Coral Reefs

, Volume 29, Issue 2, pp 247–251 | Cite as

Theme section on “Mesophotic Coral Ecosystems: Characterization, Ecology, and Management”

  • L. M. HindersteinEmail author
  • J. C. A. Marr
  • F. A. Martinez
  • M. J. Dowgiallo
  • K. A. Puglise
  • R. L. Pyle
  • D. G. Zawada
  • R. Appeldoorn


Mesophotic coral ecosystems (MCEs) are characterized by the presence of light-dependent corals and associated communities that are typically found at depths ranging from 30 to 40 m and extending to over 150 m in tropical and subtropical regions. The dominant communities providing structural habitat in the mesophotic zone can be comprised of coral, sponge, and algal species. Because working in this depth range is constrained by traditional SCUBA limits, less is known about corals and associated organisms there compared to shallower coral communities. Following the first-ever gathering of international scientists to review and discuss existing knowledge of MCEs, this issue focuses on the ecological characterization, geomorphology, and concept of MCEs as refugia for shallow-water populations. The review and research papers comprising this special issue reflect the current scientific understanding of these ecosystems and the underlying mechanisms that regulate them, as well as potential resource management implications. It is important to understand the value and role of mesophotic coral ecosystems in tropical and subtropical regions as these areas face increasing environmental change and human impacts


Mesophotic coral ecosystem Biodiversity Geomorphology Connectivity Community structure Resource management 

Mesophotic coral ecosystem workshop

On 12–15 July 2008, a scientific workshop was held in Jupiter, Florida, to identify critical research and resource management needs for mesophotic coral ecosystems (MCEs) (Puglise et al. 2009). The workshop was hosted by the Perry Institute for Marine Science (PIMS) and organized by two offices of the U.S. National Oceanic and Atmospheric Administration (NOAA): the Center for Sponsored Coastal Ocean Research (CSCOR) and the Office of Ocean Exploration and Research/NOAA’s Undersea Research Program (OER/NURP) and also by the U.S. Geological Survey (USGS). The workshop gathered scientists and managers from governmental and non-governmental organizations for the first time to discuss MCE-related topics. The inception for this workshop grew out of NOAA/CSCOR’s Coral Reef Ecosystems Studies program, which identified the need for further investigations of MCEs.

The goal of the workshop was to review and discuss current knowledge about the characterization (biodiversity, geomorphology, microbiology), ecology (connectivity, community structure/dynamics), and management of MCEs. Three primary products resulted from the workshop: (1) this special Theme Section of Coral Reefs, focused on MCEs; (2) a MCEs Research Strategy (Puglise et al. 2009); and (3) a MCE information portal found at (Bongaerts et al. 2008).

The following articles provide a description of the current state of knowledge about the science and management of MCEs. This information is intended to serve as the baseline for further research initiatives that address the characterization, ecology, and management of MCEs.

Introduction to mesophotic coral ecosystems

In the study of marine ecosystems, much attention has been devoted to the ecology of shallow-water (<30 m) coastal communities. Over the last several decades, there also has been a marked increase in the study of deep-sea environments. However, intermediate depths, particularly coastal environments in the 30–150 m depth range, have received relatively little attention.

Coral ecosystems at intermediate depths are by no means new to science; in fact, Darwin (1889) was one of the first to report the existence of hermatypic corals at depths up to 128 m, while Gardiner (1903) and Vaughan (1907) noted unique geomorphic characteristics of corals at intermediate depths. The 1960s and early 1970s saw the beginning of direct observations on deep coral environments and experimentation with new diving technologies (Stark and Stark 1972; Starck and Colin 1978). However, the physiologically imposed depth limits of sampling with conventional SCUBA, and the impracticality and expense of using submersibles in deep coral environments, limited exploration of the deeper portions of zooxanthellate coral communities (Lang 1974; James and Ginsburg 1979; Nelson and Appeldoorn 1985; Reed 1985; Hanisak and Blair 1988; Liddell and Ohlhorst 1988; Aponte and Ballantine 2001). As a consequence, fully two-thirds of the total depth range of zooxanthellate coral environments remain largely unexplored (Pyle 1996, 2000; Feitoza et al. 2005).

