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The health and continued existence of coral reef ecosystems are threatened by an increasing array of environmental and anthropogenic impacts. Coral disease is one of the prominent causes of increased mortality among reefs globally, particularly in the Caribbean. Although over 40 different coral diseases and syndromes have been reported worldwide, only a few etiological agents have been confirmed; most pathogens remain unknown and the dynamics of disease transmission, pathogenicity and mortality are not understood. Causal relationships have been documented for only a few of the coral diseases, while new syndromes continue to emerge. Extensive field observations by coral biologists have provided substantial documentation of a plethora of new pathologies, but our understanding, however, has been limited to descriptions of gross lesions with names reflecting these observations (e.g., black band, white band, dark spot). To determine etiology, we must equip coral diseases scientists with basic biomedical knowledge and specialized training in areas such as histology, cell biology and pathology. Only through combining descriptive science with mechanistic science and employing the synthesis epizootiology provides will we be able to gain insight into causation and become equipped to handle the pending crisis. One of the critical challenges faced by coral disease researchers is to establish a framework to systematically study coral pathologies drawing from the field of diagnostic medicine and pathology and using generally accepted nomenclature. This process began in April 2004, with a workshop titled Coral Disease and Health Workshop: Developing Diagnostic Criteria co-convened by the Coral Disease and Health Consortium (CDHC), a working group organized under the auspices of the U.S. Coral Reef Task Force, and the International Registry for Coral Pathology (IRCP). The workshop was hosted by the U.S. Geological Survey, National Wildlife Health Center (NWHC) in Madison, Wisconsin and was focused on gross morphology and disease signs observed in the field. A resounding recommendation from the histopathologists participating in the workshop was the urgent need to develop diagnostic criteria that are suitable to move from gross observations to morphological diagnoses based on evaluation of microscopic anatomy. As a continuation of building the foundation and framework for coral disease diagnostics, the CDHC convened the Coral Disease and Health Workshop: Coral Histopathology II in Charleston, South Carolina, July 11-14, 2005. The workshop was hosted by the Department of Pathology and Laboratory Medicine at the Medical University of South Carolina, Charleston, SC which provided expertise, facilities and equipment in support of the workshop. All of the histological slides and related photographs used in the discussions were prepared and supplied by the IRCP. This workshop brought together 15 experts in veterinary and medical pathology and coral biology from national and international research institutes and government laboratories. The mission was to devise a standardized approach to examining microscopic anatomy and pathology of corals and a standardized nomenclature to facilitate accurate descriptions of the microscopic morphology of corals and enhance communication among specialists investigating causes of coral death. 2 The participants of this workshop deliberated for 3 days to refine the nomenclature for gross and microscopic anatomy of corals and systematically described microscopic changes associated with selected coral diseases. The findings and recommendations from the deliberations will be submitted to the research community for peer review. The standardized nomenclature and descriptions produced at this workshop will ultimately be made available to the scientific community through a variety of media including the World Wide Web. An exciting highlight of this meeting was provided by Professor Robert Ogilvie (MUSC Department of Cell Biology and Anatomy) when he introduced participants to a new digital technology that is revolutionizing histology and histopathology in the medical field. The Virtual Slide technology creates digital images of histological tissue sections by computer scanning actual slides in high definition and storing the images for retrieval and viewing. Virtual slides now allow any investigator with access to a computer and the web to view, search, annotate and comment on the same tissue sections in real time. Medical and veterinary slide libraries across the country are being converted into virtual slides to enhance biomedical education, research and diagnosis. The coral health and disease researchers at this workshop deem virtual slides as a significant way to increase capabilities in coral histology and a means for pathology consultations on coral disease cases on a global scale.
