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Ecological interactions among hosts are critical to consider when predicting disease dynamics. Most theory predicts that intraguild predation (IGP) and cannibalism negatively impact parasite populations, but this is based primarily on assumptions of simple or single-host life cycles. Here we investigate the effects of cannibalism in a size-structured host population on two digenean trematodes that have complex, multihost life cycles. A high incidence of cannibalism among paratenic hosts produced higher parasite infection loads and abundance, whereas cannibalism among obligate hosts reduced parasite abundances. We attributed this difference to trophic transmission aggregating parasites in larger, potentially fitter hosts and also to transmission among paratenic hosts via cannibalism. Moreover, we found evidence of indirect competitive interactions between parasites that can also increase infections at small scales. Our results show there are multiple mechanisms through which high cannibalism environments can benefit parasites that use paratenic hosts and trophic transfer to complete their life cycles.
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Canibalismo , Interações Hospedeiro-Parasita , Animais , Trematódeos/fisiologia , Cadeia Alimentar , Doenças dos Peixes/parasitologiaRESUMO
Warming has broad and often nonlinear impacts on organismal physiology and traits, allowing it to impact species interactions like predation through a variety of pathways that may be difficult to predict. Predictions are commonly based on short-term experiments and models, and these studies often yield conflicting results depending on the environmental context, spatiotemporal scale, and the predator and prey species considered. Thus, the accuracy of predicted changes in interaction strength, and their importance to the broader ecosystems they take place in, remain unclear. Here, we attempted to link one such set of predictions generated using theory, modeling, and controlled experiments to patterns in the natural abundance of prey across a broad thermal gradient. To do so, we first predicted how warming would impact a stage-structured predator-prey interaction in riverine rock pools between Pantala spp. dragonfly nymph predators and Aedes atropalpus mosquito larval prey. We then described temperature variation across a set of hundreds of riverine rock pools (n = 775) and leveraged this natural gradient to look for evidence for or against our model's predictions. Our model's predictions suggested that warming should weaken predator control of mosquito larval prey by accelerating their development and shrinking the window of time during which aquatic dragonfly nymphs could consume them. This was consistent with data collected in rock pool ecosystems, where the negative effects of dragonfly nymph predators on mosquito larval abundance were weaker in warmer pools. Our findings provide additional evidence to substantiate our model-derived predictions while emphasizing the importance of assessing similar predictions using natural gradients of temperature whenever possible.
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Aedes , Odonatos , Animais , Ecossistema , Odonatos/fisiologia , Larva/fisiologia , Comportamento Predatório/fisiologia , Cadeia AlimentarRESUMO
To predict the impacts of environmental change on species, we must first understand the factors that limit the present-day ranges of species. Most anuran amphibians cannot survive at elevated salinities, which may drive their distribution in coastal locations. Previous research showed that coastal Hyla cinerea are locally adapted to brackish habitats in North Carolina, USA. Although Hyla squirella and Hyla chrysoscelis both inhabit coastal wetlands nearby, they have not been observed in saline habitats. We take advantage of naturally occurring microgeographic variation in coastal wetland occupancy exhibited by these congeneric tree frog species to explore how salt exposure affects oviposition site choice, hatching success, early tadpole survival, plasma osmolality and tadpole body condition across coastal and inland locations. We observed higher survival among coastal H. cinerea tadpoles than inland H. cinerea, which corroborates previous findings. But contrary to expectations, coastal H. cinerea had lower survival than H. squirella and H. chrysoscelis, indicating that all three species may be able to persist in saline wetlands. We also observed differences in tadpole plasma osmolality across species, locations and salinities, but these differences were not associated with survival rates in salt water. Instead, coastal occupancy may be affected by stage-specific processes like higher probability of total clutch loss as shown by inland H. chrysoscelis or maladaptive egg deposition patterns as shown by inland H. squirella. Although we expected salt water to be the primary filter driving species distributions along a coastal salinity gradient, it is likely that the factors dictating anuran ranges along the coast involve stage-, species- and location-specific processes that are mediated by ecological processes and life history traits.
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Predicting the combined effects of predators on shared prey has long been a focus of community ecology, yet quantitative predictions often fail. Failure to account for nonlinearity is one reason for this. Moreover, prey depletion in multiple predator effects (MPE) studies generates biased predictions in applications of common experimental and quantitative frameworks. Here, we explore additional sources of bias stemming from nonlinearities in prey predation risk. We show that in order to avoid bias, predictions about the combined effects of independent predators must account for nonlinear size-dependent risk for prey as well as changes in prey risk driven by nonlinear predator functional responses and depletion. Historical failure to account for biases introduced by well-known nonlinear processes that affect predation risk suggest that we may need to reevaluate the general conclusions that have been drawn about the ubiquity of emergent MPEs over the past three decades.
