Limited intraspecific variation in drought resistance along a pronounced tropical rainfall gradient.

Assessing within-species variation in response to drought is crucial for predicting species' responses to climate change and informing restoration and conservation efforts, yet experimental data are lacking for the vast majority of tropical tree species. We assessed intraspecific variation in response to water availability across a strong rainfall gradient for 16 tropical tree species using reciprocal transplant and common garden field experiments, along with measurements of gene flow and key functional traits linked to drought resistance. Although drought resistance varies widely among species in these forests, we found little evidence for within-species variation in drought resistance. For the majority of functional traits measured, we detected no significant intraspecific variation. The few traits that did vary significantly between drier and wetter origins of the same species all showed relationships opposite to expectations based on drought stress. Furthermore, seedlings of the same species originating from drier and wetter sites performed equally well under drought conditions in the common garden experiment and at the driest transplant site. However, contrary to expectation, wetter-origin seedlings survived better than drier-origin seedlings under wetter conditions in both the reciprocal transplant and common garden experiment, potentially due to lower insect herbivory. Our study provides the most comprehensive picture to date of intraspecific variation in tropical tree species' responses to water availability. Our findings suggest that while drought plays an important role in shaping species composition across moist tropical forests, its influence on within-species variation is limited.

Fitness surfaces and local thermal adaptation in Drosophila along a latitudinal gradient.

Local adaptation is commonly cited to explain species distribution, but how fitness varies along continuous geographical gradients is not well understood. Here, we combine thermal biology and life-history theory to demonstrate that Drosophila populations along a 2500 km latitudinal cline are adapted to local conditions. We measured how heat tolerance and viability rate across eight populations varied with temperature in the laboratory and then simulated their expected cumulative Darwinian fitness employing high-resolution temperature data from their eight collection sites. Simulations indicate a trade-off between annual survival and cumulative viability, as both mortality and the recruitment of new flies are predicted to increase in warmer regions. Importantly, populations are locally adapted and exhibit the optimal combination of both traits to maximize fitness where they live. In conclusion, our method is able to reconstruct fitness surfaces employing empirical life-history estimates and reconstructs peaks representing locally adapted populations, allowing us to study geographic adaptation in silico.

Soil microbial identity explains home-field advantage for litter decomposition.

Unraveling the mechanisms of home-field advantage (HFA) is essential to gain a complete understanding of litter decomposition processes. However, knowledge of the relationships between HFA effects and microbial communities is lacking. To examine HFA effects on litter decomposition, we identified the microbial communities and conducted a reciprocal transplant experiment, including all possible combinations of soil and litter, between sites at two elevations in cool-temperate forests. Soil origin, rather than HFA, was an important factor in controlling litter decomposition processes. Microbiome-wide association analyses identified litter fungi and bacteria specific to the source soil, which completely differed at a low taxonomic level between litter types. The relative abundance of these microbes specific to source soil was positively correlated with litter mass loss. The results indicated that the unique relationships between plant litter and soil microbes through plant-soil linkages drive litter decomposition processes. In the short term, soil disturbances resulting from land-use changes have the potential to disrupt the effect of soil origin and hinder the advancement of litter decomposition. These findings contribute to an understanding of HFA mechanisms and the impacts of land-use change on decomposition processes in forest ecosystems.

Using warming tolerances to predict understory plant responses to climate change.

Climate change is pushing species towards and potentially beyond their critical thermal limits. The extent to which species can cope with temperatures exceeding their critical thermal limits is still uncertain. To better assess species' responses to warming, we compute the warming tolerance (ΔTniche ) as a thermal vulnerability index, using species' upper thermal limits (the temperature at the warm limit of their distribution range) minus the local habitat temperature actually experienced at a given location. This metric is useful to predict how much more warming species can tolerate before negative impacts are expected to occur. Here we set up a cross-continental transplant experiment involving five regions distributed along a latitudinal gradient across Europe (43° N-61° N). Transplant sites were located in dense and open forests stands, and at forest edges and in interiors. We estimated the warming tolerance for 12 understory plant species common in European temperate forests. During 3 years, we examined the effects of the warming tolerance of each species across all transplanted locations on local plant performance, in terms of survival, height, ground cover, flowering probabilities and flower number. We found that the warming tolerance (ΔTniche ) of the 12 studied understory species was significantly different across Europe and varied by up to 8°C. In general, ΔTniche were smaller (less positive) towards the forest edge and in open stands. Plant performance (growth and reproduction) increased with increasing ΔTniche across all 12 species. Our study demonstrated that ΔTniche of understory plant species varied with macroclimatic differences among regions across Europe, as well as in response to forest microclimates, albeit to a lesser extent. Our findings support the hypothesis that plant performance across species decreases in terms of growth and reproduction as local temperature conditions reach or exceed the warm limit of the focal species.

