development of invertebrate assemblages on artificial ...

DEVELOPMENT OF INVERTEBRATE ASSEMBLAGES ON ARTIFICIAL REEF CONES OFF SOUTH CAROLINA: COMPARISON TO AN ADJACENT HARD-BOTTOM HABITAT A thesis submitted in partial fulfillment of the requirements for the degree MASTER OF SCIENCE in ENVIRONMENTAL STUDIES by DANY E. BURGESS MAY 2008 at THE GRADUATE SCHOOL AT THE COLLEGE OF CHARLESTON

Approved by: Dr. Elizabeth Wenner, Thesis Advisor Dr. Jeff Hyland Dr. Rachael King Mr. Robert Martore Dr. Amy T. McCandless, Dean of Graduate Studies

UMI Number: 1450538

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ABSTRACT

DEVELOPMENT OF INVERTEBRATE ASSEMBLAGES ON ARTIFICIAL REEF CONES OFF SOUTH CAROLINA: COMPARISON TO AN ADJACENT HARD-BOTTOM HABITAT A thesis submitted in partial fulfillment of the requirements for the degree MASTER OF SCIENCE in ENVIRONMENTAL STUDIES by DANY E. BURGESS MAY 2008 at

THE GRADUATE SCHOOL OF THE COLLEGE OF CHARLESTON Artificial reefs are often used to increase the amount of hard-bottom habitat in otherwise sandy areas, including parts of South Carolina’s continental shelf. In 1997 and 2003, the SCDNR deployed two designed concrete reefs off the coast of Charleston, SC, for use in fishing experiments. This study was conducted to assess the development of epifaunal invertebrate assemblages on both the younger (“Area 53” – 2 years old) and older (“Area 51” – 8 years old) reefs. Each artificial reef was also compared to an adjacent natural reef, “Julian’s Ledge”, in an attempt to determine whether designed structures can form habitats that resemble natural hard-bottom areas over time. Macrofaunal invertebrates from each of the three reef sites were collected during Spring/Summer 2005. A total of 24,940 individuals were found, comprising at least 384 motile and sessile species. Cluster analysis revealed that species composition between reef sites was distinct, with Julian’s Ledge displaying higher species number and diversity; however, evidence for convergence over time included a large group of species common to all three sites, and a higher level of similarity between Julian’s Ledge and Area 51 than between Julian’s Ledge and Area 53. Additional sampling at a later time period could help to elucidate whether these trends may be attributed to reef age, or other environmental variables. This study provided the first catalogue of invertebrate data for any of South Carolina’s designed experimental artificial reefs.

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ACKNOWLEGDEMENTS

First and foremost, I would like to thank my committee members, Elizabeth Wenner (advisor), Rachael King, Jeff Hyland, and Bob Martore for their time, guidance, and patience throughout the course of this project. Special thanks to Bob Martore and the Artificial Reef Program for providing financial support for as long as I needed it; to the staff of the SERTC lab for generously sharing their workspace and supplies; to Jessica Boynton for GIS assistance; and to Mike Arendt and Stephen Blackmore for database management and moral support. In addition, I would like to thank the talented scientists and fellow graduate students who dedicated many uncompensated hours to helping me collect and identify thousands of critters: David Knott, Susan DeVictor, Dale Calder, Dave Pawson, Richard Heard, Joe Cowan, Nadia Meyers, Ian Moody, Daryl Stubbs, Bryan Frasier, Chris Bradshaw, and especially Rachael King. Thanks also to the crews of the R/V Palmetto and the R/V Silver Crescent for providing field transportation, to the MARMAP program for the use of their electronic balance, and to the infinitely helpful and accommodating faculty and staff of the MES program. Finally, I would like to dedicate the completion of this thesis to my family, particularly my parents, David and Kathy Burgess, who have always believed that I could be whatever I wanted; to my sister, who never stopped speaking to me even when I truly deserved it; also to my amazing grandparents, Cynthia and Joseph Miller, who have been dragged to more ceremonies and backyard tea parties than they probably care to count, and always kept smiling. Their support over the years, both emotional and financial, has kept me on the path to achieving my dreams. Their lessons – mainly to respect and care for the environment and all its creatures – had a hand in shaping those dreams. I love you guys. Thank you all so much for sharing this experience with me. I would not have been able to do it without you.

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TABLE OF CONTENTS PAGE ABSTRACT............................................................................................................ ii LIST OF FIGURES .................................................................................................v LIST OF TABLES................................................................................................ vii INTRODUCTION ...................................................................................................1 Objectives.....................................................................................................9 MATERIALS AND METHODS...........................................................................11 Study Sites ..................................................................................................11 Preliminary Site Visit.................................................................................12 Field Sampling ...........................................................................................12 Laboratory Processing of Scraped Samples ..............................................14 Analysis of Digital Photographs and Archived Video Data ......................16 Statistical Analyses – Univariate Methods ................................................17 Statistical Analyses – Multivariate Methods..............................................19 RESULTS ..............................................................................................................21 Description of Study Sites ..........................................................................21 Species Number, Species Richness, Abundance, and Biomass..................22 Diversity Indices ........................................................................................23 Species Composition – Artificial Reefs ......................................................23 Species Composition – Natural Reef..........................................................25 Similarity Indices .......................................................................................26 DISCUSSION ........................................................................................................30 Community Composition............................................................................30 Community Indices.....................................................................................33 Age of Artificial Reefs ................................................................................35 Influence of Other Factors.........................................................................38 Implications for Fisheries Management ....................................................39 Future Directions.......................................................................................41 CONCLUSIONS....................................................................................................43 LITERATURE CITED ..........................................................................................45 FIGURES...............................................................................................................54 TABLES ................................................................................................................80 APPENDICES .......................................................................................................95

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LIST OF FIGURES FIGURE

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1.) Map of study locations, showing approximate position of artificial and natural reefs to the Charleston peninsula.....................................55 2.) Configuration of artificial and natural reefs .....................................................56 3.) Dimensions of the Swiss Cone .........................................................................57 4.) Digital photograph of a preserved scrape sample collected from Julian’s Ledge station A during August 2005...................................................58 5.) Digital photographs of ossicles, viewed with a compound microscope, and corresponding sea cucumber specimens............................................................59 6.) Digital photographs of scrape samples collected from Julian’s Ledge stations during 2005...............................................................................60 7.) Digital photographs of scrape samples collected from Area 53 during 2005 .........................................................................................61 8.) Digital photographs of scrape samples collected from Area 51 during 2005 .........................................................................................62 9 a-b.) a.) Mean number of species (s) per 15 cm² quadrat for each reef site, and b.) total number of species in all scrape samples for each reef site............................................................................63 10.) Median species richness (SR) per 15 cm² quadrat for Area 53, Area 51, and Julian’s Ledge.............................................................64 11 a-b.) a.) Median number of individuals (n) per 15 cm² quadrat for each reef site, and b.) total number of individuals in all scrape samples for each reef site........................................................65 12.) Mean biomass per 15 cm² quadrat for Area 53, Area 51, and Julian’s Ledge .........................................................................................66 13 a-b.) Median values of a.) diversity (H´) and b.) evenness (J´) per 15 cm² quadrat for Area 53, Area 51, and Julian’s Ledge..................67 14.) Percent contribution of major taxa (≥1%) to the total abundance of Area 53 ....................................................................................68

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15.) Percent contribution of major enumerated taxa (≥1%) to the total abundance of Area 51 ....................................................................................69 16.) Percent contribution of major enumerated taxa (≥1%) to the total abundance of Julian’s Ledge............................................................................70 17.) Percent contribution of major sessile taxa to the total number of sessile species from Area 53 ......................................71 18.) Percent contribution of major sessile taxa to the total number of sessile species from Area 51 ......................................72 19.) Percent contribution of major sessile taxa to the total number of sessile species from Julian’s Ledge...............................................73 20 a-b.) Dorsal views of a.) female and b.) male specimens of Limnotheres nasutus, collected from the northeast and northwest stations of Area 53...................................................................74 21.) Normal Canberra Metric cluster dendrogram (replicates pooled by sum of species abundance) ...........................................75 22.) Inverse Canberra Metric cluster dendrogram (replicates pooled by sum of species abundance).............................................................76 23.) Normal Jaccard similarity dendrogram (replicates pooled by presence/absence).........................................................77 24.) Inverse Jaccard similarity dendrogram (replicates pooled by presence/absence)...........................................................................78 25 a-b.) a.) Correlation between abundance of Syllis and Haplosyllis sp. and diversity (H’) in samples collected from Area 51 in 2005, and b.) abundance of Syllis and Haplosyllis sp. in Area 51 collections, noting presence/absence of the sponge Lissodendoryx sp ................................................................79

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LIST OF TABLES

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1.) Collection dates and hydrologic parameter measurements for artificial and natural reef stations visited in 2005 and 2006 .......................81 2.) Community structure values per 15 cm² for stations sampled at the younger artificial reef (A53), older artificial reef (A51), and Julian’s Ledge natural reef in 2005 .........................................82 3.) Summary of statistical analyses of community structure values......................83 4.) Rank by abundance, overall (> 1%) .................................................................84 5.) Rank by abundance of the top 25 most abundant species in samples collected from Area 53 and Area 51 in 2005..................................85 6.) Rank by abundance of the top 25 most abundant species collected from Julian’s Ledge in 2005..............................................................86 7.) Overall rank by frequency of occurrence (%) of major sessile taxa occurring in 10 or more samples collected from Area 53, Area 51, and Julian’s Ledge in 2005 ...............................................................87 8.) Rank by frequency of occurrence (%) of the top 25 most frequently occurring sessile species in samples collected from Area 53 and Area 51 during 2005.....................................................................88 9.) Rank by frequency of occurrence (%) of the top 25 most frequently occurring sessile species in samples collected from Julian’s Ledge during 2005...............................................................................89 10.) Species groups resulting from inverse Canberra metric cluster analysis............................................................................................................90 11.) Species groups resulting from inverse Jaccard similarity analysis.................91 12.) Results of Components of Variance analysis..................................................94

