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Biological processes play important roles in determining how global changes manifest at local scales. Primary producers can absorb increased CO2 via daytime photosynthesis, modifying pH in aquatic ecosystems. Yet producers and consumers also increase CO2 via respiration. It is unclear whether biological modification of pH differs across the year, and, if so, what biotic and abiotic drivers underlie temporal differences. We addressed these questions using the intensive study of tide pool ecosystems in Alaska, USA, including quarterly surveys of 34 pools over 1 year and monthly surveys of five pools from spring to fall in a second year. We measured physical conditions, community composition, and changes in pH and dissolved oxygen during the day and night. We detected strong temporal patterns in pH dynamics. Our measurements indicate that pH modification varies spatially (between tide pools) and temporally (across months). This variation in pH dynamics mirrored changes in dissolved oxygen and was associated with community composition, including both relative abundance and diversity of benthic producers and consumers, whose role differed across the year, particularly at night. These results highlight the importance of the time of year when considering the ways that community composition influences pH conditions in aquatic ecosystems.

Climate change manifests unevenly across space and time and produces complex patterns of stress for ecological systems. Species can also show substantial among‐population variability in response to environmental change across their geographic range due to evolutionary processes. Explanatory factors or their proxies, such as temperature and latitude, help parse these sources of environmental and intraspecific variability; however, overemphasizing latitudinal trends can obscure the role of local environmental conditions in shaping population vulnerability to climate change. Focusing on the geographic center of a species range to disentangle latitude, we test the hypothesis that populations from warmer regions of a species range are more vulnerable to ocean warming. We conducted a mesocosm experiment and field reciprocal transplant with four populations of a marine snail, Nucella lamellosa, from two regions in British Columbia, Canada, that differ in thermal characteristics: the Central Coast, a cool region, and the Strait of Georgia, one of the warmest regions of this species' range and one that is warming faster than the Central Coast. Populations from the Strait of Georgia experienced growth reductions at contemporary summertime seawater temperatures in the laboratory and showed stark reductions in survival and growth under future seawater conditions and when outplanted at their native transplant sites. This indicates a high vulnerability to ocean warming, especially given the faster rate of ocean warming in this region. In contrast, populations from the cooler Central Coast demonstrated high performance at contemporary seawater temperatures and high growth and survival in projected future seawater temperatures and at their native outplant sites. Given their position within the geographic center of N. lamellosa's range, extirpation events in the vulnerable Strait of Georgia populations could compromise connectivity within the metapopulation and lead to gaps across this species' range. Overall, our study supports predictions that populations from warm regions of species ranges are more vulnerable to environmental warming, suggests that the Strait of Georgia and other inland or coastal seas could be focal points for climate change effects and ecological transformation, and emphasizes the importance of analyzing climate change vulnerability in the context of regional environmental data and throughout a species' range. [ABSTRACT FROM AUTHOR]

Plankton live under the countervailing selective pressures of predation and ultraviolet radiation (UVR). In lakes, zooplankton are transparent reducing visibility to predatory fishes but are pigmented in the absence of fishes, hypothetically reducing UVR damage. In the sea, planktivorous fishes are widespread, so plankton typically are transparent and ascend to productive surface waters at night to forage and descend during the day to reduce visibility to predators. However, larvae of some species face the unique constraint of traveling in surface currents in the daytime during migrations between adult and larval habitats. We would expect these larvae to be transparent since companion studies demonstrated increased predation risk of pigmented larvae under strong sunlight. Paradoxically, larvae range from being darkly to lightly pigmented. We hypothesize that some larvae are more heavily pigmented to reduce UVR damage, while other species travelling in subsurface currents with low UVR might be more transparent. Linking larval morphology to depth-dependent selective pressures would add a key element to help improve predictions of larval vertical distributions, which are important for simulating larval transport trajectories. We quantitatively tested the hypothesis that selection may have favored photoprotective pigmentation for larvae in the predominantly transparent plankton community while testing the differential effects of UVA and UVB radiation. We measured larval pigmentation of 12 species of crabs and exposed them to visible light only, visible + UVA, or visible + UVA + UVB in the tropics. Controlling for phylogeny, more pigmented species survived UVR better than less pigmented species, especially on sunnier days, though intraspecific comparisons for four species were equivocal. Most species died even from UVA exposure, which has long been regarded as relatively harmless despite penetrating deeper underwater than UVB. Thus, we demonstrate with a phylogenetically controlled analysis that crab larvae are pigmented in the predominantly transparent planktonic community to protect from UVR, improving our understanding of the selective forces acting on animal coloration and the factors determining planktonic distributions, survival, and dispersal. This linkage of morphology with susceptibility will be important for developing mechanistic models of environmental stress responses to better predict larval dispersal in current and future climates.