Somatic growth dynamics are an integrated response to environmental conditions. Hawksbill sea turtles (Eretmochelys imbricata) are long-lived, major consumers in coral reef habitats that move over broad geographic areas (hundreds to thousands of kilometers). We evaluated spatio-temporal effects on hawksbill growth dynamics over a 33-yr period and 24 study sites throughout the West Atlantic and explored relationships between growth dynamics and climate indices. We compiled the largest ever data set on somatic growth rates for hawksbills – 3541 growth increments from 1980 to 2013. Using generalized additive mixed model analyses, we evaluated 10 covariates, including spatial and temporal variation, that could affect growth rates. Growth rates throughout the region responded similarly over space and time. The lack of a spatial effect or spatio-temporal interaction and the very strong temporal effect reveal that growth rates in West Atlantic hawksbills are likely driven by region-wide forces. Between 1997 and 2013, mean growth rates declined significantly and steadily by 18%. Regional climate indices have significant relationships with annual growth rates with 0- or 1-yr lags: positive with the Multivariate El Niño Southern Oscillation Index (correlation = 0.99) and negative with Caribbean sea surface temperature (correlation = −0.85). Declines in growth rates between 1997 and 2013 throughout the West Atlantic most likely resulted from warming waters through indirect negative effects on foraging resources of hawksbills. These climatic influences are complex. With increasing temperatures, trajectories of decline of coral cover and availability in reef habitats of major prey species of hawksbills are not parallel. Knowledge of how choice of foraging habitats, prey selection, and prey abundance are affected by warming water temperatures is needed to understand how climate change will affect productivity of consumers that live in association with coral reefs. Main conclusions The decadal declines in growth rates between 1997 and 2013 throughout the West Atlantic most likely resulted from warming waters through indirect negative effects on the foraging resources of hawksbills. These climatic influences are complex. With increasing temperatures, the trajectories of decline of coral cover and availability in reef habitats of major prey species of hawksbills are not parallel. Knowledge of how choice of foraging habitats, prey selection, and prey abundance are affected by warming water temperatures is needed to understand how climate change will affect productivity of consumers that live in association with coral reefs.
Coral reef ecosystem management benefits from continual, quantitative assessment of the resources being managed, plus assessment of factors that affect distribution patterns of organisms in the ecosystem. In this study, we investigated the relationships among physical, benthic, and fish variables in effort to help explain the distribution patterns of ecologically and economically important species on twelve patch reefs within Biscayne National Park (BNP), Florida, U.S.A. We visited 196 randomly-located sampling stations across twelve shallow (< 10m) patch reefs, using SCUBA to conduct our surveys. We measured physical variables (e.g., substratum type), estimated the percent cover of benthic community members (e.g., coral, algae), and counted and estimated mean size for each fish species observed. We also used high-density bathymetric data collected remotely via airborne laser surveying (Experimental Advanced Airborne Research Lidar (EAARL)) to calculate rugosity (bumpiness) of the reef habitat. Here we present our findings visually by graphing our quantitative community and physical structure data simultaneously in a GIS map format. You will see that biological organisms arrange themselves on each patch reef in a non-random manner. For example, many species of fish prefer to locate themselves in areas of the reef where the rugosity index is high. Rugose parts of the reef provide them with good hiding places from predators. These maps (and the data used to create them) are permanent records of the status of reef resources found on these twelve patch reefs in BNP as of September, 2003. The survey data found in the shapefile located on this CD product includes benthic percent cover data for algae, coral, encrusting invertebrates, and substratum type, in addition to gorgonian abundance and volume, total fish abundance and species richness, and specific counts for Acanthurids (surgeonfish), Scarids (parrotfish), Lutjanids (snappers), Haemulids (grunts), Serranids (groupers), and Pomacentrids (damselfish).
The ongoing degradation of coral reef habitats is widely acknowledged to have adverse effects on the abundance and diversity of reef fish populations, yet the direct effects on ecosystem functions remain uncertain. This study used a quantitative approach to determine the mechanistic links between fish assemblages and ecological function. We investigated the effects of 3D habitat structure and coral morphology on the ecological, behavioral, and morphological functional traits of reef fish within a protected marine national park. Fish traits such as Gregariousness, Water Column Position, and Body Shape were identified to be highly influential in shaping the multidimensional fish functional space, which was categorized into 10 Fish Functional Groups (FFG). Furthermore, habitat complexity and coral morphology significantly explained the abundances of eight out of 10 FFG. Notably, the habitat complexity metrics of Slope and Surface Complexity, along with coral morphologies of Branching and Mounding types, emerged as the most influential habitat features across FFG. Pairing Compressiform species and Schooling Short/Deep species, for example, significantly increased in abundance on substrate with higher Slopes and increased percentages of branching coral cover. Additionally, Cryptic and Nocturnal species exhibited statistically significant associations with all coral morphologies and substrates with high trait values of Slope and Curvature. Elucidating ecological drivers of specific functional groups of reef fish is critical for determining how changes in reef composition and structure will alter fish assemblages. Broad scale patterns were also detected, suggesting that although structural complexity is important, live coral morphologies have a greater positive impact on reef fish functional groups. These findings have direct implications for conservation and monitoring efforts, offering valuable insights for predicting the impacts of environmental change on community dynamics and ecosystem functioning.
