Evaluation of annual resolution coral geochemical records as climate proxies in the Great Barrier Reef of Australia

Sampling of annually banded massive coral skeletons at annual (or higher) resolutions is increasingly being used to obtain replicate long-term time series of changing seawater conditions. However, few of these studies have compared and calibrated the lower annual resolution records based on coral geochemical tracers with the corresponding instrumental climate records, although some studies have inferred the climatic significance of annual coral series derived from averages of monthly or sub-annual records. Here, we present annual resolution analysis of coral records of elemental and stable isotopic composition that are approximately 70 years long. These records were preserved in two coexisting colonies of Porites sp. from Arlington Reef, on the Great Barrier Reef in Australia, and are used to evaluate the climatic significance of annually resolved coral geochemical proxies. The geochemical records of coral sample “10AR2,” with its faster and relatively constant annual growth rate, appear to have been independent of skeletal growth rate and other vital effects. The annual resolution of Sr/Ca and Δδ¹⁸O time series was shown to be a good proxy for annual sea surface temperature (SST; r = ?0.67, n = 73, p < 0.0000001) and rainfall records (r = ?0.34, n = 67, p < 0.01). However, a slower growing coral sample, “10AR1” showed significantly lower correlations (r = ?0.20, n = 71, p = 0.05 for Sr/Ca and SST; r = ?0.19, n = 67, p = 0.06 for Δδ¹⁸O and rainfall), indicating its greater susceptibility to biological/metabolic effects. Our results suggest that while annually resolved coral records are potentially a valuable tool for determining, in particular, long timescale climate variability such as Pacific Decadal Oscillation, Interdecadal Pacific Oscillation, and other climatic factors, the selection of the coral sample is important, and replication is essential.     

Present limits to heat-adaptability in corals and population-level responses to climate extremes

Climate change scenarios suggest an increase in tropical ocean temperature by 1–3°C by 2099, potentially killing many coral reefs. But Arabian/Persian Gulf corals already exist in this future thermal environment predicted for most tropical reefs and survived severe bleaching in 2010, one of the hottest years on record. Exposure to 33–35°C was on average twice as long as in non-bleaching years. Gulf corals bleached after exposure to temperatures above 34°C for a total of 8 weeks of which 3 weeks were above 35°C. This is more heat than any other corals can survive, providing an insight into the present limits of holobiont adaptation. We show that average temperatures as well as heat-waves in the Gulf have been increasing, that coral population levels will fluctuate strongly, and reef-building capability will be compromised. This, in combination with ocean acidification and significant local threats posed by rampant coastal development puts even these most heat-adapted corals at risk. WWF considers the Gulf ecoregion as “critically endangered”. We argue here that Gulf corals should be considered for assisted migration to the tropical Indo-Pacific. This would have the double benefit of avoiding local extinction of the world''s most heat-adapted holobionts while at the same time introducing their genetic information to populations naïve to such extremes, potentially assisting their survival. Thus, the heat-adaptation acquired by Gulf corals over 6 k, could benefit tropical Indo-Pacific corals who have <100 y until they will experience a similarly harsh climate. Population models suggest that the heat-adapted corals could become dominant on tropical reefs within ∼20 years.     

Cellular responses in sea fan corals: granular amoebocytes react to pathogen and climate stressors

Background

Climate warming is causing environmental change making both marine and terrestrial organisms, and even humans, more susceptible to emerging diseases. Coral reefs are among the most impacted ecosystems by climate stress, and immunity of corals, the most ancient of metazoans, is poorly known. Although coral mortality due to infectious diseases and temperature-related stress is on the rise, the immune effector mechanisms that contribute to the resistance of corals to such events remain elusive. In the Caribbean sea fan corals (Anthozoa, Alcyonacea: Gorgoniidae), the cell-based immune defenses are granular acidophilic amoebocytes, which are known to be involved in wound repair and histocompatibility.

Methodology/Principal Findings

We demonstrate for the first time in corals that these cells are involved in the organismal response to pathogenic and temperature stress. In sea fans with both naturally occurring infections and experimental inoculations with the fungal pathogen Aspergillus sydowii, an inflammatory response, characterized by a massive increase of amoebocytes, was evident near infections. Melanosomes were detected in amoebocytes adjacent to protective melanin bands in infected sea fans; neither was present in uninfected fans. In naturally infected sea fans a concurrent increase in prophenoloxidase activity was detected in infected tissues with dense amoebocytes. Sea fans sampled in the field during the 2005 Caribbean Bleaching Event (a once-in-hundred-year climate event) responded to heat stress with a systemic increase in amoebocytes and amoebocyte densities were also increased by elevated temperature stress in lab experiments.