Genetic response of a perennial grass to warm and wet environments interacts and is associated with trait means as well as plasticity.

The potential for rapid evolution is an important mechanism allowing species to adapt to changing climatic conditions. Although such potential has been largely studied in various short-lived organisms, to what extent we can observe similar patterns in long-lived plant species, which often dominate natural systems, is largely unexplored. We explored the potential for rapid evolution in Festuca rubra, a long-lived grass with extensive clonal growth dominating in alpine grasslands. We used a field sowing experiment simulating expected climate change in our model region. Specifically, we exposed seeds from five independent seed sources to novel climatic conditions by shifting them along a natural climatic grid and explored the genetic profiles of established seedlings after 3 years. Data on genetic profiles of plants selected under different novel conditions indicate that different climate shifts select significantly different pools of genotypes from common seed pools. Increasing soil moisture was more important than increasing temperature or the interaction of the two climatic factors in selecting pressure. This can indicate negative genetic interaction in response to the combined effects or that the effects of different climates are interactive rather than additive. The selected alleles were found in genomic regions, likely affecting the function of specific genes or their expression. Many of these were also linked to morphological traits (mainly to trait plasticity), suggesting these changes may have a consequence on plant performance. Overall, these data indicate that even long-lived plant species may experience strong selection by climate, and their populations thus have the potential to rapidly adapt to these novel conditions.

Field and Laboratory Evidence That Chlorpyrifos Exposure Reduced the Population Density of a Freshwater Snail by Increasing Juvenile Mortality.

Pesticides have been frequently detected in global freshwater ecosystems, but attempts to document changes in population dynamics of organisms upon exposure to pesticides, establish a causal relationship between exposure and population effects, and identify the key toxic events within individuals under natural field conditions remain rare. Here, we used a field survey, a reciprocal cross-transplant experiment, and a laboratory toxicity experiment to build a compelling case that exposure to the insecticide chlorpyrifos was responsible for differences in snail (Bellamya aeruginosa) densities in eastern (ELL) and western basins of Liangzi Lake in China. Our field survey and reciprocal cross-transplant experiment revealed significant differences in snail densities, juvenile percentage, survival, and relative telomere length (RTL) in the two basins. The insecticide chlorpyrifos detected in snail tissues was negatively correlated with snail densities, the percentage of juvenile snails, and RTL and had an extremely high risk quotient in ELL. In the laboratory experiment, tissue concentrations of chlorpyrifos detected in ELL were associated with reduced RTL and increased juvenile mortality in B. aeruginosa. These results support the hypothesis that chlorpyrifos exposure in ELL reduced the density of snails by reducing juvenile survival and, consequently, recruitment to the adult population.

Local adaptation: Causal agents of selection and adaptive trait divergence.

Divergent selection across the landscape can favor the evolution of local adaptation in populations experiencing contrasting conditions. Local adaptation is widely observed in a diversity of taxa, yet we have a surprisingly limited understanding of the mechanisms that give rise to it. For instance, few have experimentally confirmed the biotic and abiotic variables that promote local adaptation, and fewer yet have identified the phenotypic targets of selection that mediate local adaptation. Here, we highlight critical gaps in our understanding of the process of local adaptation and discuss insights emerging from in-depth investigations of the agents of selection that drive local adaptation, the phenotypes they target, and the genetic basis of these phenotypes. We review historical and contemporary methods for assessing local adaptation, explore whether local adaptation manifests differently across life history, and evaluate constraints on local adaptation.

Ecological lags govern the pace and outcome of plant community responses to 21st-century climate change.

