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Spurdog

The spurdog, scientifically known as Squalus acanthias and commonly referred to as the , is a small to medium-sized belonging to the family , characterized by its slender, elongate body, pointed snout, large eyes, and two dorsal fins each preceded by a sharp, venomous spine. Typically reaching lengths of 70–100 cm (up to 1.6 m in females), it features a grey-brown upper body with white spots that fade with age, a pale underbelly, and no anal fin, making it one of the most abundant and widespread in temperate and waters worldwide. Found primarily in coastal and offshore waters over the continental shelves of the North Atlantic and temperate waters of the southern hemisphere, including parts of the Indian and Pacific Oceans (note: North Pacific populations off North America are a distinct species, Squalus suckleyi), the spurdog is a migratory, schooling predator that inhabits depths from shallow inshore areas to 900 m, preferring cooler temperatures below 17°C and full-strength seawater. It feeds opportunistically on a diverse diet including small bony fishes like herring and capelin, cephalopods such as squid, and crustaceans, often hunting in packs and using its spines for defense against predators by curling its body and striking. Reproduction is notably slow and complex, with ovoviviparous females carrying litters of 1–15 pups (average 6–7) for an extended gestation period of 18–24 months—the longest among vertebrates—reaching sexual maturity at 6–35 years (females later than males) and potentially living 25–100 years, which contributes to its vulnerability to overexploitation. Despite its historical abundance and commercial value for meat, fins, and oil (often used in ), the spurdog faces significant threats from targeted fisheries, , and slow recovery rates, leading to global population declines of over 50% in many regions since the mid-20th century. Classified as Vulnerable by the (assessed 2019), it is subject to various regional quotas, such as those in the and North American fisheries, with signs of partial recovery in some areas due to management efforts since the early 2000s. While generally not aggressive toward humans, its spines can inflict painful wounds, though it poses minimal direct threat.

Taxonomy

Etymology and nomenclature

The common name "spurdog" is a compound word derived from "" and "," alluding to the sharp, prominent spines preceding the fins, which resemble spurs. This name is particularly prevalent in the northeastern Atlantic and parts of , where the species is also known as the , piked dogfish, mud , or grayfish. The term "" itself originates from observations by fishermen of the species hunting in packs, akin to dogs chasing prey. The scientific name Squalus acanthias was first established by in 1758 in his . The genus name Squalus derives from the Latin "squalus," an ancient term for sharks or certain marine fishes considered unclean or unfit for consumption. The species epithet acanthias comes from the Greek "akanthias" (ἀκανθίας), meaning thorny or spiny, directly referencing the characteristic dorsal spines. While "spurdog" most commonly denotes Squalus acanthias, the name is sometimes applied more broadly to other in the Squalus, which collectively share similar smooth dorsal spines and are referred to as spurdogs.

Classification and synonyms

The spurdog, scientifically known as Squalus acanthias, is classified within the domain Eukaryota, kingdom Animalia, phylum Chordata, class , subclass , order , family , genus Squalus, and S. acanthias (Linnaeus, 1758). This placement situates it among the cartilaginous fishes, specifically within the squaloid characterized by their spindle-shaped bodies and lack of an anal fin. The species belongs to the family , commonly known as dogfish sharks, which comprises two genera and 42 species distributed globally in temperate and tropical waters, as of 2025. Within the genus Squalus, S. acanthias is one of the most widespread and well-studied members, sharing traits such as paired spines and a caudal fin with a strong lower lobe with its congeners. Historically, S. acanthias has accumulated numerous synonyms due to regional variations in early descriptions and taxonomic revisions. Notable examples include Acanthias (Risso, 1827), proposed for Mediterranean populations based on minor morphological differences, and Squalus americanus (Storer, 1839), applied to North Atlantic specimens reflecting geographic naming conventions; both are now considered junior synonyms after comprehensive reviews confirmed their conspecificity with S. acanthias. Other synonyms, such as Acanthias americanus (Storer, 1846) and Spinax mediterraneus (Gistel, 1848), arose from fragmented early classifications but were unified through morphological and molecular analyses. Taxonomic debates persist regarding the separation of S. acanthias populations, particularly between Atlantic and Pacific lineages, driven by genetic evidence. While some studies indicate low divergence between Atlantic and southern Pacific groups, suggesting panmictic populations, higher genetic differentiation across the North Pacific has led to the recognition of the northeastern Pacific form as a distinct species, Squalus suckleyi (Girard, 1854), resurrected in 2010 based on and subtle vertebral counts; this split implies that previous broad applications of S. acanthias to Pacific waters may warrant reevaluation in transitional zones.

