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Palinurus elephas

Palinurus elephas, commonly known as the European spiny lobster or common spiny lobster, is a large species of spiny lobster in the family Palinuridae, endemic to the eastern Atlantic Ocean and Mediterranean Sea. It features a robust, heavily armored body covered in sharp spines, long spiny antennae that exceed the body length, and lacks large chelipeds, instead using small hook-like claws on the first pair of walking legs. Adults typically measure 40 cm in total length, with a maximum of 60 cm, and exhibit an orange dorsal coloration with darker spines and pale spots, though variations in brown, sandy, or purple hues occur.^[1]^[2]^[1] This nocturnal, benthic species inhabits rocky and coralligenous substrates, favoring crevices, caves, and ledges for shelter during the day, at depths ranging from 5 to 200 meters, though commonly found between 5 and 70 meters. Its distribution spans from southern Norway and the British Isles southward to Morocco, including the Azores, Canary Islands, and throughout the western Mediterranean, but it is absent from the extreme eastern and southeastern Mediterranean basins. Juveniles settle in shallower, more complex habitats like Posidonia oceanica meadows, while adults prefer deeper, structured environments with ample refuges.^[1]^[2]^[1] P. elephas is gonochoristic, with females reaching sexual maturity at a carapace length of 7–12.1 cm and males at 8–8.5 cm, typically breeding annually from June to November, with egg-laying peaking in September–October. Females carry eggs under the abdomen for about 8 months, hatching in early summer as phyllosoma larvae that undergo a prolonged pelagic phase before settling as pueruli. The species is omnivorous, feeding primarily on mollusks, echinoderms, crustaceans, macroalgae, and detritus during nocturnal foraging excursions.^[1]^[2]^[1] Commercially important, P. elephas supports artisanal fisheries using trammel nets and pots, particularly in the Mediterranean and Atlantic coasts, but populations have declined due to overexploitation, leading to its classification as Vulnerable (VU) on the IUCN Red List. Management measures include minimum landing sizes (e.g., 110 mm carapace length), closed seasons (e.g., May–August in some areas), and marine protected areas to promote recovery and sustainable yields. Local extinctions occurred in parts of southwest Britain in the 1970s, though signs of recovery have been observed since 2014.^[1]^[2]^[3]

Taxonomy

Nomenclature

Palinurus elephas was first described scientifically as Astacus elephas by the Danish entomologist Johan Christian Fabricius in his 1787 work Mantissa Insectorum Sistens Eorum Species Nuper Detectas Adjectis Characteribus Genericis, Differentiis Specificis, Emendationibus, Observationibus, with the type locality erroneously listed as the "Americae meridionalis Insulis" (islands of southern America); this has been corrected to the Mediterranean Sea based on the species' known distribution and specimen origins.^[4] The genus Palinurus was subsequently established by Franziscus Hermannus Weber in 1795, into which Fabricius's species was transferred, forming the current binomial Palinurus elephas (Fabricius, 1787), which remains the valid name under the International Code of Zoological Nomenclature (ICZN).^[5] The genus name Palinurus derives from the ancient Roman Tyrrhenian Sea port of Palinurus (modern Palinuro, Campania, Italy), a site renowned for the abundance of this lobster off its promontory, itself named after the helmsman Palinurus from Virgil's Aeneid.^[6] The specific epithet elephas, Latin for "elephant," likely alludes to the species' elongated, curved antennae that resemble elephant tusks.^[4] Prior to Fabricius's description, the species lacked a formal binomial name under the Linnaean system but appeared in early natural history literature and illustrations, often grouped under broader terms like locusta marina (sea locust) in Roman texts, reflecting its spiny, locust-like appearance; no specific misclassifications are recorded before 1787.^[7] Known synonyms include Astacus elephas Fabricius, 1787 (original combination, superseded); Cancer locusta de Wulfen, 1791 (junior synonym); and Palinurus vulgaris Latreille, 1803 (also junior synonym, sometimes attributed to Risso, 1816).^[4]

