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Highlighted recent publications from the

  Marine Ecosystem Dynamics Laboratory  


  • Mesophotic communities are not immune from coral bleaching, are in danger, and are degrading in the Red Sea.

  • Heat accumulation at mesophotic depths is almost twofold higher than on adjacent shallow reefs.

  • Bleaching events affect the deep-specialist coral populations, with ~50% coral community mortality at 60 m depth.

  • The deeper benthic community lost framework builders may result in lower complexity, fewer niches, and loss of biodiversity.

  • Experimental evidence suggests that deep-specialist corals are less tolerant of heat stress and bleach more readily.

 Mesophotic Ecosystems: The Link between Shallow and Deep-Sea Habitats 


  • Mesophotic ecosystems (MEs) are characterized by the presence of light-dependent organisms, found at depths ranging from ~30 to 150 m in temperate, subtropical and tropical regions.

  • Mesophotic communities occasionally create massive reef structures with diverse but characteristic morphologies, which serve as the framework builders of those ecosystems.

  • MEs are physically linked with shallow and deep-sea habitats, and while taxa from both environments share this space, unique and endemic biodiversity is also found. 


  • Light relationships alone are sufficient to capture major ecological features across coral reef depth gradients.

  • Predicting the depths of reef zones in unusual light environments suggests that shallow-water turbid reefs should be considered as mesophotic coral ecosystems.

  • Use our App to identify your mesophotic zone boundaries: Light-Driven Model

  • See also other publications: Tamir et al. 2019; Pérez-Castro et al. 2022


  • Fluorescence is highly prevalent in reef-building corals, nevertheless, its biological role is still under ongoing debate.

  • We show that plankton exhibits preferential swimming towards green fluorescent cues and that compared to other morphs, higher predation rates are recorded in a green fluorescing morph of mesophotic corals.

  • See also other publications: Eyal et al. 2015; Ben-Zvi et al. 2015; Ben-Zvi et al. 2021

Full Publication List

Research publications (peer-reviewed):

  1. Grinham A, et al. (2024) Short and long-term impacts of a major flood on fine sediment and nitrogen loading to a large coastal embayment. Science of the Total Environment 918:170646. DOI: 10.1016/j.scitotenv.2024.170646

  2. Wiedenmann J, D'Angelo C, Mardones Velozo ML, Moore S, Benkwitt CE, Graham NAJ, Hambach B, Wilson PA, Vanstone J, Eyal G, Ben-Zvi O, Loya Y, Genin A (2023) Reef-building corals farm and feed on their photosynthetic symbionts. Nature 620:1018-1024. DOI: 10.1038/s41586-023-06442-5

  3. Gijsbers, J.C., Englebert, N., Prata, K.E., Pichon, M., Dinesen, Z., Brunner, R., Eyal, G., González-Zapata, F.L., Kahng, S.E., Latijnhouwers, K.R.W., Muir, P., Radice, V.Z., Sánchez, J.A., Vermeij, M.J.A., Hoegh-Guldberg, O., Jacobs, S.J., Bongaerts, P. (2023) Global phylogenomic assessment of Leptoseris and Agaricia reveals substantial undescribed diversity at mesophotic depths. BMC Biology 21, 147. DOI: 10.1186/s12915-023-01630-1

  4. Grinblat, M., Eyal-Shaham, L., Eyal, G., Ben-Zvi, O., Harii, S., Morita, M., Sakai, K., Hirose, M., Miller, D.J., Loya, Y. (2023) Energy allocation trade-offs as a function of age in fungiid corals. Frontiers in Marine Science, 10, 1113987. DOI: 10.3389/fmars.2023.1113987

  5. Pérez-Castro, M.A., Eyal, G., Leyte-Morales, G.E., Hinojosa-Arango, G., Enríquez S. (2023) Benthic Characterization of Mesophotic Communities Based on Optical Depths in the Southern Mexican Pacific Coast (Oaxaca). Diversity 15, 531. DOI: 10.3390/d15040531

  6. Brown, K.T., Eyal, G., Dove, S.G., and Barott, K.L. (2023). Fine-scale heterogeneity reveals disproportionate thermal stress and coral mortality in thermally variable reef habitats during a marine heatwave. Coral Reefs, 42:131-142. DOI: 10.1007/s00338-022-02328-6

  7. Rinsky, M., Weizman, E., Ben-Asher, H.W., Eyal, G., Zhu, B., and Levy, O. (2022). Temporal gene expression patterns in the coral Euphyllia paradivisa reveal the complexity of biological clocks in the cnidarian-algal symbiosis. Science Advances 8, eabo6467. DOI: 10.1126/sciadv.abo6467