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INTRODUCTION

In the waters of the South Atlantic Bight (defined here as the region between Cape Hatteras, North Carolina and Cape Canaveral, Florida; Struhsaker 1969), only a small percentage of the sea floor is hard-bottom habitat; in South Carolina alone, it is estimated that rocky ledges and outcroppings, which support more than 70% of the region’s offshore pelagic and reef fisheries, make up only 10-20% of the bottom substrate of the continental shelf (Van Dolah et al. 1994). These formations, originally referred to as ‘live-bottom’ by Cummins et al. (1962), are characterized by a variety of attached sessile invertebrates and diverse tropical and subtropical fish assemblages (Wenner et al. 1983). The majority of seafloor habitat in the South Atlantic Bight is a smooth, sandy substrate where productivity is driven by sand-dwelling infauna rather than by reef-associated fauna (Hyland et al. 2006). Artificial reefs composed of man-made materials have been used as fisheries management strategies in these predominantly sand sea floor habitats, increasing the amount of hard substratum that serves as a base for new reef communities. Artificial reefs were defined by the European Artificial Reef Research Network (EARRN; see Jensen 1997) as “submerged structures placed on the sea bed deliberately, to mimic some characteristics of natural reefs.” Since the deployment of the first artificial reef in Japan in the 1700’s (Ino 1974), the primary goal – to increase fishing success – has remained relatively unchanged (Ambrose and Swarbrick 1989). In the face of the recent decline of harvests for many of the world’s fisheries, reef project managers

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have begun to consider artificial reefs important tools for the enhancement of resource production. In many cases, however, an artificial reef will quickly develop a fish community with similar or greater abundance and diversity than the surrounding natural habitat (review by Bohnsack and Sutherland 1985; Bortone et al. 1994; Rilov and Benayahu 2000; Arena et al. 2007), giving rise to the debate on whether artificial reefs actually increase fish production, or merely attract and concentrate existing fish stocks so that they can be more easily exploited (Alevizon and Gorham 1989; Lindberg 1997; Pickering and Whitmarsh 1997; Bortone 1998; review by Svane and Petersen 2001). Also controversial are the impacts that physical and biological changes, imposed by the placement of made-made structures, can have on existing infaunal populations (Davis et al. 1982; Ambrose and Anderson 1990; Danovaro et al. 2002). In order to shed light on the attraction-production debate, minimize harm to the receiving environment, and continue to improve understanding of artificial reef function, researchers have suggested more thorough planning, monitoring, and experimental efforts associated with current and future reef projects (Bohnsack and Sutherland 1985; Carter et al. 1985; Bohnsack 1989; Seaman et al. 1989; Pickering and Whitmarsh 1997; Baine 2001; Miller 2002). One component of artificial reefs that has benefited from research is the type of structure that makes up the reefs. Historically, this has varied based on geographic region. In the United States, much emphasis has been placed on dumped waste or scrap materials including tires, old bridges, subway cars, and sunken vessels. Often referred to as “materials of opportunity,” mostly due to their availability and cost-effectiveness (Bell et al. 1989; Pickering et al. 1998), these types of materials may be structurally unstable or have the potential for leaching contaminants into the water. Although many of these

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types of reefs are still in use, increased funding in the past few decades, due largely to public support, has allowed for the advancement of reef-building technology, and has lead to the development of fabricated or designed reefs (Bell et al. 1989; McGurrin et al. 1989; Kellison and Sedberry 1998). These reefs integrate both biological and engineering principles by using informed design in an effort to fulfill the life history requirements of target species (Seaman and Jensen 2000). Prefabricated reef modules were introduced into the United States in the early 1980s, and are typically constructed from marine cement with a pH similar to that of seawater. The durability and structural options provided by these modules make them a more effective way to simulate the natural limestone outcroppings, which constitute much of the hard bottom habitat in the South Atlantic Bight region, without the environmental consequences associated with many types of industrial wastes and secondary-use materials (Bell et al. 1989; Fitzhardinge and Bailey-Brock 1989; AGSMFC 2004). After deployment, each new reef unit undergoes a successional process involving the formation of a bacterial film (which conditions the substrate for further colonization) and the recruitment of a variety of algal species and invertebrates that account for a large percentage of the reef’s productivity (Wahl 1989; Palmer-Zwahlen and Aseltine 1994). These fouling assemblages often include, but are not limited to, barnacles, octocorals, hydroids, bryozoans, tunicates, and sponges, along with a suite of motile fauna that utilize the food and habitat provided by stationary organisms (Wendt et al. 1989). Epifaunal communities are important components of successful artificial reefs because they enhance the stability and aesthetic appearance of reef structures, making them more suitable for recreational purposes - which, in turn, helps relieve natural habitats of these

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pressures (Leeworthy et al 2006). Large benthic suspension feeders can also modify the flux of food, larvae, and sediment deposition around a reef (Eckman 1983), and provide refuges for both predators and prey (Thrush and Dayton 2002). Invertebrates also provide food and habitat for both juvenile and adult fish, and often dictate the total biomass and types of fish assemblages that a reef can support (Hueckel and Stayton 1982; Palmer-Zwahlen and Aseltine 1994; Relini et al. 1994). The actual numbers and types of invertebrates that ultimately colonize an artificial reef surface may be dictated by many factors. Natural reefs constitute an important source of larvae and spores of epibenthic organisms; therefore, proximity to natural habitats may affect the composition of communities that subsequently settle on artificial reefs (Van Dolah et al. 1988; review by Svane and Petersen 2001). This type of recruitment is a function of large-scale hydrodynamic variations that include water depth, temperature, and current. Because recruitment is seasonal in temperate regions, the season in which a reef is deployed may also greatly affect the number and types of larvae that colonize it (review by Svane and Petersen 2001). Researchers have cited composition and texture of the substratum, habitat complexity and stability, and water clarity as additional reasons why invertebrate communities on artificial reefs may develop differently than their natural counterparts (Pamintuan et al 1994; Connell and Glasby 1999; Badalamenti et al. 2002; Perkol-Finkel and Benayahu 2005). Orientation of the substrate has proven to be another important factor, as vertical surfaces tend to show greater species abundance, percent cover, and biomass than horizontal surfaces (Baynes and Szmant 1989; Wendt et al. 1989; Ponti et al. 2002; Knott et al. 2004).

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For artificial reefs, upward trends in invertebrate biomass and species diversity have been shown to continue for several years (Fager 1971; Van Dolah et al. 1988; Relini et al. 1994) or for as many as 10-15 years after deployment (Aseltine-Neilson et al. 1999; Perkol-Finkel and Benayahu 2005). During this developmental period, their fouling assemblages may remain distinct from natural surroundings. Even after an epifaunal community is considered to be in equilibrium, biological interactions such as competition and predation, localized physical disturbances, and seasonal fluctuations may continually change what species are dominant (Pamintuan et al. 1994; Osman and Whitlatch 1998). These processes are crucial for maintaining diversity and species richness and preventing homogenization or monocultures, because space is often a limiting factor on hard substrata (Levin and Paine 1974). Although the dynamic nature of fouling assemblages on artificial substrata suggests that they will presumably display greater variability than those on natural reefs, it also appears that artificial habitats function similarly to natural ones in many respects, and may even converge with them over time. Wendt et al. (1989) compared five similar artificial reefs ranging from 3.5 to 10 years in age, and found that there was a group of species common to all of them - evidence of a “stable” or “climax” benthic community being achieved after a certain period of time. They also noted that many of the same types of organisms were found on nearby natural reefs. This parallels the pattern observed in the successional development of invertebrate assemblages by other artificial reef researchers, leading them to conclude that the potential exists for imitating adjacent natural communities (Hueckel and Buckley 1987; Jensen et al. 1994).

