Morphological plasticity in scleractinian corals

When describing coral shape and form the term phenotypic plasticity, i.e. environment-induced changes in morphology, is often used synonymously with intraspecific variation. Variation, however, may simply be due to genetic differentiation (polymorphism). Of the 1314 extant scleractinian coral species, less than 20 have been tested for plastic responses. Morphological plasticity has important implications for coral identification, as skeletal features used in coral systematics are directly affected by environment. Furthermore, plastic changes can indicate how corals acclimatise to environmental change. The studies that have examined phenotypic plasticity in corals experimentally can be divided into two groups, i.e. 'non-clonal'—those that have transplanted whole colonies or fragments of colonies (but not treated the fragments as clones) to new environments, and 'clonal'—those that have transplanted colony fragments and used them as clone-mates. The use of clone-mates is preferable as it facilitates the identification of among-genotype variation for plasticity. The heterogeneous nature of the reef environment makes identifying the parameters that affect coral morphology difficult in the field, but there are also many problems conducting suitable aquarium experiments. Nevertheless, evidence to date suggests light and water movement are the most important variables inducing change. As these factors are known to be axiomatic to coral growth, it is possible that associated plastic changes in corals are adaptive; however, this hypothesis is yet to be tested rigorously.     

Reduced Diversity and High Sponge Abundance on a Sedimented Indo-Pacific Reef System: Implications for Future Changes in Environmental Quality

Although coral reef health across the globe is declining as a result of anthropogenic impacts, relatively little is known of how environmental variability influences reef organisms other than corals and fish. Sponges are an important component of coral reef fauna that perform many important functional roles and changes in their abundance and diversity as a result of environmental change has the potential to affect overall reef ecosystem functioning. In this study, we examined patterns of sponge biodiversity and abundance across a range of environments to assess the potential key drivers of differences in benthic community structure. We found that sponge assemblages were significantly different across the study sites, but were dominated by one species Lamellodysidea herbacea (42% of all sponges patches recorded) and that the differential rate of sediment deposition was the most important variable driving differences in abundance patterns. Lamellodysidea herbacea abundance was positively associated with sedimentation rates, while total sponge abundance excluding Lamellodysidea herbacea was negatively associated with rates of sedimentation. Overall variation in sponge assemblage composition was correlated with a number of variables although each variable explained only a small amount of the overall variation. Although sponge abundance remained similar across environments, diversity was negatively affected by sedimentation, with the most sedimented sites being dominated by a single sponge species. Our study shows how some sponge species are able to tolerate high levels of sediment and that any transition of coral reefs to more sedimented states may result in a shift to a low diversity sponge dominated system, which is likely to have subsequent effects on ecosystem functioning.     

Evolution of fringing reefs: space and time constraints from the Gulf of Aqaba

This study documents the pattern and rate of reef growth during the late Holocene as revealed by unique geological conditions at the subsiding NW Gulf of Aqaba. We discovered that the modern fringing reef near the city Elat grows on top of a fossil submerged mid-Holocene reef platform. Four coral cores from the fossil platform were dated using the radiocarbon and U-Th methods. The fossil corals range from 5.6±0.1 to 2.4±0.03 ka, constraining the initiation of the modern reef to 2,400 years ago at most. We documented the detailed morphology of the reef using aerial photographs and scuba diving. The survey shows that at its northern end, growth of the 2-km-long reef is inhibited by an active alluvial fan, and it is composed of isolated knolls that are just approaching the sea surface. Towards the south, the knolls are progressively larger and closer together, until they form a continuous reef platform. Along this north-to-south trend we follow the evolution of reef morphology, changes in coral distribution, and the development of a lagoon separated from the open sea. Based on these observations, we suggest a four-stage reef growth model: (a) the reef initiates as coral colonies, forms knolls, and begins to grow upward, limited by the sea surface. (b) Upon reaching the surface, the knolls spread laterally, preferentially parallel to the dominant wave direction assuming an elongated morphology. (c) Continued growth results in adjacent knolls eventually coalescing to form a continuous jagged reef. We interpret the spurs-and-grooves morphology that can be traced across the reef at Elat as remnants of the original trends of knolls. (d) While reef expansion continues, the original knoll trends may be obscured as a massive reef front takes shape. Considering reef growth rates and observations from the modern reef at Elat, this evolution scheme predicts an age range of 10³ years for corals on the reef platform. The range and distribution of radiometric ages we obtained from the fossil reef platform underlying the living Elat reef confirm this hypothesis.     

