Black reefs: iron-induced phase shifts on coral reefs

The Line Islands are calcium carbonate coral reef platforms located in iron-poor regions of the central Pacific. Natural terrestrial run-off of iron is non-existent and aerial deposition is extremely low. However, a number of ship groundings have occurred on these atolls. The reefs surrounding the shipwreck debris are characterized by high benthic cover of turf algae, macroalgae, cyanobacterial mats and corallimorphs, as well as particulate-laden, cloudy water. These sites also have very low coral and crustose coralline algal cover and are call black reefs because of the dark-colored benthic community and reduced clarity of the overlying water column. Here we use a combination of benthic surveys, chemistry, metagenomics and microcosms to investigate if and how shipwrecks initiate and maintain black reefs. Comparative surveys show that the live coral cover was reduced from 40 to 60% to <10% on black reefs on Millennium, Tabuaeran and Kingman. These three sites are relatively large (>0.75 km²). The phase shift occurs rapidly; the Kingman black reef formed within 3 years of the ship grounding. Iron concentrations in algae tissue from the Millennium black reef site were six times higher than in algae collected from reference sites. Metagenomic sequencing of the Millennium Atoll black reef-associated microbial community was enriched in iron-associated virulence genes and known pathogens. Microcosm experiments showed that corals were killed by black reef rubble through microbial activity. Together these results demonstrate that shipwrecks and their associated iron pose significant threats to coral reefs in iron-limited regions.     

Could some coral reefs become sponge reefs as our climate changes?

Coral reefs across the world have been seriously degraded and have a bleak future in response to predicted global warming and ocean acidification (OA). However, this is not the first time that biocalcifying organisms, including corals, have faced the threat of extinction. The end‐Triassic mass extinction (200 million years ago) was the most severe biotic crisis experienced by modern marine invertebrates, which selected against biocalcifiers; this was followed by the proliferation of another invertebrate group, sponges. The duration of this sponge‐dominated period far surpasses that of alternative stable‐ecosystem or phase‐shift states reported on modern day coral reefs and, as such, a shift to sponge‐dominated reefs warrants serious consideration as one future trajectory of coral reefs. We hypothesise that some coral reefs of today may become sponge reefs in the future, as sponges and corals respond differently to changing ocean chemistry and environmental conditions. To support this hypothesis, we discuss: (i) the presence of sponge reefs in the geological record; (ii) reported shifts from coral‐ to sponge‐dominated systems; and (iii) direct and indirect responses of the sponge holobiont and its constituent parts (host and symbionts) to changes in temperature and pH. Based on this evidence, we propose that sponges may be one group to benefit from projected climate change and ocean acidification scenarios, and that increased sponge abundance represents a possible future trajectory for some coral reefs, which would have important implications for overall reef functioning.     

Bioeroders in fossil reefs

Summary Boring algae, fungi and bacteria have been the most constant factor in bioerosion through earth history. Their record reaches back into the middle Precambrian. The only fossil reefs specifically researched for these microendoliths are of Triassic and Upper Jurassic age. Boring worms appear in reefs in the Lower Cambrian. Boring sponges and bivalves first appear also in the lower Paleozoic, but do not become abundant in reefs until the Triassic. Effective substrate excavating grazers are relatively young geologically: Patellids and substrate excavating Echinoids evolved in the Triassic but did not become important bioeroders until the Jurassic or Cretaceous. Scarid fishes are even younger, the oldest representatives having been found in the Miocene. Thus, it seems that the intensity of bioerosion changed significantly during earth history. This may have had consequences for diversity of reef organisms, quality and quantity of reef debris, for diagenesis and record of reef rock.     

Triassic reefs in Slovenia

The paper deals with the distribution, paleogeography, age and biota of Triassic reefs in Slovenia. Most of these reefs have not been studied in detail up to now, but the paleographical distributional pattern can be outlined (Figs. 1 and 2). Triassic reefs are known from Central and Northern Slovenia, predominantly occurring at the margins of the “Slovenian trough” (which separates the northern Julian Platform and the southern Dinaric Platform) and at the margins of an intraplatform trough within the Julian platform. Reef growth started in the Ladinian and Cordevolian and continued (with interruptions during the Upper Carnian ?) to the Norian and Rhaetian. Anisian environments are characterized by the predominance of algal mats and dasycladacean algae. Cordevolian patch reefs as well as Norian and Rhaetian reefs were built during the Late Triassic by calcareous sponges and corals, which belong to different species (Tab. 1 and 2). Some smaller Cordevolian patch reefs may have been formed within deeper-water sediments. An interesting facies sequence is developed in the Norian Dachstein Limestone reef of Pokljuka (Julian Alps), starting with deeper-marine cherty limestones, which gradually succeeded by crinoidal limestones followed by reef limestones and lagoonal Dachstein Limestones.     

