Chemical diversity of scleractinian corals revealed by untargeted metabolomics and molecular networking

The chemical diversity of scleractinian corals is closely related to their physiological,ecological,and evolutionary status,and can be influenced by both genetic background and environmental variables.To investigate intraspecific variation in the metabolites of these corals,the metabolomes of four species(Pocillopora meandrina,Seriatopora hystrix,Acropora formosa,and Fungia fungites)from the South China Sea were analyzed using untargeted mass spectrometry-based metabolomics.The results showed that a variety of metabolites,including amino acids,peptides,lipids,and other small molecules,were differentially distributed among the four species,leading to their significant separation in principal component analysis and hierarchical clustering plots.The higher content of storage lipids in branching corals(P.meandrina,S.hystrix,and A.formosa)compared to the solitary coral(F.fungites)may be due to the high densities of zooxanthellae in their tissues.The high content of aromatic amino acids in P.meandrina may help the coral protect against ultraviolet damage and promote growth in shallow seawater,while nitrogen-rich compounds may enable S.hystrix to survive in various challenging environments.The metabolites enriched in F.fungites,including amino acids,dipeptides,phospholipids,and other small molecules,may be related to the composition of the coral's mucus and its life-history,such as its ability to move freely and live solitarily.Studying the chemical diversity of scleractinian corals not only provides insight into their environmental adaptation,but also holds potential for the chemotaxonomy of corals and the discovery of novel bioactive natural products.

Cembranoids of Soft Corals:Recent Updates and Their Biological Activities

Soft corals are well-known as excellent sources of marine-derived natural products.Among them,members of the genera Sar-cophyton,Sinularia,and Lobophytum are especially attractive targets for marine natural product research.In this review,we reported the marine-derived natural products called cembranoids isolated from soft corals,including the genera Sarcophyton,Sinularia,and Lobophytum.Here,we reviewed 72 reports published between 2016 and 2020,comprising 360 compounds,of which 260 are new compounds and 100 are previously known compounds with newly recognized activities.The novelty of the organic molecules and their relevant biological activities,delivered by the year of publication,are presented.Among the genera presented in this report,Sarcophyton spp.produce the most cembranoid diterpenes;thus,they are considered as the most important soft corals for marine natural product research.Cembranoids display diverse biological activities,includ-ing anti-cancer,anti-bacterial,and anti-inflammatory.As cembranoids have been credited with a broad range of biological activities,they present a huge potential for the development of various drugs with potential health and ecological benefits.

Scleractinian Corals and Reefs of Vietnam as a Part of the Pacific Reef Ecosystem

The paper analyzes both published and unpublished results of the investigations of Vietnamese reef building corals and reefs performed in the last decades of the twentieth century and first decades twenty-first. The state of the art in the study of reef-building scleractinian corals and reefs is presented. The scleractinian fauna of Vietnam is shown to match in species diversity (350 species of 80 genera) the tropical coral fauna of the Indonesian–Malacca fertile center, from which Indo-Pacific reef-building corals originated. The whole Vietnam coast from the Gulf of Tonkin to the Gulf of Siam is a biogeographically single whole and is a part of the Indo-Polynesian Province of the Indo-Pacific Area.

Cool water brought by upwelling in the Sanya Bay benefits corals in the background of global warming

It has been reported that global warming has negative effects on coral ecosystems in the past 50 years and the effects vary in different ocean environment. In order to make clear the coral reef status in the background of global warming along the south coast of Hainan Island of China, satellite and in situ data are used to retrieve the information of the coral reef status and surrounding environmental factors. The results show that cool water induced by upwelling along the south coast of Hainan Island is found in the area every summer month, especially in the relatively strong El Ni？o years（2002–2003 and 2005）. From the NOAA satellite data, degree heating week（DHW） index does not exceed 3 in Sanya Bay even in the relatively strong El Ni？o years. By comparison of a coral reef growth rate in the Sanya Bay with respect to El Ni？o events from 1957 to 2000, coral＇s growth rate is relatively greater during 1972, 1991–1994 and 1998 El Ni？o event. By analyzing the environmental factors, it is found that the cool water induced by upwelling may be the main reason for protecting corals from global warming effects.

