Coral settlement on a highly disturbed equatorial reef system.
ABSTRACT: Processes occurring early in the life stages of corals can greatly influence the demography of coral populations, and successful settlement of coral larvae that leads to recruitment is a critical life history stage for coral reef ecosystems. Although corals in Singapore persist in one the world's most anthropogenically impacted reef systems, our understanding of the role of coral settlement in the persistence of coral communities in Singapore remains limited. Spatial and temporal patterns of coral settlement were examined at 7 sites in the southern islands of Singapore, using settlement tiles deployed and collected every 3 months from 2011 to 2013. Settlement occurred year round, but varied significantly across time and space. Annual coral settlement was low (~54.72 spat m(-2) yr(-1)) relative to other equatorial regions, but there was evidence of temporal variation in settlement rates. Peak settlement occurred between March-May and September-November, coinciding with annual coral spawning periods (March-April and October), while the lowest settlement occurred from December-February during the northeast monsoon. A period of high settlement was also observed between June and August in the first year (2011/12), possibly due to some species spawning outside predicted spawning periods, larvae settling from other locations or extended larval settlement competency periods. Settlement rates varied significantly among sites, but spatial variation was relatively consistent between years, suggesting the strong effects of local coral assemblages or environmental conditions. Pocilloporidae were the most abundant coral spat (83.6%), while Poritidae comprised only 6% of the spat, and Acroporidae <1%. Other, unidentifiable families represented 10% of the coral spat. These results indicate that current settlement patterns are reinforcing the local adult assemblage structure ('others'; i.e. sediment-tolerant coral taxa) in Singapore, but that the replenishment capacity of Singapore's reefs appears relatively constrained, which could lead to less resilient reefs.
Project description:Tropical reefs are dynamic ecosystems that host diverse coral assemblages with different life-history strategies. Here, we quantified how juvenile (<50 mm) coral demographics influenced benthic coral structure in reef flat and reef slope habitats on the southern Great Barrier Reef, Australia. Permanent plots and settlement tiles were monitored every six months for three years in each habitat. These environments exhibited profound differences: the reef slope was characterised by 95% less macroalgal cover, and twice the amount of available settlement substrata and rates of coral settlement than the reef flat. Consequently, post-settlement coral survival in the reef slope was substantially higher than that of the reef flat, and resulted in a rapid increase in coral cover from 7 to 31% in 2.5 years. In contrast, coral cover on the reef flat remained low (~10%), whereas macroalgal cover increased from 23 to 45%. A positive stock-recruitment relationship was found in brooding corals in both habitats; however, brooding corals were not directly responsible for the observed changes in coral cover. Rather, the rapid increase on the reef slope resulted from high abundances of broadcast spawning Acropora recruits. Incorporating our results into transition matrix models demonstrated that most corals escape mortality once they exceed 50 mm, but for smaller corals mortality in brooders was double those of spawners (i.e. acroporids and massive corals). For corals on the reef flat, sensitivity analysis demonstrated that growth and mortality of larger juveniles (21-50 mm) highly influenced population dynamics; whereas the recruitment, growth and mortality of smaller corals (<20 mm) had the highest influence on reef slope population dynamics. Our results provide insight into the population dynamics and recovery trajectories in disparate reef habitats, and highlight the importance of acroporid recruitment in driving rapid increases in coral cover following large-scale perturbation in reef slope environments.
Project description:Ocean acidification (OA) refers to the ongoing decline in oceanic pH resulting from the uptake of atmospheric CO(2). Mounting experimental evidence suggests that OA will have negative consequences for a variety of marine organisms. Whereas the effect of OA on the calcification of adult reef corals is increasingly well documented, effects on early life history stages are largely unknown. Coral recruitment, which necessitates successful fertilization, larval settlement, and postsettlement growth and survivorship, is critical to the persistence and resilience of coral reefs. To determine whether OA threatens successful sexual recruitment of reef-building corals, we tested fertilization, settlement, and postsettlement growth of Acropora palmata at pCO(2) levels that represent average ambient conditions during coral spawning (?400 ?atm) and the range of pCO(2) increases that are expected to occur in this century [?560 ?atm (mid-CO(2)) and ?800 ?atm (high-CO(2))]. Fertilization, settlement, and growth were all negatively impacted by increasing pCO(2), and impairment of fertilization was exacerbated at lower sperm concentrations. The cumulative impact of OA on fertilization and settlement success is an estimated 52% and 73% reduction in the number of larval settlers on the reef under pCO(2) conditions projected for the middle and the end of this century, respectively. Additional declines of 39% (mid-CO(2)) and 50% (high-CO(2)) were observed in postsettlement linear extension rates relative to controls. These results suggest that OA has the potential to impact multiple, sequential early life history stages, thereby severely compromising sexual recruitment and the ability of coral reefs to recover from disturbance.
