Showing papers in "Coral Reefs in 2009"

Larval retention and connectivity among populations of corals and reef fishes: history, advances and challenges

Abstract: The extent of larval dispersal on coral reefs has important implications for the persistence of coral reef metapopulations, their resilience and recovery from an increasing array of threats, and the success of protective measures. This article highlights a recent dramatic increase in research effort and a growing diversity of approaches to the study of larval retention within (self-recruitment) and dispersal among (connectivity) isolated coral reef populations. Historically, researchers were motivated by alternative hypotheses concerning the processes limiting populations and structuring coral reef assemblages, whereas the recent impetus has come largely from the need to incorporate dispersal information into the design of no-take marine protected area (MPA) networks. Although the majority of studies continue to rely on population genetic approaches to make inferences about dispersal, a wide range of techniques are now being employed, from small-scale larval tagging and paternity analyses, to large-scale biophysical circulation models. Multiple approaches are increasingly being applied to cross-validate and provide more realistic estimates of larval dispersal. The vast majority of empirical studies have focused on corals and fishes, where evidence for both extremely local scale patterns of self-recruitment and ecologically significant connectivity among reefs at scales of tens of kilometers (and in some cases hundreds of kilometers) is accumulating. Levels of larval retention and the spatial extent of connectivity in both corals and fishes appear to be largely independent of larval duration or reef size, but may be strongly influenced by geographic setting. It is argued that high levels of both self-recruitment and larval import can contribute to the resilience of reef populations and MPA networks, but these benefits will erode in degrading reef environments.

Connectivity, biodiversity conservation and the design of marine reserve networks for coral reefs

Abstract: Networks of no-take reserves are important for protecting coral reef biodiversity from climate change and other human impacts. Ensuring that reserve populations are connected to each other and non-reserve populations by larval dispersal allows for recovery from disturbance and is a key aspect of resilience. In general, connectivity between reserves should increase as the distance between them decreases. However, enhancing connectivity may often tradeoff against a network’s ability to representatively sample the system’s natural variability. This “representation” objective is typically measured in terms of species richness or diversity of habitats, but has other important elements (e.g., minimizing the risk that multiple reserves will be impacted by catastrophic events). Such representation objectives tend to be better achieved as reserves become more widely spaced. Thus, optimizing the location, size and spacing of reserves requires both an understanding of larval dispersal and explicit consideration of how well the network represents the broader system; indeed the lack of an integrated theory for optimizing tradeoffs between connectivity and representation objectives has inhibited the incorporation of connectivity into reserve selection algorithms. This article addresses these issues by (1) updating general recommendations for the location, size and spacing of reserves based on emerging data on larval dispersal in corals and reef fishes, and on considerations for maintaining genetic diversity; (2) using a spatial analysis of the Great Barrier Reef Marine Park to examine potential tradeoffs between connectivity and representation of biodiversity and (3) describing a framework for incorporating environmental fluctuations into the conceptualization of the tradeoff between connectivity and representation, and that expresses both in a common, demographically meaningful currency, thus making optimization possible.

Coral disease following massive bleaching in 2005 causes 60% decline in coral cover on reefs in the US Virgin Islands

Abstract: In the northeast Caribbean, doldrum-like conditions combined with elevated water temperatures in the summer/fall 2005 created the most severe coral bleaching event ever documented within this region. Video monitoring of 100 randomly chosen, permanent transects at five study sites in the US Virgin Islands revealed over 90% of the scleractinian coral cover showed signs of thermal stress by paling or becoming completely white. Lower water temperatures in October allowed some re-coloring of corals; however, a subsequent unprecedented regional outbreak of coral disease affected all sites. Five known diseases or syndromes were recorded; however, most lesions showed signs similar to white plague. Nineteen scleractinian species were affected by disease, with >90% of the disease-induced lesions occurring on the genus Montastraea. The disease outbreak peaked several months after the onset of bleaching at all sites but did not occur at the same time. The mean number of disease-induced lesions increased 51-fold and the mean area of disease-associated mortality increased 13-fold when compared with pre-bleaching disease levels. In the 12 months following the onset of bleaching, coral cover declined at all sites (average loss: 51.5%, range: 42.4–61.8%) reducing the five-site average from 21.4% before bleaching to 10.3% with most mortality caused by white plague disease, not bleaching. Continued losses through October 2007 reduced the average coral cover of the five sites to 8.3% (average 2-year loss: 61.1%, range: 53.0–79.3%). Mean cover by M. annularis (complex) decreased 51%, Colpophyllia natans 78% and Agaricia agaricites 87%. Isolated disease outbreaks have been documented before in the Virgin Islands, but never as widespread or devastating as the one that occurred after the 2005 Caribbean coral-bleaching event. This study provides insight into the effects of continued seawater warming and subsequent coral bleaching events in the Caribbean and highlights the need to understand links between coral bleaching and disease.

Connectivity and resilience of coral reef metapopulations in marine protected areas: matching empirical efforts to predictive needs.