Dispersal has far‐reaching implications for individuals, populations, and communities, especially in sessile organisms. Escaping competition with conspecifics and with kin are theorized to be key factors leading to dispersal as an adaptation. However, manipulative approaches in systems in which adults are sessile but offspring have behaviors is required for a more complete understanding of how competition affects dispersal. Here, we integrate a series of experiments to study how dispersal affects the density and relatedness of neighbors, and how the density and relatedness of neighbors in turn affects fitness. In a marine bryozoan, we empirically estimated dispersal kernels and found that most larvae settled within ~1 m of the maternal colony, although some could potentially travel at least 10s of meters. Larvae neither actively preferred or avoided conspecifics or kin at settlement. We experimentally determined the effects of spreading sibling larvae by manipulating the density and relatedness of settlers and measuring components of fitness in the field. We found that settler density reduced maternal fitness when settler neighbors were siblings compared with when neighbors were unrelated or absent. Genetic markers also identified very few half sibs (and no full sibs) in adults from the natural population, and rarely close enough to directly interact. In this system, dispersal occurs over short distances (meters) yet, in contrast with expectations, there appears to be limited kinship between adult neighbors. Our results suggest that the limited dispersal increases early offspring mortality when siblings settle next to each other, rather than next to unrelated conspecifics, potentially reducing kinship in adult populations. High offspring production and multiple paternity could further dilute kinship at settlement and reduce selection for dispersal beyond the scale of 10s of meters. [ABSTRACT FROM AUTHOR]

Environmental stress can reshape trophic interactions by excluding predators or rendering prey vulnerable, depending on the relative sensitivity of species to the stressor. Classical models of food web responses to stress predict either complete predator exclusion from stressed areas or complete prey vulnerability if predators are stress tolerant. However, if the consumer response to the stress is individually variable, the result may be a distributive stress model (DSM) whereby predators distribute consumption pressure across a range of prey guilds and their trophic niche is expanded. We test these models in one of the largest hypoxic "Dead Zones" in the world, the northern Gulf of Mexico, by combining geochemical tracers of hypoxia exposure and isotope ratios to assess individual‐level trophic responses. Hypoxia‐exposed fish occupied niche widths that were 14.8% and 400% larger than their normoxic counterparts in two different years, consistent with variable displacement from benthic to pelagic food webs. The degree of isotopic displacement depended on the magnitude of hypoxia exposure. These results are consistent with the DSM and highlight the need to account for individually variable sublethal effects when predicting community responses to environmental stress. [ABSTRACT FROM AUTHOR]

Biodiversity is typically considered as a one‐dimensional metric (e.g., species richness), yet the consequences of species loss may be different depending on where extinctions occur in the food web. Here, I used a manipulative field experiment in a temperate subtidal marine system to explore the implications of diversity loss at multiple trophic levels for ecosystem functioning and food web structure. The four manipulated predators included the small painted goby and common prawn, which are also fed on by the larger black goby and shore crab. Antagonistic interactions between the manipulated predators (e.g., intraguild predation, intimidation, interference competition) limited their negative effects on the rest of the food web. Top‐down control was so suppressed at the highest level of multitrophic diversity that the resulting food webs were as complex and productive as those containing no manipulated predators. Negative interactions between the predators weakened as multitrophic diversity decreased, however, resulting in stronger consumption of lower trophic levels and a simpler food web with lower rates of two key ecosystem processes: primary production and decomposition. These results show how indirect interactions between predators on multiple trophic levels help to promote the complexity and functioning of natural systems. [ABSTRACT FROM AUTHOR]

