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Heatwave frequency and intensity will increase as climate change progresses. Intertidal sessile invertebrates, which often form thermally benign microhabitats for associated species, are vulnerable to thermal stress because they have minimal ability to behaviourally thermoregulate. Understanding what factors influence the mortality of biogenic species and how heatwaves might impact their ability to provide habitat is critical. Here, we characterize the community associated with the thatched barnacle, Semibalanus cariosus (Pallass, 1788), in British Columbia (BC), Canada. Then, we investigate what site-level and plot-level environmental factors explained variations in barnacle mortality resulting from an unprecedented regional heatwave in BC, Canada. Furthermore, we used a manipulative shading experiment deployed prior to the heatwave to examine the effect of thermal stress on barnacle survival and recruitment and the barnacle-associated community. We identified 50 taxa inhabiting S. cariosus beds, with variations in community composition between sites. Site-scale variables and algal canopy cover did not predict S. cariosus mortality, but patch-scale variation in substratum orientation did, with more direct solar irradiance corresponding with higher barnacle mortality. The shading experiment demonstrated that S. cariosus survival, barnacle recruitment, and invertebrate community diversity were higher under shades where substratum temperatures were lower. Associated community composition also differed between shaded and non-shaded plots, suggesting S. cariosus was not able to fully buffer acute thermal stress for its associated community. While habitat provisioning by intertidal foundation species is an important source of biodiversity, these species alone may not be enough to prevent substantial community shifts following extreme heatwaves. As heatwaves become more frequent and severe, they may further reduce diversity via the loss of biogenic habitat, and spatial variation in these impacts may be substantial.

In late June 2021 a heatwave of unprecedented magnitude impacted the Pacific Northwest (PNW) region of Canada and the United States. Many locations broke all-time maximum temperature records by more than 5°C, and the Canadian national temperature record was broken by 4.6°C, with the highest recorded temperature 49.6°C. Local records were broken by large margins, even when compared to local records broken during the infamous heatwaves in Europe 2003, and Russian in 2010. A region of high pressure that became stationary over the region (an atmospheric block) was the dominant cause of this heatwave; however, trajectory analysis finds that upstream diabatic heating played a key role in the magnitude of the temperature anomalies. Weather forecasts provided advanced notice of the event, while sub-seasonal forecasts showed an increased likelihood of a heat extreme with 10-20 day lead times, with an increased likelihood of a blocking event seen in forecasts initialized 3 weeks prior to the heatwave peak. The impacts of this event were catastrophic. We provide a summary of some of these impacts, including estimates of hundreds of attributable deaths across the PNW, mass-mortalities of marine life, reduced crop and fruit yields, river flooding from rapid snow and glacier melt, and a substantial increase in wildfires—the latter contributing to devastating landslides in the months following. These impacts provide examples we can learn from, and a vivid depiction of how climate change can be so devastating.

Marine heatwaves (MHWs), discrete but prolonged periods of anomalously warm seawater, can fundamentally restructure marine communities and ecosystems. Although our understanding of these events has improved in recent years, key knowledge gaps hinder our ability to predict how MHWs will affect patterns of biodiversity. Here, we outline a functional trait approach that enables a better understanding of which species and communities will be most vulnerable to MHWs, and how the distribution of species and composition of communities are likely to shift through time. Our perspective allows progress toward unifying extreme events and longer term environmental trends as co-drivers of ecological change, with the incorporation of species traits into our predictions allowing for a greater capacity to make management decisions.

Vessel hull-fouling is responsible for most bioinvasion events in the marine environment, yet it lacks regulation in most countries. Although experts advocate a preventative approach, research efforts on pre-arrival processes are limited. The performance of mobile epifauna during vessel transport was evaluated

Small-scale spatial variation in temperature is an important attribute of many habitats, as spatial thermal refugia may allow organisms to avoid some negative consequences of thermal extremes. Although the ecological importance of thermal refugia in intertidal habitats is well-known in summer, the relative availability, effectiveness, and ecological relevance of thermally benign microhabitats in the winter is poorly understood. Here, we explored small-scale thermal variability on a temperate rocky coastline during winter, and the relevance of these thermal patterns to Littorina scutulata, a common intertidal gastropod. Sheltered microhabitats, including crevices, undersides of boulders, and areas beneath algal canopy, were all warmer than surrounding exposed areas during winter low tides. L. scutulata was able to recover relatively quickly from exposure to low temperatures down to -3°C, but performance costs began to accrue at lower temperatures. However, the presence of a refuge habitat buffered the negative effects of cold temperatures, and sheltered L. scutulata regained mobility more quickly than conspecifics on exposed surfaces. Further experiments demonstrated that L. scutulata were more frequently found in thermal refugia when previously exposed to low temperatures, but not when exposed to milder temperatures. Our results suggest that realistic low temperatures can impose important costs on intertidal species such as L. scutulata, but the availability of thermal refugia coupled with behavioural adaptations can allow species to reduce many such costs. Just as in summer, thermal refugia in winter may be important for the distribution and abundance of intertidal species and the diversity and function of coastal ecosystems.

The abiotic environment varies continuously at a variety of temporal scales. While this variation is known to be ecologically important, multiple scales of variability are rarely explicitly considered in ecological studies. Here, we combine field observations and laboratory experiments to determine the individual and population level effects of short-term (tidal) and longer-term (seasonal and interannual) salinity variation on the dogwhelkNucella lamellosain the Strait of Georgia, British Columbia, Canada. The Fraser River heavily influences surface salinity in the Strait of Georgia, which varies with season, depth, and distance to the river mouth. At low salinity sites,N. lamellosapopulation size decreased following high outflow years, with fewer juveniles present, as opposed to high salinity sites, which had higher population densities in all years. Sustained salinity exposure in the laboratory caused developmental delay of encapsulated embryos and complete mortality at 9 and 12 psu. Juvenile dogwhelks (N. lamellosato survive otherwise low salinity conditions for considerably longer. Overall, our results suggest that seasonal and interannual variation in salinity have a profound influence onN. lamellosapopulations and that shorter-scale fluctuations can moderate these seasonal and interannual effects. It is likely that similar multi-scale environmental effects will determine survival and population dynamics in many species.

