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Snailfish

Snailfishes, comprising the family Liparidae within the order Cottiformes, are a diverse group of ray-finned fishes renowned for their extraordinary depth range and adaptations to environments. With over 450 across approximately 31 genera, they exhibit elongate, tadpole-like bodies covered in scaleless, gelatinous skin that provides and pressure resistance. These fishes inhabit oceans worldwide, from and polar regions to temperate waters in , Pacific, and Indian Oceans, spanning latitudes and longitudes broadly. They occupy an unparalleled bathymetric range among vertebrates, from shallow intertidal tidepools and coastal zones to the hadal depths exceeding 8,000 meters in ocean trenches, with a Pseudoliparis snailfish recorded as the deepest-living bony at a depth of 8,336 meters in the Izu-Ogasawara Trench. Physically, snailfishes feature a soft, cartilaginous lacking a , which aids survival under immense hydrostatic pressures in deep-sea habitats. Many species have pelvic fins fused into a ventral suction disk for adhering to substrates like rocks, corals, or the seafloor, while their , anal, and caudal fins often merge into a continuous marginal . Their loose, jelly-like may bear small prickles in some taxa, and body sizes vary from under 10 cm to over 30 cm, though most are small and poorly suited for sustained , preferring to or drift with bottom currents. Ecologically, snailfishes are primarily benthic or demersal, with deep-water forms showing specialized traits like translucent bodies for in low-light conditions and enzymes in muscles that function under high pressure. They feed as carnivores on small benthic and pelagic , including amphipods, copepods, worms, and decapod , which can constitute the bulk of their ; larval stages consume planktonic prey. As key components of cold-water food webs, they serve as predators of and prey for larger fishes and marine mammals, contributing to across shallow coastal and abyssal ecosystems. Recent discoveries, such as three new abyssal described in 2025, highlight ongoing explorations revealing their morphological diversity and evolutionary adaptations.

Taxonomy and Classification

Evolutionary History

The family Liparidae belongs to the suborder Cottoidei within the order Cottiformes (traditionally classified under ), where it forms a monophyletic group sister to the . This placement reflects key divergences from other cottoid families, such as the Cottidae (sculpins), which represent more basal lineages within Cottoidei characterized by armored bodies and shallower distributions. Mitogenomic analyses have firmly established this sister-group relationship, highlighting shared synapomorphies like reduced swim bladders and specialized pectoral fins adapted for benthic lifestyles. The fossil record of Liparidae is sparse, with the earliest confirmed evidence consisting of a single assigned to Liparis (?) minusculus from late deposits near Antwerpen, , dating to approximately 25 million years ago. estimates indicate that the family likely originated in the North Pacific during the late Eocene to early (around 40–20 million years ago), coinciding with tectonic changes that deepened ocean basins and facilitated initial forays into deeper waters. Evidence for early deep-sea colonization emerges from phylogenetic reconstructions showing Liparidae as one of the primary invasive families into abyssal and hadal depths, contributing disproportionately to diversity below 6,000 meters since the . Evolutionary transitions in Liparidae trace from shallow-water ancestors in coastal and shelf habitats to dominance in deep-sea environments, driven by adaptations to increasing hydrostatic and low temperatures. Key morphological shifts include the complete loss of scales, resulting in smooth, flexible , and the development of gelatinous, low-density bodies composed of mucoid tissues that provide and without rigid structures. Deeper lineages exhibit further reductions in skeletal mineralization and , such as simplified crania and reduced fin rays, hypothesized to minimize weight while maintaining functionality in high- regimes. These changes represent across multiple clades, with hadal species diverging from neritic relatives around 20 million years ago. Post-2020 molecular phylogenies, incorporating and RADseq data, have clarified intra-family relationships, resolving Liparidae as with major subclades corresponding to depth zones. Recent analyses confirm the monophyly of hadal clades, particularly within genera like Careproctus and Pseudoliparis, revealing multiple independent colonizations of trenches and suggesting across basins. A 2025 study using three mitochondrial markers demonstrated that hadal snailfishes extend phylogenetically across Pacific and Atlantic trenches, supporting taxonomic revisions and underscoring rapid diversification in extreme depths over the past 10–15 million years.

