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Scleractinian corals are colonial animals with a range of life history strategies, making up diverse species assemblages that define coral reefs. We tagged and tracked approximately 30 colonies from each of 11 species during seven trips spanning six years (2009-2015) in order to measure their vital rates and competitive interactions on the reef crest at Trimodal Reef, Lizard Island, Australia. Pairs of species were chosen from five growth forms where one species of the pair was locally rare (R) and the other common (C). The sampled growth forms were massive [Goniastrea pectinata (R) and G. retiformis (C)], digitate [Acropora humilis (R) and A. cf. digitifera (C)], corymbose [A. millepora (R) and A. nasuta (C)], tabular [A. cytherea (R) and A. hyacinthus (C)] and arborescent [A. robusta (R) and A. intermedia (C)]. An extra corymbose species with intermediate abundance, A. spathulata was included when it became apparent that A. millepora was too rare on the reef crest, making the 11 species in total. The tagged colonies were visited each year in the weeks prior to spawning. During visits, two or more observers each took 2-3 photographs of each tagged colony from directly above and on the horizontal plane with a scale plate to track planar area. Dead or missing colonies were recorded and new colonies tagged in order to maintain approximately 30 colonies per species throughout the six years of the study. In addition to tracking tagged corals, 30 fragments were collected from neighboring untagged colonies of each species for counting numbers of eggs per polyp (fecundity); and fragments of untagged colonies were brought into the laboratory where spawned eggs were collected for biomass and energy measurements. We also conducted surveys at the study site to generate size structure data for each species in several of the years. Each tagged colony photograph was digitized by at least two people. Therefore, we could examine sources of error in planar area for both photographers and outliners. Competitive interactions were recorded for a subset of species by measuring the margins of tagged colony outlines interacting with neighboring corals. The study was abruptly ended by Tropical Cyclone Nathan (Category 4) that killed all but nine of the over 300 tagged colonies in early 2015. Nonetheless, these data will be of use to other researchers interested in coral demography and coexistence, functional ecology, and parametrizing population, community and ecosystem models. The data set is not copyright restricted, and users should cite this paper when using the data. Publisher PDF

Models of dispersal potential are required to predict connectivity between populations of sessile organisms. However, to date, such models do not allow for time-varying rates of acquisition and loss of competence to settle and metamorphose, and permit only a limited range of possible survivorship curves. We collect high-resolution observations of coral larval survival and metamorphosis, and apply a piecewise modeling approach that incorporates a broad range of temporally varying rates of mortality and loss of competence. Our analysis identified marked changes in competence loss and mortality rates, the timing of which implicates developmental failure and depletion of energy reserves. Asymmetric demographic rates suggest more intermediate-range dispersal, less local retention, and less long-distance dispersal than predicted by previously employed non-piecewise models. Because vital rates are likely temporally asymmetric, at least for nonfeeding broadcast-spawned larvae, piecewise analysis of demographic rates will likely yield more reliable predictions of dispersal potential.

MD was supported by the ERC (BioTIME 250189) and the Scottish Funding Council (MASTS –HR09011). JM, AB and SC were supported by fellowships from the Australian Research Council (FT110100609, FT0990652 and DP0880544, respectively). Competition is an important determinant of assemblage structure and population regulation, often resulting in decreased growth, fecundity or survival. In corals, most studies testing for an effect of competition on demographic traits, such as growth, have been experimental and often impose very high levels of competition upon colonies. To more realistically assess the role of competition on coral traits, multispecies studies in the wild are required. Here, we use 5 yr of data that includes 11 coral species on the reef crest at Lizard Island on the Great Barrier Reef to quantify the effect of competition on growth. Additionally, we test whether species differ in their susceptibility to direct‐contact (overgrowth and digestion) and overtopping competition, and whether species from some morphological groups are more likely to compete with one another than with species from other morphological groups. We also investigate the relationships between competitive ability and three key traits: growth rate, mechanical stability and fecundity. In contrast to most previous work using field manipulations of competition, we found a negligible effect of competition on growth. Acropora species consistently won overtopping encounters but lost in direct‐contact encounters, and these results were consistent among the four Acropora morphological groups. In contrast, the massive Goniastrea spp. were poor at overtopping but generally won direct‐contact encounters. Only tabular colonies were disproportionally more likely to compete against one another than with other morphologies. This propensity increases intraspecific relative to interspecific competition, a phenomenon that can promote coexistence when it is present among dominant competitors. Good competitors grew more quickly and had higher fecundity but were less mechanically stable, implying a tradeoff between performance during disturbance vs. performance in the absence of disturbance. We conclude that competition among adults is less likely to influence community dynamics than previously thought. If competition does have an effect, it is more likely to occur at life‐stages other than adults. Publisher PDF

