Fishing for oil and meat drives irreversible defaunation of deepwater sharks and rays.

The deep ocean is the last natural biodiversity refuge from the reach of human activities. Deepwater sharks and rays are among the most sensitive marine vertebrates to overexploitation. One-third of threatened deepwater sharks are targeted, and half the species targeted for the international liver-oil trade are threatened with extinction. Steep population declines cannot be easily reversed owing to long generation lengths, low recovery potentials, and the near absence of management. Depth and spatial limits to fishing activity could improve conservation when implemented alongside catch regulations, bycatch mitigation, and international trade regulation. Deepwater sharks and rays require immediate trade and fishing regulations to prevent irreversible defaunation and promote recovery of this threatened megafauna group.

An analysis of arsenic concentrations associated with sargassum influx events in Barbados.

Tropical Atlantic blooms of pelagic Sargassum spp. present major socioeconomic and ecological challenges for Caribbean and West African nations. Valorisation of sargassum provides an opportunity to ameliorate some of the damage to national economies; however, the active uptake of arsenic by pelagic sargassum creates significant barriers to its use. When defining valorisation pathways, it is important to understand arsenic speciation in pelagic sargassum, given the different levels of toxicity associated with different arsenic species. In this study, we assess the temporal variability of total arsenic and inorganic arsenic in pelagic sargassum arriving in Barbados; and test whether arsenic concentrations are linked to oceanic sub-origins. Results indicate that inorganic arsenic, the most toxic form, represents a consistent and substantial percentage of the total arsenic present in pelagic sargassum, and that variability in arsenic concentration does not appear to be driven by sample months, years or oceanic sub-origins/transport pathways.

Seasonal variation in morphotype composition of pelagic Sargassum influx events is linked to oceanic origin.

The recent proliferation of pelagic Sargassum spp. in the Tropical Atlantic causes major ecological and socioeconomic impacts to the wider Caribbean when it washes ashore, with regional fisheries and tourism industries particularly affected. The Caribbean influxes have been tracked to a new bloom region known as the North Equatorial Recirculation Region (NERR) encompassing the area between the South Equatorial Current and the North Equatorial Counter Current and extending from Africa to South America. The vast biomass of Sargassum presents serious problems when it washes ashore but also represents significant commercial opportunities, especially with biofuel and fertilizer. The floating Sargassum mats are themselves diverse ecosystems that vary both in their biodiversity and biochemical attributes. Two major species (Sargassum fluitans and S. natans) have been identified as well as several distinguishable morphotypes of each. Oceanic mixing tends to blend the morphotypes together making it difficult to determine if there are regions of the NERR that favour bloom and growth of the distinct types. In this study, we quantify the species and morphotype composition of Sargassum strandings in Barbados and test if this is related to separate oceanic origins and routes travelled using a backtracking algorithm based on ocean drifter data. We found significant seasonal variation in the relative abundance of three morphotypes and this could be traced to two distinct easterly sub-origins and/or transport pathways; one area around 15° N that travels directly E-W across the Atlantic, and another area generally south of 10° N that takes a more meandering route coming close the coast of South America. These findings contribute towards our understanding of why the Tropical Atlantic bloom is presently occurring as well as towards addressing valorisation constraints surrounding variation in the supply of the three commonly occurring morphotypes.

DNA barcoding validates species labelling of certified seafood.

Seafood is one of the most traded food commodities in the world with demand steadily increasing [1]. There is, however, a rising concern over the vulnerability of seafood supply chains to species mislabelling and fraud [1,2]. DNA methods have been widely used to detect species mislabelling and a recent meta-analysis of 4500 seafood product tests from 51 publications found an average of 30 percent were not the species stated on the label or menu [3]. This high rate poses a serious threat to consumer trust, reputations of seafood businesses and the sustainability of fishery resources. Seafood certification schemes may help reduce this problem. Here, we use DNA barcoding [4] to validate the species identity of 1402 certified seafood products derived from 27 species across 18 countries and find that in over 99% of cases species labelling was correct.

Genomics of habitat choice and adaptive evolution in a deep-sea fish.

