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European flying squid

The European flying squid (Todarodes sagittatus) is a medium- to large-sized cephalopod mollusk in the family Ommastrephidae, distinguished by its robust, cylindrical or conical mantle that can reach up to 75 cm in length and 2.2 kg in weight, along with wide, rhomboidal fins spanning about 45% of the mantle length, biserial suckers on its arms, and long tentacles equipped with 20–30 small proximal suckers and 6–12 larger distal ones.^[1]^[2] This species exhibits a short lifespan of 1–2 years, with rapid growth rates in juveniles, and it is renowned for its "flying" behavior, gliding above the water surface to evade predators using jet propulsion.^[1]^[2] Distributed primarily in the nerito-oceanic waters of the eastern North Atlantic from Iceland and the Barents Sea (61°N) southward to the Gulf of Guinea (13°S), including the Mediterranean Sea, North Sea, Norwegian Sea, and mid-Atlantic Ridge, T. sagittatus also appears sporadically in the western Indian Ocean and western Pacific, such as off southern Australia and the South China Sea.^[1]^[3]^[2] It occupies benthopelagic habitats on continental slopes and shelves, from epipelagic surface waters down to mesopelagic depths of over 1,000 m (with records to 4,595 m), preferring temperatures of 2–14°C (mean 9.3°C) and often associating with sandy or muddy bottoms at 200–800 m during spawning, while performing diel vertical migrations and schooling in open ocean areas.^[1]^[2] Ecologically, T. sagittatus is a voracious predator with a diet dominated by fishes (such as myctophids and other Osteichthyes, comprising up to 85% frequency of occurrence), crustaceans (e.g., decapods at 49%), and cephalopods (30%), including cannibalism among conspecifics, shifting from fish-heavy intake in juveniles to more crustacean-focused in adults.^[1]^[4] It reproduces year-round on continental slopes, with peaks in late winter to spring in northern Europe, as a gonochoristic species where females release 120,000 to over 950,000 eggs (2.4 mm diameter) in gelatinous spawning masses, maturing at 170–500 mm mantle length and with males dying post-spawning.^[1]^[2] As prey for tunas, swordfish, sharks, seals, and cetaceans, and a paratenic host for anisakid nematodes, it plays a key role in pelagic food webs and supports moderate fisheries as bycatch in trawl operations, often marketed alongside related species like Illex coindetii.^[1]

Taxonomy

Classification

The European flying squid, Todarodes sagittatus, is classified within the domain Eukaryota, kingdom Animalia, phylum Mollusca, class Cephalopoda, subclass Coleoidea, superorder Decapodiformes, order Oegopsida, family Ommastrephidae, subfamily Todarodinae, genus Todarodes, and species sagittatus.^[5] This placement reflects its position among the pelagic squids, characterized by ten arms and the ability to eject water for propulsion typical of cephalopods.^[1] Todarodes sagittatus serves as the type species for the genus Todarodes, which in turn is the type genus for the subfamily Todarodinae.^[5] The subfamily Todarodinae distinguishes itself from related subfamilies like Ommastrephinae (which includes the genus Ommastrephes) through key morphological traits, including the absence of photophores on the mantle and arms, a funnel groove featuring a foveola without side pockets, and a left tentacular club with two pairs of carpal suckers, no carpal-locking apparatus, and quadriserial dactylus suckers.^[1] In contrast, genera in Ommastrephinae, such as Ommastrephes, possess photophores for bioluminescence and exhibit different tentacular club configurations with protective membranes and longitudinal ridges.^[1] The classification of Todarodinae as a distinct subfamily originated in the mid-20th century, elevated from within Ommastrephinae based on morphological analyses that highlighted these unique traits.^[1] William Adam formally established Todarodinae in 1960, defining it with Todarodes as the type genus and emphasizing features like the funnel groove structure.^[1] Subsequent revisions, including Roeleveld's 1988 reassessment of genera like Todaropsis and Nigmatullin's 1992 phylogenetic proposals, reinforced the three-subfamily system for Ommastrephidae (Illicinae, Todarodinae, Ommastrephinae), supported by later molecular studies confirming monophyly.^[1]

