An assessment of marine, estuarine, and riverine habitat vulnerability to climate change in the Northeast U.S.

Climate change is impacting the function and distribution of habitats used by marine, coastal, and diadromous species. These impacts often exacerbate the anthropogenic stressors that habitats face, particularly in the coastal environment. We conducted a climate vulnerability assessment of 52 marine, estuarine, and riverine habitats in the Northeast U.S. to develop an ecosystem-scale understanding of the impact of climate change on these habitats. The trait-based assessment considers the overall vulnerability of a habitat to climate change to be a function of two main components, sensitivity and exposure, and relies on a process of expert elicitation. The climate vulnerability ranks ranged from low to very high, with living habitats identified as the most vulnerable. Over half of the habitats examined in this study are expected to be impacted negatively by climate change, while four habitats are expected to have positive effects. Coastal habitats were also identified as highly vulnerable, in part due to the influence of non-climate anthropogenic stressors. The results of this assessment provide regional managers and scientists with a tool to inform habitat conservation, restoration, and research priorities, fisheries and protected species management, and coastal and ocean planning.

Discovery of a spawning ground reveals diverse migration strategies in Atlantic bluefin tuna (Thunnus thynnus).

Atlantic bluefin tuna are a symbol of both the conflict between preservationist and utilitarian views of top ocean predators, and the struggle to reach international consensus on the management of migratory species. Currently, Atlantic bluefin tuna are managed as an early-maturing eastern stock, which spawns in the Mediterranean Sea, and a late-maturing western stock, which spawns in the Gulf of Mexico. However, electronic tagging studies show that many bluefin tuna, assumed to be of a mature size, do not visit either spawning ground during the spawning season. Whether these fish are spawning in an alternate location, skip-spawning, or not spawning until an older age affects how vulnerable this species is to anthropogenic stressors including exploitation. We use larval collections to demonstrate a bluefin tuna spawning ground in the Slope Sea, between the Gulf Stream and northeast United States continental shelf. We contend that western Atlantic bluefin tuna have a differential spawning migration, with larger individuals spawning in the Gulf of Mexico, and smaller individuals spawning in the Slope Sea. The current life history model, which assumes only Gulf of Mexico spawning, overestimates age at maturity for the western stock. Furthermore, individual tuna occupy both the Slope Sea and Mediterranean Sea in separate years, contrary to the prevailing view that individuals exhibit complete spawning-site fidelity. Overall, this complexity of spawning migrations questions whether there is complete independence in the dynamics of eastern and western Atlantic bluefin tuna and leads to lower estimates of the vulnerability of this species to exploitation and other anthropogenic stressors.

The Effects of Sub-Regional Climate Velocity on the Distribution and Spatial Extent of Marine Species Assemblages.

Many studies illustrate variable patterns in individual species distribution shifts in response to changing temperature. However, an assemblage, a group of species that shares a common environmental niche, will likely exhibit similar responses to climate changes, and these community-level responses may have significant implications for ecosystem function. Therefore, we examine the relationship between observed shifts of species in assemblages and regional climate velocity (i.e., the rate and direction of change of temperature isotherms). The assemblages are defined in two sub-regions of the U.S. Northeast Shelf that have heterogeneous oceanography and bathymetry using four decades of bottom trawl survey data and we explore temporal changes in distribution, spatial range extent, thermal habitat area, and biomass, within assemblages. These sub-regional analyses allow the dissection of the relative roles of regional climate velocity and local physiography in shaping observed distribution shifts. We find that assemblages of species associated with shallower, warmer waters tend to shift west-southwest and to shallower waters over time, possibly towards cooler temperatures in the semi-enclosed Gulf of Maine, while species assemblages associated with relatively cooler and deeper waters shift deeper, but with little latitudinal change. Conversely, species assemblages associated with warmer and shallower water on the broad, shallow continental shelf from the Mid-Atlantic Bight to Georges Bank shift strongly northeast along latitudinal gradients with little change in depth. Shifts in depth among the southern species associated with deeper and cooler waters are more variable, although predominantly shifts are toward deeper waters. In addition, spatial expansion and contraction of species assemblages in each region corresponds to the area of suitable thermal habitat, but is inversely related to assemblage biomass. This suggests that assemblage distribution shifts in conjunction with expansion or contraction of thermal habitat acts to compress or stretch marine species assemblages, which may respectively amplify or dilute species interactions to an extent that is rarely considered. Overall, regional differences in climate change effects on the movement and extent of species assemblages hold important implications for management, mitigation, and adaptation on the U.S. Northeast Shelf.

