’s mesophotic coral reefs and associated communities.

Item Journal Contribution

Authors Reed, John K.; González-Díaz, Patricia; Busutil, Linnet; Farrington, Stephanie; Martínez- Daranas, Beatriz; Cobián Rojas, Dorka; Voss, Joshua; Diaz, Cristina; David, Andrew; Hanisak, M. Dennis; González Mendez, Juliett; García Rodríguez, Alain; González- Sánchez, Patricia M.; Viamontes Fernández, Jorge; Estrada Pérez, Daniel; Studivan, Michael; Drummond, Felicia; Jiang, Mingshun; A. Pomponi, Shirley

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Link to Item http://hdl.handle.net/1834/14977 revista de investigaciones marinas REVISTA DE INVESTIGACIONES MARINAS Centro de Investigaciones Marinas Universidad de La Habana http://www.cim.uh.cu/rim/

ARTÍCULO ORIGINAL CUBA’S MESOPHOTIC CORAL REEFS AND ASSOCIATED FISH COMMUNITIES Arrecifes de coral mesofóticos de Cuba y comunidades de peces asociadas

John K. Reed1, Patricia González-Díaz2, Linnet Busutil3, Stephanie Farrington1, Beatriz Martínez- Daranas2, Dorka Cobián Rojas4, Joshua Voss1, Cristina Diaz1, Andrew David5, M. Dennis Hanisak1, Juliett González Mendez6, Alain García Rodríguez3, Patricia M. González- Sánchez7, Jorge Viamontes Fernández8, Daniel Estrada Pérez8, Michael Studivan1, Felicia Drummond5, Mingshun Jiang1, Shirley A. Pomponi1

1 Cooperative Institute for Ocean Exploration, Research, and Technology, Harbor Branch Oceanographic Institute, Florida Atlantic University, ABSTRACT Fort Pierce, Florida, USA A joint Cuba-U.S. expedition was conducted May 14-June 12, 2017 2 Centro de Investigaciones to characterize for the first time the extent and health of mesophotic Marinas, Universidad de La coral ecosystems (MCEs) along the entire coastline of Cuba. Remote- Habana, Cuba ly Operated Vehicle (ROV) dives at 36 sites confirmed the presence of MCE habitat along all coasts of Cuba. ROV dives covered 27 km, 3 Instituto de Ciencias del Mar, La Habana, Cuba at depths of 25-188 m, and documented habitat and with 103 hours of high-definition video and 21,146 digital images. A total of 4 ECOVIDA-Sistema Nacional 477 taxa of benthic macrobiota and 178 fish taxa were identified, and de Areas Protegidas, Pinar 343 specimens of benthic and algae were collected to del Río Province, Cuba verify and assess population genetic structure. The pri- 5 NOAA, National Marine mary geomorphological features were the Deep Island Slope (125- Fisheries Service, Southeast >150 m), Deep Fore-Reef Escarpment (the ‘Wall’, 50-125 m), and Fisheries Science Center, Deep Fore-Reef Slope (30-50 m). Most vertical surfaces of the Wall Panama City, Florida, USA were covered with dense , algae, octocorals, and black corals. 6 Centro Nacional de Áreas Agaricia was the most abundant scleractinian on the Wall at Protegidas, La Habana, Cuba depths of 50-75 m, and was observed to 122 m. Of the 2,240 sclerac- tinian colonies that were counted in this study, only 12 corals (0.53%, 7 Acuario Nacional de Cuba, La Habana, Cuba mainly Agaricia spp.) showed signs of bleaching, and one Agaricia had black band disease, comprising remarkably low disease preva- 8 Geocuba Estudios Marinos, lence. The most frequently recorded genera were Xestospon- La Habana, Cuba gia, , and Agelas. At least 10 previously unknown sponge species were collected during the expedition. Sites outside of marine * Autor para correspondencia: protected areas generally had lower fish abundances, a possible in- [email protected] dicator of historical overfishing. Lionfish were observed at most sites but abundances were low compared to other Caribbean regions.

Recibido: 31.01.2017 Key words: Cuba, mesophotic reef, geomorphology, biozonation, biodi- Aceptado: 21.06.2018 versity.

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 61 Reed • González-Díaz • Busutil • et. al.

RESUMEN Entre el 14 de mayo y el 12 de junio de 2017 INTRODUCtioN se llevó a cabo una expedición conjunta entre Mesophotic coral reef ecosystems (MCEs) Cuba y EEUU. El objetivo fue mapear y carac- are light-dependent benthic communities terizar, por primera vez, la extensión y salud that occur deeper than shallow reefs and de los arrecifes de coral mesofóticos (MCE, typically range from depths of 30 m to the por sus siglas en inglés) alrededor de las cos- bottom of the photic zone, which may ex- tas de Cuba. 43 inmersiones con un Vehículo Operado por Remoto (ROV, por sus siglas en tend to >150 m in some regions (Lesser et inglés) en 36 sitios, confirmaron la presencia al., 2009; Hinderstein et al., 2010; Baker de arrecifes mesofóticos a lo largo de toda la lí- et al., 2016). MCEs represent in part an nea costera cubana. Las inmersiones del ROV extension of shallow-water coral reef eco- desde 25 a 188 m de profundidad abarcaron, systems and support a diverse assemblage en total, 27 km, y se documentaron hábitats y of habitat-building taxa, including corals, especies con 103 horas de videos de alta defi- nición y 21,146 imágenes digitales. Un total de sponges and algae, and associated commer- 477 taxa de macrobiota bentónica y 178 taxa cial and recreational fisheries. However, de peces fueron identificados. Se recolectaron many of the MCE species are depth spe- 343 especímenes de invertebrados bentónicos cialists not found on shallow reefs. Studies y algas para verificaciones taxonómicas y eva- of MCEs worldwide are expanding our un- luación de estructura de poblaciones. Los zo- derstanding of their distribution, health, nas geomorfológicas más prominentes fueron: Pendiente profunda (125- >150 m), Escarpe biodiversity, and ecology (Hinderstein et profundo del arrecife frontal (la “pared”, 50- al., 2010; Baker et al., 2016; Loya et al., 125 m), y arrecife de franja profundo (30-50 m). 2016, 2018; Roman, 2018). Many MCEs La pared presentó la mayor diversidad y den- worldwide appear to be thriving compared sidad de la macrobiota; todas las superficies to shallow reefs. The deep reef refugia hy- verticales estuvieron cubiertas por diversidad pothesis suggests that MCEs may be less de esponjas, algas, gorgonias y coral negro. Agaricia fue el género más abundante de escle- impacted from natural and anthropogenic ractíneos y dominó a profundidades entre 40- impacts than shallow coral reefs, and may 75 m. Solo 12 colonias de corales escleractíneos be more stable and resilient than shallow (principalmente Agaricia spp.) de 2,240 colo- reefs (Bongaerts et al., 2010; Bridge and nias (0,53%) registradas con el ROV mostraron Guinotte, 2012). MCEs may also act as re- signos de blanqueamiento, y una Agaricia pre- fugia for shallow reef species through the sentó enfermedad de banda negra, evidencian- do la muy baja prevalencia de enfermedades. export of fish and coral larvae (Serrano et Los géneros más frecuentes de Porifera fueron al., 2014; Vaz et al., 2016). However, it is Xestospongia, Aplysina, y Agelas. Al menos now apparent that MCEs are also vulnera- una decena de especies de esponjas desconoci- ble to disturbances from all facets of pertur- das para la ciencia fueron colectadas. Los si- bations including climate change, bottom tios que se encontraban fuera de áreas marinas trawling, invasive species, and pollution protegidas, generalmente poseían abundancia de peces baja, lo cual pudiera ser un posible (Bak et al., 2005; Andradi-Brown et al., indicador de sobrepesca histórica. Individuos 2016; Appledoorn et al., 2016a). Smith and de pez león fueron observados en la mayoría de Holstein (2016) further state that the main los sitios pero con abundancias bajas en com- premise of the deep reef refugia hypoth- paración con otras regiones del Caribe. eses is incorrect and that it now appears palabras claves: Cuba, arrecife mesofótico, that increases in temperatures above the geomorfología, biozonación, biodiversidad local mean warmest conditions can lead to

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 62 Reed • González-Díaz • Busutil • et. al.

thermal stress and bleaching. Also there is pioneers of research on Cuba’s mesophotic growing evidence that the hypothesis may reefs. Zlatarski and Estalella’s (1982) book only be applicable to depth generalists on “The Scleractinians of Cuba” (first pub- MCEs, and their contribution to replenish- lished in Russian in 1980, 1982 in French, ment of shallow reefs may not be as conse- 2008 in Spanish) provides detailed reef quential as once hoped (Schlesinger et al., profiles and coral species distributions 2018). from 44 sites around the island, some MCEs have been studied extensive- of which extend into the >40-70 m depth ly throughout the Western Atlantic re- range; however, the eastern tip and north- gions of the Caribbean, Bahamas, and the east coast lack data. Scuba dives (some to Gulf of Mexico (Hinderstein et al., 2010; depths of 70 m) for this study resulted in Baker et al., 2016; Loya et al., 2016, 2018). an extraordinary collection of corals (5,924 Whereas considerable data have been re- specimens) which are now archived in the ported regarding the distribution, ecol- National Aquarium of Cuba (in Havana). ogy, and health of Cuba’s shallow reefs González Ferrer’s (2004) book “Corales (Alcolado et al., 2001a; Alcolado et al., Petreos Jardines Sumergidos de Cuba” also 2001b; Alcolado et al., 2003; Aguilar et al., provides invaluable data on the stony cor- 2004; Aguilar Betancourt and González- als of Cuba as well as the history of Cuban Sansón, 2007 ; Alcolado and García, 2007; reef research since the 16th century. This Martínez-Daranas et al., 2008; Alcolado book provides beautiful color plates for all et al., 2009; González-Díaz et al., 2010; the hard corals found in Cuba, as well as Perera Valderrama et al., 2013; González- discussions of reef types, coral disease and Díaz et al., 2014; Alcolado et al., 2015; bleaching. Kühlmann (1974, 1983) also González-Díaz et al., 2015; Suárez et al., provided data on the depth distribution of 2015; Caballero Aragon et al., 2016; Ferrer scleractinian corals from scuba dives to 70 Rodríguez et al., 2016; González-Díaz et m in Cuba. al., 2018; Roman, 2018), relatively little In 2015, a Joint Statement was developed is known about the distribution, commu- between the United States and the Republic nity structure and health of Cuba’s deep of Cuba on Cooperation on Environmental mesophotic reefs. The book “The Ecology Protection (November 24, 2015), and a of the Marine of Cuba” (Claro et al., Memorandum of Understanding (MOU) 2001) provides a comprehensive descrip- was signed by the U.S. National Oceanic tion of physical attributes of coastal Cuba and Atmospheric Administration (NOAA), including geology, geomorphology, hydrol- the U.S. National Park Service and Cuba’s ogy, and major habitats from intertidal National Center for Protected Areas. This depths out to the shelf break at approxi- MOU establishes a ‘Sister-Sanctuary’ rela- mately 30 m. The fish fauna of what Claro tionship between Guanahacabibes National et al. (2001) termed as the slope and bathy- Park and Banco de San Antonio Marine al zone (100-600 m) was described from Sanctuary in Cuba, and the Florida Keys dives with Harbor Branch Oceanographic National Marine Sanctuary (FKNMS) Institute’s Johnson-Sea-Link submers- and Flower Garden Banks National Marine ible in 1997. In the 1970s, Vassil Zlatarski Sanctuary (FGBNMS) in the United States. and Nereida Martinez Estalella were the It is recognized that these regions are all

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 63 Reed • González-Díaz • Busutil • et. al. inextricably linked through ocean currents. Current Profiler) oceanographic data, and In support of these agreements, a joint re- water column data for benthic carbonate search expedition was conducted from May chemistry including aragonite saturation 14 to June 12, 2017 on the University of state. Data collected during this expedition Miami’s ship F.G. Walton Smith to sur- and on planned future cruises will be used vey the mesophotic reefs of Cuba. This to compare the health and connectivity of project was jointly planned and in collabo- Cuba’s MCEs to reef systems elsewhere in ration with organizations in Cuba and the the Caribbean, Gulf of Mexico, and south- U.S., including Centro de Investigaciones eastern United States. Marinas at University of Havana (CIM- This paper provides a preliminary over- UH), Centro Nacional de Áreas Protegidas view of the results of this expedition to date, (CNAP), Instituto de Ciencias del Mar including general descriptions of the ocean- (ICIMAR), Geocuba Estudios Marinos, ographic data around Cuba, MCE habitats, Guanahacabibes National Park-Sistema geomorphology, biozonation, and biodiver- Nacional de Areas Protegidas (PNG-SNAP), sity of these reefs. Future taxonomic and Acuar io Nacional de Cuba (A NC ); and for the genetic analyses of the specimens, along U.S., two NOAA Cooperative Institutes: the with quantitative analyses of the video and Cooperative Institute for Ocean Exploration, photographic data, will allow a more pre- Research, and Technology (CIOERT) at cise characterization of the diversity, zona- Harbor Branch Oceanographic Institute/ tion, health, and regional differences of the Florida Atlantic University (HBOI-FAU), mesophotic communities of Cuba, as well as and the Cooperative Institute for Marine a better understanding of the connectivity and Atmospheric Studies (CIMAS) at the of Cuban reefs with the Sister Sanctuaries University of Miami. in the U.S. and elsewhere in the Caribbean. The primary goal of this research ex- These data will also be useful for decision pedition was to survey the extent of meso- making for new and existing marine pro- photic reefs along the entire coastline of tected areas (MPAs) into these deeper wa- Cuba using a remotely operated vehicle ters of Cuba. Cuba’s strong marine policies (ROV). The objectives of the expedition and legislation has already resulted in 105 were to: 1) characterize Cuba’s MCE ben- MPAs, covering nearly 25% of its insular thic habitats, benthic macroinvertebrate shelf, yet overfishing, poaching, pollution and algal communities, and fish popula- and global warming are threats to these tions (emphasis on commercially impor- vulnerable ecosystems (Perera Valderrama tant /snapper species, spawning et al., 2018; Roman, 2018). grounds, and Lionfish); 2) understand the genetic connectivity of Cuba’s MCEs with Methods and Materials mesophotic and shallow reefs upstream Locations of proposed dive sites were com- and cross-stream (including the Sister- piled prior to the cruise, based on data from Sanctuaries in U.S. and throughout Cuba); various sources: 1) recommendations from 3) characterize the health of Cuba’s MCEs; CIM-UH, CNAP, and ICIMAR; 2) NOAA 4) conduct ROV collections of benthic spe- and regional coastal charts; and, 3) site da- cies for taxonomy and molecular analyses; ta from HBOI’s Johnson-Sea-Link manned 5) collect CTD, ADCP (Acoustic Doppler submersible 1997 expedition (Discovery

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Channel “Forbidden Depths” documen- was usually for sample collections and fish tary). Some sites were selected directly video transects. Photo transects were con- offshore of well-characterized shallow-wa- ducted using the digital still camera orient- ter reef sites or within Marine Protected ed perpendicular to the substrate (~1-2 m Areas (MPAs). Except for Banco de San off bottom), and images were haphazardly Antonio, there are no high-resolution taken 2-3 per minute. In addition, at most maps of the Cuba’s deepwater shelf and sites, at least one horizontal transect was slope. Specific sites were selected based on made along a constant depth contour wher- the above sources and plotted in ArcGIS ever Agaricia spp. and Montastraea caver- 10.3 using coastal charts at depths of 30- nosa corals first became common (usually 150 m. In order to circumnavigate the is- between 50 and 70 m depths). These hor- land in the time available, transit between izontal transects were typically along the sites at night was generally from 50-150 face of the vertical rock wall or overhang- km. Additional dives were made within ing rock buttresses, so the camera was the Guanahacabibes National Park which pointed straight forward, perpendicular is of special interest. Two 3-4 hour dives toward the wall. Time and distance of the were made each day with a Mohawk ROV. horizontal transects varied depending on At each site a fathometer profile was made currents and topography, but generally prior to the ROV dive to document the were about 15 minutes, and transited ~100 depth and location of the shelf-edge break m, for a total of 30 images. Fish were iden- and to select the exact target site for the tified throughout the dive using the video ROV. Sketches of reef profiles were drawn camera pointed forward and down ~20° to to illustrate the various dive sites. The spe- view from the horizon to close up. In addi- cific depths and slopes were drawn to scale tion, fish video transects of approximately from observations made during the ROV 30 minutes were made at each site along dives. The slope angles, inflections of the the upper wall and deep fore-reef crest. slopes at different depths, depths of zones, Direction of transects were haphazard, but and depths of shelf-edge reef crest were generally headed along slope, but also de- made from the ROV dive notes. pended on the ship's maneuverability with Video was recorded throughout each the wind and current. ROV dive with an Insite Pacific Mini Zeus The Mohawk ROV was equipped with high-definition CMOS color zoom cam- a collection skid, which consists of a small era. Digital still images were taken with a 5-function manipulator, five suction buck- Kongsberg Maritime OE14-408. Both cam- ets, and a Plexiglas box. Specimens of cor- eras had 10-cm parallel lasers for scale als, sponges, and algae were collected at (green- still; red- video). At each site a con- most sites and will be used for museum tinuous video/photo transect commenced specimens and taxonomic identification, at a maximum depth of ~150 m and contin- genetic analysis, and coral health stud- ued upslope to the upper mesophotic reef ies. Specimens were photographed in situ, zone or about 30 m. The photo transects and in the ship’s lab, and then preserved were used for habitat characterization in 95% ethanol, 5% formalin, or TRIzol. and species identifications. When a sec- ROV sensors included: Seabird Fastcat 49,

ond dive was made at the same dive site, it CO2 and pH. Shipboard oceanographic data

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 65 Reed • González-Díaz • Busutil • et. al.

