Search

Ecological patterns are determined by the interplay between abiotic factors and interactions among species. As the Earth's climate warms, interactions such as competition, predation, and mutualism are changing due to shifts in per capita interaction strength and the relative abundance of interacting species. Changes in interspecific relationships, in turn, can drive important local-scale changes in community dynamics, biodiversity, and ecosystem functioning, and can potentially alter large-scale patterns of distribution and abundance. In many cases, the importance of indirect effects of warming, mediated by changing species interactions, will be greater—albeit less well understood—than direct effects in determining the community- and ecosystem-level outcomes of global climate change. Despite considerable community-specific idiosyncrasy, ecological theory and a growing body of data suggest that certain general trends are emerging at local scales: positive interactions tend to become more prevalent with warming, and top trophic levels are disproportionately vulnerable. In addition, important ecological changes result when the geographic overlap between species changes, and when the seasonal timing of life history events of interacting species falls into or out of synchrony. We assess the degree to which such changes are predictable, and urge advancement on several high priority questions surrounding the relationships between temperature and community ecology. An improved understanding of how assemblages of multiple, interacting species will respond to climate change is imperative if we hope to effectively prepare for and adapt to its effects.

Global warming has important implications for the dynamics and ecological impacts of emerging diseases. We investigated temperature effects on scuticociliate, Orchitophrya cf. stellarum, infections in ripe testes of two Pacific northeast sea stars (Asterina miniata Brandt and Pisaster ochraceus Brandt) using laboratory and field approaches. We predicted that a small increase in temperature would result in higher ciliate growth rates and heightened infection intensities. To test this we (1) cultured free-living O. stellarum at 10 and 15 °C and quantified ciliate abundance after 3 days, and (2) housed sea stars of both species at 10 and 15 °C for durations varying from 4 to 21 days and then measured the infection intensity. Ciliate densities in cultures were two orders of magnitude higher in the warmer treatment. Infection intensity was also temperature sensitive: greater proportions of testes were infected and infection stage was more advanced at 15 versus 10 °C, leading to a reduction in spermatozoa and regression of the germinal layer within three weeks. In seven field populations surveyed, we found a tight linear correlation between infection prevalence (percent infected sea stars) and infection intensity (proportion of infected testes per individual and mean infection score). However, 45% of P. ochraceus testes exhibited heavy infections versus 8% of A. miniata testes, which may relate to the different thermal habitat of each species: P. ochraceus occurs higher on the shore and likely reaches higher body temperatures at low tide. While the sex ratio of A. miniata is unbiased, P. ochraceus populations are consistently female-biased and show no relationship to infection prevalence (ranged from 30 to 90%). O. cf. stellarum infections of testes of both sea stars are prevalent in field populations, are highly temperature sensitive, and lead to rapid loss of reproductive potential.

In many coastal environments, variation in salinity and organismal responses to that variation are important determinants of the distribution and abundance of species. This study examined the effects of acute salinity changes on sea stars (Pisaster ochraceus) collected from a high-salinity site (Bamfield, BC) and a low-salinity site (Vancouver, BC). Sea stars from both sites were exposed to salinities ranging 15–30 psu. Following a 24-h exposure, the osmolality, sodium concentrations, and chloride concentrations in the perivisceral fluid all varied directly with salinity and were very close to the treatment salinities in both the Bamfield and Vancouver sea stars. The righting response (measured as an activity coefficient) was salinity dependent, with the lowest activity levels at a salinity of 15 psu. Activity coefficients did not vary between the two source populations. Feeding rates on mussels were strongly salinity dependent, but the salinity pattern was population specific. Bamfield sea stars fed the most at 30 psu, whereas Vancouver sea stars fed the most at 20 psu. High post-experimental mortalities were observed in Bamfield sea stars that had been exposed to a salinity of 15 psu; no such mortality was observed in Vancouver animals. This study provides evidence that the sea stars from the lower salinity environment had been able to acclimatize or adapt to low-salinity conditions. However, the results also suggest that there are limits to this tolerance, and that future changes in salinity may have important consequences for marine communities via alteration of keystone predation.

Ecological studies often focus on average effects of environmental factors, but ecological dynamics may depend as much upon environmental extremes. Ecology would therefore benefit from the ability to predict the frequency and severity of extreme environmental events. Some extreme events (e.g., earthquakes) are simple events: either they happen or they don't, and they are generally difficult to predict. In contrast, extreme ecological events are often compound events, resulting from the chance coincidence of run-of-the-mill factors. Here we present an environmental bootstrap method for resampling short-term environmental data (rolling the environmental dice) to calculate an ensemble of hypothetical time series that embodies how the physical environment could potentially play out differently. We use this ensemble in conjunction with mechanistic models of physiological processes to analyze the biological consequences of environmental extremes. Our resampling method provides details of these consequences that would be difficult to obtain otherwise, and our methodology can be applied to a wide variety of ecological systems. Here, we apply this approach to calculate return times for extreme hydrodynamic and thermal events on intertidal rocky shores. Our results demonstrate that the co-occurrence of normal events can indeed lead to environmental extremes, and that these extremes can cause disturbance. For example, the limpet Lottia gigantea and the mussel Mytilus californianus are co-dominant competitors for space on wave-swept rocky shores, but their response to extreme environmental events differ. Limpet mortality can vary drastically through time. Average yearly maximum body temperature of L. gigantea on horizontal surfaces is low, sufficient to kill fewer than 5% of individuals, but on rare occasions environmental factors align by chance to induce temperatures sufficient to kill >99% of limpets. In contrast, mussels do not exhibit large temporal variation in the physical disturbance caused by breaking waves, and this difference in the pattern of disturbance may have ecological consequences for these competing species. The effect of environmental extremes is under added scrutiny as the frequency of extreme events increases in response to anthropogenically forced climate change. Our method can be used to discriminate between chance events and those caused by long-term shifts in climate.

