Positive genetic associations among fitness traits support evolvability of a reef‐building coral under multiple stressors

Climate change threatens organisms in a variety of interactive ways that requires simultaneous adaptation of multiple traits. Predicting evolutionary responses requires an understanding of the potential for interactions among stressors and the genetic variance and covariance among fitness‐related traits that may reinforce or constrain an adaptive response. Here we investigate the capacity of Acropora millepora, a reef‐building coral, to adapt to multiple environmental stressors: rising sea surface temperature, ocean acidification, and increased prevalence of infectious diseases. We measured growth rates (weight gain), coral color (a proxy for Symbiodiniaceae density), and survival, in addition to nine physiological indicators of coral and algal health in 40 coral genets exposed to each of these three stressors singly and combined. Individual stressors resulted in predicted responses (e.g., corals developed lesions after bacterial challenge and bleached under thermal stress). However, corals did not suffer substantially more when all three stressors were combined. Nor were trade‐offs observed between tolerances to different stressors; instead, individuals performing well under one stressor also tended to perform well under every other stressor. An analysis of genetic correlations between traits revealed positive covariances, suggesting that selection to multiple stressors will reinforce rather than constrain the simultaneous evolution of traits related to holobiont health (e.g., weight gain and algal density). These findings support the potential for rapid coral adaptation under climate change and emphasize the importance of accounting for corals’ adaptive capacity when predicting the future of coral reefs.     

Threatened Caribbean Coral Is Able to Mitigate the Adverse Effects of Ocean Acidification on Calcification by Increasing Feeding Rate

Global climate change threatens coral growth and reef ecosystem health via ocean warming and ocean acidification (OA). Whereas the negative impacts of these stressors are increasingly well-documented, studies identifying pathways to resilience are still poorly understood. Heterotrophy has been shown to help corals experiencing decreases in growth due to either thermal or OA stress; however, the mechanism by which it mitigates these decreases remains unclear. This study tested the ability of coral heterotrophy to mitigate reductions in growth due to climate change stress in the critically endangered Caribbean coral Acropora cervicornis via changes in feeding rate and lipid content. Corals were either fed or unfed and exposed to elevated temperature (30°C), enriched pCO₂ (800 ppm), or both (30°C/800 ppm) as compared to a control (26°C/390 ppm) for 8 weeks. Feeding rate and lipid content both increased in corals experiencing OA vs. present-day conditions, and were significantly correlated. Fed corals were able to maintain ambient growth rates at both elevated temperature and elevated CO₂, while unfed corals experienced significant decreases in growth with respect to fed conspecifics. Our results show for the first time that a threatened coral species can buffer OA-reduced calcification by increasing feeding rates and lipid content.     

Effects of the multiple stressors high temperature and reduced salinity on the photosynthesis of the hermatypic coral Galaxea fascicularis

This study investigates the physiological responses in the hermatypic coral Galaxea fascicularis exposed to salinity stress (from 37 ppt to 15 ppt) for 12 h, combined effects of reduced salinity (from 37 ppt to 20 ppt) and two temperatures (26 °C and 32 °C) for 12 h and combined effects of reduced salinity (from 37 ppt to 25 ppt) and two temperatures (26 °C and 29.5 °C) for 10 d. The results demonstrate that the coral is tolerant to 12 h exposure to extremely low salinity (15 ppt). The study also shows that combined effects of temperature and low salinity aggravate the damage on the photosynthesis of the symbiotic dinoflagellates in 12 h exposure to 20 ppt sea water. This study suggests that high temperature (29.5 °C) aggravates the damage of trivially low salinity (30 ppt) on the holobiont (the coral and its symbiotic dinoflagellates) in 10 d exposure. However, high temperature (29.5 °C) may have an antagonistic effect between temperature and low salinity (25 ppt) on metabolism of the holobiont. Based on the above results, we suggest that (1) the true mechanism of corals exposed to combined effects of low salinity and high temperature is complicated. This calls for more studies on different corals. Future studies should aim at investigating long-term low-level stress in order to simulate in situ conditions more accurately; (2) when corals exposed to extremely severe combined stressors for short-term or trivially severe stressors for relative long-term, the combined effects of two stressors (such as low salinity and high temperature) may be negative, otherwise, the effects may be additive.     

