Recurrent disturbances, recovery trajectories, and resilience of coral assemblages on a South Central Pacific reef

Coral reefs are increasingly threatened by various disturbances, and a critical challenge is to determine their ability for resistance and resilience. Coral assemblages in Moorea, French Polynesia, have been impacted by multiple disturbances (one cyclone and four bleaching events between 1991 and 2006). The 1991 disturbances caused large declines in coral cover (~51% to ~22%), and subsequent colonization by turf algae (~16% to ~49%), but this phase-shift from coral to algal dominance has not persisted. Instead, the composition of the coral community changed following the disturbances, notably favoring an increased cover of Porites, reduced cover of Montipora and Pocillopora, and a full return of Acropora; in this form, the reef returned to pre-disturbance coral cover within a decade. Thus, this coral assemblage is characterized by resilience in terms of coral cover, but plasticity in terms of community composition.     

Recruitment and larval connectivity of a remnant Acropora community in the Arabian Gulf,United Arab Emirates

Coral cover and community structure in the Arabian Gulf have changed considerably in recent decades. Recurrent bleaching events have dramatically reduced the abundance of previously dominant Acropora corals and have given space to other more thermally resistant coral taxa. The loss of Acropora spp. has reduced reef structural complexity and associated biodiversity. Sir Bu Nair Island (SBN) is a nature reserve in the United Arab Emirates that sustains some of the last dense and extensive Acropora stands in the southern Gulf. This study investigated coral recruitment at a southern coral reef on SBN and examined larval dispersal and reef connectivity between SBN and other local and regional reefs through an agent-based model coupled with a 3D hydrodynamic model. Recruitment was surveyed with settlement tiles deployed from April to September 2019. Contrary to other reefs in the Gulf, we found that Acropora is indeed the major coral recruiter settling at SBN reefs, followed by Porites. The models indicate that SBN reefs are mostly self-seeding but also connected to other reefs in the Gulf. SBN can supply coral larvae to the neighbouring islands Siri and Abu Musa, and nearby reefs along with the north-eastern Emirates, Iranian coast and Strait of Hormuz. Findings highlight the importance of SBN to protect remnant populations of the locally almost extinct Acropora in a region where natural coral recovery is increasingly sparse.

     

The Lagoon at Caroline/Millennium Atoll,Republic of Kiribati: Natural History of a Nearly Pristine Ecosystem

A series of surveys were carried out to characterize the physical and biological parameters of the Millennium Atoll lagoon during a research expedition in April of 2009. Millennium is a remote coral atoll in the Central Pacific belonging to the Republic of Kiribati, and a member of the Southern Line Islands chain. The atoll is among the few remaining coral reef ecosystems that are relatively pristine. The lagoon is highly enclosed, and was characterized by reticulate patch and line reefs throughout the center of the lagoon as well as perimeter reefs around the rim of the atoll. The depth reached a maximum of 33.3 m in the central region of the lagoon, and averaged between 8.8 and 13.7 m in most of the pools. The deepest areas were found to harbor large platforms of Favia matthaii, which presumably provided a base upon which the dominant corals (Acropora spp.) grew to form the reticulate reef structure. The benthic algal communities consisted mainly of crustose coralline algae (CCA), microfilamentous turf algae and isolated patches of Halimeda spp. and Caulerpa spp. Fish species richness in the lagoon was half of that observed on the adjacent fore reef. The lagoon is likely an important nursery habitat for a number of important fisheries species including the blacktip reef shark and Napoleon wrasse, which are heavily exploited elsewhere around the world but were common in the lagoon at Millennium. The lagoon also supports an abundance of giant clams (Tridacna maxima). Millennium lagoon provides an excellent reference of a relatively undisturbed coral atoll. As with most coral reefs around the world, the lagoon communities of Millennium may be threatened by climate change and associated warming, acidification and sea level rise, as well as sporadic local resource exploitation which is difficult to monitor and enforce because of the atoll''s remote location. While the remote nature of Millennium has allowed it to remain one of the few nearly pristine coral reef ecosystems in the world, it is imperative that this ecosystem receives protection so that it may survive for future generations.     