Diversity of symbiotic algae of the genus Symbiodinium in scleractinian corals of the Xisha Islands in the South China Sea

Symbiotic algae （Symbiodinium sp.） in scleractinian corals are important in understanding how coral reefs will respond to global climate change. The present paper reports on the diversity of Symbiodinium sp. in 48 scleractinian coral species from 25 genera and 10 families sampled from the Xisha Islands in the South China Sea, which were identified with the use of restriction fragment length polymorphism （RFLP） of the nuclear ribosomal DNA large subunit gene （rDNA）. The results showed that： （i） Symbiodinium Clade C was the dominant zooxanthellae in scleractinian corals in the Xisha Islands; （ii） Symbiodinium Clade D was found in the corals Montipora aequituberculata, Galaxea fascicularis, and Plerogyra sinuosa; and （iii） both Symbiodinium Clades C and D were found simultaneously in Montipora digitata, Psammocora contigua, and Galaxeafascicularis. A poor capacity for symbiosis polymorphism, as uncovered by RFLP, in the Xisha Islands indicates that the scleractinian corals have low adaptability to environmental changes. Further studies are needed to investigate zooxanthellae diversity using other molecular markers.

Morphology and molecular phylogeny of three black corals(Antipatharia,Schizopathidae)from seamounts in the Western Pacific Ocean,with description of a new species

Three deep-sea black corals belonging to the family Schizopathidae,are reported from two seamounts on the Caroline Ridge in the tropical Western Pacific:Umbellapathes parva sp.nov.with depth of 1488-1766 m,Telopathes cf.magna MacIsaac&Best,2013 with depth of 937-1016 m,and Stauropathes cf.punctata(Roule,1905)with depth of 942-1353 m.The latter two species are the first records in the Western Pacific Ocean.U.parva sp.nov.is characterized by monopodial corallum,relatively long unpinnulated stalk with complex pinnulated branche s developing from the lowermost primary pinnules and small spine s.It differs from two known congeners by much smaller spine s and the present of secondary pinnules.T.cf.magna MacIsaac&Best,2013 has sparsely branched corallum,relatively long and simple pinnules arranged in subopposite or alternate pairs.S.cf.punctata(Roule,1905)is characterized by the corallum with almost planar pinnulated branches,and pinnules arranged in subopposite pairs.Our results indicate that the corallum size and shape are variable in conspecific specimens particularly those at different growth stages.By contrast,the size of the polypar spines as well as the abpolypar spines have little variation and can be served as a main differing feature for schizopathid species.The phylogenetic analyses using nuclear internal transcribed spacer region(spanning partial 18 S rDNA,ITS 1,5.8 S,ITS2,and partial 28 S rDNA)and two mitochondrial fragments cox3-IGR-coxl(COI)and trnW-IGR-nad2(NAD2)showed Telopathes,Stau ropathes,and Bathypathes had close relationships,and Umbellapathes formed a sister clade with Alternatipathes.

El Ni?o/Southern Oscillation during the 4.2 ka event recorded by growth rates of corals from the North South China Sea

The 4.2 ka event that occurred during the period from 4 500–3 900 a BP was characterized by cold and dry climates and resulted in the collapse of civilizations around the world. The cause of this climatic event, however, has been under debate. We collected four corals(Porites lutea) from Yongxing Island, Xisha Islands, South China Sea, dated them with the U-series method, and measured the annual coral growth rates using X-ray technology. The dating results showed that the coral growth ages were from 4 500–3 900 a BP, which coincide well with the period of the4.2 ka event. We then reconstructed annual sea surface temperature anomaly(SSTA) variations based on the coral growth rates. The growth rate-based SSTA results showed that the interdecadal SSTA from 4 500–3 900 a BP was lower than that during modern times(1961–2008 AD). A spectral analysis showed that the SSTA variations from4 500–3 900 a BP were under the influence of El Nino-Southern Oscillation(ENSO) activities. From 4 500–4 100 a BP, the climate exhibited La Nina-like conditions with weak ENSO intensity and relatively stable and lower SSTA amplitudes. From 4 100–3 900 a BP, the climate underwent a complicated period of ENSO variability and showed alternating El Nino-or La Nina-like conditions at interdecadal time scales and large SSTA amplitudes. We speculate that during the early and middle stages of the 4.2 ka event, the cold climate caused by weak ENSO activities largely weakened social productivity. Then, during the end stages of the 4.2 ka event, the repeated fluctuations in the ENSO intensity caused frequent extreme weather events, resulting in the collapse of civilizations worldwide. Thus, the new evidence obtained from our coral records suggests that the 4.2 ka event as well as the related collapse of civilizations were very likely driven by ENSO variability.

