Incorporating adaptive responses into future projections of coral bleaching

Climate warming threatens to increase mass coral bleaching events, and several studies have projected the demise of tropical coral reefs this century. However, recent evidence indicates corals may be able to respond to thermal stress though adaptive processes (e.g., genetic adaptation, acclimatization, and symbiont shuffling). How these mechanisms might influence warming‐induced bleaching remains largely unknown. This study compared how different adaptive processes could affect coral bleaching projections. We used the latest bias‐corrected global sea surface temperature (SST) output from the NOAA/GFDL Earth System Model 2 (ESM2M) for the preindustrial period through 2100 to project coral bleaching trajectories. Initial results showed that, in the absence of adaptive processes, application of a preindustrial climatology to the NOAA Coral Reef Watch bleaching prediction method overpredicts the present‐day bleaching frequency. This suggests that corals may have already responded adaptively to some warming over the industrial period. We then modified the prediction method so that the bleaching threshold either permanently increased in response to thermal history (e.g., simulating directional genetic selection) or temporarily increased for 2–10 years in response to a bleaching event (e.g., simulating symbiont shuffling). A bleaching threshold that changes relative to the preceding 60 years of thermal history reduced the frequency of mass bleaching events by 20–80% compared with the ‘no adaptive response’ prediction model by 2100, depending on the emissions scenario. When both types of adaptive responses were applied, up to 14% more reef cells avoided high‐frequency bleaching by 2100. However, temporary increases in bleaching thresholds alone only delayed the occurrence of high‐frequency bleaching by ca. 10 years in all but the lowest emissions scenario. Future research should test the rate and limit of different adaptive responses for coral species across latitudes and ocean basins to determine if and how much corals can respond to increasing thermal stress.     

Caribbean mesophotic coral ecosystems are unlikely climate change refugia

Deeper coral reefs experience reduced temperatures and light and are often shielded from localized anthropogenic stressors such as pollution and fishing. The deep reef refugia hypothesis posits that light‐dependent stony coral species at deeper depths are buffered from thermal stress and will avoid bleaching‐related mass mortalities caused by increasing sea surface temperatures under climate change. This hypothesis has not been tested because data collection on deeper coral reefs is difficult. Here we show that deeper (mesophotic) reefs, 30–75 m depth, in the Caribbean are not refugia because they have lower bleaching threshold temperatures than shallow reefs. Over two thermal stress events, mesophotic reef bleaching was driven by a bleaching threshold that declines 0.26 °C every +10 m depth. Thus, the main premise of the deep reef refugia hypothesis that cooler environments are protective is incorrect; any increase in temperatures above the local mean warmest conditions can lead to thermal stress and bleaching. Thus, relatively cooler temperatures can no longer be considered a de facto refugium for corals and it is likely that many deeper coral reefs are as vulnerable to climate change as shallow water reefs.     

How much time can herbivore protection buy for coral reefs under realistic regimes of hurricanes and coral bleaching?

Coral reefs have been more severely impacted by recent climate instability than any other ecosystem on Earth. Corals tolerate a narrow range of physical environmental stress, and increases in sea temperature of just 1 °C over several weeks can result in mass coral mortality, often exceeding 95% of individuals over hundreds of square kilometres. Even conservative climate models predict that mass coral bleaching events could occur annually by 2050. Unfortunately, managers of coral‐reef resources have few options available to meet this challenge. Here, we investigate the role that fisheries conservation tools, including the designation of marine reserves, can play in altering future trajectories of Caribbean coral reefs. We use an individual‐based model of the ecological dynamics to test the influence of spatially realistic regimes of disturbance on coral populations. Two major sources of disturbance, hurricanes and coral bleaching, are simulated in contrasting regions of the Caribbean: Belize, Bonaire, and the Bahamas. Simulations are extended to 2099 using the HadGEM1 climate model. We find that coral populations can maintain themselves under all levels of hurricane disturbance providing that grazing levels are high. Regional differences in hurricane frequency are found to cause strikingly different spatial patterns of reef health with greater patchiness occurring in Belize, which has less frequent disturbance, than the Bahamas. The addition of coral bleaching led to a much more homogenous reef state over the seascape. Moreover, in the presence of bleaching, all reefs exhibited a decline in health over time, though with substantial variation among regions. Although the protection of herbivores does not prevent reef degradation it does delay rates of coral loss even under the most severe thermal and hurricane regimes. Thus, we can estimate the degree to which local conservation can help buy time for reefs with values ranging between 18 years in the Bahamas and over 50 years in Bonaire, compared with heavily fished systems. Ultimately, we demonstrate that local conservation measures can benefit reef ecosystem services but that their impact will vary spatially and temporally. Recognizing where such management interventions will either help or fail is an important step towards both achieving sustainable use of coral‐reef resources and maximizing resource management investments.     