Recent disturbances augment community shifts in coral assemblages in Moorea, French Polynesia

Coral reefs are often subject to disturbances that can cause enduring changes in community structure and abundance of coral reef organisms. In Moorea, French Polynesia, frequent disturbances between 1979 and 2003 caused marked shifts in taxonomic composition of coral assemblages. This study explores recent changes in live cover and taxonomic structure of coral communities on the north coast of Moorea, French Polynesia, to assess whether coral assemblages are recovering (returning to a previous Acropora-dominated state) or continuing to move towards an alternative community structure. Coral cover declined by 29.7% between July 2003 and March 2009, mostly due to loss of Acropora and Montipora spp. Coral mortality varied among habitats, with highest levels of coral loss on the outer reef slope (7–20 m depth). In contrast, there was limited change in coral cover within the lagoon, and coral cover actually increased on the reef crest. Observed changes in coral cover and composition correspond closely with the known feeding preferences and observed spatial patterns of Acanthaster planci L., though observed coral loss also coincided with at least one episode of coral bleaching, as well as persistent populations of the corallivorous starfish Culcita novaeguineae Muller & Troschel. While climate change poses an important and significant threat to the future structure and dynamics coral reef communities, outbreaks of A. planci remain a significant cause of coral loss in Moorea. More importantly, these recent disturbances have followed long-term shifts in the structure of coral assemblages, and the relative abundance of both Pocillopora and Porites continue to increase due to disproportionate losses of Acropora and Montipora. Moreover, Pocillopora and Porites dominate assemblages of juvenile corals, suggesting that there is limited potential for a return to an Acropora-dominated state, last recorded in 1979.     

Effects of disturbance on coral communities: bleaching in Moorea,French Polynesia

This study examines patterns of susceptibility and short-term recovery of corals from bleaching. A mass coral bleaching event began in March, 1991 on reefs in Moorea, French Polynesia and affected corals on the shallow barrier reef and to >20 m depth on the outer forereef slope. There were significant differences in the effect of the bleaching among common coral genera, with Acropora, Montastrea, Montipora, and Pocillopora more affected than Porites, Pavona, leptastrea or Millepora. Individual colonies of the common species of Acropora and Pocillopora were marked and their fate assessed on a subsequent survey in August, 1991 to determine rates of recovery and mortality. Ninety-six percent of Acropora spp. showed some degree of bleaching compared to 76% of Pocillopora spp. From March to August mortality of bleached colonies of Pocillopora was 17%, 38% recovered completely, and many suffered some partial mortality of the tissue. In contrast, 63% of the Acropora spp. died, and about 10% recovered completely. Generally, those colonies with less than 50% of the colony area affected by the bleaching recovered at a higher rate than did those with more severe bleaching. Changes in community composition four months after the event began included a significant decrease only in crustose algae and an increase in cover of filamentous algae, much of which occupied plate-like and branching corals that had died in the bleaching event. Total coral cover and cover of susceptible coral genera had declined, but not significantly, after the event.     

The influence of repetitive thermal stresses on the dominance of reef-building Acropora spp. (Scleractinia) on coral reefs of the Maldive Islands

The distribution and size-age structure of Acropora corals were studied in two Maldivian atolls that differ in their geographic position and sea surface temperature regimes. The frequency and strength of thermal anomalies for the last 2 decades had a significant influence on the abundance, mortality rates, and age structures of acroporid communities. The long-term temperature amplitude was higher and the maxima were more pronounced in the northernmost Ihavandippolu Atoll than those in the equatorial South Huvadhoo Atoll. These differences resulted in a 10.4% mean cover of Acropora at Ihavandippolu Atoll, whereas the Acropora cover in the South Huvadhoo Atoll reached 59.5%. In the northern atoll, the coral mortality rate after the 2010 thermal anomaly was 3 times higher than that in the southern atoll. Younger acroporid colonies (up to 2 years old) dominated the northern atoll reefs, while the southern atoll showed a high proportion of older mature colonies. In both atolls, healthy table colonies of Acropora cytherea with a disk diameter greater than 2 m were observed that apparently survived three thermal anomalies since 1998. The mechanisms of acclimatization of Acropora and the prospects for its dominance in the Maldives under changing environmental conditions are discussed.     