Forecasting the trajectories of species assemblages in response to ongoing climate change requires quantifying the time lags in the demographic and ecological processes through which climate impacts species' abundances. Since experimental climate manipulations are typically abrupt, the observed species responses may not match their responses to gradual climate change. We addressed this problem by transplanting alpine grassland turfs to lower elevations, recording species' demographic responses to climate and competition, and using these data to parameterise community dynamics models forced by scenarios of gradual climate change. We found that shifts in community structure following an abrupt climate manipulation were not simply accelerated versions of shifts expected under gradual warming, as the former missed the transient rise of species benefiting from moderate warming. Time lags in demography and species interactions controlled the pace and trajectory of changing species' abundances under simulated 21st-century climate change, and thereby prevented immediate diversity loss.

Local adaptation in a marine foundation species: Implications for resilience to future global change.

Environmental change is multidimensional, with local anthropogenic stressors and global climate change interacting to differentially impact populations throughout a species' geographic range. Within species, the spatial distribution of phenotypic variation and its causes (i.e., local adaptation or plasticity) will determine species' adaptive capacity to respond to a changing environment. However, comparatively less is known about the spatial scale of adaptive differentiation among populations and how patterns of local adaptation might drive vulnerability to global change stressors. To test whether fine-scale (2-12 km) mosaics of environmental stress can cause adaptive differentiation in a marine foundation species, eelgrass (Zostera marina), we conducted a three-way reciprocal transplant experiment spanning the length of Tomales Bay, CA. Our results revealed strong home-site advantage in growth and survival for all three populations. In subsequent common garden experiments and feeding assays, we showed that countergradients in temperature, light availability, and grazing pressure from an introduced herbivore contribute to differential performance among populations consistent with local adaptation. Our findings highlight how local-scale mosaics in environmental stressors can increase phenotypic variation among neighboring populations, potentially increasing species resilience to future global change. More specifically, we identified a range-center eelgrass population that is pre-adapted to extremely warm temperatures similar to those experienced by low-latitude range-edge populations of eelgrass, demonstrating how reservoirs of heat-tolerant phenotypes may already exist throughout a species range. Future work on predicting species resilience to global change should incorporate potential buffering effects of local-scale population differentiation and promote a phenotypic management approach to species conservation.

Drivers of local extinction risk in alpine plants under warming climate.

The scarcity of local plant extinctions following recent climate change has been explained by demographic inertia and lags in the displacement of resident species by novel species, generating an 'extinction debt'. We established a transplant experiment to disentangle the contribution of these processes to the local extinction risk of four alpine plants in the Swiss Alps. Projected population growth (λ) derived from integral projection models was reduced by 0.07/°C of warming on average, whereas novel species additionally decreased λ by 0.15 across warming levels. Effects of novel species on predicted extinction time were greatest at warming < 2 °C for two species. Projected population declines under both warming and with novel species were primarily driven by increased mortality. Our results suggest that extinction debt can be explained by a combination of demographic inertia and lags in novel species establishment, with the latter being particularly important for some species under low levels of warming.

Artificial habitats induce plasticity in colonies of the marine bryozoan Schizoporella errata.

Modular organization provides flexibility for colonial animals to deal with variable and unpredictable environmental conditions since each module has specific tasks within the colony, such as feeding, defending or reproducing. Depending on the selecting pressures, sessile organisms may phenotypically adjust the morphology of each module or modify their density, increasing individual fitness. Here we used the marine bryozoan Schizoporella errata (Cheilostomata, Schizoporellidae) to test how the divergent conditions between two artificial habitats, the location inside a marina (IM) and the external wall of the breakwater (BW), affect colony size and the density of the distinct modules. The density of avicularia and ovicells, modules related to defense and reproduction, respectively, did not differ between habitats. However, colonies growing in the turbulent waters of BW were, in general, larger and had higher density of feeding autozooids than those at IM. Reciprocal transplants of bryozoan clones indicated that trait variation is genotype-dependent but varies according to the environmental conditions at the assigned location. The occurrence of larger colonies with more zooids in BW is probably linked to the easier feeding opportunity offered by the small diffusive boundary layer around the colony at this location. Since in colonial polymorphic organisms each module (zooid) performs a specific function, the phenotypic response is not uniform across colonies, affecting only those modules that are susceptible to variations in the main selective pressures. Understanding the importance of colony-level plasticity is relevant to predict how modularity will contribute to organisms to deal with human-induced environmental changes in coastal habitats.

Adaptation and correlated fitness responses over two time scales in Drosophila suzukii populations evolving in different environments.