Results of Thirty Years of Research on Corals and Reefs of Vietnam

The author traced the history of the thirty years of study of the Vietnamese reefs since the second half of the 20th century up to the present time. This paper describes the types of reefs, taxonomic composition and structure of Scleractinian reef ecosystems. It is shown that due to a high level of diverse knowledge revealed, high species richness of coral fauna of Vietnam (376 species of 80 genera) is a single point of origin coral fauna Indo-Pacific. Analysis of knowledge of Vietnamese corals and reefs revealed the extent of their positive or negative status under anthropogenic influence or under the level of wilderness protection, experiment on artificial cultivation of corals and the re-establishment of a coral community.

Structural and functional characterization of a novel ShK peptide and its analogue from zoanthids (corals)

OBJECTIVE To identify and characterize a novel neuroprotective ShK peptide and its analogue originated from coral P.caribaeoru.METHODS P.caribaeoru was collected and subjected to transcriptome sequencing at which further bioinformatics analysis had identified a unigene encoding a putative ShK protein candidate,named as PcShK1.PcShK1 and its rhodamine derivative(PcShK1-RhoB) were synthesized and tested for the neuroprotective effect in 6-OHDA induced Parkinson disease models in vitro and in vivo.Briefly,zebrafish larvae were co-exposed to 6-OHDA and various doses of the peptides;then,dopaminergic(DA) neurons immunoreactivity and locomotion behavior of the zebrafish larvae were examined.Similarly,PC12 cells were cultured with 6-OHDA in the absence or presence of different concentrations of peptides.Cell viability was determined by MTT assay while calcium flow in the PC12 cells was monitored by Fluo-4 fluorescent dye.RESULTS Compared with control group,6-OHDA treatment could lead to DA neurons loss and locomotion deficits in PD model of zebrafish larvae(P<0.01).Both PcS hK 1(2.5,5.0 and 7.5 μmol·L^(-1)) and PcS hK 1-RhoB(0.50 and 0.75 μmol·L^(-1)) were found to protect and restore dopaminergic neurons from6-OHDA mediated injury and locomotion deficiency in the PD zebrafish respectively(P<0.01).In addition,PcShK1(2.5 to 20.0 μmol·L^(-1)) and PcShK1-RhoB(0.6 to 2.5 μmol·L^(-1)) effectively prevented against 6-OHDA toxicity in PC12 cells(P<0.01).Further study revealed that they might exert their neuroprotective effects through regulating the calcium homeostasis.CONCLUSION PcS hK 1 and PcS hK 1-RhoB show neuroprotective effects on 6-OHDA induced PD models,and the underlying protective mechanisms of these peptides probably involve calcium homeostasis regulation.

Triassic Scleractinian Corals in China：A Review of Present Knowledge

Scleractinian corals appeared during the Anisian period of the Triassic after the end-Permian mass extinction, which in Triassic were distinct from the Jurassic-Cretaceous scleractinian corals. The study on Triassic scleractinian coral fauna as a separate field is significant for exploring the development and evolution of modern corals and investigating environmental changes since the Mesozoic. The first Triassic coral in China was reported in 1925, and since then, nearly 25 articles dealing with taxonomy and 17 papers about fossil reports on Triassic scleractinian corals in China have been published, which refer to 60 genera and 312 species, 49 localities and 25 strata. In this paper, the history, taxonomy, localities, stratigraphic distribution, current research, and existing knowledge gaps of the Triassic scleractinian corals in China are reviewed. More specifically, the research findings of the Triassic scleractinain coral in China since 1925 are discussed; the species of the Triassic scleractian coral fossil reported in China, has been examined and its synonyms have been sorted out; the geographical distributions of the Middle and Late Triassic scleractinian corals in China are systematically treated; the Triassic coral biostratigraphy in China has been improved; and the stratigraphic ranges of existing genera and species are provided. The above conclusions are presented in the form of complete figures. At the same time, we analyzed shortcomings in current research and identified productive future research directions of the Triassic scleractinian corals in China.

