STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION: PUERTO RICO AND THE U.S. VIRGIN ISLANDS
Steven J. Lutz1,2 and Robert N. Ginsburg1
I. INTRODUCTION Regional Deep-sea Research
This chapter summarizes published and Louis F. de Pourtalès first discovered deep-sea unpublished information on deep corals in the life in the region, including deep-water corals,
wider Caribbean region (Caribbean Sea and in the 1860’s. Pourtalès, a scientist aboard the CARIBBEAN Bahamian Archipelago) including those in the Coast Survey steamer Bibb, found evidence of life exclusive economic zone (EEZ) of United States off Cuba at 945 m (3,100 ft). He studied regional Possessions. It summarizes the occurrences of deep-sea corals and published comprehensive the corals, associated species and their sea floor works (Pourtalès 1867, 1871). Pourtalès was accumulations; it provides basic information on also a pioneer of deep-sea dredging, a method the geography, geology and oceanography of the that was used by later research cruises. These region and how they interact to produce sea floor cruises were not focused on deep-sea corals and elevations; and it discusses stressors, including recovered specimens were incidental findings. potential fisheries interactions. Since the discoveries made by Pourtalès and the Bibb many research cruises have contributed For the purposes of this chapter deep corals are to the understanding of deep-sea corals. Early defined as those in waters below 100 m. cruises included voyages by the U.S. Coast Survey vessel Blake (late 1870’s), U.S. Fish The territorial jurisdiction of the U.S. in the Commission Albatross (1880’s) and the H.M.S. Caribbean is limited to the areas around Puerto Challenger (1872-1876). Later research cruises Rico, the U.S. Virgin Islands and the uninhabited included voyages by the R/V Oregon and R/V island of Navassa between Cuba and Hispaniola, Oregon II (1950’s to 1970’s), R/V Gerda (1960’s), illustrated in Figure 8.1. The EEZ of Puerto Rico R/V Pillsbury (1960’s and 70’s), the R/V Eastward and the U.S. Virgin Islands extends from the (1970’s), the R/V Akademik Vernadski (early edge of the territorial waters to 200 nautical miles 1970’s); and R/V Columbus Iselin (1970’s and to the north and is bounded to the south by the 80’s). EEZ of Venezuela, to the east by the British Virgin Islands and to the west by Dominican Republic. Beginning in the 1970’s, dredging was The EEZ of Navassa Island is bounded to the supplemented by the use of research north by Cuba, to the west by Jamaica and to the submersibles. These direct observations and east by Haiti. photographs of deep-sea coral communities provided a major advance in understanding the As information on deep-sea corals in the U.S. occurrences, morphology and communities of Caribbean is limited, a consideration of the wider deep-sea corals in the Straits of Florida and off Caribbean enables a broader understanding of the northeast slope of the Little Bahama Bank reported and potential deep water corals in the (Figure 8.1) (Neumann and Ball 1970; Neumann U.S. Caribbean. et al. 1977; Messing 1990; Reed et al. 2005). Other submersible research of the region includes 1Ocean Research and Education Foundation the following: to 300 m (the “sub reef habitat”) 1300 Galiano St., Coral Gables, FL 33134 at Discovery Bay Jamaica (Lang 1974; Hartman 1973; Lang et al. 1975); to 300 m in Belize 2Corresponding author addres: Marine Conserva- (Ginsburg and James 1979); research by Harbor tion Biology Institute, Public Policy Office 600 Pennsylvania Ave SE, Suite 210, Branch Oceanographic Institute (HBOI) to 1,000 Washington DC 20003 m for sites in the Greater and Lesser Antilles and Netherlands Antilles (Reed pers. comm.).
307 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION CARIBBEAN Study area showing geography and United States territories. The approximate boundaries of U.S. Caribbean EEZs are identified by the Study area showing geography and United States territories. Figure 8.1. Islands; B) Navassa Island. A) Puerto Rico and the U.S. Virgin blue dashed line:
308 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION CARIBBEAN
Figure 8.2. Regional currents, channels, and passages (adapted from Gyory et al. 2005a and b).
Submersible research for the U.S. Caribbean II. GEOLOGICAL SETTING includes dives by HBOI and NOAA (Nelson and Appeldoorn 1985). The Caribbean Sea basin, the islands within it, and the Bahamas Archipelago, consisting of In 1979 Stephen Cairns published a seminal shallow banks and inter-bank depressions, make review of deep-water stony corals occurring in the up this region (Figure 8.1). While most of the Caribbean Sea and adjacent waters, examining area is centered on the Caribbean plate several eighty-eight species of deep-water corals (both continental plates meet in this region, which are solitary and colonial species) (Cairns 1979). hotspots for seismic activity. Interactions between the Caribbean plate, North American plate and Recent regional deep-sea, coral related, the South American plate produced the major investigations include the following: research on topographic features and allow the Caribbean the distribution of deep water coral mounds in basin to be divided into four smaller basins; the the Straits of Florida by the use of high-resolution Yucatan, Colombian, Venezuelan and Granada multibeam sounding (Correa et al. 2006; McNeill basins (Figure 8.1). 2006); continued private pharmaceutical research throughout the region and especially in the Both tectonic activity and biological activity Straits of Florida interests (Reed pers. comm.); produce features significant to deep water corals research on azooxanthellate coral communities in this region. Tectonic activity is responsible for in Colombia’s Caribbean waters, off the San deep water trenches, ridges, basins and inter- Bernardo-Rosario and Santa Marta Islands island passages and channels (topographic (Reyes et al. 2005; Santodomingo et al. 2006); features are detailed in section III). Products of and an ongoing effort to characterize seafloor biological activity include deep coral mounds and habitats down to 1,000 meters within the U.S. lithoherms (discussed in section IV). Virgin Islands and Puerto (NOAA 2007).
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III. OCEANOGRAPHIC SETTING coastline of South America (Fratantoni 2001). The current then flows to the northwest, over the The Caribbean Sea is approximately 2,640,000 Colombian basin, towards the Nicaragua Rise km2 or 1,425,485 square nautical miles. It is a and a trough southwest of Jamaica, with a branch semi-enclosed basin, bounded on the west by forming an counterclockwise Panama-Colombia Central America, on the south by Central and Gyre (where the current meets Central America). South America, on the east by the Lesser Antilles Strong flow (up to 60 cm/sec) has been recorded and on the north by the Greater Antilles. With along the Panamanian and Colombian coasts a few exceptions, the Caribbean Sea generally (Gyory et al. 2005a). exceeds 1,830 m (6,000 ft) in depth with many sections exceeding 3,660 m (12,000 ft) in depth. Where currents are diverted by or constricted Its greatest known depth is 7,535 m (24,721 ft) between landmasses, current velocities increase. below sea level in the Cayman Trench located The larger and deeper inter island spaces produce between Cuba and Jamaica. The Puerto Rican major seaways; the more numerous, smaller CARIBBEAN Trench, the deepest area within the study region and shallower ones produce similar but smaller and deepest part of the Atlantic Ocean, lies north increases in velocity. Current velocities will of Puerto Rico just outside the Caribbean, with a also increase when water flow meets elevations depth of 9,200 m (30,183 ft). The only notable of the sea floor, typically platforms or banks of extensive shallow areas are the Bahamas Banks calcareous deposits. For example, the passages and the Nicaragua Rise. Several gaps between between the large and long-lived platforms of the islands on the north and east are major channels Bahamian Archipelago and the Florida Peninsula and passes for ocean currents connecting the are sites of increased high flow. Caribbean Sea with the Atlantic Ocean and Gulf of Mexico. Chief among these are the Windward At the Nicaragua Rise channel-like constrictions Passage between Cuba and Hispaniola; the between shallow banks accelerate current. The Mona Passage between Puerto Rico and flow and water is accelerated through the trough Hispaniola; the Anegada Passage south of the southwest of Jamaica and then turns west as it Virgin Islands; smaller channels between islands crosses the Cayman Ridge and enters Yucatan in the Windward Island chain; and the Grenada Channel as the Yucatan Current (Gyory et al. Passage between the Lesser Antilles and the 2005d). Strong currents (170 cm/sec) have been South American continent (Figure 8.2). reported in the Yucatan Channel (Pillsbury 1890; Gyory et al. 2005d). Water flow exits the Yucatan As a result of the exchanges with the open Atlantic, Channel into the Gulf of Mexico and doubles back there is little seasonal variation in surface water as the Loop Current (Gyory et al. 2005c) before temperatures, with a general range from 25.5o C entering the Straits of Florida between Western in the winter to 28 o C in the summer. Below the Great Bahama Bank and the Florida Reef Tract surface, water structure is highly stratified in the where it is joined by waters passing through the upper 1200 meters; weakly stratified between Old Bahama Channel to form the Florida Current 1,200 and 2,000 meters and uniform below 2000 (Gyory et al. 2005b). Gyory et al. (2005b) reports meters. This stratification is directly related to the strong surface flow in the Florida Current; “at shallow depth of passages between the Lesser times there is a 2 m/sec flow within a few miles Antilles. They act as sills and impede deep-water of the (Florida) coast.” When this Florida Current flow into the Caribbean (Gordon 1967; Gyory et exits the Straits it is joined by the Antilles Current al. 2005a). and turns eastward and becomes the Gulf Stream sensu stricto. Most water flows into the Caribbean Sea from the Atlantic through the Grenada, St. Vincent and The Antilles Current transports tropical waters St. Lucia Passages in the southeast (Figure 8.2) from the Atlantic northwestward, flowing north (Johns et al. 2002; Gyory et al. 2005a). From of the Greater Antilles and joins the Gulf Stream these passages, water flows clockwise across the off Florida past the outer Bahamas (Figure 8.2) Caribbean Sea basin as the Caribbean Current, (Rowe et al. 2005). Its waters are concentrated the main surface circulation in the Caribbean into a strong northward jet centered at 400 meters Sea. It reaches high surface velocity (70 cm/ deep about 80-100 km wide (Lee et al. 1996). It sec) along the Netherlands Antilles and adjacent is a considerable source of warm water for the
310 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Appendix 8.1. CARIBBEAN Appendix 8.2 (some locations overlap). References are listed in Distribution of major and potential structure-forming stony corals (scleractinia) deep sea bio-buildups (lithoherms, mounds, banks). Figure 8.3. The letter S identifies the locations of trawl stations referred to in
311 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION CARIBBEAN Reported distribution of potential habitat-forming corals (larger gorgonians, etc.). References include CITES 2007 and those listed in Figure 8.4. Appendix 8.1.
312 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION CARIBBEAN
Figure 8.5. Distribution of deep water features and corals found in the U.S. Caribbean and adjacent area. References include EarthRef.org 2007 and those listed in Appendix 8.1.
Gulf Stream system. However, it occurs more Deep-sea corals of interest to this chapter, are as an eddy field along the Bahamas Archipelago those termed “structure-forming” and “potential rather than as a constant flow (Gunn and Watt habitat-forming”. Structure-forming corals are 1982). associated with deep-sea coral habitat such as coral banks, bioherms and lithoherms. Both structure and non-structure forming corals occur IV. STRUCTURE AND HABITAT-FORMING within the study area (Figures 8.3, 8.4 and DEEP CORALS 8.5). Bayer (1961) describes the importance of octocorals (gorgonians and soft corals) in the reef A wide variety of deep-sea corals occur within building process and framework of shallow reefs. the wider Caribbean region. These include stony The importance of octocorals in deep water reefs corals, gorgonians, soft corals, stylasterids, is not yet understood but may be great in terms of black corals, lithotelestid coral and sea pens. contribution to framework and baffling of marine The knowledge about the distribution of these sediment, larger octocorals are expected to corals is generally considered poor, greater for contribute and baffle more than smaller species. some groups than others. Cairns (1979) notes, Appendix 8.1 details the attributes of each coral with regard to deep-sea stony corals, that the species including distribution. southern coasts of the Greater Antilles (Cuba and Hispaniola) and Costa Rica are not well sampled a. Stony corals (Class Anthozoa, Order and that the Straits of Florida (including the Scleractinia) northwest coast of Cuba) and the Lesser Antilles Ninety-nine species of azooxanthellate hard are well sampled. Many species of deep-sea corals are reported from the study area in depths corals occur within the U.S. Caribbean. Species below 100 m. Most are small single polyp species, with an Antillian distribution (occurring within the shaped like cups, which often occur solitarily (e.g. Greater and Lesser Antilles) may also occur in Figure 8.6). Solitary corals are not reported to form U.S. Caribbean waters. significant structures. Nine species of structure-
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building and branched azooxanthellate hard corals - 8.10 and 8.15 describe and illustrate structure- are reported from the study area: Dendrophyllia building corals. alternata, Desmophyllum dianthus (also known as Desmophyllum cristagalli), Enallopsammia Six species of azooxanthellate hard corals profunda, Enallopsammia rostrata, Lophelia are regarded as the major structure-forming pertusa, Madracis myriaster, Madrepora oculata, species: E. profunda, E. rostrata, L. pertusa, M. Madrepora carolina and Solenosmilia variabilis carolina, M. oculata and S. variabilis. Two genera (Cairns pers. comm.). Table 8.1 and Figures 8.7 (Lophelia and Solenosmilia) constitute two of
Table 8.1. Structure-forming deep water stony corals of the wider Caribbean region. The measurements given in brackets correspond to the approximate horizontal length for each specimen illustrated.