  8. Hammerman, N.M., Roff, G., Lybolt, T., Eyal, G., and Pandolfi, J.M. (2022). Unravelling Moreton Bay reef history: An urban high-latitude setting for coral development. Frontiers in Ecology and Evolution 10, 884850. DOI: 10.3389/fevo.2022.884850

  9. Eyal, G., Laverick, J.H., Ben-Zvi, O., Brown, K.T., Kramer, N., Tamir, R., Lindemann, Y., Levy, O., and Pandolfi, J.M. (2022). Selective deep water coral bleaching occurs through depth isolation. Science of the Total Environment 844, 157180. DOI: 10.1016/j.scitotenv.2022.157180

  10. Ben-Zvi, O., Lindemann, Y., Eyal, G., and Loya, Y. (2022). Coral fluorescence: a prey-lure in deep habitats. Communications Biology 5, 1-8. DOI: 10.1038/s42003-022-03460-3

  11. Kramer, N., Eyal, G., Tamir, R., and Loya, Y. (2022). Growth and survival dynamics of mesophotic coral juveniles in shallow reefs. Marine Ecology Progress Series 682, 237-242. DOI: 10.3354/meps13956

  12. Hammerman, N.M., Roff, G., Rodriguez-Ramirez, A., Leonard, N., Staples, T.L., Eyal, G., Rossbach, S., Havlik, M.N., Saderne, V., and Zhao, J.-x. (2022). Reef accumulation is decoupled from recent degradation in the central and southern Red Sea. Science of the Total Environment 809, 151176. DOI: 10.1016/j.scitotenv.2021.151176

  13. Baird, A.H., Edwards, A.J., Guest, J.R., Harii, S., Hatta, M., Lachs, L., Mera, H., Sinniger, F., Abrego, D., Ben-Zvi, O., et al. (2022). A coral spawning calendar for Sesoko Station, Okinawa, Japan. Galaxea, Journal of Coral Reef Studies 24, 41-49. DOI: 10.3755/galaxea.G2021_S10O

  14. Yuasa, H., Kajitani, R., Nakamura, Y., Takahashi, K., Okuno, M., Kobayashi, F., Shinoda, T., Toyoda, A., Suzuki, Y., Thongtham, N., et al. (2021). Elucidation of the speciation history of three sister species of crown-of-thorns starfish (Acanthaster spp.) based on genomic analysis. DNA Research 28, dsab012. DOI: 10.1093/dnares/dsab012

  15. Eyal, G., Eyal-Shaham, L., and Loya, Y. (2021). Symbiodiniaceae conduct under natural bleaching stress during advanced gametogenesis stages of a mesophotic coral. Coral Reefs 40, 959-964. DOI: 10.1007/s00338-021-02082-1

  16. Bongaerts, P., Cooke, I.R., Ying, H., Wels, D., den Haan, S., Hernandez-Agreda, A., Brunner, C.A., Dove, S., Englebert, N., Eyal, G., et al. (2021). Morphological stasis masks ecologically divergent coral species on tropical reefs. Current Biology 31, 2286-2298. e2288. DOI: 10.1016/j.cub.2021.03.028

  17. Ben-Zvi, O., Wangpraseurt, D., Bronstein, O., Eyal, G., and Loya, Y. (2021). Photosynthesis and bio-optical properties of fluorescent mesophotic corals. Frontiers in Marine Science, 8, 651601. DOI: 10.3389/fmars.2021.651601

  18. Eyal, G., Laverick, J.H., Bongaerts, P., Levy, O., and Pandolfi, J.M. (2021). Mesophotic coral ecosystems of the Great Barrier Reef are understudied and underexplored. Frontiers in Marine Science, 8, 622856. DOI: 10.3389/fmars.2021.622856

  19. Yuval, M., Alonso, I., Eyal, G., Tchernov, D., Loya, Y., Murillo, A.C., and Treibitz, T. (2021). Repeatable semantic reef-mapping through photogrammetry and label-augmentation. Remote Sensing 13, 659. DOI: 10.3390/rs13040659

  20. Baird, A.H., Guest, J.R., Edwards, A.J., Bauman, A.G., Bouwmeester, J., Mera, H., Abrego, D., Alvarez-Noriega, M., Babcock, R.C., Barbosa, M.B., et al. (2021). An Indo-Pacific coral spawning database. Scientific data 8, 1-9. DOI: 10.1038/s41597-020-00793-8

