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Common periwinkle

The common periwinkle (Littorina littorea), also known as the edible periwinkle, is a marine gastropod mollusc in the family Littorinidae, characterized by its sharply conical shell reaching a maximum height of 52 mm, typically dark grey-brown with spiral darker lines and a white columella.^[1] Native to the northeastern Atlantic Ocean from northern Spain to the White Sea in Russia, it inhabits rocky and estuarine shores in the intertidal to shallow sublittoral zones up to 60 m depth, where it grazes on microalgae such as Ulva and Fucus species.^[1]^[2] This species is gonochoristic, with separate sexes, reaching sexual maturity at a shell height of 10-12 mm and a lifespan of 5-10 years; breeding occurs annually from February to June, producing 10,000 to 100,000 eggs per female in gelatinous capsules that hatch into planktonic larvae.^[1] It forms dense aggregations on hard substrates and tolerates brackish conditions, playing a key role as a primary consumer in coastal ecosystems by controlling algal growth and serving as prey for birds, fish, and crabs.^[1]^[2] Introduced to North America around the 19th century—likely via transatlantic shipping—it has become invasive along the eastern seaboard from Labrador to Delaware, competing with native snails like Littorina saxatilis and altering marsh habitats through intensified herbivory on grasses such as Spartina alterniflora.^[2]^[3] Economically, it supports commercial fisheries, particularly in Ireland where annual harvests have been valued at approximately €1.6 million (based on 2014 data), and is harvested for human consumption as a delicacy often boiled and served with vinegar.^[1]^[4] Additionally, L. littorea acts as a bioindicator for marine pollutants due to its sensitivity to contaminants; its genome was sequenced in 2025, aiding research on its biology and invasiveness.^[1]^[5]

Taxonomy

Classification

The common periwinkle, Littorina littorea, belongs to the kingdom Animalia, phylum Mollusca, class Gastropoda, order Littorinimorpha, family Littorinidae, genus Littorina, and species L. littorea.^[2] The Littorinidae family, to which L. littorea belongs, underwent significant diversification during the Neogene period (approximately 23–2.6 million years ago), with fossil records from the Pliocene epoch (5.3–2.6 million years ago) documenting the trans-Arctic interchange that contributed to the origins of northern Atlantic clades, including ancestors of L. littorea.^[6] Estimates based on these fossils and molecular clock calibrations suggest that L. littorea diverged from related lineages around 5–10 million years ago, reflecting adaptations to cool-temperate intertidal environments during this period of climatic cooling and habitat shifts.^[6] Within the genus Littorina, L. littorea shares close evolutionary ties with species such as L. saxatilis and L. obtusata, all of which inhabit similar rocky intertidal zones in the North Atlantic.^[7] However, phylogenetic analyses indicate that L. littorea occupies a distinct position in the subgenus Littorina, forming a separate clade from the more closely related L. saxatilis and L. obtusata, which belong to the subgenus Neritrema.^[7] Mitochondrial DNA studies, including sequences of the cytochrome oxidase subunit I (COI) gene, further confirm these genetic distinctions, revealing unique haplotypes for L. littorea that underscore its monophyletic status relative to its congeners.^[8]^[9]

Nomenclature

The binomial name of the common periwinkle is Littorina littorea (Linnaeus, 1758).^[10] The genus name Littorina derives from the Latin littor-, a stem of littus meaning "seashore," combined with the suffix -ina, reflecting the shore-dwelling habitat of the snails in this genus.^[11] The specific epithet littorea is the feminine form of littoreus, meaning "of the shore" or "shore-related," emphasizing the species' intertidal lifestyle.^[11] The species was originally described by Carl Linnaeus in the 10th edition of Systema Naturae per regna tria naturae, published in 1758, where it was initially named Turbo littoreus on page 761.^[12] Over time, numerous synonyms accumulated due to taxonomic revisions and regional descriptions, including Nerita littorea Linnaeus, 1758; Littorina vulgaris J. Sowerby, 1832; Turbo bicarinatus S. Woodward, 1833; and Turbo ustulatus Lamarck, 1822.^[10] These synonyms were resolved under the principles of the International Code of Zoological Nomenclature (ICZN), which prioritizes the original Linnaean name and stabilizes Littorina littorea as the valid binomial within the family Littorinidae.^[10] In English, the species is commonly known as the common periwinkle, winkle, edible winkle, or wrinkled winkle.^[10]^[3] Equivalent common names in other languages include littorine commune or bigorneau commun in French.^[10]