Fig. 1. Map of Cuba showing ROV dive sites (red stars) on mesophotic reef sites (30-150 m depths) during the F.G. Walton Smith cruise, May 14 to June 12, 2017. Pink polygons- marine protected areas (UNEP-WCMC, 2017). continuously recorded surface water data separate Access databases which were lat- including temperature, salinity, florescence, er compiled with the benthic habitat data- and dissolved oxygen, and ADCP data. A base. Duplicate sets of the video/photo data shipboard CTD and water sample rosette and specimens are curated in the U.S. at were cast daily to profile the dive sites. CIOERT (HBOI-FAU), and in Cuba at Metadata and habitat dive notes were CIM-UH/ANC/ICIMAR/CNAP. recorded during the dive into a Microsoft Access database which links to the ROV Results navigation data. Teams of U.S. and Cuban Thirty-six mesophotic reef sites around scientists specializing in coral, sponge, al- the coast of Cuba were surveyed during gae, and fish taxonomy recorded data in 43 ROV dives (Fig. 1, Table 1). Total ship

Table 1. ROV dive sites on Cuba’s mesophotic reefs during the F.G. Walton Smith cruise, May 14 to June 12, 2017.

Dive Depth Station Location Latitude Longitude No. Range (m) C-04 1 Cuba, NW coast, Cayo Levisa MPA 22°53.483'N 83°34.969'W 25-188 C-04A 2,3 Cuba, NW coast, 2 nmi north of Cayo Arenas, Cayo Levisa MPA 22°52.638'N 83°39.103'W 49-172 C-06 4,5 Cuba, NW coast, Norte de Los Pretiles MPA 22°30.487'N 84°25.907'W 43-170 C-07 6,7 Cuba, NW coast, Archipiélago de Los Colorados, No MPA 22°13.405'N 84°44.837'W 45-178 C-10 8 Cuba, W coast, Banco de San Antonio MPA, east wall 22°00.901'N 84°59.947'W 25-183 C-10A 9 Cuba, W coast, Banco de San Antonio MPA, south wall 22°00.017'N 85°00.977'W 30-150 C-12 10 Cuba, W coast, Los Cayuelos, Guanahacabibes National Park 21°50.005'N 84°57.426'W 149 Cuba, W coast, Faro Roncali, off lighthouse, Guanahacabibes C-12A 11 21°52.111'N 84°57.824'W 45-156 National Park C-12B 12 Cuba, W coast, Las Tumbas, Guanahacabibes National Park 21°55.477'N 84°57.494'W 38-148 Cuba, W coast, El Almirante, Punto de Buceo, Cabo Corrientes, C-15 13 21°48.224'N 84°31.137'W 25-145 Guanahacabibes National Park

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C-16A 14 Cuba, W coast, Cabo Corrientes, Guanahacabibes National Park 21°45.512'N 84°31.874'W 28-150 Cuba, S coast, SW tip of Isla de la Juventud, Punta de C-21 15 21°35.263'N 83°10.325'W 25-158 Pedernales MPA Cuba, S coast, SW Isla de la Juventud, bay on west side of point, C-21A 16 21°36.010'N 83°10.943'W 28-150 Punta Francés MPA C-23 17 Cuba, S coast, SE Isla de la Juventud, Punta Brava MPA 21°29.419'N 82°39.228'W 37-150 Cuba, S coast, E of Isla de la Juventud, Archipiélago de Los C-24 18 21°31.924'N 82°27.480'W 40-150 Canarreos, Cayo Matías MPA Cuba, S coast, Archipiélago de Los Canarreos, Cayo Rosario C-28 19 21°33.469'N 81°44.401'W 29-150 MPA Cuba, S coast, Archipiélago de Los Canarreos, Cayo Largo del C-29 20 21°34.854'N 81°33.130'W 26-157 Sur MPA Cuba, S coast, Golfo de Cazones, Punta Brava, Sur Ciénaga de C-31A 21 21°57.650'N 81°10.355'W 18-150 Zapata, no MPA Cuba, S coast, Bahía de Cochinos, east wall, Zapata National C-33A 22 22°07.444'N 81°07.518'W 30-150 Park Cuba, S coast, south of Cienfuegos, Banco Silvertown, west wall, C-38 23,24 21°25.180'N 79°56.377'W 24-150 proposed MPA Cuba, S coast, south of Cienfuegos, Banco Silvertown, south C-38A 25 21°24.470'N 79°54.942'W 40-153 wall, proposed MPA Cuba, S coast, Cayo Caballones, south slope, Jardines de La C-41 26 20°47.495'N 78°57.323'W 30-150 Reina MPA Cuba, S coast, Cabo Cruz, Desembarco del Granma National C-48 27,28 19°49.600'N 77°44.551'W 40-154 Park C-50 29,30 Cuba, S coast, Sierra Maestra, Chivirico Proposed MPA 19°56.551'N 76°23.937'W 27-155 C-52 31 Cuba, S coast, Sierra Purial, Punta Imías, no MPA 20°02.601'N 74°41.498'W 60-161 Cuba, S coast, Sierra Purial, east tip of Punta Imías, 1 nmi east C-52A 32 20°02.513'N 74°40.345'W 43-156 of ROV 31, no MPA Cuba, eastern tip of Cuba, NW Punta Maisí, Punta Silencio, no C-53A 33 20°19.007'N 74°16.001'W 54-155 MPA Cuba, eastern tip of Cuba, NW Punta Maisí, Punta del Fraile, no C-53B 34 20°19.482'N 74°13.940'W 50-142 MPA C-54 35 Cuba, NE coast, Cabo Lucrecia, no MPA 21°05.508'N 75°39.821'W 45-151 C-54B 36 Cuba, NE coast, Cabo Lucrecia, proposed MPA 21°04.817'N 75°38.530'W 65-156 Cuba, NE coast, Archipiélago Sabana-Camagüey, Cayo Sabinal C-56 37,38 21°41.096'N 77°10.241'W 24-150 MPA Cuba, NE coast, Archipiélago Sabana-Camagüey, Cayo Coco, C-58 39 22°34.192'N 78°22.361'W 42-140 proposed MPA Cuba, NE coast, Archipiélago Sabana-Camagüey, Cayo Coco, C-58A 40 22°35.262'N 78°27.706'W 46-144 proposed MPA (just north of NW edge of MPA) Cuba, NE coast, Archipiélago Sabana-Camagüey, Cayo La Vela C-59A 41 22°54.858'N 79°41.854'W 22-143 MPA Cuba, NE coast, Archipiélago Sabana-Camagüey, Cayo Jutía, ¼ C-59C 42 22°59.446'N 79°48.188'W 30-147 nmi N of snorkel Station, no MPA Cuba, NE coast, Archipiélago Sabana-Camagüey, Cayo Cruz del C-60A 43 23°17.792'N 80°53.809'W 40-161 Padre, inside and outside MPA

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 67 Reed • González-Díaz • Busutil • et. al. transit around the island covered ~2,778 the eastern end of Cuba. Surface currents km. Dives ranged from depths of 188 m along the northeastern Cuban coast, how- to 25 m and resulted in a total dive time ever, did not have a prevailing direction, of 103 hours, covering approximately 27 and were likely influenced by the compe- km. A total of 21,146 digital still images tition between the Florida Current and documented habitat and species, and pho- Antilles Current and by tidal fluctuations. to transect images (7,063) will be used for Historical measurements by Atkinson et al. future analyses of percent cover and densi- (1995) suggested that the residual currents ty of corals, sponges, and algae. The high- in the Old Bahamas Channel were pre- definition video will be used to document dominantly westward toward the Florida species composition and density of fish. A Straits. Temperatures recorded by the total of 343 specimens of benthic macro-in- ROV showed surface temperatures rang- vertebrates and macro-algae were collec- ing from 27.09-29.52oC for all dives (Table ted with the ROV. 2). In general, temperatures were greatest along the south coast (29.52oC). The coolest Oceanographic Data temperatures were found along the north- The surface currents along the west- east coast (27.09oC). At mid-depth of the ern half of Cuba, between Havana and dives (50 m, in the deep mesophotic zone), Cienfuegos, were likely part of the Cuban temperatures ranged from 25.26-28.62oC. Countercurrent, which flows counterclock- Minimum temperatures at the deeper wise against the Caribbean and Florida depths (162-182 m) were 21.35- 25.93oC. Currents (Fig. 2). Along the southeast- A thermocline of 25-28oC was common ern coast from Archipelago Jardines de la around 80-100 m depths. Surface salinities Reina to the eastern end, which abuts the ranged from 35.11-36.73 PSU and at 50 m northern steep slope of the Cayman Trench, depths were 35.14-36.77 PSU. Visibility currents were generally eastward. Surface based on ROV video was excellent at most currents were counterclockwise around sites (estimated 30- >50 m), except sites

Fig. 2. Shelf-edge currents around Cuba from 6-hourly mean surface currents (arrows) measured with shipboard ADCP during the F.G. Walton Smith cruise, May 14 to June 12, 2017.

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Table 2. Oceanographic data collected with a Seabird Fastcat 49 on the Mohawk ROV during the F.G. Walton Smith cruise, May 14 to June 12, 2017. Maximum Depth Mid Depth (50 m) Surface (1 m) Depth (m) Temp.(oC) Salinity (PPT) Temp. (oC) Salinity (PPT) Temp. (oC) Salinity (PPT) Max Range Range Range Range Range Range Northwest Coast 177.27 22.47-24.19 36.84-36.89 26.77-27.44 36.31-36.40 27.71-28.27 36.16-36.37 West Coast 182.13 22.53-25.93 36.65-36.90 27.50-28.25 36.29-36.37 28.17-28.58 36.28-36.31 Southwest Coast 162.29 22.69-24.15 35.59-36.89 27.68-28.46 35.14-36.41 28.53-29.52 35.11-36.41 Southeast Coast 162.06 22.53-24.41 36.80-36.93 25.99-28.62 36.11-36.67 28.42-29.48 36.04-36.35 Northeast Coast 162.01 21.35-25.56 36.72-37.41 25.26-27.88 36.39-36.77 27.09-28.54 35.64-36.73 Total 182.13 21.35-25.93 35.59-37.41 25.26-28.62 35.14-36.77 27.09-29.52 35.11-36.73

near stream runoff. Surface currents at the deep reef features are likely carbonate the dives sites varied from 10 to 51 cm s-1 limestone, and none of the sites appeared to (0.2-1 kn), but occasionally were in excess be granitic, even at the southeastern coast of 100 cm s-1 during which the ROV could off the Sierra Maestra mountain range not be safely deployed and recovered. where we expected granitic rock slopes as the coastline drops from +1,974 m (moun- Geomorphology tain peak) to -7,686 m (Cayman Trough). The ROV dives documented the deep meso- The following is a general description of photic habitat surrounding Cuba. Many of the deep fore-reef and slope (Figs. 3, 4). the dive sites, especially on the eastern end Deep Island Slope (125- >150 m)- At of Cuba, had never been explored previously. many sites, this zone was a steep (70o), rel- All 43 dives confirmed the presence of MCE atively smooth rock pavement slope inter- habitat. It appears from these dives that, sected with low relief, vertical sediment just like the shallow reefs that fringe most chutes (1-m deep, 1-3 m wide), although of the Cuban coast, the deep reefs also par- some sites had slopes varying between 45o allel most of the shelf edge for nearly 2,800 and 90o at 150 m. The surface rock usually km, and include deep fringing reefs that are showed horizontal layering and had scal- close to shore (primarily along the northwest lop-shaped erosional features (10-30 cm coast, southeast coast, and northeast coast), diameter). This zone had the lowest diver- the barrier reefs along the various archipel- sity and density of biota. Fathometer pro- agos (Los Colorados, Canarreos, Jardines files showed slopes of 10-45o extending to de la Reina, and Sabana-Camagüey), and depths of 500 m. deep-water banks (Banco de San Antonio, Deep Fore-Reef Escarpment (the ‘Wall’, and Banco Silvertown) (for terminology 50-125 m)- This zone formed a near-verti- of reef types, see page 390, Zlatarski and cal wall (80-90o slope) at most sites. The Estalella, 1982; González Ferrer, 2004). base of the wall was commonly between Topographically, the most conspicuous fea- 100 and 125 m depths although at many tures of most dive sites were the deep island sites the wall continued to >150 m (>200 slope (125- >150 m), deep fore-reef escarp- m at Site C-29 off the NE coast). At some ment (vertical wall; 50-125 m), and deep sites, the upper wall (45-80 m) was formed fore-reef (30-50 m) (Fig. 3). The geology of of large rock “buttresses” which overhung

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Fig. 3. Schematic of general geomorphological features and biozonation of Cuba’s mesophotic reefs showing a well- developed fore reef zone and wall with buttresses. the plane of the wall by 3-5 m. The but- Deep Fore-Reef Crest and Slope (30-50 m)- tresses were generally 10-20 m wide and Seaward of the shallower reef zones (3-25 m), were separated by sediment chutes which a zone of shelf-edge reefs generally paral- provided a conduit for the downslope leled the shelf-edge break. At some sites transport of the shelf sediments (Hoskin the deep fore-reef crest formed a ridge or et al., 1986). Fewer macrobiota occurred sill-like feature (3-5 m relief). Generally, in the chutes. Between depths of 60 and this ridge was not continuous but a series 125 m, the wall was highly eroded at ma- of mounds (5-10 m diameter, 3-5 m high). ny sites, with karst-like topography form- The fore-reef slope of this zone was typical- ing caves (1-3 m diameter) and ledges (1-2 ly steep (45-60o) with rocky escarpments, m wide). and spurs and grooves. The fore-reef slope

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Fig. 4. Habitat and reef biota of Cuba’s mesophotic reefs. Deep fore-reef slope: A) 1-m diameter plates of Orbicella faveolata, conical dead Montastraea cavernosa, tube sponges (), and rope sponges (Agelas sceptrum, Aplysina cauliformis), Punta de Pedernales MPA, Site C-21, 33 m; B) dense shingle-like corals O. faveolata, Agaricia spp., Montastraea cavernosa, Bahía de Cochinos, Site C-33A, 30 m. Deep fore-reef escarpments (the ‘wall’): C) rock buttress with dense array of sponges (yellow sponge- Siphonodictyon coralliphagum), black corals, branching octo- corals, and stylaster corals, Guanahacabibes National Park, Site C-12B, 82 m (lasers 10 cm); D) plates of Agaricia spp., Guanahacabibes National Park, Site C-12B, 64 m (lasers 10 cm); E) bushes sp. on vertical wall, Guanahacabibes National Park, Site C-12B, 79 m; F) large barrel sponge Xestospongia sp., green alga Halimeda sp., and scleractinian Agaricia sp., Cayo Largo del Sur MPA, Site C-29, 61 m.