In late June to early July 2021 a heatwave of unprecedented magnitude impacted the Pacific Northwest region, lands colonially named British Columbia (BC) and Alberta (AB) in Canada, and Washington (WA) and Oregon (OR) in the United States. Many locations broke all-time maximum daily temperature records by more than 5°C. The standing Canadian national temperature record was broken on three consecutive days, at multiple locations, with the highest temperature of 49.6°C recorded, 4.6°C higher than the previous Canadian record. Weather forecasts provided advanced notice of the severity of the event, while sub-seasonal forecasts showed substantially increased likelihood of atmospheric blocking and high temperatures with 10-day lead times, and some skill out to 18 days. The impacts of this event were catastrophic. Estimates indicate at least 900 attributable deaths occurred across BC, WA and OR, likely the deadliest weather event in Canadian1 history. The heat contributed to mass-mortalities of marine life, reduced crop and fruit yields, river flooding from rapid snow and glacier melt, and a rapid increase in wildfires—the latter contributing to devastating landslides in the months following.

Ongoing climate change has caused rapidly increasing temperatures, and an unprecedented decline in seawater pH, known as ocean acidification. Increasing temperatures are redistributing species towards higher and cooler latitudes which are most affected by ocean acidification. Whilst the persistence of intertidal species in cold environments is related to their capacity to resist sub-zero air temperatures, studies have never considered the interacting impacts of ocean acidification and freeze stress on species survival and distribution. A full-factorial experiment was used to study whether ocean acidification increases mortality in Mytilus spp. following sub-zero air temperature exposure. We examined physiological processes behind variation in freeze tolerance using 1H NMR metabolomics, analyses of fatty acids, and amino acid composition. We show that low pH conditions (pH = 7.5) significantly decrease freeze tolerance in both intertidal and subtidal populations of Mytilus spp. Under current day pH conditions (pH = 7.9), intertidal M. trossulus were more freeze tolerant than subtidal M. trossulus and M. galloprovincialis. Opposite, under low pH conditions, subtidal M. trossulus was more freeze tolerant than the other groups. We observed a marked shift from negative to positive metabolite-metabolite correlations across species under low pH conditions, but there was no evidence that the concentration of individual metabolites or amino acids affected freeze tolerance. Finally, pH-induced changes in the composition of cell membrane phospholipid fatty acids had no effect on survival. These results suggest that ocean acidification can offset the poleward expanding facilitated by warming, and that reduced freeze tolerance could result in a niche squeeze if temperatures become lethal at the equatorward edge.

Predicted shifts in mean and extreme temperatures associated with climate change can have variable impacts on organisms, and the sign and magnitude of these impacts may depend upon local context. For Hedophyllum sessile, a habitat-forming intertidal kelp, the impacts of warming may vary with local density and position in the intertidal zone. To assess the potential context-dependence of warming, we manipulated H. sessile densities across an intertidal gradient and experimentally imposed periodic thermal stress in the field. The recruitment of H. sessile juveniles was unimodally related to shore level, peaking near the center of the species’ vertical distribution and falling off at the upper and lower distributional limits. Experimental warming tended to have mildly positive effects on recruitment lower on the shore regardless of adult density, and in upper zone, high density plots. However, warming had strongly negative effects on recruitment in upper zone, low density plots. Temperature manipulations also had context-specific effects on adult plant growth; seasonal increases in blade number and canopy cover were slightly enhanced by warming in high-density plots but greatly reduced by warming in low-density plots. Finally, experimental heating had context-dependent effects on an understory herbivore, the chiton Katharina tunicata, which increased in abundance following heating in high density plots but decreased in low density plots. Our results demonstrate that extreme temperature events can affect multiple species and multiple life history stages, and that the impacts of such events can depend upon both environmental (e.g. intertidal height) and biological (e.g. adult density) context.

Invasive species often exhibit disproportionately strong negative effects in their introduced range compared to their native range, and much research has been devoted to understanding the role of shared evolutionary history, or lack thereof, in driving these differences. Less studied is whether introduced species, particularly those that are important as facilitators in their native range, have persistent positive effects in their invaded range despite a lack of a shared evolutionary history with the invaded community. Here, we manipulated the density of a habitat-forming facilitator, the high intertidal acorn barnacle Balanus glandula, factorially with herbivore density in its native range (Bluestone Point, British Columbia, Canada) and invaded range (Punta Ameghino, Chubut Province, Argentina) to determine how this facilitator differentially affects associated species at these two locations. Given that high intertidal species at Punta Ameghino (PA) are evolutionarily naive to barnacles, we predicted that the positive effects of B. glandula at PA would be absent or weak compared to those at Bluestone Point (BP). However, we found that B. glandula had an equally positive effect on herbivore biomass at PA compared to BP, possibly because the moisture-retaining properties of barnacle bed habitats are particularly important in seasonally dry Patagonia. Barnacle presence indirectly decreased ephemeral algal cover at BP by increasing grazer pressure, but barnacles instead facilitated ephemeral algae at PA. In contrast, B. glandula increased perennial algal cover at BP, but generally decreased perennial algal cover at PA, likely due to differences in dominant algal morphology. Though our experiment was limited to one location on each continent, our results suggest that shared evolutionary history may not be a prerequisite for strong facilitation to occur, but rather that the nature and strength of novel species interactions are determined by the traits of associated species and the environment in which they occur.

Understanding and predicting responses of ectothermic animals to temperature are essential for decision-making and management. The thermal performance curve (TPC), which quantifies the thermal sensitivity of traits such as metabolism, growth and feeding rates in laboratory conditions, is often used to predict responses of wild populations. However, central assumptions of this approach are that TPCs are relatively static between populations and that curves measured under stable temperature conditions can predict performance under variable conditions. We test these assumptions using two latitudinally matched populations of the ecosystem engineer Mytilus trossulus that differ in their experienced temperature variability regime. We acclimated each population in a range of constant or fluctuating temperatures for six weeks and measured a series of both short term (feeding rate, byssal thread production) and long-term (growth, survival) metrics to test the hypothesis that performance in fluctuating temperatures can be predicted from constant temperatures. We find that this was not true for any metric, and that there were important interactions with the population of origin. Our results emphasize that responses to fluctuating conditions are still poorly understood and suggest caution must be taken in the use of TPCs generated under constant temperature conditions for the prediction of wild population responses.