Description

Shell characteristics

The shell of the common periwinkle, Littorina littorea, is solid and heavy, exhibiting an ovate-conical or turban shape with a prominent onion-like spire. It is dextrally coiled, typically featuring 6 to 7 whorls that are flat-sided with shallow sutures, and lacks an umbilicus. The aperture is ovate, with a thin and sharp outer lip; the interior is often brown, while the columella is white. Mature shells commonly reach heights of 10 to 52 mm, providing robust protection in intertidal environments.^[2]^[3] The shell surface is sculptured with fine growth lines, low narrow axial ribs, and subtle spiral lines, contributing to its textured appearance. Coloration ranges from grayish-brown to black, frequently accented by lighter spiral bands or lines (typically 8 to 25 narrow black or brown ones on a pale brown base), though variations include uniform cream, orange, or red hues. The apex is often lighter or white in juveniles. Aperture width measures approximately 10 to 12 mm in adults.^[2]^[1] L. littorea demonstrates phenotypic plasticity in shell morphology, particularly in response to environmental conditions. Shell thickness increases with greater wave exposure, with studies showing up to 20% greater thickness in high-energy sites compared to sheltered areas, enhancing resistance to mechanical stress.^[13]

Internal anatomy

The common periwinkle, Littorina littorea, possesses a horny operculum that is spiral in shape, typically consisting of 2–3 turns, which serves to close the shell aperture when the animal retracts its soft body for protection.^[14] This structure, composed primarily of chitin, fits snugly against the shell's opening, preventing desiccation and predation during low tide exposure.^[15] The foot is a large, muscular organ enabling locomotion across rocky substrates and adhesion to surfaces via a thin layer of pedal mucus, which provides both lubrication for movement and stickiness for temporary attachment against wave dislodgement.^[15] The radula, a ribbon-like structure in the mouth, features approximately 280 rows of teeth arranged in a taenioglossate pattern with seven teeth per transverse row, facilitating the grazing of microalgae and detritus from substrates.^[16] Respiration occurs primarily through a ctenidium, or gill, located within the mantle cavity, where water is drawn in for gas exchange via diffusion across the gill filaments into the hemocyanin-containing blood.^[15] The nervous system incorporates a well-developed osphradium in the mantle cavity, a chemosensory organ that detects waterborne chemical cues to guide foraging and avoid predators.^[15] This radula-mediated grazing process, as detailed in the feeding ecology section, underscores the periwinkle's role as an algal consumer in intertidal communities.^[15]

Reproduction and life history

Reproductive strategies

The common periwinkle, Littorina littorea, is gonochoristic, with separate sexes determined genetically and maintaining an approximately 1:1 sex ratio in natural populations.^[1]^[17] Males are distinguished by the presence of a penis during the breeding season, while females possess a capsule gland for egg production.^[1] Reproduction involves internal fertilization achieved through copulation, during which the male everts its penis to transfer spermatophores directly into the female's reproductive tract.^[18]^[19] Males exhibit mate preference for larger females, which are typically more fecund, influencing mating success in this promiscuous species.^[1] Breeding occurs primarily from February to June in the native North Atlantic range, triggered when seawater temperatures exceed 10°C, though the exact timing varies with latitude, local climate, and habitat (e.g., earlier maximum spawning in January in estuaries).^[20] Females are iteroparous, producing multiple broods per season and releasing eggs in gelatinous, planktonic capsules shed directly into the water; each brood contains 10,000 to 100,000 eggs across numerous capsules, each typically holding 2 to 9 eggs.^[21]^[20] True simultaneous hermaphroditism is absent, but intersex conditions—where females develop male characteristics in the reproductive system—have been reported at low frequencies in stressed populations exposed to pollutants like tributyltin, potentially reducing fertility.^[22] These anomalies are rare in unimpacted habitats and do not represent a standard reproductive strategy.^[1]