Fluorescence color diversity of great barrier reef corals

A group of variously colored proteins belonging to the green fAuorescent protein(GFP)family are responsible for coloring coral tissues.Corals of the Great Barrier Reef were studied with the custom-built fiber laser fluorescence spectrometers.Spectral analysis showed that most of the excarmined corals contained multiple fuorescent peaks ranging from 470 to 620nm.This obser-vation was attributed to the presence of multiple genes of GFP-like proteins in a single coral,as well as by the photo-induced post-translational modifcations of certain GFP-like proteins.We isolated a novel photo-convertible fuorescent protein(FP)from one of the tested corals.We:propose that two processes may explain the observed diversity of the fuorescent spectra in corals:(1)dark post-translational modifcation(maturation),and(2)color photo-conversion of certain maturated proteins in response to sunlight.

Resilience of a high latitude Red Sea corals to extreme temperature

Our research objective was to expand the very limited knowledgebase pertaining to the ecology of fringing coral reefs in the Gulf of Suez, Egypt. Specifically, determine dominant coral species and investigate why this reef is capable of surviving at such a high-latitude and extreme harsh environment. Data collection included annual reef surveys, randomized quadrat sampling, five permanent video transects and in situ seawater temperature. Of the known Gulf of Suez 35 taxa, only six (Acropora humilis, A. microclados, A. hemprichii, Litophyton arboretum, Stylophora pistillata, Porites columna, and P. plantulata), compose 94% of the reef's coral cover. Coral dominance across species shifted drastically during the study period. However, the six coral dominance remained unchanged, while some decreased others increased. These six coral taxa regularly experience daily changes in seawater temperature and seasonal variations that exceed These extreme temperatures variation and the fact that only six coral taxa dominance remained unchanged, suggest that these corals may have developed a mechanism to cope with extreme seawater temperatures as evidenced by their continued growth and survival over the study period. We speculate that species dominance shift occurred largely as a result of a local oil spill rather than exposure to extreme temperatures. Further scrutiny of these species and the mechanisms by which they are able to thrive is recommended, as they hold the potential to benefit other coral communities as a resilient transplant species and model for understanding coral survivability in extreme environmental conditions.

How many donor colonies should be cross-fertilized for nursery farming of sexually propagated corals?

Coral reef restoration approaches have often utilized adult colonies as sources for fragments (i.e. clones) to be transplanted. Although restoration through this method is fast and cheap, it has been pointed out that it may reduce genetic diversity of the restored population. Low genetic diversity is a concern for reef restoration when seed fragments are raised asexually from only a few donor colonies. This can lead to lower fertilization rates among seed fragments, and reducing the longterm benefits of reef restoration in particular areas. Additionally, low genetic diversity can compound the effects of increased ocean temperature and other environmental stressors, further jeopardizing the health of a reef. An alternative approach through sexually propagated coral cultures and out-plantings can alleviate this problem. Sexually produced offsprings are more genetically diverse. They can be produced in far greater numbers than coral fragments and do not imply destructive methods. Ongoing research at the Akajima Marine Science Laboratory in Okinawa, Japan has helped to improve the production and maintenance of sexually propagated larval cultures. Our results show that crosses between gametes from 6 or more colonies will provide the highest fertilization rate (>95%). Based on the results, we suggest the use of 6 or more donor colonies for practical gamete fertilization in sexually derived coral culture.

Polyp-Canal Reconstruction Reveals Evolution Toward Complexity in Corals

Modern scleractinian corals are classified into robust,complex,and basal clades through comparative molecular studies.However,only few morphological or biological criteria can systematically determine the evolutionary trajectories of these major scleractinian coral clades.Here,we obtained the structural information of 21 scleractinian coral species representing robust and complex clades:High-resolution micro-computed tomography was used to reconstruct the polyp-canal systems in their colonies and to visualize the dynamic polyp growth processes.We found that the emergence of mesh-like canals may distinguish representatives of complex and robust clades.The differences in polyp-canal connections suggest distinct evolutionary trajectories among coral species:The formation of the canal network promoted the development of more complex coral structures,and coral polyps within this network formed calices of very similar volume,following precise axial growth directions.The influence of individual polyps on the coral colony becomes less significant as coral structures become more complex,and coral species with more complicated polyp-canal systems occupied niches more efficiently.This work supplements current evolutionary studies on reef-building corals,providing insight for further studies on coral growth patterns.

