Patterns in the distribution of soft corals across the central Great Barrier Reef

Distribution patterns of soft coral genera were examined at 11 reefs situated in a broad transect from inshore to the Coral Sea in the central region of the Great Barrier Reef. Twenty-five genera representing the Orders Alcyonacea and Stolonifera were recorded, and the survey also included one genus of the Order Gorgonacea. Total living soft coral cover is greatest on outershelf reef slopes, and is often less than and inversely related to the cover by stony corals. Soft coral diversity is generally low on reef flats, where soft coral cover is low or nil except in protected, inshore areas. The most diverse assemblages occur on reef slopes in midshelf and outershelf areas, where Efflatounaria and nephtheid genera predominate, and widely distributed alcyoniid genera are common. These richer assemblages are less well represented in the Coral Sea, while innershelf reefs support a less diverse fauna of somewhat different generic composition. Distribution patterns of soft corals across the transect broadly match similar variations in the distributions of stony corals and fishes, inshore reefs being generally depauperate. Such variations across the continental shelf are closely associated with changes in prevailing environmental conditions, but further research will be required to elucidate the effects of environmental parameters on benthic community structure.     

Community structure of stony and soft corals on vertical unplanned artificial reefs in Eilat (Red Sea): comparison to natural reefs

In many reef ecosystems, artificial reefs (AR) have become permanent additions to the area, sustaining well-developed benthic communities. Long-term studies on the development of AR coral communities are scarce, and comparisons with their natural surroundings are limited. The present study describes the stony and soft coral community structure of unplanned vertical AR in Eilat (Red Sea) that have progressed beyond the initial successional phases, and compares these to the adjacent natural reefs (NR). Coral communities were characterized using belt transects, conducted on 34- and 14-year-old unplanned AR, and on two proximate NR. Stony corals were the major component in the NR, while soft corals, mainly Nephtheidae, accounted for up to 90% of the total living coverage in the AR. This was attributed to physical and biological features associated with the ARs vertical orientation, which was absent in the NR, and to the life history traits of these soft corals. Community differences between the two AR were related to structural stability and age. The results suggest that AR may increase local heterogeneity and space availability by adding novel habitats, increasing production and elevating species diversity in the surroundings.Communicated by Biological Editor H.R. Lasker     

Halimeda growth and diversity on the deep fore-reef of Enewetak Atoll

The deep fore-reef at Enewetak has been examined from the submersible Makali'i. Green algae grow to about-150 m at photon flux densities of approximately 1 Em^-2s^-1. Halimeda cover is 50% at many sites down to-90 m. Halimeda populations are important within the zone of scleractinian corals down to about-65 m, while a Halimeda zone with low coral cover or lacking corals between-65 m and-150 m probably is an important source of reef carbonate. Halimedas of the deep fore-reef, like those of the lagoon, constitute an important structural component in reef building. Other calcareous green algae such as Tydemania are less important on the deep fore-reef, but growth of coralline red algae continues to over-200m. Halimeda diversity is high down to near the base of the euphotic zone.     

Effects of fish predation and seaweed competition on the survival and growth of corals

On Caribbean coral reefs, high rates of grazing by herbivorous fishes are thought to benefit corals because fishes consume competing seaweeds. We conducted field experiments in the Florida Keys, USA, to examine the effects of grazing fishes on coral/seaweed competition. Initially, fragments of Porites divaracata from an inshore habitat were transplanted into full-cage, half-cage, and no-cage treatments on a fore-reef. Within 48 h, 56% of the unprotected corals in half-cage and no-cage treatments (62 of 111) were completely consumed. Stoplight parrotfish (Sparisoma viride) were the major coral predators, with redband parrotfish (S. aurofrenatum) also commonly attacking this coral. Next, we transplanted fragments of P. porites collected from the fore-reef habitat where our caging experiments were being conducted into the three cage treatments, half in the presence of transplanted seaweeds, and half onto initially clean substrates. The corals were allowed to grow in these conditions, with concurrent development of competing seaweeds, for 14 weeks. Although seaweed cover and biomass were both significantly greater in the full-cage treatment, coral growth did not differ significantly between cage treatments even though corals placed with pre-planted seaweeds grew significantly less than corals placed on initially clean substrate. This surprising result occurred because parrotfishes not only grazed algae from accessible treatments, but also fed directly on our coral transplants. Parrotfish feeding scars were significantly more abundant on P. porites from the half and no-cage treatments than on corals in the full cages. On this Florida reef, direct fish predation on some coral species (P. divaracata) can exclude them from fore-reef areas, as has previously been shown for certain seaweeds and sponges. For other corals that live on the fore-reef (P. porites), the benefits of fishes removing seaweeds can be counterbalanced by the detrimental effects of fishes directly consuming corals. Received: 31 May 1997 / Accepted: 2 September 1997     