海洋酸化对珊瑚礁生态系统的影响研究进展

目前,大气CO2浓度的升高已导致海水pH值比工业革命前下降了约0.1,海水碳酸盐平衡体系随之变化,进而影响珊瑚礁生态系统的健康。近年来的研究表明海洋酸化导致造礁石珊瑚幼体补充和群落恢复更加困难,造礁石珊瑚和其它造礁生物(Reef-building organisms)钙化率降低甚至溶解,乃至影响珊瑚礁鱼类的生命活动。虽然海洋酸化对造礁石珊瑚光合作用的影响不显著,但珊瑚-虫黄藻共生体系会受到一定影响。建议选择典型海区进行长期系统监测,结合室内与原位模拟试验,从个体、种群、群落到系统不同层面,运用生理学和分子生物学技术,结合生态学研究手段,综合研究珊瑚的相应响应,以期深入认识海洋酸化对珊瑚礁生态系统健康(例如珊瑚白化)的影响及其效应。     

Red Sea corals under Artificial Light Pollution at Night (ALAN) undergo oxidative stress and photosynthetic impairment

Coral reefs represent the most diverse marine ecosystem on the planet, yet they are undergoing an unprecedented decline due to a combination of increasing global and local stressors. Despite the wealth of research investigating these stressors, Artificial Light Pollution at Night (ALAN) or “ecological light pollution” represents an emerging threat that has received little attention in the context of coral reefs, despite the potential of disrupting the chronobiology, physiology, behavior, and other biological processes of coral reef organisms. Scleractinian corals, the framework builders of coral reefs, depend on lunar illumination cues to synchronize their biological rhythms such as behavior, reproduction and physiology. While, light pollution (POL) may mask and lead de‐synchronization of these biological rhythms process. To reveal if ALAN impacts coral physiology, we have studied two coral species, Acropora eurystoma and Pocillopora damicornis, from the Gulf of Eilat/Aqaba, Red Sea, which is undergoing urban development that has led to severe POL at night. Our two experimental design data revealed that corals exposed to ALAN face an oxidative stress condition, show lower photosynthesis performances measured by electron transport rate (ETR), as well as changes in chlorophyll and algae density parameters. Testing different lights such as Blue LED and White LED spectrum showed more extreme impact in comparison to Yellow LEDs on coral physiology. The finding of this work sheds light on the emerging threat of POL and the impacts on the biology and ecology of Scleractinian corals, and will help to formulate specific management implementations to mitigate its potentially harmful impacts.     

The use of coral nubbins in coral reef ecotoxicology testing

While there is an urgent demand to establish reliable ecotoxicological assays for reef corals, there has not been yet an available source material that can supply the high number of colony replicates needed for reliable tests. In past experiments, the major obstacle to obtaining as many fragments as possible had been the damage inflicted to donor colonies by pruning. In this paper, we present the application of coral nubbins, a novel source material for coral ecotoxicology assays. Nubbins from the branching Red Sea coral Stylophora pistillata (n>450) were used for evaluating the impacts of water soluble fractions from a crude oil, an oil dispersant and dispersed oil. Coral nubbins (minute coral fragments in the size of one to several polyps) harvested from a single colony are genetically identical to each other, may be obtained in any quantity needed and whenever research activities demand their use. Several dozens of nubbins can be obtained from a single small branch in branching coral species, a procedure that has minimal impact on donor genotypes. Nubbins production is a low cost procedure and requires limited maintenance space. Results of short and long-term acute ecotoxicological tests are revealed and discussed here, indicating the advantageous use of nubbins as ubiquitous coral material for toxicology assays and physiological studies.     

