Marine heatwaves are not a dominant driver of change in demersal fishes.

Marine heatwaves have been linked to negative ecological effects in recent decades1,2. If marine heatwaves regularly induce community reorganization and biomass collapses in fishes, the consequences could be catastrophic for ecosystems, fisheries and human communities3,4. However, the extent to which marine heatwaves have negative impacts on fish biomass or community composition, or even whether their effects can be distinguished from natural and sampling variability, remains unclear. We investigated the effects of 248 sea-bottom heatwaves from 1993 to 2019 on marine fishes by analysing 82,322 hauls (samples) from long-term scientific surveys of continental shelf ecosystems in North America and Europe spanning the subtropics to the Arctic. Here we show that the effects of marine heatwaves on fish biomass were often minimal and could not be distinguished from natural and sampling variability. Furthermore, marine heatwaves were not consistently associated with tropicalization (gain of warm-affiliated species) or deborealization (loss of cold-affiliated species) in these ecosystems. Although steep declines in biomass occasionally occurred after marine heatwaves, these were the exception, not the rule. Against the highly variable backdrop of ocean ecosystems, marine heatwaves have not driven biomass change or community turnover in fish communities that support many of the world's largest and most productive fisheries.

Are we ready to track climate-driven shifts in marine species across international boundaries? - A global survey of scientific bottom trawl data.

Marine biota are redistributing at a rapid pace in response to climate change and shifting seascapes. While changes in fish populations and community structure threaten the sustainability of fisheries, our capacity to adapt by tracking and projecting marine species remains a challenge due to data discontinuities in biological observations, lack of data availability, and mismatch between data and real species distributions. To assess the extent of this challenge, we review the global status and accessibility of ongoing scientific bottom trawl surveys. In total, we gathered metadata for 283,925 samples from 95 surveys conducted regularly from 2001 to 2019. We identified that 59% of the metadata collected are not publicly available, highlighting that the availability of data is the most important challenge to assess species redistributions under global climate change. Given that the primary purpose of surveys is to provide independent data to inform stock assessment of commercially important populations, we further highlight that single surveys do not cover the full range of the main commercial demersal fish species. An average of 18 surveys is needed to cover at least 50% of species ranges, demonstrating the importance of combining multiple surveys to evaluate species range shifts. We assess the potential for combining surveys to track transboundary species redistributions and show that differences in sampling schemes and inconsistency in sampling can be overcome with spatio-temporal modeling to follow species density redistributions. In light of our global assessment, we establish a framework for improving the management and conservation of transboundary and migrating marine demersal species. We provide directions to improve data availability and encourage countries to share survey data, to assess species vulnerabilities, and to support management adaptation in a time of climate-driven ocean changes.

The rise of a marine generalist predator and the fall of beta diversity.

Determining the importance of physical and biological drivers in shaping biodiversity in diverse ecosystems remains a global challenge. Advancements have been made towards this end in large marine ecosystems with several studies suggesting environmental forcing as the primary driver. However, both empirical and theoretical studies point to additional drivers of changes in diversity involving trophic interactions and, in particular, predation. Moreover, a more integrated but less common approach to the assessment of biodiversity changes involves analyses of spatial ß diversity, whereas most studies to date assess only changes in species richness (α diversity). Recent research has established that when cod, a dominant generalist predator, was overfished and collapsed in a northwest Atlantic food web, spatial ß diversity increased; that is, the spatial structure of the fish assemblage became increasingly heterogeneous. If cod were to recover, would this situation be reversible, given the inherent complexity and non-linear dynamics that typify such systems? A dramatic increase of cod in an ecologically similar large marine ecosystem may provide an answer. Here we show that spatial ß diversity of fish assemblages in the Barents Sea decreased with increasing cod abundance, while decadal scale changes in temperature did not play a significant role. These findings indicate a reversibility of the fish assemblage structure in response to changing levels of an apex predator and highlight the frequently overlooked importance of trophic interactions in determining large-scale biodiversity patterns. As increased cod abundance was largely driven by changes in fisheries management, our study also shows that management policies and practices, particularly those involving apex predators, can have a strong effect in shaping spatial diversity patterns, and one should not restrict the focus to effects of climate change alone.

Resource-driven colonization by cod in a high Arctic food web.

