The cover of living and dead corals from airborne remote sensing

Trends in coral cover are widely used to indicate the health of coral reefs but are costly to obtain from field survey over large areas. In situ studies of reflected spectra at the coral surface show that living and recently dead colonies can be distinguished. Here, we investigate whether such spectral differences can be detected using an airborne remote sensing instrument. The Compact Airborne Spectrographic Imager (Itres Research Ltd, Canada) was flown in two configurations: 10 spectral bands with 1-m² pixels and 6 spectral bands with 0.25-m² pixels. First, we show that an instrument with 10 spectral bands possesses adequate spectral resolution to distinguish living Porites, living Pocillopora spp., partially dead Porites, recently dead Porites (total colony mortality within 6 months), old dead (>6 months) Porites, Halimeda spp., and coralline red algae when there is no water column to confuse spectra. All substrata were distinguished using fourth-order spectral derivatives around 538 nm and 562 nm. Then, at a shallow site (Tivaru) at Rangiroa Atoll, Tuamotu Archipelago (French Polynesia), we show that live and dead coral can be distinguished from the air to a depth of at least 4 m using first- and fourth-order spectral derivatives between 562–580 nm. However, partially dead and recently dead Porites colonies could not be distinguished from an airborne platform. Spectral differences among substrata are then exploited to predict the cover of reef substrata in ten 25-m² plots at nearby Motu Nuhi (max depth 8 m). The actual cover in these plots was determined in situ using quadrats with a 0.01-m² grid. Considerable disparity occurred between field and image-based measures of substrate cover within individual 25-m² quadrats. At this small scale, disparity, measured as the absolute difference in cover between field and remote-sensing methods, reached 25% in some substrata but was always less than 10% for living coral (99% of which consisted of Porites spp.). At the scale of the reef (all ten 25-m² quadrats), however, disparities in percent cover between imagery and field data were less than 10% for all substrata and extremely low for some classes (e.g. <3% for living Porites, recently dead Porites and Halimeda). The least accurately estimated substrata were sand and coralline red algae, which were overestimated by absolute values 7.9% and 6.6%, respectively. The precision of sampling was similar for field and remote-sensing methods: field methods required 19 plots to detect a 10% difference in coral cover among three reefs with a statistical power of 95%. Remote-sensing methods required 21 plots. However, it took 1 h to acquire imagery over 92,500 m² of reef, which represents 3,700 plots of 25 m² each, compared with 3 days to survey 10 such plots underwater. There were no significant differences in accuracy between 1-m² and 0.25-m² image resolutions, suggesting that the advantage of using smaller pixels is offset by reduced spectral information and an increase in noise (noise was observed to be 1.6–1.8 times greater in 0.25-m² pixels). We show that airborne remote sensing can be used to monitor coral and algal cover over large areas, providing that water is shallow and clear, and that brown fleshy macroalgae are scarce, that depth is known independently (e.g. from sonar survey).     

Changes in Coral Reef Communities and an Associated Reef Fish Species, <Emphasis Type="Italic">Cephalopholis cruentata</Emphasis> (Lacépède), After 30 years on Curaçao (Netherlands Antilles)

Using the same methodology and identical sites, we repeat a study dating from 1973 and quantify cover of hard coral species, soft corals, sponges, hard substratum and soft substratum, and density of a commercially important reef fish species, the graysby Cephalopholis cruentata, along a depth-gradient of 3–36 m on the coral reefs of Curaçao. The objective was to determine the multi-decade change in benthic coral reef cover and structural complexity, and their effect on densities of an associated reef fish species. Total hard coral cover decreased on average from 52% in 1973 to 22% in 2003, representing a relative decline of 58%. During this time span, the cover of hard substratum increased considerably (from 11 to 58%), as did that of soft corals (from 0.1 to 2.2%), whereas the cover of sponges showed no significant change. Relative decline of hard coral cover and of reef complexity was greatest in shallow waters (near the coast), which is indicative of a combination of anthropogenic influences from shore and recent storm damage. Cover of main reef builder coral species (Agaricia spp., Siderastrea siderea, Montastrea annularis) decreased more than that of other species, and resulted in a significant decrease in reef complexity. Although density of C. cruentata was highly correlated to cover of Montastrea and Agaricia in 1973, the loss of coral cover did not show any effect on the total density of C. cruentata in 2003. However, C. cruentata showed a clear shift in density distribution from shallow water in 1973 to deep water in 2003. It can be concluded that the reefs of Curaçao have degraded considerably in the last three decades, but that this has had no major effect on the population size of one commercially important coral-associated fish species.     

