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Coral restoration is widely used around the world to address dramatic declines in coral cover; however, very few studies have looked specifically at the temporal response of fish assemblages (i.e. abundance and diversity) to coral restoration. Several critical reef functions and processes are driven by fishes, thereby making their recovery and responses around restoration structures key indicators of success. This study evaluates fish abundance and community composition on restoration plots following 8–12 years of restoration activity, in four locations (two Caribbean and two Indo‐Pacific). Responses were very complex with region‐, site‐, and body size‐specific patterns. Overall, fish abundance only increased in Indo‐Pacific sites where damselfish responded positively to increased coral cover and topographic complexity. Restoration effects on other fish families and particularly on larger bodied reef fish were negative or neutral at all locations. If restoration initiatives are going to substantively improve the condition and recovery of degraded reef fish communities, restoration efforts need to be planned, designed, and monitored based on fish‐specific habitat requirements and locally specific community dynamics.

Research into causative agents underlying coral disease have focused primarily on bacteria, whereas potential roles of viruses have been largely unaddressed. Bacteriophages may contribute to diseases through the lysogenic introduction of virulence genes into bacteria, or prevent diseases through lysis of bacterial pathogens. To identify candidate phages that may influence the pathogenicity of black band disease (BBD), communities of bacteria (16S rRNA) and T4-bacteriophages (gp23) were simultaneously profiled with amplicon sequencing among BBD-lesions and healthy-coral-tissue of Montipora hispida, as well as seawater (study site: the central Great Barrier Reef). Bacterial community compositions were distinct among BBD-lesions, healthy coral tissue and seawater samples, as observed in previous studies. Surprisingly, however, viral beta diversities based on both operational taxonomic unit (OTU)-compositions and overall viral community compositions of assigned taxa did not differ statistically between the BBD-lesions and healthy coral tissue. Nonetheless, relative abundances of three bacteriophage OTUs, affiliated to Cyanophage PRSM6 and Prochlorococcus phages P-SSM2, were significantly higher in BBD-lesions than in healthy tissue. These OTUs associated with BBD samples suggest the presence of bacteriophages that infect members of the cyanobacteria-dominated BBD community, and thus have potential roles in BBD pathogenicity.

Projected increases in sea surface temperatures will exceed corals' ability to withstand heat stress within this century. Experimental evolution of cultured symbionts (Symbiodiniaceae) at high temperatures followed by reintroduction into corals can enhance coral heat tolerance. Several studies have selected for enhanced tolerance in Cladocopium goreaui (C1) over multiple time scales and then compared the performance of coral juveniles infected with the heat-tolerant C1 selected strain (SS) to the performance of juveniles infected with the C1 wild type (WT). To derive lessons about host benefits when symbionts are experimentally selected, here we compare the performance of SS- and WT-juveniles after 21 cell generations of heat selection versus longer periods (73–131) in recently published experiments. After 21 generations, we found rapid improvement in heat tolerance of SS through an overall shift in the mean tolerance to temperature. This did not translate to improved growth and survivorship of the coral. Specifically, survival did not differ significantly between juveniles of Acropora tenuis hosting WT versus SS at any temperature. Juveniles infected with WT exhibited greater skeletal growth than those infected with SS at 27 and 31°C but not at 32.5°C. SS-juvenile symbiont cell densities increased significantly at 27°C relative to SS-juveniles in the 31 and 32.5°C. Photosynthetic efficiencies in SS-juveniles were higher compared to WT-juveniles at 31°C, equal at 27°C, and lower at 32.5°C. These results suggest that selection over longer generation (>130) times will be needed to confer host benefits and will be dependent on the stability of this association being maintained in nature.

Global increases in coral disease prevalence have been linked to ocean warming through changes in coral-associated bacterial communities, pathogen virulence and immune system function. However, the interactive effects of temperature and pathogens on the coral holobiont are poorly understood. Here, we assessed three compartments of the holobiont (host, Symbiodinium and bacterial community) of the coral Montipora aequituberculata challenged with the pathogen Vibrio coralliilyticus and the commensal bacterium Oceanospirillales sp. under ambient (27°C) and elevated (29.5 and 32°C) seawater temperatures. Few visual signs of bleaching and disease development were apparent in any of the treatments, but responses were detected in the holobiont compartments. V. coralliilyticus acted synergistically and negatively impacted the photochemical efficiency of Symbiodinium at 32°C, while Oceanospirillales had no significant effect on photosynthetic efficiency. The coral, however, exhibited a minor response to the bacterial challenges, with the response towards V. coralliilyticus being significantly more pronounced, and involving the prophenoloxidase-activating system and multiple immune system-related genes. Elevated seawater temperatures did not induce shifts in the coral-associated bacterial community, but caused significant gene expression modulation in both Symbiodinium and the coral host. While Symbiodinium exhibited an antiviral response and upregulated stress response genes, M. aequituberculata showed regulation of genes involved in stress and innate immune response processes, including immune and cytokine receptor signalling, the complement system, immune cell activation and phagocytosis, as well as molecular chaperones. These observations show that M. aequituberculata is capable of maintaining a stable bacterial community under elevated seawater temperatures and thereby contributes to preventing disease development.

