Virgin Islands

diversity Review A Unique Coral Community in the Mangroves of Hurricane Hole, St. John, US Virgin Islands Caroline S. Rogers ID Wetland and Aquatic Research Center, US Geological Survey, St. John, VI 00830, USA; caroline_rogers@usgs.gov; Tel.: +1-340-693-8950 Received: 9 June 2017; Accepted: 1 August 2017; Published: 4 August 2017 Abstract: Corals do not typically thrive in mangrove environments. However, corals are growing on and near the prop roots of red mangrove trees in Hurricane Hole, an area within the Virgin Islands Coral Reef National Monument under the protection of the US National Park Service in St. John, US Virgin Islands. This review summarizes current knowledge of the remarkable biodiversity of this area. Over 30 scleractinian coral species, about the same number as documented to date from nearby coral reefs, grow here. No other mangrove ecosystems in the Caribbean are known to have so many coral species. This area may be a refuge from changing climate, as these corals weathered the severe thermal stress and subsequent disease outbreak that caused major coral loss on the island’s coral reefs in 2005 and 2006. Shading by the red mangrove trees reduces the stress that leads to coral bleaching. Seawater temperatures in these mangroves are more variable than those on the reefs, and some studies have shown that this variability results in corals with a greater resistance to higher temperatures. The diversity of sponges and fish is also high, and a new genus of serpulid worm was recently described. Continuing research may lead to the discovery of more new species. Keywords: corals; mangroves; climate change refuge; thermal stress; US Virgin Islands 1. Introduction For hundreds of years, four bays within Hurricane Hole, St. John, US Virgin Islands, have been a refuge for boats during major storms (Figure 1). Hurricane Hole is a beautiful seascape with coral reefs, mangroves, seagrass beds and a remarkable biodiversity in corals, fishes, sponges and other organisms. This area is part of the Virgin Islands Coral Reef National Monument established in 2001 by President Clinton through a Presidential Proclamation The particular biological and physical oceanographic conditions within these four mangrove-lined bays within Hurricane Hole (which range in size from 0.06 to 0.11 km2 ) appear to be providing a refuge for corals at a time of changing climate [1], (Figure 2). Corals and the reefs they create are at risk from numerous human activities around the world [2]. Corals do not usually thrive among mangroves because of the typically turbid and low (and sometimes high) salinity waters. Here in Hurricane Hole there are no permanent streams entering the bays, with the only freshwater coming from rainfall and possibly from groundwater. The mangroves provide shade, attenuating more than 70% of the photosynthetically active radiation (PAR) [1]. In addition, the proximity of seagrass beds, mangroves, and patch reefs, and the particular hydrographic conditions and biological influences of these habitats on seawater chemistry, generate chemical conditions that buffer against lower pH (ocean acidification). Ocean acidification reduces the rates at which calcifiers such as corals and coralline algae grow and causes carbonate sediment dissolution [3]. Diversity 2017, 9, 29; doi:10.3390/d9030029 www.mdpi.com/journal/diversity Diversity 2017, 9, 29 2 of 16 Diversity 2017, 9, 29 Diversity 2017, 9, 29 2 of 16 2 of 16 Figure 1. 1.Location of St. John, US Virgin Islands, Islands, and andthe themangrove-lined mangrove-lined bays Hurricane Hole, Figure bays in in Hurricane Hole, Figure 1. Location Location of of St. St.John, John,US USVirgin Virgin Islands, and the mangrove-lined bays in Hurricane Hole, within Virgin Islands Coral Reef National Monument. within Virgin Virgin Islands within IslandsCoral CoralReef ReefNational NationalMonument. Monument. (a) (b) (a) (b) Figure 2. Cont. Diversity 2017, 9, 29 3 of 16 Diversity 2017, 9, 29 3 of 16 (c) (d) Figure 2. Red Mangroves fringe the shorelines of the bays in Hurricane Hole and their prop roots Figure 2. Red Mangroves fringe the shorelines of the bays in Hurricane Hole and their prop roots extend into the clear waters below (a,b). Corals grow on and among the prop roots and on nearby extend into the clear waters below (a,b). Corals grow on and among the prop roots and on nearby hard hard substrata (c,d). All photos in this article were taken by the author. substrata (c,d). All photos in this article were taken by the author. When