Heterotrophy in marine animal forests in an era of climate change.

Marine animal forests (MAFs) are benthic ecosystems characterised by biogenic three-dimensional structures formed by suspension feeders such as corals, gorgonians, sponges and bivalves. They comprise highly diversified communities among the most productive in the world's oceans. However, MAFs are in decline due to global and local stressors that threaten the survival and growth of their foundational species and associated biodiversity. Innovative and scalable interventions are needed to address the degradation of MAFs and increase their resilience under global change. Surprisingly, few studies have considered trophic interactions and heterotrophic feeding of MAF suspension feeders as an integral component of MAF conservation. Yet, trophic interactions are important for nutrient cycling, energy flow within the food web, biodiversity, carbon sequestration, and MAF stability. This comprehensive review describes trophic interactions at all levels of ecological organisation in tropical, temperate, and cold-water MAFs. It examines the strengths and weaknesses of available tools for estimating the heterotrophic capacities of the foundational species in MAFs. It then discusses the threats that climate change poses to heterotrophic processes. Finally, it presents strategies for improving trophic interactions and heterotrophy, which can help to maintain the health and resilience of MAFs.

Coral transplantation in urban environments: Insights from colony survival and growth on artificial frames versus the seabed.

Urban and green transitions require infrastructures that can cause pressure on coastal ecosystems. In northern Taiwan, plans to convert an oil-fired power plant to gas would necessitate a port terminal construction, threatening nearby corals and marine life. To mitigate construction impacts, the relocation of affected corals was proposed. We conducted a transplantation study, prior to such a large-scale coral relocation, to assess its feasibility and to identify potential risks associated with the marginal location of northern Taiwan for tropical corals. Five coral species, representative of the different ecological strategies, were selected. We used two methods (artificial frames and seabed cementation) to transplant 246 colony fragments to two pre-selected sites. Over a year, we monitored fragment survival and growth, in parallel with environmental conditions. We found that survival and growth were significantly influenced by transplantation methods, sites, and species. The difference between methods revealed biotic (predation by corallivorous snails) and abiotic (mechanical damage by waves) factors affecting coral survival and growth. Acropora species exhibited high growth, but also high mortality, consistent with their known ecology. Other species presented slower growth but higher survival. One site provided a better environment for corals, which we attributed to topography and reduced exposure. Overall, this study provides interesting insights into relocating corals in a high-latitude and urban coral ecosystem, highlighting risks related to mechanical damages and predation.

Macroalgae exhibit diverse responses to human disturbances on coral reefs.

Scientists and managers rely on indicator taxa such as coral and macroalgal cover to evaluate the effects of human disturbance on coral reefs, often assuming a universally positive relationship between local human disturbance and macroalgae. Despite evidence that macroalgae respond to local stressors in diverse ways, there have been few efforts to evaluate relationships between specific macroalgae taxa and local human-driven disturbance. Using genus-level monitoring data from 1205 sites in the Indian and Pacific Oceans, we assess whether macroalgae percent cover correlates with local human disturbance while accounting for factors that could obscure or confound relationships. Assessing macroalgae at genus level revealed that no genera were positively correlated with all human disturbance metrics. Instead, we found relationships between the division or genera of algae and specific human disturbances that were not detectable when pooling taxa into a single functional category, which is common to many analyses. The convention to use percent cover of macroalgae as an indication of local human disturbance therefore likely obscures signatures of local anthropogenic threats to reefs. Our limited understanding of relationships between human disturbance, macroalgae taxa, and their responses to human disturbances impedes the ability to diagnose and respond appropriately to these threats.

A molecular census of early-life stage scleractinian corals in shallow and mesophotic zones.

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.

Strong horizontal and vertical connectivity in the coral Pocillopora verrucosa from Ludao, Taiwan, a small oceanic island.

