Anthias tenuis threadnose bass X X X X

Centropristis ocyurus bank sea bass X X X X

Centropristis philadelphica rock sea bass X X X

Epinephelus drummondhayi speckled hind X X X X

Epinephelus flavolimbatus yellowedge grouper X X X X

Epinephelus niveatus snowy grouper X X X X X

Epinephelus nigritus warsaw grouper X X X

Gonioplectrus hispanus Spanish flag X X X X

Hemanthias leptus longtail bass X X X X

Hemanthias vivanus red barbier X X X X X

Liopropoma eukrines wrasse bass X X X X X

Mycteroperca microlepis gag X X X X X

Mycteroperca phenax scamp X X X X X

Paranthias furcifer creole-fish X X X X X

Pronotogrammus martinicensis* roughtongue bass X X X X X

Rypticus maculatus whitespotted soapfish X X X X

Serranus atrobranchus blackear bass X X X

Serranus notospilus saddle bass X X X

Serranus phoebe tattler X X X X

Grammatidae – basslets

Lipogramma regia1royal basslet X X

Opistognathidae – jawfishes

Opistognathus lonchurus moustache jawfish X X X X

Priacanthidae - bigeyes

Priacanthus arenatus bigeye X X X X

Pristigenys alta short bigeye X X X X

Apogonidae - cardinalfishes

Apogon pseudomaculatus twospot cardinalfish X X X X

Malacanthidae - tilefishes

Caulolatilus chrysops goldface tilefish X X X X

Caulolatilus intermedius anchor tilefish X X

Malacanthus plumieri sand tilefish X X X X X

Pomatomidae - bluefishes

Pomatomus saltatrix bluefish X X X

Echeneidae - remoras

Echeneis naucrates sharksucker X

Carangidae - jacks

Caranx crysos blue runner X X

Caranx hippos crevalle jack X

Selar crumenophthalmus bigeye scad X X

Seriola dumerili greater amberjack X X X X X

Seriola fasciata lesser amberjack X X X X

Seriola rivoliana almaco jack X X X X

Seriola zonata banded rudderfish X X

Trachurus lathami rough scad X X X

PRF – primary reef fish A – Angling As – Sabiki angling Ro – ROV observed Rc –ROV collected

 Tp – Trap Tl – Trawl P – Photograph

1

new Gulf of Mexico record

*revised nomenclature
38

Table 1.4. (continued).

Scientific Name Common Name PRF A As Ro Rc Tp Tl P

Lutjanidae - snappers

Lutjanus campechanus red snapper X X X X X

Pristipomoides aquilonaris wenchman X X X X

Rhomboplites aurorubens vermilion snapper X X X X X X

Sparidae – porgies

Calamus leucosteus whitebone porgy X X X

Calamus nodosus knobbed porgy X X X

Lagodon rhomboides pinfish X X X X

Pagrus pagrus red porgy X X X X X

Stenotomus caprinus longspine porgy X X X

Sciaenidae - drums

Cynoscion arenarius sand seatrout X X

Leiostomus xanthurus spot X X X

Micropogonias undulatus Atlantic croaker X X X

Pareques iwamotoi blackbar drum X X X X

Pareques umbrosus cubbyu X X X X

Mullidae - goatfishes

Mullus auratus red goatfish X X

Upeneus parvus dwarf goatfish X X X

Chaetodontidae - butterflyfishes

Chaetodon aya bank butterflyfish X X X X X

Chaetodon ocellatus spotfin butterflyfish X X X X

Chaetodon sedentarius reef butterflyfish X X X X X

Pomacanthidae - angelfishes

Holacanthus bermudensis blue angelfish X X X X

Holacanthus ciliaris queen angelfish X X

Holacanthus tricolor rock beauty X X

Pomacentridae - damselfishes

Chromis enchrysura yellowtail reeffish X X X X X

Chromis insolata sunshinefish X X

Chromis scotti purple reeffish X X

Labridae - wrasses

Bodianus pulchellus spotfin hogfish X X X X

Decodon puellaris red hogfish X X X X X

Halichoeres bathyphilus greenband wrasse X X X X

Uranoscopidae - stargazers

Kathetostoma albigutta lancer stargazer X X

Blenniidae - combtooth blennies

Parablennius marmoreus seaweed blenny X X

2 X

Gobiidae - gobies

Coryphopterus punctipectophorus spotted goby X X

Risor ruber tusked goby X X

 3

Sphyraenidae – barracudas

Sphyraena borealis northern sennet X X

Gempylidae* - snake mackerels

Neoepinnula americana American sackfish X

Trichiuridae - cutlassfishes

Trichiurus lepturus Atlantic cutlassfish X X X

PRF – primary reef fish A – Angling As – Sabiki angling Ro – ROV observed Rc –ROV collected

 Tp – Trap Tl – Trawl P – Photograph

2

stomach content

3

box core

*revised nomenclature
39

Table 1.4. (continued).

Scientific Name Common Name PRF A As Ro Rc Tp Tl P

Scombridae - mackerels

Acanthocybium solandri wahoo X

Auxis thazard frigate mackerel X

Euthynnus alletteratus little tunny X

Sarda sarda Atlantic bonita X

Scomberomorus cavalla king mackerel X X

Stromateidae - butterfishes

Peprilus burti gulf butterfish X

Bothidae - lefteye flounders

Engyophrys senta spiny flounder X X

Trichopsetta ventralis sash flounder X X

Paralichthyidae* - sand flounders

Ancylopsetta dilecta three-eye flounder X X X

Citharichthys cornutus horned whiff X X X

Cyclopsetta fimbriata spotfin flounder X X X

Paralichthys squamilentus broad flounder X X

Syacium papillosum dusky flounder X X X

Achiridae - American soles

Gymnachirus texae fringed sole X X

Soleidae* - soles

Symphurus diomedianus spottedfin tonguefish X X X

Symphurus parvus pygmy tonguefish X

Balistidae - triggerfishes

Balistes capriscus gray triggerfish X X X

Monacanthidae* - filefishes

Monacanthus tuckeri slender filefish X X X

Ostraciidae - boxfishes

Acanthostracion quadricornis* scrawled cowfish X

Tetraodontidae - puffers

Canthigaster rostrata sharpnose puffer X X X X X

Canthigaster jamestyleri 1goldface toby X X X X

Lagocephalus laevigatus smooth puffer X X

Sphoeroides dorsalis marbled puffer X X

Sphoeroides spengleri bandtail puffer X X X

Diodontidae - porcupinefishes

Chilomycterus antillarum web burrfish X X

Chilomycterus schoepfii striped burrfish X

Diodon holocanthus balloonfish X X

Molidae – ocean sunfishes

Mola mola ocean sunfish X

PRF – primary reef fish A – Angling As – Sabiki angling Ro – ROV observed Rc –ROV collected

 Tp – Trap Tl – Trawl P – Photograph

1

new Gulf of Mexico record

*revised nomenclature
A

C

B

D

Figure 1.17. Still frames taken from video surveys at the Pinnacles Reef Tract. A. Pronotogrammus martinicensis school 

along crevice in reef top at Yellowtail Reef, 63 m. B. Close-up of mixed group of P. martinicensis and 

Hemanthias vivanus along reef face. C. Mixed school of anthiines along reef face at CMEP-Site 5, 75 m. D. 

Single P. martinicensis along reef face at Scamp Reef, 105 m. C and D illustrate invertebrates that dominate the 

drowned reef community [ahermatypic coral colonies, Rhizopsammia manuelensis (black clusters), and 

crinoids].

40
41

A B

C D

Figure 1.18. Still frames taken from video surveys at the Pinnacles Reef Tract. A. Mixed school of Hemanthias vivanus and 

Chromis enchrysura with a single Pristigenys alta on reef top biotope at Yellowtail Reef, 62 m. B. Holacanthus 

bermudensis and Chaetodon aya along northern edge of reef top biotope at Yellowtail Reef, 63 m. C-D. Reef top 

biotope with well-developed sessile invertebrate community of gorgonians and crinoids, Yellowtail Reef, 62 m.
D

A B

C

Figure 1.19. Still frames taken from video surveys at the Pinnacles Reef Tract. A. Seriola dumerili schooling above reef flat 

top, Yellowtail Reef, 62m. B. Gonioplectrus hispanus at reef base, CMEP-Site 5, 78 m. C. Female Mycteroperca 

phenax on reef face, Scamp Reef, 105 m. D. Dominant male M. phenax exhibiting grey-head color phase along 

reef face, Scamp Reef, 105 m.

42
43

ROV Survey

Videotape data from only three cruises (USGS 97-01, 99-03 and 2000-01) were of 

sufficient quality for quantitative analysis of reef fish abundance. Fishes observed during 

other ROV surveys where visibility was poor, or while collecting fishes using the suction 

sampler (USGS 2000-01 and 2000-02), were added to the list of species occurring on the 

Pinnacles Reef Tract but not included in the analysis. Eighty-eight species were 

identified during ROV surveys, with 17 taxa observed only by this method (Table 1.4).

ROV surveys revealed the dominance of planktivorous reef fishes on high profile reef 

features (Table 1.5, Appendix A). Anthiines were the principal taxa observed by day, 

and communities were dominated by either Hemanthias vivanus or Pronotogrammus 

martinicensis, depending on reef location and biotope. Day ROV surveys yielded higher 

(9719 individuals) overall abundance of fishes than night surveys (2624 individuals) 

(Table 1.5, Appendix A1), indicating most reef species are diurnally active and hide in 

crevices or burrows at night. About equal numbers of taxa were observed during the day 

(68) and night (66) though almost twice as much effort was applied during the day. Day 

surveys had 20 unique taxa, of which none were frequent. Night surveys had 18 unique 

taxa. Three taxa, Bregmaceros cantori, myctophids, and Trichiurus lepturus, were 

common, but not observed on all night dives. These typically non-reef species may make 

a more substantial contribution to the reefs than previously expected. We compared reef 

fish abundance by survey period and reef tract simultaneously as these factors might be 

confounded. Only data from reef habitat were used in this analysis. The chi-square 

goodness of fit test showed that most taxa had a significant difference in abundance 

between survey period and reef tract (Table 1.6). The diurnal nature of the fauna is 

evident in that the total number of individuals was significantly lower at night especially 

on shallow reefs. Seven taxa were significantly more abundant on shallow reefs during 

the day (Table 1.6). Only Synodus spp. was significantly lower during the day due to its 

absence from shallow water reefs. Two taxa, P. martinicensis and Anthias tenuis were 

significantly more abundant on deep reefs during the day. Five taxa were most abundant 

on shallow and six on deep reefs (Table 1.6). Pareques umbrosus was significantly 

lower on shallow reefs at night while Chaetodon aya, Decodon puellaris, and Serranus 

phoebe were lower on deep reefs.

When day/night comparisons were made by reef there was a highly significant difference 

in total fish abundance between day and night with higher numbers during the day (Table 

1.7). However, there was no significant difference in number of taxa observed between 

day and night. Of the top six taxa, four were more abundant during the day but none 

were significantly so due to the small sample size (Table 1.7). Pronotogrammus 

martinicensis tends to lie on the reef surface at night being inactive but visible. In 

contrast H. vivanus hides in crevices at night, thus hidden from the ROV survey. 

Myctophids, a nocturnally migrating taxon, were significantly more abundant at night 

overall, as was T. lepturus, but due to patchy distributions there was no significant 

difference among reefs.
44

Table 1.5. Abundance of top 30 taxa observed in ROV surveys partitioned by survey 

period (day/night) and reef tract (shallow/deep) for reef habitat.

Percent of Total

Overall Reef Habitat

Day Night

Rank Taxa No. Shallow Deep Shallow Deep

1

Pronotogrammus 

martinicensis 5027 25.1 77.3 46.7 27.4

2 Hemanthias vivanus 3881 54.4 3.6 15.9 0.3

3 Gobiidae 499 3.9 <0.1 0.4

4 Myctophidae 335 0.1 23.1

5 Anthias tenuis 272 <0.1 8.3 0.8

6 Chromis enchrysura 240 3.0 <0.1 3.1 0.1

7 Trichiurus lepturus 239 16.5

8 Pristigenys alta 231 1.1 1.8 4.6 3.2

9 Bregmaceros cantori 179 2.6 9.9

10 Chaetodon aya 161 1.2 1.3 2.8 0.7

11 Halichoeres bathyphilus 130 1.7 0.3 0.3 0.1

12 Trachurus lathami 123 0.3 2.0 5.5

12 Decodon puellaris 123 1.0 0.3 0.4 0.1

14 Pareques umbrosus 115 1.0 0.7 0.3 1.7

15 Serranus notospilus 98

16 Serranus phoebe 84 0.8 0.2 0.9

17 Canthigaster rostrata 83 0.6 0.1 3.1 0.1

18 Corniger spinosus 80 0.5 0.5 1.8 0.5

19 Synodus spp. 77 0.2 1.2

20 Liopropoma eukrines 74 0.8 0.5 0.3 0.2

21 Scorpaena dispar 66 0.2 0.3 2.6 0.3

22 Saurida spp. 64

23 Seriola dumerili 63 0.1 0.4 0.9 2.1

24 Chaetodon sedentarius 45 0.3 0.2 1.7

25 Paranthias furcifer 39 0.6 0.1

26 Rhomboplites aurorubens 36 0.3 1.4 0.1

27 Apogon pseudomaculatus 35 0.2 0.0 1.3 0.4

27 Mycteroperca phenax 35 0.3 0.3 0.3

29 Chromis scotti 29 0.4 0.1

30 Scorpaenidae 28 0.0 0.3 0.2 1.0

Total No. Individuals 12343 6574 3145 1175 1449

No. Taxa 66 60 42 48 43

No. Unique Taxa 11 5 3 7

Video Survey Time (min) 689 329 296 313
45

Table 1.6. Results of chi-square goodness of fit test for reef fish abundance by survey 

period (day/night) and reef tract (shallow/deep). Significantly higher or 

lower abundance is indicated by factor.

Day Night Overall

Rank Taxa Shallow Deep Shallow Deep P

1 Pronotogrammus martinicensis Higher <0.0001

2 Hemanthias vivanus Higher <0.0001

3 Gobiidae Higher <0.0001

4 Myctophidae Higher <0.0001

5 Anthias tenuis Higher <0.0001

6 Chromis enchrysura Higher <0.0001

7 Trichiurus lepturus Higher <0.0001

8 Pristigenys alta ns

9 Bregmaceros cantori Higher <0.0001

10 Chaetodon aya Lower 0.0005

11 Halichoeres bathyphilus Higher <0.0001

12 Trachurus lathami Higher <0.0001

12 Decodon puellaris Lower <0.0001

14 Pareques umbrosus Lower 0.0003

15 Serranus notospilus ND

16 Serranus phoebe Lower <0.0001

17 Canthigaster rostrata Higher <0.0001

18 Corniger spinosus ns

19 Synodus spp. Lower <0.0001

20 Liopropoma eukrines Higher <0.0001

21 Scorpaena dispar Higher <0.0001

22 Saurida spp. ND

23 Seriola dumerili Higher <0.0001

24 Chaetodon sedentarius Higher <0.0001

25 Paranthias furcifer Higher <0.0001

26 Rhomboplites aurorubens Higher <0.0001

27 Apogon pseudomaculatus Higher 0.0006

27 Mycteroperca phenax ns

29 Chromis scotti Higher <0.0001

30 Scorpaenidae Higher <0.0001

No. Individuals Higher Higher <0.0001

No. Taxa Higher <0.0001

ns – not significant ND – no data for comparison
46

Table 1.7. Day-night comparison of selected community parameters and top six taxa 

(abundance) paired by reef based on ROV data. Sample size equals five. 

Significant differences are indicated in bold.

Mean per Minute Video

Day Night t P

No. Individuals 16.48 4.00 3.28 0.01

No. Taxa 0.36 0.32 0.29 0.39

P. martinicensis 6.35 1.38 1.48 0.11

H. vivanus 5.44 0.24 2.10 0.05

Myctophids 0 0.68 -1.55 0.10

A. tenuis 0.85 0.02 1.00 0.19

Gobiids 0.17 0.01 1.37 0.12

T. lepturus 0 0.33 -1.01 0.18

Taxonomic groups that were not effectively sampled or video documented during the 

course of the study included juvenile anthiines and gobiids, due to their extremely small 

size. Gobies were the third most abundant species observed by ROV censuses (Table 

1.5), and may be represented by a number of species, but only the spotted goby, 

Coryphopterus punctipectophorus, was documented by close-up video during this study 

(Table 1.4).
47

Biotopes

From ROV analyses of fish faunal composition, we recognize that species richness and 

relative abundance of reef fishes differs among biotopes [reef zones (including reef top, 

reef crest, reef face, and reef base), talus zone, and sand flat habitats, Table 1.8, Figs. 

1.20-1.23], and these differences occur across the various study reefs. All reef biotopes 

are dominated by anthiine serranids, but showed partitioning. Hemanthias vivanus was

most abundant in reef top and reef crest biotopes while P. martinicensis was increasingly 

abundant on the reef face and reef base (Table 1.8). Additional reef fish taxa that 

numerically dominated reef top and reef crest biotopes included gobiids, the pomacentrid 

C. enchrysura, the priacanthid Pristigenys alta and the labrids Halichoeres bathyphilus 

and D. puellaris. The reef face biotope was dominated by anthiine serranids. The 

schooling serranid Paranthias furcifer, D. puellaris, and the holocentrid Corniger 

spinosus also characterize this biotope. Fishes common on the reef base biotope included 

species that typically are associated with crevices and holes, including the holocentrid 

Ostichthys trachypoma, the sciaenid Pareques umbrosus and the serranid Liopropoma 

eukrines. 

Fishes observed on the reef top were represented by more taxa (28) than reef crest, face, 

and base biotopes (19, 22 and 22, respectively). High species richness is likely due to the 

diversity of microhabitats available, including rock outcrops, accumulated coarse sand, 

and dense assemblages of sessile invertebrates such as sponges, gorgonians and 

antipatharians that form spatially complex structure. The diversity of habitat present on 

the reef top also presents a wide variety of prey resources that are associated with 

unconsolidated sediments and a diverse invertebrate assemblage.

A wide range of fishes also frequents the reef top biotope during night hours. Mobile 

demersal visual piscivores (groupers, snappers) are found on the reef top during evening 

hours, and schools of amberjack commonly occur on flat-topped features at night. Night 

ROV surveys also documented the occurrence of soft bottom sciaenids ( e.g., Leiostomus 

xanthurus) and carangids (Trachurus lathami) utilizing the reef top. Midwater planktivores 

from the adjacent slope, including B. cantori, T. lepturus, and lanternfishes (Myctophidae), 

have been occasionally observed over reef features at night, and were observed in great 

abundances on deeper sites such as the Ludwick and Walton Pinnacles (Appendix A4).

While reef biotopes were similar in overall species composition, they differed greatly 

from the talus zone and sand flat (Table 1.8, Figs. 1.22-1.23). Species assemblages were 

very similar among reef biotopes (percent similarity>0.40) and dissimilar from the talus 

zone and sand flat (percent similarity = 0.14) (Fig. 1.20). Overall number of taxa was 

lower in the talus zone (15) and sand flat (16) than reef biotopes. Gobiids were 

particularly abundant in the circum-reef talus zone (70.1%), with D. puellaris, and the 

serranids Serranus phoebe and S. notospilus were also dominant (Table 1.8, Appendix 

A). Lizardfishes (Synodus spp.) were also common in this biotope.
48

Table 1.8. Biotope association of Pinnacles Reef Tract fishes based on ROV surveys 

during Cruise 97-01, combined for all study sites. Numbers of individuals 

observed at each biotope during videotape analysis, and percent of total, are 

given. Top five taxa by biotope are indicated in bold.

