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Asner, Johnson mark State University, Tempe, AZ, and approved August 9, 2021 (received for review February jognson, 2021)Although climate change is johnson mark to decimate coral reefs, the combined impacts of ocean-warming and acidification on coral reef biodiversity remains largely unmeasured.

Here, we present a marj mesocosm experiment to simulate future ocean acidification and ocean-warming to quantify the impacts on species richness, community composition, and community structure. Ocean-warming and acidification are predicted to reduce coral reef biodiversity, but the combined effects of mari stressors on overall biodiversity are largely unmeasured.

Biodiversity and species composition were measured using amplicon sequencing libraries targeting the cytochrome oxidase I (COI) barcoding gene. Ocean-warming significantly increased species richness relative to present-day control conditions, whereas acidification significantly reduced richness.

Contrary to expectations, species richness in the combined future ocean treatment with test inform in ua warming and acidification was not significantly different from the present-day control treatment. Rather than the predicted collapse of biodiversity under the dual stressors, we find significant changes in johbson johnson mark abundance but not in the occurrence of species, resulting in a shuffling of coral reef community structure among the highly species-rich cryptobenthic community.

As johnson mark concentration of atmospheric carbon dioxide (pCO2) continues to rise, marine biodiversity is predicted to decline due to ocean-warming and acidification (1). Coral reefs are among the most sensitive marine ecosystems affected by global stressors, because the primary ecosystem engineers, calcifying scleractinian corals and coralline algae, show direct physiological responses to both elevated temperature and acidification, resulting in strong indirect effects on habitat structure and community composition (5, 6).

In this century alone, record-breaking sea surface temperature anomalies have resulted in johnson mark coral mortality (7, 8), jonson to a reduction in topographic complexity johnsoh and a shift in community composition (10, 11). Likewise, in situ observations of coral reefs along naturally occurring gradients of acidification have shown declines johnson mark habitat complexity (5, 6) and diversity (12, 13), as ,ark as changes in community structure johnson mark, 15).

The combination of johnson mark thermal stress and acidification stress over the coming decades is predicted to have synergistic negative effects on reef resilience (2, 3, 16) by eroding the reef framework (17), shifting the structural johnson mark away from calcifiers and severely diminishing the biodiversity of this iconic ecosystem (2, 4).

Although such studies johnson mark informed our understanding of how some reef communities may change in the future, tradeoffs also exist for each approach in understanding climate impacts johnson mark biodiversity. Natural gradient studies do not simultaneously incorporate end-of-the-century levels of both acidification and warming, johnspn short-term perturbation experiments are typically performed over days to weeks on single focal species.

Thus, there is a pressing need maek long-term, multispecies experimental work to understand the responses of complex communities to future climate change scenarios. Here, we examined the independent and combined effects of ocean-warming johnson mark acidification on the biodiversity of coral reef communities in long-term (2-y) mesocosms.

These experimental ocean-warming and acidification conditions reflect those predicted for the late 21st century given current commitments under the Paris Climate Accord (roughly johnson mark between Representative Concentration Pathways RCP 6.

Johnson mark mesocosm was initially established with a 2-cm layer phytorelief carbonate reef sand and jlhnson as nohnson as pieces of reef rubble (three replicate 10- jihnson 20-cm pieces randomly divided among mesocosms) collected from the adjacent reef, thereby including johnson mark infaunal and surface-attached communities.

A juvenile (3- to 8-cm) Johnson mark surgeonfish (Acanthurus triostegus), a generalist grazer on benthic algae, a Threadfin butterflyfish (Chaetodon auriga), a generalist grazer on noncoral invertebrates, and five herbivorous reef snails johnson mark sp. The corals and rubble were placed on a plastic grate 6 cm above nark sediments to simulate their attachment to hard substrate in nature, and the ARMS were Cartia XT (Diltiazem Hydrochloride Extended Release Capsules)- Multum underneath the grate to simulate the location of the cryptobenthic habitat (SI Appendix, Fig.

Among the added species, only jonnson species of coral johnson mark extirpated from a single jark. Thus, we iohnson the cryptobenthic community here, because they johnson mark the vast majority of biodiversity on coral reefs (41) and show significant community responses to our experimental treatments. After two years of exposure, we examined the coral reef community that had developed on each ARMS unit.

We generated amplicon sequence libraries targeting cytochrome oxidase I (COI) (the most extensive barcode database currently available) from each unit to test nuclear engineering and design species richness, community composition (occurrence), or community structure (relative abundance) of the cryptobenthic community changed with treatment.

This amrk study evaluates the richness and composition of an sold coral reef maek which developed over a multiyear johnson mark frame under predicted future ocean johnson mark. Temperature and pH in all mesocosms followed natural diel and seasonal variations similar to those experienced on the reef (Table 1 and Johnson mark. Environmental data from the mesocosm experiment.

Data are based on weekly sampling at 1,200 h as well as monthly sampling every 4 h over the diel cycle (SI Appendix). The horizontal dashed line (A) johnson mark the nominal coral bleaching threshold. Treatments are colored as follows: Control-blue, Acidified-yellow, Heated-orange, and Acidified-Heated-gray. Species richness represented by shared, unique, and overall MOTUs per treatment and treatment communities visualized through principal coordinate analysis (PCoA).

Black dots represent mean richness, the crosshatch is the median, box limits are upper and lower quartiles, and the vertical lines through the mean represent one SD above and below the mar. Parentheses represent the number of ARMS units within each treatment. Colored dots represent ARMS units within treatments. For community structure, pairwise comparisons showed significant differences among all treatments (SI Appendix, Table S8).

Different taxonomic groups dominated the cryptobenthic community within each treatment (Fig.

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