Species Description Illustration D. alternata occurs throughout the North Atlantic. This coral CARIBBEAN forms bushy colonies up to 1 m in height and has been described as a reef builder. It occurs at depths of 279 - 900 Dendrophyllia m in the tropical western Atlantic. Regionally, it has been alternata recorded with M. oculata and M. myriaster (Cairns 1979; Cairns and Stanley 1982; Le Goff-Vitry 2003). [30 cm, specimen recovered at 290.8 m off Curacao] D. dianthus (also known as D. cristagalli) forms densely branched colonies is reported to contribute to reef formation. It occurs globally to 2,460 m and in the tropical western Atlantic Desmophyllum from 155 - 1,939 m. Regionally, it has been recorded with dianthus E. profunda, L. pertusa, M. oculata, and S. variabilis (Cairns 1979; Cairns and Stanley 1982; Rogers 1999). [10.8 cm, specimen recovered at 597 m off the east coast of Florida] E. profunda occurs globally and is endemic to the western Atlantic. It forms massive bushy colonies, up to 1 m thick, and is regarded a major structure-forming species. It occurs Enallopsammia from 403 - 1,748 m and is often associated with L. pertusa, profunda M. oculata and S. variabilis (Cairns 1979; Cairns and Stanley 1982; Rogers 1999; Reed 2002; Freiwald et al. 2004). [30.7 cm, specimen recovered at 801 m in the Straits of Florida off Cuba] E. rostrata occurs in the U.S. Caribbean and globally. This coral forms massive bushy colonies and is regarded a major structure-forming species. It occurs from 300 - 1,646 m in the tropical western Atlantic. It is associated with L. pertusa on Enallopsammia deep-water reefs in the north Atlantic and has been recorded rostrata with S. variabilis, M. oculata, and E. profunda in the wider Caribbean (Cairns 1979; Rogers 1999; Reed 2002; Freiwald et al. 2004). [31.4 cm, specimen recovered at 1,097 m off Bermuda. Specimen includes L. pertusa and D. dianthus]
L. pertusa occurs in the U.S. Caribbean, globally, and is well known from the north Atlantic. This coral forms bushy colonies up to a meter in height and is regarded the most common reef-building deep/cold-water coal. It has been reported from waters as shallow as 25 m to as deep as 3,383 m. Regionally, Lophelia it is most common between 500 - 800 m. L. pertusa is often pertusa found associated with E. profunda and has been recorded with D. dianthus, M. carolina, M. oculata, and S. variabilis in the wider Caribbean (Cairns 1979; Cairns and Stanley 1982; Rogers 1999; Dawson 2002; Reed 2002; Freiwald et al. 2004). [24.3 cm, specimen recovered at 793 m in the Straits of Florida off Cuba]
314 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
Species Description llustration
M. myriaster occurs in the U.S. Caribbean and throughout the wider Caribbean. It forms bushy colonies and is reported as a main framework builder of deep-water coral banks in Madracis Colombian waters. Regionally, it has been recorded with M. myriaster carolina. It occurs from 37 - 1220 m in the tropical western Atlantic (Cairns 1979; Reyes et al. 2005; Santodomingo et al. 2006). [30.2 cm, specimen recovered at 200 m off Jamaica]
M. carolina occurs in the U.S. Caribbean and throughout the tropical western Atlantic. This coral forms bush-like colonies with a thick base and is��������������������������������� associated with cold-water reefs as a structure-forming species���������������������������. It occurs from 53 - 801 m Madrepora (most commonly between 200 - 300 m). Regionally, it has carolina CARIBBEAN been recorded with E. profunda, E. rostrata, and D. dianthus (Cairns 1979; Dawson 2002; Freiwald et al. 2004). [27.6 cm, specimen recovered at 333 - 375 m in the northwest Providence Channel off Grand Bahama Island] M. oculata occurs in the U.S. Caribbean and globally. This coral forms large bushy colonies and is regarded a major structure-forming species. It occurs from 55 - 1,950 m. It is commonly associated with L. pertusa in the eastern and Madrepora northeastern Atlantic and Mediterranean Sea. Regionally, it oculata has been recorded to occur with E. profunda, E. rostrata, D dianthus, L. pertusa, and S. variabilis (Cairns 1979; Rogers 1999; Dawson 2002; Reed 2002; Freiwald et al. 2004). [24.1 cm, specimen recovered at 634 m in the Gulf of Mexico] S. variabilis occurs globally. This coral forms bushy colonies and is regarded a major structure-forming species. It occurs from 220 - 2,165 m and is known from the northern slope of the Little Bahama Bank from 1,000 - 1,300 m, where it is Solenosmilia associated with deep-water banks. It is often found associated variabilis with L. pertusa, Madrepora spp., and E. profunda, and has been recorded with E. rostrata and D. dianthus in the wider Caribbean (Cairns 1979; Mullins et al. 1981; Cairns and Stanley 1982; Rogers 1999; Freiwald et al. 2004). [20.7 cm, specimen recovered at 1,000 m in the North Atlantic Ocean]
Figure 8.6. Assorted solitary deep water stony corals: a) Stephanocyathus diadema, from The Bahamas at 1,779 m; b) S. laevifundus, from St. Vincent at 576-842 m; c) Caryophyllia ambrosia caribbeana, from Surinam at 1,318-1,343 m; d) Deltocyathus eccentricus, from Jamaica at 457-558 m; e) Bathypsammia tintinnabulum, from the Straits of Florida at 689-699 m (B. tintinnabulum is not known from the region but is presented here for comparison).
315 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
Close up of the potential structure- CARIBBEAN Figure 8.9. forming coral D. alternata, sampled by submersible, Figure 8.7. Deep sea coral habitat (M. oculata and from the Netherlands Antilles, at 290.8 m. Paramuricea sp.) at 2,230 m in the Bahamas (Straits of Florida off Cat Cay). Photo credit: Reed 2006.
Figure 8.10. Close up of the major structure-forming coral M. oculata, sampled by submersible from the Straits of Florida at 746.4 m. Photo credit: Reed 2006.
reported from the U.S. Caribbean. Figure 8.8. Structure-forming coral (E. profunda) providing habitat for unidentified crustaceans and a crinoid. Photographed from submersible at 2,721 m Another major structure-forming species, Oculina in Straits of Florida. Photo credit: Reed 2006 varicosa, occurs throughout the Caribbean. O. varicosa is a facultative zooxanthellate species the four major constituents of known world-wide (i.e. it occurs with zooxanthellate in shallow deep-water coral banks at depths of 400 - 700 m water and without in deeper water). O. varicosa (Cairns and Stanley 1982). Both M. oculata and is reported to 152 m and forms bushy colonies M. carolina are regarded as minor constituents of several meters in height (commonly at 45 - 91 coral banks in the Straits of Florida (Cairns 1979). m). It is described as a main framework builder Solenosmilia is the dominant structure-building of deep-water banks off Florida’s central eastern coral found on deep coral banks north of the Little coast. However, as no deep water banks of Bahama Bank (Reed 2002) (Table 8.2, Mullins O. varicosa have been reported from the wider Reefs), where Madrepora and Enallopsammia Caribbean (only shallow water species) this also occur. Lophelia and Enallopsammia are species is not considered further. common components of the lithoherms east of the Little Bahama Bank in the Straits of Florida Madracis myriaster, an azooxanthellate stony (Reed 2002) (Table 8.2, Bahamas Lithoherms). coral found throughout the wider Caribbean, Four major structure-forming species E. rostrata, is reported to form bushy shaped colonies and L. pertusa, M. oculata and M. carolina have been significantly contribute to habitat formation on 316 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
Table 8.2. Scleractinian-dominated coral habitats (“reefs”) of the wider Caribbean. Locations are identified in Figure 8.3.
Location Description This site is characterized by a region of deep coral mounds (bioherms) on the northern deep shelf edge of the Little Bahama Bank at depths of 1000-1300 m. Mounds 5-40 m in height are reported. The bioherms are dominated by the major structure forming deep Mullins Reefs water coral Solenosmilia sp., Lophelia is reportedly absent. An area of approximately 1500 km2 is described. Site profiled via. submersible, seismic reflections, and dredges (Mullins et al. 1981; Reed 2002). This site is characterized by a region of lithoherms on the western shelf edge of the Little Bahama Bank in the Straits of Florida at depths of 300-500 m. Pinnacles 30-50 m in height are reported. The lithoherms are dominated by the major structure forming deep water coral L. pertusa. E. profunda and M. oculata are also present. D. alternata, A. fecunda, and Bahamas
the larger black coral Distichopathes filix have been reported from the region. An idealized CARIBBEAN Lithoherms lithotherm from this site is illustrated in Figure 8.23. Wreckfish have been reported in association with these lithoherms. An area of approximately 330 km2 is described. Site profiled via. submersible, seismic reflections, and dredges (Neumann and Ball 1970; Neumann et al. 1977; Messing et al. 1990; Reed 2002). Deep-water mounds (banks and lithoherms) and cemented hardgrounds are described off Mullins Bimini and entering the New Providence Channel. The mounds are described to continue Mounds northward and connect to the Bahamas Lithoherms. Site profiled via. coring and seismic reflections (Mullins and Neumann 1979). Deep water mounds are described at 500-800 m along the western shelf edge of the Great Santaren Bahama Bank, running from the Santaren Channel to the Straits of Florida. Mounds >30 m Mounds were reported. Site profiled via. multi-beam side scan sonar and drop camera (Correa et al. 2006; Grasmueck et al. 2006). This site is described as a deep water coral bank on the northwestern shelf of Colombia at a depth of 200 m. The bank is dominated by the potential structure-forming deep water coral M. myriaster. Anomocora fecunda, Coenosmilia arubuscula, and Polymyces fragilis Santa Marta were reported abundant. Black corals and octocorals were reported numerous and include Bank Antipathes columnaris, Elatopathes abetina, Sticopathes spp., Chrysogorigia desbonni, and Nicella guadalupensis. Site profiled via echosounding and dredges (Reyes et al. 2005). This site is described as a deep water coral bank on the northern shelf of Colombia at a depth of 155-160 m. The bank is dominated by the potential structure-forming deep water coral M. myriaster. Other species considered to contribute to habitat include A. fecunda, C. arubuscula, Eguchipsammia cornucopia, M. oculata, Javania cailleti, Caryophyllia San Bernardo berteriana, P. fragilis, Thalamophyllia riisei, Oxysmilia rotundifolia, and other Madracis Bank species. Flounder, basslets, and scorpionfish were the most abundant species of fish recovered. Madracis thickets and coralline mounds (up to 10 m in height) are reported. Site profiled via. echosounding, dredges, and grab sampling (Reyes et al. 2005; Santodomingo et al. 2006). deep-water coral banks off Colombia (Reyes et al. community occurs at less than 100 m in depth, C. 2005; Santodomingo et al. 2006). Investigations debilis is not considered further. with sonar profiling and trawl and grab sampling suggest deep water banks from two locations Two other potential structure-forming species, D. off Columbia’s continental shelf at depths of alternata and D. dianthus are found throughout 155 - 200 m; Santa Marta and San Bernardo the wider Caribbean, but have not been reported Banks. Table 8.2 details these locations. Relief, from the U.S. Caribbean. D. alternata is found “suggestive of reef structures”, was reported. M. throughout the North Atlantic and forms bushy myriaster occurs in the U.S Caribbean (Figure colonies to 1 m in height. It has been described as 8.5). Reyes et al. (2005) also report a shallower a reef builder (Le Goff-Vitry 2003) and has been azooxanthellate coral community off Colombia at recorded with M. oculata and M. myriaster in the a depth of 70 m. Cladocora debilis was noted as wider Caribbean. D. dianthus is reported to form the main matrix builder of this community. As this densely branched colonies and to “contribute to
317 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
Figure 8.11. Large deep-sea black coral (Leiopathes glaberrima) recovered from 304 m in the North Atlantic (approximately 116 cm in height). CARIBBEAN
Figure 8.13. Bubblegum coral (Paragorgia johnsoni) sampled by submersible from 620 m west of the Little Bahama Figure 8.12. Large deep water gorgonian Bank in the area of (Paracalyptrophora sp.) and deep water sponge Bahamas Lithoherms (Pachastrellidae sp.). Photographed from (approximately 51 cm submersible at 2,193 m in The Bahamas (Straits in height). Photo credit: of Florida) (gorgonian is approximately 53.5 cm in Amy Wolfrum. height). Photo credit: Reed 2006.
Figure 8.15. Bamboo coral (Keratoisis flexibilis) on deck and utilizing a deep water structure-forming coral (E. profunda) Figure 8.14. Large deep water gorgonian as substrate (Paramuricea sp.) providing habitat for an (bamboo coral unidentified crustacean. Photographed from is approximately submersible at 2631 m in The Bahamas (Straits of 34 cm in height). Florida off Cat Cay) (gorgonian is approximately 33 Coral recovered cm in width). Photo credit: Reed 2006. from 806.5 m by submersible. Photo credit: Reed 2006.
318 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
is considered habitat-forming as colonies 1 m tall and 1.5 m across were reported. Gerardia sp. was found in dense stands on lithoherm crests, where current flow was the greatest.
d. Gorgonians (Class Anthozoa, Order Gorgonacea) Approximately one hundred and forty-seven species of gorgonians are reported from the wider Caribbean at depths below 100 m. Thirteen species are reported from the U.S. Caribbean (more are expected). Forty-four species are considered potential habitat-forming species,
with twelve occurring in U.S. Caribbean waters: CARIBBEAN Figure 8.16 Large deep water gorgonian Nicella Acanthogorgia goesi, Callogorgia americana americana, collected from 165 m off the Dominican Americana, Diodogorgia nodulifera, Ellisella Republic (approximately 53 cm in width). barbadensis, E. elongata, Narella bellissima, N. pauciflora, N. deichmannae, N. obesa, N. reef formation” (Cairns and Stanley 1982). D. guadelupensis, Riisea paniculata and Swiftia alternata and D. dianthus are detailed in Table exserta. The maximum reported height of a 8.2. wider Caribbean deep water gorgonian is 300 cm (Pseudoplexaura porosa), and 244 cm for U.S. The distribution of major and potential structure- Caribbean waters (E. barbadensis). Two species forming species in the study area generally of red or pink corals (family Coralliidae); 13 species follows the Antillian arc and the margins of the of bamboo coral (family Isididae); one species continental shelves of South and Central America of bubblegum coral (family Paragorgiidae); and (Figure 8.3). 34 species of red tree coral (family Primnoidae) are known from the region. Detailed distribution b. Black corals (Class Anthozoa, Order data (beyond occurring in the western Atlantic) Antipatharia, Families Cladopathidae and was available for only 54% (82) of known deep Schizopathidae) water gorgonians. Height and/or width data Approximately thirty-two species of antipatharia was available for 34% (50) of known deep water are reported from the wider Caribbean in depths gorgonians. It is expected that many more below 100 m, with thirteen considered potentially gorgonians are potential habitat-forming species. important habitat-forming species. Eight species Figures 8.12 - 8.16 and 8.17 illustrate deep-sea are reported from the U.S. Caribbean, with five gorgonians. considered potentially important habitat-forming species: Antipathes americana, A. caribbeana, Plumapathes pennacea, Tanacetipathes hirta, Parantipathes tetrasticha. Opresko and Sanchez (2005) list two regional black coral species of commercial importance: A. atlantica and A. gracilis. Both species are considered potentially habitat-forming. The maximum reported height of deep water antipatharians occurring in U.S. Caribbean waters is 61 cm (P. tetrasticha). Figure 8.11 illustrates a large deep-sea black coral. c. Gold corals (Class Anthozoa, Order Zoanthidea) One species of gold coral is reported from the Figure 8.17. Deep water coral habitat at 829.4 m in The Bahamas (Straits of Florida). A variety of deep- Caribbean region. Messing et al. (1990) describe sea corals are illustrated, including the soft corals a large zoanthidean tentatively identified as Anthomastus agassizi (red in color) and Pseudodrifa Gerardia sp. in Bahamian waters. This species nigra (brown in color). Photo credit: Reed (2006).
319 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION CARIBBEAN Figure 8.20. Deep-sea stylasterid (Stylaster Figure 8.18. Sea pens (Renilla reniformis), from miniatus) providing substrate for a solitary deep Curacao, Netherlands Antilles (each approximately 2 water stony coral (Paracyathus pulchellus) and other cm in width). This species occurs to 108 m or more. unidentified invertebrates, the Straits of Florida, at 191 m (stylasterid is approximately 10 cm in width). e. True soft corals (Class Anthozoa, Order Alcyonacea and Suborder Stolonifera) Caribbean: Renilla reniformis, with a maximum Twenty species of true soft corals (Alcyonacea) reported height of 6.5 cm it is not considered an are reported from the wider Caribbean in important habitat-forming species (illustrated in depths below 100 m. Only one is considered Figure 8.18). It is bioluminescent. The maximum a potentially important habitat-forming species; reported height of a deep water pennatulacean Neospongodes portoricensis, with a maximum occurring in the wider Caribbean is 210 cm, for reported height of 30 cm. Only one is reported Funiculina quadrangularis. According to Picton from the U.S. Caribbean: Stereotelesto corallina. and Howsen (2002) F. quadrangularis is often Detailed distribution data was available for 35% found in very large colonies. This order contains (7) of known deep water alcyonaceans. Figure the deepest deep-sea corals, many occur below 8.17 illustrates a deep-sea soft coral. 1,000 m, with one species (Umbellula magniflora) reported from 6,300 m in the Caribbean Sea f. Pennatulaceans (Class Anthozoa, Order basin. Most deep water sea pens are found on Pennatulacea) basin floors, rises and trenches throughout the Pennatulaceans, or sea pens, live in soft- Caribbean region including the Caribbean Sea sediment and are known to form extensive basin, Aves Ridge and Puerto Rico and Cayman groves in some areas (Morgan et al. 2006). Eight Trenches. Height and width data were available deep-water sea pens are reported from the wider for 25% (2) of known deep water sea pens. Caribbean. One species is reported from the U.S. g. Stylasterids (Class Hydrozoa, Order Anthoathecatae, family Stylasteridae) Forty-one species of stylasterids are reported from the wider Caribbean at depths below 100 m. Only one is regarded as a potential habitat- forming species; Distichopora sulcata (Figure 8.19), with a maximum reported height of 25 cm and width of 30 cm, endemic to the waters off northwest Cuba. However, additional larger stylasterids are likely - during a submersible dive James and Ginsburg (1979) recorded a larger unidentified stylasterid measuring “about 30 cm Figure 8.19. Large deep water stylasterid across” growing out from a rock wall at 290 m (Distichopora sulcata) dredged off Havana, Cuba off Belize Glovers Reef). Fifteen smaller (non (approximately 33 cm in width). habitat-forming) species (<25 cm in height or
320 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION width) occur in U.S. Caribbean waters. All wider and 8.10). These bush-like aggregations can Caribbean stylasterids are referenced by Cairns form habitat on suitable substrate and contribute (1986). to the formation of deep sea bio-buildups through their skeletal debris and by trapping Puerto Rican waters contain two stylasterid pelagic sediment. Two types of deep-sea bio- species: S. atlanticus and S. spatula. Stylasterids buildups are reported from the wider Caribbean have been recorded with many other species of region (deep coral mounds and lithoherms); their deep water corals and can serve as substrate morphologies and compositions, and deep-sea allowing invertebrates to gain a foothold (Figure coral banks are discussed below. Seamounts 8.20). are also discussed.