Mass Coral Reef Bleaching: A Recent Outcome of Increased El Niño Activity?

Coral reefs are generally considered to be the most biologically productive of all marine ecosystems, but in recent times these vulnerable aquatic resources have been subject to unusual degradation. The general decline in reefs has been greatly accelerated by mass bleaching in which corals whiten en masse and often fail to recover. Empirical evidence indicates a coral reef bleaching cycle in which major bleaching episodes are synchronized with El Niño events that occur every 3–4 years on average. By heating vast areas of the Pacific Ocean, and affecting the Indian and Atlantic Oceans as well, El Niño causes widespread damage to reefs largely because corals are very sensitive to temperature changes. However, mass bleaching events were rarely observed before the 1970s and their abrupt appearance two decades ago remains an enigma. Here we propose a new explanation for the sudden occurrence of mass bleaching and show that it may be a response to the relative increase in El Niño experienced over the last two decades.     

Short-Term Coral Bleaching Is Not Recorded by Skeletal Boron Isotopes

Coral skeletal boron isotopes have been established as a proxy for seawater pH, yet it remains unclear if and how this proxy is affected by seawater temperature. Specifically, it has never been directly tested whether coral bleaching caused by high water temperatures influences coral boron isotopes. Here we report the results from a controlled bleaching experiment conducted on the Caribbean corals Porites divaricata, Porites astreoides, and Orbicella faveolata. Stable boron (δ¹¹B), carbon (δ¹³C), oxygen (δ¹⁸O) isotopes, Sr/Ca, Mg/Ca, U/Ca, and Ba/Ca ratios, as well as chlorophyll a concentrations and calcification rates were measured on coral skeletal material corresponding to the period during and immediately after the elevated temperature treatment and again after 6 weeks of recovery on the reef. We show that under these conditions, coral bleaching did not affect the boron isotopic signature in any coral species tested, despite significant changes in coral physiology. This contradicts published findings from coral cores, where significant decreases in boron isotopes were interpreted as corresponding to times of known mass bleaching events. In contrast, δ¹³C and δ¹⁸O exhibited major enrichment corresponding to decreases in calcification rates associated with bleaching. Sr/Ca of bleached corals did not consistently record the 1.2°C difference in seawater temperature during the bleaching treatment, or alternatively show a consistent increase due to impaired photosynthesis and calcification. Mg/Ca, U/Ca, and Ba/Ca were affected by coral bleaching in some of the coral species, but the observed patterns could not be satisfactorily explained by temperature dependence or changes in coral physiology. This demonstrates that coral boron isotopes do not record short-term bleaching events, and therefore cannot be used as a proxy for past bleaching events. The robustness of coral boron isotopes to changes in coral physiology, however, suggests that reconstruction of seawater pH using boron isotopes should be uncompromised by short-term bleaching events.     

Endolithic algae: an alternative source of photoassimilates during coral bleaching

Recent reports of worldwide coral bleaching events leading to devastating coral mortality have caused alarm among scientists and resource managers. Differential survival of coral species through bleaching events has been widely documented. We suggest that among the possible factors contributing to survival of coral species during such events are endolithic algae harboured in their skeleton, providing an alternative source of energy. We studied the dynamics of photosynthetic pigment concentrations and biomass of endoliths in the skeleton of the encrusting coral Oculina patagonica throughout a bleaching event. During repeated summer bleaching events these endolithic algae receive increased photosynthetically active radiation, increase markedly in biomass, and produce increasing amounts of photoassimilates, which are translocated to the coral. Chlorophyll concentrations and biomass of endoliths were 4.6 +/- 1.57 and 1570 +/- 427 microg cm(-2) respectively, in skeletons of relatively healthy colonies (0-40% bleaching) but up to 14.8 +/- 2.5 and 4036 +/- 764 microg cm(-2) endolith chlorophyll and biomass respectively, in skeletons of bleached colonies (greater than 40% bleaching). The translocation dynamics of (14)C-labelled photoassimilates from the endoliths to bleached coral tissue showed significantly higher 14C activity of the endoliths harboured within the skeletons of bleached corals than that of the endoliths in non-bleached corals. This alternative source of energy may be vital for the survivorship of O. patagonica, allowing gradual recruitment of zooxanthellae and subsequent recovery during the following winter.     