The process of local adaptation involves differential changes in fitness over time across different environments. Although experimental evolution studies have extensively tested for patterns of local adaptation at a single time point, there is relatively little research that examines fitness more than once during the time course of adaptation. We allowed replicate populations of the fruit pest Drosophila suzukii to evolve in one of eight different fruit media. After five generations, populations with the highest initial levels of maladaptation had mostly gone extinct, whereas experimental populations evolving on cherry, strawberry and cranberry media had survived. We measured the fitness of each surviving population in each of the three fruit media after five and after 26 generations of evolution. After five generations, adaptation to each medium was associated with increased fitness in the two other media. This was also true after 26 generations, except when populations that evolved on cranberry medium developed on cherry medium. These results suggest that, in the theoretical framework of a fitness landscape, the fitness optima of cherry and cranberry media are the furthest apart. Our results show that studying how fitness changes across several environments and across multiple generations provides insights into the dynamics of local adaptation that would not be evident if fitness were analysed at a single point in time. By allowing a qualitative mapping of an experimental fitness landscape, our approach will improve our understanding of the ecological factors that drive the evolution of local adaptation in D. suzukii.

Temperature effects on forest understorey plants in hedgerows: a combined warming and transplant experiment.

BACKGROUND AND AIMS: Hedgerows have been shown to improve forest connectivity, leading to an increased probability of species tracking the shifting bioclimatic envelopes. However, it is still unknown how species in hedgerows respond to temperature changes, and whether effects differ compared with those in nearby forests. We aimed to elucidate how ongoing changes in the climate system will affect the efficiency of hedgerows in supporting forest plant persistence and migration in agricultural landscapes. METHODS: Here we report results from the first warming experiment in hedgerows. We combined reciprocal transplantation of plants along an 860-km latitudinal transect with experimental warming to assess the effects of temperature on vegetative growth and reproduction of two common forest herbs (Anemone nemorosa and Geum urbanum) in hedgerows versus forests. KEY RESULTS: Both species grew taller and produced more biomass in forests than in hedgerows, most likely due to higher competition with ruderals and graminoids in hedgerows. Adult plant performance of both species generally benefitted from experimental warming, despite lower survival of A. nemorosa in heated plots. Transplantation affected the species differently: A. nemorosa plants grew taller, produced more biomass and showed higher survival when transplanted at their home site, indicating local adaptation, while individuals of G. urbanum showed greater height, biomass, reproductive output and survival when transplanted northwards, likely owing to the higher light availability associated with increasing photoperiod during the growing season. CONCLUSIONS: These findings demonstrate that some forest herbs can show phenotypic plasticity to warming temperatures, potentially increasing their ability to benefit from hedgerows as ecological corridors. Our study thus provides novel insights into the impacts of climate change on understorey plant community dynamics in hedgerows, and how rising temperature can influence the efficiency of these corridors to assist forest species' persistence and colonization within and beyond their current distribution range.

Transcriptomic plasticity of mesophotic corals among natural populations and transplants of Montastraea cavernosa in the Gulf of Mexico and Belize.

While physiological responses to low-light environments have been studied among corals on mesophotic coral ecosystems worldwide (MCEs; 30-150 m), the mechanisms behind acclimatization and adaptation to depth are not well understood for most coral species. Transcriptomic approaches based on RNA sequencing are useful tools for quantifying gene expression plasticity, particularly in slow-growing species such as scleractinian corals, and for identifying potential functional differences among conspecifics. A tag-based RNA-Seq (Tag-Seq) pipeline was applied to quantify transcriptional variation in natural populations of the scleractinian coral Montastraea cavernosa from mesophotic and shallower environments across five sites in Belize and the Gulf of Mexico: Carrie Bow Cay, West and East Flower Garden Banks, Pulley Ridge, and Dry Tortugas. Regional site location was a stronger driver of gene expression patterns than depth. However, mesophotic corals among all sites shared similar regulation of metabolic and cell growth functional pathways that may represent common physiological responses to environmental conditions at depth. Additionally, in a transplant experiment at West and East Flower Garden Banks, colonies transplanted from mesophotic to shallower habitats diverged from the control mesophotic group over time, indicating depth-regulated plasticity of gene expression. When the shallower depth zone experienced a bleaching event, bleaching severity did not differ significantly between transplants and shallow controls, but gene expression patterns indicated variable regulation of stress responses among depth treatments. Coupled observational and experimental studies of gene expression among mesophotic and shallower M. cavernosa provide insights into the ability of this depth-generalist coral species to persist under varying environmental conditions.