the Virgin Islands Coral Reef Monument was first established by President Clinton, the When Islands Coral Reef Monument was first established Clinton, significance ofthe theVirgin mangroves in Hurricane Hole, particularly their rolebyasPresident a nursery, was the significance of the mangroves in Hurricane Hole, particularly their role as a nursery, was highlighted highlighted (see text box). However, it was not until 2009 that the extraordinary diversity of the (seewas text recognized. box). However, it was not until 2009 thatUS the National extraordinary the corals corals Active management by the Park diversity Service, of including thewas recognized. Active management by the US National Park Service, including the installation of a storm installation of a storm mooring system, is no doubt helping to protect this diverse ecosystem. The mooring system, is no doubt helping to protect this diverse ecosystem. The secure mooring system secure mooring system decreases the likelihood that boats will break free or drag their anchors and decreases likelihood that boats will break free or drag their anchors and go up into the mangroves. go up into thethe mangroves. Excerpt from Proclamation 7399 of January 17 January 2001 TextText box:box: Excerpt from Proclamation 7399 of 17 2001 Establishment of the Virgin Islands Coral Reef National Monument Establishment of the Virgin Islands Coral Reef National Monument The Virgin Islands CoralCoral ReefReef National Monument, in the lands off the island of St. John The Virgin Islands National Monument, in submerged the submerged lands off the island of St. John in theinU.S. Islands, contains all of the of a Caribbean tropicaltropical marine ecosystem. This the Virgin U.S. Virgin Islands, contains all elements of the elements of a Caribbean marine ecosystem. designation furthers the protection of the scientific included in included the Virgin National This designation furthers the protection of theobjects scientific objects inIslands the Virgin Islands Park,National created in 1956created and expanded in 1962. ---------Park, in 1956 and expanded in 1962. ———­ The submerged monument lands within Hurricane Hole include the most extensive and The submerged monument lands within Hurricane Hole include the most extensive and well-developed mangrove habitat on St. John. The Hurricane Hole area is an important nursery area well-developed mangrove habitat on St. John. The Hurricane Hole area is an important nursery area for reef-associated fish and invertebrates, instrumental in maintaining water quality by filtering and for reef-associated fish and invertebrates, instrumental in maintaining water quality by filtering trapping sediment and debris in fresh water runoff from the fast land, and essential to the overall and trapping sediment and debris in fresh water runoff from the fast land, and essential to the functioning and productivity of regional fisheries. Numerous coral reef-associated species, including overall functioning and productivity of regional fisheries. Numerous coral reef-associated species, the spiny lobster, queen conch, and Nassau grouper, transform from planktonic larvae to including the spiny lobster, queen conch, and Nassau grouper, transform from planktonic larvae to bottom-dwelling juveniles in the shallow nearshore habitats of Hurricane Hole. As they mature, they bottom-dwelling juveniles in the shallow nearshore habitats of Hurricane Hole. As they mature, move offshore and take up residence in the deeper coral patch reefs, octocoral hardbottom, and algal they move offshore and take up residence in the deeper coral patch reefs, octocoral hardbottom, and plains of the submerged monument lands to the south and north of St. John. algal plains of the submerged monument lands to the south and north of St. John. 1.1. 1.1. Stony Corals Stony Corals Remarkably, over 30 species of scleractinian corals grow on or to the prop roots of the red red Remarkably, over 30 species of scleractinian corals grow on close or close to the prop roots of the mangroves along thethe shores the bays bays(Princess (PrincessBay, Bay, Otter Creek, Water Creek). mangroves along shoresofofthree three of of the Otter Creek, andand Water Creek). (Table 1, (Table 1, Figure 3). Only a few species found on nearby coral reefs (such as Mussa angulosa, and Figure 3). Only a few species found on nearby coral reefs (such as Mussa angulosa, and Madracis spp.) Madracis spp.) have not been seen in these bays, and a few have been observed in the bays but not listed as occurring