Project description:<h4>Background</h4>The transition from planktonic planula to sessile adult corals occurs at low frequencies and post settlement mortality is extremely high. Herbivores promote settlement by reducing algal competition. This study investigates whether invertebrate herbivory might be modulated by other ecological factors such as substrata variations and coral species identity.<h4>Methodology/principal findings</h4>The experiment was conducted at the Flower Garden Banks, one of the few Atlantic reefs not experiencing considerable degradation. Tiles of differing texture and orientation were kept in bins surrounded by reef (24 m). Controls contained no herbivores while treatment bins contained urchins (Diadema antillarum) or herbivorous gastropods (Cerithium litteratum). Juvenile corals settling naturally were monitored by photography for 14 months to evaluate the effects of invertebrate herbivory and substratum properties. Herbivory reduced algae cover in urchin treatments. Two genera of brooding coral juveniles were observed, Agaricia and Porites, both of which are common but not dominant on adjacent reef. No broadcast spawning corals were observed on tiles. Overall, juveniles were more abundant in urchin treatments and on vertical, rough textured surfaces. Although more abundant, Agaricia juveniles were smaller in urchin treatments, presumably due to destructive overgrazing. Still, Agaricia growth increased with herbivory and substrata texture-orientation interactions were observed with reduced growth on rough tiles in control treatments and increased growth on vertical tiles in herbivore treatments. In contrast to Agaricia, Porites juveniles were larger on horizontal tiles, irrespective of herbivore treatment. Mortality was affected by substrata orientation with vertical surfaces increasing coral survival.<h4>Conclusions/significance</h4>We further substantiate that invertebrate herbivores play major roles in early settlement processes of corals and highlight the need for deeper understanding of ecological interactions modulating these effects. The absence of broadcast-spawning corals, even on a reef with consistently high coral cover, continues to expose the recruitment failure of these reef-building corals throughout the Caribbean.
Project description:Larval production and recruitment underpin the maintenance of coral populations, but these early life history stages are vulnerable to extreme variation in physical conditions. Environmental managers aim to minimise human impacts during significant periods of larval production and recruitment on reefs, but doing so requires knowledge of the modes and timing of coral reproduction. Most corals are hermaphroditic or gonochoric, with a brooding or broadcast spawning mode of reproduction. Brooding corals are a significant component of some reefs and produce larvae over consecutive months. Broadcast spawning corals are more common and display considerable variation in their patterns of spawning among reefs. Highly synchronous spawning can occur on reefs around Australia, particularly on the Great Barrier Reef. On Australia's remote north-west coast there have been fewer studies of coral reproduction. The recent industrial expansion into these regions has facilitated research, but the associated data are often contained within confidential reports. Here we combine information in this grey-literature with that available publicly to update our knowledge of coral reproduction in WA, for tens of thousands of corals and hundreds of species from over a dozen reefs spanning 20° of latitude. We identified broad patterns in coral reproduction, but more detailed insights were hindered by biased sampling; most studies focused on species of Acropora sampled over a few months at several reefs. Within the existing data, there was a latitudinal gradient in spawning activity among seasons, with mass spawning during autumn occurring on all reefs (but the temperate south-west). Participation in a smaller, multi-specific spawning during spring decreased from approximately one quarter of corals on the Kimberley Oceanic reefs to little participation at Ningaloo. Within these seasons, spawning was concentrated in March and/or April, and October and/or November, depending on the timing of the full moon. The timing of the full moon determined whether spawning was split over two months, which was common on tropical reefs. There were few data available for non-Acropora corals, which may have different patterns of reproduction. For example, the massive Porites seemed to spawn through spring to autumn on Kimberley Oceanic reefs and during summer in the Pilbara region, where other common corals (e.g. Turbinaria & Pavona) also displayed different patterns of reproduction to the Acropora. The brooding corals (Isopora & Seriatopora) on Kimberley Oceanic reefs appeared to planulate during many months, possibly with peaks from spring to autumn; a similar pattern is likely on other WA reefs. Gaps in knowledge were also due to the difficulty in identifying species and issues with methodology. We briefly discuss some of these issues and suggest an approach to quantifying variation in reproductive output throughout a year.
