Extreme flooding, water quality and mass bleaching compounded impacts in wild and restored Elkhorn coral (Acropora palmata) populations in contrasting urban coral reefs in Puerto Rico
DOI:
https://doi.org/10.15517/0c15pc82Keywords:
Acropora palmata; marine heatwaves; mass bleaching; mortality; restorationAbstract
Introduction: Recurrent marine heatwaves, stronger hurricanes, and degraded water quality represent increasingly compounded threats to Caribbean coral reefs. These stressors may compromise the persistence of wild endangered Elkhorn coral (Acropora palmata) populations and the demographic performance of restored populations.
Objectives: This study examined three climate-driven impact scenarios affecting A. palmata in Puerto Rico: (1) extreme rainfall and flooding from Tropical Storm Isaías affecting wild A. palmata colonies at the urban reef of El Escambrón; (2) compounded bleaching events during 2023–2024 affecting restored colonies at El Escambrón; and (3) the 2024 bleaching event affecting restored colonies at Punta Melones, Culebra, previously fragmented by Hurricane Beryl.
Methods: Case 1 used line photo-transects from El Escambrón (2005-2022) to assess long-term spatial and temporal changes in benthic community structure and wild A. palmata populations. Cases 2 and 3 monitored restored colonies monthly across ten replicate plots per site, measuring survival, live tissue cover, size metrics, and colony condition. Coral metrics were correlated with water quality parameters.
Results: Across El Escambrón and Punta Melones, A. palmata colonies experienced catastrophic mortality of 98–100% associated with recurrent thermal stress, low-salinity urban runoff pulses, and compounded bleaching. Sea surface temperature anomaly, 2024 Bleaching Alert Level 4 conditions, chlorophyll-a, particulate organic carbon, and dissolved oxygen were significantly associated with mortality.
Conclusions: Coral mortality reflects combined acute and chronic stressors, especially in shallow reefs with degraded water quality and reduced thermal tolerance. With limited recruitment, increasing colony isolation, and low genetic diversity, the species may approach functional extinction without a refugia-based restoration framework.
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Aguilar, C., Raina, J. B., Fôret, S., Hayward, D. C., Lapeyre, B., Bourne, D. G., & Miller, D. J. (2019). Transcriptomic analysis reveals protein homeostasis breakdown in the coral Acropora millepora during hypo-saline stress. BMC Genomics, 20, 148. https://doi.org/10.1186/s12864-019-5527-2
Anderson, M. J., Gorley, A. M., & Clarke, K. R. (2008). PERMANOVA+ for PRIMER: guide to software and statistical methods. Primer-E Press.
Anthony, K. R. N., & Hoegh-Guldberg, O. (2003). Variation in coral photosynthesis, respiration, and growth characteristics in contrasting light microhabitats: an analogue to plants in forest gaps and understoreys? Functional Ecology, 17(2), 246–259. https://doi.org/10.1046/j.1365-2435.2003.00731.x
Aronson, R. B., & Precht, W. F. (2001). White-band disease and the changing face of Caribbean coral reefs. Hydrobiologia, 460, 25–38. https://doi.org/10.1023/A:1013103928980
Bachoon, D. S., Markand, S., Otero, E., Perry, G., & Ramsubaugh, A. (2010). Assessment of non-point sources of fecal pollution in coastal waters of Puerto Rico and Trinidad. Marine Pollution Bulletin, 60(7), 1117–1121.
Baird, A. H., & Marshall, P. A. (2002). Mortality, growth and reproduction in scleractinian corals following bleaching on the Great Barrier Reef. Marine Ecology Progress Series, 237, 133–141. https://doi.org/10.3354/meps237133
Baker, D. M., Freeman, C. J., Wong, J. C., Fogel, M. L., & Knowlton, N. (2018). Climate change promotes parasitism in a coral symbiosis. The ISME Journal, 12(3), 921–930. https://doi.org/10.1038/s41396-018-0046-8
Banister, R. B., Viehman, T. S., Schopmeyer, S., & van Woesik, R. (2024). Environmental predictors for the restoration of a critically endangered coral, Acropora palmata, along the Florida reef tract. PLoS ONE, 19(1), e0296485. https://doi.org/10.1371/journal.pone.0296485
Barott, K. L., Rodríguez-Brito, B., Janouškovec, J., Marhaver, K. L., Smith, J. E., Keeling, P., & Rohwer, F. L. (2011). Microbial diversity associated with four functional groups of benthic reef algae and the reef-building coral Montastraea annularis. Environmental Microbiology, 13(5), 1192–1204. https://doi.org/10.1111/j.1462-2920.2010.02419.x
Baumann, J., Grottoli, A. G., Hughes, A. D., & Matsui, Y. (2014). Photoautotrophic and heterotrophic carbon in bleached and non-bleached coral lipid acquisition and storage. Journal of Experimental Marine Biology and Ecology, 461, 469–478. https://doi.org/10.1016/j.jembe.2014.09.017
Bauzá-Ortega, J. F., Villareal, S., & García García, C. (2024). Cuarto informe de la condición ambiental del Estuario de la Bahía de San Juan [Unpublished technical report]. Programa del Estuario de la Bahía de San Juan, Puerto Rico.