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Ecologia , Comportamento Predatório , Animais , Comportamento Predatório/fisiologiaRESUMO
AbstractHybridization often occurs at the parapatric range interface between closely related species, but fitness outcomes vary: hybrid offspring exhibit diverse rates of viability and reproduction compared with their parental species. The mobile hybrid zone between two chickadee congeners (Poecile atricapillus × Poecile carolinensis) has been well studied behaviorally and genetically, but the viability of hybrids and the underlying mechanisms contributing to hybrid fitness have remained unclear. To better characterize the fitness costs of hybridization in this system, we analyzed 21 years of data from four sites, including more than 1,400 breeding attempts by the two species, to show that rates of hatching success changed substantially as the zone of hybridization moved across the landscape. Admixture-associated declines in hatching success correlated with reduced proportions of heterogametic (female) offspring, as predicted by Haldane's rule. Our data support an underlying mechanism implicating genetic admixture of the homogametic (male) parent as the primary determinant of offspring sex ratio, via incompatibilities on the hemizygous Z chromosome. Our long-term study is the first to directly measure changes in fitness costs as a vertebrate hybrid zone moves, and it shows that changes in these costs are a way to track the distribution of a hybrid zone across the landscape.
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Razão de Masculinidade , Aves Canoras , Animais , Feminino , Hibridização Genética , Masculino , Reprodução , Cromossomos Sexuais , Aves Canoras/genéticaRESUMO
The dear enemy hypothesis predicts that territorial individuals will be less aggressive toward known neighbors than to strangers. This hypothesis has been well studied and there is a wealth of data demonstrating its prevalence in some taxa. However, a quantitative synthesis is needed to test the generality of the phenomenon, identify key mechanisms driving the behavior, and guide future research. In this study, we conduct a meta-analysis and, we test the importance of the location of intrusion, the type of experiment conducted (field, laboratory, or neutral arenas), and the sex and breeding status of territory holders, on the occurrence of dear enemy behavior. We also test how various ecological and life history traits, such as territory type, stimulus of intrusion, and taxonomic group, affect the magnitude of dear enemy behavior. We find that this phenomenon is common and that taxonomic class and breeding status are correlated with the expression of dear enemy behaviors. Further, we found that the way authors measure aggression influences the likelihood of identifying dear enemy responses, and thus we discuss potential pitfalls of dear enemy studies. Considering this conclusion, we discuss future lines of inquiry that could more directly examine the mechanisms of the dear enemy phenomenon.
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Agressão , Territorialidade , HumanosRESUMO
Salinization is a global phenomenon affecting ecosystems and forcing freshwater organisms to deal with increasing levels of ionic stress. However, our understanding of mechanisms that permit salt tolerance in amphibians is limited. This study investigates mechanisms of salt tolerance in locally adapted, coastal populations of a treefrog, Hyla cinerea. Using a common garden experiment, we (i) determine the extent that environment (i.e., embryonic and larval saltwater exposure) or genotype (i.e., coastal vs. inland) affects developmental benchmarks and transcriptome expression, and (ii) identify genes that may underpin differences in saltwater tolerance. Differences in gene expression, survival, and plasma osmolality were most strongly associated with genotype. Population genetic analyses on expressed genes also delineated coastal and inland groups based on genetic similarity. Coastal populations differentially expressed osmoregulatory genes including ion transporters (atp1b1, atp6V1g2, slc26a), cellular adhesion components (cdh26, cldn1, gjb3, ocln), and cytoskeletal components (odc1-a, tgm3). Several of these genes are the same genes expressed by euryhaline fish after exposure to freshwater, which is a novel finding for North American amphibians and suggests that these genes may be associated with local salinity adaptation. Coastal populations also highly expressed glycerol-3-phosphate dehydrogenase 1 (gpd1), which indicates they use glycerol as a compatible osmolyte to reduce water loss - another mechanism of saltwater tolerance previously unknown in frogs. These data signify that Hyla cinerea inhabiting coastal, brackish wetlands have evolved a salt-tolerant ecotype, and highlights novel candidate pathways that can lead to salt tolerance in freshwater organisms facing habitat salinization.