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In 2006, Perkol-Finkel et al. examined a 119-year-old shipwreck in the Red Sea and found that, given sufficient time, an artificial reef’s communities can become almost indistinguishable from those of a nearby natural reef, provided that the two habitats offer similar structural features. This was supported by the findings of a 5-year monitoring study in Miami-Dade County, Florida. An artificial mitigation reef, structurally designed to mimic local low-relief carbonate ridges that had been damaged by a beach renourishment project, developed benthic assemblages that exhibited a “high level of similarity to the natural reef in species composition and relative species representation”, with similarity increasing significantly during the study period (Thanner et al. 2006). The results of these studies emphasize the importance of using an experimental approach to standardize hydrological conditions, reef size, structure, age, or isolation when comparing natural and artificial substrata (Bohnsack and Sutherland 1985; Carr and Hixon 1997; Perkol-Finkel and Benayahu 2007). In other studies, observed differences between the invertebrate communities on natural reefs and those on urban structures or “unplanned” reefs such as pier pilings (Connell and Glasby 1999; Glasby 1999; Connell 2000, 2001; Perkol-Finkel and Benayahu 2004) have been attributed to lack of control for the above mechanisms, now known to influence reef recruitment and colonization. Artificial reefs have been the subject of extensive scrutiny and research, and fishery managers have expressed a desire to better understand differences among reef populations and communities, including those on natural reefs, in order to enhance the productivity of reef resources (Steimle and Meier 1997.) As yet, however, relatively little is known about their ecology (review by Bohnsack and Sutherland 1985). Part of the problem is that past artificial reef research has focused largely on the dynamics of fish

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populations and ignored the effects of fouling development and function (Fitzhardinge and Bailey-Brock 1989; Relini et al. 1994), despite direct evidence of links between the two ecosystems. Contemporaneous comparison with nearby natural reefs is also crucial for evaluating the ‘performance’ of an artificial reef (Carr and Hixon 1997; review by Svane and Petersen 2001); however, only a few long-term studies (greater than 5 years) have been conducted that compare artificial substrata to its natural counterparts, and the majority of these have been conducted in tropical areas, where growth of stony corals is the main concern (Thanner et al. 2006; Perkol-Finkel and Benayahu 2004, 2007). In the South Atlantic Bight, researchers have used fouling plates (Van Dolah et al. 1988) or looked at scrap reefs such as sunken vessels (Wendt et al. 1989), but designed reefs in this area present an alternate substrate, in terms of size, shape, and material, which may be governed by a completely different set of physical and biological benefits and constraints. The Artificial Reef Program of the South Carolina Department of Natural Resources has made efforts to address the deficit of designed reef data. In the last decade, two experimental reefs made up of hollow concrete modules were constructed and deployed off the coast of Charleston, S.C. These modules are cone-shaped, moderately rugose, and include large holes to increase habitat complexity for invertebrates and fish (see Figure 3). Originally deployed for the purpose of researching their potential as marine protected areas (MPAs), the coordinates for these reefs remain undisclosed to the public. “Area 53” is the younger of the two artificial sites; it was deployed in April 2003 and was approximately two years old at the time of this study. “Area 51” was deployed in May 1997, and was approximately eight years old. Originally, Area 51 was used for fishing

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experiments; the southeast and southwest corners were set up as an unfished reserve, while the northeast and northwest corners were experimentally fished with hook and line and baited traps. Fishing experiments at Area 51 ceased in 2002. Fish tagging and video monitoring studies are still conducted regularly at Areas 51 and 53, but as yet, there have been no monitoring efforts associated with the development of invertebrate assemblages on these reefs. While it is clear that enhancement of fish populations for recreational and commercial use remains the primary goal of today’s artificial reef programs, objectives of reef establishment have expanded to include relieving natural habitats of pressures from the fishing and tourism industries (Seaman et al. 1989), as well as the rehabilitation and restoration of disturbed natural reef systems (Pickering et al. 1998; review by Svane and Petersen 2001). Such disturbances may be due to severe weather events, but more often, are the result of damage caused by human economic developments in coastal areas. In the Maldives, the destructive fishing practice of coral mining has reduced many reefs to rubble, resulting in a loss of coral cover and fish diversity. Natural recovery time for these reefs can span decades, so artificial reefs have been used to restore reef fish populations and stimulate the recruitment of new coral colonies (Clark and Edwards 1999). In the South Atlantic Bight, ship groundings and increased fishing pressures specifically, the use of bottom-fishing techniques such as trawling - may also threaten the structural integrity and species assemblages of hard bottom and coral reefs. Reviews of numerous studies conducted over a wide range of geographic regions have documented measurable impacts to the seafloor as a result of bottom-fishing, including decreased biodiversity and habitat complexity, and homogenization of the substrata (Auster and

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Langton 1998; Collie et al. 2000; Veale et al. 2000; Thrush and Dayton 2002). Little is known about the recovery time of these systems, and much depends on how often the disturbance occurs (i.e., whether the effects are short-term or chronic). Although effects of chronic trawling on hard-bottom areas are not well documented, mobile bottom-fishing gear has been shown to damage or remove large epifaunal organisms after even a single use (Van Dolah et al. 1987). Dredging for beach renourishment projects and navigation channels has also been labeled as one of the most harmful human activities to hard bottom habitat, causing dislocation of live rock and corals, and stress to sessile invertebrates from elevated levels of sedimentation and turbidity (SAFMC 1998b). Some researchers believe that concrete reef structures may have the potential to support epifaunal communities similar to those on natural subtropical reefs, and if so, they may prove to be a useful tool in the restoration of portions of those reefs that have been damaged or denuded as a result of anthropogenic activities.

Objectives The main purpose of this study was to determine whether or not invertebrate communities that colonize South Carolina’s designed artificial reefs become similar to those found on natural hard-bottom reefs over time. Specific objectives were to: 1.) assess similarities and differences in the structure and composition of benthic invertebrate communities on a younger (Area 53) and older (Area 51) artificial reef, 2.) compare each of these to an adjacent natural reef, and 3.) compile invertebrate species lists for Area 53, Area 51, and the natural reef habitat. The following was the main hypothesis tested:

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H0: There are no major differences in benthic epifaunal community structure and selected indices (number of species, species richness, abundance, biomass, species diversity, and species evenness) among a 2-year-old designed artificial reef (Area 53), an 8-year-old designed artificial reef (Area 51), and an adjacent natural reef (“Julian’s Ledge”).

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MATERIALS AND METHODS

Study Sites The two artificial reef sites sampled in this study (Area 53 and Area 51) are located at undisclosed coordinates approximately 30 miles off the coast of Charleston, SC (Figure 1). Their depths are comparable (21 and 30m, respectively) and both are located in relatively close proximity to a 25-m-deep natural reef called “Julian’s Ledge”. Julian’s Ledge (32 32.986 N - 079 21.608 W) is part of “The Gardens”, a moderate-relief hardbottom area about 26 miles offshore of Charleston. The three sites form a triangular shape, with distances between each site ranging from 5 to 13 miles (Figure 2). The two artificial reefs are each set up in a 2.4 km² configuration, with 100 reef cones placed randomly at each corner. The orientation of the units was not altered after deployment, and many are situated on their sides or upside down, and some are broken into fragments. The four corners of each artificial reef area are separated by 1.5 nautical miles of sand bottom, and were thus treated as four smaller “patch” reefs. The cones (Figure 3) that make up these patches are a variation on the “Swiss Cone” designed by Stevens Towing Company in Yonge’s Island, SC (Robert Martore, personal communication).

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Preliminary Site Visit A preliminary visit to the artificial sites took place on November 2-4, 2004. Digital photographs and video clips were taken of cones on all corners of Area 53, and of the southeast and southwest corners of Area 51. Photographic data taken on this preliminary trip was not analyzed during the course of this study but was used to develop sampling techniques, and to assist with sampling design.

Field Sampling Field sampling consisted of a combination of underwater digital photography and scrape sampling. The sampling plan for this study was a multiple nested design, with fixed treatments of artificial reef (young), artificial reef (old,) and natural reef. For each treatment, four stations (i.e., the four corners of each artificial reef area, and stations A through D on Julian’s Ledge; Figure 2) were sampled for six replicates each. Thus, the number of samples resulting from this design was 24 from each reef site, for a total of 72. A single collection trip was planned for each reef site; however, sampling of Area 53 began on April 21st, 2005, and required an additional sampling day on May 10th, 2005, as well as a trip to the northeast and northwest corners in April 2006, in order to make up for insufficient material collected from these stations (see further explanation below). Sampling at Area 51 was conducted on July 13th, 2005, and Julian’s Ledge sampling was conducted on August 10th, 2005. Sampling time was limited due to safe diving practices. Thus, for reef cones at Area 53 and Area 51 that were already randomly configured, bottom time was limited to 20 minutes or less. Two types of hull-mounted depth recorders, a Furuno FCV-2000 dual

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frequency depth sounder and a Sitex dual frequency depth sounder, were used to detect the reef structures and to place an anchor as close as possible to a central location on a corner. A dive team consisting of 2-3 people descended the anchor line to collect one random sample from each of 6 cones closest to the anchor (regardless of cone orientation). If all 6 scrapings were not obtained by the first dive team, a second team was sent down at the same station to collect the remaining samples, noting not to sample the same cones twice. From each selected cone, a 15 cm2 exterior quadrat (cone interiors were not sampled) was photographed using a 5.0 megapixel Casio Exilim with attached strobe arm, and scraped with a hammer and chisel into a zippered mesh collection bag until the bare cone surface was exposed. The mesh bags were numbered 1-6, and a photograph of each bag’s number was taken by divers prior to photographing the actual quadrat. This allowed the sample photographs to later be matched to the corresponding scraped material. A 15 cm ruler was held for scale in each photograph and used periodically during the scraping process to verify the size of the sample area. The original sampling design called for one horizontal and one vertical sample from each selected cone (to give the best possible representation of organisms which might grow preferentially on different surface orientations); however, several attempts at collecting horizontal samples during the first collection trip to Area 53 revealed that there was not enough leverage to scrape from an overhead position. Also, without the assistance of gravity, scraped organisms floated in all directions and many were lost to currents and foraging reef fish. The horizontal scrapings were incomplete every time, resulting in three partial samples from Area 53’s northeast corner, and two samples with no material from the northwest corner (for a total of N = 22 instead of the intended 24).