Summary 1. Small-scale distributions of rocky intertidal organisms may be determined in part by temperature and desiccation stress during low tide and the ability of organisms to resist such stresses. Biophysical modelling techniques, coupled with data on the physiological tolerance of organisms can be used to predict the frequency and severity of abiotic stress events. 2. The limpet Lottia gigantea is a major competitor for primary space on rocky shores along the west coast of North America. The goal of this study was to quantify the thermal and desiccation tolerance of this species and to predict the frequency of stressful events at Hopkins Marine Station (HMS) in Pacific Grove, CA, USA. 3. Lottia gigantea were exposed to thermal stress regimes in the laboratory using protocols designed to mimic prolonged exposures in the field. The production of heat shock protein 70 (Hsp70) was measured as an indicator of sub-lethal stress, and mortality rates were measured. 4. Peak expression of Hsp70 occurred during exposures at 32 ° C, while the lethal limits of the limpets ranged between 30 and 42 ° C, depending on the exposure regime. Lethal temperatures were approximately 5 ° C lower in 50‐60% relative humidity conditions compared to 100% relative humidity trials. 5. Results from the physiology experiments were compared to field surveys of the upper limits of L. gigantea at HMS and with heat-budget model predictions of limpet body temperatures on a variety of substratum orientations. 6. The highest frequency of sub-lethal and lethal stress events was predicted to occur on highshore westerly and southerly slopes and horizontal surfaces, microhabitats where L. gigantea is comparatively rare at HMS. Predicted stress and mortality events were rare at low-shore sites, vertical surfaces, and sloping rock faces oriented towards the northeast and northwest. 7. The combination of direct measurements of sub-lethal and lethal temperature thresholds using realistic stress exposures and predictions from a heat-budget model give insight into the role of thermal disturbance in affecting the local-scale distributions of intertidal organisms. These methods hold great promise for enhancing our understanding of the role of abiotic factors in limiting species distributions and making predictions about responses to future climatic conditions.

Anthropogenic climate change poses a serious threat to biodiversity. In marine environments, multiple climate variables, including temperature and CO 2 concentration ([CO 2 ]), are changing simultaneously. Although temperature has well-documented ecological effects, and many heavily calcified marine organisms experience reduced growth with increased [CO 2 ], little is known about the combined effects of temperature and [CO 2 ], particularly on species that are less dependent on calcified shells or skeletons. We manipulated water temperature and [CO 2 ] to determine the effects on the sea star Pisaster ochraceus , a keystone predator. We found that sea star growth and feeding rates increased with water temperature from 5 °C to 21 °C. A doubling of current [CO 2 ] also increased growth rates both with and without a concurrent temperature increase from 12 °C to 15 °C. Increased [CO 2 ] also had a positive but nonsignificant effect on sea star feeding rates, suggesting [CO 2 ] may be acting directly at the physiological level to increase growth rates. As in past studies of other marine invertebrates, increased [CO 2 ] reduced the relative calcified mass in sea stars, although this effect was observed only at the lower experimental temperature. The positive relationship between growth and [CO 2 ] found here contrasts with previous studies, most of which have shown negative effects of [CO 2 ] on marine species, particularly those that are more heavily calcified than P. ochraceus . Our findings demonstrate that increased [CO 2 ] will not have direct negative effects on all marine invertebrates, suggesting that predictions of biotic responses to climate change should consider how different types of organisms will respond to changing climatic variables.

Summary 1On thermally stressful rocky shores, small, slow-moving ectotherms such as limpets exhibit morphological characteristics such as high-spired and heavily ridged shells which may reduce the likelihood of reaching stressful or lethal body temperatures. 2The effects of shell height and shell surface area on predicted limpet body temperatures were tested with a previously developed heat budget model. The model was parameterized with morphological data from three species (Lottia gigantea, Patella vulgata and Siphonaria gigas), which differ dramatically in their morphology and in the body temperatures they are likely to reach in the field. 3Limpet models and standard cones with higher height : length ratios lost heat to convection more readily than models with lower spired shells. 4Heavily ridged shells lost heat to convection more readily than smoother shells, but this effect was only pronounced at high wind velocities. 5When the heat budget model parameters were applied to a real environmental data set, the model predicts that maximum body temperatures and cumulative thermal stress vary among species. These differences are related primarily to the height : length ratio of the shell, and to a lesser extent to the presence of ridges. 6These results suggest that some intra- and interspecific variation in limpet morphology may be phenotypic or evolutionary responses to variation in environmental temperatures. Our findings are supported by observed patterns of limpet morphological variation across natural thermal gradients.

Temperature is among the main structuring agents on rocky intertidal shores. Although infrequent mortality events associated with high temperatures have been observed in several inter- tidal taxa, careful documentation of these events is rare, and small-scale variability in mortality patterns remains poorly understood. In Bodega Bay, California, USA, 2 mortality events occurred on exposed rocky shores during the spring of 2004 when low tides occurred around mid-day. Limpets Lottia scabra were killed during unseasonably warm weather in mid-March. In late April, record- high temperatures resulted in widespread mortality of the mussel Mytilus californianus. Levels of mortality for both species were closely associated with small-scale variability in temperature, which in turn was closely associated with substratum orientation. Invertebrates occupying surfaces facing the sun when stress was most intense were much more likely to die than those living on surfaces angled ≥45° away. Because the within-day timing of thermal stress varied seasonally, the highest mortalities were recorded on southwest-facing surfaces on the March afternoon low tide and on southeast-facing surfaces on the April morning low tide. Limpets and mussels showed little mortality during the events that were harmful to the other taxon, suggesting that these different taxa respond to different aspects of their thermal environment. If climate change results in more frequent and more severe episodes of thermal stress, future ecological change may also be heavily dependent on tidal dynamics and small-scale variation in substrate orientation.

Although global warming is acknowledged as a primary threat to populations and communities, the impact of rising temperature on community structure remains poorly understood. In this study, we investigated the direct and indirect effects of temperature on epilithic primary producers (micro- and macroalgae) and an abundant consumer, the rough limpet Lottia scabra, in the rocky intertidal zone in central and northern California, USA. We factorially manipulated temperature and limpet abundance in the field to determine the effects of temperature on herbivore growth and mortality, algal abundance, and the strength of plant–herbivore interactions. Microalgal growth was positively affected by shading at both locations, and negatively affected by limpet grazing at Pacific Grove but not at Bodega Bay. Macroalgae were only abundant at Bodega Bay, where changes in abundance were negatively related to grazing and independent of temperature. Despite temperature-related changes in microalgal food supply, there were no direct or indirect effects of temperature manipulation on L. scabra growth or mortality. Furthermore, temperature did not alter the importance of herbivory at either site. These results indicate that the influence of increasing temperature, as is predicted with climate change, will have differential effects on producers and consumers. However, thermal effects at one trophic level do not necessarily propagate through the food web to other trophic levels.