A Vulnerability Assessment of Fish and Invertebrates to Climate Change on the Northeast U.S. Continental Shelf.

Climate change and decadal variability are impacting marine fish and invertebrate species worldwide and these impacts will continue for the foreseeable future. Quantitative approaches have been developed to examine climate impacts on productivity, abundance, and distribution of various marine fish and invertebrate species. However, it is difficult to apply these approaches to large numbers of species owing to the lack of mechanistic understanding sufficient for quantitative analyses, as well as the lack of scientific infrastructure to support these more detailed studies. Vulnerability assessments provide a framework for evaluating climate impacts over a broad range of species with existing information. These methods combine the exposure of a species to a stressor (climate change and decadal variability) and the sensitivity of species to the stressor. These two components are then combined to estimate an overall vulnerability. Quantitative data are used when available, but qualitative information and expert opinion are used when quantitative data is lacking. Here we conduct a climate vulnerability assessment on 82 fish and invertebrate species in the Northeast U.S. Shelf including exploited, forage, and protected species. We define climate vulnerability as the extent to which abundance or productivity of a species in the region could be impacted by climate change and decadal variability. We find that the overall climate vulnerability is high to very high for approximately half the species assessed; diadromous and benthic invertebrate species exhibit the greatest vulnerability. In addition, the majority of species included in the assessment have a high potential for a change in distribution in response to projected changes in climate. Negative effects of climate change are expected for approximately half of the species assessed, but some species are expected to be positively affected (e.g., increase in productivity or move into the region). These results will inform research and management activities related to understanding and adapting marine fisheries management and conservation to climate change and decadal variability.

Long-Term Changes in the Distributions of Larval and Adult Fish in the Northeast U.S. Shelf Ecosystem.

Many studies have documented long-term changes in adult marine fish distributions and linked these changes to climate change and multi-decadal climate variability. Most marine fish, however, have complex life histories with morphologically distinct stages, which use different habitats. Shifts in distribution of one stage may affect the connectivity between life stages and thereby impact population processes including spawning and recruitment. Specifically, many marine fish species have a planktonic larval stage, which lasts from weeks to months. We compared the spatial distribution and seasonal occurrence of larval fish in the Northeast U.S. Shelf Ecosystem to test whether spatial and temporal distributions changed between two decades. Two large-scale ichthyoplankton programs sampled using similar methods and spatial domain each decade. Adult distributions from a long-term bottom trawl survey over the same time period and spatial area were also analyzed using the same analytical framework to compare changes in larval and adult distributions between the two decades. Changes in spatial distribution of larvae occurred for 43% of taxa, with shifts predominately northward (i.e., along-shelf). Timing of larval occurrence shifted for 49% of the larval taxa, with shifts evenly split between occurring earlier and later in the season. Where both larvae and adults of the same species were analyzed, 48% exhibited different shifts between larval and adult stages. Overall, these results demonstrate that larval fish distributions are changing in the ecosystem. The spatial changes are largely consistent with expectations from a changing climate. The temporal changes are more complex, indicating we need a better understanding of reproductive timing of fishes in the ecosystem. These changes may impact population productivity through changes in life history connectivity and recruitment, and add to the accumulating evidence for changes in the Northeast U.S. Shelf Ecosystem with potential to impact fisheries and other ecosystem services.

An Integrated Assessment Model for Helping the United States Sea Scallop (Placopecten magellanicus) Fishery Plan Ahead for Ocean Acidification and Warming.

