Coral Reefs

, Volume 33, Issue 2, pp 313–328 | Cite as

Fishes associated with mesophotic coral ecosystems in La Parguera, Puerto Rico

  • I. BejaranoEmail author
  • R. S. Appeldoorn
  • M. Nemeth


Fishes associated with mesophotic coral ecosystems (MCEs) of the La Parguera shelf-edge were surveyed between 2007 and 2011 using mixed-gas rebreather diving. Fishes were identified and counted within belt transects and roving surveys at 30, 40, 50, 60 and 70 m depth. Vertical transects from 70 to 30 m depth helped determine depth distribution ranges. One hundred and three species were identified at MCEs (40–70 m), with high abundances and species richness, though both varied greatly among transects. Most species at MCEs were common inhabitants of shallow reefs, but some were restricted to mesophotic depths. An additional 15 species were added to those previously classified as indicator species of mesophotic areas in Puerto Rico. The MCE fish assemblage was distinct from shallow areas (30 m), with taxonomic composition, abundance and the proportion of trophic guilds varying with increasing depth. The dominant trophic guild within MCEs was the zooplanktivores, while herbivores dominated shallow reefs. Both herbivores and zooplanktivores responded strongly, and oppositely, to depth. The few herbivores associated with deep MCEs are small-bodied species. The largest changes within the mesophotic fish community along the depth gradient occurred at 60 m, similar to that reported for algae and corals, and seem to represent both a response to reduced light and variations in herbivory. The presence of commercially important fishes at MCEs, many considered to be threatened by fishing pressure in shallow areas, suggests that MCEs are important for the conservation of these species. This study represents the first quantitative in situ observations and descriptions of fishes inhabiting MCEs at depths of 50–70 m in Puerto Rico and highlights the role of MCEs as valuable habitats for reef fishes. The composition and distribution of the MCEs fish community should be incorporated when planning for the spatial management of coral reef resources.


Fish assemblage Mesophotic coral ecosystems (MCEs) Depth Rebreather Trophic guilds Herbivores 



We thank deep divers Milton Carlo, Hector J. Ruíz and Clark Sherman for assisting in field sampling, and D. Mateos-Molina for his continuous help and support. This work was supported by the National Oceanic and Atmospheric Administration’s Center for Sponsored Coastal Ocean Research (NOAA/CSCOR) (Grant No. NA06NOS4780190), through the Caribbean Coral Reef Institute (CCRI). This project used the facilities of the University of Puerto Rico, and the University of North Carolina—Wilmington’s Undersea Research Center oversaw the training of rebreather divers.