The metabolic theory of ecology ( MTE) and ecological stoichiometry ( ES) are both prominent frameworks for understanding energy and nutrient budgets of organisms. We tested their separate and joint power to predict nitrogen (N) and phosphorus (P) excretion rates of ectothermic aquatic invertebrate and vertebrate animals (10,534 observations worldwide). MTE variables (body size, temperature) performed better than ES variables (trophic guild, vertebrate classification, body N:P) in predicting excretion rates, but the best models included variables from both frameworks. Size scaling coefficients were significantly lower than predicted by MTE (<0.75), were lower for P than N, and varied greatly among species. Contrary to expectations under ES, vertebrates excreted both N and P at higher rates than invertebrates despite having more nutrient-rich bodies, and primary consumers excreted as much nutrients as carnivores despite having nutrient-poor diets. Accounting for body N:P hardly improved upon predictions from treating vertebrate classification categorically. We conclude that basic data on body size, water temperature, trophic guild, and vertebrate classification are sufficient to make general estimates of nutrient excretion rates for any animal taxon or aquatic ecosystem. Nonetheless, dramatic interspecific variation in size-scaling coefficients and counter-intuitive patterns with respect to diet and body composition underscore the need for field data on consumption and egestion rates. Together, MTE and ES provide a powerful conceptual basis for interpreting and predicting nutrient recycling rates of aquatic animals worldwide. [ABSTRACT FROM AUTHOR]

A persistent challenge for ecologists is understanding the ecological mechanisms that maintain global patterns of biodiversity, particularly the latitudinal diversity gradient of peak species richness in the tropics. Spatial and temporal variation in community composition contribute to these patterns of biodiversity, but how this variation and its underlying processes change across latitude remains unresolved. Using a model system of sessile marine invertebrates across 258 of latitude, from the temperate zone to the tropics, we tested the prediction that spatial and temporal patterns of taxonomic richness and composition, and the community assembly processes underlying these patterns, will differ across latitude. Specifically, we predicted that high beta diversity (spatial variation in composition) and high temporal turnover contribute to the high species richness of the tropics. Using a standardized experimental approach that controls for several confounding factors that hinder interpretation of prior studies, we present results that support our predictions. In the temperate zone, communities were more similar across spatial scales from centimeters to tens of kilometers and temporal scales up to one year than at lower latitudes. Since the patterns at northern latitudes were congruent with a null model, stochastic assembly processes are implicated. In contrast, the communities in the tropics were a dynamic spatial and temporal mosaic, with low similarity even across small spatial scales and high temporal turnover at both local and regional scales. Unlike the temperate zone, deterministic community assembly processes such as predation likely contributed to the high beta diversity in the tropics. Our results suggest that community assembly processes and temporal dynamics vary across latitude and help structure and maintain latitudinal patterns of diversity. [ABSTRACT FROM AUTHOR]