The severity of the impact of elevated atmospheric pCO2 to coral reef ecosystems depends, in part, on how sea-water pCO2 affects the balance between calcification and dissolution of carbonate sediments. Presently, there are insufficient published data that relate concentrations of pCO 2 and CO32- to in situ rates of reef calcification in natural settings to accurately predict the impact of elevated atmospheric pCO2 on calcification and dissolution processes. Rates of net calcification and dissolution, CO32- concentrations, and pCO2 were measured, in situ, on patch reefs, bare sand, and coral rubble on the Molokai reef flat in Hawaii. Rates of calcification ranged from 0.03 to 2.30 mmol CaCO3 m-2 h-1 and dissolution ranged from -0.05 to -3.3 mmol CaCO3 m-2 h-1. Calcification and dissolution varied diurnally with net calcification primarily occurring during the day and net dissolution occurring at night. These data were used to calculate threshold values for pCO2 and CO32- at which rates of calcification and dissolution are equivalent. Results indicate that calcification and dissolution are linearly correlated with both CO32- and pCO2. Threshold pCO2 and CO32- values for individual substrate types showed considerable variation. The average pCO2 threshold value for all substrate types was 654??195 ??atm and ranged from 467 to 1003 ??atm. The average CO32- threshold value was 152??24 ??mol kg-1, ranging from 113 to 184 ??mol kg-1. Ambient seawater measurements of pCO2 and CO32- indicate that CO32- and pCO2 threshold values for all substrate types were both exceeded, simultaneously, 13% of the time at present day atmospheric pCO2 concentrations. It is predicted that atmospheric pCO2 will exceed the average pCO2 threshold value for calcification and dissolution on the Molokai reef flat by the year 2100.
Mass mortality of the dominant coral reef herbivore Diadema antillarum in the Caribbean in the early 1980s contributed to a persistent phase shift from coral- to algal-dominated reefs. In 2022, a scuticociliate most closely related to Philaster apodigitiformis caused further mass mortality of D. antillarum across the Caribbean, leading to >95% mortality at affected sites. Mortality was also reported in the related species Diadema setosum in the Mediterranean in 2022, though the causative agent of the Mediterranean outbreak has not yet been determined. In April 2023, mass mortality of Diadema setosum occurred along the Sultanate of Oman's coastline. Urchins displayed signs compatible with scuticociliatosis including abnormal behavior, drooping and loss of spines, followed by tissue necrosis and death. Here we report the detection of an 18S rRNA gene sequence in abnormal urchins from Muscat, Oman, that is identical to the Philaster strain responsible for D. antillarum mass mortality in the Caribbean. We also show that scuticociliatosis signs can be elicited in Diadema setosum by experimental challenge with the cultivated Philaster strain associated with Caribbean scuticociliatosis. These results demonstrate the Philaster sp. associated with D. antillarum mass mortality has rapidly spread to geographically distant coral reefs, compelling global-scale awareness and monitoring for this devastating condition through field surveys, microscopy, and molecular microbiological approaches, and prompting investigation of long-range transmission mechanisms.
Tropical coral reefs harbour a reservoir of enormous biodiversity that is increasingly threatened by direct human activities and indirect global climate shifts. Emerging coral diseases are one serious threat implicated in extensive reef deterioration through disruption of the integrity of the coral holobiont – a complex symbiosis between the coral animal, endobiotic alga and an array of microorganisms. In this article, we review our current understanding of the role of microorganisms in coral health and disease, and highlight the pressing interdisciplinary research priorities required to elucidate the mechanisms of disease. We advocate an approach that applies knowledge gained from experiences in human and veterinary medicine, integrated into multidisciplinary studies that investigate the interactions between host, agent and environment of a given coral disease. These approaches include robust and precise disease diagnosis, standardised ecological methods and application of rapidly developing DNA, RNA and protein technologies, alongside established histological, microbial ecology and ecological expertise. Such approaches will allow a better understanding of the causes of coral mortality and coral reef declines and help assess potential management options to mitigate their effects in the longer term.