Larval stages

The common periwinkle, Littorina littorea, is oviparous and releases egg capsules containing numerous embryos into the seawater, where they develop into planktotrophic veliger larvae that rely on phytoplankton for nutrition during their free-swimming phase.^[2] The eggs typically hatch after 5–10 days, depending on temperature and salinity, emerging as veliger larvae measuring about 180 μm in length.^[23] These larvae spend 3–6 weeks in the plankton, with the duration influenced by environmental conditions such as water temperature (optimal between 7–20°C) and food availability, allowing for extended development before competency to settle.^[23]^[1] Veliger larvae of L. littorea possess a prominent ciliated velum, a lobed structure that facilitates both locomotion through ciliary beating and filter-feeding on suspended particles, enabling survival in the water column.^[2] A shell gland within the larval mantle secretes the initial protoconch, a translucent, chitinous shell that forms the embryonic whorls and measures approximately 0.275 mm in height at settlement.^[2] Distinctive pigmentation, often red or black spots on the velar lobes, aids in identification. Metamorphosis from veliger to juvenile occurs upon encountering suitable settlement cues, such as chemical signals from macroalgae like Fucus species or bacterial films on intertidal rocks, prompting the resorption of the velum and the development of a functional foot and operculum.^[23]^[24] The extended planktonic duration provides significant dispersal potential, with larvae capable of traveling up to 100 km via ocean currents, promoting gene flow and recolonization of suitable habitats across the North Atlantic.^[25] This mobility contributes to the species' broad distribution but is tempered by high mortality rates exceeding 90% during the larval phase, largely attributable to predation by zooplankton, fish, and other planktivores, as well as environmental stressors like temperature fluctuations.^[26]

Growth patterns

Following settlement, juvenile common periwinkles (Littorina littorea) exhibit rapid post-larval growth, reaching an average shell height of approximately 14 mm by the end of their first year and 17.4 mm by the end of the second year.^[27] Growth rates vary by sex, with females typically growing faster than males.^[28] This sexual dimorphism in growth contributes to females dominating larger size classes in mature populations. Sexual maturation occurs at a shell height of 10-12 mm, generally around one year of age, though this can be influenced by environmental factors such as temperature and food availability.^[1] Optimal growth and maturation are supported at water temperatures of 15-20°C, where metabolic rates and feeding efficiency are maximized; above 20°C, growth slows due to physiological stress.^[1] Abundant food resources, particularly microalgae and macroalgae, accelerate size attainment to maturation thresholds, while scarcity delays it.^[29] Adult L. littorea have a lifespan ranging from 5 to 20 years, with most individuals living 5-10 years in natural populations, though exceptional longevity up to 20 years has been observed in aquaria.^[29] Age can be estimated from incremental growth rings on the shell, formed annually due to seasonal variations in growth rates, with clearer rings appearing during slower winter periods.^[27] In later years, senescence is evident through reduced mobility and slower growth increments, leading to increased vulnerability to environmental stressors and predation.^[28]

Distribution

Native range

The common periwinkle (Littorina littorea) is native to the rocky intertidal shores of the northeastern Atlantic Ocean, with its original geographic distribution spanning from the Bay of Biscay in northern Spain northward to the White Sea in northern Russia.^[1] This range encompasses the coastal regions of France, the British Isles, Scandinavia, and the Baltic Sea, where the species occupies a variety of subtidal and intertidal habitats on moderately exposed to sheltered rocky shores.^[30] Prior to significant human-mediated dispersal in the 19th century, populations maintained relative stability across this extent, reflecting long-term adaptation to the region's post-glacial environmental dynamics.^[1] Archaeological evidence indicates the presence of L. littorea in human settlements along the European Atlantic coast dating back to Roman times, with shells frequently recovered from middens that highlight its early role as a dietary resource.^[31] Population densities are highest in temperate zones within this native range, particularly where average water temperatures fall between 10 and 20°C, conditions that support optimal growth and reproduction while avoiding thermal stress at higher extremes.^[1]^[23] These densities contribute to the species' dominance as a key intertidal grazer in such areas, with stable abundances documented prior to the 1800s through consistent archaeological occurrences.^[31] Genetic studies reveal a cline in diversity across the native range, with the highest levels observed in southern European populations, such as those in Portugal, where gene diversity reaches up to 0.436.^[32] This southern richness contrasts with lower values in northern regions (e.g., 0.181 in Northern Europe), a pattern driven by post-glacial recolonization from Iberian refugia following the Last Glacial Maximum, leading to successive founder effects and reduced variability northward.^[32] Such phylogeographic structure underscores the species' evolutionary history tied to Pleistocene climate fluctuations within its native Atlantic domain.^[33]