Description

Physical characteristics

The spurdog, or (Squalus acanthias), possesses a slim, elongated body that is cylindrical in shape, adapted for agile swimming in environments. This body features two fins, each preceded by a prominent venomous that serves as a defensive , capable of inflicting painful injuries through mild secreted from glands at the base. The caudal fin is asymmetrical, with a larger lobe and a smaller ventral lobe, while the pectoral and pelvic s provide stability and maneuverability. Notably, the spurdog lacks an anal fin, a characteristic trait distinguishing it from many other shark species. The skin of the spurdog is covered in placoid scales, known as dermal denticles, which give it a rough, sandpaper-like and contribute to hydrodynamic efficiency by reducing drag. These denticles are small, low-lying structures with three cusps, a central ridge, and lateral extensions. Coloration is typically slate gray or grayish-brown on the surface, fading to pale gray or white on the ventral side, often with rows of white spots along the flanks that may fade with age. The head is characterized by a narrow, pointed , moderately large eyes positioned midway between the snout tip and gill slits—lacking a —and a small terminal mouth equipped with sharp, curved teeth arranged in multiple rows; upper teeth number around 28 and are obliquely pointed, while lower teeth (22-24) form a cutting edge. Internally, the spurdog's liver is large and oil-filled, providing buoyancy to compensate for the absence of a , a common in cartilaginous fishes. The digestive system includes a intestine, which enhances nutrient absorption by increasing the internal surface area. Sexual dimorphism is evident, with males possessing paired claspers for and generally attaining a slightly smaller size than females at maturity.

Size, growth, and lifespan

The spurdog (Squalus acanthias) typically reaches an average of 80–100 , though maximum recorded lengths extend to 160 total , with weights up to 9.4 . This species exhibits , with females attaining larger sizes than males. Growth is notably slow, at approximately 2–4 per year, contributing to its delayed maturation and vulnerability to exploitation. Sexual maturity occurs at lengths of 60–100 cm for males, typically around 11 years of age, while females mature later at 80–124 cm and ages of 12–21 years. Lifespan varies by sex, with females living up to 35–40 years and males up to 20–30 years, reflecting their protracted life history. Age determination relies on counting annual growth bands in vertebrae, a method analogous to in trees, which provides reliable estimates of longevity and growth increments. Regional variations in size are evident, with Mediterranean populations generally smaller than those in the North Atlantic or North Pacific, where maturity sizes and maximum lengths tend to be larger.

Distribution and habitat

Geographic range

The spurdog (Squalus acanthias), also known as the , exhibits a primarily in temperate and waters across multiple ocean basins. In the North Atlantic, populations range from and the Murmansk Coast in the northeast to in the southeast, encompassing the western Atlantic from to , as well as the Mediterranean and Seas. In the southern hemisphere, the species occurs in temperate waters off southern , , , , southern , and parts of the and South Pacific Oceans. The spurdog is typically found from coastal areas to offshore continental shelves, inhabiting depths of 10–200 m, though it has been recorded up to 750 m or more in some regions. Migratory patterns involve seasonal north-south movements, often driven by temperature preferences between 5–15°C, allowing the to track optimal thermal conditions across its range. Genetic studies indicate distinct northern hemisphere (Atlantic) and southern hemisphere (Indo-Pacific) populations, with low divergence within regions but higher differentiation between hemispheres, supported by evidence from tagging recaptures showing limited inter-oceanic movement. Historical overfishing prior to the 2000s has led to significant population declines and localized range contractions in parts of the North Atlantic and some southern regions, reducing abundance in formerly core areas.