Coral records of Mid-Holocene sea-level highstands and climate responses in the northern South China Sea

High-resolution sea-level data and high-precision dating of corals in the northern South China Sea(SCS)during the Holocene provide a reference and historical background for current and future sea-level changes and a basis for scientific assessment of the evolutionary trend of coral reefs in the SCS.Although sporadic studies have been performed around Hainan Island in the northern SCS,the reconstructed sea level presents different values or is controversial because the indicative meaning of the sea-level indicators were neither quantified nor uniform criteria.Here,we determined the quantitative relationship between modern living coral and sea level by measuring the top surfaces of 27 live Porites corals from the inner reef flat along the east coast of Hainan Island and assessed the accuracy of results obtained using coral as sea-level indicators.Additionally,three in situ fossil Porites corals were analyzed based on elevation measurements,digital X-ray radiography,and U-Th dating.The survey results showed that the indicative meanings for the modern live Porites corals is(146.09±8.35)cm below the mean tide level(MTL).It suggested that their upward growth limit is constrained by the sea level,and the lowest low water is the highest level of survival for the modern live Porites corals.Based on the newly defined indicative meanings,6 new sea-level index points(SLIPs)were obtained and 19 published SLIPs were recalculated.Those SLIPs indicated a relative sea level fluctuation between(227.7±9.8)cm to(154.88±9.8)cm MTL between(5393±25)cal a BP and(3390±12)cal a BP,providing evidences of the Mid-Holocene sea-level highstand in the northern SCS.Besides that,our analysis demonstrated that different sea-level histories may be produced based on different indicative meanings or criteria.The dataset of 276 coral U-Th ages indicates that coral reef development in the northern SCS comprised the initial development,boom growth,decline,and flourishing development again.A comparison with regional records indicated that synergistic effects of climatic and environmental factors were involved in the development of coral reefs in the northern SCS.Thus,the cessation of coral reef development during the Holocene in the northern SCS was probably associated with the dry and cold climate in South China,as reflected in the synchronous weakening of the ENSO and East Asian summer monsoon induced by the reduction of the 65°N summer insolation,which forced the migration of the Intertropical Convergence Zone.

Liquefaction susceptibility and deformation characteristics of saturated coral sandy soils subjected to cyclic loadings-a critical review

Coral sandy soils widely exist in coral island reefs and seashores in tropical and subtropical regions.Due to the unique marine depositional environment of coral sandy soils,the engineering characteristics and responses of these soils subjected to monotonic and cyclic loadings have been a subject of intense interest among the geotechnical and earthquake engineering communities.This paper critically reviews the progress of experimental investigations on the undrained behavior of coral sandy soils under monotonic and cyclic loadings over the last three decades.The focus of coverage includes the contractive-dilative behavior,the pattern of excess pore-water pressure(EPWP)generation and the liquefaction mechanism and liquefaction resistance,the small-strain shear modulus and strain-dependent shear modulus and damping,the cyclic softening feature,and the anisotropic characteristics of undrained responses of saturated coral sandy soils.In particular,the advances made in the past decades are reviewed from the following aspects:(1)the characterization of factors that impact the mechanism and patterns of EPWP build-up;(2)the identification of liquefaction triggering in terms of the apparent viscosity and the average flow coefficient;(3)the establishment of the invariable form of strain-based,stress-based,or energy-based EPWP ratio formulas and the unique relationship between the new proxy of liquefaction resistance and the number of cycles required to reach liquefaction;(4)the establishment of the invariable form of the predictive formulas of small strain modulus and strain-dependent shear modulus;and(5)the investigation on the effects of stress-induced anisotropy on liquefaction susceptibility and dynamic deformation characteristics.Insights gained through the critical review of these advances in the past decades offer a perspective for future research to further resolve the fundamental issues concerning the liquefaction mechanism and responses of coral sandy sites subjected to cyclic loadings associated with seismic events in marine environments.

High-precision U-series dating of Holocene corals from South China Sea by thermal ionization mass spectrometry(TIMS)

Using the isotopic diluting and thermal ionization mass spectrometric (TIMS) technique, the U-series dating procedure was established on the basis of measurement of national standard GBW04412. With this highly precise technique the Holocene corals from the South China Sea were dated. Comparison between this method and α-spectrometry was made. The results show that a thriving period of corals in the South China Sea appeared at 6 800—5 800 a BP, during which the paleosea level was the higher than at present in the Holecene.