Conclusions/Significance

The observed amoebocyte responses indicate that sea fan corals use cellular defenses to combat fungal infection and temperature stress. The ability to mount an inflammatory response may be a contributing factor that allowed the survival of even infected sea fan corals during a stressful climate event.     

Marine biological controls on climate via the carbon and sulphur geochemical cycles

We review aspects of the influence of the marine biota on climate, focusing particularly on their role in mediating surface temperatures via their influence on atmospheric carbon dioxide (CO₂) and dimethyl sulphide (DMS) concentrations. Variation in natural CO₂ concentrations occurring over 10³ to 10⁵ years are set by oceanic processes, and in particular by conditions in the Southern Ocean, so it is to this region that we must look to understand the glacial-interglacial changes in CO₂ concentrations. It seems likely that marine productivity in the Southern Ocean is limited by a combination of restricted iron supply to the region and insufficient light. Plankton-produced DMS is thought to influence climate by changing the numbers of cloud condensation nuclei available in remote regions; the efficiency of this mechanism is still unknown, but calculations suggest it may be a powerful influence on climate. It has a much shorter time-scale than the CO₂ effect, and as a consequence may well be a player on the ''global change'' timescale. The direction of both the CO₂ and the DMS mechanisms is such that more marine productivity would lead to lower global temperatures, and we speculate that the overall effect of the marine biota today is to cool the planet by ca. 6°C as a result of these two mechanisms, with one-third of this figure being due to CO₂ effects and two-thirds due to DMS. While the marine biota influence climate, climate also influences the marine biota, chiefly via changing atmospheric circulation. This in turn alters ocean circulation patterns, responsible for mixing up sub-surface nutrients, and also influences the transport of nutrients, such as iron, in atmospheric dust. A more vigorous atmospheric circulation would be expected to increase the productivity of the marine biota on both counts. Thus during glacial time, the colder and drier climate might be expected to stimulate greater marine productivity than occurs today. Since more production leads to greater cooling by reduction in CO₂ and increase in DMS, the marine biota-climate system appears to have been in positive feedback in the glacial-interglacial transition, with the changes in the climate system being reinforced by changes in the marine biota. In the context of anthropogenic change, we cannot at present say what sign the feedback on climate will have, because we have no clear idea whether circulation will become more or less vigorous in the future.     

Late Miocene paleoenvironmental changes in North Africa and the Mediterranean recorded by geochemical proxies (Monte Gibliscemi section,Sicily)

The astronomically tuned marls of the Monte Gibliscemi section, Sicily, constitute an archive to trace the late Miocene palaeoenvironmental conditions (~ 9.7–7.0 Ma) in North Africa. Here we have utilised carbonate content and Al-normalised geochemical proxies to trace changes in terrigenous source area and bottom-water ventilation. The terrigenous input into the section is dominated by North African river systems draining into the Eastern Mediterranean. The proxy parameters indicate that the palaeoenvironmental conditions in North Africa were humid from 9.5 Ma onward with high fluvial input to the Mediterranean. Increases in the Si/Al and Mg/Al ratios occurred from 8.4 to 8.2 Ma and from 8.05 to 7.75 Ma, with maximum values similar to those of Messinian diatomites. These peaks indicate conditions of enhanced biosiliceous productivity and the presence of authigenic clay formation. Sluggish water circulation in the Mediterranean during those times is inferred from the Mn/Al and V/Al behaviour. Late Miocene changes of the Betic (southern Spain) and Rifian (Morocco) Mediterranean–Atlantic gateways are interpreted as the driving force for the changes in water circulation and the transgression associated with the opening of the Rifian corridor is interpreted to occur at 7.8 Ma.     

Patterns of ciliary currents in Atlantic reef corals and their functional significance

Patterns of ciliary currents of 35 species of Atlantic reef corals are described and compared with currents of Pacific corals. Observations were made during the day and at night, during feeding and without food. There is a basic pattern of ciliary currents common to both Atlantic and Pacific species. In all but the family Agaricidae currents flow off the oral disk and up or out between the tentacles. In the centre of the disk region currents flow towards the mouth or the peristome. On the polyp stalk or column there was considerable variation between species in both Atlantic and Pacific forms. In some species currents flow downwards toward the coenosarc while in others, current pass up the stalk towards the tentacles.
In the Atlantic Agaricidae there may be an inward flow towards the mouth, an outward flow or a unidirectional flow across the corallum. The patterns of flow depend upon the state of contraction of the polyps or the shape and proximity of adjacent polyps.
No ciliary current reversal was observed in Atlantic species. Ciliary currents are functional as a cleansing mechanism and facilitated the ability of mucus nets and strands to gather particles.     

