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Abstract—Submersible surveys at nu- Deep reef fish surveys by submersible on merous reefs and banks in the north- western Gulf of Mexico (NWGOM) were Alderdice, McGrail, and Sonnier Banks in conducted as part of the Sustainable Seas Expedition (SSE) during July/August the Northwestern Gulf of Mexico 2002 to identify reef fish communities, characterize benthic habitats, and iden- Douglas C. Weaver tify deep coral reef ecosystems. To iden- tify the spatial extent of hard bottom reef Emma L. Hickerson communities, the Flower Garden Banks George P. Schmahl National Marine Sanctuary (FGBNMS) Flower Garden Banks National Marine Sanctuary, NOAA and the U.S. Geological Survey (USGS) 4700 Avenue U, Building 216 mapped approximately 2000 km2 of the Galveston, TX 77551 Northwestern Gulf of Mexico (NWGOM) Email (for Weaver): [email protected] continental shelf during June 2002 with high-resolution multibeam bathymetry. Previous investigations conducted on the features of interest (with the exceptions of East and West Flower Garden and Son- nier Banks, accessible by SCUBA) had Introduction The reefs and banks of the north- not been conducted since the 1970s and western Gulf have been afforded some 1980s, and did not have the use of high- resolution maps to target survey sites. The The northwestern Gulf of Mexico protection since the early 1970s. The base maps were instrumental in navigating (NWGOM) reefs and banks are asso- MMS recognized early on that these submersibles to specific features at each ciated with the surface expression of sensitive areas should not be subjected study site during the Sustainable Seas salt domes, and have been identified to the direct impacts of offshore oil and Expedition (SSE)—a submersible effort as unique biological features warrant- gas development, and defined them as culminating from a partnership between the National Atmospheric and Oceanic ing protection by Minerals Manage- “no activity” zones (Fig. 1). However, Administration (NOAA) and the National ment Service (MMS) from oil and gas these features are subject to a variety Geographic Society (NGS). We report the activities. The most recent investiga- of other potential impacts unrelat- initial findings of our submersible surveys, tions of reef fishes on the shelf-edge ed to offshore development, which including habitat and reef fish diversity at features in the northwestern Gulf may not be regulated sufficiently. McGrail, Alderdice, and Sonnier Banks. A total of 120 and 40,724 indi- (with the exceptions of East and West These impacts include those of com- viduals were identified from video surveys Flower Garden, Stetson and Sonnier mercial shipping (predominantly an- at the three banks. Planktivorous fishes Banks) were conducted during the choring), fishing, cultural resource constituted over 87% by number for the 1970s and 1980s (Rezak et al., 1985; recovery and recreational activities. three banks, ranging from 81.4% at Son- Dennis and Bright, 1988a, 1988b), There is recent emphasis on the in- nier Banks to 94.3% at Alderdice Bank, indicating a direct link to pelagic prey without the use of high-resolution creased use of marine protected areas communities, particularly in the deep multibeam bathymetry or high-resolu- (MPA’s) in many ocean regions as a reef zones. High numbers of , tion camera systems. method to consolidate management snappers, jacks, and other fishery spe- The Sustainable Seas Expedition under a coordinated mechanism. Ac- cies were observed on all three features. (SSE) was a five-year collaborative curately identifying the resources in These sites were nominated as Habitat Areas of Particular Concern (HAPC) by effort between NOAA and the Na- the northwestern Gulf will contribute the Gulf of Mexico Fishery Council in tional Geographic Society (NGS) to greatly to this effort. March 2004. Data obtained during this explore important marine ecosystems project will contribute to benthic habitat with special emphasis on the nation’s characterization and assessment of the as- National Marine Sanctuaries. Led by Background sociated fish communities through future SCUBA, ROV, and submersible missions, Sylvia Earle, Explorer-in-Residence of and allow comparisons to other deep reef the NGS, the SSE program utilized The Department of Oceanography ecosystems found throughout the Gulf of both manned and unmanned sub- at Texas A&M University (TAMU) Mexico and western Atlantic Ocean. mersible technologies to explore the conducted studies on “topographic NWGOM in 1999, 2001, and 2002. features”—Outer Continental Shelf The use of the submersibles allowed (OCS) reefs and banks—during the the Flower Garden Banks National 1970s and 1980s. Funding was pro- Marine Sanctuary (FGBNMS) team vided by the U.S. Department of the to visit several of the reefs and banks Interior (DOI), Bureau of Land Man- of the NWGOM and conduct explor- agement (BLM), out of which MMS atory dives to investigate reef fish and later emerged through reorganiza- benthic communities in July 2002. tion. The purpose of the TAMU study 70 Professional Paper NMFS 5

Figure 1 Location of outer shelf banks in the northwestern Gulf of Mexico.

was to provide the MMS with data on the geologic, sandstone outcrops on the crests of these features. The biologic, hydrologic, and chemical characteristics of the Algal-Sponge Zone covers the largest area among the reefs and banks, to be used as a basis for management reef-building zones on shelf-edge banks. The dominant decisions on tracts nominated for oil and gas leasing organisms of the zone are the coralline algae, which (Rezak et al., 1985). are the most important carbonate producers through As a comparison, the technology used for the his- algal nodules and algal reefs (Minnery et al., 1985). The toric surveys conducted by Rezak et al. (1985) included nodules range from 1 to 20 cm in size, cover up to 80 bathymetric mapping using a LORAN-C positioning sys- percent of the bottom, and generally occur between 55 tem for navigation, a 3.5kHz subbottom profiler, EG&G and 85 m. Partly Drowned Reefs are also a major biotope Uniboom seismic system for subbottom information, occurring at similar depths to the Algal-Sponge Zone, and EG&G side-scan sonar systems for bottom char- and are defined as massive reefal structures covered with acterization. Direct observations of the sea floor were living crusts of coralline algae. With increased water made using the Texas A&M submersible DRV Diaphus depth, the assemblages of the zone become less diverse, (Rezak and Tieh, 1984; Dennis and Bright, 1988a). characterized by antipatharians, comatulid crinoids, di- Each of the survey sites has a distinct biological zona- minished leafy or coralline algae, and limited fish. High tion associated with base depths and water turbidity, turbidity, sedimentation, and re-suspension occur in the and the following zones (when present), described dur- Nepheloid Zone, where exposed rocks lack encrusting ing the early TAMU/MMS studies (Rezak et al., 1985; coralline algal crusts. Drowned Reefs occurring in this 1990). The Stephanocoenia-Millepora Zone is inhabited zone are often covered with a thin veneer of sediment, by a low-diversity coral assemblage of 12 hermatypic and epifauna are scarce. This zone occurs on all banks, coral species and can be found at the FGBNMS, Mc- but its depth differs at each bank and extends to the Grail Bank, and Bright Bank to depths of 50 m. On surrounding soft bottom. midshelf banks, such as Stetson and Sonnier Banks, the The Sonnier Bank complex (historically referred Millepora-Sponge Zone occupies depths less than 36 m. to as Three Hickey Rock and Candy Mountain) is a Crusts of the hydrozoan coral, Millepora, sponges, and mid-shelf bank, defined by Rezak et al. (1985, 1990) as other epifauna occupy the tops of siltstone, claystone, or salt dome structures rising from depths of 80 m or less Weaver et al: Reef fish surveys by submersible in the Northwestern Gulf of Mexico 71