Bitten down to size: Fish predation determines growth form of the Caribbean coral reef sponge Mycale laevis

Interactions between organisms add complexity to ecosystem function, particularly on coral reefs. The Caribbean orange icing sponge Mycale laevis is semi-cryptic, often growing under coral colonies or between coral branches. This association is reportedly a mutualism, with the sponge deterring boring sponges from invading the coral skeleton and the coral providing an expanding surface for sponge growth. But is there an alternative explanation for the proximity of sponge and coral? We examined the importance of fish predation on the growth of the sponge. While the semi-cryptic growth form of M. laevis predominates on reefs off the Florida Keys and the Bahamas Islands, M. laevis grows with a non-cryptic, erect morphology off Bocas del Toro, Panama. Surveys revealed that sponge-eating fishes were rare or absent at Bocas del Toro compared to sites in the Florida Keys. Because past studies were inconsistent about the palatability of M. laevis to fish predators, we conducted feeding experiments with sponges from all three sites. Crude organic extracts of M. laevis from all three sites were palatable to generalist fish predators in aquarium assays, and field feeding assays and caging experiments conducted in the Florida Keys confirmed that spongivorous fishes readily ate exposed fragments of M. laevis. Our results suggest that M. laevis is restricted to its semi-cryptic growth form by spongivorous predators, with corals providing a physical refuge from predation. This alternative explanation supports the broader hypothesis that Caribbean reef sponges can be categorized on the basis of chemical defense into defended, palatable, and preferred species, the last of which are restricted to refugia.     

Polyp oriented modelling of coral growth

The morphogenesis of colonial stony corals is the result of the collective behaviour of many coral polyps depositing coral skeleton on top of the old skeleton on which they live. Yet, models of coral growth often consider the polyps as a single continuous surface. In the present work, the polyps are modelled individually. Each polyp takes up resources, deposits skeleton, buds off new polyps and dies. In this polyp oriented model, spontaneous branching occurs. We argue that branching is caused by a so called “polyp fanning effect” by which polyps on a convex surface have a competitive advantage relative to polyps on a flat or concave surface. The fanning effect generates a more potent branching mechanism than the Laplacian growth mechanism that we have studied previously (J. Theor. Biol. 224 (2003) 153). We discuss the application of the polyp oriented model to the study of environmentally driven morphological plasticity in stony corals. In a few examples we show how the properties of the individual polyps influence the whole colony morphology. In our model, the spacing of polyps influences the thickness of coral branches and the overall compactness of the colony. Density variations in the coral skeleton may also be important for the whole colony morphology, which we address by studying two variants of the model. Finally, we discuss the importance of small scale resource translocation in the coral colony and its effects on the morphology of the colony.     

Towards a morphological metric of assemblage dynamics in the fossil record: a test case using planktonic foraminifera

With a glance, even the novice naturalist can tell you something about the ecology of a given ecosystem. This is because the morphology of individuals reflects their evolutionary history and ecology, and imparts a distinct ‘look’ to communities—making it possible to immediately discern between deserts and forests, or coral reefs and abyssal plains. Once quantified, morphology can provide a common metric for characterizing communities across space and time and, if measured rapidly, serve as a powerful tool for quantifying biotic dynamics. Here, we present and test a new high-throughput approach for analysing community shape in the fossil record using semi-three-dimensional (3D) morphometrics from vertically stacked images (light microscopic or photogrammetric). We assess the potential informativeness of community morphology in a first analysis of the relationship between 3D morphology, ecology and phylogeny in 16 extant species of planktonic foraminifera—an abundant group in the marine fossil record—and in a preliminary comparison of four assemblages from the North Atlantic. In the species examined, phylogenetic relatedness was most closely correlated with ecology, with all three ecological traits examined (depth habitat, symbiont ecology and biogeography) showing significant phylogenetic signal. By contrast, morphological trees (based on 3D shape similarity) were relatively distantly related to both ecology and phylogeny. Although improvements are needed to realize the full utility of community morphometrics, our approach already provides robust volumetric measurements of assemblage size, a key ecological characteristic.     