Sustainably managed wild fisheries support food and nutritional security, livelihoods, and cultures (1). Harmful fisheries subsidies—government payments that incentivize overcapacity and lead to overfishing—undermine these benefits yet are increasing globally (2). World Trade Organization (WTO) members have a unique opportunity at their ministerial meeting in November to reach an agreement that eliminates harmful subsidies (3). We—a group of scientists spanning 46 countries and 6 continents—urge the WTO to make this commitment...

Freezing is an extreme stress to living cells, and so freeze-tolerant animals often accumulate protective molecules (termed cryoprotectants) to prevent the cellular damage caused by freezing. The bay mussel

Extreme environments have driven the evolution of some of the most inspiring adaptations in nature. In the intertidal zone of wave-swept shores, organisms face physical forces comparable to hurricanes and must further endure thermal and desiccation stress during low tides, compromising their physiological and biomechanical performance. We examine how these multiple stressors have influenced the evolution of tissue properties during desiccation using eight phylogenetically independent pairs of intertidal and subtidal macrophytes. Intertidal species generally lost water more slowly than their subtidal counterparts, presumably as an adaption to regular emersion. Under partial desiccation, breaking force, strength, and extensibility of intertidal species generally exceeded those of subtidal species, although important differences existed among phylogenetic pairs. This was often true even when subtidal relatives resisted greater forces or were more extensible under full hydration. The interacting effects of mechanical forces and desiccation during low tide are likely a major selective agent in determining macrophyte performance and fitness. Overall, we found that lineages that have independently evolved to occupy the wave-swept intertidal have converged on performance metrics that are likely to be adaptive to the interacting stressors associated with their extreme niches.

Ocean acidification (OA) driven by anthropogenic CO 2 emissions affects marine ecosystems, fisheries and aquaculture. Assessing the impacts of OA using projection models facilitates the development of future scenarios and potential solutions. Here, we explored various ways to incorporate OA impacts into a multi-stressor dynamic bioclimatic envelope model to project biogeographic changes of ten commercially exploited invertebrate species. We examine three dimensions of uncertainties in modelling biophysical OA effects: model structure, parameterization, and scenario uncertainty. Our results show that projected OA impacts are most sensitive to the choice of structural relationship between OA and biological responses, followed by the choice of climate change emission scenarios and parameterizations of the size of OA effects. Species generally showed negative effects to OA but sensitivity to the various sources of uncertainty were not consistent across or within species. For example, some species showed higher sensitivity to structural uncertainty and very low sensitivity to parameter uncertainty, while others showed greatest sensitivity to parameter uncertainty. This variability is largely due to geographic variability and difference in life history traits used to parameterize model simulations. Our model highlights the variability across the sources of uncertainty and contributes to the development of integrating OA impacts in species distribution models. We further stress the importance of defining the limitations and assumptions, as well as exploring the range of uncertainties associated with modelling OA impacts.

Ecologically dominant species often define ecosystem states, but as human disturbances intensify, their subordinate counterparts increasingly displace them. We consider the duality of disturbance by examining how environmental drivers can simultaneously act as a stressor to dominant species and as a resource to subordinates. Using a model ecosystem, we demonstrate that CO2-driven interactions between species can account for such reversals in dominance; i.e., the displacement of dominants (kelp forests) by subordinates (turf algae). We established that CO2 enrichment had a direct positive effect on productivity of turfs, but a negligible effect on kelp. CO2 enrichment further suppressed the abundance and feeding rate of the primary grazer of turfs (sea urchins), but had an opposite effect on the minor grazer (gastropods). Thus, boosted production of subordinate producers, exacerbated by a net reduction in its consumption by primary grazers, accounts for community change (i.e., turf displacing kelp). Ecosystem collapse, therefore, is more likely when resource enrichment alters competitive dominance of producers, and consumers fail to compensate. By recognizing such duality in the responses of interacting species to disturbance, which may stabilize or exacerbate change, we can begin to understand how intensifying human disturbances determine whether or not ecosystems undergo phase shifts. Refereed/Peer-reviewed

Connecting the nonlinear and often counterintuitive physiological effects of multiple environmental drivers to the emergent impacts on ecosystems is a fundamental challenge. Unfortunately, the disconnect between the way “stressors” (e.g., warming) is considered in organismal (physiological) and ecological (community) contexts continues to hamper progress. Environmental drivers typically elicit biphasic physiological responses, where performance declines at levels above and below some optimum. It is also well understood that species exhibit highly variable response surfaces to these changes so that the optimum level of any environmental driver can vary among interacting species. Thus, species interactions are unlikely to go unaltered under environmental change. However, while these nonlinear, species-specific physiological relationships between environment and performance appear to be general, rarely are they incorporated into predictions of ecological tipping points. Instead, most ecosystem-level studies focus on varying levels of “stress” and frequently assume that any deviation from “normal” environmental conditions has similar effects, albeit with different magnitudes, on all of the species within a community. We consider a framework that realigns the positive and negative physiological effects of changes in climatic and nonclimatic drivers with indirect ecological responses. Using a series of simple models based on direct physiological responses to temperature and ocean pCO2, we explore how variation in environment-performance relationships among primary producers and consumers translates into community-level effects via trophic interactions. These models show that even in the absence of direct mortality, mismatched responses resulting from often subtle changes in the physical environment can lead to substantial ecosystem-level change. Refereed/Peer-reviewed