Bellwood, D. R., Hughes, T. P., Folke, C., & Nyström, M. (2004). Confronting the coral reef crisis. Nature, 429, 827–833. https://doi.org/10.1038/nature02691
Bessell-Browne, P., Epstein, H. E., Hall, N., Buerger, P., & Berry, K. (2021). Severe heat stress resulted in high coral mortality on Maldivian reefs following the 2015–2016 El Niño event. Oceans, 2(1), 233–245. https://doi.org/10.3390/oceans2010014
Bonilla, S., Zabaleta, B., Vaz, P., Etchebehere, L. G., Haakonsson, S., Risso, J., D’Alessandro, B., Sienra, D., Urban, A., Hernández, T., Trasante, T., Islas, M. B:, & Aubriot, L. (2026). Exceptionally large clumps of Microcystis wesenbergii on Río de la Plata beaches driven by changes in wind direction. Limnologica, 118, 126329. https://doi.org/10.1016/j.limno.2026.126329
Bon, M., Kayal, M., Dromard, C. R., & Desrosiers, C. (2026). Impact of the 2023 coral bleaching event in Martinique, Eastern Caribbean. Coral Reefs, 45, 167–180. https://doi.org/10.1007/s00338-025-02694-x
Bonkosky, M., Hernández-Delgado, E. A., Sandoz, B., Robledo, I. E., Norat-Ramírez, J., & Mattei, H. (2009). Detection of spatial fluctuations of non-point source fecal pollution in coral reef surrounding waters in southwestern Puerto Rico using PCR-based assays. Marine Pollution Bulletin, 58(1), 45–54. https://doi.org/10.1016/j.marpolbul.2008.09.008
Briggs, N. D., Page, C. A., Giuliano, C., Alessi, C., Hoogenboom, M., Bay, L. K., & Randall, C. J. (2024). Dissecting coral recovery: bleaching reduces reproductive output in Acropora millepora. Coral Reefs, 43(3), 557–569. https://doi.org/10.1007/s00338-024-02483-y
Bruckner, A. W. (2002). Proceedings of the Caribbean. Acropora Workshop: Potential Application of the US Endangered Species Act as a Conservation Strategy [Technical Memorandum]. NOAA Technical Memorandum NMFS-OPR.
Bruckner, A. W., & Hourigan, T. (2000). Proactive management for conservation of Acropora cervicornis and Acropora palmata: Application of the U. S. Endangered Species Act. Proceedings of the 9th International Coral Reef Symposium (Bali, Indonesia), 2, 661–665.
Charpy, L., Casareto, B. E., Langlade, M. J., & Suzuki, Y. (2012). Cyanobacteria in coral reef ecosystems: a review. Journal of Marine Sciences, 2012(1), 259571. https://doi.org/10.1155/2012/259571
Chen, Y., Ma, J., Huang, H., Chen, P., Chao, F., & Luo, W. (2017). Effects of temperature, pH, ammonia nitrogen content and salinity on coral (Acropora millepora & Palythoa sp.) bleaching. Agricultural Biotechnology, 6(6), 52–59.
Clarke, K. R., Gorley, R. N., Somerfield, P. J., & Warwick, R. M. (2014). Change in marine communities: an approach to statistical analysis and interpretation [Methods manual, 3rd Ed.]. PRIMER-E. https://learninghub.primer-e.com/books/a-quick-guide-to-primer-7/page/download-manuals
Cloern, J. E. (2001). Our evolving conceptual model of the coastal eutrophication problem. Marine Ecology Progress Series, 210, 223–253. https://doi.org/10.3354/meps210223
Cordeiro, R., Luz, R., Vasconcelos, V., Fonseca, A., & Gonçalves, V. (2020). A critical review of cyanobacteria distribution and cyanotoxins occurrence in Atlantic Ocean islands. Cryptogamie, Algologie, 41(9), 73–89. https://doi.org/10.5252/cryptogamie-algologie2020v41a9
Cunning, R., & Baker, A. C. (2013). Excess algal symbionts increase the susceptibility of corals to bleaching. Nature Climate Change, 3, 259–262. https://doi.org/10.1038/nclimate1711
Cziesielski, M. J., Schmidt-Roach, S., & Aranda, M. (2019). The past, present, and future of coral heat stress studies. Ecology and Evolution, 9(17), 10055–10066. https://doi.org/10.1002/ece3.5576
Destri, G., Güth, A. Z., Luza, A. L., Ibanhez, J. Y., Dottori, M., Silveira, I. C., Braz, G. B., De La Cour, J. L., Manzello, D. P., Skirving, W. J., & Mies, M. (2025). The thermal stress history of South Atlantic reefs reveals increasing intensity, duration, frequency, and likely undocumented bleaching episodes. Global Change Biology, 31(4), e70162. https://doi.org/10.1111/gcb.70162
Díaz-Ortega, G., & Hernández-Delgado, E. A. (2014). Unsustainable land-based source pollution in a climate of change: A roadblock to the conservation and recovery of Elkhorn Coral Acropora palmata (Lamarck 1816). Natural Resources, 5(10), 561–581. https://doi.org/10.4236/nr.2014.510050
Donovan, M. K., Adam, T. C., Shantz, A. A., Speare, K. E., Munsterman, K. S., Rice, M. M., Schmitt, R:, Holbrook, S. J., & Burkepile, D. E. (2020). Nitrogen pollution interacts with heat stress to increase coral bleaching severity. Proceedings of the National Academy of Sciences, 117(10), 5351–5357. https://doi.org/10.1073/pnas.1915395117
Donovan, M. K., Burkepile, D. E., Kratochwill, C., Shlesinger, T., Sully, S., Oliver, T. A., Hodgson, G., Freiwald, J., & van Woesik, R. (2021). Local conditions magnify coral loss after marine heatwaves. Science, 372(6545), 977–980. https://doi.org/10.1126/science.abd9464