  21. Weinstein, D.K., Tamir, R., Kramer, N., Eyal, G., Berenshtein, I., Shaked, Y., Loya, Y., and Torfstein, A. (2021). Mesophotic reef geomorphology categorization, habitat identification, and relationships with surface cover and terrace formation in the Gulf of Aqaba. Geomorphology 379, 107548. DOI: 10.1016/j.geomorph.2020.107548

  22. Ben-Zvi, O., Ofer, E., Eyal, G., and Loya, Y. (2021). Experimental evidence of temperature-induced bleaching in two fluorescence morphs of a Red Sea mesophotic coral. Coral Reefs 40, 187-199. DOI: 10.1007/s00338-020-02027-0

  23. Eyal-Shaham, L., Eyal, G., Ben-Zvi, O., Sakai, K., Harii, S., Sinniger, F., Hirose, M., Cabaitan, P., Bronstein, O., and Feldman, B. (2020). A unique reproductive strategy in the mushroom coral Fungia fungites. Coral Reefs 39, 1793-1804. DOI: 10.1007/s00338-020-02004-7

  24. Wada, N., Yuasa, H., Kajitani, R., Gotoh, Y., Ogura, Y., Yoshimura, D., Toyoda, A., Tang, S.-L., Higashimura, Y., Sweatman, H., et al. (2020). A ubiquitous subcuticular bacterial symbiont of a coral predator, the crown-of-thorns starfish, in the Indo-Pacific. Microbiome 8, 1-14. DOI: 10.1186/s40168-020-00880-3

  25. Kramer, N., Tamir, R., Eyal, G., and Loya, Y. (2020). Coral morphology portrays the spatial distribution and population size-structure along a 5–100 m depth gradient. Frontiers in Marine Science 7, 615. DOI: 10.3389/fmars.2020.00615

  26. Laverick, J.H., Tamir, R., Eyal, G., and Loya, Y. (2020). A generalized light‐driven model of community transitions along coral reef depth gradients. Global Ecology and Biogeography 29, 1554-1564. DOI: 10.1111/geb.13140 (Front Cover)

  27. Tamir, R., Ben-Zvi, O., Eyal, G., Kramer, N., and Loya, Y. (2020). Reciprocal-transplantation between shallow and mesophotic stony corals. Marine Environmental Research, 105035. DOI: 10.1016/j.marenvres.2020.105035

  28. Ben-Zvi, O., Tamir, R., Keren, N., Tchernov, D., Berman-Frank, I., Kolodny, Y., Benaltabet, T., Bavli, H., Friedman, M., Glanz-Idan, N., et al. (2020). Photophysiology of a mesophotic coral 3 years after transplantation to a shallow environment. Coral Reefs 39, 903-913. DOI: 10.1007/s00338-020-01910-0

  29. Meron, D., Maor-Landaw, K., Eyal, G., Elifantz, H., Banin, E., Loya, Y., and Levy, O. (2020). The complexity of the holobiont in the Red Sea coral Euphyllia paradivisa under heat stress. Microorganisms 8, 372. DOI: 10.3390/microorganisms8030372

  30. Tamir, R., Eyal, G., Cohen, I., and Loya, Y. (2020). Effects of light pollution on the early life stages of the most abundant northern red sea coral. Microorganisms 8, 193. DOI: 10.3390/microorganisms8020193

  31. Bongaerts, P., Perez-Rosales, G., Radice, V.Z., Eyal, G., Gori, A., Gress, E., Hammerman, N.M., Hernandez-Agreda, A., Laverick, J., and Muir, P. (2019). Mesophotic. org: a repository for scientific information on mesophotic ecosystems. Database-The Journal of Biological Databases and Curation 2019, baz140. DOI: 10.1093/database/baz140

  32. Alonso, I., Yuval, M., Eyal, G., Treibitz, T., and Murillo, A.C. (2019). CoralSeg: Learning coral segmentation from sparse annotations. Journal of Field Robotics 36, 1456-1477. DOI: 10.1002/rob.21915

  33. Tamir, R., Eyal, G., Kramer, N., Laverick, J.H., and Loya, Y. (2019). Light environment drives the shallow‐to‐mesophotic coral community transition. Ecosphere 10, e02839. DOI: 10.1002/ecs2.2839

  34. Meron, D., Maor-Landaw, K., Weizman, E., Waldman Ben-Asher, H., Eyal, G., Banin, E., Loya, Y., and Levy, O. (2019). The algal symbiont modifies the transcriptome of the scleractinian coral Euphyllia paradivisa during heat stress. Microorganisms 7, 256. DOI: 10.3390/microorganisms7080256