Coral reef restoration initiatives are burgeoning in response to the need for novel management strategies to address dramatic global declines in coral cover. However, coral restoration programs typically lack rigor and critical evaluation of their effectiveness. A review of 83 peer-reviewed papers that used coral transplantation for reef restoration reveals that growth and survival of coral fragments were the most widely used indicators of restoration success, with 88% of studies using these two indicators either solely (55%) or in combination with a limited number of other ecological factors (33%). In 53% of studies, reef condition was monitored for 1 year or less, while only 5% of reefs were monitored for more than 5 years post-transplantation. These results highlight that coral reef restoration science has focused primarily on short-term experiments to evaluate the feasibility of techniques for ecological restoration and the initial establishment phase post-transplantation, rather than on longer-term outcomes for coral reef communities. Here, we outline 10 socioecological indicators that comprehensively evaluate the effectiveness of coral reef restoration across the four pillars of sustainability (i.e. environmental, sociocultural, governance, and economic contributions to sustainable communities). We recommend that evaluations of the effectiveness of coral restoration programs integrate ecological indicators with sociocultural, economic, and governance considerations. Assessing the efficacy of coral restoration as a tool to support reef resilience will help to guide future efforts and ensure the sustainable maintenance of reef ecosystem goods and services.

Despite the ecological significance of the relationship between reef-building corals and intracellular photosynthetic dinoflagellates of the genus Symbiodinium, very little is known about the molecular mechanisms involved in its establishment. Indeed, microarray-based analyses point to the conclusion that host gene expression is largely or completely unresponsive during the establishment of symbiosis with a competent strain of Symbiodinium. In this study, the use of Illumina RNA-Seq technology allowed detection of a transient period of differential expression involving a small number of genes (1073 transcripts

Symbiodinium are a diverse group of unicellular dinoflagellates that are important nutritional symbionts of reef-building corals. Symbiodinium putative species ('types') are commonly identified with genetic markers, mostly nuclear and chloroplast encoded ribosomal DNA regions. Population genetic analyses using microsatellite loci have provided insights into Symbiodinium biogeography, connectivity and phenotypic plasticity, but are complicated by: (i) a lack of consensus criteria used to delineate inter- vs. intragenomic variation within species; and (ii) the high density of Symbiodinium in host tissues, which results in single samples comprising thousands of individuals. To address this problem, Wham & LaJeunesse (2016) present a method for identifying cryptic Symbiodinium species from microsatellite data based on correlations between allele size distributions and nongeographic genetic structure. Multilocus genotypes that potentially do not recombine in sympatry are interpreted as secondary 'species' to be discarded from downstream population genetic analyses. However, Symbiodinium species delineations should ideally incorporate multiple physiological, ecological and molecular criteria. This is because recombination tests may be a poor indicator of species boundaries in Symbiodinium due to their predominantly asexual mode of reproduction. Furthermore, discontinuous microsatellite allele sizes in sympatry may be explained by secondary contact between previously isolated populations and by mutations that occur in a nonstepwise manner. Limitations of using microsatellites alone to delineate species are highlighted in earlier studies that demonstrate occasional bimodal distributions of allele sizes within Symbiodinium species and considerable allele size sharing among Symbiodinium species. We outline these issues and discuss the validity of reinterpretations of our previously published microsatellite data from Symbiodinium populations on the Great Barrier Reef (Howells et al. 2013).

Emerging infectious diseases are contributing to global declines in coral reef ecosystems, highlighting a growing need for aetiological knowledge to develop effective management strategies. In this review, we focus on black band disease (BBD), one of the most virulent diseases and the only polymicrobial disease so far known to affect corals. A multipartite microbial consortium dominated by Cyanobacteria, but also including sulfur-cycling bacteria, other bacterial groups and members of the Archaea and Eukarya, forms a sulfide-rich anaerobic mat that migrates across the surface of coral colonies, killing the underlying tissues. The polymicrobial nature of the disease challenges classic aetiological approaches to unravelling disease causation. Here, we synthesize current knowledge on the range of pathogens forming the microbial consortium with recent studies on the transmission, biogeochemistry and environmental drivers of BBD to develop a conceptual model of BBD pathogenesis. The model illustrates how the development of BBD virulence factors is linked to a cascade of microbial community shifts and associated functional roles that progressively develop the microbial consortium from comparatively benign cyanobacterial patches to virulent BBD lesions. This review showcases how an approach that integrates multiple key aspects of the disease provides insights essential to elucidating the aetiology of BBD.