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, p<0.05) between two adjacent locations (Dabaisha 23-30 and Dabaisha 38-45 m), but no differentiation elsewhere, suggesting high levels of connectivity among sites and depths. STRUCTURE analysis showed no genetic clustering among sites or depths, indicating that all Pocillopora 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.

Impacts of heat stress and storm events on the benthic communities of Kenting National Park (Taiwan).

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.

Learning from differences: Abiotic determinism of benthic communities in Northern Taiwan.

Differences in the initial configuration of ecological communities may lead to contrasting trajectories when facing environmental changes. Here, we propose to uncover the determinism of benthic communities by carrying out a detailed investigation of their response to small-scale modification of environmental conditions, including physical, chemical, and geological factors. At ten locations (confounding site and depth) in Northern Taiwan, communities were delineated using a morpho-functional classification of the organisms. A k-means clustering was used to identify k homogenous groups among transects. Their environmental determinism was examined by combining this result with 16 environmental variables of transect conditions into a regression tree framework. Biotic and abiotic data were further analyzed with a Multivariate Regression Tree (MRT) to ascertain the hierarchical environmental determinism. The classifications produced by both approaches were compared using the Adjusted Rand index (ARI) to assess the predictive power of unsupervised clustering on its missing explanatory components (abiotic variables). k-means and MRT produced five clusters, respectively, with a similarity of 0.82 in ARI. Wave motion, followed by substrate types resolved most of the variance, while chemical factors in this study were uniform throughout the region. Comparable structures for both methods (clustering groups) demonstrated that the delineated clusters matched with contrasting environmental conditions which could be explained by the existence of various benthic communities. Further consideration of these different communities and their environmental context will be important in determining their trajectories under global changes and may help in the interpretation of community modifications with changing environmental conditions.

The plasticity of gonad development of sexual reproduction in a scleractinian coral, Porites lichen.

Metazoans have evolved a complexity of sexual system and gonad development, however, sexual reproduction of scleractinian corals is not well understood. This study aimed to address the sexual system and gametogenesis in Porites lichen, a common species in the Indo-West Pacific. This study represents the first description of sexual system, which were determined by histological analysis of the samples collected in northern Taiwan. In addition, female and hermaphroditic colonies were separately cultured in aquarium to further monitor the release of eggs/larvae and thereby confirm the breeding system. The results demonstrate that P. lichen is a polygamodioecious brooder and displays seasonal gametogenesis and embryogenesis that ends in late summer. In hermaphroditic colonies, male polyps are predominant and hermaphroditic polyps make up a very small percent (1%-19.3%). In addition, two new gametogenic features were observed from the histological analysis: 1) oocytes developed within the spermaries in hermaphroditic polyps during the early stage of gametogenesis and 2) melanin granular cells were clustered in spermaries in both male and hermaphroditic colonies. This study demonstrated the plasticity of gametogenesis and melanin related cells appeared in corals, which provides an important information to explore hormones and molecular mechanism involving in gonadal arrangement and production of melanin for further studies.

Toward a standardised protocol for the stable isotope analysis of scleractinian corals.

RATIONALE: The stable isotope analysis of carbon and nitrogen is a powerful tool in many ecological studies, but different sample treatments may affect stable isotope ratios and hamper comparisons among studies. The goal of this study was to determine whether treatments that are commonly used to prepare scleractinian coral samples for stable isotope analysis yield different δ15 N and δ13 C values, and to provide guidelines toward a standardised protocol. METHODS: The animal tissues and Symbiodiniaceae of two symbiotic scleractinian coral species (Stylophora pistillata and Porites lutea) were divided into subsamples to test the effects of the drying method, lipid extraction, acidification treatment and water washing. All the subsamples were analysed for their δ15 N and δ13 C values, using continuous flow elemental analyser/isotope ratio mass spectrometry. RESULTS: The drying method and lipid extraction treatment had no substantial effects on the δ15 N and δ13 C values of Symbiodiniaceae and animal tissues. Acid treatment did cause significant differences in δ13 C values (mean differences ≤0.5‰, with individual samples becoming up to 2.0‰ more negative), whereas no ecologically significant differences were observed in δ15 N values. Animal tissue δ13 C values may vary depending on whether samples are washed or not. CONCLUSIONS: To move towards a standardised protocol in coral research, we recommend using an available drying method (as they are equally acceptable) for the stable isotope analysis of scleractinian corals, examining the need for lipid extraction on a case-by-case basis, performing a direct acidification of Symbiodiniaceae and animal tissues, and avoiding washing animal tissue with distilled water.