Statistical indices in medicine and epidemiology and their classification

Mathematical statistics deals with abstract notions, while medicine solves complicated and many-sided problems. For this reason medical statistics faces some moot points in the interpretation of a number of notions and the classification of statistical indices. In the present article the definition of variables and statistical indices is formulated and their characterization is given. An attempt is made to provide the systematization and natural classification of the latter. Statistical indices are defined as the characteristics of statistical totalities. To classify statistical indices, the most essential signs are used: the character of a variable (external relations), the trend of study (internal content), the form of expression (calculation), derived indices and characteristics (comparison and the results of analysis).     

Correlations of climate and plant ecology to leaf size and shape: potential proxies for the fossil record

The sizes and shapes (physiognomy) of fossil leaves are widely applied as proxies for paleoclimatic and paleoecological variables. However, significant improvements to leaf-margin analysis, used for nearly a century to reconstruct mean annual temperature (MAT), have been elusive; also, relationships between physiognomy and many leaf ecological variables have not been quantified. Using the recently developed technique of digital leaf physiognomy, correlations of leaf physiognomy to MAT, leaf mass per area, and nitrogen content are quantified for a set of test sites from North and Central America. Many physiognomic variables correlate significantly with MAT, indicating a coordinated, convergent evolutionary response of fewer teeth, smaller tooth area, and lower degree of blade dissection in warmer environments. In addition, tooth area correlates negatively with leaf mass per area and positively with nitrogen content. Multiple linear regressions based on a subset of variables produce more accurate MAT estimates than leaf-margin analysis (standard errors of ±2 vs. ±3°C); improvements are greatest at sites with shallow water tables that are analogous to many fossil sites. The multivariate regressions remain robust even when based on one leaf per species, and the model most applicable to fossils shows no more signal degradation from leaf fragmentation than leaf-margin analysis.     

Effect of measured chemical sympathectomy on the dynamics of the chromatin functional indices of rat sympathetic neurocytes during their differentiation

Differences have been found between age-related shifts of DNA template activity of sympathetic neuron chromatin of normal and guanethidine-desympathized rats. The mean level of DNA template activity in mature neurons was the higher the heavier the desympathization of rat. In the ganglia of guanethidine-treated rats a subpopulation of neurocytes was revealed with an unusually high nucleolar-to-nucleoplasmic label ratio, never observed in control rats. The maturational disturbances in the course of alterations in histones visualized by ammoniacal silver, as well as shifts in the percentage of specifically stained nuclei are proportional to the level of desympathization. These changes in histones can be interpreted as a preliminary acceleration and aging of guanethidine-treated rat ganglia which are proportional to the number of destructed cells.     

Evaluating alternative exposure indices in epidemiologic studies on extremely low-frequency magnetic fields

Choosing the right exposure index for epidemiological studies on 50–60 Hz magnetic fields is difficult due to the lack of knowledge about critical exposure parameters for the biological effects of magnetic fields. This paper uses data from a previously published epidemiological investigation on early pregnancy loss (EPL) to study the methods of evaluating the exposure-response relationship of 50 Hz magnetic fields. Two approaches were used. The first approach was to apply generalized additive modeling to suggest the functional form of the relationship between EPL and magnetic field strength. The second approach evaluated the goodness of fit of the EPL data with eight alternative exposure indices: the 24 h average of magnetic field strength, three indices measuring the proportion of time above specified thresholds, and four indices measuring the proportion of time within specified intensity windows. Because the original exposure data included only spot measurements, estimates for the selected exposure indices were calculated indirectly from the spot measurements using empirical nonlinear equations derived from 24 h recordings in 60 residences. The results did not support intensity windows, and a threshold-type dependence on field strength appeared to be more plausible than a linear relationship. In addition, the study produced data suggesting that spot measurements may be used as surrogates for other exposure indices besides the time average field strength. No final conclusions should be drawn from this study alone, but we hope that this exercise stimulates evaluation of alternative exposure indices in other planned and ongoing epidemiological studies. © 1996 Wiley-Liss, Inc.     

Structure-forming corals and sponges and their use as fish habitat in Bering Sea submarine canyons

Continental margins are dynamic, heterogeneous settings that can include canyons, seamounts, and banks. Two of the largest canyons in the world, Zhemchug and Pribilof, cut into the edge of the continental shelf in the southeastern Bering Sea. Here currents and upwelling interact to produce a highly productive area, termed the Green Belt, that supports an abundance of fishes and squids as well as birds and marine mammals. We show that in some areas the floor of these canyons harbors high densities of gorgonian and pennatulacean corals and sponges, likely due to enhanced surface productivity, benthic currents and seafloor topography. Rockfishes, including the commercially important Pacific ocean perch, Sebastes alutus, were associated with corals and sponges as well as with isolated boulders. Sculpins, poachers and pleuronectid flounders were also associated with corals in Pribilof Canyon, where corals were most abundant. Fishes likely use corals and sponges as sources of vertical relief, which may harbor prey as well as provide shelter from predators. Boulders may be equivalent habitat in this regard, but are sparse in the canyons, strongly suggesting that biogenic structure is important fish habitat. Evidence of disturbance to the benthos from fishing activities was observed in these remote canyons. Bottom trawling and other benthic fishing gear has been shown to damage corals and sponges that may be very slow to recover from such disturbance. Regulation of these destructive practices is key to conservation of benthic habitats in these canyons and the ecosystem services they provide.     

