Pleistocene reef environments, constituent grains, and coral community structure: Curaçao, Netherlands Antilles

We investigated the degree to which component grains vary with depositional environment in sediments from three reef habitats from the Pleistocene (125?ka) Hato Unit of the Lower Terrace, Curaçao, Netherlands Antilles: windward reef crest, windward back reef, and leeward reef crest. The windward reef crest sediment is the most distinctive, dominated by fragments of encrusting and branching coralline red algae, coral fragments and the encrusting foraminiferan Carpenteria sp. Windward back reef and leeward reef crest sediments are more similar compositionally, only showing significant differences in relative abundance of coral fragments and Homotrema rubrum. Although lacking high taxonomic resolution and subject to modification by transport, relative abundance of constituent grain types offers a way of assessing ancient skeletal reef community composition, and one which is not limited to a single taxonomic group. The strong correlation between grain type and environment we found in the Pleistocene of Curaçao suggests that constituent grain analysis may be an effective tool in delineating Pleistocene Caribbean reef environments. However, it will not be a sufficient indicator where communities vary significantly within reef environments or where evolutionary and/or biogeographical processes lead to different relationships between community composition and reef environment. Detailed interpretation of geological, biological, and physical characteristics of the Pleistocene reefs of Curaçao reveals that the abundance of the single coral species, Acropora palmata, is not a good predictor of the ecological structure of the ancient reef coral communities. This coral was the predominant species in two of the three reef habitats (windward and leeward reef crest), but the taxonomic composition (based on species relative abundance data) of the reef coral communities was substantially different in these two environments. We conclude that qualitative estimates of coral distribution patterns (presence of a key coral species or the use of a distinctive coral skeletal architecture), when used as a component in a multi-component analysis of ancient reef environments, probably introduces minimal circular reasoning into quantitative paleoecological studies of reef coral community structure.     

Composition of Archaea in Seawater,Sediment, and Sponges in the Kepulauan Seribu Reef System,Indonesia

Coral reefs are among the most diverse and productive ecosystems in the world. Most research has, however, focused on eukaryotes such as corals and fishes. Recently, there has been increasing interest in the composition of prokaryotes, particularly those inhabiting corals and sponges, but these have mainly focused on bacteria. There have been very few studies of coral reef Archaea, despite the fact that Archaea have been shown to play crucial roles in nutrient dynamics, including nitrification and methanogenesis, of oligotrophic environments such as coral reefs. Here, we present the first study to assess Archaea in four different coral reef biotopes (seawater, sediment, and two sponge species, Stylissa massa and Xestospongia testudinaria). The archaeal community of both sponge species and sediment was dominated by Crenarchaeota, while the seawater community was dominated by Euryarchaeota. The biotope explained more than 72 % of the variation in archaeal composition. The number of operational taxonomic units (OTUs) was highest in sediment and seawater biotopes and substantially lower in both sponge hosts. No “sponge-specific” archaeal OTUs were found, i.e., OTUs found in both sponge species but absent from nonhost biotopes. Despite both sponge species hosting phylogenetically distinct microbial assemblages, there were only minor differences in Kyoto Encyclopedia of Genes and Genomes (KEGG) functional pathways. In contrast, most functional pathways differed significantly between microbiomes from sponges and nonhost biotopes including all energy metabolic pathways. With the exception of the methane and nitrogen metabolic pathway, all energy metabolic pathways were enriched in sponges when compared to nonhost biotopes.     

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.     