on the reefs (see Table 1). No other mangrove ecosystems in the Caribbean are Diversity 2017, 9, 29 4 of 16 have not been seen in these bays, and a few have been observed in the bays but not listed as occurring on the reefs (see Table 1). No other mangrove ecosystems in the Caribbean are known to harbor so many coral species. The species present include major reef framework building species such as brain corals (Colpophyllia natans; Diploria labyrinthiformis) and star corals (Orbicella spp.), as well as species with smaller colonies such as Porites astreoides. Corals in the genus Orbicella are some of the most abundant corals on the reefs around St. John and in the mangroves. Surprisingly, some corals that are quite rare on the coral reefs and more often found in deeper water, such as Mycetophyllia spp. and Scolymia spp., are found growing in the shade of the mangroves. Some corals also grow on patch reefs in deeper water in the bays. The sides of the bays slope to the sea floor to a depth of about 5 m and then the floor slopes more gradually to the centers of the bays. Corals are more numerous near the entrances to the bays, but, surprisingly, can also be found in low numbers in the calmer, siltier portions at the head of the bays. The middle of the bays where depths reach 10 to 14 m can be turbid. Generally the water is quite clear right along the shore likely due to filter feeding organisms like tunicates, sponges and tree oysters which grow there [4,5]. Six of the seven Caribbean coral species listed as threatened under the US Endangered Species Act (Mycetophyllia ferox has not been reported) are found in Hurricane Hole, with four of them in these mangrove-fringed bays (Orbicella annularis, O. faveolata, O. franksi, Dendrogyra cylindrus). Three small colonies of one of the other listed species, Acropora palmata, were also observed to have settled there but did not survive to grow into larger colonies. Acropora cervicornis has not been found. Table 1. Stony corals in Princess Bay, Otter Creek, and Water Creek. Coral Species Princess Bay Otter Creek Water Creek Stephanocoenia intersepta Acropora palmata Undaria agaricites Agaricia spp. Siderastrea siderea Siderastrea radians Porites astreoides Porites porites Porites furcata Porites divaricata Favia fragum Diploria labyrinthiformis Pseudodiploria clivosa Pseudodiploria strigosa Manicina areolata Colpophyllia natans Colpophyllia amaranthus 1 Cladocora arbuscula 1 Orbicella annularis Orbicella faveolata Orbicella franksi Montastraea cavernosa Solenastrea bournoni Phyllangia americana 1 Oculina diffusa 1 Meandrina meandrites Dichocoenia stokesi Dendrogyra cylindrus Scolylmia cubensis Scolymia lacera Mycetophyllia spp. Eusmilia fastigiata Tubastrea aurea (hydrozooan) Millepora spp. Totals x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 19 x x x x x x x x x x x x x x x x x x x x x x 30 x x x x x x x x x x x x x x x x x 26 1 species not reported to date from the outer reefs. Species identifications confirmed with colleagues and based on identification guides at www.agrra.org. Tubastrea aurea (hydrozooan) Millepora spp. Totals x 19 x x 30 x x 26 1 species Diversity 2017, 9,not 29 reported to date from the outer reefs. Species identifications confirmed with colleagues 5 of 16 and based on identification guides at www.agrra.org. (a) (b) (c) (d) (e) (f) (g) (h) Figure 3. Cont. Diversity 2017, 9, 29 6 of 16 Diversity 2017, 9, 29 6 of 16 (i) (j) (k) Figure 3. Stony corals in the mangrove-lined bays of Hurricane Hole. (a) a colony of Orbicella Figure 3. Stony corals in the mangrove-lined bays of Hurricane Hole. (a) a colony of Orbicella annularis; (b)Orbicella a colonyfaveolata of Orbicella with a small Orbicella sp. recruit on in thefront rockofjust (b) annularis; a colony of withfaveolata a small Orbicella sp. recruit visible on thevisible rock just it; in of it; (c)natans Colpophyllia natans (foreground) Orbicella faveolata; (d) Siderastrea siderea;agaricites (e) Undaria (c) front Colpophyllia (foreground) and Orbicella and faveolata; (d) Siderastrea siderea; (e) Undaria agaricites and Millepora alcicornis; (f) Poritesfurcata; (g) Porites astreoides; (h) Diploria labyrinthiformis and Millepora alcicornis; (f) Poritesfurcata; (g) Porites astreoides; (h) Diploria labyrinthiformis and Orbicellaand Orbicella annularis surrounded by Porites porites; (i) Eusmilia(j) fastigiata; (j)sp.; Scolymia sp.; (k) Mycetophyllia annularis surrounded by Porites porites; (i) Eusmilia fastigiata; Scolymia (k) Mycetophyllia aliciae aliciae with Porites