Intraspecific diversity promotes evolutionary change, and when partitioned among geographic regions or habitats can form the basis for speciation. Marine species live in an environment that can provide as much scope for diversification in the vertical as in the horizontal dimension. Understanding the relevant mechanisms will contribute significantly to our understanding of eco-evolutionary processes and effective biodiversity conservation. Here, we provide an annotated genome assembly for the deep-sea fish Coryphaenoides rupestris and re-sequencing data to show that differentiation at non-synonymous sites in functional loci distinguishes individuals living at different depths, independent of horizontal spatial distance. Our data indicate disruptive selection at these loci; however, we find no clear evidence for differentiation at neutral loci that may indicate assortative mating. We propose that individuals with distinct genotypes at relevant loci segregate by depth as they mature (supported by survey data), which may be associated with ecotype differentiation linked to distinct phenotypic requirements at different depths.

The role of the Strait of Gibraltar in shaping the genetic structure of the Mediterranean Grenadier, Coryphaenoides mediterraneus, between the Atlantic and Mediterranean Sea.

Population genetic studies of species inhabiting the deepest parts of the oceans are still scarce and only until recently we started to understand how oceanographic processes and topography affect dispersal and gene flow patterns. The aim of this study was to investigate the spatial population genetic structure of the bathyal bony fish Coryphaenoides mediterraneus, with a focus on the Atlantic-Mediterranean transition. We used nine nuclear microsatellites and the mitochondrial cytochrome c oxidase I gene from 6 different sampling areas. No population genetic structure was found within Mediterranean with both marker types (mean ΦST = 0.0960, FST = -0.0003, for both P > 0.05). However, within the Atlantic a contrasting pattern of genetic structure was found for the mtDNA and nuclear markers (mean ΦST = 0.2479, P < 0.001; FST = -0.0001, P > 0.05). When comparing samples from Atlantic and Mediterranean they exhibited high and significant levels of genetic divergence (mean ΦST = 0.7171, FST = 0.0245, for both P < 0.001) regardless the genetic marker used. Furthermore, no shared haplotypes were found between Atlantic and Mediterranean populations. These results suggest very limited genetic exchange between Atlantic and Mediterranean populations of C. mediterraneus, likely due to the shallow bathymetry of the Strait of Gibraltar acting as a barrier to gene flow. This physical barrier not only prevents the direct interactions between the deep-living adults, but also must prevent interchange of pelagic early life stages between the two basins. According to Bayesian simulations it is likely that Atlantic and Mediterranean populations of C. mediterraneus were separated during the late Pleistocene, which is congruent with results for other deep-sea fish from the same region.

Functional, size and taxonomic diversity of fish along a depth gradient in the deep sea.

Biodiversity is well studied in ecology and the concept has been developed to include traits of species, rather than solely taxonomy, to better reflect the functional diversity of a system. The deep sea provides a natural environmental gradient within which to study changes in different diversity metrics, but traits of deep-sea fish are not widely known, hampering the application of functional diversity to this globally important system. We used morphological traits to determine the functional richness and functional divergence of demersal fish assemblages along the continental slope in the Northeast Atlantic, at depths of 300-2,000 m. We compared these metrics to size diversity based on individual body size and species richness. Functional richness and size diversity showed similar patterns, with the highest diversity at intermediate depths; functional divergence showed the opposite pattern, with the highest values at the shallowest and deepest parts of the study site. Species richness increased with depth. The functional implications of these patterns were deduced by examining depth-related changes in morphological traits and the dominance of feeding guilds as illustrated by stable isotope analyses. The patterns in diversity and the variation in certain morphological traits can potentially be explained by changes in the relative dominance of pelagic and benthic feeding guilds. All measures of diversity examined here suggest that the deep areas of the continental slope may be equally or more diverse than assemblages just beyond the continental shelf.

Depth as a driver of evolution in the deep sea: Insights from grenadiers (Gadiformes: Macrouridae) of the genus Coryphaenoides.