Mangrove and coral reef sponge faunas: untold stories about shallow water Porifera in the Caribbean

Sponge faunas from coral reefs and mangrove ecosystems in the Caribbean have mostly been studied from an ecological perspective, with researchers considering the effects of physical and biological factors on their species distribution. To discern evolutionary patterns, this study analyzed the systematic composition, taxonomic diversity, and ecological properties (reproductive strategies, size, shape, endosymbiosis) of mangrove and reef sponge assemblages from seven distant Caribbean localities. Species composition was compared by use of cluster analysis (Sørensen’s), and taxonomic diversity by use of the biodiversity index average taxonomic distinctness (AvTD). Mangrove and reef-associated sponge faunas were found to be statistically dissimilar, with the AvTD values suggesting stronger taxonomic bias toward specific groups in mangroves, irrespective of geographic distance. Most Demospongiae orders have 30–50% more species in coral reefs than in mangroves. The richest reef genera (Agelas, Aplysina, Callyspongia, Petrosia, and Xestospongia) rarely colonize contiguous mangrove formations. The distribution and diversity of suprageneric taxa suggest that coral reef sponge assemblages might represent an older fauna. This historical interpretation would place mangrove subtidal habitats as the youngest marine ecosystem, rather than a below-optimum ecosystem. Life history traits support a biological split discussed here from the perspective of distinct evolutionary histories and different environmental conditions.     

Modelling sponge species diversity using a morphological predictor: a tropical test of a temperate model

The contribution of sponges to marine surveys is often underestimated due to problems of identification, synonymous species and limited numbers of specialists in the field. Bell & Barnes (2001) illustrated how sponge morphological diversity (diversity of body forms) might be used as a predictor of sponge species diversity and richness. This study investigated these relationships at six tropical West Indian Ocean localities in a number of habitat types. These habitats included tropical coral reefs, soft substratum (seagrass, mangrove and sand), caves and boulders. Sampling was undertaken at three depth zones in coral reef habitats only (intertidal, 10–15 m and 20–25 m), with the other habitats sampled in less than 10m of water. Species diversity and richness were significantly correlated (P < 0.05) with morphological diversity at all localities and depths in coral reef and soft substratum habitats. However, no significant correlation was found between these variables in cave or boulder habitats. The slope of the linear regression found between morphological diversity and species diversity did not significantly differ between coral reef, soft substratum and temperate reef (data taken from Bell & Barnes 2001) habitats. Similarly coral reefs showed the same relationship between morphological diversity and species richness as temperate reefs, however the relationship between morphological diversity and species richness was significantly different at both habitats compared with soft substratum environments. Sponge morphological diversity therefore may be more useful as a predictor of sponge species diversity, rather than species richness, as the former relationship is common between more habitats than the latter.     

Co-occurrence of herbivorous fish functional groups correlates with enhanced coral reef benthic state

Aim

Biodiversity loss is impacting essential ecosystem functions and services across the globe. Recently, our interest in the benefits of biodiversity for ecosystem function has shifted focus from measurements of species richness to functional diversity and composition. However, the additional importance of other community characteristics, such as species evenness and co-occurrence, for diversity-driven ecosystem function is less known. We used herbivorous coral reef fish as a model system to investigate how co-occurrence of different functional groups, rather than purely functional diversity, within an assemblage might affect the coral reef benthic state.

Location

Western Atlantic.

Time period

2007–2017.

Major taxa studied

Herbivorous reef fish.

Methods

We analysed benthic and fish assemblage data from 601 sites across 12 countries in the western Atlantic. Using diversity–interaction models, we investigated how the composition and relative abundances of reef fish functional groups were correlated with benthic cover and estimates of coral calcification rates. We used statistical interactions to explore the importance of co-occurrence of herbivorous fish functional groups for the coral reef benthic state.

Results

We found that co-occurrence of herbivorous fish functional groups, in addition to functional diversity, was correlated with reduced algal cover and increased coral accretion. Moreover, pairwise statistical interactions between functional groups were significantly correlated with an improvement in the coral reef benthic state.

Main conclusions

Our results support the idea that functional group co-occurrence, in addition to functional diversity, within herbivorous fish offers additional benefits to the coral reef benthic state. We identify farming damselfish and excavating parrotfish as potential key determinants of the coral reef benthic state and highlight that co-occurrence of cropping and scraping herbivores might promote coral accretion. Our findings support the argument that protecting herbivore abundance without regard to the species and functional groups present is not enough to preserve coral reef health and that fine-scale community composition must be considered.     