The relationship between corallite morphology and colony shape in some massive reef-corals

Multivariate statistical tests are performed on three coral species to determine if a relationship exists between colony shape and corallite morphology. Hemispherical to branched colony shapes are emphasized. The material consists of specimens of Montastraea annularis and M. cavernosa from modern reef habitats in Jamaica. Colonies of the more branched M. limbata from the Neogene of the Dominican Republic are analyzed for comparison. The characters studied consist largely of linear measurements of colony dimensions and corallite structures in transverse thin-sections. The results show that different complexes of corallite characters vary within each species. With the possible exception of a complex describing theca thickness in M. annularis, none of these corallite complexes is related to variation in colony shape within species. Colony shape variation within species is related to upward colony growth rate. These results indicate that corallite morphology and colony shape can be represented by two different vectors of variation within species. Comparisons of intraspecific with interspecific patterns suggest that a strong genetic component explains colony shape variation in corals.     

Climate warming, marine protected areas and the ocean-scale integrity of coral reef ecosystems

Coral reefs have emerged as one of the ecosystems most vulnerable to climate variation and change. While the contribution of a warming climate to the loss of live coral cover has been well documented across large spatial and temporal scales, the associated effects on fish have not. Here, we respond to recent and repeated calls to assess the importance of local management in conserving coral reefs in the context of global climate change. Such information is important, as coral reef fish assemblages are the most species dense vertebrate communities on earth, contributing critical ecosystem functions and providing crucial ecosystem services to human societies in tropical countries. Our assessment of the impacts of the 1998 mass bleaching event on coral cover, reef structural complexity, and reef associated fishes spans 7 countries, 66 sites and 26 degrees of latitude in the Indian Ocean. Using Bayesian meta-analysis we show that changes in the size structure, diversity and trophic composition of the reef fish community have followed coral declines. Although the ocean scale integrity of these coral reef ecosystems has been lost, it is positive to see the effects are spatially variable at multiple scales, with impacts and vulnerability affected by geography but not management regime. Existing no-take marine protected areas still support high biomass of fish, however they had no positive affect on the ecosystem response to large-scale disturbance. This suggests a need for future conservation and management efforts to identify and protect regional refugia, which should be integrated into existing management frameworks and combined with policies to improve system-wide resilience to climate variation and change.     

Quantifying the response of structural complexity and community composition to environmental change in marine communities

Habitat structural complexity is a key factor shaping marine communities. However, accurate methods for quantifying structural complexity underwater are currently lacking. Loss of structural complexity is linked to ecosystem declines in biodiversity and resilience. We developed new methods using underwater stereo‐imagery spanning 4 years (2010–2013) to reconstruct 3D models of coral reef areas and quantified both structural complexity at two spatial resolutions (2.5 and 25 cm) and benthic community composition to characterize changes after an unprecedented thermal anomaly on the west coast of Australia in 2011. Structural complexity increased at both resolutions in quadrats (4 m²) that bleached, but not those that did not bleach. Changes in complexity were driven by species‐specific responses to warming, highlighting the importance of identifying small‐scale dynamics to disentangle ecological responses to disturbance. We demonstrate an effective, repeatable method for quantifying the relationship among community composition, structural complexity and ocean warming, improving predictions of the response of marine ecosystems to environmental change.     

Relative Importance of Coral Cover,Habitat Complexity and Diversity in Determining the Structure of Reef Fish Communities

The structure of coral reef habitat has a pronounced influence on the diversity, composition and abundance of reef-associated fishes. However, the particular features of the habitat that are most critical are not always known. Coral habitats can vary in many characteristics, notably live coral cover, topographic complexity and coral diversity, but the relative effects of these habitat characteristics are often not distinguished. Here, we investigate the strength of the relationships between these habitat features and local fish diversity, abundance and community structure in the lagoon of Lizard Island, Great Barrier Reef. In a spatial comparison using sixty-six 2m² quadrats, fish species richness, total abundance and community structure were examined in relation to a wide range of habitat variables, including topographic complexity, habitat diversity, coral diversity, coral species richness, hard coral cover, branching coral cover and the cover of corymbose corals. Fish species richness and total abundance were strongly associated with coral species richness and cover, but only weakly associated with topographic complexity. Regression tree analysis showed that coral species richness accounted for most of the variation in fish species richness (63.6%), while hard coral cover explained more variation in total fish abundance (17.4%), than any other variable. In contrast, topographic complexity accounted for little spatial variation in reef fish assemblages. In degrading coral reef environments, the potential effects of loss of coral cover and topographic complexity are often emphasized, but these findings suggest that reduced coral biodiversity may ultimately have an equal, or greater, impact on reef-associated fish communities.     