Advances in technical diving methods and instrumentation, such as mixed gas diving, rebreathers, and autonomous underwater vehicles, as well as in imaging techniques, e.g., see Gleason et al. (2010), are increasingly providing easier access to study coral ecosystems in the intermediate depth realm (Pyle et al. 2008). Kahng et al. (2010) review the geographic distribution of studies of MCEs and conclude that although some generalizations may be made about community structure and distribution of MCE in the Caribbean, these generalizations cannot be made for the vastly understudied Indo-Pacific regions. In all areas, major gaps of knowledge still exist.

Coral ecosystems in this depth range have sometimes been referred to as the “upper Twilight Zone” (Fricke and Knauer 1986) or “Coral-reef Twilight Zone” (Pyle 1996; Brokovich et al. 2008), or simply “deep coral reefs”. The term “twilight zone” has also been ascribed to a much deeper zone in the open ocean, depths between the euphotic zone and 1,000 m (Buesseler et al. 2007). The term “deep coral reefs” is both technically inaccurate (in the context of the geological definition of “reef”), and “deep coral” is often applied to the clearly distinct cold-water, aphotic deep-sea coral communities, which can occur at much greater depths. To avoid confusion, we follow Ginsburg (2007) in referring to the zone in deeper water where zooxanthellate corals occur as “mesophotic”.

Mesophotic coral ecosystems (MCEs) are characterized by the presence of light-dependent corals and associated communities that are typically found at depths ranging from 30 to 40 m and extending to over 150 m in tropical and subtropical regions (Figs. 1, 2). The dominant communities providing structural habitat in the mesophotic zone can be comprised of coral, sponge, and algal species. The upper boundary not only reflects the limits imposed using traditional SCUBA diving, but also the depths at or below which there is the beginning of a shift in species composition (Liddell and Ohlhorst 1988; Kahng et al. 2010) as Garcia-Sais (2010) describes for fish at Isla Desecheo, Puerto Rico. Depth records for zooxanthellate corals have been documented for Agaricia grahamae at 119 m in the Caribbean (Reed 1985), Leptoseris fragilis at 145 m in the Red Sea (Fricke et al. 1987), Leptoseris sp. at 153 m in Hawaii (Kahng and Maragos 2006), and L. hawaiiensis at 165 m in Johnston Atoll (Maragos and Jokiel 1986). The lower boundary of “over 150 m” reflects the deeper extensions of these documented observations, as well as other unpublished observations in the Pacific and the documented presence of crustose coralline algae at 268 m in the Bahamas (Littler et al. 1985, 1986). This depth boundary is necessarily imprecise due to differences in light penetration, patterns of thermal variation, and various other physical and ecological parameters that vary between and within geographic regions (Kahng et al. 2010; Smith et al. 2010). Furthermore, other processes structuring mesophotic coral reef communities are not well understood, particularly, as Smith et al. (2010) demonstrate the role of disturbance, e.g., disease.
Fig. 1

A diver using rebreather technology explores a highly diverse mesophotic coral ecosystem at 120+ m in Fiji. Photo credit Richard Pyle

Fig. 2

This mesophotic coral ecosystem at 60+ m in Puerto Rico is characterized by an assemblage of corals, algae, and sponges. Note the flattened morphology of the Montastrea coral. Photo credit Hector Ruiz

Considered as extensions of shallower coral reef ecosystems, MCEs are likely to have biological, physical, and chemical connectivity with these reefs and associated communities, as well as unique assemblages, and extensions to deep biota. Although these ecosystems harbor species found in their shallower counterparts, they may also be colonized by a number of depth-restricted species of fishes, invertebrates, and algae, and a lower diversity of corals (Hanisak and Blair 1988; Pyle 2000; Ballantine and Aponte 2003, 2005; Jarrett et al. 2005; Armstrong et al. 2006; Brokovich et al. 2008; Garcia-Sais 2010). To this end, MCEs that serve as refugia for shallow and mid-depth species (Glynn 1996; Armstrong et al. 2006) may warrant special resource management attention and protection to help maintain local and regional biodiversity (Riegl and Piller 2003), However, the review by Bongaerts et al. (2010) demonstrates how little is known about deep reefs and coral reproduction over depth, and thus they propose a list of urgent research priorities to determine the extent to which deep reefs may act as a refuge in the face of global reef decline. In addition, MCEs are thought to serve as spawning grounds and may function as a larval supply for some shallow-water species (Armstrong et al. 2006; Brokovich et al. 2007; García-Sais et al. 2008).