Aquaculture of coral offers an alternative to wild harvest for the ornamental trade and shows considerable promise for restoring reefs and preserving biodiversity. Here, we compare advantages and disadvantages of asexually derived fragments versus sexually derived propagules and in situ versus ex situ nursery phases for the ornamental trade and reef restoration. Asexual propagules, sourced from a donor coral colony that is cut into smaller parts and attached to artificial substrate, are most commonly used. The most suitable corals are typically branching species, although fragments from species with other growth forms can be successful, albeit slower growing. Sexually derived propagules are collected from the wild or from colonies in aquaria during spawning, with an artificial substrate provided for settlement. The timing of spawning is known for many broadcast spawning corals, but opportunities for collection of gametes are generally limited to only once or a few times per year. Brooding species with multiple periods of larval release provide better options for culture of sexually derived propagules. Propagation techniques have developed considerably over the past 20 years, yielding faster growth rates, reduced mortality and reduced detachment from substrates. Simple and cost–effective propagation techniques can be used to restore denuded reefs, preserve endangered species, provide live corals to the international ornamental trade, enable livelihood diversification for coastal communities and provide experimental materials for marine research. This review provides a comprehensive synthesis of recent developments in aquaculture propagation techniques for the purpose of ornamental trade and coral reef restoration, including asexual and sexual propagation, nursery and transplantation stages.

Parks and protected areas have been instrumental in reducing anthropogenic sources of damage in terrestrial and aquatic environments. Pathogen invasion often succeeds physical wounding and injury, yet links between the reduction of damage and the moderation of disease have not been assessed. Here, we examine the utility of no-take marine reserves as tools for mitigating diseases that affect reef-building corals. We found that sites located within reserves had fourfold reductions in coral disease prevalence compared to non-reserve sites (80 466 corals surveyed). Of 31 explanatory variables assessed, coral damage and the abundance of derelict fishing line best explained differences in disease assemblages between reserves and non-reserves. Unexpectedly, we recorded significantly higher levels of disease, coral damage, and derelict fishing line in non-reserves with fishing gear restrictions than in those without gear restrictions. Fishers targeting stocks perceived to be less depleted, coupled with enhanced site access from immediately adjacent boat moorings, may explain these unexpected patterns. Significant correlations between the distance from mooring sites and prevalence values for a ciliate disease known to infest wounded tissue (r = −0.65), coral damage (r = −0.64), and the abundance of derelict fishing line (r = −0.85) corroborate this interpretation. This is the first study to link disease with recreational use intensity in a park, emphasizing the need to evaluate the placement of closures and their direct relationship to ecosystem health. Since corals are modular, ecological processes that govern reproductive and competitive fitness are frequently related to colony surface area therefore, even low levels of cumulative tissue loss from progressing diseases pose significant threats to reef coral persistence. Disease mitigation through reductions in physical injury in areas where human activities are concentrated is another mechanism by which protected areas may improve ecosystem resilience in a changing climate.

Mounting evidence of cryptic species in a wide range of taxa highlights the need for careful analyses of population genetic data sets to unravel within-species diversity from potential interspecies relationships. Here, we use microsatellite loci and hierarchical clustering analysis to investigate cryptic diversity in sympatric and allopatric (separated by 450 km) populations of the widespread coral Seriatopora hystrix on the Great Barrier Reef. Structure analyses delimited unique genetic clusters that were confirmed by phylogenetic and extensive population-level analyses. Each of four sympatric yet distinct genetic clusters detected within S. hystrix demonstrated greater genetic cohesion across regional scales than between genetic clusters within regions (10 km). Moreover, the magnitude of genetic differentiation between different clusters (0.620 G"ST ) was similar to the difference between S. hystrix clusters and the congener S. caliendrum (mean G"ST 0.720). Multiple lines of evidence, including differences in habitat specificity, mitochondrial identity, Symbiodinium associations and morphology, corroborate the nuclear genetic evidence that these distinct clusters constitute different species. Hierarchical clustering analysis combined with more traditional population genetic methods provides a powerful approach for delimiting species and should be regularly applied to ensure that ecological and evolutionary patterns interpreted for single species are not confounded by the presence of cryptic species.