Latitudinal patterns of nonnative species richness suggest fewer successful invasions in the tropics, relative to temperate regions. One main hypothesis for this pattern is that biotic resistance to invasion is stronger in the tropics than at higher latitudes. Biotic resistance can limit the distribution and abundance of nonnative species and, in extreme cases, can prevent establishment. We provide the first experimental test of this hypothesis, comparing the strength of biotic resistance in a tropical and a temperate marine ecosystem. Prédation is one mechanism of biotic resistance, and since prédation can be stronger at lower latitudes, we predicted that prédation will serve to increase biotic resistance more in the tropics than at higher latitude. We conducted predator-exclusion experiments on marine epifaunal communities, a heavily invaded system, focusing on nonnative tunicates as a model fauna. The effect of prédation on species richness of nonnative tunicates was more than three times greater at sites in tropical Panama than in temperate Connecticut, consistent with the prediction of stronger biotic resistance in the tropics. In Connecticut, prédation reduced the abundance of one nonnative tunicate but did not affect the abundances of any other nonnative tunicate species, and no species were excluded from communities. In contrast, prédation resulted in striking reductions in abundance and often exclusion of nonnative tunicates from experimental communities in Panama. If proved to be general, latitudinal differences in the biotic resistance of communities to nonnative species establishment may help explain emerging patterns of global invasions. [ABSTRACT FROM AUTHOR]

Overexploitation of predators has been linked to the collapse of a growing number of shallow-water marine ecosystems. However, salt-marsh ecosystems are often viewed and managed as systems controlled by physical processes, despite recent evidence for herbivore-driven die-off of marsh vegetation. Here we use field observations, experiments, and historical records at 14 sites to examine whether the recently reported die-off of northwestern Atlantic salt marshes is associated with the cascading effects of predator dynamics and intensive recreational fishing activity. We found that the localized depletion of top predators at sites accessible to recreational anglers has triggered the proliferation of herbivorous crabs, which in turn results in runaway consumption of marsh vegetation. This suggests that overfishing may be a general mechanism underlying the consumer-driven die-off of salt marshes spreading throughout the western Atlantic. Our findings support the emerging realization that consumers play a dominant role in regulating marine plant communities and can lead to ecosystem collapse when their impacts are amplified by human activities, including recreational fishing. [ABSTRACT FROM AUTHOR]

The species composition, density, and frequency of recruitment into any given habitat are highly variable in most biological systems that rely on dispersive propagules (larvae, seeds, spores, etc.). There are few direct experimental studies of how recruitment variation between single species influences the composition and assembly of whole communities in many of these systems. We manipulated recruitment of a variety of single taxa and followed their effects on the subsequent development of hard-substrate communities of sessile animals living in temperate marine waters. The effects of recruitment on communities were complex. Patterns of recruitment of individual species influenced community structure, but these effects varied greatly depending on the identity of species recruits, the time of community development, and location across three different sites. Variable recruitment of arborescent bryozoans and didemnid ascidians had little effect on community structure. At one site, recruitment of the colonial ascidian Botryllus schlosseri had short-lived effects on community structure, while barnacles had more persistent effects. At another site, recruitment of B. schlosseri and the bryozoan Watersipora subtorquata had strong persistent effects on community structure, dominating space where they recruited and influencing the abundances of a variety of different taxa. Differences in the effects of species recruitment on communities appear to be caused by differences between the ecology and life history of recruiting species as well as differences in background processes between sites. These results demonstrate that discrete recruitment events that vary between single species can be important drivers of community composition but are likely to be heavily influenced by the local environment, even within a single species. [ABSTRACT FROM AUTHOR]

Coastal environments are among the most productive on the planet, providing a wide range of ecosystem services. Development and exploitation mean that they are faced with stresses from a number of anthropogenic sources. Such stresses are typically studied in isolation, but multiple stressors can combine in unexpected ways to alter the structure of ecological systems. Here, we experimentally explore the impacts of inorganic nutrients and organic matter on a range of food web properties. We find that these two stressors combine additively to produce significant increases in connectance and mean food chain length. Such increases are typically associated with enhanced robustness to secondary extinctions and productivity, respectively. Despite these apparent beneficial effects, we find a simplification of web structure in terms of taxon richness and diversity, and altered proportions of basal and top species. These effects are driven by a reduction in community assembly and lower consistency in a range of system properties as a result of the multiple stressors. Consequently, impacted food webs are likely to be more vulnerable to human- or climate-induced perturbations in the long term. [ABSTRACT FROM AUTHOR]

