Fear effects associated with predator presence and habitat structure interact to alter herbivory on coral reefs.

Non-consumptive fear effects are an important determinant of foraging decisions by consumers across a range of ecosystems. However, how fear effects associated with the presence of predators interact with those associated with habitat structure remain unclear. Here, we used predator fish models (Plectropomus leopardus) and experimental patches of the macroalga Sargassum ilicifolium of varying densities to investigate how predator- and habitat-associated fear effects influence herbivory on coral reefs. We found the removal of macroalgal biomass (i.e. herbivory) was shaped by the interaction between predator- and habitat-associated fear effects. Rates of macroalgal removal declined with increasing macroalgal density, likely due to increased visual occlusion by denser macroalgae patches and reduced ability of herbivorous fishes to detect the predators. The presence of the predator model reduced herbivory within low macroalgal density plots, but not within medium- and high-density macroalgal plots. Our results suggest that fear effects due to predator presence were greatest at low macroalgal density, yet these effects were lost at higher densities possibly due to greater predation risk associated with habitat structure and/or the inability of herbivorous fishes to detect the predator model.

Seasonal variation of Sargassum ilicifolium (Phaeophyceae) growth on equatorial coral reefs.

Temporal and spatial variations in Sargassum ilicifolium thallus density and length were investigated on equatorial coral reefs in Singapore from November 2011 to October 2012. Thalli density varied little throughout the year, however, we found strong seasonal patterns in thallus length and identified temperature as the significant driver. Sargassum ilicifolium reached maximum length in December (110.39 ± 2.37 cm) during periods of cooler water temperatures, and minimum length in May (9.88 ± 0.48 cm) during periods of warmer water temperatures. Significant spatial variation was also observed for both thallus density and length of S. ilicifolium among reefs. Within reefs, densities of S. ilicifolium were higher on reef flats (20.40 ± 0.40 individuals · 0.25 m-2 ) compared to upper reef slopes (5.66 ± 0.23 individuals · 0.25 m-2 ). Our findings highlight that marked seasonality in the growth of canopy-forming macroalgae can occur within equatorial reef systems where temperature ranges are restricted (<3°C).

Structural complexity and component type increase intertidal biodiversity independently of area.

Complexity is well accepted as one of the primary drivers of biodiversity, however, empirical support for such positive associations is often confounded with surface area and undermined by other potential explanatory factors, especially the type of structural component (e.g., pits, crevices, overhangs, etc.). In the present study, sample units (artificial substrates) of equal surface area (± 0.2%) were used to simultaneously examine the independent effects of complexity and different structural component types on species richness (S), abundance (N), and community composition. We created simple and complex concrete substrates of four different geometric designs using novel software. The substrates (n = 8) were mounted onto granite seawalls (at two tidal heights) on two islands south of Singapore Island. After 13 months of colonization, all 384 tiles were collected and their assemblages compared. A total of 53 744 individuals of 70 species/morphospecies were collected and identified. Our results show that greater complexity can support greater species richness and different communities that are independent of surface area. Furthermore, the type of structure (e.g., "pits," "grooves," "towers") can have an effect on richness and community composition that is independent of complexity.

Genome-wide analysis to uncover how Pocillopora acuta survives the challenging intertidal environment.

Characterisation of genomic variation among corals can help uncover variants underlying trait differences and contribute towards genotype prioritisation in coastal restoration projects. For example, there is growing interest in identifying resilient genotypes for transplantation, and to better understand the genetic processes that allow some individuals to survive in specific conditions better than others. The coral species Pocillopora acuta is known to survive in a wide range of habitats, from reefs artificial coastal defences, suggesting its potential use as a starter species for ecological engineering efforts involving coral transplantation onto intertidal seawalls. However, the intertidal section of coastal armour is a challenging environment for corals, with conditions during periods of emersion being particularly stressful. Here, we scanned the entire genome of P. acuta corals to identify the regions harbouring single nucleotide polymorphisms (SNPs) and copy number variations (CNVs) that separate intertidal colonies (n = 18) from those found in subtidal areas (n = 21). Findings revealed 74,391 high quality SNPs distributed across 386 regions of the P. acuta genome. While the majority of the detected SNPs were in non-coding regions, 12% were identified in exons (i.e. coding regions). Functional SNPs that were significantly associated with intertidal colonies were found in overrepresented genomic regions linked to cellular homeostasis, metabolism, and signalling processes, which may represent local environmental adaptation in the intertidal. Interestingly, regions that exhibited CNVs were also associated with metabolic and signalling processes, suggesting P. acuta corals living in the intertidal have a high capacity to perform biological functions critical for survival in extreme environments.

