Out of Their Depth? Isolated Deep Populations of the Cosmopolitan Coral Desmophyllum dianthus May Be Highly Vulnerable to Environmental Change

Deep sea scleractinian corals will be particularly vulnerable to the effects of climate change, facing loss of up to 70% of their habitat as the Aragonite Saturation Horizon (below which corals are unable to form calcium carbonate skeletons) rises. Persistence of deep sea scleractinian corals will therefore rely on the ability of larvae to disperse to, and colonise, suitable shallow-water habitat. We used DNA sequence data of the internal transcribed spacer (ITS), the mitochondrial ribosomal subunit (16S) and mitochondrial control region (MtC) to determine levels of gene flow both within and among populations of the deep sea coral Desmophyllum dianthus in SE Australia, New Zealand and Chile to assess the ability of corals to disperse into different regions and habitats. We found significant genetic subdivision among the three widely separated geographic regions consistent with isolation and limited contemporary gene flow. Furthermore, corals from different depth strata (shallow <600 m, mid 1000-1500 m, deep >1500 m) even on the same or nearby seamounts were strongly differentiated, indicating limited vertical larval dispersal. Genetic differentiation with depth is consistent with the stratification of the Subantarctic Mode Water, Antarctic Intermediate Water, the Circumpolar Deep and North Pacific Deep Waters in the Southern Ocean, and we propose that coral larvae will be retained within, and rarely migrate among, these water masses. The apparent absence of vertical larval dispersal suggests deep populations of D. dianthus are unlikely to colonise shallow water as the aragonite saturation horizon rises and deep waters become uninhabitable. Similarly, assumptions that deep populations will act as refuges for shallow populations that are impacted by activities such as fishing or mining are also unlikely to hold true. Clearly future environmental management strategies must consider both regional and depth-related isolation of deep-sea coral populations.     

Transcriptome‐based target‐enrichment baits for stony corals (Cnidaria: Anthozoa: Scleractinia)

Despite the ecological and economic significance of stony corals (Scleractinia), a robust understanding of their phylogeny remains elusive due to patchy taxonomic and genetic sampling, as well as the limited availability of informative markers. To increase the number of genetic loci available for phylogenomic analyses in Scleractinia, we designed 15,919 DNA enrichment baits targeting 605 orthogroups (mean 565 ± SD 366 bp) over 1,139 exon regions. A further 236 and 62 barcoding baits were designed for COI and histone H3 genes respectively for quality and contamination checks. Hybrid capture using these baits was performed on 18 coral species spanning the presently understood scleractinian phylogeny, with two corallimorpharians as outgroup. On average, 74% of all loci targeted were successfully captured for each species. Barcoding baits were matched unambiguously to their respective samples and revealed low levels of cross‐contamination in accordance with expectation. We put the data through a series of stringent filtering steps to ensure only scleractinian and phylogenetically informative loci were retained, and the final probe set comprised 13,479 baits, targeting 452 loci (mean 531 ± SD 307 bp) across 865 exon regions. Maximum likelihood, Bayesian and species tree analyses recovered maximally supported, topologically congruent trees consistent with previous phylogenomic reconstructions. The phylogenomic method presented here allows for consistent capture of orthologous loci among divergent coral taxa, facilitating the pooling of data from different studies and increasing the phylogenetic sampling of scleractinians in the future.     

Three-dimensional species distribution modelling reveals the realized spatial niche for coral recruitment on contemporary Caribbean reefs

The three-dimensional structure of habitats is a critical component of species' niches driving coexistence in species-rich ecosystems. However, its influence on structuring and partitioning recruitment niches has not been widely addressed. We developed a new method to combine species distribution modelling and structure from motion, and characterized three-dimensional recruitment niches of two ecosystem engineers on Caribbean coral reefs, scleractinian corals and gorgonians. Fine-scale roughness was the most important predictor of suitable habitat for both taxa, and their niches largely overlapped, primarily due to scleractinians' broader niche breadth. Crevices and holes at mm scales on calcareous rock with low coral cover were more suitable for octocorals than for scleractinian recruits, suggesting that the decline in scleractinian corals is facilitating the recruitment of octocorals on contemporary Caribbean reefs. However, the relative abundances of the taxa were independent of the amount of suitable habitat on the reef, emphasizing that niche processes alone do not predict recruitment rates.     

