Merluccius
Merluccius is a genus of marine ray-finned fishes belonging to the family Merlucciidae, known as hakes, which includes 16 accepted species primarily inhabiting continental shelves and slopes in temperate and subtropical waters.[1] These species are characterized by an elongated, fusiform body, two dorsal fins, a large terminal mouth with a protruding lower jaw armed with strong teeth, and a light lateral line organ.[2] The genus exhibits a widespread but disjunct distribution across the world's oceans, excluding the Indo-West Pacific region, with species found in the eastern Atlantic from Europe to southern Africa, the western Atlantic from Canada to Argentina, the southeastern Pacific from Peru to Chile, the northeastern Pacific along the coast of North America, and isolated populations around southern Australia, New Zealand, and Tasmania.[3] Most species occupy depths between 50 and 500 meters, behaving as demersal or benthopelagic predators that remain near the bottom during the day but ascend into midwater at night to feed on small fishes, cephalopods, and crustaceans.[2] The accepted species include M. albidus, M. angustimanus, M. australis, M. bilinearis, M. capensis, M. gayi, M. hernandezi, M. hubbsi, M. merluccius, M. paradoxus, M. patagonicus, M. peruanus, M. polli, M. productus, M. senegalensis, and M. tasmanicus.[1] Hakes of the genus Merluccius are among the most commercially important demersal fish groups globally, supporting major fisheries through bottom trawling, gillnetting, and longlining, with annual catches historically exceeding 1 million tonnes, particularly from species like the European hake (M. merluccius), Argentine hake (M. hubbsi), and Pacific hake (M. productus).[2] They are marketed fresh, frozen, or processed into fillets and fishmeal, contributing significantly to regional economies in countries bordering the Atlantic and Pacific Oceans, though overexploitation has led to stock declines in several populations.[2]Taxonomy
Classification
The genus Merluccius belongs to the kingdom Animalia, phylum Chordata, class Actinopterygii, order Gadiformes, and family Merlucciidae.[4] It was established by Constantine Samuel Rafinesque in 1810 as part of his early systematic revisions of fish taxa.[4] The type species designated for Merluccius is Merluccius smiridus Rafinesque, 1810, which is now considered a junior synonym of Merluccius merluccius (Linnaeus, 1758), the European hake.[4] Phylogenetically, Merluccius forms a monophyletic group within the family Merlucciidae, a clade of gadiform fishes characterized by shared osteological and genetic traits.[5] Molecular studies, including analyses of mitochondrial control regions and nuclear markers, confirm its close relationship to other hake genera in Merlucciidae, such as Macruronus, with the family itself positioned as sister to Gadidae within Gadiformes.[5][3] Historical taxonomic revisions of Merluccius have relied on both morphological and molecular data to delineate species boundaries, evolving from early descriptions to modern phylogenomic approaches.[6] Inada's 1981 osteological studies recognized 12 species, but subsequent morphological and genetic analyses have increased this to 16 valid species as currently accepted, addressing ambiguities in rare morphotypes and confirming monophyletic superclusters (e.g., Old World vs. New World lineages).[3][1] Notable revisions include molecular confirmation of distinctions between sympatric species like Merluccius capensis and M. paradoxus in the Benguela Current, supported by allozyme and mtDNA data that highlight their depth-related separation and reproductive isolation.[6]Etymology
The genus name Merluccius derives from the New Latin compound formed by mare (Latin for "sea") and lucius (Latin for "pike"), collectively meaning "sea pike," a reference to the elongated, pike-like body of these marine fishes.[7] Alternative interpretations trace it to Medieval Latin merlutius or merlucius, possibly linked to merula (Latin for a type of sea fish or blackbird), reflecting the species' brownish-gray coloration, though the "sea pike" etymology is more widely accepted in ichthyological literature.[8] The genus Merluccius was first established in 1810 by Constantine Samuel Rafinesque in his work Caratteri di alcuni nuovi generi e nuove specie di animali e piante della Sicilia, where he elevated the group from its prior placement within the genus Gadus.