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180 results for “Ocean acidification”
The vulnerability of shellfish aquaculture to climate change and ocean acidification– a global assessment
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Adaptive phenotypic evolution of Skeletonema costatum to ocean acidification and warming with trade-offs from a multi-year outdoor experiment
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Can niche plasticity mediate species persistence under ocean acidification?
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Data from: Biogeography of ocean acidification: differential field performance of transplanted mussels to upwelling-driven variation in carbonate chemistry
<p>Ocean acidification (OA) represents a serious challenge to marine ecosystems. Laboratory studies addressing OA indicate broadly negative effects for marine organisms, particularly those relying on calcification processes. Growing evidence also suggests OA combined with other environmental stressors may be even more deleterious. Scaling these laboratory studies to ecological performance in the field, where environmental heterogeneity may mediate responses, is a critical next step toward understanding OA impacts on natural communities. We leveraged an upwelling-driven pH mosaic along the California Current System to deconstruct the relative influences of pH, ocean temperature, and food availability on seasonal growth, condition and shell thickness of the ecologically dominant intertidal mussel Mytilus californianus . In 2011 and 2012, ecological performance of adult mussels from local and commonly sourced populations was measured at 8 rocky intertidal sites between central Oregon and southern California. Sites coincided with a large-scale network of intertidal pH sensors, allowing comparisons among pH and other environmental stressors. Adult California mussel growth and size varied latitudinally among sites and inter-annually, and mean shell thickness index and shell weight growth were reduced with low pH. Surprisingly, shell length growth and the ratio of tissue to shell weight were enhanced, not diminished as expected, by low pH. In contrast, and as expected, shell weight growth and shell thickness were both diminished by low pH, consistent with the idea that OA exposure can compromise shell-dependent defenses against predators or wave forces. We also found that adult mussel shell weight growth and relative tissue mass were negatively associated with increased pH variability. Including local pH conditions with previously documented influences of ocean temperature, food availability, aerial exposure, and origin site enhanced the explanatory power of models describing observed performance differences. Responses of local mussel populations differed from those of a common source population suggesting mussel performance partially depended on genetic or persistent phenotypic differences. In light of prior research showing deleterious effects of low pH on larval mussels, our results suggest a life history transition leading to greater resilience in at least some performance metrics to ocean acidification by adult California mussels. Our data also demonstrate "hot" (more extreme) and "cold" (less extreme) spots in both mussel responses and environmental conditions, a pattern that may enable mitigation approaches in response to future changes in climate.</p>
Data from: Ocean warming and acidification alter the behavioural response to flow of the sea urchin Paracentrotus lividus
Ocean warming (OW) and acidification (OA) are intensively investigated as they pose major threats to marine organism. However, little effort is dedicated to another collateral climate change stressor, the increased frequency and intensity of storm events, here referred to as intensified hydrodynamics. A 2-month experiment was performed to identify how OW and OA (temperature: 21°C; pHT: 7.7, 7.4; control: 17°C-pHT7.9) affect the resistance to hydrodynamics in the sea urchin Paracentrotus lividus using an integrative approach that includes physiology, biomechanics and behaviour. Biomechanics was studied under both no-flow condition at the tube foot (TF) scale and flow condition at the individual scale. For the former, TF disk adhesive properties (attachment strength, tenacity) and TF stem mechanical properties (breaking force, extensibility, tensile strength, stiffness, toughness) were evaluated. For the latter, resistance to flow was addressed as the flow velocity at which individuals detached. Under near- and far-future OW and OA, individuals fully balanced their acid-base status, but skeletal growth was halved. TF adhesive properties were not affected by treatments. Compared to the control, mechanical properties were in general improved under pHT7.7 while in the extreme treatment (21°C-pHT7.4) breaking force was diminished. Three behavioural strategies were implemented by sea urchins and acted together to cope with flow: improving TF attachment, streamlining and escaping. Behavioural responses varied according to treatment and flow velocity. For instance, individuals at 21°C-pHT7.4 increased the density of attached TF at slow flows or, controlled TF detachment at fast flow to compensate for weakened TF mechanical properties. They also showed an absence of streamlining favouring an escaping behaviour as they ventured in a riskier faster movement at slow flows. At faster flows, the effects of OW and OA were detrimental causing earlier dislodgment. These plastic behaviours reflect a potential scope for acclimation in the field, where this species already experiences diel temperature and pH fluctuations.