Ocean acidification, the progressive change in ocean chemistry caused by uptake of atmospheric CO2, is likely to affect some marine resources negatively, including shellfish. The Atlantic sea scallop (Placopecten magellanicus) supports one of the most economically important single-species commercial fisheries in the United States. Careful management appears to be the most powerful short-term factor affecting scallop populations, but in the coming decades scallops will be increasingly influenced by global environmental changes such as ocean warming and ocean acidification. In this paper, we describe an integrated assessment model (IAM) that numerically simulates oceanographic, population dynamic, and socioeconomic relationships for the U.S. commercial sea scallop fishery. Our primary goal is to enrich resource management deliberations by offering both short- and long-term insight into the system and generating detailed policy-relevant information about the relative effects of ocean acidification, temperature rise, fishing pressure, and socioeconomic factors on the fishery using a simplified model system. Starting with relationships and data used now for sea scallop fishery management, the model adds socioeconomic decision making based on static economic theory and includes ocean biogeochemical change resulting from CO2 emissions. The model skillfully reproduces scallop population dynamics, market dynamics, and seawater carbonate chemistry since 2000. It indicates sea scallop harvests could decline substantially by 2050 under RCP 8.5 CO2 emissions and current harvest rules, assuming that ocean acidification affects P. magellanicus by decreasing recruitment and slowing growth, and that ocean warming increases growth. Future work will explore different economic and management scenarios and test how potential impacts of ocean acidification on other scallop biological parameters may influence the social-ecological system. Future empirical work on the effect of ocean acidification on sea scallops is also needed.

Decadal changes in zooplankton of the Northeast U.S. continental shelf.

The abundance of the subarctic copepod, Calanus finmarchicus, and temperate, shelf copepod, Centropages typicus, was estimated from samples collected bi-monthly over the Northeast U.S. continental shelf (NEUS) from 1977-2010. Latitudinal variation in long term trends and seasonal patterns for the two copepod species were examined for four sub-regions: the Gulf of Maine (GOM), Georges Bank (GB), Southern New England (SNE), and Mid-Atlantic Bight (MAB). Results suggested that there was significant difference in long term variation between northern region (GOM and GB), and the MAB for both species. C. finmarchicus generally peaked in May - June throughout the entire study region and Cen. typicus had a more complex seasonal pattern. Time series analysis revealed that the peak time for Cen. typicus switched from November - December to January - March after 1985 in the MAB. The long term abundance of C. finmarchicus showed more fluctuation in the MAB than the GOM and GB, whereas the long term abundance of Cen. typicus was more variable in the GB than other sub-regions. Alongshore transport was significantly correlated with the abundance of C. finmarchicus, i.e., more water from north, higher abundance for C. finmarchicus. The abundance of Cen. typicus showed positive relationship with the Gulf Stream north wall index (GSNWI) in the GOM and GB, but the GSNWI only explained 12-15% of variation in Cen. typicus abundance. In general, the alongshore current was negatively correlated with the GSNWI, suggesting that Cen. typicus is more abundant when advection from the north is less. However, the relationship between Cen. typicus and alongshore transport was not significant. The present study highlights the importance of spatial scales in the study of marine populations: observed long term changes in the northern region were different from the south for both species.

Projecting range limits with coupled thermal tolerance - climate change models: an example based on gray snapper (Lutjanus griseus) along the U.S. east coast.

We couple a species range limit hypothesis with the output of an ensemble of general circulation models to project the poleward range limit of gray snapper. Using laboratory-derived thermal limits and statistical downscaling from IPCC AR4 general circulation models, we project that gray snapper will shift northwards; the magnitude of this shift is dependent on the magnitude of climate change. We also evaluate the uncertainty in our projection and find that statistical uncertainty associated with the experimentally-derived thermal limits is the largest contributor (∼ 65%) to overall quantified uncertainty. This finding argues for more experimental work aimed at understanding and parameterizing the effects of climate change and variability on marine species.

Role of egg predation by haddock in the decline of an Atlantic herring population.

