Despite the deep sea being the largest habitat on Earth, there are just 77 population genetic studies of invertebrates (115 species) inhabiting non-chemosynthetic ecosystems on the deep-sea floor (below 200 m depth). We review and synthesize the results of these papers. Studies reveal levels of genetic diversity comparable to shallow-water species. Generally, populations at similar depths were well connected over 100s–1,000s km, but studies that sampled across depth ranges reveal population structure at much smaller scales (100s–1,000s m) consistent with isolation by adaptation across environmental gradients, or the existence of physical barriers to connectivity with depth. Few studies were ocean-wide (under 4%), and 48% were Atlantic-focused. There is strong emphasis on megafauna and commercial species with research into meiofauna, “ecosystem engineers” and other ecologically important species lacking. Only nine papers account for ~50% of the planet's surface (depths below 3,500 m). Just two species were studied below 5,000 m, a quarter of Earth's seafloor. Most studies used single-locus mitochondrial genes revealing a common pattern of non-neutrality, consistent with demographic instability or selective sweeps; similar to deep-sea hydrothermal vent fauna. The absence of a clear difference between vent and non-vent could signify that demographic instability is common in the deep sea, or that selective sweeps render single-locus mitochondrial studies demographically uninformative. The number of population genetics studies to date is miniscule in relation to the size of the deep sea. The paucity of studies constrains meta-analyses where broad inferences about deep-sea ecology could be made.
Deep-sea ecosystems and hydrothermal vents
Microplastics are widespread in the natural environment and present numerous ecological threats. While the ultimate fate of marine microplastics are not well known, it is hypothesized that the deep sea is the final sink for this anthropogenic contaminant. This study provides a quantification and characterisation of microplastic pollution ingested by benthic macroinvertebrates with different feeding modes (Ophiomusium lymani, Hymenaster pellucidus and Colus jeffreysianus) and in adjacent deep water > 2200 m, in the Rockall Trough, Northeast Atlantic Ocean. Despite the remote location, microplastic fibres were identified in deep-sea water at a concentration of 70.8 particles m−3, comparable to that in surface waters. Of the invertebrates examined (n = 66), 48% ingested microplastics with quantities enumerated comparable to coastal species. The number of ingested microplastics differed significantly between species and generalized linear modelling identified that the number of microplastics ingested for a given tissue mass was related to species and not organism feeding mode or the length or overall weight of the individual. Deep-sea microplastics were visually highly degraded with surface areas more than double that of pristine particles. The identification of synthetic polymers with densities greater and less than seawater along with comparable quantities to the upper ocean indicates processes of vertical re-distribution. This study presents the first snapshot of deep ocean microplastics and the quantification of microplastic pollution in the Rockall Trough. Additional sampling throughout the deep-sea is required to assess levels of microplastic pollution, vertical transportation and sequestration, which have the potential to impact the largest global ecosystem.
The apparent prevalence of rare species (rarity) in the deep sea is a concern for environmental management and conservation of biodiversity. Rare species are often considered at risk of extinction and, in terrestrial and shallow water environments, have been shown to play key roles within an ecosystem. In the deep-sea environment, current research focuses primarily on abundant species and deep-sea stakeholders are questioning the importance of rare species in ecosystem functioning. This study asks whether deep-sea stakeholders (primarily scientists) view rare-species research as a priority in guiding environmental management. Delphi methodology (i.e., an iterative survey approach) was used to understand views about whether or not ‘deep-sea scientists should allocate more resources to research on rare species in the deep sea, even if this means less resources might be available for abundant-species research.’ Results suggest little consensus regarding the prioritization of resources for rare-species research. From Survey 1 to Survey 3, the average participant response shifted toward a view that rare-species research is not a priority if it comes at a cost to research on abundant species. Participants pointed to the need for a balanced approach and highlighted knowledge gaps about even the most fundamental questions, including whether rare species are truly ‘rare’ or simply under-sampled. Participants emphasized the lack of basic biological knowledge for rare and abundant species, particularly abundant meio- and microscopic species, as well as uncertainty in the roles rare and abundant species play in ecosystem processes. Approaches that jointly consider the role of rare and abundant species in ecosystem functioning (e.g., biological trait analysis) may help to clarify the extent to which rare species need to be incorporated into deep-sea environment management in order to maintain ecosystem functioning.