  1. Aguilar-Perera A, Appeldoorn RS (2008) Spatial distribution of marine fishes along a cross-shelf gradient containing a continuum of mangrove-seagrass-coral reefs off Southwestern Puerto Rico. Estuar Coast Shelf Sci 76:378–394CrossRefGoogle Scholar
  2. Aponte NE, Ballantine DL (2001) Algal species distribution and composition as a factor of depth on the deep insular slope at Lee Stocking Island, Bahamas. Deep Sea Res Part I 48:2185–2194CrossRefGoogle Scholar
  3. Ballantine DL, Ruíz H (2010) Two new deepwater Peyssonnelia species, Peyssonnelia iridescens and Peyssonnelia gigaspora (Peyssonneliaceae, Rhodophyta) from Puerto Rico, Caribbean Sea. Phycologia 49:537–544CrossRefGoogle Scholar
  4. Ballantine DL, Ruíz H (2011) A new encrusting deep-water coral reef alga, Peyssonnelia incomposita (Peyssonneliaceae, Rhodophyta), from Puerto Rico, Caribbean Sea. Cryptogam Algol 32:19–26CrossRefGoogle Scholar
  5. Ballantine DL, Ruíz H, Aponte NE (2009) Notes on the benthic marine algae of Puerto Rico IX. Additions to the flora including reports of three species for the first time in the Atlantic Ocean. Bot Mar 52:229–235CrossRefGoogle Scholar
  6. Ballantine DL, Athanasiadis A, Ruíz H (2011) Notes on the benthic marine algae of Puerto Rico. X. Additions to the flora. Bot Mar 54:293–302Google Scholar
  7. Battista TA, Stecher ML (2006) Data acquisition and processing report, NF-06-03, S-I911-NF-06, March 21-April 2, 2006, U.S. Virgin Islands and Puerto Rico, NOAA Ship Nancy Foster 104Google Scholar
  8. Bejarano I, Appeldoorn RA (2013) Seawater turbidity and fish communities on coral reefs of Puerto Rico. Mar Ecol Prog Ser 474:217–226CrossRefGoogle Scholar
  9. Bellwood DR, Hughes TP, Folke C, Nystrom N (2004) Confronting the coral reef crisis. Nature 429:827–833PubMedCrossRefGoogle Scholar
  10. Bridge TCL, Done TJ, Friedman A, Beaman RJ, Williams SB, Pizzaro O, Webster JM (2011) Variability in mesophotic coral reef communities along the Great Barrier Reef, Australia. Mar Ecol Prog Ser 428:63–75CrossRefGoogle Scholar
  11. Brock VE (1954) A preliminary report on a method of estimating reef fish populations. J Wildl Manage 18:297–308CrossRefGoogle Scholar
  12. Brokovich E, Einbinder S, Shashar N, Kiflawi M, Kark S (2008) Descending to the twilight-zone: changes in coral reef fish assemblages along a depth gradient down to 65 m. Mar Ecol Prog Ser 371:253–262CrossRefGoogle Scholar
  13. Brokovich E, Ayalon I, Einbinder S, Segev N, Shaked Y, Genin A, Kark S, Kiflawi M (2010) Grazing pressure on coral reefs decreases across a wide depth gradient in the Gulf of Aqaba, Red Sea. Mar Ecol Prog Ser 399:69–80CrossRefGoogle Scholar
  14. Clements KD, Raubenheimer D, Choat JH (2009) Nutritional ecology of marine herbivorous fishes: ten years on. Funct Ecol 23:79–92CrossRefGoogle Scholar
  15. Colin PL (1974) Observation and collection of deep reef fishes off the coasts of Jamaica and Honduras. Mar Biol 24:29–38CrossRefGoogle Scholar
  16. Colin PL (1976) Observation of deep reef fishes in the Tongue-of-the-Ocean, Bahamas. Bull Mar Sci 26:603–605Google Scholar
  17. Colin PL (1978) Caribbean reef invertebrates and plants: A field guide to the invertebrates and plants occurring on coral reefs of the Caribbean, the Bahamas and Florida. TFH Publications, Neptune City, NJGoogle Scholar
  18. Corgosinho PHC, Schizas NV (2013) Archeolourinia, a new genus and species of Louriniidae Monard, 1927 (Copepoda, Harpacticoida) from a Caribbean mesophotic zone. J Mar Biol Assoc UK 93:651–657CrossRefGoogle Scholar
  19. Duffy JE, Hay ME (1990) Seaweed adaptations to herbivory. Bioscience 40:368–375CrossRefGoogle Scholar
  20. Feitoza BM, Rosa RS, Rocha LA (2005) Ecology and zoogeography of deep-reef fishes in northeastern Brazil. Bull Mar Sci 76:725–742Google Scholar
  21. García-Sais JR (2010) Reef habitats and associated sessile-benthic and fish assemblages across a euphotic–mesophotic depth gradient in Isla Desecheo, Puerto Rico. Coral Reefs 29:277–288CrossRefGoogle Scholar