and having a relief of about 4 to 50 m. The banks are northwestern Gulf of Mexico continental shelf1. Multi- located 135 km south of the border, at posi- beam bathymetry data sets were used as base maps to tion 28°20ʹN/92°27ʹW. As noted by Rezak et al. (1985) guide submersible surveys of select topographic features the Sonnier Bank complex consists of eight separate during the 2002 Sustainable Seas Expedition to the banks or peaks associated with a single salt dome. The northern Gulf of Mexico, conducted by the FGBNMS peaks are nearly conical features with a maximum re- in conjunction with the NGS. lief of about 30 m. Observations made by SCUBA and The SSE missions initiated an ongoing multidisci- submersible have characterized the reef habitat on the plinary project led by the FGBNMS to revisit the reefs peaks as entirely encrusted with fire coral (Millepora and banks of the NWGOM and build on the descrip- sp.) and sponges (primarily Neofibularia nolitangere and tion and characterization effort conducted during past Ircinia sp.) (Rezak et al., 1985). This is the basis for the research programs. zonation name of Millepora-Sponge Zone. Rezak et al. During the SSE missions, submersible dives were con- (1985) reported three species of coral from the crests ducted on the East Flower Garden (EFGB), West Flower at Sonnier Banks: Stephanocoenia sp., Millepora sp., and Garden (WFGB), Jakkula, Sidner, Alderdice, Bright, Agaricia sp. In 1997, SCUBA surveys conducted by MMS McGrail, and Sonnier Banks. Results from the SSE and FGBNMS representatives compiled coral observa- submersible surveys at Alderdice, McGrail, and Sonnier tions on the two shallowest peaks (24 m and 18 m) of Banks are reported herein, due to the more extensive Sonnier Banks. Other hermatypic coral species added surveys conducted at these features. by MMS diver surveys include Madracis decactis, M. mirabilis, M. pharensis, Siderastrea radians, Montastraea cavernosa, and Agaricia fragilis. Materials and methods McGrail Bank (formerly referred to as 18 Fathom Bank) is located 180 km south of the coast of Louisi- The SSE dives were conducted using Nuytco Research ana, at 27°58ʹN/92°36ʹW. As described by Rezak et al. Ltd’s Deepworker 2000 (DW2000) and Deep Rover (1985), the McGrail Bank is an arcuate pair of north- (DR) submersibles (Fig. 2). Both systems are single- east-southwest trending ridges separated by a valley. Mc- person submersibles—the DW2000 rated to 610 m and Grail Bank is one of the few banks in the northwestern DR rated to 1000 m. One of the benefits of these two Gulf of Mexico that has extensive growth of reef-build- submersibles is the nearly 360° observation capability ing corals, in addition to East and West Flower Garden afforded to the pilot. Other technology that enhanced Banks. The original BLM OCS surveys documented the capabilities of the submersibles during the SSE four species of coral: the blushing star coral (Stephano- missions included accurate underwater tracking using coenia intersepta; formerly S. michilini), fire coral (Mil- the ORE International 4410C Track Point II with the lepora sp.), the great star coral (Montastraea cavernosa), Winfrog integrated navigation package, underwater and Agaricia sp. communication, and high resolution digital video and Alderdice Bank is unique among the offshore banks still camera. in that it bears outcrops of basalt associated with the Submersible surveys were conducted along selected underlying salt dome (Rezak and Tieh, 1984). Alderdice waypoints based on reef topography to survey as much Bank is located about 160 km south of Marsh Island, LA hard bottom habitat as possible during each dive. Reef at 92°00ʹW/28°05ʹN. The bank is an oval, elongate in an fishes and macroinvertebrates were only counted dur- east-west direction, and covers an area of approximately ing segments where the field of view was illuminated 9.8 km2. Rezak and Tieh (1984) states that the bank and the submersible maintained close contact to the is a classic example of the surface expression of a salt bottom. Numbers of individuals for schooling species dome. A single basalt outcropping, described by Rezak were estimated based on paused still frames and ex- and Tieh (1984), was discovered during a submersible trapolated to the entire school. were identified dive during the BLM OCS studies. Potassium-argon age to the lowest possible taxon following Humann and De- determination anaysis indicated an age of 76.8 ± 3.3 × Loach (2002), based on the taxonomic nomenclature of 106 years (late Cretaceous-Campanian). Robins et al. (1991). Trophic categories were assigned As a result of these investigations, MMS initiated regu- based on dietary information from Randall (1967), latory zones around the majority of the hard bottom Smith-Vaniz et al. (1999), or Bullock and Smith (1991) habitat in the NWGOM, regulating oil and gas activity. for individual species or closely related taxa. Regulations placed on the oil and gas industry included the mandatory shunting of discharges to within 10 m of the sea floor. 1 USGS (United States Geological Survey). 2002. Multibeam Bathy- metry Mapping of the Northwestern Gulf of Mexico. http:// During 8–28 June 2002, a multibeam mapping cruise walrus.wr.usgs.gov/pacmaps/wg-index.html. [Accessed 14 August was conducted to survey approximately 2000 km2 of the 2005]. 72 Professional Paper NMFS 5

A

B

Figure 2 Submersibles used during SSE2002. A) Deepworker 2000, piloted by G.P. Schmahl during SSE Dive A20-185 at McGrail Bank. B) Deep Rover, piloted by Kip Evans, during SSE Dive A20-186 at McGrail Bank. Photograph and still image by Kip Evans, NGS, and G. P. Schmahl, FGBNMS. Weaver et al: Reef fish surveys by submersible in the Northwestern Gulf of Mexico 73

Table 1 Submersible dives conducted at Sonnier, McGrail, and Alderdice Banks.

Depth Survey Track SSE dive no. Bank Submersible Pilot range (m) duration length (km) Habitats observed

A20-179 Alderdice Deep Worker D. Weaver 62–73 5.6 hours 1.1 basalt spires, drowned reef zone A20-180 Alderdice Deep Worker G. Schmahl 62–90 4.2 hours 0.9 basalt spires, algal nodule-sponge zone A20-181 Alderdice Deep Worker E. Hickerson 68–85 4.3 hours 1.3 Algal nodule sponge zone A20-184 McGrail Deep Worker G. Schmahl 45–78 5.2 hours 1.7 Coral cap, algal nodule-sponge zone A20-185 McGrail Deep Rover K. Evans 45–85 5.5 hours 2.5 Coral cap, algal nodule-sponge zone A20-186 Sonnier Deep Worker D. Weaver 19–60 3.2 hours 2.1 Millepora-sponge zone, drowned reef zone