Ningaloo Reef: Shallow Marine Habitats Mapped Using a Hyperspectral Sensor

Research, monitoring and management of large marine protected areas require detailed and up-to-date habitat maps. Ningaloo Marine Park (including the Muiron Islands) in north-western Australia (stretching across three degrees of latitude) was mapped to 20 m depth using HyMap airborne hyperspectral imagery (125 bands) at 3.5 m resolution across the 762 km² of reef environment between the shoreline and reef slope. The imagery was corrected for atmospheric, air-water interface and water column influences to retrieve bottom reflectance and bathymetry using the physics-based Modular Inversion and Processing System. Using field-validated, image-derived spectra from a representative range of cover types, the classification combined a semi-automated, pixel-based approach with fuzzy logic and derivative techniques. Five thematic classification levels for benthic cover (with probability maps) were generated with varying degrees of detail, ranging from a basic one with three classes (biotic, abiotic and mixed) to the most detailed with 46 classes. The latter consisted of all abiotic and biotic seabed components and hard coral growth forms in dominant or mixed states. The overall accuracy of mapping for the most detailed maps was 70% for the highest classification level. Macro-algal communities formed most of the benthic cover, while hard and soft corals represented only about 7% of the mapped area (58.6 km²). Dense tabulate coral was the largest coral mosaic type (37% of all corals) and the rest of the corals were a mix of tabulate, digitate, massive and soft corals. Our results show that for this shallow, fringing reef environment situated in the arid tropics, hyperspectral remote sensing techniques can offer an efficient and cost-effective approach to mapping and monitoring reef habitats over large, remote and inaccessible areas.     

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     

Using Agent-Based Models to Aid Reef Restoration: Enhancing Coral Cover and Topographic Complexity through the Spatial Arrangement of Coral Transplants

High coral cover and topographic complexity are favorable qualities of a healthy coral reef. Because coral reef restoration is expensive and coral growth is naturally slow, there is a need to strategically arrange coral transplants to maximize coral cover and topographic complexity. Similarly, it is important to understand how differences in the life history characteristics of coral transplants can influence changes in the structural attributes of coral reefs. This study utilizes agent‐based computer modeling to explore the different spatial scenarios of coral transplantation using corals with contrasting r‐ and K‐selected life histories. Spatial indexes are used to compare coral cover and topographic complexity at incremental time scales, within which disturbance events are of minor importance in spatial structuring. The outcomes of the model suggest that even‐spaced grided transplanting arrangements provide the fastest increase in coral cover and three‐dimensional habitat space (topographic complexity) across large temporal scales (<30 years) for corals with r‐selected life history strategies.     

Live coral repels a common reef fish ectoparasite

Coral reefs are undergoing rapid changes as living corals give way to dead coral on which other benthic organisms grow. This decline in live coral could influence habitat availability for fish parasites with benthic life stages. Gnathiid isopod larvae live in the substratum and are common blood-feeding parasites of reef fishes. We examined substrate associations and preferences of a common Caribbean gnathiid, Gnathia marleyi. Emergence traps set over predominantly live coral substrata captured significantly fewer gnathiids than traps set over dead coral substrata. In laboratory experiments, gnathiids preferred dead coral and sponge and tended to avoid contact with live coral. When live gnathiids were added to containers with dead or live coral, significantly fewer were recovered from the latter after 24 h. Our data therefore suggest that live coral is not suitable microhabitat for parasitic gnathiid isopods and that a decrease in live coral cover increases available habitat for gnathiids.     

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.

     

Linking Demographic Processes of Juvenile Corals to Benthic Recovery Trajectories in Two Common Reef Habitats

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.     