Downs, C. A., Kramarsky-Winter, E., Woodley, C. M., Downs, A., Winters, G., Loya, Y., & Ostrander, G. K. (2009). Cellular pathology and histopathology of hypo-salinity exposure on the coral Stylophora pistillata. Science of the Total Environment, 407(17), 4838–4851. https://doi.org/10.1016/j.scitotenv.2009.05.015
Eakin, C. M., Morgan, J. A., Heron, S. F., Smith, T. B., Liu, G., Alvarez-Filip, L., Baca, B., Bartels, E., Bastidas, C., Bouchon, C., Brandt, M., Bruckner, A. W., Bunkley-Williams, L., Cameron, A., Causey, B. D., Chiappone, M., Christensen, T. R. L., Crabbe, M. J. C., Day, O. … Yusuf, Y. (2010). Caribbean corals in crisis: record thermal stress, bleaching, and mortality in 2005. PLoS ONE, 5(11), e13969. https://doi.org/10.1371/journal.pone.0013969
Ellis, S. R. (1976) History of dredging and filling of lagoons in the San Juan area, Puerto Rico [Report]. Water Resources Investigations 38–76, U.S. Geological Survey. https://doi.org/10.3133/wri7638
ERDDAP-NOAA (2024). Easier access to scientific data [Data base]. National Oceanic and Atmospheric Administration. https://coastwatch.pfeg.noaa.gov/erddap/griddap/index.html?page=1&itemsPerPage=1000
Erftemeijer, P. L. A., Riegl, B., Hoeksema, B. W., & Todd, P. A. (2012). Environmental impacts of dredging and other sediment disturbances on corals: A review. Marine Pollution Bulletin, 64(9), 1737–1765. https://doi.org/10.1016/j.marpolbul.2012.05.008
Estrada-Saldívar, N., Jordán-Dalhgren, E., Rodríguez-Martínez, R. E., Perry, C., & Alvarez-Filip, L. (2019). Functional consequences of the long-term decline of reef-building corals in the Caribbean: evidence of across-reef functional convergence. Royal Society Open Science, 6(10), 190298. https://doi.org/10.1098/rsos.190298
Fabricius, K. E. (2005). Effects of terrestrial runoff on the ecology of corals and coral reefs: review and synthesis. Marine Pollution Bulletin, 50(2), 125–146. https://doi.org/10.1016/j.marpolbul.2004.11.028
Familkhalili, R., Storlazzi, C. D., Nemeth, M., & Viehman, S. (2025). Assessing the effect of coral reef restoration location on coastal flood hazard along the San Juan Coastline, Puerto Rico. Frontiers in Marine Science, 12, 1528460. https://doi.org/10.3389/fmars.2025.1528460
Faxneld, S., Jörgensen, T. L., & Tedengren, M. (2010). Effects of elevated water temperature, reduced salinity and nutrient enrichment on the metabolism of the coral Turbinaria mesenterina. Estuarine, Coastal and Shelf Science, 88(4), 482–487. https://doi.org/10.1016/j.ecss.2010.05.008
Ford, A. K., Bejarano, S., Nugues, M. M., Visser, P. M., Albert, S., & Ferse, S. C. (2018). Reefs under siege—the rise, putative drivers, and consequences of benthic cyanobacterial mats. Frontiers in Marine Science, 5, 18. https://doi.org/10.3389/fmars.2018.00018
Ford, A. K., Van Hoytema, N., Moore, B. R., Pandihau, L., Wild, C., & Ferse, S. C. (2017). High sedimentary oxygen consumption indicates that sewage input from small islands drives benthic community shifts on overfished reefs. Environmental Conservation, 44(4), 405–411. https://doi.org/10.1017/S0376892917000054
Ford, A. K., Visser, P. M., Van Herk, M. J., Jongepier, E., & Bonito, V. (2021). First insights into the impacts of benthic cyanobacterial mats on fish herbivory functions on a nearshore coral reef. Scientific Reports, 11, 7147. https://doi.org/10.1038/s41598-021-84016-z
Gardner, T. A., Côté, I. M., Gill, J. A., Grant, A., & Watkinson, A. R. (2003). Long-term region-wide declines in Caribbean corals. Science, 301(5635), 958–960. https://doi.org/10.1126/science.1086050
Glynn, P. W. (1973). Aspects of the ecology of coral reefs in the western Atlantic region. In O. A. Jones, & R. Endean (Eds.), Biology and Geology of Coral Reefs (Vol. 2, pp. 271–324). Academic Press.
Gómez-Andújar, N. X., & Hernandez-Delgado, E. A. (2020). Spatial benthic community analysis of shallow coral reefs to support coastal management in Culebra Island, Puerto Rico. PeerJ, 8, e10080. https://doi.org/10.7717/peerj.10080
Gómez-Gómez, F., & Ellis. S. R. (1983). Hydrologic characteristics of lagoons at San Juan, Puerto Rico, during an October 1974 tidal cycle [Report]. Open-File Report 82–349. U.S. Geological Survey. https://doi.org/10.3133/ofr82349
González-Barrios, F. J., Estrada-Saldívar, N., Pérez-Cervantes, E., Secaira-Fajardo, F., & Álvarez-Filip, L. (2023). Legacy effects of anthropogenic disturbances modulate dynamics in the world’s coral reefs. Global Change Biology, 29(12), 3285–3303. https://doi.org/10.1111/gcb.16686
González-Figueroa, M. C., & Hernández-Delgado, E. A. (2021). Variación espacial en los patrones de recuperación natural de los arrecifes de coral someros urbanos en Puerto Rico. Perspectivas en Asuntos Ambientales, 9, 90–111.