JM, AB and SC were supported by fellowships from the Australian Research Council (FT110100609, FT0990652 and DP0880544 respectively). MD was supported by the ERC (BioTIME 250189) and the Scottish Funding Council (MASTS - HR09011). Understanding species differences in demographic strategies is a fundamental goal of ecology. In scleractinian corals, colony morphology is tightly linked with many demographic traits, such as size-specific growth and morality. Here, we test how well morphology predicts the colony size-fecundity relationship in eight species of broadcast-spawning corals. Variation in colony fecundity is greater among morphologies than between species with a similar morphology, demonstrating that colony morphology can be used as a quantitative proxy for demographic strategies. Additionally, we examine the relationship between size-specific colony fecundity and mechanical vulnerability (i.e. vulnerability to colony dislodgment). Interestingly, the relationship between size-specific fecundity and mechanical vulnerability varied among morphologies. For tabular species, the most fecund colonies are the most mechanically vulnerable, while the opposite is true for massive species. For corymbose and digitate colonies, mechanical vulnerability remains relatively constant as fecundity increases. These results reveal strong differences in the demographic trade-offs among species of different morphologies. Using colony morphology as a quantitative proxy for demographic strategies can help predict coral community dynamics and responses to anthropogenic change. Publisher PDF

Interspecific interactions that produce nontransitive competitive networks have been proposed to promote diversity in a broad range of systems, including coral reefs. In this paper, we model the effect of size-dependent shifts in competitive ability on the coexistence of canopy-forming and understory coral species, and we predict how these shifts influence patterns of community structure along large-scale gradients in disturbance and recruitment limitation. We consider three models, representing a gradient from purely hierarchical competition in which the canopy-former is dominant, to competition involving standoffs and reversals between the understory species and juvenile canopy-formers. Analysis of these models leads to two key conclusions. First, as competition becomes less transitive, coexistence may be promoted or inhibited, depending upon the extent to which the canopy-former can sustain itself by clonal propagation. Specifically, when clonal growth alone is adequate to sustain the canopy-former, increasing nontransitivity promotes coexistence. When it is not, nontransitivity inhibits coexistence. Secondly, size-dependent nontransitivity dramatically changes how gradients in disturbance and recruitment affect species coexistence. In contrast to hierarchical interactions, standoffs and reversals do not show an “intermediate recruitment” phenomenon, in which coexistence is facilitated at intermediate levels of recruitment. Moreover, under hierarchical competition, the dominant always benefits more than the subordinate as recruitment is increasingly facilitated. Under standoffs and reversals, however, increasing recruitment often favors the canopy-former at some levels of disturbance, but the understory species at other levels of disturbance. These results differ markedly from previous models of stage-dependent competition, suggesting that promotion of coexistence by ontogenetic shifts in competitive ability depends upon the mechanisms by which competition occurs in particular ecological contexts. The results also indicate that the effects of gradients in disturbance and recruitment on community structure depend fundamentally on how species compete for space.

