Environmental conditions and herbivore biomass determine coral reef benthic community composition: implications for quantitative baselines

Coral Reefs - Our ability to understand natural constraints on coral reef benthic communities requires quantitative assessment of the relative strengths of abiotic and biotic processes across large...     

Coral bleaching, reef fish community phase shifts and the resilience of coral reefs

The 1998 global coral bleaching event was the largest recorded historical disturbance of coral reefs and resulted in extensive habitat loss. Annual censuses of reef fish community structure over a 12-year period spanning the bleaching event revealed a marked phase shift from a prebleach to postbleach assemblage. Surprisingly, we found that the bleaching event had no detectable effect on the abundance, diversity or species richness of a local cryptobenthic reef fish community. Furthermore, there is no evidence of regeneration even after 5–35 generations of these short-lived species. These results have significant implications for our understanding of the response of coral reef ecosystems to global warming and highlight the importance of selecting appropriate criteria for evaluating reef resilience.     

Impact of cyclones on hard coral and metapopulation structure,connectivity and genetic diversity of coral reef fish

Cyclones have one of the greatest effects on the biodiversity of coral reefs and the associated species. But it is unknown how stochastic alterations in habitat structure influence metapopulation structure, connectivity and genetic diversity. From 1993 to 2018, the reefs of the Capricorn Bunker Reef group in the southern part of the Great Barrier Reef were impacted by three tropical cyclones including cyclone Hamish (2009, category 5). This resulted in substantial loss of live habitat-forming coral and coral reef fish communities. Within 6–8 years after cyclones had devastated, live hard corals recovered by 50–60%. We show the relationship between hard coral cover and the abundance of the neon damselfish (Pomacentrus coelestis), the first fish colonizing destroyed reefs. We present the first long-term (2008–2015 years corresponding to 16–24 generations of P. coelestis) population genetic study to understand the impact of cyclones on the meta-population structure, connectivity and genetic diversity of the neon damselfish. After the cyclone, we observed the largest change in the genetic structure at reef populations compared to other years. Simultaneously, allelic richness of genetic microsatellite markers dropped indicating a great loss of genetic diversity, which increased again in subsequent years. Over years, metapopulation dynamics were characterized by high connectivity among fish populations associated with the Capricorn Bunker reefs (2200 km²); however, despite high exchange, genetic patchiness was observed with annual strong genetic divergence between populations among reefs. Some broad similarities in the genetic structure in 2015 could be explained by dispersal from a source reef and the related expansion of local populations. This study has shown that alternating cyclone-driven changes and subsequent recovery phases of coral habitat can greatly influence patterns of reef fish connectivity. The frequency of disturbances determines abundance of fish and genetic diversity within species.

     

Storm-induced coral rubble deposition: Pleistocene records of natural reef disturbance and community response.

Shallow-water Pleistocene coral reef facies in Barbados (dominated by Acropora palmata rubble) record evidence of deposition under contrasting non-catastrophic - (fair-weather?) and storm-induced conditions. Depositional styles are interpreted on the basis of coral rubble fabrics and calcareous encruster successions. Terrace exposures on the west of the island comprise uniform (3-4 m thick) depositional sequences. Individual coral samples exhibit similarities in encruster community composition and thickness, and a transition from photophilic to sciaphilic encrusting forms. These are indicative of colonization during gradual burial in an accumulating rubble pile. By contrast, NE coast sites comprise repetitive sequences of discrete (0.4-1 m thick) depositional units. Rubble colonization within each unit is characterized by a vertical succession from thin (1-2 mm), sciaphilic encruster-dominated sequences at the base, to progressively thicker (up to 20 mm), photophilic encruster-dominated sequences at the top. These are interpreted as multiple storm deposits, with upper surfaces colonized by opportunist coral species (primarily Agaricia agaricites). In contrast to many modern hurricane-impacted reef systems, however, there is no evidence of long-term shifts in coral community composition following physical disturbance. Colony sizes of opportunist corals at the tops of storm units are consistent with growth over timescales of <10 years. These are overlain on each occasion by a new A. palmata rubble pile, indicating successful recovery over successive physical disturbance cycles.     