Classification

Palinurus elephas belongs to the kingdom Animalia, phylum Arthropoda, subphylum Crustacea, class Malacostraca, order Decapoda, infraorder Achelata, family Palinuridae, genus Palinurus, and species elephas.^[4] This hierarchy places it among the spiny lobsters, a group of marine decapod crustaceans characterized by a lack of true claws (chelipeds) and elongated bodies adapted to benthic lifestyles.^[8] Within the family Palinuridae, which comprises about 57 species across 9 genera, P. elephas serves as the type species of the genus Palinurus, established by its original description in 1787.^[4] The genus Palinurus is distinguished from related genera such as Panulirus primarily by the morphology of the antennal flagella, which in Palinurus are broad and strap-like, lacking the long, slender, whiplike extensions typical of Panulirus species.^[9] This structural difference aids in stridulation and is a key diagnostic trait in palinurid taxonomy.^[10] Palinurids represent an ancient lineage within the Achelata, with fossil evidence and molecular phylogenies indicating that the family's crown group diverged during the Late Triassic, around 225 million years ago, though major radiations occurred in the Jurassic.^[11] This evolutionary history underscores their persistence as a distinct group through Mesozoic marine environments, predating many modern decapod families.^[12]

Description

Morphology

Palinurus elephas possesses a distinctive body structure typical of spiny lobsters in the family Palinuridae, featuring an elongated cephalothorax and abdomen that together form a robust, armored exoskeleton. The cephalothorax, resulting from the fusion of the head and thorax, is encased in a hard carapace bearing numerous forward-directed spines, including prominent supraorbital spines above the eyes, which enhance protection against predators. Unlike true lobsters in the family Nephropidae, P. elephas lacks chelae or pincers on its appendages, relying instead on the spiny carapace and other defensive features. The abdomen comprises six distinct somites, each covered by chitinous tergites and pleurae, with additional rows of sharp spines along the dorsal margins to deter attacks.^[1]^[2]^[13] The antennae of P. elephas are prominent and multifunctional, consisting of long, whip-like structures that often exceed the length of the body. These include heavy, calcified stalks lined with robust spines for lashing at threats during defense, while the tapering flagellum is equipped with sensory setae for mechanoreception and chemoreception, aiding in navigation and foraging. The antennules, shorter paired appendages, complement the antennae by detecting chemical cues in the water column.^[1]^[2] The appendages of P. elephas are adapted for life on rocky seafloors. The pereopods, or walking legs, number five pairs emerging from the cephalothorax; these are spinose and chela-free, with the distal dactyli functioning as small hooks for gripping substrates and prey, facilitating movement over uneven terrain. On the abdomen, biramous swimmerets (pleopods) are present on the first five somites, assisting in subtle locomotion and, in females, modified with setae to secure eggs during brooding.^[2]^[13] Sensory organs in P. elephas enable effective environmental interaction. Compound eyes, mounted on short movable stalks, provide a broad visual field suited to dim underwater conditions. Statocysts, internal balance organs located in the cephalothorax near the antennal bases, detect gravitational and acceleratory forces, supporting orientation and equilibrium during movement.^[2]^[13]

Size and coloration

Palinurus elephas attains a maximum carapace length of 19.5 cm in males and 15.5 cm in females in the Mediterranean, with slightly higher maxima (up to 20 cm CL) reported in the Atlantic, corresponding to a total length of up to 60 cm.^[2]^[13] Females are generally smaller than males overall, with sexual dimorphism manifested in males achieving greater maximum sizes and exhibiting heavier body mass at equivalent carapace lengths, as well as narrower abdomens compared to the broader abdomens of females.^[2]^[14] Live specimens display a mottled coloration typically ranging from reddish-brown to orange dorsally, adorned with pale yellow or white spots on the carapace and abdominal segments, and a pale white ventral surface.^[2]^[1] Upon capture and cooking, the exoskeleton undergoes a pronounced color shift to reddish hues due to the denaturation of proteins binding the carotenoid pigment astaxanthin, releasing its free red form.^[15]