[7] Rafinesque designated Merluccius smiridus—a Sicilian local name "smiriddu" for the fish—as the type species, though this is now considered a synonym of Merluccius merluccius (Linnaeus, 1758), making the genus name tautonymous with Linnaeus's earlier description of Gadus merluccius.[9] This initial taxonomic separation addressed confusions in earlier classifications, where hakes were lumped with cods in Gadus due to superficial similarities in body form and habitat.[10] Vernacular names for Merluccius species vary regionally and often reflect fishing traditions or morphology; the common English term "hake" originates from Old Norse haki (meaning "hook"), likely alluding to the fish's prominent lower jaw or the hooks used in capture.[11] In Europe, M. merluccius is known as "European hake" or "merluza" in Spanish, while Pacific species like M. productus are called "Pacific whiting" in North American fisheries, distinguishing them from smaller whiting species in the genus Merlangius.[12] Historical synonyms beyond the type include placements in genera like Epicopus (now obsolete), underscoring ongoing refinements in merluccid taxonomy.[13]Description
Morphology
Merluccius species exhibit an elongated, fusiform body that is compressed laterally, particularly toward the caudal region, with a well-differentiated caudal peduncle facilitating streamlined movement.[2] The skin is covered with small, cycloid scales that are often deciduous, giving the appearance of a relatively smooth texture.[14] They possess a large terminal mouth that is oblique, with the lower jaw protruding slightly, equipped with rows of large, hinged conical teeth adapted for capturing prey.[15] The head is large and flattened, featuring a V-shaped crest on top.[16] Two dorsal fins are present: the first is short-based and spinous, while the second is long and rayed.[2] The pelvic fins are thoracic in position, inserted subjugularly with 7-10 rays, and the anal fin is long and low, often similar in length to the second dorsal fin.[2] The pectoral fins are long and pointed, with their first ray positioned at or below the eye center.[14] A lateral line is present, typically straight with 101 to 171 scales, and the caudal fin is separate.[17][10] These fin configurations, combined with a physoclistous swim bladder that provides buoyancy control without connection to the gut, support their demersal and benthopelagic lifestyle, enabling efficient hovering and maneuvering over the seafloor.[2] Sensory adaptations include large eyes suited for vision in low-light conditions typical of deep-water environments.[16] The olfactory system is well-developed, aiding in prey detection within turbid or low-visibility waters, as is common in gadiform fishes.[15] Coloration typically features silvery sides that fade into a darker, brownish or greenish back, providing countershading for camouflage in open water.[18] Juveniles often display more pronounced patterns, such as stripes, enhancing their ability to blend with benthic structures and avoid predators.[19] Across species, body size varies, influencing proportional features like fin length relative to overall morphology.[2]Size and growth
Species in the genus Merluccius exhibit considerable variation in adult size, with typical total lengths (TL) ranging from 30 to 100 cm across most species.[20] For example, the North Pacific hake (M. productus) reaches a maximum of 91 cm TL, while the Cortez hake (M. hernandezi) attains smaller sizes, commonly up to around 30-40 cm TL.[21][22] Growth in Merluccius species is characterized by rapid initial rates in the first 2-3 years, often 20-30 cm per year, followed by a marked slowdown in later stages.[23] This pattern is well-described by the von Bertalanffy growth model, with parameters for the European hake (M. merluccius) including an asymptotic length (L∞) of approximately 85 cm and a growth coefficient (K) of about 0.15 year-1.[24] Sexual dimorphism is evident, as females generally grow faster and attain larger sizes than males in most species.[25] Development proceeds through planktonic larval stages beginning with a yolk sac, during which larvae are buoyant and passively dispersed.[26] Metamorphosis occurs at around 2-3 cm TL, marked by the development of fins and a shift to more active juvenile forms, typically after 25-30 days post-hatching.[26] Sexual maturity is reached at 25-40 cm TL, varying by species and sex, with females often maturing at larger sizes.[27] Females also tend to have longer lifespans than males, contributing to the observed size differences in adulthood.[25]Distribution and habitat