More evidence for pervasive paraphyly in scleractinian corals: Systematic study of Southeast Asian Faviidae (Cnidaria; Scleractinia) based on molecular and morphological data

Coral taxonomy and systematics continue to be plagued by a host of problems. Due to high phenotypic variability within species, morphological approaches have often failed to recognize natural taxa, and molecular techniques have yet to be applied to many groups. Here, we summarize the levels of paraphyly found for scleractinian corals and test, based on new data, whether paraphyly is also a significant problem in Faviidae, the second-most speciose hermatypic scleractinian family. Using both DNA sequence and morphological data we find that, regardless of analysis technique (maximum parsimony, maximum likelihood and Bayesian likelihood), many conventional taxonomic groups are not monophyletic. Based on two mitochondrial markers (COI and a noncoding region) that we amplified for 81 samples representing 41 faviid species and 13 genera, five genera that are represented by more than one species are paraphyletic, as is the family Faviidae. The morphological characters currently used to identify these corals similarly fail to recover many genera. Furthermore, trees based on both data types are incongruent, and total evidence analysis does little to salvage conventional taxonomic groupings. Morphological convergence, phenotypic variability in response to the environment, and recent speciation are likely causes for these conflicts, which suggest that the present classification of corals is in need of a major overhaul. We propose more detailed studies of problematic faviid taxa using standardized morphological, mitochondrial, and nuclear genetic markers to facilitate combining of data.     

Isolation of microsatellite loci from the scleractinian corals,Montastraea cavernosa and Porites astreoides

The ability to assess genetic variation is critical for determining genetic diversity and population structure. In corals, slow evolutionary rates in mitochondrial genomes have left allozymes as the only markers presently available to investigate patterns of intraspecific genetic variation. Characteristics of microsatellites render them more informative than allozymes for such analyses; however, few coral microsatellites are available. This study describes polymorphic microsatellite loci isolated from two scleractinian coral species. Most loci exhibit significant heterozygote deficiencies, likely due to nonrandom mating or Wahlund effects. These markers are being used to investigate gene flow among populations, providing insight into reef connectivity.     

Spatial study of juvenile corals in the Northern region of the Mesoamerican Barrier Reef System (MBRS)

Juvenile coral abundance and community composition depend on the spatial scale studied. To investigate this, an evaluation was made of juvenile coral density with hierarchical spatial analysis in the northern Mesoamerican Barrier Reef System (MBRS) at ~10 m depth. Study scope included semi-protected and unprotected areas located in this region. A total of 19 juvenile coral taxa were found, including 10 scleractinian species, 8 scleractinian coral genera not identified to species, and 1 Millepora species (Hydrozoa-Milleporidae). In terms of relative abundance, Agaricia spp., Siderastrea spp., and Porites spp. were the main juvenile taxa in the coral community at the surveyed sites, reefs, and regions levels. Greater variance was seen at smaller scales, at site level for taxa richness, and at the transect level for juvenile density, and lower variance was seen at larger scales (reefs and regions). The variance component contribution from each scale likely differed from other studies because of the different factors affecting the community and the different extensions of each scale used in each study. Densities (1–6.4 juvenile corals/m²) and dominant taxa found in this study agree with other studies from the Western Atlantic. Detected variability was explained by different causal agents, such as low grazing rates by herbivorous organisms, turbidity, and/or sediment suffocation and some nearby or distant localized disturbance (human settlement and a hurricane).     

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.     