Body size plays a crucial role in determining the strength of species interactions, population dynamics, and community structure. We measured how changes in body size affect the trophic relationship between the sea star Pisaster ochraceus and its prey, the mussel Mytilus trossulus. We tested the effects of a wide range of predator and prey sizes on sea stars' prey-size preference, feeding rate, and prey tissue consumption. We found that preferred prey size increased with sea star size. Pisaster consumption rate (mussels consumed per day) and tissue intake rate (grams of tissue consumed per day) also increased with sea star size. Pisaster consumption rate, but not tissue intake rate, decreased with increasing mussel size. Juvenile sea stars preferred the most profitable prey sizes-that is, those that maximized tissue consumed per unit handling time. When adult sea stars were offered larger, more profitable mussels, tissue intake rates (grams per day) tended to increase, although this relationship was not statistically significant. Our results indicate that the Pisaster-Mytilus interaction depends on the sizes of both predator and prey, that predation rates are sensitive to even small changes in body size, and that shifts in size distributions may affect predator energetics and prey numbers differently depending on the factors that limit tissue consumption rates.

PREMISE OF THE STUDY Morphology and material properties are the main components of the mechanical design of organisms, with species groups developing different optimization strategies in the context of their physical environment. For intertidal and subtidal seaweeds, possessing highly flexible and extensible tissues allows individuals to bend and reconfigure in flow, thereby reducing drag. Previous research has shown that aging may compromise these qualities. Tissue age increases with distance from the blade's meristem, which differs in its position on kelps and red algae. Here, we assess whether longitudinal patterns of blade material properties differ between these two algal groups according to tissue age. METHODS We performed tensile tests on tissues samples excised from various positions along the extent of blades in nine kelp species (basal growth) and 15 species of red algae (apical growth). KEY RESULTS We found that older tissues were less flexible and extensible than younger tissues in all species tested. As predicted, tissue near the basal meristem in kelp was more flexible and extensible than older tissue at the blade's distal end. The opposite pattern was observed for red algae, with the most flexible and extensible tissues found near the apical meristem at the distal ends of blades. CONCLUSIONS We propose that divergent patterns in the distribution of material properties along blades may have different consequences for the performance of kelps and red algae. The positioning of younger tissues at the blade base for kelps may enable these species to attain larger body sizes in wave-swept habitats.

© 2015 by the Ecological Society of America. Ocean acidification, chemical changes to the carbonate system of seawater, is emerging as a key environmental challenge accompanying global warming and other humaninduced perturbations. Considerable research seeks to define the scope and character of potential outcomes from this phenomenon, but a crucial impediment persists. Ecological theory, despite its power and utility, has been only peripherally applied to the problem. Here we sketch in broad strokes several areas where fundamental principles of ecology have the capacity to generate insight into ocean acidification's consequences. We focus on conceptual models that, when considered in the context of acidification, yield explicit predictions regarding a spectrum of population- and community-level effects, from narrowing of species ranges and shifts in patterns of demographic connectivity, to modified consumer-resource relationships, to ascendance of weedy taxa and loss of species diversity. Although our coverage represents only a small fraction of the breadth of possible insights achievable from the application of theory, our hope is that this initial foray will spur expanded efforts to blend experiments with theoretical approaches. The result promises to be a deeper and more nuanced understanding of ocean acidification and the ecological changes it portends.

Wave splash modifies the duration and timing of aerial exposure of intertidal organisms, influencing patterns of vertical zonation, thermal stress, and the consequences of climate change. Harley and Helmuth (Limnol. Oceanogr. 48:1498-1508, 2003) described a method for measuring effective shore level (ESL), a metric that combines the influence of wave splash and tidal regime on patterns of emersion and immersion. They identified immersion events as sharp drops in temperature recorded by submersible dataloggers and compared the tide height at the time of the temperature drop to the wave height recorded by an offshore buoy. Here we explore the generality of this method at 10 sites along the Pacific coast of North America spanning 14° of latitude. We deployed miniature temperature loggers at fixed intertidal heights at each site and recorded temperatures at intervals of 5 to 15 min for periods of up to 5 years. We use these data to explore the effects of different approaches to calculating temperature drops and wave heights, as well as variation in the buoy location, on ESL calculations. We present a software program (SiteParser) that can be used to identify temperature drops in a datalogger time series and also calculate daily and monthly summary statistics of temperature. We show that ESL parameters provide a useful metric for comparing the effects of wave action on immersion patterns within sites. We also introduce a metric of average wave run-up that can be used to compare the effect of wave action on immersion patterns among more distant locations.

We explicitly quantified spatial and temporal patterns in the body temperature of an ecologically important species of intertidal invertebrate, the mussel Mytilus californianus, along the majority of its latitudinal range from Washington to southern California, USA. Using long-term (five years), high-frequency temperature records recorded at multiple sites, we tested the hypothesis that local ''modifying factors'' such as the timing of low tide in summer can lead to large-scale geographic mosaics of body temperature. Our results show that patterns of body temperature during aerial exposure at low tide vary in physiologically meaningful and often counterintuitive ways over large sections of this species' geographic range. We evaluated the spatial correlations among sites to explore how body temperatures change along the latitudinal gradient, and these analyses show that ''hot spots'' and ''cold spots'' exist where temperatures are hotter or colder than expected based on latitude. We identified four major hot spots and four cold spots along the entire geographic gradient with at least one hot spot and one cold spot in each of the three regions examined (Washington- Oregon, Central California, and Southern California). Temporal autocorrelation analysis of year-to-year consistency and temporal predictability in the mussel body temperatures revealed that southern animals experience higher levels of predictability in thermal signals than northern animals. We also explored the role of wave splash at a subset of sites and found that, while average daily temperature extremes varied between sites with different levels of wave splash, yearly extreme temperatures were often similar, as were patterns of predictability. Our results suggest that regional patterns of tidal regime and local pattern of wave splash can overwhelm those of large-scale climate in driving patterns of body temperature, leading to complex thermal mosaics of temperature rather than simple latitudinal gradients. A narrow focus on population changes only at range margins may overlook climatically forced local extinctions and other population changes at sites well within a species range. Our results emphasize the importance of quantitatively examining biogeographic patterns in environ- mental variables at scales relevant to organisms, and in forecasting the impacts of changes in climate across species ranges.