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(e.g., along the SE coast off Sierra Maestra, Deep Island Slope (125- >150 m)- Bahia de Cochinos, and Guanahacabibes). Although the deep island slope often had Elsewhere the shelf-edge reefs occurred photosynthetic species such as crustose seaward of the Archipelagos (e.g., Los coralline red algae (CCA) and encrusting Colorados, Canarreos, Jardines de la green algae, there were no zooxanthel- Reina, Sabana-Camagüey) and may be late corals. Ahermatypic corals such as considered barrier reefs (Zlatarski and Madracis myriaster were common along Estalella, 1982; González Ferrer, 2004). with various azooxanthellate, branch- Also the two deep-water banks that were ing octocorals (Ellisella elongata, Nicella surveyed (San Antonio and Silvertown) goreaui, Swiftia exserta), black cor- had similar topography along the shelf- als (Antipathes sp., Tanacetipathes sp., edge break. Stichopathes sp.), and stylasterine cor- als. Sponges were common but not as di- Biozonation verse or dense as on the wall. Common Mesophotic coral ecosystems (MCEs) are sponge species include a bright yellow characterized by light-dependent corals and crustose sponge (Order ) form- associated communities typically found at ing abundant large patches (20-100 cm2), depths ranging from 30-40 m and extend- thick-walled vase sponge (Xestospongia sp. ing to >150 m in tropical and subtropical Cu-01), several species of Oceanapia with regions (Baker et al., 2016). Various re- elongated projections, and massive sponges searchers have divided what is now called from various taxa (Orders Haplosclerida, the mesophotic zone into different subzones Suberitida, and Tetractinellida). and depths (e.g., Goreau and Goreau, 1973; Deep Fore-Reef Escarpment (the ‘Wall’, Lang, 1974; James and Ginsburg, 1979; 50-125 m)- This is the most impressive Ginsburg et al., 1991; Lesser et al., 2009; and recurring feature of most sites along Locker et al., 2010; Sherman et al., 2010; the Cuban shelf edge. This zone had the Clark et al., 2014; Reed, 2016; Laverick greatest diversity and density of macrobio- et al., 2017). For this paper, we use the ta of the deep reefs. Starting at 100-125 m, depth of 30 m as the upper extent of the nearly all vertical surfaces were covered upper mesophotic zone and have used the with sponges (e.g., Agelas spp., Aplysina depth of 50 m as the transition between spp., Callyspongia spp., Ceratoporella the upper and lower mesophotic zones nicholsoni, Clathria spp., Geodia spp., where we generally saw a fairly distinct Leucetta spp., Oceanapia spp., Petrosia change in geomorphological zonation and spp., Plakortis spp., Spirastrella spp. and species. Therefore, the upper mesophot- Xestospongia spp.); algae (CCA, Halimeda ic zone is generally in the deep fore-reef, spp., Peyssonneliaceae (several species), and the lower mesophotic zone is basical- Lobophora spp., and Dictyota spp.), branch- ly on the wall and buttresses of the deep ing octocorals (Nicella spp., Ellisella spp., fore-reef escarpment. The definitive maxi- Swiftia exserta), and black corals (bushy mum depth of the mesophotic zone is prob- Antipatharia to 1-2 m, and dense fields lematic and varies from site to site based of Stichopathes sp.). In general, the domi- on light, sedimentation, and substrate nant scleractinian corals on the wall were (Baker et al., 2016). Agaricia (A. fragilis, A. grahamae, A.

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lamarcki, and possibly A. undata). These Biodiversity typically first occurred at the maximum A total of 477 taxa of benthic macrobiota and depths of 80-122 m, and were often quite 178 fish taxa were identified from the ROV common to abundant between 50 and video/photo surveys and from the specimens 70 m and on the upper slope of the but- collected (Appendices 1 and 2, respective- tresses. Therefore, the bottom of the meso- ly). These data will be quantified in the fu- photic zone generally was around 80-120 m, ture for percent cover and for documenting but this was very variable by site. Corals densities of corals and fish. A total of 343 were less common where there was less mesophotic benthic samples were collected light, e.g., under the overhanging buttress- during this expedition; preliminary analy- es, on north facing walls, or where the deep ses indicate that some are new species and fore-reef did not form a sill that blocks the records of depth or distribution. flow of sediments downslope. For example, Corals- A total of the 101 cnidarian one reef at Faro Roncali, Guanahacabibes taxa that were identified morphological- National Park (Site C-12A) had no ridge- ly included 46 Scleractinia, 32 like deep fore-reef to block sediment flow (branching octocorals and Alcyoniina), from the shelf, and as a result no sclerac- and 15 Antipatharia. The deepest occur- tinian corals or CCA were observed on the rences of identified zooxanthellate corals wall. But, Site C-12B nearby had a dis- were Agaricia sp. (122 m), Stephanocoenia tinct shelf-edge reef with 5-m relief which intersepta (104 m), and M. cavernosa blocked sediment flow except in the area of (101 m) (Table 3). These were the most the chutes, and also had dense populations common scleractinians on the wall in the of plate coral on the wall and upper slope. deeper mesophotic zone, whereas O. fave- Deep Fore-Reef (30-50 m)- This region olata was more common in the upper meso- forms the upper mesophotic zone where photic zone. The most common octocorals higher abundance and diversity of scler- were in the genera Ellisella, Nicella, and actinian species were characteristically (syn. ), found. Agaricia spp., Montastraea caver- and Swiftia exserta. The most common nosa, Orbicella faveolata, Stephanocoenia black corals included Stichopathes spp., intersepta and Porites astreoides were dom- Tanacetipathes tanacetum, Tanacetiphates inant. Several sites, including Banco de sp., and Antipathidae. In addition, 123 San Antonio (Site C-10A), Guanahacabibes scleractinian corals were collected, some of (C-12B), Bahía de Cochinos (C-33A), and which will be used for genetic analysis. Cayo Sabinal (C-54B), had dense shingles Sponges- Cuban MCEs proved to be a of plate corals (O. faveolata or Agaricia spp.) very favorable habitat for marine sponges on the fore reef slope and crest (estimat- in both species richness and abundance. ed 60-80% cover at Site C-33A). Common We identified 296 distinct taxa: 280 shallow reef sponges were abundant here Demospongiae, 9 Homoscleromorpha, and as well, such as Agelas spp, Aiolochoria 7 Calcarea, of which 115 (39%) were iden- spp., Aplysina spp., Amphimedon spp., tified to species and 107 to genus (35%). Cinachyrella spp., Cliona spp., spp., The rest were only tentatively identified Niphates spp., Smenospongia spp., Spongia to families, orders, or classes, and are the spp., spp., and Xestospongia spp. subject of ongoing taxonomic research.

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Table 3. Deepest observations of macrobiota during ROV dives at all Sites during the F.G. Walton Smith cruise, May rock (183 m deep), Xestospongia sp. Cu-01 14 to June 12, 2017. (168 m), Ceratoporella nicholsoni (159 m), X. muta (134 m), and Leucetta sp. (124 m) Max. Depth Taxa (Table 3). Peak diversity started at depths (m) of 100-125 m in which sponge morpholo- Scleractinia gies (crusts, plates, tubes, spherical, vas- Agaricia sp. 122 es, and branches) and multicolor patterns Stephanocoenia intersepta 104 abounded. From the collections, there are Montastraea cavernosa 101 at least 10 sponge species that are new to Helioseris cucullata 79 science within the genera of Aplysina and Agaricia grahamae 74 Verongula ( Verongiida); Amphimedon, Orbicella faveolata 72 Callyspongia, and Xestospongia Octocorallia (Haplosclerida); Clathria (Poecilosclerida); Swiftia exserta 188 Cinachyrella (Tetractinellida); Aaptos Porifera (Suberitida); Erylus (Tetractinellida) and Verongiida Cu-01 (yellow crust) 183 Plakortis (Homosclerophorida). Xestospongia sp. Cu-01 168 Algae- A total of 63 taxa of macroalgae (29 Ceratoporella nicholsoni (sclerosponge) 159 Chlorophyta, 25 Rhodophyta, 9 Ochrophyta) Xestospongia muta 134 and 1 Cyanobacteria (syn., Cyanophyta) Leucetta sp. (Calcarea) 124 have been identified to date. The most fre- Chlorophyta quent taxa were crustose coralline algae Chlorophyta (thin green crust) 169 (CCA), Halimeda copiosa, Lobophora spp., Halimeda sp. 127 and Dictyota spp.. In general, the diversity Rhodophyta and coverage of algae were very low on the deep island slope where only CCA and thin Crustose coralline (CCA) 169 encrusting green algae occurred. Between Peyssonneliaceae 139 50 and 100 m, the algal diversity and cover Ochrophyta increased, with H. copiosa and CCA domi- Lobophora sp. 139 nating. In the upper mesophotic zone (30-50 Dictyota sp. 100 m), the species richness and cover increased Chordata even more, with Lobophora spp. and other Pterois volitans/miles (Lionfish) 188 Dictyotales dominating. Some interesting records of maximum depth of algal occur- The ten most frequently recorded species ance include an unidentified green crust were the barrel sponge Xestospongia muta, (169 m), CCA (169 m), Peyssonneliaceae (139 Xestospongia sp. Cu-01, an encrusting un- m), Lobophora spp. (139 m), and Halimeda identified bright yellow Verongiida sp. Cu- spp. (127 m) (Table 3). Cyanobacteria were 01, large tubular Aplysina archeri, several observed at eight sites and abundant at species of Agelas (A. sceptrum, A. dilata- one described below. Detrital leaves of the ta, and A. citrina), and Mycale laxissima. marine angiosperm Thalassia testudinum Sponges colonized all depths and substrates were also observed on the deep island slope sur veyed. T he deepest records were a bright at some sites showing the vertical connec- yellow Verongiida sp. Cu-01 encrusting on tivity between coastal-marine ecosystems.

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Fish- A total of 178 species of fish a no-take during the spawning season(s) were observed throughout the survey since other commercially important spe- (Appendix 2). Between 60 and 150 m, fish cies might also there. We observed diversity and abundances were low at all Lionfish (Pterois volitans/miles) at most sites, whereas from 30-60 m the diver- sites; the greatest depth was 188 m at sity increased, as well as the frequency Cabo Lucrecia (Site C-54B). The abun- and abundance of fish species. The deep dance of the invasive Lionfish was gener- island slope zone (125-150 m) was dom- ally low, generally <10 observed during a inated by Blackcap Basslet (Gramma four hour dive at most sites. An exception melacara), Cave Basslet ( was at Cayo Coco (Site C-58) where they mowbrayi), Sunshinefish ( insola- were relatively abundant with 38 counted ta), Blackfin Snapper (Lutjanus buccanel- on one dive. However, sampling by ROV la), Squirrelfish (Holocentrus adscensionis) video cameras could bias fish counts due and (H. rufus). The to avoidance and therefore abundances of walls (lower mesophotic reef zone, 50-120 Lionfish could be greater than we observed. m) and upper deep fore-reef (30-50 m) were inhabited by species common on Cuban Regional Comparisons of Cuba’s MCEs shallow reefs, including numerous species Sketches of reef profiles were drawn to il- from the families Acanthuridae, Balistidae, lustrate various regions of Cuba (Figs. 5-8). Carangidae, Chaetodontidae, Haemulidae, These are not the specific coastal zones , Labridae, , outlined in Areces (2002), but represent , , and the reef profiles of six sites within each . Groups of jack (Caranx latus, C. of the following general regions of Cuba: ruber and C. lugubris) were frequently ob- western, southern, eastern, and northern. served. Ten species of grouper (Serranidae) Photographs taken with the ROV that rep- were identified throughout the sampling resent each of these regions are provided area including Coney (Cephalopholis ful- in Appendix 3. Four general habitats were va), Graysby (Cephalopholis cruentata), added to the sketches: rock pavement, mud/ Nassau Grouper (Epinephelus striatus) and sediment, caves/ledges, and coral shingles. Red Hind (E. guttatus). Hundreds of Ocean The zone of eroded rock forming caves or re- (Canthidermis sufflamen) were entrants and ledges was found on the wall observed at Banco de San Antonio MPA. of the deep fore-reef escarpment at many of Courtship and nest building behavior by the sites. These features are karst-like to- the triggerfish were observed leading to the pography likely formed during the low sea conclusion that this is a spawning aggrega- levels of the last glacial period. The coral tion site. A spawning aggregation of Dog shingles zone consisted mostly of O. fave- Snapper (Lutjanus jocu) was also observed olata and Agaricia spp. and was found at at a site in Archipelago de Camagüey be- some sites on the deep fore-reef crest and tween 60 and 70 m depths. This site is slope (generally 30-40 m). Each profile also part of the Centro y Oeste de Cayo Coco includes the maximum depth observed at Ecological Reserve (Perera-Valderrama et that site for the primary plate corals: M. al., 2018). We propose this site should be cavernosa, Agaricia spp., O. faveolata, and made a no-take fishery reserve or, at least, S. intersepta (abbr. in Figs. 5-8).

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Fig. 5. Sketches of reef profiles selected to illustrate the various regions of Cuba’s MCEs. (see Fig. 1). Western region (Archipelago de Los Colorados, Banco San Antonio, Guanahacabibes): A. Site C-04, B. Site C-07, C. Site C-10A, D. Site C-12B, E. Site C-15, F. Site C-16A. The specific depths and slopes were drawn to scale from observations made during the ROV dives.

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Fig. 6. Sketches of reef profiles selected to illustrate the various regions of Cuba’s MCEs (see Fig. 1). Southern region (Isla de la Juventud, Archipelago de los Canarreos, Bahia de Cochinos, Banco Silvertown, Jardines de La Reina): A. Site C-21, B. Site C-23, C. Site C-28, D. Site C-33A, E. Site C-38, F. Site C-41. The specific depths and slopes were drawn to scale from observations made during the ROV dives

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In the western region, the crest of the deep as 50-60 m at some sites, and several shelf-edge reefs was generally between sites near the eastern tip of Cuba showed 30 and 40 m (Fig. 5). This fore-reef zone extensive erosion and lacked the vertical generally consisted of a series of mounds wall (Punta Imias, C-53; Punta Maisi, (3-10 m relief) along the shelf-edge break C-53A; Cabo Lucrecia, C-54B). Sherman that were intersected with sand chutes. et al. (2010) also found lower topograph- Some had buttresses on the deep fore-reef ic relief and less pronounced escarpment slope and upper wall. Shingles of O. fave- on the southeastern region of Puerto Rico. olata were found at Sites C-10A (Banco They attribute this due to the fact that the San Antonio) and C-12B (Las Tumbas, eastern facing slopes are more exposed to Guanahacabibes). The deep fore-reef es- trade-wind generated swells and tropical carpment generally was highly eroded hurricanes that typically approach from with caves or reentrants and ledges at the southeast. They also found that MCEs depths of about 60 to 110-130 m. Agaricia are patchier and more widely spaced on the spp. generally was first observed around southeast slopes. 70 m depth, but was as deep as 100 m at The northern region of Archipelago de Banco San Antonio (Site C-10A). In addi- Sabana-Camagüey (Fig. 8) also showed tion, these shelf-edge reefs were all quite variability ranging from eroded, low slope similar even though they comprise fring- geomorphology (Sites C-58, C-58A) to the ing reefs (Guanahacabibes), barrier reefs walls and buttresses (Sites C-56, C-59C, (Archipelago de Los Colorados), and bank and C-60A) that were typically present at reef (San Antonio). the western region. One site at Cayo Coco The reefs of the southern region gen- (C-58A) was mostly smooth rock pavement, erally showed less pronounced mounds with a relatively narrow region of highly or ridges along the shelf-edge break (Fig. eroded, rock ledge habitat at 70-90 m. Also 6), and the shelf-edge break varied great- the maximum depth of Agaricia tended to ly among sites. For example, the fore-reef be shallower at several sites in this region slope at SW Isla de Juventud (C-21) and perhaps due to the northern facing walls Bahia de Cochinos (C-33A) continued up (45 m at C-58, 52 m at C-59C). to at least 20 m depth where we had to cut To date, only general comparisons of off the transect for the safety of the ship. benthic macrobiota and fish distribu- Site C-33A had the densest cover of O. fa- tions can be made for the various meso- veolata coral shingles at any site, in depths photic regions of Cuba from these data. of 20-30 m. Cayo Rosario (C-28) also had Quantitative analyses of the photo tran- dense O. faveolata shingles on the deep sects for macrobenthos cover and density fore-reef slope but these were mostly dead. as well as analyses of fish densities from Agaricia occurred in this region as deep as the video transects are planned as part 107 m (Banco Silvertown, C-38). Also the of our multinational research efforts and zone of eroded topography varied greatly will allow for more detailed comparisons among the sites. of any regional and depth differences in Shelf-edge reefs along the eastern region the future. Appendices 1 and 2 provide were highly variable (Fig. 7). The shelf- the species lists of macrobenthos (inverte- edge break of the deep fore-reef was as brates and algae) and fish, respectively, by

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Fig. 7. Sketches of reef profiles selected to illustrate the various regions of Cuba’s MCEs (see Fig. 1). Eastern region (Granma National Park, Chivirico MPA, Punta Imias, Punta Maisi, Cabo Lucrecia): A. Site C-48, B. Site C-50, C. Site C-52, D. Site C-53A, E. Site S-54, F. Site C-54B. The specific depths and slopes were drawn to scale from observations made during the ROV dives.