Validation of degree heating weeks as a coral bleaching index in the northwestern Pacific

Mass bleaching is the most significant threat to coral reefs. The United States National Oceanic and Atmospheric Administration monitors world sea surface temperature (SST) and releases warnings for bleaching based on degree heating weeks (DHW), which is the accumulation of temperature anomalies exceeding the monthly maximum mean SST for a given region. DHW values >4.0 °C-weeks are thought to induce bleaching, and those >8.0 °C-weeks are thought to result in widespread bleaching and some mortality. This study validates the effectiveness of DHW as a mass bleaching index by on-site historical observation at eight sites in the northwestern Pacific Ocean. The mass bleaching events occurred during different years at different sites. The recorded years of the bleaching events matched well with DHW values >8 °C-weeks, and the logistically projected probability of bleaching against DHW showed a positive relationship. DHW provides a reasonable threshold for bleaching.     

Marine heatwaves reveal coral reef zones susceptible to bleaching in the Red Sea

As the Earth's temperature continues to rise, coral bleaching events become more frequent. Some of the most affected reef ecosystems are located in poorly monitored waters, and thus, the extent of the damage is unknown. We propose the use of marine heatwaves (MHWs) as a new approach for detecting coral reef zones susceptible to bleaching, using the Red Sea as a model system. Red Sea corals are exceptionally heat‐resistant, yet bleaching events have increased in frequency. By applying a strict definition of MHWs on >30 year satellite‐derived sea surface temperature observations (1985–2015), we provide an atlas of MHW hotspots over the Red Sea coral reef zones, which includes all MHWs that caused major coral bleaching. We found that: (a) if tuned to a specific set of conditions, MHWs identify all areas where coral bleaching has previously been reported; (b) those conditions extended farther and occurred more often than bleaching was reported; and (c) an emergent pattern of extreme warming events is evident in the northern Red Sea (since 1998), a region until now thought to be a thermal refuge for corals. We argue that bleaching in the Red Sea may be vastly underrepresented. Additionally, although northern Red Sea corals exhibit remarkably high thermal resistance, the rapidly rising incidence of MHWs of high intensity indicates this region may not remain a thermal refuge much longer. As our regionally tuned MHW algorithm was capable of isolating all extreme warming events that have led to documented coral bleaching in the Red Sea, we propose that this approach could be used to reveal bleaching‐prone regions in other data‐limited tropical regions. It may thus prove a highly valuable tool for policymakers to optimize the sustainable management of coastal economic zones.     

Historical temperature variability affects coral response to heat stress

Coral bleaching is the breakdown of symbiosis between coral animal hosts and their dinoflagellate algae symbionts in response to environmental stress. On large spatial scales, heat stress is the most common factor causing bleaching, which is predicted to increase in frequency and severity as the climate warms. There is evidence that the temperature threshold at which bleaching occurs varies with local environmental conditions and background climate conditions. We investigated the influence of past temperature variability on coral susceptibility to bleaching, using the natural gradient in peak temperature variability in the Gilbert Islands, Republic of Kiribati. The spatial pattern in skeletal growth rates and partial mortality scars found in massive Porites sp. across the central and northern islands suggests that corals subject to larger year-to-year fluctuations in maximum ocean temperature were more resistant to a 2004 warm-water event. In addition, a subsequent 2009 warm event had a disproportionately larger impact on those corals from the island with lower historical heat stress, as indicated by lower concentrations of triacylglycerol, a lipid utilized for energy, as well as thinner tissue in those corals. This study indicates that coral reefs in locations with more frequent warm events may be more resilient to future warming, and protection measures may be more effective in these regions.     