The submersible dives were facilitated by the avail- around the coral cap region of McGrail Bank, while a ability of detailed multibeam bathymetry collected single submersible transect was conducted at Sonnier during June 2002. A Konsberg Simrad EM1000 high Bank (Fig. 3). Depths surveyed at Sonnier Banks ranged resolution multibeam echosounder was used to map from 19 to 60 m, McGrail Bank 45–85 m, and Alderdice the seafloor at specific features, following hydrographic Bank 62–90 m. Sonnier Banks were characterized by standards.1 The multibeam surveys were conducted and Millepora-sponge and drowned reef habitats, while Mc- data analyzed by James Gardner (Univ. of New Hamp- Grail and Alderdice transects were dominated by algal shire, formerly of USGS, Menlo Park). Resulting data nodule-sponge communities (Table 1). were gridded at 5 m resolution and used to produce One hundred and twenty (120) fishes were observed georeferenced images and 3D visualizations. Discussion in association with the deep reef communities of the of the earlier bathymetric surveys conducted within the three banks (Appendix A). McGrail Bank had the high- FGBNMS is presented in Gardner et al. (1998), and data est observed species richness (78 species), followed by from those surveys are also available1. Sonnier Banks (77), and Alderdice Bank (68). A total of Base maps were used to guide submersible opera- 40,724 fishes were counted from submersible videotape tions as part of the SSE mission to the northern Gulf from submersible dives (Table 2). of Mexico during 24 July–6 August 2002, co-sponsored The top 25 species observed on each bank are pre- by NGS and NOAA. Submersible dives were conducted sented in Table 3. Planktivorous fishes dominated the using the Deepworker 2000 and Deep Rover submers- reef communities of all three banks, but different taxa ibles aboard the Oceaneering International, Inc. M/V were numerically abundant at each location. Sonnier OCEAN PROJECT. Bank reef fish populations are dominated by the yel- Bathymetry maps were geo-referenced in ArcView lowtail reeffish (Chromis enchrysura), creole-fish (Paran- GIS (Version 3.2, Environmental Research Systems thias furcifer), and brown chromis (Chromis mulilineata). Institute, Redlands, CA), and used to plot waypoints McGrail Bank reef fish populations are dominated by for submersible transects. Submersible position was P. furcifer, the threadnose bass (Anthias tenuis), and the continuously logged during dives, and dive tracks were yellow goatfish ( martinicis). In contrast, later superimposed on bathymetry to identify location Alderdice Bank is dominated by the roughtongue bass of video surveys. (Pronotogrammus martinicensis), C. enchrysura, and P. furcifer. Planktivorous fishes constituted over 81.4% by num- Results ber at each bank, ranging from 81.6% at Sonnier Banks to 94.3% at Alderdice Bank. The remaining categories Six submersible dives were conducted at the three study comprised less than 10%, with the exception of ben- sites, focusing on hard bottom communities and high thic carnivores at Sonnier Banks and McGrail Bank, profile reef structures. Three submersible dives were represented by large populations of tomtate ( made on the basalt spire features and large mound of aurolineatum) and yellow goatfish, respectively. Benthic Alderdice Bank (Table 1). Two dives were conducted carnivores represented the second most abundant tro- 74 Professional Paper NMFS 5

phic guild on each bank. Epibenthic browsers, includ- trast, piscivores exhibited a slight increase from 0.9% ing invertebrate browsers such as pomacanthids and at Sonnier Banks to 1.6% at McGrail Bank to 1.9% at chaetodontids, declined in number from Sonnier Banks Alderdice Bank. (4.5%) to McGrail (1.6%) to Alderdice (0.5%). Herbi- vores also decreased in this order, and herbivores such Sonnier Banks as scarids and acanthurids made up 0.6% at Sonnier, 0.3% at McGrail, and 0.1% at Alderdice. Generalized High-resolution multibeam bathymetry of the Sonnier carnivores also followed this general pattern. In con- Banks reveals up to at least a dozen additional lower

Table 2 Reef fishes by trophic category at Sonnier, McGrail, and Alderdice Banks. Numbers of individuals observed during video surveys are given.

Sonnier Banks McGrail Bank Alderdice Bank Totals

No. Percent No. Percent No. Percent No. Percent

Planktivores 6284 81.4 12,465 81.6 16,717 94.3 35,466 87.1 Piscivores 68 0.9 249 1.6 335 1.9 652 1.6 General carnivores 351 4.5 218 1.4 226 1.3 795 2.0 Herbivores 49 0.6 39 0.3 11 0.1 99 0.2 Epibenthic browsers 350 4.5 237 1.6 97 0.5 783 1.9 Benthic carnivores 615 8.0 2067 13.5 346 2.0 3028 7.4 Totals 7717 100.0 15,275 100.0 17,732 100.0 40,724 100.0

A

Figure 3 A) Multibeam bathymetry map of Sonnier Banks, oblique view (bathymetry data courtesy James Gardner, University of New Hampshire, Center for Coastal and Ocean Mapping, 24 Colovos Road, Durham, NH 03824). Submersible surveys were conducted on Millepora-sponge zones and drowned reef areas as part of the Sustainable Seas Expeditions during July–August 2002. B) Plan view (ve=vertical exaggeration) of Sonnier Banks (bathymetry data courtesy James Gardner). C) A still video frame of the Millepora-Sponge cap at Sonnier Banks, with a queen angelfish, Holacanthus ciliaris. Assorted sponges and Millepora (tan in color) are present. Weaver et al: Reef fish surveys by submersible in the Northwestern Gulf of Mexico 75

B 28.35

28.345 -22m

-27m

-32m 28.34 -37m r" I»::-. 2 -42m Q. no -47m 28.335 -52m

-5701

28.33 -6201

28.325 -92.46 -92.455 -92.45 -92.445 Longitude C

Figure 3 (continued) 76 Professional Paper NMFS 5

Table 3 Top 25 fish species observed during submersible surveys of Sonnier, McGrail, and Alderdice Banks during SSE2002. Dominance rank is based on numerical abundance from videotaped ROV surveys. Trophic categories are based on dietary studies of Randall (1967) or Bullock and Smith (1991).