Understanding the nature and causes of global gradients in species richness is a perennial ecological problem, and recent work has highlighted the need to assess these gradients relative to an appropriate statistical expectation. This paper examines latitudinal and longitudinal gradients in species richnesses of corals and reef fishes in the Indo-Pacific domain and compares them with gradients predicted by a mid-domain model in which geographic domains are located at random between the latitudinal and longitudinal boundaries of this region. We test for significant differences between observed and predicted species-richness patterns, and we identify regions that are enriched or depauperate in species, relative to expectation. In addition, we move beyond previous mid-domain analyses by directly comparing observed spatial distributions of geographic ranges with those predicted by a mid-domain model. This comparison indicates precisely how species-richness anomalies are produced by nonrandomness in the distribution of species ranges. For both corals and fishes, large and statistically significant differences exist between observed latitudinal and longitudinal species-richness gradients and those predicted by mid-domain models. Longitudinally, species richness is markedly higher than predicted along the African coast and, to a lesser extent, within the Indo-Australian Archipelago (IAA), and it is markedly lower than expected in the eastern Pacific. Latitudinally, species richness becomes increasingly higher than predicted as one moves from the equator to the tropical margins; then it becomes sharply lower than predicted beyond the tropics. Unexpectedly, differences between observed and predicted spatial distributions of range endpoints and midpoints reveal a pattern of nonrandomness that is highly congruent with the hypothesis that gyres in the Indian and Pacific Oceans, with the IAA forming a porous boundary between them, have a major influence on Indo-Pacific species-richness patterns. Our analyses indicate that the perspective offered by a focus on explaining nonrandomness in the location of geographic ranges (rather than explaining why species numbers vary in space) is likely to dramatically alter our assessments of alternative explanations for global species-richness gradients.

A latitudinal gradient in the recruitment rates of intertidal mussels and barnacles was detected in the Northeast Pacific during 1996 and 1997. This gradient was approximately a stepcline: annual recruitment, on average, was 1-2 orders of magnitude higher in central and northern Oregon than in central and northern California. In contrast to the regional differences, large-scale gradients in recruitment within California were small: correlations of recruitment with latitude were weak, and, in all but one case, statistically insignificant. Nonetheless, trends in the data suggest that recruitment within central and northern California was highest between San Francisco and Monterey Bay, where larvae may be retained more nearshore than to the north or south. If so, apparently conflicting claims about latitudinal gradients in recruitment within California can be reconciled. The large scale transition in recruitment rates supports the hypothesis that a marked shift in the intensity of upwelling near Cape Blanco in southern Oregon is a major cause of a coincident transition in community structure. Stronger upwelling (and thus offshore flow) to the south has been hypothesized to transport larvae further offshore and thereby reduce larval supply to nearshore benthic communities. This study confirms that the predicted differences in recruitment exist, and that these differences are large. Preliminary calculations indicate that regional differences in offshore flow are likely to make a larger contribution to the recruitment transitions than several other plausible causes. In addition, recruitment transitions are larger, more abrupt, and more consistent across species than corresponding shifts in percentage cover, which favor competitive dominants. This supports model predictions that competition for space is more intense where recruitment is high. However, the absence of strong, large-scale recruitment gradients within California suggests that mesoscale processes are relatively more important than latitudinal trends in upwelling as determinants of community structure patterns at smaller scales.

Biodiversity may provide insurance against ecosystem collapse by stabilizing assemblages that perform particular ecological functions (the "portfolio effect"). However, the extent to which this occurs in nature and the importance of different mechanisms that generate portfolio effects remain controversial. On coral reefs, herbivory helps maintain coral dominated states, so volatility in levels of herbivory has important implications for reef ecosystems. Here, we used an extensive time series of abundances on 35 reefs of the Great Barrier Reef of Australia to quantify the strength of the portfolio effect for herbivorous fishes. Then, we disentangled the contributions of two mechanisms that underlie it (compensatory interactions and differential responses to environmental fluctuations ["response diversity"]) by fitting a community-dynamic model that explicitly includes terms for both mechanisms. We found that portfolio effects operate strongly in herbivorous fishes, as shown by nearly independent fluctuations in abundances over time. Moreover, we found strong evidence for high response diversity, with nearly independent responses to environmental fluctuations. In contrast, we found little evidence that the portfolio effect in this system was enhanced by compensatory ecological interactions. Our results show that portfolio effects are driven principally by response diversity for herbivorous fishes on coral reefs. We conclude that portfolio effects can be very strong in nature and that, for coral reefs in particular, response diversity may help maintain herbivory above the threshold levels that trigger regime shifts.