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.     

Community structure,social organization and ecological requirements of coral reef fish (Pomacentridae)

In the Gulf of Aqaba (Red Sea), 25 species of damselfish (Pomacentridae) which form typical interspecific communities were studied. Several species are typical for different depth zones, others are ubiquitous. Damselfish are variously specialized in feeding (herbivore, omnivore, plankton-carnivore); habitat (bottom-dwelling, midwater); home-and-shelter site (selective or unselective). Environmental factors and social structures were found to be interlinked. Species with the same social structure are similarly specialized ecologically. A hypothetical flow diagram shows the effects of abiotic and biotic factors on group size, available food, and social structure. Damselfish are extremely flexible to environmental change, showing intraspecific modification of group structure, feeding habits, reproductive strategy etc. Experimental results suggest a strong selection against redundant males. This research is considered a pilot study for future field work, aimed at a prediction of environmental influences on behaviour.Supported by the Deutsche Forschungsgemeinschaft, Bad Godesberg, and the Max-Planck-Institut für Verhaltensphysiologie, Seewiesen.     

Future coral reef habitat marginality: temporal and spatial effects of climate change in the Pacific basin

Marginal reef habitats are regarded as regions where coral reefs and coral communities reflect the effects of steady-state or long-term average environmental limitations. We used classifications based on this concept with predicted time-variant conditions of future climate to develop a scenario for the evolution of future marginality. Model results based on a conservative scenario of atmospheric CO₂ increase were used to examine changes in sea surface temperature and aragonite saturation state over the Pacific Ocean basin until 2069. Results of the projections indicated that essentially all reef locations are likely to become marginal with respect to aragonite saturation state. Significant areas, including some with the highest biodiversity, are expected to experience high-temperature regimes that may be marginal, and additional areas will enter the borderline high temperature range that have experienced significant ENSO-related bleaching in the recent past. The positive effects of warming in areas that are presently marginal in terms of low temperature were limited. Conditions of the late 21st century do not lie outside the ranges in which present-day marginal reef systems occur. Adaptive and acclimative capabilities of organisms and communities will be critical in determining the future of coral reef ecosystems.Electronic supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.     

Complex migration and the development of genetic structure in subdivided populations: an example from Caribbean coral reef ecosystems

A matrix‐based projection model is used in conjunction with the results of a coupled bio‐physical dispersal model to examine the spread of alleles through subdivided populations over time, and the associated development of genetic structural patterns. Applying this approach, it becomes possible to quantitatively evaluate the contribution of spatially explicit migration towards patterns of genetic structure observed in the field. To provide a concrete example, the model was used to examine genetic dispersal between coral reef patches of the Caribbean. Using generic life‐history parameters, the model shows the formation of a strong genetic break between eastern and western patches, as well as the development of a gradient along the length of the Bahamian archipelago, corresponding with evidence previously collected for coral and fish species. The data also suggest that Jamaica and the Cayman Islands are important stepping stones between the reefs of the northern Caribbean (Hispaniola and the Bahamas) and the Mesoamerican Barrier Reef. The model provides an effective means of evaluating regional‐scale genetic connectivity through time and identifying natural clusters of genetic exchange.     

Exploring the nature of ecological specialization in a coral reef fish community: morphology,diet and foraging microhabitat use

Patterns of ecological specialization offer invaluable information about ecosystems. Yet, specialization is rarely quantified across several ecological niche axes and variables beyond the link between morphological and dietary specialization have received little attention. Here, we provide a quantitative evaluation of ecological specialization in a coral reef fish assemblage (f. Acanthuridae) along one fundamental and two realized niche axes. Specifically, we examined ecological specialization in 10 surgeonfish species with regards to morphology and two realized niche axes associated with diet and foraging microhabitat utilization using a recently developed multidimensional framework. We then investigated the potential relationships between morphological and behavioural specialization. These relationships differed markedly from the traditional ecomorphological paradigm. While morphological specialization showed no relationship with dietary specialization, it exhibited a strong relationship with foraging microhabitat specialization. However, this relationship was inverted: species with specialized morphologies were microhabitat generalists, whereas generalized morphotypes were microhabitat specialists. Interestingly, this mirrors relationships found in plant–pollinator communities and may also be applicable to other ecosystems, highlighting the potential importance of including niche axes beyond dietary specialization into ecomorphological frameworks. On coral reefs, it appears that morphotypes commonly perceived as most generalized may, in fact, be specialized in exploiting flat and easily accessible microhabitats.     