Perspectives in coral reef hydrodynamics

Knowledge of skeletogenesis in scleractinian corals is central to reconstructing past ocean and climate histories, assessing and counteracting future climate and ocean acidification impacts upon coral reefs, and determining the taxonomy and evolutionary path of the Scleractinia. To better understand skeletogenesis and mineralogy in extant scleractinian corals, we have investigated the nature of the initial calcium carbonate skeleton deposited by newly settling coral recruits. Settling Acropora millepora larvae were sampled daily for 10 days from initial attachment, and the carbonate mineralogy of their newly deposited skeletons was investigated. Bulk analyses using Raman and infrared spectroscopic methods revealed that the skeletons were predominantly comprised of aragonite, with no evidence of calcite or an amorphous precursor phase, although presence of the latter cannot be discounted. Sensitive selected area electron diffraction analyses of sub-micron areas of skeletal regions further consolidated these data. These findings help to address the uncertainty surrounding reported differences in carbonate mineralogy between larval and adult extant coral skeletons by indicating that skeletons of new coral recruits share the same aragonitic mineralogy as those of their mature counterparts. In this respect, we can expect that skeletogenesis in both larval and mature growth stages of scleractinian corals will be similarly affected by ocean acidification and predicted environmental changes.     

Exon-primed, intron-crossing (EPIC) loci for five nuclear genes in deep-sea protobranch bivalves: primer design, PCR protocols and locus utility

We describe PCR primers and amplification protocols developed to obtain introns from conserved nuclear genes in deep-sea protobranch bivalves. Because almost no sequence data for protobranchs are publically available, mollusk and other protostome sequences from GenBank were used to design degenerate primers, making these loci potentially useful in other invertebrate taxa. Amplification and sequencing success varied across the test group of 30 species, and we present five loci spanning this range of outcomes. Intron presence in the targeted regions also varied across genes and species, often within single genera; for instance, the calmodulin and β-tubulin loci contained introns with high frequency, whereas the triose phosphate isomerase locus never contained an intron. In introns for which we were able to obtain preliminary estimates of polymorphism levels in single species, polymorphism was greater than traditional mitochondrial loci. These markers will greatly increase the ability to assess population structure in the ecologically important protobranchs, and may prove useful in other taxa as well.     

Patterns of scleractinian coral recruitment at Lord Howe Island,an isolated subtropical reef off eastern Australia

Changing oceanic conditions, particularly ocean warming and altered currents, can affect the reproductive success of corals. Improving the knowledge of coral reproductive processes at the marginal range limits of coral reefs is important for understanding the ecology of subtropical coral communities and the potential for coral species to expand their ranges in higher latitudes in the future. The extent of live coral cover around subtropical Lord Howe Island (LHI; 31°33′S, 159°05′E) approximately 600 km off the east coast of Australia, has been relatively stable over the last several decades; however, shifts in dominant species in the adult coral community have been reported. To examine the potential influences of recent altered currents and shifts in dominant scleractinian taxa within this community, this study examined spatial and seasonal variation of coral larval settlement at different habitats within the LHI reef lagoon. The study also assessed whether the assemblage of scleractinian corals settling at LHI has changed between 1990–1991 and 2011–2012. Mean densities of coral settlement in 2011–2012 (230 spat m^?2 yr^?1) were consistent with those reported in 1990–1991 and in other regions. However, changes in taxonomic composition were apparent with increases in the proportion of Acroporidae spat at some sites. Settlement of all taxa was highest over summer months, whereas during winter only one coral spat (Pocilloporidae) was detected. Coral settlement was highest and most taxonomically diverse at sites closest to the reef crest, where mortality of settled spat was also greatest. Rates of settlement were high compared with juvenile densities; hence, post-settlement mortality is also likely to be high. Post-settlement processes, influenced by local environmental conditions, are likely to be very important in structuring the adult coral communities within the LHI reef lagoon.     

Taxonomic status of Euphylliidae corals in Peninsular Malaysia based on morphological structures and phylogenetic analyses using mitochondrial COI gene

The morphological and molecular studies provide greater taxonomic resolution for the scleractinian coral identification. The Euphylliidae corals are among the scleractinian family for which their corallite and polyp morphologies have been examined for species identification. However, knowledge on the molecular study for coral identification in Malaysia is very limited. Therefore, this study aimed to examine the morphological structures and phylogenetic analyses for six Euphylliidae coral species using the mitochondrial gene of cytochrome oxidase subunit I (COI). The results showed that the Euphylliidae corals are present under both “complex” and “robust” coral clades as supported by many researchers. The result also revealed that the species phylogeny of Euphylliidae corals is in concordances with its morphological structures of corallites. It can be concluded the combination between morphological structures and phylogenetic analyses provide more accurate identification than relying on morphological study alone. Hence, it provides a future direction for the scleractinian research progress in species identification.     