Paradise lost: End‐of‐century warming and acidification under business‐as‐usual emissions have severe consequences for symbiotic corals

Despite recent efforts to curtail greenhouse gas emissions, current global emission trajectories are still following the business‐as‐usual representative concentration pathway (RCP) 8.5 emission pathway. The resulting ocean warming and acidification have transformative impacts on coral reef ecosystems, detrimentally affecting coral physiology and health, and these impacts are predicted to worsen in the near future. In this study, we kept fragments of the symbiotic corals Acropora intermedia (thermally sensitive) and Porites lobata (thermally tolerant) for 7 weeks under an orthogonal design of predicted end‐of‐century RCP8.5 conditions for temperature and pCO₂ (3.5°C and 570 ppm above present‐day, respectively) to unravel how temperature and acidification, individually or interactively, influence metabolic and physiological performance. Our results pinpoint thermal stress as the dominant driver of deteriorating health in both species because of its propensity to destabilize coral–dinoflagellate symbiosis (bleaching). Acidification had no influence on metabolism but had a significant negative effect on skeleton growth, particularly when photosynthesis was absent such as in bleached corals or under dark conditions. Total loss of photosynthesis after bleaching caused an exhaustion of protein and lipid stores and collapse of calcification that ultimately led to A. intermedia mortality. Despite complete loss of symbionts from its tissue, P. lobata maintained small amounts of photosynthesis and experienced a weaker decline in lipid and protein reserves that presumably contributed to higher survival of this species. Our results indicate that ocean warming and acidification under business‐as‐usual CO₂ emission scenarios will likely extirpate thermally sensitive coral species before the end of the century, while slowing the recovery of more thermally tolerant species from increasingly severe mass coral bleaching and mortality. This could ultimately lead to the gradual disappearance of tropical coral reefs globally, and a shift on surviving reefs to only the most resilient coral species.     

Effects of the multiple stressors high temperature and reduced salinity on the photosynthesis of the hermatypic coral Galaxea fascicularis

This study investigates the physiological responses in the hermatypic coral Galaxea fascicularis exposed to salinity stress (from 37 ppt to 15 ppt) for 12 h, combined effects of reduced salinity (from 37 ppt to 20 ppt) and two temperatures (26 °C and 32 °C) for 12 h and combined effects of reduced salinity (from 37 ppt to 25 ppt) and two temperatures (26 °C and 29.5 °C) for 10 d. The results demonstrate that the coral is tolerant to 12 h exposure to extremely low salinity (15 ppt). The study also shows that combined effects of temperature and low salinity aggravate the damage on the photosynthesis of the symbiotic dinoflagellates in 12 h exposure to 20 ppt sea water. This study suggests that high temperature (29.5 °C) aggravates the damage of trivially low salinity (30 ppt) on the holobiont (the coral and its symbiotic dinoflagellates) in 10 d exposure. However, high temperature (29.5 °C) may have an antagonistic effect between temperature and low salinity (25 ppt) on metabolism of the holobiont. Based on the above results, we suggest that (1) the true mechanism of corals exposed to combined effects of low salinity and high temperature is complicated. This calls for more studies on different corals. Future studies should aim at investigating long-term low-level stress in order to simulate in situ conditions more accurately; (2) when corals exposed to extremely severe combined stressors for short-term or trivially severe stressors for relative long-term, the combined effects of two stressors (such as low salinity and high temperature) may be negative, otherwise, the effects may be additive.     

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.     

Vulnerability of global coral reef habitat suitability to ocean warming,acidification and eutrophication

Coral reefs are threatened by global and local stressors. Yet, reefs appear to respond differently to different environmental stressors. Using a global dataset of coral reef occurrence as a proxy for the long‐term adaptation of corals to environmental conditions in combination with global environmental data, we show here how global (warming: sea surface temperature; acidification: aragonite saturation state, Ω_arag) and local (eutrophication: nitrate concentration, and phosphate concentration) stressors influence coral reef habitat suitability. We analyse the relative distance of coral communities to their regional environmental optima. In addition, we calculate the expected change of coral reef habitat suitability across the tropics in relation to an increase of 0.1°C in temperature, an increase of 0.02 μmol/L in nitrate, an increase of 0.01 μmol/L in phosphate and a decrease of 0.04 in Ω_arag. Our findings reveal that only 6% of the reefs worldwide will be unaffected by local and global stressors and can thus act as temporary refugia. Local stressors, driven by nutrient increase, will affect 22% of the reefs worldwide, whereas global stressors will affect 11% of these reefs. The remaining 61% of the reefs will be simultaneously affected by local and global stressors. Appropriate wastewater treatments can mitigate local eutrophication and could increase areas of temporary refugia to 28%, allowing us to ‘buy time’, while international agreements are found to abate global stressors.     