Biological destruction of coral reefs

The major agents of biological destruction of coral reefs can be divided into grazers, etchers and borers. Each of these groups is reviewed on a world wide basis, together with the mechanisms by which they destroy the coral substrate. Rates of bioerosion attributed to major agents of grazers, etchers and borers are given, together with limitations of some of the measurements. Recent work is highlighting the variability in rates of bioerosion both over time and space. Factors which may be responsible for this variability are discussed. Bioerosion is a major factor influencing reef morphology and the ways in which this is achieved is discussed in some detail. Although the review concentrates mainly on present day reefs, some attempt is made to consider the impact of bioerosion on older reefs.     

Middle Ordovician reefs of Norway

The Middle Ordovician reefs of Norway were the first to develop in the western part of the Balto-scandian epicontinental sea and are the earliest coral-stromatoporoid reefs so far reported in Europe. Small patch reefs in the Steinvika Limestone, Langesund-Skien district, consist mainly of algae, echinoderms, corals and stromatoporoids. Bryozoans, molluscs, arthropods and brachiopods are also present. The reefs developed on pelmatozoan-rich substrates and are organically zoned, consisting of a pioneer community of stemmed echinoderms and sheet algae, a high-diversity intermediate community dominated by fasciculate corals and a low diversity climax community of massive corals and stromatoporoids. These communities are interpreted as the seral stages of an autogenic ecological succession. Small patch reefs are also present in the laterally equivalent Mjøsa Limestone, Toten and Nes-Hamar districts. These are organically very similar to those in the Steinvika Limestone and developed in an identical way. A large complex, consisting of several reefs, is also present in the Mjøsa Limestone. Unlike the reefs elsewhere, which developed within shallow inshore areas, this complex developed at the outer edge of the inshore shelf. The outstanding feature of the complex is the main reef forming the offshore limit which is totally dominated by stromatoporoids and lacks a sequential development. This is due to the influence of the harsher environment at the shelf edge.     

Global change and coral reefs: impacts on reefs, economies and human cultures

Coral reefs have reconstituted themselves after previous large sea-level variations, and climate changes. For the past 6000 years of unusually stable sea-level, reefs have grown without serious interruptions. During recent decades, however, new stresses threaten localized devastation of many reefs. A new period of global climate change is occurring, stimulated by anthropogenic increases in greenhouse gases. Coral reefs will cope well with predicted sea-level rises of 4.5 cm per decade, but reef islands will not. Higher sea levels will provide corals with greater room for growth across reef flats, but there are no foreseeable mechanisms for reef island growth to keep pace with sea-level rise, therefore many low islands may ultimately become uninhabitable. Climate change will introduce localized variations in weather patterns, but changes to individual reefs cannot be predicted. Reefs on average should cope well with regional climate change, as they have coped with similar previous fluctuations. Air temperature increases of 0.2–0.3 °C/decade will induce slower increases in sea-surface temperatures, which may cause localized, or regional increases in coral bleaching. Changes in rainfall will impact on reefs near land masses. Likewise, increased storms and variations in El Nino Southern Oscillation (ENSO) may stress some reefs, but not others. The greatest impact of climate change will be a synergistic enhancement of direct anthropogenic stresses (excessive sediment and pollution from the land; over-fishing, especially via destructive methods; mining of coral rock and sand; and engineering modifications), which currently cause most damage to coral reefs. Many of the world's reefs have been degraded and more will be damaged as anthropogenic impacts increase under the ‘demophoric’ increases in population (demos) and economic (phoric) activity. This biotic and habitat loss will result in severe economic and social losses. Reefs, however, have considerable recovery powers and losses can be minimized by effective management of direct human impacts and reducing indirect threats of global climate change.     

Coral-algal competition on damaged reefs

Natural and anthropogenic catastrophes occurred at the end of the previous and in the beginning of the current centuries at the coral reefs of the World Ocean, and their consequences for the tropical shelf ecosystems have been described based on published data and our own investigations. It has been shown that in recent decades coral populations on reefs of tropical and subtropical regions of the World Ocean have been reduced by 80%, and in some areas have completely vanished. The biodiversity of reef ecosystems has been considerably reduced. The main reason for such changes is a 1-2°C increase in the temperature of surface waters in comparison with the monthly mean temperature in the hot season. The fate of the damaged coral reefs is under discussion. It is thought that in clean waters partially damaged coral reefs can recover, whereas in waters polluted as the result of human activity they collapse. The rate of coral reef restoration depends on the hydrological and hydrochemical conditions, frequency of natural calamities and competitive interrelation of algae and corals on the damaged sites of coral reefs. The nature of competitive interrelation between algae and corals is considered, viz., the dynamics of obliteration of damaged and dead coral colonies by various algal species, mechanisms of competitive interrelation, effects of the environment on the competitive ability of corals and algae, the internal and external conditions for victory in competitive activity. It has been suggested that coral reefs can be restored through temporary transformation into a vegetable reef. In the absence of natural calamities damaged reefs can be clearly restored to their original or altered state over several decades, but only in clean waters.     