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Ecossistema , Tolerância ao Sal , Aclimatação , Adaptação Fisiológica/genética , Animais , Anuros/genética , Salinidade , Tolerância ao Sal/genéticaRESUMO
Endocrine disrupting compounds (EDCs) are ubiquitous environmental pollutants that alter endocrine system function, induce birth defects, and a myriad of other negative health outcomes. Although the mechanism of toxicity of many EDCs have been studied in detail, little work has focused on understanding the mechanisms through which pregnant dams and fetuses protect themselves from EDCs, or if those protective mechanisms are sexually dimorphic in fetuses. In this study, we examined proteomic alterations in the livers of mouse dams and their male and female fetuses induced by vinclozolin, a model antiandrogenic EDC. Dam livers upregulated nine phase I and phase II detoxification pathways and pathway analysis revealed that more pathways are significantly enriched in dam livers than in fetal livers. Phase I and II detoxification proteins are also involved in steroid and steroid hormone biosynthesis and vinclozolin likely alters steroid levels in both the dam and the fetus. The response of the fetal liver proteome to vinclozolin exposure is sexually dimorphic. Female fetal livers upregulated proteins in xenobiotic metabolism pathways, whereas male fetal livers upregulated proteins in oxidative phosphorylation pathways. These results suggest that female fetuses increase protective mechanisms, whereas male fetuses increase ATP production and several disease pathways that are indicative of oxidative damage. Females fetuses upregulate proteins and protective pathways that were similar to the dams whereas males did not. If this sexually dimorphic pattern is typical, then males might generally be more sensitive to EDCs.
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Antagonistas de Androgênios/toxicidade , Disruptores Endócrinos/toxicidade , Fígado/efeitos dos fármacos , Oxazóis/toxicidade , Proteoma , Trifosfato de Adenosina/metabolismo , Antagonistas de Androgênios/metabolismo , Animais , Disruptores Endócrinos/metabolismo , Feminino , Fígado/embriologia , Fígado/metabolismo , Masculino , Exposição Materna , Desintoxicação Metabólica Fase I , Desintoxicação Metabólica Fase II , Camundongos , Oxazóis/metabolismo , Fosforilação Oxidativa , Gravidez , Proteômica , Caracteres Sexuais , Fatores SexuaisRESUMO
Predation is a pervasive force that structures food webs and directly influences ecosystem functioning. The relative body sizes of predators and prey may be an important determinant of interaction strengths. However, studies quantifying the combined influence of intra- and interspecific variation in predator-prey body size ratios are lacking.We use a comparative functional response approach to examine interaction strengths between three size classes of invasive bluegill and largemouth bass toward three scaled size classes of their tilapia prey. We then quantify the influence of intra- and interspecific predator-prey body mass ratios on the scaling of attack rates and handling times.Type II functional responses were displayed by both predators across all predator and prey size classes. Largemouth bass consumed more than bluegill at small and intermediate predator size classes, while large predators of both species were more similar. Small prey were most vulnerable overall; however, differential attack rates among prey were emergent across predator sizes. For both bluegill and largemouth bass, small predators exhibited higher attack rates toward small and intermediate prey sizes, while larger predators exhibited greater attack rates toward large prey. Conversely, handling times increased with prey size, with small bluegill exhibiting particularly low feeding rates toward medium-large prey types. Attack rates for both predators peaked unimodally at intermediate predator-prey body mass ratios, while handling times generally shortened across increasing body mass ratios.We thus demonstrate effects of body size ratios on predator-prey interaction strengths between key fish species, with attack rates and handling times dependent on the relative sizes of predator-prey participants.Considerations for intra- and interspecific body size ratio effects are critical for predicting the strengths of interactions within ecosystems and may drive differential ecological impacts among invasive species as size ratios shift.
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Sea level rise is mixing formerly isolated freshwater communities with saltwater communities. The structure of these new aquatic communities is jointly controlled by pre- and post-colonization processes. Similarly, since salinity is a strong abiotic determinant of post-colonization survival in coastal systems, changes in salinity will likely impact community composition. In this study, we examine how a strong abiotic gradient affects the diversity and structure of bacterial and zooplankton communities and associated ecosystem functions (decomposition and carbon mineralization). We ran a six week dispersal experiment using mesocosm ponds with four distinct salinity profiles (0, 5, 9, and 13 psu). We find that salinity is the primary driver of both bacterial and zooplankton community composition. We find evidence that as bacterial richness increases so does the amount of decomposition. A phenomenological model suggests carbon mineralization may decrease at mid-salinities; this warrants future work into possible mechanisms for this apparent loss of function. Understanding how salinization changes community structure and ecosystem function may be paramount for managing and conserving coastal plain ecosystems where salinity is increasing due to sea level rise, saltwater intrusion, storm surges, and drought.