Disentangling the complex microbial community of coral reefs using standardized Autonomous Reef Monitoring Structures (ARMS)

Autonomous Reef Monitoring Structures (ARMS) have been applied worldwide to describe eukaryotic cryptic reef fauna. Conversely, bacterial communities, which are critical components of coral reef ecosystem functioning, remain largely overlooked. Here we deployed 56 ARMS across the 2,000‐km spread of the Red Sea to assay biodiversity, composition and inferred underlying functions of coral reef‐associated bacterial communities via 16S rRNA gene sequencing. We found that bacterial community structure and diversity aligned with environmental differences. Indeed, sea surface temperature and macroalgae cover were key in explaining bacterial relative abundance. Importantly, taxonomic and functional alpha diversity decreased under more extreme environmental conditions (e.g., higher temperatures) in the southern Red Sea. This may imply a link between bacterial community diversity and functional capabilities, with implications for conservation management. Our study demonstrates the utility of ARMS to investigate the response of coral reef‐associated bacterial communities to environmental change.     

Sedimentation on three caribbean atolls: Glovers reef,lighthouse reef and turneffe Islands,belize

Summary The chief mode of carbonate sedimentation on the Belizean atolls Glovers Reef, Lighthouse Reef and Turneffe Islands is the accumulation of organically-derived particles. Variations in the distribution of the composition and grain-sizes of surface sediments, collected along transects across the atolls, are environmentally controlled. Two major sediment types may be distinguished. (1) Reef and fore reef sediments are dominated by fragments of coral, coralline algae andHalimeda. Mean grain-sizes range from 1–2 mm. (2) Back reef sediments contain more mollusk fragments, more fine-grained sediment (<125 μm) and appear to have fewerHalimeda fragments. In addition, sediments from inner platforms and shallow lagoonal parts of Glovers and Lighthouse Reefs comprise non-skeletal grains, namely fecal pellets. Sediments from lagoonal patch reefs may contain up to 20% coral fragments. Mean grain-sizes range from 0.1–1 mm and are finest on the inner platform and lagoon floor of the back reef environment. Within the reef and fore reef environments, it is not possible to distinguish sub-environments on the basis of textural and compositional differences of the sediments. Sediments from patch reefs contrast with those from back reef lagoons and inner platforms and are similar in terms of grain-sizes and compositions to reef and fore reef surface sediments. Non-skeletal grains forming in shallow parts of the back reef in Glovers and Lighthouse Reefs are interpreted to be indurated by interstitial precipitation of calcium carbonate from warm, supersaturated water flushing the sediment. The lack of hardened non-skeletal particles in the back reef sediments of Turneffe Islands is most probably due to the abundance of muddy, organic-rich sediment in the well-protected lagoon. Fine sediment is less permeable and organic films prevent cement overgrowth on particles.     

Controls on coral-ground development along the northern Mesoamerican Reef tract

Coral-grounds are reef communities that colonize rocky substratum but do not form framework or three-dimensional reef structures. To investigate why, we used video transects and underwater photography to determine the composition, structure and status of a coral-ground community located on the edge of a rocky terrace in front of a tourist park, Xcaret, in the northern Mesoamerican Reef tract, Mexico. The community has a relatively low coral, gorgonian and sponge cover (<10%) and high algal cover (>40%). We recorded 23 species of Scleractinia, 14 species of Gorgonacea and 30 species of Porifera. The coral community is diverse but lacks large coral colonies, being dominated instead by small, sediment-tolerant, and brooding species. In these small colonies, the abundance of potentially lethal interactions and partial mortality is high but decreases when colonies are larger than 40 cm. Such characteristics are consistent with an environment control whereby storm waves periodically remove larger colonies and elevate sediment flux. The community only survives these storm conditions due to its slope-break location, which ensures lack of burial and continued local recruitment. A comparison with similar coral-ground communities in adjacent areas suggests that the narrow width of the rock terrace hinders sediment stabilization, thereby ensuring that communities cannot escape bottom effects and develop into three-dimensional reef structures on geological time scales.     