For sessile organisms, dispersal and recruitment are typically spatially stochastic, but there is little understanding of how this variability scales up to influence processes such as competitive coexistence. Here we argue that coexistence of benthic marine animals is enhanced by stochastic differences between species in the spatial distribution of larval settlement. Differentiation of settlement distributions among competitors results in intraspecifically aggregated settlement, which can reduce overall interspecific competition and increase overall intraspecific competition. We test for the components of this mechanism using a pair of subtidal invertebrates, and we find that the mean interspecific effect of the dominant competitor is substantially reduced by natural settlement variability. Using a simulation parameterized with experimental data, we find that variable settlement could play an important role in long-term coexistence between these species. This mechanism may apply broadly to benthic marine communities, which can be highly diverse and typically exhibit large settlement fluctuation over a range of scales. [ABSTRACT FROM AUTHOR]

The indirect, ecosystem-level consequences of ocean fishing, and particularly the mechanisms driving them, are poorly understood. Most studies focus on density-mediated trophic cascades, where removal of predators alternately causes increases and decreases in abundances of lower trophic levels. However, cascades could also be driven by where and when prey forage rather than solely by prey abundance. Over a large gradient of fishing intensity in the central Pacific's remote northern. Line Islands, including a nearly pristine, baseline coral reef system, we found that changes in predation risk elicit strong behavioral responses in foraging patterns across multiple prey fish species These responses were observed as a function of both short-term ("acute") risk and longer-term ("chronic") risk, as well as when prey were exposed to model predators to isolate the effect of perceived predation risk from other potentially confounding factors. Compared to numerical prey responses, antipredator. behavioral responses such as these can potentially have far greater net impacts (by occurring over entire assemblages) and operate over shorter temporal scaIe (with potentially instantaneous response times) in transmitting top-down effects. A rich body of literature exists on both the direct effects of human removal of predators from ecosystems and predators' effects on prey behavior. Our results draw together these lines of research and provide the first empirical evidence that large-scale human removal of predators from a natural ecosystem indirectly alters prey behavior. These behavioral changes may, in turn, drive previously unsuspected alterations in reef food webs. [ABSTRACT FROM AUTHOR]

Intraguild predation (IGP) occurs when one predator species consumes another predator species with whom it also competes for shared prey. One question of interest to ecologists is whether multiple predator species suppress prey populations more than a single predator species, and whether this result varies with the presence of IGP. We conducted a meta-analysis to examine this question, and others, regarding the effects of IGP on prey suppression. When predators can potentially consume one another (mutual . IGP), prey suppression is greater in the presence of one predator species than in the presence of multiple predator species; however, this result was not found for assemblages with unidirectional or no IGP. With unidirectional IGP, intermediate predators were generally more effective than the top predator at suppressing the shared prey, in agreement with TOP theory. Adding a top predator to an assemblage generally caused prey to be released from predation, while adding an intermediate predator caused prey populations to be suppressed. However, the effects of adding a top or intermediate predator depended on the effectiveness of these predators when they were alone. Effects of IGP varied across different ecosystems (e.g., lentic, lotic, marine, terrestrial invertebrate, and terrestrial vertebrate), with the strongest patterns being driven by terrestrial invertebrates: Finally, although IGP theory is based on equilibrium conditions, data from short-term experiments can inform us about systems that are dominated by transient dynamics. Moreover, short-term experiments may be connected in some way to equilibrium models if the predator and prey densities used in experiments approximate the equilibrium densities in nature. [ABSTRACT FROM AUTHOR]