The ability of coral reefs to engineer complex three-dimensional habitats is central to their success and the rich biodiversity they support. In tropical reefs, encrusting coralline algae bind together substrates and dead coral framework to make continuous reef structures, but beyond the photic zone, the cold-water coral Lophelia pertusa also forms large biogenic reefs, facilitated by skeletal fusion. Skeletal fusion in tropical corals can occur in closely related or juvenile individuals as a result of non-aggressive skeletal overgrowth or allogeneic tissue fusion, but contact reactions in many species result in mortality if there is no ‘self-recognition’ on a broad species level. This study reveals areas of ‘flawless’ skeletal fusion in Lophelia pertusa , potentially facilitated by allogeneic tissue fusion, are identified as having small aragonitic crystals or low levels of crystal organisation, and strong molecular bonding. Regardless of the mechanism, the recognition of ‘self’ between adjacent L. pertusa colonies leads to no observable mortality, facilitates ecosystem engineering and reduces aggression-related energetic expenditure in an environment where energy conservation is crucial. The potential for self-recognition at a species level, and subsequent skeletal fusion in framework-forming cold-water corals is an important first step in understanding their significance as ecological engineers in deep-seas worldwide.
Coral bleaching is the largest global threat to coral reef ecosystem persistence this century. Advancing our understanding of coral bleaching and developing solutions to protect corals and the reefs they support are critical. In the present article, we, the US National Science Foundation–funded Coral Bleaching Research Coordination Network, outline future directions for coral bleaching research. Specifically, we address the need for embedded inclusiveness, codevelopment, and capacity building as a foundation for excellence in coral bleaching research and the critical role of coral-bleaching science in shaping policy. We outline a path for research innovation and technology and propose the formation of an international coral bleaching consortium that, in coordination with existing multinational organizations, could be a hub for planning, coordinating, and integrating global-scale coral bleaching research, innovation, and mitigation strategies. This proposed strategy for future coral bleaching research could facilitate a step-function change in how we address the coral bleaching crisis.
A sequence of submerged terraces (L1–L12) offshore Lanai was previously interpreted as reefal, and correlated with a similar series of reef terraces offshore Hawaii island, whose ages are known to be <500 ka. We present bathymetric, observational, lithologic and 51 87 Sr/ 86 Sr isotopic measurements for the submerged Lanai terraces ranging from −300 to −1000 m (L3–L12) that indicate that these terraces are drowned reef systems that grew in shallow coral reef to intermediate and deeper fore-reef slope settings since the early Pleistocene. Age estimates based on 87 Sr/ 86 Sr isotopic measurements on corals, coralline algae, echinoids, and bulk sediments, although lacking the precision (∼±0.23 Ma) to distinguish the age–depth relationship and drowning times of individual reefs, indicate that the L12–L3 reefs range in age from ∼1.3–0.5 Ma and are therefore about 0.5–0.8 Ma older than the corresponding reefs around the flanks of Hawaii. These new age data, despite their lack of precision and the influence of later-stage submarine diagenesis on some analyzed corals, clearly revise the previous correlations between the reefs off Lanai and Hawaii. Soon after the end of major shield building (∼1.3–1.2 Ma), the Lanai reefs initiated growth and went through a period of rapid subsidence and reef drowning associated with glacial/interglacial cycles similar to that experienced by the Hawaii reefs. However, their early Pleistocene initiation means they experienced a longer, more complex growth history than their Hawaii counterparts.
Coral reef ecosystem management benefits from continual quantitative assessment of the resources being managed, plus assessment of factors that affect distribution patterns of organisms in the ecosystem. In this study, we investigate the relationships among physical, benthic, and fish variables in an effort to help explain the distribution patterns of organisms on patch reefs within Biscayne National Park, FL, USA. We visited a total of 196 randomly selected sampling stations on 12 shallow (<10 m) patch reefs and measured physical variables (e.g., substratum rugosity, substratum type) and benthic and fish community variables. We also incorporated data on substratum rugosity collected remotely via airborne laser surveying (Experimental Advanced Airborne Research Lidar—EAARL). Across all stations, only weak relationships were found between physical, benthic cover, and fish assemblage variables. Much of the variance was attributable to a “reef effect,” meaning that community structure and organism abundances were more variable at stations among reefs than within reefs. However, when the reef effect was accounted for and removed statistically, patterns were detected. Within reefs, juvenile scarids were most abundant at stations with high coverage of the fleshy macroalgae Dictyota spp., and the calcified alga Halimeda tuna was most abundant at stations with low EAARL rugosity. Explanations for the overwhelming importance of “reef” in explaining variance in our dataset could include the stochastic arrangement of organisms on patch reefs related to variable larval recruitment in space and time and/or strong historical effects due to patchy disturbances (e.g., hurricanes, fishing), as well as legacy effects of prior residents (“priority” effects).