Here we consider the role of depth as a driver of evolution in a genus of deep-sea fishes. We provide a phylogeny for the genus Coryphaenoides (Gadiformes: Macrouridae) that represents the breadth of habitat use and distributions for these species. In our consensus phylogeny species found at abyssal depths (>4000m) form a well-supported lineage, which interestingly also includes two non-abyssal species, C. striaturus and C. murrayi, diverging from the basal node of that lineage. Biogeographic analyses suggest the genus may have originated in the Southern and Pacific Oceans where contemporary species diversity is highest. The abyssal lineage seems to have arisen secondarily and likely originated in the Southern/Pacific Oceans but diversification of this lineage occurred in the Northern Atlantic Ocean. All abyssal species are found in the North Atlantic with the exception of C. yaquinae in the North Pacific and C. filicauda in the Southern Ocean. Abyssal species tend to have broad depth ranges and wide distributions, indicating that the stability of the deep oceans and the ability to live across wide depths may promote population connectivity and facilitate large ranges. We also confirm that morphologically defined subgenera do not agree with our phylogeny and that the Giant grenadier (formerly Albatrossia pectoralis) belongs to Coryphaenoides, indicating that a taxonomic revision of the genus is needed. We discuss the implications of our findings for understanding the radiation and diversification of this genus, and the likely role of adaptation to the abyss.

The economic implications of changing regulations for deep sea fishing under the European Common Fisheries Policy: UK case study.

Economic impact assessment methodology was applied to UK fisheries data to better understand the implications of European Commission proposal for regulations to fishing for deep-sea stocks in the North-East Atlantic (EC COM 371 Final 2012) under the Common Fisheries Policy (CFP). The aim was to inform the on-going debate to develop the EC proposal, and to assist the UK fishing industry and Government in evaluating the most effective options to manage deep sea fish stocks. Results indicate that enforcing the EC proposal as originally drafted results in a number of implications for the UK fleet. Because of the proposed changes to the list of species defined as being deep sea species, and a new definition of what constitutes a vessel targeting deep sea species, a total of 695 active UK fishing vessels would need a permit to fish for deep sea species. However, due to existing and capped capacity limits many vessels would potentially not be able to obtain such a permit. The economic impact of these changes from the status quo reveals that in the short term, landings would decrease by 6540 tonnes, reducing gross value added by £3.3 million. Alternative options were also assessed that provide mitigation measures to offset the impacts of the proposed regulations whilst at the same time providing more effective protection of deep sea Vulnerable Marine Ecosystems (VMEs). The options include setting a 400m depth rule that identifies a depth beyond which vessels would potentially be classified as fishing for deep sea species and designating 'core areas' for deep sea fishing at depths>400m to minimise the risk of further impacts of bottom fishing gear on deep sea habitats. Applying a 400m depth limit and 'core fishing' area approach deeper than 400m, the impact of the EC proposal would essentially be reduced to zero, that is, on average no vessels (using the status quo capacity baseline) would be impacted by the proposal.

A trait-based metric sheds new light on the nature of the body size-depth relationship in the deep sea.

Variation within species is an often-overlooked aspect of community ecology, despite the fact that the ontogenetic structure of populations influences processes right up to the ecosystem level. Accounting for traits at the individual level is an important advance in the implementation of trait-based approaches in understanding community structure and function. We incorporate individual- and species-level traits into one succinct assemblage structure metric, fractional size, which is calculated as the length of an individual divided by its potential maximum length. We test the implementation of fractional size in demersal fish assemblages along a depth gradient in the deep sea. We use data from an extensive trawl survey at depths of 300-2030 m on the continental slope of the Rockall Trough, Northeast Atlantic, to compare changes in fractional size structure along an environmental gradient to those seen using traditional taxonomic and trait-based approaches. The relationship between fractional size and depth was particularly strong, with the overall pattern being an increase with depth, implying that individuals move deeper as they grow. Body size increased with depth at the intraspecific and assemblage levels. Fractional size, size structure and species composition all varied among assemblages, and this variation could be explained by the depth that the assemblage occupied. The inclusion of individual-level traits and population fractional size structure adds to our understanding at the assemblage level. Fractional size, or where an individual is in its growth trajectory, appears to be an especially important driver of assemblage change with depth. This has implications for understanding fisheries impacts in the deep sea and how these impacts may propagate across depths.

The Pillars of Hercules as a bathymetric barrier to gene flow promoting isolation in a global deep-sea shark (Centroscymnus coelolepis).

