Extrinsic control of species replacement on a Holocene reef in Belize: the role of coral disease

 Well-preserved, Holocene coral reefs provide the opportunity to discriminate between models of intrinsically driven succession and extrinsically driven species replacement, especially when paleontological patterns can be combined with ecological observations of the underlying mechanisms. Rhomboid shoals in the central shelf lagoon of the Belizean Barrier Reef experienced a recent and dramatic change in community composition. Agaricia tenuifolia replaced Acropora cervicornis as the dominant coral species at 3–15 m depth along the flanks of the reefs. We tested the hypothesis that shallowing upward caused this shift in dominance. A core extracted from 0.5 m water depth on one of the shoals, Channel Cay, revealed a shallowing-upward shift in dominance from Acropora to Porites divaricata. This successional sequence was quite different from the Acropora-to-Agaricia transition observed in four cores from 6–11 m water depth. Ecological observations showed that Agaricia became the dominant at ≥3 m depth after Acropora populations were decimated by a regional outbreak of white-band disease. The Acropora-to-Agaricia transition was clearly a case of extrinsically driven species replacement rather than an intrinsically driven, successional, shallowing-upward sequence. Accepted: 14 May 1998     

Macroalgae reduces survival of nursery‐reared Acropora corals in the Florida reef tract

Recent declines in coral populations along the Florida reef tract have prompted the establishment of coral restoration programs which raise coral species, such as the threatened Acropora cervicornis, in nurseries ready for outplanting. Large numbers of nursery‐reared coral colonies have been outplanted along the Florida reef tract in recent years, yet few studies have characterized benthic habitats that are considered optimal for colony survival. In 2016, we surveyed 23 A. cervicornis restoration sites, located at six different reefs in the upper Florida Keys. We examined the condition of the outplanted corals and quantified the benthic assemblages adjacent to the outplanted coral colonies. We found that where A. cervicornis survived for more than 1 year, the substrate significantly supported less brown macroalgae of the genus Dictyota than at sites where A. cervicornis had died. Coral survival was highest at sites with less than 15% Dictyota cover. These results suggest that the habitat conditions that supported Dictyota spp. were not conducive to A. cervicornis growth and survival. Restoration practitioners should avoid attaching nursery‐raised corals to substrate with Dictyota spp. cover greater than 15%.     

FIRST RECORD OF THE INDO‐PACIFIC REEF CORAL GENUS ISOPORA IN THE CARIBBEAN REGION: TWO NEW SPECIES FROM THE NEOGENE OF CURAÇAO,NETHERLANDS ANTILLES

Abstract: The coral genus Isopora, a sister group of the modern dominant Acropora until now only known from the Pliocene to Recent of the Indo‐Pacific, is recorded in the Caribbean for the first time. Two new species, Isopora ginsburgi and Isopora curacaoensis, are described from the Neogene Seroe Domi Formation of Curaçao, Netherlands Antilles. Study of large collections made systematically through the sequence indicates that Isopora first occurred in the Caribbean during the Mio–Pliocene, at approximately the same time as the origination of many modern Caribbean reef coral dominants including Acropora cervicornis. It last occurred in the region during the late Pliocene as part of a pulse of extinction, in which several genera that live today in the Indo‐Pacific became extinct in the Caribbean. Throughout its Caribbean duration, Isopora co‐occurred with the two abundant modern Caribbean species of Acropora, A. cervicornis and A. palmata. Comparisons with Neogene collections made elsewhere in the Caribbean indicate that Isopora was restricted in distribution to the southern Caribbean. Isopora species are viviparous, while Acropora are oviparous, and this difference in reproductive strategy may have played a role in the extinction of Isopora in the Caribbean. The occurrences of Isopora reported in this study are the oldest records to date of Isopora worldwide, and are important for understanding the biogeographic separation between reef coral faunas in the Caribbean and Indo‐Pacific regions.     

Rapid phase-shift reversal on a Jamaican coral reef

Many Caribbean reefs have experienced a phase-shift in community structure, the principle features being a decline in coral cover and an increase in macroalgal biomass. However, one Jamaican reef—Dairy Bull on the north shore near Discovery Bay—is once again dominated by scleractinian corals and several key species have returned. Living coral cover at 6–8 m depth at Dairy Bull has doubled over the past 9 years and is now ~54%. The absolute cover of Acropora cervicornis was <1% in 1995, but increased to ~11% by January 2004. During this time the cover of macroalgae decreased by 90%, from 45 to 6%. We speculate that long-lived colonies of Montastraea annularis may have facilitated the recovery of this reef by providing structural refugia.     

