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Coral reef ecosystems have suffered an unprecedented loss of habitat-forming hard corals in recent decades. While marine conservation has historically focused on passive habitat protection, demand for and interest in active restoration has been growing in recent decades. However, a disconnect between coral restoration practitioners, coral reef managers and scientists has resulted in a disjointed field where it is difficult to gain an overview of existing knowledge. To address this, we aimed to synthesise the available knowledge in a comprehensive global review of coral restoration methods, incorporating data from the peer-reviewed scientific literature, complemented with grey literature and through a survey of coral restoration practitioners. We found that coral restoration case studies are dominated by short-term projects, with 60% of all projects reporting less than 18 months of monitoring of the restored sites. Similarly, most projects are relatively small in spatial scale, with a median size of restored area of 100 m 2. A diverse range of species are represented in the dataset, with 229 different species from 72 coral genera. Overall, coral restoration projects focused primarily on fast-growing branching corals (59% of studies), and report survival between 60 and 70%. To date, the relatively young field of coral restoration has been plagued by similar 'growing pains' as ecological restoration in other ecosystems. These include 1) a lack of clear and achievable objectives, 2) a lack of appropriate and standardised monitoring and reporting and, 3) poorly designed projects in relation to stated objectives. Mitigating these will be crucial to successfully scale up projects, and to retain public trust in restoration as a tool for resilience based management. Finally, while it is clear that practitioners have developed effective methods to successfully grow corals at small scales, it is critical not to view restoration as a replacement for meaningful action on climate change.

Fast repetition rate (FRR) fluorescence can provide highly resolved estimates of light absorption by photosystem II (PSII), a variable that is critical to bio-optical determinations of phytoplankton productivity. We compared estimates of chlorophyll a-specific light absorption by PSII, a chl PSII , using both biophysical (FRR) and optical (chlorophyll a-specific light absorption coefficient, a chl ) techniques on cultures of phytoplankton from diverse taxa. Biophysical determinations of a chl PSII were obtained from the product of the effective light absorption cross-section of PSII (σ PSII ), measured by FRR fluorescence, and the ratio of PSII reaction centers to chlorophyll a (n PSII ), measured by oxygen flash yields. Both parameters were highly variable between individual taxa. In mixtures of algae, the estimates of σ PSII were largely determined by the taxon that dominated fluorescence intensity. Independent optical estimates of a chl PSII were obtained by weighting the light absorption spectra of photosynthetic pigments by the fluorescence excitation spectra. The biophysical and optical estimates of a chl PSII were highly correlated (r 2 = 0.94) with a slope that was not significantly different from 1 and an intercept of 0. Estimates of productivity using biophysical or optical absorption measurements should therefore be comparable when the latter are adjusted to account for the proportion of light provided for photochemistry into both photosystem I and II. Finally, we show how simultaneous measurements of σ PSII and a chl PSII can be used to derive n PSII where flash-yield measurements are impractical, as is almost universally the case in field measurements.

Tropical reefs have been impacted by thermal anomalies caused by global warming that induced coral bleaching and mortality events globally. However, there have only been very few recordings of bleaching within the Red Sea despite covering a latitudinal range of 15° and consequently it has been considered a region that is less sensitive to thermal anomalies. We therefore examined historical patterns of sea surface temperature (SST) and associated anomalies (1982-2012) and compared warming trends with a unique compilation of corresponding coral bleaching records from throughout the region. These data indicated that the northern Red Sea has not experienced mass bleaching despite intensive Degree Heating Weeks (DHW) of >15°C-weeks. Severe bleaching was restricted to the central and southern Red Sea where DHWs have been more frequent, but far less intense (DHWs <4°C-weeks). A similar pattern was observed during the 2015-2016 El Niño event during which time corals in the northern Red Sea did not bleach despite high thermal stress (i.e. DHWs >8°C-weeks), and bleaching was restricted to the central and southern Red Sea despite the lower thermal stress (DHWs < 8°C-weeks). Heat stress assays carried out in the northern (Hurghada) and central (Thuwal) Red Sea on four key reef-building species confirmed different regional thermal susceptibility, and that central Red Sea corals are more sensitive to thermal anomalies as compared to those from the north. Together, our data demonstrate that corals in the northern Red Sea have a much higher heat tolerance than their prevailing temperature regime would suggest. In contrast, corals from the central Red Sea are close to their thermal limits, which closely match the maximum annual water temperatures. The northern Red Sea harbours reef-building corals that live well below their bleaching thresholds and thus we propose that the region represents a thermal refuge of global importance.