astreoides in the background. with Porites astreoides in the background. These shallow, mangrove-associated coral communities differ from “true” or “framework” These shallow, mangrove-associated coral communities differ from “true” or “framework” coral coral reefs, which are rigid, topographically complex structures developed from carbonate accretion reefs, which are rigid, topographically complex structures developed from carbonate accretion by by corals and other cementing and calcifying organisms and the product of biological and geological corals and other cementing and calcifying organisms and the product of biological and geological processes [6,7]. The corals in the mangroves are growing on prop roots or hard non-limestone processes [6,7]. The corals in the mangroves are growing on prop roots or hard non-limestone substrata substrata nearby, and are not currently depositing layers of limestone that accrete into structurally nearby, and are not currently depositing layers of limestone that accrete into structurally complex reefs. complex reefs. Although it is not possible to rigorously quantify the abundance of corals in the complex, Although it is not possible to rigorously quantify the abundance of corals in the complex, three-dimensional space created by the red mangroves fringing the bays, because the roots make it three-dimensional space created by the red mangroves fringing the bays, because the roots make it impossible to swim above the corals and count them, it is clear that the relative abundance of coral impossible to swim above the corals and count them, it is clear that the relative abundance of coral species differs between the mangrove communities and the coral reefs. Three species of star corals species differs between the mangrove communities and the coral reefs. Three species of star corals of of the genus Orbicella grow in Hurricane Hole and on the reefs around St. John and are the most the genus Orbicella grow in Hurricane Hole and on the reefs around St. John and are the most abundant corals on the true reefs around the island. Scientists have surveyed transects along several abundant corals on the true reefs around the island. Scientists have surveyed transects along several reefs as part of an ongoing long-term monitoring program [8]. Results of these statistically rigorous reefs as part of an ongoing long-term monitoring program [8]. Results of these statistically rigorous surveys show the changes in coral cover and relative abundance of corals over time, since the inception surveys show the changes in coral cover and relative abundance of corals over time, since the of the program in 1989. The reefs have been affected by hurricanes, overfishing, sedimentation, and inception of the program in 1989. The reefs have been affected by hurricanes, overfishing, vessel groundings. However, the most dramatic changes were associated with the highest seawater sedimentation, and vessel groundings. However, the most dramatic changes were associated with temperatures on record in the Caribbean in the late summer of 2005 [9]. Temperatures exceeded 29.5 ◦ C the highest seawater temperatures on record in the Caribbean in the late summer of 2005 [9]. Temperatures exceeded 29.5 °C in August and September, resulting in over 90% of the coral cover becoming bleached [8], (Figure 4). Bleaching occurs when corals lose their symbiotic algae Diversity 2017, 9, 29 7 of 16 Diversity 2017,and 9, 29September, resulting in over 90% of the coral cover becoming bleached [8], (Figure 7 of 4). 16 in August Bleaching occurs when corals lose their symbiotic algae (“zooxanthellae”). The result is a conspicuous (“zooxanthellae”). The resulttoiscompletely a conspicuous change in color, from paling to completely white, and change in color, from paling white, and partial or complete mortality if bleaching persists partial or complete mortality if bleaching persists for too long. Bleaching episodes are predicted to for too long. Bleaching episodes are predicted to become more severe and more frequent in the future become moretemperatures severe and more frequent in [10]. the future as sea water temperatures continue to rise [10]. as sea water continue to rise Figure 4. Extensive Extensive bleaching bleaching of of Orbicella colonies off northwestern St. John in November 2005. Figure As of 2005, 2005, bleached bleached coral coral colonies colonies gradually gradually regained As water water temperatures temperatures cooled cooled in in the the fall fall of regained their normal coloration. However, an