Goreau, T. F. (1959). The ecology of Jamaican coral reefs I. Species composition and zonation. Ecology, 40(1), 67–90. https://doi.org/10.2307/1929924
Graham, N. A. J., Jennings, S., MacNeil, M. A., Mouillot, D., & Wilson, S. K. (2015). Predicting climate-driven regime shifts versus rebound potential in coral reefs. Nature, 518, 94–97. https://doi.org/10.1038/nature14140
Grottoli, A. G., Rodrigues, L. J., & Palardy, J. E. (2006). Heterotrophic plasticity and resilience in bleached corals. Nature, 440, 1186–1189. https://doi.org/10.1038/nature04565
Grottoli, A. G., Warner, M. E., Levas, S. J., Aschaffenburg, M. D., Schoepf, V., McGinley, M., Baumann, J., & Matsui, Y. (2014). The cumulative impact of annual coral bleaching can turn some coral species winners into losers. Global Change Biology, 20(12), 3823–3833. https://doi.org/10.1111/gcb.12658
Hernández, W. J., Ortiz-Rosa, S., Armstrong, R. A., Geiger, E. F., Eakin, C. M., & Warner, R. A. (2020). Quantifying the effects of Hurricanes Irma and María on coastal water quality in Puerto Rico using moderate resolution satellite sensors. Remote Sensing, 12(6), 964. https://doi.org/10.3390/rs12060964
Hernández-Delgado, E. A. (2015). The emerging threats of climate change on tropical coastal ecosystem services, public health, local economies and livelihood sustainability of small islands: Cumulative impacts and synergies. Marine Pollution Bulletin, 101(1), 5–28. https://doi.org/10.1016/j.marpolbul.2015.09.018
Hernández-Delgado, E. A. (2024). Coastal restoration challenges and strategies for small island developing states in the face of sea level rise and climate change. Coasts, 4(2), 235–286. https://doi.org/10.3390/coasts4020014
Hernández-Delgado, E. A., Alejandro-Camis, P., Cabrera-Beauchamp, G., Fonseca-Miranda, J. S., Gómez-Andújar, N. X., Gómez, P., Guzmán-Rodríguez, R., Olivo–Maldonado, I., & Suleimán-Ramos, S. E. (2024). Stronger hurricanes and climate change in the Caribbean Sea: Threats to the sustainability of endangered coral species. Sustainability, 16(4), 1506. https://doi.org/10.3390/su16041506
Hernández-Delgado, E. A., Fonseca-Miranda, J. S., Mercado-Molina, A. E., & Suleimán-Ramos, S. E. (2025). Integrating 3D-printed and natural staghorn coral (Acropora cervicornis) restoration enhances fish assemblages and their ecological functions. Diversity, 17(7), 445. https://doi.org/10.3390/d17070445
Hernández-Delgado, E. A., González-Ramos, C. M., & Alejandro-Camis, P. J. (2014b). Large-scale coral recruitment patterns on Mona Island, Puerto Rico: evidence of a transitional community trajectory after massive coral bleaching and mortality. Revista de Biología Tropical, 62(3), 49–64. https://doi.org/10.15517/rbt.v62i0.15901
Hernandez-Delgado, E. A., Hutchinson-Delgado, Y. M., Laureano, R., Hernandez-Pacheco, R., Ruiz-Maldonado, T. M., Oms, J., & Diaz, P. L. (2010a, November 1–5). Sediment stress, water turbidity, and sewage impacts on threatened elkhorn coral (Acropora palmata) stands at Vega Baja, Puerto Rico. Proceedings of the Gulf and Caribbean Fisheries Institute (San Juan, Puerto Rico), 63, 83–92.
Hernández-Delgado, E. A., & Laureano, R. (2024). Bringing back reef fish: sustainable impacts of community-based restoration of Elkhorn Coral (Acropora palmata) in Vega Baja, Puerto Rico (2008–2023). Sustainability, 16(14), 5985. https://doi.org/10.3390/su16145985
Hernandez-Delgado, E. A., Medina-Muniz, J. L., Mattei, H., & Norat-Ramirez, J. (2017). Unsustainable land use, sediment-laden runoff, and chronic raw sewage offset the benefits of coral reef ecosystems in a no-take marine protected area. Environmental Management and Sustainable Development, 6(2), 292. https://doi.org/10.5296/emsd.v6i2.10687
Hernández-Delgado, E. A., Mercado-Molina, A. E., Alejandro-Camis, P. J., Candelas-Sánchez, F., Fonseca-Miranda, J. S., González-Ramos, C. M., Guzmán-Rodríguez, R., Mège, P., Montañez-Acuña, A. A., Olivo-Maldonado, I., Otaño-Cruz, A., & S. E. Suleimán-Ramos. (2014a). Community-based coral reef rehabilitation in a changing climate: Lessons learned from hurricanes, extreme rainfall, and changing land use impacts. Open Journal of Ecology, 4(14), 918–944. https://doi.org/10.4236/oje.2014.414077
Hernández-Delgado, E. A., Mercado-Molina, A. E., & Suleimán-Ramos, S. E. (2018a). Multi-disciplinary lessons learned from low-tech coral farming and reef rehabilitation practices. I. Best management practices. In C. Duque-Beltrán, & E. Tello-Camacho (Eds.), Corals in a Changing World (pp. 213–243). InTech Publishers https://doi.org/10.5772/intechopen.73151
Hernández-Delgado, E. A., Mercado-Molina, A. E., Suleimán-Ramos, S. E., & Lucking, M. A. (2018b). Multi-disciplinary lessons learned from low-tech coral farming and reef rehabilitation practices. II. Coral demography and social-ecological benefits. In C. Duque-Beltrán, & E. Tello-Camacho (Eds.), Corals in a Changing World (pp. 245–268). InTech Publishers. https://doi.org/10.5772/intechopen.74283
Hernández-Delgado, E. A., & Ortiz-Flores, M. F. (2022). The long and winding road of shallow Caribbean marginal coral reefs recovery from disturbance in the Anthropocene: A case study from Puerto Rico. Diversity, 14(10), 804. https://doi.org/10.3390/d14100804
Hernández-Delgado, E. A., Rodríguez, J., & Murcia Eslava, J. D. (2020). The fresh face of death: Preliminary assessment of a massive coral and invertebrate mortality event at El Escambrón Beach, San Juan, Puerto Rico – August 2020 [Technical report]. Department of Natural and Environmental Resources and to the San Juan Bay Estuary Program & Sociedad Ambiente Marino.