Diversity and Genera

The family Liparidae, commonly known as snailfishes, encompasses 464 valid distributed across 31 genera (as of November 2025). This reflects their to a wide range of marine environments, from shallow coastal waters to the , though taxonomic challenges persist due to the presence of cryptic species that are morphologically indistinguishable but genetically distinct. Morphological convergence among deep-sea forms further complicates identification, often requiring integrated approaches combining , imaging, and traditional to delineate boundaries. Among the genera, Careproctus stands out as the most speciose, with around 150 inhabiting depths from shallow waters to . These snailfishes exhibit broad ecological tolerance, contributing significantly to the family's overall diversity. In contrast, Pseudoliparis comprises a smaller number of specialized for hadal environments, such as the trenches exceeding 6,000 meters, where they dominate the fish fauna. The genus Liparis, with approximately 98 , is predominantly found in shallow, temperate, and cold coastal waters of the , often among or beds. Recent explorations have continued to expand known diversity, with three new abyssal species described in 2025 from the eastern at depths of 4,000–6,000 meters near Station M. These include Careproctus colliculi, the bumpy snailfish, distinguished by its pink coloration and textured skin; Careproctus yanceyi, the dark snailfish, noted for its uniform dark hue; and Paraliparis em, a sleek form with subtle striping. Such discoveries underscore the ongoing revelation of hidden in remote deep-sea habitats, with 43 new liparid species described in the past decade alone.

Physical Description

External Morphology

Snailfishes exhibit an elongated, tadpole-like characterized by a disproportionately large head, a soft and gelatinous trunk, and a tapering posterior that lacks scales. This form is typical across the family Liparidae, with body sizes generally ranging from about 5 cm to over 30 cm in standard length, though some species like Polypera simushirae can reach up to 77 cm. The gelatinous texture arises from surrounding the body, providing flexibility but minimal structural rigidity. The fins of snailfishes are adapted for and attachment rather than rapid . Pectoral fins are notably large, often with 20–22 rays, where the uppermost rays are elongated and extend beyond the lower lobe, aiding in maneuvering over substrates. Pelvic fins are typically reduced or modified into a ventral disc formed by fused rays, which is prominent in shallow-water and many deep-sea species for adhering to surfaces; however, in some extreme deep-sea forms, these fins may be further reduced or absent. Skin in snailfishes is thin, loose, and scaleless, often translucent in deep-sea species such as , which lacks pigmentation to blend with the surrounding water column. Coloration varies by habitat and species: deep-sea forms tend toward pale or transparent hues for in low-light environments, while shallower-water species display mottled browns or darker patterns for benthic concealment. The loose, jelly-like may bear small prickles or spinules in some shallow-water taxa. A notable example is the 2025-described bumpy snailfish (Careproctus colliculi), which features pinkish skin with a distinctive bumpy . Sexual dimorphism in snailfishes is generally minor and poorly documented across the family, but some species exhibit subtle differences such as proportionally larger heads in males. For instance, in Liparis chefuensis, males display considerable dimorphism in and fin morphology compared to females. These variations are often linked to reproductive roles but do not drastically alter the overall external appearance.

Internal Anatomy

Snailfishes possess a skeletal system characterized by reduced , where many elements retain a cartilaginous structure rather than fully mineralizing into , an particularly pronounced in inhabiting greater depths. This reduction in facilitates in high-pressure environments by minimizing the overall mass of the . Micro-CT analyses across the Liparidae reveal an observed trend of declining bone mineralization with increasing habitat depth, with deeper-living taxa exhibiting visually lower skeletal densities compared to their shallower counterparts. In addition, specific cranial elements, such as the dentary, , and suborbital bones, become progressively shorter as depth increases, a pattern observed in comparative morphological studies of over 20 liparid . The digestive system of snailfishes is relatively straightforward, suited to their benthic lifestyle and of small and organic , featuring an inflated that aids in initial . In the hadal species , the stomach is particularly enlarged, potentially enhancing storage and digestion efficiency under extreme conditions. Sensory organs show variation with depth; shallower species often have larger eyes adapted for low-light environments, while hadal forms like P. swirei possess smaller eyes that, despite their reduced size, maintain functional metabolic activity indicative of preserved visual capabilities through proteomic adaptations. Unlike many shallow-water fishes, snailfishes lack a swim bladder, relying instead on extensive gelatinous subdermal tissues and watery muscle composition to achieve neutral buoyancy without gas-filled organs. These low-density gelatinous layers, composed primarily of extracellular matrix, permeate the body and reduce overall specific gravity, allowing the fish to hover effortlessly in the water column. In hadal species such as P. swirei, this buoyancy strategy is complemented by minimal skeletal density, with thin, incompletely ossified bones that resist compression while keeping body mass low. Such internal modifications highlight the family's evolutionary convergence on lightweight, pressure-tolerant anatomies across deep-sea habitats.