Pervasive overharvesting of consumers and anthropogenic nutrient loading are changing the strengths of top-down and bottom-up forces in ecosystems worldwide. Thus, identifying the relative and synergistic roles of these forces and how they differ across habitats, ecosystems, or primary-producer types is increasingly important for understanding how communities are structured. We used factorial meta-analysis of 54 field experiments that orthogonally manipulated herbivore pressure and nutrient loading to quantify consumer and nutrient effects on primary producers in benthic marine habitats. Across all experiments and producer types, herbivory and nutrient enrichment both significantly affected primary-producer abundance. They also interacted to create greater nutrient enrichment effects in the absence of herbivores, suggesting that loss of herbivores produces more dramatic effects of nutrient loading. Herbivores consistently had stronger effects than did nutrient enrichment for both tropical macroalgae and seagrasses. The strong effects of herbivory but limited effects of nutrient enrichment on tropical macroalgae suggest that suppression of herbivore populations has played a larger role than eutrophication in driving the phase shift from coral- to macroalgal-dominated reefs in many areas, especially the Caribbean. For temperate macroalgae and benthic microalgae, the effects of top-down and bottom-up forces varied as a function of the inherent productivity of the ecosystem. For these algal groups, nutrient enrichment appeared to have stronger effects in high- vs. low-productivity systems, while herbivores exerted a stronger top-down effect in low-productivity systems. Effects of herbivores vs. nutrients also varied among algal functional groups (crustose algae, upright macroalgae, and filamentous algae), within a functional group between temperate and tropical systems, and according to the metric used to measure producer abundance. These analyses suggest that human alteration of food webs and nutrient availability have significant effects on primary producers but that the effects vary among latitudes and primary producers, and with the inherent productivity of ecosystems. [ABSTRACT FROM AUTHOR]

Nitrogen and carbon stable isotope data are frequently used to describe the origins and transformations of organic matter. Nitrogen stable isotopes (δ15N) in tissue are used to estimate species' trophic levels, the extent of omnivory, food chain length, and community-wide relationships between body size and trophic level; the latter leading to estimates of predator-prey mass ratios for parameterization, calibration, and validation of food web models. Carbon stable isotopes (δ13C) are used to identify pathways linking producers and consumers and for studies of migration and movement. Collectively, δ15N and δ13C, often with other stable isotopes such as δ34S, may be used to define the contribution of different producers and pathways to consumer production, to assess the trophic impacts of invasive species and habitat modification, and to predict past habitat use, movements, and migrations. Stable isotope data often complement dietary data (e.g., from stomach contents) in food web studies, because stable isotope composition is indicative of assimilated diet over months to years, depending on species, size, environment, and tissue type. There are relatively few large-scale compilations of δ15N and δ13C data for marine species from offshore habitats, but such data facilitate comparative analysis and research into food web structure and function. The data provided comprise 5535 records for individuals of 62 species of fish and squid weighing 0.3 g to 17920 g and sampled from the northeast Atlantic shelf seas (Celtic Sea, North Sea, Irish Sea, Channel) from 2002 to 2010. For every sampled individual the record lists: species name, date of sampling, position of sampling, body mass, percentage nitrogen in muscle tissue, percentage carbon in muscle tissue, and δ15N and δ13C natural abundance in muscle tissue. Awareness of, and access to, these data should catalyze and facilitate new research with stable isotopes, to improve understanding of marine biology, food web ecology, and human impacts on the environment. [ABSTRACT FROM AUTHOR]

Knowledge of relationships between predator size and prey size are needed to describe interactions of species and size classes in food webs. Most estimates of predator and prey sizes have been based on dietary studies and apply to small numbers of species in a relatively narrow size range. These estimates may or may not be representative of values for other groups of species and body sizes or for other locations. Marine predator and prey size data associated with published literature were identified and collated to produce a single data set. If predator or prey length of mass were not measured in the original study, the length or mass was calculated using length-mass relationships. The data set consists of 34931 records from 27 locations covering a wide range of environmental conditions from the tropics to the poles and for 93 types of predator with sizes ranging from 0.1 mg to over 415 kg and 174 prey types with sizes from 75 pg to over 4.5 kg. Each record includes: predator and prey scientific names, common names, taxa, life stages and sizes (length and mass with conversion details), plus the type of feeding interaction, geographic location (with habitat description, latitude, longitude) and mean annual environ- mental data (sea surface temperature and primary productivity). [ABSTRACT FROM AUTHOR]