Relative Importance of Coral Cover,Habitat Complexity and Diversity in Determining the Structure of Reef Fish Communities

The structure of coral reef habitat has a pronounced influence on the diversity, composition and abundance of reef-associated fishes. However, the particular features of the habitat that are most critical are not always known. Coral habitats can vary in many characteristics, notably live coral cover, topographic complexity and coral diversity, but the relative effects of these habitat characteristics are often not distinguished. Here, we investigate the strength of the relationships between these habitat features and local fish diversity, abundance and community structure in the lagoon of Lizard Island, Great Barrier Reef. In a spatial comparison using sixty-six 2m² quadrats, fish species richness, total abundance and community structure were examined in relation to a wide range of habitat variables, including topographic complexity, habitat diversity, coral diversity, coral species richness, hard coral cover, branching coral cover and the cover of corymbose corals. Fish species richness and total abundance were strongly associated with coral species richness and cover, but only weakly associated with topographic complexity. Regression tree analysis showed that coral species richness accounted for most of the variation in fish species richness (63.6%), while hard coral cover explained more variation in total fish abundance (17.4%), than any other variable. In contrast, topographic complexity accounted for little spatial variation in reef fish assemblages. In degrading coral reef environments, the potential effects of loss of coral cover and topographic complexity are often emphasized, but these findings suggest that reduced coral biodiversity may ultimately have an equal, or greater, impact on reef-associated fish communities.     

The influence of coral reef benthic condition on associated fish assemblages

Accumulative disturbances can erode a coral reef's resilience, often leading to replacement of scleractinian corals by macroalgae or other non-coral organisms. These degraded reef systems have been mostly described based on changes in the composition of the reef benthos, and there is little understanding of how such changes are influenced by, and in turn influence, other components of the reef ecosystem. This study investigated the spatial variation in benthic communities on fringing reefs around the inner Seychelles islands. Specifically, relationships between benthic composition and the underlying substrata, as well as the associated fish assemblages were assessed. High variability in benthic composition was found among reefs, with a gradient from high coral cover (up to 58%) and high structural complexity to high macroalgae cover (up to 95%) and low structural complexity at the extremes. This gradient was associated with declining species richness of fishes, reduced diversity of fish functional groups, and lower abundance of corallivorous fishes. There were no reciprocal increases in herbivorous fish abundances, and relationships with other fish functional groups and total fish abundance were weak. Reefs grouping at the extremes of complex coral habitats or low-complexity macroalgal habitats displayed markedly different fish communities, with only two species of benthic invertebrate feeding fishes in greater abundance in the macroalgal habitat. These results have negative implications for the continuation of many coral reef ecosystem processes and services if more reefs shift to extreme degraded conditions dominated by macroalgae.     

Mid-Holocene coral community data as baselines for understanding contemporary reef ecological states

Land-use changes and associated deteriorations in water quality are cited as major drivers of marine ecosystem change, and can modify community abundance and diversity on coral reefs. This study uses palaeoecological data derived from a mid-Holocene age coral reef in the Wet Tropics region of Australia's Great Barrier Reef to develop a record of coral community composition and diversity, from a period that significantly pre-dates European settlement in the region. Major changes in catchment sediment and nutrient yields since European settlement have been documented, and thus the data presented provides a baseline against which to compare contemporary ecological datasets. Natural variations in coral assemblage composition, as preserved in core records, clearly occurred in this mid-Holocene reef and were associated with the reef shallowing to sea level as it accreted vertically. Comparisons between modern and mid-Holocene coral community data from equivalent water depths did not reveal marked shifts in coral community composition and diversity, suggesting the long-term persistence of a resilient coral assemblage over these time periods.     