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Methods were subsequently revised to include vertical scrapings only, and the two Area 53 stations with partial or missing samples were revisited in 2006 (material collected on this trip was not analyzed in this study, see Statistical Analysis: Univariate Methods). All scraped material was transferred on deck from the collection bags to corresponding labeled jars, and fixed with a 10% buffered formalin-seawater solution. Depth and temperature readings taken by dive computers were recorded. The same process was repeated in May 2005 for Area 51. In June 2005, four patches of the natural reef Julian’s Ledge (labeled A, B, C, and D) were sampled similarly to the four corners of the artificial reefs. All four patches were located at a single dive site, but were spaced out along a 40 m transect to simulate the distance found between the artificial reef stations (Figure 2). Six 15 cm² quadrats were randomly selected at each station along the transect, and each was digitally photographed before scraping or chiseling all organisms from the rocky substrate.

Laboratory Processing of Scraped Samples Once in the Southeastern Regional Taxonomic Center (SERTC) laboratory, samples were washed through a U.S. standard #35, 500 µm sieve to remove fine sediments, and transferred to a 70% EtOH solution. Processing began with a rough sort to remove chunks of cement (from the artificial reef samples), incidental vertebrates, and macroalgae, which were not included in any analysis. Total invertebrate biomass for each sample was then obtained by using a digital electronic balance to measure wet weights to the nearest gram. Barnacles were not included in biomass measurements due to the difficulty of distinguishing and separating them from the Julian’s Ledge samples,

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which consisted of chunks of rock rubble with attached and unattached algae and epifauna (Figure 4). Therefore, in order to standardize biomass for the two reef types, all barnacles (including soft tissue, shells, and shell fragments) were identified and removed from each sample prior to weighing. All remaining organisms were placed into one of two categories: “enumerated” or “sessile.” Enumerated organisms included all mobile taxa, as well as several groups of sessile taxa where individuals could be easily counted, including anemones, sessile bivalves, and some serpulid polychaetes (the tubiculous serpulid Spirobranchus giganteus (Pallas 1766), which would usually be considered sessile, was counted because it was large enough to be easily distinguished and removed from the substrate. It should also be noted that the burrowing barnacle Kochlorine floridana Wells and Tomlinson, 1966, which lacks a shell and did not contribute significantly to biomass due to its small size and infrequent occurrence, was separated and included with the counted organisms.) Sessile taxa included all encrusting and colonial organisms (Hydrozoa, Ectoprocta, Octocorallia, Ascidacea, Porifera, as well as several species of sessile, tube-dwelling polychates including the tubiculous serpulids Filograna implexa Berkeley and Chaetopteridae sp.). All organisms were placed into broad taxonomic categories, identified to the lowest practical taxonomic level, and counted (or, for sessile species which were not counted due to the impossibility of separating them, marked as present or absent). For groups such as polychaetes and amphipods, which can break into multiple segments during sample processing, only animals with heads were counted to minimize confusion. When necessary, large organisms such as sponges and tunicates were dissected to remove

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commensals, including polychaetes and amphipods. Tissue samples from sea cucumbers, sponges, and octocorals were dissolved in bleach and the remaining structural elements (ossicles, spicules, fibers, and sclerites) were examined using a compound microscope (Figure 5, a-b). Reference collections and taxonomic keys from the SERTC lab, Grice Marine Lab, and the Benthic Ecology labs from SCDNR and NOAA were utilized to assist with identifications, and the identifications of some specimens were confirmed by the SERTC staff. When necessary, specimens or photographs were sent to experts for identification. Samples are currently stored at the Marine Resources Research Institute at the SCDNR, and voucher specimens will eventually be accessioned into the SERTC collection.

Analysis of Digital Photographs and Archived Video Data Due to discrepancies in quality, digital photographs and videos were not consistently suitable for percent cover analysis, so observations made from these data were entirely qualitative and not statistically analyzed. Photographs were used to assist with the identification of organisms whose physical characteristics (i.e. shape, color) may have been altered during preservation. Archived videos of Area 53 and Area 51 collected by the SCDNR Artificial Reef Program, taken from deployment to the time of this study, were examined to determine duration from initial colonization to complete covering of the reef cones. Species that were present in photographs or videos but not collected in scrape samples were not included in statistical analysis.

16

Statistical Analyses: Univariate Methods Not all of the data collected in this study were used in analyses. Meiofaunal organisms (JL, 0.024

s S 0.000 JL>A53, 0.000 JL>A51, 0.001

NS 0.640*

NS 0.160*

Variable SR biomass NS S 0.000** 0.167* JL>A53, 0.000 JL>A51, 0.000 NS 0.150*

NS 0.403

H' S 0.000** JL>A53, 0.000 JL>A51, 0.000

J' S 0.000** JL>A53, 0.000 JL>A51, 0.000

NS 0.067*

NS 0.055*

83

Table 4. Ranking of most abundant species overall (> 1%), showing total abundance, percent of total abundance, and a summary of abundance of each species by reef site.

Scientific Name Syllis and Haplosyllis sp. Stenothoe sp. Elasmopus sp. Carpias bermudensis Chama congregata Pseudomedaeus agassizii Ericthonius brasiliensis Undet. Terebellidae Leucothoe spinicarpa Undet. Sipuncula Ophiactis algicola Gammaropsis sp. Undet. Leptocheliidae Undet. Actiniaria Polydora sp. Cymadusa sp. Hiatella arctica Timoclea grus Undet. Aspidosiphoniformes Lumbrineris inflata Microjassa tetradonta Micropanope sp. Caprella penantis Deutella incerta Undet. Nemertea Photis sp. Synalpheus townsendi Undet. Cirratulidae Hesionidae sp. A Ophiactis savignyi Spio sp. Eunice antennata Megalobrachium soriatum Proceraea sp. Websterinereis tridentata Undet. Sabellidae

Total Of Abundance 8847 1054 1033 854 610 563 545 545 497 472 438 434 400 353 318 306 292 284 279 271 251 236 225 206 202 186 183 183 179 173 172 169 166 162 144 136

% Total Abundance 35.47 4.23 4.14 3.42 2.45 2.26 2.19 2.19 1.99 1.89 1.76 1.74 1.60 1.42 1.28 1.23 1.17 1.14 1.12 1.09 1.01 0.95 0.90 0.83 0.81 0.75 0.73 0.73 0.72 0.69 0.69 0.68 0.67 0.65 0.58 0.55

Area 53 91 225 585 575 121 343 240 174 126 77 8 230 0 81 105 151 157 13

Area 51 7949 822 215 268 366 214 286 185 355 184 13 55 393 242 194 105 62 37

Julian’s Ledge 807 7 233 11 123 6 19 186 16 211 417 149 7 30 19 50 73 234

2 61 14 186 1 175 2 27 32 11 51 0 99 112 17 104 94 9

44 76 237 49 221 28 51 159 96 147 94 0 49 37 74 48 27 40

233 134 0 1 3 3 149 0 55 25 34 173 24 20 75 10 23 87

84

Table 5. Rank by abundance of the top 25 most abundant species in samples collected from Area 53 and Area 51 in 2005.

Area 53

Abundance

% of Total

Elasmopus sp. Carpias bermudensis Pseudomedaeus agassizii Ericthonius brasiliensis Gammaropsis sp. Stenothoe sp. Micropanope sp. Deutella incerta Undet. Terebellidae Hiatella arctica Cymadusa sp. Leucothoe spinicarpa Chama congregata Eunice antennata Polydora sp. Proceraea sp. Spio sp. Websterinereis tridentata Syllis and Haplosyllis sp. Undet. Actiniaria Undet. Sipuncula Dulichiella sp. Lumbrineris inflata Hesionidae sp. A Undet. Anamixidae

585 575 343 240 230 225 186 175 174 157 151 126 121 112 105 104 99 94 91 81 77 73 61 51 47

11.83 11.63 6.93 4.85 4.65 4.55 3.76 3.54 3.52 3.17 3.05 2.55 2.45 2.26 2.12 2.10 2.00 1.90 1.84 1.64 1.56 1.48 1.23 1.03 0.95

Area 51 Syllis and Haplosyllis sp. Stenothoe sp. Undet. Leptocheliidae Chama congregata Leucothoe spinicarpa Ericthonius brasiliensis Carpias bermudensis Undet. Actiniaria Microjassa tetradonta Caprella penantis Elasmopus sp. Pseudomedaeus agassizii Polydora sp. Undet. Terebellidae Undet. Sipuncula Photis sp. Undet. Cirratulidae Pycnogonum cessaci Cymadusa sp. Synalpheus townsendi Hesionidae sp. A Lumbrineris inflata Megalobrachium soriatum Hiatella arctica Lithophaga bisulcata

Abundance

% of Total

7949 822 393 366 355 286 268 242 237 221 215 214 194 185 184 159 147 106 105 96 94 76 74 62 57

54.87 5.67 2.71 2.53 2.45 1.97 1.85 1.67 1.64 1.53 1.48 1.48 1.34 1.28 1.27 1.10 1.01 0.73 0.72 0.66 0.65 0.52 0.51 0.43 0.39

85

Table 6. Rank by abundance of the top 25 most abundant species in samples collected from Julian’s Ledge in 2005.