REEF TOP REEF CREST REEF SLOPE REEF BASE TALUS ZONE SAND FLAT

Taxa No. Percent No. Percent No. Percent No. Percent No. Percent No. Percent

Hemanthias vivanus 1698 58.8 314 59.7 442 39.6 39 8.7 - -

Pronotogrammus martinicensis 618 21.4 135 25.7 536 48.1 308 68.8 1 0.5 4 1.2

Gobiidae 161 5.6 27 5.1 10 0.9 5 1.1 129 70.1 71 20.5

Chomis enchrysura 88 3.0 7 1.3 5 0.4 2 0.4 2 1.1 - -

Halichoeres bathyphilus 57 2.0 9 1.7 - - 3 0.7 7 3.8 - -

Pristigenys alta 43 1.5 1 0.2 8 0.7 2 0.4 1 0.5 - -

Decodon puellaris 37 1.3 1 0.2 13 1.2 8 1.8 18 9.8 19 5.5

Chaetodon aya 26 0.9 6 1.1 9 0.8 5 1.1 - - - -

Canthigaster rostrata 24 0.8 - - 2 0.2 - - - - - -

Trachurus lathami 19 0.7 - - - - - - - - 3 0.9

Serranus phoebe 15 0.5 3 0.6 3 0.3 6 1.3 8 4.3 5 1.4

Liopropoma eukrines 13 0.5 - - - - 9 2.0 1 0.5 - -

Scorpaena dispar 12 0.4 3 0.6 4 0.4 3 0.7 2 1.0 2 0.6

Rhomboplites aurorubens 6 0.2 - - 9 0.8 - - - - - -

Parablennius marmoreus 6 0.2 1 0.2 - - - - - - - -

Sargocentron bullisi 6 0.2 - - - - - - - - - -

Priacanthus arenatus 5 0.2 - - - - 3 0.7 1 0.5 - -

Pareques umbrosus 5 0.2 - - - - 26 5.8 - - - -

Mycteroperca microlepis 5 0.2 1 0.2 5 0.4 - - - - - -

Chaetodon sedentarius 5 0.2 5 1.0 3 0.3 2 0.4 - - - -

Seriola dumerili 4 0.1 3 0.6 - - - - - - - -

Holacanthus bermudensis 4 0.1 1 0.2 2 0.2 - - - - - -

Corniger spinosus 4 0.1 5 1.0 12 1.1 6 1.3 - - - -

Serranus notospilus 3 0.1 - - - - 4 0.9 3 1.6 84 24.2

Centropristis ocyurus 3 0.1 - - - - 1 0.2 - - 2 0.6

Synodus sp. 2 0.1 - - - - - - 6 3.3 40 11.5

Lutjanus campechanus 2 0.1 - - 2 0.2 - - - - - -

Mycteroperca phenax 2 0.1 - - 11 1.0 1 0.2 - - - -

Pontinus rathbuni - - 2 0.4 - - - - 1 0.5 - -

Pagrus pagrus - - 1 0.2 - - - - - - - -

Paranthias furcifer - - 1 0.2 15 1.3 - - - - - -

Pareques iwamotoi - - - - 5 0.4 2 0.4 - - - -

Rypticus maculatus - - - - 2 0.2 - - - - - -

Opsanus pardus - - - - 2 0.2 - - - - - -

Gonioplectrus hispanus - - - - 2 0.2 - - - - - -

Ostichthys trachypoma - - - - - - 11 2.5 - - - -

Hemanthias leptus - - - - - - 1 0.2 - - - -

Unidentified flatfish - - - - - - 1 0.2 - - 7 2.0

Saurida sp. - - - - - - - - - - 62 17.9

Pristipomoides aquilonaris - - - - - - - - - - 7 2.0

Hoplunnis sp. - - - - - - - - - - 7 2.0

Syacium papillosum - - - - - - - - - - 6 1.7

Upeneus parvus - - - - - - - - - - 3 0.9

Bregmaceros cantori - - - - - - - - 1 0.5 2 0.6

Ancylopsetta dilecta - - - - - - - - 1 0.5 - -

Other taxa 13 0.5 - - 13 1.2 - - 2 1.0 22 6.6

Totals 2886 100.0 526 100.0 1115 100.0 448 100.0 184 100.0 346 100.0
49

Biotope

Similarity

Figure 1.20. Typical ichthyofauna contributing to community structure on the Pinnacles Reef Tract, northeastern Gulf of Mexico, 

based on ROV data. Cluster diagram shows relationship among biotopes.

Percent Similarity

Biotope

Similarity
50

Figure 1.21. Pinnacles Shallow Reef Tract fish faunal composition for reef crest biotope.
Figure 1.22. Pinnacles Shallow Reef Tract fish faunal composition for reef base biotope.

51
Figure 1.23. Pinnacles Shallow Reef Tract fish faunal composition for talus zone biotope.

52
53

Sand flats were also dominated by gobies, with the small serranid S. notospilus, 

lizardfishes (Synodus spp. and Saurida spp.), D. puellaris and flatfishes also common 

(Table 1.8). Anthiine serranids were either absent (H. vivanus) or in low abundance (P.

martinicensis) at these biotopes.

Reef Groups and Species Associations

Fish assemblages from ROV data were compared among reefs using cluster analysis. 

Five groups of reefs were resolved at a 0.30 similarity level (Fig. 1.24). Table 1.9

compares reef type and community parameters among these groups. Taxa clustering 

resulted in eight groups at a 0.12 similarity level (Fig. 1.25).

Site groups 3 to 5 separate reefs with distinctive faunal compositions. Deeper reef night 

samples formed the most unique cluster (Site Group 5) with high constancy (faithfulness 

to a reef group) of Taxa Group G such as myctophids, T. lepturus, B. cantori, and T. 

lathami (Fig. 1.25). The Yellowtail Reef night survey was exceptional in the high 

constancy with Taxa Group G and very low abundance of P. martinicensis. This species 

is found on the bottom at night and was regularly seen during the night at other sites. Its 

low abundance at Yellowtail Reef may be due to the low relief of the area surveyed at 

night. Far Tortuga and Shark Reef (Site Group 3), both large mounds with gentle, silted 

slopes and no high vertical faces, shared high abundance of gobies and D. puellaris.

Site Group 2 contained most of the high relief shallow reef trend sites with the exception 

of Solitary Mounds. This site group had high constancy with Taxa Groups B and F the 

largest suite of taxa and strong fidelity (the degree to which taxa are limited to a reef 

group) with Taxa Group F that included the common taxa, H. vivanus and P. 

martinicensis (Fig. 1.26). Solitary Mounds fit with this group due to its high abundance 

of C. enchrysura. Site Group 2 reefs on average had high abundances and low species 

richness (Table 1.9).

Site Group 1 sites included deep reefs, night surveys of high diversity shallow water sites, 

and shallow low relief sites. This indicates that the deep reef fauna is most closely 

related to the nocturnal fauna of the shallow reef tract. No taxa group had high constancy 

for this site group, but there was strong fidelity with Group F taxa (Fig. 1.26). The low 

abundance of Taxa Group B, especially C. enchrysura and D. puellaris, in Site Group 1 

differentiates it from Site Group 2. Alabama Alps day and night surveys were very 

similar (percent similarity = 0.83) sharing 32% (20) of the taxa. In particular A. tenuis, P. 

umbrosus, and S. dumerili contributed to this similarity. Cat’s Paw and Corkscrew Reefs 

were not well resolved moving among clusters depending on similarity measure 

employed.
54

Figure 1.24. Cluster of reef sites using percent similarity and unweighted pair-group mean 

average combinatorial strategy. [N] designates night dives.
Table 1.9. Summary statistics for ROV data by site group based on cluster analysis. [N] denotes night dives at these sites.

Reef Survey Total Total Mean Mean

Site Group Reef Trend Relief Time (min) No. Fish No. Taxa No. Fish*No. Taxa*

1

Alabama Alps [N] Deep High 97.0 482 28 4.97 0.06

Alabama Alps Deep High 61.2 1697 25 27.75 0.01

Ludwick & Walton Pinnacles1 Deep High 244.8 1088 32 4.44 0.03

Scamp Reef Deep High 23.3 360 16 15.44 0.04

Cat’s Paw Reef Shallow High 27.4 81 7 2.96 0.09

Patch Reef Field Shallow Low 58.6 85 12 1.45 0.14

Roughtongue Reef [N] Shallow High 200.2 966 41 4.83 0.04

Corkscrew Reef Shallow High 40.9 363 24 8.88 0.07

Porgy Reef [N] Shallow High 58.5 130 14 2.22 0.11

2

CMEP5 Shallow High 11.3 580 21 51.42 0.04

Yellowtail Reef Shallow High 171.6 2535 40 14.77 0.02

Double Top Reef Shallow High 106.2 354 16 3.33 0.05

Roughtongue Reef Shallow High 90.9 1516 29 16.69 0.02

Solitary Mounds2Shallow Intermediate 79.6 584 25 7.34 0.04

3

Far Tortuga Shallow Intermediate 23.5 72 11 3.06 0.15

Shark Reef Shallow Low 78.6 103 17 1.31 0.17

4

Yellowtail Reef [N] Shallow High 37.2 79 24 2.12 0.30

5

Ludwick & Walton Pinnacles3 [N] Deep High 146.2 766 25 5.24 0.03

Scamp reef [N] Deep High 69.9 200 20 2.86 0.10

1

includes Pinnacles 1, A, B, and C. 2includes Mounds 1 and 2. 3includes Pinnacles 1 and A. *per minute video

55
56

Figure 1.25. Cluster analysis of taxa using percent similarity and unweighted pair-group 

mean average combinatorial strategy. Clusters were truncated at the group 

level. Taxa are represented by code, first three letters of genera plus four 

letters of species name, or first seven letters for genus and higher taxa.
57

Figure 1.26. Constancy and fidelity for site and taxa groups defined by cluster analysis.

Size of cells is related to number of taxa or sites in each group.

Low <0.1

0.1

0.3

0.5

High 0.7

Low <1

<2

<3

High >3
58

Angling

Various remote sampling gears are effective for different groups of taxa. Whereas 

underwater video documents the fauna much more holistically, some taxa are attracted to 

or repelled by the ROV light. In addition, cryptic and burrowing species are 

underrepresented. We used several additional gears types to compliment the ROV survey 

and enhance our knowledge of the system. Sixty-nine species were taken by hook and line 

(Table 1.4). The gear contributed substantially to our knowledge of community 

composition, as 21 species were only taken by this gear (particularly sharks, jacks, and 

mackerels). The numerically dominant taxa collected by baited hook and line are similar to 

previous surveys (CSA 1992), as R. aurorubens and P. pagrus were the most frequently 

caught large predators in both studies (Table 1.10). The numerical dominance of R. 

aurorubens and P. pagrus also was reported by Grimes et al. (1982) for shelf-edge reefs 

of the Carolinas. Hook and line sampling was important in estimating large predator 

abundance, since many of the larger predators are either attracted to the lights of the ROV 

(carangids, large serranids) or repelled (R. aurorubens, P. pagrus) during visual surveys 

biasing abundance estimates. While R. aurorubens and P. pagrus dominated hook and 

line samples, they were rarely observed during fish surveys, and typically swim away 

from the lights when encountering the ROV. We know from comparison of video survey 

and angling collections that the latter provide an incomplete and highly biased picture of 

faunal composition, particularly when using standard hook and line angling gear with 

large, baited hooks.

Traps

Traps collected 38 fish taxa with six species- Conger oceanicus, Epinephelus nigritus, 

Gymnothorax moringa, Hippocampus sp. (stomach contents), Urophycis floridana, and 

U. regia - unique to the gear (Table 1.4). Of the common taxa, the gear added 

substantially to the collections of Pareques spp., Brotula barbata, and eels such as 

Muraena retifera and Gymnothorax kolpos (Table 1.11). Traps are most effective 

samplers of cryptic and burrowing species and fish that locate prey by chemosensory 

means (e.g., eels).

Trawls

Trawls could only be deployed on low relief bottom and thus sampled near-reef sand flat 

biotope. Fifty-five taxa were collected (Table 1.4) with 27 unique to the gear. The 

faunal separation of the reef and flat bottom communities can be seen in the increased 

importance of ophidiids (cusk eels), triglids (sea robins), and flatfish (Bothidae and 

Paralichthyidae) in the trawl catch (Table 1.12). Saurida spp. and Serranus notospilus

were dominate in the trawl catch and in ROV surveys on surrounding bottom. Few 

gobies were taken by the trawl due to mesh size and their propensity to rapidly flee to 

nearby burrows. Trawls were the only method effective in sampling juvenile D. puellaris 

and S. notospilus.
59

Table 1.10. Abundance of top 15 predatory fishes caught by baited hook and line 

angling from 1997-2000.

Rank Species No. Percent

1 Rhomboplites aurorubens 450 60.4

2 Pagrus pagrus 114 15.3

3 Lutjanus campechanus 94 12.6

4 Seriola dumerili 22 3.0

5 Centropristis ocyurus 9 1.2

6 Caulolatilus chrysops 9 1.2

7 Malacanthus plumieri 7 0.9

8 Opsanus pardus 6 0.8

9 Paranthias furcifer 4 0.5

10 Mycteroperca phenax 4 0.5

11 Seriola rivoliana 3 0.4

12 Mycteroperca microlepis 3 0.4

13 Muraena retifera 3 0.4

14 Epinephelus niveatus 3 0.4

15 Epinephelus flavolimbatus 3 0.4

Total 745 100
60

Table 1.11. Abundance of top 15 fish taxa caught in traps from 1997-2000.

Rank Species No. Percent

1 Rhomboplites aurorubens 233 34.9

2 Pagrus pagrus 126 18.9

3 Pareques umbrosus 71 10.6

4 Centropristis ocyurus 42 6.3

5 Lutjanus campechanus 33 4.9

6 Brotula barbata 31 4.6

7 Seriola dumerili 15 2.2

8 Muraena retifera 15 2.2

9 Gymnothorax saxicola 14 2.1

10 Mycteroperca phenax 13 1.9

11 Micropogonias undulatus 10 1.5

12 Mycteroperca microlepis 7 1.0

13 Gymnothorax kolpos 7 1.0

14 Pareques iwamotoi 5 0.7

15 Holacanthus bermudensis 5 0.7

Total 668 100
61

Table 1.12. Abundance of top 15 fish taxa caught by trawl from 1997.

Rank Species No. Percent

1 Saurida spp. 140 14.0

2 Serranus notospilus 122 12.2

3 Prionotus paralatus 79 7.9

4 Pristipomoides aqualonaris 76 7.6

5 Stenotomus caprinus 72 7.2

6 Syacium papillosum 70 7.0

7 Prionotus stearnsi 47 4.7

8 Serranus atrobranchus 43 4.3

9 Centropristis philadelphica 37 3.7

10 Synodus spp. 36 3.6

11 Citharichthys cornutus 34 3.4

12 Ophidion holbrooki 27 2.7

13 Trichopsetta ventralis 20 2.0

14 Lepophidium brevibarbe 20 2.0

15 Halieuthichthys aculeatus 18 1.8

Total 998 100
62

DISCUSSION

Within the depth range of the Pinnacles reef tract (60-110 m), previous studies in the 

northeastern Gulf have indicated that the fish fauna includes components of the PanCaribbean reef fish fauna, the Carolinian inshore shelf fauna, and the soft-bottom deepwater outer shelf/upper slope fauna (Shipp and Hopkins 1977, Darnell 1991b, Gittings 

et al. 1991, McEachran 1991, Snyder 2001). Faunal composition and habitat affinities 

were comparatively evaluated with previous studies on Gulf of Mexico and western 

Atlantic reef and offshore continental shelf fish species (e.g., Springer and Bullis 1956, 

Bullis and Thompson 1965, Bright and Cashman 1974, Smith et al. 1975, Walls 1975, 

Hastings et al. 1976, Smith 1976, Sonnier et al. 1976, Gilmore 1977, Shipp and Hopkins 

1978, Williams and Shipp 1980, Boland et al. 1983, Clarke 1986, Dennis and Bright 

1988a, b, Hoese and Moore 1998, Thompson et al. 1999, Snyder 2001). At least 250 reefassociated fishes have been documented from offshore banks, reefs, and other hard-bottom 

habitats in the northern Gulf of Mexico based on previous literature (Appendix B). The 

suite of primary reef fishes inhabiting the Pinnacles Reef Tract (Table 1.4) represents a 

depauperate, winnowed fraction of the total reef-fish fauna (Appendix B). The 

selectively winnowed reef-fish fauna of the Pinnacles Reef Tract appears to represent a 

characteristic assemblage of deep-reef fishes broadly distributed in the Pan-Caribbean area 

(Colin, 1974, Colin, 1976). It is important to establish the reasons for this winnowing, 

particularly if the potential effects of anthropogenic activities on the OCS are to be 

rationally assessed and monitored. It can be hypothesized that the known Pinnacles 

reef fish diversity is diminished relative to the Caribbean due to: 1) distance from the 

source fauna, 2) depth limitation, 3) temperature limitation, 4) habitat limitation due to 

turbidity, 5) trophic limitations, 6) fishing pressure, or 7) incomplete sampling. 

However, the first alternative can largely be eliminated as mid-shelf and shelf edge 

banks off Texas and Louisiana are as far or farther from the Caribbean source region, 

but support a large number of reef species (Appendix B). Over 150 reef fish species 

reported to occur in the northwestern Gulf and the Florida Middle Ground have not 

been documented in the Pinnacles Reef Tract (Appendix B). Thus, a large number of 

Caribbean reef-fish species do arrive on reefs in the northern Gulf either by larval 

transport on ocean currents (e.g., Loop Current intrusions), or by active migration. 

Therefore, habitat or other limitations, and not geographic location alone, appear to be 

responsible for the absence (or rarity) of many reef fishes on the Pinnacles.

Many Caribbean reef-fish species do not occur at increasing depths within a given 

geographic area, even in regions of high water clarity and warm temperatures that 

extend beyond 100 m (Colin 1974, 1976, Dennis and Bright 1988a). Depth per se does 

not have a direct effect on fish assemblages but covaries with light, temperature, and 

other physical parameters. The effects of temperature versus depth are difficult to 

distinguish, and highly variable according to geographic location. The northern Gulf of 

Mexico is exposed to winter cold fronts that can drastically lower the temperature of 

inshore waters (Kelly and Bender 2001). The position of the Pinnacles Reef Tract on 

the OCS provides temperature buffering, with deep Gulf waters moderating

temperature fluctuations. Winter bottom temperature averaged 18.1°C with a low of 

15.4°C during the study period (Kelly and Bender, 2001). These temperatures are just 
63

below levels required for reef development and are similar to conditions at mid-shelf 

banks in the northwestern Gulf of Mexico that lack active reef growth (Bright et al. 

1984, Rezak et al. 1985). While reef fish recruits can colonize outside their normal 

geographic range they often do not persist due to limiting environmental conditions. 

However, water temperatures are not low enough at the Pinnacles Reef Tract to be a 

limiting factor to colonization for reef fishes. Chaetodon ocellatus (spotfin 

butterflyfish), a tropical reef species, does not survive winter temperatures below 10°C 

though numerous recruits colonize New Jersey estuaries annually (McBride and Able 

1998). Similarly, reef fishes in the northern Adriatic can tolerate temperatures from 7-

10°C during winter months (Kotrschal and Reynolds 1982). These temperatures are 

well below winter minima recorded at the Pinnacles Reef Tract and the temperature 

range becomes less variable with depth contributing to stable environmental conditions 

(Rezak et al. 1985, Kelly and Bender 2001).

A persistent nepheloid layer characterized by high turbidity was identified as a 

controlling factor for hard bottom communities in the northwestern Gulf (Rezak et al. 

1985). The nepheloid layer increases light attenuation resulting in decreased epibiota 

and reef fish species richness and abundance below 80 m (Dennis and Bright, 1988a, 

Rezak et al. 1990). Previous studies have suggested that the Mississippi River plume 

influences the distribution and abundance of sessile invertebrates within 70 km of the 

river delta, and may produce a gradient of sedimentation and water column turbidity 

throughout the Pinnacles Reef Tract (Gittings et al. 1992a, Hardin et al. 2001, Kelly 

and Bender 2001). In addition, turbidity may affect the feeding efficiency of 

planktivores (Cuker, 1993). In the northeastern Gulf, nepheloid layers are infrequent, 

though in conjunction with episodic Mississippi freshwater plumes and upwelling result 

in increased light attenuation (Kelly and Bender, 2001).