  22. García-Sais JR, Castro R, Sabater J, Carlo M (2004) Monitoring of coral reef communities from Isla de Vieques, Puerto Rico. Final Report submitted to the Department of Natural and Environmental Resources (DNER), US Coral Reef National Monitoring Program, NOAA.
  23. García-Sais JR, Castro R, Sabater J, Carlo M (2007) Characterization of benthic habitats and associated reef communities at Bajo de Sico Seamount, Mona Passage, Puerto Rico. Final Report submitted to Caribbean Fishery Management Council (CFMC/NOAA)Google Scholar
  24. García-Sais JR, Appeldoorn RS, Battista T, Bauer L, Bruckner A, Caldow C, Carrubba L, Corredor J, Diaz E, Lilyestrom C, García-Moliner G, Hernández-Delgado E, Menza C, Morell J, Pait A, Sabater J, Weil E, Williams E, Williams S (2008) The state of coral reef ecosystems of the commonwealth of Puerto Rico. In: Waddell J, Clarke A (eds) The state of coral reef ecosystems of the United States and Pacific Freely Associated States: 2008 NOAA Technical Memorandum NOS NCCOS 78. NOAA/NCCOS Center for Coastal Monitoring and Assessment’s Biogeography Team, Silver Spring, MD, pp 75–116Google Scholar
  25. Gilmartin M (1960) The ecological distribution of the deep water algae of Eniwetok Atoll. Ecology 41:210–221CrossRefGoogle Scholar
  26. Hanisak MD, Blair SM (1988) Deep-water benthic macroalgal communities with emphasis on Florida and the Bahamas. In: Agegian CR (ed) Biochemical cycling and fluxes between the deep euphotic zone and other oceanic realms. National Undersea Research Program Research Report 88-1. National Oceanic and Atmospheric Administration (NOAA). Department of Commmerce, Washington, DC, pp 61–83Google Scholar
  27. Hinderstein LM, Marr JCA, Martinez FA, Dowgiallo MJ, Puglise KA, Pyle RL, Zawada DG, Appeldoorn RA (2010) Theme section on “Mesophotic Coral Ecosystems: Characterization, Ecology, and Management”. Coral Reefs 29:247–251CrossRefGoogle Scholar
  28. Itzikowitz M, Haley M, Otis C, Evers D (1991) A reconnaissance of the deeper Jamaican coral reef fish communities. NE Gulf Sci 12:25–34Google Scholar
  29. Ives AR, Carpenter SR (2007) Stability and diversity of ecosystems. Science 317:58–62PubMedCrossRefGoogle Scholar
  30. Kahng SE, García-Sais JR, Spalding HL, Brokovich E, Wagner D, Weil E, Hinderstein L, Toonen RJ (2010) Community ecology of mesophotic coral reef ecosystems. Coral Reefs 29:255–275CrossRefGoogle Scholar
  31. Leichter JJ, Genovese SJ (2006) Intermittent upwelling and subsidized growth of the scleractinian coral Madracis mirabilis on the deep fore-reef slope of Discovery Bay, Jamaica. Mar Ecol Prog Ser 316:95–103CrossRefGoogle Scholar
  32. Leichter JJ, Stewart HL, Miller SL (2003) Episodic nutrient transport to Florida coral reefs. Limnol Oceanogr 48:1394–1407CrossRefGoogle Scholar
  33. Leichter JJ, Stokes MD, Genovese SJ (2008) Deep water macroalgal communities adjacent to the Florida Keys reef tract. Mar Ecol Prog Ser 356:123–138CrossRefGoogle Scholar
  34. Leichter JJ, Wing SR, Miller SL, Denny MW (1996) Pulsed delivery of subthermocline water to Conch Reef (Florida Keys) by internal tidal bores. Limnol Oceanogr 41:1490–1501CrossRefGoogle Scholar
  35. Lesser MP (2006) Benthic–pelagic coupling on coral reefs: feeding and growth of Caribbean sponges. J Exp Mar Biol Ecol 328:277–288CrossRefGoogle Scholar
  36. Lesser MP, Slattery MP (2011) Phase shift to algal dominated communities at mesophotic depths associated with lionfish (Pterois volitans) invasion on a Bahamian coral reef. Biol Invasions 13:1855–1868CrossRefGoogle Scholar
  37. Lesser MP, Slattery MP, Stat M, Ojimi M, Gates R, Grottoli A (2010) Photoacclimatization by the coral Montastraea cavernosa in the mesophotic zone: light, food, and genetics. Ecology 91:990–1003PubMedCrossRefGoogle Scholar
  38. Liddell WD, Ohlhorst SL (1988) Hard substrata community patterns, 1–120 m, North Jamaica. Palaios 3:413–423CrossRefGoogle Scholar
  39. Matos-Caraballo D (2004) Puerto Rico/NMFS cooperative fisheries statistics program. April 2001-March 2004. Final report submitted to the National Marine Fisheries Service by the fisheries research laboratory, Department of Natural and Environmental Resources, San JuanGoogle Scholar
