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The decline of coral reefs has fueled interest in determining whether mesophotic reefs can shield against disturbances and help replenish deteriorated shallower reefs. In this study, we characterized spatial (horizontal and vertical) and seasonal patterns of diversity in coral recruits from Dabaisha and Guiwan reefs at Ludao, Taiwan. Concrete blocks supporting terra‐cotta tiles were placed at shallow (15m) and mesophotic (40m) depths, during 2016–2018. Half of the tiles were retrieved and replaced biannually over three 6‐month surveys (short‐term); the remainder retrieved at the end of the 18‐month (long‐term) survey. 451 recruits were located using fluorescent censusing and identified by DNA barcoding. Barcoding the mitochondrial cytochrome oxidase I (COI) gene resulted in 17 molecular operational taxonomic units (MOTUs). To obtain taxonomic resolution to the generic level, Pocillopora were phylotyped using the mitochondrial open reading frame (ORF), resolving eight MOTUs. Acropora, Isopora, and Montipora recruits were identified by the nuclear PaxC intron, yielding ten MOTUs. Overall, 35 MOTUs were generated and were comprised primarily of Pocillopora and, in fewer numbers, Acropora, Isopora, Pavona, Montipora, Stylophora, among others. 40% of MOTUs recruited solely within mesophotic reefs while 20% were shared by both depth zones. MOTUs recruiting across a broad depth distribution appear consistent with the hypothesis of mesophotic reefs acting as a refuge for shallow‐water coral reefs. In contrast, Acropora and Isopora MOTUs were structured across depth zones representing an exception to this hypothesis. This research provides an imperative assessment of coral recruitment in understudied mesophotic reefs and imparts insight into the refuge hypothesis., We characterized spatial (horizontal and vertical) and seasonal patterns of diversity in coral recruits at shallow and mesophotic reefs in Ludao, Taiwan, using DNA barcoding of the COI, PAXC, and ORF regions. Thirty‐five MOTUs were generated and were comprised primarily of Pocillopora and, in fewer numbers, Acropora, Isopora, Pavona, Montipora, Stylophora, among others. MOTUs recruiting across a broad depth distribution appear consistent with the hypothesis of mesophotic reefs acting as a refuge for shallow‐water coral reefs.

Polycyathus chaishanensis is a symbiotic caryophyllid coral described from a single population in a tidal pool off Chaishan, Kaohsiung, Taiwan. Due to its rarity, P. chaishanensis was declared a critically-endangered species under the Taiwan Wildlife Protection Act. In May 2017, a P. chaishanensis colony was discovered in the intertidal area of the Datan Algal Reef, Taoyuan, Taiwan. To determine whether this is a stable population in the algal reef, a demographic census—including data on occurrence, distribution, and colony size—was carried out in the algal reef in southern Taoyuan. Intertidal censuses and sediment collections were conducted at five different sections—Baiyu, Datan G1, Datan G2, Yongxing, and Yongan algal reefs—during the monthly spring low tide from July 2018 to January 2019. In total, 84 colonies—23 in Datan G1 and 61 in Datan G2—were recorded from a tidal range of − 160 to − 250 cm, according to the Taiwan Vertical Datum 2001 compiled by the Central Weather Bureau. No P. chaishanensis was found in Baiyu, Yongxing, or Yongan. The P. chaishanensis colony sizes ranged from 2.55 to 81.5 cm in diameter, with the larger P. chaishanensis present in the lower intertidal zone. Sediment was extremely high, with monthly site averages ranging from 3,818.26 to 29,166.88 mg cm−2 day−1, and there was a significant difference between sites and months, both of which affected the distribution of P. chaishanensis in the algal reef. Our study confirms the existence of a second population of P. chaishanensis in Taiwan, highlighting the importance of the Datan Algal Reef for the survival and protection of this critically-endangered caryophyllid coral and why it is so urgent that the reef should be conserved.

Over the past few decades, extreme events—such as ocean warming, typhoons, and coral bleaching—have been increasing in intensity and frequency, threatening coral reefs from the physiological to ecosystem level. In the present study, the impacts of rising seawater temperatures, typhoons, and coral bleaching events on benthic communities were seasonally assessed over a 21 month-period, using photo-transects at 11 sites in Kenting National Park (KNP), Taiwan. Between August 2015 and April 2017, seven typhoon events were recorded and in situ seawater temperatures in KNP reached a maximum of 31.2 °C, as opposed to an average maximum SST of 28.8 °C (2007–2016). The state and response of benthic communities to these events were interpreted based on the environmental conditions of KNP. The repeated storms lowered the levels of thermal stress during the 2015–2016 El Niño event and may have mitigated its impact on the Taiwanese coral reefs. However, storm-induced local shifts from coral to macro-algae dominance were observed. Storms may mitigate the negative effects of heatwaves, but the mechanical damage induced by the storms may also decrease the structural complexity of reefs and their associated diversity. Eventually, despite reef persistence, the composition and function of remnant communities may profoundly diverge from those in regions with less active storms.

The discovery of multi-species synchronous spawning of scleractinian corals on the Great Barrier Reef in the 1980s stimulated an extraordinary effort to document spawning times in other parts of the globe. Unfortunately, most of these data remain unpublished which limits our understanding of regional and global reproductive patterns. The Coral Spawning Database (CSD) collates much of these disparate data into a single place. The CSD includes 6178 observations (3085 of which were unpublished) of the time or day of spawning for over 300 scleractinian species in 61 genera from 101 sites in the Indo-Pacific. The goal of the CSD is to provide open access to coral spawning data to accelerate our understanding of coral reproductive biology and to provide a baseline against which to evaluate any future changes in reproductive phenology., info:eu-repo/semantics/published

As evolutionary relationships among some coral species still remain unclear, studies on unstudied area such as the Persian Gulf (PG), as part of the western Indo-Pacific, may reveal a better understanding of phylogenetic positions and relationships of corals. In the present study, the phylogenetic relationships of eight common coral species (Favites pentagona, Platygyra daedalea, Cyphastrea microphthalma, Siderastrea savignyana, Pavona decussata, Pavona cactus, Goniopora columna, and Goniopora djiboutiensis) collected from two Iranian Islands were compared with the congeneric sequences from the Indo-Pacific (IP) using rDNA region. The result shows that some coral species which were hitherto considered as representatives of widespread species from IP are related to distinct lineages. Further, it appears that morphological convergence between the taxa leads to an underestimation of the real coral species diversity in the PG. The current study is the first attempt to investigate the phylogenetic position of coral species from the PG in comparison to their counterparts from the IP. As conservation planning hinges on the identification of species, taxonomic revisions have to be undertaken in order to obtain a more reliable picture of coral species diversity in the PG.

Spatial patterns of coral reef benthic communities vary across a range of broad-scale biogeographical levels to fine-scale local habitat conditions. This study described spatial patterns of coral reef benthic communities spanning across the 536-km coast of Kenya. Thirty-eight reef sites representing different geographical zones within an array of habitats and management levels were assessed by benthic cover, coral genera and coral colony size classes. Three geographical zones were identified along the latitudinal gradient based on their benthic community composition. Hard coral dominated the three zones with highest cover in the south and Porites being the most abundant genus. Almost all 15 benthic variables differed significantly between geographical zones. The interaction of habitat factors and management levels created a localised pattern within each zone. Four habitats were identified based on their similarity in benthic community composition; 1. Deep-Exposed Patch reef in Reserve areas (DEPR), 2. Deep-Exposed Fringing reefs in Unprotected areas (DEFU), 3. Shallow Fringing and Lagoon reefs in Protected and Reserve areas (SFLPR) and 4. Shallow Patch and Channel reefs (SPC). DEPR was found in the north zone only and its benthic community was predominantly crustose coralline algae. DEFU was found in central and south zones mainly dominated by soft corals, Acropora, Montipora, juvenile corals and small colonies of adult corals. SFLPR was dominated by macroalgae and turf algae and was found in north and central zones. SPC was found across all geographical zones with a benthic community dominated by hard corals of mostly large colonies of Porites and Echinopora. The north zone exhibits habitat types that support resistance properties, the south supports recovery processes and central zone acts as an ecological corridor between zones. Identifying habitats with different roles in reef resilience is useful information for marine spatial planning and supports the process of designing effective marine protected areas.