Spatial variations in the trophic status of Favia palauensis corals in the South China Sea:Insights into their different adaptabilities under contrasting environmental conditions

Scleractinian corals exhibit a dual trophic pattern of autotrophic photosynthesis and heterotrophic predation.However,whether corals can adjust their trophic status under contrasting environmental conditions remains unclear.In our study,70 scleractinian corals(Favia palauensis)were collected from Sanya and the Xisha and Nansha Islands in the South China Sea.We measured the zooxanthellae density(ZD)andδ^(13)C of zooxanthellae(δ^(13)C_(z))and host tissue(δ^(13)C_(h))and analyzed the difference betweenδ^(13)C_(h) andδ^(13)C_(z)(i.e.,△^(h-z)^(13)C).The relatively high ZD andδ^(13)C_(z) values in the samples from Sanya indicate that these corals might have higher photosynthetic rates and autotrophic abilities than those from Xisha and Nansha.In contrast,the relatively lowδ^(13)C_(h) and△^(h-z)^(13)C values in the samples from Xisha and Nansha suggest that these corals might have a higher heterotrophic ability than those from Sanya.In addition,we tested the coral tissue biomass and skeletalδ^(13)C(δ^(13)C_(s))in the samples from Sanya and examined their correlations with△^(h-z)^(13)C.The results showed a negative correlation,indicating that the more the organic material produced by the coral,the stronger its heterotrophic ability.Our results show that corals can adjust their trophic status under different environmental and physiological conditions,which is essential for increasing their adaptability to different environmental conditions.

Palaeobiogeographical constraints on the distribution of foraminifers and rugose corals in the Carboniferous Tindouf Basin,South Morocco

The northern flank of the Tindouf Syncline in southern Morocco exhibits a continuous, well exposed Carboniferous succession with limestones of Late Asbian to Early Bashkirian age containing rich and diverse foraminiferal and rugose coral assemblages. Analysis of these assemblages provides new data on the relatively poorly known Saharan basins. The palaeobiogeographical relationship of the Tindouf Basin with other Palaeotethyan basins is complex. Although there is a predominance of cosmopolitan taxa for the Palaeotethys, it is recognized that there was an influence of basins from NW Europe, such as the UK and Ireland. Some taxa are recorded in both NW Europe and Tindouf without any characteristic contributions from intermediate basins in northern Morocco. The neighbouring Béchar Basin in Algeria presents distinct assemblages. The bulk of the data analyzed suggest that this sector of the western Palaeotethys can be subdivided into four palaeobiogeographical subprovinces: the Atlantic Subprovince (UK, Ireland, N France and Belgium), the Mediterranean Subprovince (Pyrenees, Montagne Noire, Betic Cordillera, Rif (N Morocco) and Balearic Islands), and the Saharan Subprovince (Béchar, Reggan, Ahnet-Mouydir and Tindouf). In between, mobile belts of mixed faunal assemblage characteristics are observed (e.g., SW Spain and Central Meseta) forming the West peri-Gondwanan Subprovince.Analysis of the Tindouf Basin faunas shows that, as in other Saharan basins, there is a high diversity and abundance of foraminiferal taxa, with a higher proportion of survivors and longer stratigraphic ranges; these features also are mirrored by rugose corals. This emphasizes the longevity of the carbonate platform in a tropical setting, where periodic transgressions introduced new assemblages, and oceanic currents are interpreted as one of the main controlling factors for the distribution of the taxa in these subprovinces. Moreover, not only were water temperatures on the platform higher, but also tectonic stability greater. It is considered that the effects of the first phases of the Gond-wanan glaciation were minimal on the Tindouf faunas.

Different mechanisms underlie similar species-area relationships in two tropical archipelagoes

Despite much research in the field of island biogeography,mechanisms regulating insular diversity remain elusive.Here,we aim to explore mechanisms underlying plant species-area relationships in two tropical archipelagoes in the South China Sea.We found positive plant species-area relationships for both coral and continental archipelagoes.However,our results showed that different mechanisms contributed to similar plant species-area relationships between the two archipelagoes.For coral islands,soil nutrients and spatial distance among communities played major roles in shaping plant community structure and species diversity.By contrast,the direct effect of island area,and to a lesser extent,soil nutrients determined plant species richness on continental islands.Intriguingly,increasing soil nutrients availability(N,P,K)had opposite effects on plant diversity between the two archipelagoes.In summary,the habitat quality effect and dispersal limitation are important for regulating plant diversity on coral islands,whereas the passive sampling effect,and to a lesser extent,the habitat quality effect are important for regulating plant diversity on continental islands.More generally,our findings indicate that island plant species-area relationships are outcomes of the interplay of both niche and neutral processes,but the driving mechanisms behind these relationships depends on the type of islands.