Species assemblages vary in structure due to a wide variety of processes operating at ecological and much broader biogeographical scales. Cross-scale studies of assemblage structure are necessary to fully understand this variability. Here, we evaluate the abundance and occupancy patterns of hierarchically sampled coral assemblages in three habitats (reef flat, crest, and slope) and five regions (Indonesia, Papua New Guinea, the Solomon Islands, American Samoa, and the Society Islands) across the west-central Pacific Ocean. Specifically, we compare two alternative models that unify spatial variance and occupancy via the negative binomial distribution. The first assumes a power-law scaling between the mean and variance of abundance; the second assumes a quadratic variance–mean relationship and a constant abundance-invariant aggregation parameter. Surprisingly, the well-established power-law model performs worse than the model assuming abundance-invariant aggregation, for both variance–mean and occupancy–abundance relationships. We also find strong evidence for regional and habitat variation in these relationships and in the levels of aggregation estimated by the abundance-invariant aggregation model. Among habitats, corals on reef flats exhibited lower occupancy and higher levels of aggregation compared to reef crests and slopes. Among regions, low occupancy and high aggregation were most pronounced across all habitats in American Samoa. These patterns may be related to habitat and regional differences in disturbance and recovery processes. Our results suggest that the spatial scaling of abundance and occupancy is sensitive to processes operating among these habitats and at regional scales. However, the consistency of these relationships across species within assemblages suggests that a theoretical unification of spatial variance and occupancy patterns is indeed possible.

Dispersal influences ecological dynamics, evolution, biogeography, and biodiversity conservation, but models of larval dispersal in marine organisms make simplifying assumptions that are likely to approximate poorly the temporal dynamics of larval survival and capacity for settlement. In particular, larval mortality rates are typically assumed to be constant throughout larval life; and all larvae are frequently assumed to acquire and lose competence at the same time. To improve upon these assumptions, we here develop simple models of dispersal potential that incorporate rates of mortality, and acquisition and loss of settlement competence. We fit these models to empirical competence and survival data for five scleractinian coral species, to test the models' ability to characterize empirical survival and competence patterns, and to estimate the dispersal potential implied by those patterns. The models fit the data well, incorporating qualitative features of competence and survival that traditional approaches to modeling dispersal do not, with important implications for dispersal potential. Most notably, there was high within-cohort variation in the duration of the competent period in all species, and this variation increases both self-recruitment and long-distance dispersal compared with models assuming a fixed competent period. These findings help to explain the seeming paradox of high genetic population structure, coupled with large geographic range size, observed in many coral species. More broadly, our approach offers a way to parsimoniously account for variation in competence dynamics in dispersal models, a phenomenon that our results suggest has important effects on patterns of connectivity in marine metapopulations.

Patterns in the commonness and rarity of species are a fundamental characteristic of ecological assemblages; however, testing between alternative models for such patterns remains an important challenge. Conventional approaches to fitting or testing species abundance models often assume that species, not individuals, are the units that are sampled and that species' abundances are independent of one another. Here we test three different models (the Poisson lognormal, the negative binomial, and the neutral, “zero-sum multinomial” [ZSM]) against species abundance distributions of Indo-Pacific corals and reef fishes. We derive and apply several alternative bootstrap analyses of model fit, each of which makes different assumptions about how species abundance data are sampled, and we assess the extent to which tests of model fit are sensitive to such assumptions. For all models, goodness of fit is remarkably consistent, regardless of whether one assumes that species or individuals are the units that are sampled or whether or not one assumes that species' abundances are statistically independent of one another. However, goodness-of-fit estimates are approximately twice as precise and detect lack of model fit more frequently, when based on sampling of individuals, rather than species. Bootstrap analyses indicate that the Poisson lognormal distribution exhibits substantially better fit to species abundance patterns, consistent with model selection analyses. In particular, heterogeneity in species abundances (many rare and few highly abundant species) is too great to be captured by the ZSM model or the negative binomial model and is best explained by models that predict species abundance patterns that are much closer, but not identical, to the lognormal distribution. More broadly, our bootstrap analyses suggest that estimates of model fit are likely to be robust to assumptions about the statistical interdependence of species abundances, but that tests of model fit are more powerful when they assume sampling of individuals, rather than species. Such individual-based tests therefore may be able to identify lack of model fit where previous tests have been inconclusive.