Adaptation and/or acclimatization through various mechanisms have been suggested to help some tropical coral species to overcome temperature-induced bleaching that is intensifying with climate change; however, while much research has been done on the physiological responses of tropical and subtropical corals to stress, little is known about these responses in corals in marginal environments—e.g., high-latitude and non-reefal communities. In this study, we examined the thermal-tolerant physiology of the flowerpot coral, Alveopora japonica, endemic to the high-latitude Jeju Island (33.39°N), South Korea and Oulastrea crispata and Coelastrea aspera from the subtropical non-reefal coral community on the Penghu Islands (23.34°N), Taiwan. Analysis of physiological parameters; photochemical efficiency, Chlorophyll pigment, Symbiodiniaceae cell number and host soluble proteins - showed that A. japonica can survive through a wide range of temperature stresses (10–32 °C) over a period of 8 days without showing signs of bleaching. In addition, corals O. crispata and C. aspera withstood temperature stresses of up to 33 °C and repeated temperature fluctuations without bleaching. Our results indicate that, under large seasonal variations and asymmetrical daily fluctuations in temperature, corals currently living in marginal environments could have thermal plasticity, allowing them to survive in the future climate change scenarios. This study reiterates the importance of studying the eco-physiology of corals that are generally ignored because of their neutral or positive responses to stress.

Symbiodiniaceae communities in some corals often shuffle or switch after severe bleaching events, one of the major threats to coral survival in a world with climate change. In this study we reciprocally transplanted five Leptoria phrygia colonies between two sites with significantly different temperature regimes and monitored them for 12 months. Our ITS2 amplicon deep sequencing demonstrated that L. phrygia acclimatized to maintain a strong and stable association with Durusdinium D17, D. trenchii, and D. glynnii, but also remained flexible and formed a short-term association with different Cladocopium. Most interestingly, two colonies shuffled between Durusdinium and Cladocopium without the occurrence of bleaching; one colony even switched its dominant Cladocopium after generic shuffling. Both dominant Cladocopium were originally rare with relative abundances as low as 0.024%. This is the first record of adult corals switching dominant symbiont without bleaching.

This study monitored symbiont communities bi-monthly in native coral cores used in a reciprocal transplantation of the coralPlatygyra verweyiover two years (2014–2016) and samples of mother colonies from three locations with variable thermal regimes; our results show that associating with multiple Symbiodiniaceae genera (Cladocopiumspp. andDurusdiniumspp.) is not a prerequisite for symbiont shuffling.Platygyra verweyiassociates with certain Symbiodiniaceae genera based on location. Results of quantitative real-time PCR indicated small-scale temporal changes in Symbiodiniaceae genera compositions from 2014 to 2016; however, these changes were not enough to invoke shuffling or switching, despite degree heating weeks exceeding 6 °C-weeks in 2014 and 4 °C-weeks in 2015, which usually resulted in substantial coral bleaching. Microsatellite analysis of theP. verweyihost showed no genetic differences among the study locations. Our results suggest thatP. verweyiundergoes long-term acclimatization and/or adaptation based on microgeographic and local environmental conditionsby altering its combinations of associated Symbiodiniaceae. Results also suggest that shuffling might not be as common a phenomenon as it has been given credit for; corals thrive through specific associations, and many corals could still be vulnerable to climate change-induced stress, despite being promiscuous or able to associate with rare and background Symbiodiniaceae genera.

Mesophotic habitats could be sheltered from natural and anthropogenic disturbances and act as reproductive refuges, providing propagules to replenish shallower populations. Molecular markers can be used as proxies evaluating the connectivity and inferring population structure and larval dispersal. This study characterizes population structure as well as horizontal and vertical genetic connectivity of the broadcasting coral Pocillopora verrucosa from Ludao, a small oceanic island off the eastern coast of Taiwan. We genotyped 75 P. verrucosa specimens from three sites (Gongguan, Dabaisha, and Guiwan) at three depth ranges (Shallow: 7–15 m, Mid-depth: 23–30 m, and Deep: 38–45 m), spanning shallow to upper mesophotic coral reefs, with eight microsatellite markers. F-statistics showed a moderate differentiation (FST = 0.106, pPocillopora individuals could be drawn from a single panmictic population. Simulations of recent migration assigned 30 individuals (40%) to a different location from where they were collected. Among them, 1/3 were assigned to deeper locations, 1/3 to shallower populations and 1/3 were assigned to the right depth but a different site. These results suggest high levels of vertical and horizontal connectivity, which could enhance the recovery of P. verrucosa following disturbances around Ludao, a feature that agrees with demographic studies portraying this species as an opportunistic scleractinian.

Colony morphological features is among the best predictor of the scleractinian coral’s function in reef ecosystems. However, morphological traits are categorical and to convert this information into a quantitative value as well as estimate their influence on ecosystem process remain a challenge. Here, we propose a trait-based approach to quantify morphological diversity and assess the structural complexity of the habitat provided by corals. We used a previously published dataset that is related to a bleaching event that affected the coral reef off Tikus Island in Indonesia in 1983. We found clear signs of recovery of the coral assemblage’s complexity toward pre El Niño conditions five years after the event. Independent of the change observed in species richness, this return in structural complexity was accompanied by a global decrease in species number associated with each particular morphological entity (Functional Redundancy) and an increase in the number of single-species entities (Functional Vulnerability). Together with species loss, we show an overall functional erosion of the coral assemblage and suggest that the role of the coral reef habitat could be strongly imperiled under repeated or synergistic disturbances. This approach offers an opportunity for a better understanding of coral responses to natural and anthropogenic disturbances.

Coral reefs in the Anthropocene are being subjected to unprecedented levels of stressors, including local disturbances—such as overfishing, habitat destruction, and pollution—and large-scale destruction related to the global impacts of climate change—such as typhoons and coral bleaching. Thus, the future of corals and coral reefs in any given community and coral-Symbiodiniaceae associations over time will depend on their level of resilience, from individual corals to entire ecosystems. Herein we review the environmental settings and long-term ecological research on coral reefs, based on both coral resilience and space, in Kenting National Park (KNP), Hengchun Peninsula, southern Taiwan, wherein fringing reefs have developed along the coast of both capes and a semi-closed bay, known as Nanwan, within the peninsula. These reefs are influenced by a branch of Kuroshio Current, the monsoon-induced South China Sea Surface Current, and a tide-induced upwelling that not only shapes coral communities, but also reduces the seawater temperature and creates fluctuating thermal environments which over time have favoured thermal-resistant corals, particularly those corals close to the thermal effluent of a nuclear power plant in the west Nanwan. Although living coral cover (LCC) has fluctuated through time in concordance with major typhoons and coral bleaching between 1986 and 2019, spatial heterogeneity in LCC recovery has been detected, suggesting that coral reef resilience is variable among subregions in KNP. In addition, corals exposed to progressively warmer and fluctuating thermal environments show not only a dominance of associated, thermally-tolerant Durusdinium spp. but also the ability to shuffle their symbiont communities in response to seasonal variations in seawater temperature without bleaching. We demonstrate that coral reefs in a small geographical range with unique environmental settings and ecological characteristics, such as the KNP reef, may be resilient to bleaching and deserve novel conservation efforts. Thus, this review calls for conservation efforts that use resilience-based management programs to reduce local stresses and meet the challenge of climate change.