Assessing the impact of static and fluctuating ocean acidification on the behavior of Amphiprion percula
<p>Attached is the complete raw data from Vaughan and Dixson 2021 ‘Assessing the impact of static and fluctuating ocean acidification on the behavior of <em>Amphiprion percula</em>’. </p> <p>Data collected from the behavioral lateralization trials has been inputted into the file ‘Vaughan_2020_Lateralization_Raw’. Column A indicate the CO<sub>2</sub> treatment group, where “SPD” = Static Present Day, “SFD” = Static Future Day, “FPD” = Fluctuating Present Day, and “FFD” = Fluctuating Future Day. Each individual fish used from each treatment group (n=30) is displayed in Column B. Column C shows the binary results, in order, of each fish’s turns in the T-maze, and was scored as 0 (right turn) or 1 (left turn) for a total of 10 turns. The total number of turns to the right and left are provided in Column D-E. The relative lateralization (<em>L</em><sub>R</sub>) of each fish was calculated {L<sub>R</sub> = [(Turn to the right – Turn to the left)/(Turn to the right + Turn to the left)] ∗ 100} in Column F. Absolute lateralization (<em>L</em><sub>A</sub>) is provided in Column G.</p> <p>Chemosensory response data has been inputted into the file ‘Vaughan_2020_Chemosensory_Raw’. Column A and B display treatment group and fish ID (n=20) as outlined above. The cue used in trial of either Tang (nonpredator) or Cod (predator) is provided in Column C, and the control in Column D. Numbers in these are used solely for the purpose of data analysis. The side of the cue in the flume is provided in Column E, and corresponds with the cue labelled in Column C. Buckets containing either the cue or control were placed above the flume and color coded as “BS” (blue side) and “RS” (red side), as the person scoring the trials was blinded. This also helped account for the switch (from one side of the flume to the other) that occurs halfway through each trial. Columns F-G represent results from the first 2min recording period, and Columns H-I represent results from the second 2min recording period. The total tallies from each fish are provided in Column J; the totals from each side are calculated in Columns K-L, and then sorted by either cue or control in Columns M-N. Proportions and percentages in cue and control are calculated and provided in Columns O-P and Q-R, respectively. </p> <p>Carbonate chemistry data is compiled and provided in the ‘Vaughan_2020_Carbonate_Chemistry’. Measurements were taken each week (Column A) of each treatment group (as stated above, Column B). Column C reflects the time recordings were taken in the fluctuating treatments to hit the high, mid and low CO<sub>2</sub> points at “6:30”, “12:30” and “18:30”. Measurements of static treatment groups were taken at randomly selected times to get the reflection of the carbonate chemistry of these treatments, but for the purpose of clarity in this document they are listed as “Static”. Measurements were taken from a subset of tanks each that rotated each week (Column D). Our target pH<sub>NBS</sub> values (i.e. what was programmed into the APEX System) are listed in Column E. Columns F-H displayed pH<sub>NBS</sub> (taken with APEX probes), temperature °C (taken with a portable Mettler Toledo probe) and salinity (taken with a refractometer). Water samples were analyzed spectrophotometrically to provide pH<sub>T</sub> and dissolved inorganic carbon, with values provided in Columns I-J. Using the program CO2SYS, total alkalinity and <em>p</em>CO<sub>2</sub> were calculated, with values provided in Columns K-L.</p>
Data from: Adult exposure to ocean acidification is maladaptive for larvae of the Sydney rock oyster Saccostrea glomerata in the presence of multiple stressors
Parental effects passed from adults to their offspring have been identified as a source of rapid acclimation that may allow marine populations to persist as our surface oceans continue to decrease in pH. Little is known, however, whether parental effects are beneficial for offspring in the presence of multiple stressors. We exposed adults of the oyster Saccostrea glomerata to elevated CO2 and examined the impacts of elevated CO2 (control = 392; 856 µatm) combined with elevated temperature (control = 24; 28°C), reduced salinity (control = 35; 25) and reduced food concentration (control = full; half diet) on their larvae. Adult exposure to elevated CO2 had a positive impact on larvae reared at elevated CO2 as a sole stressor, which were 8% larger and developed faster at elevated CO2 compared with larvae from adults exposed to ambient CO2. These larvae, however, had significantly reduced survival in all multistressor treatments. This was particularly evident for larvae reared at elevated CO2 combined with elevated temperature or reduced food concentration, with no larvae surviving in some treatment combinations. Larvae from CO2-exposed adults had a higher standard metabolic rate. Our results provide evidence that parental exposure to ocean acidification may be maladaptive when larvae experience multiple stressors.
