Unprecedented bleaching-induced mortality in Porites spp. at Rangiroa Atoll, French Polynesia

In April-May 1998, mass coral bleaching was observed in the lagoon of Rangiroa Atoll, Tuamotu Archipelago, French Polynesia. Six months later, the extent of bleaching-induced coral mortality was assessed at three sites. Corals in the fast-growing genus Pocillopora had experienced >99% mortality. Many large colonies of the slow-growing genus Porites (mean horizontal cross-sectional area 5.8 m²) had also died - a phenomenon not previously observed in French Polynesia and virtually unprecedented world-wide. At one site, 25% of colonies, or 44% of the pre-bleaching cover of living Porites, experienced whole-colony mortality. At the two other sites, recently dead Porites accounted for 41% and 82% of the pre-bleaching live cover. Mortality in Porites was negatively correlated with depth between 1.5 and 5 m. Using a 50-year dataset of mean monthly sea surface temperature (SST), derived from ship- and satellite-borne instruments, we show that bleaching occurred during a period of exceptionally high summer SST. 1998 was the first year in which mean monthly SSTs exceeded the 1961-1990 upper 95% confidence limit (29.4°C) for a period of three consecutive months. We suggest that the sustained 3-month anomaly in local summer SST was a major cause of coral mortality, but do not discount the synergistic effect of solar radiation. Recovery of the size-frequency distribution of Porites colonies to pre-bleaching levels may take at least 100 years.     

Model of coral population response to accelerated bleaching and mass mortality in a changing climate

We model coral community response to bleaching and mass mortality events which are predicted to increase in frequency with climate change. The model was parameterized for the Arabian/Persian Gulf, but is generally applicable. We assume three species groups (Acropora, faviids, and Porites) in two life-stages each where the juveniles are in competition but the adults can enter a size-refuge in which they cannot be competitively displaced. An aggressive group (Acropora species) dominates at equilibrium, which is not reached due to mass mortality events that primarily disadvantage this group (compensatory mortality, >90% versus 25% in faviids and Porites) roughly every 15 years. Population parameters (N individuals, carrying capacity) were calculated from satellite imagery and in situ transects, vital rates (fecundity, mortality, and survival) were derived from the model, field observations, and literature. It is shown that populations and unaltered community structure can persist despite repeated 90% mortality, given sufficiently high fecundity of the remaining population or import from connected populations. The frequency of disturbance determines the dominant group—in low frequency Acropora, in high frequency Porites. This is congruent with field observations. The model of an isolated population was more sensitive to parameter changes than that of connected populations. Highest sensitivity was to mortality rate and recruitment rate. Community composition was sensitive to spacing of disturbances and level of catastrophic mortality. Decreased mortality led to Acropora dominance, increased mortality led to Acropora extinction. In nature, closely spaced disturbances have severely disadvantaged Acropora populations over the last decade. Unless a longer (>10 years) disturbance-free interval can be maintained, a permanent shift away from Acropora dominance will be observed. A mortality rate of 99% in Acropora, as observed in 1996, is not sustainable if repetitive and neither is a disturbance frequency <15 years—each leading to population collapse. This shows that the severity and/or the spacing of the 1996–1998–2002 disturbances were unusual in frequency and duration.     

The relationship between bleaching and mortality of common corals

Reef corals are likely to have many subtle but four gross responses to anomalous warm water. These are (1) not bleach and live (mortality <10%), (2) not bleach and die (mortality >20%), (3) bleach and live, and (4) bleach and die. The frequency of these four possible gross responses was determined for 18 common coral taxa over an exceptionally warm 1998 El Niño where intense bleaching was observed, and mortality determined from line transects averaged 41.2±34.7 (±SD). Field studies included (1) recording the loss of color (bleaching) and observing recently dead individuals among 6,803 colonies during five sampling periods and (2) estimating mortality based on 180 m of line-intercept transects completed 4 months before and near the end of the bleaching episode. There was no clear relationship between the loss of color and either direct observation or transect-based estimates of mortality for the 18 taxa. The morphology of the taxa did not influence color loss but branching and encrusting taxa had higher mortality than massive and submassive taxa. Loss of color and mortality are the most common responses to warm water as only Pavona did not lose color or die and only two taxa, Cyphastrea and Millepora, did not significantly lose color but died. Of the 15 taxa that lost color, five taxa, Astreopora, Favia, Favites, Goniopora, and Leptoria, did not die. These taxa are those most likely to have reduced potential mortality by the loss of pigments and associated algal symbionts. Death of the branching taxa was detected reasonably by direct field observation but some taxa were underestimated when compared with mortality estimates based on line transects. Death of encrusting and massive taxa including Echinopora, Galaxea, Hydnophora, Montipora, Platygyra, and massive Porites was poorly detected from direct observations but they proved to have modest to high mortality (20–80%) based on line transects. There was no single response of these common corals to warm water but these data, collected during an extreme warm-water anomaly, indicate that the loss of color is most frequently a sign of morbidity, particularly for branching and encrusting taxa.Communicated by P.W. Sammarco, Chauvin     

