Species redistribution creates unequal outcomes for multispecies fisheries under projected climate change.

Climate change drives species distribution shifts, affecting the availability of resources people rely upon for food and livelihoods. These impacts are complex, manifest at local scales, and have diverse effects across multiple species. However, for wild capture fisheries, current understanding is dominated by predictions for individual species at coarse spatial scales. We show that species-specific responses to localized environmental changes will alter the collection of co-occurring species within established fishing footprints along the U.S. West Coast. We demonstrate that availability of the most economically valuable, primary target species is highly likely to decline coastwide in response to warming and reduced oxygen concentrations, while availability of the most abundant, secondary target species will potentially increase. A spatial reshuffling of primary and secondary target species suggests regionally heterogeneous opportunities for fishers to adapt by changing where or what they fish. Developing foresight into the collective responses of species at local scales will enable more effective and tangible adaptation pathways for fishing communities.

Recommendations for quantifying and reducing uncertainty in climate projections of species distributions.

Projecting the future distributions of commercially and ecologically important species has become a critical approach for ecosystem managers to strategically anticipate change, but large uncertainties in projections limit climate adaptation planning. Although distribution projections are primarily used to understand the scope of potential change-rather than accurately predict specific outcomes-it is nonetheless essential to understand where and why projections can give implausible results and to identify which processes contribute to uncertainty. Here, we use a series of simulated species distributions, an ensemble of 252 species distribution models, and an ensemble of three regional ocean climate projections, to isolate the influences of uncertainty from earth system model spread and from ecological modeling. The simulations encompass marine species with different functional traits and ecological preferences to more broadly address resource manager and fishery stakeholder needs, and provide a simulated true state with which to evaluate projections. We present our results relative to the degree of environmental extrapolation from historical conditions, which helps facilitate interpretation by ecological modelers working in diverse systems. We found uncertainty associated with species distribution models can exceed uncertainty generated from diverging earth system models (up to 70% of total uncertainty by 2100), and that this result was consistent across species traits. Species distribution model uncertainty increased through time and was primarily related to the degree to which models extrapolated into novel environmental conditions but moderated by how well models captured the underlying dynamics driving species distributions. The predictive power of simulated species distribution models remained relatively high in the first 30 years of projections, in alignment with the time period in which stakeholders make strategic decisions based on climate information. By understanding sources of uncertainty, and how they change at different forecast horizons, we provide recommendations for projecting species distribution models under global climate change.

The role of continental shelf bathymetry in shaping marine range shifts in the face of climate change.

As a consequence of anthropogenic climate change, marine species on continental shelves around the world are rapidly shifting deeper and poleward. However, whether these shifts deeper and poleward will allow species to access more, less, or equivalent amounts of continental shelf area and associated critical habitats remains unclear. By examining the proportion of seabed area at a range of depths for each large marine ecosystem (LME), we found that shelf area declined monotonically for 19% of LMEs examined. However, the majority exhibited a greater proportion of shelf area in mid-depths or across several depth ranges. By comparing continental shelf area across 2° latitudinal bands, we found that all coastlines exhibit multiple instances of shelf area expansion and contraction, which have the potential to promote or restrict poleward movement of marine species. Along most coastlines, overall shelf habitat increases or exhibits no significant change moving towards the poles. The exception is the Southern West Pacific, which experiences an overall loss of area with increasing latitude. Changes in continental shelf area availability across latitudes and depths are likely to affect the number of species local ecosystems can support. These geometric analyses help identify regions of conservation priority and ecological communities most likely to face attrition or expansion due to variations in available area.

Range edges of North American marine species are tracking temperature over decades.