Data from: Ocean acidification during pre-fertilization chemical communication affects sperm success
Ocean acidification (OA) poses a major threat to marine organisms, particularly during reproduction when externally shed gametes are vulnerable to changes in seawater pH. Accordingly, several studies on OA have focused on how changes in seawater pH influence sperm behaviour and/or rates of in vitro fertilization. By contrast, few studies have examined how pH influences pre-fertilization gamete interactions, which are crucial during natural spawning events in most externally fertilizing taxa. One mechanism of gamete interaction that forms an important component of fertilization in most taxa is communication between sperm and egg-derived chemicals. These chemical signals, along with the physiological responses in sperm they elicit, are likely to be highly sensitive to changes in seawater chemistry. In this study, we experimentally tested this possibility using the blue mussel, Mytilus galloprovincialis, a species in which females have been shown to use egg-derived chemicals to promote the success of sperm from genetically compatible males. We conducted trials in which sperm were allowed to swim in gradients of egg-derived chemicals under different seawater CO2 (and therefore pH) treatments. We found that sperm had elevated fertilization rates after swimming in the presence of egg-derived chemicals in low pH (pH 7.6) compared to ambient (pH 8.0) seawater. This observed effect could have important implications for the reproductive fitness of external fertilizers, where gamete compatibility plays a critical role in modulating reproduction in many species. For example, elevated sperm fertilization rates might disrupt the eggs' capacity to avoid fertilizations by genetically incompatible sperm. Our findings highlight the need to understand how OA affects the multiple stages of sperm-egg interactions, and to develop approaches that disentangle the implications of OA for female, male and population fitness.
Data from: Ocean acidification alters the response of intertidal snails to a key sea star predator
Organism-level effects of ocean acidification (OA) are well recognized. Less understood are OA's consequences for ecological species interactions. Here, we examine a behaviourally mediated predator–prey interaction within the rocky intertidal zone of the temperate eastern Pacific Ocean, using it as a model system to explore OA's capacity to impair invertebrate anti-predator behaviours more broadly. Our system involves the iconic sea star predator, Pisaster ochraceus, that elicits flee responses in numerous gastropod prey. We examine, in particular, the capacity for OA-associated reductions in pH to alter flight behaviours of the black turban snail, Tegula funebralis, an often-abundant and well-studied grazer in the system. We assess interactions between these species at 16 discrete levels of pH, quantifying the full functional response of Tegula under present and near-future OA conditions. Results demonstrate the disruption of snail anti-predator behaviours at low pH, with decreases in the time individuals spend in refuge locations. We also show that fluctuations in pH, including those typical of rock pools inhabited by snails, do not materially change outcomes, implying little capacity for episodically benign pH conditions to aid behavioural recovery. Together, these findings suggest a strong potential for OA to induce cascading community-level shifts within this long-studied ecosystem.
Data from: Impacts of ocean acidification on sea urchin growth across the juvenile to mature adult life-stage transition is mitigated by warming
Understanding how growth trajectories of calcifying invertebrates are affected by changing climate requires acclimation experiments that follow development across life history transitions. In a long-term acclimation study, the effects of increased acidification and temperature on survival and growth of the tropical sea urchin Tripneustes gratilla from the early juvenile (5 mm test diameter- TD) through the developmental transition to the mature adult (60 mm TD) were investigated. Juveniles were reared in a combination of three temperature and three pH/pCO2 treatments, including treatments commensurate with global change projections. Elevated temperature and pCO2/pH both affected growth, but there was no interaction between these factors. The urchins grew more slowly at pH 7.6, but not at pH 7.8. Slow growth may be influenced by the inability to compensate coelomic fluid acid-base balance at pH 7.6. Growth was faster at +3 and +6 °C compared to that in ambient temperature. Acidification and warming had strong and interactive effects on reproductive potential. Warming increased the gonad index, but acidification decreased it. At pH 7.6 there were virtually no gonads in any urchins regardless of temperature. The T. gratilla were larger at maturity under combined near-future warming and acidification scenarios (+3 °C/pH 7.8). Although the juveniles grew and survived in near-future warming and acidification conditions, chronic exposure to these stressors from an early stage altered allocation to somatic and gonad growth. In the absence of phenotypic adjustment, the interactive effects of warming and acidification on the benthic life phases of sea urchins may compromise reproductive fitness and population maintenance as global climatic change unfolds.