Global range
The genus Merluccius exhibits a circumglobal distribution in temperate to subtropical waters of the Atlantic and eastern Pacific Oceans, as well as southern regions of the western Pacific including New Zealand and Tasmania, with species such as M. australis and M. tasmanicus occurring off southern Australia, New Zealand, and Tasmania,[1] but is absent from the Indo-Pacific region encompassing the Indian Ocean and central-western Pacific.[28] Species typically inhabit continental shelves and slopes at depths ranging from 50 to 1,000 m, with some extending beyond 1,000 m in certain areas.[29] This distribution spans both hemispheres, reflecting an anti-tropical pattern where northern and southern populations are separated by tropical zones.[6] In the Atlantic Ocean, Merluccius species occupy both eastern and western basins. Eastern Atlantic representatives, such as M. merluccius, range from Norway southward to Mauritania, including the Mediterranean Sea and extending into the Gulf of Guinea.[30] Western Atlantic species include M. bilinearis, distributed along the North American coast from Newfoundland and the Gulf of St. Lawrence to the Mid-Atlantic Bight near Cape Hatteras.[31] Southern African populations, exemplified by M. capensis, occur off the southeastern Atlantic coast from Angola (near Baie Farte at 12°30'S) around the Cape of Good Hope to Natal, South Africa.[32] Pacific Ocean distributions are confined to the eastern and southern sectors. In the eastern Pacific, M. productus inhabits waters from the Gulf of Alaska to Baja California, Mexico.[33] Southern Pacific species include M. peruanus off Peru and M. gayi off the coasts of Chile from Arica (18°S) to Chiloé Island (42°S).[34][35] Historical expansions of Merluccius ranges have been influenced by post-glacial climate shifts following the Pleistocene, with genetic analyses revealing isolation between northern and southern populations due to barriers during glacial maxima that forced northward recolonization.[6] Allozyme data support this, showing greater genetic divergence across tropical divides than within hemispheres, indicative of vicariant events during ice ages.[6]Environmental preferences
Merluccius species are primarily demersal fishes, occupying depths of 70–350 m during the day and often undertaking diel vertical migrations to mid-water layers at night; they show a preference for muddy or sandy substrates on continental slopes.[15][2] These hakes favor water temperatures between 8 and 18°C, with salinity levels of 35–36 ppt, though subtropical species such as Merluccius gayi can tolerate warmer conditions up to 22°C.[36][37][38] Merluccius avoid low-oxygen zones where dissolved oxygen falls below 2 ml/L, as seen in distributions influenced by oxygen minima in upwelling systems.[39] They are commonly associated with upwelling areas, such as the Benguela and California currents, where nutrient-rich waters support high productivity essential for their food resources.[40][2] Seasonal migrations in Merluccius involve both vertical and horizontal movements tied to temperature fronts, with temperate species like Merluccius merluccius typically shifting northward and inshore during summer.[10][41]Biology
Reproduction
Species of the genus Merluccius are oviparous, gonochoristic fishes that reproduce through external fertilization, producing pelagic eggs and larvae that drift in ocean currents.[42] They exhibit a batch-spawning strategy with indeterminate fecundity, releasing multiple batches of eggs over extended spawning seasons that vary by species and region; for example, M. merluccius typically spawns from winter to spring in the Atlantic but shows year-round activity with peaks in spring-summer in the Mediterranean.[42][43] This protracted spawning enhances larval dispersal and survival by aligning with favorable environmental conditions.[44] Fecundity in Merluccius is high, with females producing 100,000 to over 1,000,000 eggs annually depending on body size and condition, as indeterminate fecundity allows ongoing oocyte recruitment during spawning.[45] Sexual maturity is generally reached at 2–5 years of age and 25–50 cm total length (TL), though these thresholds vary across species; males often mature earlier and at smaller sizes than females, such as at approximately 33 cm and 2.5 years for male M. merluccius.