Relative predation intensity of an intertidal gastropod on artificial coastal defense structures.

Despite seawalls becoming ubiquitous coastal features, and having some physical similarities to natural rocky shores, it remains unclear how these urban habitats influence predator-prey interactions. Predators can affect intertidal mobile prey densities through two pathways: (1) successful predation directly influences prey mortality rates, and (2) direct and indirect effects of predation can scare and induce motile prey to seek safer areas. In this study, we investigated whether intertidal predators affect the density of the marine gastropod, Nerita undata, at four seawall sites in Singapore. Using a tethering method that we developed, we monitored the mortality and other evidence of predation (shell state) of tethered N. undata. Field experiments revealed high (22.5%-82.5%) predation potential of N. undata across the four sites, with significantly higher predation risk at lower shore heights and for snails with mixed shell coloration. Observations and analysis of the shell state after 3 days showed that predation on seawalls was primarily by crushing predators such as fish. Other predators of N. undata include predatory snails, with various feeding methods that left behind different predator signatures. Our results add substantially to the limited knowledge on predator-prey interactions on seawalls, particularly for Nerita undata, and suggest that seawall systems are more dynamic than previously thought. This further highlights the role of these artificial structures as important habitats and feeding grounds in urban coastal ecosystems.

The effect of coral colony morphology, coral surface condition, particle size, and seeding point on the trapping and deposition of microplastics.

Coral reefs are increasingly identified as microplastic sinks. Understanding the trapping and deposition effects on microplastics among coral colonies of different morphologies can help identify which corals and coral reefs are at higher risk of microplastic exposure. Here, we used a current-generating saltwater flume to explore microplastic trapping and deposition among branching coral, Pocillopora acuta, colonies with contrasting morphologies (open and compact), together with varying coral surface conditions (live, dead, and waxed), microplastic sizes (400 to 500 µm and 900 to 1000 µm), and seeding points (above-colony and mid-colony). Results revealed that more microplastics were trapped by, and deposited nearer to, compact colonies compared to those with a more open morphology-likely due to differences in flow dynamics. More of the larger microplastics were trapped, as were those introduced at the mid seeding point, but coral surface condition had no significant effect. These findings add to the growing evidence that corals are effective at trapping and facilitating deposition of microplastics. Branching corals with compact structures are potentially at high risk of microplastic pollution impact. We posit that coral composition, i.e. the relative abundance of compact branching colonies, will affect microplastic accumulation in natural reef environments. SYNOPSIS: This study demonstrates the effects of coral morphology on microplastic trapping and deposition, providing mechanistic insights into the factors that contribute to coral reefs acting as microplastic sinks.

Regional decline in growth rates of massive Porites corals in Southeast Asia.