The cosmopolitan nonindigenous sea anemone, Diadumene lineata, has been recorded from the Pacific coast of Canada since 1859. Despite its long historical presence in the region, the factors influencing the distribution of this species in British Columbia, Canada, are poorly understood, as are the factors that may restrict its local abundance. We conducted large‐scale surveys of D. lineata during the winter and summer of 2016, coupled with higher frequency surveys at one focal site. Furthermore, we used laboratory and field experiments to determine the potential importance of a large‐scale driver, salinity, and a small‐scale driver, microhabitat within boulders, in setting distributional limits at various spatial scales. Our results confirm that D. lineata is widely distributed within the Salish Sea. This broad distribution may be facilitated by post‐introduction dispersal; we observed anemones attached to floating substrata, and a few mature specimens contained eggs, suggesting that reproduction is perhaps not limited to asexual fission. We found that D. lineata was abundant throughout the year, although absent from the lowest salinity sites. This corresponded with our laboratory experiments, in which survival of individuals of D. lineata declined at salinities below 10 psu, and no survivors were observed at or below 2 psu. Manipulative experiments in the field demonstrated that survival was high under rocks where D. lineata is commonly found, but was reduced on the tops of rocks, highlighting the importance of microhabitat refugia in buffering temperature and reducing desiccation. A better understanding of the factors that limit the broad‐scale and local distribution of this anemone are an important step in determining potential ecological impacts of this organism and how distributions may shift into the future.

Human activities have caused an increase in atmospheric CO2 over the last 250 years, leading to unprecedented rates of change in seawater pH and temperature. These global scale processes are now commonly referred to as ocean acidification and warming, and have the potential to substantially alter the physiological performance of many marine organisms. It is vital that the effects of ocean acidification and warming on marine organisms are explored so that we can predict how marine communities may change in future. In particular, the effect of ocean acidification and warming on host-parasite dynamics is poorly understood, despite the ecological importance of these relationships. Here, we explore the response of one himasthlid trematode, Himasthla sp., an abundant and broadly distributed species of marine parasite, to combinations of elevated temperature and pCO2 that represent physiological extremes, pre-industrial conditions, and end of century predictions. Specifically, we quantified the life span of the free-living cercarial stage under elevated temperature and pCO2, focussing our research on functional life span (the time cercariae spend actively swimming) and absolute life span (the period before death). We found that the effects of temperature and pCO2 were complex and interactive. Overall, increased temperature negatively affected functional and absolute life span, e.g. across all pCO2 treatments the average time to 50% cessation of active swimming was approximately 8 h at 5 °C, 6 h at 15 °C, 4 h at 25 °C, and 2 h at 40 °C. The effect of pCO2, which significantly affected absolute life span, was highly variable across temperature treatments. These results strongly suggest that ocean acidification and warming may alter the transmission success of trematode cercariae, and potentially reduce the input of cercariae to marine zooplankton. Either outcome could substantially alter the community structure of coastal marine systems.

Thermal performance curves (TPCs), which quantify how an ectotherm's body temperature (T-b) affects its performance or fitness, are often used in an attempt to predict organismal responses to climate change. Here, we examine the key - but often biologically unreasonable - assumptions underlying this approach; for example, that physiology and thermal regimes are invariant over ontogeny, space and time, and also that TPCs are independent of previously experienced T-b. We show how a critical consideration of these assumptions can lead to biologically useful hypotheses and experimental designs. For example, rather than assuming that TPCs are fixed during ontogeny, one can measure TPCs for each major life stage and incorporate these into stage-specific ecological models to reveal the life stage most likely to be vulnerable to climate change. Our overall goal is to explicitly examine the assumptions underlying the integration of TPCs with T-b, to develop a framework within which empiricists can place their work within these limitations, and to facilitate the application of thermal physiology to understanding the biological implications of climate change.

Summary Increasing levels of CO2 in the atmosphere are affecting ocean chemistry, leading to increased acidification (i.e. decreased pH) and reductions in calcium carbonate saturation state. Many species are likely to respond to acidification, but the direction and magnitude of these responses will be based on interspecific and ontogenetic variation in physiology and the relative importance of calcification. Differential responses to ocean acidification (OA) among species will likely result in important changes in community structure and diversity. To characterize the potential impacts of OA on community composition and structure, we examined the response of a marine fouling community to experimental CO2 enrichment in field-deployed flow-through mesocosm systems. Acidification significantly altered the community structure by altering the relative abundance of species and reduced community variability, resulting in more homogenous biofouling communities from one experimental tile to the next both among and within the acidified mesocosms. Mussel (Mytilus trossulus) recruitment was reduced by over 30% in the elevated CO2 treatment compared to the ambient treatment by the end of the experiment. Strong differences in mussel cover (up to 40% lower in acidified conditions) developed over the second half of the 10-week experiment. Acidification did not appear to affect the mussel growth, as average mussel sizes were similar between treatments at the end of the experiment. Hydroid (Obelia dichotoma) cover was significantly reduced in the elevated CO2 treatment after 8 weeks. Conversely, the percentage cover of bryozoan colonies (Mebranipora membranacea) was higher under acidified conditions with differences becoming apparent after 6 weeks. Neither recruitment nor final size of barnacles (Balanus crenatus) was affected by acidification. By the end of the experiment, diversity was 41% lower in the acidified treatment relative to ambient conditions. Overall, our findings support the general expectation that OA will simplify marine communities by acting on important ecological processes that ultimately determine the community structure and diversity.

Fifty years ago, Robert Treat Paine published his seminal paper on food web complexity and biodiversity (Paine 1966). In this work, Paine used sea stars to illustrate how relationships among species were key determinants of emergent ecological patterns, and in so doing he injected a healthy dose of testable mechanism into a field that was largely phenomenological at the time. The example of the sea star Pisaster ochraceus as a keystone predator (Paine 1966, 1969) remains a standard example in Ecology textbooks, and undergraduates who take ecology courses today are likely to associate the name Paine with starfish. This article is protected by copyright. All rights reserved.