The contribution of corals to reef structural complexity in Kāne‘ohe Bay

Coral Reefs - The structural complexity of coral reefs provides important ecosystem functions, such as wave attenuation for coastal protection, surfaces for coral growth, and habitat for other...     

Fine-scale time series surveys reveal new insights into spatio-temporal trends in coral cover (2002–2018), of a coral reef on the Southern Great Barrier Reef

Reef monitoring programmes often focus on limited sites, predominantly on reef slope areas, which do not capture compositional variability across zones. This study assessed spatial and temporal changes in hard coral cover at four hierarchical spatial scales. ~ 55,000, geo-referenced photoquadrats were collected annually from 2002 to 2018 and analysed using artificial intelligence for 31 sites across reef flat and reef slope zones on Heron Reef, Southern Great Barrier Reef, Australia. Trends in hard coral cover were examined at three spatial scales: (1) “reef scale”, all data; (2) “geomorphic zone scale”—north/south reef slope, inner/outer reef flat; and (3) “site scale”—31 sites. Coral cover trajectories were also examined at: (4) “sub-site scale”—sub-division of sites into 567 sub-sites, to estimate variability in coral cover trajectories via spatial statistical modelling. At reef scale coral cover increased over time to 25.6 ± 0.4 SE % in 2018 but did not recover following disturbances caused by disease (2004–2008), cyclonic conditions (2009) or severe storms (2015) to the observed pre-disturbance level (44.0 ± 0.7 SE %) seen in 2004. At geomorphic zone scale, the reef slope had significantly higher coral cover than the reef flat. Trends of decline and increase were visible in the slope zones, and the southern slope recovered to pre-decline levels. Variable coral cover trends were visible at site scale. Furthermore, sub-site spatial modelling captured eight years of coral recovery that occurred at different times and magnitudes across the four geomorphic zones, effectively estimating variability in the trajectory of the reef’s coral community. Derived spatial predictions for the entire reef show patchy coral recovery, particularly on the southern slope, and that recovery hotspots are distributed across the reef. These findings suggest that to fully understand and interpret coral decline or recovery on a reef, more accurate assessment can be achieved by examining sites distributed within different geomorphic zones to capture variation in exposure, depth and consolidation.

     

Predicting coral community recovery using multi‐species population dynamics models

Predicting whether, how, and to what degree communities recover from disturbance remain major challenges in ecology. To predict recovery of coral communities we applied field survey data of early recovery dynamics to a multi‐species integral projection model that captured key demographic processes driving coral population trajectories, notably density‐dependent larval recruitment. After testing model predictions against field observations, we updated the model to generate projections of future coral communities. Our results indicated that communities distributed across an island landscape followed different recovery trajectories but would reassemble to pre‐disturbed levels of coral abundance, composition, and size, thus demonstrating persistence in the provision of reef habitat and other ecosystem services. Our study indicates that coral community dynamics are predictable when accounting for the interplay between species life‐history, environmental conditions, and density‐dependence. We provide a quantitative framework for evaluating the ecological processes underlying community trajectory and characteristics important to ecosystem functioning.     

Reciprocal facilitation and non‐linearity maintain habitat engineering on coral reefs

Ecosystem engineers that create habitats facilitate the coexistence of many interacting species. This biotic response to habitat engineering may result in non‐intuitive cascading interactions, potentially including feedbacks to the engineer. Such feedback mechanisms, either positive or negative, may be especially important for the maintenance of biogenic habitats and their community‐wide facilitation. Here, we describe the complex interactions and feedbacks that link marine habitat‐forming engineers, the reef‐building corals, and a group of herbivores, the parrotfishes; the latter preventing the overgrowth of macroalgae, a major competitor of corals. Using density data of eight parrotfish species on a Caribbean reef, we first describe the form of the response of parrotfish abundance to increasing topographic complexity generated by coral growth. Topographic complexity enhanced parrotfish abundance by promoting habitat suitability, but the shape (linear vs asymptotic) and strength of this response varied across species and size. Parrotfish grazing intensity, estimated from data on abundance and species‐, size‐ and life phase‐specific grazing rates also increased with topographic complexity despite an increase in the surface area over which parrotfish graze. Depending on fish species, this functional response was found to be linear or asymptotic. Using a simple analytical model we then explored the effects of topographic complexity and fishing pressure on coral‐algal competition, with particular emphasis on the implications of non‐linearities in the intensity of grazing. Simulations demonstrate that fishing and habitat degradation impair the performance of grazing, but that an asymptotic response of grazing intensity to topographic complexity increases the ecological resilience of coral reefs. Parrotfish and corals are mutually beneficial by creating a loop of positive, indirect feedbacks that maintain their own structure and function: coral growth promotes habitat suitability for parrotfish, concordantly enhancing grazing intensity, which in turn facilitates coral growth by reducing competitive exclusion by macroalgae. We conclude that the resilience of biogenic habitats is enhanced by non‐linear biotic responses to engineering and by the emergence of reciprocal facilitation linking habitat engineering and response organisms.     