Regional‐scale dominance of non‐framework building corals on Caribbean reefs affects carbonate production and future reef growth

Coral cover on Caribbean reefs has declined rapidly since the early 1980's. Diseases have been a major driver, decimating communities of framework building Acropora and Orbicella coral species, and reportedly leading to the emergence of novel coral assemblages often dominated by domed and plating species of the genera Agaricia, Porites and Siderastrea. These corals were not historically important Caribbean framework builders, and typically have much smaller stature and lower calcification rates, fuelling concerns over reef carbonate production and growth potential. Using data from 75 reefs from across the Caribbean we quantify: (i) the magnitude of non‐framework building coral dominance throughout the region and (ii) the contribution of these corals to contemporary carbonate production. Our data show that live coral cover averages 18.2% across our sites and coral carbonate production 4.1 kg CaCO₃ m^?2 yr^?1. However, non‐framework building coral species dominate and are major carbonate producers at a high proportion of sites; they are more abundant than Acropora and Orbicella at 73% of sites; contribute an average 68% of the carbonate produced; and produce more than half the carbonate at 79% of sites. Coral cover and carbonate production rate are strongly correlated but, as relative abundance of non‐framework building corals increases, average carbonate production rates decline. Consequently, the use of coral cover as a predictor of carbonate budget status, without species level production rate data, needs to be treated with caution. Our findings provide compelling evidence for the Caribbean‐wide dominance of non‐framework building coral taxa, and that these species are now major regional carbonate producers. However, because these species typically have lower calcification rates, continued transitions to states dominated by non‐framework building coral species will further reduce carbonate production rates below ‘predecline’ levels, resulting in shifts towards negative carbonate budget states and reducing reef growth potential.     

Wave exposure shapes reef community composition and recovery trajectories at a remote coral atoll

In a time of unprecedented ecological change, understanding natural biophysical relationships between reef resilience and physical drivers is of increasing importance. This study evaluates how wave forcing structures coral reef benthic community composition and recovery trajectories after the major 2015/2016 bleaching event in the remote Chagos Archipelago, Indian Ocean. Benthic cover and substrate rugosity were quantified from digital imagery at 23 fore reef sites around a small coral atoll (Salomon) in 2020 and compared to data from a similar survey in 2006 and opportunistic surveys in intermediate years. Cluster analysis and principal component analysis show strong separation of community composition between exposed (modelled wave exposure > 1000 J m⁻³) and sheltered sites (< 1000 J m⁻³) in 2020. This difference is driven by relatively high cover of Porites sp., other massive corals, encrusting corals, soft corals, rubble and dead table corals at sheltered sites versus high cover of pavement and sponges at exposed sites. Total coral cover and rugosity were also higher at sheltered sites. Adding data from previous years shows benthic community shifts from distinct exposure-driven assemblages and high live coral cover in 2006 towards bare pavement, dead Acropora tables and rubble after the 2015/2016 bleaching event. The subsequent recovery trajectories at sheltered and exposed sites are surprisingly parallel and lead communities towards their respective pre-bleaching communities. These results demonstrate that in the absence of human stressors, community patterns on fore reefs are strongly controlled by wave exposure, even during and after widespread coral loss from bleaching events.

     

Effects of the 2000 southern Mozambique floods on a marginal coral community: the case at Xai-Xai

In early 2000 the southern part of Mozambique suffered the worst flooding in 50 years, causing fatalities and considerable material loss. This study aimed to investigate the impact of this flood on the coral communities in Xai-Xai lagoon. Benthic cover was assessed in January 2000 (before the floods) and September 2000 (after the floods) using the line intercept transect technique. A decrease in hard coral cover of the order of 58.5% was observed. The soft coral community was significantly affected, with a decrease in percentage cover of 90.4%. Coralline algae also decreased by 85.1%. All other categories increased in percentage cover: turf algae (164.4%), other invertebrates (e.g. sponges, sea urchins — 111.1%), fleshy algae (80.4%), rubble (34.4%) and dead coral (379.0%). The main causes of this degradation were the reduced water salinity and the large amount of sediment discharged by the Limpopo River. Some massive (e.g. Porites, Favia, Favites and Goniopora) and encrusting (e.g. Echinopora) hard coral genera seemed less affected, suggesting an elevated capacity to cope with this kind of stress through mucus-sheet formation. The extent of the flood effect on other reefs on the southern Mozambique coast is discussed and a monitoring programme is proposed.     