Julian's Ledge Syllis and Haplosyllis sp. Ophiactis algicola Timoclea grus Elasmopus sp. Undet. Aspidosiphoniformes Undet. Sipuncula (other orders) Undet. Terebellidae Ophiactis savignyi Gammaropsis sp. Undet. Nemertea Lumbrineris inflata Chama congregata Undet. Sabellidae Gregariella coralliophaga Eunice filamentosa Megalobrachium soriatum Hiatella arctica Undet. Capitellidae Amphipholis squamata Liljeborgia sp. Eunice sp. A Synalpheus townsendi Pagurus carolinensis Cymadusa sp. Undet. Nebaliopsididae

Abundance 807 417 234 233 233 211 186 173 149 149 134 123 87 85 83 75 73 70 69 68 67 55 53 50 46

% of Total 14.65 7.57 4.25 4.23 4.23 3.83 3.38 3.14 2.71 2.71 2.43 2.23 1.58 1.54 1.51 1.36 1.33 1.27 1.25 1.23 1.22 1.00 0.96 0.91 0.84

86

Table 7. Overall rank by frequency of occurrence (%) of major sessile taxa occurring in 10 or more samples collected from Area 53, Area 51, and Julian’s Ledge in 2005.

Species Balanus trigonus Sertularella conica Clathrina sp. Crisulipora orientalis Bugula fulva Schizoporella unicornis Dynamena quadridentata Eudendrium carneum Thyroscyphus marginatus Tricellaria sp. A Filograna implexa Monostaechas quadridens Undet. Chaetopteridae Didemnum candidum Trididemnum savignii Undet. Demospongiae sp. E Chaperia sp. A Amathia vidovici Schizoporella violacea Tetraplaria dichotoma Dynamena cornicina Spongiidae sp. A Corydendrium parasiticum Halecium tenellum Mycale sp.

Count of Collections 70 37 32 29 27 22 21 21 20 20 19 19 19 19 18 18 17 15 14 13 12 11 10 10 10

Frequency of Occurrence (%) 100.00 52.86 45.71 41.43 38.57 31.43 30.00 30.00 28.57 28.57 27.14 27.14 27.14 27.14 25.71 25.71 24.29 21.43 20.00 18.57 17.14 15.71 14.29 14.29 14.29

87

Table 8. Rank by frequency of occurrence (%) of the top 25 most frequently occurring sessile species in samples collected from Area 53 (A53) and Area 51 (A51) during 2005.

A53 Balanus trigonus Sertularella conica Tricellaria sp. A Trididemnum savignii Bugula fulva Didemnum candidum Mycale sp. Schizoporella unicornis Undet. Demospongiae sp. E Halecium tenellum Clathrina sp. Aplidium stellatum (?) Clytia linearis Eusynstyela floridana Leucetta sp. Dynamena cornicina Spongiidae sp. A Leucosolenida sp. A Undet. Chondropsidae Symplegma viride Obelia dichotoma Dynamena quadridentata Leucosolenida sp. B Beania mirabilis

Occurrence (%) 100.00 86.36 77.27 63.64 54.55 50.00 45.45 40.91 40.91 40.91 36.36 31.82 31.82 27.27 27.27 22.73 22.73 22.73 22.73 18.18 18.18 13.64 13.64 13.64

A51 Balanus trigonus Eudendrium carneum Monostaechas quadridens Sertularella conica Bugula fulva Amathia vidovici Corydendrium parasiticum Undet. Demospongiae sp. E Crisulipora orientalis Schizoporella unicornis Dynamena cornicina Bugula neritina Halocordyle disticha Undet. Petrosina Lissodendoryx sp. Molgula occidentalis Clathrina sp. Dynamena quadridentata Didemnum candidum Trididemnum savignii Spongiidae sp. A Bugula turrita Caulibugula pearsei Symplegma viride Spongia sp.

Occurrence (%) 100.00 83.33 79.17 75.00 54.17 54.17 41.67 37.50 29.17 29.17 29.17 29.17 29.17 25.00 20.83 20.83 16.67 16.67 16.67 16.67 16.67 16.67 16.67 12.50 12.50

88

Table 9. Rank by frequency of occurrence (%) of the top 25 most frequently occurring sessile species in samples collected from Julian’s Ledge during 2005.

Julian’s Ledge Balanus trigonus Crisulipora orientalis Clathrina sp. Thyroscyphus marginatus Filograna implexa Undet. Chaetopteridae Chaperia sp. A Dynamena quadridentata Schizoporella violacea Tetraplaria dichotoma Ciocalypta sp. Turbicellepora dichotoma Arthropoma cecilii Eudistoma carolinense Leucosolenida sp. C Microporella sp. A Petraliella bisinuata Schizoporella unicornis Scypha sp. Leucosolenida sp. E Ircinia felix Cinachyra sp. Thesea nivea Didemnum candidum Craniella sp. Ectoprocta sp. A Fangophilina sp. Halichondrida sp. A Parasmittina spathulata

Occurrence (%) 100.00 91.67 83.33 83.33 79.17 79.17 70.83 58.33 58.33 54.17 37.50 33.33 33.33 33.33 29.17 29.17 29.17 25.00 25.00 25.00 25.00 20.83 20.83 16.67 16.67 16.67 16.67 16.67 16.67

89

Table 10. Species groups resulting from inverse Canberra metric cluster analysis of samples collected from Area 53, Area 51, and Julian’s Ledge during 2005 (Am = Amphipoda; Bi = Bivalvia; D = Decapoda; E = Echinodermata; G = Gastropoda; I = Isopoda; Mx = Maxillopoda; P = Polychaeta; Ph = Phyllocarida; Py = Pycnogonida; S = Sipuncula; Ta = Tanaidacea).

Group A Ampelisca sp. (Am) Hornellia tequestae (Am) Turbonilla sp. A (G) Barbatia domingensis (Bi) Kochlorine floridana (Mx) Synelmis sp. (P) Eunice sp. A (P) Ophiostigma isocanthum (E) Undet. Nebaliopsididae (Ph) Amphipholis januarii (E) Paracerceis caudata (I) Prionospio sp. (P) Eunice filamentosa (P) Ophiactis savignyi (E) Marshallora nigrocincta (G) Spirobranchus giganteus (P) Cumingia coarctata (Bi) Brania sp. (P) Marginellidae sp. A (Bi) Maldanidae sp. B (P) Pandora sp. A (Bi) Liljeborgia sp. (Am) Pterocirrus macrocerus (P) Mithraculus forceps (D) Amphipholis squamata (E) Arca zebra (Bi) Thyone pseudofusus (E) Autolytus sp. (P) Stramonita haemastoma floridana (G) Diodora cayenensis (G) Nereiphylla fragilis (P) Parapinnixa hendersoni (D) Ampithoe sp. (Am) Chama macerophylla (Bi) Lysidice ninetta (P) Astyris lunata (G) Cerithiopsis greenii (G) Nassarina glypta (G) Costoanachis avara (G)

Group B Caprella penantis (Am) Periclimenes iridescens (D) Diplodonta punctata (Bi) Undet. Leptocheliidae (Ta) Maera sp. (Am) Colomastix sp. (Am) Pycnogonum cessaci (Py) Hypsicomus sp. (G) Lithophaga bisulcata (Bi) Odontosyllis fulgurans (P) Group C Dendostrea frons (Bi) Polynoidae sp. A (P) Dulichiella sp. (Am) Synalpheus minus (D) Vitrinellidae sp. A (G) Undet. Trematoda Group D Carpias bermudensis (I) Ericthonius brasiliensis (Am) Stenothoe sp. (Am) Pseudomedaeus agassizii (D) Leucothoe spinicarpa (Am) Chama congregata (Bi) Undet. Sipuncula (S) Undet. Terebellidae (P) Elasmopus sp. (Am) Lumbrineris inflata (P) Syllis and Haplosyllis sp. (P) Cymadusa sp. (Am) Hesionidae sp. A (P) Polydora sp. (P) Eunice antennata (P) Spio sp. (P) Hiatella arctica (Bi) Gammaropsis sp. (Am) Megalobrachium soriatum (D) Ophiothrix angulata (E) Synalpheus townsendi (D)

Proceraea sp. (P) Undet. Actiniaria Websterinereis tridentata (P) Pagurus carolinensis (D) Timoclea grus (Bi) Undet. Capitellidae (P) Gastrochaena hians (Bi) Undet. Cirratulidae (P) Micropanope nuttingi (D) Polynoidae sp. B (P) Undet. Aoridae (Am) Pherusa sp. A (P) Nicon sp. A (P) Trypanosyllis zebra (P) Group E Gregariella coralliophaga (Bi) Joeropsis coralicola (I) Lithophaga bisulcata (Bi) Undet. Aspidosiphoniformes (S) Undet. Nemertea Ophiactis algicola (E) Group F Deutella incerta (Am) Megalomma sp. (P) Microjassa tetradonta (Am) Photis sp. (Am) Latreutes parvulus (D) Pilumnus dasypodus (D) Undet. Turbellaria Musculus lateralis (Bi) Undet. Anamixidae (Am) Thor manningi (D) Podocerus sp. (Am) Pododesmus rudis (Bi) Group G Eulalia sanguinea (P) Pectinariidae sp. A (P) Pelia mutica (D)