Deep Coral Habitat Types Deep Coral Mounds A variety of terms have been used for the
Deep sea corals often occur as individual and/ unconsolidated sea floor elevations with CARIBBEAN or scattered colonies on the sea floor (Cairns which deep-sea corals occur. In general these 1979, 2000). More numerous colonies can form structures are described as “capping mounds of clumps or thickets of varying density (Mullins et unconsolidated sediment and coral rubble, which al. 1981; Messing et al. 1990) (Figures 8.7, 8.8 are often built on an underlying lithified substrate”
Figure 8.21. Idealized regional deep coral mound (not to scale). Associated benthos as follows: 1) Etmopterus sp. (dogfish sharks); 2) pteropods; 3) Brotulid sp. (fangtooths); 4) Diretmus sp. (dorys); 5) Myctophid sp. (lanternfish); 6) Macrouid sp. (grenadiers); 7) Stomias sp. (dragonfish); 8) Hexactinelida sp.; 9) Cirripathes sp.; 10) S. varibilis; 11) M. oculata; 12) Galatheid sp. (deep-sea crabs); 13) crinoids; 14) gorgonians; 15) Iridogorgia sp.; 16) Antipathes sp.; 17) Macruan sp.; 18) Keratoisis sp.; 19) Munida sp. (squat-lobsters); 20) Stylasteria sp.; 21) Nematocarcinidae sp. (deep-sea shrimp); 22) Glyphocrangon sp.; 23) solitary corals; 24) Chrysogorgia sp.; 25) Synaphobranchus sp.; 26) Lithodid sp.; 27) Nymphaster sp.; 28) porifera; 29) Alcyonaria sp.; 30) Nephropsis rosea; 31) Holothurian sp. 32) Ophiuroid sp.; 33) Turrida sp.; 34) Cruriraja sp. (skates). Sources listed in Appendix 8.2.
321 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
The most extensive known occurrence of deep coral mounds reported in the Caribbean is found on the northern slope of Little Bahama Bank at depths between 1,000 to 1,300 m (Mullins et al. 1981). Here, where bottom currents were reported to be as much as 50 cm/sec and temperatures of 4-6o C, numerous banks are inferred to occur in an area of 2,500 km2 with relief over the surrounding sea floor ranging from 5-40 m (ibid). The surface sediments of these mounds consist predominantly of pelagic organisms such as planktonic foraminifera and pteropods, with small amounts of coral skeletal debris and other invertebrates in the coarser CARIBBEAN grain sizes (ibid). These mounds appear to be developed by the baffling action of the thickets of deep corals that traps pelagic sediments and build on unconsolidated mounds. Other coelenterates found on these mounds include gorgonians, alcyonaceans and antipatharians (Figure 8.21). Mullins et al. (ibid) present a useful schematic model of the sequential development of these mounds (Figure 8.22) based on previous publications (Squires 1964; Neumann et al. 1977). Figure 8.22. Hypothetical development sequence of deep coral mounds (not to scale). Figure adapted Figure 8.3 illustrates the distribution of deep coral from Mullins et al. 1981, based in part on Squires mounds. Table 8.2 details these occurrences. 1964 and Neumann et al. 1977. Lithoherms In contrast to the unconsolidated deep coral (Reed 2002). These structures have been called mounds, lithoherms (Neumann et al. 1977; deep-water coral banks; deep-sea coral mounds; Messing et al. 1990; Reed 2002) are composed colony thickets; coral thickets; coral structures, of varied communities of benthic organisms deep-water coral reefs; and bioherms (Stetson growing on successive crust-like layers of friable et al. 1962; Squires 1964; Neumann et al. 1977; limestone. Lithoherms have also been referred Mullins et al. 1981; Reed 2002). The accepted geological term for mounds rich in unbound skeletal debris is biostrome (Bates and Jackson 1987). In what follows we use “deep coral mounds” for this type of buildup.
In the study region these deep coral mounds occur in areas with strong currents and/or upwelling where they accrete by a combination of coral skeletal debris and trapped suspended sediment swept into coral structures (Mullins et al. 1981). The resulting structures range in height over the surrounding sea floor from a few meters to 50 m Figure 8.23. Idealized lithoherm of the Straits of Florida (not to scale). (Reed 2002). Figure adapted from Messing et al. 1990.
322 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION to as bioherms, deep-water biohermal buildups Seamount; Mona Seamount; and Dos Ni’os Knoll. and muddy carbonate buildups (Neumann et Figure 8.5 illustrates their locations. The potential al. 1977; Reed 2002). Regionally, they are that seamounts throughout the region are home reported only from the eastern side of the Straits to deep water coral ecosystems deserves further of Florida at depths between 500 and 700 m attention. below sea level. According to Messing et al. (1990) the lithoherms are elongate mounds up to Relevant Geological and Oceanographic 300 m long and 50 m high. Their surfaces show Processes successive friable limestone layers composed of fine-grained carbonate skeletal sediment with Five processes play significant roles in the variable amounts of coral debris. The cement of distribution and composition of bio-buildups: (1) the crust-like layers of lithoherms is composed of foremost is amplification of current velocities magnesium calcite with 10-14 mole % magnesium by submarine topography; (2) the biological
(Wilbur and Neumann 1993). production of skeletal debris; (3) physical trapping CARIBBEAN and stabilization of sediment by thickets of corals, Messing et al. (1990) describe and illustrate (Figure sponges and alcyonarians to build elevated 8.23) a distinct, tripartite biozonation of one of accumulations; (3) availability of hard substrate these lithoherms, which they ascribe to variations for recruitment of corals; and (4) the submarine in current velocity produced by the mounds, also cementation that produces lithoherms. Process 3 described by Genin et al. (1986). On the steeper is the most significant for deep water banks. margin or side facing into the current (upcurrent margin) where velocities may reach 100 cm/ Increased current velocities appear essential for sec, corals (L. pertusa) predominate (“corals” in flourishing deep corals both because they can Figure 8.23). A zone of zoanthids forms along locally sweep away sediment to expose the hard the upcurrent crest (gold coral = “zoanthids” in substrate that encourages settlement of coral Figure 8.23) and crinoids and alcyonarians occur larvae and they provide a supply of plankton behind this zone (“crinoids and alcyonarians” in on which the corals feed. In addition, increased Figure 8.23) and on the flanks of the lithoherms currents bring a continued supply of pelagic (Genin et al. 1986). Figure 8.3 illustrates the sediment to help build local relief (mounds and distribution of known lithoherms. Table 8.2 details lithoherms). these occurrences. Two kinds of submarine topography can Deep-sea Coral Banks accelerate current velocities. One, where water Reyes et al. (2005) describes two azooxanthellate flow meets or is constricted between land masses, deep water coral habitats (banks) at depths for example, the seaways or channels between greater than 100 m off Colombia’s Caribbean islands of the Lesser Antilles or Windward shelf. Substrate included hard bottom limestone Islands (Figure 8.2). The larger and deeper inter (fossilized reef) and metamorphic rocks. It is island spaces produce the major seaways; the unclear to what extent these corals are building more numerous, smaller and shallower ones upon themselves (i.e. as with bio-buildups) or produce similar but smaller increases in velocity. simply opportunistically taking advantage of A second kind of topographic constriction occurs suitable substrate (limestone platform). Reyes et when water flow meets elevations of the sea al. (ibid) suggested similarities with shallow water floor, typically platforms or banks of calcareous reefs and stated a need for further investigations. deposits. The spaces between large and long- Figure 8.3 illustrates the distribution of known lived platforms of the Bahamian Archipelago and deep water coral banks. Table 8.2 details these the Florida Peninsula are sites of increased flow occurrences. velocities between adjacent platforms. A smaller scale example of topographic constriction of Seamounts currents is present in the series of shallow banks Deep sea corals are known as intimate associates of calcareous deposits, which have developed of seamounts throughout the world (Freiwald et on the Nicaragua rise and channel flow from the al. 2004; Clark et al. 2007). EarthRef.org (2007) Caribbean Sea contributing to the initiation of lists four seamounts occurring within the U.S. the swift Yucatan Current. Localized sediment Caribbean waters: Grappler Seamount; Whiting accumulation, either physical or as a result of
323 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
the trapping by thickets of erect invertebrates, is needed to produce sea floor relief.
The role of hard substrate in initiating coral settlement is ambiguous. On the one hand it is repeatedly cited as necessary for settlement of coral larvae (Freiwald et al. 2004, pp. 18, 21). However, in the same publication (ibid, pp 27- 28) occurrences are cited of deep-sea corals on mounds, which presumably are unconsolidated sediment. Mullins et al. (1981) described and illustrated a compelling example of corals growing on mounds of unconsolidated pelagic sediment from the margin of Little Bahama Banks. One CARIBBEAN Figure 8.24. Cross-section of a “typical” continental explanation for this apparent contradiction may slope for Puerto Rico and the U.S. Virgin Islands to be the minimal size of hard substrate required to approximately 400 m. Deep-sea research has been promote settlement. If for example, something limited and, to date, deep corals have been only as small as a bivalve shell is sufficient then sporadically reported from the region. Figure adapted deep-sea corals could establish on mounds of from Nelson and Appeldoorn 1985. unconsolidated sediments that included bivalve skeletons. Once a few corals are established includes limited ROV and submersible work off their continued production of skeletal debris Navassa Island (Littler and Littler 1999, Reed provides continually expanding hard substrate pers. comm.) and limited submersible work off for larval settlement (Figures 8.15 and 8.20 Puerto Rico and the U.S. Virgin Islands (Nelson illustrate the substrate use of deep-sea corals). and Appeldoorn 1985). Figure 8.24 illustrates Syndepositional, subsea cementation triggered a cross section of a “typical” continental slope perhaps by persistent strong currents (Straits of for Puerto Rico and the U.S. Virgin Islands, Florida) is required to produce current-resistant adapted from Nelson and Appeldoorn (1985). lithoherms and it may also be responsible for Intermittent zones of rubble, rocks and barren developing hard substrates on which deep coral sand with occasional sponges, hydroids, soft mounds develop. corals, hard corals, crinoids and gorgonians attached to rocky substrate are described. “Sea whip” type gorgonians (Ellisella sp.), often V. SPATIAL DISTRIBUTION OF CORAL wrapped with basket stars, were noted common SPECIES AND HABITAT at one location off the south coast of Puerto Rico. Rugged rock habitat, described as large, closely- The distribution of deep water corals generally packed boulders with little or no sand, was noted follows the Antilles and continental shelves of to primarily occur in the Mona Passage. Most Central, South and North America (Figures 8.3 - notably, an abundance of the potential structure- 8.5). Dawson (2002) analyzed the distribution of forming coral M. myriaster was reported at a 129 species of azooxanthellate (deep-sea) stony depth of 197-212 m on the shelf margin south of corals in the Caribbean and surrounding areas. the southwest corner of Puerto Rico. His findings include the following: the greatest diversity of species occurs at around 200-350 Ongoing deep water research in the U.S. m in depth; and the highest diversity of species Caribbean includes the exploration and is located at the northern islands of the Lesser characterization of seafloor habitats, with Antilles. ROVs, down to 1,000 meters within the U.S. Virgin Islands and Puerto Rico by NOAA (NOAA To date, no significant deep scleractinian coral 2007). This effort aims to characterize and map habitats have been discovered in the U.S. moderate to deep water coral reef ecosystem Caribbean. Deep-sea corals have been reported habitats within targeted study areas. Priority from submersible and ROV research, but only areas include the deep waters of the Buck Island sporadically. However, deep-sea research for Reef National Monument, the Escollo Grappler the region is still in its infancy. Previous research seamount to the southeast of Puerto Rico and
324 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION the La Parguera region along the southwestern larvae, the cosmopolitan distribution of shallow- coast of Puerto Rico. water benthos of coral reefs as well as many deep water coral species (including the major Inspection of regional unpublished research structure-forming corals) indicates long term reports and dredge station sorting sheets connectivity throughout the basin. illuminate interesting possibilities for deep coral habitat. Selected citations from dredge reports (individual station records) and notes include the VI. SPECIES ASSOCIATIONS WITH DEEP following: CORAL COMMUNITIES For the U.S. Caribbean: in Navassa Island waters two major structure forming corals (M. oculata and Fauna of deep-sea bio-buildups are among E. rostrata) were recovered, at 1,200 m, along with the least studied of marine assemblages. solitary corals (Stephanocyathus laevifundus and For example, the identification of deep-sea of the families Deltocyathus and Desmophyllum), gorgonians (of the genus Acanella, Iridogorgia gorgonians, collected in the Caribbean and the CARIBBEAN and Chrysogorigia) and deep water fish (Staiger Bahamas in the early 1960’s, are, to date, being and Voss 1970). revised (e.g., Cairns 2007); deep-sea Gold corals collected on lithoherms in the Bahamas in the For waters adjacent to the U.S. Caribbean: in 1970’s still await full identification (Messing pers. British Virgin Islands waters, at 205 - 380 m, comm.), and crustaceans collected in the 1880’s one potential structure-forming coral species have long waited proper scientific descriptions (M. myriaster) was recovered from the same (Larsen 1999). location in a trawl net “festooned with stalked crinoids”. Deep water fish and crustaceans were Inspection of archival data and grey literature, also recovered (Staiger 1969); in Saba waters, in 512 m off the northern edge of the Saba Bank, including original station data sheets, cruise a large deep water gorgonian (C. multiflora) was narratives and biological survey reports from recovered along with deep water fishes (Anon. 28 locations within the region identify a wide 1956); south of Hispaniola in 135-169 m, the assortment of associated species occurring with potential structure forming coral species (M. the major structure-building deep-water corals; E. myriaster) was recovered along with numerous profunda, L. pertusa, M. carolina, M. oculata and sponges, gorgonians and other invertebrates S. varibilis (Appendix 8.2). Figure 8.21 illustrates in a net noted as “We had to use the crane to several of these species in an idealized deep lift it aboard…. The catch was very good”. Deep coral mound. Many of the reported associated water fish and crustaceans were also recovered. species may grow to almost one meter in height Similar “excellent” and “magnificent” catches were also noted from two nearby trawl sets that also (larger cnidarians and porifera). recovered M. myriaster (Staiger and Voss 1970; Voss 1971). Large deep sea sponges (phylum Porifera) are a potentially important megafauna component of For waters throughout the region: in Venezuelan deep sea coral habitats. Deep sea sponges have waters with L. pertusa at 406 m - “it was the best been recorded with structure-forming and potential 10 ft. otter trawl tow I have seen in six years. habitat-forming corals (Appendix 8.2) and on There was a greater variety of organisms and a deep water mounds, lithoherms and banks. It is greater number than we usually get in a 41 ft. otter unknown what habitat role they may play on deep trawl tow” (Anon 1968); in the Straits of Florida, water coral habitats. Colin (1978) reports that in in waters just off just off Havana, Cuba, where M. carolina, M. oculata, and M. myriaster have been some areas of shallow water reefs the biomass recovered - “the “pentacrinus grounds” [crinoid of sponges “can exceed that of any other group, grounds] discovered by the Blake off Havana are including reef-building corals.” Lang et al. (1975) world famous” (Voss 1966b); and in the Straits of reported that sponges (sclerosponges) can act Florida just southeast of the area of lithoherms on as primary framework builders of reefs at depths the Little Bahama Bank) with L. pertusa, S. varibilis of 70-105 m off Jamaica. Of primary interest are and E. profunda at 738 m - “Tremendous amount the larger calcareous sponges (class Calcarea), of coral in net. Needed crane to lift it aboard. Good deep water glass sponges (class Hexatinellida), catch of fish and invertebrates” (Anon. 1973). demosponges (class Demospongiae) and Though no direct information was found sclerosponges (class Sclerospongiae). Deep-sea regarding the regional transport of deep coral sponges have been recorded up to 122 cm wide
325 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
Table 8.3. Potential fishing gear impacts on deep sea corals for the U.S. Caribbean.