This study reports the first well-replicated analysis of continuous coral growth records from warmer water reefs (mean annual sea surface temperatures (SST) >28.5 °C) around the Thai-Malay Peninsula in Southeast Asia. Based on analyses of 70 colonies sampled from 15 reefs within six locations, region-wide declines in coral calcification rate (ca. 18.6%), linear extension rate (ca. 15.4%) and skeletal bulk density (ca. 3.9%) were observed over a 31-year period from 1980 to 2010. Decreases in calcification and linear extension rates were observed at five of the six locations and ranged from ca. 17.2-21.6% and ca. 11.4-19.6%, respectively, whereas decline in skeletal bulk density was a consequence of significant reductions at only two locations (ca. 6.9% and 10.7%). A significant link between region-wide growth rates and average annual SST was found, and Porites spp. demonstrated a high thermal threshold of ca. 29.4 °C before calcification rates declined. Responses at individual locations within the region were more variable with links between SST and calcification rates being significant at only four locations. Rates of sea temperature warming at locations in the Andaman Sea (Indian Ocean) (ca. 1.3 °C per decade) were almost twice those in the South China Sea (Pacific Ocean) (ca. 0.7 °C per decade), but this was not reflected in the magnitude of calcification declines at corresponding locations. Considering that massive Porites spp. are major reef builders around Southeast Asia, this region-wide growth decline is a cause for concern for future reef accretion rates and resilience. However, this study suggests that the future rates and patterns of change within the region are unlikely to be uniform or dependent solely on the rates of change in the thermal environment.

The comparative effects of chronic microplastic and sediment deposition on the scleractinian coral Merulina ampliata.

Despite increasing research into the effects of microplastics on corals, no study to date has compared this relatively novel pollutant with a well-established stressor such as downwelling sediments. Here, Merulina ampliata coral fragments were exposed to polyethylene terephthalate (PET) and calcium carbonate particles (200-300 µm) at two deposition levels, high (115.20 ± 5.83 mg cm-2 d-1, mean ± SE) and low (22.87 ± 1.90 mg cm-2 d-1) in specially-designed Flow-Through Resuspension (FloTR) chambers. After 28 d, there were no significant differences between fragments exposed to sediments and microplastics for coral skeletal growth, Symbiodiniaceae density, and areal or cellular chlorophyll a concentrations. There were also no significant differences between levels of treatments, or with the control fragments. More PET microplastic particles were incorporated into the coral skeletons of fragments exposed to microplastics compared to those exposed to sediment and the control fragments, but there was no difference between fragments exposed to high and low microplastic levels. Together, the results show that M. ampliata appears to be able to cope with both microplastic and sediment stress, and suggests that microplastics do not represent a more serious threat than downwelling sediments at the levels tested.

EmerSense: A low-cost multiparameter logger to monitor occurrence and duration of emersion events within intertidal zones.

The intertidal zone is a harsh environment for marine life as conditions are often both extreme and variable. A wide range of sessile organisms are partially or fully emersed (exposed above the water line) during low tide. In the tropics, when corals are emersed, high light and temperature can be detrimental to their survival. To date, there is no commercially-available logger that can detect periods of emersion, information that is useful for marine research and for coastal resource management. Here, we present a low-cost Arduino-based multi-parameter logger called 'EmerSense' which can detect instances of emersion while simultaneously recording light and temperature profiles. We describe the different steps involved in fabricating EmerSense, including hardware construction and software design, and discuss the results of our field testing at an intertidal coral reef in Singapore.

Chemically Mediated Interactions with Macroalgae Negatively Affect Coral Health but Induce Limited Changes in Coral Microbiomes.

Allelopathic chemicals facilitated by the direct contact of macroalgae with corals are potentially an important mechanism mediating coral-macroalgal interactions, but only a few studies have explored their impacts on coral health and microbiomes and the coral's ability to recover. We conducted a field experiment on an equatorial urbanized reef to assess the allelopathic effects of four macroalgal species (Bryopsis sp., Endosiphonia horrida, Hypnea pannosa and Lobophora challengeriae) on the health and microbiomes of three coral species (Merulina ampliata, Montipora stellata and Pocillopora acuta). Following 24 h of exposure, crude extracts of all four macroalgal species caused significant coral tissue bleaching and reduction in effective quantum yield. The corals were able to recover within 72 h of the removal of extracts, except those that were exposed to L. challengeriae. While some macroalgal extracts caused an increase in the alpha diversity of coral microbiomes, there were no significant differences in the composition and variability of coral microbiomes between controls and macroalgal extracts at each sampling time point. Nevertheless, DESeq2 differential abundance analyses showed species-specific responses of coral microbiomes. Overall, our findings provide insights on the limited effect of chemically mediated interactions with macroalgae on coral microbiomes and the capacity of corals to recover quickly from the macroalgal chemicals.