Coral mortality following extreme low tides and high solar radiation

Extreme tidal events are one of the most predictable natural disturbances in marine benthic habitats and are important determinants of zonation patterns in intertidal benthic communities. On coral reefs, spring low tides are recurrent disturbances, but are rarely reported to cause mass mortality. However, in years when extremely low tides coincide with high noon irradiances, they have the potential to cause widespread damage. Here, we report on such an event on a fringing coral reef in the central Great Barrier Reef (Australia) in September 2005. Visual surveys of colony mortality and bleaching status of more than 13,000 corals at 14 reef sites indicated that most coral taxa at wave-protected sites were severely affected by the event. Between 40 and 75% of colonies in the major coral taxa (Acropora, Porites, Faviidae, Mussidae and Pocilloporidae) were either bleached or suffered partial mortality. In contrast, corals at wave-exposed sites were largely unaffected (<1% of the corals were bleached), as periodic washing by waves prevented desiccation. Surveys along a 1–9 m depth gradient indicated that high coral mortality was confined to the tidal zone. However, 20–30% of faviid colonies were bleached throughout the depth range, suggesting that the increase in benthic irradiances during extreme low tides caused light stress in deeper water. Analyses of an 8-year dataset of tidal records for the area indicated that the combination of extended periods of aerial exposure and high irradiances occurs during May–September in most years, but that the event in September 2005 was the most severe. We argue that extreme low-tide, high-irradiance events are important structuring forces of intertidal coral reef communities, and can be as damaging as thermal stress events. Importantly, they occur at a time of year when risks from thermal stress, cyclones and monsoon-associated river run-off are minimal.     

Lag Effects in the Impacts of Mass Coral Bleaching on Coral Reef Fish, Fisheries, and Ecosystems

Abstract:  Recent episodes of coral bleaching have led to wide-scale loss of reef corals and raised concerns over the effectiveness of existing conservation and management efforts. The 1998 bleaching event was most severe in the western Indian Ocean, where coral declined by up to 90% in some locations. Using fisheries-independent data, we assessed the long-term impacts of this event on fishery target species in the Seychelles, the overall size structure of the fish assemblage, and the effectiveness of two marine protected areas (MPAs) in protecting fish communities. The biomass of fished species above the size retained in fish traps changed little between 1994 and 2005, indicating no current effect on fishery yields. Biomass remained higher in MPAs, indicating they were effective in protecting fish stocks. Nevertheless, the size structure of the fish communities, as described with size-spectra analysis, changed in both fished areas and MPAs, with a decline in smaller fish (<30 cm) and an increase in larger fish (>45 cm). We believe this represents a time-lag response to a reduction in reef structural complexity brought about because fishes are being lost through natural mortality and fishing, and are not being replaced by juveniles. This effect is expected to be greater in terms of fisheries productivity and, because congruent patterns are observed for herbivores, suggests that MPAs do not offer coral reefs long-term resilience to bleaching events. Corallivores and planktivores declined strikingly in abundance, particularly in MPAs, and this decline was associated with a similar pattern of decline in their preferred corals. We suggest that climate-mediated disturbances, such as coral bleaching, be at the fore of conservation planning for coral reefs.     

Social feeding in ladybird beetles: adaptive significance and mechanism

Summary. Young larvae of Adalia bipunctata search an area more intensively when exposed to the odour from other larvae feeding on aphids than when exposed only to the odour of aphids. In an olfactometer young larvae were significantly attracted either to the odour of crushed aphids or larvae feeding on aphids, but not to that of aphids, larvae, larvae plus aphids or larvae feeding on an artificial diet. That is, the change in searching behaviour appears in response to a volatile released by aphids when attacked. The odour released by crushed aphids is made up entirely of aphid alarm pheromone, β-farnesene. It is likely that the adaptive significance of this response is that it increases the ability of larvae to locate larvae that have already caught prey. By sharing the aphid kill of another larva it is likely that a first instar ladybird larva greatly increases its probability of surviving to the next instar. It is suggested that this social feeding is facilitated by egg clustering, which also may additionally account for why aphidophagous ladybirds lay their eggs in clusters. Received 28 February 2000; accepted 24 March 2000     