Physical ecosystem engineers play dominant roles in a wide variety of communities. While many of the direct, positive effects of ecosystem engineers are readily apparent, the roles of engineers are often mediated by indirect interactions stemming from the facilitation of one or a few key species. Although direct and indirect effects are both critical drivers of community dynamics, they are rarely considered together with regards to ecosystem engineering. In the present study barnacle and herbivorous gastropod densities are experimentally manipulated to investigate the direct positive effects of habitat provision by barnacles as well as indirect effects mediated by mollus- can grazers. Molluscan grazers (Littorina spp.) and herbivorous arthropods were positively influ- enced by the presence of barnacles. Arthropod abundance and species richness were lower when Littorina spp. were present. This pattern was not influenced by barnacle cover, suggesting that competition among herbivore functional groups was strong but independent of biogenic habitat complexity. In addition, Littorina spp. had strong negative effects on the filamentous alga Urospora penicilliformis, but this effect was only seen in the absence of barnacles. Finally, Littorina spp. reduced the recruitment of the principal habitat-forming barnacle Balanus glandula, suggesting that Littorina spp. may mediate a negative feedback loop in B. glandula population dynamics. Given the ubiquity of ecosystem engineers, similar combinations of direct and indirect influences may have far-reaching consequences for community dynamics and species richness in a wide range of systems.

Anthropogenically induced global climate change has profound implications for marine ecosystems and the economic and social systems that depend upon them. The relationship between temperature and individual performance is reasonably well understood, and much climate-related research has focused on potential shifts in distribution and abundance driven directly by temperature. However, recent work has revealed that both abiotic changes and biological responses in the ocean will be substantially more complex. For example, changes in ocean chemistry may be more important than changes in temperature for the performance and survival of many organisms. Ocean circulation, which drives larval transport, will also change, with important consequences for population dynamics. Furthermore, climatic impacts on one or a few 'leverage species' may result in sweeping community-level changes. Finally, synergistic effects between climate and other anthropogenic variables, particularly fishing pressure, will likely exacerbate climate-induced changes. Efforts to manage and conserve living marine systems in the face of climate change will require improvements to the existing predictive framework. Key directions for future research include identifying key demographic transitions that influence population dynamics, predicting changes in the community-level impacts of ecologically dominant species, incorporating populations' ability to evolve (adapt), and understanding the scales over which climate will change and living systems will respond.

The ecological roles of small (1–1000 mg) predators in benthic marine systems are poorly understood. We investigated the natural history and predatory impact of one group of such mesopredators—larvae of dipteran flies in the genus Oedoparena—which prey on intertidal barnacles. We 1) quantified patterns of larval Oedoparena distribution and abundance in the Northwest Straits of Washington State, USA, 2) determined larval physiological tolerance limits in the laboratory, and 3) conducted a manipulative field experiment to assess the role of microhabitat temperature on predation rates in Oedoparena. Members of Oedoparena in Washington are univoltine, with peak larval abundance in late spring and early summer. Infestation frequencies in the barnacles Balanus glandula and Chthamalus dalli were as high as 22% and 35%, respectively. In laboratory studies, larvae of O. glauca were able to tolerate temperatures up to 37°C; however, this temperature is often exceeded in high intertidal habitats. In a field manipulation using experimental shades, we demonstrate that the alleviation of physiological stress greatly increased the abundance of larvae of Oedoparena spp. As a result of increased larval densities under shades, adult B. glandula mortality increased from 5% to nearly 30%, and C. dalli mortality increased from less than 20% to over 60%. Because high intertidal barnacles serve as food and habitat for a diverse array of species, Oedoparena spp. have the potential to play a major role in structuring high intertidal communities, particularly in cooler microhabitats.

Understanding the role of scale is critical to ecologists' ability to make appropriate measurements, to ''scale up'' from local, short-term experiments to larger areas and longer times, to formulate models of community structure, and to address important conservation problems. Although these issues have received increased attention in recent years, empirical measurements of the scales of ecologically important variables are still rare. Here, we measure the spatial and temporal scales of variability of 15 ecologically relevant physical and biological processes in the wave-swept intertidal zone at Mussel Point, near Hopkins Marine Station in California. We analyze temporal variability in wave height, ocean temperature, upwelling intensity, solar irradiance, and body temperature for periods ranging from ten minutes to fifty years. In addition, we measure spatial variation in shoreline topography, wave force, wave-induced disturbance, body temperature, species diversity, recruitment, primary productivity, and the abundances of grazers, predators, and the competitive dominant occupier of space. Each of these spatial variables is measured along three horizontal transects in the upper intertidal zone: a short transect (44 m long with sampling locations spaced at ;0.5-m intervals), a medium transect (175 m long with sampling locations spaced at ;1.7-m intervals), and a long transect (334 m long with sampling locations spaced at ;3.4-m intervals). Six different methods are used to quantify the scale of each variable. Distinct scales are evident in all but one of our temporal variables, demonstrating that our methods for quantifying scale can work effectively with relatively simple, periodic phenomena. However, our spatial results reveal basic problems that arise when attempting to measure the scale of variability for more complex phenomena. For a given variable and length of transect, different methods of calculating scale seldom agree, and in some cases estimates differ by more than an order of magnitude. For a given variable and method of calculating spatial scale, measurements are sensitive to the length of a transect; the longer the transect, the larger the estimate of scale. We propose that the ''1/ f noise'' nature of the data can explain both the variability among methods for calculating scale and the length dependence of spatial scales of variation, and that the 1/ f noise character of the data may be driven by the fractal geometry of shoreline topography. We conclude that it may not be possible to define a meaningful spatial scale of variation in this system. As an alternative to the boiled-down concept of ''scale,'' we suggest that it is more appropriate to examine explicitly the pattern in which variability changes with the extent of measurement (e.g., the spectrum). Knowledge of this pattern can provide useful ecological scaling ''rules'' even when a well-defined scale (or hierarchy of scales) cannot be discerned.

