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            Patterns and implications of spatial covariation in herbivore functions on resilience of coral reefsAbstract Persistent shifts to undesired ecological states, such as shifts from coral to macroalgae, are becoming more common. This highlights the need to understand processes that can help restore affected ecosystems. Herbivory on coral reefs is widely recognized as a key interaction that can keep macroalgae from outcompeting coral. Most attention has been on the role ‘grazing’ herbivores play in preventing the establishment of macroalgae, while less research has focused on the role of ‘browsers’ in extirpating macroalgae. Here we explored patterns, environmental correlates and state shift consequences of spatial co-variation in grazing and browsing functions of herbivorous fishes. Grazing and browsing rates were not highly correlated across 20 lagoon sites in Moorea, French Polynesia, but did cluster into 3 (of 4) combinations of high and low consumption rates (no site had low grazing but high browsing). Consumption rates were not correlated with grazer or browser fish biomass, but both were predicted by specific environmental variables. Experiments revealed that reversibility of a macroalgal state shift was strongly related to spatial variation in browsing intensity. Our findings provide insights and simple diagnostic tools regarding heterogeneity in top-down forcing that influences the vulnerability to and reversibility of shifts to macroalgae on coral reefs.more » « lessFree, publicly-accessible full text available December 1, 2026
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            ABSTRACT MotivationHere, we make available a second version of the BioTIME database, which compiles records of abundance estimates for species in sample events of ecological assemblages through time. The updated version expands version 1.0 of the database by doubling the number of studies and includes substantial additional curation to the taxonomic accuracy of the records, as well as the metadata. Moreover, we now provide an R package (BioTIMEr) to facilitate use of the database. Main Types of Variables IncludedThe database is composed of one main data table containing the abundance records and 11 metadata tables. The data are organised in a hierarchy of scales where 11,989,233 records are nested in 1,603,067 sample events, from 553,253 sampling locations, which are nested in 708 studies. A study is defined as a sampling methodology applied to an assemblage for a minimum of 2 years. Spatial Location and GrainSampling locations in BioTIME are distributed across the planet, including marine, terrestrial and freshwater realms. Spatial grain size and extent vary across studies depending on sampling methodology. We recommend gridding of sampling locations into areas of consistent size. Time Period and GrainThe earliest time series in BioTIME start in 1874, and the most recent records are from 2023. Temporal grain and duration vary across studies. We recommend doing sample‐level rarefaction to ensure consistent sampling effort through time before calculating any diversity metric. Major Taxa and Level of MeasurementThe database includes any eukaryotic taxa, with a combined total of 56,400 taxa. Software Formatcsv and. SQL.more » « lessFree, publicly-accessible full text available May 1, 2026
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            ABSTRACT Spatial processes, particularly scale‐dependent feedbacks, may play important and underappreciated roles in the dynamics of bistable ecosystems. For example, self‐organised spatial patterns can allow for stable coexistence of alternative states outside regions of bistability, a phenomenon known as a Busse balloon. We used partial differential equations to explore the potential for such dynamics in coral reefs, focusing on how herbivore behaviour and mobility affect the stability of coral‐ and macroalgal‐dominated states. Herbivore attraction to coral resulted in a Busse balloon that enhanced macroalgal resilience, with patterns persisting in regions of parameter space where nonspatial models predict uniform coral dominance. Thus, our work suggests herbivore association with coral (e.g., for shelter) can prevent reefs from reaching a fully coral‐dominated state. More broadly, this study illustrates how consumer space use can prevent ecosystems from undergoing wholesale state transitions, highlighting the importance of explicitly accounting for space when studying bistable systems.more » « lessFree, publicly-accessible full text available March 1, 2026
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            Abstract Understanding the mechanisms underlying nutrient (nitrogen and phosphorus) and carbon cycling in reefs is critical for effective management. Research on reef nutrient and carbon cycling needs to account for (i) the contributions of multiple organisms, (ii) abiotic and biotic drivers, and (iii) a social-ecological perspective. In this paper, we review the mechanisms underlying nutrient and carbon cycling in reef social-ecological systems and analyse them using causal loop analysis. We identify direct and indirect pathways and feedback loops through nutrient and carbon cycles that shape the dominant benthic state of reefs: coral, algal, and sponge-dominated states. We find that two of three anthropogenic impact scenarios (size-selective fishing and land use change) have primarily negative consequences for coral and macroalgae via the nutrient and carbon cycles. A third scenario (runoff) has fewer negative impacts on sponges compared to other benthos. In all scenarios, frequent positive feedback loops (size-selective fishing: 7 of 12 loops; runoff: 6 of 9 loops; land use change: 8 of 11 loops) lead to system destabilization; however, the presence of multiple loops introduces avenues whereby reefs may retain coral dominance despite anthropogenic pressures. Context-specific information on the relative strength of loops will be necessary to predict future reef state.more » « less