Abstract: Design and decision-making for marine protected areas (MPAs) on coral reefs require prediction of MPA effects with population models. Modeling of MPAs has shown how the persistence of metapopulations in systems of MPAs depends on the size and spacing of MPAs, and levels of fishing outside the MPAs. However, the pattern of demographic connectivity produced by larval dispersal is a key uncertainty in those modeling studies. The information required to assess population persistence is a dispersal matrix containing the fraction of larvae traveling to each location from each location, not just the current number of larvae exchanged among locations. Recent metapopulation modeling research with hypothetical dispersal matrices has shown how the spatial scale of dispersal, degree of advection versus diffusion, total larval output, and temporal and spatial variability in dispersal influence population persistence. Recent empirical studies using population genetics, parentage analysis, and geochemical and artificial marks in calcified structures have improved the understanding of dispersal. However, many such studies report current self-recruitment (locally produced settlement/settlement from elsewhere), which is not as directly useful as local retention (locally produced settlement/total locally released), which is a component of the dispersal matrix. Modeling of biophysical circulation with larval particle tracking can provide the required elements of dispersal matrices and assess their sensitivity to flows and larval behavior, but it requires more assumptions than direct empirical methods. To make rapid progress in understanding the scales and patterns of connectivity, greater communication between empiricists and population modelers will be needed. Empiricists need to focus more on identifying the characteristics of the dispersal matrix, while population modelers need to track and assimilate evolving empirical results.

Climate change and coral reef connectivity

Abstract: This review assesses and predicts the impacts that rapid climate change will have on population connectivity in coral reef ecosystems, using fishes as a model group. Increased ocean temperatures are expected to accelerate larval development, potentially leading to reduced pelagic durations and earlier reef-seeking behaviour. Depending on the spatial arrangement of reefs, the expectation would be a reduction in dispersal distances and the spatial scale of connectivity. Small increase in temperature might enhance the number of larvae surviving the pelagic phase, but larger increases are likely to reduce reproductive output and increase larval mortality. Changes to ocean currents could alter the dynamics of larval supply and changes to planktonic productivity could affect how many larvae survive the pelagic stage and their condition at settlement; however, these patterns are likely to vary greatly from place-to-place and projections of how oceanographic features will change in the future lack sufficient certainty and resolution to make robust predictions. Connectivity could also be compromised by the increased fragmentation of reef habitat due to the effects of coral bleaching and ocean acidification. Changes to the spatial and temporal scales of connectivity have implications for the management of coral reef ecosystems, especially the design and placement of marine-protected areas. The size and spacing of protected areas may need to be strategically adjusted if reserve networks are to retain their efficacy in the future.

Juvenile corals can acquire more carbon from high-performance algal symbionts

Abstract: Algal endosymbionts of the genus Symbiodinium play a key role in the nutrition of reef building corals and strongly affect the thermal tolerance and growth rate of the animal host. This study reports that 14C photosynthate incorporation into juvenile coral tissues was doubled in Acropora millepora harbouring Symbiodinium C1 compared with juveniles from common parentage harbouring Symbiodinium D in a laboratory experiment. Rapid light curves performed on the same corals revealed that the relative electron transport rate of photosystem II (rETRMAX) was 87% greater in Symbiodinium C1 than in Symbiodinium D in hospite. The greater relative electron transport through photosystem II of Symbiodinium C1 is positively correlated with increased carbon delivery to the host under the applied experimental conditions (r2 = 0.91). This may translate into a competitive advantage for juveniles harbouring Symbiodinium C1 under certain field conditions, since rapid early growth typically limits mortality. Both symbiont types exhibited severe reductions in 14C incorporation during a 10-h exposure to the electron transport blocking herbicide diuron (DCMU), confirming the link between electron transport through PSII and photosynthate incorporation within the host tissue. These findings advance the current understanding of symbiotic relationships between corals and their symbionts, providing evidence that enhanced growth rates of juvenile corals may result from greater translocation of photosynthates from Symbiodinium C1.

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

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

Phase shifts and the stability of macroalgal communities on Caribbean coral reefs

Abstract: Caribbean coral reefs are widely thought to exhibit two alternate stable states with one being dominated by coral and the other by macroalgae. However, the observation of linear empirical relationships among grazing, algal cover and coral recruitment has led the existence of alternate stable states to be questioned; are reefs simply exhibiting a continuous phase shift in response to grazing or are the alternate states robust to certain changes in grazing? Here, a model of a Caribbean forereef is used to reconcile the existence of two stable community states with common empirical observations. Coral-depauperate and coral-dominated reef states are predicted to be stable on equilibrial time scales of decades to centuries and their emergence depends on the presence or absence of a bottleneck in coral recruitment, which is determined by threshold levels of grazing intensity and other process variables. Under certain physical and biological conditions, corals can be persistently depleted even while increases in grazing reduce macroalgal cover and enhance coral recruitment; only once levels of recruitment becomes sufficient to overwhelm the population bottleneck will the coral-dominated state begin to emerge. Therefore, modest increases in grazing will not necessarily allow coral populations to recover, whereas large increases, such as those associated with recovery of the urchin Diadema antillarum, are likely to exceed threshold levels of grazing intensity and set a trajectory of coral recovery. The postulated existence of alternate stable states is consistent with field observations of linear relationships between grazing, algal cover and coral recruitment when coral cover is low and algal exclusion when coral cover is high. The term ‘macroalgal dominated’ is potentially misleading because the coral-depauperate state can be associated with various levels of macroalgal cover. The term ‘coral depauperate’ is preferable to ‘macroalgal dominated’ when describing alternate states of Caribbean reefs.