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Fig. 8. Sketches of reef profiles selected to illustrate the various regions of Cuba’s MCEs (see Fig. 1). Northern region (Archipelago de Sabana-Camagüey): A. Site C-56, B. Site C-58, C. Site C-58A, D. Site C-59A, E. Site C-59C, F. Site C-60A. The specific depths and slopes were drawn to scale from observations made during the ROV dives.

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site and region based on the ROV annota- Corals- Cnidarian species richness of tions and sample collections. In general, the mesophotic reefs based on the ROV an- there were some regional and site specific notations was greatest on Cuba’s south- differences, but overall the MCE commu- east coast, where a total of 146 taxa nities and species distributions were rela- were observed. The sites with the great- tively similar around the island. For these est individual coral species richness (in- comparisons we have divided the dive sites cluding Scleractinia, Octocorallia, and into 6 regions (see Fig. 1): Northwest coast- Antipatharia) were located on the south- Archipelago de Los Colorados, Sites C-04 east coast; 47 and 46 coral taxa having to C-07; West coast- Banco de San Antonio, been recorded at Sites C-48 and C-50, re- Guanahacabibes, Sites C-10 to C-16A; spectively. The northwest coast had the Southwest coast- Isla de la Juventud, fewest observed coral taxa (57 taxa), per- Archipelago de los Canarreos, Golfo de haps due in part to fewer stations within Cazones, Bahia de Cochinos, Sites C-21 to this region as compared to all other regions. C-31A; Southeast coast- Banco Silvertown, The fewest coral taxa were observed at Site Jardines de La Reina, Granma National C-12A (11 taxa) at Guanahacabibes. This Park, Chivirico MPA, Punta Imias, station was clearly an outlier in terms of Sites C-38 to C-52A; Northeast Coast- observed coral species. Within octocorals, Punta Imias [north side], Cabo Lucrecia, Ellisella sp. was observed at all sites while Archipelago de Sabana-Camagüey, Sites Nicella sp. was present in all sites except C-53 to C-60A). Based on the presence/ C-12A. Likewise, among the Scleractinia, absence species data of Appendix 1, both Agaricia sp. were observed at all stations the northeastern and southeastern regions except C-12A. M. cavernosa was also com- had the greatest number of macrobenthos monly reported in the ROV annotations taxa (259 taxa each). The west and south- (32 of the 35 sites). Frequent Antipatharia west regions had 229 and 228 taxa, respec- taxa included Stichopathes sp. which was tively, and the northwest had the least (205 observed at all sites and Tanacetipathes taxa). However, the sites were not evenly sp. which occurred at 29 sites. Twenty distributed among these regions, so com- taxa were only observed at single sites, the parisons of species richness are only tenta- majority of which were either rare species tive. However, the number of taxa per site (e.g., the scleractinians Dendrogyra cylin- is more relevant, typically, each site had a drus and Madracis myriaster). In terms of similar amount of ROV time, basically one present/absence, few strong regional dif- dive for the macrobenthic surveys. Sites ferences were observed among coral taxa, C-48 (Granma) and C-50 (Chivirico) on the although the southeast coast showed the southeast coast had the most taxa (126 and greatest species richness for scleractin- 125 taxa, respectively), but C-52 (south ian species and may represent a hotspot Punta Imias), which is also on the south- of reef-building species diversity. All other east coast, had the least (57 taxa). Unlike regions were generally similar in terms of most of the other sites in that region, Site observed coral taxa, with the exception of C-52 was muddy from 160 to 130 m, and the northwest coast. did not have the vertical rock wall and but- Sponges- Only two sponge species were tresses found at C-48 and C-50 (Fig. 7). present at all sites. The first, Xestospongia

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 81 Reed • González-Díaz • Busutil • et. al. sp. Cu-01, may be a new species and resem- Peyssonneliaceae in the Rhodophyta; some bles X. muta, but with thicker walls and is green unidentified crustose forms, the gen- relatively shorter and chubby in appearance. era Halimeda (particularly H. copiosa, H. The second, as yet unknown, is a thick bright goreaui, H. tuna), Avrainvillea, Penicillus, yellow verongiid crust (Verongiida sp. Cu- Udotea (particularly U. cyathiformis) in the 01) that lacks any fiber skeleton. Three spe- Chlorophyta; and the genera Dictyota and cies were found at 33 sites (Agelas sceptrum, Lobophora in the Ochrophyta. Site C-10 Aiolochroia crassa, and X. muta). The most (Banco San Antonio, west coast) had the species-rich sites for sponges were found at greatest number of taxa (24), whereas C-52 Site C-07 (Archipelago Colorados) on the (southeast coast) which lacked the typical northwest coast with 72 taxa and, on the vertical wall had the least (4 taxa). The south coast, C-28 (Archipielago Canarreos) northeastern region had the highest num- with 68 taxa and C-50 (Chivirico) with 66 ber of macroalgal taxa identified so far (38 taxa. However, overall the most species-rich taxa), whereas the western regions had the region for sponges was the northeastern re- least (24 taxa in the northwest region, and gion (153 taxa), followed by the southeast- 25 in the southwest). These results could ern region (148 taxa) and western region indicate better conditions for the develop- (142 taxa). The fewest taxa were found in ment of macroalgae in the northeastern re- the northwest region (113 taxa) and south- gion of Cuba, and/or lower herbivory levels, west (131 taxa). By station the fewest but further work is still needed. sponges were found at Site C-52 (33 taxa) Fish- There were 178 fish taxa identified and C-52A (37 taxa) at Punta Imias on the from the ROV annotations (Appendix 2), southeast coast, and C-58A (36 taxa) at however only one, , was Archipelago de Camagüey on the northeast present at all 36 sites. There were three coast. Surprisingly, 99 sponge taxa were species present at 33 sites; Gramma melac- found exclusively at single sites (33% of the ara, Halichoeres garnoti, and Stegastes 296 morphotypes) which might reflect high partitus. An additional three species were level of endemism or widespread but patchy found at 32 sites; Holacanthus tricolor, distributions. The most diversified genera and the invasive Lionfish were: 1) Agelas with 16 identified species (6 Pterois volitans/miles. Similar to sponges, of which occur at least in 15 sites) and 10 the most species-rich site for fish was C-07 unidentified morphotypes, and 2) Aplysina on the northwest coast, with 67 taxa ob- with 10 identified species (6 taxa occur at served. Site C-12B at Guanahacabibes had more than 15 stations) and 7 unidentified 63 taxa and C-48 on the southeast coast morphotypes. The interpretation of the re- had 62 taxa. Regionally, the southeast gional and local patterns of sponge diversity coast was the richest with 112 fish taxa will require further analysis that will study present; however, the site with the fewest factors such as habitat, current patterns, taxa was on the southeast coast (Site C-52 and biogeographic affinities. with 17 taxa), which was also true for the Algae- Macroalgal taxa that were com- sponges and algae, followed by Site C-52A mon to all regions of the Cuban mesophotic (26 taxa) and C-04 on the northwest coast coral reefs included the coralline red algae (27 taxa). Overall, the northwest coast had ( S u b c l a s s C or a l l i nophy c id ae) a nd t he f a m i ly the lowest species richness with 91 taxa;

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however, this region had the fewest num- Caribbean, Gulf of Mexico, and southeast- ber of sites. Thirty-nine taxa were found ern U.S., and include deep island slopes, only at single sites while 13 were found at off-shore banks, and shelf-edge reefs. only two sites. The number of species found However, the MCE habitat and biota are at single sites will likely decrease after de- quite variable within these regions. The tailed analyses of ROV videotapes are general geomorphology and mesophotic completed and fishes overlooked during re- zonation of the deep fore-reef slope and al-time dive annotations are added to the escarpment along the Cuban coast are database. Fish assemblages were not sig- strikingly similar to those described for nificantly different from one another in- the Bahamas (Porter, 1973; Reed, 1985; side and outside the MPAs based on the Ginsburg et al., 1991; Lesser and Slattery, non-metric multidimensional scaling ordi- 2011); Jamaica (Goreau and Goreau, 1973; nation (MDS) plot of the presence/absence Goreau and Land, 1974; Lang, 1974; Land transformed fish data from the ROV ob- and Moore, 1977; Dustan and Lang, 2018), servations. With the exception of one dive, and the Belize barrier and atoll reefs fish assemblages from all dives were 40% (James and Ginsburg, 1979). The ‘Wall’ similar to one another, regardless if they at Belize also forms spectacular vertical were inside or outside the MPAs. Based on cliffs from depths of 65 to 120 m, with ver- the SIMPER (Similarity Percentages), the tical fissures, ledges and caves, and in the species that had the most effect on fish as- 1970s, the upper brow of the wall (45-65 semblage differences inside and outside the m) had coral growth consisting of large col- MPAs were higher abundances of Caranx onies of plate corals up to 50% cover. At lugubris, Haemulon plumierii, Scaridea, Discovery Bay, Jamaica, the wall forms an and Gramma loreto inside the MPAs and escarpment from 55 to 114 m, also with higher abundances of Holocentrus ru- caves and ledges, and in the 1970s-1980s fus outside the MPAs. In the western was dominated by sponges, algae, and cor- area, sites that coincided with marine als (its current condition is unknown, J. protected areas (Banco de San Antonio, Lang, pers. comm.). At that time, the deep Guanahacabibes National Park, and Cayo fore-reef slope (30-65 m) at Jamaica had Rosario) had greater abundances of com- similar dominant coral, sponge, and al- mercially important species, i.e., snapper gal species as do the Cuban deep reefs. (Lutjanidae), grouper (Serranidae), jack The corals were predominantly Orbicella (Carangidae) and mackerel (Scombridae). in high abundance to 45 m at some sites, The sites outside of these marine protected Agaricia lamarcki on upper slope, A. unda- areas had a lower abundance of these spe- ta, A. grahamae, Helioseris cuculata, and cies, which could be an indicator of histori- M. cavernosa on the lower slope; and the cal overfishing. algae were dominated by crustose coralline algae, Halimeda copiosa, H. cryptica, plus Discussion algal turfs and fleshy macroalgae (Goreau Mesophotic Coral Habitats of Caribbean, Gulf and Goreau, 1973; Lang, 1974; Dustan and of Mexico and Southeastern U.S. Lang, 2018). Hermatypic corals (primarily MCE habitats are quite common through- Agaricia spp.) dominated the wall around out the Western Atlantic regions of the 70 m. The deep island slope (> 120 m) in

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Jamaica was also similar to the Cuban habitat around St. Croix is mostly on the reefs, and had a similar heterotrophic com- steep island walls, which is typical of most munity consisting of abundant sponges, Caribbean Islands and Bahamas, whereas including coralline sponges, the azooxan- St. Thomas and St. John have a broader thellate scleractinians, alcyonaceans, fan shelf and deep banks. In the upper meso- and whip antipatharians, and stylaster- photic zone (35 m), the dominant coral are ine corals. In the Bahamas, Reed (1985) Orbicella spp. and Agaricia spp. (Smith compared several locations including San and Holstein, 2016). At deeper depths Salvador Island that has a narrow shelf, (65 m), the lettuce coral Agaricia undata and the windward and leeward sides of dominates. Grand and Little Bahama Banks. The wall Off southwestern Puerto Rico, deep of the deep fore-reef escarpment ranges reefs occur at depths of 45-160 m; deep from 45-143 m. The base generally ranges buttresses are common at depths of 45-65 from 100-122 m. Some sites also have large m below the shallow barrier reef system buttresses occurring every 20-60 m along (Appledoorn et al., 2016b). A terrace occurs the top edge of the deep fore-reef slope, with at 80-90 m, and a steep wall extends down sediment chutes incised between the prom- to 160 m (Sherman et al., 2010). This ter- ontories (Hoskin et al., 1986). Notches, race feature which Sherman et al. (2010) ledges and reentrants or caves are common suggests as being formed during the last between 75 and 107 m, similar to Cuba. At deglaciation (14-15 ka) is not apparent at San Salvador Island, Bahamas, these re- Cuba. Agaricia spp. and Madracis spp. cor- entrants likely connect to the brackish als dominate the MCE zone, and large col- lakes on the island which show tidal influ- onies of A. undata are common. Seventeen ence. These karst-like features associat- species of scleractinian coral in the MCE ed with deep fore reefs of Jamaica, Belize, zone (50-90 m). Bahamas and Cuba, likely formed during Down current from Cuba are the meso- Late Quaternary low stands of sea lev- photic reefs of the northern Gulf of Mexico el which were at least -120 m below pres- (GOM), including the Sister Sanctuaries ent level during the last glacial maximum (FGBNMS, FKNMS), and southeastern (Fairbanks, 1989). The major topographic U.S. The geomorpholog y of these U.S. meso- features are apparently controlled by un- photic reefs are quite different from those derlying Plio-Pleistocene features that are off Cuba. The shelf-edge reefs and banks in mantled by 1-10 m of Holocene reef accre- the northwestern GOM off Texas are built tions (James and Ginsburg, 1979; Ginsburg upon salt domes but also have a relative- et al., 1991). ly steep mesophotic reef habitat which ex- The Virgin Islands also support fair- tends over a wide depth zone. FGBNMS is ly extensive and diverse MCEs, predomi- 180 km off shore Texas and is described as nately on steep walls around the islands one of the most pristine coral ecosystems and some areas of deep banks (Smith in U.S. waters (Hickerson et al., 2008). and Holstein, 2016). Extensive areas of Although the top of the dome is a shallow the shelf that are potential MCE habitat water reef at depths of 18 m, the flanks ex- (1,918 km2) at depths of 25-65 m may ex- tend into mesophotic depths down to 152 m ceed the area of the shallow reefs. MCE (Clark et al. 2014). The mesophotic reefs at

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FGBNMS have been surveyed with ROV and supports a MCE of photosynthetic by some of our team (JV, JR) using similar hard corals (17 species), macroalgae (95), methods as in Cuba (Voss et al., 2014). The sponges (92), and a large variety of tropi- mesophotic scleractinian coral communi- cal fishes (86) (Harter et al., 2017; Reed et ties on FGB and nearby banks outside the al., 2017a; Reed et al., 2018). A total of 12 sanctuary have diverse hard coral popula- hard coral species were identified at Pulley tions (21 species) and are dominated by O. Ridge. Plate coral species at Pulley Ridge franksi, M. cavernosa, S. intersepta, and include Agaricia fragilis, A. lamarcki/gra- Agaricia spp. with Pseudodiploria strigo- hamae, A. undata, H. cucullata, and M. sa and Colpophyllia natans also being rela- cavernosa, all of which are also common tively common. Also common at FGBNMS on the Cuban reefs. Unfortunately, Reed at mid-mesophotic depths (40-80 m) are et al. (2018) found a shocking 93.6% loss algal nodules (rhodoliths) which we did of coral cover since 2004 within the Pulley not find at Cuba. The deep coral zone at Ridge Habitat Area of Particular Concern FGBNMS from 85-150 m was primarily (PR HAPC). Recent taxonomic studies of on the steep slopes of the banks, and domi- sponges collected at Pulley Ridge (Reed et nated by azooxanthellate corals, antipath- al. 2017a), show a rich sponge fauna with arian and gorgonian corals, , and several new species; however, there seems sponges. Many of these species are similar to be important differences between Pulley to Cuba on the lower mesophotic zone and ridge and Cuban mesophotic sponge fauna deep island slope. which must await for comparative biodiver- In the northeastern GOM, mesophotic- sity studies. depth reefs are common all along the 70 Just 60 km east of Pulley Ridge and 225 m contour (60-90 m) from Alabama to km west of Key West, Florida, at the termi- the panhandle of Florida, and along the nus of the Florida Keys island chain, are the northwestern Florida shelf (Locker et al., Tortugas Ecological Reserves (TER) which 2010; Locker et al., 2016; see Reed, 2016 are part of the ‘Sister Sanctuaries’ within for map). However, for the most part, these the FKNMS that have shallow mesophotic lack zooxanthellate corals and are domi- reef habitat. TER North protects coral nated by azooxanthellate corals such as reef banks known as Tortugas Bank (30- Madracis sp. and Oculina sp., octocorals, 50 m depths). TER South includes Riley’s black corals, sponges, and algae. In con- Hump (30-50 m) which is an isolated off- trast to these reefs, Pulley Ridge, off the shore mound that has shallow MCE hab- southwest Florida shelf, is one of the clos- itat (Weaver et al., 2006). Scleractinians est downstream mesophotic reefs to Cuba. include M. cavernosa, O. annularis, and It is a submerged intact barrier island that Siderastrea siderea. Individual colonies of formed during the early Holocene marine M. cavernosa are mostly small, ranging transgression, and consists of the deep- from 10-50 cm in diameter but some were est photosynthetic coral reef in continen- observed to 2 m in height. At deeper meso- tal U.S. (Hine et al., 2008; Reed, 2016). photic depths, the benthic community is The southern terminus of this geological dominated by rope sponges (Aplysina cau- feature is relatively flat limestone pave- liformis) and calcareous algae including ment and rubble at depths of 60-94 m, Halimeda spp. and Penicillus spp., similar