Turbid reefs moderate coral bleaching under climate‐related temperature stress

Thermal‐stress events that cause coral bleaching and mortality have recently increased in frequency and severity. Yet few studies have explored conditions that moderate coral bleaching. Given that high light and high ocean temperature together cause coral bleaching, we explore whether corals at turbid localities, with reduced light, are less likely to bleach during thermal‐stress events than corals at other localities. We analyzed coral bleaching, temperature, and turbidity data from 3,694 sites worldwide with a Bayesian model and found that K_d490, a measurement positively related to turbidity, between 0.080 and 0.127 reduced coral bleaching during thermal‐stress events. Approximately 12% of the world's reefs exist within this “moderating turbidity” range, and 30% of reefs that have moderating turbidity are in the Coral Triangle. We suggest that these turbid nearshore environments may provide some refuge through climate change, but these reefs will need high conservation status to sustain them close to dense human populations.     

Larval connectivity across temperature gradients and its potential effect on heat tolerance in coral populations

Coral reefs are increasingly exposed to elevated temperatures that can cause coral bleaching and high levels of mortality of corals and associated organisms. The temperature threshold for coral bleaching depends on the acclimation and adaptation of corals to the local maximum temperature regime. However, because of larval dispersal, coral populations can receive larvae from corals that are adapted to very different temperature regimes. We combine an offline particle tracking routine with output from a high‐resolution physical oceanographic model to investigate whether connectivity of coral larvae between reefs of different thermal regimes could alter the thermal stress threshold of corals. Our results suggest that larval transport between reefs of widely varying temperatures is likely in the Coral Triangle and that accounting for this connectivity may be important in bleaching predictions. This has important implications in conservation planning, because connectivity may allow some reefs to have an inherited heat tolerance that is higher or lower than predicted based on local conditions alone.     

Life histories predict coral community disassembly under multiple stressors

Climate change is reshaping biological communities against a background of existing human pressure. Evaluating the impacts of multiple stressors on community dynamics can be particularly challenging in species‐rich ecosystems, such as coral reefs. Here, we investigate whether life‐history strategies and cotolerance to different stressors can predict community responses to fishing and temperature‐driven bleaching using a 20‐year time series of coral assemblages in Kenya. We found that the initial life‐history composition of coral taxa largely determined the impacts of bleaching and coral loss. Prior to the 1998 bleaching event, coral assemblages within no‐take marine reserves were composed of three distinct life histories – competitive, stress‐tolerant and weedy– and exhibited strong declines following bleaching with limited subsequent recovery. In contrast, fished reefs had lower coral cover, fewer genera and were composed of stress‐tolerant and weedy corals that were less affected by bleaching over the long term. Despite these general patterns, we found limited evidence for cotolerance as coral genera and life histories were variable in their sensitivities to fishing and bleaching. Overall, fishing and bleaching have reduced coral diversity and led to altered coral communities of ‘survivor’ species with stress‐tolerant and weedy life histories. Our findings are consistent with expectations that climate change interacting with existing human pressure will result in the loss of coral diversity and critical reef habitat.     

A coral reef refuge in the Red Sea

The stability and persistence of coral reefs in the decades to come is uncertain due to global warming and repeated bleaching events that will lead to reduced resilience of these ecological and socio‐economically important ecosystems. Identifying key refugia is potentially important for future conservation actions. We suggest that the Gulf of Aqaba (GoA) (Red Sea) may serve as a reef refugium due to a unique suite of environmental conditions. Our hypothesis is based on experimental detection of an exceptionally high bleaching threshold of northern Red Sea corals and on the potential dispersal of coral planulae larvae through a selective thermal barrier estimated using an ocean model. We propose that millennia of natural selection in the form of a thermal barrier at the southernmost end of the Red Sea have selected coral genotypes that are less susceptible to thermal stress in the northern Red Sea, delaying bleaching events in the GoA by at least a century.     