Kelp systems dominate nearshore marine environments in upwelling zones characterized by cold temperatures and high nutrients. Worldwide, kelp population persistence and recruitment success generally decreases with rising water temperatures coupled with low nutrients, making kelp populations vulnerable to impending warming of the oceans. This response to climate change at a global scale, however, may vary due to regional differences in temperature variability, acclimation, and differential responses of kelp species to changing conditions. Culture experiments were conducted on 12 eastern Pacific kelp taxa across geographic regions (British Columbia, central California, and southern California) under three nitrate levels (1, 5, and 10 μmol/L) and two temperatures (12°C and 18°C) to determine sporophyte production (i.e., recruitment success). For all taxa from all locations, sporophytes were always present in the 12°C treatment and when recruitment failure was observed, it always occurred at 18°C, regardless of nitrate level, indicating that temperature is the driving factor limiting recruitment, not nitrate. Rising ocean temperatures will undoubtedly cause recruitment failure for many kelp species; however, the ability of species to acclimatize or adapt to increased temperatures at the warmer edge of their species range may promote a resiliency of kelp systems to climate change at a global scale.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has been fixed in the paper.

Because many of the negative effects of ocean acidification on marine life may result from underlying energetic short-falls associated with increased metabolic demands, several studies have hypothesized that negative responses to high CO2 could be reduced by energy input. Although this hypothesis was supported by a recent meta-analysis, we believe that the meta-analytic calculation used was not appropriate to test the stated hypothesis. Here, we first clarify the hypothesis put forward, the crux being that the effects of increased food supply and CO2 interact statistically. We then test this hypothesis by examining the available data in a more appropriate analytical framework. Using factorial meta-analysis, we confirm that food addition has a positive effect and CO2 has a negative effect on both growth and calcification. For calcification, food addition did indeed reduce CO2 impacts. Surprisingly, however, we found that food addition actually exacerbated the effects of acidification on growth, perhaps due to increased scope upon which CO2 effects can act in food-replete situations. These interactive effects were undetectable using a multilevel meta-analytic approach. Ongoing changes in food supply and carbonate chemistry, coupled with under-described, poorly understood, and potentially surprising interactive outcomes for these two variables, suggest that the role of food should remain a priority in ocean acidification research.Arising from: L. Ramajo et al., Sci. Rep. 6: 19374 (2016).

Trophic cascades are indirect positive effects of predators on resources via control of intermediate consumers. Larger-bodied predators appear to induce stronger trophic cascades (a greater rebound of resource density toward carrying capacity), but how this happens is unknown because we lack a clear depiction of how the strength of trophic cascades is determined. Using consumer resource models, we first show that the strength of a trophic cascade has an upper limit set by the interaction strength between the basal trophic group and its consumer and that this limit is approached as the interaction strength between the consumer and its predator increases. We then express the strength of a trophic cascade explicitly in terms of predator body size and use two independent parameter sets to calculate how the strength of a trophic cascade depends on predator size. Both parameter sets predict a positive effect of predator size on the strength of a trophic cascade, driven mostly by the body size dependence of the interaction strength between the first two trophic levels. Our results support previous empirical findings and suggest that the loss of larger predators will have greater consequences on trophic control and biomass structure in food webs than the loss of smaller predators.

As the climate warms, there is little doubt that ecosystems of the future will look diff erent from those we see today. However, community responses to warming in the fi eld are poorly understood. We examined the eff ects of fi eld-based warming on intertidal communities in the Salish Sea, which is a regional thermal ‘ hot spot ’ and therefore a model system for studying thermally stressed communities. We manipulated temperature at three tidal heights by deploying black- and white-bordered settlement plates. Black plates increased in situ substratum temperature by an average of 2.6 ° C (maximum temperature, 40.9 ° C). Barnacles fared poorly on black plates in all zones. When overall thermal stress was highest (summer in the high intertidal zone) herbivores were absent. In lower tidal zones, herbivores were abundant on white plates but were scarce on black plates. Th e total percent cover of algae was unaff ected by the temperature treatment, despite the fact that macroalgae were expected to be the least thermally tolerant functional group. However, we did fi nd that ephemeral green algae exhibited a delay in phenology on black plates. We also found that species richness declined and invertebrate assemblage structure was altered due to warming. Results from this year long experiment suggest that communities in thermally stressful habitats respond to warming via the interplay between species-specifi c thermal responses and secondary adaptive strategies such as behavioral microhabitat selection. Declines in diversity and changes in the invertebrate assemblage were due to the decline of local thermally-stressed species and the lack of replacement by warm-adapted species. Given the low variation in the species pool along the northeast Pacifi c coastline, the arrival of warm-adapted species to the Salish Sea may not occur over relevant time scales, leaving local communities depauperate.

© This is an open access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication. Disease outbreaks can have substantial impacts on wild populations, but the often patchy or anecdotal evidence of these impacts impedes our ability to understand outbreak dynamics. Recently however, a severe disease outbreak occurred in a group of very well-studied organisms–sea stars along the west coast of North America. We analyzed nearly two decades of data from a coordinated monitoring effort at 88 sites ranging from southern British Columbia to San Diego, California along with 2 sites near Sitka, Alaska to better understand the effects of sea star wasting disease (SSWD) on the keystone intertidal predator, Pisaster ochraceus. Quantitative surveys revealed unprecedented declines of P. ochraceus in 2014 and 2015 across nearly the entire geographic range of the species. The intensity of the impact of SSWD was not uniform across the affected area, with proportionally greater population declines in the southern regions relative to the north. The degree of population decline was unrelated to pre-outbreak P. ochraceus density, although these factors have been linked in other well-documented disease events. While elevated seawater temperatures were not broadly linked to the initial emergence of SSWD, anomalously high seawater temperatures in 2014 and 2015 might have exacerbated the disease’s impact. Both before and after the onset of the SSWD outbreak, we documented higher recruitment of P. ochraceus in the north than in the south, and while some juveniles are surviving (as evidenced by transition of recruitment pulses to larger size classes), post-SSWD survivorship is lower than during pre-SSWD periods. In hindsight, our data suggest that the SSWD event defied prediction based on two factors found to be important in other marine disease events, sea water temperature and population density, and illustrate the importance of surveillance of natural populations as one element of an integrated approach to marine disease ecology. Low levels of SSWD-symptomatic sea stars are still present throughout the impacted range, thus the outlook for population recovery is uncertain.