outbreak of white plague disease peaked 2 their normal coloration. However, an outbreak of white plague disease peaked 2 to to 66 months months (at (at five five different different locations) locations) after after the the peak peak in in bleaching bleaching [8]. [8]. The The disease disease affected affected all all coral coral species species and and resulted resulted in living coral cover by by thethe endend of 2007. The The decline was associated with in an an average averagedecline declineofof60% 60%inin living coral cover of 2007. decline was associated partial and complete mortality of coral colonies. Orbicella spp. were affected more than other species with partial and complete mortality of coral colonies. Orbicella spp. were affected more than other but remained the mostthe abundant following the outbreak. Corals on reefson in reefs the other Virgin species but remained most abundant following the outbreak. Corals in theUS other US Islands, British Virgin Islands, and Puerto Rico also bleached severely in 2005 with subsequent Virgin Islands, British Virgin Islands, and Puerto Rico also bleached severely in 2005 with subsequent disease disease outbreaks. outbreaks. The The relationships relationships among among high high sea sea water water temperatures, temperatures, bleaching, bleaching, and and diseases diseases are are complex. complex. Bleaching episodes are not always followed by disease outbreaks, which are sometimes not Bleaching episodes are not always followed by disease outbreaks, which are sometimes not preceded preceded by bleaching. compelling evidence manyare diseases are associated with by bleaching. However,However, compelling evidence exists thatexists manythat diseases associated with unusually unusually warm temperatures, and ocean temperatures are predicted to continue increasing in the warm temperatures, and ocean temperatures are predicted to continue increasing in the future. future. Diseases are increasing in prevalence and becoming more severe. They are challenging to Diseases are increasing in prevalence and becoming more severe. They are challenging to study study and few pathogens or causes haveidentified been identified conclusively and few pathogens or causes have been conclusively [11]. [11]. The significance of the mangrove-lined bays as habitats corals not recognized The significance of the mangrove-lined bays as habitats for for corals waswas not recognized until until 2009, 2009, and no baseline studies of corals in the mangroves in Hurricane Hole were undertaken before and no baseline studies of corals in the mangroves in Hurricane Hole were undertaken before or or during the 2005 to 2007 bleaching/disease event. However, the sizes (over 0.5 to 0.75 m across) during the 2005 to 2007 bleaching/disease event. However, the sizes (over 0.5 to 0.75 m across) of of many many colonies colonies of of the the major major reef-building reef-building corals corals in in the the mangroves mangroves and and their their generally generally intact intact condition condition with few dead dead portions portions indicate indicate that that they they were were present present in in 2005 2005and andsurvived survivedthe thebleaching/disease bleaching/disease with few episode. Some of the largest and most numerous corals there are colonies of boulder episode. Some of the largest and most numerous corals there are colonies of boulder brain brain corals corals (Colpophyllia natans) and grooved brain corals (Diploria labyrinthiformis). Both of these species seem to (Colpophyllia natans) and grooved brain corals (Diploria labyrinthiformis). Both of these species seem be relatively more abundant in inthe to be relatively more abundant themangroves mangrovesthan thanon onthe thereefs reefs nearby. nearby. Although Although no no rigorous rigorous comparison has been made, most C. natans in the mangroves appear to be in better condition (i.e., comparison has been made, most C. natans in the mangroves appear to be in better condition (i.e., dead dead portions are rare) than on the reefs (Figure 5). portions are rare) than on the reefs (Figure 5). Diversity 2017, 9, 29 8 of 16 Diversity 2017, 9, 29 8 of 16 Diversity 2017, 9, 29 8 of 16 Figure 5. Some of the Colpophyllia natans (above) and Orbicella spp. colonies in the mangroves are Figure 5. Some of the Colpophyllia natans (above) and Orbicella spp. colonies in the mangroves are close close to 1 m across. to 1 m across. Figure 5. Some of the Colpophyllia natans (above) and Orbicella spp. colonies in