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            ABSTRACT Dissolved organic matter (DOM) comprises diverse compounds with variable bioavailability across aquatic ecosystems. The sources and quantities of DOM can influence microbial growth and community structure with effects on biogeochemical processes. To investigate the chemodiversity of labile DOM in tropical reef waters, we tracked microbial utilisation of over 3000 untargeted mass spectrometry ion features exuded from two coral and three algal species. Roughly half of these features clustered into over 500 biologically labile spectral subnetworks annotated to diverse structural superclasses, including benzenoids, lipids, organic acids, heterocyclics and phenylpropanoids, comprising on average one‐third of the ion richness and abundance within each chemical class. Distinct subsets of these labile compounds were exuded by algae and corals during the day and night, driving differential microbial growth and substrate utilisation. This study expands the chemical diversity of labile marine DOM with implications for carbon cycling in coastal environments.more » « less
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            Abstract Cryptic species (evolutionarily distinct lineages that do not align with morphologically defined species) are being increasingly discovered but are poorly integrated into ecological theory. In particular, we still lack a useful understanding of if and how cryptic species differ in ways that affect community recovery from disturbances and responses to anthropogenic stressors, such as the removal of consumers and pollution from nutrients. On coral reefs, nutrient pollution increases the growth of macroalgae that displace corals. Reductions in herbivorous fishes reduce the suppression of macroalgae, while reductions in coralivorous fishes reduce predation on corals. An unresolved question is if and how cryptic coral species respond differently to these impacts, thereby differing in their ability to influence coral community dynamics and maintain coral dominance. Therefore, we assessed how the response of crypticPocilloporaspecies over a period of three years following a simulated disturbance from a cyclone depended on the experimental reduction of fish consumer pressure and nutrient addition. After three years, five morphologically cryptic, but genetically distinct,Pocilloporaspecies recruited to the reef. However, recruitment was dominated by two species:Pocillopora tuahiniensis(46%) andPocillopora meandrina(43%). Under ambient conditions, recruitment ofP. tuahiniensisandP. meandrinawas similar, but experimentally reducing consumer pressure increased recruitment ofP. tuahiniensisby up to 73% and reduced recruitment ofP. meandrinaby up to 49%. In both species, nutrient enrichment increased recruitment and colony growth rates equally, but colonies ofP. tuahiniensisgrew faster and were up to 25% larger after three years than those ofP. meandrina,and growth was unaffected by reduced consumer pressure. Predation by excavating corallivorous fish was higher forP. meandrinathan forP. tuahiniensis, especially under nutrient enrichment. In contrast, polyp extension (an indicator of elevated heterotrophic feeding as well as susceptibility and attractiveness to corallivores) was lower forP. meandrinathan forP. tuahiniensis, especially under low to medium consumer pressure. Overall, we uncovered ecological differences in the response of morphologically cryptic foundation species to two pervasive stressors on coral reefs. Our results demonstrate how cryptic species respond differently to key anthropogenic stressors, which may contribute to response diversity that can support ecological resilience or increase extinction risk.more » « less
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            Abstract Understanding how foundation species recover from disturbances is key for predicting the future of ecosystems in the Anthropocene. Coral reefs are dynamic ecosystems that can undergo rapid declines in coral abundance following disturbances. Understanding why some reefs recover quickly from these disturbances whereas others recover slowly (or not at all) gives insight into the drivers of community resilience. From 2006 to 2010 coral reefs on the fore reef of Moorea, French Polynesia, experienced severe disturbances that reduced coral cover from ~46% in 2005 to <1% in 2010. Following these disturbances, coral cover increased from 2010 to 2018. Although there was a rapid and widespread recovery of corals, reefs at 17 m depth recovered more slowly than reefs at 10 m depth. We investigated the drivers of different rates of coral recovery between depths from 2010 to 2018 using a combination of time‐series data on coral recruitment, density, growth, and mortality in addition to field experiments testing for the effects of predation. Propagule abundance did not influence recovery, as the density of coral recruits (spat <6 months old) did not differ between depths. However, mortality of juvenile corals (≤5 cm diameter) was higher at 17 m, leading to densities of juvenile corals 3.5 times higher at 10 m than at 17 m depth. Yet, there were no differences in the growth of corals between depths. These results point to an early life stage bottleneck after settlement, resulting in greater mortality at 17 m than at 10 m as the likely driver of differential coral recovery between depths. We used experiments and time‐series data to test mechanisms that could drive different rates of juvenile coral mortality across depths, including differences in predation, competition, and the availability of suitable substratum. The results of these experiments suggested that increased coral mortality at 17 m may have been influenced by higher intensity of fish predation, and higher mortality of corals attached to unfavorable substratum. In contrast, the abundance of macroalgae, a coral competitor, did not explain differences in coral survival. Our work suggests that top‐down processes and substratum quality can create bottlenecks in corals that can drive rates of coral recovery after disturbance.more » « lessFree, publicly-accessible full text available January 1, 2026