An analysis of the loss of acroporid corals at Looe Key,Florida, USA: 1983–2000

The Caribbean reef-building corals Acropora palmata and Acropora cervicornis have undergone widespread declines in the past two decades, leading to their designation as candidates for listing under the United States Endangered Species Act. Whole-reef censuses in 1983 and 2000 at Looe Key National Marine Sanctuary in the Florida Keys provide estimates of areal loss of live Acropora spp. cover. Area (square meters) of live coral cover was quantified from depiction on scaled base maps of extent of coral cover observed by a snorkeler on each reef spur at each census. Certain thickets appear to have been persistent (though none expanded), but the total area of live A. palmata at Looe Key is estimated to have declined by 93% and A. cervicornis by 98% during this 17-year interval. It is likely that acroporid populations may have already undergone substantial decline prior to our initial census in 1983.     

Millennial-scale dynamics of staghorn coral in Discovery Bay, Jamaica

Populations of the staghorn coral, Acropora cervicornis, collapsed throughout the Caribbean region from the late 1970s through the 1990s. We tested the hypothesis that this recent, multidecadal interruption in coral growth was a novel event in the late Holocene. Eight cores, extracted from a lagoonal reef in Discovery Bay, Jamaica dated to 440–1260 CalBP and consisted almost entirely of A. cervicornis rubble. The A. cervicornis in the cores showed significantly less internal bioerosion than A. cervicornis from modern death assemblages in Discovery Bay, indicating generally shorter post‐mortem exposure at the sediment–water interface in the past. A. cervicornis grew continuously and was buried rapidly during the millennium preceding the 1980s, with the exception of a possible hiatus in growth and burial at some point 300–600 years ago. In the 1980s, a combination of perturbations, which included overfishing and (possibly) other forms of human interference, caused an unprecedented disruption in the growth and burial of staghorn coral populations in Discovery Bay.     

Algal contact as a trigger for coral disease

Diseases are causing alarming declines in reef‐building coral species, the foundation blocks of coral reefs. The emergence of these diseases has occurred simultaneously with large increases in the abundance of benthic macroalgae. Here, we show that physical contact with the macroalga Halimeda opuntia can trigger a virulent disease known as white plague type II that has caused widespread mortality in most Caribbean coral species. Colonies of the dominant coral Montastraea faveolata exposed to algal transplants developed the disease whereas unexposed colonies did not. The bacterium Aurantimonas coralicida, causative agent of the disease, was present on H. opuntia sampled close to, and away from diseased corals, indicating that the alga serves as a reservoir for this pathogen. Our results suggest that the spread of macroalgae on coral reefs could account for the elevated incidence of coral diseases over past decades and that reduction of macroalgal abundance could help control coral epizootics.     

Shifting ecological baselines and the demise of Acropora cervicornis in the western North Atlantic and Caribbean Province: a Pleistocene perspective

 In recent years, marine scientists have become increasingly alarmed over the decline of live coral cover throughout the Caribbean and tropical western Atlantic region. The Holocene and Pleistocene fossil record of coral reefs from this region potentially provides a wealth of long-term ecologic information with which to assess the historical record of changes in shallow water coral reef communities. Before fossil data can be applied to the modern reef system, critical problems involving fossil preservation must be addressed. Moreover, it must be demonstrated that the classic reef coral zonation patterns described in the early days of coral reef ecology, and upon which “healthy” versus “unhealthy” reefs are determined, are themselves representative of reefs that existed prior to any human influence. To address these issues, we have conducted systematic censuses of life and death assemblages on modern “healthy” patch reefs in the Florida reef tract that conform to the classic Caribbean model of reef coral zonation, and a patch reef in the Bahamas that is currently undergoing a transition in coral dominance that is part of a greater Caribbean-wide phenomenon. Results were compared to censuses of ancient reef assemblages preserved in Pleistocene limestones in close proximity to each modern reef. We have determined that the Pleistocene fossil record of coral reefs may be used to calibrate an ecological baseline with which to compare modern reef assemblages, and suggest that the current and rapid decline of Acropora cervicornis observed on a Bahamian patch reef may be a unique event that contrasts with the long-term persistence of this taxon during Pleistocene and Holocene time. Accepted: 19 May 1998     