Aims Gradients of environmental variability have been pro- posed to explain spatial variation in patterns of geographical range size. We explore this relationship in NE Pacific algae and NW Atlantic gastropods by using the characteristics of species' bathymetric distributions as a proxy for tolerance of environmental variability.

Rocky intertidal ecosystems are characterized by marked zonation patterns in which species replace one another along the vertical gradient of emersion time. Yet, we still do not fully understand the reasons that zonation patterns are variable in space and time. Here, we use effective shore level (ESL), a metric that incorporates the modifying influence of wave splash, to describe the relationship between uninterrupted emersion time and the zonation patterns of two ecologically important species: the mussel Mytilus californianus and the barnacle Balanus glandula. At the local scale (10s to 100s of meters) on Tatoosh Island, the upper limits of both species are closely related to ESL, regardless of substrate aspect or maximum temperature. At larger spatial scales (10s to 100s of kilometers), the upper limit of Balanus is related to ESL at cool sites but not at hotter sites. Thus, although ESL explains most of the local-scale variation in zonation at a cool site, other factors (temperature, desiccation) likely become important as spatial scale increases to incorporate warmer sites. Our results emphasize that an understanding of where and when specific ecological factors are limiting is crucial for our ability to explain and predict large-scale biological patterns in space and time.

Light availability and consumer pressure are fundamental factors that structure aquatic communities, but their integrated effects are rarely studied in marine benthic systems. Using a factorial experiment, I investigated the effects of light availability and grazing by limpets (Lottia digitalis) to determine the relative influence of each on the growth and abundance of producers and consumers in a rocky intertidal community during the winter. Light reduction via shading reduced the abundance of filamentous algae and reduced the abundance of the herbivorous periwinkle Littorina sitkana. The effects of limpet grazing reduced the abundance of filamentous algae, Porphyra spp., and diatom mats. Higher limpet densities were also associated with significantly lower densities of adult, but not juvenile, Littorina spp. Light and limpet density interacted to determine Lottia growth, which was high in unshaded, single-limpet enclosures but was negligible in plots with two limpets, shades, or both. Variation in bottom-up (resource-driven) and top-down (consumer-driven) forces are important determinants of abundance at both trophic levels, but the effects are complex, taxon-specific, and, for littorine snails, size-specific. Because the study species accomplish most of their growth and reproduction during the winter, the interplay of light and herbivory during this season will likely influence long-term community dynamics.

Biological responses to climate change are typically communicated in generalized terms such as poleward and altitudinal range shifts, but adaptation efforts relevant to management decisions often require forecasts that incorporate the interaction of multiple climatic and nonclimatic stressors at far smaller spatiotemporal scales. We argue that the desire for generalizations has, ironically, contributed to the frequent conflation of weather with climate, even within the scientific community. As a result, current predictions of ecological responses to climate change, and the design of experiments to understand underlying mechanisms, are too often based on broad-scale trends and averages that at a proximate level may have very little to do with the vulnerability of organisms and ecosystems. The creation of biologically relevant metrics of environmental change that incorporate the physical mechanisms by which climate trains patterns of weather, coupled with knowledge of how organisms and ecosystems respond to these changes, can offer insight into which aspects of climate change may be most important to monitor and predict. This approach also has the potential to enhance our ability to communicate impacts of climate change to nonscientists and especially to stakeholders attempting to enact climate change adaptation policies.

The consequences of disturbance are known to be non-linear functions of disturbance intensity. In this study, we investigate the effects of a range of disturbance intensities by experimentally manipulating the cover of Fucus gardneri , a dominant intertidal alga. The effects of disturbance on subsequent Fucus recovery were similar for canopy reductions of 0–80%. However, complete removal of all Fucus individuals delayed Fucus recovery, as measured by canopy cover, by at least several months. Despite changes in the dominant alga, the abundance of invertebrates and other algae did not respond to cover manipulation. Our results suggest that the recovery from small-scale disturbances is enhanced if at least a few individuals survive the event. Only by examining a variety of levels of natural or anthropogenic disturbance will ecologists be able to understand observed recovery patterns and design management strategies.

Nutrients can structure communities by influencing both plant interactions and plant herbivore interactions, though rarely do studies integrate these processes. In this study we examined how nitrogen fertilization influenced (1) the positive interaction between the marsh elder, Iva frutescens, and the black rush, Juncusgerardi, and (2) the quality of Iva as a host plant for the aphid, Uroleuconambrosiae. Previous studies have shown that by mitigating soil salt accumulation and hypoxia, Juncus is essential to the survival of Iva and its aphid herbivore at mid-marsh elevations. To address the effects of nitrogen on this interaction, we compared fertilized and unfertilized Iva plants subject to Juncus removal and control treatments in the field. Additionally, we measured the monthly population growth rates of aphids transplanted onto these Iva plants. Iva leaf biomass and flower number results indicated that fertilizing Iva eliminated its dependence upon Juncus, such that fertilized plants grown without Juncus were not different from unmanipulated plants. Aphid monthly population growth rates through mid-summer revealed that fertilization also eliminated the indirect dependency of aphids on Juncus, so that aphid growth rates on fertilized Iva without Juncus neighbors were similar to rates on unmanipulated Iva. Results also indicated that fertilizing Iva grown with Juncus increased Iva size, potentially enabling these plants to support larger aphid populations. Our results suggest that only under conditions of nitrogen limitation are the positive effects of Juncus essential to the mid-marsh persistence of Iva and its aphid herbivore. Furthermore, we found that nitrogen effects on aphid populations may arise not only from a direct effect of nutrients on Iva size but also through the indirect effects of nitrogen on the interaction between Juncus and Iva. We argue that studies integrating processes occurring both within and between trophic levels, are important to fully understanding the community-wide effects of nutrients.