Measure of Impact Current Overlap Overall Potential Potential Extent Current Fishery Geographic of use Rating Gear Type Severity of of Impact from Use in Region Extent of Use with coral of Gear Impact Fishing Gear in Region habitat Impact
Bottom Trawl None reported High High NA Uncertain NA Mid-water Trawl None reported Low Low NA Uncertain NA Dredge None reported High Low NA Uncertain NA Bottom-set Longline to 109 m Medium Low Unknown Uncertain Low Bottom-set Gillnet None reported Medium Medium Unknown Uncertain Low CARIBBEAN Longline traps or pots to 183 m High Medium To shelf edge Uncertain Low Single pots to 183 m Low Medium To shelf edge Uncertain Low Hook and line to 600 m Low Low Unknown Uncertain Low
at deep-sea coral depths in the Straits of Florida. not currently found in the U.S. Caribbean. No Figure 8.12 illustrates a deep-sea sponge. Deep- current examples were found of significant sea sponges are subject of pharmacological threats to deep water corals from the following: investigations. harvesting of precious coral, cable laying, oil and gas exploration, sedimentation, invasive species A number of fishes have been observed or and destructive fishing practices (such as bottom collected in association with deep coral habitats. trawling). While most of these are demersal species (Appendix 8.2), certain commercially important Fishing occurs to deep coral depths (>100 m) in pelagic species may also associate with deep- the U.S. Caribbean and throughout the region. sea coral habitat. Swordfish Xiphias ( gladius) This activity is mainly focused on deep water have been encountered by submersibles snapper and grouper and includes commercial visiting deep sea corals (Lophelia) in the Straits and recreational fishing effort. An extensive of Florida (Nizinski and Ross 2002) and in deep water fishery, primarily for snappers, occurs Bahamian deep waters at 600 m (Harbison and around Puerto Rico (Appeldoorn pers. comm.). Janssen 1987). Larger deep coral mounds and Target species include black snapper (Lutjanus lithoherms may play similar roles to seamounts, griseus), blackfin snapperL. ( buccanella), which are thought to attract pelagic fish because vermilion snapper (Rhomboplites aurorubens) of an “enhanced primary productivity due to and silk snapper (L. vivanus) (FAO 1993; Kojis particular oceanographic conditions supporting 2004; Cummings and Matos-Caraballo 2003). a rich ecosystem” (Allain et al. 2006). In effect, seamounts influence oceanic currents in a way Gear utilized in commercial deep water snapper that promotes plankton to develop, acting as a and grouper fishing in the U.S. Caribbean includes feeding source for resident fish, attracting larger vertical set line, bottom longline, handlines, pelagic predators. These pelagic species are electric or hydraulic reels and traps (Kojis 2004; thought to use the seamounts unique underwater Tobias 2004). Similar effort is reported throughout features primarily for food but also potentially as the region (FAO 1993). Figure 8.25 illustrates spawning grounds and/or nursery areas. a typical commercial fishing vessel capable of fishing offshore for deepwater snapper and grouper in the U.S. Virgin Islands. The maximum VII. STRESSORS ON DEEP CORAL reported depth for line fishing is 366 m (Swingle COMMUNITIES et al. 1970). The maximum reported depth for trap fishing is 183 m (Sheridan et al. 2006). Serious potential threats to deep water coral During submersible observations, Nelson and communities found in other U.S. waters are Appeldoorn (1985) encountered ghost fishing
326 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION traps at a depth of 121 m off the west end of St. Croix and a discarded longline at 236 m off the harbor entrance to San Juan, Puerto Rico.
Recreational fishing for deep water species in the U.S. Caribbean and throughout the region is termed “deep drop” fishing (also referred to as cannonball fishing). Gear includes expensive electric reels and heavy sash weights (commonly weighing 5 to 10 lbs) which allow fishing to potentially >600 m (Sword Fishing Central 2006). The Melton International Tackle Catalog (2007) lists electric deep drop reels up to $4,500 in
price. Reels “capable of landing fish as deep CARIBBEAN as 500 fathoms [914 m]” are advertised (Melton 2007, pg 95). Heavy weights are needed for optimal deep drop fishing performance; to take Figure 8.26. Typical electric reels used in deep drop the line straight down to the sea floor and keep fishing. This gear is marketed in the U.S. Caribbean. the tackle as close to the bottom as possible, in a process called “bouncing,” where the weight hold one type of fish and the base of that same bounces up off the sea floor and the whole rig mound will hold another. Some deep water remains under the boat (Brooks 2007). Non- mounds in the Bahamas go up to a couple of conventional weights, fashioned from pipes or hundred ft. [61 m] from the base. The Pomfret other containers filled with concrete are also is one of my deep water favorites. We catch used. Deep drop fishing has recently gained this species on the peaks of mounds. If your in popularity with sport fishers, the 3nd annual drop is off the mounds peak, you will have a ‘Lords of the Deep’ Fishing Tournament was held tough time catching them.” (Offshore Fishing in September 2007 in Nassau, Bahamas. Figure Forum 2006). 26 illustrates electric fishing reels utilized in deep- Table 8.3 details potential fishing gear impacts on drop fishing. Such gear is available in the U.S. deep-sea corals for the U.S. Caribbean with deep Caribbean. A small level of effort of deep-drop drop fishing included under “hook and line”. fishing is reported from the U.S. Caribbean and throughout the Caribbean region. When compared to potential threats deep coral habitats experience in other U.S. waters (chiefly The association between deep-sea fishing and destructive bottom trawling) this effort does not deep-sea coral habitat is unknown. However, a currently pose a significant threat to deep corals. review of web-based fishing forums reveals that However, the potential for damage to deep corals recreational sport fishers may be aware of and from such fishing gear is well documented, and fishing on such: may increase as the amount of fishing effort “You want to look for mounds, ridges, humps, increases (Chuenpagdee et al. 2003). Traps etc. The key is learning the currents and exact and weighted lines can crush structure-forming location of your drop especially in very deep corals and entangle or snare softer octocorals. water (1000-2000 ft) [305-610 m]. You will find The potential impacts of regional commercial that on one particular mound, the peaks will and recreational deep-sea fishing warrants further investigation. Additionally, the impact Figure 8.25. Bandit rigged boat that the removal of large deep water - a type of commercial fishing predators may have on deep coral vessel used to fish offshore for ecosystems is unknown. However, it deepwater snapper and grouper is well documented that the overfishing in the U.S. Virgin Islands and of these species can have negative capable of fishing to depths effects on shallow reef health (Dulvy where deep water coral reside. et al. 2004; Steneck and Sala 2005; Photo credit: Kojis 2005. Mumby et al. 2006).
327 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
Current efforts to manage fisheries the Caribbean Oil and gas exploration does currently occur in the waters are exacerbated by a regional ‘lack of to U.S. Caribbean. However, this activity occurs capacity’ in fisheries management (Haughton et al. in the southeast Caribbean, notably off Trinidad 2004). Traditional management strategies (such and Tobago, and in Straits of Florida waters off as catch and release) may not work for deep-water northwest Cuba. It is unknown what effect these species, as fish brought up from deep-sea coral activities have on deep coral communities. It is depths expire en-route. Additionally, remoteness also unknown to what extent regional underwater of location may play an important role with regard cables and lines affect deep coral communities. the management of deep-sea coral habitats. For example, the major structure forming corals The harvest of precious corals, red and black M. oculata and E. rostrata have been found in corals, is prohibited in the U.S. Caribbean and the waters surrounding Navassa Island (Figure there are no reports of current harvest activities. 8.5), an extremely remote uninhabited U.S. However, black coral jewelry, reportedly territory and illegal fishing activities are reported manufactured from corals harvested in the CARIBBEAN from Navassa’s waters (Miller 2003). However, Cayman Islands, is available for purchase in the the closest inhabited United States territory, U.S. Virgin islands (Greenberg and Greenberg Guantanamo Bay Naval Base (Cuba), is located 2006; Bernard K. Passman Gallery, St. Thomas, approximately 170 km distant. pers. comm.). Black coral sold in the U.S. Caribbean is reportedly harvested in the western Throughout the region, future fishing activities Caribbean from coral beds located at depths pose a significant potential threat, as shallow of 61-122 m (200-400 ft) (Bernard K. Passman water fisheries resources are depleted and Gallery, St. Thomas, pers. comm.). Regional commercial and recreational effort moves countries listed by the Convention on International into deeper water (Prado and Drew 1999; Trade in Endangered Species of Wild Fauna and FAO 2000; Koslow et al. 2000; Roberts 2002). Flora (CITES) trade database as exporting black Inspection of research station sorting sheets and coral (Antipatharia spp.) since 2001 include the current literature reveal numerous fish species Bahamas, British Virgin Islands, Cayman Islands, associated with deep corals that belong to taxa Cuba, Colombia, Dominican Republic, Mexico, currently harvested in other regions and may be Panama, and Trinidad and Tobago (Figure 8.4, harvested in the Caribbean in the future. Fishes some locations may be transshipment points) include targeted and by-catch deep-sea species (CITES 2007). The basil axis of black coral is such as deep-sea codfish, rattails or grenadiers, used in the manufacture of jewelry (Opresko and hake, dory, sea robins, wreakfish (Polyprion Sanchez 2005). americanus) and sharks (Merrett and Haedrich 1997; Carpenter 2002; Freiwald 2004; Gordon 2004) (locations of these sites are illustrated in VIII. CURRENT CONSERVATION AND Figure 8.3, species are listed in Appendix 8.2). MANAGEMENT ACTIVITIES Wreakfish (Sedberry et al. 1994) and dogfish sharks are currently harvested in U.S. continental In the U.S. Caribbean deep-water corals waters. Wreckfish and have been observed on and other marine resources are under the deep coral habitat (mounds and lithoherms) in the jurisdiction of territory (USVI: 0-3 nmi from Straits of Florida (Messing et al. 1990). Dogfish shore), commonwealth (Puerto Rico: 0-12 nmi sharks are reported on deep coral mounds from shore) and federal management authorities and lithoherms (Nizinski and Ross 2002) and (U.S. EEZ and the waters around Navassa associated with occurrences of structure-forming Island). Fisheries in the U.S. EEZ around USVI corals throughout the region (Appendix 8.2). and Puerto Rico are managed under fishery management plans developed by the Caribbean No significant threat of deep water trawling was Fishery Management Council (CFMC), based found for the wider Caribbean. However, deep in Puerto Rico. Deep sea coral language in the water shrimp trawlers are currently exploring recently reauthorized Magnuson-Stevens Act fishing beyond 70 m in depth off Colombia (Gracia includes an affirmation of the Council’s authority pers. comm.) and Reyes et al. (2005) note the designate zones to protect deep sea corals from need to asses the impacts of trawl fishing on the physical damage from fishing gear as part of their deep water coral banks off Colombia. fishery management plans, without having to
328 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION prove that corals constitute essential fish habitat IX. REGIONAL PRIORITIES TO [as amended by the Magnuson-Stevens Fishery UNDERSTAND AND CONSERVE DEEP Conservation and Management Reauthorization CORAL COMMUNITIES Act (Public Law 109-479), see Chapter 1]. Navassa Island is managed by the U.S. Fish and Morgan et al. (2006) advocate that ecosystem- Wildlife Service as a National Wildlife Refuge based management, research and mapping and that includes a 12 mile radius of marine habitats a ban on destructive fishing practices in deep sea around the island. Commonwealth and territorial coral areas are necessary to provide sufficient management authorities include the Puerto protection for deep-sea coral ecosystems. Rico Department of Natural and Environmental Resources and the United States Virgin Islands Ongoing projects that may generate information Department of Planning and Natural Resources. regarding the distribution deep coral communities include the previously mentioned benthic habitat
Current efforts to manage fisheries in Caribbean characterization effort by NOAA (NOAA 2007) CARIBBEAN waters are focused on shallow-water coral reef and a coral reef assessment and habitat mapping areas and associated fisheries (conch, grouper, effort of Navassa Island National Wildlife Refuge Caribbean spiny lobster, etc.) (Sala et al. 2001; by NOAA and the National Park Service (Miller Theile 2001; FAO 2003). However, the Territory pers. comm.). The Navassa Island effort, of the U.S. Virgin Islands includes antipatharians although currently focused on shallow water (black corals) in its local Endangered Species reefs, could include deep-water investigations. Act (ESA), thus granting the authority to protect Currently, no activities in the U.S. Caribbean are black corals (no matter the depth) within USVI focused on the biology and ecology of deep coral territorial waters. Puerto Rico implements the communities. Federal Endangered Species Act (Federal ESA) in local waters. However, currently, no deep Mapping the locations of deep coral habitat coral species are listed under the Federal ESA. would be a valuable component of any true The Caribbean Council manages corals under ecosystem based management program for the Corals and Reef Associated Invertebrates of the U.S. Caribbean. Mapping the locations of Puerto Rico and the U.S. Virgin Islands Fishery coral habitats gives a crucial spatial framework Management Plan (Coral FMP) (Caribbean for managers to better understand the effects Fishery Management Council 1994). It lists many of ocean uses on the environment. Figure 8.5 coral genera and species that are found in both illustrates existing managed areas in the U.S. shallow and deep waters. Those species found Caribbean and surrounding waters that have in deep water (including Iciligorgia schrammi potential deep water coral habitat (waters >100 (deep sea fan), Telesto spp., Ellisella spp. (sea m). Navassa Island Wildlife Refuge is the only whips) and antipatharians (black corals) would managed area that includes documented deep be protected under the Coral FMP. There are no water structure-forming and potential habitat- conservation or management efforts specifically forming scleractinian corals (E. rostrata, M. targeting deep-water corals for the U.S. Caribbean oculata and M. myriaster). or the wider Caribbean region. Examples of archival data awaiting further An assessment of regional deep-sea recreational inspection include the following: video footage and commercial fishing effort, especially deep and photographs recorded during the submersible drop fishing, would provide managers with observations around Puerto Rico and the U.S. information concerning the potential impact Virgin Islands reported by Nelson and Appeldoorn of this activity has on deep-sea corals. The (1985), stored at the National Marine Fisheries collection of socioeconomic information and Service (NMFS) Southeast Fisheries Science perceptions and attitudes to policy options, along Center in Pascagoula, Florida (Appeldoorn pers. with standard fisheries information (such as comm.); video footage and dive records recorded fishing effort, frequency of activity, landing and during submersible observations at Navassa bycatch information, etc.), could help minimize Island by the Center for Marine Conservation (now any negative impacts of policies directed at the The Ocean Conservancy); station records from conservation of deep sea coral ecosystems numerous research cruises kept on microfilm at the NMFS South East Fisheries Science Center
329 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
in Miami, Florida; and cruise reports and other mounds and/or lithoherms and banks. Similarly, information kept at the academic institutions that it seems likely that other areas throughout the carried out much original deep sea research Caribbean, including United States territories, - the University of Miami’s Marine Invertebrate with stronger currents may also have deep coral Museum in Miami, Florida, is one such location. ecosystems. The possibility that deep-sea coral habitat in these areas harbor fish, which are An assessment of regional deep-sea recreational exploitable (Appendix 8.2 and 8.3), deserves and commercial fishing effort, especially deep drop further attention. fishing, would provide managers with information concerning the potential impact of this activity. Technical note The collection of socioeconomic information and All images by Steven J. Lutz unless otherwise perceptions and attitudes to policy options, along noted. Images may be modified for clarification. with standard fisheries information (such as fishing effort, frequency of activity, landing and CARIBBEAN bycatch information, etc.), could help minimize any negative impacts of policies directed at the conservation of deep sea coral ecosystems.