Sonnier Banks McGrail Bank Alderdice Bank

Species ID Common name Trophic guild1 Rank Percent Rank Percent Rank Percent

Chromis enchrysura yellowtail reeffish PL 1 23.1 5 6.19 2 24.61 furcifer creole-fish PL 2 17.7 1 43.3 3 18.49 Chromis multilineatus brown chromis PL 3 13.6 12 0.58 — — Chromis insolata sunshine fish PL 4 6.36 4 7.64 5 5.67 Thallasoma bifasciatum bluehead PL 5 5.73 18 0.22 24 0.07 Haemulon aurolineatum tomtate GC 6 5.55 — — — — Chromis scotti purple reeffish PL 7 5.34 10 0.69 8 0.97 Stegastes variabilis cocoa damselfish H 8 2.80 — — — — Clepticus parrae creole PL 9 2.34 15 0.48 14 0.19 Stegastes partitus bicolor damselfish FL 10 2.03 7 1.04 — — Kyphosus sp. /yellow chub PL 11 1.80 — — — — Rhomboplites aurorubens vermilion snapper PL 12 1.42 — — 6 3.29 Balistes capriscus gray triggerfish BC 13 1.03 — — — — Lutjanus griseus gray snapper GC 14 1.03 9 0.73 9 0.52 Lutjanus campechanus red snapper GC 15 0.90 — — — — Bodianus pulchellus spotfin hogfish GC 16 0.66 21 0.18 10 0.42 Bodianus rufus Spanish hogfish GC 17 0.58 24 0.14 — — Lutjanus buccanella blackfin snapper GC 18 0.55 — — — — Abudefduf saxatalis PL 19 0.53 — — — — Myripristis jacobus blackbar soldierfish PL 20 0.40 — — — — Chaetodon sedentarius reef butterflyfish EB 21 0.32 13 0.54 13 0.28 Ptereleotris calliurus blue goby PL 22 0.28 — — — — adscensionis rock hind GC 23 0.27 — — — — Canthigaster rostrata sharpnose puffer EB 24 0.25 — — — — Holacanthus tricolor rock beauty EB 25 0.25 — — 22 0.08 Mulloidichthys martinicus yellow goatfish BC 26 0.25 3 13.0 7 1.60 Halichoeres bathyphilus greenband wrasse BC 28 0.24 — — 25 0.06 Holacanthus bermudensis blue angelfish EB 29 0.23 — — 23 0.07 interstitialis yellowmouth PI 33 0.18 20 0.19 15 0.17 Caranx latus horse-eye jack PI 34 0.16 14 0.53 — — Pareques umbrosus cubbyu BC 36 0.16 — — 18 0.10 Chromis cyanea blue chromis PL 38 0.15 8 0.93 — — Pseudupeneus maculates spotted goatfish BC 39 0.15 — — 20 0.09 Sparisoma atomarium greenblotch parrotfish H 42 0.14 23 0.15 — — Seriola dumerili greater amberjack PI 44 0.12 — — 11 0.30 Holocentrus rufus longfin squirrelfish PL 45 0.11 22 0.18 — — Anthias tenuis threadnose bass PL — — 2 17.9 4 13.13 Apogon affinis bigtooth cardinalfish PL — — — — 12 0.29 Aulostomus maculates trumpetfish PI — — — — 16 0.14 Caranx bartholomaei yellow jack PI — — 17 0.24 — — Caranx hippos crevalle jack PI — — 19 0.19 — — Centropyge argi pygmy angelfish EB — — 11 0.62 — — Chaetodon aya bank butterflyfish PL — — — — 17 0.10 Gonioplectrus hispanus Spanish flag PI — — — — 19 0.09 Pronotogrammus martinicensis roughtongue bass PL — — 16 0.48 1 28.45 Schultzea beta school bass PL — — 6 1.43 — — Seriola rivoliana almaco jack PI — — — — 21 0.09

1 BC-Benthic carnivores (benthic and infauma), EB-Epibenthic browsers (sessile invertebrates), GC-General carnivore (mobile invertebrates and fishes), H-Herbivore, PL-Planktivore, PI-Piscivore (fishes and cephalopods). Weaver et al: Reef fish surveys by submersible in the Northwestern Gulf of Mexico 77

relief peaks associated with the feature, ranging in depth range)—a crest approximately 0.07 km2 (0.40 km depth from 18 to 55 m (Fig. 3). Deeper peaks surveyed × 0.28 km), was documented to be populated by up to during the SSE mission had reduced coverage of fire 30% coverage of S. intersepta coral heads, averaging 1m coral and a more extensive and diverse community of width × 0.75 m tall (Fig. 4B). Numerous heads reached encrusting and massive sponges. upwards of 2 m in height. In addition to the S. intersepta The 2002 SSE submersible dive covered three of the colonies, numerous 1–2 m tall M. cavernosa colonies major peaks and two of the lower relief peaks. No addi- were encountered. tional species of hermatypic coral were observed during Reef fishes observed at McGrail Bank were over- these surveys. Astropyga magnifica, a deep water echinoid, whelmingly dominated by P. furcifer, which made up was observed in crevices along the rocky reef slopes in almost half of the individuals observed (43%). Thread- 2002. Submersible investigations revealed a diverse as- nose bass comprised 18% of fishes observed, followed by semblage of fishes and invertebrates, from the species- the yellow goatfish. The occurrence of a large school of rich reef fish assemblages on the Millepora sponge zones goatfish has not been previously reported in the litera- associated with the two shallowest banks, to species- ture. While this species was observed on all three banks, depauperate drowned reef assemblages on the smaller they were in much lower abundance at Alderdice and features within the region (Fig. 3). Results of our surveys Sonnier Banks. Sunshinefish (C. insolata), school bass indicate unique biological assemblages associated with (Schultzea beta), bicolor damselfish (Stegastes partitus), each bank within the region, related to the depth of the and the blue chromis (Chromis cyanea), were the next bank crest and the extent of the turbid Nepheloid Zone, most dominant reef fish taxa, and all are planktivorous. which was encountered over surrounding soft bottom McGrail Bank lacked many of the species associated with areas at each peak during submersible surveys. the turbid drowned reef areas of Sonnier Bank, includ- Reef fishes observed at Sonnier were numerically ing H. aurolineatum and L. campechanus. dominated by planktivores, representing the top five A notable observation made during the submersible species, eight of the top ten numerically abundant surveys is that up to five aggregations of longspine species. In addition to C. enchrysura, P. furcifer, and C. urchins (Astropyga magnifica) were encountered in the multilineatus, the sunshinefish (Chromis insolata) and sandy valley between the ridges of McGrail Bank. These bluehead (Thallasoma bifasciata) were next most abun- aggregations were made up of at least 100 individuals. dant. Dense schools of planktivores, including P. furcifer, Associated with each aggregation were upwards of six C. multilineata, T. bifasciata, and creole wrasse (Clepticus juvenile ( inermis) indi- parrae), characterize the shallow crest areas of the larger viduals to each A. magnifica aggregation (Fig. 4A). peaks. Deeper reef communities were characterized by an assemblage of H. aurolineatum, red snapper (L. Alderdice Bank campechanus), greater amberjack (Seriola dumerili), and gray triggerfish (Balistes capriscus). Soft bottom regions While previous bathymetric surveys of the area identi- surrounding the main peaks had few fishes, with the ex- fied a single “spire” at Alderdice Bank, the multibeam ception of the blue goby (Ptereleotris calliurus), living in data set resolved two distinct spires with associated talus burrows in the circum-reef talus zones, and tattler (Ser- fields (Fig. 5). The second basalt spire is located ap- ranus phoebe), associated with talus zones and carbonate proximately 220 m NNE of the outcropping described debris surrounding hard bottom features. by Rezak and Tieh (1984). Both spires provide high profile structure that attracts large schools of creole- McGrail Bank fish, vermilion snapper (Rhomboplites aurorubens), and several species of grouper, snapper, and jacks. In 2002, Submersible surveys conducted during SSE on McGrail the original basalt feature was surveyed and character- Bank revealed extensive growth of a hard coral com- ized by a similar biota as originally described by Rezak et munity dominated by S. intersepta, large brain corals al. (1985). Through direction from a topside navigation (Diploria strigosa), M. cavernosa, and a species of Agaricia. technician, the submersible pilot was able to navigate Estimated coral coverage reached 30% in some areas, at both of the spires, confirming that the second feature a depth range of 45 m–60 m, while the base of the bank was indeed an additional basalt outcrop (Figs. 5A, B). is approximately 85 m (Fig. 4A). Two SSE 2002 submers- Reef fishes observed at Alderdice Bank were over- ible dives targeted the crest of the southernmost ridge whelmingly dominated by roughtongue bass, with to identify the extent of the deep coral reef community. yellowtail reeffish and creole-fish also dominating the In addition to these four previously documented spe- reef fish community. These three species made up over cies of hermatypic corals, a 2 m tall colony of Diploria 71% of the individual fishes observed. A notable ob- strigosa was noted, as was a more extensive S. intersepta servation during the SSE mission is the occurrence of reef than previously described. One reef site (45–60 m the rare marbled grouper (Dermatolepis inermis), both at 78 Professional Paper NMFS 5