Climate change is commonly associated with many species redistributions and the influence of other factors may be marginalized, especially in the rapidly warming Arctic.The Barents Sea, a high latitude large marine ecosystem in the Northeast Atlantic has experienced above-average temperatures since the mid-2000s with divergent bottom temperature trends at subregional scales.Concurrently, the Barents Sea stock of Atlantic cod Gadus morhua, one of the most important commercial fish stocks in the world, increased following a large reduction in fishing pressure and expanded north of 80°N.We examined the influence of food availability and temperature on cod expansion using a comprehensive data set on cod stomach fullness stratified by subregions characterized by divergent temperature trends. We then tested whether food availability, as indexed by cod stomach fullness, played a role in cod expansion in subregions that were warming, cooling, or showed no trend.The greatest increase in cod occupancy occurred in three northern subregions with contrasting temperature trends. Cod apparently benefited from initial high food availability in these regions that previously had few large-bodied fish predators.The stomach fullness in the northern subregions declined rapidly after a few years of high cod abundance, suggesting that the arrival of cod caused a top-down effect on the prey base. Prolonged cod residency in the northern Barents Sea is, therefore, not a certainty.

Connectivity, persistence, and loss of high abundance areas of a recovering marine fish population in the Northwest Atlantic Ocean.

In the early 1990s, the Northwest Atlantic Ocean underwent a fisheries-driven ecosystem shift. Today, the iconic cod (Gadus morhua) remains at low levels, while Atlantic halibut (Hippoglossus hippoglossus) has been increasing since the mid-2000s, concomitant with increasing interest from the fishing industry. Currently, our knowledge about halibut ecology is limited, and the lack of recovery in other collapsed groundfish populations has highlighted the danger of overfishing local concentrations. Here, we apply a Bayesian hierarchical spatiotemporal approach to model the spatial structure of juvenile Atlantic halibut over 36 years and three fisheries management regimes using three model parameters to characterize the resulting spatiotemporal abundance structure: persistence (similarity of spatial structure over time), connectivity (coherence of temporal pattern over space), and spatial variance (variation across the seascape). Two areas of high juvenile abundance persisted through three decades whereas two in the northeast are now diminished, despite the increased abundance and landings throughout the management units. The persistent areas overlap with full and seasonal area closures, which may act as refuges from fishing. Connectivity was estimated to be 250 km, an order of magnitude less than the distance assumed by the definition of the Canadian management units (~2,000 km). The underlying question of whether there are distinct populations within the southern stock unit cannot be answered with this model, but the smaller ~250 km scale of coherent temporal patterns suggests more complex population structure than previously thought, which should be taken into consideration by fishery management.

Local overfishing may be avoided by examining parameters of a spatio-temporal model.

Spatial erosion of stock structure through local overfishing can lead to stock collapse because fish often prefer certain locations, and fisheries tend to focus on those locations. Fishery managers are challenged to maintain the integrity of the entire stock and require scientific approaches that provide them with sound advice. Here we propose a Bayesian hierarchical spatio-temporal modelling framework for fish abundance data to estimate key parameters that define spatial stock structure: persistence (similarity of spatial structure over time), connectivity (coherence of temporal pattern over space), and spatial variance (variation across the seascape). The consideration of these spatial parameters in the stock assessment process can help identify the erosion of structure and assist in preventing local overfishing. We use Atlantic cod (Gadus morhua) in eastern Canada as a case study an examine the behaviour of these parameters from the height of the fishery through its collapse. We identify clear signals in parameter behaviour under circumstances of destructive stock erosion as well as for recovery of spatial structure even when combined with a non-recovery in abundance. Further, our model reveals the spatial pattern of areas of high and low density persists over the 41 years of available data and identifies the remnant patches. Models of this sort are crucial to recovery plans if we are to identify and protect remaining sources of recolonization for Atlantic cod. Our method is immediately applicable to other exploited species.

Marine species in ambient low-oxygen regions subject to double jeopardy impacts of climate change.

We have learned much about the impacts of warming on the productivity and distribution of marine organisms, but less about the impact of warming combined with other environmental stressors, including oxygen depletion. Also, the combined impact of multiple environmental stressors requires evaluation at the scales most relevant to resource managers. We use the Gulf of St. Lawrence, Canada, characterized by a large permanently hypoxic zone, as a case study. Species distribution models were used to predict the impact of multiple scenarios of warming and oxygen depletion on the local density of three commercially and ecologically important species. Substantial changes are projected within 20-40 years. A eurythermal depleted species already limited to shallow, oxygen-rich refuge habitat (Atlantic cod) may be relatively uninfluenced by oxygen depletion but increase in density within refuge areas with warming. A more stenothermal, deep-dwelling species (Greenland halibut) is projected to lose ~55% of its high-density areas under the combined impacts of warming and oxygen depletion. Another deep-dwelling, more eurythermal species (Northern shrimp) would lose ~4% of its high-density areas due to oxygen depletion alone, but these impacts may be buffered by warming, which may increase density by 8% in less hypoxic areas, but decrease density by ~20% in the warmest parts of the region. Due to local climate variability and extreme events, and that our models cannot project changes in species sensitivity to hypoxia with warming, our results should be considered conservative. We present an approach to effectively evaluate the individual and cumulative impacts of multiple environmental stressors on a species-by-species basis at the scales most relevant to managers. Our study may provide a basis for work in other low-oxygen regions and should contribute to a growing literature base in climate science, which will continue to be of support for resource managers as climate change accelerates.

