Process-based models of species distributions and the mid-domain effect

Null models that place species ranges at random within a bounded geographical domain produce hump-shaped species richness gradients (the “Mid-Domain Effect”, or MDE). However, there is debate about the extent to which these models are a suitable null expectation for effects of environmental gradients on species richness. Here, I present a process-based framework for modeling species distributions within a bounded geographical domain. Analysis of null models consistent with the mid-domain hypothesis shows that MDEs are indeed likely to be ubiquitous consequences of geographical domain boundaries. Comparing the probability distributions of range locations for the process-based and randomization-based models reveals that randomization models probably overestimate the contribution of MDEs to empirical patterns of species richness, but it also indicates that other, testable predictions from randomization models are likely to be robust. I also show how this process-based framework can be extended beyond null models, to incorporate effects of environmental gradients within the domain. This study provides a first step towards an ecological theory of species distributions in geographical space that can incorporate both “geometric constraints” and effects of environmental gradients, and shows how such a theory can inform our understanding of species richness gradients in nature

Synthesizing Larval Competence Dynamics and Reef-Scale Retention Reveals a High Potential for Self-recruitment in Corals

Many organisms have a complex life-cycle in which dispersal occurs at the propagule stage. For marine environments, there is growing evidence that high levels of recruitment back to the natal population (self-recruitment) are common in many marine organisms. For fish, swimming behavior is frequently invoked as a key mechanism allowing high self-recruitment. For organisms with weak-swimming larvae, such as many marine invertebrates, the mechanisms behind self-recruitment are less clear. Here, we assessed whether the combination of passive retention of larvae due to re-circulation processes near reefs, and the dynamics of settlement competence, can produce the high levels of self-recruitment previously estimated by population genetic studies for reef-building corals. Additionally, we investigated whether time to motility, which is more readily measurable than competence parameters, can explain the between-species variation in self-recruitment. We measured the larval competence dynamics of broadcast-spawning and brooding corals and incorporated these in a model of larval retention around reefs to estimate the potential for self-recruitment and assess its variation among species and reefs. Our results suggest that the larvae of many corals, even those with an obligate planktonic phase, develop with sufficient rapidity to allow high levels of self-recruitment, particularly for reefs with long water retention times. Time to motility explained 77–86% of the between-species variation in potential self-recruitment in scenarios with a realistic range of retention times. Among broadcast spawners, time to motility was strongly and positively correlated with egg size, i.e., broadcast spawner species with small eggs developed more rapidly and exhibited greater potential for self-recruitment. These findings suggest that, along with water retention estimates, easy-to-measure species traits, such as egg size and time to motility, may be good predictors of potential self-recruitment, and therefore may be used to characterize the spectrum of self-recruitment in corals

Benthic composition changes on coral reefs at global scales

Globally, ecosystems are being reconfigured by a range of intensifying human-induced stressors. Coral reefs are at the forefront of this environmental transformation, and if we are to secure their key ecosystem functions and services, it is important to understand the likely configuration of future reefs. However, the composition and trajectory of global coral reef benthic communities is currently unclear. Here our global dataset of 24,468 observations spanning 22 years (1997–2018) revealed that particularly marked declines in coral cover occurred in the Western Atlantic and Central Pacific. The data also suggest that high macroalgal cover, widely regarded as the major degraded state on coral reefs, is a phenomenon largely restricted to the Western Atlantic. At a global scale, the raw data suggest decreased average (± standard error of the mean) hard coral cover from 36 ± 1.4% to 19 ± 0.4% (during a period delineated by the first global coral bleaching event (1998) until the end of the most recent event (2017)) was largely associated with increased low-lying algal cover such as algal turfs and crustose coralline algae. Enhanced understanding of reef change, typified by decreased hard coral cover and increased cover of low-lying algal communities, will be key to managing Anthropocene coral reefs

A unified model explains commonness and rarity on coral reefs

Abundance patterns in ecological communities have important implications for biodiversity maintenance and ecosystem functioning. However, ecological theory has been largely unsuccessful at capturing multiple macroecological abundance patterns simultaneously. Here, we propose a parsimonious model that unifies widespread ecological relationships involving local aggregation, species-abundance distributions, and species associations, and we test this model against the metacommunity structure of reef-building corals and coral reef fishes across the western and central Pacific. For both corals and fishes, the unified model simultaneously captures extremely well local species-abundance distributions, interspecific variation in the strength of spatial aggregation, patterns of community similarity, species accumulation, and regional species richness, performing far better than alternative models also examined here and in previous work on coral reefs. Our approach contributes to the development of synthetic theory for large-scale patterns of community structure in nature, and to addressing ongoing challenges in biodiversity conservation at macroecological scales

Indo-Pacific biodiversity of coral Rreefs: deviations from a mid-domain model

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

Enhancing student engagement to positively impact mathematics anxiety, confidence and achievement for interdisciplinary science subjects

Contemporary science educators must equip their students with the knowledge and practical know-how to connect multiple disciplines like mathematics, computing and the natural sciences to gain a richer and deeper understanding of a scientific problem. However, many biology and earth science students are prejudiced against mathematics due to negative emotions like high mathematical anxiety and low mathematical confidence. Here, we present a theoretical framework that investigates linkages between student engagement, mathematical anxiety, mathematical confidence, student achievement and subject mastery. We implement this framework in a large, first-year interdisciplinary science subject and monitor its impact over several years from 2010 to 2015. The implementation of the framework coincided with an easing of anxiety and enhanced confidence, as well as higher student satisfaction, retention and achievement. The framework offers interdisciplinary science educators greater flexibility and confidence in their approach to designing and delivering subjects that rely on mathematical concepts and practices