The combination of low winter temperatures and low light levels resulting from 

increased water depth and turbidity on the Pinnacles Reef Tract results in a reduction of 

algal and hard coral growth, and subsequent habitat availability for reef fishes. While 

planktivorous reef fishes dominate Pinnacles reefs, the absence of Chromis multilineata

(brown chromis), Chromis cyanea (blue chromis), and Clepticus parrae (creole 

wrasse), planktivores that form dense schools on reefs in the northwestern Gulf, 

suggests that it is a result of factors associated with depth and not food limitation that 

accounts for the absence of such species on the Pinnacles Reef Tract. Of the 150 

species listed in Appendix B, only 25 are known from below 60 m depths, equivalent to 

the shallowest Pinnacles reefs. Conversely, a number of Pinnacles species, notably the 

dominant anthiine serranids, are absent from the shallow coral cap regions of the 

Flower Garden Banks (18-45 m) and also are not present at the Florida Middle Ground, 

where depths range from 26-48 m (Smith et al. 1975, Clarke 1986, Brooks and Doyle 

1991). Increased water depth and associated light attenuation may induce food 

limitations that account for the absence or rarity of other taxa. For example, 

herbivorous scarids and acanthurids, which form large, free-roaming schools that graze 

algae from the reef substratum on shallow coral reefs throughout the Caribbean, and 

occur at both the Flower Gardens and Florida Middle Ground, are absent from the 

Pinnacles Reef Tract. Similarly, many species of pomacentrids (Stegastes spp.) that 
64

propagate algal gardens on dead coral surfaces are absent from the Pinnacles Reef Tract 

fauna, but are present on shallower banks in the northwestern Gulf where light levels 

are sufficient for growth of filamentous and leafy algae (Dennis and Bright, 1988a). 

Reduced light levels also results in the absence of a well-developed algal-sponge zone, 

a dominant biotope occurring at the Flower Gardens at similar depth range (50-80 m) to 

the shallow Pinnacles reefs (Rezak et al. 1985). Reef fishes that are abundant on the 

algal-sponge zone at the Flower Gardens, including Centropyge argi (cherubfish), 

Sparisoma atomarium (greenblotch parrotfish), and Serranus annularis (orangeback 

bass) are absent on the Pinnacles Reef Tract.

Substrate complexity or rugosity contributes substantially to patterns of reef fish 

distribution and abundance (Syms and Jones 2000). Friedlander and Parrish (1998) 

found that epifauna played little role in determining fish assemblages; substrate 

complexity was a much more important factor. In low relief soft bottom habitats, pits 

and burrows are particularly important in determining fish distribution (Felley et al. 

1989). On deep reefs, the dominant planktivores are visual predators active only during 

daylight hours. They retreat into crevices and shelter holes at night, and also during the 

day upon approach of potential predators. The relative abundance of small planktivores 

and benthivores may depend not only on food availability, but also on relative 

abundance of shelter holes (= substrate rugosity). While many studies have found 

correlations between hole size/number and structural complexity with reef fish 

abundance or habitat selection (Friedlander and Parrish 1998, Syms and Jones 2000, 

Snyder 2001), when empirically tested no limitation on shelter holes was found for a 

shallow water reef fish (Robertson and Sheldon 1979).

Most Pinnacle Reef Tract features are high relief drowned reefs giving rise to a range of 

biotopes (Gardner et al. 2001). Geological features associated with diaparism, 

including gas and brine seeps that greatly influence the biotic communities in the 

northwestern Gulf are rare in the Pinnacles Reef Tract (Gittings et al. 1992a, Gardner et 

al. 2001). Gittings et al. (1992a) and Hardin et al. (2001) have observed different 

assemblages of sessile macro-invertebrates in the Pinnacles Reef Tract related to 

structural form (e.g., horizontal summits versus vertical or rugged features). Study sites 

that have extensive flat-top areas including Roughtongue Reef, Yellowtail Reef, and 

Alabama Alps, develop a high diversity invertebrate assemblage that is dominated by 

octocorals, antipatharians, sponges, bryozoans, coralline algae (on shallower reef sites) 

and crinoids, and is typically characterized by relatively high percent cover of 

invertebrates (Hardin et al. 2001) (Fig. 1.18). Vertical reef faces, overhangs, and rock 

outcrops and boulders occurring at the reef base are dominated by the solitary coral R. 

manuelensis (Fig. 1.17). Results of our ROV analyses indicate strong fish faunal 

differentiation based on physically defined biotope type (Table 1.5; Figs. 1.21-1.23, 

Appendix A). Fine-scale factors such as habitat complexity (Friedlander and Parrish 

1998) may control absolute fish abundance within a particular biotope type, but relative 

abundance appears more a function of basic biotope type. 

Fishes observed on the reef top included the highest number of taxa across all biotopes, 

and this species richness is likely due to the diversity of microhabitats available, 
65

including rock outcrops, sand pockets, and dense assemblages of invertebrates such as 

sponges, octocorals (including large sea fans), and antipatharians that form spatially 

complex structures. Also the reef top represents the greatest reef area; thus the speciesarea relationship may be a factor (MacArthur and Wilson, 1963). The diversity of 

habitat present on the reef top also presents a wide variety of prey resources that are 

associated with unconsolidated sediments and a diverse invertebrate assemblage. This 

biotope not only supports planktivores (including C. enchrysura), but also benthic 

carnivores (H. bathyphilus and D. puellaris), epibenthic browsers (C. rostrata, C. 

sedentarius and H. bermudensis), and generalized carnivores (S. phoebe and L. 

eukrines) (see Chapter 2 for more details on trophic categories). 

The reef crest and face are important foraging habitat not only for planktivores, but also 

for epibenthic carnivores, and mobile piscivores that feed on both planktivores and 

benthivores. The rugose, highly sculptured substrate provides shelter for small fishes. 

Stony corals largely replace soft corals, crinoids, antipatharians, and sponges on vertical 

reef faces, limiting the structural complexity of the reef surface. The reef base, composed 

of caves and undercuts, is often occupied by species such as G. hispanus, P. umbrosus, 

C. spinosus, O. trachypoma, and L. eukrines. Extending out from the reef base is the 

circum-reef talus apron. This biotope is characterized by small to medium-size benthic

carnivores (e.g., labrids, serranids, P. umbrosus) that forage for benthic and epibenthic 

prey, and not plankton. Once on the open sand flat habitat of the shelf proper, fish 

density is low and fish are primarily active at night and aggregated around any structure 

(Felley et al. 1989). The distinct fish assemblages within each biotope and the different 

areal extent of biotopes among reefs needs further study to provide a more detailed 

picture of factors controlling deep reef community structure. We are extending our 

future research in this area.

Reef profile and topographic complexity remain somewhat enigmatic in their respective 

roles in controlling community structure of reef fishes. Beyond a critical threshold, the 

overall height of a pinnacle structure may be fairly unimportant. What is critical is that 

the structure be of sufficient height to reduce exposure to turbidity and sedimentation, 

and perturb flow on the reef face and crest, and over the reef top (Gittings et al. 1992a, 

MacDonald and Peccini 2001). Messing et al. (1990) found that lithoherm banks in the 

northeastern Straits of Florida hydrodynamically accelerated ambient 2-7 cm/s bottom 

currents to greater than 100 cm/s over the upcurrent reef crest. These authors 

schematically illustrate both the compression of current streamlines over a lithoherm 

resulting in current acceleration (Messing et al. 1990, Fig. 3), and the resultant 

upcurrent to downcurrent biozonation of suspension-feeding sessile macroinvertebrates 

(Messing et al. 1990, Fig. 4). Fairly continuous currents from the southwest sweep the 

steep-sided Pinnacle reefs as reflected in the orientation of soft corals (MacDonald and 

Peccini 2001). Flow models indicate that acceleration of currents over shelf-edge 

pinnacles may increase availability of pelagic prey in the vicinity of high profile 

features (Kelly and Bender 2001). The "wall of mouths" concept developed on the Great 

Barrier Reef (Hamner et al. 1988) suggests an upcurrent to downcurrent pattern of faunal 

differentiation due to plankton removal from the impinging current and the importance of 

plankton availability in controlling fish faunal assemblages among biotopes. The 
66

characterization of biotopes and their associated fish communities on a finer scale that 

couples water currents and biological/geological characteristics of the benthic habitat may 

ultimately be necessary to explain the occurrence and composition of the reef fish 

assemblage. We are presently following this line of inquiry.

Although hermatypic corals with zooxanthellae and leafy algae are a major carbon 

source for fishes on shallow reefs (Ogden 1982), recent research has revealed that 

plankton also may be of great importance in certain regions (Hamner et al. 1988). A 

characteristic set of deep-water planktivores, together with a suite of geographically 

ubiquitous, deep-water browsers and benthic carnivores, dominate the Pan-Caribbean 

deep-water fish fauna (Colin, 1974, Colin, 1976; Dennis and Bright, 1988a, this study). 

Indeed, a model of trophic partitioning of the reef-fish fauna in deep water, based on the 

Pinnacles Reef Tract example, may have broad application throughout deep reef 

communities of the world’s oceans (see Chapter 2 for details). Trophic partitioning of the 

Pinnacles fauna closely parallels the earlier findings of Thresher and Colin (1986) for the 

deep (>75-90 m) reef community of Enewetak Atoll in the southwestern Pacific. Below 

90 m, the Enewetak fish fauna was similarly dominated by planktivorous anthiine 

serranids (>90% of all individuals), together with planktivorous labrids and pomacentrids 

(Chromis spp.). All other trophic guilds were very poorly represented in deep water, 

except for piscivores (1-10% by number) that feed on the abundant planktivores. Patterns 

are similar in the Pinnacles Reef Tract except for the low abundance of piscivores.

Both the dominant sessile macro-invertebrates (e.g., gorgonians and antipatharians) and 

reef fishes on Pinnacles Reef Tract reefs are planktivores, and spatial and temporal 

access to plankton appear to be major factors in the control of community structure and 

organism distribution (Hardin et al. 2001, MacDonald and Peccini 2001). Consistency 

of supply and volume of the planktonic food source may be a strong controlling factor 

for these fish assemblages. Intercepting the incoming current, the upcurrent reef face 

and crest should be the most favorable biotopes for swarms of diurnal planktivores 

(predominantly anthiine serranids). We hypothesize that this is true based on ROV 

observations, but a quantitative assessment is needed.

Direct anthropogenic effects, such as fishing, also may have altered the reef fish 

community in the Pinnacles Reef Tract. Commercial fisheries for Lutjanus 

campechanus began in the 1870’s, first based in Pensacola, Florida, and expanding to 

numerous fleets based throughout the Gulf by the early 1930’s (Jarvis 1935), targeting 

snappers and groupers on offshore banks. Darnell (1991b) reports declines in biomass 

of bottom fishes on the shelf and decreased harvest rates of L. campechanus, and 

attributes these patterns to overfishing and habitat loss. Snyder (2001) points out 

declining catch rates of large groupers (Epinephelus spp. and Mycteroperca spp.), L. 

campechanus, and S. dumerili throughout the northeastern Gulf since the early 1970’s, 

and a rise in catch rates of R. aurorubens during this same time period. These patterns 

have been linked to the widespread use of bottom longlines in outer shelf waters of the 

Gulf (Richards and McGowan 1989) that target large demersal fishes, and may explain 

the low abundance of these species in our collections and ROV surveys. The apparent 

decrease in the abundance of large piscivores in the Pinnacles Reef Tract due to fishing 
67

activities may have had the further consequence of increasing the absolute and relative 

abundance of their primary prey, the anthiine serranid planktivores or other species of 

lower trophic levels, such as R. aurorubens, and lead to alteration of ecosystem 

structure and function (Parsons 1992, 1996, Pauly et al. 1998). Thus, caution should be 

exercised in generalizing patterns in community structure for ecosystems that have been 

subject to anthropogenic alteration.

Fishes and invertebrates have been characterized at the Pinnacles Reef Tract by three 

major studies and all major reef features have been precisely mapped. This area may 

be the most well-known deep hard bottom area in the world. Thus we postulate that the 

majority of the reef fish species that occur at depths between 60 and 110 m are now 

documented. Yet due to the large area and diversity of habitat further scientific inquiry 

will continue to add to our knowledge of the fish fauna. Our present knowledge of the 

Pinnacles ichthyofauna can be used to characterize the fundamental taxonomic 

composition and trophic structure of deep reef-fish communities throughout southeastern 

United States and Gulf of Mexico OCS areas and therefore be useful in assessing future 

anthropogenic impacts on faunal structure.
68
69

CHAPTER 2

TROPHIC ECOLOGY OF PINNACLES TRACT REEF FISHES

INTRODUCTION

Defining trophic pathways is the key to developing an understanding of the ecological 

framework in the reef fish community. Analysis of food web structure also identifies 

relative importance of resident reef fishes (autochthonous) versus off-reef trophic 

resources (allochthonous) such as adjacent soft-substrate and pelagic prey in the diet of 

large predatory fishes, predator-prey relationships among fishes and invertebrates, and 

may also identify potential pathways of energy transfer from fishes to the benthic 

invertebrate assemblage through the release of fecal material. Trophic guild categories 

are general terms to describe the location of their main source of prey and identify 

pelagic versus benthic trophic pathways. Within each feeding guild most members will 

utilize a wide spectrum of prey based on availability/encounter rates, prey size, and 

prey mobility. Detailed studies of the food habits of deep reef fishes have not been 

reported in the literature, and few reports of shallow reef fish diets in the Gulf of 

Mexico are available (e.g., Bullock and Smith 1991, Nelson and Bortone 1996, Weaver 

1996). Investigations on food habits of reef fishes in the South Atlantic Bight primarily 

have focused on commercially important species (Manooch 1977, Grimes 1979, Ross 

1982, Matheson et al. 1986, Dodrill et al. 1993, Lindquist and Clavijo 1993, Lindquist 

et al. 1994, Sedberry and Cuellar 1993). The reef fish, invertebrate, and plant 

assemblages that characterize live bottom communities in the Gulf of Mexico vary 

dramatically with depth (Rezak et al. 1985, 1990), limiting comparisons among shallow 

and deep reef communities from published literature. Therefore, identifying food 

habits of deep reef fishes and developing food web models from the literature would 

yield an inaccurate, simplistic view. A full understanding requires sampling of fishes 

from deepwater reef communities to document dietary patterns. Diets of marine fishes 

are known to vary dramatically with body size, local microhabitat, and season (Crabtree 

et al. 1991, Mullaney and Gale 1995, Sheridan and Trimm 1983, Starck 1971), and 

therefore collection of fishes provides the only method to develop a detailed description 

of trophic pathways in the deep reef environment. Trophodynamic investigations 

undertaken in the present study include feeding habits and food web structure of 

Pinnacles Reef Tract fishes.
70

OBJECTIVES

· Delineate food habits of numerically dominant fish species. 

· Construct a trophic model based on quantitatively-defined food webs that model the 

Pinnacles fish community structure and function, and convey predictions about 

primary sources of forage base and energy for reef and reef-associated biotopes.

METHODS

There is an extensive body of literature regarding methods for analysis of fish stomach 

contents and food habits, and conventional quantitative parameters of broad application 

have been reviewed (Hyslop 1980). Specimens sampled for food habits from USGS 97-

01 through 2000-01 cruises were kept on ice, frozen, or preserved in formalin until 

processed in the laboratory at the Florida Caribbean Science Center, Gainesville, FL 

(FCSC). Only specimens from hook and line sampling were analyzed and specimens 

with empty or everted stomachs were excluded. Trap specimens were not included in 

the analysis. Fishes were dissected in the laboratory and the whole digestive tracts 

(stomachs plus intestines) were removed. Prey items were identified to lowest possible 

taxonomic level. For simplicity, dietary analyses in this report are given as percent 

number, calculated by dividing the number of prey within each taxon by the total 

number of prey items for each of the characteristic reef fishes collected excluding 

empty stomachs. Stomach contents were used to identify the ecological role of each taxon 

in the reef fish community and used to identify feeding guilds following previous authors 

(Hobson 1982, Bohnsack et al. 1999).

Food Web Model

A food web model was developed for the Pinnacles fish community. A quantitative 

model was based on numerical contribution of prey to the diet of each predator. 

Primary goals of food web models are: 1) to compare relative contribution of 

allochthonous materials (i.e., plankton and adjacent soft-bottom off-reef organisms) to 

the energy budget of the dominant members of the reef fish community; and 2) to 

determine the relative importance of reef fishes in the diet of large reef-associated 

carnivores. Diets of the characteristic reef fishes were incorporated into a working food 

web model following the methods of Winemiller (1990) to identify the dominant 

trophic pathways among the reef fish community. Trophic position of each predator 

was calculated using the formula:
71

where Ti is the trophic level of an individual predator species i, summing relative 

importance of prey taxon j where pij is the proportion of the total diet (based on 

numerical abundance) for species iconsisting of prey taxon j. Primary producers are 

assigned a value of T=0.0, herbivores T=1.0, and a consumer eating half plant and half 

herbivore tissue would be T=1.5, and so forth, as Ti’s are computed sequentially from 

bottom to top. Calculated Ti values were used as the y axis for placement of consumer 

nodes of the food web. The x coordinate of each consumer was weighted according to 

the proportion of prey from different habitat compartments. Proportions of pelagic prey 

(plankton) were multiplied by 0.0 (far left), reef-based prey by 0.5 (middle), and soft 

bottom prey by 1.0 (far right), to determine relative dependence on each habitat for 

prey (Winemiller 1990). This method gives estimates of relative dependence on 

allochthonous versus autochthonous materials (i.e., a position close to the left portion of 

the web is dependent on pelagic prey, the middle portion resident reef prey, and the 

right side off-reef soft bottom prey). Links connecting each compartment were then 

calculated with respect to movement of energy from prey to predator (uplinks) 

(Winemiller 1990). Links among the different nodes (predator-prey relation) can be 

expressed with uniform width and relative strength (qualitative models) or line width 

varied to reflect proportional contribution of prey in the diet (quantitative models). 

Terminology of plankton size classes and constituent members follow Christensen 

(1995).

RESULTS AND DISCUSSION 

Over 1000 individuals were examined for food habits with approximately 50% 

containing prey items. Many specimens everted their stomachs due to the change in 

pressure when brought to the surface and the majority of the large predators (jacks, 

snappers, and groupers) had empty stomachs. In addition, many resident reef fishes 

were poorly sampled by hook and line angling, reflected by low sample sizes. Feeding 

guilds were defined by characteristic prey (Table 2.1). Dietary patterns of the 

characteristic reef fish taxa are presented in this report (Table 2.2, Appendix C). Fish 

were assigned to feeding guilds based on stomach contents to identify similarity in prey 

resources among species, sources of prey, and overall patterns in trophic structure 

(Table 2.3). Use of the conventional terms, such as macroinvertivores and 

microinvertivores, by other authors to describe feeding guilds is avoided due to the 

wide size range of prey selected by most species (e.g., Serranus phoebe feeds on adult 

crabs and fishes as well as decapod and fish larvae).

The numerically dominant species (greater than 80% of the individuals observed by 

ROV) on high profile reefs, Pronotogrammus martinicensis and Hemanthias vivanus, 

have diets dominated by calanoid copepods, gastropod larvae, and a variety of other 

pelagic mesoplankton organisms (Table 2.2, Appendix C). Macroplanktivores that 

prey on larger size classes of plankton include Rhomboplites aurorubens, Pristigenys 

alta, Apogon pseudomaculatus, and Chaetodon aya. Benthic feeders included Pagrus 

pagrus, Pareques umbrosus, Decodon puellaris, and Halichoeres bathyphilus. 
72

Table 2.1. Feeding guilds defined by characteristic prey type.

Feeding Guild

 

Mesoplanktivore- Mixed zooplankton in the microzooplankton to mesoplankton (20-2000μ) 

size classes. Includes copepods, ostracods, larvaceans, gastropod veligers, pteropods, 

crustacean larvae, invertebrate eggs, and fish eggs.

Macroplanktivore- Mixed zooplankton in the macroplankton (>2000μ) size class. 

Includes hyperiid amphipods, shrimp zoea, stomatopod zoea, crab megalopa, fish 

larvae, salps, and squids.