To predict community-level responses to climate change, we must understand how variation in environmental conditions drives changes in an organism's ability to acquire resources and translate those resources into growth, reproduction, and survival. This challenge can be approached mechanistically by establishing linkages from biophysics to community ecology. For example, body temperature can be predicted from environmental conditions and species-specific morphological and behavioral traits. Variation in body temperature within and among species dictates physiological performance, rates of resource acquisition, and growth. These ecological characteristics, along with population size, define the strength with which species interact. Finally, the direct (individual level) and indirect (community level) effects of temperature jointly determine community structure. This mechanistic framework can complement correlational approaches to better predict ecological responses to climate change and identify which characteristics of a species or community act as leverage points for change. Research priorities for further development of the mechanistic approach include documentation and prediction of relevant spatial and temporal variation in body temperature and the relationships between body temperature, individual performance, and interspecific interactions.

▪ Abstract The Exxon Valdez oil spill was the largest in US maritime history. We review post-spill research and set it in its legal context. The Exxon Corporation, obviously responsible for the spill, focused on restoration, whereas the Trustees, a coalition of state and federal entities, focused on damage and its assessment. Despite billions of dollars expended, little new understanding was gained about the recovery dynamics of a high latitude marine ecosystem subject to an anthropogenic pulse perturbation. We discuss a variety of case studies that highlight the limitations to and shortcomings of the research effort. Given that more spills are inevitable, we recommend that future studies address spatial patterns in the intertidal, and focus on the abundances of long-lived species and on organisms that preserve a chronological record of growth. Oil spills, while tragic, represent opportunities to gain insight into the dynamics of marine ecosystems and should not be wasted.

Most studies that forecast the ecological consequences of climate change target a single species and a single life stage. Depending on climatic impacts on other life stages and on interacting species, however, the results from simple experiments may not translate into accurate predictions of future ecological change. Research needs to move beyond simple experimental studies and environmental envelope projections for single species towards identifying where ecosystem change is likely to occur and the drivers for this change. For this to happen, we advocate research directions that (i) identify the critical species within the target ecosystem, and the life stage(s) most susceptible to changing conditions and (ii) the key interactions between these species and components of their broader ecosystem. A combined approach using macroecology, experimentally derived data and modelling that incorporates energy budgets in life cycle models may identify critical abiotic conditions that disproportionately alter important ecological processes under forecasted climates.

The global acidification of the earth's oceans is predicted to impact biodiversity via physiological effects impacting growth, survival, reproduction, and immunology, leading to changes in species abundances and global distributions. However, the degree to which these changes will play out critically depends on the evolutionary rate at which populations will respond to natural selection imposed by ocean acidification, which remains largely unquantified. Here we measure the potential for an evolutionary response to ocean acidification in larval development rate in two coastal invertebrates using a full-factorial breeding design. We show that the sea urchin species Strongylocentrotus franciscanus has vastly greater levels of phenotypic and genetic variation for larval size in future CO(2) conditions compared to the mussel species Mytilus trossulus. Using these measures we demonstrate that S. franciscanus may have faster evolutionary responses within 50 years of the onset of predicted year-2100 CO(2) conditions despite having lower population turnover rates. Our comparisons suggest that information on genetic variation, phenotypic variation, and key demographic parameters, may lend valuable insight into relative evolutionary potentials across a large number of species.

As CO2levels increase in the atmosphere, so too do they in the sea. Although direct effects of moderately elevated CO2in sea water may be of little consequence, indirect effects may be profound. For example, lowered pH and calcium carbonate saturation states may influence both deposition and dissolution rates of mineralized skeletons in many marine organisms. The relative impact of elevated CO2on deposition and dissolution rates are not known for many large-bodied organisms. We therefore tested the effects of increased CO2levels—those forecast to occur in roughly 100 and 200 years—on both shell deposition rate and shell dissolution rate in a rocky intertidal snail,Nucella lamellosa. Shell weight gain per day in live snails decreased linearly with increasing CO2levels. However, this trend was paralleled by shell weight loss per day in empty shells, suggesting that these declines in shell weight gain observed in live snails were due to increased dissolution of existing shell material, rather than reduced production of new shell material. Ocean acidification may therefore have a greater effect on shell dissolution than on shell deposition, at least in temperate marine molluscs.

This study investigates wasting disease in the northeast Pacific keystone predatory sea star Pisaster ochraceus on the outer west coast of Vancouver Island (British Columbia, Canada). To quantify the effects of temperature, season and locality on the vulnerability of P. ochraceus to wasting disease, we conducted surveys and experiments in early and late summer. To test the prediction that a small increase in temperature would result in heightened infection intensities, we housed sea stars at different temperatures in the laboratory and caged sea stars subtidally at 2 depths. Prevalence and infection intensity were always higher in warm temperature treatments and did not differ between the sexes or with increasing size. Disease effects also varied with season and locality. Specimens held in aquaria displayed significantly higher disease prevalence and infection intensity in June versus August. Furthermore, sea stars from a sheltered inlet showed markedly higher prevalence of the disease in late summer, while wave-exposed sites had consistently low disease prevalence. Seasonal changes in reproductive potential, host condition and/or physiological acclimation, as well as differences in environmental regime among localities, may impact the dynamics of wasting disease. These results demonstrate that small increases in temperature could drive mass mortalities of Pisaster due to wasting disease, with vulnerability possibly reaching a peak in spring and in populations from sheltered localities. This is the most northern report of wasting disease in the class Asteroidea on the west coast of North America.

Ecosystem engineering is an important process in a variety of ecosystems. However, the relationship between engineer density and engineering impact remains poorly understood. We used experiments and a mathematical model to examine the role of engineer density in a rocky intertidal community in northern California. In this system, the whelk Nucella ostrina preys on barnacles (Balanus glandula and Chthamalus dalli), leaving empty barnacle tests as a resource (favorable microhabitat) for other species. Field experiments demonstrated that N. ostrina predation increased the availability of empty tests of both barnacle species, reduced the density of the competitively dominant B. glandula, and indirectly increased the density of the competitively inferior C. dalli. Empty barnacle tests altered microhabitat humidity, but not temperature, and presumably provided a refuge from wave action. The herbivorous snail Littorina plena was positively associated with empty test availability in both observational comparisons and experimental manipulations of empty test availability, and L. plena density was elevated in areas with foraging N. ostrina. To explore the effects of variation in N. ostrina predation, we constructed a demographic matrix model for barnacles in which we varied predation intensity. The model predicted that number of available empty tests increases with predation intensity to a point, but declines when predation pressure was strong enough to severely reduce adult barnacle densities. The modeled number of available empty tests therefore peaked at an intermediate level of N. ostrina predation. Non-linear relationships between engineer density and engineer impact may be a generally important attribute of systems in which engineers influence the population dynamics of the species that they manipulate.