Coral reef distribution, status and geomorphology–biodiversity relationship in Kuna Yala (San Blas) archipelago, Caribbean Panama

Most of the knowledge of the reef geomorphology and benthic communities of Kuna Yala coral reefs (Caribbean Panama) comes from the western side of the archipelago, a few tens of kilometers around Punta San Blas (Porvenir). To bridge the gap between Porvenir and the Colombia–Panama border, we investigated with Landsat images the extent and geomorphological diversity of the entire Kuna Yala to provide geomorphologic maps of the archipelago in 12 classes. In addition to remote sensing data, in situ survey conducted in May–June 2001 provided a Kuna Yala-wide first synoptic vision of reef status, in terms of benthic diversity (number of species of coral, octocorals, and sponges) and reef health (coral versus algal cover). For a total reef system estimated to cover 638 km² along 480 km of coastline, 195 km² include coral dominated areas and only 35 km² can be considered covered by corals. A total of 69 scleractinian coral, 38 octocoral, and 82 sponge species were recorded on the outer slopes of reef formations, with a slightly higher diversity in the area presenting the most abundant and diverse reef formations (western Kuna Yala). Attempts to relate benthic diversity and geomorphological diversity provided only weak relationships regardless of the taxa, and suggest that habitat heterogeneity within geomorphological areas explain better the patterns of coral diversity. This study confirms the potential of combined remote sensing and in situ surveys for regional scale assessment, and we suggest that similar approaches should be generalized for reef mapping and assessment for other reef sites.     

Review of coral reef ecosystem remote sensing

Coral reef communities face unprecedented pressures at local, regional and global scales as a consequence of climate change and anthropogenic disturbance. Remote sensing, from satellites or aircraft, is possibly the only means to measure the effects of such stresses at appropriately large spatial scales. In the past 30 years, remote sensing of coral reefs has made rapid progress. However, the current technology is still not mature enough to monitor complicated coral reef ecosystems. Compared with foreign research in this field, our work lags far behind. There are still deficiencies in many aspects, such as basic data collection, theoretical research and platform construction. In our nation, it is even unclear how coral reefs disperse and where they may be unhealthy. In this paper, general characteristics of coral reef ecosystems and spectral features of different reef benthos have been summarized, based initially on a review of relevant literature in recent years. Based on the spectral separability of different reef types or benthos, remote sensing can be used to monitor two aspects of coral reefs: (1) Measurement of the ecological properties of reefs. (2) Health assessment of the coral reef ecosystem. In the first part, optical remote sensing methods are widely used to map reef geomorphology and habitats or biotopes. The investigation of geomorphologic zonation has proven to be one of the most successful applications, as different geomorphologic zones are associated with characteristic benthic community structures and occur at spatial scales of tens to hundreds of meters, they are amenable to remote detection by moderate to high resolution sensors. With more and more attention on the ecological problems of coral reefs, a number of studies have used high resolution sensors to map reef communities. The number of classes distinguishable depends on many factors, including the platforms, resolution (spectral, spatial and temporal resolution) and environmental conditions (water depth, water clarity, surface roughness, etc.). Compared with deep water color remote sensing, or terrestrial remote sensing, three techniques for the measurement of reef ecological properties are examined in this paper: (1) Coral reef classification system using remote sensing. (2) Techniques of sea surface correction and water column correction. (3) Techniques of coral reef information extraction from images. In terms of the complexity of coral reef ecosystems, the current techniques still need further improvement or optimization. In the health assessment of coral reef ecosystems, there are two ways to carry out the monitoring using remote sensing: (1) Monitoring the pigment or symbiotic zooxanthellae contents in corals. (2) Measuring the environmental properties of reefs. The first way is theoretically feasible, but difficult to achieve in practice. Currently, most reef health assessments are carried out by measuring environmental parameters, including sea surface temperature, solar radiation, ultraviolet radiation, water color, wind speed and direction, rainfall, ocean acidification, sea level, etc., of which sea surface temperature has been routinely measured by NOAA to monitor coral bleaching. In addition to the contents above, this article puts forward five main prospects for development in the future: (1) Establishment of a coral reef classification system using remote sensing. (2) Satellite launch for monitoring coral reefs. (3) Theoretical and methodological development. (4) Establishment of a spectral database for different reef benthos. (5) Integrated application of multi-source remote sensing data. It is hoped that the information provided here will be a reference for subsequent similar studies.     

