No fish have been found in the deepest 25% of the ocean (8,400-11,000 m). This apparent absence has been attributed to hydrostatic pressure, although direct evidence is wanting because of the lack of deepest-living species to study. The common osmolyte trimethylamine N-oxide (TMAO) stabilizes proteins against pressure and increases with depth, going from 40 to 261 mmol/kg in teleost fishes from 0 to 4,850 m. TMAO accumulation with depth results in increasing internal osmolality (typically 350 mOsmol/kg in shallow species compared with seawater's 1,100 mOsmol/kg). Preliminary extrapolation of osmolalities of predicted isosmotic state at 8,000-8,500 m may indicate a possible physiological limit, as greater depths would require reversal of osmotic gradients and, thus, osmoregulatory systems. We tested this prediction by capturing five of the second-deepest known fish, the hadal snailfish (Notoliparis kermadecensis; Liparidae), from 7,000 m in the Kermadec Trench. We found their muscles to have a TMAO content of 386 ± 18 mmol/kg and osmolality of 991 ± 22 mOsmol/kg. These data fit previous extrapolations and, combined with new osmolalities from bathyal and abyssal fishes, predict isosmotic state at 8,200 m. This is previously unidentified evidence that biochemistry could constrain the depth of a large, complex taxonomic group.deep-sea | piezolyte | chemical chaperone | osmoregulation

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Baited landers were deployed at 83 stations at four locations in the west Pacific Ocean from bathyal to hadal depths: The Kermadec Trench, the New Hebrides Trench, the adjoining South Fiji Basin and the Mariana Trench. Forty-seven putative fish species were observed. Distinct fish faunal groups were identified based on maximum numbers and percentage of observations. Both analyses broadly agreed on the community structure: A bathyal group at <3000 m in the New Hebrides and Kermadec trenches, an abyssal group (3039-4692 m) in the Kermadec Trench, an abyssal-hadal transition zone (AHTZ) group (Kermadec: 4707-6068 m, Mariana: 4506-6198 m, New Hebrides: 2578-6898 m, South Fiji Basin: 4074-4101 m), and a hadal group of endemic snailfish in the Kermadec and Mariana trenches (6750-7669 m and 6831-8143 m respectively). The abyssal and hadal groups were absent from the New Hebrides Trench. Depth was the single factor that best explained the biological variation between samples (16%), the addition of temperature and average surface primary production for the previous year increased this to 36% of variation. The absence of the abyssal group from the New Hebrides Trench and South Fiji Basin was due to the absence of macrourids (Coryphaenoides spp.), which defined the group. The macrourids may be energetically limited in these areas. In their absence the species of the AHTZ group appear released of competition with the macrourids and are found far shallower at these sites. The fish groups had distinct feeding strategies while attending the bait: The bathyal and abyssal groups were almost exclusively necrophagous, the AHTZ group comprised predatory and generalist feeders, while the hadal snailfishes were exclusively predators. With increasing depth, predation was found to increase while scavenging decreased. The data suggest scavenging fish fauna do not extend deeper than the hadal boundary.

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Pseudoliparis swirei sp. nov. is described from 37 individuals collected in the Mariana Trench at depths 6898-7966 m. The collection of this new species is the deepest benthic capture of a vertebrate with corroborated depth data. Here, we describe P. swirei sp. nov. and discuss aspects of its morphology, biology, distribution, and phylogenetic relationships to other hadal liparids based on analysis of three mitochondrial genes. Pseudoliparis swirei sp. nov. is almost certainly endemic to the Mariana Trench, as other hadal liparids appear isolated to a single trench/ trench system in the Kermadec, Macquarie, South Sandwich, South Orkney, Peru-Chile, Kurile-Kamchatka and Japan trenches. The discovery of another hadal liparid species, apparently abundant at depths where other fish species are few and only found in low numbers, provides further evidence for the dominance of this family among the hadal fish fauna.

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Synopsis Deep-sea trenches, depths 6000–11,000 m, are characterized by high pressures, low temperatures, and absence of sunlight. These features make up the majority of the deepest marine habitat—the hadal zone—home to distinct communities from those in the surrounding abyssal plains. The snailfishes, family Liparidae (Scorpaeniformes), have found notable success in the hadal zone from ∼6000 to 8200 m, comprising the dominant ichthyofauna in at least six trenches worldwide. The hadal fish community is distinct from the abyssal community where elongate, scavenging fishes such as rattails (Macrouridae), cutthroat eels (Synaphobranchidae), tripodfishes (Ipnopidae), eelpouts (Zoarcidae), and cusk eels (Ophidiidae) are most common. Until recently, little was known about the biology of these deepest-living fishes, or the factors that drive their success at hadal depths. Here, I review recent investigations spanning the abyssal–hadal boundary and discuss the factors structuring these communities, including the roles of pressure adaptation, feeding ecology, and life history. Hadal fishes show specialized adaptation to hydrostatic pressure both in accumulation of the pressure-counteractant trimethylamine n-oxide and in intrinsic changes to enzymes. Stomach content and amino acid isotope analyses, and jaw morphology suggest that suction-feeding predatory fishes like hadal liparids may find an advantage to descending into the trench where amphipods are increasingly abundant. Analysis of otolith growth zones suggest that snailfishes may be adapted to a seismically active, high-disturbance hadal environment by having relatively short life-spans. This review synthesizes the known literature on the planet’s deepest-living fishes and informs new understanding of adaptations to life in the trenches.

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