Early establishment of coral-microbial symbioses is fundamental to the fitness of corals, but comparatively little is known about the onset and succession of bacterial communities in their early life history stages. In this study, bacterial associates of the coral Acropora millepora were characterized throughout the first year of life, from larvae and 1-week-old juveniles reared in laboratory conditions in the absence of the dinoflagellate endosymbiont Symbiodinium to field-outplanted juveniles with established Symbiodinium symbioses, and sampled at 2 weeks and at 3, 6 and 12 months. Using an amplicon pyrosequencing approach, the diversity of both nitrogen-fixing bacteria and of bacterial communities overall was assessed through analysis of nifH and 16S rRNA genes, respectively. The consistent presence of sequences affiliated with diazotrophs of the order Rhizobiales (23-58% of retrieved nifH sequences; 2-12% of 16S rRNA sequences), across all samples from larvae to 12-month-old coral juveniles, highlights the likely functional importance of this nitrogen-fixing order to the coral holobiont. Dominance of Roseobacter-affiliated sequences (55% of retrieved 16S rRNA sequences) in larvae and 1-week-old juveniles, and the consistent presence of sequences related to Oceanospirillales and Altermonadales throughout all early life history stages, signifies their potential importance as coral associates. Increased diversity of bacterial communities once juveniles were transferred to the field, particularly of Cyanobacteria and Deltaproteobacteria, demonstrates horizontal (environmental) uptake of coral-associated bacterial communities. Although overall bacterial communities were dynamic, bacteria with likely important functional roles remain stable throughout early life stages of Acropora millepora.

Extremes in seasonal environmental conditions can significantly impact the health and physiological functioning of reef corals, underscoring the need for knowledge of seasonally specific baselines from which to monitor and forecast impending stress. Increases above summertime means in seawater temperature, sunlight intensity, turbidity, or sedimentation may reduce coral immunocompetency and increase disease and bleaching susceptibility. We analyzed temporal patterns in innate immunity parameters over nine time points throughout one year to establish baseline levels from which anomalies might be detected for representative species from three major reef‐building coral families (Acroporidae, Faviidae, and Poritidae). Temporal patterns in both phenoloxidase activity and expression of green fluorescent protein‐like proteins varied among the three families, as did overall constitutive levels. For example, Porites cylindrica had 2.8‐fold higher yearly average levels of phenoloxidase activity than Acropora millepora, which had the lowest levels. In contrast, mean fluorescence was lowest in Acropora millepora and highest in Echinopora mammiformis. Relationships between the potential physical drivers (seasonal variation in seawater temperature, rainfall, salinity) and temporal patterns in these parameters also differed among the three species. For example, phenoloxidase activity was positively correlated with seawater temperature in A. millepora, but negatively correlated in both E. mammiformis and P. cylindrica. Distinctions in constitutive levels and temporal patterns in these parameters among species suggest that corals from these three families have evolved different strategies for investing resources into innate immune parameters. Such differences highlight the need for species‐specific baselines and long‐term assessments to accurately predict coral reef trajectories in rapidly changing environments.

The dinoflagellate photosymbiont Symbiodinium plays a fundamental role in defining the physiological tolerances of coral holobionts, but little is known about the dynamics of these endosymbiotic populations on coral reefs. Sparse data indicate that Symbiodinium populations show limited spatial connectivity; however, no studies have investigated temporal dynamics for in hospite Symbiodinium populations following significant mortality and recruitment events in coral populations. We investigated the combined influences of spatial isolation and disturbance on the population dynamics of the generalist Symbiodinium type C2 (ITS1 rDNA) hosted by the scleractinian coral Acropora millepora in the central Great Barrier Reef. Using eight microsatellite markers, we genotyped Symbiodinium in a total of 401 coral colonies, which were sampled from seven sites across a 12-year period including during flood plume-induced coral bleaching. Genetic differentiation of Symbiodinium was greatest within sites, explaining 70-86% of the total genetic variation. An additional 9-27% of variation was explained by significant differentiation of populations among sites separated by 0.4-13 km, which is consistent with low levels of dispersal via water movement and historical disturbance regimes. Sampling year accounted for 6-7% of total genetic variation and was related to significant coral mortality following severe bleaching in 1998 and a cyclone in 2006. Only 3% of the total genetic variation was related to coral bleaching status, reflecting generally small (8%) reductions in allelic diversity within bleached corals. This reduction probably reflected a loss of genotypes in hospite during bleaching, although no site-wide changes in genetic diversity were observed. Combined, our results indicate the importance of disturbance regimes acting together with limited oceanographic transport to determine the genetic composition of Symbiodinium types within reefs.

Aim: Understanding patterns in a species' occupancy and abundance across multiple scales is important for management purposes, particularly for protecting threatened species. Here, we develop a new quantitative, multiscale model of occupancy and abundance that characterizes seven types of rarity and one of commonness in high-diversity communities of staghorn coral. We examine how rare species influence coral community structure and explore spatial variability in underlying patterns of community structure in the context of optimizing the data needed to protect biodiversity. Location: North-west Pacific Ocean. Methods: We present categorical abundance data for 87 species of staghorn corals occurring within 100 sites across five reefs in the north-west Pacific Ocean. We develop a new model that combines measures of global distribution, local distribution and local abundance to describe eight mutually exclusive occupancy–abundance patterns, which can be used to prioritize regional species conservation. Traditional and new analytical approaches are compared to explore how rare species influence multidimensional space and community structure. Results: We show that five types of occupancy–abundance relationships exist in staghorn coral assemblages, including four patterns of rarity. The overwhelming majority of species (73%) are rare according to local abundance and/or distribution criteria. Occupancy–abundance patterns are spatially variable in staghorn coral communities, and no single underlying distribution fits all assemblages. Our findings suggest that 54 species are at risk of regional extinction, 30 of which are also classified as 'vulnerable' by the IUCN. Main conclusions: Our model demonstrates that multiple occupancy–abundance patterns exist in staghorn coral assemblages. We conclude that 66% of the pool of staghorn coral fauna in the north-west Pacific is at risk of regional extinction. At the locations and scales examined here, occupancy–abundance patterns and the underlying distributions of coral communities are spatially variable, suggesting that it may not be appropriate to apply unified ecological theory to communities with a large proportion of threatened species because this may jeopardize biodiversity conservation.