Because MCEs are deep and can occur in remote localities (Kahng et al. 2010; Locker et al. 2010), there is a common assumption that they are less likely to be impacted by anthropogenic (e.g., overfishing, pollution) or natural (e.g., hurricanes, tsunamis, elevated temperatures) disturbances. To the extent to which such assumptions are borne out, MCEs may serve as a reference point for ecosystem condition in comparison with adjacent compromised, shallower coral reefs. For example, Bak et al. (2005) documented a lack of anthropogenically driven declines on reefs at 30–40 m in Curacao and Bonaire in contrast to the reefs at 10–20 m depth.

However, there is reason to suspect that this assumption is not valid for certain MCEs. For example, some fishing industries specifically target the larger predatory fishes that inhabit these depths. A small increase in shallow-water turbidity due to coastal development, watershed runoff, and pollution may have a greater and more devastating impact on MCEs (at the lower limits of photosynthetically viable light levels) than it would on the shallow reefs that are more directly exposed to the disturbance. While hurricanes and tsunamis have a smaller direct impact at greater depths, they may wash limestone rubble down the reef slope, potentially smothering MCEs (Bak et al. 2005).

Moreover, there is some indication that oscillations in sea temperatures, such as those caused by internal waves, cold-water intrusion, and down-welling of warmer waters, may extend to deeper depths and cause depauperate zones, stress, bleaching, and eventually death (Hickey et al. 2000; Smith 2001; Wolanski et al. 2004; Colin 2009; Smith et al. 2010). Although such potential threats are mostly speculative at this time, in certain locations encroaching threats have begun to adversely affect the condition of MCEs (Menza et al. 2007). Coral mortality events and shifting baselines at mesophotic depths have been documented by scientists, and the causes and consequences may be quite different from shallow-water reefs (Bak et al. 2005; Leichter and Genovese 2006; Menza et al. 2007; Smith et al. 2010). A better understanding of these environments is needed and will likely offer potential findings of major interest for conservation and resource management.

To determine where potential MCEs may exist and to ascertain their underlying geomorphology, Locker et al. (2010) describe regional mapping efforts that have been initiated that use a variety of manned and unmanned survey techniques. These have resulted in an increase in knowledge of the geographic extent of MCEs and a glimpse of the community structure of the MCEs found, e.g., in Hawaii, Rooney et al. (2010); and in Tutuila, American Samoa, Bare et al. (2010). Sherman et al. (2010) provide a more detailed analysis of the evolution of the underlying geological structure supporting the extant MCEs in southwestern Puerto Rico, relating their results to Caribbean sea level change models.

The results of this workshop give a glimpse of the complexity of MCEs but also emphasize the large gaps in our knowledge that currently exist. It is important to understand the value and role of mesophotic coral ecosystems in tropical and subtropical regions as these areas face increasing environmental change and human impacts.



NOAA, USGS, and PIMS wish to thank all the individuals and organizations that participated in the workshop and the writing, editing, and reviewing of this collection of manuscripts. Publication of this issue is supported by the offices of NOAA/NCCOS/CSCOR and NOAA/OER/NURP. We hope the time and effort committed to this initiative will result in increased collaboration between countries, institutions, and agencies with interests in understanding more about mesophotic coral ecosystems.