Social-environmental drivers inform strategic management of coral reefs in the Anthropocene.

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.

Temporal variation and photochemical efficiency of species in Symbiodinaceae associated with coral Leptoria phrygia (Scleractinia; Merulinidae) exposed to contrasting temperature regimes.

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.

Metagenomic, phylogenetic, and functional characterization of predominant endolithic green sulfur bacteria in the coral Isopora palifera.

BACKGROUND: Endolithic microbes in coral skeletons are known to be a nutrient source for the coral host. In addition to aerobic endolithic algae and Cyanobacteria, which are usually described in the various corals and form a green layer beneath coral tissues, the anaerobic photoautotrophic green sulfur bacteria (GSB) Prosthecochloris is dominant in the skeleton of Isopora palifera. However, due to inherent challenges in studying anaerobic microbes in coral skeleton, the reason for its niche preference and function are largely unknown. RESULTS: This study characterized a diverse and dynamic community of endolithic microbes shaped by the availability of light and oxygen. In addition, anaerobic bacteria isolated from the coral skeleton were cultured for the first time to experimentally clarify the role of these GSB. This characterization includes GSB's abundance, genetic and genomic profiles, organelle structure, and specific metabolic functions and activity. Our results explain the advantages endolithic GSB receive from living in coral skeletons, the potential metabolic role of a clade of coral-associated Prosthecochloris (CAP) in the skeleton, and the nitrogen fixation ability of CAP. CONCLUSION: We suggest that the endolithic microbial community in coral skeletons is diverse and dynamic and that light and oxygen are two crucial factors for shaping it. This study is the first to demonstrate the ability of nitrogen uptake by specific coral-associated endolithic bacteria and shed light on the role of endolithic bacteria in coral skeletons.

Biogeography of functional trait diversity in the Taiwanese reef fish fauna.

The richness of Taiwanese reef fish species is inversely correlated to latitude as a direct consequence of the abiotic environment and its effects on benthic habitats. However, to date, no studies have investigated the variations in the diversity of traits (FD) linked with the role of these fishes in the ecosystem. FD is usually considered more sensitive than species richness in detecting early changes in response to disturbances, and therefore could serve as an indicator of ecological resilience to environmental changes. Here, we aim to characterize FD in the Taiwanese reef fish fauna and to document its regional variations. Six traits were used to categorize the 1,484 reef fish species occurring in four environmentally contrasted regions around Taiwan. The number of unique trait combinations (FEs), their richness (FRic), their redundancy (FR), their over-redundancy (FOR), and their vulnerability (FV) were compared among these regions. Overall, 416 FEs were identified. Their number decreased from south to north in step with regional species richness but FRic remained similar among regions. FR and FOR were higher to the south. At the local scale, variations in FEs and FRic are in concordance with the worldwide pattern of FD. High-latitude, impoverished fish assemblages, offer a range of trait combinations similar to diversified tropical assemblages. Increasing diversity in the latter mainly contributes to raising FR and supports already over-redundant entities. High vulnerability makes many combinations highly sensitive to species loss, and was higher at intermediate latitudes when using a fine resolution in trait categories. It suggests that the loss of FEs may first be characterized by an increase in their vulnerability, a pattern that could have been overlooked in previous global scale analyses. Overall, this study provides new insights into reef fish trait biogeography with potential ramifications for ecosystem functioning.