Latitudinal variation in coral communities in eastern Australia: a qualitative biophysical model of factors regulating coral reefs

The processes underlying the distributional limits of both corals and coral reefs can be elucidated by examining coral communities at high latitudes. Coral-dominated communities in eastern Australia cover a latitudinal range of >2,500 km, from the northern Great Barrier Reef (11°S) to South West Rocks (31.5°S). Patterns of coral species richness from 11 locations showed a clear separation between the Great Barrier Reef and subtropical sites, with a further abrupt change at around 31°S. Differences in community structure between the Great Barrier Reef and more southern sites were mainly attributable to higher cover of massive corals, branching Acropora, dead coral and coralline algae on the Great Barrier Reef, and higher cover of macroalgae and bare rock at more southern sites. The absence of some major reef-building taxa (i.e., staghorn Acropora and massive Porites) from most subtropical sites coincided with the loss of reef accretion capacity. Despite high cover of hard corals in communities at up to 31°S, only Lord Howe Island contained areas of reef accretion south of the Great Barrier Reef. Factors that have been hypothesized to account for latitudinal changes in coral community structure include water temperature, aragonite saturation, light availability, currents and larval dispersal, competition between corals and other biota including macroalgae, reduced coral growth rates, and failure of coral reproduction or recruitment. These factors do not operate independently of each other, and they interact in complex ways.     

Benthic Composition of a Healthy Subtropical Reef: Baseline Species-Level Cover,with an Emphasis on Algae,in the Northwestern Hawaiian Islands

The Northwestern Hawaiian Islands (NWHI) are considered to be among the most pristine coral reef ecosystems remaining on the planet. These reefs naturally contain a high percent cover of algal functional groups with relatively low coral abundance and exhibit thriving fish communities dominated by top predators. Despite their highly protected status, these reefs are at risk from both direct and indirect anthropogenic sources. This study provides the first comprehensive data on percent coverage of algae, coral, and non-coral invertebrates at the species level, and investigates spatial diversity patterns across the archipelago to document benthic communities before further environmental changes occur in response to global warming and ocean acidification. Monitoring studies show that non-calcified macroalgae cover a greater percentage of substrate than corals on many high latitude reef sites. Forereef habitats in atoll systems often contain high abundances of the green macroalga Microdictyon setchellianum and the brown macroalga Lobophora variegata, yet these organisms were uncommon in forereefs of non-atoll systems. Species of the brown macroalgal genera Padina, Sargassum, and Stypopodium and the red macroalgal genus Laurencia became increasingly common in the two northernmost atolls of the island chain but were uncommon components of more southerly islands. Conversely, the scleractinian coral Porites lobata was common on forereefs at southern islands but less common at northern islands. Currently accepted paradigms of what constitutes a “healthy” reef may not apply to the subtropical NWHI, and metrics used to gauge reef health (e.g., high coral cover) need to be reevaluated.     

Analysis of complete mitochondrial DNA sequences of three members of the Montastraea annularis coral species complex (Cnidaria, Anthozoa, Scleractinia)

Complete mitochondrial nucleotide sequences of two individuals each of Montastraea annularis, Montastraea faveolata, and Montastraea franksi were determined. Gene composition and order differed substantially from the sea anemone Metridium senile, but were identical to that of the phylogenetically distant coral genus Acropora. However, characteristics of the non-coding regions differed between the two scleractinian genera. Among members of the M. annularis complex, only 25 of 16,134 base pair positions were variable. Sixteen of these occurred in one colony of M. franksi, which (together with additional data) indicates the existence of multiple divergent mitochondrial lineages in this species. Overall, rates of evolution for these mitochondrial genomes were extremely slow (0.03–0.04% per million years based on the fossil record of the M. annularis complex). At higher taxonomic levels, patterns of genetic divergence and synonymous/nonsynonymous substitutions suggest non-neutral and unequal rates of evolution between the two lineages to which Montastraea and Acropora belong.     