Introduced ranges

The common periwinkle, Littorina littorea, was introduced to the northwestern Atlantic coast of North America by the mid-19th century, with the first documented record occurring in the 1840s at Pictou, Nova Scotia.^[2] This introduction likely originated from European populations, facilitated by transatlantic shipping vectors such as ballast rock discharge and hull fouling from vessels arriving from Great Britain and Ireland.^[34] High propagule pressure from frequent ship arrivals—over 800 European vessels entering Pictou Harbor between 1773 and 1861—contributed to successful establishment, with the species rapidly spreading southward at an initial rate of approximately 25–50 km per year.^[34] By 1873, populations had reached Maine, extending to Massachusetts by 1875, New Jersey by 1892, and further south to Delaware by 1970 and Maryland/Virginia between 1959 and 1989; today, established populations span from Red Bay, Labrador, to Lewes, Delaware.^[2]^[3] Beyond the northwestern Atlantic, L. littorea has seen sporadic introductions elsewhere, though most have failed to establish self-sustaining populations. On the Pacific coast of North America, individuals were first noted in 1937, with larger groups appearing in San Francisco Bay (1968) and Anaheim Bay (2002), likely transported via the live seafood trade from East Coast sources; however, these consist solely of adults without evidence of reproduction or larval recruitment, preventing long-term establishment from California to Washington.^[2]^[35] In the Mediterranean, introductions occurred in Italy and Sicily in 1978, but populations became extinct by 1988.^[2] Introductions to the southern hemisphere remain unconfirmed and unestablished, with no verified records of persistence in regions like Australia or South America.^[36] The species is classified as invasive in North America by the IUCN Global Invasive Species Database, reflecting its widespread establishment and displacement of native intertidal communities.^[36] Genetic analyses of introduced populations reveal a history of multiple colonization events, with East Coast North American populations exhibiting relatively high mitochondrial haplotype diversity—suggesting repeated introductions possibly dating back to the late 18th or early 19th century, or even earlier Viking-era translocations—though overall genetic diversity is reduced compared to native European stocks, indicative of founder effects and bottlenecks.^[2]^[37] In contrast, Pacific Coast introductions show unexpectedly high genetic diversity without significant bottlenecks but lack the reproductive success needed for persistence.^[35]

Ecology

Habitat requirements

The common periwinkle, Littorina littorea, primarily inhabits the intertidal zone of rocky shores, where it is most abundant from the mid- to low-tide levels, corresponding to depths of approximately 0-2 m during low tide.^[1] This species extends into the shallow sublittoral zone but becomes less common beyond the immediate intertidal area, with rare occurrences up to 60 m depth in low-density populations.^[1] It thrives on hard substrates such as bedrock, boulders, and stones, as well as biogenic structures like mussel beds (Mytilus edulis), which provide stable attachment points in sedimentary intertidal environments.^[1]^[38] L. littorea exhibits broad tolerances to environmental fluctuations typical of its intertidal habitat, including salinity levels from approximately 14 to 40 ppt, enabling it to occupy both fully marine and brackish conditions in estuaries.^[23] Temperature tolerance spans sub-zero conditions, where adults can survive freezing of over 50% of their extracellular body fluids, up to around 30°C, though growth and reproduction are optimal between 7 and 20°C.^[1]^[23] For refuge, individuals frequently associate with macroalgal canopies such as Fucus vesiculosus and Ascophyllum nodosum, which offer protection from desiccation and predation in the upper intertidal.^[3]^[39] To endure prolonged aerial exposure during low tides, L. littorea seals its shell operculum with mucus, minimizing water loss and maintaining internal hydration.^[1] Population densities vary by exposure, reaching up to 900 individuals per m² in sheltered estuarine habitats with sedimentary substrates, while averaging around 300 per m² on exposed rocky coasts.^[40] These higher densities in protected areas reflect the species' preference for microhabitats with reduced wave action and greater moisture retention.^[40]