Phytoplankton primary productivity is most commonly measured by 14 C assimilation although less direct methods, such as O 2 exchange, have also been employed. These methods are invasive, requiring bottle incubation for up to 24 h. As an alternative, Fast Repetition Rate fluorometry (FRRf) has been used, on wide temporal and spatial scales within aquatic systems, to estimate photosystem II (PSII) electron flux per unit volume (JV PSII ), which generally correlates well with photosynthetic O 2 evolution. A major limitation of using FRRf arises from the need to employ an independent method to determine the concentration of functional photosystem II reaction centers ([RCII]); a requirement that has prevented FRR fluorometers being used, as stand-alone instruments, for the estimation of electron transport. Within this study, we have taken a new approach to the analysis of FRRf data, based on a simple hypothesis; that under a given set of environmental conditions, the ratio of rate constants for RCII fluorescence emission and photochemistry falls within a narrow range, for all groups of phytoplankton. We present a simple equation, derived from the established FRRf algorithm, for determining [RCII] from dark FRRf data alone. We also describe an entirely new algorithm for estimating JV PSII , which does not require determination of [RCII] and is valid for a heterogeneous model of connectivity among RCIIs. Empirical supporting evidence is presented. These data are derived from FRR measurements across a diverse range of microalgae, in parallel with independent measurements of [RCII]. Possible sources of error, particularly under nutrient stress conditions, are discussed. *Corresponding author: E-mail: koxborough@chelsea.co.uk AcknowledgmentsWe would like to thank Tania Cresswell-Maynard (University of Essex) for providing many of the cultures used within this study and Kimberley Walrond (University of Manchester) for helpful discussions. Contributions of DJS and RJG were supported by the EU project PRO-TOOL (EU-226880). The contribution of CMM was supported by the Natural Environment Research Council, UK (NE/G009155/1). The authors owe extreme thanks to Hugh MacIntyre (Dalhousie University, Halifax, Canada) and Marie-Hélène Forget (Bedford Institute of Oceanography, Dartmouth, Canada) for their assistance and contributions in facilitating the Mk I FASTtracka data sets evaluated here.

Occurrences whereby cnidaria lose their symbiotic dinoflagellate microalgae (Symbiodinium spp.) are increasing in frequency and intensity. These so-called bleaching events are most often related to an increase in water temperature, which is thought to limit certain Symbiodinium phylotypes from effectively dissipating absorbed excitation energy that is otherwise used for photochemistry. Here, we examined photosynthetic characteristics and hydrogen peroxide (H2 O2 ) production, a possible signal involved in bleaching, from two Symbiodinium types (a thermally "tolerant" A1 and "sensitive" B1) representative of cnidaria-Symbiodinium symbioses of reef-building Caribbean corals. Under steady-state growth at 26°C, a higher efficiency of PSII photochemistry, rate of electron turnover, and rate of O2 production were observed for type A1 than for B1. The two types responded very differently to a period of elevated temperature (32°C): type A1 increased light-driven O2 consumption but not the amount of H2 O2 produced; in contrast, type B1 increased the amount of H2 O2 produced without an increase in light-driven O2 consumption. Therefore, our results are consistent with previous suggestions that the thermal tolerance of Symbiodinium is related to adaptive constraints associated with photosynthesis and that sensitive phylotypes are more prone to H2 O2 production. Understanding these adaptive differences in the genus Symbiodinium will be crucial if we are to interpret the response of symbiotic associations, including reef-building corals, to environmental change.

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The bleaching of corals in response to increases in temperature has resulted in significant coral reef degradation in many tropical marine ecosystems. This bleaching has frequently been attributed to photoinhibition of photosynthetic electron transport and the consequent photodamage to photosystem II (PSII) and the production of damaging reactive oxygen species (ROS) in the zooxanthellae (Symbiodinium spp.). However, these events may be because of perturbations of other processes occurring within the zooxanthellae or the host cells, and consequently constitute only secondary responses to temperature increase. The processes involved with the onset of photoinhibition of electron transport, photodamage to PSII and pigment bleaching in coral zooxanthellae are reviewed. Consideration is given to how increases in temperature might lead to perturbations of metabolic processes in the zooxanthellae and/or their host cells, which could trigger events leading to bleaching. It is concluded that production of ROS by the thylakoid photosynthetic apparatus in the zooxanthellae plays a major role in the onset of bleaching resulting from photoinhibition of photosynthesis, although it is not clear which particular ROS are involved. It is suggested that hydrogen peroxide generated in the zooxanthellae may have a signalling role in triggering the mechanisms that result in expulsion of zooxanthellae from corals.