Data from: Low recruitment due to altered settlement substrata as primary constraint for coral communities under ocean acidification
The future of coral reefs under increasing CO2 depends on their capacity to recover from disturbances. To predict the recovery potential of coral communities that are fully acclimatized to elevated CO2, we compared the relative success of coral recruitment and later life stages at two volcanic CO2 seeps and adjacent control sites in Papua New Guinea. Our field experiments showed that the effects of ocean acidification (OA) on coral recruitment rates were up to an order of magnitude greater than the effects on the survival and growth of established corals. Settlement rates, recruit and juvenile densities were best predicted by the presence of crustose coralline algae, as opposed to the direct effects of seawater CO2. Offspring from high CO2 acclimatized parents had similarly impaired settlement rates as offspring from control parents. For most coral taxa, field data showed no evidence of cumulative and compounding detrimental effects of high CO2 on successive life stages, and three taxa showed improved adult performance at high CO2 that compensated for their low recruitment rates. Our data suggest that severely declining capacity for reefs to recover, due to altered settlement substrata and reduced coral recruitment, is likely to become a dominant mechanism of how OA will alter coral reefs.
Data from: Bioturbation determines the response of benthic ammonia oxidising microorganisms to ocean acidification
Ocean acidification (OA), caused by the dissolution of increasing concentrations of atmospheric carbon dioxide (CO2) in seawater, is projected to cause significant changes to marine ecology and biogeochemistry. Potential impacts on the microbially driven cycling of nitrogen are of particular concern. Specifically, under seawater pH levels approximating future OA scenarios, rates of ammonia oxidation (the rate-limiting first step of the nitrification pathway) have been shown to dramatically decrease in seawater, but not in underlying sediments. However, no prior study has considered the interactive effects of microbial ammonia oxidation and macrofaunal bioturbation activity, which can enhance nitrogen transformation rates. Using experimental mesocosms, we investigated the responses to OA of ammonia oxidizing microorganisms inhabiting surface sediments and sediments within burrow walls of the mud shrimp Upogebia deltaura. Seawater was acidified to one of four target pH values (pHT 7.90, 7.70, 7.35 and 6.80) in comparison with a control (pHT 8.10). At pHT 8.10, ammonia oxidation rates in burrow wall sediments were, on average, fivefold greater than in surface sediments. However, at all acidified pH values (pH ≤ 7.90), ammonia oxidation rates in burrow sediments were significantly inhibited (by 79–97%; p < 0.01), whereas rates in surface sediments were unaffected. Both bacterial and archaeal abundances increased significantly as pHT declined; by contrast, relative abundances of bacterial and archaeal ammonia oxidation (amoA) genes did not vary. This research suggests that OA could cause substantial reductions in total benthic ammonia oxidation rates in coastal bioturbated sediments, leading to corresponding changes in coupled nitrogen cycling between the benthic and pelagic realms.
Data from: Evolutionary change during experimental ocean acidification
Rising atmospheric carbon dioxide (CO2) conditions are driving unprecedented changes in seawater chemistry, resulting in reduced pH and carbonate ion concentrations in the Earth's oceans. This ocean acidification has negative but variable impacts on individual performance in many marine species. However, little is known about the adaptive capacity of species to respond to an acidified ocean, and as a result, predictions regarding future ecosystem responses remain incomplete. Here we demonstrate that ocean acidification generates striking patterns of genome-wide selection in purple sea urchins (Strongylocentrotus purpuratus) cultured under different CO2 levels. We examined genetic change at 19,493 loci in larvae from seven adult populations cultured under realistic future CO2 levels. Although larval development and morphology showed little response to elevated CO2, we found substantial allelic change in 40 functional classes of proteins involving hundreds of loci. Pronounced genetic changes, including excess amino acid replacements, were detected in all populations and occurred in genes for biomineralization, lipid metabolism and ion homeostasis, gene classes that build skeletons and interact in pH regulation. Such genetic change represents a neglected and important impact of ocean acidification that may influence populations that show few outward signs of response to acidification. Our results demonstrate the capacity for rapid evolution in the face of ocean acidification and show that standing genetic variation could be a reservoir of resilience to climate change in this coastal upwelling ecosystem. However, effective response to strong natural selection demands large population sizes and may be limited in species impacted by other environmental stressors.