Benthic Carnivores (~microinvertivores)-primarily infauna/epifaunal invertebrates including 

crabs, bivalves, gastropods, brittle stars, sea urchins, and polychaetes.

Epibenthic Browsers- encrusting, sessile invertebrates including hydrozoans, bryozoans, soft 

corals, crinoids, and sponges.

Generalized Carnivores (~macroinvertivores)- a variety of mobile, epibenthic prey that may 

include fishes, crabs, shrimps, and gastropods.

Piscivores- includes fishes and squids.
73

Table 2.2. Dietary analysis of characteristic reef fishes of the Pinnacles Reef Tract by 

trophic guild. Values given are based on numerical contribution to the diet 

(%N).

Mesoplanktivores Macroplanktivores Benthic Carnivores

Sample Size 2 20 47 1 120 5 13 15 9 72 4 2 2 10 28 1 4

Phylum Prey Item Percent Number

Invertebrate eggs - 19.1 3.0 - 11.0 - - - - - - - - - - -

Protoctista Foraminiferans - - - - - - - - - - - - - 6.9 - - -

Radiolarians - - - - 1.5 - - - - - - - - - - - -

Porifera Sponges - - - - - - - - - - - - - - 1.1 - -

Cnidaria Hydrozoans - - - - - - - - - - - - - - - - -

Soft Corals - - - - - - 4.8 - - - - - - - - - -

Mollusca Heteropods - - - - - - - - - - - - - - - - -

Gastropods - - - - - - - - - - 14.3 - 31.3 - 16.0 33.3 -

Pelecypods - - - - - - - - - - 38.1 20.0 37.5 48.3 3.2 - -

Pteropods - - - - - - - 1.3 - 6.9 - - - - - - -

Chitons - - - - - - - - - - - - - - - - -

Squids - - - - - - - 32.9 16.0 0.2 - - - 3.4 - - -

Annelida Polychaetes - 13.3 17.7 35.7 10.3 - - - - 14.3 - 45.0 6.3 3.4 4.3 - 9.1

Arthropoda Copepods 60.0 32.4 78.8 28.6 69.7 - - - - 15.2 - - - - - - -

Amphipods 20.0 - - - 0.2 - 5.1 - - 15.2 - - - - - - -

Barnacle larvae - - - - 0.1 - - - - 1.4 - - - - - - -

Crab larvae - 1.1 - 14.3 0.4 12.5 90.1 14.5 16.0 3.7 - - 12.5 13.8 2.1 - -

Crabs - - - - 25.0 - 1.3 2.0 - 19.0 5.0 6.3 6.9 34.0 33.3 36.4

Isopods - - - - - 37.5 - - - - - - - - - - -

Ostracods - - 0.1 - 1.7 - - - - 2.1 - - - - - - -

Shrimp larvae 20.0 2.1 - 21.4 - 25.0 - 1.3 16.0 10.1 - - - 6.9 - - 9.1

Shrimps - - - - - - - 3.9 40.0 - - - - - 2.1 - 36.4

Stomatopod larvae - - - - - - - 27.6 6.0 8.0 - - 6.3 - 6.4 - -

Stomatopods - - - - - - - - - - - 25 - - 4.3 - -

Bryozoa Bryozoans - - - - - - - - - - - - - - - - -

Echindermata Echinoids - - - - - - - - - - - - - - 20.2 - -

Ophiuroids - - - - - - - - - - 9.5 5.0 - 6.9 - 33.3 9.1

Holothuroids - - - - - - - - - - - - - - 1.1 - -

Crinoids - - - - - - - - - - - - - - - - -

Chordata Larvaceans - 32.0 0.1 - 1.9 - - - - 1.1 - - - - - - -

Salps - - - - 0.5 - - - - 9.9 - - - - - - -

Fish eggs - - - - 2.4 - - - - 1.4 - - - - - - -

Fish larvae - - - - - - - 14.5 - 10.6 - - - - - - -

Fishes - - - - - - - 2.6 2.0 - 19.0 - - 3.4 5.3 - -
74

Table 2.2. (continued).

 Epibenthic browsers Generalized Carnivores Piscivores Holacanthus bermudensis Chaetodon sedentarius Canthigaster rostratus Centropristis ocyurus Epinephelus flavolimbatus Holocentrus adscensionis Liopropoma eukrines Lutjanus campechanus Malacanthus plumieri Opsanus pardus Pontinus rathbuni Sargocentron bullisi Serranus phoebe Aulostomus maculatus Gymnothorax saxicola Seriola dumerili

Sample Size 4 8 2 6 2 3 1 26 7 5 1 2 44 1 1 6

Phylum Prey Item Percent Number

Invertebrate eggs - - - - - - - - - - - - - - - -

Protoctista Foraminiferans - - - - - - - - - - - - - - - -

Radiolarians - - - - - - - - - - - - - - - -

Porifera Sponges 8.3 - - - - - - - - - - - - - - -

Cnidaria Hydrozoans - 71.4 - - - - - - - - - - - - - -

Soft Corals 41.7 14.4 - - - - - - - - - - - - - -

Mollusca Heteropods - - - - - - - 5.1 - - - - - - - -

Gastropods - - 3.2 11.8 - - - - 25.0 33.3 - - - - - -

Pelecypods - - 7.8 - - - - - 18.8 - - - - - - -

Pteropods - - - - - - - - - - - - - - - -

Chitons - - 2.1 - - - - - - - - 16.7 - - - -

Squids - - - - - - - 2.6 - - - - 1.4 - - -

Annelida Polychaetes - - - - - - - - - - 16.7 1.4 - - -

Arthropoda Copepods 5.2 - - - - - - - - - - - - - -

Amphipods - - - - - - - - - - - - 1.4 - - -

Barnacle larvae - - - - - - - - - - - - - - - -

Crab larvae - - - - - 42.9 - - - - - 33.3 5.6 - - -

Crabs - - - 76.5 - - - 5.1 18.8 25.0 - 41.7 - - -

Isopods - - - - - - - 10.3 - - - - - - - -

Ostracods - 2.2 - - - - - - - - - - - - - -

Shrimp larvae - 2.1 - - - - - - - - - - - - - -

Shrimps - - - 5.9 33.3 14.3 - 7.7 - - 60.0 - 5.6 - - -

Stomatopod larvae - - - - - - - - - - - 33.3 - - - -

Stomatopods - - - - - - - 2.6 - - - - 1.4 - - -

Bryozoa Bryozoans 41.7 - - - - - - - 6.3 - - - - - - -

Echinoids - 1.5 - - - - - - 8.3 - - - - - -

Ophiuroids - - - 5.9 - - - - 12.5 - - - 1.4 - - -

Holothuroids - - - - - - - - - - - - - - - -

Crinoids 8.3 - 78.6 - - - - - - 8.3 - - - - - -

Chordata Larvaceans - - - - - - - - - - - - - - - -

Salps - - - - - - - 10.3 - - - - - - - -

Fish eggs - - - - - - - - - - - - - - - -

Fish larvae - - - - - - - - - - - - 2.8 - - -

Fishes - - - - 66.7 42.9 100 56.4 18.8 25.0 40.0 - 37.5 100 100 100
75

Table 2.3. Feeding guilds of fishes along the Pinnacles Reef Tract. Determination of 

feeding guild is modified from Hobson (1982) and Bohnsack et al. (1997, 

1999) and based on stomach contents identified in this study, or from food 

habits by Randall (1967) for closely related taxa.

Feeding Guild

Mesoplanktivores- Anthias tenuis (threadnose bass), Chromis enchrysura (yellowtail reeffish), 

Hemanthias vivanus (red barbier), Paranthias furcifer (creole-fish), Pronotogrammus 

martinicensis (roughtongue bass), Trachurus lathami (rough scad).

Macroplanktivores- Apogon pseudomaculatus (twospot cardinalfish), Chaetodon aya (bank 

butterflyfish), Priacanthus arenatus (bigeye), Pristigenys alta (short bigeye),

Rhomboplites aurorubens (vermilion snapper).

Benthic Carnivores- Balistes capriscus (gray triggerfish), Calamus leucosteus (whitebone 

porgy), Calamus nodosus (knobbed porgy), Caulolatilus chrysops (goldface tilefish),

Decodon puellaris (red hogfish), Halichoeres bathyphilus (greenband wrasse), Pagrus 

pagrus (red porgy), Pareques iwamotoi (blackbar drum), and P. umbrosus (cubbyu).

Epibenthic Browsers –Canthigaster rostrata (sharpnose puffer), Chaetodon sedentarius (reef 

butterflyfish), Chaetodon ocellatus (spotfin butterflyfish), Holacanthus bermudensis 

blue angelfish), Parablennius marmoreus (seaweed blenny).

Generalized Carnivores- Bodianus pulchellus (spotfin hogfish), Centropristis ocyurus (bank 

sea bass), Corniger spinosus (spinycheek soldierfish), Epinephelus flavolimbatus 

(yellowedge grouper), Epinephelus nigritus (Warsaw grouper), Epinephelus niveatus 

(snowy grouper), Holocentrus adscensionis (squirrelfish), Liopropoma eukrines 

(wrasse bass), Lutjanus campechanus (red snapper),Muraenidae, Opsanus pardus 

(leopard toadfish), Malacanthus plumieri (sand tilefish), Sargocentron bullisi

(deepwater squirrelfish), Pontinus rathbuni (highfin scorpionfish), Rypticus maculatus 

(whitespotted soapfish), Scorpaena dispar (hunchback scorpionfish), Serranus phoebe 

(tattler).

Piscivores- Aulostomus maculatus (trumpetfish), Fistularia petimba (red cornetfish), 

Muraenidae, Mycteroperca microlepis (gag), M. phenax (scamp), Seriola dumerili 

(greater amberjack), Seriola fasciata (lesser amberjack), Seriola rivoliana (almaco 

jack).
76

Epibenthic browsers included Chaetodon sedentarius and Holacanthus bermudensis

whose diets were composed of sessile invertebrates. Piscivores were represented by 

Mycteroperca spp., Seriola spp., Lutjanus campechanus, muraenids, Aulostomus 

maculatus, and Fistularia petimba. The generalized carnivore guild was represented by 

numerous taxa including S. phoebe, and a variety of serranids, scorpaenids, muraenids, 

and holocentrids. As discussed by Snyder (2001), this guild contains the most taxa of 

deep reef fishes, although individual species are not as numerically abundant as 

planktivores and benthic carnivores in reef communities.

Planktivorous anthiine fishes may serve as a keystone predator by foraging in the water 

column and transferring energy to the deep reef ecosystem. Anthiines in turn become 

prey for a variety of reef predators. Stomach contents indicate that newly settled 

juveniles and young-of-the-year anthiines are an important seasonal component of the 

diet for a variety of reef fishes, including L. campechanus, carangids, and serranids. 

Trophic pathways among the deep reef community indicate primary links to the pelagic 

community through direct consumption of planktonic organisms and may be 

supplemented by surrounding benthic communities through consumption of eggs and 

larvae of soft-bottom fishes, a variety of invertebrates, and potentially by spawning 

populations of migratory species from nearshore reef communities (e.g., Mycteroperca 

microlepis). In contrast, Darnell (1991a) concluded that zooplankton contributed only 

2% of the overall foods for the soft-bottom fish community of the Mississippi-Alabama 

shelf, and that small epibenthic organisms, infauna, and larger demersal organisms 

constituted the majority of the food resources for the soft bottom assemblage.

Initial analysis of food web structure (Figs. 2.1, 2.2) for the deep reef community further 

identifies potential pathways of energy flow and trophic links between hard- and softbottom habitats that may be useful in identifying changes in trophic structure associated 

with anthropogenic activities (e.g., the oil/gas exploration and fishing) in areas that are of 

special concern to resource managers. The current reef fish community exhibits trophic 

pathways that are primarily supported by planktonic prey, as planktivorous reef fishes 

dominated all reef biotopes observed during ROV surveys and R. aurorubens, a 

macroplanktivore, was the numerically dominant predator taken by angling and traps.

While generalized carnivores are the most speciose feeding guild on the Pinnacles Reef 

Tract, they were relatively rare in ROV surveys and collections made during this study. 

Planktivores (primarily anthiines and the pomacentrid C. enchrysura) numerically 

dominated all reef biotopes during ROV surveys with benthic carnivores (Gobiidae, H. 

bathyphilus, D. puellaris, and P. umbrosus) second in importance. Generalized 

carnivores were represented by S. phoebe and L. eukrines, but individual taxa made up

2% or less by number on reef biotopes. Food web models based on relative abundance of 

fishes indicate that the primary pathway of energy to the deep reef community is by 

direct consumption of copepods, amphipods, fish larvae, decapod larvae, and gelatinous 

zooplankton that prey on smaller planktonic prey groups in the water column (Figs. 2.1, 

2.2). Secondary pathways that support benthic carnivores are presumably based on 

detrital pathways that support infaunal and epifaunal invertebrates. The numerically 

dominant predators, R. aurorubens and P. pagrus, feed directly on planktonic and soft-
Figure 2.1. Qualitative food web model for reef fishes of the Pinnacles Reef Tract constructed from stomach content analysis. 

Position on the food web model is based on relative proportion of pelagic prey (x axis) and trophic position (y axis). 

77
Figure 2.2. Quantitative food web model for reef fishes of the Pinnacles Reef Tract constructed from stomach content analysis. 

Position on the food web model is based on relative proportion of pelagic prey (x axis) and trophic position (y axis). 

Relative sizes of nodes (circles) depict abundance of reef fishes based on ROV data analysis and hook and line 

collections and relative widths of links depicted are based on numerical contribution of prey in the diet of each predator.

78
79

bottom organisms, respectively, and do not have diets characterized by reef-associated

fishes or invertebrates. Seriola spp., L. campechanus, and S. phoebe prey upon anthiine 

serranids, and therefore also appear to be supported primarily by pelagic trophic 

pathways.

While few groupers examined during this study retained identifiable stomach contents, 

anthiine serranids presumably form a significant part of their diet, as A. tenuis, H. 

vivanus, and P. martinicensis have been reported from the stomach contents of deepwater 

groupers (Bullock and Smith 1991). Predation on anthiines would provide a direct link to 

pelagic food webs.

Trophic pathways within reef systems have been the subject of study since the 1950's 

(Odum and Odum 1955, Hiatt and Strasburg 1960), yet trophic interactions and 

mechanisms of energy transfer within the reef fish community are still poorly known 

(Bohnsack and Sutherland 1985, Nelson and Bortone 1996). In their original work on the 

trophic structure of coral reefs, Odum and Odum (1955) concluded that the coral reef 

community at Enewetak Atoll was primarily supported by benthic and zooxanthellate 

algae, and suggested that incoming plankton was not a significant component of organic 

matter to the community. Tropical oceans that bathe most coral reefs are oligotrophic, 

therefore most shallow reef systems might be self-supporting, although more recent 

studies have identified the important trophic links to zooplankton in tropical coral reef 

communities (Hamner et al. 1988). Benthic algae typically supports a large population of 

grazing reef fishes. These herbivores are dominant members of many shallow coral reef 

systems (Hiatt and Strasburg 1960, Ogden 1982, Ebeling and Hixon 1991), yet are absent 

from the Pinnacles Reef Tract. Porter and Porter (1977) suggested that coral reef 

communities support a self-generated food web, and tightly recycle nutrients. Pinnacles 

deep reef communities are characterized by an abundance of planktivorous fishes and 

suspension-feeding invertebrates (Hardin et al. 2001), utilizing allochthonous prey 

resources in the water column to sustain populations, and are therefore very different in 

trophic structure when compared to many shallow coral reef communities.

The relative importance of plankton in food web structure of reef communities has 

received limited attention, particularly for more eutrophic warm-temperate reef 

ecosystems in the Atlantic and Gulf of Mexico. Advection of plankton from open ocean 

water masses to the continental shelf has been noted (Hopkins et al. 1981). Bottom 

currents also bring plankton from adjacent shelf waters to the shelf break (Kelly and 

Bender 2001). Copepods and gelatinous zooplankton are the common grazers of 

phytoplankton and nanoplankton (<5 μ) in the open water column and may be important 

in transfer of open water primary production to other components of marine ecosystems 

such as the benthic fish assemblage (Fourtier et al. 1994, Kleppel et al. 1996). Along a 

shallow windward reef face in Australia, transport of plankton from the open ocean was 

considered the major input of organic material to the fish and invertebrate community 

(Hamner et al. 1988). On temperate reefs in California, over 67% of fishes are obligate 

or facultative planktivores (Ebeling and Hixon 1991). Planktivorous fishes also dominate 

fish communities on New Zealand rocky reefs (Kingsford and MacDiarmid 1988, 

Kingsford 1989). Feeding on plankton is an important trophic pathway in many reef fish 
80

communities, particularly those exposed to strong ocean currents. Planktivory is 

considered an important route of energy flow to the reef community by two pathways: 1) 

direct consumption of zooplankton by planktivorous fishes and sessile 

macroinvertebrates, and 2) consumption of planktivores by larger piscivores or 

deposition of fecal material to the reef (Hobson 1991). On Indo-Pacific coral reefs, fecal 

material provides an alternative source of nutrients to reef inhabitants as many fishes 

ingest fecal material of Chromis atropectoralis, a planktivore that releases feces of high 

energy content (Bailey and Robertson 1982). Hamner et al. (1988) also noted that fecal 

pellets of planktivores were observed on the reef surface more frequently than 

zooplankton. Chromis punctipinnis, a common member of the reef fish community in 

California, has been shown to deposit substantial quantities of fecal material to rocky 

reefs each night (Bray et al. 1981). Many invertebrates (e.g., crabs, shrimps, and brittle 

stars) rapidly ingest deposited fecal material deposited by planktivorous fishes (Rothans 

and Miller 1991) and in turn these mobile invertebrates are preyed upon by fishes. 

Sedberry and Cuellar (1993) discussed the potential role in energy transfer by R. 

aurorubens through release of feces over the reef surface. This pathway may be 

important in trophic connections between planktivorous reef fishes and benthic 

invertebrate assemblages in deep reef communities throughout the western Atlantic. 

Reef-dwelling planktivores that dominate the Pinnacles reef community may supplement 

the diet of many motile reef invertebrates that are the dominant prey of other resident 

fishes.

Hard-bottom communities are generally characterized by much higher fish biomass than 

surrounding, unconsolidated sediments (Sedberry and Van Dolah 1984, Parker and Ross 

1986). However, studies of productivity on shallow (<38 m) hard-bottom reef systems 

along the southeastern U. S. reveal that these reef communities are heterotrophic and not 

nearly as productive as their tropical coral reef counterparts. Typical productivity/ 

respiration ratios for the southeastern U.S. live bottom reefs average 0.68 versus 1.45 on 

Indo-Pacific coral reefs (Longhurst and Pauly 1987, Hopkinson et al. 1991). Shallow 

hard bottom communities require secondary production via pelagic zooplankton or 

benthic primary production in adjacent sandy habitats as a food source to support reef 

organisms (Hopkinson et al. 1991). Deep reefs in the Gulf of Mexico should exhibit even 

lower primary productivity/respiration ratios than shallow water communities. 

Therefore, dense populations of Pinnacles fishes appear to exceed levels that could be 

supported solely by locally produced prey resources. Fishes on deep reefs must therefore 

be “subsidized” by passive transport of prey from surrounding benthic or pelagic habitats 

on water currents, or via active feeding migrations of predators to adjacent areas of the 

water column or the underwater landscape (Polis et al. 1997). On shallow coral reef 

ecosystems, grunts of the genus Haemulon migrate away from the reef to forage on 

benthic invertebrates in adjacent sea grass beds and sand flat areas at night, and form 

inactive schools over coral reefs by day (Meyer et al. 1983, Parrish 1989). Temperate 

reef fishes also forage in off-reef habitats and consume invertebrates in neighboring softbottom habitats (Posey and Ambrose 1994, Weaver 1996). In both tropical and 

temperate reef systems, planktivorous reef fishes aggregate at discrete areas and rely on 

water currents to transport prey resources from the surrounding water column (Bray 

1981, Hamner et al. 1988). Passive transport of planktonic prey to resident fishes, or 
81

active migration away from reefs to forage on benthic or pelagic prey resources 

represents allochthonous trophic subsidies. Spatial subsidies, where primary and 

secondary productivity of adjacent habitats provides prey resources that are transported to 

the reef community enhance populations of resident predators, result in increased 

predator density at levels well above those sustainable from local resources (Polis et al. 