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Environmental change and habitat fragmentation will affect population densities for many species. For those species that have locally adapted to persist in changed or stressful habitats, it is uncertain how density dependence will affect adaptive responses. Anurans (frogs and toads) are typically freshwater organisms, but some coastal populations of green treefrogs (Hyla cinerea) have adapted to brackish, coastal wetlands. Tadpoles from coastal populations metamorphose sooner and demonstrate faster growth rates than inland populations when reared solitarily. Although saltwater exposure has adaptively reduced the duration of the larval period for coastal populations, increases in densities during larval development typically increase time to metamorphosis and reduce rates of growth and survival. We test how combined stressors of density and salinity affect larval development between salt-adapted ("coastal") and nonsalt-adapted ("inland") populations by measuring various developmental and metamorphic phenotypes. We found that increased tadpole density strongly affected coastal and inland tadpole populations similarly. In high-density treatments, both coastal and inland populations had reduced growth rates, greater exponential decay of growth, a smaller size at metamorphosis, took longer to reach metamorphosis, and had lower survivorship at metamorphosis. Salinity only exaggerated the effects of density on the time to reach metamorphosis and exponential decay of growth. Location of origin affected length at metamorphosis, with coastal tadpoles metamorphosing slightly longer than inland tadpoles across densities and salinities. These findings confirm that density has a strong and central influence on larval development even across divergent populations and habitat types and may mitigate the expression (and therefore detection) of locally adapted phenotypes.
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A multicellular host and its microbial communities are recognized as a metaorganism-a composite unit of evolution. Microbial communities have a variety of positive and negative effects on the host life history, ecology, and evolution. This study used high-throughput amplicon sequencing to characterize the complete skin and gut microbial communities, including both bacteria and fungi, of a terrestrial salamander, Plethodon glutinosus (Family Plethodontidae). We assessed salamander populations, representing nine mitochondrial haplotypes ('clades'), for differences in microbial assemblages across 13 geographic locations in the Southeastern United States. We hypothesized that microbial assemblages were structured by both host factors and geographic distance. We found a strong correlation between all microbial assemblages at close geographic distances, whereas, as spatial distance increases, the patterns became increasingly discriminate. Network analyses revealed that gut-bacterial communities have the highest degree of connectedness across geographic space. Host salamander clade was explanatory of skin-bacterial and gut-fungal assemblages but not gut-bacterial assemblages, unless the latter were analyzed within a phylogenetic context. We also inferred the function of gut-fungal assemblages to understand how an understudied component of the gut microbiome may influence salamander life history. We concluded that dispersal limitation may in part describe patterns in microbial assemblages across space and also that the salamander host may select for skin and gut communities that are maintained over time in closely related salamander populations.
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Fenômenos Fisiológicos Bacterianos , Fungos/fisiologia , Trato Gastrointestinal/microbiologia , Microbiota , Pele/microbiologia , Urodelos/microbiologia , Distribuição Animal , Animais , Bactérias/isolamento & purificação , Fungos/isolamento & purificação , Microbioma Gastrointestinal , Micobioma , Sudeste dos Estados Unidos , Análise Espacial , TennesseeRESUMO
Organisms often respond to environmental change via phenotypic plasticity, in which an individual modulates its phenotype according to the environment. Highly variable or changing environments can exceed physiological limits and generate maladapted plastic phenotypes, which is termed nonadaptive plasticity. In some cases, selection may reduce the negative or disruptive impacts of environmental stress and produce locally adapted populations. Salt is an increasingly prevalent contaminant of freshwater systems and can induce nonadaptive plastic phenotypes for freshwater organisms like amphibians. Hyla cinerea is a frog species with populations inhabiting brackish, coastal habitats, so we use this species to test whether coastal populations are locally adapted to tolerate saltwater by determining how salt exposure during the embryonic and larval stages alters mortality and plastic developmental and metamorphic phenotypes of coastal and inland populations. Coastal frogs have higher survival, faster growth rates, and metamorphose sooner than inland frogs across salinities. Coastal frogs also metamorphose smaller (likely a consequence of earlier metamorphosis) yet maintain constant size, while higher salinities reduce metamorphic size for inland frogs. Coastal frogs evolved to minimize nonadaptive and disruptive impacts of saltwater during larval development and accelerate the larval period to reduce time spent in a stressful environment.