Molecular assessment of Pocillopora verrucosa (Scleractinia; Pocilloporidae) distribution along a depth gradient in Ludao, Taiwan.

It can be challenging to identify scleractinian corals from the genus Pocillopora Lamarck 1816 in the field because of their large range of inter- and intra-specific morphological variation that co-occur with changes in the physical environment. This task is made more arduous in the context of a depth gradient, where light and water current could greatly affect the morphology of the corallum. Pocillopora verrucosa (Ellis & Solander 1786) in Taiwan was previously reported exclusively from shallow waters (<10 m in depth), but a recent observation of this species in the mesophotic zone (>40 m in depth) questions this bathymetric distribution. We used the mitochondrial open reading frame and the histone 3 molecular markers to investigate the vertical and horizontal spatial distribution of P. verrucosa around Ludao (Green Island), Taiwan. We genotyped 101 P. verrucosa-like colonies collected from four depth zones, ranging from 7 to 45 m, at three sites around the island. Of the 101 colonies sampled, 85 were genotyped as P. verrucosa, 15 as P. meandrina, and one specimen as an undescribed Pocillopora species. P. verrucosa was found at all depths, while P. meandrina and the undescribed Pocillopora specimen were limited to 15 m depth. P. verrucosa has a large bathymetric distribution around Ludao and could benefit from the refuge that the mesophotic zone offers. This study illustrates the difficulty of identifying Pocillopora corals in the field and emphasizes the relevance of molecular taxonomy as an important and complementary tool to traditional taxonomy for clarifying vertical and horizontal species distribution. Our results also illustrate the need in conservation biology to target species genetic diversity rather than just species diversity.

Spatial variation in the morphological traits of Pocillopora verrucosa along a depth gradient in Taiwan.

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.

Geographical variations in bacterial communities associated with soft coral Scleronephthya gracillimum.

Environmental impacts can alter relationships between a coral and its symbiotic microbial community. Furthermore, changes in the microbial community associated with increased seawater temperatures can cause opportunistic infections, coral disease and death. Interactions between soft corals and their associated microbes are not well understood. The species Scleronephthya gracillimum is distributed in tropical to temperate zones in coral assemblages along the Kuroshio Current region. In this study we collected S. gracillimum from various sites at different latitudes, and compared composition of their bacterial communities using Next Generation Sequencing. Coral samples from six geographically distinct areas (two sites each in Taiwan, Japan, and Korea) had considerable variation in their associated bacterial communities and diversity. Endozoicimonaceae was the dominant group in corals from Korea and Japan, whereas Mycoplasma was dominant in corals from Taiwan corals. Interestingly, the latter corals had lower relative abundance of Endozoicimonaceae, but greater diversity. These biogeographic differences in bacterial composition may have been due to varying environmental conditions among study locations, or because of host responses to prevailing environmental conditions. This study provided a baseline for future studies of soft coral microbiomes, and assessment of functions of host metabolites and soft coral holobionts.

A functional approach to the structural complexity of coral assemblages based on colony morphological features.

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.

Shifting communities after-- typhoon damage on an upper mesophotic reef in Okinawa, Japan.

Very few studies have been conducted on the long-term effects of typhoon damage on mesophotic coral reefs. This study investigates the long-term community dynamics of damage from Typhoon 17 (Jelawat) in 2012 on the coral community of the upper mesophotic Ryugu Reef in Okinawa, Japan. A shift from foliose to bushy coral morphologies between December 2012 and August 2015 was documented, especially on the area of the reef that was previously recorded to be poor in scleractinian genera diversity and dominated by foliose corals. Comparatively, an area with higher diversity of scleractinian coral genera was observed to be less affected by typhoon damage with more stable community structure due to less change in dominant coral morphologies. Despite some changes in the composition of dominant genera, the generally high coverage of the mesophotic coral community is facilitating the recovery of Ryugu Reef after typhoon damage.

Structure of Benthic Communities along the Taiwan Latitudinal Gradient.

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.