运用PCR-RFLP方法研究三亚鹿回头岸礁造礁石珊瑚共生藻的组成

造礁石珊瑚共生藻的系统分类研究对于理解珊瑚礁生态系统对全球变化的响应具有十分重要的意义.本研究利用PCR技术扩增核糖体基因人亚基片段以及限制性片段长度多态性(RFLP)的方法,对海南三亚鹿回头岸礁的8科14属22种造礁石珊瑚的共生藻组成进行了研究.结果表明鹿回头岸礁造礁石珊瑚共生藻以C系群为优势系群,偶尔发现D系群与鹿角杯形珊瑚(Pocilolpora damieornis)和黄癣蜂巢珊瑚(Favia favus)共生:另外丑鹿角珊瑚(Acropora horrida)和丛生盔形珊瑚(Galaxea fascicularis)可以同时与C系群和D系群共生.共生多型性的发现暗示珊瑚与共生藻的共生关系具有灵活性.研究结果同样显示共生藻的核糖体基因人片段的DNA多态性偏低.未来应该结合其他的分子标记对鹿回头岸礁造礁石珊瑚共生藻的DNA多态性进行更深入的研究.     

造礁石珊瑚对低温的耐受能力及响应模式

通过实验室生态模拟,研究了低温胁迫下三亚湾5种造礁石珊瑚（十字牡丹珊瑚、佳丽鹿角珊瑚、花鹿角珊瑚、强壮鹿角珊瑚、澄黄滨珊瑚）的耐受性,分析了造礁石珊瑚对低温的响应模式.结果表明：造礁石珊瑚耐受低温能力与其骨骼类型有关,枝状珊瑚最先死亡,块状珊瑚的耐受能力明显高于枝状珊瑚;14 ℃持续3 d是三亚湾枝状造礁石珊瑚的致死低温;14 ℃持续3 d为块状澄黄滨珊瑚的致白化低温;12 ℃持续10 d为叶片状十字牡丹珊瑚的致死温度;块状澄黄滨珊瑚受到低温胁迫时表面形成粘膜,阻止了珊瑚进一步排出共生虫黄藻. 耐高温的珊瑚对低温也表现出较强的耐受能力,珊瑚对低温胁迫的响应模式与对高温的响应模式基本一致, 即珊瑚首先不伸展触手,紧接着不断释放粘液并排出共生藻,最后白化、死亡.     

西沙群岛海域造礁石珊瑚物种多样性与分布特点

2006年3-5月,采用国际通用的截线样带法对海南省西沙群岛主要珊瑚礁海域的18个岛礁造礁石珊瑚进行了实地调查,共记录13科45属188种,其中62个新记录种.整合文献资料,该海域共记录造礁石珊瑚13科50属204种,基本包括分布在我国(不含台湾省)的所有造礁石珊瑚科属和绝大多数种类.造礁石珊瑚以生长速度快的分枝状珊瑚为主.物种多样性最高的是华光礁海域、金银岛海域和永兴岛海域.聚类分析可以把18个调查岛礁分成3个类群:有人类居住的大型环礁群、独立岛礁群和无人居住的大型环礁群(华光礁).为了有效保护珍稀的造礁石珊瑚资源,按照其种类多样性和分布特点重点保护关键海域是十分必要的.     

珊瑚及共生藻在白化过程中的适应机制研究进展

珊瑚礁生态系统具有非常重要的生态学功能。但是随着全球气候变暖和CO2浓度的升高,珊瑚白化事件越来越频繁,珊瑚礁生态系统面临严重的危机。影响珊瑚白化的重要因子主要有海水温度的异常（过高或过低）,太阳辐射与紫外线辐射,海水盐度的偏离,珊瑚疾病,海洋污染,长棘海星的爆发,人类的过度捕鱼和全球CO2浓度升高等。其中,海洋表面水体温度（SST）的异常升高为珊瑚白化的主要因素。珊瑚主要是通过珊瑚与共生藻的生理适应机制以及更换共生藻基因型机制两种方式来适应环境胁迫的。生理适应机制主要通过叶黄素循环、珊瑚色素荧光（热）、活性氧清除系统（自由基）、分泌紫外线吸收物质MAAs（紫外光）、产生热休克蛋白HspS（热）来实现的。珊瑚共生藻基因型更换适应机制是指珊瑚的适应性白化假说。珊瑚的适应性白化假说还有很多争议,还需要更多的实验证据提供支持。未来的研究重点将在珊瑚白化过程中共生藻-珊瑚共生功能体作为整体性的研究,尤其是珊瑚宿主在白化过程中对共生功能体作出贡献的研究。     