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Adaptação Fisiológica , Anuros/fisiologia , Pressão Osmótica , Tolerância ao Sal , Animais , Anuros/genética , Feminino , Água Doce , Variação Genética , Larva/genética , Larva/fisiologia , Masculino , Metamorfose Biológica , North Carolina , Fenótipo , Salinidade , Estresse Fisiológico , Áreas AlagadasRESUMO
Amphibians host a community of microbes on their skin that helps resist infectious disease via the dual influence of anti-pathogenic microbial species and emergent community dynamics. Many frogs rely on freshwater habitats, but salinization is rapidly increasing saltwater concentrations in wetlands around the globe, increasing the likelihood that frogs will come into contact with salt-contaminated habitats. Currently, we know little about how increased salt exposure will affect the symbiotic relationship between the skin microbes and frog hosts. To better understand how salt exposure in a natural context affects the frog skin microbiome, we use Hyla cinerea, a North American treefrog species that can inhabit brackish wetlands, to explore three questions. First, we determine the extent that microbial communities in the environment and on frog skin are similar across populations. Second, we assess the microbial species richness and relative abundance on frogs from habitats with different salinity levels to determine how salinity affects the microbiome. Third, we test whether the relative abundances of putatively pathogen-resistant bacterial species differ between frogs from inland and coastal environments. We found that the frog microbiome is more similar among frogs than to the microbial communities found in surface water and soil, but there is overlap between frog skin and the environmental samples. Skin microbial community richness did not differ among populations, but the relative abundances of microbes were different across populations and salinities. We found no differences in the relative abundances of the anti-fungal bacteria Janthinobacterium lividum, the genus Pseudomonas, and Serratia marcescens, suggesting that environmental exposure to saltwater has a limited influence on these putatively beneficial bacterial taxa.
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Anuros/microbiologia , Bactérias/isolamento & purificação , Microbiota , Pele/microbiologia , Animais , Anuros/classificação , Bactérias/classificação , Bactérias/genética , Ecossistema , Ilhas , Áreas AlagadasRESUMO
Resource pulses are well documented and have important consequences for population dynamics relative to continuous inputs. However, pulses of top-down factors (e.g. predation) are less explored and appreciated in the ecological literature. Here, we use a simple differential equation population model to show how pulsed removals of individuals from a population alter population size relative to continuous dynamics. Pulsed removals result in lower equilibrium population sizes relative to continuous removals, and the differences are greatest at low population growth rates, high removal rates, and with large, infrequent pulses. Furthermore, the timing of the removal pulses (either stochastic or cyclic) affects population size. For example, cyclic removals are less likely than stochastic removals to result in population eradication, but when eradication occurs, the time until eradication is shorter for cyclic than with stochastic removals.
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We investigated how recent changes in the distribution and abundance of a fouling organism affected the strength of interactions between a commercially important foundation species and a common predator. Increases in the abundance of boring sponges that bioerode the calcified shells of oysters and other shelled organisms have been attributed to increased salinization of estuarine ecosystems. We tested the hypothesis that fouling by boring sponges will change the interaction strength between oysters and a common predator (stone crabs). We generated five oyster density treatments crossed with two sponge treatments (sponge and no sponge). We contrasted the interaction strength between stone crabs and fouled and non-fouled oysters by comparing the parameters of fitted functional response curves based on Rogers random predation model. We found that fouled oysters suffered higher predation from stone crabs, and that the increased predation risk stemmed from a reduction in the handling time needed to consume the fouled oysters. These findings highlight the importance of understanding the effects of abiotic changes on both the composition of ecological communities, and on the strengths of direct and indirect interactions among species. Global climate change is altering local ecosystems in complex ways, and the success of restoration, management, and mitigation strategies for important species requires a better appreciation for how these effects cascade through ecosystems.