Data from: Parasitic infection: a buffer against ocean acidification?
Recently, there has been a concerted research effort by marine scientists to quantify the sensitivity of marine organisms to ocean acidification (OA). Empirical data generated by this research have been used to predict changes to marine ecosystem health, biodiversity and productivity that will be caused by continued acidification. These studies have also found that the effects of OA on marine organisms can be significantly modified by additional abiotic stressors (e.g. temperature or oxygen) and biotic interactions (e.g. competition or predation). To date, however, the effects of parasitic infection on the sensitivity of marine organisms to OA have been largely ignored. We show that parasitic infection significantly altered the response of a marine gastropod to simulated OA conditions by reducing the mortality of infected individuals relative to uninfected conspecifics. Without the inclusion of infection data, our analysis would not have detected the significant effect of pH on host mortality. These results strongly suggest that parasitic infection may be an important confounding factor in OA research and must be taken into consideration when assessing the response of marine species to OA.
Data from: Novel reverse radioisotope labelling experiment reveals carbon assimilation of marine calcifiers under ocean acidification conditions
<p><span>1. Ocean acidification by anthropogenic carbon dioxide emissions is projected to depress metabolic and physiological activity in marine calcifiers. To evaluate the sensitivity of marine organisms against ocean acidification, the assimilation of nutrients into carbonate shells and soft tissues must be examined.</span></p> <p><span>2. We designed a novel experimental protocol, reverse radioisotope labelling, to trace partitioning of nutrients within a single bivalve species under ocean acidification conditions. Injecting CO<sub>2</sub> gas, free from radiocarbon, can provide a large contrast between carbon dissolved in the water and the one assimilated from atmosphere. By culturing modern aquifer organisms in acidified seawater, we were able to determine differences in the relative contributions of the end members, dissolved inorganic carbon (DIC) in seawater and metabolic CO<sub>2</sub>, to shell carbonate and soft tissues. </span></p> <p><span>3. Under all pCO<sub>2</sub> conditions (463, 653, 872, 1137, and 1337 μatm), radiocarbon (Δ<sup>14</sup>C) values of the bivalve (<i>Scapharca broughtonii</i>) shell were significantly correlated with seawater DIC values; therefore, shell carbonate was derived principally from seawater DIC. The Δ<sup>14</sup>C results together with stable carbon isotope (δ<sup>13</sup>C) data suggest that in <i>S. broughtonii</i> shell δ<sup>13</sup>C may reflect the kinetics of isotopic equilibration as well as end-member contributions; thus, care must be taken when analyzing end-member contributions by a previous method using δ<sup>13</sup>C. The insensitivity of <i>S. broughtonii</i> to perturbations in pCO<sub>2</sub> up to at least 1337 µatm indicates that this species can withstand ocean acidification.</span></p> <p><span>4. Usage of radioisotope to dope for tracer experiments requires strict rules to conduct any operations. Yet, reverse radioisotope labeling proposing in this study has a large advantage and is a powerful tool to understanding physiology of aquifer organisms that can be applicable to various organisms and culture experiments, such as temperature, salinity, and acidification experiments, to improve understanding of the proportions of nutrients taken in by marine organisms under changing environments.</span></p>
Data from: Differential responses to ocean acidification between populations of Balanophyllia elegans corals from high and low upwelling environments
Ocean acidification (OA), the global decrease in surface water pH from absorption of anthropogenic CO2, may put many marine taxa at risk. However, populations that experience extreme localized conditions, and are adapted to these conditions predicted in the global ocean in 2100, may be more tolerant to future OA. By identifying locally adapted populations, researchers can examine the mechanisms used to cope with decreasing pH. One oceanographic process that influences pH, is wind driven upwelling. Here we compare two Californian populations of the coral Balanophyllia elegans from distinct upwelling regimes, and test their physiological and transcriptomic responses to experimental seawater acidification. We measured respiration rates, protein and lipid content, and gene expression in corals from both populations exposed to pH levels of 7.8 and 7.4 for 29 days. Corals from the population that experiences lower pH due to high upwelling, maintained the same respiration rate throughout the exposure. In contrast, corals from the low upwelling site had reduced respiration rates, protein content, and lipid-class content at low pH exposure, suggesting they have depleted their energy reserves. Using RNA-Seq, we found that corals from the high upwelling site upregulated genes involved in calcium ion binding and ion transport, most likely related to pH homeostasis and calcification. In contrast, corals from the low upwelling site downregulated stress response genes at low pH exposure. Divergent population responses to low pH observed in B. elegans highlight the importance of multi-population studies for predicting a species' response to future OA.