Ecological responses to climate change may occur gradually with changing conditions, or they may occur rapidly once some threshold or “tipping point” has been reached. Here, we use a high-resolution, 30-year data set on the upper vertical limit of a high intertidal alga to demonstrate that distributional shifts in this species do not keep pace with gradual trends in air temperature or sea level, but rather occur in sudden, discrete steps. These steps occur when unusually warm air temperatures are associated with unusually calm seas and are contingent in the sense that neither atmospheric nor sea conditions by themselves were sufficient to generate the underlying physiological challenge. Shifts in the upper limit did not correlate with large environmental perturbations such as El Niños; rather, they appeared to be associated with stochastic departures from otherwise gradual environmental trends. Our results exemplify the view that multiple environmental factors should be considered when attempting to understand ecological responses to climate change. Furthermore, punctuated responses such as those we have identified urge caution when attempting to infer causal mechanisms and project future distributional shifts using data of limited temporal resolution or scope.

Abiotic conditions in the oceans change naturally across a wide range of spatial and temporal scales. Recently, it has become abundantly clear that anthropogenic changes ranging from large-scale warming and ocean acidification to smaller-scale eutrophication and pollution are having important impacts on marine ecosystems as well. Whilst the relative importance of variation in abiotic factors depends upon the scale of interest, ecological responses are often determined by the combined influence of broader-scale (e.g. temperature and storm frequency) and local-scale interactions (e.g. catchment type×coastal morphology). Although this complexity continues to yield ecological surprises for conservation biologists and managers, continuing research on the importance of multiple and potentially interacting abiotic variables is helping to improve predictions regarding the ecological ramifications of human activities.

The sea star Pisaster ochraceus is one of the more striking species on the rocky shores of the Northeast Pacific, in part due to the dramatic color polymorphism of the adults. Along the open Pacific coast, Pisaster populations are 6%-28% orange, with a small percentage of brilliant purple stars and a large percentage of reddish-brown to dull purple ones. However, populations in the San Juan Island Archipelago (Washington, USA) and the southern Strait of Georgia (British Columbia, Canada) are almost entirely brilliant purple. The factors that maintain the color polymorphism, and those that contribute to among-site variation in color frequencies, remain unknown. We examined the relationships between color frequencies and several ecological and morphological variables, and conducted a large-scale phylogeographic survey of Pisaster populations. We found very low population genetic structure, suggesting that gene flow is high and geographic variation in color frequencies is not a vestige of Pleistocene glacial refugia. Color frequencies are also unrelated to adult size and to the frequency of injury within a population. However, there are suggestive relationships between color frequency and diet, and with areas of potentially low salinity. We propose that, although the color polymorphism may have an underlying genetic component, the regional-scale variation in color frequency is ecologically controlled.

Anthropogenically induced global climate change has profound implications for marine ecosystems and the economic and social systems that depend upon them. The relationship between temperature and individual performance is reasonably well understood, and much climate-related research has focused on potential shifts in distribution and abundance driven directly by temperature. However, recent work has revealed that both abiotic changes and biological responses in the ocean will be substantially more complex. For example, changes in ocean chemistry may be more important than changes in temperature for the performance and survival of many organisms. Ocean circulation, which drives larval transport, will also change, with important consequences for population dynamics. Furthermore, climatic impacts on one or a few 'leverage species' may result in sweeping community-level changes. Finally, synergistic effects between climate and other anthropogenic variables, particularly fishing pressure, will likely exacerbate climate-induced changes. Efforts to manage and conserve living marine systems in the face of climate change will require improvements to the existing predictive framework. Key directions for future research include identifying key demographic transitions that influence population dynamics, predicting changes in the community-level impacts of ecologically dominant species, incorporating populations' ability to evolve (adapt), and understanding the scales over which climate will change and living systems will respond.

SUMMARYLiving at the interface between the marine and terrestrial environments,intertidal organisms may serve as a bellwether for environmental change and a test of our ability to predict its biological consequences. However, current models do not allow us to predict the body temperature of intertidal organisms whose heat budgets are strongly affected by conduction to and from the substratum. Here, we propose a simple heat-budget model of one such animal,the limpet Lottia gigantea, and test the model against measurements made in the field. Working solely from easily measured physical and meteorological inputs, the model predicts the daily maximal body temperatures of live limpets within a fraction of a degree, suggesting that it may be a useful tool for exploring the thermal biology of limpets and for predicting effects of climate change. The model can easily be adapted to predict the temperatures of chitons, acorn barnacles, keyhole limpets, and encrusting animals and plants.

SUMMARYWhen coupled with long-term meteorological records, a heat-budget model for the limpet, Lottia gigantea, provides a wealth of information regarding environmental and topographic controls of body temperature in this ecologically important species. (1) The maximum body temperature predicted for any site (37.5°C) is insufficient to kill all limpets, suggesting that acute thermal stress does not set an absolute upper limit to the elevation of L. gigantea on the shore. Therefore, the upper limit must be set by behavioral responses, sublethal effects or ecological interactions. (2)Temperatures sufficient to kill limpets are reached at only a small fraction of substratum orientations and elevations and on only three occasions in 5 years. These rare predicted lethal temperatures could easily be missed in field measurements, thereby influencing the interpretation of thermal stress.(3) Body temperature is typically higher than air temperature, but maximum air temperature can nonetheless be used as an accurate predictor of maximum body temperature. Warmer air temperatures in the future may thus cause increased mortality in this intertidal species. Interpretation of the ecological effects of elevated body temperature depends strongly on laboratory measurements of thermal stress, highlighting the need for additional research on the temporal and spatial variability of thermal limits and sublethal stress. The lengthy time series of body temperatures calculated from the heat-budget model provides insight into how these physiological measurements should be conducted.