Distribution and habitat

Geographic range

Palinurus elephas, commonly known as the European spiny lobster, has a primary geographic range in the northeastern Atlantic Ocean, extending from the British Isles (including records from southern Norway) and the Madeira Islands southward to Mauritania, including the Azores and Canary Islands. This distribution encompasses coastal waters along the Iberian Peninsula, North Africa, and associated archipelagos, where the species occupies rocky habitats in temperate to subtropical conditions.^[2]^[16] The Mediterranean Sea serves as the main stronghold for P. elephas, with dense populations concentrated in the western and central basins, such as around the Balearic Islands, Sardinia, and Sicily. In contrast, populations are sparser in the eastern Mediterranean and the Adriatic Sea, where environmental conditions and historical exploitation have limited abundance.^[2]^[13] The species' range is primarily limited by temperature preferences, thriving in waters between 15 and 25°C, and is absent from colder northern Atlantic regions where temperatures drop below this threshold.^[1]^[17]

Habitat preferences

Palinurus elephas primarily inhabits coastal waters at depths ranging from 5 to 70 meters, although records extend to 200 meters in some areas. Juveniles settle and reside in shallower waters, typically up to 20 meters, while adults prefer deeper zones within this range.^[18]^[19]^[20] The species shows a strong preference for hard substrates such as rocky reefs, boulders, and coralligenous formations that provide ample crevices and micro-caves for shelter, while it avoids soft sediments like sand or mud. These structured habitats are essential for protection and are commonly found along exposed coasts.^[18]^[16] P. elephas requires temperate marine conditions with temperatures between 13 and 24°C, where it is sensitive to acute elevations above 24°C that can cause mortality; salinity levels of 35 to 38 ppt in well-oxygenated waters are optimal, as the species exhibits low tolerance to deoxygenation. Juveniles often utilize seagrass beds like Posidonia oceanica in shallower, well-aerated zones for additional cover.^[18]^[1]^[16] To evade predators including the common octopus (Octopus vulgaris) and various fish species, P. elephas employs diurnal sheltering behavior, retreating into vertical crevices, ledges, or caves during the day and emerging at night; this preference for shelters that minimize light entry enhances survival in predator-rich environments.^[18]^[21]^[22]

Life history

Reproduction

Palinurus elephas exhibits seasonal reproduction, with mating occurring from August to October in the Mediterranean Sea and June to November in the Atlantic Ocean.^[2] During this period, mature individuals aggregate in suitable habitats, where females extrude eggs that are fertilized externally by spermatophores deposited by males on the female's sternum. Larger females tend to mate and spawn earlier in the season compared to smaller ones.^[2] Females demonstrate high fecundity, producing clutches ranging from approximately 14,000 to over 200,000 eggs depending on body size, with mean values around 30,000 to 90,000 eggs reported across Mediterranean geographic sub-areas. Fecundity increases linearly or allometrically with carapace length (CL), as described by relationships such as F = 2428 × CL - 148,988 in some populations. Eggs are fertilized and immediately attached to the setose pleopods under the female's abdomen, forming a brood mass that she carries externally.^[2]^[23]^[24] Brooding lasts 4-5 months in the Mediterranean, with eggs developing through stages marked by increasing pigmentation until hatching in late winter (typically December to February); in the Atlantic, brooding extends to about 8 months, with hatching in early summer (April to June). Throughout incubation, females actively maintain the brood by grooming and fanning with pleopods to ensure oxygenation and prevent fungal growth, though natural egg loss can reach 14-26% due to various factors.^[2]^[1]^[23]^[24] The population sex ratio is generally close to 1:1, though it varies by region and fishing method, with some areas showing slight biases toward males or females. Sexual maturity is attained at a carapace length of 70-90 mm for females and 74-85 mm for males, differing by sex and location; this corresponds to an age of approximately 3-5.5 years, based on growth models from protected populations.^[2]^[25]^[24]^[1]