Este artículo contiene 10 páginas, 5 figuras., Without drastic efforts to reduce carbon emissions and mitigate globalized stressors, tropical coral reefs are in jeopardy. Strategic conservation and management requires identification of the environmental and socioeconomic factors driving the persistence of scleractinian coral assemblages—the foundation species of coral reef ecosystems. Here, we compiled coral abundance data from 2,584 Indo-Pacific reefs to evaluate the influence of 21 climate, social and environmental drivers on the ecology of reef coral assemblages. Higher abundances of framework-building corals were typically associated with: weaker thermal disturbances and longer intervals for potential recovery; slower human population growth; reduced access by human settlements and markets; and less nearby agriculture. We therefore propose a framework of three management strategies (protect, recover or transform) by considering: (1) if reefs were above or below a proposed threshold of >10% cover of the coral taxa important for structural complexity and carbonate production; and (2) reef exposure to severe thermal stress during the 2014–2017 global coral bleaching event. Our findings can guide urgent management efforts for coral reefs, by identifying key threats across multiple scales and strategic policy priorities that might sustain a network of functioning reefs in the Indo-Pacific to avoid ecosystem collapse.

Climate change threatens coral survival by causing coral bleaching, which occurs when the coral’s symbiotic relationship with algal symbionts (Symbiodiniaceae) breaks down. Studies on thermal adaptation focus on symbionts because they are accessible both in vitro and in hospite. However, there is little known about the physiological and biochemical response of adult corals (without Symbiodiniaceae) to thermal stress. Here we show acclimatization and/or adaptation potential of menthol-bleached aposymbiotic coral Platygyra verweyi in terms of respiration breakdown temperature (RBT) and malate dehydrogenase (MDH) enzyme activity in samples collected from two reef sites with contrasting temperature regimes: a site near a nuclear power plant outlet (NPP-OL, with long-term temperature perturbation) and Wanlitong (WLT) in southern Taiwan. Aposymbiotic P. verweyi from the NPP-OL site had a 3.1 °C higher threshold RBT than those from WLT. In addition, MDH activity in P. verweyi from NPP-OL showed higher thermal resistance than those from WLT by higher optimum temperatures and the activation energy required for inactivating the enzyme by heat. The MDH from NPP-OL also had two times higher residual activity than that from WLT after incubation at 50 °C for 1 h. The results of RBT and thermal properties of MDH in P. verweyi demonstrate potential physiological and enzymatic response to a long-term and regular thermal stress, independent of their Symbiodiniaceae partner.

BackgroundThe chemically-rich seaweed Galaxaura is not only highly competitive with corals, but also provides substrate for other macroalgae. Its ecology and associated epiphytes remain largely unexplored. To fill this knowledge gap, we undertook an ecological assessment to explore the spatial variation, temporal dynamics, and diversity of epiphytic macroalgae of Galaxaura divaricata on patch reefs in the lagoon of Dongsha Atoll, a shallow coral reef ecosystem in the northern South China Sea that has been repeatedly impacted by mass coral bleaching events.MethodsTwelve spatially independent patch reefs in the Dongsha lagoon were first surveyed to assess benthic composition in April 2016, and then revisited to determine G. divaricata cover in September 2017, with one additional Galaxaura-dominated reef (site 9). Four surveys over a period of 17 months were then carried out on a degraded patch reef site to assess the temporal variation in G. divaricata cover. Epiphytic macroalgae associated with G. divaricata were quantified and identified through the aid of DNA barcoding at this degraded site.ResultsPatch reefs in the Dongsha lagoon were degraded, exhibiting relatively low coral cover (5-43%), but high proportions of macroalgae (13-58%) and other substrate (rubble and dead corals; 23-69%). The distribution of G. divaricata was heterogeneous across the lagoon, with highest abundance (16-41%) in the southeast area. Temporal surveys showed consistently high covers (mean ± SD = 16.9 ± 1.21%) of G. divaricata for 17 months. Additional photographic evidence suggested that overgrowth of G. divaricata can persist for 3.5 years. Yet, G. divaricata provides substrate to other macroalgae (e.g., Lobophora sp.) that also limit the growth of corals.ConclusionsOur study demonstrates that an allelopathic seaweed, such as G. divaricata, can overgrow degraded coral reefs for extended periods of time. By providing habitat for other harmful macroalgae, a prolonged Galaxaura overgrowth could further enhance the spread of macroalgae, and strengthen negative feedback loops, decreasing the recovery potential of degraded reefs.

BackgroundThe chemically-rich seaweed Galaxaura is not only highly competitive with corals, but also provides substrate for other macroalgae. Its ecology and associated epiphytes remain largely unexplored. To fill this knowledge gap, we herein undertook an ecological assessment to explore the spatial variation, temporal dynamics, and epiphytic macroalgae of G. divaricata on patch reefs in the lagoon of Dongsha Atoll, a shallow coral reef ecosystem in the northern South China Sea, repeatedly impacted by mass coral bleaching events.MethodsTwelve spatially independent patch reefs in the Dongsha lagoon were first surveyed to assess the benthic composition in April 2016, and then revisited to determine G. divaricata percent cover in September 2017, with one additional Galaxaura-dominated reef (site 9). Four surveys over a period of 17 months were carried out on a degraded patch reef (site 7) to assess the temporal variation in G. divaricata cover. Epiphytic macroalgae associated with G. divaricata were quantified and identified through the aid of DNA barcoding.ResultsPatch reefs in the Dongsha lagoon were degraded, exhibiting relatively low live coral cover (5-43%), but high proportions of macroalgae (13-58%) and other substrates (rubble and dead corals; 23-69%). The distribution of G. divaricata was heterogeneous across the lagoon, with highest abundance (16-41%) in the southeast area. Temporal surveys from site 7 and photo-evidence from site 9 suggested that an overgrowth by G. divaricata was still present to a similar extend after 17 months and 3.5 years. Yet, G. divaricata provides a suitable substrate some allelopathic macroalgae (e.g., Lobophora sp.).ConclusionsOur study demonstrates that an allelopathic seaweed, such as G. divaricata, can overgrow degraded coral reefs for extended periods of time. By providing habitat for harmful macroalgae, a prolonged Galaxaura overgrowth could strengthen negative feedback loops on degraded coral reefs, further decreasing their recovery potential.

Most studies on endosymbiotic dinoflagellate algae (genus Symbiodinium) associated with scleractinian corals focus on tropical and sub-tropical reefs. Their diversity in outlying, non-reef coral communities at high latitudes is still not fully documented. In this study, we analyzed the Symbiodinium diversity associated with five scleractinian species collected at eight sites around Jeju Island (South Korea, 33.4°N) between 5 and 15 m depth. Denaturing gradient gel electrophoresis of amplified internal transcribed spacer region 2 distinguished five Symbiodinium types. We observed a high level of specificity between host genera and Symbiodinium spp. despite existing in an environment with large seasonal oscillations in temperature and light. Psammocora albopicta and Psammocora profundacella were associated with C1 and Montipora millepora with C17. Alveopora japonica was associated exclusively with an unusual F-type, the only known clade F representative functionally important to a scleractinian coral. Oulastrea crispata was associated with Symbiodinium boreum (type D15), occasionally co-occurring with type C3 (in 4 % of specimens). In addition to increasing the knowledge of Symbiodinium diversity in high-latitude coral communities, this study constitutes an important baseline upon which the effects of projected environmental change in the near future can be assessed. A better understanding of high-latitude coral communities is critical for understanding how a warming planet will affect the tempo and mode of shifts in the composition of temperate marine communities.