  40. McClain CR, Barry JP (2010) Habitat heterogeneity, biogenic disturbance, and resource availability work in concert to regulate biodiversity in deep submarine canyons. Ecology 91:964–976PubMedCrossRefGoogle Scholar
  41. McCune B, Mefford M (2006). PC-ORD. Multivariate analysis of ecological data. Version 5.0. MjM Software, Gleneden Beach, Oregon, USAGoogle Scholar
  42. Morelock J, Schneiderman N, Bryant W (1977) Shelf reefs, southwestern Puerto Rico. Studies in Geology 4:17–25Google Scholar
  43. Morrison D (1988) Comparing fish and urchin grazing in shallow and deeper coral reef algal communities. Ecology 69:1367–1382CrossRefGoogle Scholar
  44. Nelson JS (2006) Fishes of the world. John Wiley and Sons, Inc. New York. 4th edition, p601Google Scholar
  45. Nelson WR, Appeldoorn RS (1985) A submersible survey of the continental slope of Puerto Rico and the US Virgin Islands, 1–23 October 1985. Cruise report, R/V Seward Johnson, National Marine Fisheries Service, Pascagoula, Mississippi LaboratoriesGoogle Scholar
  46. Nemeth M, Appeldoorn RS (2009) The distribution of herbivorous coral reef fishes within fore-reef habitats: the role of depth, light and rugosity. Caribb J Sci 45:247–253Google Scholar
  47. Pesic V, Chatterjee T, Schizas NV (2012) A new species of Pontarachna (Acari, Hydrachnidia, Pontarachnidea) from a mesophotic coral ecosystem off Vieques Island, Puerto Rico, Caribbean Sea. Zootaxa 3440:63–67Google Scholar
  48. Petrescu I, Chatterjee T, Schizas NV (2012) New genus and new species of Cumaceans (Crustacea: Peracarida: Cumacea) from the mesophotic coral ecosystem of SW Puerto Rico, Caribbean Sea. Zootaxa 3476:55–61Google Scholar
  49. Pyle RL (2000) Assessing undiscovered fish biodiversity on deep coral reefs using advanced self-contained diving technology. Mar Tech Soc J 34:82–91CrossRefGoogle Scholar
  50. Pyle RL, Earle JL, Greene BD (2008) Five new species of the damselfish genus Chromis (Perciformes: Labroidei: Pomacentridae) from deep coral reefs in the tropical western Pacific. Zootaxa 1671:3–31Google Scholar
  51. Randall JE (1967) Food habits of reef fishes of the West Indies. Stud Trop Oceanogr 5:665–847Google Scholar
  52. Riegl B, Piller WE (2003) Possible refugia for reefs in times of environmental stress. Int J Earth Sci 92:520–531CrossRefGoogle Scholar
  53. Rooney J, Donham E, Montogmery A, Spalding H, Parrish F, Boland R, Fenner D, Gove J, Vetter O (2010) Mesophotic coral ecosystems in the Hawaiian Archipelago. Coral Reefs 29:361–367CrossRefGoogle Scholar
  54. Runcie LW, Gurgel CFD, Mcdermid KJ (2008) In situ photosynthetic rates of tropical marine macroalgae at their lower depth limit. Eur J Phycol 43:377–388CrossRefGoogle Scholar
  55. Russ GR (2003) Grazer biomass correlates more strongly with production than biomass of algal turfs on coral reefs. Coral Reefs 22:63–67Google Scholar
  56. Sherman C, Nemeth M, Ruíz H, Bejarano I, Appeldoorn RS, Pagan F, Sharer M, Weil E (2010) Geomorphology and benthic cover of mesophotic coral ecosystems of the upper insular slope of southwest Puerto Rico. Coral Reefs 29:347–360CrossRefGoogle Scholar
  57. Thresher RE, Colin PL (1986) Trophic structure, diversity and abundance of fishes of the deep reef (30–300 m) at Enewetak, Marshall Islands. Bull Mar Sci 38:253–272Google Scholar
  58. Van den Hoek C, Breeman AM, Bak RPM, van Buurt G (1978) The distribution of algae, corals, and gorgonians in relation to depth, light attenuation, water movement and grazing pressure in the fringing reef of Curacao, Netherlands Antilles. Aquat Bot 5:1–46CrossRefGoogle Scholar
  59. Walker BH, Salt D (2006) Resilience thinking: Sustaining ecosystems and people in a changing world. Island Press. Washington, D.C., USAGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2014

Authors and Affiliations

  1. 1.Department of Marine SciencesUniversity of Puerto RicoMayagüezUSA

Personalised recommendations