X. CONCLUSIONS
It is clear from this review that the U.S. Caribbean and the wider Caribbean region contain a huge diversity of deep water corals. However, information on regional distribution of deep corals is limited and needs great expansion if deep corals can be part of any meaningful ecosystem based management strategy. Nevertheless, the thorough description of deep coral mounds and lithoherms in the Bahamas and in the Straits of Florida and deep coral habitat off Colombia provides valuable information on the bathymetric and hydrographic conditions favoring the development of deep coral communities. From this information inferences may be made regarding the morphology, composition and organism zonation of regional deep coral communities and the potential for deep coral habitat in the U.S. Caribbean.
The lithoherms of the Straits of Florida are remarkable examples of the interaction of strong currents with a varied suite of benthos (corals, zoanthids, crinoids, alcyonarians, etc.) a rain of pelagic sediments and syndepositional cementation of the sediments and skeletal debris. This interaction produces current- resistant structures with a continual supply of new hard substrate for an attached and boring benthic community. It is these same interrelated processes that allow the development of ocean facing, wave-resistant shallow reefs. There are strong indications that the Straits of Florida contain undiscovered large areas of deep coral
330 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
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339 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
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Theile S (2001) Queen Conch fisheries and Viada ST, Cairns SD (1987) Range extensions their management in the Caribbean. of ahermatypic Scleractinia in the Gulf Technical Report to the CITES of Mexico. Northeast Gulf Science 9 Secretariat. Brussels, TRAFFIC (2): 131-134 Europe, 95 pp Watling L, Auster PJ (2005) Distribution Tobias W (2004) Status of the Reefs: Reef of deep-water Alcyonacea off the Fish Resources. U.S. Virgin Islands Northeast coast of the United States. Department of Planning and Natural In: Freiwald A, Roberts JM (eds) Resources, Division of Fish and Cold-Water Corals and Ecosystems, Wildlife: 11 pp Springer-Verlag: 259-276
UNEP-WCMC (United Nations Environment Warner GF (1981) Species descriptions and Programme-World Conservation ecological observations of black corals Monitoring Centre) (2003) Checklist (Antipatharia) from Trinidad. Bulletin of fish and invertebrates listed in of Marine Science 31: 147-163 the CITES appendices and in EC Wilber JR, Neumann AC (1993) Effects of Regulation 338/97. 6th Edition. JNCC submarine cementation on microfabrics Report 341 and physical properties of carbonate van der Land J, Opresko DM (2001) slope deposits, Northern Bahamas. In: Antipatharia, in: Costello MJ et al. Rezak R, Lavoie DL (eds). Carbonate (eds.) (2001). European register Microfabrics, Springer, New York: 79- of marine species: a check-list of 94 the marine species in Europe and Wilkinson C (ed) (2004) Status of coral a bibliography of guides to their reefs of the world: 2004. Volume 2. identification. Collection Patrimoines Australian Institute of Marine Science, Naturels 50: 109 pp Townsville, Queensland, Australia: Voss GL (1966a) Narrative of the Cruise p- 301 pp 6607 to the Southwestern Caribbean, July 7 - 22. Processed Report, RSMAS, University of Miami: 38 pp
340 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 1979, 2000, UNEP-WCMC 2003; Reyes et al. 2005 Cairns 1979, 2000; and Zibrowis 1997 Cairns 1979, 2000; Reyes et al. 2005 Cairns 1979 Cairns 1999, 2000 Cairns 1979; Reyes et al. 2005 Cairns 1979, 2000; Reyes et al. 2005 Cairns 2000 Cairns 2000 Cairns 1979 Cairns 1979, 2000; Reyes et al. 2005 CARIBBEAN 0-263 37-640 35-229 30-329 45-494 27-274 13-220 53-708 274-311 500-700 238-274 Depth Range U.S. Virgin U.S. Virgin Distribution ♠ Widespread Caribbean and Bahamas, including the Islands Straits of Florida, southern Caribbean Caribbean, southern Florida, of Straits Antilles Lesser Straits of Florida, Bahamas, Lesser Antilles Throughout Caribbean, including Puerto Rico Mexico, Colombia Channel, Yucatan Straits of Florida, Antilles; Venezuela Lesser Southeastern Caribbean, Lesser Antilles Straits of Florida, southeastern and western Caribbean Nicaragua Only off Channel, Yucatan Antilles, Bahamas, Colombia Species ▲ Anomocora fecunda Anomocora marchadi Anomocora prolifera Anthemiphyllia patera Astrangia poculata Balanophyllia bayeri Balanophyllia cyathoides Balanophyllia dineta Balanophyllia floridana Balanophyllia hadros Balanophyllia palifera Family Deep-sea corals of the wider Caribbean region, including those located in U.S. Caryophyllidae Caryophyllidae Caryophyllidae Anthemiphylliidae Rhizangiidae Dendrophylliidae Dendrophylliidae Dendrophylliidae Dendrophylliidae Dendrophylliidae Dendrophylliidae Taxon Higher Phylum Cnidaria Anthozoa Class Subclass Hexacorallia (Zoantharia) Order Scleractinia Appendix 8.1. ▲ Bold denotes major and potential structure-forming scleractinia (species associated with deep water banks, lithoherms, etc.) potential habitat-forming antipatharia, octocorallia, pennatulacea, and hydrozoa (species greater than or equal to 25 cm species greater than or equal to 25 cm in height width ♠ Bold denotes distribution including U.S. Caribbean territories ‡ For material examined: HBOI = Harbor Branch Oceanographic Institute; USNM United States National Museum
341 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 1979, UNEP-WCMC 2003; Reyes et al. 2005 Cairns 1979 Cairns 1979; Reyes et al. 2005 Cairns 1979, 2000 Cairns 1979, 2000; Reyes et al. 2005 Cairns 1979, 2000; Reyes et al. 2005 Cairns 1979 Cairns 2000 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979, 2000 Cairns 2000; UNEP-WCMC 2003; Reyes et al. 2005 Cairns 1979, 2000 Cairns 2000 CARIBBEAN 5-100 12-183 97-399 73-618 70-366 50-100) 412-505 244-805 150-730 129-249 99-1033 183-380 714-843 183-1646 700-1817 Depth Range 32-480 (common Puerto U.S. U.S. territories Distribution ♠ U.S. territories US territories U.S. territories U.S. territories Bahamas, Cuba, Grenada, Jamaica, Colombia Straits of Florida, St. Lucia Widespread Caribbean, Bahamas, including Antilles, Straits of Florida, Bahamas, including Barbados; Colombia Widespread Caribbean, Bahamas, including Islands to Cuba, Antilles, Virgin including Bahamas and Caribbean, Greater Antilles, southern and western Caribbean, including Antilles, Venezuela Antilles, including Lesser territories Antilles Straits of Florida, Antilles Straits of Florida, eastern and southern Caribbean, including Rico Straits of Florida, St. Lucia Bahamas and Caribbean Species ▲ Balanophyllia wellsi Bathypsammia fallosocialis Caryophyllia ambrosia caribbeana Caryophyllia antillarum Caryophyllia barbadendis Caryophyllia berteriana Caryophyllia corrugata Caryophyllia crypta Caryophyllia parvula Caryophyllia paucipalata Caryophyllia polygona Caryophyllia zopyros Cladocora debilis Cladopsammia manuelensis Coenocyathus caribbeana Family Dendrophylliidae Dendrophylliidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Dendrophylliidae Caryophyllidae Taxon Higher
342 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 1979, 2000; Reyes et al. 2005 Cairns 1979, 2000; Reyes et al. 2005 Cairns 2000 Cairns 1979 Cairns 1979 Cairns 1979, 2000, UNEP-WCMC 2003 Cairns 1979, 2000 Cairns 1979, 2000 Cairns 1979 Cairns 1979, 2000; Reyes et al. 2005 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979, 2000, UNEP-WCMC 2003 Cairns 1979, 2000 Cairns 1979, 2000; Reyes et al. 2005 Cairns 1979, 2000 CARIBBEAN 97-399 74-622 37-366 81-675 93-300 97-505 0.5-347 110-421 110-553 311-567 183-800 183-443 494-907 183-907 201-777 276-900 277-823 403-2634 155-2200 Depth Range U.S. Virgin Islands U.S. Virgin Distribution ♠ U.S. territories U.S. territories U.S. territories U.S. territories U.S. territories Bahamas and Caribbean, including U.S. territories Widespread Caribbean, Bahamas, including Throughout Caribbean, Bahamas, including Channel Yucatan Straits of Florida, Havana, Cuba Only off Straits of Florida, Cuba, Grenada, Venezuela Straits of Florida, southern Lesser Antilles Antilles Straits of Florida, Lesser Anguilla Straits of Florida, Widespread Caribbean, Bahamas, including Widespread Caribbean, Bahamas, including Widespread Caribbean, Bahamas, including Straits of Florida, Belize, Lesser Antilles Straits of Florida, Bahamas Antilles, Bahamas Straits of Florida, Bahamas, Martinique, Antilles, Bahamas, including Greater U.S. territories Yucatan Bahamas, Straits of Florida, Channel, southeastern Caribbean Channel, Venezuela Yucatan D. (=
Species ▲ ) Coenocyathus parvulus Coenosmilia arbuscula Colangia immersa Concentrotheca laevigata Cylicia inflata Dasmosmilia lymani Dasmosmilia variegata Deltocyathoides stimpsonii Deltocyathus agassizii Deltocyathus calcar Deltocyathus eccentricus Deltocyathus italicus Deltocyathus moseleyi Deltocyathus pourtalesi Dendrophyllia alternata Desmophyllum dianthus cristagalli Desmophyllum striatum Eguchipsammia cornucopia Eguchipsammia gaditana Family Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Turbinoliidae Caryophyllidae Caryophyllidae Turbinoliidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Dendrophylliaidae Caryophyllidae Caryophyllidae Dendrophylliaidae Dendrophylliaidae Taxon Higher
343 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 1979 Cairns 1979, UNEP-WCMC 2003 Cairns 1979, UNEP-WCMC 2003 Cairns 1979, 2000; UNEP-WCMC 2003 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979, 2000 Cairns 1979, 2000 Cairns 2000 Cairns 1979, 2000 Cairns 1979, 2000; Reyes et al. 2005 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979, 2000, UNEP-WCMC 2003 Cairns 2000; Reyes et al. 2005 Cairns 2000 CARIBBEAN 2-146 24-311 80-366 97-700 46-241 30-653 51-130 357-618 285-439 30-1809 695-810 403-1748 300-1646 216-1097 183-1664 146-1200 1450-2745 1089-1234 2842-3475 Depth Range Puerto (absent Bahamas) Distribution ♠ Straits of Florida, St. Lucia Antilles, Straits of Florida, Bahamas, Antilles, including western Caribbean, Islands U.S. Virgin Bahamas, Cuba Straits of Florida, Mexico, Western Panama Widespread Caribbean, including U.S. territories Bahamian archipelago Lesser Yucatan, Straits of Florida, Antilles Widespread Caribbean, Bahamas. Throughout Caribbean, Bahamas Channel, Straits of Yucatan Antilles, Florida Channel Yucatan Antilles, Bahamas, Bahamas, Straits of Florida, western Antilles Caribbean, Widespread Caribbean, Bahamas, including U.S. territories Bahamas, Jamaica Antilles, Straits of Florida, Bahamas, western Caribbean south east Cuba Only off Straits of Florida, Bahamas, southern Antilles, including Caribbean, Islands Rico and U.S. Virgin Widespread Caribbean, including U.S. territories Southern Caribbean Species ▲ Enallopsammia profunda Enallopsammia rostrata Flabellum (pavoninum) atlanticum Flabellum floridanum Flabellum moseleyi Fungiacyathus marenzelleri Fungiacyathus pusillus Fungiacyathus symmetricus Gardineria minor Gardineria paradoxa Gardineria simplex Guynia annulata Javania cailleti Javania pseudoalabastra Labyrinthocyathus langae Leptopenus discus Lophelia pertusa Madracis asperula Madracis brueggemanni Family Dendrophylliaidae Dendrophylliaidae Flabellidae Flabellidae Flabellidae Fungiacyathidae Fungiacyathidae Fungiacyathidae Gardineriidae Gardineriidae Gardineriidae Guyniidae Flabellidae Flabellidae Caryophyllidae Micrabaciidae Caryophyllidae Pocilloporidae Pocilloporidae Taxon Higher
344 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 1979, 2000, UNEP-WCMC 2003 Cairns 2000; Reyes et al. 2005 Cairns 1979, 2000; UNEP-WCMC 2003; Reyes et al. 2005 Cairns 1979 Cairns 2000; UNEP-WCMC 2003 Cairns 1979, 2000; Reyes et al. 2005 Cairns 1979, 2000; Reyes et al. 2005 Cairns 1979 Cairns 1979, 2000 Cairns 1979 Cairns 2000; Reyes et al. 2005 Cairns 2000 Cairns 1979, 2000; Reyes et al. 2005 Cairns 1979 Cairns 1979, 2000 CARIBBEAN 40-60) 11-333 46-640 17-250 20-355 12-143 75-822 20-1220 284-457 497-907 137-309 137-931 100-300) 144-1391 349-1200 Depth Range 53-801 (common 25-159 (common , , Puerto U.S. territories Distribution ♠ Puerto Rico and U.S. Puerto Rico and U.S. U.S. territories U.S. territories Widespread Caribbean, Bahamas, including Islands Virgin Widespread Caribbean, Bahamas, including U.S. territories Tobago, Antilles, Bahamas, Greater western Caribbean, including Islands Rico and U.S. Virgin Venezuela Widespread Caribbean, Bahamas, including Islands Virgin Straits of Florida, Cuba Widespread Caribbean, Bahamas, including Widespread Caribbean, including U.S. territories Antilles Straits of Florida, Caribbean western Antilles, Bahamas, Antilles Lesser Straits of Florida, Bahamas, northern Caribbean, including Barbados Disjunct distribution, Florida, southern Caribbean Straits of Florida, Bahamas, western and southern Caribbean, Antilles Lesser Antilles, Straits of Florida, Bahamas, including Straits of Florida, Bahamas, northern and eastern Caribbean
Species ▲ pharensis forma Madracis myriaster Madracis pharensis Madrepora carolina Madrepora oculata Oculina tenella Oxysmilia rotundifolia Paracyathus pulchellus Peponocyathus folloculus Phacelocyathus flos Placotrochides frusta Polycyathus mayae Polycyathus senegalensis Polymyces fragilis Pourtalocyathus hispidus Premocyathus cornuformis Family Pocilloporidae Pocilloporidae Oculinidae Oculinidae Oculinidae Caryophyllidae Caryophyllidae Turbinoliidae Caryophyllidae Flabellidae Caryophyllidae Caryophyllidae Flabellidae Guyniidae Caryophyllidae Taxon Higher
345 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 1977, 2000, UNEP-WCMC 2003 Cairns 1979, 2000 Cairns 2000 Cairns 1979, 2000 Cairns 1979 Cairns 1979, 2000 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979 Cairns 1979, 2000; Reyes et al. 2005 Cairns 1979; UNEP-WCMC 2003 Cairns 1979 CARIBBEAN 4-914 88-640 4.5-119 0.5-508 123-549 165-835 299-715 155-649 320-488 320-488 155-220 214-878 400-576 795-2113 300-1158 220-1383 543-1250 Depth Range