Concentrating tourism activities can be an effective way to closely manage high-use parks and minimize the extent of the effects of visitors on plants and animals, although considerable investment in permanent tourism facilities may be required. On coral reefs, a variety of human-related disturbances have been associated with elevated levels of coral disease, but the effects of reef-based tourist facilities (e.g., permanent offshore visitor platforms) on coral health have not been assessed. In partnership with reef managers and the tourism industry, we tested the effectiveness of concentrating tourism activities as a strategy for managing tourism on coral reefs. We compared prevalence of brown band disease, white syndromes, black band disease, skeletal eroding band, and growth anomalies among reefs with and without permanent tourism platforms within the Great Barrier Reef Marine Park. Coral diseases were 15 times more prevalent at reefs with offshore tourism platforms than at nearby reefs without platforms. The maximum prevalence and maximum number of cases of each disease type were recorded at reefs with permanently moored tourism platforms. Diseases affected 10 coral genera from 7 families at reefs with platforms and 4 coral genera from 3 families at reefs without platforms. The greatest number of disease cases occurred within the spatially dominant acroporid corals, which exhibited 18-fold greater disease prevalence at reefs with platforms than at reefs without platforms. Neither the percent cover of acroporids nor overall coral cover differed significantly between reefs with and without platforms, which suggests that neither factor was responsible for the elevated levels of disease. Identifying how tourism activities and platforms facilitate coral disease in marine parks will help ensure ongoing conservation of coral assemblages and tourism.

Understanding the effects of elevated seawater temperatures on each member of the coral holobiont (the complex comprised of coral polyps and associated symbiotic microorganisms, including Bacteria, viruses, Fungi, Archaea and endolithic algae) is becoming increasingly important as evidence accumulates that microbial members contribute to overall coral health, particularly during thermal stress. Here we use a metagenomic approach to identify metabolic and taxonomic shifts in microbial communities associated with the hard coral Acropora millepora throughout a natural thermal bleaching event at Magnetic Island (Great Barrier Reef). A direct comparison of metagenomic data sets from healthy versus bleached corals indicated major shifts in microbial associates during heat stress, including Bacteria, Archaea, viruses, Fungi and micro-algae. Overall, metabolism of the microbial community shifted from autotrophy to heterotrophy, including increases in genes associated with the metabolism of fatty acids, proteins, simple carbohydrates, phosphorus and sulfur. In addition, the proportion of virulence genes was higher in the bleached library, indicating an increase in microorganisms capable of pathogenesis following bleaching. These results demonstrate that thermal stress results in shifts in coral-associated microbial communities that may lead to deteriorating coral health.

This study compared the effect of heat stress on coral-associated bacterial communities among juveniles of the coral, Acropora tenuis, hosting different Symbiodinium types. In comparison to a control temperature treatment (28 degrees C), we documented dramatic changes in bacterial associates on juvenile corals harbouring ITS 1 type D Symbiodinium when placed in a high (32 degrees C) temperature treatment. In particular, there was a marked increase in the number of retrieved Vibrio affiliated sequences, which coincided with a 44% decline in the photochemical efficiency of the D-juveniles. Interestingly, these Vibrio sequences affiliated most closely with the coral pathogen, Vibrio coralliilyticus, which has been implicated in some coral disease outbreaks. In contrast, A. tenuis hosting ITS 1 type C1 Symbiodinium did not exhibit major bacterial shifts in the elevated temperature treatment, indicating a more stable bacterial community during thermal stress; concomitantly a decline (10%) in photochemical efficiency was minimal for this group. D juveniles that had been exposed to moderately elevated sea temperatures (30 degrees C) in the field before being placed in the control temperature treatment displayed a decrease in the number of Vibrio affiliated sequences and bacterial profiles shifted to become more similar to profiles of corals harbouring type C1 Symbiodinium. In combination, these results demonstrate that thermal stress can result in shifts in coral-associated bacterial communities, which may lead to deteriorating coral health. The lower resilience of A. tenuis to thermal stress when harbouring Symbiodinium D highlights the importance of inter-kingdom interactions among the coral host, dinoflagellate endosymbiont and bacterial associates for coral health and resilience.