Understanding the dynamics of species range edges in the modern era is key to addressing fundamental biogeographic questions about abiotic and biotic drivers of species distributions. Range edges are where colonization and extirpation processes unfold, and so these dynamics are also important to understand for effective natural resource management and conservation. However, few studies to date have analyzed time series of range edge positions in the context of climate change, in part because range edges are difficult to detect. We first quantified positions for 165 range edges of marine fishes and invertebrates from three U.S. continental shelf regions using up to five decades of survey data and a spatiotemporal model to account for sampling and measurement variability. We then analyzed whether those range edges maintained their edge thermal niche-the temperatures found at the range edge position-over time. A large majority of range edges (88%) maintained either summer or winter temperature extremes at the range edge over the study period, and most maintained both (76%), although not all of those range edges shifted in space. However, we also found numerous range edges-particularly poleward edges and edges in the region that experienced the most warming-that did not shift at all, shifted further than predicted by temperature alone, or shifted opposite the direction expected, underscoring the multiplicity of factors that drive changes in range edge positions. This study suggests that range edges of temperate marine species have largely maintained the same edge thermal niche during periods of rapid change and provides a blueprint for testing whether and to what degree species range edges track temperature in general.

Climate-Driven Shifts in Marine Species Ranges: Scaling from Organisms to Communities.

The geographic distributions of marine species are changing rapidly, with leading range edges following climate poleward, deeper, and in other directions and trailing range edges often contracting in similar directions. These shifts have their roots in fine-scale interactions between organisms and their environment-including mosaics and gradients of temperature and oxygen-mediated by physiology, behavior, evolution, dispersal, and species interactions. These shifts reassemble food webs and can have dramatic consequences. Compared with species on land, marine species are more sensitive to changing climate but have a greater capacity for colonization. These differences suggest that species cope with climate change at different spatial scales in the two realms and that range shifts across wide spatial scales are a key mechanism at sea. Additional research is needed to understand how processes interact to promote or constrain range shifts, how the dominant responses vary among species, and how the emergent communities of the future ocean will function.

Changes in higher trophic level productivity, diversity and niche space in a rapidly warming continental shelf ecosystem.

There is long-standing ecological and socioeconomic interest in what controls the diversity and productivity of ecosystems. That focus has intensified with shifting environmental conditions associated with accelerating climate change. The U.S. Northeast Shelf (NES) is a well-studied continental shelf marine ecosystem that is among the more rapidly warming marine systems worldwide. Furthermore, many constituent species have experienced significant distributional shifts. However, the system response of the NES to climate change goes beyond simple shifts in species distribution. The fish and macroinvertebrate communities of the NES have increased in species diversity and overall productivity in recent decades, despite no significant decline in fishing pressure. Species distribution models constructed using random forest classification and regression trees were fit for the dominant species in the system. Over time, the areal distribution of occupancy habitat has increased for approximately 80% of the modeled taxa, suggesting most species have significantly increased their range and niche space. These niche spaces were analyzed to determine the area of niche overlap between species pairs. For the vast majority of species pairs, interaction has increased over time suggesting greater niche overlap and the increased probability for more intense species interactions, such as between competitors or predators and prey. Furthermore, the species taxonomic composition and size structure indicate a potential tropicalization of the fish community. The system and community changes are consistent with the view that the NES may be transitioning from a cold temperate or boreal ecoregion to one more consistent with the composition of a warm temperate or Carolinian system.

Opportunities for behavioral rescue under rapid environmental change.

Laboratory measurements of physiological and demographic tolerances are important in understanding the impact of climate change on species diversity; however, it has been recognized that forecasts based solely on these laboratory estimates overestimate risk by omitting the capacity for species to utilize microclimatic variation via behavioral adjustments in activity patterns or habitat choice. The complex, and often context-dependent nature, of microclimate utilization has been an impediment to the advancement of general predictive models. Here, we overcome this impediment and estimate the potential impact of warming on the fitness of ectotherms using a benefit/cost trade-off derived from the simple and broadly documented thermal performance curve and a generalized cost function. Our framework reveals that, for certain environments, the cost of behavioral thermoregulation can be reduced as warming occurs, enabling behavioral buffering (e.g., the capacity for behavior to ameliorate detrimental impacts) and "behavioral rescue" from extinction in extreme cases. By applying our framework to operative temperature and physiological data collected at an extremely fine spatial scale in an African lizard, we show that new behavioral opportunities may emerge. Finally, we explore large-scale geographic differences in the impact of behavior on climate-impact projections using a global dataset of 38 insect species. These multiple lines of inference indicate that understanding the existing relationship between thermal characteristics (e.g., spatial configuration, spatial heterogeneity, and modal temperature) is essential for improving estimates of extinction risk.