Long-term monitoring of benthic macroinvertebrate community structure: a perspective from a Colorado river*

Temporal and spatial trends were examined in benthic macroinvertebrate and physical-chemical data collected for at least ten years at ten sites along the plains reach of the Cache la Poudre River, Colorado, USA. A distinct longitudinal environment gradient was found as many of the water chemistry parameter levels changed downstream from the reference site. Seasonal Kendall analyses on individual sites indicated that several chemical parameters, including conductivity, un-ionized NH₃-N and NO₂-N have increased since the beginning of the study within most sites. Levels of some parameters (e.g., dissolved oxygen, un-ionized NH₃-N) violated aquatic life standards a few times during the study.Over 175 taxa of macroinvertebrates (primarily insects) were collected in the study reach from 1981–1996. Results from detrended correspondence analyses (DCA) on macroinvertebrate data indicated that this stretch of the river exhibited little longitudinal change beyond the two farthest upstream sites. There was a decline in macroinvertebrate density and total number of taxa within most individual sites during 1983–1984, corresponding with the highest recorded discharge in 75 years (1983) and a prolonged, heavy spring runoff in 1984. Taxa richness and density recovered to pre-1983 levels within a few months to a year following the high flows at most sites. These findings suggested that the macroinvertebrate assemblages had low resistance to disturbance, but high resilience. However, the results from DCAs and Kendall's Coefficient of Concordance (W) on individual sites for the entire study period suggested a similar macroinvertebrate community structure through time. It would appear that the composition and abundance of the lotic macroinvertebrate assemblages in the Poudre River has remained relatively constant over the long-term. This has occurred even with some potentially negative changes in water chemistry and increased urban development.     

Uptake of picophytoplankton,bacterioplankton and virioplankton by a fringing coral reef community (Ningaloo Reef,Australia)

We examined the importance of picoplankton and virioplankton to reef trophodynamics at Ningaloo Reef, (north-western Australia), in May and November 2008. Picophytoplankton (Prochlorococcus, Synechococcus and picoeukaryotes), bacterioplankton (inclusive of bacteria and Archaea), virioplankton and chlorophyll a (Chl a) were measured at five stations following the consistent wave-driven unidirectional mean flow path of seawater across the reef and into the lagoon. Prochlorococcus, Synechococcus, picoeukaryotes and bacterioplankton were depleted to similar levels (~40% on average) over the fore reef, reef crest and reef flat (=‘active reef’), with negligible uptake occurring over the sandy bottom lagoon. Depletion of virioplankton also occurred but to more variable levels. Highest uptake rates, m, of picoplankton occurred over the reef crest, while uptake coefficients, S (independent of cell concentration), were similarly scaled over the reef zones, indicating no preferential uptake of any one group. Collectively, picophytoplankton, bacterioplankton and virioplankton accounted for the uptake of 29 mmol C m⁻² day⁻¹, with Synechococcus contributing the highest proportion of the removed C. Picoplankton and virioplankton accounted for 1–5 mmol N m⁻² day⁻¹ of the removed N, with bacterioplankton estimated to be a highly rich source of N. Results indicate the importance of ocean–reef interactions and the dependence of certain reef organisms on picoplanktonic supply for reef-level biogeochemistry processes.     