-45 A 9km -50 -55 ve=10x -60 -65 -70 -75 -80

27 -85 -90 -95 -100 -1 05 -1 10 27. 97 -1 15 -120 -1 25 -130 -135 -140 -145 -150 -1 55

27.94 _92.56 .92.59 .92.62

Figure 4 A) Top: Multibeam bathymetry map of McGrail Banks, oblique view (bathymetry data courtesy James Gardner; see Fig. 3 for contact info.). Bottom: Marble grouper, Dermatolepis inermis, resting at the base of sea urchin aggregation. B) Extensive growth of hard corals, including Stephanocoenia, Montastraea, and Diploria spp., on the summits of the bank were documented during submersible surveys during the Sustainable Seas Expedition during summer 2002. Video still courtesy SSE. Weaver et al: Reef fish surveys by submersible in the Northwestern Gulf of Mexico 79

B

Figure 4 (continued)

the basalt outcroppings and the eastern algal/sponge lepora-sponge zones on the shallow peaks of the main mound. features is reflected in the abundance of creole-fish, creole wrasse, brown chromis, and other planktivores that typically associate with the shallow crests of other Discussion reefs and banks in the Gulf of Mexico, such at East Flower Garden Bank, West Flower Garden Bank, and Reef fishes of the northwestern Gulf of Mexico, as previ- Stetson Bank (Rezak et al., 1985; Pattengill-Semmens ous authors have noted, are represented by a tropical et al., 1997). Sonnier Banks also have limited deep reef assemblage on mid to outer shelf banks where suitable habitat, due to the dominance of a nepheloid layer at habitat occurs (Rezak et al., 1985, 1990; Dennis and the deeper areas surrounding the main peaks (Rezak et Bright, 1988a). Results of this survey provide the first al., 1990). This results in extensive soft bottom commu- detailed look at reef fish communities associated with nities around the banks, a lack of algal nodule-sponge the chosen features using modern submersibles, and zones, and limited distribution of partly drowned and identify differences among the fish communities as- drowned reef structures and associated reef fish com- sociated with their unique benthic assemblages. The munities. number of reef-associated species at each bank was In addition to general comments on the reef fish very similar, ranging from 68 at Sonnier Banks to 78 at community at McGrail Bank, Rezak et al. (1985) noted McGrail Bank. While total species richness was similar, significant populations of common black sea urchins the numerically dominant taxa vary between sites and (Diadema sp.) and spiny lobsters (Panulirus sp.) occur- reflect the distribution of habitat at each of the chosen ring on the ledges and along the slopes of many of the study sites. Sonnier Banks are comprised of steep sided, shelf edge banks. These observations are inconsistent relatively flat-topped features that have limited talus with the 2002 observations. This can be accounted in aprons and shallow peaks. The predominance of Mil- part by the well-documented Caribbean-wide mass die- 80 Professional Paper NMFS 5

A

Figure 5 A) Multibeam bathymetry map of Alderdice Bank, plan view. B) Multibeam bathymetry, oblique view (ve=vertical exaggeration) of basalt spires and isolated mound (bathymetry data courtesy James Gardner, see Fig. 3 for contact info.). C) A school of roughtongue bass, Pronotogrammus martinicensis, the most abundant species observed at Alderdice Bank. This small, planktivorous sea bass is one of the most common members of the deep reef fish community, and a common prey species of groupers and snappers. Photograph courtesy FGBNMS/NURC-UNCW.

off of D. antillarum in the early 1980s (Lessios et al., rable biological assemblages as reported by Rezak et 1984). This die-off was reported at the Flower Garden al. (1985). Rezak et al. (1985) report encountering an Banks NMS in 1984 (Gittings and Hickerson, 1998). exceptionally large number of yellowtail reeffish (Chro- Spiny lobster observations were rare in 2002, with a total mis enchrysura) on the basalt peak—this is consistent of three individuals observed on all three banks during with observations made on the eastern algal/sponge submersible dives. mound in 2002 Observations of biological communities at Alderdice The domination of planktivores in the deep reef fish Bank were very similar to those reported in the earlier community is both a regional and worldwide pattern studies. Both basalt spires appear abruptly out of the (Thresher and Colin, 1986; Hamner et al., 1988). Pat- soft bottom sediments, and the rock surfaces are clearly tengill-Semmens et al. (1997) reported planktivores to visible. Rezak and Tieh (1984) speculated that the up- be the dominant trophic guild on Stetson Bank and lifting of the basalt is a geologically recent occurrence, the East and West Flower Garden Banks. Percentages based on the lack of growth on the basalt blocks. The of planktivores were lower in their study (36 to 73%), 2002 observations of the basalt spires gave no indication reflecting the shallower depths surveyed by SCUBA, lim- that growth on the blocks had increased significantly in ited to the shallow crests of each bank. Deep-water reef the 25 years since the last observations. Additional dives fishes at Enewetak, Marshall Islands (Thresher and Colin, on the eastern algal/sponge mound revealed compa- 1986), became dominated by planktivores and piscivores Weaver et al: Reef fish surveys by submersible in the Northwestern Gulf of Mexico 81

B

C

Figure 5 (continued)

with increasing depths between 90 and 120 m, and were planktivorous damselfishes (Chromis spp.). Planktivores numerically dominated by anthiine serranids (streamer abundant on the shallower peaks of Sonnier Bank, T. basses), and the pomacentrids (Chromis spp.). The results bifasciata, C. multilineata, and S. partitus, decreased in of our study directly parallel this pattern, as planktivores abundance at Alderdice Bank, and were replaced by P. became increasingly abundant on the deeper reef site martinicensis and A. tenuis, two species that dominate oth- (Alderdice), and were also numerically dominated by er deepwater areas of the Flower Garden Banks National anthiine serranids (A. tenuis and P. martinicensis) and Marine Sanctuary (Dennis and Bright, 1988a). 82 Professional Paper NMFS 5