Habitat preferences

The spurdog (Squalus acanthias) is primarily a demersal species, inhabiting the bottom layers of coastal and waters where it favors sandy or muddy substrates for resting and foraging. These soft-bottom environments provide suitable conditions for the species to maintain its position near the , with individuals often observed in close with structural features such as rocky reefs or beds that offer and microhabitats. Spurdogs thrive in full-strength salinity, typically ranging from 32 to 36 , though they exhibit tolerance for slightly reduced salinities in brackish estuarine margins without venturing into freshwater. Water temperatures between 5 and 15°C are optimal, with peak abundances recorded in regions where conditions stabilize around 6–11°C, supporting their metabolic and behavioral needs. In terms of vertical distribution, spurdogs occupy nearshore waters during summer months when temperatures are favorable, shifting to deeper areas exceeding 200 m in winter to track cooler conditions. They frequently associate with zones along continental margins, where nutrient-rich waters influence their preferred depth ranges of 10–200 m, though records extend to over 700 m. Buoyancy regulation is facilitated by a large liver containing high levels of low-density oils, such as , enabling the species to hover efficiently over benthic substrates without constant swimming. Regionally, habitat preferences vary within the spurdog's temperate global distribution across the North Atlantic and southern oceans. In European waters, such as along the northwest continental shelf and estuarine edges of the Irish Sea and North Sea, spurdogs utilize shallow, soft-bottom bays and inlets up to 150 m deep. In North America, they predominantly occupy open continental shelves, including the Scotian Shelf and U.S. Northeast coast, favoring broader shelf habitats from nearshore mudflats to slopes at 100–300 m.

Biology and behavior

Reproduction and development

The spurdog (Squalus acanthias) exhibits ovoviviparous reproduction, in which eggs are fertilized internally and develop within the female's without a placental connection to the mother. Embryos are initially nourished by a substantial , which sustains them for the first several months; once depleted, they receive additional nutrients from uterine secretions known as histotroph, produced by the maternal reproductive tract. This aplacental viviparity allows for the protection and development of offspring inside the female until they are fully formed and ready for live birth. Mating involves , with males using one of their paired —modified pelvic fins—to insert sperm into the female's . The features specialized structures, including cartilaginous anchors at the tip, that secure it during , facilitating efficient sperm transfer. Breeding seasonality varies regionally; for instance, in the North Atlantic, fertilization often occurs during winter months from October to February, while in the , mating and pupping overlap in late winter to early spring. The gestation period lasts 18 to 24 months, the longest documented among shark species, reflecting the spurdog's slow life-history strategy. During this extended development, embryos grow at rates of approximately 1 cm per month in their second year, reaching full term without residual yolk. Females typically produce litters every two years due to the prolonged gestation, limiting their reproductive output. Recent studies in the North Pacific (as of 2023) report average uterine fecundity of about 4.1, with high reproductive potential across the region, though varying by population. Litter sizes range from 2 to 15 pups, with averages of 6 to 8 depending on the population and maternal condition; larger females bear more offspring. Pups are born live at lengths of 22 to 33 cm, fully independent and resembling miniature adults with functional spines and teeth. Fecundity, measured as the number of mature ovarian eggs or uterine embryos, increases linearly with female length and age, adding roughly one egg per 4 cm of additional body length. This positive relationship underscores the species' K-selected traits, where smaller, younger females contribute fewer offspring, resulting in an overall low reproductive rate that heightens vulnerability to population declines. The combination of small litters, biennial cycles, and delayed maturity thus constrains recovery potential in exploited populations.