1997). Local enhancement of fish populations at the Pinnacles occurs through two major 

pathways: 1) consumption of plankton by numerically dominant anthiines and R. 

aurorubens as they actively forage into the water column or 2) foraging in adjacent softbottom sediments by benthic carnivores such as P. pagrus, S. phoebe, and labrids.

While small diurnal planktivores, such as P. martinicensis and H. vivanus, are considered 

to be an important trophic link between open water plankton and the reef community, 

resident nocturnal planktivores may play a relatively minor role in the overall energetics 

of the reef community. Although nocturnal planktivores are widely distributed on reefs, 

most resident species occur in low abundance (i.e., P. alta), and are not considered to be 

important prey for piscivorous fishes due to their deep bodies and well-developed fin 

spines (Hobson 1991). Hobson (1991) also noted that reef-dwelling nocturnal 

planktivores were likely to be less important in transferring energy between open-water 

communities and the reef ecosystem, since they forage close to the reef on emergent prey 

at night. Many of the emergent plankters such as amphipods and crab zoea collected at 

night were common prey in the diet of nocturnal planktivores (Alldredge and King 1977, 

Porter and Porter 1977). Amphipods, isopods, and a variety of other small crustaceans 

make up substantial populations of cryptofauna that live within the reef substrata during 

the day (Klumpp et al. 1988).

However, on the deep reef tract of the Pinnacles, other nocturnal planktivores appear to 

play an important role in reef trophodynamics. Small midwater species such as B. 

cantori, T. lathami, and myctophids are periodically abundant on high profile reefs, and 

would provide an additional prey resource for nocturnal carnivores. While rarely 

observed in ROV surveys, R. aurorubens appears to be the numerically dominant 

predator and forages by day and night on plankton (Grimes 1979). Other species that 

feed on the reef surface at night include Leiostomus xanthurus, a species normally 

associated with soft bottom habitats. Foraging activities of these species may transport 

considerable energy between hard bottom, soft bottom and pelagic habitats during 

feeding migrations.

Large predators, including S. dumerili and L. campechanus, also utilize prey from 

pelagic, reef, and soft-bottom habitats and provide pathways of energy transfer. The 

presence of B. cantori, T. lathami and urochordates in the diet of L. campechanus

indicates direct links to the pelagic food web, while the presence of soft bottom fishes, 

including lizardfishes and sea robins in the diet of S. dumerili indicates foraging activities 

in soft bottom habitats. Large predators, including groupers, will often migrate between 

reef structures and opportunistically consume soft-bottom and pelagic prey (Weaver 

1996) and provide conduits of energy flow between reef and off-reef communities.
82

Potential impacts of the petroleum industry in outer shelf regions may differentially affect 

pelagic versus benthic feeders if drilling wastes and other discharges influence nearbottom turbidity, sedimentation, and prey availability. Identification of trophic structure 

may also assist in evaluation of overfishing of large predators and the associated shift in 

the reef fish community. Shelf-edge reef sites are the primary spawning locations for 

many commercially important groupers and snappers, and have been heavily fished in 

recent years (Richards and McGowan 1989, Gilmore and Jones 1992, Coleman et al. 

1996, Koenig et al. 2000). Koenig et al. (2000) reported a decrease in large predator 

abundance (primarily Mycteroperca phenax and S. dumerili), and attributed this change 

to increased fishing activity in this region. Based on ROV surveys, P. martinicensis and 

H. vivanus comprised 6% (total planktivores made up 6.4%) of the reef fish community 

in 1980, and 71% (total planktivores ~84%) of the reef fish community in 1997 surveys 

on the east coast of Florida (Koenig et al. 2000), a pattern that parallels the current reef 

fish community at the Pinnacles. This increase in lower trophic levels in the deep reef 

fish community also parallels a reported decline in the mean trophic level of commercial 

fisheries worldwide (Pauly et al. 1998). The numerical dominance by small 

planktivorous reef fish taxa in shelf-edge reef communities and R. aurorubens in our 

hook and line samples in the Pinnacles Reef Tract have likely been magnified by removal 

of piscivores through recent increases in commercial fishing activity in this region and 

historical fishing practices throughout the northwestern Gulf of Mexico since the mid 

1800’s. Snyder (2001) discusses the decline in catch rates of the large predatory species 

that characterize offshore reef ecosystems since the 1970’s, and an increase in catch rates 

of R. aurorubens during this same time period. While our observations of trophic 

structure in the deep reef community closely resemble those observed in the Indo-Pacific 

(Thresher and Colin 1986), the influence on fishing activity will be difficult to assess in 

determining a natural community, and the current patterns of reef fish community 

structure may have been greatly altered by human activity through fishing during the past 

century.
83

ACKNOWLEDGMENTS

Outer Continental Shelf Ecosystem Program (1997-1999)

(K. Sulak, PI)

We thank the following individuals for invaluable collaboration and assistance in the 

field during research cruises: G. Yeargin, D. DeVries, T. Glancy, P. Schofield, S. 

Meister, B. Eleby, S. Ross, M. Randall, P. Stevens, S. Szedlmayer, R. McBride, J. 

Caruso, C. Harper, L. Parker, J. Barichivich, G. Hill, G. Fitzhugh, C. Gurshin, J. 

Lancaster, C. Furman and M. Zatcoff. We also thank Captain Paul Beaugez and the 

Crew of the R/V TOMMY MUNRO, and Captain Jerry Love and the Crew of the R/V 

SUNCOASTER for their logistical support in research cruises. ROV support was 

provided by L. Horn and T. Potts, National Undersea Research Center (NURC) 

University of North Carolina-Wilmington (UNCW), and C. Harper, National 

Oceanographic and Atmospheric Administration/National Marine Fisheries Service 

(NOAA/NMFS)-Stennis Space Center and NMFS Mississippi laboratories. Additional 

logistical support was provided by the NURC-UNCW Research Center, NOAA-NMFS, 

the Marine Resources Research Institute of the South Carolina Department of Natural 

Resources, the Florida Marine Research Institute of the Florida Fish and Wildlife 

Conservation Commission, the University of Florida (UF), the University of South 

Florida, the University of North Carolina at Wilmington, Auburn University, and the 

Louisiana Universities Marine Consortium (LUMCON). Maps, data, and general 

information on the Pinnacles region was provided by the Continental Shelf Associates 

(CSA), I. MacDonald, K. Spring, D. Snyder, and W. Schroeder. Detailed bathymetry 

maps for this report were provided by J. Gardner and P. Dartnell, USGS-Menlo Park. 

Research on the Pinnacles Reef Tract was funded by the Outer Continental Shelf 

Ecosystem Program (K. Sulak, PI) of the U.S. Geological Survey, in Cooperation with 

the Minerals Management Service (MMS), Gulf of Mexico Region. High-resolution 

bathymetric mapping was supported by USGS Interactive BRD/GD Science Program (K. 

Sulak, J. Gardner, and C. Sherwood, Co-PI’s) with supplementary support from USGS 

Geologic Division, NMFS, and MMS. Additional funding and logistical support for 

doctoral research were provided to D. Weaver by the Walt Disney Conservation Fund, 

the National Fish and Wildlife Foundation, and the Department of Fisheries and Aquatic 

Sciences-UF during cruises 2000-01 and 2000-2. G. Brewer provided advice during all 

phases of this project, and reviewed the earlier drafts of this report. T. Ahlfeld, S. Childs, 

and G. Boland of the Minerals Management Service provided review and comments of 

the draft report.
84
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APPENDIX A

Summary of ROV surveys from 1997-2001

Legend:

A=Alabama Alps

C=Cat's Paw Reef

D=Double Top Reef

CS=Corkscrew Reef

C5=CMEP Monitoring Site 5

F=Far Tortuga Reef

L=Ludwick&Walton Pinnacle

P=Porgy Reef

PF=Patch Reef Field

R=Roughtongue Reef

S=Shark Reef

SC=Scamp Reef

SM=Solitary Mound

T=Triple Top Reef

Y=Yellowtail Reef
A-2

Station No. 04 08 09 11 12 20 22 27 33 35 36 44 45 50

Base Depth (m) 68 68 78 78 78 78 106 104 110 110 110 77 71 72

Study Site Y Y R R R C L-A L-B L-C L-1 L-1 PF SM-1 SM-2

Taxa

Antigonia capros - - - - - - - - - 0.05 - - - -

Apogonidae - 0.09 - - - - - - - - - - - -

Apogon pseudomaculatus - - - - - - - - - - - - - -

Aulostomus maculatus - - - - - - - - - - - - - -

Bodianus pulchellus - - - - - - - - - - - - - -

Bregmaceros cantori - - - - - - - - - - - - - -

Canthigaster rostrata 0.14 - 0.08 0.10 - - - - - - - - 0.04 -

Carangidae - - - - - - - - - 0.05 - - - -

Centropristis ocyurus - - - - - - - - - - - - - -

Chaetodon aya - 0.09 - 0.36 0.09 - 0.03 - 0.06 0.05 0.18 0.03 - -

Chaetodon ocellatus - - - - - - - - - - - - - 0.03

Chaetodon sedentarius - 0.18 - 0.05 0.09 0.04 - - - - - - - 0.06

Chilomycterus schoepfii - - - - - - - - - - - - - -

Chromis enchrysura 4.58 - 0.08 0.57 0.19 - - - - - - - 0.09 0.51

Corniger spinosus - 0.18 - - - 0.04 0.03 - 0.01 0.19 - 0.03 0.02 -

Cyclopsetta fimbriata - - - - - - - - - - - - - -

Decodon puellaris 0.14 - 0.31 - 0.09 - 0.11 - 0.01 0.09 - 0.03 0.35 -

Diodon holocanthus - - - - - - - - - - - - - -

Epinephelus flavolimbatus - - - - - - - - - 0.05 - - - -

Epinephelus niveatus - - - - - - - - - - - - - -

Fistularia petimba - - - - - - - - - - - - - -

Gobiidae 0.56 0.18 2.33 1.20 - 0.04 - - 0.01 - - 0.09 1.56 -

Gonioplectrus hispanus - 0.09 - - - - - - 0.02 - 0.18 - - -

Gymnothorax kolpos - - - 0.05 - - - - - - - - - -

Halichoeres bathyphilus 0.28 0.35 0.93 1.30 0.09 - - - - - 0.04 0.02 - -

Hemanthias leptus - - - - - - 0.03 - 0.04 - 0.04 - - -

Hemanthias vivanus 28.75 - 10.11 25.08 31.50 - - - 1.30 0.97 0.41 7.78 -

Holacanthus bermudensis - - - - - - - - - - - - 0.02 -

Holacanthus tricolor - - - - - - - - - - - - - -

Holocentrus adsensionus - - - 0.05 - - - - - - - - - -

Lactophrys quadricornis - - - - - - - - - - - - 0.02 -

Leiostomus xanthurus - - - - - - - - - - -

Liopropoma eukrines - 0.09 - 0.26 0.19 - 0.06 - - - 0.04 0.02 0.06 -

Lutjanus campechanus - - - - 0.19 - - 0.04 - - - 0.09 0.03

Monacanthus spp. - - - - - - - - - - - - - -

Muraenidae - - - - - - - - - - 0.04 - - -

Mycteroperca microlepis - - - - - - - - 0.12 - - - 0.02 -

Mycteroperca phenax - - - - 0.37 - - - - 0.05 0.04 - 0.06 -

Myctophidae - - - - - - - - - - - - - -

Ogcocephalus spp. - - 0.08 - - - 0.03 - - - - - - 0.03

Opsanus pardus - - - - - - - - - - - - - 0.03

Ostichthys trachypoma - - - - - - 0.31 - - - 0.04 - - -

Pagrus pagrus - - - - - - - - 0.02 - - - - -

Parablennius marmoreus - - 0.08 0.05 - - - - - - - - - -

Paranthias furcifer - - - - 1.12 - - - - - - - - -

Pareques iwamotoi - - - - - - 0.06 - 0.07 - 0.04 - - -

Pareques umbrosus - 0.62 - - - - - - 0.02 - - 0.02 - 0.03

Peprilus burti - - - - - - - - - - - - - -

Pontinus spp. - - - - - - - - - - - - - -

Priacanthus arenatus - - - - - 0.07 0.08 - - - - - - -

Pristigenys alta 0.28 0.27 0.08 0.10 0.09 0.04 - - 0.05 0.51 0.09 0.02 0.11 0.45

Pronotogrammus martinicensis 0.42 16.93 0.86 3.02 2.52 2.66 7.18 - 3.33 1.72 6.52 0.73 0.84 0.03

Rhomboplites aurorubens - - - - - - - - - - - - 0.02 0.12

Rypticus maculatus - - - - - - - - - - - - - -

Sargocentron bullisi - - - 0.10 0.09 - - - - - - - - -

Scorpaena dispar - - - 0.05 - 0.07 0.14 - 0.05 - - 0.02 - 0.06

Seriola dumerili 0.14 0.09 - - - - - - 0.04 - - - 0.04 -

Seriola fasciata - - - - 0.09 - - - - - - - - -

Seriola rivoliana - - - - - - - - - - - - - -

Serranus notospilus - - - - - - 0.11 - - - - - - -

Serranus phoebe 0.14 0.09 - - 0.19 - 0.03 - - - - 0.03 0.04 0.03

Sphoeroides spengleri - - - - - - - - - - - - - -

Synodus sp. - - - - - - - - - - - - - -

Trachurus lathami - - - - - - - - - - - - - 0.57

Trichiuridae - - - - - - - - - - - -

Unidentified flatfish - - - - - - 0.03 - - - - - - -

Upeneus parvus - - - - - - - - - - - - - 0.03

Total Fish per Minute 35.42 19.24 14.93 32.36 36.90 2.95 8.21 - 3.92 4.10 8.24 1.45 11.17 2.01

No. Minutes 7.2 11.3 12.9 19.2 10.7 27.3 35.9 ND 84.4 21.5 22.7 58.6 46.3 33.3

Appendix A1. Summary of reef-associated fishes observed during 1997 (97-01) ROV

surveys by habitat. Data are expressed as number of fish observed per 

minute of video for each site. ND-No Data and (n)-night survey.
A-3

53 54 56 63 64 65 69 72 76 81 83 86 87 90 Station No.

76 78 78 77 77 78 78 78 78 70 68 68 68 68 Base Depth (m)

T D D S S R R(n) CS P(n) F Y(n) Y Y Y Study Site

Taxa

- - - - - - - - - - - - - - Antigonia capros

- - - - - - - - - - - - - - Apogonidae

- - - - - - 0.04 0.02 0.07 - 0.08 - - - Apogon pseudomaculatus

- - - - - - 0.01 - - - - - - - Aulostomus maculatus

- - - - - 0.02 - - - - - 0.03 0.02 - Bodianus pulchellus

- - - - - - 0.09 - 0.32 - - - - - Bregmaceros cantori

- - - - - 0.19 0.06 0.02 0.03 - 0.05 - 0.21 0.07 Canthigaster rostrata

- - - - - - - - - - - - - - Carangidae

- 0.09 - - - - - - - 0.09 - - - - Centropristis ocyurus

- 0.06 0.02 0.02 - 0.15 0.06 0.29 0.03 0.17 0.11 0.17 0.19 0.28 Chaetodon aya

- - - - - - 0.01 - - - - - - - Chaetodon ocellatus

- - - 0.02 0.10 0.02 0.06 0.02 0.03 - 0.05 0.14 0.02 - Chaetodon sedentarius

- - - - - - - - - - - - - 0.07 Chilomycterus schoepfii

- 0.03 - 0.03 - 0.35 0.08 0.05 - - 0.21 0.28 0.11 0.21 Chromis enchrysura

- - - 0.02 - 0.06 0.14 0.15 - - 0.05 0.10 0.13 0.07 Corniger spinosus

- - - - - - - - - - 0.03 - - - Cyclopsetta fimbriata

- 0.14 0.02 0.17 0.10 0.10 - 0.10 0.02 0.04 0.03 0.17 0.06 0.28 Decodon puellaris

- - - - - - - - - - 0.03 - - - Diodon holocanthus

- - - - - - - - - - - - - - Epinephelus flavolimbatus

- - - - - - - 0.02 - - - - - - Epinephelus niveatus

- - - - - - 0.01 - - - - - - - Fistularia petimba

- 0.43 0.02 0.17 0.97 - - 0.10 0.05 0.89 0.05 0.38 0.49 0.07 Gobiidae 

- - - - - - - 0.02 - - - - - - Gonioplectrus hispanus 

- - - - - - - - 0.05 - - - - - Gymnothorax kolpos

- 0.02 - - - 0.37 - 0.27 - 0.26 0.03 0.10 0.19 0.28 Halichoeres bathyphilus

- - - - - - - - - - - - - - Hemanthias leptus

- 1.89 1.43 0.19 - 8.24 0.09 1.34 - - 0.08 1.59 4.79 8.78 Hemanthias vivanus

- - - - - 0.08 - 0.07 - - - - - - Holacanthus bermudensis

- - - - - 0.02 0.01 - - - - - - - Holacanthus tricolor

- - - - - - 0.02 - - - - - - - Holocentrus adsensionus

- - - - - - - - - - - - - - Lactophrys quadricornis

0.10 - - - - - - - Leiostomus xanthurus

- - 0.10 - - 0.02 - 0.15 - 0.13 - 0.03 0.15 0.50 Liopropoma eukrines

- - - - - - - - - - 0.05 - - - Lutjanus campechanus

- - - - - - - - - - 0.03 - - - Monacanthus spp.

- - - - - - - 0.02 - - - - - - Muraenidae

- - - - - - - - - - - - - - Mycteroperca microlepis

- - - - - - - 0.12 - - - - 0.15 0.07 Mycteroperca phenax

- - - - - - 0.01 - - - - - - - Myctophidae

- - - - - - - - - - 0.08 - - - Ogcocephalus spp.

- - - - - - 0.01 - 0.02 0.09 0.05 - 0.02 0.07 Opsanus pardus

- - - - - - - - - - - - - - Ostichthys trachypoma

- - - - - - - - - - - - - - Pagrus pagrus

- - - - - 0.04 - - - - - - 0.04 - Parablennius marmoreus

- - - - - - 0.01 - - - - - 0.08 0.14 Paranthias furcifer

- - - - - - - - - - - - - - Pareques iwamotoi

- 0.14 - 0.03 - 0.02 0.02 0.90 - - 0.03 - - - Pareques umbrosus

- - - - - - 0.01 - - - - - - - Peprilus burti

- - 0.02 - 0.05 - 0.01 - 0.03 - - 0.03 - - Pontinus spp.

- - - - - 0.06 0.01 0.02 - - - - - - Priacanthus arenatus

- 0.08 0.02 0.03 - 0.08 0.28 0.20 - 0.17 0.21 0.24 0.04 - Pristigenys alta

- 0.68 0.82 0.32 0.20 2.56 1.29 4.74 1.20 0.72 0.05 3.42 3.84 6.79 Pronotogrammus martinicensis

- 0.03 0.17 0.02 0.15 - 0.05 - 0.17 - - - - - Rhomboplites aurorubens

- - - 0.03 - - - - - - - - - Rypticus maculatus

- - - - - 0.06 - - - - 0.13 - 0.02 - Sargocentron bullisi

- - 0.10 0.05 - 0.02 0.14 0.02 - - 0.21 - - - Scorpaena dispar

- - - - - - - - - - - - - - Seriola dumerili

- - - - - - - - - - - - - - Seriola fasciata

- - - - - - - 0.05 - - - - - - Seriola rivoliana

- - - 0.05 - - - 0.07 - - - - - - Serranus notospilus

- 0.03 0.10 - 0.15 0.10 0.04 0.10 - 0.47 0.08 - 0.04 0.14 Serranus phoebe

- - - 0.02 - - - - - - - - - 0.07 Sphoeroides spengleri

- 0.03 - - - 0.02 0.02 - - 0.04 - - - - Synodus sp.