Data from: Can larvae of a marine fish adapt to ocean acidification? Evaluating the evolutionary potential of California Grunion (Leuresthes tenuis)
Ocean acidification can reduce the growth and survival of marine species during their larval stages. However, if populations have the genetic capacity to adapt and increase their tolerance of low pH and high pCO2 levels, this may offset the harmful effects of ocean acidification. By combining controlled breeding experiments with laboratory manipulations of seawater chemistry, we evaluated genetic variation in tolerance of ocean acidification conditions for a nearshore marine fish, the California Grunion (Leuresthes tenuis). Our results indicated that acidification conditions increased overall mortality rates of grunion larvae, but did not have a significant effect on growth. Groups of larvae varied widely with respect to mortality and growth rates in both ambient and acidified conditions. We demonstrate that the potential to evolve in response to ocean acidification is best described by considering additive genetic variation in fitness-related traits under both ambient and acidified conditions, and by evaluating the genetic correlation between traits expressed in these environments. We used a multivariate animal model to estimate additive genetic (co)variance in larval growth and mortality rates under both ambient and acidified conditions (low pH/high pCO2). Our results suggest appreciable genetic variation in larval mortality rates (h2Ambient = 0.120; h2Acidified = 0.183; rG = 0.460), but less genetic variation in growth (h2Ambient = 0.092; h2Acidified = 0.101; rG = 0.135). Maternal effects on larval mortality rates accounted for 26-36% of the variation in phenotypes, but maternal effects accounted for only 8% of the variation in growth. Collectively, our estimates of genetic variation and covariation suggest that populations of California Grunion have the capacity to adapt relatively quickly to long-term changes in ocean chemistry.
Data from: A quantitative genetic approach to assess the evolutionary potential of a coastal marine fish to ocean acidification
Assessing the potential of marine organisms to adapt genetically to increasing oceanic CO2 levels requires proxies such as heritability of fitness-related traits under ocean acidification (OA). We applied a quantitative genetic method to derive the first heritability estimate of survival under elevated CO2 conditions in a metazoan. Specifically, we reared offspring, selected from a wild coastal fish population (Atlantic silverside, Menidia menidia), at high CO2 conditions (~2,300 μatm) from fertilization to 15 days post hatch, which significantly reduced survival compared to controls. Perished and surviving offspring were quantitatively sampled and genotyped along with their parents, using 8 polymorphic microsatellite loci, to reconstruct a parent-offspring pedigree and estimate variance components. Genetically related individuals were phenotypically more similar (i.e., survived similarly long at elevated CO2 conditions) than unrelated individuals, which translated into a significantly non-zero heritability (0.20 ± 0.07). The contribution of maternal effects was surprisingly small (0.05 ± 0.04) and non-significant. Survival among replicates was positively correlated with genetic diversity, particularly with observed heterozygosity. We conclude that early life survival of M. menidia under high CO2 levels has a significant additive genetic component that could elicit an evolutionary response to OA, depending on the strength and direction of future selection.
Data for Responses of Marine Trophic Levels to the Combined Effects of Ocean Acidification and Warming
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Fouling communities from the South African west coast are vulnerable to cooling and ocean acidification
<p>Data was generated during a laboratory study considering the responses of fouling communities to altered acidificationa nd cooling.. All methods are detailed within Matikinca & Robinson (2024) Fouling communities from the South African west coast are vulnerable to cooling and ocean acidifcation. Marine Biodiversity 54: 27 DOI: 10.1007/s12526-024-01420-0</p> <p>Data layout: Community data is provided for plates subjected to different treatments. Additionally, species richness and diverstity are provided. </p>
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