Large-scale stress factors affecting coral reefs: open ocean sea surface temperature and surface seawater aragonite saturation over the next 400 years

One-third of the world’s coral reefs have disappeared over the last 30 years, and a further third is under threat today from various stress factors. The main global stress factors on coral reefs have been identified as changes in sea surface temperature (SST) and changes in surface seawater aragonite saturation (Ω_arag). Here, we use a climate model of intermediate complexity, which includes an ocean general circulation model and a fully coupled carbon cycle, in conjunction with present-day observations of inter-annual SST variability to investigate three IPCC representative concentration pathways (RCP 3PD, RCP 4.5, and RCP 8.5), and their impact on the environmental stressors of coral reefs related to open ocean SST and open ocean Ω_arag over the next 400 years. Our simulations show that for the RCP 4.5 and 8.5 scenarios, the threshold of 3.3 for zonal and annual mean Ω_arag would be crossed in the first half of this century. By year 2030, 66–85% of the reef locations considered in this study would experience severe bleaching events at least once every 10 years. Regardless of the concentration pathway, virtually every reef considered in this study (>97%) would experience severe thermal stress by year 2050. In all our simulations, changes in surface seawater aragonite saturation lead changes in temperatures.     

When depth is no refuge: cumulative thermal stress increases with depth in Bocas del Toro,Panama

Coral reefs are increasingly affected by high-temperature stress events and associated bleaching. Monitoring and predicting these events have largely utilized sea surface temperature data, due to the convenience of using large-scale remotely sensed satellite measurements. However, coral bleaching has been observed to vary in severity throughout the water column, and variations in coral thermal stress across depths have not yet been well investigated. In this study, in situ water temperature data from 1999 to 2011 from three depths were used to calculate thermal stress on a coral reef in Bahia Almirante, Bocas del Toro, Panama, which was compared to satellite surface temperature data and thermal stress calculations for the same area and time period from the National Oceanic and Atmospheric Administration Coral Reef Watch Satellite Bleaching Alert system. The results show similar total cumulative annual thermal stress for both the surface and depth-stratified data, but with a striking difference in the distribution of that stress among the depth strata during different high-temperature events, with the greatest thermal stress unusually recorded at the deepest measured depth during the most severe bleaching event in 2005. Temperature records indicate that a strong density-driven temperature inversion may have formed in this location in that year, contributing to the persistence and intensity of bleaching disturbance at depth. These results indicate that depth may not provide a stress refuge from high water temperature events in some situations, and in this case, the water properties at depth appear to have contributed to greater coral bleaching at depth compared to near-surface locations. This case study demonstrates the importance of incorporating depth-stratified temperature monitoring and small-scale oceanographic and hydrologic data for understanding and predicting local reef responses to elevated water temperature events.     

Boring sponges,an increasing threat for coral reefs affected by bleaching events

Coral bleaching is a stress response of corals induced by a variety of factors, but these events have become more frequent and intense in response to recent climate‐change‐related temperature anomalies. We tested the hypothesis that coral reefs affected by bleaching events are currently heavily infested by boring sponges, which are playing a significant role in the destruction of their physical structure. Seventeen reefs that cover the entire distributional range of corals along the Mexican Pacific coast were studied between 2005/2006, and later between 2009/2010. Most of these coral reefs were previously impacted by bleaching events, which resulted in coral mortalities. Sponge abundance and species richness was used as an indicator of bioerosion, and coral cover was used to describe the present condition of coral reefs. Coral reefs are currently highly invaded (46% of the samples examined) by a very high diversity of boring sponges (20 species); being the coral reef framework the substrate most invaded (56%) followed by the rubbles (45%), and the living colonies (36%). The results also indicated that boring sponges are promoting the dislodgment of live colonies and large fragments from the framework. In summary, the eastern coral reefs affected by bleaching phenomena, mainly provoked by El Niño, present a high diversity and abundance of boring sponges, which are weakening the union of the colony with the reef framework and promoting their dislodgment. These phenomena will probably become even more intense and severe, as temperatures are projected to continue to rise under the scenarios for future climate change, which could place many eastern coral reefs beyond their survival threshold.     

Effect of colony size and surrounding substrate on corals experiencing a mild bleaching event on Heron Island reef flat (southern Great Barrier Reef,Australia)

In January–May 2006, Heron Island in the Great Barrier Reef experienced a mild bleaching event. The effect of colony size, morphology and surrounding substrate on the extent of bleaching was explored. In contrast with previous studies, colony size did not influence bleaching sensitivity, suggesting that there may be a threshold of light and temperature stress beyond which size plays a role. Also contrasting with previous studies, massive corals were more affected by bleaching than branching corals. Massive corals surrounded by sand were more affected than the ones surrounded by rubble or dead coral. It is hypothesized that light reflectance from sand increases stress levels experienced by the colonies. This effect is maximized in massive corals as opposed to branching corals that form dense thickets on Heron Island. These results emphasize the importance of the ecological dynamics of coral communities experiencing low, moderate and high levels of bleaching for the understanding of how coral communities may change under the stress of climate change.     