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.     

Heterotrophy in Tropical Scleractinian Corals

The dual character of corals, that they are both auto- and heterotrophs, was recognized early in the twentieth Century. It is generally accepted that the symbiotic association between corals and their endosymbiotic algae (called zooxanthellae) is fundamental to the development of coral reefs in oligotrophic tropical oceans because zooxanthellae transfer the major part of their photosynthates to the coral host (autotrophic nutrition). However, numerous studies have confirmed that many species of corals are also active heterotrophs, ingesting organisms ranging from bacteria to mesozooplankton. Heterotrophy accounts for between 0 and 66% of the fixed carbon incorporated into coral skeletons and can meet from 15 to 35% of daily metabolic requirements in healthy corals and up to 100% in bleached corals. Apart from this carbon input, feeding is likely to be important to most scleractinian corals, since nitrogen, phosphorus, and other nutrients that cannot be supplied from photosynthesis by the coral's symbiotic algae must come from zooplankton capture, particulate matter or dissolved compounds. A recent study showed that during bleaching events some coral species, by increasing their feeding rates, are able to maintain and restore energy reserves.
This review assesses the importance and effects of heterotrophy in tropical scleractinian corals. We first provide background information on the different food sources (from dissolved organic matter to meso- and macrozooplankton). We then consider the nutritional inputs of feeding. Finally, we review feeding effects on the different physiological parameters of corals (tissue composition, photosynthesis and skeletal growth).     

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。     

Competitive strategies of soft corals (Coelenterata: Octocorallia). II. Variable defensive responses and susceptibility to scleractinian corals

Interactions involving competition for space between several species of alcyonacean and scleractinian corals were assessed experimentally on Britomart Reef, central region of the Great Barrier Reef, Australia. Colonies of three soft coral species, Sarcophyton ehrenbergi Marenzeller, Nephthea brassica Kukenthal, and Capnella lacertiliensis Macfayden Forskal (Coelenterata:Alcyonacea) were relocated within stands of two scleractinian corals, Parités andrewsi Vaughan (= P. cylindrica Dana) and Pavona cactus Förskal (Coelenterata:Scleractinia). Undisturbed scleractinian and relocated alcyonacean controls were also monitored.Alcyonacean corals induced necrosis of tissue in scleractinian corals. Necrosis was significantly more pronounced when colonies were in contact but was also observed in the absence of contact, implicating the presence of active allelopathic agents. Scleractinian coral species varied in their susceptibility to the ill effects of alcyonaceans, with Pontes andrewsi being more susceptible than Pavona cactus. Of the soft corals, Nephthea caused the highest degree of mortality in the two scleractinian corals examined and Sarcophyton the least. Some soft corals appear to retain their toxins while others release them, implying a combination of anti-predatory and anti-competitor roles for the secondary metabolites. Scleractinian corals were often overgrown by soft corals.Both species of scleractinian corals were found to cause approximately equal amounts of tissue necrosis in alcyonaceans. These effects were more pronounced when colonies were in direct contact. The necrotic effects among alcyonacean corals were species-specific. Alcyonaceans also overgrew scleractinian corals and secreted a protective polysaccharide layer in areas proximal to scleractinians. Secretion of this layer was stimulated differentially by the two scleractinian species and also varied in frequency of occurrence among the alcyonaceans.High levels of tissue necrosis were observed in both groups of organisms within 3 wk of initiation of the experiment. Necrosis increased with time in the scleractinian corals and decreased in the alcyonaceans. The development of a protective polysaccharide layer in the alcyonaceans increased with time.     

Laboratory models for the study of coral pathologies

Worldwide, the scleractinian corals that characterize contemporary coral reef communities are exhibiting a variety of pathological conditions. These conditions range from diseases linked with specific pathogens to the syndrome known as bleaching. The latter phenomenon involves the loss or reduction of the symbiotic zooxanthellae on which the corals depend. Bleaching appears to be a generalized stress syndrome, but in some cases it may be due to pathogenic infections. A full understanding of coral pathologies requires the development of laboratory models. We have developed two complementary protocols that will facilitate the study of coral pathologies at a number of levels. The first method involves the induction of bleaching by exposing the coral to an acute period of reduced temperature. The second protocol allows the dissociation of coral polyps into a number of cell types that can be maintained long-term in primary culture. Among these are multicellular endothelial isolates (MEI) that contain zooxanthellae and show a high rate of motility. The bleaching protocol will enable investigators to study the processes by which corals recover from bleaching, and it will offer a standard that can be compared to other conditions that lead to bleaching. The cell culture technique will enable the study of mechanisms underlying pathological conditions at the cellular level, and permit studies of how pathological conditions disrupt the relationship between corals and their zooxanthellae.     