Competition between corals and algal turfs along a gradient of terrestrial influence in the nearshore central Great Barrier Reef

Competition between benthic algae and corals is a key process in the community ecology of reefs, especially during reef degradation. However, there have been very few experimental tests for competition between corals and benthic algae, despite widespread assumptions that algae are generally superior competitors, especially in eutrophic conditions. This study tested for competition for space between the massive coral Porites lobata and algal filamentous turfs on three reefs along a cross-shelf gradient of terrestrial influence, by experimentally removing or damaging either corals or algae. The corals and algae were competing for space, but, significantly, the algae appeared to have little effect on coral growth. In contrast, corals significantly inhibited algal growth, suggesting Porites was the competitive superior. Importantly, coral growth was generally positive, even on the reef with the greatest terrestrial influence. Competitive outcomes did not support the argument that algae are more successful competitors in more eutrophic conditions.     

Influence of the coral reef assemblages on the spatial distribution of echinoderms in a gradient of human impacts along the tropical Mexican Pacific

Fourteen species of echinoderms and their relationships to the benthic structure of the coral reefs were assessed at 27 sites—with different levels of human disturbances—along the coast of the Mexican Central Pacific. Diadema mexicanum and Phataria unifascialis were the most abundant species. The spatial variation of the echinoderm assemblages showed that D. mexicanum, Eucidaris thouarsii, P. unifascialis, Centrostephanus coronatus, Toxopneustes roseus, Holothuria fuscocinerea, Cucumaria flamma, and Echinometra vanbrunti accounted for the dissimilarities among the sites. The spatial variation among the sites was mainly explained by the cover of the hard corals (Porites, Pocillopora, Pavona, Psammocora), different macroalgae species (turf, encrusting calcareous algae, articulated calcareous algae, fleshy macroalgae), sponges, bryozoans, rocky, coral rubble, sand, soft corals (hydrocorals and octocorals), Tubastrea coccinea coral, Balanus spp., and water depth. The coverage of Porites, Pavona, and Pocillopora corals, soft coral, rock, and Balanos shows a positive relationship with the sampling sites included within the natural protected area with low human disturbances. Contrary, fleshy macroalgae, sponges, and soft coral show a positive relationship with higher disturbance sites. The results presented here show the importance of protecting the structural heterogeneity of coral reef habitats because it is a significant factor for the distribution of echinoderm species and can contribute to the design of conservation programs for the coral reef ecosystem.     

Book review of Littler DM. Littler MM (2000) Caribbean Reef Plants An Identification Guide to the Reef Plants of the Caribbean,Bahamas, Florida and Gulf of Mexico