90

Table 11. Species groups resulting from inverse Jaccard similarity analysis of samples collected from Area 53, Area 51, and Julian’s Ledge during 2005 (Am = Amphipoda; As = Ascidiacea; Bi = Bivalvia; Br = Brachiopoda; D = Decapoda; E = Echinodermata; Ec = Ectoprocta; G = Gastropoda; Hy = Hydrozoa; I = Isopoda; Mx = Maxillopoda; My = Mysida; N = Nudibranchia; O = Octocorallia; P = Polychaeta; Ph = Phyllocarida; Po = Porifera; Py = Pycnogonida; S = Sipuncula; St = Stomatopoda, Ta = Tanaidacea). Group A Undet. Terebellidae (P) Websterinereis tridentata (P) Balanus trigonus (Mx) Undet. Capitellidae (P) Undet. Sipuncula (S) Timoclea grus (Bi) Undet. Actiniaria Syllis and Haplosyllis sp. (P) Synalpheus townsendi (D) Proceraea sp. (P) Spio sp. (P) Ophiothrix angulata (E) Polydora sp. (P) Megalobrachium soriatum (D) Micropanope nuttingi (D) Hiatella arctica (Bi) Lumbrineris inflata (P) Gammaropsis sp. (Am) Hesionidae sp. A (P) Elasmopus sp. (Am) Eunice antennata (P) Chama congregata (Bi) Cymadusa sp. (Am) Carpias bermudensis (I) Clathrina sp. (Po) Schizoporella unicornis (Ec) Polynoidae sp. B (P) Undet. Aoridae (Am) Pagurus carolinensis (D) Ericthonius brasiliensis (Am) Stenothoe sp. (Am) Bugula fulva (Ec) Pherusa sp. A (P) Leucothoe spinicarpa (Am) Pododesmus rudis (Bi) Ophiactis algicola (E) Dynamena quadridentata (Hy) Undet. Cirratulidae (P)

Gastrochaena hians (Bi) Undet. Nemertea Joeropsis coralicola (I) Lithophaga bisulcata (Bi) Undet. Aspidosiphoniformes (S) Nicon sp. A (P) Trypanosyllis zebra (P) Eulalia sanguinea (P) Pelia mutica (D) Latreutes parvulus (D) Pilumnus dasypodus (D) Undet. Turbellaria Deutella incerta (Am) Megalomma sp. (P) Pseudomedaeus agassizii (D) Sertularella conica (Hy) Undet. Demospongiae sp. E (Po) Photis sp. (Am) Dynamena cornicina (Hy) Microjassa tetradonta (Am) Trididemnum savignii (As) Doto sp. (N) Undet. Anamixidae (Am) Didemnum candidum (As) Podocerus sp. (Am) Tricellaria sp. A (Ec) Spongiidae sp. A (Po) Musculus lateralis (Bi) Thor manningi (D) Marphysa sp. A (P) Pectinariidae sp. A (P) Group B Diplodonta punctata (Bi) Undet. Leptocheliidae Maera sp. (Am) Periclimenes iridescens (D)

Caprella penantis (Am) Eudendrium carneum (Hy) Lissodendoryx sp. (Po) Halocordyle disticha (Hy) Monostaechas quadridens (Hy) Corydendrium parasiticum (Hy) Colomastix sp. (Am) Spongia sp. (Po) Lysianopsis alba (Am) Amathia vidovici (Ec) Odontosyllis fulgurans (P) Lithophaga aristata (Bi) Pycnogonum cessaci (Py) Hypsicomus sp. (P) Group C Sertularella unituba (Hy) Leucosolenida sp. A (Po) Group D Bugula uniserialis (Ec) Astyris lunata (Bi) Group E Mithrax hispidus (D) Parapinnixa hendersoni (D) Nereiphylla fragilis (P) Alderina sp. A (Ec) Arca imbricata (Bi) Gemmotheres chamae (D) Group F Vermiliopsis annulata (P) Scalibregmatidae sp. A (P)

91

Table 11. Continued. Group F (continued) Pteria colymbus (Bi) Conopea merrilli (Mx) Chama macerophylla (Bi) Undet. Cumacea Group G Arbacia punctulata (E) Stenorhynchus seticornis (D) Erichsonella filiformis (I) Ampithoe sp. (Am) Aplysilla sulfurea (Po) Group H Anaitides sp. A (P) Glycera sp. A (P) Undet. Echiura Scypha sp. (Po) Costoanachis avara (G) Cerithiopsis greenii (G) Nassarina glypta (G) Group I Pagurus brevidactylus (D) Thracia morrisoni (Bi) Malleus candeanus (Bi) Natacidae sp. A (G) Boonea seminuda (G) Cantharus multangulus (G) Diplodonta semiaspera (Bi) Ectoprocta sp. A (Ec) Halichondrida sp. A (Po) Anchialina typica (My) Parasmittina trispinosa (Ec) Spirobranchus giganteus (P) Chicoreus pomum (G) Mytilidae sp. A (Bi) Paguristes tortugae (D)

Marshallora nigrotincta (G) Modiolus americanus (Bi) Crepidula aculeata (G) Gonodactylus bredini (St) Craniella sp. (Po) Cerithiopsis emersonii (Bi) Kalliapseudidae sp. (Ta) Chrysopetalidae sp. B (P) Pitho sp. (D) Amphipholis januarii (E) Ceratoneries mirabilis (P) Macrocoeloma trispinosum (D) Microporella sp. A (Ec) Vitrinellidae sp. B (G) Brania sp. (P) Maldanidae sp. B (P) Marginellidae sp. A (G) Ciocalypta sp. (Po) Gouldia cerina (Bi) Cinachyra sp. (Po) Crisulipora orientalis (Ec) Diodora cayenensis (G) Turbicellepora dichotoma (Ec) Undet. Inarticulata (Br) Leucosolenida sp. C (Po) Liljeborgia sp. (Am) Pterocirrus macroceros (P) Undet. Chaetopteridae (P) Undet. Nebaliopsididae (Ph) Ampelisca sp. (Am) Thyroscyphus marginatus (Hy) Turbonilla sp. A (G) Synelmis sp. (P) Tetraplaria dichotoma (Ec) Prionospio sp. (P) Schizoporella violacea (Ec) Paracerceis caudata (I) Petraliella bisinuata (Ec) Ophiactis savignyi (E) Ophiostigma isocanthum (E)

Ircinia felix (Po) Kochlorine floridana (Mx) Filograna implexa (P) Hornellia tequestae (Am) Eunice sp. A (P) Exogone sp. (P) Eudistoma carolinense (As) Eunice filamentosa (P) Barbatia domingensis (Bi) Chaperia sp. A (Ec) Arthropoma cecilii (Ec) Cumingia coarctata (Bi) Pandora sp. A (Bi) Hyattella sp. (Po) Thyone pseudofusus (E) Amphipholis squamata (E) Arca zebra (Bi) Leucosolenida sp. E (Po) Mithraculus forceps (D) Autolytus sp. (P) S. haemastoma floridana (G) Gregariella coralliophaga (Bi) Lysidice ninetta (P) Parasmittina spathulata (Ec) Trypanosyllis sp. A (P) Thesea nivea (O) Undet. Apseudidae (Ta) Arcopsis adamsi (Bi) Microphrys antillensis (D) Scrupocellaria regularis (Ec) Fangophilina sp. (Po) Processa sp. (D) Amphiura sp. A (E) Rullerinereis sp. A (P) Group J Synalpheus minus (D) Leucosolenida sp. B (Po) Vitrinellidae sp. A (G)

92

Table 11. Continued. Group J (continued) Typton sp. A (D) Periclimenes americanus (D) Achelia sawayai (Py) Ircinia strobilina (Po) Facelina sp. (N) Neopontonides beaufortensis (D) Caulibugula pearsei (Ec) Molgula occidentalis (As) Undet. Petrosina (Po) Amblyosyllis formosa (P)

Aplidium stellatum? (As) Pilumnus sayi (D) Limnotheres nasutus (D) Eusynstyela floridana (As) Undet. Trematoda Beania mirabilis (Ec)

Group G Chrysopetalidae sp. A (P) Group H Bugula turrita (Ec) Bugula neritina (Ec) Obelia dichotoma (Hy) Tumidotheres maculatus (D) Leucosolenia sp. (Po) Lumbrineridae sp. A (P) Parvilucina multilineata (Bi) Dendostrea frons (Bi) Doridacea sp. B (N) Polycera chilluna (N) Aeolidacea sp. A (N) Doridacea sp. A (N) Mycale sp. (Po) Polynoidae sp. A (P) Turritopsis nutricula (Hy) Symplegma viride (As) Undet. Chondropsidae (Po) Dulichiella sp. (Am) Halecium tenellum (Hy) Clytia linearis (Hy) Pomatoceros sp. A (P) Leucetta sp. (Po) Ostreidae sp. A (Bi)

93

Table 12. Results of Components of Variance analysis, showing contribution of percent variance among reef stations, and percent variance due to error, to total variance.