A corrosive concoction: The combined effects of ocean warming and acidification on the early growth of a stony coral are multiplicative

Survival of coral planulae, and the successful settlement and healthy growth of primary polyps are critical for the dispersal of scleractinian corals and hence the recovery of degraded coral reefs. It is therefore important to explore how the warmer and more acidic oceanic conditions predicted for the future could affect these processes. This study used controlled culture to investigate the effects of a 1 °C increase in temperature and a 0.2-0.25 unit decrease in pH on the settlement and survival of planulae and the growth of primary polyps in the Tropical Eastern Pacific coral Porites panamensis. We found that primary polyp growth was reduced only marginally by more acidic seawater but the combined effect of high temperature and lowered pH caused a significant reduction in growth of primary polyps by almost a third. Elevated temperature was found to significantly reduce the amount of zooxanthellae in primary polyps, and when combined with lowered pH resulted in a significant reduction in biomass of primary polyps. However, survival and settlement of planula larvae were unaffected by increased temperature, lowered acidity or the combination of both. These results indicate that in future scenarios of increased temperature and oceanic acidity coral planulae will be able to disperse and settle successfully but primary polyp growth may be hampered. The recovery of reefs may therefore be impeded by global change even if local stressors are curbed and sufficient sources of planulae are available.     

Century‐scale records of coral growth rates indicate that local stressors reduce coral thermal tolerance threshold

Coral bleaching, during which corals lose their symbiotic dinoflagellates, appears to be increasing in frequency and geographic extent, and is typically associated with abnormally high water temperatures and solar irradiance. A key question in coral reef ecology is whether local stressors reduce the coral thermal tolerance threshold, leading to increased bleaching incidence. Using tree‐ring techniques, we produced master chronologies of growth rates in the dominant reef builder, massive Montastraea faveolata corals, over the past 75–150 years from the Mesoamerican Reef. Our records indicate that the 1998 mass bleaching event was unprecedented in the past century, despite evidence that water temperatures and solar irradiance in the region were as high or higher mid‐century than in more recent decades. We tested the influence on coral extension rate from the interactive effects of human populations and thermal stress, calculated here with degree‐heating‐months (DHM). We find that when the effects of chronic local stressors, represented by human population, are taken into account, recent reductions in extension rate are better explained than when DHM is used as the sole predictor. Therefore, the occurrence of mass bleaching on the Mesoamerican reef in 1998 appears to stem from reduced thermal tolerance due to the synergistic impacts of chronic local stressors.     

Functional genomic analysis of corals from natural CO2‐seeps reveals core molecular responses involved in acclimatization to ocean acidification

Little is known about the potential for acclimatization or adaptation of corals to ocean acidification and even less about the molecular mechanisms underpinning these processes. Here, we examine global gene expression patterns in corals and their intracellular algal symbionts from two replicate population pairs in Papua New Guinea that have undergone long‐term acclimatization to natural variation in pCO₂. In the coral host, only 61 genes were differentially expressed in response to pCO₂ environment, but the pattern of change was highly consistent between replicate populations, likely reflecting the core expression homeostasis response to ocean acidification. Functional annotations highlight lipid metabolism and a change in the stress response capacity of corals as key parts of this process. Specifically, constitutive downregulation of molecular chaperones was observed, which may impact response to combined climate change‐related stressors. Elevated CO₂ has been hypothesized to benefit photosynthetic organisms but expression changes of in hospite Symbiodinium in response to acidification were greater and less consistent among reef populations. This population‐specific response suggests hosts may need to adapt not only to an acidified environment, but also to changes in their Symbiodinium populations that may not be consistent among environments, adding another challenging dimension to the physiological process of coping with climate change.     