Distribution and Habitat

Global Occurrence

Snailfishes (family Liparidae) exhibit a , inhabiting all major ocean basins from the to the Antarctic. They are recorded in , Pacific, , and Southern Oceans, with species spanning from coastal shallows to extreme depths. The family demonstrates highest in the North Pacific, where numerous genera and approximately 60 in the genus Careproctus alone have been documented, reflecting the region's role as a center of origin and diversification. Notably, snailfishes are largely absent from tropical and subtropical shallow waters, with no adapted to warm, surface environments in those regions. Latitudinal gradients in snailfish distribution show a strong dominance in temperate and polar regions, where most species—predominantly in cold waters—thrive due to their physiological affinities for lower temperatures. This pattern underscores a reverse latitudinal diversity gradient compared to many shallow-water fish groups, with polar and subpolar zones supporting significant assemblages. Recent discoveries in 2025 off the coast, including three new abyssal species (Careproctus colliculi, C. yanceyi, and Paraliparis em) at depths exceeding 3,000 meters in the eastern Pacific, highlight emerging hotspots and ongoing revelations of biodiversity in these areas. Snailfishes primarily occupy benthic or bentho-pelagic zones, ranging from rare intertidal occurrences to the hadal depths of trenches. Their distribution often aligns with genus-specific ranges; for instance, species of the genus Liparis, such as L. fabricii and L. tunicatus, are commonly found in coastal waters, including the Chukchi and East Siberian Seas. These preferences enable snailfishes to exploit a broad vertical niche across global seascapes. Migration and dispersal in snailfishes are generally limited, constrained by their demersal lifestyles and lack of extensive pelagic phases in many , leading to high in isolated basins. In the , for example, several liparid exhibit regional endemism, with distributions confined to Antarctic shelves and trenches, facilitating localized adaptations and .

Environmental Preferences

Snailfishes (family Liparidae) are primarily cold-water specialists, thriving in environments with temperatures typically ranging from 0 to 10°C. Shallow-water species, such as the slimy snailfish (Liparis mucosus), prefer temperatures between 8.7 and 17.9°C, while deep-sea and hadal forms endure near-freezing conditions around 1–2°C. These fishes generally avoid warm tropical surface waters, occurring instead in polar, subpolar, and temperate regions, or at bathyal depths in tropical areas where temperatures remain suitably low. In terms of substrate and microhabitats, snailfishes closely associate with benthic environments, including soft sediments, rocky outcrops, and areas influenced by currents. Many species utilize a ventral sucking disc formed by modified pelvic fins to adhere to rocks or other hard substrates, facilitating stability in flow-prone microhabitats. Deeper-dwelling forms often rest directly on soft mud or silt bottoms, exploiting these substrates for camouflage and foraging proximity to the seafloor. Snailfishes exhibit adaptations to varying oxygen and levels, reflecting their diverse ecological niches. Hadal tolerate low-oxygen conditions prevalent in deep trenches, aided by genetic enhancements such as tandem duplications in the gene that boost resistance to . Most are strictly with full oceanic , but certain coastal , like the kelp snailfish (Liparis tunicatus), are and inhabit brackish waters. Occasionally, snailfishes form associations with chemosynthetic communities at hydrothermal vents and cold seeps, where they exploit the elevated nutrient availability in these dynamic, sulfide-rich microhabitats.