Strong stabilizing selection on timing of germination in a Mediterranean population of Arabidopsis thaliana.

PREMISE: Timing of germination can strongly influence plant fitness by affecting seedling survival and by having cascading effects on later life-history traits. In seasonal environments, the period favorable for seedling establishment and growth is limited, and timing of germination is likely to be under stabilizing selection because of conflicting selection through survival and fecundity. Moreover, optimal germination time may vary among genotypes because of inherent differences in later life-history traits. METHODS: To examine how germination time affects survival, fecundity, and the relative fitness of two genotypes differing in time to first flower, we conducted a field experiment in an Italian population of the winter annual Arabidopsis thaliana, in which seedling establishment occurs mainly in November. We transplanted seedlings of the local genotype and of a Swedish genotype monthly from August to December and monitored survival and fecundity. RESULTS: Only seedlings transplanted in November and December survived until reproduction, and fitness of the November cohort was 35 times higher than that of the December cohort, indicating strong stabilizing selection on timing of germination. There was no evidence of conflicting selection: seedling survival, adult survival, and fecundity were all highest in the November cohort. Moreover, the relative fitness of the two genotypes did not differ significantly between cohorts. CONCLUSIONS: The very narrow window of opportunity for seedling establishment was related to rapid seasonal changes in soil moisture and temperature, suggesting that rate of seasonal change is an important aspect to consider for understanding spatiotemporal variation in selection on phenological traits.

Contrasting environmental factors drive local adaptation at opposite ends of an environmental gradient in the yellow monkeyflower (Mimulus guttatus).

PREMISE: Identifying the environmental factors responsible for natural selection across different habitats is crucial for understanding the process of local adaptation in plants. Despite its importance, few studies have successfully isolated the environmental factors driving local adaptation in nature. In this study, we evaluated the agents of selection responsible for local adaptation of the monkeyflower Mimulus guttatus to California's coastal and inland habitats. METHODS: We implemented a manipulative reciprocal transplant experiment at coastal and inland sites, where we excluded aboveground stressors in an effort to elucidate their role in the evolution of local adaptation. RESULTS: Excluding aboveground stressors, most likely a combination of salt spray and herbivory, completely rescued inland annual plant fitness when transplanted to coastal habitat. The exclosures in inland habitat provided a benefit to the performance of coastal perennial plants. However, the exclosures are unlikely to provide much fitness benefit to the coastal plants at the inland site because of their general inability to flower in time to escape from the summer drought. CONCLUSIONS: Our study demonstrates that a distinct set of selective agents (aboveground vs. belowground) are responsible for local adaptation at opposite ends of an environmental gradient.

Ecological niche differences between two polyploid cytotypes of Saxifraga rosacea.

PREMISE: Different cytotypes of a species may differ in their morphology, phenology, physiology, and their tolerance of extreme environments. We studied the ecological niches of two subspecies of Saxifraga rosacea with different ploidy levels: the hexaploid Central European endemic subspecies sponhemica and the more widely distributed octoploid subspecies rosacea. METHODS: For both cytotypes, we recorded local environmental conditions and mean plant trait values in populations across their areas of distribution, analyzed their distributions by niche modeling, studied their performance at two transplant sites with contrasting conditions, and experimentally tested their cold resistance. RESULTS: Mean annual temperature was higher in hexaploid than in octoploid populations and experiments indicated that frost tolerance of the hexaploid is lower than that of the octoploid. Reproduction of octoploids from Central Europe was higher than that of hexaploids at a transplant site in subarctic Iceland, whereas the opposite was true in temperate Luxembourg, indicating adaptation of the octoploids to colder conditions. Temperature variables were also most important in niche models predicting the distribution of the two cytotypes. Genetic differences in survival among populations were larger for the octoploids than for the hexaploids in both field gardens, suggesting that greater genetic variability may contribute to the octoploid's larger distributional range. CONCLUSIONS: Our results support the hypotheses that different cytotypes may have different niches leading to spatial segregation, and that higher ploidy levels can result in a broader ecological niche and greater tolerance of more extreme conditions.

Tradeoffs between phenotypic plasticity and local adaptation influence the ecophysiology of the moss, Sphagnum magellanicum.