Management under uncertainty: guide-lines for incorporating connectivity into the protection of coral reefs

Abstract: The global decline in coral reefs demands urgent management strategies to protect resilience. Protecting ecological connectivity, within and among reefs, and between reefs and other ecosystems is critical to resilience. However, connectivity science is not yet able to clearly identify the specific measures for effective protection of connectivity. This article aims to provide a set of principles or practical guidelines that can be applied currently to protect connectivity. These ‘rules of thumb’ are based on current knowledge and expert opinion, and on the philosophy that, given the urgency, it is better to act with incomplete knowledge than to wait for detailed understanding that may come too late. The principles, many of which are not unique to connectivity, include: (1) allow margins of error in extent and nature of protection, as insurance against unforeseen or incompletely understood threats or critical processes; (2) spread risks among areas; (3) aim for networks of protected areas which are: (a) comprehensive and spread—protect all biotypes, habitats and processes, etc., to capture as many possible connections, known and unknown; (b) adequate—maximise extent of protection for each habitat type, and for the entire region; (c) representative—maximise likelihood of protecting the full range of processes and spatial requirements; (d) replicated—multiple examples of biotypes or processes enhances risk spreading; (4) protect entire biological units where possible (e.g. whole reefs), including buffers around core areas. Otherwise, choose bigger rather than smaller areas; (5) provide for connectivity at a wide range of dispersal distances (within and between patches), emphasising distances <20–30 km; and (6) use a portfolio of approaches, including but not limited to MPAs. Three case studies illustrating the application of these principles to coral reef management in the Bohol Sea (Philippines), the Great Barrier Reef (Australia) and Kimbe Bay (Papua New Guinea) are described.

Record densities of Indo-Pacific lionfish on Bahamian coral reefs

Abstract: First documented off Florida more than a decade ago, populations of Indo-Pacific lionfish (Pterois volitans and P. miles) have rapidly spread throughout the Bahamas and northern Caribbean (Whitfield et al. 2007; Hamner et al. 2007; USGS Nonindigenous Aquatic Species Database 2007). Here, we report lionfish densities from reefs off the southwest coast of New Providence, Bahamas (25 04.6†N, 77 20.6†W) which far exceed the highest densities documented for this species in both its invaded and native ranges. Densities of P. volitans were measured on 10 · 50 m transects from May to July 2008, at depths of 12 to 20 m. At three sites, each separated by more than 1 km, we found >390 lionfish per hectare (mean ± 1 SD; 393.3 ± 144.4 lionfish ha, n 1⁄4 4 transects per site). These densities are more than 18 times higher than those reported by Whitfield et al. (2007) from invaded habitats off the coast of North Carolina, USA (21.2 ± 5.1 ha). Fishelson (1997) provides the only rough estimate of lionfish density of which we are aware for the native range: ~80 adult lionfish in a 1-km stretch of Red Sea reef. Assuming a mean search width of 10 m, this would yield a density of ~80 lionfish ha, which is five times lower than that in New Providence. Caribbean sightings have now been confirmed as far west as Cuba and the Cayman Islands and southeast to St. Croix (Lad Akins, REEF, pers. comm.). The impacts of this invasion are generating great concern, as individual lionfish have been shown to reduce recruitment of Bahamian native fish by 79% on small experimental reefs (Albins and Hixon 2008). Given the high densities noted here (Fig. 1), the impacts of lionfish on natural reefs are expected to be extreme.

Photobiology of Symbiodinium revisited: bio-physical and bio-optical signatures

Abstract: Light is often the most abundant resource within the nutrient-poor waters surrounding coral reefs. Consequently, zooxanthellae (Symbiodinium spp.) must continually photoacclimate to optimise productivity and ensure coral success. In situ coral photobiology is becoming dominated by routine assessments using state-of-the-art non-invasive bio-optical or chlorophyll a fluorescence (bio-physical) techniques. Multiple genetic types of Symbiodinium are now known to exist; however, little focus has been given as to how these types differ in terms of characteristics that are observable using these techniques. Therefore, this investigation aimed to revisit and expand upon a pivotal study by Iglesias-Prieto and Trench (1994) by comparing the photoacclimation characteristics of different Symbiodinium types based on their bio-physical (chlorophyll a fluorescence, reaction centre counts) and bio-optical (optical absorption, pigment concentrations) ‘signatures’. Signatures described here are unique to Symbiodinium type and describe phenotypic responses to set conditions, and hence are not suitable to describe taxonomic structure of in hospiteSymbiodinium communities. In this study, eight Symbiodinium types from clades and sub-clades (A–B, F) were grown under two PFDs (Photon Flux Density) and examined. The photoacclimation response by Symbiodinium was highly variable between algal types for all bio-physical and for many bio-optical measurements; however, a general preference to modifying reaction centre content over effective antennae-absorption was observed. Certain bio-optically derived patterns, such as light absorption, were independent of algal type and, when considered per photosystem, were matched by reaction centre stoichiometry. Only by better understanding genotypic and phenotypic variability between Symbiodinium types can future studies account for the relative taxonomic and physiological contribution by Symbiodinium to coral acclimation.