Information on spatial variability and distribution patterns of organisms in coral reef environments is necessary to evaluate the increasing anthropogenic disturbance of marine environments (Richmond 1993; Wilkinson 1993; Dayton 1994). Therefore different types of subtidal, reef-associated hard substrata (reef flats, reef slopes, coral carpets, coral patches, rock grounds), each with different coral associations, were investigated to determine the distribution pattern of molluscs and their life habits (feeding strategies and substrate relations). The molluscs were strongly dominated by taxa with distinct relations to corals, and five assemblages were differentiated. The Dendropoma maxima assemblage on reef flats is a discrete entity, strongly dominated by this encrusting and suspension-feeding gastropod. All other assemblages are arranged along a substrate gradient of changing coral associations and potential molluscan habitats. The Coralliophila neritoidea–Barbatia foliata assemblage depends on the presence of Porites and shows a dominance of gastropods feeding on corals and of bivalves associated with living corals. The Chamoidea–Cerithium spp. assemblage on rock grounds is strongly dominated by encrusting bivalves. The Drupella cornus–Pteriidae assemblage occurs on Millepora–Acropora reef slopes and is strongly dominated by bivalves associated with living corals. The Barbatia setigera–Ctenoides annulata assemblage includes a broad variety of taxa, molluscan life habits and bottom types, but occurs mainly on faviid carpets and is transitional among the other three assemblages. A predicted degradation of coral coverage to rock bottoms due to increasing eutrophication and physical damage in the study area (Riegl and Piller 2000) will result in a loss of coral-associated molluscs in favor of bivalve crevice dwellers in dead coral heads and of encrusters on dead hard substrata.     

Long-term persistence of low coral cover and abundance on a disturbed coral reef flat in the northern Red Sea

Levels of coral cover and abundance on a coral reef flat in Eilat (Israeli Red Sea) were estimated in 2001 by surveying nineteen 10-m transects, and compared to the levels reported in the same area between 1966 and 1973. Lower values compared to 1966 levels are evident, and there has been only a modest recovery following a catastrophic low tide that killed a large proportion of the corals in 1970. Percent cover of soft and stony corals (16.1%) was less than half of that reported for 1969 (35%), when a sharp decrease in coral abundance had already been observed. The total number of soft and stony coral colonies observed was 300, compared to 541 in 1966. In contrast to 1966, when half of the transects surveyed contained more than 30 coral colonies, no transects with this number of corals were observed. The cover of seven of the most common stony coral species was 841 cm, which is twice the coral cover of that in 1973, but only 22% of the 1969 level. Millepora dichotoma, an abundant species before 1970, has almost disappeared, and the soft coral Litophyton, abundant in 1972, was not observed. Anthropogenic nutrient enrichment is apparently among the causes for the lack of coral recovery in the studied reef flat. Reefs located further away from sources of pollution have recovered quickly after natural and anthropogenic disturbances and have retained their coral abundance and diversity.     

Caribbean corals exhibit species-specific differences in competitive abilities with an aggressive encrusting alga,Ramicrusta textilis

Caribbean coral cover has decreased substantially in recent decades, with much of the live coral being replaced by macroalgae. Encrusting red algae in the genus Ramicrusta have become abundant throughout the region and have demonstrated widespread harm to corals by overgrowing living tissue, causing colony mortality, and impairing coral recruitment. In this research, Ramicrusta textilis was identified by morpho-anatomy and DNA sequencing from nine sites around St. Thomas, US Virgin Islands, and 3D photogrammetry was used to measure the rate of algal growth on stony corals. 3D models of individual coral colonies (five species plus controls, N = 72) competing with R. textilis revealed differential competitive abilities among taxa, with Siderastrea siderea being the only species capable of inhibiting overgrowth by the alga (mean linear algal growth − 1.1 mm yr⁻¹). Important reef building coral species such as Orbicella annularis and Orbicella faveolata were poor competitors (mean linear algal growth + 15 mm yr⁻¹ and + 7.7 mm yr⁻¹, respectively), indicating that the emergence of the alga could have significant impacts on Caribbean coral reef species diversity, community composition, and structural complexity.

     

Differential thermal tolerance between algae and corals may trigger the proliferation of algae in coral reefs

Marine heatwaves can lead to rapid changes in entire communities, including in the case of shallow coral reefs the potential overgrowth of algae. Here we tested experimentally the differential thermal tolerance between algae and coral species from the Red Sea through the measurement of thermal performance curves and the assessment of thermal limits. Differences across functional groups (algae vs. corals) were apparent for two key thermal performance metrics. First, two reef‐associated algae species (Halimeda tuna and Turbinaria ornata) had higher lethal thermal limits than two coral species (Pocillopora verrucosa and Stylophora pistillata) conferring those species of algae with a clear advantage during heatwaves by surpassing the thermal threshold of coral survival. Second, the coral species had generally greater deactivation energies for net and gross primary production rates compared to the algae species, indicating greater thermal sensitivity in corals once the optimum temperature is exceeded. Our field surveys in the Red Sea reefs before and after the marine heatwave of 2015 show a change in benthic cover mainly in the southern reefs, where there was a decrease in coral cover and a concomitant increase in algae abundance, mainly turf algae. Our laboratory and field observations indicate that a proliferation of algae might be expected on Red Sea coral reefs with future ocean warming.     