Bryophytes are a diverse plant group and are functionally different from vascular plants. Yet, their peculiarities are rarely considered in the theoretical frameworks for plants. Currently, we lack information about the magnitude and the importance of intraspecific variability in the ecophysiology of bryophytes and how these might translate to local adaptation-a prerequisite for adaptive evolution. Capitalizing on two ecologically distinct (hummock and hollow) phenotypes of Sphagnum magellanicum, we explored the magnitude and pattern of intraspecific variability in this species and asked whether the environmental-mediated changes in shoot and physiological traits are due to phenotypic plasticity or local adaptation. Size, pigmentation, and habitat type that distinguished the phenotypes in the field did not influence the trait responses under a transplant and factorial experiment. In addition, the magnitude and pattern of trait variability (e.g., branch, stem and capitulum mass) changed with the treatments, which suggest that trait responses were due largely to phenotypic plasticity. The trait responses also suggest that the ecophysiological needs for mosses to grow in clumps, where they maintain a uniform growth may have an overriding effect over the potential for a fixed adaptive response to environmental heterogeneity, which would constrain local adaptation. We conclude that extending the trait-based framework to mosses or making comparisons between mosses and vascular plants under any theoretical framework would only be meaningful to the extent that growth form and dispersal strategies are considered.

Transcriptome analysis reveals plasticity in gene regulation due to environmental cues in Primula sikkimensis, a high altitude plant species.

BACKGROUND: Studying plasticity in gene expression in natural systems is crucial, for predicting and managing the effects of climate change on plant species. To understand the contribution of gene expression level variations to abiotic stress compensation in a Himalaya plant (Primula sikkimensis), we carried out a transplant experiment within (Ambient), and beyond (Below Ambient and Above Ambient) the altitudinal range limit of species. We sequenced nine transcriptomes (three each from each altitudinal range condition) using Illumina sequencing technology. We compared the fitness variation of transplants among three transplant conditions. RESULTS: A large number of significantly differentially expressed genes (DEGs) between below ambient versus ambient (109) and above ambient versus ambient (85) were identified. Transcripts involved in plant growth and development were mostly up-regulated in below ambient conditions. Transcripts involved in signalling, defence, and membrane transport were mostly up-regulated in above ambient condition. Pathway analysis revealed that most of the genes involved in metabolic processes, secondary metabolism, and flavonoid biosynthesis were differentially expressed in below ambient conditions, whereas most of the genes involved in photosynthesis and plant hormone signalling were differentially expressed in above ambient conditions. In addition, we observed higher reproductive fitness in transplant individuals at below ambient condition compared to above ambient conditions; contrary to what we expect from the cold adaptive P. sikkimensis plants. CONCLUSIONS: We reveal P. sikkimensis's capacity for rapid adaptation to climate change through transcriptome variation, which may facilitate the phenotypic plasticity observed in morphological and life history traits. The genes and pathways identified provide a genetic resource for understanding the temperature stress (both the hot and cold stress) tolerance mechanism of P. sikkimensis in their natural environment.

Species-specific calcification response of Caribbean corals after 2-year transplantation to a low aragonite saturation submarine spring.

Coral calcification is expected to decline as atmospheric carbon dioxide concentration increases. We assessed the potential of Porites astreoides, Siderastrea siderea and Porites porites to survive and calcify under acidified conditions in a 2-year field transplant experiment around low pH, low aragonite saturation (Ωarag) submarine springs. Slow-growing S. siderea had the highest post-transplantation survival and showed increases in concentrations of Symbiodiniaceae, chlorophyll a and protein at the low Ωarag site. Nubbins of P. astreoides had 20% lower survival and higher chlorophyll a concentration at the low Ωarag site. Only 33% of P. porites nubbins survived at low Ωarag and their linear extension and calcification rates were reduced. The density of skeletons deposited after transplantation at the low Ωarag spring was 15-30% lower for all species. These results suggest that corals with slow calcification rates and high Symbiodiniaceae, chlorophyll a and protein concentrations may be less susceptible to ocean acidification, albeit with reduced skeletal density. We postulate that corals in the springs are responding to greater energy demands for overcoming larger differences in carbonate chemistry between the calcifying medium and the external environment. The differential mortality, growth rates and physiological changes may impact future coral species assemblages and the reef framework robustness.