The influence of multiple factors upon reef fish abundance and species richness in a tropical coral complex

The present study was conducted on Tamandaré reefs, northeast Brazil and aimed to analyse the importance of different factors (e.g. tourism activity, fishing activity, coral abundance and algal abundance) on reef fish abundance and species richness. Two distinct reef areas (A ver o mar and Caieiras) with different levels of influence were studied. A total of 8239 reef fish individuals were registered, including 59 species. Site 1 (A ver o mar) presented higher reef fish abundance and richness, with dominance of roving herbivores (29.9 %) and mobile invertebrate feeders (28.7 %). In contrast, at Site 2 (Caieiras) territorial herbivores (40.9 %) predominated, followed by mobile invertebrate feeders (24.6 %). Concerning the benthic community, at Site 1 macroalgae were recorded as the main category (49.3 %); however, Site 2 was dominated by calcareous algae (36.0 %). The most important variable explaining more than 90 % of variance on reef fish abundance and species richness was macroalgae abundance, followed by fishing activity. Phase shifts on coral reefs are evident, resulting in the replacement of coral by macroalgae and greatly influencing reef fish communities. In this context, it is important to understand the burden of the factors that affect reef fish communities and, therefore, influence the extinction vulnerability of coral reef fishes.     

A unified model explains commonness and rarity on coral reefs

Abundance patterns in ecological communities have important implications for biodiversity maintenance and ecosystem functioning. However, ecological theory has been largely unsuccessful at capturing multiple macroecological abundance patterns simultaneously. Here, we propose a parsimonious model that unifies widespread ecological relationships involving local aggregation, species‐abundance distributions, and species associations, and we test this model against the metacommunity structure of reef‐building corals and coral reef fishes across the western and central Pacific. For both corals and fishes, the unified model simultaneously captures extremely well local species‐abundance distributions, interspecific variation in the strength of spatial aggregation, patterns of community similarity, species accumulation, and regional species richness, performing far better than alternative models also examined here and in previous work on coral reefs. Our approach contributes to the development of synthetic theory for large‐scale patterns of community structure in nature, and to addressing ongoing challenges in biodiversity conservation at macroecological scales.     

Factors influencing spatial distribution of fish communities on a fringing reef at Mauritius, S.W. Indian Ocean

This paper analyses the spatial patterns of fish communities on a narrow fringing reef of Mauritius and identifies some controlling factors. Canonical Correspondence Analysis revealed that the location in the reef ecosystem, the living coral coverage, and the proximity to a dredged area were the major factors controlling the distribution and abundance of fish species. Two distinct communities were observed across the fringing reef, representing a land-ocean gradient. The low diversity of fish species, and also of invertebrates such as corals, molluscs, and echinoderms is probably a consequence of the narrowness of the fringing reef and of anthropogenic impacts, notably the high pollution by nutrients.     

Parrotfish abundance and selective corallivory on a Belizean coral reef

Parrotfish are important members of coral reef communities because they consume macroalgae that would otherwise outcompete reef-building corals for space. However, some Caribbean parrotfish species also feed directly on live corals, and thus have the potential to negatively impact coral fitness and survival. This study investigates selective grazing by parrotfish on particular coral species, differences in grazing incidence among reef habitats and intraspecific discrimination among colonies of several coral species. We also investigate spatial and temporal patterns of parrotfish species abundance across habitats on the Belize barrier reef, and examine correlations between parrotfish abundance and grazing intensity across reef habitats. We found that members of the Montastraea annularis species complex, major builders of Caribbean reefs, were preferred targets of parrotfish grazing across all reef habitats, while M. cavernosa, Agaricia agaricites, Diploria strigosa, Porites astreoides and Porites porites were not preferred; Siderastrea siderea was preferentially grazed only in the spur and groove habitats. Parrotfish grazing preferences varied across habitats; M. annularis was grazed most often in shallow habitats, whereas M. franksi was consumed more at depth. Although it was not possible to directly observe parrotfish grazing on corals, we did find a positive correlation between Sparisoma aurofrenatum abundance and M. franksi grazing incidence across habitats. Finally, when we compared our results to parrotfish abundances measured by a previous study, we found that Sparisoma viride and Sp. aurofrenatum, two species known to be corallivorous, had increased abundances between 1982 and 2004. In light of escalating threats on Caribbean reef corals, it would be important for future studies to evaluate the impact of parrotfish corallivory on coral survival.     