Knowledge of the mechanisms limiting connectivity and gene flow in deep-sea ecosystems is scarce, especially for deep-sea sharks. The Portuguese dogfish (Centroscymnus coelolepis) is a globally distributed and near threatened deep-sea shark. C. coelolepis population structure was studied using 11 nuclear microsatellite markers and a 497-bp fragment from the mtDNA control region. High levels of genetic homogeneity across the Atlantic (Φ(ST) = -0.0091, F(ST) = 0.0024, P > 0.05) were found suggesting one large population unit at this basin. The low levels of genetic divergence between Atlantic and Australia (Φ(ST) = 0.0744, P < 0.01; F(ST) = 0.0015, P > 0.05) further suggested that this species may be able to maintain some degree of genetic connectivity even across ocean basins. In contrast, sharks from the Mediterranean Sea exhibited marked genetic differentiation from all other localities studied (Φ(ST) = 0.3808, F(ST) = 0.1149, P < 0.001). This finding suggests that the shallow depth of the Strait of Gibraltar acts as a barrier to dispersal and that isolation and genetic drift may have had an important role shaping the Mediterranean shark population over time. Analyses of life history traits allowed the direct comparison among regions providing a complete characterization of this shark's populations. Sharks from the Mediterranean had markedly smaller adult body size and size at maturity compared to Atlantic and Pacific individuals. Together, these results suggest the existence of an isolated and unique population of C. coelolepis inhabiting the Mediterranean that most likely became separated from the Atlantic in the late Pleistocene.

A Scientific Basis for Regulating Deep-Sea Fishing by Depth.

The deep sea is the world's largest ecosystem, with high levels of biodiversity and many species that exhibit life-history characteristics that make them vulnerable to high levels of exploitation. Many fisheries in the deep sea have a track record of being unsustainable. In the northeast Atlantic, there has been a decline in the abundance of commercial fish species since deep-sea fishing commenced in the 1970s. Current management is by effort restrictions and total allowable catch (TAC), but there remain problems with compliance and high levels of bycatch of vulnerable species such as sharks. The European Union is currently considering new legislation to manage deep-sea fisheries, including the introduction of a depth limit to bottom trawling. However, there is little evidence to suggest an appropriate depth limit. Here we use survey data to show that biodiversity of the demersal fish community, the ratio of discarded to commercial biomass, and the ratio of Elasmobranchii (sharks and rays) to commercial biomass significantly increases between 600 and 800 m depth while commercial value decreases. These results suggest that limiting bottom trawling to a maximum depth of 600 m could be an effective management strategy that would fit the needs of European legislations such as the Common Fisheries Policy (EC no. 1380/2013) and the Marine Strategy Framework Directive (2008/56/EC).

Halogenated persistent organic pollutants in relation to trophic level in deep sea fish.

The bioaccumulation of persistent organic pollutants (POPs) in deep sea fish from the Rockall fishing area was investigated. Predator and prey species were analysed for stable isotopes, fatty acids, polychlorinated biphenyls (PCBs) and polybrominated diphenyl ethers (PBDEs). δ(15)N indicated that black scabbard was at the highest trophic level and the prey the lowest. The fatty acid signatures indicated that black scabbard and black dogfish fed at a higher trophic level compared to the roundnose grenadier. PCBs and PBDEs were detected in the liver of all three predator species. PCB concentrations were significantly higher in the roundnose grenadier, possibly due to their longer life span. PCB concentrations were compared to OSPAR assessment criteria, concentrations were above background but below Environmental Assessment Criteria for all but one congener. PCB concentrations were below food safety levels in the flesh, but exceeded the limit for liver in the roundnose grenadier and black dogfish.

A tale of two seas: contrasting patterns of population structure in the small-spotted catshark across Europe.

Elasmobranchs represent important components of marine ecosystems, but they can be vulnerable to overexploitation. This has driven investigations into the population genetic structure of large-bodied pelagic sharks, but relatively little is known of population structure in smaller demersal taxa, which are perhaps more representative of the biodiversity of the group. This study explores spatial population genetic structure of the small-spotted catshark (Scyliorhinus canicula), across European seas. The results show significant genetic differences among most of the Mediterranean sample collections, but no significant structure among Atlantic shelf areas. The data suggest the Mediterranean populations are likely to have persisted in a stable and structured environment during Pleistocene sea-level changes. Conversely, the Northeast Atlantic populations would have experienced major changes in habitat availability during glacial cycles, driving patterns of population reduction and expansion. The data also provide evidence of male-biased dispersal and female philopatry over large spatial scales, implying complex sex-determined differences in the behaviour of elasmobranchs. On the basis of this evidence, we suggest that patterns of connectivity are determined by trends of past habitat stability that provides opportunity for local adaptation in species exhibiting philopatric behaviour, implying that resilience of populations to fisheries and other stressors may differ across the range of species.