Diet and feeding habits

The spurdog (Squalus acanthias) is a carnivorous predator whose diet primarily consists of small to medium-sized bony fishes such as herring (Clupea harengus) and whiting (Merlangius merlangus), cephalopods including squid (Loligo spp.), and crustaceans like euphausiids and crabs. Stomach content analyses from large-scale studies, including examinations of over 14,000 individuals, indicate that teleost fishes typically comprise 50-55% of the diet by occurrence, with crustaceans at 35% and cephalopods or other molluscs at around 5%, reflecting high fish biomass in consumed prey. As opportunistic feeders, spurdogs also prey on smaller , , and occasionally comb jellies or sea anemones, with diet composition exhibiting seasonal shifts driven by prey availability; fishes predominate in winter diets, while become more significant in summer. These shifts are evident in regional studies, such as those in waters, where pelagic prey accounts for about 80% of consumption overall. Spurdogs are active hunters that employ bursts of speed in ram-feeding to pursue and capture prey, often using to transport it, and function as predators in some contexts by striking from cover or schools. Their teeth, arranged in bands suited for grasping and cutting, facilitate processing of elusive or elongated prey through lateral head shaking, which effectively severs items into manageable pieces. Daily ration estimates, derived from metabolic rate and stomach evacuation models, are approximately 0.5% of body weight, supporting their energetic demands as mobile predators. Spurdogs occupy a mid-level trophic position of approximately 4.0, calculated from isotope and content analyses across populations. Ontogenetic diet shifts occur, with juveniles primarily consuming smaller invertebrates and crustaceans, transitioning to a fish-dominated diet around 650-700 mm total length as they grow.

Ecology

Social behavior and migration

Spiny dogfish (Squalus acanthias), also known as spurdogs, exhibit highly social behavior characterized by the formation of large schools comprising hundreds to thousands of individuals. These schools are often segregated by size, sex, and maturity stage, with groups typically consisting of either mature males, mature females, or immature sharks; mature females are frequently observed in inshore areas, while immature individuals tend to school offshore. This segregation pattern persists across populations, including in the western North Atlantic where adult females and males show distinct spatial distributions during certain seasons. Spurdog migrations follow annual cycles driven primarily by temperature changes, with populations moving northward along coastal areas in summer to avoid warmer temperatures and southward in winter to deeper offshore habitats. Tagging studies, including telemetry, have revealed distinct movement patterns: northern populations in the display localized residency with limited seasonal shifts, while southern groups off exhibit oscillatory inshore-offshore cycles, with overlap between groups occurring mainly in autumn from to . These migrations are tracked effectively through conventional and pop-up archival tags, confirming north-south traversals along the U.S. East Coast over distances of several hundred kilometers. In terms of daily activity, spurdog demonstrate patterns, occupying deeper waters during the day (mean depths of 25–100 m depending on population) and shallower areas at night (mean depths of 24–80 m), which aligns with increased nocturnal activity. Within , individuals display coordinated swimming, and the is known for aggressive interactions, particularly when pursuing prey in groups, enhancing group hunting efficiency. Regional variations include observations in waters, where spurdog form aggregations in sea lochs like , showing site fidelity during winter residency without strong evidence of size- or sex-based segregation in catches.

Predators and ecological role

Spurdogs occupy a mid-to-upper trophic position in marine food webs and are preyed upon by a variety of larger predators. Larger sharks, including the (Lamna nasus), actively consume as part of their diet, particularly in the North Atlantic where form a notable component of stomach contents. Other elasmobranchs, such as conspecific adults, also prey on smaller individuals. Piscivorous fish like (Gadus morhua), red hake (Urophycis chuss), and (Lophius americanus) target , especially in coastal and shelf habitats. Marine mammals, including seals and orcas (Orcinus orca), further contribute to predation pressure, with documented instances of these species feeding on in the North Atlantic and Pacific. As opportunistic predators, spurdogs play a key role in regulating prey populations, exerting top-down control on small , cephalopods, and such as , , and , which aligns with their generalist diet of benthic and pelagic species. In some coastal ecosystems, particularly along the northeast U.S. , function as a due to their high and broad prey overlap with commercially important groundfish like , helping to stabilize fluctuations in lower trophic levels through their voracious and flexible feeding habits. Their generalist nature allows them to buffer ecosystem variability by switching prey based on availability, thereby maintaining balance in dynamic environments. Spurdogs commonly host metazoan parasites, including cestodes (tapeworms) such as Griphobothrium species and copepods like Lernaeenicus spp., which attach to the gills, skin, or digestive tract; these infections are prevalent but typically exert low impact on host health and population dynamics.