Seasonal variation in temperature fluctuations may provide corals and their algal symbionts varying abilities to acclimate to changing temperatures. We hypothesized that different temperature ranges between seasons may promote temperature-tolerance of corals, which would increase stability of a bacterial community following thermal stress. Acropora muricata coral colonies were collected in summer and winter (water temperatures were 23.4–30.2 and 12.1–23.1 °C, respectively) from the Penghu Archipelago in Taiwan, then exposed to 6 temperature treatments (10–33 °C). Changes in coral-associated bacteria were determined after 12, 24, and 48 h. Based on 16S rRNA gene amplicons and Illumina sequencing, bacterial communities differed between seasons and treatments altered the dominant bacteria. Cold stress caused slower shifts in the bacterial community in winter than in summer, whereas a more rapid shift occurred under heat stress in both seasons. Results supported our hypothesis that bacterial community composition of corals in winter are more stable in cold temperatures but changed rapidly in hot temperatures, with opposite results for the bacterial communities in summer. We infer that the thermal tolerance ranges of coral-associated bacteria, with a stable community composition, are associated with their short-term (3 mo) seawater thermal history. Therefore, seasonal acclimation may increase tolerance of coral-associated bacteria to temperature fluctuations.

Dongsha Atoll (also known as Pratas) in Taiwan is the northernmost atoll in the South China Sea and a designated marine national park since 2007. The marine park’s scope of protection covers the bio-resources of its waters in addition to uplands, so it is important to have data logging information and analyses of marine flora and fauna, including their physiology, ecology, and genetics. As part of this effort, we investigatedSymbiodiniumassociations in scleractinian corals from Dongsha Atoll through surveys carried out at two depth ranges (shallow, 1–5 m; and deep, 10–15 m) in 2009 and during a bleaching event in 2010.Symbiodiniumcomposition was assessed using restriction fragment length polymorphism (RFLP) of 28S nuclear large subunit ribosomal DNA (nlsrDNA). Our results showed that the 796 coral samples from seven families and 20 genera collected in 2009 and 132 coral samples from seven families and 12 genera collected in 2010 were associated withSymbiodiniumC, D and C+D. Occurrence of clade D in shallow water (24.5%) was higher compared to deep (14.9%). Due to a bleaching event in 2010, up to 80% of coral species associated withSymbiodiniumC underwent moderate to severe bleaching. Using the fine resolution technique of denaturing gradient gel electrophoresis (DGGE) of internal transcribed spacer 2 (ITS2) in 175 randomly selected coral samples, from 2009 and 2010, eightSymbiodiniumC types and twoSymbiodiniumD types were detected. This study is the first baseline survey onSymbiodiniumassociations in the corals of Dongsha Atoll in the South China Sea, and it shows the dominance ofSymbiodiniumclade C in the population.

Corallimorpharia is a small Order of skeleton-less animals that is closely related to the reef-building corals (Scleractinia) and of fundamental interest in the context of understanding the potential impacts of climate change in the future on coral reefs. The relationship between the nominal Orders Corallimorpharia and Scleractinia is controversial—the former is either the closest outgroup to the Scleractinia or alternatively is derived from corals via skeleton loss. This latter scenario, the “naked coral” hypothesis, is strongly supported by analyses based on mitochondrial (mt) protein sequences, whereas the former is equally strongly supported by analyses of mt nucleotide sequences. The “naked coral” hypothesis seeks to link skeleton loss in the putative ancestor of corallimorpharians with a period of elevated oceanic CO2 during the Cretaceous, leading to the idea that these skeleton-less animals may be harbingers for the fate of coral reefs under global climate change. In an attempt to better understand their evolutionary relationships, we examined mt genome organization in a representative range (12 species, representing 3 of the 4 extant families) of corallimorpharians and compared these patterns with other Hexacorallia. The most surprising finding was that mt genome organization in Corallimorphus profundus, a deep-water species that is the most scleractinian-like of all corallimorpharians on the basis of morphology, was much more similar to the common scleractinian pattern than to those of other corallimorpharians. This finding is consistent with the idea that C. profundus represents a key position in the coral corallimorpharian transition.

The zooxanthellate scleractinian coral, Alveopora japonica Eguchi, 1968, inhabits high-latitude waters from southern Taiwan to mainland Japan. This species may have benefited from recent increase in seawater temperature, and it has been hypothesized that a shift from kelp forest to coral dominance may be occurring at some locations at the edge of its distribution. Here, we examined the coral diversity and composition of the benthic community associated with extensive A. japonica stands from Biyangdo, northeast of Jeju Island (33.41°N), South Korea. Coral recruit density was further examined by estimating the number of juvenile corals in photographs taken directly from the benthos. The depauperate coral assemblage (five species in total) was dominated by A. japonica, which contributed to 64.0% ( SE 1.4%) of the benthic cover. Together with local reports documenting recent development of A. japonica populations, the exceptionally high density of recruits, 7590 (SE 660) recruits per m 2 , suggests a population that could be increasing. Additional data are needed to correlate the trend observed in this population with rising seawater temperature. However, this "weedy" and opportunistic coral species may be colonizing the region more aggressively than expected by chance, possibly to the detriment of the original ecosystem dominated by macroalgae. Overall, this work constitutes an important baseline against which future changes recorded in the temperate benthic communities can be assessed.

Coral reef ecosystems are under the direct threat of increasing atmospheric greenhouse gases, which increase seawater temperatures in the oceans and lead to bleaching events. Global bleaching events are becoming more frequent and stronger, and understanding how corals can tolerate and survive high-temperature stress should be accorded paramount priority. Here, we review evidence of the different mechanisms that corals employ to mitigate thermal stress, which include association with thermally tolerant endosymbionts, acclimatisation, and adaptation processes. These differences highlight the physiological diversity and complexity of symbiotic organisms, such as scleractinian corals, where each species (coral host and microbial endosymbionts) responds differently to thermal stress. We conclude by offering some insights into the future of coral reefs and examining the strategies scientists are leveraging to ensure the survival of this valuable ecosystem. Without a reduction in greenhouse gas emissions and a divergence from our societal dependence on fossil fuels, natural mechanisms possessed by corals might be insufficient towards ensuring the ecological functioning of coral reef ecosystems.

The Symbiodinaceae are paradoxical in that they play a fundamental role in the success of scleractinian corals, but also in their dismissal when under stress. In the past decades, the discovery of the endosymbiont's genetic and functional diversity has led people to hope that some coral species can survive bleaching events by associating with a stress-resistant symbiont that can become dominant when seawater temperatures increase. The variety of individual responses encouraged us to scrutinize each species individually to gauge its resilience to future changes. Here, we analyse the temporal variation in the Symbiodinaceae community associated with Leptoria phrygia, a common scleractinian coral from the Indo-Pacific. Coral colonies were sampled from two distant reef sites located in southern Taiwan that differ in temperature regimes, exemplifying a 'variable site' (VS) and a 'steady site' (SS). We investigated changes in the relative abundance of the dominant symbiont and its physiology every 3-4 months from 2016-2017. At VS, 11 of the 12 colonies were dominated by the stress-resistant Durusdinium spp. (>90% dominance) and only one colony exhibited co-dominance between Durusdinium spp. and Cladocopium spp. Every colony displayed high photochemical efficiency across all sampling periods, while showing temporal differences in symbiont density and chlorophyll a concentration. At SS, seven colonies out of 13 were dominated by Cladocopium spp., five presented co-dominance between Durusdinium spp./Cladocopium spp. and only one was dominated by Durusdinium spp. Colonies showed temporal differences in photochemical efficiency and chlorophyll a concentration during the study period. Our results suggest that VS colonies responded physiologically better to high temperature variability by associating with Durusdinium spp., while in SS there is still inter-colonial variability, a feature that might be advantageous for coping with different environmental changes.