the mangroves are In 2010, another less severe bleaching episode occurred allowing comparison of the responses of close to 1 m across. these two species.less Successive observations (and occurred photographs) of colonies in theofmangroves In 2010, another severe bleaching episode allowing comparison the responses documented much less bleaching in Colpophyllia natans than in Diploria labyrinthiformis (Figure 6), and of these two species. Successive observations (and photographs) of colonies in the mangroves In 2010, another less severe bleaching episode occurred allowing comparison of the responses of only low amounts of complete mortality (0 to 7.1%, respectively). Most colonies recovered completely documented much less bleaching in Colpophyllia natans than in Diploria labyrinthiformis (Figure 6), these two species. Successive observations (and photographs) of colonies in the mangroves [12]. Because of the low these species on reefinrespectively). transects, no rigorous could documented much less bleaching inofmortality Colpophyllia natans than Diploria labyrinthiformis (Figure 6), and and only low amounts ofabundance complete (0 to the 7.1%, Mostcomparison colonies recovered be made of the severity of or responses to thermal stress between these two habitats. only low[12]. amounts of complete (0 to 7.1%, Most completely completely Because of the mortality low abundance ofrespectively). these species oncolonies the reefrecovered transects, no rigorous Bleaching results high temperatures andon high [13].noInrigorous 2010, thecomparison temperatures in [12]. Because themade lowfrom abundance of these the irradiance reef transects, could comparison couldofbe of the severity of species or responses to thermal stress between these two habitats. the mangroves as recorded by in situ thermometers at a depth of less than 1 m reached both higher be made of results the severity ofhigh or responses to thermal stress between these[13]. two habitats. Bleaching from temperatures and high irradiance In [1]. 2010, in and lower values than thosehigh recorded on the reefs at a depth of about Atthe firsttemperatures glance, the Bleaching results from temperatures and high irradiance [13]. 10 In m 2010, the temperatures in the mangroves as recorded by in situ thermometers at a depth of less than 1 m reached both higher and mangroves would not seembytoinbesitu offering a refuge at from higher temperatures. However, the mangroves as recorded thermometers a depth of less than 1 m reached bothvariable higher lowertemperatures values than those recorded on thewith reefscorals at a depth of about 10 mresistance [1]. At first glance, theinmangroves have been associated that have greater to bleaching some and lower values than those recorded on the reefs at a depth of about 10 m [1]. At first glance, the studies [14]. Research is underway to characterize the genotypes and microbial communities would not seem to be offering a refuge from higher temperatures. However, variable temperatures mangroves would not seem to be offering a refuge from higher temperatures. However, variable with colonies fromthat Hurricane Hole resistance to seehave if they provide clues to bleaching their responses to have associated been associated with corals have greater to bleaching in some studies [14]. Research temperatures have been associated with corals that greater resistance to in some temperature increases or other parameters such as irradiance levels [15]. is underway to characterize genotypes microbial associated colonies from studies [14]. Research isthe underway to and characterize thecommunities genotypes and microbialwith communities associated colonies from Hurricane Hole to see responses if they provide clues to their responsesortoother Hurricane Holewith to see if they provide clues to their to temperature increases temperature increases or other parameters such as irradiance levels [15]. parameters such as irradiance levels [15]. Figure 6. A bleached colony of Diploria labyrinthiformis next to an unbleached Colpophyllia natans. Figure A bleached colony Diplorialabyrinthiformis labyrinthiformis next Colpophyllia natans. Figure 6. A6.bleached colony ofof Diploria nexttotoananunbleached unbleached Colpophyllia natans. Diversity 2017, 9, 29 9 of 16 1.2. Fishes About 80 species of fish have been recorded from Hurricane Hole [16] (Table 2, Figure 7). Gray Snappers (Lutjanus griseus) are among the largest and most abundant. Large parrotfishes are noticeably scarce. Queen Angelfishes (Holocanthus ciliarus) are conspicuous, perhaps