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            Abstract This study adopts a curvature dynamics approach to understand and predict the trajectory of an idealized depth-averaged barotropic outflow onto a slope in shallow water. A novel equation for streamwise curvature dynamics was derived from the barotropic vorticity equation and applied to a momentum jet subject to bottom friction, topographic slope, and planetary rotation. The terms in the curvature dynamics equation have a natural geometric interpretation whereby each physical process can influence the flow direction. It is shown that a weakly spreading jet onto a steep slope admits the formulation of a 1D ordinary differential equation system in a streamline coordinate system, yielding an integrable ordinary differential equation system that predicts the kinematical behavior of the jet. The 1D model was compared with a set of high-resolution idealized depth-averaged circulation model simulations where bottom friction, planetary rotation, and bottom slope were varied. Favorable performance of the 1D reduced physics model was found, especially in the near field of the outflow. The effect of nonlinear processes such as topographic stretching and bottom torque on the fate of the jet outflow is explained using curvature dynamics. Even in the tropics, planetary rotation can have a surprisingly strong influence on the near-field deflection of an intermediate-scale jet, provided that it flows across steep topography.more » « less
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            Abstract The coral-dinoflagellate endosymbiosis is based on nutrient exchanges that impact holobiont energetics. Of particular concern is the breakdown or dysbiosis of this partnership that is seen in response to elevated temperatures, where loss of symbionts through coral bleaching can lead to starvation and mortality. Here we extend a dynamic bioenergetic model of coral symbioses to explore the mechanisms by which temperature impacts various processes in the symbiosis and to enable simulational analysis of thermal bleaching. Our model tests the effects of two distinct mechanisms for how increased temperature impacts the symbiosis: 1) accelerated metabolic rates due to thermodynamics and 2) damage to the photosynthetic machinery of the symbiont caused by heat stress. Model simulations show that the model can capture key biological responses to different levels of increased temperatures. Moderately increased temperatures increase metabolic rates and slightly decrease photosynthesis. The slightly decreased photosynthesis rates cause the host to receive less carbon and share more nitrogen with the symbiont. This results in temporarily increased symbiont growth and a higher symbiont/host ratio. In contrast, higher temperatures cause a breakdown of the symbiosis due to escalating feedback that involves further reduction in photosynthesis and insufficient energy supply for$$\hbox {CO}_2$$ concentration by the host. This leads to the accumulation of excess light energy and the generation of reactive oxygen species, eventually triggering symbiont expulsion and coral bleaching. Importantly, bleaching does not result from accelerated metabolic rates alone; it only occurs as a result of the photodamage mechanism due to its effect on nutrient cycling. Both higher light intensities and higher levels of DIN render corals more susceptible to heat stress. Conversely, heterotrophic feeding can increase the maximal temperature that can be tolerated by the coral. Collectively these results show that a bioenergetics model can capture many observed patterns of heat stress in corals, such as higher metabolic rates and higher symbiont/host ratios at moderately increased temperatures and symbiont expulsion at strongly increased temperatures.more » « less
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            Abstract Hundreds of studies now document positive relationships between biodiversity and critical ecosystem processes, but as ecological communities worldwide shift toward new species configurations, less is known regarding how the biodiversity of undesirable species will shape the functioning of ecosystems or foundation species. We manipulated macroalgal species richness in experimental field plots to test whether and how the identity and diversity of competing macroalgae affected the growth, survival, and microbiome of a common coral in Mo'orea, French Polynesia. Compared to controls without algal competitors, coral growth was significantly suppressed across three macroalgal monocultures, a polyculture of the same three macroalgae, and plots containing inert seaweed mimics; coral mortality was limited and did not differ significantly among treatments. One macroalga suppressed coral growth significantly less than the other two, but none differed from the inert mimic in terms of coral suppression. The composition, dispersion, and diversity of coral microbiomes in treatments with live macroalgae or inert plastic mimics did not differ from controls experiencing no competition. Microbiome composition differed between two macroalgal monocultures and a monoculture versus plastic mimics, but no other microbiome differences were observed among macroalgal or mimic treatments. Together, these findings suggest that algal diversity does not alter harmful impacts of macroalgae on coral performance, which could be accounted for by physical structure alone in these field experiments. While enhancing biodiversity is a recognized strategy for promoting desirable species, it would be worrisome if biodiversity also enhanced the negative impacts of undesirable species. We documented no such effects in this investigation.more » « less
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