The role of a dominant predator in shaping biodiversity over space and time in a marine ecosystem.

1. Exploitation of living marine resources has resulted in major changes to populations of targeted species and functional groups of large-bodied species in the ocean. However, the effects of overfishing and collapse of large top predators on the broad-scale biodiversity of oceanic ecosystems remain largely unexplored. 2. Populations of the Atlantic cod (Gadus morhua) were overfished and several collapsed in the early 1990s across Atlantic Canada, providing a unique opportunity to study potential ecosystem-level effects of the reduction of a dominant predator on fish biodiversity, and to identify how such effects might interact with other environmental factors, such as changes in climate, over time. 3. We combined causal modelling with model selection and multimodel inference to analyse 41 years of fishery-independent survey data (1970-2010) and quantify ecosystem-level effects of overfishing and climate variation on the biodiversity of fishes across a broad area (172 000 km(2) ) of the Scotian Shelf. 4. We found that alpha and beta diversity increased with decreases in cod occurrence; fish communities were less homogeneous and more variable in systems where cod no longer dominated. These effects were most pronounced in the colder north-eastern parts of the Scotian Shelf. 5. Our results provide strong evidence that intensive harvesting (and collapse) of marine apex predators can have large impacts on biodiversity, with far-reaching consequences for ecological stability across an entire ecosystem.

The potential for spatial distribution indices to signal thresholds in marine fish biomass.

The frequently observed positive relationship between fish population abundance and spatial distribution suggests that changes in distribution can be indicative of trends in abundance. If contractions in spatial distribution precede declines in spawning stock biomass (SSB), spatial distribution reference points could complement the SSB reference points that are commonly used in marine conservation biology and fisheries management. When relevant spatial distribution information is integrated into fisheries management and recovery plans, risks and uncertainties associated with a plan based solely on the SSB criterion would be reduced. To assess the added value of spatial distribution data, we examine the relationship between SSB and four metrics of spatial distribution intended to reflect changes in population range, concentration, and density for 10 demersal populations (9 species) inhabiting the Scotian Shelf, Northwest Atlantic. Our primary purpose is to assess their potential to serve as indices of SSB, using fisheries independent survey data. We find that metrics of density offer the best correlate of spawner biomass. A decline in the frequency of encountering high density areas is associated with, and in a few cases preceded by, rapid declines in SSB in 6 of 10 populations. Density-based indices have considerable potential to serve both as an indicator of SSB and as spatially based reference points in fisheries management.

Thermal habitat index of many northwest Atlantic temperate species stays neutral under warming projected for 2030 but changes radically by 2060.

Global scale forecasts of range shifts in response to global warming have provided vital insight into predicted species redistribution. We build on that insight by examining whether local warming will affect habitat on spatiotemporal scales relevant to regional agencies. We used generalized additive models to quantify the realized habitat of 46 temperate/boreal marine species using 41+ years of survey data from 35°N-48°N in the Northwest Atlantic. We then estimated change in a "realized thermal habitat index" under short-term (2030) and long-term (2060) warming scenarios. Under the 2030 scenario, ∼10% of species will lose realized thermal habitat at the national scale (USA and Canada) but planktivores are expected to lose significantly in both countries which may result in indirect changes in their predators' distribution. In contrast, by 2060 in Canada, the realized habitat of 76% of species will change (55% will lose, 21% will gain) while in the USA, the realized habitat of 85% of species will change (65% will lose, 20% will gain). If all else were held constant, the ecosystem is projected to change radically based on thermal habitat alone. The magnitude of the 2060 warming projection (∼1.5-3°C) was observed in 2012 affirming that research is needed on effects of extreme "weather" in addition to increasing mean temperature. Our approach can be used to aggregate at smaller spatial scales where temperate/boreal species are hypothesized to have a greater loss at ∼40°N. The uncertainty associated with climate change forecasts is large, yet resource management agencies still have to address climate change. How? Since many fishery agencies do not plan beyond 5 years, a logical way forward is to incorporate a "realized thermal habitat index" into the stock assessment process. Over time, decisions would be influenced by the amount of suitable thermal habitat, in concert with gradual or extreme warming.

Spatial scale of similarity as an indicator of metacommunity stability in exploited marine systems.

