Francis Neat

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 km2 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.

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.

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.

A comparison of otolith microchemistry and otolith shape analysis for the study of spatial variation in a deep-sea teleost, Coryphaenoides rupestris

The study of the morphological and chemical characteristics of otoliths have recently been proposed as complementary tools for fish stock identification. However, their effectiveness remains to be fully assessed, especially in species whose life history is still poorly understood. The relative efficiency of otolith microchemistry and otolith shape analysis in discriminating samples of the deep-sea macrourid Coryphaenoides rupestris collected in different areas in the North Atlantic was examined. Otolith microchemistry based on LA/ICP-MS found significant differences in the concentrations of lithium, manganese and barium among sample sites. This allowed for very high classification accuracy (92%), when using discriminant function analysis. Otolith shape analysis based both on linear shape measurements and elliptical fourier analysis revealed a contrasting weak discrimination, with only 43% classification success. Otolith microchemistry appears to be a more effective tool in identifying individual fish from different locations. The implications for the study of population structure are discussed taking into account the limitations of the methodologies employed.

Analysing migrations of Atlantic cod Gadus morhua in the north-east Atlantic Ocean: then, now and the future

The application of data storage tags bears the potential for a quantum leap in the research on fish migrations, because not only first-capture and recapture positions are known, but at least theoretically, the migration path during the period at large can be reconstructed. Position, however, cannot be measured directly but has to be estimated using the available data on light, temperature, pressure and salinity. The reconstructed locations based on advanced estimation techniques have been termed geolocations. Examples are discussed which illustrate the applicability of geolocations in individual path descriptions, separation of reproductively isolated populations, timing and areas of spawning, tidal transport and use of protected areas. The examples are based on archival tag data from the North Sea, the Baltic Sea, the Barents Sea and Faroese and Icelandic Waters. Besides presenting the state-of-the-art geolocations for cod Gadus morhua in the north-east Atlantic Ocean, the major aim of this review is to raise awareness of gaps in knowledge and to identify ideas for new research.

A field survey of the bredding habits of Eretmodus cyanostictus, a biparental mouthbrooding cichlid in Lake Tanganyika

Oral incubation of young or ‘mouthbrooding’ reduces the selective advantages of care by two parents and thus biparental care is rare among ‘mouthbrooding’ fish. We surveyed the breeding biology of Eretmodus cyanostictus, a biparental mouthbrooder from Lake Tanganyika, to understand what factors maintain biparental care. We found larger males than females, a male-biased sex ration and indications that spawning is synchronized around the full moon. These preliminary findings suggest that the benefits of desertion for males are low; males may maximize their reproductive success by helping raise young while females regain reproductive condition.

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 (ΦST = −0.0091, FST = 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 (ΦST = 0.0744, P < 0.01; FST = 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 (ΦST = 0.3808, FST = 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 sharks 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 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.

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.