悬浮物对造礁石珊瑚营养方式的影响及其适应性研究进展

营养方式是造礁石珊瑚获取能量与营养物质的基础,影响其生长与分布。近年来珊瑚礁区悬浮物含量与组分结构发生显著变化,其对造礁石珊瑚营养方式的诸多影响正成为当前研究热点。研究系统综述了珊瑚礁区悬浮物变化特征、悬浮物对造礁石珊瑚营养方式的影响及其适应性研究现状。发现近年来人类活动加剧与强降雨事件频发是驱动珊瑚礁,尤其是近岸珊瑚礁区悬浮物含量递增、组分改变与变频加剧的主因;悬浮物变化对造礁石珊瑚光合自养与异养营养的影响存在显著的种间差异,这主要与悬浮物消光效应、生物可利用性及造礁石珊瑚种类密切相关。虽然少数种类造礁石珊瑚具光合可塑性或异养可塑性,能在高含量悬浮物存在的弱光环境中较好生长。然而对绝大多数造礁石珊瑚而言,其营养方式适应性较差,无法在悬浮物影响下较好地获取生命活动所需的能量与营养物质,进而难以生存。总体来说,悬浮物被认为是近年来影响造礁石珊瑚生长与分布的重要环境因子之一,而关于造礁石珊瑚营养方式对悬浮物变化的响应及其适应机制,当前研究仍较薄弱,需要进一步加强相关研究。     

造礁石珊瑚记录现代海洋生态环境信息研究进展

造礁石珊瑚因其独特的生物学和生态学特性,已被广泛用作环境因子的代用指标.过去国内外的研究主要侧重于地质年代环境信息的记录,随着样品分析测试精度的提高,用造礁石珊瑚骨骼研究现代热带的生态环境也逐渐增多,尤其对于影响海洋生态环境的事件信息的记录.许多研究都证明了造礁石珊瑚骨骼中各种示踪剂在监测过去环境变化方面的有效性,对目前和将来的生态监控以及环境研究都具有广阔的前景.目前,珊瑚基础生理学的机制尚未完全探明,进一步了解珊瑚微观结构层次的生理机制是该领域研究的关键所在.我国学者已开展了南海西沙、南沙、台湾岛、海南岛、徐闻和涠洲岛等地区的造礁石珊瑚骨骼多种指标的研究.     

海洋酸化对珊瑚礁生态系统的影响研究进展

目前,大气CO2浓度的升高已导致海水pH值比工业革命前下降了约0.1,海水碳酸盐平衡体系随之变化,进而影响珊瑚礁生态系统的健康。近年来的研究表明海洋酸化导致造礁石珊瑚幼体补充和群落恢复更加困难,造礁石珊瑚和其它造礁生物(Reef-building organisms)钙化率降低甚至溶解,乃至影响珊瑚礁鱼类的生命活动。虽然海洋酸化对造礁石珊瑚光合作用的影响不显著,但珊瑚-虫黄藻共生体系会受到一定影响。建议选择典型海区进行长期系统监测,结合室内与原位模拟试验,从个体、种群、群落到系统不同层面,运用生理学和分子生物学技术,结合生态学研究手段,综合研究珊瑚的相应响应,以期深入认识海洋酸化对珊瑚礁生态系统健康(例如珊瑚白化)的影响及其效应。     