- - - - - - 0.10 - 0.12 - 0.08 - - - Trachurus lathami

- - - - - - - - 0.07 - 0.29 - - - Trichiuridae

- - - - - - - - - - - - - - Unidentified flatfish

- - - - - - - - - - - - - - Upeneus parvus 

- 3.68 2.86 1.17 1.79 12.61 2.82 8.89 2.22 3.06 2.12 6.71 10.61 17.91 Total Fish per Minute

ND 65.0 41.3 59.1 19.6 48.1 123.2 41.0 58.5 23.5 37.2 28.9 97.2 14.1 No. Minutes

Appendix A1. (continued).
A-4

Station No. 04 08 09 11 12 20 22 27 33 35 36 44 45 50

Base Depth (m) 68 68 78 78 78 78 106 104 110 110 110 77 71 72

Study Site Y Y R R R C L-A L-B L-C L-1 L-1 PF SM-1 SM-2

Taxa

Bregmaceros cantori - - - - - - - - - - - - - -

Centropristis ocyurus - 0.21 - - - - - - - - - - - -

Citharichthys cornutus - - - - - - - - - - - - - -

Decodon puellaris - - - - - - - - 0.03 - - 0.29 1.00 -

Gobiidae - 0.63 - - - - 0.08 0.04 0.31 0.56 - 0.18 0.25 -

Halichoeres bathyphilus - - - - - - - - - - - - - -

Halieutichthys aculeatus - - - - - - - - - - - - - -

Hemanthias vivanus - - - - - - - - - - - - - -

Hoplunnis sp. - - - - - - - 0.08 - - - - - -

Lagodon rhomboides - - - - - - - - - - - - - -

Leiostomus xanthurus - - - - - - - - - - - - -

Lepophidium brevibarbe - - - - - - - 0.01 - - - - - -

Ogcocephalus spp. - - - - - - - 0.01 0.03 - - - - -

Ophichthidae - - - - - - - 0.01 - - - - - -

Opistognathidae - - - - - - - - - - - - - -

Pagrus pagrus - - - - - - - - - - - - - -

Paralichthys squamilentus - - - - - - - - 0.03 - - - - -

Parequetus umbrosus - - - - - - - - - - - - - -

Pontinus sp. - - - - - - - - - - - - - -

Prionotus stearnsi - - - - - - - 0.01 - - - - - -

Pristipomoides aquilonaris - - - - - - - - - - - - - -

Raja sp. - - - - - - - - - - - - - -

Rhynchoconger sp. - - - - - - - 0.01 - - - - - -

Saurida sp. - - - - - - - - - - - - - -

Scorpaena dispar - - - - - - - - - - - - - -

Serranus atrobranchus - - - - - - - - - - - - - -

Serranus notospilus - 1.26 - - - - 0.17 0.33 0.23 0.07 - 0.11 1.24 -

Serranus phoebe - - - - - - 0.17 0.02 - - - 0.04 - -

Sphoeroides sp. - 0.21 - - - - - - - - - - - -

Syacium papillosum - - - - - - - 0.01 0.05 - - - - -

Symphurus sp. - - - - - - - 0.01 - - - - - -

Synodus sp. - - - - - - - 0.02 0.03 0.07 - - - -

Trachurus lathami - - - - - - - 0.04 - - - - - -

Trichiuridae - - - - - - - 0.01 - - - - - -

Unidentified flatfish - - - - - - - 0.01 0.03 - - - - -

Upeneus parvus - - - - - - - 0.04 - - - - - -

Total Fish per Minute - 2.09 - - - - 0.42 0.73 0.75 0.70 - 0.73 2.49 -

No. Minutes ND 4.8 ND ND ND ND 12.0 84.7 38.5 14.3 ND 27.5 4.0 ND

Station No. 04 08 09 11 12 20 22 27 33 35 36 44 45 50

Base Depth (m) 68 68 78 78 78 78 106 104 110 110 110 77 71 72

Study Site Y Y R R R C L-A L-B L-C L-1 L-1 PF SM-1 SM-2

Taxa

Ancylopsetta dilecta - - - - - - - - - - - - - -

Chromis enchrysura - - - - - - - - - - - - - -

Decodon puellaris - 2.20 - - - - - - - - - - - -

Gobiidae - 8.81 - - - - - - 0.68 - - - - -

Halichoeres bathyphilus - 0.31 - - - - - - - - - - - -

Liopropoma eukrines - 0.16 - - - - - - - - - - - -

Opistognathidae - - - - - - - - - - - - - -

Pontinus sp. - - - - - - - - 0.14 - - - - -

Priacanthus arenatus - 0.16 - - - - - - - - - - - -

Pristigenys alta - 0.16 - - - - - - - - - - - -

Pronotogrammus martinicensis - - - - - - - - - - - - - -

Scorpaena dispar - - - - - - - - - - - - - -

Serranus notospilus - 0.16 - - - - - - 0.27 - - - - -

Serranus phoebe - 0.16 - - - - - - - - - - - -

Synodus sp. - - - - - - - - - - - - - -

Total Fish per Minute - 12.74 - - - - - - 1.35 - - - - -

No. Minutes ND 6.4 ND ND ND ND ND ND 7.4 ND ND ND ND ND

Appendix A2. Summary of fishes associated with the talus zone observed during 1997 

(97-01) ROV surveys. Data are as number of fish observed per minute 

of video. ND-No Data and (n)-night survey.

Appendix A3. Summary of fishes associated with the sand flats observed during 1997 

(97-01) ROV surveys. Data are as number of fish observed per minute 

of video. ND-No Data and (n)-night survey.
A-5

53 54 56 63 64 65 69 72 76 81 83 86 87 90 Station No.

76 78 78 77 77 78 78 78 78 70 68 68 68 68 Base Depth (m)

T D D S S R R(n) CS P(n) F Y(n) Y Y Y Study Site

Taxa

- - - - - - - - - 0.05 - - - - Ancylopsetta dilecta

- - - - - - - - - - - 0.25 - - Chromis enchrysura

- - - - - - - 0.10 - 0.05 - 0.25 0.21 0.86 Decodon puellaris 

- - - - - - - 3.12 - 1.03 - 4.43 0.21 - Gobiidae

- - - - - - - - - - - - 0.64 - Halichoeres bathyphilus

- - - - - - - - - - - - - - Liopropoma eukrines

- - - - - - - 0.10 - 0.05 - - - - Opistognathidae

- - - - - - - - - - - - - - Pontinus sp.

- - - - - - - - - - - - - - Priacanthus arenatus

- - - - - - - - - - - - - - Pristigenys alta

- - - - - - - - - - - 0.12 - - Pronotogrammus martinicensis

- - - - - - - - - 0.05 - 0.12 - - Scorpaena dispar

- - - - - - - 0.31 - - - - - - Serranus notospilus

- - - - - - - - - 0.05 - 0.74 - - Serranus phoebe

- - - - - - - 0.21 - 0.09 - 0.37 - - Synodus sp. 

- - - - - - - 3.84 - 1.30 - 6.28 1.06 0.86 Total Fish per Minute

ND ND ND ND ND ND ND 9.6 ND 21.6 ND 8.1 4.7 1.2 No. Minutes

53 54 56 63 64 65 69 72 76 81 83 86 87 90 Station No.

76 78 78 77 77 78 78 78 78 70 68 68 68 68 Base Depth (m)

T D D S S R R(n) CS P(n) F Y(n) Y Y Y Study Site

Taxa

0.07 - - - - - - 0.33 - - - - - Bregmaceros cantori

- - - 0.08 - - - - - - - - - - Centropristis ocyurus

- - - - - - - - - - - 0.18 - - Citharichthys cornutus

- - - - - - - 0.19 - - - - - - Decodon puellaris

0.16 - 0.18 1.48 - - - 0.55 0.83 - - 1.06 - - Gobiidae 

- - - - - - - 0.08 - - - 0.18 - - Halichoeres bathyphilus

0.03 - - - - - - - - - - - - - Halieutichthys aculeatus

- - - - - - - 0.04 - - - - - - Hemanthias vivanus 

- - - - - - - - - - - - - - Hoplunnis sp.

- - - 0.08 - - - - - - - - - - Lagodon rhomboides

- - - 0.08 - - - - 0.08 - - - - - Leiostomus xanthurus

- - - - - - - - - - - - - - Lepophidium brevibarbe

- - - - - - - - - - - - - - Ogcocephalus spp.

- - - - - - - - - - - - - - Ophichthidae

- - - - - - - 0.04 - - - - - - Opistognathidae

- - 0.08 - - - - - Pagrus pagrus

- - - 0.08 - - - - - - - - - - Paralichthys squamilentus

- - - - - - - - 0.08 - - - - - Parequetus umbrosus

- - - - - - - - 0.17 - - - - - Pontinus sp.

- - - - - - - - - - - - - - Prionotus stearnsi

0.07 - - - - - - 0.16 - - - - - - Pristipomoides aquilonaris

- - - - - - - 0.04 - - - - - - Raja sp.

- - - - - - - - 0.08 - - 1.94 - - Rhynchoconger sp.

2.03 - - - - - - - - - - 0.35 - - Saurida sp.

- - - 0.08 - - - 0.08 - - - 0.18 - - Scorpaena dispar

0.03 - - - - - - - - - - 0.18 - - Serranus atrobranchus

0.03 1.43 0.18 - - - - 0.51 0.42 - - 1.06 - - Serranus notospilus

- - - - - - - 0.04 - - - - - - Serranus phoebe

- - - - - - - - - - - - - - Sphoeroides sp.

0.13 - - - - - - - - - - - - - Syacium papillosum

- - - - - - - - - - - - - - Symphurus sp.

0.07 1.43 0.73 - - - - 0.04 0.50 - - 5.11 - - Synodus sp.

- - 0.18 - - - - - - - - - - - Trachurus lathami

- - - - - - - - 0.08 - - 0.18 - - Trichiuridae

- - 0.18 - - - - - - - - 0.35 - - Unidentified flatfish

- - - - - - - - - - - - - - Upeneus parvus

2.63 2.86 1.45 1.88 - - - 1.75 2.67 - - 10.74 - - Total Fish per Minute

30.5 0.7 5.5 12.8 ND ND ND 25.7 11.8 ND ND 5.7 ND ND No. Minutes

Appendix A2. (continued).

Appendix A3. (continued).
Station No. 0001-028 0001-029 0001-036 0001-055 0001-057 0001-060 9903-2132 0001-016 0001-018 0001-019 0001-033 0001-052 0001-059

Base Depth (m) 69 69 88 110 105 79 109 80 110 110 88 110 105

Study Site Y Y A L-H SC C5 L-1(n) R (n) L-2 (n) L-2 (n) A (n) L-A (n) SC (n)

Taxa

Anthias tenuis - 0.02 - - - - - - - - 0.11 - -

Apogon pseudomaculatus - 0.22 0.05 - - - - 0.04 - - 0.05 - 0.01

Aulostomus maculatus - 0.04 0.05 - - - - - - - - - -

Bodianus pulchellus - 0.11 - - - - - - - - - - -

Bregmaceros cantori - - - - - - - - - 2.22 0.03 - -

Brotula barbata - - - - 0.04 0.09 - - - - 0.01 - -

Canthigaster jamestyleri 0.11 - - - - - - 0.01 - - - - -

Canthigaster rostrata 0.32 0.22 0.15 - - - - 0.32 - - 0.01 - -

Carapidae - - - - - - - - - 0.02 - - -

Centropristis ocyurus - - - - - 0.44 - - - - - - -

Chaetodon aya 0.43 0.20 0.70 0.04 0.13 0.44 - 0.26 - 0.03 0.05 0.18 0.03

Chaetodon sedentarius 0.11 0.04 0.10 - - - - 0.10 - - - - -

Chilomycterus antillarum - - - - - - - - - 0.02 - - -

Chromis enchrysura 1.51 1.40 0.05 - - 0.27 - 0.23 - - 0.02 - -

Chromis insolata - 0.04 - - - - - - - - - - -

Chromis scotti - 0.47 - - - 0.27 - 0.01 - - - - -

Corniger spinosus - 0.11 0.15 - 0.04 0.27 - 0.03 - - 0.07 - -

Coryphopterus punctipectophorus 0.22 0.15 - - - - - 0.03 - - - - -

Decodon puellaris - 0.02 - 0.05 - - - 0.04 - - - - 0.01

Epinephelus niveatus - - - - - 0.09 - - - - - - -

Fistularia petimba - - - - - - - - - - - - 0.01

Gonioplectrus hispanus - - - - 0.04 0.09 - - - - - - -

Gymnothorax hubbsi 0.11 - - - - - - - - - - - -

Gymnothorax kolpos - - - - - - - - - - - 0.18 -

Halichoeres bathyphilus 0.11 0.20 0.05 0.10 - - - 0.04 - - 0.01 - -

Hemanthias leptus - - - - 0.26 - - - - - - - -

Hemanthias vivanus 36.97 6.74 0.10 - 2.57 25.71 - 2.25 - - 0.04 0.18 -

Holacanthus bermudensis 0.11 0.04 - 0.01 0.09 0.09 - - - - 0.01 - -

Holocentrus adscensionus 0.11 - - - - - - - - - - - -

Lactophrys quadricornis - - - - - 0.09 - - - - - - -

Liopropoma eukrines - 0.13 0.05 0.09 0.09 0.18 - 0.05 - - 0.02 0.18 -

Lutjanus campechanus - - 0.05 - - - - - - - 0.01 - -

Malacanthus plumieri - - - - - - - 0.01 - - - - -

Muraenidae - 0.02 - - - - - - - - 0.01 - -

Appendix A4. Summary of Cruise USGS 99-03 and 2000-01(0001) ROV surveys. Values given are number of individuals observed

per minute of tape time. Night surveys are indicated by (n).

A-6
Station No. 0001-028 0001-029 0001-036 0001-055 0001-057 0001-060 9903-2132 0001-016 0001-018 0001-019 0001-033 0001-052 0001-059

Base Depth (m) 69 69 88 110 105 79 109 80 110 110 88 110 105

Study Site Y Y A L-H SC C5 L-1(n) R (n) L-2 (n) L-2 (n) A (n) L-A (n) SC (n)

Taxa

Mustelus sp. - - - - - - 0.04 - - 0.02 0.01 - 0.01

Mycteroperca microlepis - - - - 0.04 - - - - - - - -

Mycteroperca phenax - 0.02 - 0.01 0.13 - - - - 0.02 0.01 0.18 0.03

Myctophidae - - - - - - 0.24 - 0.65 0.10 - 27.37 2.15

Neoepinnula americana - - - - - - - - 0.03 - - -

Ogcocephalus sp. - - - - - 0.09 - 0.01 - - 0.01 - 0.01

Opistognathus lonchurus - 0.02 - - - - - - - - - -

Opsanus pardus - - - - - - - 0.01 - - - - -

Ostichthys trachypoma - - - - 0.09 - - - 0.13 0.03 - 1.46 -

Pagrus pagrus - - - 0.11 0.09 - 0.04 0.01 - - - - 0.01

Paranthias furcifer - 0.37 - - - - - - - - - - -

Pareques iwamotoi - - - - - 0.09 - - - - 0.02 0.36 0.10

Pareques umbrosus - 0.02 0.15 0.04 0.21 0.44 - - - - 0.20 0.55 0.04

Pomatomus saltatrix - - - - - - - - - - - - 0.01

Priacanthus arenatus - - - - - 0.27 - 0.01 - - - - -

Pristigenys alta 0.32 0.02 0.20 0.32 0.09 0.53 0.05 0.14 0.13 0.32 0.10 0.91 0.07

Pronotogrammus martinicensis 0.97 2.79 14.01 1.48 11.41 20.83 0.02 4.13 - 0.22 3.74 3.10 0.03

Rhomboplites aurorubens - - - - - - - - 0.04 0.02 - - -

Rypticus maculatus - - - - - - - - - - - - -

Sargocentron bullisi 0.11 0.07 - 0.01 - - - 0.08 - - - - -

Scomberomorus cavalla - - - - - - - - 0.04 - - - -

Scorpaena dispar - 0.02 - 0.01 - - - 0.06 - - 0.04 - -

Scorpaenidae - - - 0.06 0.13 0.18 0.02 0.03 0.04 0.11 0.01 - 0.07

Seriola dumerili - - 0.25 - - 0.35 - 0.14 - 0.10 0.22 0.55 0.01

Seriola rivoliana 0.22 0.02 - - - - - - - - 0.02 - -

Serranus phoebe 0.22 0.04 0.05 0.04 - 0.62 - 0.04 - - - - -

Sphoeroides spengleri - - - - - - - 0.01 - - - - -

Stenotomus caprinus - - - - - - - - - - - - 0.01

Synodus sp. - - - - - - - - - 0.11 0.02 - 0.13

Trachurus lathami - - - - - - - 0.03 0.52 0.87 0.09 0.73 -

Trichiurus lepturus - - - - - - 4.31 - - - - 0.18 0.01

Upeneus parvus - - - - - - - - - 0.05 - 0.91 0.07

Total Fish per Minute 1.84 3.39 14.66 2.09 12.18 23.40 4.71 4.71 1.57 1.98 4.49 36.31 2.79

No. Minutes 9.3 53.7 20.1 80.3 23.3 11.3 55.0 77.0 22.6 63.1 97.0 5.5 69.9

Appendix A4. (continued).

A-7

APPENDIX B

A list of reef-fish species from natural reefs in the northern Gulf of Mexico
Appendix B. A list of reef-fish species from natural reefs in the northern Gulf of Mexico with a comparison among the northwestern 

Gulf, Pinnacles, and West Florida shelf. Data sources: NW Gulf: Ca-Cashman 1973, Br-Bright et al. 1974, So-Sonnier 

et al. 1976, Bo-Boland et al. 1983, Pu-Putt et al. 1986, DB-Dennis and Bright 1988a, b, Pa-Pattengill et al. 1997, and 

HB-Hoese and Moore 1998. West Florida: SW-Springer and Woodburn, 1960, Po-Powell et al. 1972, Sm-Smith et al. 

1975, Sm-Smith 1976, SH-Shipp and Hopkins 1978, Ha-Hastings 1979, WS-Williams and Shipp 1980, BG-Bullock 

and Godcharles 1982, DG-Darcy and Gutherz 1984, Cl-Clarke 1981, DK-Darnell and Kleypas 1987, and BS-Bullock 

and Smith 1991.

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Orectolobidae – carpet sharks

Ginglymostoma cirratum nurse shark So Sm76

Muraenidae – morays

Enchelycore carychroa chestnut moray Ca

Enchelycore nigricans viper moray Ca

Gymnothorax funebris green moray Cl

Gymnothorax hubbsi lichen moray X

Gymnothorax kolpos blacktail moray X

Gymnothorax miliaris goldentail moray Pa

Gymnothorax moringa spotted moray Ca DB X Sm75

Gymnothorax vicinus purplemouth moray Ca

Muraena retifera reticulated moray X Sm75 SH

Synodontidae – lizardfishes

Synodus intermedius sand diver DB DB X Sm75

Synodus saurus bluestripe lizardfish Pa Sm75

Synodus synodus red lizardfish Ca DB

Bythiditidae – viviparous brotulas

Ogilbia cayorum key brotula SW DK

Ophidiidae - cusk-eels 

Brotula barbata bearded brotula HM X

Batrachoididae

Opsanus pardus leopard toadfish HM X SW/Sm75 SH

B-2
Appendix B. (continued).