Fragmentation and pre-existing species turnover determine land-snail assemblages of tropical rain forest

Aim The main aims of the study were: (1) to investigate the effect of fragment age in relation to other patch‐ and landscape‐scale measures of forest fragmentation, and (2) to assess the relative importance of fragmentation, habitat degradation (i.e. degradation caused by selective logging and past shifting cultivation) and putative pre‐existing species turnover in structuring current land‐snail assemblages. Location South‐western Sri Lanka. Methods The land‐snail fauna was sampled using standardized belt transects. Fifty‐seven transects were sampled in 21 lowland rain forest fragments (c. 1–33,000 ha). The spatial arrangement of fragments in the study area was explicitly considered in an effort to take into account the non‐random nature of fragmentation and degradation and the possibility that current species composition may reflect patterns of species turnover that existed prior to fragmentation. The data set of 57 land‐snail species and 28 environmental and spatial variables was analysed using canonical correspondence analysis and partial canonical correspondence analysis. Results Fragment age, mean shape complexity (i.e. a landscape‐scale measure of shape complexity), altitude, and the spatial variables x (longitude), y (latitude) and y² explained significant variation in land‐snail species composition. None of the three nominal variables quantifying habitat degradation was significantly correlated with variation in species composition. The independent effects of fragment age and mean shape complexity were similar. The combined effect of the spatial variables alone was larger than the independent effects of fragment age, mean shape complexity or altitude, but was of the same order of magnitude. The total variation explained by the spatial variables was comparable to the total non‐spatial variation accounted for by fragment age, mean shape complexity and altitude. Main conclusions Fragment age was found to be one of only two key determinants (the other was shape complexity at the landscape scale) driving fragmentation‐related changes in community composition. The influence of pre‐fragmentation patterns of species turnover on assemblage structure can be stronger than the effects of fragmentation measures, such as age, and may override the effects of forest degradation. Thus, strong patterns of pre‐existing turnover may potentially confound interpretation of the effects of forest fragmentation and degradation.     

Heritability of morphology in brook trout with variable life histories

Distinct morphological variation is often associated with variation in life histories within and among populations of both plants and animals. In this study, we examined the heritability of morphology in three hatchery strains of brook trout (Salvelinus fontinalis), which were historically or are currently used for stocking and supplementation of both migratory and resident ecotypes in the upper Great Lakes region. In a common garden experiment, significant variation in body morphology was observed within and across populations sampled at three time periods. The most notable differences among strains were differences in dorso-ventral body depth and the shape of the caudal peduncle, with some differences in the anterior-posterior placement of the dorsal and ventral fins. Variation with and among 70 half-sib families indicates that heritabilities of morphology and body size were significant at most developmental time points both within and across strains. Heritabilities for morphological characters within strains ranged from 0 to 0.95 across time points. Significant within-strain heritabilities for length ranged from 0 to 0.93 across time points and for weight ranged from 0 to 0.88. Significant additive genetic variation exists within and across hatchery brook trout strains for morphology and size, indicating that these traits are capable of responding to natural or artificial selection.     

Effects of corallivory and coral colony density on coral growth and survival

A suite of processes drive variation in coral populations in space and time, yet our understanding of how variation in coral density affects coral performance is limited. Theory predicts that reductions in density can send coral populations into a predator pit, where concentrated corallivory maintains corals at low densities. In reality, how variation in coral density alters corallivory rates is poorly resolved. Here, we experimentally quantified the effects of corallivory and coral density on growth and survival of small colonies of the staghorn coral Acropora pulchra. Our findings suggest that coral density and corallivory have strong but independent effects on coral performance. In the presence of corallivores, corals suffered high but density-independent mortality. When corallivores were excluded, however, vertical extension rates of colonies increased with increasing densities. While we found no evidence for a predator pit, our results suggest that spatio-temporal variation in corallivore and coral densities can fundamentally alter population dynamics via strong effects on juvenile corals.