  35. Lindemann, Y., Eyal, G., and Genin, A. (2019). Intense capture of swarming pteropods by large-polyp corals. Galaxea, Journal of Coral Reef Studies 21, 9-10. DOI: 10.3755/galaxea.21.1_9

  36. Pinheiro, H.T., Eyal, G., Shepherd, B., and Rocha, L.A. (2019). Ecological insights from environmental disturbances in mesophotic coral ecosystems. Ecosphere 10, e02666. DOI: 10.1002/ecs2.2666

  37. Ben-Zvi, O., Eyal, G., and Loya, Y. (2019). Response of fluorescence morphs of the mesophotic coral Euphyllia paradivisa to ultra-violet radiation. Scientific Reports 9, 5245. DOI: 10.1038/s41598-019-41710-3

  38. Eyal-Shaham, L., Eyal, G., Sakai, K., Nozawa, Y., Harii, S., Sinniger, F., Bronstein, O., Ben-Zvi, O., Shlesinger, T., and Loya, Y. (2019). Repetitive sex change in the stony coral Herpolitha limax across a wide geographic range. Scientific Reports 9, 2936. DOI: 10.1038/s41598-018-37619-y

  39. Kramer, N., Eyal, G., Tamir, R., and Loya, Y. (2019). Upper mesophotic depths in the coral reefs of Eilat, Red Sea, offer suitable refuge grounds for coral settlement. Scientific Reports 9, 2263. DOI: 10.1038/s41598-019-38795-1

  40. Eyal, G., Cohen, I., Eyal-Shaham, L., Ben-Zvi, O., Tikochinski, Y., and Loya, Y. (2019). Photoacclimation and induction of light-enhanced calcification in the mesophotic coral Euphyllia paradivisa. Royal Society Open Science 6, 180527. DOI: 10.1098/rsos.180527

  41. Andradi-Brown, D.A., Vermeij, M.J., Slattery, M., Lesser, M., Bejarano, I., Appeldoorn, R., Goodbody-Gringley, G., Chequer, A.D., Pitt, J.M., Eddy, C., et al. (2017). Large-scale invasion of western Atlantic mesophotic reefs by lionfish potentially undermines culling-based management. Biological Invasions 19, 939-954. DOI: 10.1007/s10530-016-1358-0

  42. Hume, B.C., Voolstra, C.R., Arif, C., D’Angelo, C., Burt, J.A., Eyal, G., Loya, Y., and Wiedenmann, J. (2016). Ancestral genetic diversity associated with the rapid spread of stress-tolerant coral symbionts in response to Holocene climate change. Proceedings of the National Academy of Sciences 113, 4416-4421. DOI: 10.1073/pnas.1601910113

  43. Beijbom, O., Treibitz, T., Kline, D.I., Eyal, G., Khen, A., Neal, B., Loya, Y., Mitchell, B.G., and Kriegman, D. (2016). Improving automated annotation of benthic survey images using wide-band fluorescence. Scientific Reports 6, 23166. DOI: 10.1038/srep23166

  44. Eyal-Shaham, L., Eyal, G., Tamir, R., and Loya, Y. (2016). Reproduction, abundance and survivorship of two Alveopora spp. in the mesophotic reefs of Eilat, Red Sea. Scientific Reports 6, 20964. DOI: 10.1038/srep20964

  45. Eyal, G., Eyal-Shaham, L., Cohen, I., Tamir, R., Ben-Zvi, O., Sinniger, F., and Loya, Y. (2016). Euphyllia paradivisa, a successful mesophotic coral in the northern Gulf of Eilat/Aqaba, Red Sea. Coral Reefs 35, 91-102. DOI: 10.1007/s00338-015-1372-1

  46. Eyal, G., Wiedenmann, J., Grinblat, M., D’Angelo, C., Kramarsky-Winter, E., Treibitz, T., Ben-Zvi, O., Shaked, Y., Smith, T.B., Harii, S., et al. (2015). Spectral diversity and regulation of coral fluorescence in a mesophotic reef habitat in the Red Sea. PloS One 10, e0128697. DOI: 10.1371/journal.pone.0128697

  47. Katz, T., Ginat, H., Eyal, G., Steiner, Z., Braun, Y., Shalev, S., and Goodman-Tchernov, B. (2015). Desert flash floods form hyperpycnal flows in the coral-rich Gulf of Aqaba, Red Sea. Earth and Planetary Science Letters 417, 87-98. DOI: 10.1016/j.epsl.2015.02.025