Bioindicators of changes in water quality on coral reefs: review and recommendations for monitoring programmes

Abstract: Effective environmental management requires monitoring programmes that provide specific links between changes in environmental conditions and ecosystem health. This article reviews the suitability of a range of bioindicators for use in monitoring programmes that link changes in water quality to changes in the condition of coral-reef ecosystems. From the literature, 21 candidate bioindicators were identified, whose responses to changes in water quality varied spatially and temporally; responses ranged from rapid (hours) changes within individual corals to long-term (years) changes in community composition. From this list, the most suitable bioindicators were identified by determining whether responses were (i) specific, (ii) monotonic, (iii) variable, (iv) practical and (v) ecologically relevant to management goals. For long-term monitoring programmes that aim to quantify the effects of chronic changes in water quality, 11 bioindicators were selected: symbiont photophysiology, colony brightness, tissue thickness and surface rugosity of massive corals, skeletal elemental and isotopic composition, abundance of macro-bioeroders, micro- and meiobenthic organisms such as foraminifera, coral recruitment, macroalgal cover, taxonomic richness of corals and the maximal depth of coral-reef development. For short-term monitoring programmes, or environmental impact assessments that aim to quantify the effects of acute changes in water quality, a subset of seven of these bioindicators were selected, including partial mortality. Their choice will depend on the specific objectives and the timeframe available for each monitoring programme. An assessment framework is presented to assist in the selection of bioindicators to quantify the effects of changing water quality on coral-reef ecosystems.

Coral surface area quantification-evaluation of established techniques by comparison with computer tomography

Abstract: The surface area of scleractinian corals represents an important reference parameter required for various aspects of coral reef science. However, with advancements in detection accuracy and novel approaches for coral surface area quantification, evaluation of established techniques in comparison with state-of-the-art technology gains importance to coral researchers. This study presents an evaluation of methodological accuracy for established techniques in comparison to a novel approach composed of computer tomography (CT) and 3-dimensional surface reconstruction. The skeleton surface area of reef corals from six genera representing the most common morphological growth forms was acquired by CT and subsequently measured by computer-aided 3-dimensional surface reconstruction. Surface area estimates for the same corals were also obtained by application of four established techniques: Simple and Advanced Geometry, Wax Coating and Planar Projection Photography. Comparison of the resulting area values revealed significant differences between the majority (82%) of established techniques and the CT reference. Genus-specific analysis assigned the highest accuracy to geometric approximations (Simple or Advanced Geometry) for the majority of assessed coral genera (maximum accuracy: 104%; Simple Geometry with Montipora sp.). The commonly used and invasive Wax Coating technique reached intermediate accuracy (47–74%) for the majority of genera, but performed outstanding in the measurement of branching Acropora spp. corals (maximum accuracy: 101%), while the Planar Projection Photography delivered genera-wide low accuracy (12–36%). Comparison of area values derived from established techniques and CT additionally yielded approximation factors (AFs) applicable as factors in the mathematical improvement of surface area estimates by established techniques in relation to CT reference accuracy.

Decline in skeletal growth of the coral Porites lutea from the Andaman Sea, South Thailand between 1984 and 2005

Abstract: Of the few studies that have examined in situ coral growth responses to recent climate change, none have done so in equatorial waters subject to relatively high sea temperatures (annual mean >27°C). This study compared the growth rate of Porites lutea from eight sites at Phuket, South Thailand between two time periods (December 1984–November 1986 and December 2003–November 2005). There was a significant decrease in coral calcification (23.5%) and linear extension rates (19.4–23.4%) between the two sampling periods at a number of sites, while skeletal bulk density remained unchanged. Over the last 46 years, sea temperatures (SST) in the area have risen at a rate of 0.161°C per decade (current seasonal temperature range 28–30°C) and regression analysis of coral growth data is consistent with a link between rising temperature and reduced linear extension in the order of 46–56% for every 1°C rise in SST. The apparent sensitivity of linear extension in P. lutea to increased SST suggests that corals in this part of the Andaman Sea may already be subjected to temperatures beyond their thermal optimum for skeletal growth.

Are artificial reefs surrogates of natural habitats for corals and fish in Dubai, United Arab Emirates?

Abstract: Artificial reefs are often promoted as mitigating human impacts in coastal ecosystems and enhancing fisheries; however, evidence supporting their benefits is equivocal. Such structures must be compared with natural reefs in order to assess their performance, but past comparisons typically examined artificial structures that were too small, or were immature, relative to the natural reefs. We compared coral and fish communities on two large (>400,000 m3) and mature (>25 year) artificial reefs with six natural coral patches. Coral cover was higher on artificial reefs (50%) than in natural habitats (31%), but natural coral patches contained higher species richness (29 vs. 20) and coral diversity (H′ = 2.3 vs. 1.8). Multivariate analyses indicated strong differences between coral communities in natural and artificial habitats. Fish communities were sampled seasonally for 1 year. Multivariate fish communities differed significantly among habitat types in the summer and fall, but converged in the winter and spring. Univariate analysis indicated that species richness and abundance were stable throughout the year on natural coral patches but increased significantly in the summer on artificial reefs compared with the winter and spring, explaining the multivariate changes in community structure. The increased summer abundance on artificial reefs was mainly due to adult immigration. Piscivores were much more abundant in the fall than in the winter or spring on artificial reefs, but had low and stable abundance throughout the year in natural habitats. It is likely that the decreased winter and spring abundance of fish on the artificial reefs resulted from both predation and emigration. These results indicate that large artificial reefs can support diverse and abundant coral and fish communities. However, these communities differ structurally and functionally from those in natural habitats, and they should not be considered as replacements for natural coral and fish communities.

Herbivory versus corallivory: are parrotfish good or bad for Caribbean coral reefs?