The distribution and the structure of benthic assemblages vary with latitude. However, few studies have described benthic communities along large latitudinal gradients, and patterns of variation are not fully understood. Taiwan, lying between 21.90°N and 25.30°N, is located at the center of the Philippine-Japan arc and lies at the northern margin of coral reef development. A wide range of habitats is distributed along this latitudinal gradient, from extensive fringing coral reefs at the southern coast to non-reefal communities at the north. In this study, we examined the structure of benthic communities around Taiwan, by comparing its assemblages in four regions, analyzing the effects of the latitudinal gradient, and highlighting regional characteristics. A total of 25 sites, 125 transects, and 2,625 photographs were used to analyze the benthic communities. Scleractinian corals present an obvious gradient of increasing diversity from north to south, whereas macro-algae diversity is higher on the north-eastern coast. At the country scale, Taiwanese coral communities were dominated by turf algae (49%). At the regional scale, we observed an important heterogeneity that may be caused by local disturbances and habitat degradation that smooths out regional differences. In this context, our observations highlight the importance of managing local stressors responsible for reef degradation. Overall, this study provides an important baseline upon which future changes in benthic assemblages around Taiwan can be assessed.

The scleractinian family Lobophylliidae is undergoing a major taxonomic revision thanks to the combination of molecular and morphological data. In this study, we investigate the evolutionary relationships and the macro- and micromorphology of six nominal coral species belonging to two of the nine molecular clades of the Lobophylliidae, clades A and B, and of Symphyllia wilsoni, a lobophylliid species analyzed from a molecular point of view for the first time. Sequence data from mitochondrial DNA (COI and the intergenic spacer between COI and l-rRNA), and nuclear DNA (histone H3 and ITS region) are used to generate robust molecular phylogenies and a median-joining haplotype network. Molecular results are strongly in agreement with detailed observations of gross- and fine-scale morphology of skeletons, leading to the formal revision of the genera Micromussa and Homophyllia and the description of two newly discovered zooxanthellate shallow-water species, Micromussa pacifica sp. nov. Benzoni & Arrigoni and Micromussa indiana sp. nov. Benzoni & Arrigoni, and a new genus, Australophyllia gen. nov. Benzoni & Arrigoni. In particular, Acanthastrea lordhowensis and Montastraea multipunctata are moved into Micromussa, A. hillae is synonymized with A. bowerbanki and is transferred to Homophyllia, and a revised diagnosis for both genera is provided. Micromussa pacifica sp. nov. is described from the Gambier Islands with its distribution spanning New Caledonia and eastern Australia. Despite a superficial resemblance with Homophyllia australis, it has distinctive macroand micromorphological septal features. Micromussa indiana sp. nov., previously identified as M. amakusensis, is here described from the Gulf of Aden and the southern Red Sea as a distinct species that is genetically separated from M. amakusensis and is morphologically distinct from the latter due to its smaller corallite size and lower number of septa. Finally, molecular trees show that S. wilsoni is closely related, but molecularly separated from clades A and B, and, also based on a unique combination of corallite and sub-corallite characters, the species is moved into Australophyllia gen. nov. These findings confirm the need for using both genetic and morphological datasets for the ongoing taxonomic revision of scleractinian corals.

Scleractinian corals are affected by environment-induced phenotypic plasticity and intraspecific morphological variation caused by genotype. In an effort to identify new strategies for resolving this taxonomic issue, we applied a molecular approach for species evaluation to two closely related genera, Echinophyllia and Oxypora, for which few molecular data are available. A robust multi-locus phylogeny using DNA sequence data across four loci of both mitochondrial (COI, ATP6-NAD4) and nuclear (histone H3, ITS region) origin from 109 coral colonies was coupled with three independent putative species delimitation methods based on barcoding threshold (ABGD) and coalescence theory (PTP, GMYC). Observed overall congruence across multiple genetic analyses distinguished two traditional species (E. echinoporoides and O. convoluta), a species complex composed of E. aspera, E. orpheensis, E. tarae, and O. glabra, whereas O. lacera and E. echinata were indistinguishable with the sequenced loci. The combination of molecular species delimitation approaches and skeletal character observations allowed the description of two new reef coral species, E. bulbosa sp. n. from the Red Sea and E. gallii sp. n. from the Maldives and Mayotte. This work demonstrated the efficiency of multi-locus phylogenetic analyses and recently developed molecular species delimitation approaches as valuable tools to disentangle taxonomic issues caused by morphological ambiguities and to re-assess the diversity of scleractinian corals.

In the context of rising seawater temperatures associated with climate change, the issue of whether coral holobionts deal with this challenge by shuffling their associations with stress- and/or heat-tolerant Symbiodinium, by generating heat-resistant host genotypes, or both is important for coral survival. In this study, the composition of communities of the endosymbiont Symbiodinium and the population genetics of the coral host Platygyra verweyi were examined in a reef impacted by hot-water discharged from the outlet of a nuclear power plant in operation in Kenting, Southern Taiwan since 1984. The water at this site is 2.0-3.0 °C warmer than adjacent reefs in summer, which have an average seawater temperature of 29.0 °C. The data were compared with those for the same species at other sites within 12 km of the outlet site. Platygyra verwyei was associated with one or both of Symbiodinium types C3 (heat sensitive) and D1a (heat tolerant) at all sites with the latter being the dominant at the nuclear power plant outlet. The proportion of C3 in populations increased gradually with increasing distance from the hot-water discharge. Genetic analysis of the Platygyra verweyi host using mitochondrial and nuclear markers showed no genetic differentiation among sites. Changes in the composition of Symbiodinium types associated with P. verweyi among closely located sites in Kenting suggested that this coral might have acclimatized to the constant thermal stress by selective association with heat-tolerant Symbiodinium types, whereas the role of the host in adaptation was inconclusive.

The scale and intensity of marine cage culture have increased in the Asian-Pacific region, particularly in oligotrophic waters where coral reef organisms flourish. In this study, the influence of marine cage culture on subtropical coral communities in turbid waters was evaluated by measuring environmental parameters and benthic community compositions at Magongwan in the Penghu Islands, Taiwan. A canonical discriminant analysis of environmental parameters revealed that elevated levels of ammonium, nitrite, and chlorophyll a (chl a) released from the cages were the main pollution indicators that, in addition to sedimentation and turbidity, distinguished Impact Zone 1 (cage-culture zone) from the other 2 zones - Impact Zone 2 (800 m away from the cages) and the reference zone - in these turbid waters. Results of the canonical correlation analysis indicated that the coverage extents of macroalgae, sponges, and zoanthids were strongly correlated with levels of ammonium, nitrite, phosphate, chl a, and dissolved oxygen. Coral communities in Impact Zone 1 were mostly composed of stress-tolerant massive and submassive corals, but were lacking branching Acropora corals. In contrast, coral communities in the other zones, with high habitat complexity and species richness, were dominated by coral species with diverse morphologies, including branching Acropora coral communities. These results suggest that marine cage culture has been causing chronic nutrient enrichment in the surrounding waters at Magongwan, which may have resulted in a deterioration of suitable habitats for coral reef organisms. Nevertheless, intermediate levels of nutrients and particulate organic matter relative to the other zones might have been caused by the adjacent cage culture, resulting in the high coral coverage and diversity in the Impact Zone 2 in particular.