Great Barrier Reef butterflyfish community structure: the role of shelf position and benthic community type

The extent to which fish communities are structured by spatial variability in coral reef habitats versus stochastic processes (such as larval supply) is very important in predicting responses to sustained and ongoing habitat degradation. In this study, butterflyfish and benthic communities were surveyed annually over 15 years on 47 reefs (spanning 12° of latitude) of the Great Barrier Reef (GBR). Spatial autocorrelation in the structure of butterflyfish communities versus key differences in reef habitats was investigated to assess the extent to which the structure of these fish communities is influenced by habitat conditions. Benthic communities on each of the 47 reefs were broadly categorised as either: 1. Poritidae/Alcyoniidae, 2. mixed taxa, 3. soft coral or 4. Acropora-dominated habitats. These habitat types most reflected increases in water clarity and wave exposure, moving across the GBR shelf from coastal to outer-shelf environments. In turn, each habitat type also supported very distinct butterflyfish communities. Hard coral feeders were always the dominant butterflyfish species in each community type. However, the numerically dominant species changed according to habitat type, representing spatial replacement of species across the shelf. This study reveals clear and consistent differences in the structure of fish communities among reefs associated with marked differences in habitat structure.     

Temporal settlement patterns of larvae of the broadcast spawning reef coral Favites chinensis and the broadcast spawning and brooding reef coral Goniastrea aspera from Okinawa, Japan

The faviid corals, Favites chinensis and Goniastrea aspera are widely distributed in the Indo-Pacific region. Both corals are hermaphroditic broadcast spawners, but G. aspera is also known to brood planula larvae in Okinawa. This study investigated the temporal settlement patterns of planula larvae of the scleractinian corals F. chinensis and G. aspera that developed from spawned gametes, and planula release and settlement of brooded larvae of G. aspera from Okinawa, Japan. Some of the broadcast-spawned larvae of F. chinensis and G. aspera had very short pre-competency periods of 1–2 and 2–3 days after spawning, and relatively long maximum settlement-competency periods of 56–63 and 63–70 days after spawning, respectively. These pre-competency periods are among the shortest reported for larvae of broadcast spawning coral species, and appear to be negatively correlated with seawater temperature. F. chinensis larvae tended to settle rapidly with 34–39% of larvae settling in the first week after spawning, while broadcast-spawned G. aspera larvae had a slower settlement pattern with 11–15% of larvae settling in the first week after spawning. Brooded larvae of G. aspera settled more rapidly, with settlement rates of 27–31% within the first 24 h and 45–65% within the first week after the start of the experiment. The production of planula larvae with rapid settlement capabilities may enable F. chinensis and G. aspera to establish and maintain populations in shallow reef sites at Okinawa. The release of the brooded planulae for up to 2 months may explain why G. aspera is locally more dominant on shallow reefs in Okinawa than F. chinensis. On a broader scale, the longer settlement competency periods of some of the broadcast-spawned larvae of these species increase their potential for longer-distance dispersal and may partly explain the wide biogeographic distribution of these species in the Indo-Pacific region.     

Ribosomal DNA, species structure, and biogeography of the cactophilic yeast Clavispora opuntiae

The ribosomal DNA of the cactophilic yeast species Clavispora opuntiae was studied in order to clarify the global distribution of the yeast. Over 500 strains, including isolates from several new localities worldwide, were characterized by rDNA restriction mapping. An unusual restriction pattern previously encountered only in one strain, from Conception Island in the Bahamas, was found in several Brazilian isolates. Sequences of the D1/D2 and D7/D8 divergent domains of the large subunit (LSU) and of the intergenic spacers (IGS) confirmed that these strains represent a genetically distinct variety of Clavispora opuntiae. This divergence had previously been hypothesized on the basis of reduced genetic recombination in inter-varietal crosses and the presence of a polymorphic ApaI restriction site located in the LSU. The exact position of the ApaI site in the D8 divergent domain and the nature of the variation that it reveals were determined. The complete sequences of 12 intergenic spacers clarified the significance of the species-wide variation uncovered by restriction mapping. Most of the polymorphic sites occur in the IGS1 and IGS2 regions, on either side of the 5S gene, and the variation is largely due to differences in the numbers and the sequences of internal repeats. Two other polymorphic sites are located in the external transcribed spacer (ETS) region. The reliability of various sites as indicators of overall spacer sequence divergence differed from one case to another. Variety-specific probes were devised and used to screen 120 strains for the presence of recombinant rDNA spacers. Three strains gave ambiguous results, but these did not constitute evidence that inter-varietal recombination has taken place in nature. The hypothesis that the global movement of Clavispora opuntiae has been influenced by the worldwide biological control of prickly pear with Cactoblastis cactorum, a moth of Argentinian origin, has received additional support from the demonstration that Argentinian strains have rDNAs similar to those found where the moth has been introduced. A dramatic founder effect was identified in a yeast population collected in cacti (Maui, Hawaii) in a site where the moth had been recently introduced.     