Character release following extinction in a Caribbean reef coral species complex

Abstract The Pleistocene extinction of the widespread organ‐pipe Montastraea coral had measurable morphological and ecological effects on surviving lineages of the Montastraea“annularis” species complex. Extinction of the organ‐pipe Montastraea occurred after more than 500,000 years of dominance in the shallow‐water reef habitat of Barbados. Extinction resulted in a morphological shift of the columnar Montastraea lineage from thick to thin columns in modern reef environments. Pleistocene colonies of the columnar morphotype sympatric with organ‐pipe Montastraea showed greater column widths than those in allopatry. We subjected our data to a number of criteria for interpreting the morphological shift as character release following lifting of competitive pressure after extinction. The morphological differences do not appear to be due either to chance or to physical properties of the marine environment. Differential local extinction and recolonization of four members of the species complex did not occur on Barbados, so that the species coexisted and appear to have coevolved between more than 600,000 and 82,000 years ago. The morphological shift is related to coral growth form and growth rate, and thus reflects the acquisition of a primary resource in corals‐light. Character release occurred at the same oceanic Caribbean island (Barbados) where environments have fluctuated with similar variance throughout the period of coexistence. Not only has competition among living members of the Montastraea“annularis” species complex been convincingly demonstrated, but trends in relative abundance among fossil members of the species complex strongly suggest that a competitive hierarchy was operating during their Pleistocene coexistence on Barbados. We also observed an ecological analogue to character release on another Caribbean island, Curaçao. The distribution and abundance of living columnarM. annularis s.s. and massive M. faveolata from the leeward reef crest in Curaçao is greater now than in the Pleistocene, when organ‐pipe Montastraea dominated this shallow‐water reef habitat. Extinction of the faster growing, shallow‐water organ‐pipe Montastraea resulted in higher abundance of the columnar Montastraea lineage in shallow‐water habitats, where it shifted its morphology to one adapted to high light levels. The species extinction released surviving lineages from a competitive network that had resulted in lower rank abundance in the Pleistocene community and enhanced abundance of both columnar M. annularis s.s. and M. faveolata in modern communities. Full validation of our interpretation of character release must await experiments that demonstrate whether phenotypic differences between populations have a genetic basis. However, we believe the results of this study point to the important, yet heretofore neglected, role that biological interactions have played in the evolution of closely related reef coral species.     

Changes in coral assemblages during an outbreak of Acanthaster planci at Lizard Island, northern Great Barrier Reef (1995–1999)

Population outbreaks of crown-of-thorns starfish (Acanthaster planci L.) represent one of the most significant biological disturbances on tropical coral reefs and have the potential to devastate coral communities, thereby altering the biological and physical structure of reef habitats. This study reports on changes in area cover, species diversity and taxonomic composition of corals during an outbreak of A. planci at Lizard Island, in the northern Great Barrier Reef, Australia. Mean coral cover declined by 28.8% across ten locations studied. However, densities of A. planci, and their effects on local coral assemblages, were very patchy. Declines in coral cover were mostly due to the selective removal of certain coral taxa (mainly Acropora and Pocilloporidae corals); such that the greatest coral loss occurred at locations with highest initial cover of preferred coral prey. Most notably, coral assemblages in back-reef locations were transformed from topographically complex staghorn Acropora-dominated habitats, to relatively depauperate assemblages dominated by alcyonacean soft corals. Although coral loss was greatest among formerly dominant taxa (especially Acropora), effects were sufficiently widespread across different coral taxa, such that overall coral diversity tended to decline. Clearly, moderate outbreaks of A. planci have the potential to greatly alter community structure of coral communities even if they do not devastate live corals. Recovery in this instance is expected to be very rapid given that all coral taxa persisted, and effects were greatest among fast growing corals.     