Nomenclature

The European flying squid bears the scientific name Todarodes sagittatus (Lamarck, 1798), with the species originally described by Jean-Baptiste Lamarck in 1798 under the binomial Loligo sagittata in the genus Loligo, based on specimens from the Mediterranean Sea.^[3] This initial classification placed it among the loliginid squids, but subsequent taxonomic revisions recognized its distinct characteristics, leading to its transfer to the genus Ommastrephes established by Alcide d'Orbigny in 1835–1847, as Ommastrephes sagittatus.^[6] Further refinement occurred in 1880 when Japetus Steenstrup erected the genus Todarodes, designating T. sagittatus as the type species, reflecting its placement within the ommastrephid family due to features like the funnel groove and tentacular club structure.^[6] The genus name Todarodes derives from the Ancient Greek todaros (oblong) combined with the suffix -odes, meaning "resembling" or "like."^[6] The specific epithet sagittatus comes from the Latin sagitta, meaning "arrow," alluding to the species' elongated, arrow-shaped body form.^[3] Several junior synonyms have been proposed for Todarodes sagittatus over time, reflecting historical misclassifications and regional descriptions. Key synonyms include Loligo sagittata Lamarck, 1798 (the original combination); Ommastrephes sagittatus (Lamarck, 1798) d'Orbigny, 1835–1847; Loligo aequipoda Vérany, 1851; Loligo todarus Rafinesque, 1814 (nomen nudum); and Ommatostrephes sagittatus Lamarck, 1798, among others such as Sepia media Barbut, 1783 and Loligo brasiliensis Férussac, 1823.^[3]^[6] The World Register of Marine Species (WoRMS) maintains a comprehensive list of these synonyms, confirming Todarodes sagittatus as the valid name.^[3] Common names for the species vary by region and language, often emphasizing its arrow-like form or flying behavior. In English, it is known as the European flying squid, arrow squid, or red squid; in French as toutenon commun or calmar; in German as Pfeilkalmare or großer Pfeilkalmar; and in Spanish as pota europea or simply pota in fisheries contexts.^[3]^[6]

Description

Morphology

The European flying squid, Todarodes sagittatus, possesses a streamlined body adapted for agile swimming in pelagic environments. Its mantle is elongated, cylindrical, and robustly muscular, tapering posteriorly to a pointed end, which facilitates rapid propulsion via jet ejection from the funnel. The fins are broad and rhomboidal, spanning approximately half the mantle length and forming a distinctive shape that aids in stability and maneuverability during flight-like leaps above the water surface.^[1] The species features ten circumoral appendages: eight arms arranged in two rows around the mouth, each equipped with biserial suckers for manipulation and grasping, and two longer tentacles that extend well beyond the arms. The tentacles terminate in expanded, club-shaped ends bearing four longitudinal rows of suckers, bordered by protective membranes and keels that enhance prey capture efficiency; in males, the right ventral arm (arm IV) is hectocotylized in its distal third, modified with fleshy papillae and altered suckers to facilitate spermatophore transfer during mating.^[1] The head is broad and robust, bearing large, prominent eyes suited to low-light oceanic depths, with a strong chitinous beak positioned centrally for tearing prey.^[1] Internally, the gladius—a thin, arrow-shaped chitinous structure—provides skeletal support along the dorsal midline of the mantle, featuring a narrow rachis without prominent vanes. Females possess paired nidamental glands within the mantle cavity, which secrete jelly-like coatings to encase eggs during reproduction.^[1]

Size and coloration

The European flying squid, Todarodes sagittatus, exhibits a size range typical of medium-sized ommastrephid squids, with maximum recorded mantle lengths reaching 750 mm for unsexed specimens, though commonly observed lengths fall between 250 and 350 mm dorsal mantle length (DML).^[7]^[2] Females attain larger sizes than males, with maximum DML up to 750 mm compared to 500 mm for males, while total length, including tentacles, can extend to approximately 1 m.^[7]^[8] Weights typically reach up to 2.2 kg, reflecting the species' robust build.^[2] Growth in T. sagittatus is rapid, enabling individuals to reach sexual maturity at 200–300 mm DML within about one year of hatching.^[8] This accelerated development supports a short lifespan of slightly over one year, with daily weight increments averaging 2.22 g for males and 3.66 g for females during subadult and adult phases.^[8] In terms of coloration, the mantle is dark purplish-brown dorsally with silvery sides and a ventral sheen, featuring reddish chromatophores that facilitate camouflage through rapid color adjustments.^[9] The fins are translucent with darker edges, and the overall pigmentation can pale during stress or mating activities.^[9] Sexual dimorphism is pronounced in T. sagittatus, with females not only larger overall but also possessing proportionally larger fins than males.^[7]^[8] Males are further distinguished by the enlarged hectocotylus on one arm, used in reproduction.^[7]

