In situ observation of a macrourid fish at 7259†m in the Japan Trench: swimbladder buoyancy at extreme depth.

A macrourid, Coryphaenoides yaquinae sp. inc., was observed to be attracted to bait and exhibiting normal foraging behaviour during a period of 80 min within view of a baited video camera on the sea floor at 7259 m - the deepest ever observation of a fish species with a swim bladder. The buoyancy provided by an oxygen-filled swim bladder at 74.4 MPa pressure was estimated to be 0.164 N, at a theoretical energy cost of 20 kJ, 200 times less than the cost of equivalent lipid buoyancy. During normal metabolism, 192 days would be required to fill the swimbladder. At these depths, oxygen is very incompressible, so changes in volume during ascent or descent are small. However, swimbladder function is crucially dependent on a very low rate of diffusion of oxygen across the swimbladder wall. The oxygen in the swimbladder could theoretically sustain aerobic metabolism for over 1 year but is unlikely to be used as a reserve.

Near-equal compressibility of liver oil and seawater minimises buoyancy changes in deep-sea sharks and chimaeras.

Whereas upper ocean pelagic sharks are negatively buoyant and must swim continuously to generate lift from their fins, deep-sea sharks float or swim slowly buoyed up by large volumes of low-density oils in their livers. Investigation of the pressure, volume, temperature (PVT) relationships for liver oils of 10 species of deep-sea Chondrichthyes shows that the density difference between oil and seawater, Δρ, remains almost constant with pressure down to full ocean depth (11 km, 1100 bar), theoretically providing buoyancy far beyond the maximum depth of occurrence (3700 m) of sharks. However, Δρ does change significantly with temperature and we show that the combined effects of pressure and temperature can decrease buoyancy of oil by up to 10% between the surface and 3500 m depth across interfaces between warm southern and cold polar waters in the Rockall Trough in the NE Atlantic. This increases drag more than 10-fold compared with neutral buoyancy during horizontal slow swimming (0.1 m s-1), but the effect becomes negligible at high speeds. Chondrichthyes generally experience positive buoyancy change during ascent and negative buoyancy change during descent, but contrary effects can occur at interfaces between waters of different densities. During normal vertical migrations buoyancy changes are small, increasing slow-speed drag no more than 2- to 3-fold. Equations and tables of density, pressure and temperature are provided for squalene and liver oils of Chimaeriformes (Harriotta raleighana, Chimaera monstrosa, Hydrolagus affinis), Squaliformes (Centrophorus squamosus, Deania calcea, Centroscymnus coelolepis, Centroscyllium fabricii, Etmopterus spinax) and Carcharhiniformes (Apristurus laurussonii, Galeus murinus).

New High-Tech Flexible Networks for the Monitoring of Deep-Sea Ecosystems.

Increasing interest in the acquisition of biotic and abiotic resources from within the deep sea (e.g., fisheries, oil-gas extraction, and mining) urgently imposes the development of novel monitoring technologies, beyond the traditional vessel-assisted, time-consuming, high-cost sampling surveys. The implementation of permanent networks of seabed and water-column-cabled (fixed) and docked mobile platforms is presently enforced, to cooperatively measure biological features and environmental (physicochemical) parameters. Video and acoustic (i.e., optoacoustic) imaging are becoming central approaches for studying benthic fauna (e.g., quantifying species presence, behavior, and trophic interactions) in a remote, continuous, and prolonged fashion. Imaging is also being complemented by in situ environmental-DNA sequencing technologies, allowing the traceability of a wide range of organisms (including prokaryotes) beyond the reach of optoacoustic tools. Here, we describe the different fixed and mobile platforms of those benthic and pelagic monitoring networks, proposing at the same time an innovative roadmap for the automated computing of hierarchical ecological information on deep-sea ecosystems (i.e., from single species' abundance and life traits to community composition, and overall biodiversity).

Bathyal and abyssal demersal bait-attending fauna of the Eastern Mediterranean Sea.

Baited cameras were deployed over a depth range of 532-5111 m in the Ionian Sea to characterise the large mobile fauna. The planned installation of a neutrino telescope also offers the potential for biological observatories. The current study was intended to aid observatory placement. At increasing depths, sediment was observed to become more uniform and animal burrows and tracks reduced. A total of 10 species of deep-sea fishes were identified from images; four elasmobranchs, which were not recorded deeper than 1841 m, and six teleosts. At depths > 3000 m, including Calypso Deep, the deepest point in the Mediterranean, only one fish species was observed; the Mediterranean grenadier, Coryphaenoides mediterraneus (3400-5111 m), extending this species' maximum recorded depth to 5111 m. Four species of decapod crustacea could be identified from images. The dressed deep-sea shrimp, Acanthephyra eximia (1346-5111 m) was the only invertebrate recorded at abyssal depths, including the deepest point. A faunal change was detected at ~ 1000 m depth. Incorporating other studies from the Eastern Mediterranean identified additional faunal boundaries at ~ 1500 m and ~ 2500 m. The time from landing the observation equipment to the arrival of the first fish increased exponentially with depth at a slower rate to that observed in the Atlantic Ocean. The estimated density of bait-attending deep-sea fish was, therefore, significantly impoverished compared to the Atlantic Ocean at equivalent depth. Barriers to colonisation, low resource input, and high temperature at depth relative to the Atlantic Ocean are probable causes of the impoverished fauna.