Variance - station (%) Variance - error (%)

n s 9.8 90.2

SR 50.7 49.3

Variable biomass H´ J´ 49.8 8.3 22.3 23.5 50.2 91.7 77.7 76.5

94

APPENDICES

95

Appendix A. List of species collected from each sampling site during 2005. Area 53 Amphipoda Americorophium sp. Ampelisca sp. Ampithoe sp. Caprella penantis Colomastix sp. Cymadusa sp. Deutella incerta Dulichiella sp. Elasmopus sp. Ericthonius brasiliensis Gammaropsis sp. Hornellia tequestae Leucothoe spinicarpa Liljeborgia sp. Lysianopsis alba Maera sp. Microjassa tetradonta Photis sp. Podocerus sp. Stenothoe sp. Undet. Amphilochidae Undet. Anamixidae Undet. Aoridae Undet. Eusiridae Undet. Iphimediidae

x x x x x x x x x x

x x x x x x

Area 51

Julian’s Ledge

x

x x x x x x x x x x x x x x x x

x x x x x x x x x x x x x x x x x x x

x x x x x x

Anthozoa Leptogorgia cardinalis Oculina arbuscula Telesto fruticulosa Thesea nivea Undet. Actiniaria Undet. Zoanthidia

x x

x

Ascidiacea Aplidium sp.

x

x

x x x x x

96

Aplidium stellatum (?) Clavelina sp. Didemnidae sp. A Didemnum candidum Eudistoma carolinense Eusynstyela floridana Molgula occidentalis Pyura vittata Styelidae sp. A Symplegma viride Trididemnum savignii Undet. Ascidiacea - colonial Undet. Ascidiacea - solitary Bivalvia Americardia media Arca imbricata Arca zebra Arcopsis adamsi Barbatia candida Barbatia domingensis Chama congregata Chama macerophylla Chama sp. A Crassinella lunulata Cumingia coarctata Dendostrea frons Diplodonta punctata Diplodonta semiaspera Gastrochaena hians Gouldia cerina Gregariella coralliophaga Hiatella arctica Isognomon radiatus Lithophaga aristata Lithophaga bisulcata Lyonsiidae sp. A Malleus candeanus Modiolus americanus Musculus lateralis

x

x

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97

Mytilidae sp. A Nuculana sp. A Ostrea equestris Ostreidae sp. A Pandora sp. A Parvilucina multilineata Plicatula gibbosa Pododesmus rudis Pteria colymbus Thracia morrisoni Timoclea grus Undet. Bivalvia Cumacea Undet. Cumacea Decapoda Alpheus formosus Alpheus normanni Alpheus sp. Anchistioides antiguensis Gemmotheres chamae Hippolyte obliquimanus Hippolyte sp. Hippolyte zostericola Latreutes fucorum Latreutes parvulus Leptochela papulata Lucifer faxoni Macrocoeloma trispinosum Megalobrachium soriatum Micropanope nuttingi Micropanope sp. Microphrys antillensis Mithraculus forceps Mithrax hispidus Mithrax pleuracanthus Mithrax sp. Munida sp. A Neopontonides beaufortensis

x x x x x x

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98

Paguristes sp. A Paguristes tortugae Pagurus brevidactylus Pagurus carolinensis Parapinnixa hendersoni Pelia mutica Periclimenaeus sp. Periclimenes americanus Periclimenes iridescens Periclimenes sp. Petrolisthes galathinus Pilumnus dasypodus Pilumnus sayi Pilumnus sp. Pinnixa chaetopterana Limnotheres nasutus Pitho sp. Podochela gracilipes Podochela sidneyi Processa sp. Pseudomedaeus agassizii Sicyonia laevigata Speloeophorus pontifer Stenocionops furcatus coelata Stenocionops sp. Stenorhynchus seticornis Synalpheus fritzmuelleri Synalpheus minus Synalpheus sp. Synalpheus townsendi Thor dobkini Thor manningi Thor sp. Tumidotheres maculatus Typton sp. A Undet. Brachyura Undet. Caridea Undet. Pinnotheridae Zaops ostreum

x x x

x x x

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x

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Echinoidea Arbacia punctulata Echiura Undet. Echiura Ectoprocta Aeverrillia setigera Alderina sp. A Amathia vidovici Arthropoma cecilii Beania hirtissima Beania mirabilis Bugula fulva Bugula neritina Bugula turrita Bugula uniserialis Caberea boryi Caulibugula pearsei Caulibugula sp. Celleporaria magnifica Chaperia sp. A Crisulipora orientalis Diaperoecia floridana Ectoprocta sp. A Hippoliosina rostrigera Hippopleurifera mucronata Hippoporina contracta Lichenopora radiata Microporella sp. A Parasmittina spathulata Parasmittina trispinosa Petraliella bisinuata Reptadeonella costulata Rhynchozoon rostratum Schizoporella unicornis Schizoporella violacea Scrupocellaria regularis Tetraplaria dichotoma Tricellaria sp. A

x

x

x

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x

x x x

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x

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x

Turbicellepora dichotoma Gastropoda Aeolidacea sp. A Aeolidacea sp. B Aeolidacea sp. C Aeolidacea sp. D Aeolidacea sp. E Aeolidacea sp. F Aeolidacea sp. G Aeolidacea sp. H Astyris lunata Boonea seminuda Cantharus multangulus Cerithiopsis emersonii Cerithiopsis greenii Cerithiopsis sp. A Cerithium atratum Chicoreus pomum Costoanachis avara Crepidula aculeata Crepidula plana Diodora cayenensis Doridacea sp. A Doridacea sp. B Doridacea sp. C Doridacea sp. D Doridacea sp. E Doto sp. Facelina sp. Marginellidae sp. A Marshallora nigrotincta Melanella jamaicensis Melanella sp. A Muricidae sp. A Nassarina glypta Natacidae sp. A Odostomia sp. A Olivella mutica Pollia tincta

x

x x x x x x

x

x x x x

x x

x

x

x

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Polycera chilluna Seila adamsi Stramonita haemastoma floridana Terebra dislocata Trachypollia nodulosa Triphora triserialis Turbo castanea Turbonilla sp. A Turridae sp. A Undet. Aeolidacea Undet. Prosobranchia Vexillum sykesi Vitrinellidae sp. A Vitrinellidae sp. B

x x x

x x x x

x x x

x

Holothuroidea Istichopus badionotus Ocnus pygmaeus Thyone deichmannae Thyone pseudofusus Hydroida Bougainvillia muscus Campanularia hincksii Clytia linearis (?) Clytia noliformis (?) Corydendrium parasiticum Dynamena cornicina Dynamena quadridentata Eudendrium carneum Eudendrium sp. Halecium tenellum Halocordyle disticha Macrorhynchia allmani Monostaechas quadridens Obelia dichotoma Plumularia setacea Sertularella conica Sertularella unituba Thyroscyphus marginatus

x x

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x x x x x x x

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Turritopsis nutricula

x

Inarticulata Undet. Inarticulata Isopoda Arcturella spinata Arcturidae sp. A Carpias bermudensis Carpias minutus Carpias sp. A Edotia triloba Erichsonella filiformis Joeropsis coralicola Paracerceis caudata Undet. Anthuridae Undet. Bopyridae Undet. Gnathiidae Maxillopoda Balanus trigonus Conopea merrilli Copepoda - Harpacticoida Kochlorine floridana Mysidacea Anchialina typica Ophiuroidea Amphipholis januarii Amphipholis squamata Amphiura sp. A Ophiactis algicola Ophiactis savignyi Ophiocominae sp. A Ophioderma appressum Ophiopsila riisei Ophiostigma isocanthum Ophiothrix angulata Undet. Ophiuroidea

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x

x

x x

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Nemertea Undet. Nemertea

x

x

Phyllocarida Undet. Nebaliopsididae Polychaeta Amblyosyllis formosa Anaitides sp. A Aphroditidae sp. A Aphroditidae sp. B Autolytus sp. Brania sp. Ceratoneries mirabilis Chone sp. A Chrysopetalidae sp. A Chrysopetalidae sp. B Eulalia sanguinea Eunice antennata Eunice filamentosa Eunice sp. A Exogone sp. Filograna implexa Glycera sp. A Hesionidae sp. A Hypsicomus sp. Laonice cirrata Lumbrineridae sp. A Lumbrineris inflata Lysidice ninetta Maldanidae sp. A Maldanidae sp. B Marphysa sp. A Megalomma sp. Neanthes sp. A Nereiphylla fragilis Nereis riisei Nicon sp. A Odontosyllis fulgurans

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x

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Oenonidae sp. A Paranaitis sp. A Pectinariidae sp. A Pherusa sp. A Polydora sp. Polynoidae sp. A Polynoidae sp. B Pomatoceros sp. A Prionospio sp. Proceraea sp. Pseudovermilia occidentalis Pterocirrus macroceros Rullerinereis sp. A Scalibregmatidae sp. A Serpulidae sp. A Spio sp. Spirobranchus giganteus Syllis and Haplosyllis sp. Synelmis sp. Trypanosyllis sp. A Trypanosyllis zebra Undet. Capitellidae Undet. Chaetopteridae Undet. Cirratulidae Undet. Eunicidae Undet. Nereididae Undet. Phyllodocidae Undet. Polychaeta Undet. Sabellidae Undet. Serpulidae Undet. Syllidae Undet. Terebellidae Vermiliopsis annulata Websterinereis tridentata Xenosyllis sp. Porifera Aplysilla sulfurea Aplysina sp. Axinella bookhouti

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Axinella polycapella Chelonaplysilla erecta Cinachyra sp. Ciocalypta sp. Clathrina sp. Cliona sp. Craniella sp. Dysidea fragilis Dysidea (?) sp. Fangophilina sp. Geodia sp. Halichondrida sp. A Haliclona (?) sp. B Hyattella sp. Ircinia felix (?) Ircinia strobilina (?) Leucetta sp. Leucosolenia sp. Leucosolenida sp. A Leucosolenida sp. B Leucosolenida sp. C Leucosolenida sp. D Leucosolenida sp. E Lissodendoryx sp. Microciona prolifera Mycale sp. Samus anonymus Scypha sp. Spongia sp. Spongiidae sp. A Undet. Ancorinidae (?) Undet. Calcarea Undet. Chondropsidae (?) Undet. Demospongiae Undet. Demospongiae sp. A Undet. Demospongiae sp. B Undet. Demospongiae sp. C Undet. Demospongiae sp. D Undet. Demospongiae (?) sp. E Undet. Dictyoceratida

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Undet. Hemiasterellidae Undet. Leucosolenida Undet. Niphatidae (?) Undet. Pachastrellidae Undet. Petrosina Undet. Poecilosclerida Undet. Suberitidae (?) Undet. Tethyidae Pycnogonida Achelia sawayai Achelia sp. A Anoplodactylus lentus Ascorhynchus sp. A Pycnogonum cessaci Undet. Ammotheidae Sipuncula Undet. Aspidosiphoniformes Undet. Sipuncula (other orders)

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Stomatopoda Gonodactylus bredini Gonodactylus sp.