[27][46] Spawning typically occurs in offshore areas with strong currents that facilitate larval dispersal, such as the shelf break (50–100 m depth) for M. gayi off central Chile or the Bay of Biscay for M. merluccius.[43][47] Hermaphroditism is rare in the genus but has been documented in isolated cases, including synchronous functional hermaphrodites in M. merluccius from the Mediterranean and M. productus from the Pacific.[48][49] Eggs of Merluccius measure 1–1.5 mm in diameter and hatch in 5–7 days at typical sea temperatures (10–15°C), yielding larvae around 2.2–3.5 mm in length with yolk sacs for initial nourishment.[50][51] Early larval stages are highly vulnerable, experiencing elevated mortality from predation and passive advection away from suitable nursery habitats.[43]Diet
Merluccius species are carnivorous mid-level predators occupying trophic levels between 3.0 and 4.5, typically around 4.0–4.4, depending on age and region, positioning them as key energy transfer agents in marine food webs.[52][53] Their diet undergoes significant ontogenetic shifts: larvae are primarily planktivorous, consuming copepod eggs, nauplii, and calanoid copepods, which constitute up to 74% of their diet by volume.[54] Juveniles transition to small crustaceans like euphausiids and mysids, dominating their intake (e.g., 98% by weight in early juveniles of M. productus).[54][55] As adults, Merluccius become predominantly piscivorous, with fish comprising 70–94% of their diet by weight in larger individuals, alongside decapods, cephalopods, and euphausiids; for instance, M. merluccius preys heavily on schooling species such as Trachurus trachurus (47% index of relative importance) and Sardina pilchardus, while M. productus targets herring, sand lance, and eulachon.[54][52] Cannibalism occurs frequently in dense populations, accounting for up to 30% of the diet in some ecosystems, particularly when juvenile abundance overlaps with adults.[56][57] These hakes employ an opportunistic ambush feeding strategy, targeting patchily distributed schooling prey in mid-water or near the surface, with a pronounced nocturnal pattern evidenced by higher stomach fullness at night and daily vertical migrations to access pelagic resources.[58][59] Stomach content analyses reveal seasonal variations, such as increased crustacean consumption in summer and fish dominance in winter, reflecting prey availability fluctuations.[52][60]Ecology
Behavior
Merluccius species exhibit pronounced diel vertical migration patterns, remaining closer to the bottom during the day and ascending into pelagic waters at night to facilitate feeding activities. Archival tagging studies on the European hake (M. merluccius) have revealed frequent vertical excursions across the thermocline, with individuals experiencing rapid temperature fluctuations exceeding 7°C during these movements, typically occupying depths of 100–350 m overall.[61] Juveniles follow a similar rhythm, with those measuring 9–15 cm in total length shifting 10–20 m off the bottom at night while distributing between pelagic and benthic zones on the continental shelf, often forming schools in mid-water layers to enhance foraging efficiency.[62] Seasonal migrations in Merluccius involve latitudinal shifts driven by reproductive cycles and environmental cues, such as temperature gradients. For instance, in M. merluccius, adults tend to concentrate in southern regions during winter for spawning and disperse northward in summer, with juveniles recruiting to northern nursery areas as they grow; this pattern aligns with ontogenetic changes, where larger individuals prefer deeper, cooler waters (9.65–19.23°C).[63] Acoustic tracking and survey data on Pacific hake (M. productus) further demonstrate temperature-mediated responses, with distributions shifting poleward in warmer conditions and non-stationary effects influencing migration routes along the continental slope.[64] Socially, Merluccius species form loose aggregations rather than highly polarized tight schools, allowing flexible grouping for resource access while maintaining individual mobility. These aggregations facilitate coordinated predator avoidance through acoustic signaling, where individuals detect and respond to conspecific sounds or vibrations to evade threats in open water.[65] In low-visibility depths typical of their habitat (often below 100 m), Merluccius rely on the lateral line system to detect hydrodynamic vibrations and water movements, enabling navigation, prey localization, and predator evasion where visual cues are limited. This mechanosensory adaptation is crucial for benthic-pelagic transitions, supporting behaviors like diel migrations in stratified environments.[66]Predation and symbiosis