The deep-sea is a large source of marine genetic resources (MGR), which have many potential uses and are a growing area of research. Much of the deep-sea lies in areas beyond national jurisdiction (ABNJ), including 65% of the global ocean. MGR in ABNJ occupy a significant gap in the international legal framework. Access and benefit sharing of MGR is a key issue in the development of a new international legally-binding instrument under the United Nations Convention on the Law of the Sea (UNCLOS) for the conservation and sustainable use of marine biological diversity in ABNJ. This paper examines how this is relevant to deep-sea scientific research and identifies emerging challenges and opportunities. There is no internationally agreed definition of MGR, however, deep-sea genetic resources could incorporate any biological material including genes, proteins and natural products. Deep-sea scientific research is the key actor accessing MGR in ABNJ and sharing benefits such as data, samples and knowledge. UNCLOS provides the international legal framework for marine scientific research, international science cooperation, capacity building and marine technology transfer. Enhanced implementation could support access and benefit sharing of MGR in ABNJ. Deep-sea scientific researchers could play an important role in informing practical new governance solutions for access and benefit sharing of MGR that promote scientific research in ABNJ and support deep-sea stewardship. Advancing knowledge of deep-sea biodiversity in ABNJ, enhancing open-access to data and samples, standardisation and international marine science cooperation are significant potential opportunity areas.
Cold-water coral (CWC) habitats can form complex structures which provide refuge, nursery grounds and physical support for a diversity of other living organisms, but despite their ecological significance, CWCs are still vulnerable to human pressures such as fishing, pollution, ocean acidification and global warming
Providing coherent and representative conservation of vulnerable marine ecosystems including CWCs is one of the aims of the Marine Protected Areas networks being implemented across European seas and oceans under the EC Habitats Directive, the Marine Strategy Framework Directive and the OSPAR Convention. In order to adequately represent ecosystem diversity these initiatives require a standardised habitat classification that organises the variety of biological assemblages and provides consistent and functional criteria to map them across European Seas (Howell 2010). One such classification system, EUNIS, enables a broad level classification of the deep sea based on abiotic and geomorphological features. More detailed lower biotope-related levels are currently under-developed, particularly with regards deep-water habitats (>200 m depth).
This paper proposes a hierarchical CWC biotope classification scheme that could be incorporated by existing classification schemes such as EUNIS. The scheme was developed within the EU FP7 project CoralFISH to capture the variability of CWC habitats identified using a wealth of seafloor imagery datasets from across European seas and oceans. Depending on the resolution of the imagery being interpreted, this hierarchical scheme allows data to be recorded from broad CWC biotope categories down to detailed taxonomy-based levels, thereby providing a flexible yet valuable information level for management. The CWC biotope classification scheme identifies 81 biotopes and highlights the limitations of the classification framework and guidance provided by EUNIS, the EC Habitats Directive, OSPAR and FAO; with limited categories for identifying and classifying these CWC habitats.
The deep sea encompasses the largest ecosystems on Earth. Although poorly known, deep seafloor ecosystems provide services that are vitally important to the entire ocean and biosphere. Rising atmospheric greenhouse gases are bringing about significant changes in the environmental properties of the ocean realm in terms of water column oxygenation, temperature, pH and food supply, with concomitant impacts on deep-sea ecosystems. Projections suggest that abyssal (3000–6000 m) ocean temperatures could increase by 1°C over the next 84 years, while abyssal seafloor habitats under areas of deep-water formation may experience reductions in water column oxygen concentrations by as much as 0.03 mL L–1 by 2100. Bathyal depths (200–3000 m) worldwide will undergo the most significant reductions in pH in all oceans by the year 2100 (0.29 to 0.37 pH units). O2 concentrations will also decline in the bathyal NE Pacific and Southern Oceans, with losses up to 3.7% or more, especially at intermediate depths. Another important environmental parameter, the flux of particulate organic matter to the seafloor, is likely to decline significantly in most oceans, most notably in the abyssal and bathyal Indian Ocean where it is predicted to decrease by 40–55% by the end of the century. Unfortunately, how these major changes will affect deep-seafloor ecosystems is, in some cases, very poorly understood. In this paper, we provide a detailed overview of the impacts of these changing environmental parameters on deep-seafloor ecosystems that will most likely be seen by 2100 in continental margin, abyssal and polar settings. We also consider how these changes may combine with other anthropogenic stressors (e.g., fishing, mineral mining, oil and gas extraction) to further impact deep-seafloor ecosystems and discuss the possible societal implications.
The exploration of deep-sea mineral resources on continental margins is increasing worldwide. In the SW Atlantic, Campos Basin has been Brazil's main deep-sea area for oil and gas extraction since the 1980′s, with currently over 11,000 km2 of leased blocks below 200 m depth. The historical record of exploration and the lack of a basin-wide management for the offshore industry in the SW Atlantic threaten the biodiversity and ecological function of vulnerable deep-sea ecosystems. This study identified habitats of biological interest on the Campos Basin and proposes relevant areas for conservation (EBSAs) that could be included in the first deep-sea Marine Protected Area (MPA) network in the South Atlantic. A total of 42 benthic habitats were mapped including cold-water coral reefs, submarine canyons, soft sediment slope and a seamount. Those habitats fill conservation criteria to be proposed as EBSAs along Campos Basin and could support a MPA network with a 5.5% overlap (2330 km2) to current leased blocks. If implemented, the MPA network would cover 31% of the Campos Basin and offer 31–100% protection of EBSAs with minimal interference on industry. This study is the first to identify EBSAs in a deep-sea basin of major economic importance in Brazil's EEZ and their conservation would also protect areas at two biogeographic provinces in the South Atlantic. Furthermore, the methods demonstrated here could be widely applied to other offshore oil and gas areas that lack environmental management measures at early stages of bidding rounds or during the process of environmental licensing.