The adaptive significance of male polymorphism in the acarid mite Caloglyphus berlesei

Summary Two forms of males occur in the acarid mite, Caloglyphus berlesei. One of them uses its thickened third pair of legs to kill other males; the other, without modified legs, does not attack other males. Previous studies have shown that the form of the male is environmentally determined: fighter males develop only at low population densities and their development can be suppressed by substances emanating from dense colonies. In this study, the duration of development, longevity and virility of the two male morphs were measured. The only significant difference was that fighters had a shorter development time between the two last moults. In another experiment, two groups of colonies were maintained under the same conditions but had different numbers of individuals: small colonies contained 2 fighters and 2 non-fighters, whereas large colonies contained 30 males of each type. The relative reproductive success of both morphs was estimated from numbers of matings. In small colonies, the estimated reproductive success (ERS) was significantly higher for fighters than for non-fighters, mainly because in over 50% of these colonies a single fighter male managed to kill all the rival males and monopolize the females. The opposite was true in large colonies, in which non-fighters achieved significantly higher ERS, mainly because the fighters were killed in fights more often than the non-fighters. This implies that the ratio of costs to benefits obtained by adopting the fighter-male strategy increases with the number of rival males, which may explain conditional male development in C. berlesei.     

Increased mortality and photoinhibition in the symbiotic dinoflagellates of the Indo–Pacific coral Stylophora pistillata (Esper) after summer bleaching

Coral bleaching (the loss of symbiotic dinoflagellates from reef-building corals) is most frequently caused by high-light and temperature conditions. We exposed the explants of the hermatypic coral Stylophora pistillata to four combinations of light and temperature in late spring and also in late summer. During mid-summer, two NOAA bleaching warnings were issued for Heron Island reef (Southern Great Barrier Reef, Australia) when sea temperature exceeded the NOAA bleaching threshold, and a ‘mild’ (in terms of the whole coral community) bleaching event occurred, resulting in widespread S. pistillata bleaching and mortality. Symbiotic dinoflagellate biomass decreased by more than half from late spring to late summer (from 2.5×10⁶ to 0.8×10⁶ dinoflagellates cm² coral tissue), and those dinoflagellates that remained after summer became photoinhibited more readily (dark-adapted F _V : F _M decreased to (0.3 compared with 0.4 in spring), and died in greater numbers (up to 17% dinoflagellate mortality compared with 5% in the spring) when exposed to artificially elevated light and temperature. Adding exogenous antioxidants (d-mannitol and l-ascorbic acid) to the water surrounding the coral had no clear effect on either photoinhibition or symbiont mortality. These data show that light and temperature stress cause mortality of the dinoflagellate symbionts within the coral, and that susceptibility to light and temperature stress is strongly related to coral condition. Photoinhibitory mechanisms are clearly involved, and will increase through a positive feedback mechanism: symbiont loss promotes further symbiont loss as the light microenvironment becomes progressively harsher.     

Coral Reef Habitats as Surrogates of Species, Ecological Functions, and Ecosystem Services

Abstract: Habitat maps are often the core spatially consistent data set on which marine reserve networks are designed, but their efficacy as surrogates for species richness and applicability to other conservation measures is poorly understood. Combining an analysis of field survey data, literature review, and expert assessment by a multidisciplinary working group, we examined the degree to which Caribbean coastal habitats provide useful planning information on 4 conservation measures: species richness, the ecological functions of fish species, ecosystem processes, and ecosystem services. Approximately one‐quarter to one‐third of benthic invertebrate species and fish species (disaggregated by life phase; hereafter fish species) occurred in a single habitat, and Montastraea‐dominated forereefs consistently had the highest richness of all species, processes, and services. All 11 habitats were needed to represent all 277 fish species in the seascape, although reducing the conservation target to 95% of species approximately halved the number of habitats required to ensure representation. Species accumulation indices (SAIs) were used to compare the efficacy of surrogates and revealed that fish species were a more appropriate surrogate of benthic species (SAI = 71%) than benthic species were for fishes (SAI = 42%). Species of reef fishes were also distributed more widely across the seascape than invertebrates and therefore their use as a surrogate simultaneously included mangroves, sea grass, and coral reef habitats. Functional classes of fishes served as effective surrogates of fish and benthic species which, given their ease to survey, makes them a particularly useful measure for conservation planning. Ecosystem processes and services exhibited great redundancy among habitats and were ineffective as surrogates of species. Therefore, processes and services in this case were generally unsuitable for a complementarity‐based approach to reserve design. In contrast, the representation of species or functional classes ensured inclusion of all processes and services in the reserve network.     