Hernández-Delgado, E. A., & Rodríguez-González, Y. M. (2025). Runaway climate across the wider Caribbean and eastern tropical Pacific in the Anthropocene: Threats to coral reef conservation, restoration, and social–ecological resilience. Atmosphere, 16(5), 575. https://doi.org/10.3390/atmos16050575
Hernández-Delgado, E. A., & Rosado-Matías, B. J. (2017). Long-lasting impacts of beach renourishment on nearshore urban coral reefs: A glimpse of future impacts of shoreline erosion, climate change and sea level rise. Annals of Marine Biology and Research, 4(1), 1021.
Hernández-Delgado, E. A., Sandoz, B., Bonkosky, M., Mattei, H., & Norat, J. (2010b, July 7–11). Impacts of non-point source sewage pollution in Elkhorn coral, Acropora palmata (Lamarck), assemblages of the southwestern Puerto Rico shelf. Proceedings 11th International Coral Reefs Symposium (Fort Lauderdale, Florida), 747–751.
Hughes, T. P., Anderson, K. D., Connolly, S. R., Heron, S. F., Kerry, J. T., Lough, J. M., Baird, A. H., Baum, J. K., Berumen, M. L., Bridge, T. C., Claar, D. C., Eakin, C. M., Gilmour, J. P., Graham, N. A. J., Harrison, H., Hobbs, J-P. A., Hoey, A. S., Hoogennboom, M., Lowe, R. K. … Wilson, S. K. (2018). Spatial and temporal patterns of mass bleaching of corals in the Anthropocene. Science, 359(6371), 80–83. https://doi.org/10.1126/science.aan8048
Hughes, T. P., Kerry, J. T., Álvarez-Noriega, M., Álvarez-Romero, J. G., Anderson, K. D., Baird, A. H., Babcock, R. C., Beger, M., Bellwood, D. R., Berkelmans, R., Bridge, T. C., Butler, I. R., Byrne, M., Cantin, N. E., Comeau, S., Connolly, S. R., Cumming, G. S., Dalton, S. J., Diaz-Pulido, G., … Wilson, S. K. (2017). Global warming and recurrent mass bleaching of corals. Nature, 543, 373–377. https://doi.org/10.1038/nature21707
Huntington, B. E., Miller, M. W., Pausch, R., & Richter, L. (2017). Facilitation in Caribbean coral reefs: high densities of staghorn coral foster greater coral condition and reef fish composition. Oecologia, 184, 247–257. https://doi.org/10.1007/s00442-017-3859-7
Huntington, B., Weible, R., Halperin, A., Winston, M., McCoy, K., Amir, C., Asher, J., & Vargas-Angel, B. (2022). Early successional trajectory of benthic community in an uninhabited reef system three years after mass coral bleaching. Coral Reefs, 41, 1087–1096. https://doi.org/10.1007/s00338-022-02246-7
Inoue, M., Shinmen, K., Kawahata, H., Nakamura, T., Tanaka, Y., Kato, A., Shinzato, C., Iguchi, A., Kan, H., Suzuki, A., & Sakai, K. (2012). Estimate of calcification responses to thermal and freshening stresses based on culture experiments with symbiotic and aposymbiotic primary polyps of a coral, Acropora digitifera. Global and Planetary Change, 92–93, 1–7.
Jackson, J., Donovan, M., Cramer, K., & Lam, V. (2014). Status and trends of Caribbean coral reefs: 1970-2012. Global Coral Reef Monitoring Network & International Union for the Conservation of Nature (IUCN).
Johnston, E. C., Counsell, C. W., Sale, T. L., Burgess, S. C., & Toonen, R. J. (2020). The legacy of stress: coral bleaching impacts reproduction years later. Functional Ecology, 34(11), 2315–2325. https://doi.org/10.1111/1365-2435.13653
Kerswell, A. P., & Jones, R. J. (2003). Effects of hypo-osmosis on the coral Stylophora pistillata: nature and cause of low-salinity bleaching. Marine Ecology Progress Series, 253, 145–154. https://doi.org/10.3354/meps253145
Kjerfve, B., McField, M., Thattai, D., & Giró, A. (2021). Coral reef health in the Gulf of Honduras in relation to fluvial runoff, hurricanes, and fishing pressure. Marine Pollution Bulletin, 172, 112865. https://doi.org/10.1016/j.marpolbul.2021.112865
Kuanui, P., Chavanich, S., Viyakarn, V., Omori, M., & Lin, C. (2015). Effects of temperature and salinity on survival rate of cultured corals and photosynthetic efficiency of zooxanthellae in coral tissues. Ocean Science Journal, 50(2), 263–268. https://doi.org/10.1007/s12601-015-0023-3
Kuffner, I. B., & Paul, V. J. (2004). Effects of the benthic cyanobacterium Lyngbya majuscula on larval recruitment of the reef corals Acropora surculosa and Pocillopora damicornis. Coral Reefs, 23(3), 455–458. https://doi.org/10.1007/s00338-004-0416-8
Kuffner, I. B., Walters, L. J., Becerro, M. A., Paul, V. J., Ritson-Williams, R., & Beach, K. S. (2006). Inhibition of coral recruitment by macroalgae and cyanobacteria. Marine Ecology Progress Series, 323, 107–117. https://doi.org/10.3354/meps323107
Lapointe, B. E., Barile, P. J., Littler, M. M., & Littler, D. S. (2004). Macroalgal blooms on southeast Florida coral reefs: II. Cross-shelf discrimination of nitrogen sources indicates widespread assimilation of sewage nitrogen. Harmful Algae, 4(6), 1106–1122. https://doi.org/10.1016/j.hal.2005.06.002
Larsen, M. C., & Webb, R. M. (2009). Potential effects of runoff, fluvial sediment, and nutrient discharges on the coral reefs of Puerto Rico. Journal of Coastal Research, 25(1), 189–208. https://doi.org/10.2112/07-0920.1
Leggat, W., Camp, E. F., Suggett, D. J., Heron, S. F., Fordyce, A. J., Gardner, S., Deakin, L., Turner, M., Beeching, L. J., Kuzhiumparambil, U., Eakin, C. M., & Ainsworth, T. D. (2019). Rapid coral decay is associated with marine heatwave mortality events on reefs. Current Biology, 29(16), 2723–2730. https://doi.org/10.1016/j.cub.2019.06.077
Mann, M. E. (2024). Cat 6 hurricanes have arrived. PNAS, 121(7), e2322597121. https://doi.org/10.1073/pnas.2322597121