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BACKGROUND: In many regions, freshwater wetlands are increasing in salinity at rates exceeding historic levels. Some freshwater organisms, like amphibians, may be able to adapt and persist in salt-contaminated wetlands by developing salt tolerance. Yet adaptive responses may be more challenging for organisms with complex life histories, because the same environmental stressor can require responses across different ontogenetic stages. Here we investigated responses to salinity in anuran amphibians: a common, freshwater taxon with a complex life cycle. We conducted a meta-analysis to define how the lethality of saltwater exposure changes across multiple life stages, surveyed wetlands in a coastal region experiencing progressive salinization for the presence of anurans, and used common garden experiments to investigate whether chronic salt exposure alters responses in three sequential life stages (reproductive, egg, and tadpole life stages) in Hyla cinerea, a species repeatedly observed in saline wetlands. RESULTS: Meta-analysis revealed differential vulnerability to salt stress across life stages with the egg stage as the most salt-sensitive. Field surveys revealed that 25% of the species known to occur in the focal region were detected in salt-intruded habitats. Remarkably, Hyla cinerea was found in large abundances in multiple wetlands with salinity concentrations 450% higher than the tadpole-stage LC50. Common garden experiments showed that coastal (chronically salt exposed) populations of H. cinerea lay more eggs, have higher hatching success, and greater tadpole survival in higher salinities compared to inland (salt naïve) populations. CONCLUSIONS: Collectively, our data suggest that some species of anuran amphibians have divergent and adaptive responses to salt exposure across populations and across different life stages. We propose that anuran amphibians may be a novel and amenable natural model system for empirical explorations of adaptive responses to environmental change.
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Understanding how climate change and other environmental stressors will affect species is a fundamental concern of modern ecology. Indeed, numerous studies have documented how climate stressors affect species distributions and population persistence. However, relatively few studies have investigated how multiple climate stressors might affect species. In this study, we investigate the impacts of how two climate change factors affect an important foundation species. Specifically, we tested how ocean acidification from dissolution of CO 2 and increased sea surface temperatures affect multiple characteristics of juvenile eastern oysters (Crassostrea virginica). We found strong impacts of each stressor, but no interaction between the two. Simulated warming to mimic heat stressed summers reduced oyster growth, survival, and filtration rates. Additionally, we found that CO 2-induced acidification reduced strength of oyster shells, which could potentially facilitate crab predation. As past studies have detected few impacts of these stressors on adult oysters, these results indicate that early life stages of calcareous marine organisms may be more susceptible to effects of ocean acidification and global warming. Overall, these data show that predicted changes in temperature and CO 2 can differentially influence direct effects on individual species, which could have important implications for the nature of their trophic interactions.
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Research in behavioral ecology routinely quantifies individual variation in behavior using transitions between discrete environments, for example prey moving from a no predator to predator treatment. This research often ignores behavioral variation in response to temporal fluctuations in environmental conditions around an unchanging mean environment. In this study we evaluate the effects of temporal fluctuations in predation risk (predator cue concentration of Procambarus spp.), without the confounding effects of a changing mean, on among-individual variation in anti-predator behavior in freshwater snails (Physa acuta). We also evaluate how the interaction between environmental and individual variation affects snail survival and reproduction by exposing snails to lethal predators following the behavioral assays. Our analyses revealed a trend towards higher among-individual variation in mean behavior when snails were exposed to intermediate levels of environmental variation compared to highly variable or constant environments. However, because of large uncertainty in estimates of among-individual variation, differences among treatments were indistinguishable from noise for most, but not all behaviors. In the lethal predator trials, snail survival and time to mortality was the lowest in the high variation environment. Also, as environmental variation increased snail egg production decreased and snails laid more of their eggs underneath a provided shelter.
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To adaptively express inducible defenses, prey must gauge risk based on indirect cues of predation. However, the information contained in indirect cues that enable prey to fine-tune their phenotypes to variation in risk is still unclear. In aquatic systems, research has focused on cue concentration as the key variable driving threat-sensitive responses to risk. However, while risk is measured as individuals killed per time, cue concentration may vary with either the number or biomass killed. Alternatively, fine-grained variation in cue, that is, frequency of cue pulses irrespective of concentration, may provide a more reliable signal of risk. Here, we present results from laboratory experiments that examine the relationship between red-eyed treefrog tadpole growth and total cue, cue per pulse, and cue pulse frequency. We also reanalyze an earlier study that examined the effect of fine-grained variation in predator cues on wood frog tadpole growth. Both studies show growth declines with increasing cue pulse frequency, even though individual pulses in high-frequency treatments contained very little cue. This result suggests that counter to earlier conclusions, tadpoles are using fine-grained variation in cue arising from the number of predation events to assess and respond to predation risk, as predicted by consumer-resource theory.