because they feed on the numerous sponges. Schools of grunts are often seen among the roots, and the three species of trunkfishes are present. Table 2. Fish species in Hurricane Hole, based on Boulon (1992) [16] with recent additions. Source of names is Fishbase or World Register of Marine Species. Scientific Name Abudefduf saxatilis Acanthurus bahianus Acanthurus chirurgus Acanthurus coeruleus Aluterus monocerus Amblycirrhitus pinos Anisotremus virginicus Apogon binotatus Archosargus rhomboidalis Atherinomorus stipes Aulostomus maculatus Balistes vetula Calamus bajonado Cantherhines pullus Canthigaster rostrata Carangoides bartholomaei Caranx ruber Chaetodon capistratus Chaetodon sedentarius Chilomycterus spp. Coryphopterus glaucofraen Coryphopterus personatus Diodon hystrix Epinephelus striatus Epinephelus guttatus Eucinostomus argenteus Haemulon aurolineatum Haemulon chrysargyreum Haemulon flavolineatum Haemulon macrostomum Haemulon plumierii Haemulon sciurus Halichoeres bivittatus Halichoeres garnoti Halichoeres radiatus Histrio histrio Holacanthus ciliaris Holocentrus adscensionis Hypoplectrus spp. Hyporamphus unifasciatus Jenkinsia lamprotaenia Lactophrys bicaudalis Lactophrys trigonus Lutjanus analis Lutjanus buccanella Lutjanus griseus Lutjanus jocu Lutjanus mahogoni Lutjanus synagris Malacoctenus macropus Common Name Sergeant Major Ocean Surgeonfish Doctorfish Blue Tang Unicorn Filefish Redspotted Hawkfish Porkfish Barred Cardinalfish Sea Bream Hardhead Silverside Trumpetfish Queen Triggerfish Jolthead Porgy Orangespotted Filefish Sharpnose Puffer Yellow Jack Bar Jack Foureye Butterflyfish Reef Butterflyfish Burrfish Bridled Goby Masked Goby Porcupinefish Nassau Grouper Red Hind Spotfin/Silver Mojarra Tomtate Smallmouth Grunt French Grunt Spanish Grunt White Grunt Bluestriped Grunt Slippery Dick Yellowhead Wrasse Puddingwife Sargassum Frogfish Queen Angelfish Squirrelfish Hamlets Common halfbeak Dwarf herring Spotted Trunkfish Trunkfish Mutton Snapper Blackfin Snapper Gray Snapper Dog Snapper Mahogany Snapper Lane Snapper Rosy Blenny Diversity 2017, 9, 29 10 of 16 Diversity 2017, 9, 29 10 of 16 Table 2. Cont. Malacoctenus macropus Scientific Name Malacoctenus triangulatus Microphis brachyurus lineatus Malacoctenus triangulatus Monacanthus Microphis tuckeri brachyurus lineatus Mulloidichthys martinicus tuckeri Monacanthus Nicholsina usta Mulloidichthys martinicus Ocyurus chrysurus Nicholsina usta Pomacanthus Ocyurus arcuatus chrysurus arcuatus Pomacanthus Pomacanthus paru PseudupeneusPomacanthus maculatus paru maculatus SargocentrumPseudupeneus coruscum Scarus iseri Sargocentrum coruscum Scarus iseri Scarus taeniopterus Scarus taeniopterus Sparisoma atomarium Sparisoma atomarium Sparisoma chrysopterum Sparisoma chrysopterum Sparisoma radians Sparisoma radians Sparisoma viride Sparisoma viride Sphoeroides spengleri Sphoeroides spengleri Sphoeroides testudineus Sphoeroides testudineus Sphyraena barracuda Sphyraena barracuda Stegastes planifrons Stegastes planifrons Stegastes adustus Stegastes adustus Stegastes leucostictus Stegastes leucostictus Stegastes partitus Stegastes partitus Stegastes variabilis Stegastes variabilis Synodus intermedius Synodus intermedius Thalassoma bifasciatum Thalassoma bifasciatum Trachinotus spp. Trachinotus spp. Rosy Blenny Common Saddled Blenny Name Pipefish Saddled Blenny Slender Filefish Pipefish Yellow Goatfish Slender Filefish Emerald Parrotfish Yellow Goatfish Yellowtail Snapper Emerald Parrotfish Gray Angelfish Yellowtail Snapper Gray Angelfish French Angelfish French Angelfish Spotted Goatfish Spotted Goatfish Reef Squirrelfish Reef Squirrelfish Striped Parrotfish Striped Parrotfish Princess Parrotfish Princess Parrotfish Greenblotch Parrotfish Greenblotch Parrotfish Redtail Parrotfish Redtail Parrotfish Bucktooth Parrotfish Bucktooth StoplightParrotfish Parrotfish Stoplight Parrotfish Bandtail Puffer Bandtail Puffer Checkered Puffer Checkered Puffer Great Barracuda Great Barracuda Threespot Damselfish Threespot Damselfish Dusky Damselfish Dusky Damselfish Beaugregory Beaugregory Bicolor Damselfish Bicolor Damselfish Cocoa Damselfish Cocoa Damselfish Sand Diver Sand Diver Bluehead Bluehead Pompano Pompano (a) (b) (c) (d) Figure 7. Cont. Diversity 2017, 9, 29 11 of 16 Diversity 2017, 9, 29 11 of 16 (e) (f) (g) Figure 7. Fishes observed in Hurricane Hole’s mangrove-lined bays. (a) Gray Snappers (Lutjanus Figure 7. Fishes observed in Hurricane Hole’s mangrove-lined bays. (a) Gray Snappers (Lutjanus griseus); (b) Queen Angelfish (Ho