Effects of territorial damselfish on an algal-dominated coastal coral reef

Territorial damselfish are important herbivores on coral reefs because they can occupy a large proportion of the substratum and modify the benthic community to promote the cover of food algae. However, on coastal coral reefs damselfish occupy habitats that are often dominated by unpalatable macroalgae. The aim of this study was to examine whether damselfish can maintain distinctive algal assemblages on a coastal reef that is seasonally dominated by Sargassum (Magnetic Island, Great Barrier Reef). Here, three abundant species (Pomacentrus tripunctatus, P. wardi and Stegastes apicalis) occupied up to 60% of the reef substrata. All three species promoted the abundance of food algae in their territories. The magnitudes of the effects varied among reef zones, but patterns were relatively stable over time. Damselfish appear to readily co-exist with large unpalatable macroalgae as they can use it as a substratum for promoting the growth of palatable epiphytes. Damselfish territories represent patches of increased epiphyte load on macroalgae, decreased sediment cover, and enhanced cover of palatable algal turf.     

不同生长状态珊瑚光谱特征

珊瑚礁生态系统迅速退化是目前重要的生态环境问题之一,应用遥感技术监测大范围珊瑚礁的结构组成和变迁有很大的潜力。珊瑚光谱响应特征受珊瑚生态习性影响,在光学上相似而容易造成混淆误判。采集了西沙群岛大量石珊瑚样品的光谱,对其光谱特征进行分析及成因探讨。通过导数光谱、主成分分析研究了不同生长状态珊瑚的光谱差异,并建立珊瑚生长状态高光谱遥感判别准则。结果表明,珊瑚的光谱特性及其变化均较为复杂,受珊瑚种类和生长环境影响,光谱形状主要由共生藻色素吸收决定的。结合520—530 nm、564—574 nm和600—605 nm的导数光谱可以区分健康珊瑚、白化珊瑚和藻类覆盖的死珊瑚。总体判定准确度优于80%,误判的主要来源是种内珊瑚反射率差异。研究表明珊瑚礁环境高光谱遥感可以定量评估珊瑚状态的变化。     

Mesophotic communities of the insular shelf at Tutuila, American Samoa

An investigation into the insular shelf and submerged banks surrounding Tutuila, American Samoa, was conducted using a towed camera system. Surveys confirmed the presence of zooxanthellate scleractinian coral communities at mesophotic depths (30–110 m). Quantification of video data, separated into 10-m-depth intervals, yielded a vertical, landward-to-seaward and horizontal distribution of benthic assemblages. Hard substrata composed a majority of bottom cover in shallow water, whereas unconsolidated sediments dominated the deep insular shelf and outer reef slopes. Scleractinian coral cover was highest atop mid-shelf patch reefs and on the submerged bank tops in depths of 30–50 m. Macroalgal cover was highest near shore and on reef slopes approaching the bank tops at 50–60 m. Percent cover of scleractinian coral colony morphology revealed a number of trends. Encrusting corals belonging to the genus Montipora were most abundant at shallow depths with cover gradually decreasing as depth increased. Massive corals, such as Porites spp., displayed a similar trend. Percent cover values of plate-like corals formed a normal distribution, with the highest cover observed in the 60–70 m depth range. Shallow plate-like corals belonged mostly to the genus Acropora and appeared to be significantly prevalent on the northeastern and eastern banks. Deeper plate-like corals on the reef slopes were dominated by Leptoseris, Pachyseris, or Montipora genera. Branching coral cover was high in the 80–110 m depth range. Columnar and free-living corals were also occasionally observed from 40–70 m.     

Specificity of larval settlement of the Caribbean Orange Icing Sponge,Mycale laevis

The Caribbean sponge Mycale laevis is often found growing in close proximity to living scleractinian corals. This commonly observed sponge–coral association has been considered a mutualism, with the coral providing substratum for the sponge, and the sponge protecting the coral skeleton from boring organisms. We examined the specificity of sponge recruitment to live corals, expecting a positive and specific settlement response if a mutualism exists. Benthic surveys conducted off Key Largo, Florida, and Bocas del Toro, Panama, revealed that individuals of M. laevis grew on substrata that included dead coral and other species of sponges. Selectivity analysis indicated that at three of the four survey sites, M. laevis was not randomly distributed, but associated with live corals more frequently than expected from proportional coral cover. However, settlement assays demonstrated that larvae of M. laevis did not preferentially respond to the presence of live coral. We have previously demonstrated that adults of M. laevis are chemically undefended and readily eaten by spongivorous fishes unless protected by adjacent substrata such as live corals. In overfished areas, where spongivore density is low, the sponge is not selectively distributed near corals. Initial results of settlement experiments with different substrata suggested that larvae of M. laevis responded positively to the presence of the chemically defended sponge Amphimedon compressa, perhaps indicating an associational defense. Further experiments revealed that larvae were reacting to artificially high concentrations of exudates from cut surfaces of Am. compressa; settlement was not enhanced in response to healed pieces of Am. compressa. In addition, the larvae of M. laevis did not selectively respond to live coral or to chemically defended heterospecifics. These results indicate that the commonly observed proximity of M. laevis to live corals is not driven by larval settlement behavior, but instead by post‐settlement mortality due to predation.     