Merluccius species are subject to predation by a range of larger marine predators, reflecting their position as mid-trophic level consumers in coastal and shelf ecosystems. For European hake (M. merluccius), documented predators include larger fishes such as Atlantic cod (Gadus morhua), swordfish (Xiphias gladius), greater amberjack (Seriola dumerili), and spiny dogfish (Squalus acanthias), as well as marine mammals like bottlenose dolphins (Tursiops truncatus) and harbor porpoises (Phocoena phocoena).[67] Similarly, Pacific hake (M. productus) are preyed upon by California sea lions, small cetaceans, and spiny dogfish, with predation intensity varying seasonally and linked to hake migrations.[68][69] Juveniles across the genus show particularly high vulnerability to predation, including significant cannibalism by older conspecifics, which drives elevated early-life mortality and shapes population recruitment dynamics.[57] In their ecosystems, Merluccius serve as essential prey, channeling energy to apex predators and maintaining trophic balance. Pacific hake biomass, for instance, supports key predators like marine mammals, seabirds, sharks, and groundfishes in the northeastern Pacific, contributing to the stability of fisheries for species such as Pacific cod.[70] European hake fulfill a comparable role in the northeastern Atlantic and Mediterranean, where their abundance influences the foraging success of demersal predators and marine mammals.[52] Symbiotic interactions in Merluccius primarily involve parasitic relationships, with anisakid nematodes such as Anisakis simplex and A. pegreffii commonly infecting viscera and musculature across species. These parasites exhibit high prevalence—up to 80% in some populations—and can negatively impact host condition and energy allocation, particularly during reproduction, though effects on proximate composition and fatty acids are minimal.[71][72][73] Overfishing of Merluccius has triggered trophic cascades in upwelling-dominated systems like the Benguela Current, where depletion of hake stocks (e.g., Cape hake M. capensis) has reduced top-down control, leading to expansions in mesopredators, invertebrates, and algal blooms while altering overall food web structure.[74] In the Mediterranean and northeastern Atlantic, similar overexploitation of European hake has contributed to regime shifts by disrupting energy transfer and predator-prey balances in demersal communities.[75]Species
Recognized species
The genus Merluccius comprises 16 valid species according to current taxonomic assessments, such as WoRMS (Fricke et al. 2025).[1] These species are primarily demersal fishes found in temperate and subtropical waters of the Atlantic and Pacific Oceans, with varying maximum sizes and regional distributions. Key examples include the European hake (M. merluccius), which inhabits the Northeast Atlantic and Mediterranean up to 1.35 m in length, and the Pacific whiting (M. productus), occurring in the Northeast Pacific and reaching 105 cm.[17][69] The following table summarizes the recognized species, their common names, primary distributions, and maximum reported lengths:| Scientific Name | Common Name | Primary Distribution | Max Length |
|---|---|---|---|
| M. albidus | Offshore hake | NW Atlantic | 60 cm |
| M. angustimanus | Panama hake | Eastern Central Pacific | 40 cm |
| M. australis | Southern hake | Southern Pacific/Atlantic | 160 cm |
| M. bilinearis | Silver hake | NW Atlantic | 60 cm |
| M. capensis | Shallow-water Cape hake | SE Atlantic | 140 cm |
| M. gayi | Chilean hake | SE Pacific | 70 cm |
| M. hernandezi | Cortez hake | Eastern Central Pacific | 18 cm |
| M. hubbsi | Argentine hake | SW Atlantic | 100 cm |
| M. merluccius | European hake | NE Atlantic/Mediterranean | 135 cm |
| M. paradoxus | Deep-water Cape hake | SE Atlantic | 120 cm |
| M. patagonicus | Patagonian hake | SW Atlantic | 80 cm |
| M. peruanus | Peruvian hake | SE Pacific | 115 cm |
| M. polli | Benguela hake | SE Atlantic | 50 cm |
| M. productus | Pacific whiting | NE Pacific | 105 cm |
| M. senegalensis | Senegalese hake | E Atlantic | 90 cm |
| M. tasmanicus | Tasmanian hake | Southern Australia/Tasmania | 37 cm SL |