Conservation and human interactions

Conservation status and threats

The spurdog (Squalus acanthias) is classified as Vulnerable on the globally, an assessment first made in 2006 and reaffirmed in 2020 due to ongoing population declines driven by exploitation and other pressures. Regional assessments vary, with the Northeast Atlantic population initially rated as in 2006 and updated to Endangered in 2015, reflecting severe historical depletions, while the former Pacific populations—now recognized as a distinct species (S. suckleyi)—are assessed as Least Concern as of 2024, indicating relative stability. Natural threats to spurdog populations include , which can alter preferred temperature ranges of 6–15°C and potentially disrupt migration patterns, though direct impacts on the species appear limited. , projected to lower pH by 0.8–1.2 units in northern seas by mid-century, indirectly affects spurdog by reducing prey availability through disruptions and impairing sensory behaviors like prey detection in related shark species. Bycatch remains a significant anthropogenic threat, with spurdog frequently captured incidentally in demersal fisheries such as gillnets and trawls, resulting in high post-capture mortality rates often exceeding 20% due to their slow recovery from stress. Habitat degradation exacerbates vulnerability, as bottom trawling damages benthic environments where spurdog aggregate, while pollutants like mercury bioaccumulate in their tissues—reaching levels up to 0.5 ppm in mature individuals—and microplastics are ingested by over half of examined specimens in the North-East Atlantic, posing risks to health and reproduction. Signs of recovery are evident in protected areas, particularly the Northeast Atlantic, where strict fishing bans since 2010 have led to a increase from approximately 316,000 tonnes in 2010 to 373,000 tonnes by 2017, alongside improved indices. This rebound, supported by the species' slow but persistent reproductive rate, underscores the potential for population stabilization under reduced fishing pressure, though full recovery may span decades.

Fisheries and management

Spurdog (Squalus acanthias) fisheries have a long history of exploitation, particularly in the North Atlantic, where targeted fishing intensified in the 20th century. Landings peaked in the 1990s, with annual catches exceeding 50,000 tonnes in the Northeast Atlantic, driven by demand for meat in Europe (such as for fish and chips) and byproducts like liver oil rich in vitamin A. In the Northwest Atlantic, U.S. landings reached approximately 27,000 tonnes in the mid-1990s, primarily targeting large females for export markets. These high-volume fisheries contributed to significant stock depletions by the early 2000s. Current spurdog fisheries are heavily regulated or prohibited in many regions to promote . Targeted was banned in the from 2011 to 2022, following a 90% reduction in total allowable catch (TAC) in 2010. The ban was lifted in 2023 with the introduction of shared TACs between the and , set at 22,309 tonnes for 2025, alongside measures for monitored in avoidance programs. In the United States, the Atlantic operates under quotas, with the 2024/2025 season allocated at 4,852 metric tons (about 10.7 million pounds), reduced to 4,236 metric tons for 2025/2026 to maintain . The U.S. Atlantic holds Marine Stewardship Council (MSC) certification for , covering harvests from to . Management measures emphasize precautionary approaches, including TAC systems, minimum landing sizes, and seasonal closures. In the Northeast Atlantic, the North East Atlantic Fisheries Commission (NEAFC) aligns with EU-UK agreements prohibiting directed fisheries beyond allocated TACs and requiring live release of where possible, supporting the 2025 TAC of 22,309 tonnes. U.S. regulations under the Magnuson-Stevens Act mandate federal permits, trip limits, and gear restrictions like prohibitions on overnight gillnet soaks to minimize bycatch of such as , effective from May 2025. Internationally, the International Council for the Exploration of the Sea (ICES) provides annual advice on stock status to inform EU and UK quotas, projecting scenarios to keep spawning biomass above critical thresholds. Economically, spurdog remains a low-value but supports high-volume , with exports from the U.S. primarily to for smoked or fresh fillets (known as "schillerlocken" in ). Byproducts, especially liver oil, historically provided for supplements, though synthetic alternatives have reduced this market; today, the focus is on meat for human consumption. Monitoring relies on stock assessments through trawl surveys, tagging studies, and fishery-dependent data to track abundance and fishing mortality. In the U.S., the Northeast Fisheries Science Center conducts biennial assessments, confirming the stock as not overfished in 2023, with spawning biomass at 191 million pups. ICES uses survey indices and landing reports for Northeast Atlantic evaluations, noting improving but ongoing vulnerability to illegal trade, addressed via genetic tracing in enforcement efforts.

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