A review of macrosymbiosis in the coral reef ecosystem

Symbiotic associations are very prevalant in the coral reef ecosystem. Reasons may include the innate high species diversity, the dominance of the substratum by animals, and the complex nutrient recycling which characterize the system. Corals in particular host many symbiont species. They occupy much of the substratum, their branches and folds provide shelter, nematoblasts and enveloping skeletons provide protection, and mucus and rapidly regenerating tissues provide food.Some corals may benefit from the associations. For example, commensal worms and small crustanceans may attack the coral predator Acanthaster planciand drive them away from their hosts. The solitary coral Heteropsammia has gained mobility from an association with a worm, and can thereby exploit soft muddy bottoms which smother most other corals.In the barnacles, seemingly bound by a rigid morphology, is illustrated the great range of symbiotic associations to be found on the reef. Many species exploit the corals for substrata and continue to feed on plankton but one species has taken advantage of the proximity of fast-growing host tissue and become parasitic.Other barnacles have exploited motile species of the benthos and nekton. These gain not only a substratum but mobility, freedom from predators and competitors, feeding currents, and even scraps of food from their hosts. Platylepas ophiophilus. the sea snake barnacle, appears to have been forced into its specialized niche by more efficient competitors. Some have been parasitic; the Ascothoracids and Rhizocephalans are among the most specialized and degenerative of all parasites.     

The influence of habitat and adults on the spatial distribution of juvenile corals

Population distributions are affected by a variety of spatial processes, including dispersal, intraspecific dynamics and habitat selection. Within reef‐building coral communities, these processes are especially important during the earliest life stages when reproduction provides mobility among sessile organisms and populations experience the greatest mortality bottlenecks both before and immediately after settlement. Here, we used large‐area imaging to create photomosaics that allowed us to identify and map the location of 4681 juvenile (1–5 cm diameter) and 25 902 adult (>5 cm diameter) coral colonies from eight 100‐m² plots across the forereef of Palmyra Atoll. Using metrics of density, percent cover and the relative location of each colony within each plot, we examined abundance and spatial relationships between juvenile and adult coral taxa. Within coral taxa, juvenile density was generally positively related to the numerical density and percent cover of adults. Nearest neighbor analyses showed aggregation of juveniles near adults of the same taxon for two of the focal taxa (Pocillopora and Fungiids), while all other taxa showed random spatial patterning relative to adults. Three taxa had clustered distributions of juveniles overall. Additionally, we found that on a colony level, juveniles for five of nine focal taxa (accounting for >98% of all identified juveniles) associated with a specific habitat type, with four of those five taxa favoring unconsolidated (e.g. rubble) over consolidated substrata. The general lack of clustering in juvenile corals contrasts with consistent clustering patterns seen in adult corals, suggesting that adult spatial patterns are largely driven by processes occurring after maturity such as partial colony mortality, including fission and fragmentation. The association of many taxa with unconsolidated habitat also suggests that corals may play an important role in colonizing natural rubble patches that could contribute to reef stabilization over time.     

Suppressed recovery of functionally important branching Acropora drives coral community composition changes following mass bleaching in Indonesia