A newly discovered sequence of lower to middle Miocene rocks from the eastern Bocas del Toro archipelago, western Panama, reveals the timing and environment of the earliest stages in the rise of the Isthmus of Panama in this region. Two new formations, the Punta Alegre Formation (lower Miocene, Aquitanian to Burdigalian) and the Valiente Formation (middle Miocene, Langhian to Serravallian), are here named and formally described. The Punta Alegre Formation contains a diagnostic microfauna of benthic and planktic foraminifera and calcareous nannofossils that indicate deposition in a 2000-m-deep pre-isthmian neotropical ocean from as old as 21.5–18.3 Ma. Its lithology varies from silty mudstone to muddy foraminiferal ooze with rare thin microturbidite layers near the top. The Valiente Formation, which ranges in age from 16.4 to ca. 12.0 Ma, lies with slight angular unconformity on the Punta Alegre Formation and consists of five lithofacies: (1) columnar basalt and flow breccia, (2) pyroclastic deposits, (3) coarse-grained volcaniclastic deposits, (4) coral-reef limestone with diverse large coral colonies, and (5) marine debris-flow deposits and microturbidites. These lithofacies are interpreted to indicate that after ca. 16 Ma a volcanic arc developed in the region of Bocas del Toro and that by ca. 12 Ma an extensively emergent archipelago of volcanic islands had formed. 39 Ar/ 40 Ar dating of basalt flows associated with the fossiliferous sedimentary rocks in the upper part of the Valiente Formation strongly confirms the ages derived from planktic foraminifera and nannofossils. Paleobathymetric analysis of the two new formations in the Valiente Peninsula and Popa Island, in the Bocas del Toro archipelago, shows a general shallowing from lower- through upper-bathyal to upper-neritic and emergent laharic and fluviatile deposits from ca. 19 to 12 Ma. The overlying nonconformable Bocas del Toro Group contains a lower transgressive sequence ranging from basal nearshore sandstone to upper-bathyal mudstone (ca. 8.1–5.3 Ma) and an upper regressive sequence (5.3–3.5 Ma). A similar paleobathymetric pattern is observed from the Gatun to Chagres Formations (12–6 Ma) in the Panama Canal Basin area and in the Uscari, Rio Banana, Quebrada Chocolate, and Moin Formations (8–1.7 Ma) in the southern Limón Basin of Costa Rica.
Climate plays a central role in coral-reef development, especially in marginal environments. The high-latitude reefs of southeast Florida are currently non-accreting, relict systems with low coral cover. This region also did not support the extensive Late Pleistocene reef development observed in many other locations around the world; however, there is evidence of significant reef building in southeast Florida during the Holocene. Using 146 radiometric ages from reefs extending ~ 120 km along Florida’s southeast coast, we test the hypothesis that the latitudinal extent of Holocene reef development in this region was modulated by climatic variability. We demonstrate that although sea-level changes impacted rates of reef accretion and allowed reefs to backstep inshore as new habitats were flooded, sea level was not the ultimate cause of reef demise. Instead, we conclude that climate was the primary driver of the expansion and contraction of Florida’s reefs during the Holocene. Reefs grew to 26.7° N in southeast Florida during the relatively warm, stable climate at the beginning of the Holocene Thermal Maximum (HTM) ~ 10,000 years ago, but subsequent cooling and increased frequency of winter cold fronts were associated with the equatorward contraction of reef building. By ~ 7800 years ago, actively accreting reefs only extended to 26.1° N. Reefs further contracted to 25.8° N after 5800 years ago, and by 3000 years ago reef development had terminated throughout southern Florida (24.5–26.7° N). Modern warming is unlikely to simply reverse this trend, however, because the climate of the Anthropocene will be fundamentally different from the HTM. By increasing the frequency and intensity of both warm and cold extreme-weather events, contemporary climate change will instead amplify conditions inimical to reef development in marginal reef environments such as southern Florida, making them more likely to continue to deteriorate than to resume accretion in the future.