Abstract: With coral cover in decline on many Caribbean reefs, any process of coral mortality is of potential concern. While sparisomid parrotfishes are major grazers of Caribbean reefs and help control algal blooms, the fact that they also undertake corallivory has prompted some to question the rationale for their conservation. Here the weight of evidence for beneficial effects of parrotfishes, in terms of reducing algal cover and facilitating demographic processes in corals, and the deleterious effects of parrotfishes in terms of causing coral mortality and chronic stress, are reviewed. While elevated parrotfish density will likely increase the predation rate upon juvenile corals, the net effect appears to be positive in enhancing coral recruitment through removal of macroalgal competitors. Parrotfish corallivory can cause modest partial colony mortality in the most intensively grazed species of Montastraea but the generation and healing of bite scars appear to be in near equilibrium, even when coral cover is low. Whole colony mortality in adult corals can lead to complete exclusion of some delicate, lagoonal species of Porites from forereef environments but is only reported for one reef species (Porites astreoides), for one habitat (backreef), and with uncertain incidence (though likely <<10%). No deleterious effects of predation on coral growth or fecundity have been reported, though recovery of zooxanthellae after bleaching events may be retarded. The balance of evidence to date finds strong support for the herbivory role of parrotfishes in facilitating coral recruitment, growth, and fecundity. In contrast, no net deleterious effects of corallivory have been reported for reef corals. Corallivory is unlikely to constrain overall coral cover but contraints upon dwindling populations of the Montastraea annularis species complex are feasible and the role of parrotfishes as a vector of coral disease requires evaluation. However, any assertion that conservation practices should guard against protecting corallivorous parrotfishes appears to be unwarranted at this stage.

A map of human impacts to a “pristine” coral reef ecosystem, the Papahānaumokuākea Marine National Monument

Abstract: Effective and comprehensive regional-scale marine conservation requires fine-grained data on the spatial patterns of threats and their overlap. To address this need for the Papahānaumokuākea Marine National Monument (Monument) in Hawaii, USA, spatial data on 14 recent anthropogenic threats specific to this region were gathered or created, including alien species, bottom fishing, lobster trap fishing, ship-based pollution, ship strike risks, marine debris, research diving, research equipment installation, research wildlife sacrifice, and several anthropogenic climate change threats i.e., increase in ultraviolet (UV) radiation, seawater acidification, the number of warm ocean temperature anomalies relevant to disease outbreaks and coral bleaching, and sea level rise. These data were combined with habitat maps and expert judgment on the vulnerability of different habitat types in the Monument to estimate spatial patterns of current cumulative impact at 1 ha (0.01 km2) resolution. Cumulative impact was greatest for shallow reef areas and peaked at Maro Reef, where 13 of the 14 threats overlapped in places. Ocean temperature variation associated with disease outbreaks was found to have the highest predicted impact overall, followed closely by other climate-related threats, none of which have easily tractable management solutions at the regional scale. High impact threats most tractable to regional management relate to ship traffic. Sensitivity analyses show that the results are robust to both data availability and quality. Managers can use these maps to (1) inform management and surveillance priorities based on the ranking of threats and their distributions, (2) guide permitting decisions based on cumulative impacts, and (3) choose areas to monitor for climate change effects. Furthermore, this regional analysis can serve as a case study for managers elsewhere interested in assessing and mapping region-specific cumulative human impacts.

Thinking and managing outside the box: coalescing connectivity networks to build region-wide resilience in coral reef ecosystems

Abstract: As the science of connectivity evolves, so too must the management of coral reefs. It is now clear that the spatial scale of disturbances to coral reef ecosystems is larger and the scale of larval connectivity is smaller than previously thought. This poses a challenge to the current focus of coral reef management, which often centers on the establishment of no-take reserves (NTRs) that in practice are often too small, scattered, or have low stakeholder compliance. Fished species are generally larger and more abundant in protected reserves, where their reproductive potential is often greater, yet documented demographic benefits of these reproductive gains outside reserves are modest at best. Small reproductive populations and limited dispersal of larvae play a role, as does the diminished receptivity to settling larvae of degraded habitats that can limit recruitment by more than 50%. For “demographic connectivity” to contribute to the resilience of coral reefs, it must function beyond the box of no-take reserves. Specifically, it must improve nursery habitats on or near reefs and enhance the reproductive output of ecologically important species throughout coral reef ecosystems. Special protection of ecologically important species (e.g., some herbivores in the Caribbean) and size-regulated fisheries that capitalize on the benefits of NTRs and maintain critical ecological functions are examples of measures that coalesce marine reserve effects and improve the resilience of coral reef ecosystems. Important too is the necessity of local involvement in the management process so that social costs and benefits are properly assessed, compliance increased and success stories accrued.

Stability of coral–endosymbiont associations during and after a thermal stress event in the southern Great Barrier Reef

Abstract: Shifts in the community of symbiotic dinoflagellates to those that are better suited to the prevailing environmental condition may provide reef-building corals with a rapid mechanism by which to adapt to changes in the environment. In this study, the dominant Symbiodinium in 10 coral species in the southern Great Barrier Reef was monitored over a 1-year period in 2002 that coincided with a thermal stress event. Molecular genetic profiling of Symbiodinium communities using single strand conformational polymorphism of the large subunit rDNA and denaturing gradient gel electrophoresis of the internal transcribed spacer 2 region did not detect any changes in the communities during and after this thermal-stress event. Coral colonies of seven species bleached but recovered with their original symbionts. This study suggests that the shuffling or switching of symbionts in response to thermal stress may be restricted to certain coral species and is probably not a universal feature of the coral-symbiont relationship.