Feeding ecology

The common periwinkle, Littorina littorea, functions primarily as an herbivorous grazer in intertidal ecosystems, using its radula—a chitinous, tooth-bearing ribbon in the mouth—to scrape microalgae, diatoms, and macroalgae from rock and sediment surfaces. Preferred macroalgae include ephemeral green species such as Ulva lactuca and Enteromorpha intestinalis (now often classified under Ulva), which provide high-quality nutrition and are selectively consumed over tougher perennials like fucoids. This feeding method targets biofilms rich in diatoms and other unicellular algae, allowing the snail to efficiently harvest microbial layers that form on hard substrates.^[41]^[42]^[1]^[39] While predominantly herbivorous, L. littorea displays omnivorous tendencies by incidentally ingesting detritus, bacteria, and small invertebrates, such as juvenile barnacles or polychaetes, during radula scraping. These supplementary food sources contribute to its nutritional intake, particularly in nutrient-poor environments, and underscore its role as a generalist consumer that processes a broad spectrum of organic matter. Feeding rates vary with age, size, and conditions, but young individuals can consume up to approximately 28 mg of food per gram of body weight per day under optimal circumstances, supporting rapid growth and metabolic demands.^[3]^[43] Foraging behavior in L. littorea is closely synchronized with tidal cycles, with most grazing occurring during submersion when movement is facilitated and desiccation risk is low; activity often persists nocturnally under constant conditions, reflecting an underlying circadian rhythm. On exposed shores, individuals tend to forage more nocturnally to minimize daytime heat and desiccation stress, whereas in sheltered habitats, diurnal activity may predominate due to reduced wave action and more stable moisture. This temporal patterning influences energy allocation and predator avoidance.^[44]^[45] Through its grazing, L. littorea exerts significant control over algal community structure, preventing the overgrowth of fast-colonizing ephemeral species like Ulva and Enteromorpha while promoting diversity by clearing space for slower-growing algae. High densities can bulldoze sediments and inhibit mat formation, maintaining bare substrates that favor diatom biofilms over dense canopies; this top-down pressure is particularly evident in moderately exposed intertidal zones, where snail populations shape habitat heterogeneity.^[46]^[47]^[48]

Interactions with other organisms

The common periwinkle, Littorina littorea, faces significant predation pressure from a variety of marine and avian predators across its range. The European green crab, Carcinus maenas, is a primary predator, exerting strong lethal and sublethal effects by crushing shells and inducing behavioral changes such as reduced foraging in periwinkles.^[49] Shorebirds, including oystercatchers (Haematopus ostralegus and Haematopus palliatus), actively forage on periwinkles by hammering or stabbing their shells, particularly targeting exposed individuals in the intertidal zone. Various fish species, such as those in coastal waters, also consume periwinkles, contributing to overall mortality, especially among juveniles post-settlement.^[50] Predation collectively accounts for substantial adult mortality, with rates estimated at 20-30% annually in some populations, influencing periwinkle distribution and density.^[1] Parasitism represents another key biotic interaction for L. littorea, with several trematode species altering host physiology and behavior. The trematode Renicola roscovita infects periwinkles as an intermediate host, often leading to parasitic castration that impairs reproduction and reduces macroalgal consumption rates.^[51] Similarly, Cryptocotyle lingua causes sterility, retarded growth, altered zonation patterns, and decreased heat tolerance in infected snails, with metacercariae encysting in tissues and correlating with foot discoloration.^[52] The shell-boring polychaete Polydora ciliata further impacts periwinkles by excavating cavities in shells, weakening structural integrity and increasing vulnerability to predators; prevalence varies by habitat but affects 10-50% of individuals in sedimentary shores.^[53] Interspecific competition occurs prominently with the rough periwinkle Littorina saxatilis, particularly for intertidal space and resources, where L. littorea inhibits the growth and abundance of the native species in overlapping zones.^[54] Symbiotic relationships include mutualism with epibiotic algae, such as encrusting species that provide camouflage against visual predators while gaining a stable substrate on the periwinkle's shell.^[55] As an invasive species in North America, L. littorea outcompetes native snails like Littorina saxatilis and mud snails (Ilyanassa obsoleta), displacing them through resource monopolization and altering community structure in the Northwest Atlantic.^[56]^[30]