Reef corals and the communities they form evidently possess effective mechanisms of adaptation and acclimation that have ensured their survival and recurrence over geologic time. Current reef degradation suggests that these mechanisms are being taxed beyond their limits; understanding of the problem is hampered by serious inadequacies in our understanding of physiological stress responses, the range and implications of reproductive strategies, and the mechanisms of calcification and algal symbiosis. Reef community and population responses to environmental change appear substantially different on different time scales, and a combination of short-term perspectives and definitional confusion complicates interpretation and prediction of reef responses. Calcium carbonate saturation state is now recognized as a potentially important control of reef calcification, which means that rising atmospheric CO 2 represents a direct threat to reef ecosystems on a global scale.
Coral disease is a growing problem for coral reefs globally and diseases have been linked to thermal stress, excess nutrients, overfishing and other human impacts. The Red Sea is a unique environment for corals with a strong environmental gradient characterized by temperature extremes and high salinities, but minimal terrestrial runoff or riverine input and their associated pollution. Yet, relatively little is known about coral diseases in this region. Disease surveys were conducted at 22 reefs within three regions (Yanbu, Thuwal, Al Lith) in the central Red Sea along the Saudi Arabian coast. Surveys occurred in October 2015, which coincided with a hyperthermal-induced bleaching event. Our objectives were to 1) document types, prevalence, and distribution of coral diseases in a region with minimal terrestrial input, 2) compare regional differences in diseases and bleaching along a latitudinal gradient of environmental conditions, and 3) use histopathology to characterize disease lesions at the cellular level. Coral reefs of the central Red Sea had a widespread but a surprisingly low prevalence of disease (<0.5%), based on the examination of >75,750 colonies. Twenty diseases were recorded affecting 16 coral taxa and included black band disease, white syndromes, endolithic hypermycosis, skeletal eroding band, growth anomalies and focal bleached patches. The three most common diseases were Acropora white syndrome (59.1% of the survey sites), Porites growth anomalies (40.9%), and Porites white syndrome (31.8%). Sixteen out of 30 coral genera within transects had lesions and Acropora , Millepora and Lobophyllia were the most commonly affected. Cell-associated microbial aggregates were found in four coral genera including a first report in Stylophora . Differences in disease prevalence, coral cover, amount of heat stress as measured by degree heating weeks (DHW) and extent of bleaching was evident among sites. Disease prevalence was not explained by coral cover or DHW, and a negative relationship between coral bleaching and disease prevalence was found. The northern-most sites off the coast of Yanbu had the highest average disease prevalence and highest average DHW values but no bleaching. Our study provides a foundation and baseline data for coral disease prevalence in the central Red Sea, which is projected to increase as a consequence of increased frequency and severity of ocean warming.
Coral-reef degradation is disrupting the balance between reef accretion and erosion and threatening the persistence of essential coral-reef habitats. In south Florida, most reefs are already net eroding, and without intervention, valuable ecosystem services may be lost. Coral restoration holds the potential to reverse those trends; however, typical restoration monitoring does not adequately capture key geo-ecological functions. We addressed this knowledge gap using carbonate budgets and Structure-from-Motion models to evaluate the impact of coral restoration on reef-accretion potential and structural complexity at eight offshore and three inshore coral reefs in the Lower Florida Keys. Within 2–6 years following outplanting, restoration of rapidly growing A. cervicornis populations increased reef-accretion potential to 2.8 mm y − 1 and drove significant increases in structural complexity. There was no measurable impact of restoring slower-growing, massive corals on reef-accretion potential inshore; however, whereas the severe 2023 coral-bleaching event immediately following our study caused near-complete mortality of A. cervicornis , 59% of massive corals survived, highlighting potential trade-offs between coral growth and survival on future restoration efficacy. We conclude that although restoration can produce rapid, small-scale increases in reef-accretion potential, there remain important uncertainties about how and whether ecosystem-scale benefits of restoration on important geo-ecological reef functions can persist long term.