Comparative analysis of energy allocation to tissue and skeletal growth in corals

In aquatic invertebrates that form exoskeletons, the partitioning of energy between skeletal and tissue growth is an important tradeoff, especially under resource limitation or physiological stress. Here, we provide the first comparative analysis of energy investment into tissue and skeleton in corals. We develop a mathematical growth model based on colony geometry, tissue mass and quality (enthalpy), and predicted cost of calcification. For hemispherical colonies, the model predicts greater investment in tissue at small sizes, but a shift to skeletal-dominated growth at colony sizes greater than 5-14 cm radius, depending on tissue mass and quality. A similar transition occurs in branches, but is a function of radius and length. An experimental study to assess the impact of resource (light) limitation and physiological stress (sediment load) on energy partitioning in small hemispherical colonies (Goniastrea retiformis Lamarck) and branches (Porites cylindrica Dana) showed that tissue mass and quality varies greatly over small increments in colony or branch size. In particular, allocations to tissue growth varied tenfold (from positive to negative) more across sediment treatments than did allocations to skeletal growth. A model of energy acquisition versus loss (scope for growth) indicated that tissue growth is more responsive to resource variation and physiological stress than skeletal growth. These results suggest that (1) skeletal and tissue growth rates are weakly correlated across environmental conditions, and that (2) variation in tissue properties is a better proxy for coral health or stress than skeletal growth

Allometric growth in reef-building corals

Funding: ARC Centre of Excellence for Coral Reef Studies and the Australian Research Council for fellowship and research support; Scottish Funding Council (MASTS, grant reference HR09011) and the ERC project bioTIME.Predicting demographic rates is a critical part of forecasting the future of ecosystems under global change. Here, we test if growth rates can be predicted from morphological traits for a highly diverse group of colonial symbiotic organisms: scleractinian corals. We ask whether growth is isometric or allometric among corals, and whether most variation in coral growth rates occurs at the level of the species or morphological group. We estimate growth as change in planar area for 11 species, across five morphological groups and over 5 years. We show that coral growth rates are best predicted from colony size and morphology rather than species. Coral size follows a power scaling law with a constant exponent of 0.91. Despite being colonial organisms, corals have consistent allometric scaling in growth. This consistency simplifies the task of projecting community responses to disturbance and climate change.PostprintPeer reviewe

Volatility in coral cover erodes niche structure, but not diversity, in reef fish assemblages

The world’s coral reefs are experiencing increasing volatility in coral cover, largely because of anthropogenic environmental change, highlighting the need to understand how such volatility will influence the structure and dynamics of reef assemblages. These changes may influence not only richness or evenness but also the temporal stability of species’ relative abundances (temporal beta-diversity). Here, we analyzed reef fish assemblage time series from the Great Barrier Reef to show that, overall, 75% of the variance in abundance among species was attributable to persistent differences in species’ long-term mean abundances. However, the relative importance of stochastic fluctuations in abundance was higher on reefs that experienced greater volatility in coral cover, whereas it did not vary with drivers of alpha-diversity. These findings imply that increased coral cover volatility decreases temporal stability in relative abundances of fishes, a transformation that is not detectable from static measures of biodiversity

Prevalence of multimodal species abundance distributions is linked to spatial and taxonomic breadth

We thank the University of St Andrews MHD Cluster and the Bioinformatics Unit (Wellcome Trust ISSF grant 105621/Z/14/Z). L.H.A. was supported by Fundação para a Ciência e Tecnologia, Portugal (POPH/FSE SFRH/BD/90469/2012). A.E.M. acknowledges the ERC (BioTIME 250189). M.D. acknowledges funding from the Marine Alliance for Science and Technology Scotland (MASTS), funded by the Scottish Funding Council (grant reference HR09011) and contributing institutions.Aim. Species abundance distributions (SADs) are a synthetic measure of biodiversity and community structure. Although typically described by unimodal logseries or lognormal distributions, empirical SADs can also exhibit multiple modes. However, we do not know how prevalent multimodality is, nor do we have an understanding of the factors leading to this pattern. Here we quantify the prevalence of multimodality in SADs across a wide range of taxa, habitats and spatial extents. Location. Global. Methods. We used the second-order Akaike information criterion for small sample sizes (AICc) and likelihood ratio tests (LRTs) to test whether models with more than one mode accurately describe the empirical abundance frequency distributions of the underlying communities. We analysed 117 empirical datasets from intensely sampled communities, including taxa ranging from birds, plants, fish and invertebrates, from terrestrial, marine and freshwater habitats. Results. We find evidence for multimodality in 14.5% of the SADs when using AICc and LRT. This is a conservative estimate, as AICc alone estimates a prevalence of multimodality of 22%. We additionally show that the pattern is more common in data encompassing broader spatial scales and greater taxonomic breadth, suggesting that multimodality increases with ecological heterogeneity. Main conclusions. We suggest that higher levels of ecological heterogeneity, underpinned by larger spatial extent and higher taxonomic breadth, can yield multimodal SADs. Our analysis shows that multimodality occurs with a prevalence that warrants its systematic consideration when assessing SAD shape and emphasizes the need for macroecological theories to include multimodality in the range of SADs they predict.PostprintPeer reviewe