Mass coral bleaching events may have disproportionate effects on branching corals, leading to coral community restructuring, reduced biodiversity, and decreased structural complexity. This affects overall reef health and resilience. Functionally important, fast-growing branching Acropora corals were a historically dominant and vital component of Indonesian reefs throughout the twentieth century, yet the genus is also one of the most vulnerable to external stressors. This study used long-term annual reef monitoring data from Indonesia’s Wakatobi Marine National Park (WMNP) to investigate the effects of a mass bleaching event in 2010 on Acropora and other branching corals, evaluate their post-disturbance recovery trajectories, and analyse shifts in coral community composition. Post-bleaching scleractinian coral cover decreased across study sites, with losses in branching corals especially evident. Long-term branching Acropora cover decreased significantly and failed to demonstrate the significant post-disturbance recovery of other branching corals (especially Porites). In areas characterised by relatively high branching Acropora cover (> 15% mean cover) prior to bleaching, long-term coral community composition changes have trended predominately towards branching and massive Porites and branching Montipora. The novelty and key contribution of this study is that results suggest suppressed recovery of Acropora in the WMNP. Contributing factors may include the Allee effect (inhibition of reproduction at low population densities), other forms of inhibited larval recruitment, direct and indirect spatial competition, and changes in the physical reef habitat. These findings have critical implications for this functionally important taxon, future reef conservation efforts, and overall reef health and resilience in the park.

     

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.     

Depth distributions of coral reef fishes: the influence of microhabitat structure,settlement, and post-settlement processes

Many coral reef fishes have restricted depth ranges that are established at settlement or soon after, but the factors limiting these distributions are largely unknown. This study examines whether the availability of microhabitats (reef substrata) explains depth limits, and evaluates whether juvenile growth and survival are lower beyond these limits. Depth-stratified surveys of reef fishes at Kimbe Bay (Papua New Guinea) showed that the abundance of new settlers and the cover of coral substrata differed significantly among depths. A field experiment investigated whether settling coral reef fishes preferred particular depths, and whether these depth preferences were dependent on microhabitat. Small patch reefs composed of identical coral substrata were set up at five depths (3, 6, 10, 15 and 20 m), and settlement patterns were compared to those on unmanipulated reef habitat at the same five depths. For all species, settlement on patch reefs differed significantly among depths despite uniform substratum composition. For four of the six species tested, depth-related settlement patterns on unmanipulated habitat and on patch reefs did not differ, while for the other two, depth ranges were greater on the patch reefs than on unmanipulated habitat. A second experiment examined whether depth preferences reflected variation in growth and survival when microhabitat was similar. Newly settled individuals of Chrysiptera parasema and Dascyllus melanurus were placed, separately, on patch reefs at five depths (as above) and their survival and growth monitored. D. melanurus, which is restricted to shallow depths, had highest survival and growth at the shallowest depth. Depth did not affect either survival or growth of C. parasema, which has a broader depth range than D. melanurus (between 6 and 15 m). This suggests that the fitness costs potentially incurred by settling outside a preferred depth range may depend on the strength of the depth preference.     

Recruitment of scleractinians onto the skeletons of corals killed by black band disease

To determine what happens to scleractinian corals that have been killed by black band disease (BBD), massive corals with BBD were monitored for 11 years on a shallow reef (<10 m depth) in St. John, US Virgin Islands. Small quadrats (0.039 m²) were used to compare the rates of scleractinian recruitment to the skeletons of corals killed by either BBD or physical disturbance (Hurricane Hugo 1989). Coral recruitment was also quantified on the adjacent fringing reef using larger quadrats (0.25 m²) to detect possible biases associated with using small, permanent quadrats to assess recruitment to BBD-killed corals. Of 28 tagged colonies with BBD in 1988, 43% were lost to Hurricane Hugo in 1989, 7% were lost to unknown causes between 1991 and 1992, and 14 were monitored annually for 11 years; of these, 71% were dead and still in their original growth position in 1998. Between 1988 and 1997, corals recruited to the BBD-killed surfaces at a rate of 1.1 ± 0.3 recruits · 0.039 m⁻² · decade⁻¹ (mean ± SE, n = 14), although mortality reduced the density to 0.3 ± 0.2 recruits · 0.039 m⁻² by 1997. The rate of recruitment and the taxonomic composition of the coral recruits to BBD-killed corals were indistinguishable statistically from those to corals killed by Hurricane Hugo. This demonstrates that BBD creates space that is functionally the same as other dead coral surfaces in providing a substratum for coral recruitment. However, because coral recruits are dispersed widely, clumped in distribution and temporally variable in density on the fringing reef as a whole, it is unlikely that they will be found on monitored coral colonies that have been killed by BBD. While this hypothesis is consistent with the higher density of recruits on the fringing reef compared with BBD-killed corals, further studies are required to investigate alternative explanations such as the role of substratum age in favoring recruitment to surfaces other than those killed recently by BBD. Accepted: 26 August 1999