Correction: Eco-Label Conveys Reliable Information on Fish Stock Health to Seafood Consumers.

[This corrects the article DOI: 10.1371/journal.pone.0043765.].

Projecting shifts in thermal habitat for 686 species on the North American continental shelf.

Recent shifts in the geographic distribution of marine species have been linked to shifts in preferred thermal habitats. These shifts in distribution have already posed challenges for living marine resource management, and there is a strong need for projections of how species might be impacted by future changes in ocean temperatures during the 21st century. We modeled thermal habitat for 686 marine species in the Atlantic and Pacific oceans using long-term ecological survey data from the North American continental shelves. These habitat models were coupled to output from sixteen general circulation models that were run under high (RCP 8.5) and low (RCP 2.6) future greenhouse gas emission scenarios over the 21st century to produce 32 possible future outcomes for each species. The models generally agreed on the magnitude and direction of future shifts for some species (448 or 429 under RCP 8.5 and RCP 2.6, respectively), but strongly disagreed for other species (116 or 120 respectively). This allowed us to identify species with more or less robust predictions. Future shifts in species distributions were generally poleward and followed the coastline, but also varied among regions and species. Species from the U.S. and Canadian west coast including the Gulf of Alaska had the highest projected magnitude shifts in distribution, and many species shifted more than 1000 km under the high greenhouse gas emissions scenario. Following a strong mitigation scenario consistent with the Paris Agreement would likely produce substantially smaller shifts and less disruption to marine management efforts. Our projections offer an important tool for identifying species, fisheries, and management efforts that are particularly vulnerable to climate change impacts.

Diversity in thermal affinity among key piscivores buffers impacts of ocean warming on predator-prey interactions.

Asymmetries in responses to climate change have the potential to alter important predator-prey interactions, in part by altering the location and size of spatial refugia for prey. We evaluated the effect of ocean warming on interactions between four important piscivores and four of their prey in the U.S. Northeast Shelf by examining species overlap under historical conditions (1968-2014) and with a doubling in CO2 . Because both predator and prey shift their distributions in response to changing ocean conditions, the net impact of warming or cooling on predator-prey interactions was not determined a priori from the range extent of either predator or prey alone. For Atlantic cod, an historically dominant piscivore in the region, we found that both historical and future warming led to a decline in the proportion of prey species' range it occupied and caused a potential reduction in its ability to exert top-down control on these prey. In contrast, the potential for overlap of spiny dogfish with prey species was enhanced by warming, expanding their importance as predators in this system. In sum, the decline in the ecological role for cod that began with overfishing in this ecosystem will likely be exacerbated by warming, but this loss may be counteracted by the rise in dominance of other piscivores with contrasting thermal preferences. Functional diversity in thermal affinity within the piscivore guild may therefore buffer against the impact of warming on marine ecosystems, suggesting a novel mechanism by which diversity confers resilience.

Gradual changes in range size accompany long-term trends in species richness.

Species richness has long been used as an indicator of ecosystem functioning and health. Global richness is declining, but it is unclear whether sub-global trends differ. Regional trends are especially understudied, with most focused on island regions where richness is strongly impacted by novel colonisations. We addressed this knowledge gap by testing for multi-decade trends in species richness in nine open marine regions around North America (197 region-years) while accounting for imperfect observations and grounding our findings in species-level range dynamics. We found positive richness trends in eight of nine regions, four of which were statistically significant. Species' range sizes generally contracted pre-extinction and expanded post-colonisation, but the ranges of transient species expanded over the long-term, slowly increasing their regional retention and driving increasing richness. These results provide more evidence that sub-global richness trends are stable or increasing, and highlight the utility of range size for understanding richness dynamics.

Protection of large predators in a marine reserve alters size-dependent prey mortality.