  1. Aponte NE, Ballantine DL (2001) Depth distribution of algal species on the deep insular fore reef at Lee Stocking Island, Bahamas. Deep Sea Res Part I 48:2185–2194CrossRefGoogle Scholar
  2. Armstrong RA, Singh H, Torres J, Nemeth RS, Can A, Roman C, Eustice R, Riggs L, Garcia-Moliner G (2006) Characterizing the deep insular shelf coral reef habitat of the Hind Bank Marine Conservation District (US Virgin Islands) using the Seabed autonomous underwater vehicle. Cont Shelf Res 26:194–205CrossRefGoogle Scholar
  3. Bak RPM, Nieuwland G, Meesters EH (2005) Coral reef crisis in deep and shallow reefs: 30 years of constancy and change in reefs of Curacao and Bonaire. Coral Reefs 24:475–479CrossRefGoogle Scholar
  4. Ballantine DL, Aponte NE (2003) An annotated checklist of deep-reef benthic marine algae from Lee Stocking Island, Bahamas (western Atlantic), I. Chlorophyta and Heterokontophyta. Nova Hedwigia 76:113–127CrossRefGoogle Scholar
  5. Ballantine DL, Aponte NE (2005) An annotated checklist of deep-reef benthic marine algae from Lee Stocking Island, Bahamas (western Atlantic) II. Rhodophyta. Nova Hedwigia 80:147–171CrossRefGoogle Scholar
  6. Bare AY, Grimshaw KL, Rooney JJ, Sabater MG, Fenner D, Carroll B (2010) Mesophotic communities of the insular shelf at Tutuila, American Samoa. Coral Reefs 29 (this issue). doi: 10.1007/s00338-010-0600-y
  7. Bongaerts P, Martinez FA, Hinderstein LM (2008) Mesophotic Coral Ecosystems Database. Center for Sponsored Coastal Ocean Research (CSCOR/NOAA), Perry Institute of Marine Science (PIMS) and the Centre for Marine Studies (CMS/University of Queensland)
  8. Bongaerts P, Ridgeway T, Sampayo EM, Hoegh-Guldberg O (2010) Assessing the ‘deep reef refugia’ hypothesis: focus on Caribbean reefs. Coral Reefs 29 (this issue). doi: 10.1007/s00338-009-0581-x
  9. Brokovich E, Einbinder S, Kark S, Shashar N, Kiflawi M (2007) A deep nursery for juveniles of the zebra angelfish Genicanthus caudovittatus. Environ Biol Fish 80:1–6CrossRefGoogle Scholar
  10. 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
  11. Buesseler KO, Lamborg CH, Boyd PW, Lam PJ, Trull TW, Bidigare RR, Bishop JKB, Casciotti KL, Dehairs F, Elskens M, Honda M, Karl DM, Siegel DA, Silver MW, Steinberg DK, Valdes J, Van Mooy B, Wilson S (2007) Revisiting carbon flux through the ocean’s twilight zone. Science 316:567–570CrossRefPubMedGoogle Scholar
  12. Colin PL (2009) Marine environments of Palau. Coral Reef Research Foundation, Koror, p 410Google Scholar
  13. Darwin C (1889) The structure and distribution of coral reefs. Smith, Elder, & Co., LondonGoogle Scholar
  14. Feitoza BM, Rosa RS, Rocha LA (2005) Ecology and zoogeography of deep reef fishes in northeastern Brazil. Bull Mar Sci 76:725–742Google Scholar
  15. Fricke HW, Knauer B (1986) Diversity and spatial pattern of coral communities in the Red Sea upper twilight zone. Oecologia 71:29–37CrossRefGoogle Scholar
  16. Fricke HW, Vareschi E, Schlichter D (1987) Photoecology of the coral Leptoseris fragilis in the Red Sea twilight zone (an experimental study by submersible). Oecologia 73:371–381CrossRefGoogle Scholar
  17. Garcia-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 (this issue). doi: 10.1007/s00338-009-0582-9
  18. García-Sais J, Appeldoorn R, 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 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, Silver Spring, MD, pp 75–117Google Scholar