Human interactions

Culinary and historical uses

The common periwinkle (Littorina littorea) has been exploited by humans since prehistoric times, with its shells frequently appearing in archaeological shell middens along the Atlantic coasts of Europe, including sites in Spain, Portugal, Denmark, and the United Kingdom.^[31] These middens indicate early human collection for food, dating back to the mid-Holocene period in regions like Denmark's Limfjord, around 5000 BCE, where periwinkle shells served as a dietary staple in coastal settlements.^[57] In North America, periwinkle shells have been found in Norse settlements in Newfoundland dating to approximately 1000 CE and in Indigenous sites in Nova Scotia around 1300 CE, reflecting consumption following its introduction from Europe.^[31] During medieval and early modern periods in Europe, periwinkles were a common food source, particularly in coastal communities where they were harvested seasonally, often in early spring as a "starvation food" to supplement dwindling winter supplies.^[31] Archaeological evidence from sites like Skaill Farm in Orkney, Scotland, dated to the 14th–15th centuries and 17th–19th centuries, shows concentrated collection during April, suggesting their role as an accessible protein source amid food scarcity.^[31] While Roman records primarily document consumption of land snails, marine periwinkles like L. littorea likely contributed to broader shellfish diets in ancient Atlantic Europe, as inferred from similar midden assemblages.^[4] In culinary traditions, common periwinkles are typically prepared by boiling or steaming for 5–10 minutes in salted water to loosen the meat, which is then extracted from the shell using a pin, toothpick, or by sucking directly from the aperture.^[58] This method yields a tender, mildly sweet flesh often served in escargot-style dishes with garlic, butter, wine, or herbs, akin to preparations in French and West African cuisines.^[59] Nutritionally, periwinkle meat is high in protein at approximately 15.2 g per 100 g and provides significant iron content of about 3.5 mg per 100 g, making it a valuable source of essential nutrients for coastal diets.^[60]^[61] Beyond direct consumption, periwinkles have held cultural significance as fishing bait since at least the 19th century, particularly in European and North American fisheries where they were collected to attract species like cod and lobster.^[3] Recent 2025 archaeological research utilizing periwinkle shells from middens in the broader European Atlantic, including sites in Orkney, has confirmed their prehistoric and medieval exploitation patterns, employing sclerochronology to reconstruct seasonal harvesting and environmental conditions.^[31]

Commercial harvesting

Commercial harvesting of the common periwinkle (Littorina littorea) primarily involves hand-picking from intertidal zones during low tides, particularly spring tides, by part-time collectors who target lower shore areas where densities are highest.^[62] In some regions, such as parts of the northeastern United States, small lightweight drag nets (known as wrinkle drags, limited to 6 feet or less) are used in shallow waters to collect periwinkles from rocks and seaweed beds.^[63] Harvesting is largely unregulated in many areas, including Ireland, with no quotas or closed seasons enforced, though participants must comply with EU hygiene standards under Regulations (EC) No 852/2004 and 853/2004, requiring registration and record-keeping.^[62]^[64] Historically, annual yields from Scotland and Ireland exceeded 2,000 tonnes combined, with Ireland alone exporting around 3,651 tonnes in the late 1990s, though recent figures show a mean of 808 tonnes per year in Ireland from 2004 to 2015 due to fluctuating landings and reduced collector participation. As of 2024, periwinkle landings are not separately reported in national shellfish stock reviews, reflecting limited monitoring of this fishery.^[62]^[4]^[65] The supply chain begins with collectors, often numbering around 500 part-time pickers in Ireland, who sort periwinkles by size on-site or at collection points, discarding undersized individuals (typically below 13-16 mm, depending on local guidelines).^[62]^[4] Larger specimens (>16 mm in England) fetch higher prices and are prioritized for export markets.^[4] Collected periwinkles are transported to approximately 26 wholesalers or processors in Ireland, where they undergo cleaning, purging in clean seawater to expel sand and reduce microbial loads, and sometimes depuration processes to minimize contaminants like heavy metals or radionuclides.^[62]^[66] Distributors then package them live in ventilated containers for export, primarily as fresh product, with some frozen options for longer transport.^[67] Major markets are export-oriented, with Europe (especially France, Belgium, the Netherlands, and Spain) receiving the bulk for human consumption as a delicacy in seafood dishes.^[62] Shipments from northeastern North America, including Canada and Maine, also reach Asia and other European countries via hubs like Boston, supporting both food and emerging bait uses in the US and Canada for recreational fishing.^[67]^[3] Retail occurs through fishmongers, seafood markets, and restaurants, often sold live or boiled for immediate preparation. Value is enhanced through size grading and depuration, which ensures compliance with food safety standards and improves marketability, though challenges persist from overharvesting, evidenced by a more than 20% decline in Irish landings and picker numbers since 2000, alongside localized stock reductions.^[4]^[68]