Elevated temperature reduces survivorship and settlement of the larvae of the Caribbean scleractinian coral, Favia fragum (Esper)

Abstract: The effect of elevated seawater temperatures, such as those plaguing tropical seas during the summers of anomalously warm years, on early life stages of reef corals remains poorly studied. To redress this situation, survivorship of larvae of the brooding coral, Favia fragum, was studied in the laboratory, using both short term (48 h) and long term (156–191 h) exposures to 28, 29, and 31°C. Ability to settle when presented with induction substrates and survival after settlement, at the same exposure temperature and after reciprocal transfers to the other experimental temperatures, were also measured. No significant effect of temperature on survivorship was detected after 48 h of exposure, but larvae incubated for 156 h at the highest temperature (31°C) exhibited a 13% reduced survivorship compared to larvae at 28°C. Induction of settlement further increased mortality at the highest temperature (31°C); survivorship after settlement at 31°C was 27% lower than when larvae were simply maintained at the elevated temperature. These results indicate that elevated temperatures are more detrimental to coral larvae undergoing the developmentally complex settlement process than to the swimming planula stage. This may bode poorly for Caribbean corals with late summer reproductive seasons.

Reef-associated crustacean fauna: biodiversity estimates using semi-quantitative sampling and DNA barcoding

Abstract: The cryptofauna associated with coral reefs accounts for a major part of the biodiversity in these ecosystems but has been largely overlooked in biodiversity estimates because the organisms are hard to collect and identify. We combine a semi-quantitative sampling design and a DNA barcoding approach to provide metrics for the diversity of reef-associated crustacean. Twenty-two similar-sized dead heads of Pocillopora were sampled at 10 m depth from five central Pacific Ocean localities (four atolls in the Northern Line Islands and in Moorea, French Polynesia). All crustaceans were removed, and partial cytochrome oxidase subunit I was sequenced from 403 individuals, yielding 135 distinct taxa using a species-level criterion of 5% similarity. Most crustacean species were rare; 44% of the OTUs were represented by a single individual, and an additional 33% were represented by several specimens found only in one of the five localities. The Northern Line Islands and Moorea shared only 11 OTUs. Total numbers estimated by species richness statistics (Chao1 and ACE) suggest at least 90 species of crustaceans in Moorea and 150 in the Northern Line Islands for this habitat type. However, rarefaction curves for each region failed to approach an asymptote, and Chao1 and ACE estimators did not stabilize after sampling eight heads in Moorea, so even these diversity figures are underestimates. Nevertheless, even this modest sampling effort from a very limited habitat resulted in surprisingly high species numbers.

Maintenance of fish diversity on disturbed coral reefs

Abstract: Habitat perturbations play a major role in shaping community structure; however, the elements of disturbance-related habitat change that affect diversity are not always apparent. This study examined the effects of habitat disturbances on species richness of coral reef fish assemblages using annual surveys of habitat and 210 fish species from 10 reefs on the Great Barrier Reef (GBR). Over a period of 11 years, major disturbances, including localised outbreaks of crown-of-thorns sea star (Acanthaster planci), severe storms or coral bleaching, resulted in coral decline of 46–96% in all the 10 reefs. Despite declines in coral cover, structural complexity of the reef framework was retained on five and species richness of coral reef fishes maintained on nine of the disturbed reefs. Extensive loss of coral resulted in localised declines of highly specialised coral-dependent species, but this loss of diversity was more than compensated for by increases in the number of species that feed on the epilithic algal matrix (EAM). A unimodal relationship between areal coral cover and species richness indicated species richness was greatest at approximately 20% coral cover declining by 3–4 species (6–8% of average richness) at higher and lower coral cover. Results revealed that declines in coral cover on reefs may have limited short-term impact on the diversity of coral reef fishes, though there may be fundamental changes in the community structure of fishes.

Highly restricted gene flow and deep evolutionary lineages in the giant clam Tridacna maxima

Abstract: The tropical Indo-West Pacific is the biogeographic region with the highest diversity of marine shallow water species, with its centre in the Indo-Malay Archipelago. However, due to its high endemism, the Red Sea is also considered as an important centre of evolution. Currently, not much is known about exchange among the Red Sea, Indian Ocean and West Pacific, as well as connectivity within the Indo-Malay Archipelago, even though such information is important to illuminate ecological and evolutionary processes that shape marine biodiversity in these regions. In addition, the inference of connectivity among populations is important for conservation. This study aims to test the hypothesis that the Indo-Malay Archipelago and the Red Sea are important centres of evolution by studying the genetic population structure of the giant clam Tridacna maxima. This study is based on a 484-bp fragment of the cytochrome c oxidase I gene from 211 individuals collected at 14 localities in the Indo-West Pacific to infer lineage diversification and gene flow as a measure for connectivity. The analysis showed a significant genetic differentiation among sample sites in the Indo-West Pacific (Φst = 0.74, P < 0.001) and across the Indo-Malay Archipelago (Φst = 0.72, P < 0.001), indicating restricted gene flow. Hierarchical AMOVA revealed the highest fixation index (Φct = 0.8, P < 0.001) when sample sites were assigned to the following regions: (1) Red Sea, (2) Indian Ocean and Java Sea, (3) Indonesian throughflow and seas in the East of Sulawesi, and (4) Western Pacific. Geological history as well as oceanography are important factors that shape the genetic structure of T. maxima in the Indo-Malay Archipelago and Red Sea. The observed deep evolutionary lineages might include cryptic species and this result supports the notion that the Indo-Malay Archipelago and the Red Sea are important centres of evolution.