Depth Records

Snailfishes occupy an exceptionally wide vertical range in the ocean, from the shallow to the extreme hadal depths. The shallowest records include species like Liparis florae, the tidepool snailfish, which inhabits waters from 0 to 10 meters depth along the northeastern Pacific coast, often in rocky intertidal habitats exposed during low . At the deepest extremes, snailfishes represent the limit of vertebrate life, with verified observations pushing beyond 8,000 meters in multiple trenches. This range underscores their ecological versatility across pressure gradients, from near-surface to abyssal and hadal environments. The most notable hadal records highlight snailfishes as the deepest-living fish. In 2017, was documented at 7,966 meters in the using a baited trap deployed during a submersible expedition. This was eclipsed in 2023 by video footage of an unidentified Pseudoliparis species at 8,336 meters in the Izu-Ogasawara Trench, captured via a free-fall lander from the Falkor too. Similar depths have been recorded in the , where snailfishes, including Notoliparis kermadecensis, were observed up to 7,554 meters in 2014 during baited camera deployments. In the hadal zone (depths greater than 6,000 meters), snailfishes dominate the fish assemblage, comprising the dominant ichthyofauna in surveyed trenches and serving as key consumers in these sparse ecosystems. Recent surveys have expanded knowledge of their distributions without setting new hadal depth records since 2020. For instance, in 2025, three new abyssal snailfish species (Careproctus colliculi or Bumpy Snailfish at 3,268 meters, C. yanceyi or Dark Snailfish at 4,100 meters, and Paraliparis em or Sleek Snailfish at 4,100 meters) were described from the eastern Pacific, based on remotely operated vehicle (ROV) collections from using vehicles like Doc Ricketts. These findings, along with ROV observations using vehicles like SuBastian, have confirmed broader trench-spanning ranges for hadal species across the Pacific and Indian Oceans, emphasizing their prevalence in deep-sea hotspots.

Reproduction and Life History

Reproductive Biology

Snailfishes (family Liparidae) exhibit oviparous reproduction, with females laying demersal eggs that are typically attached to substrates such as rocks, , or the cavities of host organisms like king crabs in parasitic species. Clutch sizes generally range from 100 to 1,000 eggs in shallower-water , such as Paraliparis bullacephalus, where females may carry approximately 100 eggs per , with egg diameters of 2–4 mm. These eggs are demersal and , ensuring they remain in place on the for . In hadal species, such as , is markedly reduced, with clutches consisting of up to 23 large eggs (up to 9.4 mm in diameter), reflecting an adaptation for direct in extreme depths where larval dispersal is limited. This depth-related pattern shows a : fewer eggs but increased size to support advanced embryonic without free-living larvae, and no has been reported across the family. Mating behaviors involve , often occurring over suitable substrates where females deposit masses. In like Careproctus pellucidus, males establish territories and remain near the spawning site post-fertilization, providing indirect guarding by defending the area, though direct of eggs is absent. is typically reached at 2–5 years, varying with and environmental conditions. Breeding in high-latitude species is seasonal, aligned with peaks in polar productivity; for instance, Liparis fabricii spawns from summer to autumn (September–October), coinciding with enhanced blooms that support post-hatching nutrition.

Development and Lifespan

Snailfishes undergo direct , hatching from demersal eggs as large larvae capable of initial independent survival without external feeding. In species such as Careproctus pallidus, embryos develop within gelatinous egg masses attached to substrates, with hatching occurring after several weeks at low temperatures around 0–4°C; the provides essential energy reserves, depleting over several weeks post-hatching as the larvae transition to exogenous feeding. These yolk-sac stages are characterized by prominent fin folds and underdeveloped pectoral s, enabling limited mobility in shallow or coastal environments. The larval phase typically lasts 1–3 months in shallow-water species like those in the genus Liparis, during which larvae are often pelagic, dispersing in surface or mid-water layers before settling. marks a critical transition, involving resorption of the , development of fully formed pectoral and pelvic fins, and a shift to benthic or benthopelagic lifestyles; this process coincides with increased pigmentation and body compression suited to adult habitats. In deep-sea forms, larval development may occur closer to the , with reduced pelagic duration due to constrained dispersal. during these early stages is rapid relative to later life, but overall rates slow in deep-sea species post-metamorphosis, reflecting adaptation to resource-limited environments. Lifespan in snailfishes varies from 5 to 20 years, determined primarily through annuli analysis, with longer durations observed in deeper-dwelling species. For instance, hadal species such as from the and Notoliparis kermadecensis from the reach ages of 5–16 years, while shallower Careproctus melanurus can attain up to 25 years. Depth plays a key role in , as greater pressures and lower temperatures reduce metabolic demands, minimizing predation and extending life; growth zones confirm annual increments in these taxa. Senescence in snailfishes is minimal, particularly in deep-sea and hadal forms, owing to low metabolic rates that limit cellular damage and over time. These adaptations, including reduced basal metabolism at high pressures, contribute to prolonged post-reproductive survival without pronounced aging signs, such as tissue degradation, allowing individuals to persist in stable, low-energy deep environments for decades.