Symbiont specificity and bleaching susceptibility among soft corals in the 1998 Great Barrier Reef mass coral bleaching event

Considerable variability in bleaching was observed within and among soft coral taxa in the order Alcyonacea (Octocorallia: Cnidaria) on the central Great Barrier Reef (GBR, latitude 18.2°–19.0°S, longitude 146.4°–147.3°E) during the 1998 mass coral bleaching event. In April 1998, during a period of high sea surface temperatures, tissue samples were taken from bleached and unbleached colonies representative of 17 soft coral genera. The genetic identities of intracellular dinoflagellates (Symbiodinium spp.) in these samples were analyzed using PCR-denaturing gradient gel electrophoresis fingerprinting analysis of the internal transcribed spacer regions 1 and 2. Alcyonaceans from the GBR exhibited a high level of symbiont specificity for Symbiodinium types mostly in clade C. A rare clade D type (D3) was associated only with Clavularia koellikeri, while Nephthea sp. hosted symbionts in clade B (B1n and B36). Homogenous Symbiodinium clade populations were detected in all but one colony. Colonies that appeared bleached possessed symbiont types that were genetically indistinguishable from those in nonbleached conspecifics. These data suggest that parameters other than the resident endosymbionts such as host identity and colony acclimatization are important in determining bleaching susceptibility among soft corals. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Climate change, humidity, and mortality in the United States

This paper estimates the effects of humidity and temperature on mortality rates in the United States (c. 1973-2002) in order to provide an insight into the potential health impacts of climate change. I find that humidity, like temperature, is an important determinant of mortality. Coupled with Hadley CM3 climate-change predictions, I project that mortality rates are likely to change little on the aggregate for the United States. However, distributional impacts matter: mortality rates are likely to decline in cold and dry areas, but increase in hot and humid areas. Further, accounting for humidity has important implications for evaluating these distributional effects.     

Reproduction,growth and mortality of the holothurian Cucumaria pseudocurata

Certain phases of the ecology of a sea cucumber (Cucumaria pseudocurata Deichmann, 1938) population at Shell Beach, Sonoma County, California (USA), were investigated. The sea cucumber is a brooder, lays large (1 mm diameter) yolky eggs, which undergo direct development. It forms large aggregations in the rocky intertidal zone, extending from the lower boundary of the Mytilus californianus beds to about zero tide-level. The gonad cycle was followed from September, 1970 to January, 1972. Spawning occurred in January, 1971 and January, 1972. The holothurians produce from 1 to as many as 340 eggs, depending on the size of the individual; they completely spawn-out in a few days. Hatching takes place about 1 month after egg laying. The main sources of mortality are predation by the sea star Pycnopodia helianthoides and wave action. Survival of the young is negatively correlated with density of the aggregation. C. pseudocurata increases in weight 10 to 30 fold in the first year, but much more slowly in later years. Individuals live perhaps 5 years or more, and do not become reproductive until about the third year. Dispersal is accomplished by the young being washed away to a suitable new habitat. The upper range of the intertidal distribution is determined by exposure to air, and the lower range by P. helianthoides predation.Contribution No. 1 of the Bodega Marine Sciences Association.     

Coral recruitment: a spatio-temporal analysis along the coastline of Eilat,northern Red Sea

Recruitment rates of stony corals to artificial substrates were monitored for 2 years at 20 sites along the coast of Eilat, northern Red Sea, to compare with those recorded at other coral reef locations and to assess variation in recruitment at several spatial scales. Coral recruitment was low compared to that observed on the Great Barrier Reef in Australia, but was similar to levels reported from other high-latitude reef locations. Pocilloporids were the most abundant coral recruits in all seasons. Recruitment was twofold higher during the first year than during the second year of study. There was considerable spatial variability, with the largest proportion of variance, apart from the error term, attributable to differences between sites, at a scale of 10² m. Spearmans ranked correlation showed consistency in spatial patterns of recruitment of pocilloporid corals between years, but not of acroporid corals. During spring, when only the brooding pocilloporid coral Stylophora pistillata reproduces at this locality, most coral recruitment occurred at central and southern sites adjacent to well-developed coral reefs. During summer, recruitment patterns varied significantly between years, with wide variation in the recruitment of broadcasting acroporid corals at northern sites located distant from coral reefs. Settlement was low at all sites during autumn and winter. This work is the first detailed analysis of coral recruitment patterns in the Red Sea, and contributes to the understanding of the spatial and temporal scales of variation in this important reef process.Communicated by O. Kinne, Oldendorf/Luhe     