Theoretical studies suggest that the abrupt and substantial changes in the productivity of some fisheries species may be explained by predation-driven alternate stable states in their population levels. With this hypothesis, an increase in fishing or a natural perturbation can drive a population from an upper to a lower stable-equilibrium population level. After fishing is reduced or the perturbation ended, this low population level can persist due to the regulatory effect of the predator. Although established in theoretical studies, there is limited empirical support for predation-driven alternate stable states in exploited marine fish populations. We present evidence that egg predation by haddock (Melanogrammus aeglefinus) can cause alternate stable population levels in Georges Bank Atlantic herring (Clupea harengus). Egg predation by haddock explains a substantial decoupling of herring spawning stock biomass (an index of egg production) from observed larval herring abundance (an index of egg hatching). Estimated egg survival rates ranged from <2-70% from 1971 to 2005. A population model incorporating egg predation and herring fishing explains the major population trends of Georges Bank herring over four decades and predicts that, when the haddock population is high, seemingly conservative levels of fishing can still precipitate a severe decline in the herring population. These findings illustrate how efforts to rebuild fisheries can be undermined by not incorporating ecological interactions into fisheries models and management plans.

Biodiversity and ecosystem function in the Gulf of Maine: pattern and role of zooplankton and pelagic nekton.

This paper forms part of a broader overview of biodiversity of marine life in the Gulf of Maine area (GoMA), facilitated by the GoMA Census of Marine Life program. It synthesizes current data on species diversity of zooplankton and pelagic nekton, including compilation of observed species and descriptions of seasonal, regional and cross-shelf diversity patterns. Zooplankton diversity in the GoMA is characterized by spatial differences in community composition among the neritic environment, the coastal shelf, and deep offshore waters. Copepod diversity increased with depth on the Scotian Shelf. On the coastal shelf of the western Gulf of Maine, the number of higher-level taxonomic groups declined with distance from shore, reflecting more nearshore meroplankton. Copepod diversity increased in late summer, and interdecadal diversity shifts were observed, including a period of higher diversity in the 1990s. Changes in species diversity were greatest on interannual scales, intermediate on seasonal scales, and smallest across regions, in contrast to abundance patterns, suggesting that zooplankton diversity may be a more sensitive indicator of ecosystem response to inter annual climate variation than zoo plankton abundance. Local factors such as bathymetry, proximity of the coast, and advection probably drive zooplankton and pelagic nekton diversity patterns in the GoMA, while ocean-basin scale diversity patterns probably contribute to the increase in diversity at the Scotian Shelf break, a zone of mixing between the cold-temperate community of the shelf and the warm-water community offshore. Pressing research needs include establishment of a comprehensive system for observing change in zooplankton and pelagic nekton diversity, enhanced observations of "underknown" but important functional components of the ecosystem, population and metapopulation studies, and development of analytical modeling tools to enhance understanding of diversity patterns and drivers. Ultimately, sustained observations and modeling analysis of biodiversity must be effectively communicated to managers and incorporated into ecosystem approaches for management of GoMA living marine resources.

Forecasting the dynamics of a coastal fishery species using a coupled climate--population model.

Marine fisheries management strives to maintain sustainable populations while allowing exploitation. However, well-intentioned management plans may not meet this balance as most do not include the effect of climate change. Ocean temperatures are expected to increase through the 21st century, which will have far-reaching and complex impacts on marine fisheries. To begin to quantify these impacts for one coastal fishery along the east coast of the United States, we develop a coupled climate-population model for Atlantic croaker (Micropogonias undulatus). The model is based on a mechanistic hypothesis: recruitment is determined by temperature-driven, overwinter mortality of juveniles in their estuarine habitats. Temperature forecasts were obtained from 14 general circulation models simulating three CO2 emission scenarios. An ensemble-based approach was used in which a multimodel average was calculated for a given CO2 emission scenario to forecast the response of the population. The coupled model indicates that both exploitation and climate change significantly affect abundance and distribution of Atlantic croaker. At current levels of fishing, the average (2010-2100) spawning biomass of the population is forecast to increase by 60-100%. Similarly, the center of the population is forecast to shift 50 100 km northward. A yield analysis, which is used to calculate benchmarks for fishery management, indicates that the maximum sustainable yield will increase by 30 100%. Our results demonstrate that climate effects on fisheries must be identified, understood, and incorporated into the scientific advice provided to managers if sustainable exploitation is to be achieved in a changing climate.