U.S. U.S. Puerto Rico Distribution ♠ U.S. Virgin Islands U.S. Virgin U.S. territories U.S. territories U.S. territories U.S. territories Navassa Island Lesser Antilles, Colombia, Bahamas, Lesser including Yucatan Straits of Florida, Bahamas, Antilles, Channel, northeastern including Widespread Caribbean, Bahamas, including Straits of Florida, Bahamas, western Antilles (from Caribbean, to Grenada) Antilles, Straits of Florida, Lesser Jamaica, southern Caribbean Straits of Florida, western Caribbean, Antilles, including Bahamas, territories Throughout Caribbean, Bahamas, including Widespread Caribbean, including U.S. territories Antilles, Panama, Straits of Florida, and off Widespread Caribbean, Bahamas Straits of Florida, Jamaica, Bahamas, Barbados. Channel Yucatan Jamaica, Yucatan, Straits of Florida, including Florida Terrace, Only from Pourtales Antilles, including Bahamas, territories, Colombia Keys, Bahamas Gerogia to FL Only northwest Cuba Species ▲ Rhizopsammia goesi Rhizosmilia gerdae Rhizosmilia maculata Schizocyathus fissilis Solenosmilia variabilis Stenocyathus vermiformis Stephanocyathus coronatus Stephanocyathus diadema Stephanocyathus laevifundus Stephanocyathus paliferus cylindraceus Tethocyathus recurvatus Tethocyathus variabilis Tethocyathus Thalamophyllia gombergi Thalamophyllia riisei Thecopsammia socialis corbicula Trematotrochus Family Dendrophylliaidae Caryophyllidae Caryophyllidae Guyniidae Caryophyllidae Guyniidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Caryophyllidae Dendrophylliidae Turbinoliidae Taxon Higher
346 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 1979 Cairns 1979, 2000 Cairns 1979 Messing et al. 1990 Opresko 1972; UNEP-WCMC 2003; Opresko 2006 Opresko 1972; UNEP-WCMC 2003; Opresko 2006 Opresko and Cairns 1994; UNEP- WCMC 2003; Opresko 2006 Brook 1889; Opresko 1972; 1981; OBIS 2006 Warner 1981; UNEP-WCMC Warner 2003; Reyes et al. 2005; Opresko 2006 Opresko 1996; UNEP-WCMC 2003; Opresko and Sanchez 2005 Opresko 1974; UNEP-WCMC 2003; Reyes et al. 2005; Opresko 2006 Opresko 1972; UNEP-WCMC 2003; Opresko and Sanchez 2005; Reyes et al. OBIS 2006; Opresko 2006 CARIBBEAN 30-60) 55-700 37-532 50-200 10 - 115 73 - 567 205-380 129-161 ~40-240 532-1372 129 - 491 Depth Range >100 (~580-630) 11->100 (common 11->100
, Puerto Rico , Grenada, , Venezuela Distribution ♠ U.S. territories Bahamas, Virgin Islands, including Bahamas, Virgin U.S. territories Widespread Caribbean, Bahamas, including Antilles northwestern Cuba, Lesser Bahamas (Straits of Florida) Bahamas, Barbados, Cuba, Grenada, Mexico, Montserrat, Saint Vincent and the Grenadines and the Cuba, Saint Vincent Grenadines Barbados, Cuba, Guadeloupe, Montserrat Islands U.S. Virgin Antilles Netherlands Trinidad Colombia, Jamaica, Mexico, Tobago and Bahamas, Colombia, Antilles Greater and Lesser Antilles, including Greater and Lesser Islands U.S. Virgin Straits of Florida, Cuba, Saint Vincent and Trinidad and the Grenadines, Barbados, Honduras, Tobago, Panama, Colombia, Venezuela Species ▲ Trochocyathus fossulus Trochocyathus rawsonii Trochocyathus infundibulum Trochopsammia Gerardia sp. Acanthopathes humilis Acanthopathes thyoides Allopathes desbonni Antipathes americana? Antipathes atlantica Antipathes caribbeana Antipathes columnaris Antipathes lenta Family Caryophyllidae Caryophyllidae Dendrophylliaidae Gerardiidae Aphanipathidae Aphanipathidae Antipathidae Antipathidae Antipathidae Antipathidae Antipathidae Antipathidae Taxon Higher Order Zoanthidea Antipatharia Order
347 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Brook 1889, Warner 1981, STRI Brook 1889, Warner 2006; Reyes et al. 2005 UNEP-WCMC 2003 UNEP-WCMC 2003 Opresko 1974; 2006 Opresko 1972; UNEP-WCMC 2003; Reyes et al. 2005; Opresko 2006 Brook 1889 ; Opresko 1974; Opresko 2002; 2006 Brook 1889; Opresko 1974; et al. 1975; Opresko 2002; Keller, UNEP-WCMC 2003; OBIS 2006; Opresko 2006 Opresko 2006 UNEP-WCMC 2003 Opresko 2004; CITES 2006; Opresko 2006 Opresko 2006; 1972; UNEP-WCMC 2003 Opresko 1972; Rezak et al. 1985; UNEP-WCMC 2003; Reyes et al. 2005; Opresko 2006 Opresko 2003; 2006 Opresko 1974; UNEP-WCMC 2003; Opresko 2006; Material examined USNM 1026305 CARIBBEAN Unk Unk >100 >100 20-160 95-190 31 - 310 107-333 ~60-310 176 - 549 100-5000 ~175-450 ~100-5000 Depth Range , Saint Kitts and
Distribution ♠ Puerto Rico from Colombia north throughout the Caribbean, including Panama, and Colombia Tobago, and Trinidad Islands U.S. Virgin Barbados, Guadeloupe, Martinique, Montserrat, Saint Lucia Widespread Caribbean Guadeloupe, Colombia Yucatan Straits of Florida (Bahamas), Channel Mexico, Nevis, Caribbean Sea Basin, Cayman Basin Yucatan Trench, Widespread Caribbean Barbados, Cuba, Martinique, and the Montserrat, St. Vincent Grenadines Martinique, Mexico Bahamas, Barbados, Cuba, Dominica, Guadeloupe, Martinique, Montserrat, Saint and the Grenadines Vincent Bahamas, Barbados, Cuba, Martinique, Mexico, Nicaragua, and the Grenadines; Saint Vincent Colombia Widespread Caribbean Straits of Florida, Jamaica, Mexico; Bahamas Venezuela, Species ▲ Antipathes gracilis Antipathes rhipidion? Antipathes tristis? Aphanipathes pedata Aphanipathes salix Bathypathes alternata Bathypathes patula Chrysopathes sp. Cirrhipathes spiralis? Distichopathes disticha Distichopathes filix Elatopathes abietina Heliopathes americana Leiopathes glaberrima Family Antipathidae Antipathidae Antipathidae Aphanipathidae Aphanipathidae Schizopathidae Schizopathidae Cladopathidae Antipathidae Aphanipathidae Aphanipathidae Aphanipathidae Cladopathidae Leiopathidae Taxon Higher
348 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Opresko 1972; 2002; UNEP-WCMC 2003; Opresko 2006 Opresko 1972; 2002; UNEP-WCMC 2003; Opresko 2006 Opresko 1974; Colin 1978; 2006; CITES 2003, UNEP-WCMC Opresko 2006 Brook 1889; Humann 1993; UNEP-WCMC 2003; Reyes et al. 2005; Opresko 2006 Brook 1889; Opresko 2006; Reyes et al. 2005 Pourtalès 1874, 1880; Brook 1889; Reyes et al. 2005; CITES 2006; Opresko 2006 Brook 1889; van der Land and 1981; Opresko 2001; Warner UNEP-WCMC 2003; Opresko 2006 Opresko 1972; Colin 1978; 1981; UNEP-WCMC Warner 2003; Opresko 2006 UNEP-WCMC 2003; OBIS 2006; Opresko 2006 1981; Opresko 1970; Warner UNEP-WCMC 2003; Opresko 2006 Cairns 2005 Cairns 2005 CARIBBEAN >100 >200 3-229 ~14-115 64 - 640 13 - 357 46 - 915 ~20-160 ~60-345 ~45-345 175 - 428 320-3186 Depth Range , U.S. Puerto Rico Distribution ♠ Cuba, Guyana, Mexico, Saint Lucia Bahamas, Barbados, Colombia, Cuba, Guadeloupe, Venezuela Throughout Caribbean, including Islands & Puerto Rico Virgin Barbados, Colombia, Dominican Tobago and Trinidad Republic, Caribbean region Barbados, Colombia, Cuba, Grenada, Martinique, Montserrat, Saint Vincent and the Grenadines Tobago and Trinidad Barbados, Throughout Caribbean, including Puerto Rico Throughout Caribbean, including Puerto Rico Tobago and Trinidad Atlantic Western Straits of Florida, Bahamas Species ▲ Parantipathes tetrasticha Phanopathes rigida Plumapathes pennacea Stichopathes lutkeni Stichopathes occidentalis Stichopathes pourtalesi barbadensis Tanacetipathes hirta Tanacetipathes tanacetum Tanacetipathes thamnea Tanacetipathes Alcyonium digitatum Anthomastus agassizi Family Schizopathidae Schizopathidae Myriopathidae Antipathidae Antipathidae Antipathidae Myriopathidae Myriopathidae Myriopathidae Myriopathidae Alcyoniidae Alcyoniidae Taxon Higher Subclass Octocorallia Alcyonacea Order
349 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Humann 1993 Cairns 2005 Cairns 2005; Reyes et al. 2005 Cairns 2005; Reyes et al. 2005 Cairns 2005 Atlantic Fishery South Management Council 1998; Cairns 2005; Reed 2006 Cairns 2005 Cairns 2005 Bayer 1961; Humann 1993 Atlantic Bayer 1961; South Fishery Management Council 1998; Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 CARIBBEAN 805 >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 37-311 24-329 60-298 37-503 60-878 23-188 27-1153 175-586 ~82->200 Depth Range Puerto Distribution ♠ Western Atlantic Western Throughout Caribbean Havana, Cuba), Straits of Florida (off Antilles probably throughout Atlantic Western Atlantic Western Atlantic Western Bahamas, Caribbean Atlantic Western Atlantic Western Throughout Caribbean Atlantic Western Havana, Cuba), Straits of Florida (off Bahamas Throughout Caribbean Atlantic Western Antilles, including Bahamas, Rico Havana, Cuba), Straits of Florida (off Bahamas Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western delta corallina ) var. var. operculata ?) Telesto (= Species ▲ Telesto (= Anthomastus robusta Bellonella rubistella Carijora Drifta glomerata Duva florida Gersemia rubiformia Neospongodes portoricensis Nidalia deichmannae Nidalia dissidens Nidalia occidentalis Nidalia rubrapunctata Pseudodrifa capnella nigra Scleranthelia rugosa Stereonepthya portoricensis Stereotelesto nelleae Telesto septentrionalis Telesto rudis Trachythela Acanella arbuscula Acanella eburnea Acanthactis austera Family Alcyoniidae Alcyoniidae Clavulariidae Nephtheidae Nephtheidae Nephtheidae Nephtheidae Nidaliiae Nidaliiae Nidaliiae Nidaliiae Nephtheidae Clavulariidae Nephtheidae Clavulariidae Clavulariidae Clavulariidae Nephtheidae Isididae Isididae Plexauridae Taxon Higher Order Gorgonacea
350 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 2005 Cairns 2005 Auster 2005; Bayer and Watling 1961 and 1987; Watling Veronique Auster 2005; Reyes et al. 2005 Cairns and Bayer 2004b; 2005 Cairns and Bayer 2004b; 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns & Bayer 2002; 2005 Cairns & Bayer 2002; 2005 Cairns & Bayer 2002; 2005 Bayer 2001; Cairns 2005 Bayer 2001; Cairns 2005 Bayer 2001; Cairns 2005 Bayer 2001; Cairns 2005 Cairns and Bayer 2004b; Auster 2005; and 2005; Watling Cairns personal Comm. CARIBBEAN 1116 1399 >200 >200 >183 >200 >200 >200 >200 >200 >200 >200 >200 82-514 593-911 137-595 194-686 183-732 229-556 686-1125 514-2063 Depth Range U.S. Puerto Distribution ♠ U.S. territories , Virgin Islands, including , Virgin Western Atlantic Western Atlantic Western Throughout Caribbean, including Puerto Rico Straits of Florida; Guadeloupe; Colombia Straits of Florida, Bahamas, Rico Islands Virgin Antilles, lesser Yucatan, northeastern Straits of Florida Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western Antilles, Straits of Florida, Lesser including Throughout Caribbean, Bahamas, Antilles Only Bahamas Windward Passage Straits of Florida Antilles Lesser Straits of Florida, Bahamas Antilles Straits of Florida, Bahamas, Species ▲ Acanthactis scabra Acanthogorgia armata Acanthogorgia aspera Acanthogorgia schrammi Acanthoprimnoa goesi Acanthoprimnoa pectinata Anthothela bathybius Anthothela grandiflora Anthothela grandiflora sensu Bebryce cinerea Bebryce grandis Bebryce parastellata Caliacis nutans Callogorgia americana americana Callogorgia gracilis Callogorgia linguimaris Calyptrophora antilla Calyptrophora gerdae Calyptrophora pillsburyae Calyptrophora trilepis Candidella imbricata Family Plexauridae Acanthogorgiidae Acanthogorgiidae Acanthogorgiidae Primnoidae Primnoidae Anthothelidae Anthothelidae Anthothelidae Plexauridae Plexauridae Plexauridae Plexauridae Primnoidae Primnoidae Primnoidae Primnoidae Primnoidae Primnoidae Primnoidae Primnoidae Taxon Higher
351 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 1987; Cairns 2001; Veronique Cairns 2005; Reyes et al. 2005 Cairns 2001; 2005 Cairns 2001; 2005 Cairns 2001; 2005 Cairns 2001; 2005 Cairns 2001; 2005 Cairns 2001; 2005 Bayer 1961; Cairns 2005; Watling Auster 2005 and Bayer 1961; Cairns 2005; Watling Auster 2005 and Cairns 2005 Cairns 2005 Cairns 2005 1987; Bayer 1961; Veronique Humann 1993 Auster and Cairns 2005; Watling 2005 Cairns 2005 1987; Bayer 1961; Veronique Humann 1993; Cairns 2005; Armstrong et al. 2006 CARIBBEAN >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 14-183 20-488 155-595 146-526 659-677 128-1716 403-1200 320-1354 914-2256 431-1046 Depth Range , Bahamas, Guadeloupe Distribution ♠ Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western Antilles, Greater and Lesser Bahamas, Guadeloupe, Martinique, Colombia Southeastern Caribbean, Lesser Antilles, Martinique Antilles, Caribbean Antilles, Bahamas, Straits of Florida Antilles Gulf of Mexico, Lesser Yucatan Antilles, Antilles, Bahamas Greater and Lesser Straits of Florida Straits of Florida Atlantic Western Atlantic Western Atlantic Western Throughout Caribbean, including Puerto Rico Straits of Florida Atlantic Western Antilles, including Striats of Florida, Islands U.S. Virgin atlantica ) Ellisella Species ▲ ( Caribisis simplex Chalcogorgia pellucida Chelidonisis aurantiaca mexicana Chrysogorigia agassizii Chrysogorigia desbonni Chrysogorigia elegans Chrysogorigia fewkesii Chrysogorigia multiflora Chrysogorigia spiculosa Chrysogorigia squamata Chrysogorigia thrysiformis Corallium medea Corallium niobe Ctenocella Dasystenella acanthina Dendrobrachia multispina Diodogorgia nodulifera? Distichogorgia sconsa Echinomuricea atlantica Ellisella barbadensis Family Isididae Chrysogorgiidae Isididae Chrysogorgiidae Chrysogorgiidae Chrysogorgiidae Chrysogorgiidae Chrysogorgiidae Chrysogorgiidae Chrysogorgiidae Chrysogorgiidae Coralliidae Coralliidae Ellisellidae Primnoidae Dendrobrachiidae Anthothelidae Chrysogorgiidae Plexauridae Ellisellidae Taxon Higher
352 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Humann 1993; Bayer 1961; Caribbean Fishery Management Council 2004; Cairns 2005 Cairns 2005 1987; Cairns 2005 Veronique Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Auster 2005; Reed and Watling 2006 Cairns 2005 1987; Cairns 2005; Veronique Material examined HBOI 26-V-06- 1-17 (Reed 2006) Humann 1993; Cairns 1977; Cairns 2005 1987 Cairns 2005; Veronique Reed 2006 Cairns 2005 Auster and Cairns 2005; Watling 2005 Bedford Institute of Oceanography 2003; Cairns Auster 2005 and 2005; Watling Cairns 2005 1987 Cairns 2005; Veronique 1987 Cairns 2005; Veronique Auster and Cairns 2005; Watling 2005 CARIBBEAN 611 350 176 1343 >200 >200 >200 >200 >200 >200 to 667 11-366 15-219 170-878 92~>200 52~>200 61~>200 743-1125 274-3236 274-3236 607-1067 Depth Range , Bahamas, Distribution ♠ Puerto Rico Antilles, southeastern Caribbean Atlantic Western Guadeloupe Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western Straits of Florida, Bahamas Atlantic Western Guadeloupe, Martinique Antilles, Straits of Florida, southeastern Caribbean Guadeloupe Straits of Florida Atlantic Western Straits of Florida, Bahamas, Mexico, Guadeloupe, Colombia, Venezuela Straits of Florida, Bahamas, Cuba Atlantic Western Martinique Martinique, Guadeloupe Straits of Florida, Caribbean Species ▲ Ellisella elongata Ellisella funiculina Ellisella grandiflora Ellisella grandis Ellisella nivea Ellisella rosea Ellisella schmitti Eunicella albatrossae Eunicella modesta Eunicella tenuis Hypnogorgia pendula Iciligorgia schrammi Iridogorgia pourtalesi Isidella longiflora? Junceella antillarum Keratoisis flexibilis Keratoisis ornata Keratoisis siemensii Keratosis simplex Lepidisis caryophyllia Lepidisis longiflora Family Ellisellidae Ellisellidae Ellisellidae Ellisellidae Ellisellidae Ellisellidae Ellisellidae Gorgoniidae Gorgoniidae Gorgoniidae Plexauridae Anthothelidae Chrysogorgiidae Isididae Ellisellidae Isididae Isididae Isididae Isididae Isididae Isididae Taxon Higher