In Situ Coral Nurseries Serve as Genetic Repositories for Coral Reef Restoration after an Extreme Cold‐Water Event

During an unusual cold‐water event in January 2010, reefs along the Florida Reef Tract suffered extensive coral mortality, especially in shallow reef habitats in close proximity to shore and with connections to coastal bays. The threatened staghorn coral, Acropora cervicornis, is the focus of propagation and restoration activities in Florida and one of the species that exhibited high susceptibility to low temperatures. Complete mortality of wild staghorn colonies was documented at 42.9% of donor sites surveyed after the cold event. Remarkably, 72.7% of sites with complete A. cervicornis mortality had fragments surviving within in situ coral nurseries. Thus, coral nurseries served as repositories for genetic material that would have otherwise been completely lost from donor sites. The location of the coral nurseries at deeper habitats and distanced from shallow nearshore habitats that experienced extreme temperature conditions buffered the impacts of the cold‐water event and preserved essential local genotypes for future Acropora restoration activities.     

Evaluating Methods for Transplanting Endangered Elkhorn Corals in the Virgin Islands

Restoration of rare corals is desirable and restoration projects are fairly common, but scientific evaluation of this approach is limited. We tested several techniques for transplant and restabilization of Acropora palmata (the elkhorn coral), an ecologically important Caribbean coral whose populations have suffered severe declines. In rough weather, fragments break‐off colonies of branching corals like A. palmata as a normal form of asexual reproduction. We transplanted naturally produced coral fragments from remnant populations to nearby restoration sites. Untouched control fragments at the donor site died faster and grew slower than fragments attached to the reef, so attaching fragments to the reef is beneficial. Transplanted fragments grew and died at a rate similar to fragments left at the donor site (both groups were attached to the reef), so there were no effects of moving fragments or differences in habitat quality between donor and restoration sites. Growth and survival were similar using four methods of attaching fragments to the reef: cable ties, two types of epoxy resin, and hydrostatic cement. Corals sometimes compete with the macroalgae that dominate degraded reefs, and clearing surrounding algae improved the growth of fragments. After 4 years, transplanted fragments grew to 1,450 cm² in area and so were potentially sexually active. Because the methods tested are simple and cheap, they could be used by volunteer recreational divers to restore locally extirpated A. palmata populations or to enhance reefs where natural recovery is slow.     

Regional‐scale dominance of non‐framework building corals on Caribbean reefs affects carbonate production and future reef growth

Coral cover on Caribbean reefs has declined rapidly since the early 1980's. Diseases have been a major driver, decimating communities of framework building Acropora and Orbicella coral species, and reportedly leading to the emergence of novel coral assemblages often dominated by domed and plating species of the genera Agaricia, Porites and Siderastrea. These corals were not historically important Caribbean framework builders, and typically have much smaller stature and lower calcification rates, fuelling concerns over reef carbonate production and growth potential. Using data from 75 reefs from across the Caribbean we quantify: (i) the magnitude of non‐framework building coral dominance throughout the region and (ii) the contribution of these corals to contemporary carbonate production. Our data show that live coral cover averages 18.2% across our sites and coral carbonate production 4.1 kg CaCO₃ m^?2 yr^?1. However, non‐framework building coral species dominate and are major carbonate producers at a high proportion of sites; they are more abundant than Acropora and Orbicella at 73% of sites; contribute an average 68% of the carbonate produced; and produce more than half the carbonate at 79% of sites. Coral cover and carbonate production rate are strongly correlated but, as relative abundance of non‐framework building corals increases, average carbonate production rates decline. Consequently, the use of coral cover as a predictor of carbonate budget status, without species level production rate data, needs to be treated with caution. Our findings provide compelling evidence for the Caribbean‐wide dominance of non‐framework building coral taxa, and that these species are now major regional carbonate producers. However, because these species typically have lower calcification rates, continued transitions to states dominated by non‐framework building coral species will further reduce carbonate production rates below ‘predecline’ levels, resulting in shifts towards negative carbonate budget states and reducing reef growth potential.     