  19. Gardiner JS (1903) The Maldive and Laccadive groups, with notes on other coral formations in the Indian Ocean. In: Gardiner JS (ed) The fauna and geography of the Maldive and Laccadive archipelagoes. Cambridge University Press, Cambridge, pp 146–183Google Scholar
  20. Ginsburg R (2007) Mesophotic coral reefs are the frontier of reef exploration and research. In: Proceedings of the 33rd scientific meeting of the Association of Marine Laboratories of the Caribbean (AMLC) 56 (Suppl. 1): xiiGoogle Scholar
  21. Gleason ACR, Gracias N, Lirman D, Gintert BE, Smith TB, Dick MC, Reid RP (2010) Landscape video mosaic from a mesophotic coral ecosystem. Coral Reefs 29 (this issue). doi: 10.1007/s00338-009-0544-2
  22. Glynn PW (1996) Coral reef bleaching: facts, hypotheses and implications. Global Chang Biol 2:495–509CrossRefGoogle Scholar
  23. Hanisak MD, Blair SM (1988) The deep-water macroalgal community of the East Florida continental shelf (USA). Helgol Meeresunters 42:133–163CrossRefGoogle Scholar
  24. Hickey BM, MacCready P, Elliott E, Kachel NB (2000) Dense saline plumes in Exuma Sound, Bahamas. J Geophys Res C 105:11471–11488CrossRefGoogle Scholar
  25. James NP, Ginsburg RN (1979) The seaward margin of the Belize barrier and atoll reefs. Blackwell Scientific Publications, Oxford, p 191Google Scholar
  26. Jarrett BD, Hine AC, Halley RB, Naar DF, Locker SD, Neumann AC, Twichell D, Hu C, Donahue BT, Jaap WC, Palandro D, Ciembronowicz K (2005) Strange bedfellows—a deep-water hermatypic coral reef superimposed on a drowned barrier island; southern Pulley Ridge, SW Florida platform margin. Mar Geol 214:295–307CrossRefGoogle Scholar
  27. Kahng SE, Maragos JE (2006) The deepest zooxanthellate, scleractinian corals in the world? Coral Reefs 25:254CrossRefGoogle Scholar
  28. Kahng SE, Garcia-Sais JR, Spalding HL, Brokovich E, Wagner D, Weil E, Hinderstein L, Toonen RJ (2010) A review of community ecology of mesophotic coral ecosystems. Coral Reefs 29 (this issue). doi: 10.1007/s00338-010-0593-6
  29. Lang JC (1974) Biological zonation at the base of the reef. Am Sci 62:272–281Google Scholar
  30. 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
  31. Liddell WD, Ohlhorst SL (1988) Hard substrata community patterns, 1–120 m, North Jamaica. Palaios 3:413–423CrossRefGoogle Scholar
  32. Littler MM, Littler DS, Blair SM, Norris JN (1985) Deepest known plant life discovered on an uncharted seamount. Science 227:57–59CrossRefPubMedGoogle Scholar
  33. Littler MM, Littler DS, Blair SM, Norris JN (1986) Deep-water plant communities from an uncharted seamount off San Salvador Island, Bahamas: distribution, abundance, and primary productivity. Deep Sea Res 33:881–892CrossRefGoogle Scholar
  34. Locker SD, Armstrong RA, Battista TA, Rooney JJ, Sherman C, Zawada DG (2010) Geomorphology of mesophotic coral ecosystems: current perspectives on morphology, distribution, and mapping strategies. Coral Reefs 29 (this issue). doi: 10.1007/s00338-010-0613-6
  35. Maragos JE, Jokiel PL (1986) Reef corals of Johnston Atoll—one of the World’s most isolated reefs. Coral Reefs 4:141–150CrossRefGoogle Scholar
  36. Menza C, Kendall M, Rogers C, Miller J (2007) A deep reef in deep trouble. Cont Shelf Res 27:2224–2230CrossRefGoogle Scholar
  37. Nelson WR, Appeldoorn RS (1985) Cruise Report R/V Seward Johnson. A submersible survey of the continental slope of Puerto Rico and the U. S. Virgin Islands. NOAA, NMFS, SEFC, Mississippi Laboratories. University of Puerto Rico, Department of Marine Sciences 76Google Scholar