Distribution and habitat

Geographic range

The European flying squid (Todarodes sagittatus) primarily inhabits the eastern Atlantic Ocean, ranging from the waters off Greenland and Iceland southward to the Gulf of Guinea at approximately 13°S and 40°W longitude.^[1] This distribution encompasses the continental slope and oceanic waters, including the North Sea, Norwegian Sea, and Celtic Sea, as well as the mid-Atlantic Ridge. The species extends from about 61°N near the Shetland Islands southward to around 36°S off the Cape of Good Hope, South Africa, with records along the west coasts of the British Isles, France, Spain, Portugal, and northwest Africa.^[1] It also occurs sporadically in the western Indian Ocean (e.g., Saya de Malha and Nazareth Banks) and western Pacific (e.g., South China Sea and waters off southern Australia, including the Timor Sea and Tasmania).^[1] In the Mediterranean Sea and adjacent regions, T. sagittatus is widespread, occurring throughout the western, central, and eastern basins, including the Tyrrhenian, Ionian, Adriatic, and Aegean Seas, as well as the Strait of Sicily and Levant Basin. It is present in the Sea of Marmara, though not recorded in recent surveys there, and appears sporadically in the Black Sea via the Bosphorus Strait.^[10] Recent northward range expansions have brought T. sagittatus into the Barents Sea, with foraging shoals recorded there in 2010 for the first time in 25 years, attributed to Arctic warming and Atlantification of marine communities.^[11] Studies from 2020–2024, analyzing trawl data from 2005–2022, show increased abundance in the western Barents Sea south of 75°N since its re-emergence around 2010, following a historical absence after 1983.^[12] Originally a subtropical-boreal species, T. sagittatus has exhibited poleward migration linked to climate change, with seasonal immigrants becoming regular residents in the northern North Sea.^[13] A 2022 biodiversity study on the North Sea shelf documented its shift from rare, sporadic occurrences in the early 20th century to consistent winter presence of juveniles and maturing individuals in central and northern areas by 2016–2020.^[13]

Preferred habitats

The European flying squid (Todarodes sagittatus) occupies oceanic and neritic zones across a broad depth range, from epipelagic waters near the surface (0 m) to bathypelagic depths exceeding 1000 m. Adults predominantly inhabit mid-depths of 200–800 m along continental slopes, where they are commonly captured by commercial trawlers. Juveniles, in contrast, are more frequently encountered in shallower waters less than 100 m deep, often near the surface in coastal and bank areas.^[1]^[2] This species thrives in water temperatures ranging from 9–18°C and salinities typically between 35–36 psu in its marine environments.^[1] It shows a strong association with upwelling areas, particularly along western Iberian coasts, where nutrient-rich waters enhance prey availability and support higher abundances. Habitat preferences include areas over sandy or muddy substrates on continental slopes and oceanic seamounts, facilitating access to benthic and pelagic food resources. The squid generally avoids shallow coastal waters, except during seasonal incursions linked to migrations.^[1] Habitat use varies by life stage: paralarvae are confined to the upper 100 m of the water column, benefiting from warmer surface temperatures (15–22°C) for early development. Subadults transition to shelf edges at depths of 80–200 m, as observed in Norwegian waters where they aggregate in coastal banks during certain seasons.^[1]^[2]

Biology and ecology

Life cycle and reproduction

The European flying squid (Todarodes sagittatus) exhibits a short life span of 1–2 years, typically approximately one year in most populations, during which individuals grow rapidly before reaching maturity and reproducing once in a semelparous manner, after which adults typically die.^[14]^[15] This annual cycle is supported by statolith increment analysis, which indicates hatching throughout the year with a peak in late autumn, followed by accelerated growth that slows as squid approach senescence.^[16]^[14] The life cycle progresses through distinct developmental stages beginning with eggs that hatch into planktonic paralarvae measuring 1–2 mm in dorsal mantle length.^[10] These paralarvae transition to juveniles of 10–50 mm mantle length within 1–2 months, characterized by high daily growth rates of up to 1.8 mm.^[16] Sexual maturity is attained at 6–9 months of age, when individuals reach 200–300 mm mantle length, with females maturing slightly later than males.^[14] There is no parental care, and early life stages remain poorly understood due to challenges in sampling planktonic forms, though recent observations have documented spawning activity in the North Sea as of 2021.^[17] Reproduction involves internal fertilization, with males using a specialized arm called the hectocotylus to transfer spermatophores into the female's mantle cavity.^[2] Females produce 100,000–500,000 eggs per spawning event, releasing them in gelatinous strands or masses at depths of 200–800 m, often near the seafloor.^[16] Spawning occurs year-round in the Mediterranean Sea but is more seasonal in the North Atlantic, with peaks in spring and autumn, and females may release multiple batches intermittently.^[14]^[15] Maturity is fully reached by one year of age, aligning with the species' overall semelparous strategy.^[14]