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 85 Reed • González-Díaz • Busutil • et. al. to Cuba. The southern edge of the Hump is for the Oculina reefs that extend 250 km a sharp escarpment from 84-124 m called at depths of 60-90 m off eastern Florida, Miller’s Ledge. The benthic assemblages of and then continues as shelf-edge rock ledg- Miller’s Ledge are not MCEs but are domi- es from north Florida to North Carolina. nated by small sponges, bryozoans, small The deep Oculina coral reefs, however, are solitary corals, and the corkscrew sea entirely formed by azooxanthellate O. vari- whip, Stichopathes sp. This is more simi- cosa (Avent et al., 1977; Reed, 1980; Reed lar to what we found deeper in Cuba on the et al., 2007; Harter et al., 2009). The shelf deep island slope of 120->150 m. edge from Florida to North Carolina is pri- Further downstream are shallow shelf- marily hard-bottom habitat that consists of edge reefs which extend along most of the extensive ledges and escarpments at meso- Florida Keys and southeastern Florida but photic depths, and have dense populations mostly only extend to 20-25 m depths. This of sponges, algae and octocorals, but lim- region of Florida has very limited MCE hab- ited Scleractinia, mostly Oculina varicosa itat, primarily scattered patch reefs, ledg- and Madracis sp. (Harter et al., 2015). es, or rubble habitat at depths of 30-100 m. However, the mesophotic depth zone (30- Depth Records 150 m) has not been adequately surveyed in The maximum depth of zooxanthel- this region. The earliest survey of its meso- late scleractinians in the world is con- photic reefs occurred in 1979 off Key Largo sidered to be 165 m in the Pacific, where in the upper Florida Keys when Jameson the plate coral Leptoseris hawaiiensis oc- (1981) documented deep coral reef habitat curs (Kahng and Maragos, 2006). In the to 39 m and coral algal habitat to 55 m. Atlantic, the maximum depth is now 122 In 2015, Reed et al. found some deep reefs m for Agaricia sp. in Cuba. In the Atlantic (125 m) with ROV off Key West, Florida, Ocean and Caribbean, nine genera and at but these lacked zooxanthellate hard cor- least 18 species of hermatypic scleractin- als and were dominated by the octocoral ia occur to depths exceeding 70 m (Reed, genera (Diodogorgia, Thesea, Villogorgia, 1985). In the eastern GOM, the maximum and Nidalia) and sponges. However, it did depth range for 22 mesophotic scleractin- provide essential fish habitat for sever- ian species is 31-60 m and 15 species have al commercially important species includ- a maximum depth of 100 m (Jaap, 2015). ing snowy grouper, tilefish and snapper. Reed (1985) had found ten new depth re- Based on the presence of paleo-shorelines cords, including Agaricia grahamae (119 elsewhere, we could expect to find ledges m), Helioseris (syn. Leptoseris) cucullata or hard bottom habitat especially along (108 m), Montastraea cavernosa (113 m), the 70 and 125 m contours off the Florida and Madracis sp. (probably M. decactis Keys and southeastern U.S. (Locker et al., or M. pharensis f. luciphila, 115 m), all 2010). Some deep-water algal studies have on the deep fore-reef escarpments off San also been conducted along the shelf off Salvador Island, Bahamas. In comparison, southeastern Florida (Hanisak and Blair, in Cuba (Table 3) we found Agaricia sp. to 1988; Leichter et al., 2008) which could a maximum depth of 122 m, which is now a imply the presence of mesophotic habitat. new depth record in the Tropical Western This paleo-shoreline also forms the basis Atlantic, along with the depth record for

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Stephanocoenia intersepta (104 m) which in the category of ‘White Syndromes” due was previously 100 m (Reed, 1985). In to the characteristic white areas of recent the Caribbean, Kühlmann (1983) had re- tissue loss, similar to white plague disease ported S. intersepta (S. michelini, syn.) to (Weil and Hooten, 2008; Andradi-Brown, depths of 95 m, and Agaricia spp. to 79 m. 2016). White syndromes disease was al- The deepest records for algae in the west- so reported on mesophotic reefs off south- ern Atlantic include Halimeda copiosa (130 western Puerto Rico in Agaricia undata, m) and crustose coralline algae (268 m), al- A. lamarcki, A. grahamae, and A. agaric- so reported from San Salvador by Littler ites (Appledoorn et al., 2016b). Six percent et al. (1985). In comparison, maximum of the coral community showed signs of the depth of algal observations on our Cuba disease. Dustan and Lang (2018) have com- dives include an unidentified green crust piled detailed descriptions of mesophotic (169 m), crustose coralline algae (169 m), reefs on the fore-reef slope at Discovery Peyssonneliaceae (139 m), and Halimeda Bay, Jamaica, comparing their conditions sp. (likely H. copiosa; 127 m). from the 1960s with the present. Extensive changes have occurred, especially since Coral Disease/ Bleaching the 1980s due to the mass mortality of the Regarding coral health of Cuba’s meso- herbivorous Diadema (to 40 m depths), photic reefs, of the 2,240 scleractinian coral disease, bleaching, loss of other her- colonies counted in this study, only 12 cor- bivores (from overfishing), reduced water als (0.53%, mainly Agaricia spp.) showed quality, and hurricanes. Although the pin- signs of bleaching. One Agaricia had black nacles still retain their three-dimensional, band disease, and one had an unidenti- “shingled” morphology to ~45 m, partial fied white syndrome disease (0.009%). In mortality had partitioned many of the orig- shallow water, the presence of black band, inal Orbicella colonies into smaller isolates white band, and plague diseases have been by 2013. This appears somewhat similar recorded in Cuba (González Ferrer, 2004); to the upper mesophotic zone for some of white band was found in isolated colonies the Cuban reefs where the slopes are cov- of Acropora cervicornis, and black band ered with many smaller plate corals rather was found with relative frequency in iso- than the extensive sheets that may have lated colonies (Alcolado et al., 2000). In been common in the 1960s-70s. Zlatarski general, the Cuban mesophotic corals ap- and Estalella (1982) reported colonies of peared quite healthy compared to many Agaricia (f. unifaciata, likely A. lamarcki) shallow Caribbean reef sites. In the past to diameters of 3.5 to 5.6 m on the Cuban decade disease in shallow corals has in- reefs in the 1970s. creased as seawater temperatures increase Few MCEs worldwide have long-term re- (Andradi-Brown, 2016) and reports now cords monitoring changes of temperature show mesophotic coral communities are over time or shorter-term events such as not immune to disease. In the Caribbean cold-water upwelling or internal waves suf- U.S. Virgin Islands, disease in mesophotic ficient to cause coral impacts. Long-term re- corals has been documented down to 100 cords at some sites do show bleaching events m depths (Smith et al., 2010). The type of and there is evidence that some mesophotic disease in the Virgin Islands was placed corals may have lower bleaching thresholds

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 87 Reed • González-Díaz • Busutil • et. al. than shallow colonies (Baker et al., 2008, Guajimico) but showed a decline of 1.75% / Andradi-Brown et al., 2016). In the region of year which is slightly lower than estimated the Gulf of Mexico, a mass bleaching event at for the wider Caribbean (2.5-2.7% decline) the FGBNMS in 2005 impacted 45% of the (Alcolado et al., 2009). coral community (Precht et al., 2006; Clark et al., 2014). But the reefs showed resiliency Direct Human Impacts and the bleaching had been reduced to 4% In general, most dive sites appeared rel- of the population within a year. Since the atively pristine with little signs of direct millennium, coral reefs in the northeastern human impact. One site at Cayo Largo Caribbean have had many bleaching events del Sur MPA (Site 29) showed evidence of including two major bleaching events, in nutrient pollution. Landward of the shelf- 2005 and 2010 (Smith and Holstein, 2016). edge reef crest at Cayo Largo were exten- In the U.S. Virgin Islands, mesophotic cor- sive cyanobacterial mats on sediment, and al cover in some areas dropped from >25% the reef crest (35-40 m) had dense areas cover (Aronson et al., 1994) to current- of 1-m conical mounds, which appeared to ly less than 10% (García-Sais et al., 2014) be old dead coral, probably M. cavernosa. which has been attributed to both bleaching Alcolado et al. (2001b) reported high nitro- events and several large hurricanes (Smith gen nutrients and dead coral at this site at- and Holstein, 2016). Temperature records tributed from a tourist resort on nearby on Puerto Rico’s mesophotic reefs show a Cayo Largo where a sewage plant dumped seasonal range of 26-29.8oC (Appledoorn partially treated waste water. On Cuba’s et al., 2016a). Temperatures recorded by shallow reefs, areas exposed to intense our ROV dives around Cuba in May-June fishing and pollution, such as along the showed a maximum surface temperature of northwest coast and especially off Havana 29.52oC along the south coast, whereas at show reduced coral density and diversity 50 m depth, in the mesophotic zone, temper- (Duran et al., 2018). Increased nutrients atures ranged from 25.26-28.62oC, and a have been linked to algal and cyanobacteri- thermocline of 25-28oC was common around al blooms elsewhere and on reefs in Florida 80-100 m depths. Although we do not have and Belize (Lapointe, 2004; Lapointe et temperature records over the year, this up- al., 2004; Littler et al., 2006). In addition, per temperature range would not be suffi- lost or discarded fishing gear were rela- cient to cause bleaching typically for shallow tively uncommon; in total, fishing line or corals, but the tolerance levels of mesophotic long line were found at eight sites and one corals are less well understood. Since 1983, lost net was observed. Most of these were when the first bleaching event in Cuba was found on the substrate on the upper meso- recorded, there have been 7 major events photic reefs but some were found at >100 in Cuba (Jackson et al., 2014). Recent sur- m depths. veys of Cuba’s shallow reefs show a loss of reef-building species, such as O. annularis Invasive Species and M. cavernosa (Aguilar Betancourt and Several invasive species have impact- González-Sansón, 2007). Average coral cov- ed Western Atlantic MCEs; in particular, er on the fore reefs (2003-2009) was 13.4% various macroalgae such as Codium and (maximum of 27% at Bahia de Cochinos and Caulerpa, the gorgonian Carijoa riisei,

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and Lionfish Pterois volitans (Lapointe et makes it difficult to effectively control the al., 2005; Andradi-Brown et al., 2016). In species in shallow waters due to the contri- the past decade Lionfish have become a bution of larvae, juveniles and adults from major invasive on most MCE reefs in the deeper areas, where they are also difficult Caribbean, Gulf of Mexico and southeast- to catch. For this reason, there would be ern U.S. (Schofield, 2010; Andradi-Brown an intact reservoir of its populations in the et al. 2016). The first live Lionfish reported mesophotic reefs which tentatively could in northern Gulf of Mexico were document- populate shallow reefs. ed at Pulley Ridge mesophotic reef west of the Florida Keys during submersible dives Conclusions in 2010 where six fish were observed (Reed This Joint Cuba-U.S. Expedition provides and Rogers, 2011). Since then, exponential for the first time data on the extent and increases in the abundance of Lionfish have health of mesophotic coral reefs around the been observed at Pulley Ridge mesophotic entire coast of Cuba, covering nearly 2,778 reef (Andradi-Brown et al., 2016; Harter et km. This research cruise is of special sig- al., 2017). Thousands of Lionfish also have nificance to understanding the distribution been recently documented at mesophotic of the deep reefs and the basic oceano- depths in protected area reefs along the graphic, biological and ecological processes southeastern U.S. from Florida to North that take place in MCEs around Cuba and Carolina at depths of 50->100 m (Harter at local and regional levels. Future anal- et al., 2015). Recent genetic analyses indi- yses of the taxonomy and genetics of the cate western Atlantic Lionfish most com- specimens, along with quantitative analy- monly cited as Pterois volitans are hybrids ses of the video and photo data, will allow a between P. miles and P. lunulata/russelii more precise characterization of the diver- (Wilcox et al., 2017); however, all of our iden- sity and relative abundance of the meso- tifications were based upon photographic photic communities of Cuba, as well as a data which precluded morphological or ge- better understanding of the connectivity of netic analyses. On the Cuban mesophotic Cuban reefs with the Sister Sanctuaries in reefs, we observed Lionfish at most sites, the U.S. and elsewhere in the Caribbean. but the abundance of Lionfish was gener- Also these data may lead to expanding or ally low (<10 observed per site). Impacts of adding new Marine Protected Areas in Lionfish on MCEs could result in declines Cuba or more fully protecting some exist- of prey fish biomass and major shifts in the ing MPAs. The full cruise report (Reed et mesophotic benthic community. In stud- al., 2017b) provides a detailed characteriza- ies of mesophotic reefs in the Bahamas, tion of each dive site which provides bench- Lionfish predation on herbivorous fish has mark data for comparisons with future caused a shift in the benthic community studies and the effects of climate change. to an increase of algae relative to corals (Lesser and Slattery, 2011). However, from Acknowledgements ROV video surveys at Pulley Ridge, there This research was supported by the NOAA has been no significant Lionfish effect on Office of Ocean Exploration and Research the fish assemblages (Harter et al., 2017). under award number NA14OAR4320260 The presence of Lionfish at greater depths to the Cooperative Institute for Ocean

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 89 Reed • González-Díaz • Busutil • et. al.

Exploration, Research and Technology Center of Marine Research (CIM-UH); and (CIOERT) at Harbor Branch Oceanographic Yusimí A lfonso from the National Aquarium Institute-Florida Atlantic University (ANC). This paper is Contribution Number (HBOI-FAU), and by the NOAA Pacific 2151 from Harbor Branch Oceanographic Marine Environmental Laboratory un- Institute at Florida Atlantic University. der award number NA150AR4320064 to the Cooperative Institute for Marine References and Atmospheric Studies (CIMAS) at Aguilar Betancourt, C. and González- the University of Miami. This expedi- Sansón, G. (2007). Composición de la tion was conducted in support of the Joint ictiofauna costera de Ciudad de la Ha- Statement between the United States bana y evaluación preliminar de los fac- of America and the Republic of Cuba on tores que la determinan. Rev. Invest. Cooperation on Environmental Protection Mar., 28(1), 43-56. (November 24, 2015) and the Memorandum Aguilar, C., González-Sansón, G., Munkit- of Understanding between the United trick, K. R. and MacLatchy, D. L. States National Oceanic and Atmospheric (2004). Fish assemblages on fringe cor- Administration, the U.S. National Park al reefs of the northern coast of Cuba Service, and Cuba’s National Center for near Havana Harbor. Ecotoxicol. En- Protected Areas. We give special thanks to viron. Saf., 58(1), 126-138. doi:10.1016/ Carlos Díaz Maza (Director of the National S0147-6513(03)00104-0 Center of Protected Areas) and Ulises Alcolado, P., Claro, R., Martínez-Daranas, Fernández Gomez (International Relations B., Menéndez-Macías, G., García-Parra- Officer, Ministry of Science, Technology do, P., Cantelar-Ramos, K., … Neff, T. and Environment; CITMA) for assistance (2000). Estado general de los arrecifes co- in securing the necessary permits to con- ralinos de Cuba y propuestas de manejo duct the expedition and for their tremen- ambiental. (Informe Final del Proyecto). dous hospitality and logistical support in La Habana: Instituto Oceanología. Cuba. We thank the Captain and crew of Alcolado, P. M., Caballero Aragón, H., the University of Miami R/V F.G. Walton and Perera, S. (2009). Tendencia del Smith and ROV operators Lance Horn cambio en el cubrimiento vivo por cora- and Jason White, University of North les pétreos en los arrecifes coralinos de Carolina at Wilmington (UNCW-CIOERT), Cuba. Ser. Oceanol., 5, 1-14. Undersea Vehicle Program for their excel- Alcolado, P. M., Caballero, H. and Rey- lent work at sea during the expedition. Villiers, N. (2015). Understanding Cor- The Mohawk ROV is owned by the NOAA al Reef Resilience in Cuba. Congress National Marine Sanctuaries Foundation. MARCUBA 2015, Havana, Cuba. Taxonomic identifications were provided Alcolado, P. M., Claro-Madruga, R., Mar- primarily by: - JR, JV, PGD, JGM, tínez-Daranas, B., Menéndez-Macía, G., MS; Porifera- CD, LBL, SP; Fish- AD, FD, García-Parrado, P., Cantelar, K. … del DCR, AGR; macroalgae- BMD, MDH, PGS. Valle, R. (2001a). Evaluación ecológica For assistance with macroalgal identifica- de los arrecifes coralinos del oeste de tions, we also thank Ana M. Suárez, Emma Cayo Largo del Sur, Cuba: 1998-1999. E. Gómez, and Amanda Ramos from the Bol. Invest. Mar. Cost., 30, 109-132.