Implications of coral harvest and transplantation on reefs in northwestern Dominica

In June, 2002, the government of Dominica requested assistance in evaluating the coral culture and transplantation activities being undertaken by Oceanographic Institute of Dominica (OID), a coral farm culturing both western Atlantic and Indo-Pacific corals for restoration and commercial sales. We assessed the culture facilities of OID, the condition of reefs, potential impacts of coral collection and benefits of coral transplantation. Coral reefs (9 reefs, 3-20 m depth) were characterized by 35 species of scleractinian corals and a live coral cover of 8-35%. Early colonizing, brooders such as Porites astreoides (14.8% of all corals), P. porites (14.8%), Meandrina meandrites (14.7%) and Agaricia agaricites (9.1%) were the most abundant corals, but colonies were mostly small (mean = 25 cm diameter). Montastraea annularis (complex) was the other dominant taxa (20.8% of all corals) and colonies were larger (mean = 70 cm). Corals (pooled species) were missing an average of 20% of their tissue, with a mean of 1.4% recent mortality. Coral diseases affected 6.4% of all colonies, with the highest prevalence at Cabrits West (11.0%), Douglas Bay (12.2%) and Coconut Outer reef (20.7%). White plague and yellow band disease were causing the greatest loss of tissue, especially among M. annularis (complex), with localized impacts from corallivores, overgrowth by macroalgae, storm damage and sedimentation. While the reefs appeared to be undergoing substantial decline, restoration efforts by OlD were unlikely to promote recovery. No Pacific species were identified at OID restoration sites, yet species chosen for transplantation with highest survival included short-lived brooders (Agaricia and Porites) that were abundant in restoration sites, as well as non-reef builders (Palythoa and Erythropodium) that monopolize substrates and overgrow corals. The species of highest value for restoration (massive broadcast spawners) showed low survivorship and unrestored populations of these species were most affected by biotic stressors and human impacts, all of which need to be addressed to enhance survival of outplants. Problems with culture practices at OID, such as high water temperature, adequate light levels and persistent overgrowth by macroalgae could be addressed through simple modifications. Nevertheless, coral disease and other stressors are of major concern to the most important reef builders, as these species are less amenable to restoration, collection could threaten their survival and losses require decades to centuries to replace.     

Coral reefs modify their seawater carbon chemistry – implications for impacts of ocean acidification

Reviews suggest that that the biogeochemical threshold for sustained coral reef growth will be reached during this century due to ocean acidification caused by increased uptake of atmospheric CO₂. Projections of ocean acidification, however, are based on air‐sea fluxes in the open ocean, and not for shallow‐water systems such as coral reefs. Like the open ocean, reef waters are subject to the chemical forcing of increasing atmospheric pCO₂. However, for reefs with long water residence times, we illustrate that benthic carbon fluxes can drive spatial variation in pH, pCO₂ and aragonite saturation state (Ω_a) that can mask the effects of ocean acidification in some downstream habitats. We use a carbon flux model for photosynthesis, respiration, calcification and dissolution coupled with Lagrangian transport to examine how key groups of calcifiers (zooxanthellate corals) and primary producers (macroalgae) on coral reefs contribute to changes in the seawater carbonate system as a function of water residence time. Analyses based on flume data showed that the carbon fluxes of corals and macroalgae drive Ω_ain opposing directions. Areas dominated by corals elevate pCO₂ and reduce Ω_a, thereby compounding ocean acidification effects in downstream habitats, whereas algal beds draw CO₂ down and elevate Ω_a, potentially offsetting ocean acidification impacts at the local scale. Simulations for two CO₂ scenarios (600 and 900 ppm CO₂) suggested that a potential shift from coral to algal abundance under ocean acidification can lead to improved conditions for calcification in downstream habitats, depending on reef size, water residence time and circulation patterns. Although the carbon fluxes of benthic reef communities cannot significantly counter changes in carbon chemistry at the scale of oceans, they provide a significant mechanism of buffering ocean acidification impacts at the scale of habitat to reef.     