Feeding and diet

The European flying squid, Todarodes sagittatus, is an opportunistic predator with a diet dominated by fish, which constitute 50–70% of its prey by numerical importance across various regions. In the western Mediterranean, fish such as lanternfish (Ceratoscopelus maderensis) and ribbonfish (Lepidopus caudatus) account for approximately 71% of the diet, while in the North Atlantic and North Sea, herring (Clupea harengus), capelin (Mallotus villosus), and sand eels (Ammodytidae) are primary targets. Crustaceans, including euphausiids (Meganyctiphanes norvegica) and deep-sea shrimps (Aristeus antennatus, Pasiphaea spp.), form a significant secondary component at 20–35%, and cephalopods contribute 10–20%, often through cannibalism on conspecifics or other species like Loligo forbesii. Juveniles exhibit an ontogenetic shift in diet, initially consuming planktonic prey such as small crustaceans and detritus during their early paralarval stage, primarily feeding on fish as they grow, with adults shifting to include more crustaceans and cephalopods in some regions like the Mediterranean, where fish reliance decreases slightly but remains over 60%. This shift reflects increasing predatory capability and access to deeper prey. The squid employs a predatory feeding mechanism, capturing prey with long tentacles and processing it via a powerful chitinous beak, consuming a daily ration equivalent to 6–12% of its body weight to support its high metabolic demands. As a mid-level predator, T. sagittatus occupies a trophic level of approximately 4.0, positioning it as a key link between primary consumers and top predators in pelagic food webs. Recent stable isotope analyses of beaks from North Atlantic specimens indicate that contemporary populations (post-2016) exhibit a broader isotopic niche, suggesting a more generalist diet incorporating boreal fish compared to more specialized Mediterranean diets focused on mesopelagic species. As of 2024, these studies confirm ongoing expansion of the dietary niche due to climate change, with increased consumption of boreal fish species in northern populations.^[18] Seasonal variations occur, with higher fish intake during summer in North Sea schools, where up to 60% of stomachs contain fish versus a more balanced fish-cephalopod mix in winter; cannibalism is prevalent in dense populations, comprising up to 5% of cephalopod prey in the Mediterranean. Schooling behavior facilitates coordinated foraging on aggregated prey schools.

Behavior and migration

The European flying squid, Todarodes sagittatus, exhibits schooling behavior primarily during warmer months, forming loose aggregations of 10 to 100 individuals, often with a female-biased sex ratio, to facilitate collective feeding on pelagic prey such as fish and crustaceans.^[1] These schools are most prominent in summer off the coasts of Iceland and Norway, where the squid congregate in surface or near-surface waters for opportunistic hunting.^[19] In contrast, during winter, individuals tend to be more solitary or occur in smaller, dispersed groups as they shift to deeper offshore habitats.^[1] This seasonal variation in social structure aligns with the species' high metabolic demands, enabling efficient prey capture in productive zones while reducing competition in less resource-rich areas.^[20] Migration in T. sagittatus involves both diel vertical movements and broader seasonal patterns driven by feeding and maturation needs. Diurnally, the squid perform vertical migrations, descending to depths of 70–800 m during the day to avoid predators and access mesopelagic prey, then ascending to surface or epipelagic layers at night for active foraging.^[21] Seasonally, they undertake large-scale horizontal migrations, following prey distributions and temperature gradients in a roughly annual circuit; for instance, populations move northward into warming Atlantic waters during summer, with spawning groups shifting from oceanic to neritic zones along continental slopes.^[20] Recent analyses indicate these patterns have intensified with climate-driven shifts, such as increased northward incursions observed in North Atlantic surveys.^[7] Ontogenetic progression further influences these routes, with juveniles remaining in shallower, coastal areas before larger adults migrate to bathypelagic depths exceeding 500 m.^[8] For evasion and sensory adaptations, T. sagittatus relies on rapid jet propulsion, achieving burst speeds over 25 km/h to escape threats, often combined with ink release to create visual and chemical distractions for predators.^[1] Chromatophores in the skin enable quick color changes for camouflage against varying oceanic backgrounds, enhancing survival during migrations through diverse habitats.^[16] While not possessing true bioluminescent photophores, the species may use subtle reflective properties or environmental light cues for hunting and communication in low-light conditions.^[1] These behaviors are responsive to environmental stimuli, such as temperature and salinity gradients, which guide vertical positioning and overall migratory fidelity.^[20]