By maintaining interaction webs in the face of warming, herbivores dampen the effects of warming on succession and stability., Climate change can influence ecosystems via both direct effects on individual organisms and indirect effects mediated by species interactions. However, we understand little about how these changes will ripple through ecosystems or whether there are particular ecological characteristics that might make ecosystems more susceptible—or more resistant—to warming. By combining in situ experimental warming with herbivore manipulations in a natural rocky intertidal community for over 16 months, we show that herbivory regulates the capacity of marine communities to resist warming. We found that limpet herbivores helped to preserve trophic and competitive interactions under experimental warming, dampening the impact of warming on overall community composition. The presence of limpets facilitated the survival of the main habitat modifier (barnacles) under warmer conditions, which, in turn, facilitated the presence of a consumer guild. When limpets were removed, environmental warming altered trophic, competitive, and facilitative interactions, with cascading impacts on community succession and stability. We conclude that conserving trophic structure and the integrity of interaction networks is vitally important as Earth continues to warm.

Ocean acidification may have far-reaching consequences for marine community and ecosystem dynamics, but its full impacts remain poorly understood due to the difficulty of manipulating pCO2 at the ecosystem level to mimic realistic fluctuations that occur on a number of different timescales. It is especially unclear how quickly communities at various stages of development respond to intermediate-scale pCO2 change and, if high pCO2 is relieved mid-succession, whether past acidification effects persist, are reversed by alleviation of pCO2 stress, or are worsened by departures from prior high pCO2 conditions to which organisms had acclimatized. Here, we used reciprocal transplant experiments along a shallow water volcanic pCO2 gradient to assess the importance of the timing and duration of high pCO2 exposure (i.e., discrete events at different stages of successional development vs. continuous exposure) on patterns of colonization and succession in a benthic fouling community. We show that succession at the acidified site was initially delayed (less community change by 8Âweeks) but then caught up over the next 4Âweeks. These changes in succession led to homogenization of communities maintained in or transplanted to acidified conditions, and altered community structure in ways that reflected both short- and longer-term acidification history. These community shifts are likely a result of interspecific variability in response to increased pCO2 and changes in species interactions. High pCO2 altered biofilm development, allowing serpulids to do best at the acidified site by the end of the experiment, although early (pretransplant) negative effects of pCO2 on recruitment of these worms were still detectable. The ascidians Diplosoma sp. and Botryllus sp. settled later and were more tolerant to acidification. Overall, transient and persistent acidification-driven changes in the biofouling community, via both past and more recent exposure, could have important implications for ecosystem function and food web dynamics.

Changes in the Earth's environment are now sufficiently complex that our ability to forecast the emergent ecological consequences of ocean acidification (OA) is limited. Such projections are challenging because the effects of OA may be enhanced, reduced or even reversed by other environmental stressors or interactions among species. Despite an increasing emphasis on multifactor and multispecies studies in global change biology, our ability to forecast outcomes at higher levels of organization remains low. Much of our failure lies in a poor mechanistic understanding of nonlinear responses, a lack of specificity regarding the levels of organization at which interactions can arise, and an incomplete appreciation for linkages across these levels. To move forward, we need to fully embrace interactions. Mechanistic studies on physiological processes and individual performance in response to OA must be complemented by work on population and community dynamics. We must also increase our understanding of how linkages and feedback among multiple environmental stressors and levels of organization can generate nonlinear responses to OA. This will not be a simple undertaking, but advances are of the utmost importance as we attempt to mitigate the effects of ongoing global change.

Reduction in seawater pH due to rising levels of anthropogenic carbon dioxide (CO2) in the world's oceans is a major force set to shape the future of marine ecosystems and the ecological services they provide [1,2]. In particular, ocean acidification is predicted to have a detrimental effect on the physiology of calcifying organisms [3]. Yet, the indirect effects of ocean acidification on calcifying organisms, which may counter or exacerbate direct effects, is uncertain. Using volcanic CO2 vents, we tested the indirect effects of ocean acidification on a calcifying herbivore (gastropod) within the natural complexity of an ecological system. Contrary to predictions, the abundance of this calcifier was greater at vent sites (with near-future CO2 levels). Furthermore, translocation experiments demonstrated that ocean acidification did not drive increases in gastropod abundance directly, but indirectly as a function of increased habitat and food (algal biomass). We conclude that the effect of ocean acidification on algae (primary producers) can have a strong, indirect positive influence on the abundance of some calcifying herbivores, which can overwhelm any direct negative effects. This finding points to the need to understand ecological processes that buffer the negative effects of environmental change.

The effects of ocean acidification (OA) on the structure and complexity of coastal marine biogenic habitat have been broadly overlooked. Here we explore how declining pH and carbonate saturation may affect the structural complexity of four major biogenic habitats. Our analyses predict that indirect effects driven by OA on habitat-forming organisms could lead to lower species diversity in coral reefs, mussel beds and some macroalgal habitats, but increases in seagrass and other macroalgal habitats. Available in situ data support the prediction of decreased biodiversity in coral reefs, but not the prediction of seagrass bed gains. Thus, OA-driven habitat loss may exacerbate the direct negative effects of OA on coastal biodiversity; however, we lack evidence of the predicted biodiversity increase in systems where habitat-forming species could benefit from acidification. Overall, a combination of direct effects and community-mediated indirect effects will drive changes in the extent and structural complexity of biogenic habitat, which will have important ecosystem effects. How ocean acidification will impact coastal biogenic habitats is unclear. This study predicts that indirect effects on habitat-forming organisms, combined with direct effects on biodiversity, will cause changes in structural complexity and extent of these habitats.