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Holocentridae - squirrelfishes

Corniger spinosus spinycheek soldierfish X

Holocentrus adscensionis squirrelfish Ca DB X Sm75

Holocentrus rufus longspine squirrelfish Ca DB

Myripristis jacobus blackbar soldierfish Ca Sm75 DG

Neoniphon marianus longjaw squirrelfish DB DB DK

Ostichthys trachypoma bigeye soldierfish X SH/DG

Plectrypops retrospinis cardinal soldierfish Ca

Sargocentron bullisi deepwater squirrelfish X Sm75 SH/DK

Sargocentron coruscum reef squirrelfish Po

Sargocentron poco saddle squirrelfish Ca

Sargocentron vexillarium dusky squirrelfish Pa Ha DG

Caproidae - boarfishes

Antigonia capros deepbody boarfish X

Aulostomidae – trumpetfishes

Aulostomus maculatus trumpetfish Ca X Sm75

Fistulariidae – cornetfishes

Fistularia petimba red cornetfish X

Fistularia tabacaria bluespotted cornetfish Bo

Scorpaenidae – scorpionfishes

Neomerinthe hemingwayi spinycheek scorpionfish DK DK

Pontinus rathbuni highfin scorpionfish X

Scorpaena bergi goosehead scorpionfish DG

Scorpaena dispar hunchback scorpionfish DB DK Po DK

Scorpaena inermis mushroom scorpionfish DK

Scorpaena plumieri spotted scorpionfish So Sm75

Scorpaenodes caribbaeus reef scorpionfish Ca

B-3
Appendix B. (continued).

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Serranidae - sea basses

Anthias nicholsi yellowfin bass SH/WS1

Anthias tenuis threadnose bass ? X

Bathyanthias mexicana yellowtail bass HM DG/BS

Cephalopholis cruentata graysby Ca Sm75 Cl

Cephalopholis fulva coney DB Sm75

Dermatolepis inermis marbled grouper Ca DB BG

Epinephelus adscensionis rock hind Ca DB Sm75 BS

Epinephelus drummondhayi speckled hind X Sm75

Epinephelus guttatus red hind Ca DB Sm75 BS

Epinephelus itajara jewfish SW/Sm75

Epinephelus morio red grouper Bo/Pa Sm75/DK BS

Epinephelus mystacinus misty grouper BS

Epinephelus niveatus snowy grouper DB X

Epinephelus nigritus warsaw grouper DB X

Epinephelus striatus Nassau grouper DB

Gonioplectrus hispanus Spanish flag Br X

Hemanthias aureorubens streamer bass HM ?

2 BS

Hemanthias leptus longtail bass DB X

Hemanthias vivanus red barbier DB X DG

Hypoplectrus puella barred hamlet ? Sm75 Cl/DK

Liopropoma eukrines wrasse bass Ca DB X Po SH/Cl

Liopropoma rubre peppermint bass Ca

Mycteroperca acutirostris comb grouper Ca/So

Mycteroperca bonaci black grouper DB Sm75 DG

Mycteroperca interstitialis yellowmouth grouper Ca DB Sm75

?

presence uncertain.

1

photo from DeSoto Canyon indicates P. martinicensis whereas specimen from west Florida Shelf is A. nicholsi.

2

reported by Gittings et al. 1991 but no specimens known to confirm its occurrence.

B-4
Appendix B. (continued).

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Serranidae (continued)

Mycteroperca microlepis gag HM X Sm75 DG

Mycteroperca phenax scamp DB X Sm75

Mycteroperca tigris tiger grouper DB BG

Mycteroperca microlepis gag HM X Sm75 DG

Mycteroperca phenax scamp DB X Sm75

Mycteroperca tigris tiger grouper DB BG

Mycteroperca venenosa yellowfin grouper So SW/Sm75

Paranthias furcifer creole-fish Ca X Sm75 SH

Plectranthias garrupellus apricot bass BS

Pronotogrammus martinicensis roughtongue bass DB X Po SH1

Rypticus bistrispinus freckled soapfish DK WS SH/WS

Rypticus maculatus whitespotted soapfish DB X Sm75 SH

Rypticus subbifrenatus spotted soapfish Ca

Serraniculus pumilio pygmy sea bass SW

Schultzea beta school bass HM DK DG

Serranus annularis orangeback bass DB DB BS

Serranus notospilus saddle bass X DK/DG

Serranus phoebe tattler DB DB X Cl/SH

Serranus subligarius belted sandfish Pu Sm75/DK Cl

Serranus tabacarius tobaccofish Pa Sm75 BS

Serranus tigrinus harlequin bass Sm75

Serranus tortugarum chalk bass DG

Grammatidae – basslets

Lipogramma regia royal basslet X

Opistognathidae – jawfishes

Opistognathus aurifrons yellowhead jawfish DB Sm75 Cl

Opistognathus lonchurus moustache jawfish DB X Po

B-5
Appendix B. (continued).

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Priacanthidae – bigeyes

Heteropriacanthus cruentatus glasseye snapper Ca

Priacanthus arenatus bigeye Ca DB X Sm76 SH

Pristigenys alta short bigeye DB X Sm75 SH

Apogonidae – cardinalfishes

Apogon affinis bigtooth cardinalfish Pa Po WS

Apogon aurolineatus bridle cardinalfish HM DB SW/Sm76 WS/DK

Apogon binotatus barred cardinalfish Sm75

Apogon maculatus flamefish Ca DB Sm75 Cl/DK

Apogon pseudomaculatus twospot cardinalfish So DB X Sm75 SH

Apogon quadrisquamatus sawcheek cardinalfish Pa Po/DK WS/DK

Apogon townsendi belted cardinalfish Ca

Astrapogon alutus bronze cardinalfish SW/Sm76

Astrapogon stellatus conchfish DK

Phaeoptyx conklini freckled cardinalfish Ca

Phaeoptyx pigmentaria dusky cardinalfish Ca

Phaeoptyx xenus sponge cardinalfish SW/Sm75

Malacanthidae - tilefishes

Caulolatilus chrysops goldface tilefish HM X

Caulolatilus cyanops blackline tilefish DB Po/DG

Malacanthus plumieri sand tilefish Ca Br X Sm75 Po

Carangidae – jacks

Caranx bartholomaei yellow jack So

Caranx latus horse-eye jack Ca

Caranx lugubris black jack So

Caranx ruber bar jack Ca Sm75

Seriola dumerili greater amberjack Ca Br X Sm75

Seriola rivoliana almaco jack Ca X Sm75

B-6
Appendix B. (continued).

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Lutjanidae – snappers

Apsilus dentatus black snapper HM

Etelis oculatus queen snapper DG

Lutjanus apodus schoolmaster SW

Lutjanus campechanus red snapperCa BrX Sm75 SH

Lutjanus cyanopterus cubera snapper Pa Sm75

Lutjanus griseus gray snapper Ca SW/Sm75 Cl

Lutjanus jocu dog snapper So

Lutjanus synagris lane snapper HM DB Sm75

Ocyurus chrysurus yellowtail snapper Ca SW/Sm75

Rhomboplites aurorubens vermilion snapper Pu HMX Sm75 SW/DG

Haemulidae – grunts

Anisotremus virginicus porkfish HM SW/DK

Anisotremus surinamensis black margate Pa

Haemulon aurolineatum tomtate Ca SW/Sm75

Haemulon melanurum cottonwick Ca Br

Haemulon macrostomum Spanish grunt Pa SW/DK

Haemulon plumieri white grunt HM SW/Sm75

Haemulon sciurus bluestriped grunt Po

Haemulon striatum striped grunt DB Po/DG

Inermiidae – bonnetmouths

Emmelichthyops atlanticus bonnetmouth Pa

Inermia vittata boga Pa DG

Sparidae – porgies

Archosargus probatocephalus sheepshead Pu Sm76

Calamus bajonado jolthead porgy Sm75 DK

Calamus calamus saucereye porgy Pa DG

Calamus nodosus knobbed porgy CaX Sm75 DK

B-7
Appendix B. (continued).

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Sparidae (continued)

Calamus penna sheepshead porgy DG

Pagrus pagrus red porgy DB X Sm75 DK

Sciaenidae – drums

Equetus lanceolatus jackknife fish Ca Br Sm75 Cl

Equetus punctatus spotted drum So DB

Odontoscion dentex reef croaker HM Sm76

Pareques iwamotoi blackbar drum HM X SH

Pareques umbrosus cubbyu Pu DB X Sm75 SH/DK

Mullidae – goatfishes

Mulloidichthys martinicus yellow goatfish Ca

Pseudupeneus maculatus spotted goatfish Ca Br Sm76

Chaetodontidae –butterflyfishes

Chaetodon aculeatus longsnout butterflyfish Ca DB

Chaetodon aya bank butterflyfish Pa Ca X Po SH/DK

Chaetodon capistratus foureye butterflyfish So Sm75 DK

Chaetodon ocellatus spotfin butterflyfish Ca X SW/Sm75 SH/DK

Chaetodon sedentarius reef butterflyfish Ca Br X Sm75 SH/DK

Chaetodon striatus banded butterflyfish So Sm76/DK

Pomacanthidae – angelfishes

Centropyge argi cherubfish Ca DB Sm75

Holacanthus bermudensis blue angelfish Ca Br X SW/Sm5 SH/DK

Holacanthus ciliaris queen angelfish Ca X SW/Sm75 SH

Holacanthus tricolor rock beauty Ca Br X WS

Pomacanthus arcuatus gray angelfish DB Sm75/DK

Pomacanthus paru French angelfish Ca DB SW/Sm75

B-8
Appendix B. (continued).

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Kyphosidae - sea chubs

Kyphosus incisor Yellow chub So

Kyphosus sectatrix Bermuda chub Ca Po/Cl

Pomacentridae – damselfishes

Abudefduf saxatilis sergeant major Pu SW/Sm76

Chromis cyanea blue chromis Ca Sm75

Chromis enchrysura yellowtail reeffish Ca BrX Sm75 SH/DK

Chromis insolata sunshinefish DBX Po

Chromis multilineata brown chromis Ca

Chromis scotti purple reeffish CaX Sm75 SH/DK

Microspathodon chrysurus yellowtail damselfish Ca DK?

Stegastes adustus dusky damselfish Bo

Stegastes diencaeus longfin damselfish Pa

Stegastes leucostictus beaugregory Pu DK

Stegastes partitus bicolor damselfish Ca Sm75 DK

Stegastes planifrons threespot damselfish Ca WS

Stegastes variabilis cocoa damselfish Ca SW/Sm75 Cl

Cirrhitidae – hawkfishes

Amblycirrhitus pinos redspotted hawkfish Ca DB

Labridae – wrasses

Bodianus pulchellus spotfin hogfish Ca BrX Cl SH/DG

Bodianus rufus Spanish hogfish Ca Sm75

Clepticus parrae creole wrasse Ca

Decodon puellaris red hogfish HMX DK

Doratonotus megalepis dwarf wrasse Ha

Halichoeres bathyphilus greenband wrasseX Po

Halichoeres bivittatus slippery dick Ca DB SW/Sm75 Cl

Halichoeres caudalis painted wrasse So SW/Sm75 Cl/DK

B-9
Appendix B. (continued).

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Labridae (continued)

Halichoeres garnoti yellowhead wrasse Ca

Halichoeres maculipinna clown wrasse Ca

Halichoeres pictus rainbow wrasse Po Cl/DK

Halichoeres radiatus puddingwife Ca DB

Lachnolaimus maximus hogfish So SW/Sm75 Cl

Thalassoma bifasciatum bluehead Ca Sm75

Xyrichtys martinicensis rosy razorfish Po

Xyrichtys splendens green razorfish Pa Po

Scaridae – parrotfishes

Scarus coelestinus midnight parrotfish Pa Ha

Scarus iserti striped parrotfish So/Pa Sm75

Scarus taeniopterus princess parrotfish Ca

Scarus vetula queen parrotfish Ca Sparisoma atomarium greenband parrotfish DB DB WS

Sparisoma aurofrenatum redband parrotfish Ca Sm75

Sparisoma chrysopterum redtail parrotfish Ha DG

Sparisoma rubripinne redfin parrotfish Ha

Sparisoma radians bucktooth parrotfish HM Sm75 DK

Sparisoma viride stoplight parrotfish Ca Ha

Labrisomidae - labrisomids

Labrisomus haitiensis longfin blenny Sm75

Labrisomus nuchipinnis hairy blenny HM

Starksia ocellata checkered blenny Ca SW/Cl DK

Nemaclinus atelestos threadfin blenny Ca

Paraclinus fasciatus banded blenny Po

B-10
Appendix B. (continued).

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Chaenopsidae – pikeblennies

Chaenopsis limbaughi yellowface pikeblenny Po Po

Chaenopsis ocellata bluethroat pikeblenny SW SW/Po

Emblemaria atlantica banner blenny DB Cl Cl

Emblemaria pandonis sailfin blenny Ca Sm76

Emblemaria piratula pirate blenny WS WS

Blenniidae- combtooth blennies

Hypleurochilus bermudensis barred blenny Ca Po

Hypleurochilus multifilis plumed blenny HM

Hypleurochilus caudovittatus zebratail blenny SW/DK

Hypsoblennius invemar tessellated blenny HM

Ophioblennius atlanticus redlip blenny Ca

Parablennius marmoreus seaweed blenny HM/DB X Sm75 DK

Scartella cristata molly miller HM

Gobiidae – gobies

Coryphopterus eidolon pallid goby Pa

Coryphopterus glaucofraenum bridled goby Sm75

Coryphopterus punctipectophorus spotted goby HM/Pa X SW/Cl

Coryphopterus thrix sand goby Ca

Evermannichthys spongicola sponge goby Po

Gnatholepis thompsoni goldspot goby Ca

Gobiosoma grosvenori rockcut goby DK

Gobiosoma horsti yellowline goby Sm75 DK

Gobiosoma macrodon tiger goby SW/DK

Gobiosoma oceanops neon goby Ca Sm75 Cl

Gobiosoma xanthiprora yellowprow goby X

3 X3 Cl Cl

Gobulus myersi paleback goby Cl

3

new record in NW Gulf

B-11
Appendix B. (continued).

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Gobiidae (continued)

Lythrypnus elasson dwarf goby Cl/WS WS

Lythrypnus nesiotes island goby Ca Po WS/DK

Lythrypnus phorellus convict goby Ca

Lythrypnus spilus bluegold goby Ca

Priolepis hipoliti rusty goby Ca

Psilotris celsus highspine goby Cl

Risor ruber tusked goby CaX Po WS/DK

Microdesmidae - dartfishes

Ptereleotris calliurus blue dartfish DB DB SW/Sm75 Cl

Ephippidae – spadefish

Chaetodipterus faber Atlantic spadefish Pu DK Sm75

Bothidae – lefteye flounder

Bothus lunatus peacock flounder Pa

Bothus ocellatus eyed flounder Pa DK DK

Acanthuridae – surgeonfishes

Acanthurus bahianus ocean surgeonfish Ca

Acanthurus chirurgus doctorfish So Sm75

Acanthurus coeruleus blue tang Ca SW/Sm76

Sphyraenidae – barracudas

Sphyraena barracuda great barracuda Ca DB Sm76

Balistidae – leatherjackets

Balistes capriscus gray triggerfish CaX Sw/Sm75 DG

Balistes vetula queen triggerfish Ca DB Sm76

Canthidermis maculata rough triggerfish Ca

Canthidermis sufflamen ocean triggerfish Ca DB Po

Melichthys niger black durgon Ca

Xanthichthys ringens sargassum triggerfish Ca DB DK DG

B-12
Appendix B. (continued).

NW Gulf Pinnacles West Florida Shelf

Scientific Name Common Name <60 m >60 m 60-110 m <30 m >30 m

Monacanthidae – filefishes

Aluterus heudelotii dotteral filefish DK DK

Aluterus monoceros unicorn filefish Bo/Pa

Aluterus scriptus scrawled filefish Ca DK Cl DK

Cantherhines macrocerus whitespotted filefish So DK

Cantherhines pullus orangespotted filefish Ca

Monacanthus tuckeri slender filefish Ca X

Stephanolepis setifer pygmy filefish Ca Sm75 DK

Ostraciidae – boxfishes

Acanthostracion polygonia honeycomb cowfish Pa DG

Lactophrys bicaudalis spotted trunkfish Pa

Lactophrys trigonus trunkfish SW

Lactophrys triqueter smooth trunkfish Ca Po

Tetraodontidae - puffers

Canthigaster rostrata sharpnose puffer Ca DB X Sm75 DK

Canthigaster jamestyleri goldfaced puffer X

Sphoeroides spengleri bandtail puffer Pa ? X SW/Sm75 SH

Diodontidae – porcupinefishes

Chilomycterus antennatus bridled burrfish Pa

Chilomycterus antillarum web burrfish X

Diodon holocanthus balloonfish So DB X Po

Diodon hystrix porcupinefish Pa

Triacanthodidae

Parahollardia lineata jambeau DK DK/DG

B-13

APPENDIX C

Food habits of reef fishes of the Pinnacles Reef Tract
C-2

Appendix C. Food habits of reef fishes of the Pinnacles Reef Tract by trophic guild. 

Number of specimens examined that contained prey are given, along with 

range of standard length (SL). Values given are based on relative abundance 

(% number) of prey items in the diet as determined by stomach content 

analysis.

Trophic Guild: Mesoplanktivores

Bregmaceros cantori codlet Paranthias furcifer creole-fish

2 specimens: 51 mm SL 1 specimen: 245 mm SL

FOOD FOOD

Copepods 60.0 Polychaetes 35.7

Amphipods 20.0 Copepods 28.6

Shrimp Larvae 20.0 Shrimp larvae 21.4

Crab larvae 14.3

Chromis enchrysura yellowtail reeffish

20 specimens: 66-90 mm SL Pronotogrammus martinicensis roughtongue bass

120 specimens: 70-152 mm SL 

FOOD

Copepods 32.4 FOOD

Larvaceans 32.0

Invertebrate eggs 19.1 Copepods 69.7

Polychaetes 13.3 Invertebrate eggs 11.0

Shrimp and shrimp larvae 2.1 Polychaetes 10.3

 sergestids Fish eggs 2.4

Crab larvae 1.1 Larvaceans 1.9

Ostracods 1.7

Radiolarians 1.5

Hemanthias vivanus red barbier Urochordates 0.5

47 specimens: 67-110 mm SL salps

Crab larvae 0.4

FOOD Amphipods 0.2

Copepods 78.8 hyperiids

Polychaetes 17.7 Barnacle larvae 0.1

Invertebrate eggs 3.0 Fish larvae 0.05

Larvaceans 0.14 squid 0.05

Ostracods 0.14

Urochordates 0.07

 salps

Amphipods 0.04

 hyperiids

Radiolarians 0.03

Pteropods 0.03

Chaetognaths 0.02

Crab larvae 0.01

Fishes 0.01

 Hemanthias vivanus

Fish larvae 0.01

Fish eggs 0.01
C-3

Appendix C. (continued).

Trophic Guild: Macroplanktivores

Apogon pseudomaculatus twospot cardinalfish Priacanthus arenatus bigeye

5 specimens: 62-71 mm SL 15 specimens: 196-295 mm SL 

FOOD FOOD

Isopods 37.5 Squids 32.9

Crabs 25.0 Stomatopod larvae 27.6

Shrimps and shrimp larvae 25.0 Fish larvae 14.5

Crab larvae 12.5 Scomberomorus sp.

 Bothus sp.

 Synodus sp.

Chaetodon aya bank butterflyfish Crab larvae 14.5

13 specimen: 67-94 mm SL Shrimp 3.9

Fishes 2.6

FOOD nettostomatid

Shrimp larvae 1.3

Crab and shrimp larvae 90.1 Pteropods 1.3

Amphipods 5.1 Crabs 1.3

Gorgonians 4.8

Rhomboplites aurorubens vermilion snapper

Pristigenys alta short bigeye 72 specimens: 120-372 mm SL 

9 specimens: 134-257mm SL 

FOOD

FOOD Amphipods 15.2

Shrimp 40.0 hyperiids

 Lucifer faxoni Copepods 15.2

Squids 16.0 Polychaetes 14.3

Shrimp larvae 16.0 Fish and fish larvae 10.6

Crab larvae 16.0 Bregmaceros cantori

Stomatopod larvae 6.0 myctophid

Fishes 2.0 Prionotus sp.(larvae)

Crabs 2.0 Synodus sp. (larvae)

Shrimps and shrimp larvae 10.1

Lucifer faxoni

Urochordates 9.9

 salps

Stomatopod larvae 8.0

Pteropods 6.9

Crab larvae 3.7

Ostracods 2.1

Fish eggs 1.4

Barnacle larvae 1.4

Larvaceans 1.1

Squids 0.2
C-4

Appendix C. (continued).