Wave and tidal flushing in a near-equatorial mesotidal atoll

Most of the atolls found worldwide are under microtidal regimes, and their circulation mechanisms are widely documented and well known. Here, we describe the flushing mechanisms of a small-sized mesotidal atoll, based on water-level, wave and current data obtained during two different periods (total of 60 d). Rocas is the only atoll in the South Atlantic Ocean and is built primarily of coralline algae. Two reef passages connect the atoll lagoon to the ocean. Synchronous current profilers were deployed at the two reef passages, one inside and one outside the atoll, to characterize the influence of tides and waves on the circulation. Results showed that wind waves drove a setup on the exposed side of the atoll and that currents were predominately downwind, causing outflow at both reef passages. Waves breaking on the windward side supplied water to the atoll causing the lagoon water level to rise above ocean water level, driving the outflow. However, unlike microtidal atolls, at Rocas Atoll the water level drops significantly below the reef rim during low tides. This causes the reef rim to act as a barrier to water pumping into the lagoon by waves, resulting in periodic activation of the wave pumping mechanism throughout a tidal cycle. As result, inflow occurs in the wider passage during 27% of each tidal cycle, starting at low tides and reversing direction during mid-flood tide when the water level exceeded approximately 1.6 m (while overtopping the atoll’s rim). Our findings show that tides play a direct role in driving circulation on a mesotidal atoll, not only by modulating wave setup but also by determining the duration of wave pumping into the lagoon.

     

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.     

Impact of Herbivore Identity on Algal Succession and Coral Growth on a Caribbean Reef

Background

Herbivory is an important top-down force on coral reefs that regulates macroalgal abundance, mediates competitive interactions between macroalgae and corals, and provides resilience following disturbances such as hurricanes and coral bleaching. However, reductions in herbivore diversity and abundance via disease or over-fishing may harm corals directly and may indirectly increase coral susceptibility to other disturbances.

Methodology and Principal Findings

In two experiments over two years, we enclosed equivalent densities and masses of either single-species or mixed-species of herbivorous fishes in replicate, 4 m² cages at a depth of 17 m on a reef in the Florida Keys, USA to evaluate the effects of herbivore identity and species richness on colonization and development of macroalgal communities and the cascading effects of algae on coral growth. In Year 1, we used the redband parrotfish (Sparisoma aurofrenatum) and the ocean surgeonfish (Acanthurus bahianus); in Year 2, we used the redband parrotfish and the princess parrotfish (Scarus taeniopterus). On new substrates, rapid grazing by ocean surgeonfish and princess parrotfish kept communities in an early successional stage dominated by short, filamentous algae and crustose coralline algae that did not suppress coral growth. In contrast, feeding by redband parrotfish allowed an accumulation of tall filaments and later successional macroalgae that suppressed coral growth. These patterns contrast with patterns from established communities not undergoing primary succession; on established substrates redband parrotfish significantly reduced upright macroalgal cover while ocean surgeonfish and princess parrotfish allowed significant increases in late successional macroalgae.

Significance

This study further highlights the importance of biodiversity in affecting ecosystem function in that different species of herbivorous fishes had very different impacts on reef communities depending on the developmental stage of the community. The species-specific effects of herbivorous fishes suggest that a species-rich herbivore fauna can be critical in providing the resilience that reefs need for recovery from common disturbances such as coral bleaching and storm damage.     

Coral and sea urchin assemblage structure and interrelationships in Kenyan reef lagoons