The effect of substratum on the growth of Terpios,an encrusting sponge which kills corals

The effects of substratum on the growth of Terpios was demonstrated using experimental and observational data at Guam, Mariana Islands. Terpios growth was measured on live coral, reef rock, and red calcareous algae in the field. In addition, Terpios was transplanted onto live coral, air-blasted (clean) coral, reef rock, and plexiglass plates, and subsequent growth measured. Terpios grows fastest on clean substrata followed by live coral, reef rock and red calcareous algae in decreasing order. Terpios is sometimes overgrown by Montipora, Porites and red calcareous algae. Since Terpios grows fastest when living coral tissue is removed, it is not likely that Terpios ingests coral tissue as previously suggested in the literature. Instead, Terpios is probably an efficient competitor of corals for space. Terpios overgrows most hard, stable reef substrata, and the growth rate on all sample substrata is substantial. Therefore Terpios has a great potential for covering a reef and may be one of the most important causes of disturbance on some coral reefs.Contribution no. 206 from the University of Guam Marine Laboratory     

Competition between corals and algae on coral reefs: a review of evidence and mechanisms

Despite widespread acceptance that competition between scleractinian corals and benthic algae is important to the structure of coral reef communities, there is little direct experimental evidence that corals and algae do compete, and very little data on the processes and causality of their interactions. Most available evidence is observational or correlative, with intrinsic risks of confounded causality. This paper reviews and categorises the available evidence, concluding that competition between corals and algae probably is widespread on coral reefs, but also that the interaction varies considerably. Widespread replacement of corals by algae may often indicate coral mortality due to external disturbances, rather than competitive overgrowth, but may lead to competitive inhibition of coral recruitment, with consequences for reef recovery. We list eight specific processes by which corals and algae may affect each other, and suggest life history properties that will influence which of these interactions are possible. We propose a matrix for algal effects on corals, which lists the subset of processes possible for each combination of coral life form and algal functional group. This table provides a preliminary framework for improved understanding and interpretation of coral-algal interactions.     

Upper Pleistocene cold-water corals from the Inner Sea of the Maldives: taphonomy and environment

Cold-water corals of the Late Pleistocene (21,400–22,500 BP) are recorded from the sea-bottom of two inter-atoll channels (Kardiva Channel at 457-m depth and Malé Vaadhoo Channel at 443-m depth) of the eastern row of the Maldives archipelago. Coral assemblages are composed mainly by Lophelia pertusa and secondarily by Madrepora oculata and Enallopsammia rostrata. These cold-water coral patches are places where the benthic life, mainly sessile, is concentrated, which is clearly absent off-rubble patches. The main epibionts are tube-dwelling polychaetes (mainly Spirorbis and Serpula), bryozoans, siliceous sponges, barnacles, gorgonids, solitary corals, encrusting foraminifera, and microbial mats. The analysis of epibionts assemblages shows different biocoenoses between both studied sites as well as a dependency of the epibiont coverage with regard to the coral genus. Some living benthic organisms such as brachiopods, bivalves, gastropods, barnacles, and ophiuroids find refuge among coral branches. The common record of juvenile specimens of vagile organisms such as small ophiuroids, is probably related to the nursery function of the cold-water corals in spite they are fossils. Environmental requirements of Recent cold-water corals (Lophelia, Madrepora and Enallopsammia) differ of conditions at both sampling sites with sensibly lower oxygenation degree and density of waters than needed for cold-water corals. Therefore, it is proposed that the present-day oxygen and density conditions are the factors which inhibit modern cold-water coral growth in the inter-atoll channels.     