Effects of extreme seasonality on community structure and functional group dynamics of coral reef algae in the southern Red Sea (Eritrea)

Spatial and temporal variation in the biomass of four functional groups of coral reef algae (canopy algae, foliose algae, turf algae and crustose corallines) was investigated in the southern Red Sea. This region is characterised by extremely high summer temperatures (ca. 35°C). Strong seasonal shifts in the relative contribution of each group to the total macroalgal biomass were observed. On the reef flat, canopy and foliose algae dominated in winter, retaining low biomass in summer. On the fore reef, crustose corallines accounted for most of the macroalgal biomass throughout the year. Turf algae contributed least to the total biomass in all reef zones; biomass peaks shifted from midsummer on the inner reef flat to winter in the deeper zones. Biomass correlated negatively with seawater temperature in most groups, but the correlation was positive for turf algae on the shallow reef flat. We hypothesise both direct and indirect effects of the strong seasonality.

Small-sized invertebrates in a gravel stream: community structure and variability of benthic rotifers

1. The Rotifera assemblage inhabiting the streambed surface and the hyporheic zone of a gravel stream was investigated between October 1991 and October 1992. Forty-two species of Monogononta and 27 of Bdelloidea were identified. Within these two classes, dominant species differed between the surface and the hyporheic zone. At the streambed surface, the abundance of monogonont rotifers showed a seasonal pattern with significantly higher densities in pools, whereas bdelloids showed no clear temporal trend and did not differ significantly among sites. In the hyporheic zone, the depth distribution differed among the two rotifer groups, bdelloids occurred in highest densities between 0 and 30 cm sediment depth, while monogononts were most abundant at greater depths.
2. Species composition differed greatly between successive sampling dates (min. 5 to max. 26 days) at both the streambed surface and the hyporheos. At the streambed surface and in the shallow hyporheos a significantly higher percentage of species was replaced in riffles than in pools.
3. Few measured hydrophysical variables were associated with the Rotifera assemblage structure. At the streambed surface, species richness was negatively correlated with water temperature and substratum heterogeneity, and Monogononta rotifer densities declined with water depth and substratum roughness.
4. Permutation tests carried out on temporal serial correlations showed that, at riffle sites at the streambed surface, bdelloid rotifer densities, rotifer species richness and diversity did not differ significantly from a temporal, near-random pattern. The hyporheic rotifer assemblage followed similar near-random patterns.     

Kenyan coral reef lagoon fish: effects of fishing,substrate complexity,and sea urchins

Population density, number of species, diversity, and species-area relationships of fish species in eight common coral reef-associated families were studied in three marine parks receiving total protection from fishing, four sites with unregulated fishing, and one reef which recently received protection from fishing (referred to as a transition reef). Data on coral cover, reef topographic complexity, and sea urchin abundance were collected and correlated with fish abundance and species richness. The most striking result of this survey is a consistent and large reduction in the population density and species richness of 5 families (surgeonfish, triggerfish, butterflyfish, angelfish, and parrotfish). Poor recovery of parrotfish in the transition reef, relative to other fish families, is interpreted as evidence for competitive exclusion of parrotfish by sea urchins. Reef substrate complexity is significantly associated with fish abundance and diversity, but data suggest different responses for protected versus fished reefs, protected reefs having higher species richness and numbers of individuals than unprotected reefs for the same reef complexity. Sea urchin abundance is negatively associated with numbers of fish and fish species but the interrelationship between sea urchins, substrate complexity, coral cover, and management make it difficult to attribute a set percent of variance to each factor-although fishing versus no fishing appears to be the strongest variable in predicting numbers of individuals and species of fish, and their community similarity. Localized species extirpation is evident for many species on fished reefs (for the sampled area of 1.0 ha). Fifty-two of 110 species found on protected reefs were not found on unprotected reefs.