涠洲岛造礁石珊瑚群落变化特征及其环境影响因子

以涠洲岛造礁石珊瑚群落为研究对象,分析其群落物种组成、多样性、Raunkiaer频度和种间Spearman轶相关,探讨群落组成与水环境因子的相关性,并结合近10年来涠洲岛造礁石珊瑚变化情况,找出涠洲岛造礁石珊瑚群落的主导影响因素。结果表明：（1）近10年来涠洲岛造礁石珊瑚覆盖率显著降低,珊瑚形态组成趋于块状化,尽管珊瑚群落多样性较高,但群落分布较松散,群落结构较不稳定,部分优势种种间竞争较激烈。（2）悬浮物含量是影响石珊瑚群落最显著的环境因子。石珊瑚优势种群在不同水深中分布差异显著,泥沙覆盖率、营养盐对不同石珊瑚种群影响差异较大,大型海藻覆盖率在局部区域对优势珊瑚形成较强的竞争关系。（3）营养盐和泥沙沉积物的增加与涠洲岛近海养殖业及生活排污、海岸工程及海岸侵蚀密切相关。     

中国造礁石珊瑚分类厘定

造礁石珊瑚是珊瑚礁框架建造者, 具有维持珊瑚礁生态系统功能和稳定性的重要作用, 其分类对于造礁石珊瑚和珊瑚礁的研究与保护至关重要。目前, 随着分子系统学的不断发展, 造礁石珊瑚的分类体系发生改变, 伴随着出现大量同物异名。近年来也出现许多无中文学名的中国造礁石珊瑚新记录种, 这些都给物种认定和命名带来困难, 阻碍了中国造礁石珊瑚的研究与保护工作。为此, 本文收集了中国造礁石珊瑚物种记录文献资料, 采用最新的造礁石珊瑚分类体系, 确认同物异名, 形成中国造礁石珊瑚物种名录, 并对中国造礁石珊瑚物种的中文名进行统一的规范和命名。结果表明, 中国共有造礁石珊瑚2个类群16科77属445种。与《中国动物志·腔肠动物门·珊瑚虫纲·石珊瑚目·造礁石珊瑚》相比, 科级分类阶元新增7个科, 变更5个科; 属级分类阶元新增26属, 变更1属, 合并3属; 种级分类阶元新增291种, 变更13种, 合并20种, 新命名305个物种的中文名。并且筛选出187个同物异名。此外, 由于造礁石珊瑚分类体系现仍有部分争议, 文章也进行了讨论说明。     

珊瑚礁的建造者

珊瑚礁是一种独特的海洋生态资源,对于调节和优化热带海洋环境具有重要意义。目前世界上的海洋被珊瑚礁覆盖的大约有200万平方公里,为全球1/4的海洋生物提供美好的家园。珊瑚礁的主要建造者是一群称为造礁石珊瑚的原始多细胞"后生动物"——珊瑚虫。现存大约有1300种,我国已记录的有174种。大部分造礁石珊瑚以群体方式生活在一起,借助     

Resistance to thermal stress in corals without changes in symbiont composition

Discovering how corals can adjust their thermal sensitivity in the context of global climate change is important in understanding the long-term persistence of coral reefs. In this study, we showed that short-term preconditioning to higher temperatures, 3°C below the experimentally determined bleaching threshold, for a period of 10 days provides thermal tolerance for the symbiosis stability between the scleractinian coral, Acropora millepora and Symbiodinium. Based on genotypic analysis, our results indicate that the acclimatization of this coral species to thermal stress does not come down to simple changes in Symbiodinium and/or the bacterial communities that associate with reef-building corals. This suggests that the physiological plasticity of the host and/or symbiotic components appears to play an important role in responding to ocean warming. The further study of host and symbiont physiology, both of Symbiodinium and prokaryotes, is of paramount importance in the context of global climate change, as mechanisms for rapid holobiont acclimatization will become increasingly important to the long-standing persistence of coral reefs.     

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.     