Symbiodiniaceae probiotics for use in bleaching recovery

Coral reefs are currently under threat as a consequence of local and global stressors, in particular, mass coral bleaching induced by climate warming. In conjunction with global cuts to carbon emissions, active restoration interventions are being investigated as an additional option to buy time while these stressors are mitigated. One intervention with the potential to improve recovery during or postbleaching involves the addition of probiotic treatments, that is the addition of microorganisms that provide benefits to the host. Fragments of the branching coral, Acropora millepora, were experimentally exposed to a bleaching event coupled with the inoculation of Symbiodiniaceae probiotics (Durusdinium trenchii and Cladocopium goreaui) to determine if these probiotic treatments could ameliorate bleaching related stress and mortality. Fragments inoculated with C. goreaui and exposed to 32°C for 6 days exhibited significantly less mortality (9.1 ± 5%) compared to corals exposed to 32°C without probiotics (66.7 ± 8%) or with D. trenchii (41.7 ± 9%). Fragments in the C. goreaui probiotic treatment also bleached less and exhibited the highest photosynthetic efficiency compared to fragments inoculated with the D. trenchii at 32°C. Internal transcribed spacer‐2 amplicon sequencing did not detect the inoculated D. trenchii and C. goreaui cells within A. millepora tissues at the end of the experiment, suggesting the corals did not reestablish symbiosis but instead used inoculated cells as a nutritional supplement, although other factors such as shuffling conditions may have had an effect. This study highlights that nutritional supplementation can possibly aid coral resilience to temperature stress, though a far more detailed understanding of the factors that influence host regulation during symbiosis establishment is required.     

Improved sustainable maintenance for mid-water coral nursery by the application of an anti-fouling agent

The ‘gardening concept’ for reef restoration focuses on coral colonies farming in mid-water nurseries before their transplantation onto denuded reef areas. Nurseries situated in a nutrient-enriched environment significantly curtail nursery time, but extend labor, as nursery construction and farmed corals must be frequently cleaned from competing fouling organisms. Mass farming of corals calls, therefore, for efficient and cheap maintenance methodologies, which we here tested by employing Aqua-guard M250, an anti-fouling agent used in the fish farming industry. We found that this anti-fouling paint, while reducing fouling organisms on nursery components during the crucial phase of coral ramets' development from nubbins and small fragments sizes to colony sizes suitable for transplantation, is not toxic to maricultured coral fragments that staged more than 2 cm away from the paint. Applying small quantities of such antifouling paint to coral nurseries, while restricting its use to nursery components that are not in direct contact with farmed coral material, reduces fouling coverage and cleaning procedures by 90%.     

Evidence for multiple stressor interactions and effects on coral reefs

Concern is growing about the potential effects of interacting multiple stressors, especially as the global climate changes. We provide a comprehensive review of multiple stressor interactions in coral reef ecosystems, which are widely considered to be one of the most sensitive ecosystems to global change. First, we synthesized coral reef studies that examined interactions of two or more stressors, highlighting stressor interactions (where one stressor directly influences another) and potentially synergistic effects on response variables (where two stressors interact to produce an effect that is greater than purely additive). For stressor‐stressor interactions, we found 176 studies that examined at least 2 of the 13 stressors of interest. Applying network analysis to analyze relationships between stressors, we found that pathogens were exacerbated by more costressors than any other stressor, with ca. 78% of studies reporting an enhancing effect by another stressor. Sedimentation, storms, and water temperature directly affected the largest number of other stressors. Pathogens, nutrients, and crown‐of‐thorns starfish were the most‐influenced stressors. We found 187 studies that examined the effects of two or more stressors on a third dependent variable. The interaction of irradiance and temperature on corals has been the subject of more research (62 studies, 33% of the total) than any other combination of stressors, with many studies reporting a synergistic effect on coral symbiont photosynthetic performance (n = 19). Second, we performed a quantitative meta‐analysis of existing literature on this most‐studied interaction (irradiance and temperature). We found that the mean effect size of combined treatments was statistically indistinguishable from a purely additive interaction, although it should be noted that the sample size was relatively small (n = 26). Overall, although in aggregate a large body of literature examines stressor effects on coral reefs and coral organisms, considerable gaps remain for numerous stressor interactions and effects, and insufficient quantitative evidence exists to suggest that the prevailing type of stressor interaction is synergistic.