  38. Puglise KA, Hinderstein LM, Marr JCA, Dowgiallo MJ, Martinez FA (2009) Mesophotic coral ecosystems research strategy: International workshop to prioritize research and management needs for Mesophotic Coral Ecosystems. Jupiter, Florida, 12–15 July 2008. Silver Spring, MD: NOAA National Centers for Coastal Ocean Science, Center for Sponsored Coastal Ocean Research, and Office of Ocean Exploration and Research, NOAA Undersea Research Program. NOAA Technical Memorandum NOS NCCOS 98 and OAR OER 2. p 24Google Scholar
  39. Pyle RL (1996) The twilight zone. Nat Hist 105:59–62Google Scholar
  40. 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
  41. 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
  42. Reed JK (1985) Deepest distribution of Atlantic hermatypic corals discovered in the Bahamas. In: Proceedings of the 5th international coral reef symposium, vol 6, pp 249–254Google Scholar
  43. Riegl B, Piller WE (2003) Possible refugia for reefs in times of environmental stress. Int J Earth Sci 92:520–531CrossRefGoogle Scholar
  44. Rooney J, Donham E, Montgomery A, Spalding H, Parrish F, Boland R, Fenner D, Gove J, Vetter O (2010) Mesophotic coral ecosystems in the Hawaiian Archipelago. Coral Reefs 29 (this issue). doi: 10.1007/s00338-010-0596-3
  45. Sherman C, Nemeth M, Ruíz H, Bejarano I, Appeldoorn R, Pagán F, Schärer M, Weil E (2010) Geomorphology and benthic cover of mesophotic coral ecosystems of the upper insular slope of southwest Puerto Rico. Coral Reefs 29 (this issue). doi: 10.1007/s00338-010-0607-4
  46. Smith NP (2001) Weather and hydrographic conditions associated with coral bleaching: Lee stocking Island, Bahamas. Coral Reefs 20:415–422CrossRefGoogle Scholar
  47. Smith TB, Blondeau J, Nemeth RS, Pittman SJ, Calnan JM, Kadison E, Gass J (2010) Benthic structure and cryptic mortality in a Caribbean mesophotic coral reef bank system, the Hind Bank Marine Conservation District, U.S. Virgin Islands. Coral Reefs 29 (this issue). doi: 10.1007/s00338-009-0575-8
  48. Starck WA II, Colin PL (1978) Gramma linki: a new species of grammid fish from the tropical western Atlantic. Bull Mar Sci 28(1):146–152Google Scholar
  49. Starck WA II, Starck JD (1972) Probing the deep reef’s hidden realm. Natl Geogr Mag 142(6):867–886Google Scholar
  50. Vaughan TW (1907) Recent Madreporaria of the Hawaiian Islands and Laysan. Bull US Natl Mus 59:1–427Google Scholar
  51. Wolanski E, Colin PL, Naithani J, Deleersnijder E, Golbuu Y (2004) Large amplitude, leaky, island-generated, internal waves around Palau, Micronesia. Estuar Coast Shelf Sci 60:705–716CrossRefGoogle Scholar

Copyright information

© US Government Employee 2010

Authors and Affiliations

  • L. M. Hinderstein
    • 1
    Email author
  • J. C. A. Marr
    • 2
  • F. A. Martinez
    • 1
  • M. J. Dowgiallo
    • 1
  • K. A. Puglise
    • 3
    • 1
  • R. L. Pyle
    • 4
  • D. G. Zawada
    • 5
  • R. Appeldoorn
    • 6
  1. 1.Center for Sponsored Coastal Ocean ResearchNational Oceanic and Atmospheric AdministrationSilver SpringUSA
  2. 2.Perry Institute for Marine ScienceJupiterUSA
  3. 3.Office of Ocean Exploration and Research/NOAA’s Undersea Research ProgramNational Oceanic and Atmospheric AdministrationSilver SpringUSA
  4. 4.Department of Natural SciencesBishop MuseumHonoluluUSA
  5. 5.Florida Integrated Science Center, U.S. Geological SurveySt. PetersburgUSA
  6. 6.Department of Marine SciencesUniversity of Puerto RicoMayaguezUSA

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