Comparing the invasibility of experimental "reefs" with field observations of natural reefs and artificial structures

Natural systems are increasingly being modified by the addition of artificial habitats which may facilitate invasion. Where invaders are able to disperse from artificial habitats, their impact may spread to surrounding natural communities and therefore it is important to investigate potential factors that reduce or enhance invasibility. We surveyed the distribution of non-indigenous and native invertebrates and algae between artificial habitats and natural reefs in a marine subtidal system. We also deployed sandstone plates as experimental 'reefs' and manipulated the orientation, starting assemblage and degree of shading. Invertebrates (non-indigenous and native) appeared to be responding to similar environmental factors (e.g. orientation) and occupied most space on artificial structures and to a lesser extent reef walls. Non-indigenous invertebrates are less successful than native invertebrates on horizontal reefs despite functional similarities. Manipulative experiments revealed that even when non-indigenous invertebrates invade vertical "reefs", they are unlikely to gain a foothold and never exceed covers of native invertebrates (regardless of space availability). Community ecology suggests that invertebrates will dominate reef walls and algae horizontal reefs due to functional differences, however our surveys revealed that native algae dominate both vertical and horizontal reefs in shallow estuarine systems. Few non-indigenous algae were sampled in the study, however where invasive algal species are present in a system, they may present a threat to reef communities. Our findings suggest that non-indigenous species are less successful at occupying space on reef compared to artificial structures, and manipulations of biotic and abiotic conditions (primarily orientation and to a lesser extent biotic resistance) on experimental "reefs" explained a large portion of this variation, however they could not fully explain the magnitude of differences.     

Marine pollution and coral reefs

Coral reefs are exposed to many anthropogenic stresses increasing in impact and range, both on local and regional scales. The main ones discussed here are nutrient enrichment, sewage disposal, sedimentation, oil-related pollution, metals and thermal pollution. The stress comprising the main topic of this article, eutrophication, is examined from the point of view of its physiological and ecological mechanisms of action, on a number of levels. Nutrient enrichment can introduce an imbalance in the exchange of nutrients between the zooxanthellae and the host coral, it reduces light penetration to the reef due to nutrient- stimulated phytoplankton growth, and, most harmful of all, may bring about proliferation of seaweeds. The latter rapidly outgrow, smother and eventually replace, the slow-growing coral reef, adapted to cope with the low nutrient concentrations typical in tropical seas.
Eutrophication seldom takes place by itself. Sewage disposal invariably results in nutrient enrichment, but it also enriches the water with organic matter which stimulates proliferation of oxygen-consuming microbes. These may kill corals and other reef organisms, either directly by anoxia, or by related hydrogen sulfide production. Increased sediment deposition is in many cases associated with other human activities leading to eutrophication, such as deforestation and topsoil erosion.
Realistically achievable goals to ensure conservation, and in some instances, rehabilitation of coral reefs are listed.     

Coral communities of barrier reefs of Vietnam

The composition and spatial distribution of the coral communities of the barrier reefs of Jiang Bo and of Re Island were described in detail for the first time for Vietnamese waters. Their comparability to the ribbon reefs of the Great Barrier Reef in Australia and to the barrier reefs of the Philippines and Indian Ocean was revealed by morphological parameters, species diversity and zonal distribution. Their geomorphological status, the presence of fore reef, epi-reef and back reef complexes with their specific composition of flora and fauna, and an obligatory lagoon separating the reef from fringing inshore reefs, enabled the attribution of the surveyed reefs to the barrier type of reef.     

Caribbean reefs of the Anthropocene: Variance in ecosystem metrics indicates bright spots on coral depauperate reefs

Dramatic coral loss has significantly altered many Caribbean reefs, with potentially important consequences for the ecological functions and ecosystem services provided by reef systems. Many studies examine coral loss and its causes—and often presume a universal decline of ecosystem services with coral loss—rather than evaluating the range of possible outcomes for a diversity of ecosystem functions and services at reefs varying in coral cover. We evaluate 10 key ecosystem metrics, relating to a variety of different reef ecosystem functions and services, on 328 Caribbean reefs varying in coral cover. We focus on the range and variability of these metrics rather than on mean responses. In contrast to a prevailing paradigm, we document high variability for a variety of metrics, and for many the range of outcomes is not related to coral cover. We find numerous “bright spots,” where herbivorous fish biomass, density of large fishes, fishery value, and/or fish species richness are high, despite low coral cover. Although it remains critical to protect and restore corals, understanding variability in ecosystem metrics among low‐coral reefs can facilitate the maintenance of reefs with sustained functions and services as we work to restore degraded systems. This framework can be applied to other ecosystems in the Anthropocene to better understand variance in ecosystem service outcomes and identify where and why bright spots exist.