Feeding Ecology

Diet Composition

Snailfishes of the Liparidae primarily consume benthic , with amphipods forming the dominant prey across many , alongside polychaetes, mollusks, and other small crustaceans such as isopods and decapods. In hadal like from the , stomach content analyses reveal that amphipods constitute over 80% of the diet by volume, supplemented by occasional polychaetes and , reflecting a specialized predatory strategy on mobile benthic . Larger shallow-water , such as Liparis tanakae, exhibit opportunistic piscivory, incorporating small fish and natantian decapods into their diet, particularly in individuals exceeding 100 mm total length. Depth influences dietary composition markedly, with shallow and abyssal forms favoring more diverse assemblages, including copepods and gammarids, while hadal snailfishes like Notoliparis kermadecensis rely heavily on scavenging-associated prey such as detritus-laden amphipods. Larval snailfishes in nearshore environments, such as those in the , consume planktonic copepods and amphipods, transitioning to benthic diets in juveniles. As secondary to tertiary consumers, snailfishes occupy trophic levels typically ranging from 3.5 to 4.5, with compound-specific of indicating δ¹⁵N enrichment in deep-sea due to their position above primary like amphipods. This enrichment, averaging 4.15 ± 0.22 for snailfishes and 4.48 ± 0.13 for populations, underscores their role in transferring energy from invertebrate detritivores to higher predators.

Foraging Strategies

Snailfishes (family Liparidae) primarily employ benthic strategies, relying on their modified pelvic fins, which form an adhesive disc, to anchor onto substrates while using enlarged pectoral fins for precise maneuvering over soft sediments. This allows for slow cruising or hovering in close proximity to the seafloor. In shallower benthic habitats, this fin-based locomotion enables efficient navigation through complex terrains like rocky outcrops or muddy bottoms, minimizing energy use during foraging bouts. In the deep-sea and hadal zones, snailfishes adapt their tactics to the sparse food availability, often scavenging at organic falls such as bait deployments or natural carrion sinks. Hadal species like Pseudoliparis swirei from the Mariana Trench exhibit swift bottom-dwelling movements, foraging accurately and quickly across the seabed to capture mobile prey without high-speed chases. Their gelatinous, flabby bodies provide neutral buoyancy, a strategy suited to the nutrient-limited hadal environment. Sensory reliance in snailfish foraging shifts toward chemosensation in the perpetual of waters, where plays a minimal role. Genomic analyses reveal that hadal snailfishes, such as , have experienced massive losses in olfactory receptor genes but retain specialized trace amine-associated receptors, enabling detection of chemical cues from distant food sources despite the simplified system. This chemosensory adaptation allows interception of ephemeral food signals, such as those from scavenging events, without reliance on active visual hunting. These strategies align with the energy efficiency of snailfishes, characterized by low metabolic rates that support infrequent feeding in oligotrophic deep-sea habitats. Hadal in particular exhibit a "slow life" history, with reduced activity levels and metabolic demands that conserve resources during prolonged periods between meals. This efficiency minimizes interactions with competitors or predators at sites, enhancing in extreme depths.