Growth and development

The larval phase of Palinurus elephas begins upon hatching from eggs carried by the female, with newly hatched individuals emerging as phyllosoma larvae. These planktonic larvae are leaf-like in form and spend 5–6 months in the Mediterranean Sea or up to 10–12 months in the Atlantic Ocean drifting in ocean currents, making them highly vulnerable to dispersal and environmental variability.^[1]^[26] During this period, phyllosomas undergo approximately 9–10 molts to progress through their instars, feeding on plankton and growing while remaining transparent to evade predators.^[27]^[26] Metamorphosis occurs when the final-stage phyllosoma transforms into a puerulus postlarva, a transparent, leaf-shaped stage adapted for active swimming toward suitable settlement habitats. This postlarval phase lasts about 2–3 weeks, during which the puerulus descends from the water column and seeks benthic substrates, marking the transition from pelagic to demersal life.^[28]^[26] Juveniles settle on coastal reefs, particularly limestone structures with crevices and empty bivalve holes, at a carapace length of 7.5–14 mm, typically between May and August in the Mediterranean when sea temperatures rise. Initial growth is rapid, driven by frequent molting—occurring 8–12 times in the first year post-settlement, with each molt increasing carapace length by about 2 mm—allowing juveniles to reach 17–19 mm by late autumn as they shift to larger shelters.^[19]^[1] The lifespan of P. elephas extends up to 25 years, with growth incrementing steadily through molting until sexual maturity at around 4–5 years, after which the rate slows significantly as energy is allocated to reproduction.^[1]^[26]

Ecology

Diet

Palinurus elephas is an omnivorous species that primarily consumes hard-shelled benthic organisms, including molluscs such as gastropods and bivalves, echinoderms like sea urchins, and crustaceans.^[29]^[2] Its diet is supplemented by macroalgae, coralline algae, polychaete worm tubes, bryozoans, and detritus, reflecting its opportunistic feeding strategy that adapts to local prey abundance.^[2]^[30] Diet composition shows ontogenetic shifts, with smaller individuals favoring gastropods and crustaceans, while larger ones incorporate more fish remains.^[29] The species engages in both active predation and scavenging, using its robust mouthparts—including mandibles and maxillipeds—to tear and crush food items, as it lacks chelae (claws) for grasping typical of clawed lobsters.^[1]^[31] This mechanism allows it to process a variety of prey, from soft-bodied invertebrates to shelled organisms, often detected through chemosensory cues during nocturnal foraging.^[2] As a mid-level carnivore and scavenger, P. elephas occupies a trophic level of approximately 2.6, positioning it as a secondary consumer within marine food webs.^[32] Its feeding habits contribute to benthic community dynamics by controlling populations of molluscs and echinoderms.^[1] Daily food intake varies with environmental factors such as temperature and prey availability, typically comprising a small percentage of body weight to support metabolic demands and pre-moult calcium accumulation.^[2] In controlled settings, related palinurids exhibit rations around 1-5% of body weight per day, suggesting similar patterns for P. elephas under optimal conditions.^[33]

Behavior

Palinurus elephas displays a strictly nocturnal activity rhythm, emerging from shelters approximately one hour before sunset to forage and retreating before dawn to avoid diurnal predators. During daylight hours, individuals shelter in rocky crevices, fissures, or under overhangs, often in small groups consisting of pairs or up to several individuals, which provides communal protection. This gregarious sheltering behavior is typical, though overall activity levels decrease during full moon phases due to heightened visibility risks.^[1]^[34] Locomotion in P. elephas primarily involves walking or crawling along rocky substrata using the pereiopods for slow, deliberate movement during foraging. For rapid escape from threats, the species employs tail-flip swimming, a burst propulsion mechanism that propels the lobster backward at high speeds. Additionally, seasonal migrations occur, with post-reproductive individuals moving to deeper waters in winter, triggered by declining temperatures and light intensity, before returning to shallower areas in spring.^[1]^[35]^[36] The social structure of P. elephas is characterized by solitary foraging or loose aggregations in shelters, with no evidence of strong territoriality among adults. Within shared shelters, subtle hierarchies may emerge based on size or dominance, influencing access to optimal positions, though interactions remain non-aggressive overall. Sensory behaviors rely heavily on chemoreceptors located on the antennae, which detect food odors and mate pheromones through specialized sniffing strategies involving high-frequency antennular flicking to sample water currents efficiently.^[37]^[38]