Despite efforts to understand marine organismal responses to ocean acidification (gradual change in pH/ $${p_{{\text{C}}{{\text{O}}_2}}}$$ over decades), there is a lack of information about the capabilities of coastal organisms to endure rapid and extreme pH change (often full units within hours). We predicted that gastropods faced with estuarine acidification avoid extreme pH exposure through isolation and/or escape behavior, and energetically compensate for feeding and energy uptake limitations by facultative metabolic depression (FMD). To test this, we studied behavioral (organism activity) and aerobic (cardiac performance) responses to acidification in two closely related tropical intertidal species, the estuarine Indothais gradata (two populations) and the open-shore Reishia bitubercularis. Snails were exposed in the laboratory to either acutely declining or stable low pH conditions, using two acidification modes (HNO3-acidification and CO2-aeration). Under acutely declining pH, aerobic performance was regulated to unexpectedly low pH levels (4.5), effectively extending the field pH range for activity. This pH performance threshold marked the onset of behavioral isolation and FMD (as opposed to respiratory stress) and was lower in Indothais than Reishia snails during mineral acidification. Behavioral (in isolated gastropods) and environmental hypercapnic acidosis complicates interpretation of lowered metabolic performance. Stable reduced pH exposures resulted in different behavioral and physiological responses by the Indothais populations, including more prominent escape from water in the seaward population. Overall, these results suggest that aerobic and behavioral flexibility are crucial to organismal fitness in widely fluctuating pH circumstances. They further warn against overgeneralizing marine acidification consequences across physiological dispositions, taxonomic levels, and ecological systems.

Changing temperature can substantially shift ecological communities by altering the strength and stability of trophic interactions. Because many ecological rates are constrained by temperature, new approaches are required to understand how simultaneous changes in multiple rates alter the relative performance of species and their trophic interactions. We develop an energetic approach to identify the relationship between biomass fluxes and standing biomass across trophic levels. Our approach links ecological rates and trophic dynamics to measure temperature-dependent changes to the strength of trophic interactions and determine how these changes alter food web stability. It accomplishes this by using biomass as a common energetic currency and isolating three temperature-dependent processes that are common to all consumer-resource interactions: biomass accumulation of the resource, resource consumption and consumer mortality. Using this framework, we clarify when and how temperature alters consumer to resource biomass ratios, equilibrium resilience, consumer variability, extinction risk and transient vs. equilibrium dynamics. Finally, we characterise key asymmetries in species responses to temperature that produce these distinct dynamic behaviours and identify when they are likely to emerge. Overall, our framework provides a mechanistic and more unified understanding of the temperature dependence of trophic dynamics in terms of ecological rates, biomass ratios and stability. © 2014 John Wiley & Sons Ltd/CNRS.

Thermal stresses, such as those associated with global warming, have the potential to change the strength and even the direction of interspecific interactions. On rocky shores, bio‐eroding microbial endoliths infect the mussel Mytilus californianus. Infestation by microbial endoliths can weaken the shell and lead to mechanical failure and death, increased vulnerability to predation, mechanical damage, etc. However, endolith infestation is associated with the loss of the dark‐colored periostracum, exposing the underlying light gray prismatic layer of the shell and potentially reducing stressful heat gain during low tide. We explore the consequences of the mussel–endolith relationship on mussel tolerance to high environmental temperature through a series of experimental manipulations and comparative observations. Experimental sterilization of the shell surface reduces the rate of periostracum loss, suggesting that the change in shell surface color is indeed caused by microbes residing on/in the shell. Eroded shells absorbed less solar energy than shells with their periostracum intact, and mussel mimics with infested shells remained cooler on sunny days. Manipulation of shell color in the field resulted in higher mortality in black‐painted mussels relative to gray‐painted or naturally eroded mussels. Furthermore, following exceedingly hot weather, mortality was significantly lower for heavily infested, light‐colored mussels than for lightly infested, dark‐colored mussels. Thus, although shell‐boring microbes have the potential to act as parasites under the majority of conditions experienced by mussels, they may be critical mutualists during periods of intense thermal stress. This context‐dependent symbiosis may allow mussels to occupy higher shore levels than would otherwise be possible, and thus indirectly benefit the hundreds of species which use mussel beds as habitat.

Summary In addition to metabolic costs associated with reproduction, morphological and mechanical changes accompanying reproductive effort can affect an organism's performance. Reproductive effort may have unavoidable costs in plants and plant-like taxa (i.e. extra mass or drag); however, the extent to which an organism can ameliorate these consequences by additional modifications of shape or tissue properties remains unknown in seaweed and plant taxa. We investigated mechanical and morphological changes associated with reproduction and how these shifts influence functional performance in the winged kelp, Alaria marginata. Compared to vegetative blades, reproductive blades were similar in width but were longer and had greater surface area. Reproductive sporophylls were also thicker and less ruffled. Tissue extensibility and breaking stress were not different in reproductive vs. vegetative blades. However, reproductive tissue exhibited greater tensile stiffness, flexural stiffness and force to break. Reproductive blades experienced greater drag (despite decreased flapping) than did vegetative blades, but did not experience greater size-specific drag. Tissues cut into experimental blades of the same size and shape experienced greater drag when cut from reproductive tissue suggesting that the change in shape associated with the onset of reproduction ameliorates the cost of increased tissue stiffness. Nonetheless, increased blade breaking force in reproductive individuals resulted in increased blade safety factors (breaking force/drag experienced) in reproductive compared to non-reproductive sporophylls. Synthesis: In A. marginata, decreased flexibility and increased surface area are mechanical costs associated with reproduction. Decreased blade ruffliness and increased strength associated with the onset of reproduction in A. marginata ameliorate the concomitant mechanical costs of decreased flexibility and increased size. Shifts in mechanical and morphological traits among plants and plant-like taxa may allow them to increase reproductive output without decreasing functional performance.