Terpios hoshinota, a dark encrusting sponge, is known to be a competitor for space in coral reef environments, and facilitates the death of corals. Although numerous cyanobacteria have been detected in the sponge, little is known of the sponge-associated bacterial community. This study examined the sponge-associated bacterial community and the difference between the bacterial communities in the sponge and the coral partially covered by the sponge by analysis of 16S rRNA gene sequences of samples isolated from the sponge covering the corals Favia complanata, Isopora palifera, Millepora sp., Montipora efflorescens and Porites lutea. The sponge-associated bacterial community was mainly (61-98%) composed of cyanobacteria, with approximately 15% of these alphaproteobacteria and gammaproteobacteria, although the proportions varied in different sponge samples. The dominant cyanobacteria group was an isolated group closely related to Prochloron sp. The comparison of the bacterial communities isolated from sponge-free and the sponge-covered P. lutea showed that covering by the sponge caused changes in the coral-associated bacterial communities, with the presence of bacteria similar to those detected in black-band disease, suggesting the sponge might benefit from the presence of bacteria associated with unhealthy coral, particularly in the parts of the coral closest to the margin of the sponge.

Pocillopora verrucosa is a widely distributed depth-generalist coral that presents plasticity in its skeletal macro- and microstructure in response to environmental gradients. Light and water movement, which covary with depth, are the main environmental drivers of morphological plasticity in this genus; however, assessing environmentally-induced plasticity may be confounded by the extent of interspecific variation in Pocillopora. We examine the morphology of 8 typed P. verrucosa specimens collected along a depth gradient ranging from 7 to 45 meters and comprising 3 sites throughout Ludao, Taiwan. We measured 36 morphological characters, 14 which are novel, in 3 regions on the corallum—the apex, branch and base—in order to quantify their relationship to site and depth. We found significant correlation between depth and 19 morphological characters, notably branch verruca area, branch verruca height, base verruca spacing, base spinule length, and branch corallite area. 60% of microstructural characters and 25% of macrostructural characters showed a correlative relation to depth, suggesting that depth acclimatization is manifested primarily at the microstructural level. Canonical discriminant analysis of all morphometric characters by depth supports clustering into 3 groups: an overlapping 7m and 15m group, a 23-30m group, and a 38-45m group. Canonical discriminant analysis by site supports clustering into low- and high-current sites, differentiated primarily by branch septa width, base septa width, pre-terminal branch width, terminal branch maximum length, and terminal branch minimum length. We conclude that distinctive patterns of morphological variation in mesophotic specimens of P. verrucosa could reflect the effects of abiotic parameters such as light and water flow. Elucidating the mechanisms behind the morphological changes that occur in response to environmental gradients can help clarify the role that physiological plasticity plays in the acclimatization of corals to the unique environmental settings of mesophotic coral ecosystems.

A Gram-negative, aerobic, rod-shaped bacterium, designated strain CL-33T, was isolated from the encrusting pore coral Montipora aequituberculata collected from seawater off the coast of southern Taiwan. 16S rRNA gene sequence analysis showed that the strain clustered closely with Endozoicomonas elysicola MKT110T (96.7 % similarity). The novel strain required NaCl for growth and exhibited optimal growth at 25 °C and in the presence of 2–3 % NaCl. Predominant cellular fatty acids were summed feature 3 (C16 : 1 ω7c and/or C16 : 1 ω6c; 39.6 %), summed feature 8 (C18 : 1 ω7c and/or C18 : 1 ω6c; 32.8 %) and C16 : 0 (12.0 %). The DNA G+C content of strain CL-33T was 50.0 mol%. The results of physiological and biochemical tests allowed the clear phenotypic differentiation of this isolate from E. elysicola. It is evident from the genotypic, phenotypic and chemotaxonomic data presented that strain CL-33T represents a novel species of the genus Endozoicomonas, for which the name Endozoicomonas montiporae sp. nov. is proposed. The type strain is CL-33T (=LMG 24815T =BCRC 17933T).

Species of bacteria associated with Stylophora pistillata were determined by analyses of 16S ribosomal genes. Coral samples were taken from two distinct sites at Kenting, in the far south of Taiwan; three coral colonies at each site were tagged and sampled in the winter and summer of 2007. Six hundred 16S rRNA gene clones were selected and sequenced for diversity analysis and community comparison. LIBSHUFF and nonparametric multiple dimensional scaling analyses showed variations in the composition of the coral-associated bacteria in the different samples, suggesting that seasonal and geographic factors and variations in individual coral colonies were all vital drivers of the structure of the S. pistillata -associated bacterial community. To examine the association between species specificity and environmental impacts on the structure of the coral-associated bacterial community, we conducted an integrated, comparative analysis of 44 coral-associated bacterial data sets, including the present study's data. The clustering analysis suggests that the influence of spatial and temporal factors on the coral-associated bacteria population structure is considerable; nonetheless, the effect of species specificity is still detectable in some coral species, especially those from the Caribbean Sea.

Biogeographic investigations have suggested that coral-symbiont associations can adapt to higher temperatures by hosting a heat-tolerant Symbiodinium, phylotype D. It is hypothesized that phylotype D is absent in high latitudes due to its heat-tolerant characteristics. In this study, this hypothesis was tested by examining the symbiont diversity in a scleractinian coral, Oulastrea crispata, throughout its entire latitudinal distribution range in the West Pacific. Molecular phylotyping of the 5′-end of the nuclear large subunit of ribosomal DNA (lsu rDNA) indicated that phylotype D was the dominant Symbiodinium in O. crispata from the tropical reefs to the marginal non-reefal coral communities. Several colonies of tropical populations were associated with phylotype C, either alone or simultaneously with phylotype D. Analysis of the polymerase chain reaction products using single-strand conformation polymorphism (SSCP) detected relatively low densities of phylotype C in most of the O. crispata colonies surveyed. These results provide evidence for the occurrence of phylotype D in cold-water outlying coral communities. The dominant occurrence of phylotype C in some O. crispata colonies on tropical reefs and the relatively low densities of phylotype C identified by SSCP in subtropical and temperate populations show that the dominant symbiont type can vary in this coral species and that multiple symbionts can co-occur in the same host.

Seasonal dynamics of algal symbiont communities from the reef flat-dwelling coral Acropora palifera was monitored from January 2000 to July 2001 at Tantzei Bay, Kenting National Park, southern Taiwan. Zooxanthellae density and pigment contents monitored throughout the 18 mo sampling period indicated that no significant bleaching occurred in A. palifera. Molecular phylo- typing showed that 2 distinct symbionts, Phylotypes C and D, were associated with A. palifera, either individually or simultaneously. Population surveys throughout the bimonthly sampling period dem- onstrated that the symbiont community within A. palifera was highly dynamic with significant fluctu- ations; a drop in the Phylotype D percentage occurred in May, which was correlated with a rise in seawater temperature in the late spring/early summer at the sampled reef. Direct tracking of tagged corals provided evidence that seasonal fluctuations in the algal symbiont communities not only involve changes in zooxanthellae densities and pigment contents, but also a reshuffling of different Symbiodinium phylotypes. Our study highlights that stress tolerance among phylotypes urgently needs to be re-evaluated through a better understanding of the physiological traits of symbionts.