Specificity is rarely absolute in coral-algal symbiosis: implications for coral response to climate change

Some reef-building corals have been shown to respond to environmental change by shifting the composition of their algal symbiont (genus Symbiodinium) communities. These shifts have been proposed as a potential mechanism by which corals might survive climate stressors, such as increased temperatures. Conventional molecular methods suggest this adaptive capacity may not be widespread because few (～25%) coral species have been found to associate with multiple Symbiodinium clades. However, these methods can fail to detect low abundance symbionts (typically less than 10-20% of the total algal symbiont community). To determine whether additional Symbiodinium clades are present, but are not detected using conventional techniques, we applied a high-resolution, real-time PCR assay to survey Symbiodinium (in clades A-D) from 39 species of phylogenetically and geographically diverse scleractinian corals. This survey included 26 coral species thought to be restricted to hosting a single Symbiodinium clade ('symbiotic specialists'). We detected at least two Symbiodinium clades (C and D) in at least one sample of all 39 coral species tested; all four Symbiodinium clades were detected in over half (54%) of the 26 symbiotic specialist coral species. Furthermore, on average, 68 per cent of all sampled colonies within a given coral species hosted two or more symbiont clades. We conclude that the ability to associate with multiple symbiont clades is common in scleractinian (stony) corals, and that, in coral-algal symbiosis, 'specificity' and 'flexibility' are relative terms: specificity is rarely absolute. The potential for reef corals to adapt or acclimatize to environmental change via symbiont community shifts may therefore be more phylogenetically widespread than has previously been assumed.     

Variation in Symbiodinium ITS2 sequence assemblages among coral colonies

Endosymbiotic dinoflagellates in the genus Symbiodinium are fundamentally important to the biology of scleractinian corals, as well as to a variety of other marine organisms. The genus Symbiodinium is genetically and functionally diverse and the taxonomic nature of the union between Symbiodinium and corals is implicated as a key trait determining the environmental tolerance of the symbiosis. Surprisingly, the question of how Symbiodinium diversity partitions within a species across spatial scales of meters to kilometers has received little attention, but is important to understanding the intrinsic biological scope of a given coral population and adaptations to the local environment. Here we address this gap by describing the Symbiodinium ITS2 sequence assemblages recovered from colonies of the reef building coral Montipora capitata sampled across Kāne'ohe Bay, Hawai'i. A total of 52 corals were sampled in a nested design of Coral Colony(Site(Region)) reflecting spatial scales of meters to kilometers. A diversity of Symbiodinium ITS2 sequences was recovered with the majority of variance partitioning at the level of the Coral Colony. To confirm this result, the Symbiodinium ITS2 sequence diversity in six M. capitata colonies were analyzed in much greater depth with 35 to 55 clones per colony. The ITS2 sequences and quantitative composition recovered from these colonies varied significantly, indicating that each coral hosted a different assemblage of Symbiodinium. The diversity of Symbiodinium ITS2 sequence assemblages retrieved from individual colonies of M. capitata here highlights the problems inherent in interpreting multi-copy and intra-genomically variable molecular markers, and serves as a context for discussing the utility and biological relevance of assigning species names based on Symbiodinium ITS2 genotyping.     

Azooxanthellate? Most Hawaiian black corals contain Symbiodinium

The ecological success of shallow-water reef-building corals (Hexacorallia: Scleractinia) is framed by their intimate endosymbiosis with photosynthetic dinoflagellates in the genus Symbiodinium (zooxanthellae). In contrast, the closely related black corals (Hexacorallia: Anthipatharia) are described as azooxanthellate (lacking Symbiodinium), a trait thought to reflect their preference for low-light environments that do not support photosynthesis. We examined 14 antipatharian species collected between 10 and 396 m from Hawai'i and Johnston Atoll for the presence of Symbiodinium using molecular typing and histology. Symbiodinium internal transcribed spacer-2 (ITS-2) region sequences were retrieved from 43 per cent of the antipatharian samples and 71 per cent of the examined species, and across the entire depth range. The ITS-2 sequences were identical or very similar to those commonly found in shallow-water scleractinian corals throughout the Pacific. Histological analyses revealed low densities of Symbiodinium cells inside antipatharian gastrodermal tissues (0-92 cells mm(-3)), suggesting that the Symbiodinium are endosymbiotic. These findings confirm that the capacity to engage in endosymbiosis with Symbiodinium is evolutionarily conserved across the cnidarian subclass Hexacorallia, and that antipatharians associate with Symbiodinium types found in shallow-water scleractinians. This study represents the deepest record for Symbiodinium to date, and suggests that some members of this dinoflagellate genus have extremely diverse habitat preferences and broad environmental ranges.     