Manzello, D. P. (2015). Rapid recent warming of coral reefs in the Florida Keys. Scientific Reports, 5(1), 16762. https://doi.org/10.1038/srep16762
Manzello, D. P., Cunning, R., Karp, R. F., Baker, A. C., Bartels, E., Bonhag, R., Borreil, A., Bourque, A., Brown, K. T., Bruckner, A:. W., Corbett, B., D’Alessandro, M., Dahlgren, C., Dilworth, J., Geiger, E., Gilliam, D. S., Gomez, M., Hanson, G., … Williamson, O. M. (2025). Heat-driven functional extinction of Caribbean Acropora corals from Florida’s coral reefs. Science, 390(6771), 361–366. https://doi.org/10.1126/science.adx7825
Mejias-Rivera, C. L., Ricaurte-Chica, M., Carballeira-Martínez, J., Hernández, W. J., Armstrong, R. A., Cruz-Motta, J. J., & Courtney, T. A. (2026). Assessment of coral reef eutrophication thresholds across Puerto Rico using in-situ and satellite-derived chlorophyll-a. Estuaries and Coasts, 49, 26. https://doi.org/10.1007/s12237-025-01644-8
Méndez-Tejeda, R., & Hernández-Ayala, J. J. (2023). Links between climate change and hurricanes in the North Atlantic. PLOS Climate, 2(4), e0000186. https://doi.org/10.1371/journal.pclm.0000186
Mercado-Molina, A. E., Sabat, A. M., & Hernández-Delgado, E. A. (2020). Population dynamics of diseased corals: Effects of a shutdown reaction outbreak in Puerto Rican Acropora cervicornis. Advances in Marine Biology, 87(1), 61–82. https://doi.org/10.1016/bs.amb.2020.08.001
Milliman, J. D. (1973). Caribbean coral reefs. In O.A. Jones, & R. Endean (Eds.), Biology and Geology of Coral Reefs Volume 1: Geology 1 (pp. 1–50). Elsevier Science.
Morikawa, M. K., & Palumbi, S. R. (2019). Using naturally occurring climate-resistant corals to construct bleaching-resistant nurseries. PNAS, 116(21), 10586–10591. https://doi.org/10.1073/pnas.1721415116
Mumby, P. J., Hastings, A., & Edwards, H. J. (2007). Thresholds and the resilience of Caribbean coral reefs. Nature, 450, 98–101. https://doi.org/10.1038/nature06252
Murakami, H., Levin, E., Delworth, T. L., Gudgel, R., & Hsu, P. C. (2018). Dominant effect of relative tropical Atlantic warming on major hurricane occurrence. Science, 362(6416), 794–799. https://doi.org/10.1126/science.aat6711
Nakano, Y., Tsuchiya, M., Rungsupa, S., & Yamazato, K. Y. K. (2009). Influence of severe freshwater flooding during the rainy season on the coral community around Khang Khao Island in the inner Gulf of Thailand. Galaxea, Journal of Coral Reef Studies, 11(2), 131–138. https://doi.org/10.3755/galaxea.11.131
Osorio-Cano, J. D., Osorio, A. F., Alcérreca-Huerta, J. C., & Oumeraci, H. (2019). Drag and inertia forces on a branched coral colony of Acropora palmata. Journal of Fluids and Structures, 88, 31–47. https://doi.org/10.1016/j.jfluidstructs.2019.04.001
Otaño-Cruz, A., Montañez-Acuña, A. A., García-Rodríguez, N. M., Díaz-Morales, D. M., Benson, E., Cuevas, E., Ortiz-Zayas, J., & Hernández-Delgado, E. A. (2019). Caribbean near-shore coral reef benthic community response to changes on sedimentation dynamics and environmental conditions. Frontiers in Marine Science, 6, 551. https://doi.org/10.3389/fmars.2019.00551
Otaño-Cruz, A., Montañez-Acuña, A. A., Torres-López, V., Hernández-Figueroa, E. M., & Hernández-Delgado, E. A. (2017). Effects of changing weather, oceanographic conditions, and land uses on spatio-temporal variation of sedimentation dynamics along near-shore coral reefs. Frontiers in Marine Science, 4, 249. https://doi.org/10.3389/fmars.2017.00249
Pait, A. S., Whitall, D. R., Jeffrey, C. F., Caldow, C., Mason, A. L., Lauenstein, G. G., & Christensen, J. D. (2008). Chemical contamination in southwest Puerto Rico: an assessment of organic contaminants in nearshore sediments. Marine Pollution Bulletin, 56(3), 580–587. https://doi.org/10.1016/j.marpolbul.2007.11.007
Pérez-Villalona, H., Cornwell, J. C., Ortiz-Zayas, J.R., Cuevas, E. (2015). Sediment denitrification and nutrient fluxes in the San José Lagoon, a tropical lagoon in the highly urbanized San Juan Bay Estuary, Puerto Rico. Estuaries and Coasts, 38, 2259–2278. https://doi.org/10.1007/s12237-015-9953-3
Perry, C. T. (2003). Reef development at Inhaca Island, Mozambique: coral communities and impacts of the 1999/2000 southern African floods. AMBIO: A Journal of the Human Environment, 32(2), 134–139. https://doi.org/10.1579/0044-7447-32.2.134
Ramos-Scharrón, C. E., Amador, J. M., & Hernández-Delgado, E. A. (2012). An interdisciplinary erosion mitigation approach for coral reef protection – A case study from the eastern Caribbean. In A. Cruzado (Ed.), Marine Ecosystems, Intech Publications. (127–160). https://doi.org/10.5772/35709
Ramos-Scharrón, C., Torres-Pulliza, D., & Hernández-Delgado, E. A. (2015). Watershed- and island-scale land cover changes in Puerto Rico (1930s-2004) and their potential effects on coral reef ecosystems. Science of the Total Environment, 506–507, 241–251. https://doi.org/10.1016/j.scitotenv.2014.11.016
Randall, C. J., & van Woesik, R. (2015). Contemporary white-band disease in Caribbean corals driven by climate change. Nature Climate Change, 5, 375–379. https://doi.org/10.1038/nclimate2530
Randall, C. J., & Van Woesik, R. (2017). Some coral diseases track climate oscillations in the Caribbean. Scientific Reports, 7, 5719. https://doi.org/10.1038/s41598-017-05763-6