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Como citar este artículo Reed, J.K., González-Díaz, P., Busutil, L., Farrington, S., Martínez-Daranas, B., Cobián Rojas, D., ... Pomponi, S.A. (2018). Cuba’s Mesophotic Coral Reefs and Associated Fish Communities. Rev. Invest. Mar., 38(1), 60-129.

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Appendices

Appendix 1. Species list of benthic macrobiota (invertebrates and macroalgae) identified at Cuba’s mesophotic reefs from ROV dives during the F.G. Walton Smith Cuba cruise, May 14 to June 12, 2017. Dive Sites (see Fig. 1) compiled by regions: Northwest coast- Archipelago de Los Colorados, Sites C-04 to C-07; West coast- Banco de San Antonio, Guanahacabibes, Sites C-10 to C-16A; Southwest coast- Isla de la Juventud, Archipelago de los Canarreos, Golfo de Cazones, Bahia de Cochinos, Sites C-21 to C-31A; Southeast coast- Banco Silvertown, Jardines de La Reina, Granma National Park, Chivirico MPA, Punta Imias, Sites C-38 to C-52A; Northeast Coast- Punta Imias (north side), Cabo Lucrecia, Archipelago de Sabana-Camagüey, Sites C-53 to C-60A. Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast /Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Algae Cyanobacteria X X X X X X X X Chlorophyta Anadyomene stellata (Wulfen) C. X X X X X X X Agardh, 1823 Avrainvillea sp. X X X X X X Caulerpa chemnitzia (Esper) J.V. X X X X Lamouroux, 1809 Chaetomorpha sp. X Chlorophyta- unid. spp. 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 Cladophora sp. X X Cladophoropsis sp. X X Dictyosphaeria cavernosa X (Forsskål) Børgesen, 1932 Halimeda copiosa Goreau & 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 E.A.Graham, 1967 Halimeda discoidea Decaisne, X X X X X X X X X X X X X 1842 Halimeda goreaui W.R.Taylor, X X X X X X X X X X X X X X 1962 Halimeda incrassata (J.Ellis) X X J.V.Lamouroux, 1816 Halimeda opuntia (Linnaeus) J. X V. Lamouroux, 1816 Halimeda sp. 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 Halimeda tuna (J.Ellis & X X X X X X X X X X X X X X X X X X X Solander) J.V.Lamouroux, 1816 Microdictyon marinum (Bory) X P.C.Silva, 1955

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Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Microdictyon sp. X X X X X Microdictyon umbilicatum X X X X X (Velleg) Zanardini, 1862 Penicillus dumetosus Børgesen, X X X X X 1913 Penicillus pyriformis A.Gepp & X E.S.Gepp, 1905 Penicillus sp. X X X X X X Rhipocephalus oblongus X X X X (Decaisne) Kützing, 1849 Rhipocephalus phoenix (J. Ellis X X X X X X X X X & Solander) Kützing, 1843 Udotea cyathiformis (J. Agardh) D. S. Littler & M. M. Littler, X X X X X X X 1990 Udotea dixonii D. S. Littler & M. X X M. Littler, 1990 Udotea sp. X X X X X X X X X X X Valonia macrophysa Kützing, X X 1843 Valonia sp. X Valonia ventricosa J. Agardh, X X 1887 Ochrophyta Canistrocarpus cervicornis De Paula & De Clerck in De Clerck X et al., 2006 Dictyota menstrualis (Hoyt) Schnetter, Hörning & Weber- X Peukert, 1987 Dictyota pulchella Hörnig & X X Schnetter, 1988 Dictyota sp. 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 Lobophora sp. 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 Phaeophyceae- unid. spp. X X X X X Sargassum hystrix J.Agardh, X X X X X X X 1847 Sargassum sp. X X X X

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Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Stypopodium zonale (J. V. Lamouroux) Papenfuss , X X X X 1940 Rhodophyta Amphiroa fragilissima (Linnaeus) X X X J. V. Lamouroux, 1816 Amphiroa rigida J.V.Lamouroux, X 1816 Amphiroa sp. X Botryocladia pyriformis X (Børgesen) Kylin, 1931 Botryocladia sp. X Chrysymenia enteromorpha X Harvey, 1853 Chrysymenia sp. X Corallinophycidae (crustose 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 coralline algae) Cryptonemia sp. X Dasya sp. X Dichotomaria marginata (W. R. X Taylor) M. J. Wynne, 2005 Dichotomaria obtusata (J. Ellis & X Solander) Lamarck, 1816 Gracilaria sp. X Hypoglossum involvens (Harvey) X J. Agardh, 1898 Jania adhaerens J. V. X Lamouroux, 1812 Jania capillacea Harvey, 1853 X Jania pumila J.V.Lamouroux, X 1816 Jania sp. X Laurencia sp. X Martensia pavonia (J.Agardh) X J.Agardh, 1863 Melanothamnus pseudovillum (Hollenb.) Díaz-Tapia & Maggs, X 2017 REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 101 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Peyssonneliaceae 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 Rhodophyta- unid. spp. X X X X X Scinaia complanata (F.S.Collins) X A.D.Cotton, 1907 Wrangelia sp. X Porifera Calcarea Calcarea sp. Cu-01 X X X Calcarea sp. Cu-02 X X Calcarea sp. Cu-03 X Calcarea unid. sp. X X Leucetta cf. floridana (Haeckel, X X X X 1872) Leucetta floridana (Haeckel, 1872) X X X X X Leucetta sp. Cu-01 X X Leucetta sp. Cu-02 X Demospongiae Aaptos sp. Cu-01 X X X X Agelas cerebrum Assmann, van X X X X X X X X X X X X Soest & Köck, 2001 Agelas cervicornis (Schmidt, X X X X X X X X X X X X X X X X X X X X X X X 1870) Agelas cf. flabelliformis (Carter, X X X 1883) Agelas cf. tubulata Lehnert & X van Soest, 1996 Agelas citrina Gotera & Alcolado, 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 1987 (Schmidt, X X X X X X X X X X X X X X X X X 1870) (Schmidt, 1870) X X X X X X X X X X X X X X X X X X X X X X Agelas dilatata Duchassaing & X X X X X X X X X X X X X X X X Michelotti, 1864 Agelas dispar Duchassaing & X X X X X X X X X Michelotti, 1864 Agelas flabelliformis (Carter, X X X X X X X X X X X X X X 1883)

REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 102 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Agelas repens Lehnert & van X X Soest, 1998 Agelas sceptrum (Lamarck, 1815) 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 Wilson, 1902 X X Agelas sp. Cu-01 X Agelas sp. Cu-02 X X Agelas sp. Cu-03 X X X Agelas sp. Cu-04 X Agelas sp. Cu-05 X X Agelas sp. Cu-06 X X X Agelas sp. Cu-07 X X X Agelas sp. Cu-08 X X X X X X X X X X X Agelas sp. Cu-09 X Agelas sp. Cu-10 X Agelas sventres Lehnert & van X X X X Soest, 1996 Lehnert & van X X X X X X X X Soest, 1996 Agelas wiedenmayeri Alcolado, X X X X X 1984 Aiolochroia crassa (Hyatt, 1875) 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 Aiolochroia crassa var. dark blue X X (Hyatt, 1875) Aiolochroia crassa var. gray X X X X (Hyatt, 1875) Aiolochroia crassa var. purple X X X X (Hyatt, 1875) Aiolochroia crassa var. purple- X X X X X blue (Hyatt, 1875) Aiolochroia sp. Cu-01 X X X X X X X Aiolochroia sp. Cu-02 X X X X Amphimedon cf. caribica X X X X X X X (Pulitzer-Finali, 1986) Amphimedon compressa 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 Duchassaing & Michelotti, 1864 Amphimedon sp. Cu-01 X X X X X X X X X X Amphimedon sp. Cu-02 X X X X X REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 103 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Amphimedon sp. Cu-03 X Amphimedon sp. Cu-04 X Amphimedon sp. Cu-05 X Amphimedon sp. Cu-06 X Aplysina aff. lacunosa (Lamarck, X X 1814) Aplysina archeri (Higgin, 1875) 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 Aplysina bathyphila Maldonado 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 & Young, 1998 Aplysina cauliformis (Carter, 1882) 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 Aplysina cf. archeri (Higgin, 1875) X Aplysina cf. fulva (Pallas, 1766) X X X X X X X (Pallas, 1766) X X X X X X X X X X X X X X X X X X X X Aplysina fulva (Pallas, 1766) X X X X X Aplysina insularis (Duchassaing X & Michelotti, 1864) Aplysina lacunosa (Lamarck, X X X X X X X X X X X 1814) Aplysina sciophila Rützler, X X X X X X X X X X X X X X X X Piantoni, van Soest & Díaz, 2014 Aplysina sp. Cu-01 X X X X Aplysina sp. Cu-02 X Aplysina sp. Cu-03 X X X X X X X X X Aplysina sp. Cu-04 X X X X X X X X X X X X X X X X X X X X X X Aplysina sp. Cu-05 X Aplysina sp. Cu-06 X Asteropus sp. Cu-01 X X X X X X Auletta cf. tuberosa Alvarez, van X X X X X Soest & Rützler, 1998 Auletta sp. Cu-01 X Axinella cf. corrugata (George & X Wilson, 1919) Axinella corrugata (George & X X X X X X X Wilson, 1919) Axinellidae Cu-01 X X X X X X Callyspongia armigera X X X X (Duchassaing & Michelotti, 1864) REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 104 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Callyspongia cf. armigera X (Duchassaing & Michelotti, 1864) Callyspongia cf. fallax X X Duchassaing & Michelotti, 1864 Callyspongia cf. plicifera X X (Lamarck, 1814) Callyspongia fallax Duchassaing X X X & Michelotti, 1864 Callyspongia pallida Hechtel, 1965 X Callyspongia plicifera (Lamarck, X X X X X X X X X X X X X X X X X 1814) Callyspongia sp. Cu-01 X Callyspongia sp. Cu-02 X X X X X X X X X X Callyspongia sp. Cu-03 X X Callyspongia sp. Cu-04 X Callyspongia sp. Cu-05 X Callyspongia sp. Cu-06 X Callyspongia vaginalis (Lamarck, X X X X X X 1814) Ceratoporella nicholsoni X X X X X X X X X X X X X X X X X X X X X X X (Hickson, 1911) Chondrosia sp. X Cinachyrella cf. alloclada X (Uliczka, 1929) Cinachyrella kuekenthali X X X X X X X X X X (Uliczka, 1929) Cinachyrella sp. Cu-01 X X X X Cinachyrella sp. Cu-02 X X Cinachyrella sp. Cu-03 X X X X X X X X X X X X X X X Cinachyrella sp. Cu-04 X X X Cinachyrella sp. Cu-05 X X Ciocalypta cf. porrecta (Topsent, X X X X X 1928) Clathria echinata (Alcolado, 1984) X X X X X Clathria sp. Cu-01 X X Clathria sp. Cu-02 X Clathria sp. Cu-03 X X REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 105 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Clathria sp. Cu-04 X Clathria venosa (Alcolado, 1984) X X X X X X Cliona caribbaea Carter, 1882 X X X Cliona delitrix Pang, 1973 X X X X X X X X X X X X X X X X X X X X X X Cliona sp. X X Corallistes sp. X X X X X X X X X X X X Cribrochalina vasculum X X X X X X X X X X X X X X X X X X X X X X (Lamarck, 1814) Demospongiae sp. Cu-01 X X Demospongiae sp. Cu-02 X X X Demospongiae sp. Cu-03 X Demospongiae sp. Cu-04 X X X Demospongiae sp. Cu-05 X Demospongiae sp. Cu-06 X X X X Demospongiae sp. Cu-07 X Demospongiae sp. Cu-08 X X Demospongiae sp. Cu-09 X X Demospongiae sp. Cu-10 X X Demospongiae sp. Cu-11 X X X X X X X X X X X Demospongiae sp. Cu-12 X Demospongiae sp. Cu-13 X X Demospongiae sp. Cu-14 X X X X X X X X X X Demospongiae sp. Cu-15 X Demospongiae sp. Cu-16 X X Demospongiae sp. Cu-17 X Demospongiae sp. Cu-18 X Demospongiae sp. Cu-19 X Demospongiae sp. Cu-20 X X X Demospongiae sp. Cu-21 X X Demospongiae sp. Cu-22 X X X X X X X X X X X Demospongiae sp. Cu-23 X X Demospongiae sp. Cu-24 X Demospongiae sp. Cu-25 X X X X X X X X X X X Demospongiae unid. sp. 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

REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 106 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Desmapsamma anchorata X X (Carter, 1882) Desmapsamma sp. X sp. Cu-01 X Dictyoceratida unid. sp. X X X X Diplastrella megastellata X X Hechtel, 1965 Diplastrella sp. X X Discodermia sp. Cu-01 X X X X Dragmacidon alvarezae Zea & Pulido, 2016 X X X X Dragmacidon cf. alvarezae Zea & X X X X X X X Pulido, 2016 Dragmacidon sp. Cu-01 X Dysidea sp. X X Ectyoplasia ferox (Duchassaing & Michelotti, 1864) X X X X X X Erylus cf. formosus Sollas, 1886 X X Erylus new sp. X Erylus sp. Cu-01 X Geodia cf. cribata Rützler, X X X X X X X X X X X X Piantoni, van Soest & Díaz, 2014 Geodia neptuni (Sollas, 1886) 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 Geodia sp. Cu-01 X X Geodia sp. Cu-02 X X X Geodia sp. Cu-03 X X X X X X X X X X X X X Geodia sp. Cu-04 X Geodia sp. Cu-05 X Geodia sp. Cu-06 X Haliclona sp. Cu-01 X Halisarca caerulea Vacelet & Donadey, 1987 X X X Haplosclerida unid. sp. X X Hymeniacidon caerulea Pulitzer- X X Finali, 1986 Igernella sp. X X X X X

REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 107 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Iotrochota birotulata (Higgin, 1877) X X X X X X X X X X X X X X Ircinia campana (Lamarck, 1814) X X X X X Ircinia cf. strobilina (Lamarck, X 1816) Ircinia felix (Duchassaing & X X X X X X X Michelotti, 1864) Ircinia sp. Cu-01 X Ircinia sp. Cu-02 X X X X Ircinia sp. Cu-03 X X X X X Ircinia sp. Cu-04 X X Ircinia sp. Cu-05 X Ircinia strobilina (Lamarck, 1816) X X X X X X X X X X X X X X X X X X X X X X Leiodermatium sp. X X X X Lissodendoryx colombiensis Zea X X & van Soest, 1986 Melophlus ruber Lehnert & van X Soest, 1998 Microcionidae unid. sp. X X Monanchora arbuscula X X X (Duchassaing & Michelotti, 1864) Mycale cf. laevis (Carter, 1882) X X X X Mycale laevis (Carter, 1882) X X X X X X X X Mycale laxissima (Duchassaing X X X X X X X X X X X X X X X X X X X X X X X & Michelotti, 1864) Myrmekioderma gyroderma X (Alcolado, 1984) Myrmekioderma sp. Cu-01 X Myrmekioderma sp. Cu-02 X X X X X X X X X (Duchassaing & Michelotti, 1864) Neopetrosia carbonaria X (Lamarck, 1814) Neopetrosia cf. dutchi Van X X Soest, Meesters & Becking, 2014 Neopetrosia subtriangularis X (Duchassaing, 1850) Niphates alba van Soest, 1980 X X X X X X X X X

REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 108 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Niphates arenata Rützler, X X X X X X X X X X X X X X X X X X X X X Piantoni, van Soest & Díaz, 2014 Niphates cf. erecta Duchassaing X & Michelotti, 1864 Niphates digitalis (Lamarck, X X X X X X X X X X X X X X X X X X X X 1814) Niphates erecta Duchassaing & X X X X X X X X X Michelotti, 1864 Niphates sp. Cu-01 X X X X X X X Niphates sp. Cu-02 X X X X Niphatidae Cu-01 X Oceanapia bartschi (de Laubenfels, 1934) X Oceanapia peltata (Schmidt, 1870) X X Oceanapia sp. Cu-01 X X X X X X X X X X X X X X X X X X X X X X Oceanapia sp. Cu-02 X X X X X X X X X X X X X Oceanapia sp. Cu-03 X X Oceanapia sp. Cu-04 X X X X X X X Oceanapia sp. Cu-05 X X X X X Oceanapia sp. Cu-06 X Oceanapia sp. Cu-07 X Petrosia weinbergi van Soest, X X X X X X X X X X X X X X X X X 1980 Petrosiidae Cu-01 X X Petrosiidae Cu-02 X Petrosiidae Cu-03 X X Petrosiidae Cu-04 X Petrosiidae Cu-05 X Petrosiidae Cu-06 X X X X X X X X X X Petrosiidae Cu-07 X Petrosiidae Cu-08 X Petrosiidae Cu-09 X Petrosiidae Cu-10 X X X Petrosiidae Cu-11 X Petrosiidae Cu-12 X X X X X REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 109 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Petrosiidae Cu-13 X Petrosiidae Cu-14 X Petrosiidae Cu-15 X X X X X Petrosiidae Cu-16 X Petrosiidae Cu-17 X Petrosiidae Cu-18 X X Petrosiidae Cu-19 X Petrosiidae Cu-20 X X Petrosiidae unid. sp. X X X X X X X X X Phakellia folium Schmidt, 1870 X X X X X X X X Phorbas sp. X Polymastia sp. Cu-01 X X X X X X X X X X X X X Polymastia sp. Cu-02 X X Polymastia sp. Cu-03 X Polymastia sp. Cu-04 X X X X Ptilocaulis walpersi (Duchassaing X X X X X X X X X X X X X X X X & Michelotti, 1864) Scopalina ruetzleri (Wiedenmayer, X X X X X 1977) Siphonodictyon brevitubulatum X X X X Pang, 1973 Siphonodictyon coralliphagum X X X X X X X X X X X X X X X X X X X X X X Rützler, 1971 Siphonodictyon sp. Cu-01 X X X X X X Smenospongia aurea (Hyatt, 1875) X X X X X X X Smenospongia conulosa X X Pulitzer-Finali, 1986 Smenospongia echina (de X X X X X X Laubenfels, 1934) Smenospongia sp. X X Spheciospongia vesparium X X (Lamarck, 1815) Spirastrella coccinea (Duchassaing X X X X X X X X X X X X X X X X X X X X X X X X & Michelotti, 1864) Spirastrella hartmani Boury- Esnault, Klautau, Bézac, Wulff & X X X X X X X X X X X X X X X X X X X X X X X Solé-Cava, 1999 REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 110 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Spirastrella sp. Cu-01 X X X X X Spirastrella sp. Cu-02 X Spirastrellidae unid. sp. X X X X X X X X X X X X X X X X X Spongia sp. X Spongia sp. Cu-01 X Spongia sp. Cu-02 X Stylissa caribica Lehnert & van X Soest, 1998 Stylissa sp. X X X X Svenzea zeai (Alvarez, van Soest X X X X X X X X X X X X X X X X X X X X X X & Rützler, 1998) Terpios belindae Rützler & X X X X Smith, 1993 Terpios cf. belindae Rützler & X Smith, 1993 Tetillidae unid. sp. X Tetractinellida X X Tetractinellida Cu-01 X X X X X Tetractinellida Cu-02 X Tetractinellida Cu-03 X Tetractinellida Cu-04 X Tetractinellida Cu-05 X Tetractinellida Cu-06 X X Theonella atlantica van Soest & X X Stentoft, 1988 Topsentia sp. X X X X X Vansoestia caribensis Díaz, Thacker, Redmond, Pérez & X Collins, 2015 Verongiida Cu-01 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 Verongiida Cu-02 X Verongiida Cu-03 X Verongiida Cu-04 X X X Verongiida Cu-05 X X X X X X X X X X X X X X X X X X X X Verongiida Cu-06 X X X Verongiida Cu-07 X

REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 111 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Verongiida Cu-08 X X X X X Verongiida Cu-09 X Verongula cf. rigida (Esper, 1794) X X X X X Verongula gigantea (Hyatt, 1875) X X X X X X X X X X X X X X X Verongula reiswigi Alcolado, 1984 X X X X X X X X X X X X (Esper, 1794) X X X X X X X X X X X X X X X X X Verongula sp. Cu-01 X X X X X Verongula sp. Cu-02 X X Xestospongia deweerdtae X X X X X Lehnert & van Soest, 1999 Xestospongia muta (Schmidt, 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 1870) Xestospongia purpurea Rützler, X X X X X X X X Piantoni, van Soest & Díaz, 2014 Xestospongia sp. Cu-01 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 Xestospongia sp. Cu-02 X Xestospongia sp. Cu-03 X X Homoscleromorpha Homosclerophorida unid. sp. X X X Oscarella sp. Cu-01 X X Oscarella sp. Cu-02 X X Oscarellidae unid. sp. X X Plakinastrella onkodes Uliczka, X 1929 Plakinidae unid. sp. X X Plakortis dariae Ereskovsky, X X X Lavrov & Willenz, 2014 Plakortis sp. Cu-01 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 Plakortis sp. Cu-02 X Cnidaria Hydrozoa Hydroidolina 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 Millepora alcicornis Linnaeus, X X X X X X X X X X X X X X 1758 Stylasteridae 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 - non coral

REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 112 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Actiniaria X X X X X X Condylactis gigantea (Weinland, X X X 1860) Lebrunia neglecta Duchassaing X & Michelotti, 1860 Palythoa grandis (Verrill, 1900) X Zoanthidae X X Alcyonacea - Alcyoniina Chironephthya caribaea X X X X X X X X X X X X X X X X X X X X X X (Deichmann, 1936) Nidalia sp. X Alcyonacea - gorgonian Acanthogorgia sp. X Bebryce sp. X X X X Briareum sp. X Callogorgia gracilis (Milne Edwards & Haime, 1857) X X X Ellisella barbadensis (Duchassaing & Michelotti, X X X X X X X X 1864) syn. elongata (Pallas, 1766) Ellisella elongata (Pallas, 1766) X X X Ellisella sp. 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 Eunicea sp. X X X X X X X X X X X X X Gorgonia mariae Bayer, 1961 X Gorgonia ventalina Linnaeus, 1758 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Hypnogorgia sp. 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 Duchassaing, 1870 Lignella richardi (Lamouroux, 1816) X Muricea sp. X X X X Nicella goreaui Bayer, 1973 X X X X Nicella sp. 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 Paramuricea sp. X X X X X X X Paramuriceidae X Placogorgia sp. X X

REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 113 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Plexaura sp. X X X Plexaurella sp. X X X X X X X X Plexauridae 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 Primnoidae X X X X Pseudoplexaura sp. X X Pseudopterogorgia (syn. Antillogorgia) bipinnata (Verrill, X 1864) Pseudopterogorgia sp. 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 Pterogorgia sp. X Swiftia exserta (Ellis & Solander, 1786) X X X X X X X X X X X X X X X X X X X X X X X X Villogorgia sp. X Antipatharia Antipatharia unid. sp. X X Antipathes atlantica Gray, 1857 X X X X X X X X X X Antipathes furcata Gray, 1857 X X X X X X Antipathes sp. X X X X X X X X X X Antipathidae 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 Aphanipathes sp. X X X Elatopathes abietina (Pourtalès, 1874) X X X Elatopathes sp. X X Leiopathes sp. X Plumapathes pennacea (Pallas, X X X X X X 1766) Plumapathes sp. X X X Stichopathes lutkeni Brook, 1889 X X Stichopathes sp. 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 Tanacetipathes sp. 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 Tanacetipathes tanacetum X X X X X X X X X X X (Pourtalès, 1880) Scleractinia Acropora cervicornis (Lamarck, 1816) X X X X X X Agaricia agaricites (Linnaeus, 1758) X X X X X X X X X X X X X X X X X X REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 114 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Agaricia grahamae Wells, 1973 X X Agaricia lamarcki Milne Edwards X X X X X X X & Haime, 1851 Agaricia sp. 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 Colpophyllia natans (Houttuyn, X X X X X X X X X X X X X X X X X 1772) Dendrogyra cylindrus Ehrenberg, X 1834 Dichocoenia stokesii Milne X Edwards & Haime, 1848 Diploria labyrinthiformis (Linnaeus, 1758) X X X Eusmilia fastigiata (Pallas, 1766) X X X X X X X X X Helioseris cucullata (Ellis & X X X X X X X X X X X X X Solander, 1786) Isophyllia rigida (Dana, 1848) X X Isophyllia sinuosa (Ellis & X X X X X Solander, 1786) Isophyllia sp. X Madracis asperula Milne X X X Edwards & Haime, 1849 Madracis auretenra Locke, Weil & Coates, 2007 X X X Madracis decactis (Lyman, 1859) X X X Madracis formosa Wells, 1973 X X X X X X X X X X Madracis myriaster (Milne Edwards & Haime, 1850) X Madracis sp. X X X X X X X X X X Madrepora sp. X X X X Meandrina meandrites X X X X X X X X X X X X X X X X X X X X X (Linnaeus, 1758) Meandrina sp. X Montastraea cavernosa 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 (Linnaeus, 1767) Mussa angulosa (Pallas, 1766) X X X X Mycetophyllia aliciae Wells, 1973 X Mycetophyllia lamarkiana Milne X X Edwards & Haime, 1848 REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 115 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Mycetophyllia sp. X X X X X X X X X X X X X X X X Orbicella annularis (Ellis & X X X X X Solander, 1786) Orbicella faveolata (Ellis & X X X X X X X X X X X X X X X X X X X X X X X X Solander, 1786) Orbicella franksi (Gregory, 1895) X X X X X X X Oxysmilia rotundifolia (Milne X Edwards & Haime, 1848) Porites astreoides Lamarck, 1816 X X X X X X X X X X X X X X X X X X X X X Porites divaricata Le Sueur, 1820 X X X X Porites furcata Lamarck, 1816 X X X X X Porites porites (Pallas, 1766) X X X X X X X Porites sp. X X X X X X X X X X Pseudodiploria strigosa (Dana, X X X X X X X 1846) Scleractinia- unid colonial X X X X Scleractinia- unid cup X X X X X X X X X X X X X X X Scolymia cubensis (Milne X X X X X X X X X X X X X X X X X X X X Edwards & Haime, 1848) Scolymia lacera (Pallas, 1766) X Scolymia sp. X X X X X X X X Siderastrea siderea (Ellis & X X X X X X X X X X X X X X X X X Solander, 1768) Solenastrea bournoni Milne X X X X X X X X X X X Edwards & Haime, 1849 Stephanocoenia intersepta X X X X X X X X X X X X X X X X X X X X X X X (Lamarck, 1836) Other Annelida Filograna sp. X Polychaeta X Entemnotrochus adansonianus X X (Crosse & P. Fischer, 1861) Gastropoda X Strombus (syn. Lobatus) gigas X Linnaeus, 1758 REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 116 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Scientific Name C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Bryozoa Bryozoa unid. sp. X X X X X Colatooecia serrulata (Smitt, 1 X 873) Arthropoda Majidae X Panulirus argus (Latreille, 1804) X X X Penaeidae X Echinodermata Asteroidea X Comatulida X X X X X Crinoidea X Davidaster discoideus X (Carpenter, 1888) Ophiuroidea X X X X X X Chordata - Ascidiacea X X X X X X X Non-Fauna Human debris Human debris- fishing line X X X X X X Human debris- long line X Human debris- net X Disease Black Band Disease X Bleaching X X X X X X X Dead Coral X X Disease- unid. X

REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 117 Reed • González-Díaz • Busutil • et. al. Appendix 2. Species list of fish identified at Cuba’s mesophotic reefs from ROV dives during the F.G. Walton Smith Cuba cruise, May 14 to June 12, 2017. Dive sites (see Fig. 1) compiled by regions (see Appendix 1 caption).

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Order/ Common C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Scientific Name Name Chordata Actinopterygii Unid Fish 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 Larval Unid Actinopterygii- Larval X Fish Anguilliformes Gymnothorax funebris Green X Ranzani, 1839 Moray Eel Gymnothorax sp. Moray Eel X X Aulopiformes Synodus sp. Lizardfish X X X X Synodus synodus (Linnaeus, Red X X 1758) Lizardfish Holocentrus adscensionis Squirrelfish X X X X X X X X X X X X X (Osbeck, 1765) Holocentrus rufus (Walbaum, Longspine X X X X X X X X X X X X X X X X X 1792) Squirrelfish Holocentrus sp. Squirrelfish X X X X X X X X X X X X X X X X X X X X Myripristis jacobus Cuvier, Blackbar X X X X X 1829 Soldierfish marianus (Cuvier, Longjaw X X X X X X X X X X X X X X X X X X X X 1829) Squirrelfish Plectrypops retrospinis Cardinal X X (Guichenot, 1853) Soldierfish Sargocentron vexillarium Dusky X X (Poey, 1860) Squirrelfish Lophiiformes Ogcocephalus nasutus Shortnose X (Cuvier, 1829) Batfish Abudefduf saxatilis (Linnaeus, Sergeant X X X 1758) Major Acanthurus chirurgus (Bloch, Doctorfish X X X X X X X X X 1787) REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 118 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Order/ Common C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Scientific Name Name Acanthurus coeruleus Bloch & Blue Tang 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 Schneider, 1801 Acanthurus sp. Surgeonfish X Anisotremus surinamensis Black X X (Bloch, 1791) Margate Anisotremus virginicus Porkfish X X X X X X X X X (Linnaeus, 1758) Apogon pseudomaculatus Twospot X Longley, 1932 Cardinalfish Apogon sp. Cardinalfish X X X dentatus Guichenot, Black X X 1853 Snapper Bodianus pulchellus (Poey, Spotfin X X X X X X X 1860) Hogfish Bodianus rufus (Linnaeus, Spanish X X X X X X X X X X 1758) Hogfish Calamus calamus Saucereye X (Valenciennes, 1830) Porgy Calamus sp. Porgy X X X X Carangidae Jack X Carangoides bartholomaei Yellow Jack X (Cuvier, 1833) Caranx lugubris Poey, 1860 Black Jack X X X X X X X X X X X X X X X X X X Caranx ruber (Bloch, 1793) Bar Jack 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 Caranx sp. Jack X X X X Centropristis ocyurus (Jordan Bank Sea X & Evermann, 1887) Bass argi Woods & Cherubfish X X X X X Kanazawa, 1951 Cephalopholis cruentata Graysby X X X X X X X X X X X X X X X X X X X X X X X X (Lacepède, 1802) Cephalopholis fulva (Linnaeus, Coney X X X X X X X X X X X 1758) Chaetodon capistratus Foureye 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 Linnaeus, 1758 Chaetodon ocellatus Bloch, Spotfin X X X X X X X X X X X X X 1787 Butterflyfish Chaetodon sedentarius Poey, Reef X X X X X X X X X X X X 1860 Butterflyfish

REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 119 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Order/ Common C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Scientific Name Name Chaetodon striatus Linnaeus, Banded X X X X X X X 1758 Butterflyfish Chaetodontidae Butterflyfish X X X X X X X X X X X X X Chloroscombrus chrysurus Atlantic X (Linnaeus, 1766) Bumper Blue Chromis cyanea (Poey, 1860) 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 Chromis Chromis enchrysura Jordan & Yellowtail X X X X X X Gilbert, 1882 Reeffish Chromis insolata (Cuvier, 1830) Sunshinefish 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 Chromis multilineata Brown X X X X (Guichenot, 1853) Chromis Purple Chromis scotti Emery, 1968 X X X X X X Reeffish Clepticus parrae (Bloch & Creole 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 Schneider, 1801) Wrasse Coryphopterus glaucofraenum Bridled X Gill, 1863 Goby Coryphopterus personatus Masked/ X X X X X X X X X X X (Jordan & Thompson, 1905) Glass Goby Diplectrum formosum Sand Perch X (Linnaeus, 1766) Elacatinus genie (Böhlke & Cleaning X X X X Robins, 1968) Goby Elacatinus horsti (Metzelaar, Yellowline X X 1922) Goby Elacatinus louisae (Böhlke & Spotlight X Robins, 1968) Goby Elacatinus sp. Goby X Elagatis bipinnulata (Quoy & Rainbow X X Gaimard, 1825) Runner Epinephelus guttatus Red Hind X X X X X X X X X X X X X X (Linnaeus, 1758) Epinephelus morio Red X X X X X X (Valenciennes, 1828) Grouper Epinephelus striatus (Bloch, Nassau X X X X X X X X X X 1792) Grouper Equetus lanceolatus Jacknife X X X (Linnaeus, 1758) Fish REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 120 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Order/ Common C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Scientific Name Name Equetus punctatus (Bloch & Spotted X Schneider, 1801) Drum Gobiidae Goby X X X X X X X X X Gobiosoma sp. Goby X X Fairy Gramma loreto Poey, 1868 X X X X X X X X X X X X X X X X X X X X X X X Basslet Gramma melacara Böhlke & Blackcap 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 Randall, 1963 Basslet Haemulon aurolineatum Tomtate X X X X X X Cuvier, 1830 Haemulon carbonarium Poey, Caesar X X X 1860 Grunt Haemulon flavolineatum French X X X X X X X X X X X X X X (Desmarest, 1823) Grunt Haemulon melanurum Cottonwick X X (Linnaeus, 1758) Haemulon parra (Desmarest, Sailors X 1823) Choice Haemulon plumierii White Grunt 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 (Lacepède, 1801) Haemulon sciurus (Shaw, Bluestriped X X X X X X X X X X X X X X 1803) Grunt Haemulon sp. Grunt X X Haemulon striatum (Linnaeus, Striped X X X X X 1758) Grunt Haemulon vittatum (Poey, Boga X X X 1860) Halichoeres bathyphilus Greenband X (Beebe & Tee-Van, 1932) Wrasse Halichoeres bivittatus (Bloch, Slippery X 1791) Dick Halichoeres garnoti Yellowhead 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 (Valenciennes, 1839) Wrasse Halichoeres maculipinna Clown X (Müller & Troschel, 1848) Wrasse Halichoeres pictus (Poey, Rainbow X 1860) Wrasse Holacanthus bermudensis Blue X X X X X X X X X X X X X X Goode, 1876 Angelfish REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 121 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Order/ Common C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Scientific Name Name Holacanthus ciliaris (Linnaeus, Queen X X X X X X X X X X X X X X X X 1758) Angelfish Holacanthus sp. Angelfish X X X X X X X X Holacanthus tricolor (Bloch, Rock 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 1795) Beauty guttavarius Shy Hamlet X X (Poey, 1852) (Poey, Indigo X X X 1851) Hamlet Hypoplectrus nigricans (Poey, Black X X X X X X X X X X 1852) Hamlet Hypoplectrus puella (Cuvier, Barred X X X X X X X X X X X X X X X X 1828) Hamlet Hypoplectrus sp. Hamlet X X X X Hypoplectrus unicolor Butter X X X X X X X X (Walbaum, 1792) Hamlet Kyphosus sp. Chub X X X X X X Labridae Wrasse X X X X X X X X X X X X X X X X X X Labrisomus filamentosus Quillfin X Springer, 1960 Blenny Lachnolaimus maximus Hogfish X X X X X X X X X (Walbaum, 1792) Liopropoma mowbrayi Woods Cave X X X X X X X X X X X X X X X X X X X X & Kanazawa, 1951 Basslet Peppermint Liopropoma rubre Poey, 1861 X Bass Liopropoma sp. Basslet X Lucayablennius zingaro Arrow X (Böhlke, 1957) Blenny Mutton Lutjanus analis (Cuvier, 1828) X X X Snapper Lutjanus apodus (Walbaum, School- X X X X X X X X X X X X X X X X X X 1792) master Lutjanus buccanella (Cuvier, Blackfin X X X X X X X X X X X X 1828) Snapper Lutjanus campechanus (Poey, Red X 1860) Snapper Lutjanus cyanopterus (Cuvier, Cubera X 1828) Snapper REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 122 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Order/ Common C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Scientific Name Name Lutjanus griseus (Linnaeus, Gray X X 1758) Snapper Lutjanus jocu (Bloch & Dog X X X X X X X X X Schneider, 1801) Snapper Lutjanus mahogoni (Cuvier, Mahogany X X X X X X X X X 1828) Snapper Lutjanus sp. Snapper 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 Lutjanus vivanus (Cuvier, Silk X 1828) Snapper Malacanthus plumieri (Bloch, Sand X X 1786) Tilefish Microspathodon chrysurus Yellowtail X X X (Cuvier, 1830) Damselfish Mulloidichthys martinicus Yellow X X X X X X X X X X (Cuvier, 1829) Goatfish Mycteroperca bonaci (Poey, Black X X X X X X X X X 1860) Grouper Mycteroperca interstitialis Yellowmouth X X (Poey, 1860) Grouper Mycteroperca phenax Jordan Scamp X X X X X X X & Swain, 1884 Mycteroperca tigris Tiger X X X X X X X X X X (Valenciennes, 1833) Grouper Mycteroperca venenosa Yellowfin X X X X X (Linnaeus, 1758) Grouper Ocyurus chrysurus (Bloch, Yellowtail 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 1791) Snapper Pagrus sp. Porgy X Paranthias furcifer Atlantic X (Valenciennes, 1828) Creolefish Pomacanthus arcuatus Gray X X X X X X X X X X X X X X X X X X X X X X X X (Linnaeus, 1758) Angelfish Pomacanthus paru (Bloch, French X X X X X X X X X 1787) Angelfish Pomacentrus sp. Damselfish X X X X X X X X X X X X X Short alta (Gill, 1862) X X Bigeye aculeatus Longsnout X X X X X X X X X X X X X X X X X X X X X (Poey, 1860) Butterflyfish REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 123 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Order/ Common C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Scientific Name Name Prognathodes aya (Jordan, Bank X X X 1886) Butterflyfish Prognathodes guyanensis French X (Durand, 1960) Butterflyfish Pronotogrammus martinicensis Rough- X X (Guichenot, 1868) tongue Bass Pseudupeneus maculatus Spotted X X X X X X X X X X X X X X (Bloch, 1793) Goatfish Ptereleotris calliura (Jordan & Blue X Gilbert, 1882) Dartfish Ptereleotris helenae (Randall, Hovering X 1968) Dartfish Rypticus maculatus Holbrook, Whitespotted X 1855 Soapfish Rypticus saponaceus (Bloch & Greater X Schneider, 1801) Soapfish Scaridae Parrotfish X X X X X X X X X X X X X X X X X X X X X X X X Scarus coelestinus Midnight X Valenciennes, 1840 Parrotfish Striped Scarus iseri (Bloch, 1789) 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 Parrotfish Scarus taeniopterus Lesson, Princess X X X X X X X X X X X 1829 Parrotfish Scarus vetula Bloch & Queen X Schneider, 1801 Parrotfish Scomberomorus regalis Cero X X X X X X (Bloch, 1793) Scomberomorus sp. Mackeral X Seriola sp. Amberjack X Serranidae Grouper X X X X X X X X X X X X X X Serranus annularis (Günther, Orangeback X X X X X X 1880) Bass Serranus notospilus Longley, Saddle X 1935 Bass Serranus phoebe Poey, 1851 Tattler X X X X X X Serranus sp. Sea Bass X X X Serranus tabacarius (Cuvier, Tobaccofish X X X X 1829)

REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 124 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Order/ Common C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Scientific Name Name Serranus tigrinus (Bloch, Harlequin X X X X X X X X 1790) Bass Serranus tortugarum Longley, Chalk Bass X X X X 1935 Sparisoma atomarium (Poey, Greenblotch X X X 1861) Parrotfish Sparisoma aurofrenatum Redband X X X X X X X X X X X X X X X (Valenciennes, 1840) Parrotfish Sparisoma rubripinne Yellowtail X (Valenciennes, 1840) Parrotfish Sparisoma viride (Bonnaterre, Stoplight X X X X X X X X X X X X X X X X X X X X 1788) Parrotfish Sphyraena barracuda Great 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 (Edwards, 1771) Barracuda Stegastes adustus (Troschel, Dusky X X X X X X 1865) Damselfish Stegastes diencaeus (Jordan Longfin X & Rutter, 1897) Damselfish Stegastes leucostictus (Müller Beaugregory X X X X X X X X X & Troschel, 1848) Stegastes partitus (Poey, Bicolor 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 1868) Damselfish Stegastes planifrons (Cuvier, Threespot X X 1830) Damselfish Thalassoma bifasciatum Bluehead X X X X X X X X X X X X X X X X X X X X X X (Bloch, 1791) Wrasse Trachinotus blochii (Lacepède, Permit X X 1801) Trachinotus goodei Jordan & Palometa X Evermann, 1896 Scorpaeniformes Pterois volitans (Linnaeus, Lionfish 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 1758) Syngnathiformes Aulostomus maculatus Atlantic X Valenciennes, 1841 Trumpetfish Acanthostracion polygonius Honeycomb X Poey, 1876 Cowfish REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 125 Reed • González-Díaz • Busutil • et. al.

Northwest Coast West Coast Southwest Coast Southeast Coast Northeast Coast Phylum/Class/Order/ Common C-04 C-04A C-06 C-07 C-10 C-10A C-12A C-12B C-15 C-16A C-21 C-21A C-23 C-24 C-28 C-29 C-31A C-33A C-38 C-38A C-41 C-48 C-50 C-52 C-52A C-53A C-53B C-54 C-54B C-56 C-58 C-58A C-59A C-59C C-60A Scientific Name Name (Osbeck, Scrawled X 1765) Queen Balistes vetula Linnaeus, 1758 X X X X X X X Triggerfish Balistidae Triggerfish X X X X X macrocerus Whitespot- X (Hollard, 1853) ted Filefish Canthidermis sufflamen Ocean X X X X X X X X X X X X X X X X (Mitchill, 1815) Triggerfish Canthigaster rostrata (Bloch, Sharpnose X X X X X X X X X X X X X X X X X X X X X X X 1786) Puffer Lactophrys triqueter Smooth X X (Linnaeus, 1758) Trunkfish Melichthys niger (Bloch, Black X X X X X X X X X X 1786) Durgon Sphoeroides spengleri (Bloch, Bandtail X X 1785) Puffer ringens Sargassum X X X X X X X X X X X X X X X X X X Linnaeus, 1758) Triggerfish Elasmobranchii Carcharhiniformes Carcharhinidae Shark X Carcharhinus leucas (Müller & Bull Shark X Henle, 1839) Myliobatiformes Dasyatis americana Southern X X Hildebrand & Schroeder, 1928 Stingray Orectolobiformes Ginglymostoma cirratum Nurse X X X (Bonnaterre, 1788) Shark Reptilia Testudines Caretta caretta (Linnaeus, Loggerhead X X 1758) Turtle Eretmochelys imbricata Hawksbill X (Linnaeus, 1766) Sea Turtle

REVISTA INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 126 Reed • González-Díaz • Busutil • et. al. Appendix 3-1. Images from ROV dives to illustrate various regions of Cuba’s MCEs. Western region: A. Site C-04, 49 m, Agaricia spp., octocorals (Antillogorgia spp., Ellisella spp.), black coral- (Stichopathes sp.), sponges (tan finger- Agelas sceptrum, vase- Niphates digitalis); B. Site C-07, 112 m, Lionfish, sclerosponge (Ceratoporella nicholso- ni); C. Site C-12B, 66 m, Agaricia sp, octocoral (Icilogorgia schrammi), algae (Halimeda sp.), sponge (branching- Agelas sceptrum, encrusting tan- Spirastrella hartmani, encrusting red- S. coccinea); D. Site C-10, 35 m, large Xestospongia muta; E. Site C-10A, 104 m, algae (crustose coralline algae, Halimeda copiosa), sponges (spikey- Oceanapia bartschi, tan with zoanthids- Petrosida, thin rope- Agelas repens, encrusting yellow- Calcarea, tan sphere-Geodia cribata, yellow tubes- Verongula rigida); F. Site C-10A, 38 m, Orbicella faveolata, Montastraea cavernosa.

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 127 Reed • González-Díaz • Busutil • et. al. Appendix 3-2. Images from ROV dives to illustrate various regions of Cuba’s MCEs. Southern region: A. Site C-21, 32 m, Orbicella faveolata, sponges (orange lumpy-Ectyoplasia ferox, large sphere- unid. demospon- ge), algae (Halimeda copiosa), fish (Blackcap Basslet, Gramma melacara); B. Site C-24, 73 m, Agaricia lamarcki, sponges (purple tube- Aplysina acheri, plate- Agelas dilatata, sphere- Geodia sp.); C. Site C-29, 69 m, Agaricia lamarcki, sponge (tube- Aplysina archeri); D. Site C-38A, 108 m. sponges (elkhorn- Agelas cervicornis, tan fingers- Agelas sceptrum, black- Asteropus sp., purple lobate- Aiolochroia crassa, yellow encrusting- Siphonodictyon coralliophagum; E. Site C-41, 70 m, Agaricia sp., sponges (tan rope- A. sceptrum, red orange finger- Amphimedon compressa, orange rope- Aplysina cauliformis, A. fulva, orange branching- Agelas cervicornis, purple lobate- A. crassa); F. Site C-41, 142 m, scalloped rock, deep island slope.

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 128 Reed • González-Díaz • Busutil • et. al. Appendix 3-3. Images from ROV dives to illustrate various regions of Cuba’s MCEs. Eastern region: A. Site C-48, depth 96 m, eroded wall, crustose coralline algae, sponges (fan- Agelas? sp., yellow encrusting- Verongiida sp.); B. Site C-50, 78 m, octocoral (Nicella goreaui); C. Site C-52, 105 m, sponges (vase- Xestospongia sp. Cu-01, fan- Agelas clathrodes), fish (Longspine Squirrelfish, Holocentrus rufus); D. Site C-52, 81 m, sponges (large orange- unid. , fan- Cribrochalina vasculum, yellow- Agelas cervicornis, sphere- Cinachyrella sp., tan lobate- Smenospongia conulosa); E. Site C-50, 92 m, sponges (yellow- undescribed Verongiida, tan vase- Xestospongia sp. 1, yellow sphere- Cinachyrella sp.); F. Site C-50, 70 m; large Agaricia lamarcki (lasers 10 cm), sponge (encrusting- Spirastrella coccinea).

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129 Cuba’s Mesophotic Reefs 129 Reed • González-Díaz • Busutil • et. al. Appendix 3-4. Images from ROV dives to illustrate various regions of Cuba’s MCEs. Northern region: A. Site C-56, 67 m, school of Silversides or Anchovies at mouth of cave; B. Site C-58, 44 m, algae (Halimeda spp., Udotea sp.); C. Site C-58A, 67 m, school of Dog Snapper (Lutjanus jocu); D. Site C-59A, 65 m, longline wrapped on wall; E. Site C-56, 88 m, overhang of buttress on wall; F. Site C-60A, 44 m, shelf-edge reef crest with dense octocorals and sponges.

REVISTA DE INVESTIGACIONES MARINAS RNPS: 2096 • ISSN: 1991-6086• VOL. 38 • No. 1 • ENERO-JUNIO • 2018 • pp. 60-129