We examined zooxanthellae diversity in scleractinian corals from southern Taiwan and the Penghu Archipelago, a tropical coral reef and a subtropical non-reefal community, respectively. Zooxanthellae diversity was investigated in 52 species of scleractinian corals from 26 genera and 13 families, using restriction fragment length polymorphism (RFLP), and phylogenetic analyses of the nuclear small-subunit ribosomal DNA (nssrDNA) and large-subunit ribosomal DNA (nlsrDNA). RFLP and phylogenetic analyses of nuclear-encoded ribosomal RNA genes showed that Symbiodinium clade C was the dominant zooxanthellae in scleractinian corals in the seas around Taiwan; Symbiodinium clade D was also found in some species. Both Symbiodinium clade C and D were found in colonies of seven species of scleractinian corals. Symbiodinium clade D was associated with corals that inhabit either shallow water or the reef edge in deep water, supporting the hypothesis that Symbiodinium clade D is a relatively stress-tolerant zooxanthellae found in marginal habitats.

International audience; Marine invertebrates are particularly vulnerable to climatic anomalies in early life history stages because of the time spent in the water column. Studies have focused on the effect of seawater temperature on fertilization, development, and larval stages in corals; however, none of them show comparative results along an environmental gradient. In this study, we show that temperatures in the range of 15–336C have strong effects on fertilization rates and embryonic stages of two coral species, Acropora muricata in the subtropical environment and Acropora hyacinthus in subtropical and temperate environments. Deformations after the first cleavage stages were observed at low (156C) and high (336C) temperatures. Development was delayed by 6–7 h in the slightly non-optimal temperature of 206C. We found significant differences in fertilization rates and responses of embryos from different latitudes, with temperate corals being more sensitive to extremely hot temperatures and vice versa. We hypothesize that the coral development is restricted to a narrow temperature range and deviation outside this window could inhibit a species' continuance and ecological success. Thus, it would have significant negative effects on adult populations and communities, playing a role in future of coral reef survival.

Climate change has led to a decline in the health of corals and coral reefs around the world. Studies have shown that, while some corals can cope with natural and anthropogenic stressors either through resistance mechanisms of coral hosts or through sustainable relationships with Symbiodinium clades or types, many coral species cannot. Here, we show that the corals present in a reef in southern Taiwan, and exposed to long-term elevated seawater temperatures due to the presence of a nuclear power plant outlet (NPP OL), are unique in terms of species and associated Symbiodinium types. At shallow depths (

The resilience of coral reefs relies significantly on the ability of corals to recover successfully in algal-dominated environments. Larval settlement is a critical but highly vulnerable stage in the early life history of corals. In this study, we analyzed how the presence of two upright fleshy algae, Sargassum mcclurei (SM) and Padina australis (PA), and one crustose coralline algae, Mesophyllum simulans (MS), affects the settlement of Acropora muricata larvae. Coral larvae were exposed to seawater flowing over these algae at two concentrations. Larval settlement and mortality were assessed daily through four variables related to their behavior: swimming, substratum testing, metamorphosis, and stresses. Temperature, dissolved oxygen, pH, algal growth, and photosynthetic efficiency were monitored throughout the experiment. Results showed that A. muricata larvae can settle successfully in the absence of external stimuli (63 ± 6 % of the larvae settled in control treatments). While algae such as MS may stimulate substrate testing and settlement of larvae in the first day after competency, they ultimately had a lower settlement rate than controls. Fleshy algae such as PA, and in a lesser measure SM, induced more metamorphosis than controls and seemed to eventually stimulate settlement. A diverse combination of signals and/or modifications of microenvironments by algae and their associated microbial communities may explain the pattern observed in coral settlement. Overall, this study contributes significantly to the knowledge of the interaction between coral and algae, which is critical for the resilience of the reefs.

Only a few scleractinian species have been reported from the high-latitudine outlying coral communities of South Korea (Song 2004). Here, we provide the first record of the reef coral Psammocora albopicta (Fig. 1a; see Online Resource, Fig. S1 for details of the skeletal structure) from Jeju Island (33.4°N), located in the southwest of the Korean peninsula. This species, recently described by Benzoni (2006), has previously been recorded from the Arabian Gulf, the Gulf of Aden, Indonesia, Japan, Malaysia, Philippines, and southeast Australia, but this is to date the first record in such a high-latitude area. P. albopicta corals were observed in October 2012 between 5 and 10 m depth at 3 sites around Jeju: Biyangdo, Jigwido, Seogwipo. At each site, only a few colonies ( 50 cm for the maximum length), suggests that their settlement was not recent. Nevertheless, the population could be in expansion which deserves further investigations. Typical circular white patterns on the surface of P. albopicta (see Benzoni 2006), were only observed at one of the sites surveyed, Seogwipo, and on a limited number of colonies. On the specimens observed, this suggests that the presence of this discoloration may have a different origin than previously thought. White rings refer to circular depigmentation of ∼1 cm in diameter (Fig. 1b), without any direct evidence of partial mortality of the colonies. They look very similar to the white rings reported on the

Acropora species distribution patterns at Larak, Farur, and Khark Islands along the Iranian coast were investigated from October 2008 to November 2009 based on a previous study conducted on the taxonomy of Acropora by examining 1,169 colonies. The analysis showed that the species diversity and evenness were low at the three islands, but the total percentage cover of Acropora was relatively high at Larak (38.52%) with lower cover at Khark (24.01%). The Acropora community at Larak Island was significantly dissimilar to those at Farur and Khark Islands, whereas the latter two possessed similar community structures. Chlorophyll a differences of Larak with Farur and Khark Islands coincided with Acropora community differences among these islands. Relatively high coral coverage found at the islands off the Iranian coast suggests important implications for coral reef resilience and mitigation (e.g., assisted migration) in response to changing environmental conditions, not only for the Persian Gulf but also for the world's coral reefs under the impacts of climate change.

Stylophora pistillata is a widely used coral "lab-rat'' species with highly variable morphology and a broad biogeographic range (Red Sea to western central Pacific). Here we show, by analysing Cytochorme Oxidase I sequences, from 241 samples across this range, that this taxon in fact comprises four deeply divergent clades corresponding to the Pacific-Western Australia, Chagos-Madagascar-South Africa, Gulf of Aden-Zanzibar-Madagascar, and Red Sea-Persian/Arabian Gulf-Kenya. On the basis of the fossil record of Stylophora, these four clades diverged from one another 51.5-29.6 Mya, i.e., long before the closure of the Tethyan connection between the tropical Indo-West Pacific and Atlantic in the early Miocene (16-24 Mya) and should be recognised as four distinct species. These findings have implications for comparative ecological and/or physiological studies carried out using Stylophora pistillata as a model species, and highlight the fact that phenotypic plasticity, thought to be common in scleractinian corals, can mask significant genetic variation.