The interaction of climate and the timing of low tides along the West Coast of the United States creates a complex mosaic of thermal environments, in which northern sites can be more thermally stressful than southern sites. Thus, climate change may not lead to a poleward shift in the distribution of intertidal organisms, as has been proposed, but instead will likely cause localized extinctions at a series of “hot spots.” Patterns of exposure to extreme climatic conditions are temporally variable, and tidal predictions suggest that in the next 3 to 5 years “hot spots” are likely to appear at several northern sites.

Increases in the frequency, severity and duration of temperature extremes are anticipated in the near future. Although recent work suggests that changes in temperature variation will have disproportionately greater effects on species than changes to the mean, much of climate change research in ecology has focused on the impacts of mean temperature change. Here, we couple fine-grained climate projections (2050–2059) to thermal performance data from 38 ectothermic invertebrate species and contrast projections with those of a simple model. We show that projections based on mean temperature change alone differ substantially from those incorporating changes to the variation, and to the mean and variation in concert. Although most species show increases in performance at greater mean temperatures, the effect of mean and variance change together yields a range of responses, with temperate species at greatest risk of performance declines. Our work highlights the importance of using fine-grained temporal data to incorporate the full extent of temperature variation when assessing and projecting performance.

Sea otters (Enhydra lutris) are keystone predators that consume a variety of benthic invertebrates, including the intertidal mussel, Mytilus californianus. By virtue of their competitive dominance, large size, and longevity, M. californianus are ecosystem engineers that form structurally complex beds that provide habitat for diverse invertebrate communities. We investigated whether otters affect mussel bed characteristics (i.e. mussel length distributions, mussel bed depth, and biomass) and associated community structure (i.e. biomass, alpha and beta diversity) by comparing four regions that varied in their histories of sea otter occupancy on the west coast of British Columbia and northern Washington. Mussel bed depth and average mussel lengths were 1.5 times lower in regions occupied by otters for >20 years than those occupied for

Because intertidal organisms live close to their environmental tolerances, intertidal species are considered early harbingers of the effects of global change. Using a combination of spatial environmental gradients and long-term datasets, I examine the likely consequences of global warming on the distribution of Mazzaella parksii (formerly Mazzaella cornucopiae), a turf-forming red alga. On Tatoosh Island, WA, M. parksii is restricted almost entirely to north-facing surfaces. On south-facing slopes, the upper limit of M. parksii, which is set by abiotic stress, occurs much lower on the shore. However, its lower limit, which is set by herbivores, does not vary with aspect. Because its upper and lower limits are independent across the aspect gradient, M. parksii is effectively “squeezed out” of the system as one moves from a north-facing to a south-facing slope. This general pattern is observed at several spatial scales across gradients of wave exposure and air temperature. The vertical distributional patterns of M. parksii also shift across temporal gradients. Over the past 25 years, the upper limit of M. parksii on Tatoosh Island has shifted downshore by approximately 25 cm. Most of this change was driven by three consecutive harsh summers in the mid 1990's. These temporal patterns, combined with the insight gained from spatial patterns, suggest that M. parksii may go locally extinct in many parts of its range as its upper and lower limits converge. Furthermore, because vertical distributional shifts are linked to stochastic climatic effects, temporal change in marine algal populations may be rapid and unexpected.

Anthropogenic carbon dioxide (CO2) emissions and the resultant acidification of surface ocean waters are predicted to have far-reaching consequences for biological processes in the marine environment. For example, because changes in pH and pCO2 can alter sperm performance, ocean acidification may be accompanied by reductions in the success of fertilization in marine broadcast spawners. Several studies have attempted to determine the effects of elevated pCO2 on marine invertebrate fertilization success, albeit with differing results. These conflicts may stem from the use of inappropriate sperm–egg contact times and, in several cases, the lack of measurements over a range of sperm concentrations extending from sperm-limited conditions to polyspermy scenarios. In our study, we used biologically realistic sperm–egg contact times and a full range of sperm concentrations to assess the effect of elevated pCO2 on fertilization in the broadcast spawning sea urchin, Strongylocentrotus franciscanus. Fertilization experiments were carried out in seawater bubbled with CO2 to 400 (control), 800, and 1800 ppm. Using a fertilization kinetics model, we estimate that elevated pCO2 levels both increased sperm limitation and reduced the efficiency of fast blocks to polyspermy. Thus, elevated pCO2 decreased the range of sperm concentrations over which high fertilization success was likely. Given the inherent difficulties in achieving high fertilization success in broadcast spawners, raised pCO2 levels are likely to exacerbate low fertilization success in low-density populations or in areas with high water turbulence.

1. Crowding is a common ecological phenomenon that has been widely studied from the perspective of competition for limiting resources. The extent to which crowded plants and sessile animals are positively affected by their neighbours, however, has received much less attention. Here, we use four salt-marsh plant species to examine the hypothesis that crowding leads to morphologically modified individuals that are dependent on their neighbours for structural support. 2. In the spring, crowded and isolated treatments were created for the four species (Juncus gerardi, Spartina alterniflora, Ivafrutescens and Salicornia europaea). By the end of the growing season, crowded plants (a) were generally taller, thinner and produced less above-ground biomass, (b) were more susceptible to breaking and (c) grew closer to their theoretical maximum heights. 3. To explicitly test the hypothesis that crowding leads to plants that are dependent on their neighbours for support, we thinned crowded plant stands at the end of the growing season and compared the survivorship of these secondarily isolated plants with that of permanently isolated plants. For all four species, a significant number of previously crowded plants fell over, whereas plants grown in isolation remained upright. 4. Our results show that crowded plants often develop morphologies that cause them to be structurally dependent on their neighbours for support. Thus, the continuing presence of neighbours provides a positive benefit that may mitigate the negative effects of crowding. Our findings contribute to a growing body of evidence that positive interactions among crowded plants and sessile animals are a more pervasive feature of natural assemblages than is generally acknowledged.