Project description:Following disturbances, corals recolonize space through the process of recruitment consisting of the three phases of propagule supply, settlement, and post-settlement survival. Yet, each phase is influenced by biophysical factors, leading to recruitment success variability through space. To resolve the relative contributions of biophysical factors on coral recruitment, the recovery of a 150?km long coral reefs in Palau was investigated after severe typhoon disturbances. Overall, we found that benthic organisms had a relatively weak interactive influence on larval settlement rates at the scale of individual tiles, with negative effects mainly exerted from high wave exposure for Acropora corals. In contrast, juvenile coral densities were well predicted by biophysical drivers, through both direct and indirect pathways. High densities of Acropora and Poritidae juveniles were directly explained by the availability of substrata free from space competitors. Juvenile Montipora were found in higher densities where coralline algae coverage was high, which occurred at reefs with high wave exposure, while high densities of juvenile Pocilloporidae occurred on structurally complex reefs with high biomass of bioeroder fish. Our findings demonstrate that strengths of biophysical interactions were taxon-specific and had cascading effects on coral recruitment, which need consideration for predicting reef recovery and conservation strategies.
Project description:Coral reefs in Moorea, French Polynesia, suffered catastrophic coral mortality through predation by Acanthaster planci from 2006 to 2010, and Cyclone Oli in 2010, yet by 2015 some coral populations were approaching pre-disturbance sizes. Using long-term study plots, we quantified population dynamics of spawning Pocillopora spp. along the north shore of Moorea between 2010 and 2014, and considered evidence that population recovery could be supported by self-seeding. Results scaled up from study plots and settlement tiles suggest that the number of Pocillopora spp. colonies on the outer reef increased 1,890-fold between 2010 and 2014/2015, and in the back reef, 8-fold between 2010 and 2014/2015. Assuming that spawning Pocillopora spp. in Moorea release similar numbers of eggs as con-generics in Hawaii, and fertilization success is similar to other spawning corals, the capacity of Pocillopora spp. to produce larvae was estimated. These estimates suggest that Pocillopora spp. in Moorea produced a large excess of larvae in 2010 and 2014 relative to the number required to produce the recruits found in the back reef and outer reef in 2010 and 2014, even assuming that ?99.9% of the larvae do not recruit in Moorea. Less than a third of the recruits in one year would have to survive to produce the juvenile Pocillopora spp. found in the back and outer reefs in 2010 and 2014/2015. Our first order approximations reveal the potential for Pocillopora spp. on the north shore of Moorea to produce enough larvae to support local recruitment and population recovery following a catastrophic disturbance.
Project description:Coral endosymbionts in the dinoflagellate genus Symbiodinium are known to impact host physiology and have led to the evolution of reef-building, but less is known about how symbiotic communities in early life-history stages and their interactions with host parental identity shape the structure of coral communities on reefs. Differentiating the roles of environmental and biological factors driving variation in population demographic processes, particularly larval settlement, early juvenile survival and the onset of symbiosis is key to understanding how coral communities are structured and to predicting how they are likely to respond to climate change. We show that maternal effects (that here include genetic and/or effects related to the maternal environment) can explain nearly 24% of variation in larval settlement success and 5-17% of variation in juvenile survival in an experimental study of the reef-building scleractinian coral, Acropora tenuis. After 25 days on the reef, Symbiodinium communities associated with juvenile corals differed significantly between high mortality and low mortality families based on estimates of taxonomic richness, composition and relative abundance of taxa. Our results highlight that maternal and familial effects significantly explain variation in juvenile survival and symbiont communities in a broadcast-spawning coral, with Symbiodinium type A3 possibly a critical symbiotic partner during this early life stage.