A reappraisal of the diversity of geomorphological and genetic processes of New Caledonian coral reefs: a synthesis from optical remote sensing, coring and acoustic multibeam observations

Abstract: The diversity of oceanic and continental reef structures of New Caledonia is reviewed, taking into account their geological history and in particular, that since the last interglacial period. To guide this review, a new path is provided by following the diversity of units that have been mapped and characterized using high spatial resolution optical remote sensing data for the main New Caledonian coral reef complexes (banks, atolls, uplifted reefs, drowned reefs, fringing reefs, barrier reefs, patch reefs) and their individual reef-forming units. This interpretation, based on geomorphology, depth, and exposure has provided 161 unit types distributed across 4,537 km2 of reef area and 31,336 km2 of non-reef area. In addition to shallow reefs (0–30 m) described by optical remote sensing, the bathymetry of deep slopes between −20 to −1,000 m were recently mapped using multibeam acoustic data providing additional data to explain the morphological diversity. With the detailed three-dimensional topographic information acquired, hitherto unrecognized marine terraces and faulting became visible, indicating different episodes of formation of the barrier reef and of sea level variations. Finally, dating and coring corals provided a more accurate understanding of the genesis of the present reef structures. In contrast with the synoptic remote sensing data, cores provided only point data, but allowed the addition of a precise temporal dimension to the description of New Caledonian reefs. Cores provided a significant body of the information necessary for the establishment of models of reef settlement and development during the last interglacial ages in the New Caledonian region. The combined examination of the different sources of data, and the exhaustive description of remotely sensed reef units, allow a qualitative synoptic parallel to be drawn between the morphology of modern reefs and the contrasting patterns of reef growth, subsidence, and uplift rates occurring around New Caledonia.

High genetic differentiation and cross-shelf patterns of genetic diversity among Great Barrier Reef populations of Symbiodinium

Abstract: The resilience of Symbiodinium harboured by corals is dependent on the genetic diversity and extent of connectivity among reef populations. This study presents genetic analyses of Great Barrier Reef (GBR) populations of clade C Symbiodinium hosted by the alcyonacean coral, Sinularia flexibilis. Allelic variation at four newly developed microsatellite loci demonstrated that Symbiodinium populations are genetically differentiated at all spatial scales from 16 to 1,360 km (pairwise ΦST = 0.01–0.47, mean = 0.22); the only exception being two neighbouring populations in the Cairns region separated by 17 km. This indicates that gene flow is restricted for Symbiodinium C hosted by S. flexibilis on the GBR. Patterns of population structure reflect longshore circulation patterns and limited cross-shelf mixing, suggesting that passive transport by currents is the primary mechanism of dispersal in Symbiodinium types that are acquired horizontally. There was no correlation between the genetic structure of Symbiodinium populations and their host S. flexibilis, most likely because different factors affect the dispersal and recruitment of each partner in the symbiosis. The genetic diversity of these Symbiodinium reef populations is on average 1.5 times lower on inshore reefs than on offshore reefs. Lower inshore diversity may reflect the impact of recent bleaching events on Sinularia assemblages, which have been more widespread and severe on inshore reefs, but may also have been shaped by historical sea level fluctuations or recent migration patterns.

Connectivity, regime shifts and the resilience of coral reefs

Abstract: Connectivity of larvae among metapopulations in open marine systems can be a double-edged sword, allowing for the colonization and replenishment of both desirable and undesirable elements of interacting species-rich assemblages. This article studies the effect of recruitment by coral and macroalgae on the resilience of grazed reef ecosystems. In particular, we focus on how larval connectivity affects regime shifts between alternative assemblages that are dominated either by corals or by macroalgae. Using a model with bistability dynamics, we show that recruitment of coral larvae erodes the resilience of a macroalgae-dominated ecosystem when grazing is high, but has negligible effect when grazing is low. Conversely, recruitment by macroalgae erodes the resilience of a coral-dominated ecosystem when grazing is low, leading to a regime shift to macroalgae. Thus, spillover of coral recruits from highly protected areas will not restore coral cover or prevent flips to macroalgae in the surrounding seascape if grazing levels in these areas are depleted, but may be pivotal for re-building coral populations if grazing is high. Fishing restrictions and the re-introduction of herbivores should therefore be a prime conservation objective for preventing undesirable regime shifts. Connectivity by some components of coral reef assemblages (e.g., macroalgae, pathogens, crown-of-thorns starfish) may be detrimental to sustaining reefs, especially where overfishing and other drivers have eroded their resilience, making them more vulnerable to a regime shift.