Variability of Symbiodinium Communities in Waters,Sediments, and Corals of Thermally Distinct Reef Pools in American Samoa

Reef-building corals host assemblages of symbiotic algae (Symbiodinium spp.) whose diversity and abundance may fluctuate under different conditions, potentially facilitating acclimatization to environmental change. The composition of free-living Symbiodinium in reef waters and sediments may also be environmentally labile and may influence symbiotic assemblages by mediating supply and dispersal. The magnitude and spatial scales of environmental influence over Symbiodinium composition in different reef habitat compartments are, however, not well understood. We used pyrosequencing to compare Symbiodinium in sediments, water, and ten coral species between two backreef pools in American Samoa with contrasting thermal environments. We found distinct compartmental assemblages of clades A, C, D, F, and/or G Symbiodinium types, with strong differences between pools in water, sediments, and two coral species. In the pool with higher and more variable temperatures, abundance of various clade A and C types differed compared to the other pool, while abundance of D types was lower in sediments but higher in water and in Pavona venosa, revealing an altered habitat distribution and potential linkages among compartments. The lack of between-pool effects in other coral species was due to either low overall variability (in the case of Porites) or high within-pool variability. Symbiodinium communities in water and sediment also showed within-pool structure, indicating that environmental influences may operate over multiple, small spatial scales. This work suggests that Symbiodinium composition is highly labile in reef waters, sediments, and some corals, but the underlying drivers and functional consequences of this plasticity require further testing with high spatial resolution biological and environmental sampling.     

Coral–algal phase shifts alter fish communities and reduce fisheries production

Anthropogenic stress has been shown to reduce coral coverage in ecosystems all over the world. A phase shift towards an algae‐dominated system may accompany coral loss. In this case, the composition of the reef‐associated fish assemblage will change and human communities relying on reef fisheries for income and food security may be negatively impacted. We present a case study based on the Raja Ampat Archipelago in Eastern Indonesia. Using a dynamic food web model, we simulate the loss of coral reefs with accompanied transition towards an algae‐dominated state and quantify the likely change in fish populations and fisheries productivity. One set of simulations represents extreme scenarios, including 100% loss of coral. In this experiment, ecosystem changes are driven by coral loss itself and a degree of habitat dependency by reef fish is assumed. An alternative simulation is presented without assumed habitat dependency, where changes to the ecosystem are driven by historical observations of reef fish communities when coral is lost. The coral–algal phase shift results in reduced biodiversity and ecosystem maturity. Relative increases in the biomass of small‐bodied fish species mean higher productivity on reefs overall, but much reduced landings of traditionally targeted species.     

Biologic response to environmental stress in tropical reefs: Lessons from modern Polynesian coralgal atolls and Middle Permian sponge and Shamovella-microbe reefs (Capitan Limestone USA)

Despite prejudices that comparisons of paleoecological patterns in modern and fossil reef communities are of doubtful validity, we compare the biologic response of living coralgal reefs in French Polynesia to environmental stress with an exceptionally well exposed Middle Permian sponge reef and Shamovella-microbial reef of the Capitan Limestone in New Mexico. In the western Tuamotu Archipelago, reef margins are characterized by depth-related changes of biodiversity. The subtidal basic reefbuilding community contains the highest diversity (23 coral and 6 calcareous algal species). With decreasing water depth and increasing environmental stress, diversity reaches a minimum of five taxa on the reef flat. The Capitan consists of two reef stages. Reefbuilders of the lowermost exposed part of Stage 1 formed a cement-rich sponge reef with 42 taxa (28 sponge species). Decreasing water depth along the reef face is accompanied by loss of five taxa, variations in the gross morphology of sponges and changes in framework architecture. Stage 2, dominated by Shamovella obscura, one bryozoan species and microbes, is sandwiched between two unconformities suggesting much shallower water and higher environmental stress. Despite differences in shelf profile and taxonomy, both the modern and Permian reefbuilders respond to increasing environmental stress with diversity impoverishment and dominance of binders.