Where predator-prey interactions are size-dependent, reductions in predator size owing to fishing has the potential to disrupt the ecological role of top predators in marine ecosystems. In southern California kelp forests, we investigated the size-dependence of the interaction between herbivorous sea urchins and one of their predators, California sheephead (Semicossyphus pulcher). Empirical tests examined how differences in predator size structure between reserve and fished areas affected size-specific urchin mortality. Sites inside marine reserves had greater sheephead size and biomass, while empirical feeding trials indicated that larger sheephead were required to successfully consume urchins of increasing test diameter. Evaluations of the selectivity of sheephead for two urchin species indicated that shorter-spined purple urchins were attacked more frequently and successfully than longer-spined red urchins of the same size class, particularly at the largest test diameters. As a result of these size-specific interactions and the higher biomass of large sheephead inside reserves, urchin mortality rates were three times higher inside the reserve for both species. In addition, urchin mortality rates decreased with urchin size, and very few large urchins were successfully consumed in fished areas. The truncation of sheephead size structure that commonly occurs owing to fishing will probably result in reductions in urchin mortality, which may reduce the resilience of kelp beds to urchin barren formation. By contrast, the recovery of predator size structure in marine reserves may restore this resilience, but may be delayed until fish grow to sizes capable of consuming larger urchins.

Eco-label conveys reliable information on fish stock health to seafood consumers.

Concerns over fishing impacts on marine populations and ecosystems have intensified the need to improve ocean management. One increasingly popular market-based instrument for ecological stewardship is the use of certification and eco-labeling programs to highlight sustainable fisheries with low environmental impacts. The Marine Stewardship Council (MSC) is the most prominent of these programs. Despite widespread discussions about the rigor of the MSC standards, no comprehensive analysis of the performance of MSC-certified fish stocks has yet been conducted. We compared status and abundance trends of 45 certified stocks with those of 179 uncertified stocks, finding that 74% of certified fisheries were above biomass levels that would produce maximum sustainable yield, compared with only 44% of uncertified fisheries. On average, the biomass of certified stocks increased by 46% over the past 10 years, whereas uncertified fisheries increased by just 9%. As part of the MSC process, fisheries initially go through a confidential pre-assessment process. When certified fisheries are compared with those that decline to pursue full certification after pre-assessment, certified stocks had much lower mean exploitation rates (67% of the rate producing maximum sustainable yield vs. 92% for those declining to pursue certification), allowing for more sustainable harvesting and in many cases biomass rebuilding. From a consumer's point of view this means that MSC-certified seafood is 3-5 times less likely to be subject to harmful fishing than uncertified seafood. Thus, MSC-certification accurately identifies healthy fish stocks and conveys reliable information on stock status to seafood consumers.

Do behavioral foraging responses of prey to predators function similarly in restored and pristine foodwebs?

Efforts to restore top predators in human-altered systems raise the question of whether rebounds in predator populations are sufficient to restore pristine foodweb dynamics. Ocean ecosystems provide an ideal system to test this question. Removal of fishing in marine reserves often reverses declines in predator densities and size. However, whether this leads to restoration of key functional characteristics of foodwebs, especially prey foraging behavior, is unclear. The question of whether restored and pristine foodwebs function similarly is nonetheless critically important for management and restoration efforts. We explored this question in light of one important determinant of ecosystem function and structure--herbivorous prey foraging behavior. We compared these responses for two functionally distinct herbivorous prey fishes (the damselfish Plectroglyphidodon dickii and the parrotfish Chlorurus sordidus) within pairs of coral reefs in pristine and restored ecosystems in two regions of these species' biogeographic ranges, allowing us to quantify the magnitude and temporal scale of this key ecosystem variable's recovery. We demonstrate that restoration of top predator abundances also restored prey foraging excursion behaviors to a condition closely resembling those of a pristine ecosystem. Increased understanding of behavioral aspects of ecosystem change will greatly improve our ability to predict the cascading consequences of conservation tools aimed at ecological restoration, such as marine reserves.