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Fish Assemblages Associated with Red Grouper Pits at Pulley Ridge, A

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Fish Assemblages Associated with Red Grouper Pits at Pulley Ridge, A 419 Abstract—Red grouper (Epineph- elus morio) modify their habitat by Fish assemblages associated with red grouper excavating sediment to expose rocky pits, providing structurally complex pits at Pulley Ridge, a mesophotic reef in the habitat for many fish species. Sur- Gulf of Mexico veys conducted with remotely op- erated vehicles from 2012 through 2015 were used to characterize fish Stacey L. Harter (contact author)1 assemblages associated with grouper Heather Moe1 pits at Pulley Ridge, a mesophotic 2 coral ecosystem and habitat area John K. Reed of particular concern in the Gulf Andrew W. David1 of Mexico, and to examine whether invasive species of lionfish (Pterois Email address for contact author: [email protected] spp.) have had an effect on these as- semblages. Overall, 208 grouper pits 1 Southeast Fisheries Science Center were examined, and 66 fish species National Marine Fisheries Service, NOAA were associated with them. Fish as- 3500 Delwood Beach Road semblages were compared by using Panama City, Florida 32408 several factors but were considered 2 Harbor Branch Oceanographic Institute to be significantly different only on Florida Atlantic University the basis of the presence or absence 5600 U.S. 1 North of predator species in their pit (no Fort Pierce, Florida 34946 predators, lionfish only, red grou- per only, or both lionfish and red grouper). The data do not indicate a negative effect from lionfish. Abun- dances of most species were higher in grouper pits that had lionfish, and species diversity was higher in grouper pits with a predator (lion- The red grouper (Epinephelus morio) waters (>70 m) of the shelf edge and fish, red grouper, or both). These re- has been harvested in the United have been known to modify their sults may indicate that grouper pits States since the 1880s and is cur- habitat by excavating sediment to are a favorable habitat for both lion- rently the most common grouper spe- expose rocky depressions (or pits) on fish and native fish species or that cies landed in both commercial and the seafloor in areas where vertical the presence of lionfish is too recent recreational fisheries of the Gulf of relief is not already present (Cole- to have caused changes to fish com- Mexico (Fisheries Statistics Division, man et al., 2010). Red grouper use munity structure. National Marine Fisheries Service, these pits as their home territories Annual Commercial Landing Statis- (Scanlon et al., 2005). These exca- tics, website, and Saltwater Recre- vations increase the architectural ational Data and Statistics, website). complexity of the habitat, attract- Like other grouper species, the red ing many reef-associated species and grouper is a slow growing, late ma- providing shelter for juveniles of turing, relatively stationary, and long some economically important species lived fish (Moe1; Jory and Iversen2). and, thereby, increasing biodiversity Adult red grouper inhabit the deeper (Coleman et al., 2010). By excavat- ing the sediments, red grouper act as 1 Moe, M. A., Jr. 1969. Biology of the “ecological engineers” and may play Manuscript submitted 15 August 2016. red grouper Epinephelus morio (Valen- an important role in influencing com- Manuscript accepted 2 June 2017. ciennes) from the eastern Gulf of Mex- munity dynamics (Jones et al., 1994; Fish. Bull. 115:419–432 (2017). ico. Fla. Dep. Nat. Resour. Mar. Res. Coleman and Williams, 2002; Cole- Online publication date: 23 June 2017. Lab. Prof. Pap. Ser. 10, 95 p. 2 man et al., 2010). doi: 10.7755/FB.115.3.11 Jory, D. E., and E. S. Iversen. 1989. Species profiles: life histories and en- Grouper pits have been described vironmental requirements of coastal for 2 marine protected areas (MPAs) The views and opinions expressed or fishes and invertebrates (South Flori- implied in this article are those of the in the northeastern Gulf of Mexico: da)—black, red, and Nassau groupers, Madison-Swanson and Steamboat author (or authors) and do not necessarily 21 p. U.S. Fish Wildl. Serv. Biol. Rep. reflect the position of the National 82(11.110). U.S. Army Corps Eng., TR Lumps MPAs, which were estab- Marine Fisheries Service, NOAA. EL-82-4. [Available from website.] lished in 2000 on the shelf break (at 420 Fishery Bulletin 115(3) depths of 50–120 m) to alleviate fishing pressure on tively influence abundance and diversity of fish spe- aggregations of gag (Mycteroperca microlepis). Grouper cies (McClanahan, 1994; Öhman and Rajasuriya, 1998; pits at Steamboat Lumps consist of carbonate rocks Gratwicke and Speight, 2005; Harter et al., 2009). Most that have been scoured out from a thick veneer of of Pulley Ridge is a low relief, low rugosity, rock and carbonate-derived sediment. The pits are on average