The Occurrence of Mixed Infections of Symbiodinium (Dinoflagellata) within Individual Hosts

Coral reef ecosystems depend on symbiosis between dinoflagellates of the genus Symbiodinium Freudenthal and their various hosts. The physiological characteristics associated with a particular lineage or species of Symbiodinium can determine a host's susceptibility to harmful bleaching. Therefore, the threat posed by global climate change on a host may be reduced if it can switch or shuffle its dominant algal symbiont type. An important prerequisite to this potential to switch or shuffle is the ability to host multiple alternative dominant symbiont genotypes. To examine the distribution of this trait, we review reports of mixed Symbiodinium infections in corals and nonscleractinian hosts from a phylogenetic perspective. Hosts showing evidence of mixed infection are broadly distributed across the most deeply divergent host lineages, including foraminifera, mollusks, sponges, and cnidarians. The occurrence of mixed infections is also broadly distributed across most clades of scleractinian corals. Individual colonies of certain well‐studied cosmopolitan coral genera, such as Acropora, Montastraea, and Pocillopora, yield many reports of mixed infection, while other genera, such as Porites, do not. We further discuss mixed Symbiodinium infections in the context of evolutionary ecology theory. Selection pressures that affect the prevalence of mixed infection may be exerted by variation in host environment, host ontogeny, symbiont transmission strategy, host regulation of symbiont populations, availability of free‐living symbiont lineages, competition between symbiont lineages, and niche partitioning of the internal host environment.     

Diversity of algal endosymbionts (zooxanthellae) in octocorals: the roles of geography and host relationships

The presence, genetic identity and diversity of algal endosymbionts (Symbiodinium) in 114 species from 69 genera (20 families) of octocorals from the Great Barrier Reef (GBR), the far eastern Pacific (EP) and the Caribbean was examined, and patterns of the octocoral-algal symbiosis were compared with patterns in the host phylogeny. Genetic analyses of the zooxanthellae were based on ribosomal DNA internal transcribed spacer 1 (ITS1) region. In the GBR samples, Symbiodinium clades A and G were encountered with A and G being rare. Clade B zooxanthellae have been previously reported from a GBR octocoral, but are also rare in octocorals from this region. Symbiodinium G has so far only been found in Foraminifera, but is rare in these organisms. In the Caribbean samples, only Symbiodinium clades B and C are present. Hence, Symbiodinium diversity at the level of phylogenetic clades is lower in octocorals from the Caribbean compared to those from the GBR. However, an unprecedented level of ITS1 diversity was observed within individual colonies of some Caribbean gorgonians, implying either that these simultaneously harbour multiple strains of clade B zooxanthellae, or that ITS1 heterogeneity exists within the genomes of some zooxanthellae. Intracladal diversity based on ITS should therefore be interpreted with caution, especially in cases where no independent evidence exists to support distinctiveness, such as ecological distribution or physiological characteristics. All samples from EP are azooxanthellate. Three unrelated GBR taxa that are described in the literature as azooxanthellate (Junceella fragilis, Euplexaura nuttingi and Stereonephthya sp. 1) contain clade G zooxanthellae, and their symbiotic association with zooxanthellae was confirmed by histology. These corals are pale in colour, whereas related azooxanthellate species are brightly coloured. The evolutionary loss or gain of zooxanthellae may have altered the light sensitivity of the host tissues, requiring the animals to adopt or reduce pigmentation. Finally, we superimposed patterns of the octocoral-algal symbiosis onto a molecular phylogeny of the host. The data show that many losses/gains of endosymbiosis have occurred during the evolution of octocorals. The ancestral state (azooxanthellate or zooxanthellate) in octocorals remains unclear, but the data suggest that on an evolutionary timescale octocorals can switch more easily between mixotrophy and heterotrophy compared to scleractinian corals, which coincides with a low reliance on photosynthetic carbon gain in the former group of organisms.     