Sponge aggregations have been recognised as key component of shallow benthic ecosystems providing several important functional roles including habitat building and nutrient recycling. Within the deep-sea ecosystem, sponge aggregations may be extensive and available evidence suggests they may also play important functional roles, however data on their ecology, extent and distribution in the North Atlantic is lacking, hampering conservation efforts. In this study, we used Maximum Entropy Modelling and presence data for two deep-sea sponge aggregation types, Pheronema carpenteriaggregations and ostur aggregations dominated by geodid sponges, to address the following questions: 1) What environmental factors drive the broad-scale distribution of these selected sponge grounds? 2) What is the predicted distribution of these grounds in the northern North Atlantic, Norwegian and Barents Sea? 3) How are these sponge grounds distributed between Exclusive Economic Zones (EEZs) and High Seas areas? 4) What percentage of these grounds in High Seas areas are protected by the current High Seas MPA network? Our results suggest that silicate concentration, temperature, depth and amount of particulate organic carbon are the most important drivers of sponge distribution. Most of the sponge grounds are located within national EEZs rather than in the High Seas. Coordinated conservation planning between nations with significant areas of sponge grounds such as Iceland, Greenland and Faroes (Denmark), Norway (coastal Norway and Svalbard), Portugal and the UK, should be implemented in order to effectively manage these communities in view of the increasing level of human activity within the deep-sea environment.
The deep soft sediment Black Sea benthic community is dominated by cold seep habitats formed by the microbial breakdown of phytoplankton. The deep Black Sea benthic ecosystem is chemosynthetic with methanogenesis and the sulfate-driven anaerobic oxidation of methane acting as the primary metabolic pathways. Due to the depth and lack of metazoan life the deep Black Sea benthic ecosystem is generally regarded to be at low risk from anthropogenic impact and has little legislation pertaining directly to the preservation of the chemosynthetic habitats. The principal ecosystem services provided by the Black Sea include carbon sequestration and preservation of historical artefacts. Compared to other ecosystems, information on microbial biodiversity and ecosystem services in the deep Black Sea is lacking, and we highlight a need to plan and implement research programmes to address significant gaps and to enhance scientific understanding of this environment.
With anthropogenic impacts rapidly advancing into deeper waters, there is growing interest in establishing deep-sea marine protected areas (MPAs) or reserves. Reserve design depends on estimates of connectivity and scales of dispersal for the taxa of interest. Deep-sea taxa are hypothesized to disperse greater distances than shallow-water taxa, which implies that reserves would need to be larger in size and networks could be more widely spaced; however, this paradigm has not been tested. We compiled population genetic studies of deep-sea fauna and estimated dispersal distances for 51 studies using a method based on isolation-by-distance slopes. Estimates of dispersal distance ranged from 0.24 km to 2028 km with a geometric mean of 33.2 km and differed in relation to taxonomic and life-history factors as well as several study parameters. Dispersal distances were generally greater for fishes than invertebrates with the Mollusca being the least dispersive sampled phylum. Species that are pelagic as adults were more dispersive than those with sessile or sedentary lifestyles. Benthic species from soft-substrate habitats were generally less dispersive than species from hard substrate, demersal or pelagic habitats. As expected, species with pelagic and/or feeding (planktotrophic) larvae were more dispersive than other larval types. Many of these comparisons were confounded by taxonomic or other life-history differences (e.g. fishes being more dispersive than invertebrates) making any simple interpretation difficult. Our results provide the first rough estimate of the range of dispersal distances in the deep sea and allow comparisons to shallow-water assemblages. Overall, dispersal distances were greater for deeper taxa, although the differences were not large (0.3–0.6 orders of magnitude between means), and imbalanced sampling of shallow and deep taxa complicates any simple interpretation. Our analyses suggest the scales of dispersal and connectivity for reserve design in the deep sea might be comparable to or slightly larger than those in shallow water. Deep-sea reserve design will need to consider the enormous variety of taxa, life histories, hydrodynamics, spatial configuration of habitats and patterns of species distributions. The many caveats of our analyses provide a strong impetus for substantial future efforts to assess connectivity of deep-sea species from a variety of habitats, taxonomic groups and depth zones.