Human interactions

Fisheries

The European flying squid (Todarodes sagittatus) is primarily harvested as bycatch in pelagic and demersal trawl fisheries targeting other species, such as finfish, across the Northeast Atlantic and Mediterranean Sea.^[22] Targeted fisheries employ jigging techniques, where lures mimic prey to attract squid, particularly in the Mediterranean and parts of the North Atlantic.^[23] Purse seining is occasionally used for schools in surface waters, though less commonly than trawling or jigging.^[24] Commercial catches of T. sagittatus have historically averaged around 3,000 tonnes annually in the Mediterranean prior to 2020, with global totals remaining relatively low compared to other squid species due to its incidental capture.^[25] Since the 2000s, landings have shown an upward trend in several regions, linked to increased abundance from warmer water inflows, with peaks exceeding 2,600 tonnes in Celtic Seas areas during 2017.^[26] Recent data from 2020–2024 indicate shifts due to range expansion, including higher bycatch in the North Sea (e.g., up to 13 tonnes in Division 4c in 2021, with catch per unit effort rising until 2019) and sporadic increases in the Barents Sea (from near-zero to occasional tonnes, tied to Atlantification).^[26]^[12] Major producing countries include Spain (dominant in Divisions 8abd and 9a, with 243 tonnes in 2021), Portugal (key in Division 9a), France (leading in Celtic Seas and Bay of Biscay, ~220 tonnes in 2021), and Morocco (in northwest African waters).^[26]^[27] The species is marketed fresh, frozen, salted, or canned, often under the name "calamar" in southern European markets, and exported for human consumption or as bait in other fisheries.^[28] There are no dedicated total allowable catches or quotas for T. sagittatus, as cephalopod fisheries are excluded from such regulations under the EU Common Fisheries Policy, though general effort controls and minimum landing sizes apply in member states.^[29]^[30] Reports from 2023–2025 highlight elevated bycatch in the Barents Sea, prompting monitoring under ICES frameworks amid ongoing range shifts.^[12]

Conservation status

The European flying squid (Todarodes sagittatus) is classified as Least Concern on the IUCN Red List, with the assessment conducted in 2010 and reaffirmed in the 2025-1 version, owing to its extensive geographic range across the eastern Atlantic and Mediterranean, high fecundity with females producing a few thousand to over 950,000 eggs, and overall population resilience despite localized pressures.^[1]^[2] Key threats to T. sagittatus include bycatch in non-selective trawl and jigging fisheries targeting other species such as redfish and herring, which can result in substantial incidental mortality, particularly for juveniles.^[7] Climate change is driving range shifts, with warming waters facilitating northward expansion into the North Sea since the 1980s, as evidenced by increased biomass and occurrence rates correlated with rising sea surface temperatures (r² = 0.45–0.50).^[31] Additionally, ocean acidification poses risks to early life stages, impairing statolith formation and development in paralarvae, potentially reducing survival rates across cephalopod species including ommastrephids like T. sagittatus.^[32] Natural predators of T. sagittatus encompass seabirds such as fulmars, marine mammals including seals and cetaceans (e.g., pilot whales), and larger fish like tuna, swordfish, and sharks, contributing to incidental mortality through predation on all life stages.^[33] There are no species-specific conservation measures for T. sagittatus, but populations are monitored through the International Council for the Exploration of the Sea (ICES) Working Group on Cephalopod Fisheries and Life History (WGCEPH) and Food and Agriculture Organization (FAO) reporting frameworks, which track landings and biomass indices across FAO Area 27.^[34] Recent ICES and FAO studies emphasize bycatch reduction strategies, such as gear modifications and temporal closures in high-interaction zones, to mitigate fishery impacts without targeted quotas.^[26] Population trends for T. sagittatus appear stable to increasing in northern ranges, including the North Sea and Icelandic waters, attributed to warming-induced habitat suitability and recruitment pulses, though high interannual variability persists. In contrast, southern core areas like the western Mediterranean may face potential declines due to overexploitation and environmental stressors, though data remain limited and no overall global reduction is evident.^[31]

References

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