Macroalgae reduces survival of nursery‐reared Acropora corals in the Florida reef tract

Recent declines in coral populations along the Florida reef tract have prompted the establishment of coral restoration programs which raise coral species, such as the threatened Acropora cervicornis, in nurseries ready for outplanting. Large numbers of nursery‐reared coral colonies have been outplanted along the Florida reef tract in recent years, yet few studies have characterized benthic habitats that are considered optimal for colony survival. In 2016, we surveyed 23 A. cervicornis restoration sites, located at six different reefs in the upper Florida Keys. We examined the condition of the outplanted corals and quantified the benthic assemblages adjacent to the outplanted coral colonies. We found that where A. cervicornis survived for more than 1 year, the substrate significantly supported less brown macroalgae of the genus Dictyota than at sites where A. cervicornis had died. Coral survival was highest at sites with less than 15% Dictyota cover. These results suggest that the habitat conditions that supported Dictyota spp. were not conducive to A. cervicornis growth and survival. Restoration practitioners should avoid attaching nursery‐raised corals to substrate with Dictyota spp. cover greater than 15%.     

Carbonate production by scleractinian corals at Aqaba,Gulf of Aqaba,Red Sea

Summary In a fringing reef at Aqaba at the northern end of the Gulf of Aqaba (29°26′N) growth rates, density, and the calcification rate ofPorites were investigated in order to establish calculations of gross carbonate production for the reefs in this area. Colony accretion ofPorites decreases with depth as a function of decreasing growth rates. The calcification rate ofPorites is highest in shallow water (0–5 m depth) with 0.9 g·cm⁻²·yr⁻¹ and falls down to 0.5 g·cm⁻²·yr⁻¹ below 30 m. Scleractinian coral gross production is calculated from potential productivity and coral coverage. It is mainly dependent on living coral cover and to a lesser extent on potential productivity. Total carbonate production on the reef ranged from 0 to 2.7 kg/m² per year, with a reef-wide average of 1.6 kg/m² perycar. Maximum gross carbonate production by corals at Aqaba occurs at the reef crest and in the middle fore-reef from 10 to 15 m water depth. Production is low in sandy reef parts. Below 30 m depth values still reach ca. 50% of shallow water values. Mean potential production of colonies and gross carbonate production of the whole reef community at Aqaba is lower than in tropical reefs. However, carbonate production is higher than in reef areas at the same latitude in the Pacific, indicating a northward shift of reef production in the Red Sea.     

Seasonal monitoring of coral–algae interactions in fringing reefs of the Gulf of Aqaba, Northern Red Sea

This paper presents seasonal in situ monitoring data on benthic coverage and coral–algae interactions in high-latitude fringing reefs of the Northern Red Sea over a period of 19 months. More than 30% of all hermatypic corals were involved in interaction with benthic reef algae during winter compared to 17% during summer, but significant correlation between the occurrence of coral–algae interactions and monitored environmental factors such as temperature and inorganic nutrient availability was not detected. Between 5 and 10-m water depth, the macroalgae Caulerpa serrulata, Peyssonnelia capensis and filamentous turf algae represented almost 100% of the benthic algae involved in interaction with corals. Turf algae were most frequently (between 77 and 90% of all interactions) involved in interactions with hermatypic corals and caused most tissue damage to them. Maximum coral tissue loss of 0.75% day⁻¹ was observed for Acropora-turf algae interaction during fall, while an equilibrium between both groups of organisms appeared during summer. Slow-growing massive corals were more resistant against negative algal influence than fast-growing branching corals. Branching corals of the genus Acropora partly exhibited a newly observed phenotypic plasticity mechanism, by development of a bulge towards the competing organism, when in interaction with algae. These findings may contribute to understand the dynamics of phase shifts in coral reefs by providing seasonally resolved in situ monitoring data on the abundance and the competitive dynamic of coral–algae interactions.     