The ‘‘fundamental niche’’ is the range of conditions under which an organism can survive and reproduce, measured in the absence of biotic interactions. Niche measurements are often based on statistical relationships between species presence and measured environmental variables, or inferred from measured responses of species along hypothesized niche axes. In this study, we use novel, process-based models of how irradiance and gas diffusion influence photosynthesis and respiration to predict niche dimensions for three coral species: Acropora nasuta, Montipora foliosa, and Leptoria phrygia. We use a combination of mathematical modeling, laboratory experiments, and field observations to establish the link between energy acquisition and the dominant environmental gradients on reefs: light intensity and water flow velocity. Our approach allows us to quantify how the shape of the niche varies in response to light and flow conditions. The model predicts that, due to its higher photosynthetic capacity, the branching coral A. nasuta has a positive energy balance over a wider range of conditions than both a massive species (L. phrygia) and a foliose species (M. foliosa). Moreover, colony size influences niche width, with larger colonies of all three species achieving a positive energy balance over a broader range of conditions than small colonies. Comparison of model predictions with field data demonstrated that tissue biomass and reproductive output are significantly and positively correlated with predicted energy acquisition. These results show how interactions between light and flow determine organism performance along environmental gradients on coral reefs. In addition, this study demonstrates the utility of process-based models for quantifying how physiology influences ecology, and for predicting the ecological consequences of varying environmental conditions.

The size of the regional species pool may influence local patterns of diversity. However, it is unclear whether certain spatial scales are less sensitive to regional influences than others. Additive partitioning was used to separate coral-dwelling fish diversity to its alpha and beta components, at multiple scales, in several regions across the Indo-Pacific. We then examined how the relative contribution of these components changes with increased regional diversity. By employing specific random-placement null models, we overcome methodological problems with local-regional regressions. We show that, although alpha and beta diversities within each region are consistently different from random-placement null models, the increase in beta diversities among regions was similar to that predicted once heterogeneity in coral habitat was accounted for. In contrast, alpha diversity within single coral heads was limited and increased less than predicted by the null models. This was correlated with increased intraspecific aggregation in more diverse regions and is consistent with ecological limitations on the number of coexisting species at the local scale. These results suggest that, apart from very small spatial scales, variation in the partitioning of fish diversity along regional species richness gradients is driven overwhelmingly by the corresponding gradients in coral assemblage structure.

Morphological plasticity in response to environmental heterogeneity may be performance enhancing or may simply result from an intrinsic instability in morphology during development. Although patterns of morphological change are well documented for numerous taxa, it is often unclear whether this plasticity enhances the performance of organisms in the habitat to which they have acclimatized. Reef-building corals are an ideal model system in which to investigate this question. We here develop a three-dimensional geometric model and present a comprehensive photosynthesis data set with experimentally calibrated photosynthesis models that predicts energy acquisition by foliose corals as a function of colony shape. This allows us to assess the extent to which changes in colony morphology along an environmental gradient track the predicted optimal colony morphologies. Our results provide strong evidence that phenotypic plasticity in foliose corals optimizes photosynthetic energy acquisition and is not simply a mechanism to increase light capture. We show that the optimal morphology is constrained at the boundaries of the environmental gradient, with non-optimal morphologies in these habitats having greatly reduced energy acquisition. However, at the center of the environmental gradient, flexibility in photophysiology allows energy acquisition to be very similar for multiple morphologies. Our results highlight the importance of phenotypic plasticity at multiple scales. Variation in overall morphology is important at niche boundaries at which conditions are consistently more stressful, whereas physiological flexibility is important in intermediate and less predictable habitats in which a rapid and reversible response to environmental fluctuations is required.