Project description:For many corals, the timing of broadcast spawning correlates strongly with a number of environmental signals (seasonal temperature, lunar, and diel cycles). Robust experimental studies examining the role of these putative cues in triggering spawning have been lacking until recently because it has not been possible to predictably induce spawning in fully closed artificial mesocosms. Here, we present a closed system mesocosm aquarium design that utilizes microprocessor technology to accurately replicate environmental conditions, including photoperiod, seasonal insolation, lunar cycles, and seasonal temperature from Singapore and the Great Barrier Reef (GBR), Australia. Coupled with appropriate coral husbandry, these mesocosms were successful in inducing, for the first time, broadcast coral spawning in a fully closed artificial ex situ environment. Four Acropora species (A. hyacinthus, A. tenuis, A. millepora, and A. microclados) from two geographical locations, kept for over 1 year, completed full gametogenic cycles ex situ. The percentage of colonies developing oocytes varied from ~29% for A. hyacinthus to 100% for A. millepora and A. microclados. Within the Singapore mesocosm, A. hyacinthus exhibited the closest synchronization to wild spawning, with all four gravid colonies releasing gametes in the same lunar month as wild predicted dates. Spawning within the GBR mesocosm commenced at the predicted wild spawn date but extended over a period of 3 months. Gamete release in relation to the time postsunset for A. hyacinthus, A. millepora, and A. tenuis was consistent with time windows previously described in the wild. Spawn date in relation to full moon, however, was delayed in all species, possibly as a result of external light pollution. The system described here could broaden the number of institutions on a global scale, that can access material for broadcast coral spawning research, providing opportunities for institutions distant from coral reefs to produce large numbers of coral larvae and juveniles for research purposes and reef restoration efforts.
Project description:Recovery of coral reefs after disturbance relies heavily on replenishment through successful larval settlement and their subsequent survival. As part of an integrated study to determine the potential effects of water quality changes on the resilience of inshore coral communities, scleractinian coral settlement was monitored between 2006 and 2012 at 12 reefs within the inshore Great Barrier Reef. Settlement patterns were only analysed for the family Acroporidae, which represented the majority (84%) of settled larvae. Settlement of Acroporidae to terracotta tiles averaged 0.11 cm-2, representing 34 ± 31.01 (mean ± SD) spat per tile, indicating an abundant supply of competent larvae to the study reefs. Settlement was highly variable among reefs and between years. Differences in settlement among locations partly corresponded to the local cover of adult Acroporidae, while substantial reductions in Acroporidae cover caused by tropical cyclones and floods resulted in a clear reduction in settlement. Much of the observed variability remained unexplained, although likely included variability in both connectivity to, and the fecundity of, adult Acroporidae. The responsiveness of settlement patterns to the decline in Acroporidae cover across all four regions indicates the importance of supply and connectivity, and the vulnerability towards region-wide disturbance. High spatial and temporal variability, in addition to the resource-intensive nature of sampling with settlement tiles, highlights the logistical difficulty of determining coral settlement over large spatial and temporal scales.
Project description:In the Central Red Sea, healthy coral reefs meet intense coastal development, but data on the effects of related stressors for reef functioning are lacking. This in situ study therefore investigated the independent and combined effects of simulated overfishing through predator/grazer exclusion and simulated eutrophication through fertilizer addition on settlement of reef associated invertebrates on light-exposed and -shaded tiles over 4 months. At the end of the study period invertebrates had almost exclusively colonized shaded tiles. Algae were superior settling competitors on light-exposed tiles. On the shaded tiles, simulated overfishing prevented settlement of hard corals, but significantly increased settlement of polychaetes, while simulated eutrophication only significantly decreased hard coral settlement relative to controls. The combined treatment significantly increased settlement of bryozoans and bivalves compared to controls and individual manipulations, but significantly decreased polychaetes compared to simulated overfishing. These results suggest settlement of polychaetes and hard corals as potential bioindicators for overfishing and eutrophication, respectively, and settlement of bivalves and bryozoans for a combination of both. Therefore, if the investigated stressors are not controlled, phase shifts from dominance by hard corals to that by other invertebrates may occur at shaded reef locations in the Central Red Sea.