Sediment trap height affects mass, particle size, and biogeochemical composition of material collected in an equatorial coral reef.

Anthropogenic sedimentation is a major contributor to the worldwide decline in coral cover. Resuspension of benthic material can exacerbate the threat to corals, but evidence of vertical sediment gradients is limited. Here, we installed sediment traps at 10, 20, 30, 40, and 50 cm above the substrate at three equatorial reef sites for three months and determined the dry mass, particle size, and biogeochemical composition of the material collected. As the trap mouth height increased from 10 to 50 cm, dry mass decreased as sediments became finer, poorer in carbonate, and richer in silicate. Despite among-site differences in collected sediment, this vertical gradient was present in all trap arrays, likely driven by resuspension mechanisms on seabed sediments. These results have implications for coral vertical ecology and underline the importance of standardising sediment collection protocols.

Behaviour broadens thermal safety margins on artificial coastal defences in the tropics.

Tropical species are predicted to be among the most vulnerable to climate change as they often live close to their upper limits to thermal tolerance and in many cases, behavioural thermoregulation is required to persist in the thermal extremes of tropical latitudes. In concert with warming temperatures, near-shore species are faced with the additional threat of shoreline hardening, leading to a reduction in microhabitats that can provide thermal refuges. This situation is exemplified in Singapore, which lies almost on the equator and so experiences year-round hot temperatures, and much of its coastline is now seawall. To investigate the thermal ecology of a common intertidal gastropod, Nerita undata, on these artificial structures, we measured thermal conditions on two seawalls, the temperatures of habitats occupied by the snail, and compared these with the snail's thermal tolerance by measuring heart rate and behavioural thermoregulation (as preferred temperature, Tpref). At one of the two seawalls (Tanjong Rimau), temperatures experienced by N. undata exceeded all measures of thermal tolerance in the sun, while at the other (Palawan Beach), they did not. Temperatures in habitats occupied by the snails on the seawalls were similar to their measured Tpref in the laboratory and were lower than all measures of thermal tolerance. Behavioural thermoregulation by the snails, therefore, significantly increased the thermal safety margins of N. undata on the relatively homogenous seawalls in Singapore, and at one of the two seawalls were necessary to allow snails to survive. Accordingly, to facilitate motile species to maintain broad thermal safety margins through behavioural regulation, the provision of additional refuges from thermal stress is recommended on artificial coastal defences such as seawalls.

Bacterial biofilm colonization and succession in tropical marine waters are similar across different types of stone materials used in seawall construction.

Seawalls are important in protecting coastlines from currents, erosion, sea-level rise, and flooding. They are, however, associated with reduced biodiversity, due to their steep orientation, lack of microhabitats, and the materials used in their construction. Hence, there is considerable interest in modifying seawalls to enhance the settlement and diversity of marine organisms, as microbial biofilms play a critical role facilitating algal and invertebrate colonization. We assessed how different stone materials, ranging from aluminosilicates to limestone and concrete, affect biofilm formation. Metagenomic assessment of marine microbial communities indicated no significant impact of material on microbial diversity, irrespective of the diverse surface chemistry and topography. Based on KEGG pathway analysis, surface properties appeared to influence the community composition and function during the initial stages of biofilm development, but this effect disappeared by Day 31. We conclude that marine biofilms converged over time to a generic marine biofilm, rather than the underlying stone substrata type playing a significant role in driving community composition.

Spatio-temporal dynamics of coral-macroalgal interactions and their impacts on coral growth on urbanised reefs.