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Tanaidacea Undet. Apseudidae Undet. Kalliapseudidae Undet. Leptocheliidae Turbellaria Undet. Turbellaria

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Trematoda Undet. Trematoda

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Undet. Phyla

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Appendix B. List of meifaunal or pelagic organisms collected during spring/summer 2005.

Scientific Name

A53

Copepoda - Calanoida Copepoda - Cyclopoida

x

Ostracoda sp. A

x

Ostracoda sp. B

A51

NR

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x

x x

x

Ostracoda sp. C

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Ostracoda sp. D

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Ostracoda sp. E

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Ostracoda sp. F

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Sarsiellidae sp. A

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Sarsiellidae sp. B

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Undet. Myodocopida

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Undet. Nemata

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Appendix C. List of species collected exclusively from Area 53’s northeast and northwest corners during April 2006.

Species Ascidia sp. A Genetyllis castanea Haliclona sp. A Hypselodoris edenticulata Inachoides forceps Lima sp. A Lindapecten muscuosus Metoporhaphis calcarata Pachycheles rugimanus Pagurus sp. Pilumnus floridanus Polynoidae sp. C Polynoidae sp. D Smittina sp. A Undet. Ammotheidae Undet. Clathrinida Undet. Xanthidae

A53-NE x x

A53-NW x x

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Appendix D. Lists of species collected exclusively from Area 53 and Area 51 during 2005.

Area 53 Aeolidacea sp. A Aeolidacea sp. B Aeolidacea sp. C Aeolidacea sp. D Aeolidacea sp. E Aeolidacea sp. F Anchistioides antiguensis Beania mirabilis Bougainvillia muscus Clytia linearis Clytia noliformis Doridacea sp. A Doridacea sp. B Isognomon radiatus Leptochela papulata Leucetta sp. Leucosolenida sp. D Neanthes sp. A Paranaitis sp. A Limnotheres nasutus Plumularia setacea Podochela sidneyi Polycera chilluna Pyura vittata Terebra dislocata Undet. Trematoda Undet. Zoanthidia

Area 51 Achelia sp. A Aeolidacea sp. G Aeolidacea sp. H Amblyosyllis formosa Bugula turrita Campanularia hincksii Carpias minutus Caulibugula pearsei Corydendrium parasiticum Doridacea sp. C Dysidea fragilis Halocordyle disticha Hippolyte obliquimanus Hippolyte zostericola Latreutes fucorum Molgula occidentalis Monostaechas quadridens Oculina arbuscula Olivella mutica Pinnixa chaetopterana Plicatula gibbosa Spongia sp. Synalpheus fritzmuelleri Turbo castanea Undet. Eusiridae Undet. Suberitidae

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Appendix E. List of species collected exclusively from Julian’s Ledge during 2005.

Julian's Ledge Alpheus formosus Alpheus normanni Americardia media Ampelisca sp. Amphipholis januarii Amphiura sp. A Anoplodactylus lentus Aphroditidae sp. B Aplysilla sulfurea Aplysina sp. Arbacia punctulata Arcopsis adamsi Arcturella spinata Arcturidae sp. A Arthropoma cecilii Ascorhynchus sp. A Axinella bookhouti Axinella polycapella Barbatia candida Barbatia domingensis Beania hirtissima Boonea seminuda Brania sp. Caberea boryi Carpias sp. A Celleporaria magnifica Ceratoneries mirabilis Cerithiopsis sp. A Cerithium atratum Chaperia sp. A Chelonaplysilla erecta Chicoreus pomum Chone sp. A

Chrysopetalidae sp. B Cinachyra sp. Ciocalypta sp. Clavelina sp. Cliona sp. Craniella sp. Crassinella lunulata Crepidula aculeata Crepidula plana Diaperoecia floridana Didemnidae sp. A Doridacea sp. E Ectoprocta sp. A Edotia triloba Eudistoma carolinense Eunice filamentosa Eunice sp. A Fangophilina sp. Filograna implexa Geodia sp. Gonodactylus bredini Gouldia cerina Halichondrida sp. A Haliclona sp. B Hippoliosina rostrigera Hippoporina contracta Ircinia felix Istichopus badionotus Kalliapseudidae sp. Kochlorine floridana Leptogorgia cardinalis Lichenopora radiata Lucifer faxoni

Lyonsiidae sp. A Macrorhynchia allmani Maldanidae sp. B Malleus candeanus Marginellidae sp. A Melanella jamaicensis Melanella sp. A Microciona prolifera Microporella sp. A Modiolus americanus Muricidae sp. A Mytilidae sp. A Natacidae sp. A Nereis riisei Nuculana sp. A Ocnus pygmaeus Odostomia sp. A Oenonidae sp. A Ophiactis savignyi Ophiocominae sp. A Ophioderma appressum Ophiopsila riisei Ophiostigma isocanthum Ostrea equestris Paguristes sp. A Paguristes tortugae Pagurus brevidactylus Parasmittina spathulata Parasmittina trispinosa Periclimenaeus sp. Petraliella bisinuata Petrolisthes galathinus Pitho sp.

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Appendix E. Continued. Julian’s Ledge Prionospio sp.

Undet. Demospongiae sp. A

Processa sp.

Undet. Demospongiae sp. B

Pseudovermilia occidentalis

Undet. Demospongiae sp. C

Reptadeonella costulata

Undet. Demospongiae sp. D

Rhynchozoon rostratum

Undet. Gnathiidae

Samus anonymus

Undet. Hemiasterellidae

Schizoporella violacea

Undet. Iphimediidae

Scrupocellaria regularis

Undet. Nebaliopsididae

Serpulidae sp. A

Undet. Niphatidae

Speloeophorus pontifer

Undet. Pachastrellidae

Spirobranchus giganteus

Undet. Tethyidae

Styelidae sp A

Vexillum sykesi

Synelmis sp.

Vitrinellidae sp. B

Telesto fruticulosa

Xenosyllis sp.

Tetraplaria dichotoma Thesea nivea Thracia morrisoni Thyone deichmannae Trachypollia nodulosa Triphora triserialis Trypanosyllis sp. A Turridae sp. A Undet. Amphilochidae Undet. Ancorinidae Undet. Anthuridae Undet. Apseudidae Undet. Bopyridae Undet. Chaetopteridae

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Appendix F. List of species common to Area 53, Area 51, and Julian’s Ledge during 2005.

Achelia sawayai

Hiatella arctica

Proceraea sp.

Amathia vidovici

Joeropsis coralicola

Pseudomedaeus agassizii

Astyris lunata Autolytus sp.

Latreutes parvulus Leucosolenida sp. A

Rullerinereis sp. A

Balanus trigonus

Leucosolenida sp. B

Bugula fulva

Leucosolenida sp. E

Sertularella unituba Spio sp.

Caprella penantis

Spongiidae sp. A

Carpias bermudensis

Leucothoe spinicarpa Lissodendoryx sp.

Chama congregata

Lithophaga bisulcata

Chama macerophylla Clathrina sp.

Lumbrineris inflata

Stramonita haemastoma floridana Syllis and Haplosyllis sp.

Conopea merrilli Costoanachis avara Crisulipora orientalis Cymadusa sp.

Lysidice ninetta Marphysa sp. A Megalobrachium soriatum Megalomma sp. Micropanope nuttingi

Dendostrea frons

Musculus lateralis

Deutella incerta

Nereiphylla fragilis Nicon sp. A

Didemnum candidum Doto sp.

Schizoporella unicornis

Stenothoe sp.

Synalpheus minus Synalpheus townsendi Thor manningi Timoclea grus Tricellaria sp. A Trypanosyllis zebra Typton sp. A Undet. Actiniaria

Ophiactis algicola

Undet. Anamixidae

Dulichiella sp.

Ophiothrix angulata

Undet. Aoridae

Dynamena quadridentata Elasmopus sp.

Pagurus carolinensis Pectinariidae sp. A

Undet. Aspidosiphoniformes

Ericthonius brasiliensis

Pelia mutica

Undet. Cirratulidae

Eulalia sanguinea

Periclimenes americanus Pherusa sp. A

Undet. Cumacea

Pilumnus dasypodus

Undet. Sipuncula

Pilumnus sayi Podocerus sp.

Undet. Terebellidae

Eunice antennata Gammaropsis sp. Gastrochaena hians Gemmotheres chamae Glycera sp. A Gregariella coralliophaga Hesionidae sp. A

Pododesmus rudis Polydora sp.

Undet. Capitellidae

Undet. Nemertea

Undet. Turbellaria Vitrinellidae sp. A Websterinereis tridentata

Polynoidae sp. B

113