Does presence of a mid-ocean ridge enhance biomass and biodiversity?

In contrast to generally sparse biological communities in open-ocean settings, seamounts and ridges are perceived as areas of elevated productivity and biodiversity capable of supporting commercial fisheries. We investigated the origin of this apparent biological enhancement over a segment of the North Mid-Atlantic Ridge (MAR) using sonar, corers, trawls, traps, and a remotely operated vehicle to survey habitat, biomass, and biodiversity. Satellite remote sensing provided information on flow patterns, thermal fronts, and primary production, while sediment traps measured export flux during 2007-2010. The MAR, 3,704,404 km(2) in area, accounts for 44.7% lower bathyal habitat (800-3500 m depth) in the North Atlantic and is dominated by fine soft sediment substrate (95% of area) on a series of flat terraces with intervening slopes either side of the ridge axis contributing to habitat heterogeneity. The MAR fauna comprises mainly species known from continental margins with no evidence of greater biodiversity. Primary production and export flux over the MAR were not enhanced compared with a nearby reference station over the Porcupine Abyssal Plain. Biomasses of benthic macrofauna and megafauna were similar to global averages at the same depths totalling an estimated 258.9 kt C over the entire lower bathyal north MAR. A hypothetical flat plain at 3500 m depth in place of the MAR would contain 85.6 kt C, implying an increase of 173.3 kt C attributable to the presence of the Ridge. This is approximately equal to 167 kt C of estimated pelagic biomass displaced by the volume of the MAR. There is no enhancement of biological productivity over the MAR; oceanic bathypelagic species are replaced by benthic fauna otherwise unable to survive in the mid ocean. We propose that globally sea floor elevation has no effect on deep sea biomass; pelagic plus benthic biomass is constant within a given surface productivity regime.

Halogenated persistent organic pollutants in deep water fish from waters to the west of Scotland.

Halogenated persistent organic pollutants [polychlorinated biphenyls (PCBs) and polybrominated diphenyl ethers (PBDEs)] along with total lipid, were measured in the liver and muscle of three species of deep water fish (black scabbard, black dogfish (liver only) and roundnose grenadier) collected from the Rockall fishing area, to the west of Scotland, between 2006 and 2008. Both contaminant groups were detected in the muscle and liver, with concentrations of PCBs being higher than PBDEs. There were no significant differences in the PCB or PBDE concentrations between the three species, or different sampling locations in the Rockall fishing area. PCB concentrations (ΣICES (International Council for the Exploration of the Sea)7 PCBs) greater than 500 µg kg(-1) lipid weight were found in 26 of the 106 liver samples. PCB concentrations were compared to OSPAR assessment criteria, concentrations were above background but below Environmental Assessment Criteria. Estimated Toxic Equivalent (TEQ) concentrations, calculated using published models, in the fish muscle and liver indicated that consumption of deep water fish is unlikely to represent a risk to human health. The high squalene content in some of the black dogfish liver necessitated an additional clean-up step, involving gel permeation chromatography, when analyzing for PBDEs. Concentrations of PBDEs were low with many congeners being below detection limits, particularly in the muscle. There are currently no assessment criteria available for PBDEs. Furthermore, there is only very limited data on PBDEs in deep water fish. However, the concentrations observed in this study were similar to the concentrations recently reported in Mediterranean deep water fish.

Molecular markers reveal spatially segregated cryptic species in a critically endangered fish, the common skate (Dipturus batis).