Interactions between corals and macroalgae are important in influencing benthic community structures on coral reefs and have become increasingly common occurrences. However, little is known about their temporal variation as most studies have only documented them from single surveys. To investigate the dynamics of coral-macroalgal interactions, we surveyed three urbanised reefs in Singapore bi-monthly for three years. We found that the frequency of coral-macroalgal interactions varied greatly across sites and seasons. The extent of coral-macroalgal contact was positively correlated with macroalgal abundance, but the correlation differed significantly among macroalgal genera. The growth rates of Goniopora, Montipora and Pavona corals, but not Platygra, were also negatively correlated with the extent of macroalgal interactions. Overall, our results highlight that coral-macroalgal interactions are spatially and temporally dynamic, with varying effects among coral species. It is critical to consider seasonal fluctuations of macroalgae if the overall long-term impacts of macroalgae are to be understood.

Antagonistic effects of seawalls and urban sedimentation on epilithic algal matrix (EAM)-feeding fishes.

Marine urbanisation often results in the proliferation of artificial coastal defences and heavy sedimentation, adversely impacting coral reef systems in tropical coastal cities. Knowledge of how motile organisms, such as reef fish, respond to novel human-made habitats and high sedimentation is limited. Here, we examine the role of sloping granite seawalls in supporting reef fishes that utilise the epilithic algal matrix (EAM) as a food resource. We surveyed fish assemblages and feeding activities on seawalls and reef flats, and conducted a field experiment to examine the effects of sediment on EAM feeding rates. Seawalls and reef flats supported distinct fish assemblage composition with significantly greater feeding activity on seawalls. However, reduced feeding activity on EAM with elevated sediment loads suggests that urban sedimentation may limit the utility of this novel feeding ground for nearshore communities. These findings illustrate the complexities and interactive effects of anthropogenic changes driven by coastal urbanisation.

Among-genotype responses of the coral Pocillopora acuta to emersion: implications for the ecological engineering of artificial coastal defences.

Stony corals are promising transplant candidates for the ecological engineering of artificial coastal defences such as seawalls as they attract and host numerous other organisms. However, seawalls are exposed to a wide range of environmental stressors associated with periods of emersion during low tide such as desiccation and changes in salinity, temperature, and solar irradiance. All of these variables have known deleterious effects on coral physiology, growth, and fitness. In this study, we performed parallel experiments (in situ and ex situ) to examine among-genotype responses of Pocillopora acuta to emersion by quantifying growth, photophysiological metrics (Fv/Fm, non-photochemical quenching [NPQ], endosymbiont density, and chlorophyll [chl] a concentration) and survival, following different emersion periods. Results showed that coral fragments emersed for longer durations (>2 h) exhibited reduced growth and survival. Endosymbiont density and NPQ, but not Fv/Fm and chl a concentration, varied significantly among genotypes across different durations of emersion. Overall, the ability of P. acuta to tolerate emersion for up to 2 h suggests its potential to serve as a 'starter species' for transplantation efforts on seawalls. Further, careful characterisation and selection of genotypes with a high capacity to withstand emersion can help maximise the efficacy of ecological engineering using coral transplants.

Growth and carbonate production of crustose coralline algae on a degraded turbid reef system.

Crustose coralline algae (CCA) and other encrusting calcifiers drive carbonate production on coral reefs. However, little is known about the rates of growth and calcification of these organisms within degraded turbid reef systems. Here we deployed settlement cards (N = 764) across seven reefs in Singapore for two years to examine spatio-temporal variation in encrusting community composition and CCA carbonate production. Our results showed that CCA was the dominant encrusting taxa (63.7% ± 18.3SD) across reefs. CCA carbonate production rates (0.009-0.052 g cm-2 yr-1) were less than half of those reported for most Indo-Pacific reefs, but similar to other turbid reef systems. Highest CCA carbonate production rates were observed furthest from Singapore's main shipping port, due to a relative increase in CCA cover on the offshore reefs. Our results suggest that proximity to areas of high industrialisation and ship traffic may reduce the cover of encrusting calcifying organisms and CCA production rates which may have negative, long-term implications for the stabilisation of nearshore reefs in urbanised settings.