Human interactions

Fisheries and uses

Palinurus elephas has been targeted by fisheries since Roman times, when it was valued as a delicacy in the Mediterranean region.^[39] Historical records indicate that exploitation peaked in the mid-20th century, with Mediterranean landings reaching approximately 1,000 metric tons annually during the 1960s and 1970s, primarily driven by demand in coastal communities.^[25] These fisheries have since experienced declines due to intensified harvesting, though the species remains a key economic resource. Commercial harvesting of P. elephas primarily occurs in the Mediterranean Sea using baited wooden traps known as nasas, as well as hoop nets, pots, creels, trammel nets, and tangle nets.^[40]^[41] These methods are deployed by small-scale artisanal fleets, particularly in Spain, Italy, and other EU countries, with the highest landings reported from Spain and Italy.^[18] To manage stocks, seasonal closures are implemented in some areas, such as the EU-wide prohibition from January 1 to April 30, which partially overlaps with the breeding season. Current annual EU landings range from 300 to 400 metric tons, contributing to an export value exceeding €20 million, mainly to high-end markets in Europe and Asia.^[25]^[42] The species is prized for its firm, sweet meat, commonly featured in Mediterranean cuisine such as Spanish paella and grilled preparations seasoned with olive oil and lemon.^[43] Live specimens are often transported to premium markets to preserve freshness, fetching prices up to €120 per kg.^[44]^[45] This high demand underscores its economic importance, though capture typically targets individuals above a minimum size of 8-9 cm carapace length in major fishing nations like Spain and Italy.^[18]

Conservation status

Palinurus elephas is classified as Vulnerable (VU) on the IUCN Red List, with the assessment conducted on 26 September 2013 under criterion A2bd, indicating a population reduction inferred from observed declines in catch rates and habitat quality over the past three generations.^[3] The species' status reflects ongoing overexploitation and habitat loss, with a decreasing population trend observed across its range in the Mediterranean and adjacent Atlantic waters.^[46] The primary threats to P. elephas include intensive trap-based fisheries that target adults, leading to overfishing, as well as incidental bycatch in other gears where captured individuals suffer high mortality from handling or predation.^[47] Habitat degradation from coastal development and pollution further endangers the species by destroying rocky reef habitats essential for shelter and reproduction.^[48] Additionally, climate change poses risks by altering ocean temperatures and currents, which can disrupt larval dispersal patterns and reduce recruitment success, as warmer waters may shorten the pelagic larval duration and limit connectivity between populations.^[49] Conservation efforts include the establishment of Marine Protected Areas (MPAs) such as the Medes Islands and Cabrera Archipelago in the western Mediterranean, where no-take zones have demonstrated positive spillover effects, increasing adjacent fishery yields through emigration of adults and enhanced larval export.^[50] At the European Union level, Council Regulation (EC) No 1967/2006 prohibits the capture of berried females and sets a minimum landing size of 90 mm carapace length (CL) to protect immature individuals, though enforcement varies and no EU-wide quotas are currently imposed.^[47] National measures in some regions supplement these with seasonal closures during peak reproduction. Experimental aquaculture and restocking initiatives offer potential for population enhancement, with trials in Spain, such as those in the Columbretes Islands, releasing hatchery-reared juveniles into MPAs to boost local stocks.^[39] In France, facilities like the Stella Mare platform in Corsica have advanced larval rearing techniques, producing juveniles in controlled conditions despite persistent challenges in achieving high survival rates during the prolonged phyllosoma stage, which requires precise feeding and environmental control.^[51] These efforts highlight the feasibility of restocking but underscore the need for further research to overcome biological hurdles for scalable farming.^[44]

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### Summary of Palinurus elephas (Fabricius, 1787)
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[47]
Effect of light intensity on the survival of European spiny lobster ...
Consequently, the European spiny lobster is listed as a “vulnerable” species on the International Union for Conservation of Nature (IUCN) red list (Goñi, 2014).
[48]
[PDF] Press Kit 2023 - Université de Corse
Nov 7, 2023 · ... degraded by coastal development. It concerns the design of specific habitats for the various species studied within the other Stella Mare ...
[49]
Genetic Insights into the Population Connectivity, Biogeography ...
Jul 10, 2024 · Finally, the palinurids are well-known for the long duration of their larval phase, which ranges from four to over 12 months, with Panulirus ...
[50]
Spillover from six western Mediterranean marine protected areas
ABSTRACT: This study investigated spillover (biomass export) around 6 marine protected areas (MPAs) in the western Mediterranean based on catch and effort ...
[51]
European Spiny Lobster (PALINURUS ELEPHAS): The Stella Mare ...
Classified as a « vulnerable species », it is on the Red List of Threatened Species by the International Union for the Conservation of Nature (IUCN).Missing: status | Show results with:status