Sugar enrichment provides evidence for a role of nitrogen fixation in coral bleaching

The disruption of the coral–algae symbiosis (coral bleaching) due to rising sea surface temperatures has become an unprecedented global threat to coral reefs. Despite decades of research, our ability to manage mass bleaching events remains hampered by an incomplete mechanistic understanding of the processes involved. In this study, we induced a coral bleaching phenotype in the absence of heat and light stress by adding sugars. The sugar addition resulted in coral symbiotic breakdown accompanied by a fourfold increase of coral‐associated microbial nitrogen fixation. Concomitantly, increased N:P ratios by the coral host and algal symbionts suggest excess availability of nitrogen and a disruption of the nitrogen limitation within the coral holobiont. As nitrogen fixation is similarly stimulated in ocean warming scenarios, here we propose a refined coral bleaching model integrating the cascading effects of stimulated microbial nitrogen fixation. This model highlights the putative role of nitrogen‐fixing microbes in coral holobiont functioning and breakdown.     

Predicting coral bleaching hotspots: the role of regional variability in thermal stress and potential adaptation rates

Sea surface temperature fields (1870–2100) forced by CO₂-induced climate change under the IPCC SRES A1B CO₂ scenario, from three World Climate Research Programme Coupled Model Intercomparison Project Phase 3 (WCRP CMIP3) models (CCSM3, CSIRO MK 3.5, and GFDL CM 2.1), were used to examine how coral sensitivity to thermal stress and rates of adaption affect global projections of coral-reef bleaching. The focus of this study was two-fold, to: (1) assess how the impact of Degree-Heating-Month (DHM) thermal stress threshold choice affects potential bleaching predictions and (2) examine the effect of hypothetical adaptation rates of corals to rising temperature. DHM values were estimated using a conventional threshold of 1°C and a variability-based threshold of 2σ above the climatological maximum Coral adaptation rates were simulated as a function of historical 100-year exposure to maximum annual SSTs with a dynamic rather than static climatological maximum based on the previous 100 years, for a given reef cell. Within CCSM3 simulations, the 1°C threshold predicted later onset of mild bleaching every 5 years for the fraction of reef grid cells where 1°C > 2σ of the climatology time series of annual SST maxima (1961–1990). Alternatively, DHM values using both thresholds, with CSIRO MK 3.5 and GFDL CM 2.1 SSTs, did not produce drastically different onset timing for bleaching every 5 years. Across models, DHMs based on 1°C thermal stress threshold show the most threatened reefs by 2100 could be in the Central and Western Equatorial Pacific, whereas use of the variability-based threshold for DHMs yields the Coral Triangle and parts of Micronesia and Melanesia as bleaching hotspots. Simulations that allow corals to adapt to increases in maximum SST drastically reduce the rates of bleaching. These findings highlight the importance of considering the thermal stress threshold in DHM estimates as well as potential adaptation models in future coral bleaching projections.     

Coral bleaching: causes and consequences

It has been over 10 years since the phenomenon of extensive coral bleaching was first described. In most cases bleaching has been attributed to elevated temperature, but other instances involving high solar irradiance, and sometimes disease, have also been documented. It is timely, in view of our concern about worldwide reef condition, to review knowledge of physical and biological factors involved in bleaching, the mechanisms of zooxanthellae and pigment loss, and the ecological consequences for coral communities. Here we evaluate recently acquired data on temperature and irradiance-induced bleaching, including long-term data sets which suggest that repeated bleaching events may be the consequence of a steadily rising background sea temperature that will in the future expose corals to an increasingly hostile environment. Cellular mechanisms of bleaching involve a variety of processes that include the degeneration of zooxanthellae in situ, release of zooxanthellae from mesenterial filaments and release of algae within host cells which become detached from the endoderm. Photo-protective defences (particularly carotenoid pigments) in zooxanthellae are likely to play an important role in limiting the bleaching response which is probably elicited by a combination of elevated temperature and irradiance in the field. The ability of corals to respond adaptively to recurrent bleaching episodes is not known, but preliminary evidence suggests that phenotypic responses of both corals and zooxanthellae may be significant.