Reguero, B. G., Beck, M. W., Agostini, V. N., Kramer, P., & Hancock, B. (2018). Coral reefs for coastal protection: A new methodological approach and engineering case study in Grenada. Journal of Environmental Management, 210, 146–161. https://doi.org/10.1016/j.jenvman.2018.01.024
Ritson-Williams, R., Arnold, S. N., & Paul, V. J. (2020). The impact of macroalgae and cyanobacteria on larval survival and settlement of the scleractinian corals Acropora palmata, A. cervicornis and Pseudodiploria strigosa. Marine Biology, 167, 31. https://doi.org/10.1007/s00227-019-3639-5
Ritson-Williams, R., Ross, C., & Paul, V. J. (2016). Elevated temperature and allelopathy impact coral recruitment. PLoS One, 11(12), e0166581. https://doi.org/10.1371/journal.pone.0166581
Rodrigues, R. R., Gonçalves Neto, A. H., Vieira, E. A., & Longo, G. O. (2025). The severe 2020 coral bleaching event in the tropical Atlantic linked to marine heatwaves. Communications Earth & Environment, 6, 208. https://doi.org/10.1038/s43247-025-02195-3
Rodríguez-Martínez, R. E., Banaszak, A. T., McField, M. D., Beltran-Torres, A. U., & Alvarez-Filip, L. (2014). Assessment of Acropora palmata in the Mesoamerican reef system. Plos One, 9(4), e96140. https://doi.org/10.1371/journal.pone.0096140
Rodríguez-Sierra, C. J., Adelman, D., Vojta, Š., Mansilla-Rivera, I., & Lohmann, R. (2019). Passive sampling of persistent organic pollutants in four coastal aquatic systems of Puerto Rico: a pilot study. Bulletin of Environmental Contamination and Toxicology, 103, 770–775. https://doi.org/10.1007/s00128-019-02731-w
Rogers, C. S. (1990). Responses of coral reefs and reef organisms to sedimentation. Marine Ecology Progress Series, 62, 185–202. https://doi.org/10.3354/MEPS062185
Santos-Flores, C. J., & Martínez-Rodríguez, G. A. (2024). A preliminary characterization of the phytoplankton in San José Lagoon, Puerto Rico [Unpublished technical report]. Technical report submitted to the San Juan Bay Estuary Program, San Juan, Puerto Rico.
Schoepf, V., Stat, M., Falter, J. L., & McCulloch, M. T. (2015). Limits to the thermal tolerance of corals adapted to a highly fluctuating, naturally extreme temperature environment. Scientific Reports, 5, 17639. https://doi.org/10.1038/srep17639
Skirving, W. J., Heron, S. F., Marsh, B. L., Liu, G., De La Cour, J. L., Geiger, E. F., & Eakin, C. M. (2019). The relentless march of mass coral bleaching: a global perspective of changing heat stress. Coral Reefs, 38, 547–557. https://doi.org/10.1007/s00338-019-01799-4
Smith, J. E., Shaw, M., Edwards, R. A., Obura, D., Pantos, O., Sala, E., Sandin, S. A., Smriga, S., Hatay, M., & Rohwer, F. L. (2006). Indirect effects of algae on coral: algae-mediated, microbe-induced coral mortality. Ecology Letters, 9(7), 835–845. https://doi.org/10.1111/j.1461-0248.2006.00937.x
Stat, M., Pochon, X., Franklin, E. C., Bruno, J. F., Casey, K. S., Selig, E. R., & Gates, R. D. (2013). The distribution of the thermally tolerant symbiont lineage (Symbiodinium clade D) in corals from Hawaii: correlations with host and the history of ocean thermal stress. Ecology and Evolution, 3(5), 1317–1329. https://doi.org/10.1002/ece3.556
Stoyneva-Gärtner, M., Stefanova, K., Descy, J. P., Uzunov, B., Radkova, M., Pavlova, V., Mitreva, M., & Gärtner, G. (2020). Microcystis aeruginosa and M. wesenbergii were the primary planktonic microcystin producers in several Bulgarian waterbodies. Applied Sciences, 11(1), 357. https://doi.org/10.3390/app11010357
Stuart-Smith, R. D., Brown, C. J., Ceccarelli, D. M., & Edgar, G. J. (2018). Ecosystem restructuring along the Great Barrier Reef following mass coral bleaching. Nature, 560, 92–96. https://doi.org/10.1038/s41586-018-0359-9
Sully, S., Burkepile, D. E., Donovan, M. K., Hodgson, G., & Van Woesik, R. (2019). A global analysis of coral bleaching over the past two decades. Nature Communications, 10, 1264. https://doi.org/10.1038/s41467-019-09238-2
Sully, S., Hodgson, G., & van Woesik, R. (2022). Present and future bright and dark spots for coral reefs through climate change. Global Change Biology, 28(15), 4509–4522. https://doi.org/10.1111/gcb.16083
Sutherland, K. P., Shaban, S., Joyner, J. L., Porter, J. W., & Lipp, E. K. (2011). Human pathogen shown to cause disease in the threatened eklhorn coral Acropora palmata. PloS One, 6(8), e23468. https://doi.org/10.1371/journal.pone.0023468
Tatters, A. O., Smith, J., Kudela, R. M., Hayashi, K., Howard, M. D., Donovan, A. R., Loftin, K. A., & Caron, D. A. (2021). The tide turns: episodic and localized cross-contamination of a California coastline with cyanotoxins. Harmful Algae, 103, 102003. https://doi.org/10.1016/j.hal.2021.102003
Torres-Pérez, J. L., Hernández, W. J., Ramos-Scharrón, C., Armstrong, R. A., Guild, L. S., Barreto-Orta, M., Ortiz-Zayas, J. R., Cardona-Maldonado, M., Infante, R., & Viqueira-Rios, R. (2026). Bio-optical water quality trends and back-reef benthic community structure in southwestern Puerto Rico. Marine Pollution Bulletin, 228, 119557. https://doi.org/10.1016/j.marpolbul.2026.119557
Toth, L. T., Stathakopoulos, A., Kuffner, I. B., Ruzicka, R. R., Colella, M. A., & Shinn, E. A. (2019). The unprecedented loss of Florida’s reef-building corals and the emergence of a novel ecological assemblage. Ecology, 100(9), e02781. https://doi.org/10.1002/ecy.2781
True, J. D. (2012, 9–13 July). Salinity as a structuring force for near shore coral communities. Proceedings of the 12th International Coral Reef Symposium (Cairns, Australia), 1–7.