Dennis and Bright (1988a) reported three distinct under discussion were identified as areas warranting reef fish assemblages based on cluster analysis, a Mille- regulatory action through MMS, in regards to oil and pora-sponge community associated with the crest of mid- gas development. These areas are potential ecological shelf banks, an algal nodule-sponge fish assemblage, and biological significance, harboring juvenile popula- and a drowned reef fish assemblage. These categories tions of grouper, recovering populations of the long- are the dominant habitats characterizing Sonnier, Mc- spine urchins, and hermatypic deep coral reefs. These Grail, and Alderdice Banks, respectively, and results of reefs and banks have also been adopted by the Gulf this study parallel those observed by these authors on of Mexico Fisheries Management Council as Habitat previous submersible dives. While Dennis and Bright Areas of Particular Concern (HAPC). No regulations (1988a) did not provide an extensive species list for are currently in place to protect Sonnier and Alderice each of the banks surveyed, their observations of reef from over-fishing, anchoring, or excavation. Designa- fish/habitat associations remain valid. Increased resolu- tion of McGrail Bank as a Coral HAPC zone prohibits tion of video cameras, including zoom capabilities and trawling, bottom longlines, deployment of fish traps, clarity of digital videotape, allows for identification of and bottom anchoring. a larger component of the reef fish fauna, including The availability of accurate, high-resolution bathy- smaller cryptic species such as juvenile anthiines, blen- metric base maps has allowed the development of nies, and gobies. Many of the species observed here, comprehensive datasets that can be integrated via a including the roughtongue bass (P. martinicensis), have GIS database. The FGBNMS multidisciplinary effort to only recently been reported to occur in the NWGOM, characterize the reefs and banks of the NWGOM con- and the threadnose bass (A. tenuis) has not previously tinues to benefit through partnerships such as SSE, and been reported to occur in our area (Dennis and Bright, through the advancement of the available technology. 1988a, b). Recent deepwater surveys at the Flower Gar- While our studies were exploratory in nature, the basic den Banks NMS have indicated that it is one of the most patterns of habitat distribution and reef fish assem- abundant species in the deep reef community between blages will allow more rigorous, quantitative surveys of 50 and 100 m. reef fish densities and habitat association during future Rezak et al. (1990) comment on the similarity of geol- studies. Information on habitat distribution and associ- ogy and biological communities of Sonnier and Stetson ated biological assemblages is also being used to revise Banks. Submersible observations conducted during the current benthic habitat scheme for the reefs and 2002 SSE surveys support this observation, and provides banks of the NWGOM, including the Flower Garden data on habitat distribution and fish community struc- Banks National Marine Sanctuary. The data acquired ture to compare to ongoing surveys being conducted will be used to make well-informed management deci- by FGBNMS staff at Stetson Bank. Dennis and Bright sions regarding the diverse biological resources of the (1988a) reported the unique reef fish assemblage as- Northwestern Gulf of Mexico. sociated with the bank crests of the Millepora sponge zone. On the Pacific Coast, Yoklavich et al. (2000) found higher numbers of rockfishes associated with abrupt, Acknowledgments steep sided rock features surrounded by mud and sand habitats. Both Sonnier Banks and Alderdice Bank are The authors would like to particularly thank Sylvia Earle characterized by steep sided rocky features surrounded for the opportunity to be part of the SSE missions, to by relatively flat soft bottom communities. Bohnsack train as pilots, and the use of submersible during the (1989) commented on the behavioral attraction that research cruises. The SSE program was funded by grants many fishes have for high profile rocky structure, and from the Richard and Rhoda Goldman Foundation and that as a result they aggregate in large numbers. High NOAA. We also thank the NGS assistants and Nuytco numbers of reef fishes, both small and large species, Research Ltd. personnel for logistical and technical were observed at the crests of all three banks. support during cruises in the Northwestern Gulf of In general, the observations made between the his- Mexico. We thank the captain and crew of the M/V torical surveys and the 2002 SSE mission are similar. OCEAN PROJECT for hospitality and support during The fish assemblages in 2002 appear to be similar the extended research cruise. Multibeam surveys were to those observed in earlier surveys, although large funded by NOAA’s Office of Ocean Exploration and the schools of yellow that were documented Minerals Management Service (MMS). We acknowledge in the 2002 surveys were not listed as representa- the dedication and cooperation of James Gardner of tive assemblages in the earlier surveys at any of the UNH, in his efforts to expediate the extensive multi- shelf edge features. Variations are noted, but do not beam surveys prior to our submersible mission and to indicate deterioration of habitat or major shift in accelerate the processing of the data for use during our associated biological communities. The three banks research cruises. Weaver et al: Reef fish surveys by submersible in the Northwestern Gulf of Mexico 83

Literature cited In Paleoalgology: Contemporary Research and Applications (D.F. Toomey and M. H. Nitecki, eds.), p. 237–246. Springer- Bohnsack, J. A. Verlag, Berlin. 1989. Are the high densities of fishes at artificial reefs the result Pattengill-Semmens, B. X. Semmens, and S. R. Gittings. of habitat limitation or behavioral preference? Bull. Mar. Sci. 1997. Reef fish trophic structure at the Flower Gardens and Stet- 44:631–645. son Bank, NW Gulf of Mexico. Proc. 8th Int. Coral Reef Symp. Bullock, L. H., and G. B. Smith. 1:1023–1028. 1991. Seabasses (Pisces:). Memoirs of the Hourglass Randall, J. E. cruise. Florida Marine Research Institute. Dept. Nat. Res., St. 1967. Food habits of reef fishes of the West Indies. Stud. Trop. Petersburg, FL. Vol. 8, 243 p. Ocean. 5:665–847. Dennis, G. D., and T. J. Bright. Rezak, R., and T. T. Tieh. 1988a. Reef fish assemblages on hard banks in the northwestern 1984. Basalt from Louisiana Continental Shelf. Geo-Marine Let- Gulf of Mexico. Bull. Mar. Sci. 43(2): 280–307. ters 4:69–76. 1988b. New records of fishes in the northwestern Gulf of Mex- Rezak, R., T. J. Bright, and D.W. McGrail. ico, with notes on some rare species. Northeast Gulf Sci. 1985. Reefs and banks of the Northwestern Gulf of Mexico: Their 10(1):1–18. geological, biological, and physical dynamics. John Wiley and Gardner, J. V., L. A. Mayer, J. E. Hughes Clarke, and A. Kleiner. Sons, New York, 259 p. 1998. High-resolution multibeam bathymetry of East and West Rezak, R., S. R. Gittings, and T. R. Bright. Flower Gardens and Stetson Banks, Gulf of Mexico. Gulf of 1990. Biotic assemblages and ecological controls on reefs and Mexico Sci. 16(2):131–143. banks of the northwest Gulf of Mexico. Amer. Zool. 30:23–35. Gittings, S. R., and E. L. Hickerson. Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, 1998. Introduction, Flower Garden Banks National Marine Sanc- R. N. Lea, and W. B. Scott. tuary dedicated issue. Gulf of Mexico Sci. 16(2):128–130. 1991. Common and scientific names of fishes from the United Hamner, W. M., M. S. Jones, J. H. Carleton, I. R. Hauri, and D. McB. States and Canada (5th ed.). Am. . Soc. Spec. Publ. 20, Williams. 183 p. 1988. Zooplankton, piscivorous fish, and water currents on a Smith-Vaniz, W. F., B. B. Collette, and B. E. Luckhurst. windward reef face: Great Barrier Reef, Australia. Bull. Mar. 1999. The Fishes of Bermuda. Amer. Soc. Ichthy. Herpet. Spec. Sci. 42(3):459–479. Publ. 4, 424 p. Humann, P., and N. DeLoach. Thresher, R. E., and P. L. Colin. 2002. Reef fish identification (3rd ed.). New World Publications, 1986. Trophic structure, diversity and abundance of fishes of the Jacksonville, FL. 481 p. deep reef (30–300m) at Enewetak, Marshall Islands. Bull. Mar. Lessios, H. A., D. R. Robertson, and J. D. Cubit. Sci. 38:253–272. 1984. Spread of Diadema mass mortality through the Caribbean. Yoklavich, M. M., H. G. Greene, G. M. Caillet, D. E. Sullivan, R. N. Science 226:335–337. Lea, and M. S. Love. Minnery, G. A., R. Rezak, and T. J. Bright. 2000. Habitat association of deep-water rockfishes in a sub- 1985. Depth zonation and growth form of crustose coralline marine canyon: an example of a natural refuge. Fish. Bull. algae: Flower Garden Banks, Northwestern Gulf of Mexico. 98:625–641. 84 Professional Paper NMFS 5 continued Alderdice Bank McGrail Bank Appendix A Appendix Sonnier Banks Rank No. Percent Rank No. Percent Rank No. Percent 1 EB EB 27 PL 52 PL 19 PI 5 57 60 BC 0.25 BC 3 0.06 — 2 13 36 PL — — 0.04 — 0.03 80 9 EB — — 54 2 — PI — 1.04 PI 0.06 — 59 PL — 3 — — PI — 27 53 PI 31 3 — PI — 44 — 0.02 54 PI — 19 — 5 — 8 — 0.04 34 29 6 4 — — PI — — — 0.12 40 — 0.06 13 0.05 PI — 18 — — — 0.04 0.05 PI 11 17 28 — — — — — 0.17 — 0.12 17 50 PL — — 25 — — 10 0.14 — EB — — 19 14 EB 38 — — 6 — 0.14 — 39 0.06 — 30 BC — 30 61 81 — — 0.25 21 — 0.08 — 17 — 8 — 77 0.20 2 34 0.53 25 — 39 — — 59 0.11 0.05 — — 11 1 64 0.03 — — — 0.32 4 — 2 — — — — 0.07 — 0.01 1 — 66 13 — 0.02 0.01 — — — — 22 0.01 — 1 — 84 — 58 16 — — — — — 0.01 0.55 1 0.09 — 18 60 — 14 0.01 18 49 — 1 — 0.10 0.28 — 0.01 — GC PI 56 44 4 10 0.05 0.13 — 35 — 10 — 0.07 — 12 — 53 0.30 — guild Trophic Fishes observed during SSE2002 Submersible Dives at Sonnier, McGrail, and Alderdice Banks. Fishes observed during SSE2002 Submersible Dives at Sonnier, bigtooth cardinalfish PL queen triggerfish — whitespotted filefish — EB — African pompano — blue runner — crevalle jack — horse-eye jack — black jack bar jack — — almaco jack 38 13 8 50 bank butterflyfish 0.05 0.29 unidentified blenny 56 1 0.01 lookdown greater amberjack Common name