Re-evaluating the health of coral reef communities: baselines and evidence for human impacts across the central Pacific

Numerous studies have documented declines in the abundance of reef-building corals over the last several decades and in some but not all cases, phase shifts to dominance by macroalgae have occurred. These assessments, however, often ignore the remainder of the benthos and thus provide limited information on the present-day structure and function of coral reef communities. Here, using an unprecedentedly large dataset collected within the last 10 years across 56 islands spanning five archipelagos in the central Pacific, we examine how benthic reef communities differ in the presence and absence of human populations. Using islands as replicates, we examine whether benthic community structure is associated with human habitation within and among archipelagos and across latitude. While there was no evidence for coral to macroalgal phase shifts across our dataset we did find that the majority of reefs on inhabited islands were dominated by fleshy non-reef-building organisms (turf algae, fleshy macroalgae and non-calcifying invertebrates). By contrast, benthic communities from uninhabited islands were more variable but in general supported more calcifiers and active reef builders (stony corals and crustose coralline algae). Our results suggest that cumulative human impacts across the central Pacific may be causing a reduction in the abundance of reef builders resulting in island scale phase shifts to dominance by fleshy organisms.     

Detriments to post-bleaching recovery of corals

Predicting the response of coral reefs to large-scale mortality induced by climate change will depend greatly on the factors that influence recovery after bleaching events. We experimentally transplanted hard corals from a shallow reef with highly variable seawater temperature (23–36°C) to three unfished marine parks and three fished reefs with variable coral predator abundance and benthic cover. The transplanted corals were fragmented colonies collected from a reef that was relatively undisturbed by the 1997–1998 warm-water temperature anomaly, one of the most extreme thermal events of the past century, and it was assumed that they would represent corals likely to succeed in the future temperature environment. We examined the effects of four taxa, two fragment sizes, an acclimation period, benthic cover components, predators and tourists on the survival of the coral fragments. We found the lowest survival of transplants occurred in the unfished marine parks and this could be attributed to predation and not tourist damage. The density of small coral recruits approximately 6 months after the spawning season was generally moderate (~40–60/m²), and not different on fished and unfished reefs. Coral recovery between 1998 and 2002 was variable (0–25%), low (mean of 6.5%), and not different between fished and unfished reefs. There was high variability in coral mortality among the three unfished areas despite low variation in estimates of predator biomass, with the highest predation occurring in the Malindi MNP, a site with high coralline algal cover. Stepwise multiple regression analysis with 14 variables of coral predators and substratum showed that coralline algae was positively, and turf algae negatively associated with mortality of the transplants, with all other variables being statistically insignificant. This suggests that alternate food resources and predator choices are more important than predator biomass in determining coral survival. Nonetheless, large predatory fish in areas dominated by coralline algae may considerably retard recovery of eurythermal corals. This will not necessarily retard total hard coral recovery, as other more predator-tolerant taxa can recover. Based on the results, global climate change will not necessarily favor eurythermal over stenothermal coral taxa in remote or unfished reefs, where predation is a major cause of coral mortality.     

How corals made rocks through the ages

Hard, or stony, corals make rocks that can, on geological time scales, lead to the formation of massive reefs in shallow tropical and subtropical seas. In both historical and contemporary oceans, reef‐building corals retain information about the marine environment in their skeletons, which is an organic–inorganic composite material. The elemental and isotopic composition of their skeletons is frequently used to reconstruct the environmental history of Earth's oceans over time, including temperature, pH, and salinity. Interpretation of this information requires knowledge of how the organisms formed their skeletons. The basic mechanism of formation of calcium carbonate skeleton in stony corals has been studied for decades. While some researchers consider coral skeletons as mainly passive recorders of ocean conditions, it has become increasingly clear that biological processes play key roles in the biomineralization mechanism. Understanding the role of the animal in living stony coral biomineralization and how it evolved has profound implications for interpreting environmental signatures in fossil corals to understand past ocean conditions. Here we review historical hypotheses and discuss the present understanding of how corals evolved and how their skeletons changed over geological time. We specifically explain how biological processes, particularly those occurring at the subcellular level, critically control the formation of calcium carbonate structures. We examine the different models that address the current debate including the tissue–skeleton interface, skeletal organic matrix, and biomineralization pathways. Finally, we consider how understanding the biological control of coral biomineralization is critical to informing future models of coral vulnerability to inevitable global change, particularly increasing ocean acidification.