Although the establishment and spread of non-indigenous species depends upon survival in the face of novel environmental conditions and novel biological interactions, relatively little attention has been focused on the specific role of native predators in limiting invasion success. The European common periwinkle, Littorina littorea, was recently introduced to the Pacific coast of Canada and provides a case study of an introduction into an area with an important predator guild (sea stars) that is functionally minor in the invader’s native habitat. Here, we assess the likelihood of establishment, spread, and negative ecological impact of this introduced gastropod, with an emphasis on the role of native sea stars as agents of biotic resistance. Size frequency distributions and local market availability suggest that L. littorea was most likely introduced via the live seafood trade. Non-native hitchhikers (e.g., the trematode Cryptocotyle lingua) were found on/in both market and field specimens. Laboratory studies and field observations confirmed that L. littorea can survive seasonal low salinity in Vancouver, British Columbia. Periwinkles also readily consumed native Ulva, suggesting that periwinkles could impact native communities via herbivory or resource competition. Unlike native gastropods, however, L. littorea lacked behavioural avoidance responses to Northeast Pacific predatory sea stars (Pisaster ochraceus and Pycnopodia helianthoides), and sea star predation rates on L. littorea were much higher than predation rates on native turban snails (Chlorostoma funebralis) in common garden experiments. We therefore expect periwinkle establishment in British Columbia to be limited to areas with low predator density, as is seen in its field distribution to date. We caution that this conclusion may understate the importance of the L. littorea introduction if it also serves as a vector for additional non-indigenous species such as C. lingua.

Summary Organisms' ability to withstand the physical forces of their environment is a key determinant of their success. Mechanical performance of organisms is often associated with the properties of the tissues that compose them. In mechanically stressful habitats, intraspecific variation in tissue properties may result in differential survivorship and enable natural selection to act on material performance. We tested the hypothesis that tissue mechanical properties affect survivorship (a fitness component) of the perennial kelp, Egregia menziesii, in a mechanically stressful, wave-swept intertidal habitat. We measured intraspecific variation in strength and flexibility in 38 E. menziesii and tracked their survivorship in the field over the winter storm season to determine whether variation in mechanical properties led to differential survivorship. Significant interindividual variation was found in most mechanical properties, including strength and flexibility. Individuals with increased flexibility and decreased strength were more likely to survive the duration of our study, although this effect was more pronounced in individuals with smaller holdfasts. Increased frond strength was also associated with a reduction in self-thinning, potentially explaining the observed increase in whole-plant mortality with increasing frond strength. Results from this study demonstrate that variation in tissue mechanical properties among conspecifics can influence survivorship and this may have important evolutionary implications.

At a proximal level, the physiological impacts of global climate change on ectothermic organisms are manifest as changes in body temperatures. Especially for plants and animals exposed to direct solar radiation, body temperatures can be substantially different from air temperatures. We deployed biomimetic sensors that approximate the thermal characteristics of intertidal mussels at 71 sites worldwide, from 1998-present. Loggers recorded temperatures at 10–30 min intervals nearly continuously at multiple intertidal elevations. Comparisons against direct measurements of mussel tissue temperature indicated errors of ~2.0–2.5 °C, during daily fluctuations that often exceeded 15°–20 °C. Geographic patterns in thermal stress based on biomimetic logger measurements were generally far more complex than anticipated based only on ‘habitat-level’ measurements of air or sea surface temperature. This unique data set provides an opportunity to link physiological measurements with spatially- and temporally-explicit field observations of body temperature.

Seaweeds are ecologically important primary producers, competitors, and ecosystem engineers that play a central role in coastal habitats ranging from kelp forests to coral reefs. Although seaweeds are known to be vulnerable to physical and chemical changes in the marine environment, the impacts of ongoing and future anthropogenic climate change in seaweed-dominated ecosystems remain poorly understood. In this review, we describe the ways in which changes in the environment directly affect seaweeds in terms of their physiology, growth, reproduction, and survival. We consider the extent to which seaweed species may be able to respond to these changes via adaptation or migration. We also examine the extensive reshuffling of communities that is occurring as the ecological balance between competing species changes, and as top-down control by herbivores becomes stronger or weaker. Finally, we delve into some of the ecosystem-level responses to these changes, including changes in primary productivity, diversity, and resilience. Although there are several key areas in which ecological insight is lacking, we suggest that reasonable climate-related hypotheses can be developed and tested based on current information. By strategically prioritizing research in the areas of complex environmental variation, multiple stressor effects, evolutionary adaptation, and population, community, and ecosystem-level responses, we can rapidly build upon our current understanding of seaweed biology and climate change ecology to more effectively conserve and manage coastal ecosystems.

Climate change can affect organisms both directly via physiological stress and indirectly via changing relationships among species. However, we do not fully understand how changing interspecific relationships contribute to community- and ecosystem-level responses to environmental forcing. I used experiments and spatial and temporal comparisons to demonstrate that warming substantially reduces predator-free space on rocky shores. The vertical extent of mussel beds decreased by 51% in 52 years, and reproductive populations of mussels disappeared at several sites. Prey species were able to occupy a hot, extralimital site if predation pressure was experimentally reduced, and local species richness more than doubled as a result. These results suggest that anthropogenic climate change can alter interspecific interactions and produce unexpected changes in species distributions, community structure, and diversity.

Increasing levels of anthropogenic carbon dioxide in the world's oceans are resulting in a decrease in the availability of carbonate ions and a drop in seawater pH. This process, known as ocean acidification, is a potential threat to marine populations via alterations in survival and development. To date, however, little research has examined the effects of ocean acidification on rare or endangered species. To begin to assess the impacts of acidification on endangered northern abalone (Haliotis kamtschatkana) populations, we exposed H. kamtschatkana larvae to various levels of CO2 [400 ppm (ambient), 800 ppm, and 1800 ppm CO2] and measured survival, settlement, shell size, and shell development. Larval survival decreased by ca. 40% in elevated CO2 treatments relative to the 400 ppm control. However, CO2 had no effect on the proportion of surviving larvae that metamorphosed at the end of the experiment. Larval shell abnormalities became apparent in approximately 40% of larvae reared at 800 ppm CO2, and almost all larvae reared at 1800 ppm CO2 either developed an abnormal shell or lacked a shell completely. Of the larvae that did not show shell abnormalities, shell size was reduced by 5% at 800 ppm compared to the control. Overall, larval development of H. kamtschatkana was found to be sensitive to ocean acidification. Near future levels of CO2 will likely pose a significant additional threat to this species, which is already endangered with extinction due in part to limited reproductive output and larval recruitment.

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.