Immunity is a key life history trait that may explain hierarchies in the susceptibility of corals to disease and thermal bleaching, two of the greatest current threats to coral health and the persistence of tropical reefs. Despite their ongoing and rapid global decline, there have been few investigations into the immunity mechanisms of reef-building corals. Variables commonly associated with invertebrate immunity, including the presence of melanin, size of melanin-containing granular cells, and phenoloxidase (PO) activity, as well as concentrations of fluorescent proteins (FPs), were investigated in hard (Scleractinia) and soft (Alcyonacea) corals spanning 10 families from the Great Barrier Reef. Detectable levels of these indicators were present in all corals investigated, although relative investment differed among coral taxa. Overall levels of investment were inversely correlated to thermal bleaching and disease susceptibility. In addition, PO activity, melanin-containing granular cell size, and FP concentration were each found to be significant predictors of susceptibility and thus may play key roles in coral immunity. Correlative evidence that taxonomic (family-level) variation in the levels of these constituent immunity parameters underpins susceptibility to both thermal bleaching and disease indicates that baseline immunity underlies the vulnerability of corals to these two threats. This reinforces the necessity of a holistic approach to understanding bleaching and disease in order to accurately determine the resilience of coral reefs.

A phenotypic and phylogenetic comparison of geographically disparate isolates of the coral pathogen Vibrio coralliilyticus was conducted to determine whether the bacterium exists as a single cosmopolitan clonal population, which might indicate rapid spread of a pandemic strain, or is grouped into endemic and genotypically distinct strains. All strains included in this study displayed similar phenotypic characteristics to those of the typed V. coralliilyticus strain LMG 20984T. Five phylogenetic marker genes (16S, rpoA, recA, pyrH and dnaJ) frequently used for discriminating closely related Vibrio species and a zinc-metalloprotease gene (vcpA) linked to pathogenicity were sequenced in 13 V. coralliilyticus isolates collected from corals, bivalves, and their surrounding seawater in the Red and Caribbean Seas, and Indian, Pacific and Atlantic Oceans. A high level of genetic polymorphism was observed with all isolates possessing unique genotypes at all six genetic loci examined. No consistent lineage structure was observed within the marker genes and homologous recombination was detected in the 16S and vcpA genes, suggesting that V. coralliilyticus does not possess a highly clonal population structure. Interestingly, two geographically distinct (Caribbean/south-Atlantic and Indo-Pacific/north-Atlantic) and highly divergent clades were detected within the zinc-metalloprotease gene, but it is not known if these clades correspond to phenotypic differences in virulence. These findings stress the need for a multi-locus approach for inferring V. coralliilyticus phylogeny and indicate that populations of this bacterium are likely an endemic component of coral reef ecosystems globally.

The majority of reef-building corals acquire their obligate algal symbionts (Symbiodinium) from the environment. However, factors shaping the initial establishment of coral-algal symbioses, including parental effects, local environmental conditions and local availability of symbionts, are not well understood. This study monitored the uptake and maintenance of Symbiodinium in juveniles of two common corals, Acropora tenuis and Acropora millepora, that were reciprocally explanted between sites where adult colonies host different types of Symbiodinium. We found that coral juveniles were rapidly dominated by type D Symbiodinium, even though this type is not found in adult colonies (including the parental colonies) in four out of the five study populations. Furthermore, type D Symbiodinium was found in less than one-third of a wide range of coral species (n > 50) sampled at the two main study sites, suggesting that its dominance in the acroporid juveniles is not because it is the most abundant local endosymbiotic type. Moreover, dominance by type D was observed irrespective of the light intensity to which juveniles were exposed in a field study. In summary, despite its relatively low abundance in coral assemblages at the study sites and irrespective of the surrounding light environment, type D Symbiodinium is the main symbiont type initially acquired by juveniles of A. millepora and A. tenuis. We conclude that during early ontogeny in these corals, there are few barriers to the uptake of Symbiodinium types which differ from those found in parental colonies, resulting in dominance by a highly infectious and potentially opportunistic symbiont.

Symbiotic relationships have contributed greatly to the evolution and maintenance of biological diversity. On the Great Barrier Reef, species of obligate coral-dwelling fishes (genus Gobiodon) coexist by selectively recruiting to colonies of Acropora nasuta that differ in branch-tip colour. In this study, we investigate genetic variability among sympatric populations of two colour morphs of A. nasuta ('blue-tip' and 'brown-tip') living in symbiosis with two fish species, Gobiodon histrio and G. quinquestrigatus, respectively, to determine whether gobies are selecting between intraspecific colour polymorphisms or cryptic coral species. We also examine genetic differentiation among coral populations containing both these colour morphs that are separated by metres between local sites, tens of kilometres across the continental shelf and hundreds of kilometres along the Great Barrier Reef. We use three nuclear DNA loci, two of which we present here for the first time for Acropora. No significant genetic differentiation was detected between sympatric colour morphs at these three loci. Hence, symbiotic gobies are selecting among colour morphs of A. nasuta, rather than cryptic species. Significant genetic geographical structuring was observed among populations, independent of colour, at regional (i.e. latitudinal separation by < 500 km) and cross-shelf (< 50 km) scales, alongside relative homogeneity between local populations on within reef scales (< 5 km). This contrasts with the reported absence of large-scale genetic structuring in A. valida, which is a member of the same species group as A. nasuta. Apparent differences in biogeographical structuring between species within the A. nasuta group emphasize the need for comparative sampling across both spatial (i.e. within reefs, between reefs and between regions) and taxonomic scales (i.e. within and between closely related species).