In situ measured seasonal variations in F v/F m of two common Red Sea corals

Pulse amplitude modulated (PAM) fluorometry has been suggested as a tool for estimating environmental stresses on corals. However, information regarding natural changes in maximal quantum yields (F _v/F _m) of corals during “normal” (i.e. non-bleaching) years has been limited. In this study, seasonal variations in F _v/F _m for Stylophora pistillata and Favia favus, measured in situ, correlated with seasonal changes in solar irradiance but not in sea temperature. Interactions between sea temperature and irradiance were further studied by growing these corals and Pocillopora damicornis under controlled conditions. Exposure to high light with normal or high temperatures resulted in lower F _v/F _m values than exposure to low light at both temperatures. Thus, high irradiances may cause decreased F _v/F _m values in corals at least as much as, if not more than, high temperatures. Such seasonal variations should be taken into account when using PAM fluorometry as a diagnostic tool for predicting coral bleaching.     

Beyond corals and fish: the effects of climate change on noncoral benthic invertebrates of tropical reefs

Climate change is threatening tropical reefs across the world, with most scientists agreeing that the current changes in climate conditions are occurring at a much faster rate than in the past and are potentially beyond the capacity of reefs to adapt and recover. Current research in tropical ecosystems focuses largely on corals and fishes, although other benthic marine invertebrates provide crucial services to reef systems, with roles in nutrient cycling, water quality regulation, and herbivory. We review available information on the effects of environmental conditions associated with climate change on noncoral tropical benthic invertebrates, including inferences from modern and fossil records. Increasing sea surface temperatures may decrease survivorship and increase the developmental rate, as well as alter the timing of gonad development, spawning, and food availability. The broad latitudinal distribution and associated temperature ranges of several pantropical taxa suggest that some reef communities may have an in‐built adaptive capacity. Tropical benthic invertebrates will also show species‐specific sublethal and lethal responses to sea‐level rise, ocean acidification, physical disturbance, runoff, turbidity, sedimentation, and changes in ocean circulation. In order to accurately predict a species' response to these stressors, we must consider the magnitude and duration of exposure to each stressor, as well as the physiology, mobility, and habitat requirements of the species. Stressors will not act independently, and many organisms will be exposed to multiple stressors concurrently, including anthropogenic stressors. Environmental changes associated with climate change are linked to larger ecological processes, including changes in larval dispersal and recruitment success, shifts in community structure and range extensions, and the establishment and spread of invasive species. Loss of some species will trigger economic losses and negative effects on ecosystem function. Our review is intended to create a framework with which to predict the vulnerability of benthic invertebrates to the stressors associated with climate change, as well as their adaptive capacity. We anticipate that this review will assist scientists, managers, and policy‐makers to better develop and implement regional research and management strategies, based on observed and predicted changes in environmental conditions.     

Atmospheric dimethysulphide production from corals in the Great Barrier Reef and links to solar radiation, climate and coral bleaching

Coral zooxanthellae contain high concentrations of dimethylsulphoniopropionate (DMSP), the precursor of dimethylsulphide (DMS), an aerosol substance that could affect cloud cover, solar radiation and ocean temperatures. Acropora intermedia a dominant staghorn coral in the Indo-Pacific region, contain some of the highest concentrations of DMSP reported in the literature but no studies have shown that corals produce atmospheric DMS in situ and thus could potentially participate in sea surface temperature (SST) regulation over reefs; or how production varies during coral bleaching. We show that A. intermedia from the Great Barrier Reef (GBR) produces significant amounts of atmospheric DMS, in chamber experiments, indicating that coral reefs in this region could contribute to an “ocean thermostat” similar to that described for the western Pacific warm pool, where significantly fewer coral reefs have bleached during the last 25?years because of a cloud-SST feedback. However, when Acropora intermedia was stressed with higher light levels and seawater temperatures DMSP production, an indicator of zooxanthellae expulsion, increased markedly in the chamber, whilst atmospheric DMS emissions almost completely shut down. These results suggest that during increased light levels and seawater temperatures in the GBR coral shut-down atmospheric DMS aerosol production, potentially increasing solar radiation levels over reefs and exacerbating coral bleaching.