BackgroundSeawater temperature is the main factor restricting shallow-water zooxanthellate coral reefs to low latitudes. As temperatures increase, coral species and perhaps reefs may move into higher-latitude waters, increasing the chances of coral reef ecosystems surviving despite global warming. However, there is a growing need to understand the structure of these high-latitude coral communities in order to analyze their future dynamics and to detect any potential changes.Methodology/principal findingsThe high-latitude (32.75°N) community surveyed was located at Tatsukushi, Shikoku Island, Japan. Coral cover was 60±2% and was composed of 73 scleractinian species partitioned into 7 functional groups. Although only 6% of species belonged to the 'plate-like' functional group, it was the major contributor to species coverage. This was explained by the dominance of plate-like species such as Acropora hyacinthus and A. solitaryensis. Comparison with historical data suggests a relatively recent colonization/development of A. hyacinthus in this region and a potential increase in coral diversity over the last century. Low coverage of macroalgae (2% of the benthic cover) contrasted with the low abundance of herbivorous fishes, but may be reasonably explained by the high density of sea urchins (12.9±3.3 individuals m⁻²).Conclusions/significanceThe structure and composition of this benthic community are relatively remarkable for a site where winter temperature can durably fall below the accepted limit for coral reef development. Despite limited functionalities and functional redundancy, the current benthic structure might provide a base upon which a reef could eventually develop, as characterized by opportunistic and pioneer frame-building species. In addition to increasing seawater temperatures, on-going management actions and sea urchin density might also explain the observed state of this community. A focus on such 'marginal' communities should be a priority, as they can provide important insights into how tropical corals might cope with environmental changes.

Although coral reefs are facing severe challenges from a variety of natural and anthropogenic stresses, there is anecdotal evidence that the high-latitude coral species Alveopora japonica Eguchi, 1968, has increased its population over the past two decades around Jeju Island, off the southern coast of Korea. The present study provides the first ecological data on this species. Alveopora japonica is opportunistically occupying the empty space left vacant following the recent kelp-forest decline. Colony abundance, age- and size-frequency distributions, lifespan, growth rates and biological characteristics such as surface area, weight and volume of two A. japonica populations in Jeju Island were investigated. Alveopora japonica around Jeju Island is characterised by a mean colony size of 30cm2, a slow growth rate (4.8mm year–1), and a short lifespan of 12–13 years, as determined by X-radiographic measurements. Alveopora japonica presented a dense population of 120 colonies m–2 on average. Population-age and -size structures at both sites reflected a healthy status and indicated a local stability, with a stationary size structure allowing population maintenance over time. The present study provided data to develop population-dynamics models to predict the potential outcomes of A. japonica populations to alternative management scenarios in Jeju Island.

The scleractinian fauna of Taiwan is well-known and accounts for 316 species (Huang et al. 2014). However, most of the coral diversity surveys here have been limited to shallow waters, and benthic biodiversity below the 30-m depth limit of recreational diving remains largely unexplored. In February 2014, the starry coral Psammocora stellata (Verrill, 1866) was recorded for the first time in Taiwanese waters from a mesophotic reef off Green Island (121.47948° E, 22.63889° N). A free-living colony was observed lying on a sandy bottom, by scuba diving at 51 m depth (Fig. 1a). The brown colony was ~10 cm at its largest diameter and was composed of contorted branches with rounded tips. It was colonized by a variety of epibionts, mainly hydrozoans and bryozoans. Microscopic observations (Fig. 1b–c) showed the elongated shape of septa and multiple columellar processes, thus confirming the specimen (ASIZC0000991, Biodiversity Research Museum of Academia Sinica, Taiwan) identification based on the description given by Stefani et al. (2008). For a long time, the distribution of P. stellata has been considered as limited to Hawaii, Micronesia and the tropical Eastern Pacific (Veron 2000). Extension of the presence of this species to Australia, the Indian Ocean, the Red Sea, the Persian Gulf (see Stefani et al. 2008; but also Hoeksema and Best 1994) and now to Taiwan, suggests that this species has been overlooked or confused with other species (such as P. contigua), possibly due to the ecophenotypic variation in colony morphology (Stefani et al. 2008). In Taiwan, P. stellata could actually be restricted to mesophotic environments. This could explain why this species has gone unnoticed until now. Aside from extending the known distribution range of P. stellata and adding this species to the present knowledge of the scleractinian fauna of Taiwan, the observation presented here also draws attention to the possible importance of mesophotic benthic assemblages for the coral reef diversity in this area.

Both bacteria and algal symbionts (genus Symbiodinium), the two major microbial partners in the coral holobiont, respond to fluctuations in the environment, according to current reports; however, little evidence yet indicates that both populations have any direct interaction with each other in seasonal fluctuation. In this study, we present field observations of a compositional change in bacteria and Symbiodinium in the coral Isopora palifera in three separate coral colonies following monthly sampling from February to November in 2008. Using massively parallel pyrosequencing, over 200 000 bacterial V6 sequences were classified to build the bacterial community profile; in addition, the relative composition and quantity of Symbiodinium clades C and D were determined by real-time PCR. The results showed that coral-associated bacterial and Symbiodinium communities were highly dynamic and dissimilar among the tagged coral colonies, suggesting that the effect of host specificity was insignificant. The coral-associated bacterial community was more diverse (Shannon index up to 6.71) than previous estimates in other corals and showed rapid seasonal changes. The population ratios between clade C and D groups of Symbiodinium varied in the tagged coral colonies through the different seasons; clade D dominated in most of the samples. Although significant association between bacteria and symbiont was not detected, this study presents a more detailed picture of changes in these two major microbial associates of the coral at the same time, using the latest molecular approaches.

Terpios hoshinota, an encrusting cyanosponge, is known as a strong substrate competitor of reef-building corals that kills encountered coral by overgrowth. Terpios outbreaks cause significant declines in living coral cover in Indo-Pacific coral reefs, with the damage usually lasting for decades. Recent studies show that there are morphological transformations at a sponge's growth front when confronting corals. Whether these morphological transformations at coral contacts are involved with physiological outperformance (e.g., higher metabolic activity or nutritional status) over other portions of Terpios remains equivocal. In this study, we compared the indicators of photosynthetic capability and nitrogen status of a sponge-cyanobacteria association at proximal, middle, and distal portions of opponent corals. Terpios tissues in contact with corals displayed significant increases in photosynthetic oxygen production (ca. 61%), the δ13C value (ca. 4%), free proteinogenic amino acid content (ca. 85%), and Gln/Glu ratio (ca. 115%) compared to middle and distal parts of the sponge. In contrast, the maximum quantum yield (Fv/Fm), which is the indicator usually used to represent the integrity of photosystem II, of cyanobacteria photosynthesis was low (0.256~0.319) and showed an inverse trend of higher values in the distal portion of the sponge that might be due to high and variable levels of cyanobacterial phycocyanin. The inconsistent results between photosynthetic oxygen production and Fv/Fm values indicated that maximum quantum yields might not be a suitable indicator to represent the photosynthetic function of the Terpios-cyanobacteria association. Our data conclusively suggest that Terpios hoshinota competes with opponent corals not only by the morphological transformation of the sponge-cyanobacteria association but also by physiological outperformance in accumulating resources for the battle.

Analyses of mitochondrial sequences revealed the existence of a group I intron in the cytochrome oxidase subunit 1 (cox1) gene in 13 of 41 genera (20 out of 73 species) of corals conventionally assigned to the suborder Faviina. With one exception, phylogenies of the coral cox1 gene and its intron were concordant, suggesting at most two insertions and many subsequent losses. The coral introns were inferred to encode a putative homing endonuclease with a LAGLI-DADG motif as reported for the cox1 group I intron in the sea anemone Metridium senile. However, the coral and sea anemone cox1 group I introns differed in several aspects, such as the intron insertion site and sequence length. The coral cox1 introns most closely resemble the mitochondrial cox1 group I introns of a sponge species, which also has the same insertion site. The coral introns are also more similar to the introns of several fungal species than to that of the sea anemone (although the insertion site differs in the fungi). This suggests either a horizontal transfer between a sponge and a coral or independent transfers from a similar fungal donor (perhaps one with an identical insertion site that has not yet been discovered). The common occurrence of this intron in corals strengthens the evidence for an elevated abundance of group I introns in the mitochondria of anthozoans.