Symbiodiniaceae-bacteria interactions: rethinking metabolite exchange in reef-building corals as multi-partner metabolic networks

The intimate relationship between scleractinian corals and their associated microorganisms is fundamental to healthy coral reef ecosystems. Coral-associated microbes (Symbiodiniaceae and other protists, bacteria, archaea, fungi and viruses) support coral health and resilience through metabolite transfer, inter-partner signalling, and genetic exchange. However, much of our understanding of the coral holobiont relationship has come from studies that have investigated either coral-Symbiodiniaceae or coral-bacteria interactions in isolation, while relatively little research has focused on other ecological and metabolic interactions potentially occurring within the coral multi-partner symbiotic network. Recent evidences of intimate coupling between phytoplankton and bacteria have demonstrated that obligate resource exchange between partners fundamentally drives their ecological success. Here, we posit that similar associations with bacterial consortia regulate Symbiodiniaceae productivity and are in turn central to the health of corals. Indeed, we propose that this bacteria-Symbiodiniaceae-coral relationship underpins the coral holobiont's nutrition, stress tolerance and potentially influences the future survival of coral reef ecosystems under changing environmental conditions. Resolving Symbiodiniaceae-bacteria associations is therefore a logical next step towards understanding the complex multi-partner interactions occurring in the coral holobiont.     

The Roles and Interactions of Symbiont,Host and Environment in Defining Coral Fitness

Background

Reef-building corals live in symbiosis with a diverse range of dinoflagellate algae (genus Symbiodinium) that differentially influence the fitness of the coral holobiont. The comparative role of symbiont type in holobiont fitness in relation to host genotype or the environment, however, is largely unknown. We addressed this knowledge gap by manipulating host-symbiont combinations and comparing growth, survival and thermal tolerance among the resultant holobionts in different environments.

Methodology/Principal Findings

Offspring of the coral, Acropora millepora, from two thermally contrasting locations, were experimentally infected with one of six Symbiodinium types, which spanned three phylogenetic clades (A, C and D), and then outplanted to the two parental field locations (central and southern inshore Great Barrier Reef, Australia). Growth and survival of juvenile corals were monitored for 31–35 weeks, after which their thermo-tolerance was experimentally assessed. Our results showed that: (1) Symbiodinium type was the most important predictor of holobiont fitness, as measured by growth, survival, and thermo-tolerance; (2) growth and survival, but not heat-tolerance, were also affected by local environmental conditions; and (3) host population had little to no effect on holobiont fitness. Furthermore, coral-algal associations were established with symbiont types belonging to clades A, C and D, but three out of four symbiont types belonging to clade C failed to establish a symbiosis. Associations with clade A had the lowest fitness and were unstable in the field. Lastly, Symbiodinium types C1 and D were found to be relatively thermo-tolerant, with type D conferring the highest tolerance in A. millepora.

Conclusions/Significance

These results highlight the complex interactions that occur between the coral host, the algal symbiont, and the environment to shape the fitness of the coral holobiont. An improved understanding of the factors affecting coral holobiont fitness will assist in predicting the responses of corals to global climate change.     

Conservation genetics and the resilience of reef-building corals

Coral reefs have suffered long-term decline due to a range of anthropogenic disturbances and are now also under threat from climate change. For appropriate management of these vulnerable and valuable ecosystems it is important to understand the factors and processes that determine their resilience and that of the organisms inhabiting them, as well as those that have led to existing patterns of coral reef biodiversity. The scleractinian (stony) corals deposit the structural framework that supports and promotes the maintenance of biological diversity and complexity of coral reefs, and as such, are major components of these ecosystems. The success of reef-building corals is related to their obligate symbiotic association with dinoflagellates of the genus Symbiodinium. These one-celled algal symbionts (zooxanthellae) live in the endodermal tissues of their coral host, provide most of the host's energy budget and promote rapid calcification. Furthermore, zooxanthellae are the main primary producers on coral reefs due to the oligotrophic nature of the surrounding waters. In this review paper, we summarize and critically evaluate studies that have employed genetics and/or molecular biology in examining questions relating to the evolution and ecology of reef-building corals and their algal endosymbionts, and that bear relevance to coral reef conservation. We discuss how these studies can focus future efforts, and examine how these approaches enhance our understanding of the resilience of reef-building corals.