Coral reef crisis in deep and shallow reefs: 30 years of constancy and change in reefs of Curacao and Bonaire

Coral reefs are thought to be in worldwide decline but available data are practically limited to reefs shallower than 25 m. Zooxanthellate coral communities in deep reefs (30–40 m) are relatively unstudied. Our question is: what is happening in deep reefs in terms of coral cover and coral mortality? We compare changes in species composition, coral mortality, and coral cover at Caribbean (Curacao and Bonaire) deep (30–40 m) and shallow reefs (10–20 m) using long-term (1973–2002) data from permanent photo quadrats. About 20 zooxanthellate coral species are common in the deep-reef communities, dominated by Agaricia sp., with coral cover up to 60%. In contrast with shallow reefs, there is no decrease in coral cover or number of coral colonies in deep reefs over the last 30 years. In deep reefs, non-agaricid species are decreasing but agaricid domination will be interrupted by natural catastrophic mortality such as deep coral bleaching and storms. Temperature is a vastly fluctuating variable in the deep-reef environment with extremely low temperatures possibly related to deep-reef bleaching. An erratum to this article can be found at     

Genetic Diversity and Connectivity in the Threatened Staghorn Coral (Acropora cervicornis) in Florida

Over the past three decades, populations of the dominant shallow water Caribbean corals, Acropora cervicornis and A. palmata, have been devastated by white-band disease (WBD), resulting in the listing of both species as threatened under the U.S. Endangered Species Act. A key to conserving these threatened corals is understanding how their populations are genetically interconnected throughout the greater Caribbean. Genetic research has demonstrated that gene flow is regionally restricted across the Caribbean in both species. Yet, despite being an important site of coral reef research, little genetic data has been available for the Florida Acropora, especially for the staghorn coral, A. cervicornis. In this study, we present new mitochondrial DNA sequence data from 52 A. cervicornis individuals from 22 sites spread across the upper and lower Florida Keys, which suggest that Florida''s A. cervicornis populations are highly genetically interconnected (F_ST = −0.081). Comparison between Florida and existing mtDNA data from six regional Caribbean populations indicates that Florida possesses high levels of standing genetic diversity (h = 0.824) relative to the rest of the greater Caribbean (h = 0.701±0.043). We find that the contemporary level of gene flow across the greater Caribbean, including Florida, is restricted ( = 0.117), but evidence from shared haplotypes suggests the Western Caribbean has historically been a source of genetic variation for Florida. Despite the current patchiness of A. cervicornis in Florida, the relatively high genetic diversity and connectivity within Florida suggest that this population may have sufficient genetic variation to be viable and resilient to environmental perturbation and disease. Limited genetic exchange across regional populations of the greater Caribbean, including Florida, indicates that conservation efforts for A. cervicornis should focus on maintaining and managing populations locally rather than relying on larval inputs from elsewhere.     

Overgrowth and killing of corals by the brown alga Lobophora hederacea (Dictyotales,Phaeophyceae) on healthy reefs in New Caledonia: A new case of the epizoism syndrome

Coral reef degradation is often associated with regime shifts from coral‐ to macroalgal‐dominated reefs. These shifts demonstrate that under certain conditions (e.g. coral mortality, decrease in herbivory, increased nutrients supply) some macroalgae may overgrow corals. The outcome of the competition is dependent on algal aggressiveness and the coral susceptibility. In undisturbed reefs, herbivore grazing is regulating macroalgal cover, thus preventing the latter from overgrowing corals. However, some macroalgae have evolved strategies not only to outcompete corals but also to escape herbivory to some extent, allowing overgrowth of some coral species in undisturbed reefs. Epizoism represents one of those successful strategies, and has been previously documented with red algae, cyanobacteria and Lobophora variegata (Dictyotales, Phaeophyceae). Here we report a new case of epizoism leading to coral mortality, involving a recently described species of Lobophora, L. hederacea, overgrowing the coral Seriatopora caliendrum (Pocilloporidae) in undisturbed reefs in New Caledonia.     