Trophic Guild: Benthic Carnivores

Bodianus pulchellus spotfin hogfish Decodon puellaris red hogfish

4 specimens: 155-161 mm SL 2 specimens: 125-132 mm SL 

FOOD FOOD

Pelecypods 38.1 Pelecypods 37.5

Crabs 19.0 Gastropods 31.3

Fishes 19.0 Crab larvae 12.5

 H. vivanus Polychaetes 6.3

 P. martinicensis Crabs 6.3

Gastropods 14.3 pagurid

Ophiuroids 9.5 Stomatopod larvae 6.3

Caulolatilus chrysops goldface tilefish Halichoeres bathyphilus greenband wrasse

2 specimens: 331-443 mm SL 10 specimens: 97-107 mm SL

FOOD FOOD

Polychaetes 45.0 Pelecypods 48.3

Stomatopods 25.0 Crab larvae 13.8

Pelecypods 20.0 Ophiuroids 6.9

Ophiuroids 5.0 Crabs 6.9

Crabs 5.0 Shrimps and shrimp larvae 6.9

Foraminifera 6.9

Fishes 3.4

Pareques iwamotoi blackbar drum Polychaetes 3.4

1 specimen: 234 mm SL Squids 3.4

FOOD Pagrus pagrus red porgy

Ophiuroids 33.3 28 specimens: 147-455 mm SL 

Gastropods 33.3

Crabs 33.3 FOOD

Crabs 34.0

Pareques umbrosus cubbyu pagurids

4 specimens: 110-177 mm SL Munida sp.

 Calappa sp.

FOOD Petrolisthes sp.

Crabs 36.4 Echinoids 20.2

 Munida sp. spatangoids

 Calappa sp. Gastropods 16.0

 xanthid Stomatopod larvae 6.4

Shrimps 36.4 Fishes 5.3

Shrimp larvae 9.1 Pristopomoides aquilonaris

Ophiuroids 9.1 Kathetostoma albigutta

Polychaetes 9.1 Hemanthias vivanus

Stomatopods 4.3

Polychaetes 4.3

Pelecypods 3.2

Crab Larvae 2.1

Shrimps 2.1

 alpheid

Holothiuroids 1.1

Sponges 1.1
C-5

Appendix C. (continued).

Trophic Guild: Generalized Carnivores

Centropristis ocyurus bank sea bass Malacanthus plumieri sand tilefish

6 specimens: 142-212 mm SL 7 specimens: 345-376 mm SL 

FOOD FOOD

Crabs 72.2 Gastropods 25.0

 pagurids Fishes 18.8

 portunids Hemanthias vivanus

 calappid Pronotogrammus martinicensis

 Munida sp. Crabs 18.8

Gastropods 11.1 Pelecypods 18.8

Ophiuroids 5.6 Ophiuroids 12.5

Shrimps and shrimp larvae 5.6 Bryozoans 6.3

Coral fragment 5.6

Opsanus pardus leopard toadfish

Epinephelus flavolimbatus yellowedge grouper 5 specimens: 194-215 mm SL 

2 specimens: 156-694 mm SL 

FOOD

FOOD Gastropods 33.3

Fishes 66.7 Crabs 25.0

 myctophid Fishes 25.0

Shrimps 33.3 Pronogrammus martinicensis

Crinoids 8.3

Echinoids 8.3

Holocentrus adscensionis squirrelfish

3 specimens: 168-228 mm SL 

Sargocentron bullisi deepwater squirrelfish

FOOD 2 specimens: 129-134 mm SL 

Fishes 42.9

 Hemanthias vivanus FOOD

Crab larvae 42.9 Crabs and crab larvae 33.3

Shrimps 14.3 Stomatopod larvae 33.3

 alpheid Polychaetes 16.7

Chitons 16.7

Liopropoma eukrines wrasse bass

1 specimen: 70 mm SL Serranus phoebe tattler

44 specimens: 107-168 mm SL 

FOOD

Fishes 100.0 FOOD

 Coryphopterus sp. Crabs 41.7

 Munida sp.

 Xanthids

Lutjanus campechanus red snapper Fishes 37.5

26 specimens: 192-465 mm SL Hemanthias vivanus

 Coryphopterus sp.

FOOD Crab larvae 5.6

Fishes 56.4 Shrimps 5.6

 Hemanthias vivanus alpheids

 Serranus sp. Fish larvae 2.8

 Bregmaceros cantori Ophiuroids 1.4

 Hoplunnus sp. Squids 1.4

 Trachurus lathami Stomatopods 1.4

Isopods 10.3 Amphipods 1.4

Urochordates 10.3 Polychaetes 1.4

Shrimps 7.7

 Sicyona sp.

Heteropods 5.1

Crabs 5.1

Stomatopods 2.6

Squids 2.6
C-6

Appendix C. (continued).

Trophic Guild: Piscivores

Aulostomus maculatus trumpetfish Seriola dumerili greater amberjack

1 Specimen: 160 mm SL 6 specimens: 561-744 mm SL 

FOOD FOOD

Fishes 100.0 Fishes 100.0

Coryphopterus sp. Synodus sp.

 Prionotus sp.

 Hemanthias vivanus

Gymnothorax saxicola honeycomb moray Pronotogrammus martinicensis

1 specimen: 453 mm SL Trachurus lathami

FOOD

Fishes 100.0

 Hippocampus sp.

Trophic Guild: Epibenthic Browsers

Chaetodon sedentarius reef butterflyfish Holacanthus bermudensis blue angelfish

8 specimens: 92-107mm SL 4 specimens: 210-291mm SL 

FOOD FOOD

Hydrozoans 71.4 Bryozoans 41.7

Gorgonians 14.4 Gorgonians 41.7

Copepods 5.2 Crinoids 8.3

Urochordates 3.2 Sponges 8.3

 tunicates

Ostracods 2.2

Shrimp larvae 2.1

Echinoid spines 1.5

Canthigaster rostrata sharpnose puffer

2 specimens: 37-40 mm SL 

FOOD

Crinoids 78.6

Bivalves 7.8

Gastropods 3.2

Chitons 2.1
APPENDIX D

Characteristic ichthyofauna of Pinnacles Reef Tract,

displaying the biodiversity of fishes by trophic guild.
Appendix D. Characteristic ichthyofauna of Pinnacles Reef Tract, displaying the biodiversity of fishes by trophic guild. Maximum body 

size given follows Smith-Vaniz et al. (1999) or Robins and Ray (1986). Depth range is based on videotape analysis or 

site of collection recorded for each species.

Maximum Body Activity Study Depth

Species Common Name Size (mm) Pattern Sites Range(m)

Mesoplanktivores

1. Anthias tenuis threadnose bass 90 SL D A, R, Y 61- 88

2. Bregmaceros cantori antenna codlet 60 SL N R, S 65-110

3. Chromis enchrysura yellowtail reeffish 105 SL D D, R, T, Y 61- 80

4. Hemanthias leptus longtail bass (adult) 450 SL D L, S 100-110

5. Hemanthias leptus longtail bass (juvenile)

6. Hemanthias vivanus red barbier 120 SL D All Sites 61-110 

FL=fork length, SL=standard length, and TL=total length.

D=Diurnal, N=Nocturnal, C=Crepuscular.

A=Alabama Alps, D=Double Top Reef, L=Ludwick and Walton Pinnacles, R=Roughtongue Reef, 

S=Scamp Reef, T=Triple Top Reef, Y=Yellowtail Reef.

D-2
456

123

D-3
Appendix D. (continued).

Maximum Body Activity Study Depth

Species Common Name Size (mm) Pattern Sites Range(m)

Mesoplanktivores (continued)

 7. Paranthias furcifer creole-fish 350 SL D R, Y 61- 80

 8. Pronotogrammus martinicensis roughtongue bass 160 SL D All Sites 61-110

Macroplanktivores

 9 Apogon pseudomaculatus twospot cardinalfish 105 TL N All Sites 61-110

10. Chaetodon aya bank butterflyfish 150 TL D All Sites 61-110

11. Opistognathus lonchurus moustache jawfish 100 TL D Y, R 61- 80

12. Priacanthus arenatus bigeye 360 SL N A, L, R, Y 61-110

FL=fork length, SL=standard length, and TL=total length.

D=Diurnal, N=Nocturnal, C=Crepuscular.

A=Alabama Alps, D=Double Top Reef, L=Ludwick and Walton Pinnacles, R=Roughtongue Reef, 

S=Scamp Reef, T=Triple Top Reef, Y=Yellowtail Reef. 

D-4
10

11

12

789

D-5
Appendix D. (continued).

Maximum Body Activity Study Depth

Species Common Name Size (mm) Pattern Sites Range(m)

Macroplanktivores (continued)

13. Pristigenys alta short bigeye 330 FL N All Sites 61-110

14. Rhomboplites aurorubens vermilion snapper 600 TL D/N All Sites 61-110

Generalized Carnivores

15. Bodianus pulchellus spotfin hogfish 330 FL N R, Y 61- 80

16. Brotula barbata bearded brotula 330 FL N L, S 100-110

17. Centropristis ocyurus bank sea bass 300 TL D/N D, Y 61- 80

18. Corniger spinosus spinycheek soldierfish 200 TL N All Sites 61-110

FL=fork length, SL=standard length, and TL=total length.

D=Diurnal, N=Nocturnal, C=Crepuscular.

A=Alabama Alps, D=Double Top Reef, L=Ludwick and Walton Pinnacles, R=Roughtongue Reef, 

S=Scamp Reef, T=Triple Top Reef, Y=Yellowtail Reef. 

111 D-6
13

14

16

17

15 18

D-7
Appendix D. (continued).

Maximum Body Activity Study Depth

Species Common Name Size (mm) Pattern Sites Range(m)

Generalized Carnivores (continued)

19. Epinephelus drummondhayi speckled hind (adult) 1100 TL D/N Y 61- 80

20. Epinephelus drummondhayi speckled hind (juvenile)

21. Epinephelus flavolimbatus yellowedge grouper 1150 TL D/N L 100-110

22. Epinephelus niveatus snowy grouper 1200 TL D/N D, L 70-110

23. Epinephelus nigritus warsaw grouper 2300 TL D/N A 70- 88

24. Gonioplectrus hispanus Spanish flag (adult) 230 TL D/N All Sites 61-110

FL=fork length, SL=standard length, and TL=total length.

D=Diurnal, N=Nocturnal, C=Crepuscular.

A=Alabama Alps, D=Double Top Reef, L=Ludwick and Walton Pinnacles, R=Roughtongue Reef, 

S=Scamp Reef, T=Triple Top Reef, Y=Yellowtail Reef. 

D-8
19

20

21

22

23

24

D-9
Appendix D. (continued).

Maximum Body Activity Study Depth

Species Common Name Size (mm) Pattern Sites Range(m)

Generalized Carnivores (continued)

25. Gonioplectrus hispanus Spanish flag (juvenile) 230 TL D/N All Sites 61-110

26. Gymnothorax hubbsi lichen moray 300 TL N A, Y 61- 88

27. Holocentrus adscensionis squirrelfish 345 SL N R, Y 61- 80

28. Liopropoma eukrines wrasse bass 130 TL D/N All Sites 61-110

29. Lutjanus campechanus red snapper 1000 TL D/N All Sites 61-110

30. Malacanthus plumieri sand tilefish 600 TL D R, Y 61- 80

FL=fork length, SL=standard length, and TL=total length.

D=Diurnal, N=Nocturnal, C=Crepuscular.

A=Alabama Alps, D=Double Top Reef, L=Ludwick and Walton Pinnacles, R=Roughtongue Reef, 

S=Scamp Reef, T=Triple Top Reef, Y=Yellowtail Reef. 

D-10
26

27

28

29

30

25

D-11
Appendix D. (continued).

Maximum Body Activity Study Depth

Species Common Name Size (mm) Pattern Sites Range(m)

Generalized Carnivores (continued)

31. Opsanus pardus leopard toadfish 380 TL D/N R, Y 61- 80

32. Pontinus rathbuni highfin scorpionfish 250 TL D/N R, Y 61- 80

33. Rypticus maculatus whitespotted soapfish 200 TL D/N R, Y 61- 80

34. Sargocentron bullisi deepwater squirrelfish 130 SL N L, R, Y 61-110

35. Scorpaena dispar hunchback scorpionfish 300 TL N All Sites 61-110

36. Serranus phoebe tattler 215 SL D All Sites 61-110

FL=fork length, SL=standard length, and TL=total length.

D=Diurnal, N=Nocturnal, C=Crepuscular.

A=Alabama Alps, D=Double Top Reef, L=Ludwick and Walton Pinnacles, R=Roughtongue Reef, 

S=Scamp Reef, T=Triple Top Reef, Y=Yellowtail Reef.

D-12
31

33

34

35

36

32

117 D-13
Appendix D. (continued)

Maximum Body Activity Study Depth

Species Common Name Size (mm) Pattern Sites Range(m)

Piscivores

37. Aulostomus maculatus trumpetfish 750 TL D A, R, Y 61- 88

38. Fistularia petimba red cornetfish 1800 TL D/N R, S 65-110

39. Gymnothorax kolpos blacktail moray 900 TL D/N All Sites 61-110

40. Muraena retifera reticulated moray 900 TL D/N All Sites 61-110

41. Mycteroperca microlepis gag 1200 TL D/C All Sites 61-110

42. Mycteroperca phenax scamp 900 TL D/C All Sites 61-110

FL=fork length, SL=standard length, and TL=total length.

D=Diurnal, N=Nocturnal, C=Crepuscular.

A=Alabama Alps, D=Double Top Reef, L=Ludwick and Walton Pinnacles, R=Roughtongue Reef, 

S=Scamp Reef, T=Triple Top Reef, Y=Yellowtail Reef.

D-14
D-15

37

38

39

40

41 42
Appendix D. (continued).

Maximum Body Activity Study Depth

Species Common Name Size (mm) Pattern Sites Range(m)

Piscivores (continued)

43. Seriola fasciata lesser amberjack 680 FL D/N All Sites 61-110

44. Synodus intermedius sand diver 380 SL D/N Y, L 61-110

Benthic Carnivores 

45. Caulolatilus chrysops goldface tilefish 600 TL D/N L 100-110

46. Caulolatilus intermedius anchor tilefish 600 TL D/N L 100-110

47. Calamus leucosteus whitebone porgy 460 TL D/N Y, R 61- 80

48. Calamus nodosus knobbed porgy 460 TL D/N Y, R 61- 80

FL=fork length, SL=standard length, and TL=total length.

D=Diurnal, N=Nocturnal, C=Crepuscular.

A=Alabama Alps, D=Double Top Reef, L=Ludwick and Walton Pinnacles, R=Roughtongue Reef, 

S=Scamp Reef, T=Triple Top Reef, Y=Yellowtail Reef. 

D-16
43 46

44

45

47

48

D-17
Appendix D. (continued)

Maximum Body Activity Study Depth

Species Common Name Size (mm) Pattern Sites Range(m)

Benthic Carnivores (continued)

49. Decodon puellaris red hogfish 300 TL D All Sites 61-110

50. Halichoeres bathyphilus greenband wrasse (f) 230 TL D All Sites 61-110

51. Halichoeres bathyphilus greenband wrasse (m)

52. Ogcocephalus declivirostris slantbrow batfish 165 TL D/N All Sites 61-110

53. Pagrus pagrus red porgy 910 TL D/N All Sites 61-110

54. Pareques iwamotoi blackbar drum 300 TL D/N A, L, S 70-110

FL=fork length, SL=standard length, and TL=total length.

D=Diurnal, N=Nocturnal, C=Crepuscular.

A=Alabama Alps, D=Double Top Reef, L=Ludwick and Walton Pinnacles, R=Roughtongue Reef, 

S=Scamp Reef, T=Triple Top Reef, Y=Yellowtail Reef. 

122 D-18
49 52

53

54

50

51

D-19
Appendix D. (continued)

Maximum Body Activity Study Depth

Species Common Name Size (mm) Pattern Sites Range(m)

Benthic Carnivores (continued)

55. Pareques umbrosus cubbyu 250 TL D/N All Sites 61-110

Epibenthic Browsers 

56. Chaetodon sedentarius reef butterflyfish 150 TL D A, D, R, Y 61- 88

57. Chaetodon ocellatus spotfin butterflyfish 200 TL D R, Y 61- 80

58. Canthigaster jamestyleri goldface sharpnose puffer 110 TL D R, Y 61- 80

59. Canthigaster rostrata sharpnose puffer 110 TL D A, R, Y 61- 88

60. Holacanthus bermudensis blue angelfish 440 TL D All Sites 61-110

FL=fork length, SL=standard length, and TL=total length.

D=Diurnal, N=Nocturnal, C=Crepuscular.

A=Alabama Alps, D=Double Top Reef, L=Ludwick and Walton Pinnacles, R=Roughtongue Reef, 

S=Scamp Reef, T=Triple Top Reef, Y=Yellowtail Reef.

D-20
55

56

57

58

59

60

D-21

REPORT DOCUMENTATION PAGE Form Approved

OMB No. 074-0188

Public reporting burden for this collection of information is estimated to average 1 hour per response, including the time for reviewing instructions, searching existing data sources, 

gathering and maintaining the data needed, and completing and reviewing this collection of information. Send comments regarding this burden estimate or any other aspect of this 

collection of information, including suggestions for reducing this burden to Washington Headquarters Services, Directorate for Information Operations and Reports, 1215 Jefferson 

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1. AGENCY USE ONLY (Leave blank) 2. REPORT DATE

 June 2002

3. REPORT TYPE AND DATES COVERED

 Final Report 04/15/1997-09/30/2001

4. TITLE AND SUBTITLE

Northeastern Gulf of Mexico Coastal and Marine Ecosystem Program

Community Structure and Trophic Ecology of Fishes on the Pinnacles

Reef Tract

5. FUNDING NUMBERS

6. AUTHOR(S)

Douglas C. Weaver, George D. Dennis, and Kenneth J. Sulak

7. PERFORMING ORGANIZATION NAME(S) AND ADDRESS(ES) 8. PERFORMING ORGANIZATION

 REPORT NUMBER

U.S. Geological Survey

Florida Caribbean Science Center

7920 NW 71st Str.

Gainesville, Florida 32653

 

9. SPONSORING / MONITORING AGENCY NAME(S) AND ADDRESS(ES) 10. SPONSORING / MONITORING

 AGENCY REPORT NUMBER

 U.S. Geological Survey Minerals Management Service

 Eastern Regional Office Gulf of Mexico OCS Region

 1700 Leetown Road 1201 Elmwood Park Boulevard

 Kearneysville, WV 25430 New Orleans, LA 701213-2394

USGS BSR 2001-0008

OCS Study MMS 2002-0034 

11. SUPPLEMENTARY NOTES

12a. DISTRIBUTION / AVAILABILITY STATEMENT 12b. DISTRIBUTION CODE

13. ABSTRACT (Maximum 200 Words)

Topographic prominences form reef-like features along the outer continental shelf (OCS) edge and continental 

slope of the southeastern United States and Gulf of Mexico. An extensive set of reef features, known as the 

Pinnacles Reef Tract, occurs off Mississippi and Alabama at depths between 60 and 110 m. Eighteen sites within 

the combined Mississippi-Alabama Marine Ecosystems Study and Mississippi-Alabama Shelf Pinnacle Trend Habitat 

Mapping Study area were surveyed. All sampling combined returned over 6,000 specimens for food habits 

analyses, taxonomic verification and documentation, and subsequent life history analyses. The results of this study 

have expanded the number of fishes documented from the Pinnacles Reef Tract, and is the first extensive collection 

of deep (greater than 60 m) reef fishes made in this region, and possibly worldwide. Our present knowledge of the 

Pinnacles ichthyofauna can be used to characterize the fundamental taxonomic composition and trophic structure of 

deep reef-fish communities throughout southeastern United States and Gulf of Mexico OCS areas and therefore be 

useful in assessing future anthropogenic impacts on faunal structure.

14. SUBJECT TERMS 15. NUMBER OF PAGES

94 pp. + apps

Pinnacles, geology, deep reef, hard bottom communities, fish, trophic ecology, food web, 

community ecology

16. PRICE CODE

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