353 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Bayer 1952; Keller et al. 1975 Bayer 1952; Keller et al. 1975 Atlantic Bayer 1961; South Fishery Management Council 1998; Cairns 2005 Cairns 2005 Bayer 1961; Cairns 1977; Humann 1993; Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005; Bayer 1961; 1987 Veronique Cairns and Bayer 2003; 2005 Cairns and Bayer 2003; 2005 1987; Cairns and Veronique Bayer 2003; Cairns 2005; Watling Auster 2005 and Cairns and Bayer 2003; 2005 Cairns and Bayer 2003; 2005 Cairns 2007 Cairns 2007 Cairns 2007 Cairns 2005; 2007 Cairns 2007 CARIBBEAN 1485 >200 >200 >200 26-123 18-128 67-455 45-146 62-237 27-403 27-327 161-792 366-792 677-900 229-244 738-1473 Depth Range 1060-3000 1060-3000 Puerto U.S. territories U.S. territories Distribution ♠ , Bahamas, Mexico Southern Lesser Antilles Southern Lesser Antilles Southern Lesser Straits of Florida (Cuba) Atlantic Western Antilles Straits of Florida, Bahamas, Atlantic Western Atlantic Western Florida, Bahamas, Guadeloupe Straits of Florida, Bahamas, Lesser Antilles, including Antilles, Straits of Florida, Rico Antilles and Straits of Florida, Lesser Bahama. Only Bahamas. Straits of Florida, Cuba. Bahamas and southern Caribbean Antilles, southern Bahamas, Caribbean Bahamas, northweatern Caribbean, Antilles, incluuding Atlantic, including Bahamas Western and southern coast of Caribbean to Nicaragua Only Grand Cayman Island hirta ) Trachymuricea (= Species ▲ Leptogorgia euryale? Leptogorgia stheno? Lophogorgia cardinalis Metallogorgia splendens Muricea laxa Muriceides Muriceides kukenthali Muriceopsis petila Narella bellissima Narella pauciflora Narella regularis Narella spectabilis Narella versluysi Nicella goreaui Nicella americana Nicella deichmannae Nicella hebes Nicella lanceolata Family Gorgoniidae Gorgoniidae Gorgoniidae Chrysogorgiidae Plexauridae Plexauridae Plexauridae Plexauridae Primnoidae Primnoidae Primnoidae Primnoidae Primnoidae Ellisellidae Ellisellidae Ellisellidae Ellisellidae Ellisellidae Taxon Higher
354 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 2007; 2005; 1987 Veronique Cairns 2007 Cairns 2007 Cairns 2007 Cairns 2007 Cairns and Bayer 2004a; 2005 Cairns and Bayer 2004a; 2005 Cairns and Bayer 2004a; 2005 Auster 2005; material and Watling examined USNM 35928; Cairns personal comm. Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005; Reyes et al. 2005 Cairns 2005 Cairns and Bayer 2004b; 2005 Cairns and Bayer 2004b; 2005 Cairns and Bayer 2004b; 2005 CARIBBEAN 514 >200 >200 >200 >200 >200 >200 >200 27-395 55-329 60-481 110-259 174-819 374-555 165-706 522-608 247-805 400-900 310-878 494-1065 Depth Range , U.S. U.S. territories Distribution ♠ , Venezuela Bahamas, Antilles from Cuba to Bahamas, Barbados, incluuding Venezuela Antilles, including Bahamas, territories Atlantic Western Antilles, Venezuela Bahamas, Bahamas, Caribbean Only St. Lucia Antilles, and Straits of Florida, Cuba, Bahamas Yucatan Straits of Florida, Bahamas, Straits of Florida, Bahamas Atlantic Western Atlantic Western Atlantic Western Havana) Straits of Florida, Cuba (off Atlantic Western Atlantic Western Atlantic Western Atlantic Western Bahanas and northern Straits of Florida Straits of Florida, Cuba Straits of Florida
) boschmai (= Species ▲ sp. A. sp. Nicella obesa Nicella guadelupensis Nicella Nicella gracilis Nicella robusta Paracalyptrophora carinata Paracalyptrophora duplex Paracalyptrophora simplex Paragorgia johnsoni Paramuricea echinata Paramuricea grandis Paramuricea multispina Paramuricea placomus Placogorgia mirabilis Placogorgia rudis Placogorgia tenius Placogorgia tribuloidea Plumarella aculeata Plumarella aurea Plumarella dichotoma Family Ellisellidae Ellisellidae Ellisellidae Ellisellidae Ellisellidae Primnoidae Primnoidae Primnoidae Paragorgiidae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Primnoidae Primnoidae Primnoidae Taxon Higher
355 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns and Bayer 2004b; 2005 Cairns and Bayer 2004b; 2005 Cairns and Bayer 2004b; 2005; Cairns personal comm. Cairns and Bayer 2004b; 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 1977; Kapela and Lasker 1999 et al. 1975; Bedford Keller, Institute of Oceanography 2003; Auster 2005; Cairns and Watling 2005 Cairns 2007 1987; Cairns 2005 Veronique 1987; Cairns 2005 Veronique Cairns 2005 Cairns 2005 1987; Cairns 2005 Veronique 1987; Cairns 2005 Veronique CARIBBEAN >200 >200 >200 >200 >200 >200 >200 3-283 18-494 110-704 196-882 183-743 183-850 176-350 40-1953 221-858 549-1160 275-1607 Depth Range U.S. Puerto Distribution ♠ , Mexico, Panama, southern Straits of Florida, Bahamas Straits of Florida, Cuba Straits of Florida Straits of Florida Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western Antilles, Straits of Florida, southeastern Caribbean Straits of Florida, southern Lesser Antilles Bahamas, Caribbean, including territories Guadeloupe, Martinique Guadeloupe, Martinique Atlantic Western Havana), Straits of Florida, Cuba (off Channel Yucatan Straits of Florida, Bahamas, Rico Caribbean Straits of Florida, Guadeloupe typical var. Species ▲ Plumarella pellucida Plumarella pourtalesii Plumarella pourtalesii var. obtusa Plumarella pourtalesii robusta Primnoa resedaeformis Primnoella delicatissima Primnoella divaricata Primnoella polita Primnosis rigida Pseudoplexaura porosa? Radicipes gracilis Riisea paniculata Scleracis guadelupensis Scleracis petrosa Stenisis humilis Swiftia casta Swiftia exserta Swiftia koreni Family Primnoidae Primnoidae Primnoidae Primnoidae Primnoidae Primnoidae Primnoidae Primnoidae Isididae Plexauridae Chrysogorgiidae Ellisellidae Plexauridae Plexauridae Isididae Plexauridae Plexauridae Plexauridae Taxon Higher
356 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Humann 1993; Veronique 1987; Humann 1993; Veronique Cairns 2005 Auster and Cairns 2005; Watling 2005 Cairns 2005 Cairns 2005 1987; Bayer 1961; Veronique Cairns 2005 Cairns 2005; Reyes et al. 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 1987 Cairns 2005; Veronique 1987 Cairns 2005; Veronique Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005; and Bayer 2006 Cairns 2005; and Bayer 2006 Cairns 2005; and Bayer 2006 CARIBBEAN 275 >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 >200 23-366 180->200 Depth Range Distribution ♠ Straits of Florida, Guadeloupe Straits of Florida Atlantic Western Atlantic Western Guadeloupe, Barbados Saint Vincent, Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western Guadeloupe Guadeloupe Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western Atlantic Western ) rugulosa Lingnella (= var. Thesa nivea (=
Species ▲ ) Swiftia pallida Swiftia pourtalesii Swiftia sp. Sensu Thelogorgia longiflora Thelogorgia studeri richardii Thelogorgia vossi Thesea antiope Thesea bicolor Thesea gracilis Thesea grandiflora Thesea grandiflora Thesea grandulosa Thesea guadelupensis Thesea hebes Thesea parviflora Thesea rugosa Thesea solitaria Thesea sp. Sensu Thouarella bipinnata Thouarella diadema Thouarella grasshoffi Family Plexauridae Plexauridae Plexauridae Keroeididae Keroeididae Keroeididae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Plexauridae Primnoidae Primnoidae Primnoidae Taxon Higher
357 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ South Atlantic Fishery South Management Council, 1998; Cairns 2005 Cairns 2005 Cairns 2005 Cairns 2005 1987; Cairns 2005; Veronique Reyes et al. 2005 Bayer & Muzik 1977, 1979 Keller et al. 1975 Keller et al. 1975; Picton and Howsen 2002 Keller et al. 1975 Gosner 1978; Bayer 1961; NOAA 2005 Pasternak 1975 Pasternak 1975; Keller et al. 1975 Pasternak 1975 Pasternak 1975; Keller et al. 1975 Cairns 1986 Cairns 1986 Cairns 1986 CARIBBEAN 1600 4160 >200 >200 >200 15-238 50-400 176-275 198-864 159-720 161-545 ~30-4000 1336-6200 ~100-6300 ~1012-4400 ~2610-3500 Depth Range to 108 or more U.S. Distribution ♠ Straits of Florida, Cuba Atlantic Western Atlantic Western Atlantic Western Martinique, Guadeloupe, Colombia Barbados Antilles Southern Lesser Venezuela Southeastern Caribbean Sea Basin Throughout Caribbean, including Puerto Rico Caribbean Sea Trench Puerto Rico Basin, Cayman Islands Southeastern Caribbean (Venezuela) Antilles, Caribbean Southern Lesser Sea Basin Caribbean Sea Basin Channel, Yucatan Straits of Florida, Antilles, including Lesser territories Antilles Southernmost Lesser Antilles Lesser T. T. (=
Species ▲ ) Titanideum frauenfeldii Titanideum suberosum n. sp. Trichogorgia antillarum Villogorgia n. sp. Villogorgia nigrescens Villogorgia Epiphaxum micropora Anthoptilum grandiflorum Funiculina quadrangularis Kophobelemnon irregulatus Renilla reniformis Umbellula thomsoni Umbellula durissima Umbellula hemigymna Umbellula magniflora Crypthelia glossopoma Crypthelia insolita Crypthelia papillosa Family Anthothelidae Chrysogorgiidae Plexauridae Plexauridae Plexauridae Lithotelestidae Anthoptilidae Funiculinidae Kophobelemnidae Renillidae Umbellulidae Umbellulidae Umbellulidae Umbellulidae Stylasteridae Stylasteridae Stylasteridae Taxon Higher Order Helioporacea Order Pennatulacea Class Hydrozoa Order Stylasterina
358 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 CARIBBEAN 1097 68-384 39-261 60-494 139-311 102-311 159-837 125-368 183-366 165-198 333-366 198-309 194-534 196-370 150-282 270-670 545-864 123-368 164-708 150-400 419-708 300-514 267-1170 761-1061 Depth Range U.S. U.S. territories Puerto Rico Distribution ♠ Puerto Rico Greater and Lesser Antilles, including Greater and Lesser U.S. territories Antilles, including Lesser territories Antilles Lesser Antilles Southernmost Lesser Antilles, Lesser Havana Only off Channel Yucatan Straits of Florida, Caribbean Western Havana, Cuba Only off Havana, Cuba Only off Caribbean Western Channel Yucatan Only Straits of Florida, Bahamas Straits of Florida Havana Only off Barbados Only off Antilles Lesser Havana Only off Dominica, Martinique, Montserrat, Saint Kitts and Nevis Straits of Florida Antilles and Mona Passage, Lesser including Throughout Caribbean, including Puerto Rico Antilles, including St. Lucia Only off Species ▲ Crypthelia peircei Crypthelia tenuisepata Distichopora anomala Distichopora barbadensis Distichopora cervina Distichopora contorta Distichopora foliacea Distichopora rosalindae Distichopora sulcata Distichopora uniserialis Distichopora yucatanensis Errina altispina Errina cochleata Lepidopora biserialis Lepidopora carinata Lepidopora clavigera Lepidopora decipiens Lepidopora glabra Lepidotheca brochi Lepidotheca pourtalesi Pliobothrus echinatus Pliobothrus symmetricus Pliobothrus tubulatus Stenohelia pauciseptata Family Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Taxon Higher
359 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Reference ‡ Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 Cairns 1986 CARIBBEAN 823 13-298 0.5-30) 112-377 174-653 183-707 146-965 183-274 198-309 146-530 384-549 200-400) 300-400) 159-2021 Depth Range 42-692 (common 0.5-373 (common 123-759 (common (common 200-650)
Puerto Rico Puerto Rico (Isla de , Bahamas Puerto Rico Distribution ♠ Puerto Rico Lesser Antilles and the Mona Lesser Passage, including Only off Culebra) Cuba Antilles, off Greater Antilles, including Straits of Florida, U.S. territories Barbados, Dominica, Grenada, Martinique, Saint Lucia Widespread Caribbean, including Islands U.S. Virgin Continental slopes of the southeasetern United States to the Peninsula Yucatan Straits of Florida Peninsula Yucatan the Only off Channel Yucatan Straits of Florida, Straits of Florida Antilles, including Lesser Widespread Caribbean, including Puerto Rico southeastern Only off Species ▲ Stylaster antillarum Stylaster atlanticus Stylaster auranticus Stylaster complanatus Stylaster corallium Stylaster duchassaingi Stylaster erubescens Stylaster filogranus Stylaster inornatus Stylaster laevigatus Stylaster miniatus Stylaster profunda Stylaster roseus Stylaster spatula Family Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Stylasteridae Taxon Higher
360 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION S. , and Other Polychaetes; porifera; pyncnogonids Polychaetes; porifera M. oculata , (1 Station); Submersible Alvin Submersible Station); (1 M. carolina Mollusks , Gastropods; pelecypods; scaphopods; brachiopods Arca sp.; Columbarium sp.; Gastropods; Gaza sp.; Natica Turridae sp.; sp.; sp.; Voluta Xenophora sp. Gilliss CARIBBEAN L. pertusa , Echinoderms Holothurians (sea cucumbers); Nymphaster sp.; Ophiuroids; Phormosoma sp.; Zoroaster sp. (6 Stations); R/V Stations); (6 E. profunda
sp.; Gerda sp. sp.; sp. sp.;
; sp (cod) sp. Macrourid Brotulid sp. Dibranchus sp. sp. (dogfish sp. (ray); Merluccius Chaunax pictus Breviraja sp. (eels) Gadidae sp. (deepsea Etmopterus Benthobatis Fish Cruriraja Nettastomid sp. (hake); Chlorophthalmus sp. (deepsea herring); sp.; Neoscopelus (5 Stations); R/V Stations); (5 sp.; plutonia Etmopterus Sources include station data from 28 locations in the wider Caribbean, sampled
Laemonema sp. (skates); sp. (benthic sharks);
(ray); Synaphobranchus Moridae Promyllanter schmitti Galeus Benthobatis plutonia (skates); sharks); (hake); Nettastomatidae Bathyclupea Bembrops Breviraja sp. (fangtooths); (anglerfish); Cruriraja sp. (batfishes); (dogfish shark); ; Laemonema sp. (grenadiers); (hake); codfishes); (lanternfish); sp.; Synaphobranchus
;
; Columbus Iselin Columbus
sp. ; ; sp.; sp.; sp.; Rochinia Stylirostris Galatheid sp.; Palicus
sp.; sp.; Xanthidae Pylocheles Polycheles sp.; sp.; sp.; Munidopsis sp.; sp.; sp.; sp.; lobsters) Nephropsis rosea Glyphocrangon ; sp.; ; (14 Stations); R/V Stations); (14 Nematocarcinua cursor Bathyplax Hymenopenaeus Raninoides Stereomastis crabs, shrimp and Crustacean (deep-seas Homolodramia paradoxa Galatheid scutata longleyi Plesionika acanthonotus Pasiphaeidae sp.; isopods Aristaeomorpha sp.; Benthochascon Crangonidae sp.; Munida Nephropsis Phoberus sp.; sp.; Stylirostris Pillsbury (multiple stations) (Anon 1963; 1964, 1965, 1966, 1968, 1969, 1970, 1972a, 1972b, 1973, 1974; Bayer 1966; Staiger 1968a, Staiger 1966; Bayer 1974; 1973, 1972b, 1972a, 1970, 1969, 1968, 1966, 1965, 1964, 1963; (Anon stations) (multiple Other Eastward coelenterata "Solitary corals"; "Medusae" "Solitary corals"; actinarians An aggregate inventory of the benthos associated with the major structure-forming corals ;
; ; ; throughout the wider Caribbean. Benthos is grouped by coral species. species Potential structure and habitat- forming E. profunda L. pertusa S. varibilis gorgonians; hydroids; octocorals L. pertusa S. varibilis E. profunda gorgonians; "corals" Appendix 8.2. varibilis R/V vessels: research following the by R/V and station); (multiple species many that note *Please overlap). locations station (some S letter the by 3 Figure in identified are Locations 1966b). and 1966a Voss 1971; 1969, 1968b, noted were first identifications and spelling is presented as it appears on the original station sorting sheets.