rubble habitat. The highest densities of fish reside on 5–6 m in diameter but can become as large as 25 m in 2 biologically derived habitat features that provide diameter and 1–2 m deep with a density of about 250 more structural complexity: mounds of rock rubble and pits/km2 (Scanlon et al., 2005; Coleman et al., 2010; cobble created by sand tilefish (Malacanthus plumieri) Wall et al., 2011). Pits in the Madison-Swanson MPA and grouper pits (Halley et al.7). The grouper pits are differ in their level of relief; some have a thin veneer large enough (8–15 m in diameter and 1–2 m deep) to of carbonate-derived sediments and some have more be visible in high-resolution bathymetric images col- relief (Coleman et al., 2010). lected with a multibeam sonar during a research cruise Our study area was at Pulley Ridge, a mesophotic in September 2011 (NOAA8), and up to 340 pits/km2 coral ecosystem (MCE) in the northeastern Gulf of are visible in those images. Approximately 90 species Mexico, which has large populations of red grouper and of fish, both shallow-water and deepwater species, have a high density of grouper pits. This geological feature been observed on Pulley Ridge (Jaap et al., 2015). The is a carbonate ridge that extends for nearly 300 km fish communities of Pulley Ridge have been character- along the southwestern shelf of Florida and lies about ized previously, but communities associated with the 250 km west of the Florida coastline (Hine et al., 2008). grouper pits specifically have not. The southern end of Pulley Ridge supports an MCE at Species of lionfish (Pterois spp.) first were discov- depths of 60–100 m and covers an area of ~600 km2 ered on Pulley Ridge during submersible dives in 2010 (Fig. 1; Jarrett et al., 2005; Cross et al.3; USGS4; Hine (Reed and Rogers9) when 6 fish were observed. Since et al., 2008; Reed, 2016). Mesophotic coral ecosystems then, exponential increases in the abundance of this are characterized by the presence of light-dependent population of lionfish have been observed (Andradi- corals and their associated communities, and Pulley Brown et al., 2016). It is unknown at this time wheth- Ridge is the deepest MCE on the U.S. continental shelf er red lionfish P.( volitans), devil firefishP. ( miles), or (Halley et al.5). The hard-bottom substrate along this both species exist at Pulley Ridge. At this time, posi- ridge consists of rock and coral pavements and cement- tive identification can be achieved only through genet- ed conglomerates of carbonate rubble and cobble rock ic analysis; therefore, lionfish were identified only to (5–15 cm in diameter) that provide habitat for herma- genus level for this study. The invasion of lionfish is typic corals (primarily Agaricia spp., great star coral regarded as one of the most successful colonizations of [Montastraea cavernosa], and Madracis spp.), macroal- a marine species ever documented (Albins and Hixon, gae, sponges, and a large variety of species of tropical 2008; Green and Côte, 2009; Albins, 2013). Lionfish fish (Phillips et al., 1990; Halley et al.5; Reed, 2016). first were recorded in waters of the Atlantic Ocean in The Pulley Ridge Habitat Area of Particular Concern the mid-1980s, but their range has expanded to include (HAPC) was established in 2005 by the Gulf of Mexico the Gulf of Mexico, Caribbean, and much of the tropi- Fishery Management Council6 to provide protection cal and subtropical western Atlantic Ocean (Schofield, from bottom longlines and trawls. Hook-and-line fish- 2009, 2010). Over the years, densities of lionfish in the ing, however, is still allowed. The established HAPC western Atlantic Ocean have expanded rapidly to the provides protection for 348 km2, an area that is rough- point that they are nearly 5 times more abundant in ly half the total MCE area (Reed, 2016). the invaded range (Green and Côte, 2009) than in the Structural complexity has often been shown to posi- Pacific Ocean (Kulbicki et al., 2012). High individual growth and reproductive rates (Morris and Whitfield, 2009) have contributed to the rapid increase of the 3 Cross, V., D. C. Twichell, R. B. Halley, K. T. Ciembronowicz, B. D. Jarrett, E. S. Hammar-Klose, A. C. Hine, S. D. Locker, and population in the western Atlantic Ocean. Many stud- D. F. Naar. 2005. GIS compilation of data collected from the Pulley Ridge deep coral reef region. U.S. Geol. Surv. Open-File Rep. 2005-1089. [Available from website.] 7 Halley, R., G. P. Dennis, D. Weaver, and F. Coleman. Halley, 4 USGS (U.S. Geological Survey). 2005. Recently discovered R., G. P. Dennis, D. Weaver, and F. Coleman. 2005. Part reef is deepest known off continental U.S. ScienceDaily, 5 II: characterization of the Pulley Ridge coral and fish fau- January 2005. [Internet press release available from web- na. Final report to the National Oceanic and Atmospheric site.] Administration Coral Reef Conservation Grant Program, 25 5 Halley, R.
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