Effects of Aquarium Collectors on Coral Reef Fishes in Kona, Hawaii

Abstract:   No previous studies have conclusively documented the magnitude of the effect of aquarium collecting on natural populations. In Hawaii concern over the effects on reef fish populations of collecting for the aquarium trade began in the early 1970s, primarily in response to multiple-use conflicts between aquarium-fish collectors and recreational dive-tour operators. In 1997–1998 we used a paired control-impact design to estimate the effect of aquarium collectors. We compared differences in fish abundance along visual belt transects between collection sites, where collecting was known to occur, and control sites, where collecting was prohibited. To test the assumptions of our observational design, we surveyed a combination of species captured by aquarium collectors and those not captured. The extent of bleaching, broken coral, and coral cover was also surveyed. Seven of the 10 aquarium species surveyed were significantly reduced by collecting. The abundance of aquarium fish at collection sites ranged from 38% lower ( Chaetodon multicinctus ) to 75% lower ( C. quadrimaculatus ) than that at control sites. In contrast, only two of the nonaquarium species displayed a significant collection effect. There were no significant differences in damaged coral between control and collection sites to indicate the presence of destructive fishing practices. In addition, there were no increases in the abundance of macroalgae where the abundance of herbivores was reduced by aquarium collecting. Although our results suggest that aquarium collectors have a significant effect on the abundance of targeted aquarium fishes, better knowledge of the intensity and location of collecting activities is required to make a rigorous assessment of the effects of collecting on nearshore fish populations. Several lines of evidence suggest that the current system of catch reporting underestimates actual removals.     

Ecosystem ecology meets adaptive management: food web response to a controlled flood on the Colorado River, Glen Canyon

Large dams have been constructed on rivers to meet human demands for water, electricity, navigation, and recreation. As a consequence, flow and temperature regimes have been altered, strongly affecting river food webs and ecosystem processes. Experimental high-flow dam releases, i.e., controlled floods, have been implemented on the Colorado River, U.S.A., in an effort to reestablish pulsed flood events, redistribute sediments, improve conditions for native fishes, and increase understanding of how dam operations affect physical and biological processes. We quantified secondary production and organic matter flows in the food web below Glen Canyon dam for two years prior and one year after an experimental controlled flood in March 2008. Invertebrate biomass and secondary production declined significantly following the flood (total biomass, 55% decline; total production, 56% decline), with most of the decline driven by reductions in two nonnative invertebrate taxa, Potamopyrgus antipodarum and Gammarus lacustris. Diatoms dominated the trophic basis of invertebrate production before and after the controlled flood, and the largest organic matter flows were from diatoms to the three most productive invertebrate taxa (P. antipodarum, G. lacustris, and Tubificida). In contrast to invertebrates, production of rainbow trout (Oncorhynchus mykiss) increased substantially (194%) following the flood, despite the large decline in total secondary production of the invertebrate assemblage. This counterintuitive result is reconciled by a post-flood increase in production and drift concentrations of select invertebrate prey (i.e., Chironomidae and Simuliidae) that supported a large proportion of trout production but had relatively low secondary production. In addition, interaction strengths, measured as species impact values, were strongest between rainbow trout and these two taxa before and after the flood, demonstrating that the dominant consumer-resource interactions were not necessarily congruent with the dominant organic matter flows. Our study illustrates the value of detailed food web analysis for elucidating pathways by which dam management may alter production and strengths of species interactions in river food webs. We suggest that controlled floods may increase production of nonnative rainbow trout, and this information can be used to help guide future dam management decisions.     

Coral architecture affects the habitat choice and form of associated gobiid fishes

Gobiid fishes of the genus Gobiodon live in strong association with certain reef-building corals that vary considerably in size and architecture. These fishes hence are excellent model systems for studying evolutionary adaption to specific microhabitats. Using a sample of Gobiodon histrio and G. rivulatus and their most important host corals (Acropora digitifera and A. gemmifera) from the northern Red Sea, we assess (1) how corals that are occupied by gobies differ in their architecture from colonies that are not occupied and (2) how fish body shape is associated with the architecture of their host coral. Fish body shape was assessed by geometric morphometric techniques. Coral measurements included colony size, branch length (BL), and interbranch as well as branch tip distance of adjacent branches, for which we applied a new and non-destructive measurement technique based on casts of two-component epoxy resin. The most important factor influencing the occupation of corals was a BL of more than 5 cm. The distance between coral branches was clearly related to the width of the fishes and hence constrained overall fish size. G. histrio and G. rivulatus differ in adult body shape as well in their allometric development of lateral body compression, resulting in different maximum body sizes attainable in the restricted interbranch space of corals. The strong dependence of coral-associated fishes on large coral colonies with specific architectures increases the extinction risk of fishes within deteriorating coral reefs.