elene vomer eriola dumerili Acanthurus coeruleus Apogon affinis blue tang Apogon maculatus Apogon psuedomaculatus twospot cardinalfish Aulostomus maculatus flamefish Balistes capriscus trumpetfish Balistes vetula Cantherhines macrocerus Cantherhines pullus gray triggerfish Canthidermis sufflamen Xanthichtys ringens ocean triggerfish orangespotted filefish Parablennius marmoreus sargassum triggerfish seaweed blenny Alectis ciliaris EB Caranx bartholomaei Caranx chrysos PL Caranx hippos yellow jack — Caranx latus Caranx lugubris Caranx ruber 42 — S S Seriola rivoliana 7 — plumbeus Carcharhinus sp. Carcharhinus 0.05 sandbar shark — Chaetodon aculeatus Chaetodon aya — Chaetodon ocellatus — longsnout butterflyfish Chaetodon sedentarius — Starksia sp. spotfin butterflyfish reef butterflyfish EB — — — 55 — — 1 — 0.01 — — 45 6 0.04 59 1 0.10 Acanthuridae Acanthurus chirurgus Apogonidae doctorfish Aulostomidae Balistidae Blenniidae Carangidae Carcharhinidae Chaetodontidae Clinidae Species Family Weaver et al: Reef fish surveys by submersible in the Northwestern Gulf of Mexico 85 continued Alderdice Bank McGrail Bank Sonnier Banks 9 181 2.35 5 442 15 5.73 74 0.48 18 35 15 0.23 34 0.19 25 12 0.07 Appendix A (continued) Appendix Rank No. Percent Rank No. Percent Rank No. Percent 1 GC GC — GC — BC — BC 69 BC — PL 63 — 1 32 GC 47 — 2 22 16 — 0.01 PL 7 6 PL 22 — 0.03 PL — 0.21 — 428 43 0.09 — 0.29 45 49 — 20 — — 5.55 10 68 GC 9 — — GC 5 31 — PL 61 0.13 — — BC 1 16 — 0.12 BC 62 0.03 0.40 17 BC — 1 — 46 — BC 51 0.01 — BC 45 22 1 — 28 — 28 PL 0.01 — 41 — 6 — — 0.66 — — 28 GC 19 0.58 0.01 — 19 — — GC 11 — PI 0.04 — — — — 21 GC — 0.18 — 0.25 — 18 24 0.12 GC — 0.14 15 GC — — 49 28 — — PL 43 — 65 22 — 14 — 33 — PL 70 — 48 — 55 2 — 0.18 — 80 — 0.56 37 — 1 0.14 72 8 0.91 56 — 7 12 4 0.01 11 1.04 — 9 0.01 — 1 54 110 — 0.05 — 3 — 0.09 — 73 0.05 — 71 0.06 0.01 1 9 1.43 45 — 26 0.02 26 — 0.42 — 112 1 47 0.01 73 — 11 — 2 20 — — — — 0.73 0.01 2 — 1 0.06 41 0.01 — 0.13 29 — — 10 0.01 — 0.01 3 — 30 10 — 91 — — 0.02 — 10 0.06 6 0.52 — — — 576 0.06 — 3.29 — guild Trophic conger eel unidentified gobies Bermuda/yellow chub Spanish hogfish PL creole wrasse 11 139 1.80 gray snapper — dog snapper — — — — — Common name