Conservation and impacts

In North America, the common periwinkle (Littorina littorea) is considered an invasive species, particularly along the northeastern Atlantic coast, where it has significantly altered intertidal community structures by grazing on ephemeral algae and facilitating the dominance of slower-growing species like Chondrus crispus over native macroalgae such as Fucus vesiculosus.^[2] This herbivory has led to shifts in algal composition, with studies indicating significant reductions in Fucus cover in affected areas due to competitive exclusion and changes in succession dynamics. Additionally, L. littorea has displaced or suppressed native gastropods, including species like Littorina saxatilis and Littorina obtusata, through resource competition and predation on eggs, thereby reshaping food webs and parasite communities.^[2] The species is monitored by the Centre for Agriculture and Bioscience International (CABI) as a high-risk invader in North American ecosystems, with its range extending from Labrador to Delaware.^[3] Globally, L. littorea has no IUCN Red List assessment and is not evaluated for conservation status, reflecting its widespread abundance in native European ranges where it is considered of least concern.^[61] However, local population declines have been observed in both native and introduced ranges due to overharvesting and climate-driven stressors; for instance, intertidal gastropod abundances, including periwinkles, have decreased steadily over the past two decades in the Gulf of Maine, linked to ocean warming and reduced recruitment.^[69] Climate change is prompting northward range shifts in native populations, with fossil and subfossil evidence suggesting historical expansions into cooler waters like Siberia under warmer conditions, a pattern that may accelerate with ongoing global temperature rises.^[3] Management efforts focus on regulatory measures and monitoring rather than widespread eradication, as the species is well-established in many areas. In the European Union, harvesting is regulated with minimum size limits—typically 13–16 mm shell height in regions like the UK—to protect juveniles and sustain stocks, though no quotas exist for commercial quantities.^[70] In the United States, eradication trials on the Pacific coast, including in California, have largely failed to establish persistent populations due to reproductive limitations and environmental mismatches, resulting in transient adult-only aggregations that naturally decline without intervention.^[71]