361 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Other Algae; "Large sponge 18" across"; oolite chunk; polychaetes; bryozoans; tunicates Mollusks Gastropods; pelecypods; brachiopods CARIBBEAN Echinoderms Asteroidea; crinoids; echinoids; holothurians (sea cucumbers); ophiuroids; starfish; brittle stars; urchins ; Fish Calamopteryx goslinei elasmobranch egg cases ;
; ;; Munida ; sp.; Nephropsis Majidae ; Cancellus Homaridae Myropsis sp.; ; sp.; Paguristes sp.; lobsters) ; Cirripedia ; Iliacantha subglobosa Munida irrasa crabs, shrimp and Crustacean (deep-seas Anomalothir furcillatus Brachyura oratus Clibanarius anomalus Galatheid sp.; Leucosiidae sp.; schroederi quinquespinosa aculeate spinipes; Pandalidae sp.; Plesionika acanthonotus; Podochela curvirostris ; Pyromia cuspidate; Sicyonia stimpsoni; stomatopods; Sympagurus pictus
sp. sp. Other coelenterata "Solitary corals"; Actiniaria (anemones); Alcyonaria (soft corals); Hydroids; Urochordata sp. (encrusting gorgonians); zoanthids sp. sp. species Potential structure and habitat- forming M. carolina "Corals"; Antipatharia (black corals); Stylasterina (hydrocorals)
362 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
Other "Grapefruit sponges"; Cladorhiza sp (potato sponges); Hexactinellida sp. (glass sponges); Hyalonema sp.; polychaetes
;
sp.; Spirula sp.; sp.; sp.; Turridae sp. ; (bivalve, Limopsis Melongena Mitra Mollusks "Squid, unidentified"; Acesta (Clams); Arenatus Gaza superba Leptothyra Leucosyrinx sp.; sp.; sp; Pseudamusium dallil clam?); scaphopods; Solariella solenogaster (shelless mollusk); spirula sp. CARIBBEAN Echinoderms "Pancake urchin"; "Soft urchin"; asteroids; Benthopectinids; Crinoids; Goniaster sp.; holothurians (sea cucumbers); Molpadia sp. (sea cucumbers); Nymphaster sp.; ophuiroids; Phormosoma sp. (sea biscuts); Psilaster squameus; Zoroaster fulgens
sp.
Chaunax sp.; Apristurus Dibranchus Diaphus Barathronus Bathypterois Fish sp.; sp. (batfishes); Benthobatis sp. (surgeonfish); (Atlantic batfish) sp. (fangtooths); (anglerfish); Dibranchus Acanthurus Alepocephalus sp. (cat sharks); sp. (cusk-eels); (tripod fish); Brotulid pictus sp. (headlightfish); atlanticus ; Dicrolene intronigra; Diretmus sp. (dorys); Gonostomatidae sp. (bristlemouth); Halosaurs sp.; Hariotta sp. (chimaera); Ilyophis sp. (eels); Macrourid sp. (grenadiers); Monomitopus sp. (cusk-eels); Myctophid sp. (lanternfish ); Myxini (hagfish); Neoscopelus sp. (lanternfish); Nesiarchus sp. (scabbardfish); Notacanthid sp. (spiny eel); Oxygadus (rattails); Promyllantor schmitti; Saurida sp. (lizardfish); Searsid sp.; Sternoptychidae sp. (hatchetfish); Stomias sp. (dragon fish ); Synaphobranchus oregoni Synaphobranchus sp.
sp.;
sp.; sp.;
sp.; ; Gennadas Uroptychus Pontophilus sp.; sp.; Axiidae sp.; Pandalidae Sergestes sp.; sp. sp.; sp.; sp.; Bathysquilla sp.; sp.; sp.; isopods; sp.; Majidae sp.; Aristaeinae sp.; Munidopsis sp.; crucifer ; sp.;
sp.; lobsters) sp.; Uroptychus sp.; ; Aristeus Bathymunida Galatheid Glyphocrangon Nematocarcinus Nephropsis Parapagurus Prionocrangon Spongicoloides Stereomastis crabs, shrimp and Uroptychus Crustacean (deep-seas "Various crustaceans"; "Various Acanthephyra Amphipods sp.; sp.; Bathypalaemonella Bathyplax sp.; sp.; Gnathophausia Heterocarpus Lithodid Munida sp.; sp.; Oplophorus sp.; Plesiopenaeus Polycheles Polychelidae sp.; Scalpellum sp.; sp.; Stylodactylus nitidus sp; Verrucomorphs Other coelenterata "Solitary corals"; Deltocyathus sp.; Desmophyllum sp.; Hydroids; Odontocyathus coronatus; Pennatulidae sp. (sea pens); Stephanocyathus diadema; sp. Telestula (anemones); aoanthids
sp. sp. sp. ; "pink sp.; (sea sp. Cirripathes ; species Potential structure and habitat- forming M. oculata S. varibilis gorgonians"; "various coelenterates"; Acanella (bamboo corals); Bathypathes patula Calytrophora pourtalesi fan / sea whip); Chrysogorgia (sea fan / sea whips); sp. (sea whips); gorgonians; Iridogorgia (spiral corals); Keratoisis Plumarella
363 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION Other Cladorhiza sp. (potato sponges); polychaetes; pyncnogonids
sp. sp.; ; sp.; sp.; Acesta Turridae ; pteropods; Mollusks "Finned octopod"; "unidentified squid"; sp. (Clams); brachiopods; Dentalium Naticidae Ornithoteuthis antillarum Propeamusium dalli scaphopods; Solariella Solenogaster sp.; ;
;
CARIBBEAN ;
; Ophidiaster Solaster Echinoderms Ceramaster elongatus (Cookie starfish); Ceramaster sp.; cidarids; Circeaster americanus Lophaster verrilli Nymphaster sp.; sp.; caribbea Spatangoids Zoroaster ackleyi sp.
sp.
Saurida ; sp. (rattails); Gadomus Chaunax Fish sp. (cusk-eels); Chlorophthalmus sp.; sp. (sea robins); sp. (grenadiers); (Bathygadid family); sp. (fangtooths); Dibranchus atlanticus Brotulid Callionymus (Anglerfish); sp.; (Atlantic batfish); longifilis Hymenocephalus Macrourid Monomitopus Peristedion Promyllantor schmitti (lizardfish)
; sp.;
sp.; sp.;
sp.; sp.;
sp.; sp.; Sergestes Uroptychus Oplophorus sp.; Gennadas Pylopagurus sp.; sp.; isopods; sp.; Polycheles Majidae sp.; Aristaeinae Chirostylus Munidopsis sp.; sp.; Glyphocrangon sp.; lobsters) sp.; Glyphocrangon Polychelidae ; sp.; sp.; ; sp.; sp.; ; Glyphocrangon Nematocarcinus Paguristes Prionocrangon Spongicoloides Stereomastis crabs, shrimp and Rochinia Crustacean (deep-seas Acanthephyra amphipods; Axiidae Galatheid sp.; aculeate alispina Heterocarpus Lithodid Mixtopagurus paradoxus Munida sp.; Nephropsis sp.; Penaeid crucifer sp.; Processa sp; sp.; sp.; Stylodactylus Systellaspis sp. sp. ; sp. ; Other coelenterata "Solitary corals"; Odontocyathus coronatus Stephanocyathus diadema Telestula (anemones); Clavulariidae
sp. sp.; E. sp. ; ;
; "corals"; Nicella ; Keratoisis species Potential structure and . pertusa habitat- forming S. varibilis M. oculata L profunda Acanella sp. (bamboo corals); Callogorgia Chrysogorgia (sea fan / sea whips) sp. (bamboo corals); sp.; octocorals; Primnosis
364 STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION
Appendix 8.3. Commercially important deep-sea fish of the wider Caribbean (FAO 1993).
Other Species Depth (m) Snappers: Cladorhiza sp. (potato sponges); polychaetes; pyncnogonids Apsilus dentatus (black snapper) 70 – 290
sp. Etelis oculatus (queen or green snapper) 180 – 463 sp.; ; sp.; sp.; Lutjanus buccanella (blackfin snapper) 80 – 180 Acesta Turridae
; pteropods; L. vivanus (silk snapper) 60 – 210 Mollusks Rhomboplites aurorubens (vermilion snapper) 80 – 230 "Finned octopod"; "unidentified squid"; sp. (Clams); brachiopods; Dentalium Naticidae Ornithoteuthis antillarum Propeamusium dalli scaphopods; Solariella Solenogaster sp.; ;
Sharks: ;
CARIBBEAN
; Centrophorus granulosus (gulper shark) to 210
; Eugomphodus taurus (grey nurse shark) to 210 Hexanchus vitulus (bigeyed sixgill shark) to 250 Ophidiaster Solaster
Echinoderms Mustelus canis (smooth dogfish) to 250 Ceramaster elongatus (Cookie starfish); Ceramaster sp.; cidarids; Circeaster americanus Lophaster verrilli Nymphaster sp.; sp.; caribbea Spatangoids Zoroaster ackleyi Scyliorhinus boa (catshark) to 270 sp.
Sphyma lewini (scalloped hammerhead shark) to 170
sp. Squalus cubensis (Cuban dogfish) to 300
Promthicthys Prometheus (rabbit-fish) to 170 Saurida ; Groupers & hinds: sp. (rattails); Gadomus Chaunax Epinephelus adscensionis (rock hind) 0 – 120 Fish sp. (cusk-eels); Chlorophthalmus sp.; E. flavolimbatus (yellowedge grouper) to 132 sp. (sea robins); sp. (grenadiers);
(Bathygadid family); E. fulvus (coney) 0 – 230 sp. (fangtooths); E. guttatus (red hind) 0 – 200 Dibranchus atlanticus E. morio (red grouper) to 174 Brotulid Callionymus (Anglerfish); sp.; (Atlantic batfish); longifilis Hymenocephalus Macrourid Monomitopus Peristedion Promyllantor schmitti (lizardfish)
E. mystacinus (misty grouper) 180 – 270
;
sp.; Mycteroperca phenax (scamp) to 174
sp.; sp.;
sp.;
sp.; Neoscombrops sp. to 190
sp.; sp.; Sergestes Other: Uroptychus Oplophorus sp.; Gennadas Pylopagurus sp.; sp.; isopods;
sp.; Erythrocles monody (Atlantic rubyfish) to 300 Polycheles Majidae sp.; Aristaeinae Chirostylus Munidopsis sp.; sp.; Glyphocrangon
sp.; Gephyroberix darwini to 310 lobsters) sp.; Glyphocrangon Polychelidae ; sp.; sp.; ; sp.; sp.; ; Ostichthys trachypoma (bigeye soldierfish) to 290 Glyphocrangon Nematocarcinus Paguristes Prionocrangon Spongicoloides Stereomastis crabs, shrimp and Rochinia Polymixia lowel (beardfish) to 260 Crustacean (deep-seas
Acanthephyra amphipods; Axiidae Galatheid sp.; aculeate alispina Heterocarpus Lithodid Mixtopagurus paradoxus Munida sp.; Nephropsis sp.; Penaeid crucifer sp.; Processa sp; sp.; sp.; Stylodactylus Systellaspis sp. Pristipomoides sp. (jobfish) 180 – 300+
sp. ; sp. ; Other coelenterata "Solitary corals"; Odontocyathus coronatus Stephanocyathus diadema Telestula (anemones); Clavulariidae
sp. sp.; E. sp. ; ;
; "corals"; Nicella ; Keratoisis species Potential structure and . pertusa habitat- forming S. varibilis M. oculata L profunda Acanella sp. (bamboo corals); Callogorgia Chrysogorgia (sea fan / sea whips) sp. (bamboo corals); sp.; octocorals; Primnosis
365