Body Wave Generation for Anguilliform Locomotion Using a Fiber-Reinforced Soft Fluidic Elastomer Actuator Array Toward the Development of the Eel-Inspired Underwater Soft Robot.

In the field of marine biology-inspired robotics, anguilliform locomotion, as one of the most common underwater propulsion types, has been widely studied and implemented in many robot prototypes. Most of these robots consisted of rigid parts, and they were able to generate smooth sine waves along the robot body using a great number of rigid segments, electric motors, and complicated control. To simplify the robot structures and improve body compliance, anguilliform robots with a full soft body are highly desirable for better biomimicry. In this article, we propose a serial soft-actuator array consisting of four fiber-reinforced, bidirectionally bending, fluidic elastomer actuators (FEAs) to achieve the generation of anguilliform body waves. The FEA is fabricated using the typical soft lithography method with dual-chamber configuration and fiber reinforcement. The bending performance of the single FEA in free space with static loads was measured and compared with the three-dimensional simulation results using the finite element method. After that, the FEA array was assembled using the through-chamber tube connection and tested in water to investigate the bending performance with dynamic loads. Finally, with coordinated control of each segment, this FEA array achieved the generation of anguilliform body waves with different frequencies and amplitudes, which unveils a novel and promising approach to develop an anguilliform underwater robot. By attaching this FEA array to a miniaturized stand-alone station for control and actuation, an untethered swimming robot was built and successfully propelled by the FEA array, which demonstrates the capabilities and potentials of this soft and slender robot body design.

Population dynamics and growth rates of free-living mushroom corals (Scleractinia: Fungiidae) in the sediment-stressed reefs of Singapore.

The reefs of Singapore provide an excellent opportunity to study the population dynamics and growth rates of free-living mushroom corals (Fungiidae) under sediment-stressed conditions. Transect surveys at four study sites revealed a total of 11 free-living mushroom coral species-the same 11 species as those found by local studies since the 1980s. The abundance of the four most common species ranged from 1.0 to 68.3 Ind. per 100m2, while their population size-structure showed a common pattern of a higher proportion of small-sized corals than large-sized ones (i.e. positively skewed size-structure), although very few individuals of the smallest-size classes were recorded for any of the four species. A more positively skewed size-structure for each of the four most common species was observed at the reef slope (5-6m depth) than at the reef crest (2-3m depth), possibly due to a slower growth rate caused by light reduction with depth. All the mushroom corals studied exhibited a decline in growth rate with increasing size and weight, indicating determinate growth. Growth rate of each of the four most common species was similar among the study sites, despite variation in environmental conditions. Our results demonstrate species richness stability over the past three decades, suggesting that these free-living mushroom coral assemblages comprise species that are well-adapted to the chronic high sedimentation characteristic of Singapore's reefs. However, if the paucity of individuals of the smallest-size classes reflects poor recruitment and/or early mortality, there may be some cause for concern. Our robust baseline data can contribute to a long-term monitoring strategy for determination of changes in mushroom coral population dynamics.

Effects of macroalgae on coral fecundity in a degraded coral reef system.

Global declines in live coral cover and the proliferation of macroalgae on coral reefs is leading to increased coral-macroalgal interactions that impact reef recovery. However, the effects of macroalgae on coral sexual reproduction-a fundamental life-history process for maintaining population abundances-have rarely been quantified. Here, we examined the direct effects of macroalgae contact on the fecundity (eggs mesentery-1) of two coral species, Echinopora lamellosa and Merulina ampliata, across three degraded reefs in Singapore. Increasing macroalgae contact from 5% to 25% significantly reduced fecundity in colonies of both species by 67-82%, and also reduced M. ampliata egg sizes by 11.4%. These results suggest the diversion of energy from reproduction towards other processes such as repair and defence, and also reveal potential differential energy allocation strategies among coral taxa. While corals on Singapore's impacted reefs continue to produce eggs, increasing macroalgae that suppresses coral fecundity may constrain future reef recovery.