Species boundaries among five sympatric coral species of the Indo-Pacific Acropora aspera group were examined by a combination of in vitro breeding trials, comparisons of spawning times and DNA sequence analysis of ribosomal DNA internal transcribed spacer (rDNA ITS) and 5.8S regions. The breeding trials showed that reproductive compatibility exists between at least some colonies of all the species pairs tested, suggesting a large potential for natural hybridization and introgression. The Acropora ITS regions exhibited extremely high levels of variability (up to approximately 62% for ITS1, approximately 11% for 5.8S and approximately 43% for ITS2), but most of the variation was shared among four of the five species, A. millepora, A. papillare, A. pulchra and A. spathulata, consistent with extensive introgression. Phylogenetic analyses did not resolve these four species as distinct clusters across a wide biogeographic region stretching from the southern Great Barrier Reef to Papua New Guinea. However, most colonies of the fifth species, A. aspera, constituted a distinct clade in phylogenetic analyses. This is consistent with our observations of a semi-permeable temporal barrier involving differences in spawning times between this and the other four species. Although the majority of colonies of all five species generally spawned within 90 min of each other, in two out of four years, gametes were absent prior to mass spawning episodes from at least some A. aspera colonies. Hence, our data suggest that transient reproductive barriers may be the result of year-to-year variation in the date of spawning and that this difference in spawning time contributes to the genetic structure detected among Acropora species in this group. Occasional leakage through the reproductive barrier was confirmed by the observation of A. aspera xA. pulchra F1 hybrids, identified based on additivity of ITS sequences.

A major challenge for understanding the evolutionary genetics of mass-spawning corals is to explain the maintenance of discrete morphospecies in view of high rates of interspecific fertilization in vitro and nonmonophyletic patterns in molecular phylogenies. In this study, we focused on Acropora cytherea and A. hyacinthus, which have one of the highest potentials for interspecific fertilization. Using sequences of a nuclear intron, we performed phylogenetic and nested clade analyses (NCA). Both species were polyphyletic in molecular phylogenies, but the NCA indicated that they constitute statistically distinguishable lineages. Phylogenetic analysis using an intergenic region of the mitochondrial DNA (mtDNA), was inconclusive because of low levels of variability in this marker. The position of these two species differed between the nuclear DNA (nDNA) and mtDNA phylogenies and was also at odds with a cladistic analysis based on morphology. We conclude that despite the potential for high levels of hybridization and introgression, A. cytherea and A. hyacinthus constitute statistically distinguishable lineages and their taxonomic status is consistent with the cohesion species concept.

Regional-scale variation of recruitment by marine organisms may reflect geographic patterns in adult stock sizes or fecundities, large-scale hydrodynamic features that influence the transport of larvae (e.g., currents, upwelling), and patterns of early mortality. In turn, recruitment may play a vital role in determining patterns of adult abundance and community structure, from local to biogeographic scales. We examined spatial variation in recruitment by corals at a regional scale, along 3300 km of the tropical and subtropical coast of eastern Australia (10°–31° S). We used two complementary approaches: (1) a metaanalysis of 21 different studies undertaken over a 16-yr period, each of which was generally conducted at a single reef, and (2) a large-scale sampling effort in which recruitment was measured in two years on 33 reefs arrayed along the length of the Great Barrier Reef (GBR). Our goal is to compare the emergent large-scale picture derived from many small-scale studies with patterns revealed by shorter-term regional sampling. The two approaches show very similar large-scale patterns. Recruitment by spawning corals (mainly acroporids) was highest in the central GBR and declined steadily with increasing latitude by up to more than 20-fold. A smaller decline occurred on the northern GBR between Australian and Papua New Guinea. Recruitment by brooding corals (mostly pocilloporids) was greatest in the northern GBR and also declined to the south. The latitudinal decline in brooders was three-to fivefold, i.e., not as great as for spawners. Consequently, the proportion of brooded recruits increased to the south, and they generally exceeded spawners on the southern GBR and on isolated subtropical reefs at higher latitudes. Our meta-analysis shows that fully half of the variation in the ratio of spawners to brooders is attributable to one of 11 variables that we extracted from the published studies: the month when the recruitment panels were deployed. This result suggests that the intensity and timing of spawning have a crucial impact on large-scale patterns of recruitment. Elsewhere, we tested this hypothesis in the field, and confirmed that regional variation in recruitment by spawning acroporid corals was driven by spatial and temporal variation in the extent of mass spawning. Together, large-scale sampling and meta-analyses provide a powerful, combined approach for investigating large-scale patterns and the mechanisms underlying them.