Conger oceanicus Dasyatis americana Echeneis naucrates southern stingray Gnatholepis thompsoni sharksucker Gobiosoma xanthiprora goldspot goby calliurus Ptereleotris prownose goby Haemulon aurolineatum blue goby tomtate Corniger spinosus Holocentrus adscensionis Holocentrus rufus squirrelfish Myripristis jacobus spinycheek soldierfish bullisi Sargocentron longfin squirrelfish Kyphosus sp. blackbar soldierfish GC deepwater squirrelfish Bodianus pulchellus Bodianus rufus Clepticus parrae PL — spotfin hogfish Decodon puellaris bathyphilus Halichoeres bivitattus Halichoeres — garnoti 66 greenband wrasse Halichoeres red hogfish maculipinna Halichoeres slippery dick Thallasoma bifasciatum clown wrasse — 2 yellowhead wrasse bluehead Lutjanus buccanella Lutjanus campechanus Lutjanus cyanopterus 0.03 — blackfin snapper Lutjanus griseus red snapper Lutjanus jocu cubera snapper Lutjanus synagris — 75 Ocyurus chrysurus Rhomboplites aurorubens vermilion snapper lane snapper 1 — yellowtail snapper 0.01 44 38 2 5 0.01 0.03 Congridae Dasyatidae Echeneidae Gobiidae Gobiidae Haemulidae Holocentridae Kyphosidae Labridae Lutjanidae Species Family 86 Professional Paper NMFS 5 continued Alderdice Bank McGrail Bank Sonnier Banks 1 4 1782 3 7 491 1055 23.11 412 8 6.37 13.68 216 5 5.34 12 4 947 2.80 10 1168 90 6.20 106 62 7.64 0.59 0.69 2 2 5 — 4311 9 0.01 993 — 24.61 170 — 5.67 — 0.97 — — Appendix A (continued) Appendix Rank No. Percent Rank No. Percent Rank No. Percent 1 BC BC BC — GC 26 — 39 GC 20 64 PL 12 — — BC 0.26 2 0.16 — EB — 57 EB 58 0.03 EB 3 — 31 EB — 3 EB — 2000 29 3 69 35 15 PL — — 25 PL 0.02 13.09 18 — 30 0.04 PL 13 1 0.10 PL 20 62 19 PL — 18 66 0.23 38 7 1 PL 70 0.17 0.01 PL 21 41 0.26 2 280 12 0.23 11 1 40 EB 1 — 52 — 16 0.53 10 25 PL 0.01 1.60 96 0.16 0.01 53 8 0.01 — 157 — GC 4 0.09 22 — — 74 0.63 GC 4 8 0.05 — — 2.04 0.03 65 — — EB 0.14 — 1 143 34 0.03 70 24 — EB 2 — 48 7 23 68 — EB 0.01 0.94 — 6 13 1 — 159 67 — 0.03 2 14 1 — 40 0.03 — — 49 0.07 0.01 1.04 2 0.01 — 0.08 — 0.01 — 3 7 — — — 0.03 — — 0.02 — 0.09 — — — — — — 51 27 — — — — — 11 5 64 — — — 0.06 0.03 1 — — — — 0.01 — — — — — guild Trophic pygmy angelfish blue chromis sunshinefish purple reeffish bicolor damselfish unidentified parrotfish EB 75 1 0.01 — — — — — — Common name

Mulloidichthys martinicus Pseudupeneus maculatus yellow goatfish spotted goatfish Gymnothorax moringa Myrichthys breviceps spotted moray Opistognathus aurifrons sharptail eel yellowhead jawfish Lactophrys triqueter argi Centropyge Holacanthus bermudensis smooth trunkfish Holacanthus ciliaris blue angelfish Holacanthus tricolor Pomacanthus paru queen angelfish rock beauty Abudefduf saxatalis cyanea Chromis French angelfish enchrysuraChromis insolata Chromis sargeant major multilineatus Chromis scotti Chromis yellowtail reeffish Stegastes partitus brown chromis Stegastes planifrons Stegastes variabilis Priacanthus arenatus threespot damselfish cocoa damselfish canadum Rachycentron bigeye Ginglymostoma cirratum cobia EB nurse shark Cryptotomus roseus 31 Scarus taeniopterus Sparisoma atomarium bluelip parrotfish Sparisoma aurofrenatum 18 princess parrotfish greenblotch parrotfish redband parrotfish 0.23 EB — 42 — 11 — 0.14 — 23 — 23 0.15 — 42 3 0.02 Malacanthidae Malacanthus plumieri Mullidae sand tilefish Muraenidae Ophichthidae Opistognathidae Ostraciidae Pomacanthidae Pomacentridae Priacanthidae Rachycentridae Rhincodontidae Scaridae Scaridae Species Family Weaver et al: Reef fish surveys by submersible in the Northwestern Gulf of Mexico 87 69 17,519 Alderdice Bank No. Sp. N Total McGrail Bank 79 15,288 No. Sp. N Total Sonnier Banks 77 7,717 No. Sp. N Total 2 1370 17.77 1 6625 43.35 3 3239 18.49 Appendix A (continued) Appendix Rank No. Percent Rank No. Percent Rank No. Percent 1 BC BC BC 62 PI — 36 2 GC — — GC 13 0.03 GC 37 — GC — 0.17 — GC GC 12 — 23 PI — — — — — — — PI 21 0.16 71 PI — — — — — — — PL 0.27 1 48 — — — — 46 — PL — 65 — — — 67 0.01 5 GC — 9 GC 55 37 — — 1 19 — — 1 — BC 0.03 — — 76 BC 0.12 — 17 51 3 5 33 0.01 8 — — 0.01 — 1 — — — 61 200 11 6 0.10 73 0.03 0.02 — PI — PI — — — — — 0.01 — 1.14 1 2 0.07 0.08 — — EB — 6 20 1 — — 36 76 — — 0.01 0.01 — 50 77 220 — — — 16 — 0.01 5 1 47 51 52 — 2 1 — 1.44 — 0.09 — 0.03 — 0.01 5 2 — 0.01 2 0.01 — — — 68 79 0.03 0.01 0.01 — 53 — — — 1 1 43 2 — — — — 0.01 0.01 2 — 0.01 — 69 0.01 — — 57 1 — 1 0.01 — 0.01 Trophic spotted drum blackbar drum cubbyu threadnose bass coney Marble grouper PL GC wrasse bass — black grouper — — scamp — creole-fish — school bass orangeback bass — saddle bass 2 tattler GC 2745 41 red porgy 17.96 great barracuda — 7 sand diver — red lizardfish 4 0.05 PI 2300 sharpnose puffer — 46 — 13.13 EB 32 2 — 15 24 0.01 — 20 0.10 78 0.26 34 1 59 6 0.01 2 0.03 — 0.01 — 32 — 8 0.05 Common name guild

Pareques iwamotoi Pareques umbrosus Pareques cavalla Scomberomorus Anthias tenuis cruentata king mackerel Cephalopholis fulva Dermatolepis inermis Epinephelus adscensionis Epinephelus guttatus Gonioplectrus hispanus eukrines Liopropoma rock hind bonaci Mycteroperca interstitialis Mycteroperca Spanish flag microlepis Mycteroperca phenax Mycteroperca venenosa Mycteroperca yellowmouth grouper Paranthias furcifer gag PI martinicensisPronotogrammus Schultzea beta roughtongue bass yellowfin grouper Serranus annularis Serranus notospilus 33 Serranus phoebe PL PI Calamus sp. 14 — — Sphyraena barracuda 0.18 — Synodus intermedius — Synodus synodus 20 Canthigaster jamestyleri — Canthigaster rostrata — 30 goldface toby 16 50 0.20 74 5 16 0.48 29 0.03 1 0.17 — 4984 — 28.45 — PL-Planktivore, PI-Piscivore, PL-Planktivore, GC-General carnivore (mobile invertebrates and fishes), H-Herbivore, EB-Epibenthic browsers (sessile BC-Benthic invertebrates), carnivores (benthic crustaceans and infauna).

Sciaenidae punctatus Scombridae Serranidae Sparidae Pagrus pagrus Sphyraenidae Synodontidae Tetraodontidae 1 Species Family