Patterns of hard coral and sea urchin assemblage structure (species richness, diversity, and abundance) were studied in Kenyan coral reef lagoons which experienced different types of human resource use. Two protected reefs (Malindi and Watamu Marine National Parks) were protected from fishing and coral collection, but exposed to heavy tourist use. One reef (Mombasa MNP) received protection from fishermen for one year and was exploited for fish and corals prior to protection and was defined as a transitional reef. Three reefs (Vipingo, Kanamai, and Diani) were unprotected and experienced heavy fishing and some coral collection. Protected and unprotected reefs were distinct in terms of their assemblage structure with the transitional reef grouping with unprotected reefs based on relative and absolute abundance of coral genera. Protected reefs had slightly higher (p<0.01) coral cover (23.6 ± 8.3 % ± S.D.) than unprotected reefs (16.7 ± 8.5), but the transitional reef had the highest coral cover (30.8 ± 6.4) which increased by 250% since measured in 1987: largely attributable to a large increase inPorites nigrescens cover. Protected reefs had higher coral species richness and diversity and a greater relative abundance ofAcropora, Montipora andGalaxea than unprotected reefs. The transitional reef had high species richness, but lower diversity due to the high dominance ofPorites. Sea urchins showed the opposite pattern with highest diversity in most unprotected reefs. Coral cover, species richness, and diversity were negatively associated with sea urchin abundance, but the relative abundance ofPorites increased with sea urchin abundance to the point wherePorites composed >90% of the coral cover at sites with the highest sea urchin abundance. Effects of coral overcollection was only likely for the genusAcropora (staghorn corals). A combination of direct and indirect effects of human resource use may reduce diversity, species richness, and abundance of corals while increasing the absolute abundance of sea urchins and the relative cover ofPorites.     

Reef habitats and associated sessile-benthic and fish assemblages across a euphotic–mesophotic depth gradient in Isla Desecheo, Puerto Rico

Quantitative surveys of sessile benthos and fish populations associated with reef habitats across a 15–50 m depth gradient were performed by direct diver observations using rebreathers at Isla Desecheo, Puerto Rico. Statistically significant differences between depths were found for total live coral, total coral species, total benthic algae, total sponges and abiotic cover. Live coral cover was higher at the mid-shelf (20 m) and shelf-edge (25 m) stations, whereas benthic algae and sponges were the dominant sessile-benthic assemblage at mesophotic stations below 25 m. Marked shifts in the community structure of corals and benthic algae were observed across the depth gradient. A total of 119 diurnal, non-cryptic fish species were observed across the depth gradient, including 80 species distributed among 7,841 individuals counted within belt-transects. Fish species richness was positively correlated with live coral cover. However, the relationship between total fish abundance and live coral was weak. Abundance of several numerically dominant fish species varied independently from live coral cover and appeared to be more influenced by depth and/or habitat type. Statistically significant differences in the rank order of abundance of fish species at euphotic vs mesophotic stations were detected. A small assemblage of reef fishes that included the cherubfish, Centropyge argi, sunshine chromis, Chromis insolata, greenblotch parrotfish, Sparisoma atomarium, yellowcheek wrasse, Halichoeres cyanocephalus, sargassum triggerfish, Xanthichthys ringens, and the longsnout butterflyfish, Chaetodon aculeatus was most abundant or only present from stations deeper than 30 m, and thus appear to be indicator species of mesophotic habitats.     

Herbivorous fish assemblages and herbivory pressure on Laamu Atoll, Republic of Maldives

While herbivory is recognized as a fundamental process structuring coral reef communities, herbivore assemblages and processes are poorly described for reefs in the Indian Ocean region. We quantified herbivorous fish assemblage structure (abundance and diversity) in Laamu Atoll, Republic of Maldives, in four reef habitat types: faro reef flats, faro reef slopes, inner and outer atoll reef slopes (20 sites in total). Herbivorous fish assemblages, representing a total of 30 species, grouped strongly by habitat type, with the highest absolute abundance observed on faro reef flats and lowest abundance on inside atoll rim reef slopes. Removal of Thalassia seagrass blades by ambient herbivore assemblages was used in a bioassay to assess relative herbivory pressure among four habitat types (eight sites). Also, at one site a choice herbivory assay was performed to assess herbivore preference among four benthic plants across three depth zones. Relative herbivory, as indicated by Thalassia assays, was highest on inside atoll rim reef slopes and lowest on outside atoll rim reef slopes. Thalassia consumption did not correspond to overall herbivorous fish abundance, but corresponded more closely with parrotfish abundance. In the choice assays, herbivores showed strong preferences among plant types and consumption of most plant types was higher at mid-depth than in the shallow reef flat or deep reef knoll zones.