Although Acropora is the most species-rich genus of the scleractinian (stony) corals, only three species occur in the Caribbean: A. cervicornis, A. palmata and A. prolifera. Based on overall coral morphology, abundance and distribution patterns, it has been suggested that A. prolifera may be a hybrid between A. cervicornis and A. palmata. The species boundaries among these three morphospecies were examined using DNA sequence analyses of the nuclear Pax-C 46/47 intron and the ribosomal DNA Internal Transcribed Spacer (ITS1 and ITS2) and 5.8S regions. Moderate levels of sequence variability were observed in the ITS and 5.8S sequences (up to 5.2% overall sequence difference), but variability within species was as large as between species and all three species carried similar sequences. Since this is unlikely to represent a shared ancestral polymorphism, the data suggest that introgressive hybridization occurs among the three species. For the Pax-C intron, A. cervicornis and A. palmata had very distinct allele frequencies and A. cervicornis carried a unique allele at a frequency of 0.769 (although sequence differences between alleles were small). All A. prolifera colonies examined were heterozygous for the Pax-C intron, whereas heterozygosity was only 0.286 and 0.333 for A. cervicornis and A. palmata, respectively. These data support the hypothesis that A. prolifera is the product of hybridization between two species that have a different allelic composition for the Pax-C intron, i.e. A. cervicornis and A. palmata. We therefore suggest that A. prolifera is a hybrid between A. cervicornis and A. palmata, which backcrosses with the parental species at low frequency.

"Supply-side" ecology recognizes the potential role that recruitment plays in the local population dynamics of open systems. Apart from the applied fisheries literature, the converse link between adults and the production of cohorts of recruits has received much less attention. We used a hierarchical sampling design to investigate the relationships between adult abundance, fecundity, and rates of larval recruitment by acroporid corals on 33 reefs in five sectors (250-400 km apart) stretching from north to south along the length of the Great Barrier Reef, Australia. Our goal was to quantify patterns of recruitment at multiple scales, and to explore the underlying mechanisms. Specifically, we predicted that large-scale patterns of recruitment could be driven by changes in the abundance of adults and/or their fecundity, i.e., that corals exhibit a stock-recruitment relationship. The amount of recruitment by acroporids in each of two breeding seasons varied by more than 35-fold among the five sectors. Adult density varied only twofold among sectors and was not correlated with recruitment at the sector or reef scale. In contrast, fecundity levels (the proportion of colonies on each reef that contained ripe eggs) varied from 15% to 100%, depending on sector, year, and species. Spatial and temporal variation in the fecundity of each of three common Acropora species explained most of the variation (72%) in recruitment by acroporids, indicating that the production of larvae is a major determinant of levels of recruitment at large scales. Once fecundity was accounted for, none of the other variables we examined (sector, reef area, abundance of adults, or year) contributed significantly to variation in recruitment. The relationship between fecundity and recruitment was nonlinear, i.e., rates of recruitment increased disproportionately when and where the proportion of gravid colonies approached 100%. This pattern is consistent with the hypothesis that enhanced fertilization success and/or predator satiation occurs during mass-spawning events. Furthermore, it implies that small, sublethal changes in fecundity of corals could result in major reductions in recruitment.

Black band disease (BBD) is a microbial consortium that creates anoxic, sulfide-rich microenvironments and kills underlying coral tissues as it rapidly migrates across colonies. Although bacterial communities associated with BBD have been studied extensively, the presence and roles of archaea are unexplored. Using amplicon-pyrosequencing of 16S ribosomal RNA genes, we investigated the community structure of both archaea and bacteria within microbial lesions of BBD and the less-virulent precursor stage, 'cyanobacterial patches' (CP), affecting the coral Montipora hispida. We detected characteristic shifts in microbial communities during the development of BBD from CP, reflecting microenvironmental changes within lesions. Archaeal profiles in CP suggested a diverse assemblage affiliated with the Thaumarchaeota and Euryarchaeota, similar to communities described for oxic marine environments. In contrast, a novel ribotype, distantly affiliated to the Euryarchaeota, dominated up to 94% of archaeal sequences retrieved from BBD. The physiological characteristics of this dominant archaeal ribotype are unknown because of the novelty of its 16S ribosomal RNA gene sequences; however, their prominent associations with BBD lesions suggest the ability to thrive in the organic- and sulfide-rich anoxic microenvironment characteristic of BBD lesions. Discovery of this novel archaeal ribotype provides new insights into the microbial ecology and aetiology of BBD.