Vargas-Angel, B., Huntington, B., Brainard, R. E., Venegas, R., Oliver, T., Barkley, H., & Cohen, A. (2019). El Niño-associated catastrophic coral mortality at Jarvis Island, central Equatorial Pacific. Coral Reefs, 38, 731–741. https://doi.org/10.1007/s00338-019-01838-0
Vega Thurber, R. L., Burkepile, D. E., Fuchs, C., Shantz, A. A., McMinds, R., & Zaneveld, J. R. (2014). Chronic nutrient enrichment increases prevalence and severity of coral disease and bleaching. Global Change Biology, 20(2), 544–554. https://doi.org/10.1111/gcb.12450
Viehman, T. S., Reguero, B. G., Lenihan, H. S., Rosman, J. H., Storlazzi, C. D., Goergen, E. A., Canals Silander, M. F., Groves, S. H., Holstein, D. M., Bruckner, A. W., Carrick, J. V., Haus, B. K., Royster, J. B., Duval, M. S., Torres, W. I., & Hench, J. L. (2023). Coral restoration for coastal resilience: Integrating ecology, hydrodynamics, and engineering at multiple scales. Ecosphere, 14(5), e4517. https://doi.org/10.1002/ecs2.4517
Vizon, C., Urbanowiez, A., Raviglione, D., Bonnard, I., & Nugues, M. M. (2024). Benthic cyanobacterial metabolites interact to reduce coral larval survival and settlement. Harmful Algae, 132, 102582. https://doi.org/10.1016/j.hal.2024.102582
Webb, R. M. T., & Gómez-Gómez, F. (1998). Synoptic survey of water quality and bottom sediments, San Juan Bay Estuary System, Puerto Rico, December 1994-July 1995 [Report]. Water Resources Investigations Report 97-4144. U.S. Geological Survey.
Weijerman, M., Birkeland, C., Piniak, G. A., Miller, M. W., Eakin, C. M., McElhany, P., Dunlap, M. J., Patterson, M., & Brainard, R. E. (2014). Endangered Species Act listing: three case studies of data deficiencies and consequences of ESA ‘threatened’ listing on research output. Current Opinion in Environmental Sustainability, 7, 15–21. https://doi.org/10.1016/j.cosust.2013.11.026
Weis, V. M. (2008). Cellular mechanisms of cnidarian bleaching: stress caused the collapse of symbiosis. Journal of Experimental Biology, 211(19), 3059–3066. https://doi.org/10.1242/jeb.009597
Wilkinson, C., & Souter, D. (2008). Status of Caribbean coral reefs after bleaching and hurricanes in 2005 [Technical report]. Global Coral Reef Monitoring Network, and Reef and Rainforest Research Centre.
Williams, D. E., & Miller, M. W. (2005). Coral disease outbreak: pattern, prevalence and transmission in Acropora cervicornis. Marine Ecology Progress Series, 301, 119–128. https://doi.org/10.3354/meps301119
Williams, D. E., & Miller, M. W. (2006, July 28–June 2 July). Importance of disease and predation to the growth and survivorship of juvenile Acropora palmata and Acropora cervicornis: a demographic approach. Proceedings 10th International Coral Reef Symposium (Okinawa, Japan), 1096–1104.
Wilson, S. K., Adjeroud, M., Bellwood, D. R., Berumen, M. L., Booth, D., Bozec, Y. M., Chabanet, P., Cheal, A., Cinner, J., Depczynski, M., Feary, D. A., Gagliano, M., Graham, N. A. J., Halford, A. R., Halpern, B. S., Harborne, A. R., Hoey, A. S., Holbrook, S. J., Jones, G. P., … Syms, C. (2010). Crucial knowledge gaps in current understanding of climate change impacts on coral reef fishes. Journal of Experimental Biology, 213(6), 894–900. https://doi.org/10.1242/jeb.037895
Wu, Y., Li, L., Gan, N., Zheng, L., Ma, H., Shan, K., Liu, J., Xiao, B., Song, L. (2014). Seasonal dynamics of water bloom-forming Microcystis morphospecies and the associated extracellular microcystin concentrations in large, shallow, eutrophic Dianchi Lake. Journal of Environmental Sciences, 26(9), 1921–1929. https://doi.org/10.1016/j.jes.2014.06.031
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