The Physiological Mechanisms of Acclimatization in Tropical Reef Corals

SYNOPSIS. The ability of scleractinian corals to survive changesthat are predicted in the global environment over the next centurywill lie in their physiological mechanisms of acclimatization.Corals display rapid modifications in behavior, morphology andphysiology enabling them to photoacclimate to changing lightconditions, a scenario that demonstrates considerable biologicalflexibility. Here we argue that the acclimatization mechanismsin corals are fundamentally similar to those exhibited by otherinvertebrate taxa. We discuss protein metabolism as a mechanismunderlying acclimatization responses in reef corals, and explorethe relationship between protein turnover, metabolic rate, growthrate, and acclimatization capacity. Our preliminary analysessuggest that corals with low growth rates (µCa/mgN/h)and high metabolic rates (µO₂/cm²/hr), such as the massivespecies, acclimatize more effectively than those with high growthrates and low metabolic rates, a feature that is characteristicof branching species. We conclude that studies of protein turnover,combined with temporally relevant investigations into the dynamicaspects of coral dinoflagellate symbioses will provide considerableinsight into why corals exhibit such a high level of variationin response to the same environmental challenge. Furthermore,a more detailed understanding of acclimatization mechanismsis essential if we are to predict how a coral assemblage willrespond to present and future environmental challenges.     

Long distance dispersal and connectivity in amphi-Atlantic corals at regional and basin scales

Among Atlantic scleractinian corals, species diversity is highest in the Caribbean, but low diversity and high endemism are observed in various peripheral populations in central and eastern Atlantic islands and along the coasts of Brazil and West Africa. The degree of connectivity between these distantly separated populations is of interest because it provides insight into processes at both evolutionary and ecological time scales, such as speciation, recruitment dynamics and the persistence of coral populations. To assess connectivity in broadly distributed coral species of the Atlantic, DNA sequence data from two nuclear markers were obtained for six coral species spanning their distributional ranges. At basin-wide scales, significant differentiation was generally observed among populations in the Caribbean, Brazil and West Africa. Concordance of patterns in connectivity among co-distributed taxa indicates that extrinsic barriers, such as the Amazon freshwater plume or long stretches of open ocean, restrict dispersal of coral larvae from region to region. Within regions, dispersal ability appears to be influenced by aspects of reproduction and life history. Two broadcasting species, Siderastrea siderea and Montastraea cavernosa, were able to maintain gene flow among populations separated by as much as 1,200 km along the coast of Brazil. In contrast, brooding species, such as Favia gravida and Siderastrea radians, had more restricted gene flow along the Brazilian coast.     

Symbiont diversity in scleractinian corals from tropical reefs and subtropical non-reef communities in Taiwan

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.Communicated by Biological Editor H.R. Lasker     

Evaluating life‐history strategies of reef corals from species traits

Classifying the biological traits of organisms can test conceptual frameworks of life‐history strategies and allow for predictions of how different species may respond to environmental disturbances. We apply a trait‐based classification approach to a complex and threatened group of species, scleractinian corals. Using hierarchical clustering and random forests analyses, we identify up to four life‐history strategies that appear globally consistent across 143 species of reef corals: competitive, weedy, stress‐tolerant and generalist taxa, which are primarily separated by colony morphology, growth rate and reproductive mode. Documented shifts towards stress‐tolerant, generalist and weedy species in coral reef communities are consistent with the expected responses of these life‐history strategies. Our quantitative trait‐based approach to classifying life‐history strategies is objective, applicable to any taxa and a powerful tool that can be used to evaluate theories of community ecology and predict the impact of environmental and anthropogenic stressors on species assemblages.