  48. Ben-Zvi, O., Eyal, G., and Loya, Y. (2015). Light-dependent fluorescence in the coral Galaxea fascicularis. Hydrobiologia 759, 15-26. DOI: 10.1007/s10750-014-2063-6

  49. Lavy, A., Eyal, G., Neal, B., Keren, R., Loya, Y., and Ilan, M. (2015). A quick, easy and non‐intrusive method for underwater volume and surface area evaluation of benthic organisms by 3D computer modelling. Methods in Ecology and Evolution 6, 521-531. DOI: 10.1111/2041-210X.12331

  50. Pasternak, G., Ziv, R., Eyal, G., Shefer, S., Mienis, H.K., Rittner, O., and Galil, B.S. (2011). On the population of Chromodoris annulata Eliot, 1904 (Mollusca: Opistobranchia: Chromodorididae) off the Mediterranean coast of Israel. Aquatic Invasions 6, S91-S93. DOI: 10.3391/ai.2011.6.S1.021

  51. Eyal, G., Eyal-Shaham, L., and Loya, Y. (2011). “Teeth-anchorage”: sleeping behavior of a Red Sea filefish on a branching coral. Coral Reefs 30, 707-707. DOI: 10.1007/s00338-011-0766-y

Review & editorial publications (peer-reviewed):

  1. Pérez-Castro, M.Á., Schubert, N., De Oca, G.A.-M., Leyte-Morales, G.E., Eyal, G., and Hinojosa-Arango, G. (2022). Mesophotic Coral Ecosystems in the Eastern Tropical Pacific: The current state of knowledge and the spatial variability of their depth boundaries. Science of The Total Environment 806, 150576. DOI: 10.1016/j.scitotenv.2021.150576

  2. Eyal, G., and Pinheiro, H.T. (2020). Mesophotic ecosystems: the link between shallow and deep-sea habitats. Diversity 12, 411. DOI: 10.3390/d12110411

  3. Loya, Y., Eyal, G., Treibitz, T., Lesser, M.P., and Appeldoorn, R. (2016). Theme section on mesophotic coral ecosystems: advances in knowledge and future perspectives. Coral Reefs 35, 1-9. DOI: 10.1007/s00338-016-1410-7

Book chapters (peer-reviewed):

  1. Eyal, G., Yuval, M., and Treibitz, T. (2022). The use of advanced technology for monitoring coral reefs. In Coral Reefs of Australia: Perspectives from Beyond the Water's Edge (CSIRO), pp. 343-354. ISBN: 9781486315499

  2. Eyal, G., Tamir, R., Kramer, N., Eyal-Shaham, L., and Loya, Y. (2019). The Red Sea: Israel. In Mesophotic coral ecosystems (Springer), pp. 199-214. DOI: 10.1007/978-3-319-92735-0_11

  3. Turner, J.A., Andradi-Brown, D.A., Gori, A., Bongaerts, P., Burdett, H.L., Ferrier-Pagès, C., Voolstra, C.R., Weinstein, D.K., Bridge, T.C., Costantini, F., et al. (2019). Key questions for research and conservation of mesophotic coral ecosystems and temperate mesophotic ecosystems. In Mesophotic coral ecosystems (Springer), pp. 989-1003. DOI: 10.1007/978-3-319-92735-0_52

  4. Watanabe, T., Watanabe, T.K., Yamazaki, A., Yoneta, S., Sowa, K., Sinniger, F., Eyal, G., Loya, Y., and Harii, S. (2019). Coral sclerochronology: similarities and differences in the coral isotopic signatures between mesophotic and shallow-water reefs. In Mesophotic Coral Ecosystems (Springer), pp. 667-681. DOI: 10.1007/978-3-319-92735-0_36

  5. Andradi-Brown, D., Laverick, J., Bejarano, I., Bridge, T., Colin, P., Eyal, G., Jones, R., Kahng, S., Reed, J., and Smith, T. (2016). Threats to mesophotic coral ecosystems and management options. In Mesophotic coral ecosystems - a lifeboat for coral reefs? (UN Environment, GRID-Arendal), pp. 67-82. ISBN: 978-82-7701-150-9

  6. Baker, E., Thygesen, K., Harris, P., Andradi-Brown, D., Appeldoorn, R.S., Ballantine, D., Bejarano, I., Bridge, T.C., Colin, P.L., Eyal, G., et al. (2016). Mesophotic coral ecosystems: a lifeboat for coral reefs? (UN Environment, GRID-Arendal), pp. 1-98. ISBN: 978-82-7701-150-9

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