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.

Locomotory activity and feeding strategy of the hadal munnopsid isopod Rectisura cf. herculea (Crustacea: Asellota) in the Japan Trench.

Benthic fauna in the hadal zone (6500-11,000 m) rely on maintaining sufficient locomotory activity to exploit a low, patchy and uniquely distributed food supply while exposed to high pressure, low temperatures and responding to predator-prey interactions. Very little is currently known about the locomotory capabilities of hadal fauna. In situ video footage of the isopod Rectisura cf. herculea (Birstein 1957) (Asellota, Munnopsidae) was obtained from 6945 and 7703 m deep in the Japan Trench (NW Pacific Ocean). Measurements of locomotion revealed routine walking speeds of 0.19 ± 0.04 BL s(-1) (mean ± s.d.), increasing to 0.33 ± 0.04 BL s(-1) if naturally perturbed by larger organisms. When immediately threatened by the presence of predators (decapod crustaceans), the isopods are capable of eliciting backward escape jumps and burst swimming escape responses of 2.6 ± 1.5 BL s(-1) and 4.63 ± 0.9 BL s(-1), respectively. These data suggest no significant reduction in locomotory capability despite the extreme depths in which they inhabit. These observations also revealed the isopod to be a bait-attending and aggregative species and suggest that it may not be obligatorily selecting infaunal food sources as previously thought.

Diversification of acorn worms (Hemichordata, Enteropneusta) revealed in the deep sea.

Enteropneusts (phylum Hemichordata), although studied extensively because of their close relationship to chordates, have long been considered shallow-water, burrowing animals. The present paper more than doubles the number of enteropneust species recorded in the deep sea based on high-resolution imaging and sampling with remotely operated vehicles. We provide direct evidence that some enteropneusts are highly mobile-using changes in posture and currents to drift between feeding sites-and are prominent members of deep, epibenthic communities. In addition, we provide ecological information for each species. We also show that despite their great morphological diversity, most deep-living enteropneusts form a single clade (the rediagnosed family Torquaratoridae) on the basis of rDNA sequences and morphology of the proboscis skeleton and stomochord. The phylogenetic position of the torquaratorids indicates that the group, after evolving from near-shore ancestors, radiated extensively in the deep sea.

Hadal trenches: the ecology of the deepest places on Earth.

Hadal trenches account for the deepest 45% of the oceanic depth range and host active and diverse biological communities. Advances in our understanding of hadal community structure and function have, until recently, relied on technologies that were unable to document ecological information. Renewed international interest in exploring the deepest marine environment on Earth provides impetus to re-evaluate hadal community ecology. We review the abiotic and biotic characteristics of trenches and offer a contemporary perspective of trench ecology. The application of existing, rather than the generation of novel, ecological theory offers the best prospect of understanding deep ocean ecology.

The absence of sharks from abyssal regions of the world's oceans.

The oceanic abyss (depths greater than 3000 m), one of the largest environments on the planet, is characterized by absence of solar light, high pressures and remoteness from surface food supply necessitating special molecular, physiological, behavioural and ecological adaptations of organisms that live there. Sampling by trawl, baited hooks and cameras we show that the Chondrichthyes (sharks, rays and chimaeras) are absent from, or very rare in this region. Analysis of a global data set shows a trend of rapid disappearance of chondrichthyan species with depth when compared with bony fishes. Sharks, apparently well adapted to life at high pressures are conspicuous on slopes down to 2000 m including scavenging at food falls such as dead whales. We propose that they are excluded from the abyss by high-energy demand, including an oil-rich liver for buoyancy, which cannot be sustained in extreme oligotrophic conditions. Sharks are apparently confined to ca 30% of the total ocean and distribution of many species is fragmented around sea mounts, ocean ridges and ocean margins. All populations are therefore within reach of human fisheries, and there is no hidden reserve of chondrichthyan biomass or biodiversity in the deep sea. Sharks may be more vulnerable to over-exploitation than previously thought.

High swimming and metabolic activity in the deep-sea eel Synaphobranchus kaupii revealed by integrated in situ and in vitro measurements.

Several complementary studies were undertaken on a single species of deep-sea fish (the eel Synaphobranchus kaupii) within a small temporal and spatial range. In situ experiments on swimming and foraging behaviour, muscle performance, and metabolic rate were performed in the Porcupine Seabight, northeast Atlantic, alongside measurements of temperature and current regime. Deep-water trawling was used to collect eels for studies of animal distribution and for anatomical and biochemical analyses, including white muscle citrate synthase (CS), lactate dehydrogenase (LDH), malate dehydrogenase (MDH), and pyruvate kinase (PK) activities. Synaphobranchus kaupii demonstrated whole-animal swimming speeds similar to those of other active deep-sea fish such as Antimora rostrata. Metabolic rates were an order of magnitude higher (31.6 mL kg(-1) h(-1)) than those recorded in other deep-sea scavenging fish. Activities of CS, LDH, MDH, and PK were higher than expected, and all scaled negatively with body mass, indicating a general decrease in muscle energy supply with fish growth. Despite this apparent constraint, observed in situ burst or routine swimming performances scaled in a similar fashion to other studied species. The higher-than-expected metabolic rates and activity levels, and the unusual scaling relationships of both aerobic and anaerobic metabolism enzymes in white muscle, probably reflect the changes in habitat and feeding ecology experienced during ontogeny in this bathyal species.