Physiological and Molecular Adaptations

Pressure Tolerance Mechanisms

Hadal snailfish demonstrate remarkable piezophilic adaptations that allow them to endure hydrostatic pressures exceeding 800 atmospheres, equivalent to depths beyond 8,000 meters. These adaptations primarily involve biochemical and physiological strategies to counteract protein denaturation, membrane compression, and mechanical stress without relying on rigid skeletal reinforcements. Central to this is the role of trimethylamine oxide (TMAO), a compatible osmolyte that acts as a piezolyte to stabilize macromolecular structures under by counteracting compression and maintaining shells around proteins. In hadal species such as from the , TMAO concentrations in muscle tissue reach up to 386 mmol/kg, far surpassing the 40–50 mmol/kg typical in shallow-water teleosts, enabling enzymatic function at pressures that would otherwise disrupt cellular processes. To preserve membrane integrity, snailfish incorporate elevated levels of unsaturated fatty acids, particularly polyunsaturated ones like (DHA), into their bilayers. High pressure tends to rigidify membranes by reducing molecular motion and increasing packing , but these unsaturated chains introduce kinks that maintain fluidity and permeability, preventing transitions to states even at 800 atm or greater. This adjustment ensures efficient transport and signaling across cell membranes in the cold, pressurized hadal environment. Buoyancy regulation in snailfish further supports pressure tolerance by eliminating vulnerabilities associated with gas-filled structures. Unlike many shallow-water fishes, hadal snailfish lack swim bladders, avoiding from compression during descent or ascent. Instead, they rely on voluminous gelatinous tissues—primarily extracellular matrices rich in mucopolysaccharides—and low-density that provide with densities lower than surrounding (approximately 1.025–1.028 g/cm³). These soft, compressible materials allow the body to conform to without structural failure, facilitating energy-efficient hovering and maneuvering in the . Experimental studies underscore these mechanisms' efficacy. For instance, enzymes extracted from deep-sea snailfish retain near-full activity up to 1,000 in chambers, in contrast to shallow-water counterparts that lose function above 500 , demonstrating intrinsic resistance at the molecular level. Observations of live P. swirei captured from 8,000 m depths and briefly maintained under simulated conditions further confirm whole-organism to such extremes, with no immediate physiological collapse upon pressure equilibration.

Genomic and Biochemical Features

Snailfish genomes, particularly in hadal species like Pseudoliparis swirei from the Mariana Trench, are notably compact, with an assembly size of approximately 633 Mb and around 21,000 protein-coding genes. This reduced genome size facilitates efficient cellular function under extreme conditions, including high hydrostatic pressure. Comparative genomic analyses highlight significant gene losses related to bone formation, such as a frameshift mutation in the osteocalcin gene (bglap), which disrupts cartilage calcification and results in softer, gelatinous skeletons that better withstand deep-sea pressures without fracturing. Similar reductions in expression or functionality of genes like SPARC (secreted protein acidic and rich in cysteine), involved in extracellular matrix assembly and bone mineralization, further contribute to these skeletal adaptations in hadal lineages. Key molecular adaptations in snailfish genomes center on the upregulation and expansion of genes in the biosynthesis pathway, a critical for counteracting pressure-induced protein denaturation. For instance, the (fmo3) gene, which catalyzes TMAO production from dietary precursors, shows positive selection and —up to five copies in Yap Trench snailfish (Pseudoliparis cf. swirei)—enabling elevated TMAO levels (up to 261 mmol/kg in muscle tissue) that stabilize enzymes and membranes. Hypotheses of have been proposed for certain pressure-resistant enzymes, such as those enhancing (rad51 expansions observed in hadal genomes), though direct evidence remains limited and requires further validation through comparative phylogenomics. These genetic changes underscore TMAO's role as a primary biochemical , with proteomic studies confirming its dominance in maintaining under gigapascal pressures. Molecular phylogenetic analyses using (mtDNA) sequences, such as subunit I (), alongside nuclear loci like 2 (RAG2), have elucidated rapid radiations of hadal snailfishes across global trenches. These studies reveal multiple independent colonizations of the (depths >6,000 m) within the Liparidae family, with divergence times estimated at 2–5 million years ago based on mitochondrial genomes. A 2025 investigation into hadal snailfishes from the Kermadec and Peru-Chile Trenches confirmed cryptic divergence through COI barcoding, identifying distinct lineages despite morphological similarities and highlighting ongoing driven by trench . Such phylogenies emphasize the role of genomic plasticity in enabling these adaptations without requiring extensive morphological overhaul. Biochemical profiling of hadal snailfish reveals proteomic signatures of pressure tolerance, including enzymes with enhanced , such as those in and , which retain activity at pressures exceeding 1,000 atm due to TMAO-mediated chaperoning. Proteomic surveys of P. swirei muscle and liver tissues show upregulated heat-shock proteins and enzymes, but no novel hadal-specific metabolites beyond TMAO; instead, TMAO concentrations 2–3 times higher than in shallow-water relatives suffice to prevent . These traits align with genomic predictions, where expanded pathways minimize mutation accumulation from pressure-induced stress, ensuring long-term viability in extreme environments. A 2025 genomic study of deep-sea fishes, including multiple snailfish , further revealed convergent adaptations such as losses in vision-related genes (e.g., opsins) and skeletal mineralization pathways, alongside expansions in TMAO synthesis and response genes, confirming the evolutionary timeline of hadal colonization around 2–5 million years ago.

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