Differential survival of nursery‐reared Acropora cervicornis outplants along the Florida reef tract

In recent decades, the Florida reef tract has lost over 95% of its coral cover. Although isolated coral assemblages persist, coral restoration programs are attempting to recover local coral populations. Listed as threatened under the Endangered Species Act, Acropora cervicornis is the most widely targeted coral species for restoration in Florida. Yet strategies are still maturing to enhance the survival of nursery‐reared outplants of A. cervicornis colonies on natural reefs. This study examined the survival of 22,634 A. cervicornis colonies raised in nurseries along the Florida reef tract and outplanted to six reef habitats in seven geographical subregions between 2012 and 2018. A Cox proportional hazards regression was used within a Bayesian framework to examine the effects of seven variables: (1) coral‐colony size at outplanting, (2) coral‐colony attachment method, (3) genotypic diversity of outplanted A. cervicornis clusters, (4) reef habitat, (5) geographical subregion, (6) latitude, and (7) the year of monitoring. The best models included coral‐colony size at outplanting, reef habitat, geographical subregion, and the year of monitoring. Survival was highest when colonies were larger than 15 cm (total linear extension), when outplanted to back‐reef and fore‐reef habitats, and when outplanted in Biscayne Bay and Broward–Miami subregions, in the higher latitudes of the Florida reef tract. This study points to several variables that influence the survival of outplanted A. cervicornis colonies and highlights a need to refine restoration strategies to help restore their population along the Florida reef tract.     

Changes in Coral Reef Communities and an Associated Reef Fish Species, <Emphasis Type="Italic">Cephalopholis cruentata</Emphasis> (Lacépède), After 30 years on Curaçao (Netherlands Antilles)

Using the same methodology and identical sites, we repeat a study dating from 1973 and quantify cover of hard coral species, soft corals, sponges, hard substratum and soft substratum, and density of a commercially important reef fish species, the graysby Cephalopholis cruentata, along a depth-gradient of 3–36 m on the coral reefs of Curaçao. The objective was to determine the multi-decade change in benthic coral reef cover and structural complexity, and their effect on densities of an associated reef fish species. Total hard coral cover decreased on average from 52% in 1973 to 22% in 2003, representing a relative decline of 58%. During this time span, the cover of hard substratum increased considerably (from 11 to 58%), as did that of soft corals (from 0.1 to 2.2%), whereas the cover of sponges showed no significant change. Relative decline of hard coral cover and of reef complexity was greatest in shallow waters (near the coast), which is indicative of a combination of anthropogenic influences from shore and recent storm damage. Cover of main reef builder coral species (Agaricia spp., Siderastrea siderea, Montastrea annularis) decreased more than that of other species, and resulted in a significant decrease in reef complexity. Although density of C. cruentata was highly correlated to cover of Montastrea and Agaricia in 1973, the loss of coral cover did not show any effect on the total density of C. cruentata in 2003. However, C. cruentata showed a clear shift in density distribution from shallow water in 1973 to deep water in 2003. It can be concluded that the reefs of Curaçao have degraded considerably in the last three decades, but that this has had no major effect on the population size of one commercially important coral-associated fish species.     

Distribution and abundance of elkhorn coral, Acropora palmata, and prevalence of white-band disease at Buck Island Reef National Monument, St. Croix, US Virgin Islands

In the 1970s and 1980s elkhorn coral, Acropora palmata, declined dramatically throughout the Caribbean primarily due to white-band disease (WBD). In 2005, elkhorn coral was proposed for listing as threatened under the US Endangered Species Act. WBD was first documented at Buck Island Reef National Monument (BIRNM). Together with hurricanes WBD reduced live elkhorn coral coverage by probably over 90%. In the past decade some recovery has been observed at BIRNM. This study assessed the distribution and abundance of elkhorn coral and estimated the prevalence of WBD at the monument. Within an area of 795 ha, we estimated 97,232–134,371 (95% confidence limits) elkhorn coral colonies with any dimension of connected live tissue greater than one meter, about 3% of which were infected by WBD. Despite some recovery, the elkhorn coral density remains low and WBD may continue to present a threat to the elkhorn coral population.     