Understanding diversity in coral-algal symbiosis: A cluster-based approach to interpreting fine-scale genetic variation in the genus Symbiodinium

Abstract: Reef corals associate with an extraordinary diversity of dinoflagellate endosymbionts (genus Symbiodinium), and this diversity has become critical to understanding how corals respond to environmental changes. A popular molecular marker for Symbiodinium diversity, the Internal Transcribed Spacer-2 (ITS-2) region of ribosomal DNA, has revealed hundreds of distinct variants that are generally interpreted as representing different species, even though many have not been systematically tested for functional or ecological differentiation. Many of these variants are only minimally divergent from one another (1 bp or less), and others occupy basal nodes of traditional species phylogenies (“living ancestors”), indicating that some Symbiodinium ITS-2 diversity may represent intraspecific sequence variation. This hypothesis was tested for Symbiodinium clades A–D (the dominant symbionts of reef corals) through the construction of statistical parsimony networks of ITS-2 sequence diversity, and identification of clusters of closely related sequences within these networks. Initial assessments indicated that ecological differentiation exists between, but not within, these clusters. This approach, although imperfect in its ability to identify species boundaries in all cases, nevertheless dramatically reduces “species” diversity in Symbiodinium (from ~175 to 35). This testable alternative hypothesis indicates that, in Symbiodinium, “species” consist of clusters of closely related ITS-2 sequences diverging from ancestral variants that are typically ecologically dominant. A cluster-based view of Symbiodinium ITS-2 diversity improves our ability to: (1) construct well-supported symbiont phylogenies; (2) establish functional niches for symbiont species; and (3) understand flexibility and specificity within coral-algal symbioses. This cluster-based approach can ultimately be integrated with emerging population-level datasets (microsatellites and microsatellite flanking regions) to improve understanding of species diversity in Symbiodinium.

Sea-surface temperature and thermal stress in the Coral Triangle over the past two decades

Abstract: Increasing ocean temperature has become one of the major concerns in recent times with reports of various related ecological impacts becoming commonplace. One of the more notable is the increased frequency of mass coral bleaching worldwide. This study focuses on the Coral Triangle region and utilizes the National Oceanic and Atmospheric Administration-Coral Reef Watch (NOAA-CRW) satellite-derived sea surface temperature (SST) and Degree Heating Weeks (DHW) products to investigate changes in the thermal regime of the Coral Triangle waters between 1985 and 2006. Results show an upward trend in SST during this period with an average rate of 0.2°C/decade. However, warming within this region is not uniform, and the waters of the northern and eastern parts of the Coral Triangle are warming fastest. Areas in the eastern part have experienced more thermal stress events, and these stress events appear to be more likely during a La Nina.

Effects of herbivores, nutrient enrichment, and their interactions on macroalgal proliferation and coral growth

Abstract: We conducted a 20-week manipulative field experiment on shallow forereefs of the Florida Keys to assess the separate and interactive effects of herbivory and nutrient enrichment on the development of macroalgal communities and the fitness of the corals Porites porites and Siderastrea siderea. Excluding large herbivorous fishes produced macrophyte blooms both with and without nutrient enrichment. In contrast, there were no direct effects of nutrient enrichment. There were, however, small, but significant, interactive effects of herbivory and enrichment on macroalgal cover. Following nutrient enrichment, total macroalgae and the common seaweeds Dictyota spp. were suppressed in the presence, but not in the absence, of large herbivorous fishes—suggesting that fishes were selectively feeding on nutrient-enriched macrophytes. Access by large herbivores prevented algal overgrowth of corals, but these large fishes also directly grazed both corals. Excluding fishes did not alter survivorship of either coral species, but did decrease parrotfish grazing scars on both corals and increased the net growth of P. porites. Nutrient additions had no direct effects on the survivorship of corals, but there was a trend (P = 0.097) for nutrients to stimulate the growth of P. porites. The preponderance of experiments available to date indicates that loss of key herbivores is a major factor driving macroalgal blooms on coral reefs; anthropogenic nutrient pollution generally plays a more minor role.

Genetic structure of the Caribbean giant barrel sponge Xestospongia muta using the I3-M11 partition of COI

Abstract: In recent years, reports of sponge bleaching, disease, and subsequent mortality have increased alarmingly. Population recovery may depend strongly on colonization capabilities of the affected species. The giant barrel sponge Xestospongia muta is a dominant reef constituent in the Caribbean. However, little is known about its population structure and gene flow. The 5′-end fragment of the mitochondrial gene cytochrome oxidase subunit I is often used to address these kinds of questions, but it presents very low intraspecific nucleotide variability in sponges. In this study, the usefulness of the I3-M11 partition of COI to determine the genetic structure of X. muta was tested for seven populations from Florida, the Bahamas and Belize. A total of 116 sequences of 544 bp were obtained for the I3-M11 partition corresponding to four haplotypes. In order to make a comparison with the 5′-end partition, 10 sequences per haplotype were analyzed for this fragment. The 40 resulting sequences were of 569 bp and corresponded to two haplotypes. The nucleotide diversity of the I3-M11 partition (π = 0.00386) was higher than that of the 5′-end partition (π = 0.00058), indicating better resolution at the intraspecific level. Sponges with the most divergent external morphologies (smooth vs. digitate surface) had different haplotypes, while those with the most common external morphology (rough surface) presented a mixture of haplotypes. Pairwise tests for genetic differentiation among geographic locations based on F ST values showed significant genetic divergence between most populations, but this genetic differentiation was not due to isolation by distance. While limited larval dispersal may have led to differentiation among some of the populations, the patterns of genetic structure appear to be most strongly related to patterns of ocean currents. Therefore, hydrological features may play a major role in sponge colonization and need to be considered in future plans for management and conservation of these important components of coral reef ecosystems.