Many sharks and skates are particularly vulnerable to overfishing because of their large size, slow growth, late maturity and low fecundity. In Europe dramatic population declines have taken place in common skate (Dipturus batis L.), one of the largest demersal fish in regional shelf seas, leading to extirpations from substantial parts of its former range. Here we report the discovery of cryptic species in common skate collected from the northeast Atlantic continental shelf. Data from nuclear microsatellite markers indicated two clearly distinct clades and phylogenetic analysis of mitochondrial DNA sequences demonstrated monophyly of each one of them. Capture locations showed evidence of strong spatial segregation, with one taxon occurring mainly in waters off the southern British Isles and around Rockall, while the other was restricted to more northerly shelf waters. These apparently cryptic species showed overlapping substrate and depth preferences, but distributional limits were closely related to temperature gradients, potentially indicating thermal limits to their distributions. This discovery of hidden diversity within a large, critically endangered marine vertebrate demonstrates how marine biodiversity can be underestimated, even in such a relatively well-studied and heavily exploited region.

Halogenated persistent organic pollutants in Scottish deep water fish.

Halogenated persistent organic pollutants (chlorobiphenyls (CBs), polybrominated diphenyl ethers (PBDEs), hexabromocyclododecane (HBCD) and tetrabromobisphenol-A (TBBP-A)) and total lipid content were measured in the liver and muscle of three species of deep water fish (black scabbard, roundnose grenadier and black dogfish) collected from the Rockall Trough, to the west of Scotland, in 2006. CB concentrations (SigmaICES (International Council for the Exploration of the Seas)7 CBs) >500 microg kg(-1) lipid weight) were found in 9 of the 31 deep water fish liver samples. Non-ortho CBs were measured in samples with the highest ortho CB concentrations. Non-ortho CBs (CB81, 77, 126 and 169) were not detected in any of the fish muscle samples. In liver, CB81 was not detected in any of the samples while CB169 was detected in all but one sample. The total 'dioxin-like' CB concentration was calculated based on the 5 mono-ortho and 4 non-ortho CBs measured. The non-ortho CB concentration made a very small contribution to the total 'dioxin-like' CB concentrations (<1%). Concentrations for the individual ICES7 CBs in fish liver were above OSPAR Background Assessment Concentrations (BACs) in all three species, except for CB28 and 101 in black dogfish. Toxic Equivalent (TEQs) concentrations calculated for the five mono-ortho and four non-ortho CBs measured, and estimated TEQs calculated using published models in the fish muscle indicated that consumption of deep water fish muscle is unlikely to represent a risk to human health. However, dioxins and furans were not measured and the contribution to the calculated TEQs from these compounds was not taken into account. Calculated and estimated TEQs for some roundnose grenadier liver samples exceeded the 25 pg g(-1) wet weight limit for fish liver and, therefore, there may be a health risk if consumed. PBDEs were detected in both the liver and muscle of the deep water fish, whilst HBCD and TBBP-A were not detected in any of the deep water fish.

Warm water occupancy by North Sea cod.

The North Sea has warmed in recent years and there is an ongoing debate into how this is affecting the distribution of fishes and other marine organisms. Of particular interest is the commercially important Atlantic cod (Gadus morhua L.), which has declined sharply in abundance in the North Sea over the past 20 years. Observations of the temperature experienced by 129 individual cod throughout the North Sea were made during a large-scale electronic tagging programme conducted between 1999 and 2005. We asked whether individual cod fully occupied the thermal habitat available to them. To this end, we compared the temperature experience of cod with independently measured contemporaneous sea-bottom temperature data. The majority of cod experienced a warmer fraction of the sea than was potentially available to them. By summer, most of the individuals in the south experienced temperatures considered superoptimal for growth. Cooler waters were within the reach of the cod and a small number of individuals migrated to areas that allowed them to experience lower temperatures, indicating that the cod had the capacity to find cooler water. Most did not, however, suggesting that the changing thermal regime of the North Sea is not yet causing adult cod to move to cooler waters.

Reproductive allocation in Aidablennius sphynx (Teleostei, Blenniidae): females lay more eggs faster when paired with larger males.

Individuals are expected to invest more in current reproductive effort when paired with a partner of higher than average quality. Aidablennius sphynx is an external fertilizing fish with paternal care in which females gain direct benefits from spawning with large males, but often 'make do' with small males. In this study, female reproductive responses to large and small males were investigated. When paired with large males, females spawned more eggs per unit time (i.e., at a faster rate). There was no difference in the size of the eggs spawned by females in relation to partner size. By ovipositing at a faster rate, females may have allocated more reproductive effort to large males. In addition, since small males are known to release far fewer sperm than large males, females may have reduced their spawn rate with small males as a tactic to ensure fertilization.