Spatial patterns and ecology of benthic communities on a high-latitude South Florida (Broward County,USA) reef system

High-latitude reef communities consisting of typical Caribbean fauna of variable composition and density exist on four parallel ridges at varying depths along the Broward County (Florida, USA) coast. Two of these ridges, at 7–13 and 15–30 m in depth, are drowned early Holocene coral reefs of 5 and 7 ky uncorrected radiocarbon age. In this study, community data were collected on each reef using 50-m, line-intercept transects. Using multivariate non-metric statistics, the data show distinct differences in benthic community structure across several spatial gradients. Diversity indices revealed that while all values were low, species diversity (H) was consistent throughout the county, and species richness (d) and evenness (J) increased along a north-south gradient. Scleractinian coral (27 species) cover was low (<6%) in all areas, and Montastrea cavernosa dominated as the major hermatypic scleractinian. Notably absent was the major Caribbean reef-builder Acropora palmata. A rich alcyonacean fauna was present on all studied reefs, and these were typically the most important faunal group determining community structure. Although during the 2-year period 2000–2002 mean water temperatures never fell below 20 °C, scleractinia were small in size.     

棘冠海星暴发及其对珊瑚礁的生态影响研究进展

棘冠海星的反复暴发是导致印度—太平洋区域珊瑚礁生态系统退化的最主要原因之一。然而,我国对棘冠海星的研究非常有限。本文综述了国内外关于棘冠海星及其暴发的生态影响和应对策略的研究进展,得出以下主要结论：1)雌性棘冠海星个体每年产卵数量高达50万—2亿个,环境因素变化只要导致幼虫和幼体存活率的轻微提高,成体就将得到大量补充;2)棘冠海星暴发的阈值为1000—1500个/km²,暴发周期为10—27 a,每次暴发持续1—10 a,最终可能以“种群集体感染疾病而崩溃”结束;3)棘冠海星暴发对印度洋及太平洋东部和北部珊瑚礁的破坏性非常小,却直接导致太平洋的西部和南部珊瑚礁90%以上的珊瑚死亡,并通过改变珊瑚群落组成、减少珊瑚和鱼类多样性而对珊瑚礁产生间接影响;4)关于棘冠海星暴发原因的假说中“陆地营养物质输入假说”和“捕食者过度捕捞假说”得到了最普遍的认可,但都不能解释所有的暴发事件;5)应对棘冠海星暴发的主要策略有改善水质、设立保护区、投放天敌和人工清理等,其中人工清理是最直接有效的策略,但迄今并没有发现可长期抑制棘冠海星暴发的方法。因此,急需加强对棘冠海星的深入研究,探查...     

Coral population dynamics across consecutive mass mortality events

Annual coral mortality events due to increased atmospheric heat may occur regularly from the middle of the century and are considered apocalyptic for coral reefs. In the Arabian/Persian Gulf, this situation has already occurred and population dynamics of four widespread corals (Acropora downingi, Porites harrisoni, Dipsastrea pallida, Cyphastrea micropthalma) were examined across the first‐ever occurrence of four back‐to‐back mass mortality events (2009–2012). Mortality was driven by diseases in 2009, bleaching and subsequent diseases in 2010/2011/2012. 2009 reduced P. harrisoni cover and size, the other events increasingly reduced overall cover (2009: ?10%; 2010: ?20%; 2011: ?20%; 2012: ?15%) and affected all examined species. Regeneration was only observed after the first disturbance. P. harrisoni and A. downingi severely declined from 2010 due to bleaching and subsequent white syndromes, while D. pallida and P. daedalea declined from 2011 due to bleaching and black‐band disease. C. microphthalma cover was not affected. In all species, most large corals were lost while fission due to partial tissue mortality bolstered small size classes. This general shrinkage led to a decrease of coral cover and a dramatic reduction of fecundity. Transition matrices for disturbed and undisturbed conditions were evaluated as Life Table Response Experiment and showed that C. microphthalma changed the least in size‐class dynamics and fecundity, suggesting they were ‘winners’. In an ordered ‘degradation cascade’, impacts decreased from the most common to the least common species, leading to step‐wise removal of previously dominant species. A potentially permanent shift from high‐ to low‐coral cover with different coral community and size structure can be expected due to the demographic dynamics resultant from the disturbances. Similarities to degradation of other Caribbean and Pacific reefs are discussed. As comparable environmental conditions and mortality patterns must be expected worldwide, demographic collapse of many other coral populations may soon be widespread.