The spatial scale of similarity among fish communities is characteristically large in temperate marine systems: connectivity is enhanced by high rates of dispersal during the larval/juvenile stages and the increased mobility of large-bodied fish. A larger spatial scale of similarity (low beta diversity) is advantageous in heavily exploited systems because locally depleted populations are more likely to be "rescued" by neighboring areas. We explored whether the spatial scale of similarity changed from 1970 to 2006 due to overfishing of dominant, large-bodied groundfish across a 300 000-km2 region of the Northwest Atlantic. Annually, similarities among communities decayed slowly with increasing geographic distance in this open system, but through time the decorrelation distance declined by 33%, concomitant with widespread reductions in biomass, body size, and community evenness. The decline in connectivity stemmed from an erosion of community similarity among local subregions separated by distances as small as 100 km. Larger fish, of the same species, contribute proportionally more viable offspring, so observed body size reductions will have affected maternal output. The cumulative effect of nonlinear maternal influences on egg/larval quality may have compromised the spatial scale of effective larval dispersal, which may account for the delayed recovery of certain member species. Our study adds strong support for using the spatial scale of similarity as an indicator of metacommunity stability both to understand the spatial impacts of exploitation and to refine how spatial structure is used in management plans.

Decline in top predator body size and changing climate alter trophic structure in an oceanic ecosystem.

Globally, overfishing large-bodied groundfish populations has resulted in substantial increases in their prey populations. Where it has been examined, the effects of overfishing have cascaded down the food chain. In an intensively fished area on the western Scotian Shelf, Northwest Atlantic, the biomass of prey species increased exponentially (doubling time of 11 years) even though the aggregate biomass of their predators remained stable over 38 years. Concomitant reductions in herbivorous zooplankton and increases in phytoplankton were also evident. This anomalous trophic pattern led us to examine how declines in predator body size (approx. 60% in body mass since the early 1970s) and climatic regime influenced lower trophic levels. The increase in prey biomass was associated primarily with declines in predator body size and secondarily to an increase in stratification. Sea surface temperature and predator biomass had no influence. A regression model explained 65 per cent of prey biomass variability. Trait-mediated effects, namely a reduction in predator size, resulted in a weakening of top predation pressure. Increased stratification may have enhanced growing conditions for prey fish. Size-selective harvesting under changing climatic conditions initiated a trophic restructuring of the food chain, the effects of which may have influenced three trophic levels.

Temporal dynamics within a contemporary latitudinal diversity gradient.

Poleward declines in species diversity [latitudinal diversity gradients (LDG)] remain among the oldest and most widespread of macroecological patterns. However, their contemporary dynamics remain largely unexplored even though changing ecological conditions, including global change, may modify LDG and their respective ecosystems. Here, we examine temporal variation within a temperate Northwest Atlantic LDG using 31 years of annual fisheries-independent surveys and explore its dynamics in relation to a dominant climate signal [the wintertime North Atlantic Oscillation (NAO)] that varies interannually and alters the latitudinal gradient of Northwest Atlantic continental shelf bottom water temperatures. We found that the slopes of the annual LDG vary dramatically due to changes in geographic distributions of 100+ species, variations that are concealed within the cumulative, static LDG. These changes are strongly associated with changes in NAO sign and strength. This is the first illustration of temporal dynamics in a contemporary LDG and the first demonstration of the speed at which local environmental variations can alter an LDG. Our findings underscore the need to investigate factors that modify LDG separately from those that contribute to their origins.

The ups and downs of trophic control in continental shelf ecosystems.

Traditionally, marine ecosystem structure was thought to be determined by phytoplankton dynamics. However, an integrated view on the relative roles of top-down (consumer-driven) and bottom-up (resource-driven) forcing in large-scale, exploited marine ecosystems is emerging. Long time series of scientific survey data, underpinning the management of commercially exploited species such as cod, are being used to diagnose mechanisms that could affect the composition and relative abundance of species in marine food webs. By assembling published data from studies in exploited North Atlantic ecosystems, we found pronounced geographical variation in top-down and bottom-up trophic forcing. The data suggest that ecosystem susceptibility to top-down control and their resiliency to exploitation are related to species richness and oceanic temperature conditions. Such knowledge could be used to produce ecosystem guidelines to regulate and manage fisheries in a sustainable fashion.

Reconciling differences in trophic control in mid-latitude marine ecosystems.

The dependence of long-term fishery yields on primary productivity, largely based on cross-system comparisons and without reference to the potential dynamic character of this relationship, has long been considered strong evidence for bottom-up control in marine systems. We examined time series of intensive empirical observations from nine heavily exploited regions in the western North Atlantic and find evidence of spatial variance of trophic control. Top-down control dominated in northern areas, the dynamics evolved from bottom-up to top-down in an intermediate region, and bottom-up control governed the southern areas. A simplified, trophic control diagram was developed accounting for top-down and bottom-up forcing within a larger region whose base state dynamics are bottom-up and can accommodate time-varying dynamics. Species diversity and ocean temperature co-varied, being relatively high in southern areas and lower in the north, mirroring the shifting pattern of trophic control. A combination of compensatory population dynamics and accelerated demographic rates in southern areas seems to account for the greater stability of the predator species complex in this region.