References

[1]
Common periwinkle (Littorina littorea) - MarLIN
This the largest British periwinkle, with the shell reaching a maximum height of 52 mm. The shell is sharply conical with a pointed apex and surface ...Description · Habitat preferences · Physical pressures · Chemical pressures
[2]
Littorina littorea - Marine Invasions research at SERC
Littorina littorea is known as the Common Periwinkle. It is native to Europe from the White Sea, Russia to Gibraltar. It has been introduced to the West and ...
[3]
Littorina littorea (common periwinkle) | CABI Compendium
L. littorea is invasive in North America with a current range from Red Bay, Labrador, to Lewes, Delaware.
[4]
phylogeny and evolutionary rates of the gastropod genus Littorina
We have used fossil evidence and geological events to estimate the ages of some clades, and thus to calculate average rates of molecular evolution, which in ...
[5]
[PDF] USING PARASITES TO INFORM ECOLOGICAL HISTORY
littorea is classified into the subgenus Littorina; thus phylogenetically, L. saxatilis and L. obtusata are more closely related to one another than to L.Missing: position | Show results with:position
[6]
Mitochondrial DNA CoI haplotype variation in sibling species of ...
Jul 1, 2000 · These data show that there is some sharing of haplotypes between species, with L. arcana haplotypes paraphyletic with respect to L. saxatilis haplotypes.
[7]
Species and gene divergence in Littorina snails detected by array ...
Genetic differentiation of L. saxatilis ecotypes was not detected in this study, despite pronounced innate phenotypic differences.
[8]
WoRMS - World Register of Marine Species - Littorina littorea (Linnaeus, 1758)
- **Binomial Name**: Littorina littorea
[9]
LITTORINA Definition & Meaning - Merriam-Webster
Word History. Etymology. New Latin, from Latin littor-, littus seashore + New Latin -ina ...
[10]
https://www.marinespecies.org/aphia.php?p=taxdetails&id=140262
[11]
Phenotypic Plasticity in an Intertidal Snail - jstor
1981) indicated that wave exposure accounted for 84.4% of the variance in shell thickness, whereas sites within wave exposure accounted for only 4.2% of the ...
[12]
Habitat-specific size structure variations in periwinkle populations ...
Oct 15, 2008 · Mean (9.8 ± 3.9 mm) and maximum (22 mm) shell size of L. littorea on the sedimentary shore were smaller than on the rocky shore (21.5 ± 4.2 and ...Missing: dimensions | Show results with:dimensions
[13]
[PDF] the prosobranch
Littorina littorea (Linnaeus). No locality given. Shell. Solid and opaque, not glossy. The spire is rather prominent and pointed (apical angle c. 85° in ...
[14]
[PDF] SNAILS - the Etter Lab
FIGURE 1 An illustration of a typical marine snail showing the major features of the external and internal anatomy. ... The common periwinkle Littorina littorea.
[15]
Elemental analyses reveal distinct mineralization patterns in radular ...
in R. horei, and K — in Paramelania damoni ... Here the length and width of each radula were measured and the quantity of tooth rows ...
[16]
[PDF] INTERNAL DEFENCE REACTIONS OF LITTORINA LITTOREA G.J. ...
1.3 Littorina littorea ... from the crystalline style of the stomach. Waste ... stomach and intestine by emptying the blood into the visceral haemocoel.
[17]
Effects of mucus trail following on the distance between individuals ...
Mar 10, 2020 · ... sex ratio of 1:1. Individuals were distributed randomly across ... Littorina littorea. Marine Ecology Progress Series. 1999;179:247–257 ...
[18]
[PDF] THE BIOLOGY OF TERRESTRIAL MOLLUSCS - Regulations.gov
Cyclophoroideans are dioecious, with internal fertilization. There is only ... Littorina littorea (Linnaeus) (Littorinidae: Caenogastropoda). (forward ...
[19]
Multiple Paternity in Littorina obtusata (Gastropoda, Littorinidae ...
Abstract. Parental identity for juvenile Littorina obtusata was determined from three egg masses by means of micro- satellite DNA markers.
[20]
[PDF] Common perivvinkle (Littorina littorea)
Mar 21, 2019 · Size and growth rate measurements apply to shell height. Most work suggests that maturity is reached at between 10-12mm shell height.
[21]
[PDF] Summary Report of Nonindigenous Aquatic Species in U.S. Fish ...
Feb 29, 2004 · females lay 10,000-100,000 eggs contained in a horny capsule from ... Water Resources, Sacramento, 66 pp. Jackson, A., 2002. Littorina littorea.
[22]
Comparative assessment of reproductive impairment in the ...
Apr 1, 2009 · In this study we present the results of an intersex survey of Littorina littorea along the Belgian coast. Levels of female intersex and ...Missing: hermaphroditism | Show results with:hermaphroditism
[23]
Effects of temperature, salinity and diet on embryonic and early ...
Figure 3 Exemplary images of Littorina littorea larvae hatched from eggs incubated at (A) 4.8°C, (B) 6.9°C, and (C) 11.3°C highlighting typical abnormal ...Missing: breeding | Show results with:breeding
[24]
Settlement cues and their effect on gene flow in sibling ... - SICB
Jan 5, 2021 · We cultured larvae in concentrations of 50-200/L in still flasks. We fed the larvae daily concentrations of Isochrysis galbana and ...
[25]
[PDF] Do differences in developmental mode shape the potential for local ...
Mar 1, 2020 · ... dispersal over 10 -100 km. 166 ... potential Littorina littorea; a relationship not observed in the low dispersal-potential Littorina.<|control11|><|separator|>
[26]
Semibalanus balanoides and Littorina spp. on exposed to ...
Aug 1, 2024 · The strong muscular foot and a thin film of mucus ... Evaluation of the significance of metal-binding proteins in the gastropod Littorina littorea ...
[27]
The Biology of Littorina littorea. Part I. Growth of the Shell and ...
Part I. Growth of the Shell and Tissues, Spawning, Length of Life and Mortality. Published online by Cambridge University Press: 11 May 2009.
[28]
Snail shape and growth rates: Evidence for plastic shell allometry in ...
globose shells. We experimentally demonstrate that this shell variation is a function of snail growth rate. Rapidly growing.
[29]
[PDF] Periwinkle Ecology and Size of Maturity Study - NIFCA
Sexual selection on female size in a marine snail, Littorina littorea. (L.). Journal of Experimental Marine Biology and Ecology, 181(2), pp.145-157. Fish ...<|control11|><|separator|>
[30]
Littorina littorea - The Exotics Guide
Littorina littorea is a European periwinkle, around 38mm tall, with a brown, gray-brown or blackish shell, found in eastern North America.
[31]
Shells of Littorina littorea (Gastropoda) - PubMed Central - NIH
Jul 10, 2025 · Littorina littorea—the common periwinkle—is biogeographically widely distributed and occurs frequently in archaeological shell middens along ...<|control11|><|separator|>
[32]
Unique genetic variation at a species' rear edge is under threat from ...
May 18, 2011 · Gene diversity values by population ranged from 0.105 (Øygarden) to 0.436 (Portugal), and by region ranged from 0.181 (Northern Europe) to 0.343 ...1. Introduction · 2. Material And Methods · 3. Results
[33]
Genomic population structure and local adaptation in the emerging ...
Jun 25, 2025 · littorea had a post-glacial history ... In this study, we applied genomic tool to assess genetic heterogeneity in Littorina littorea.
[34]
Historical invasions of the intertidal zone of Atlantic North America ...
May 19, 2009 · ... Littorina littorea. Both are conspicuous, cooccurring members of ... 100 km. This high level of spatial precision cannot be achieved ...
[35]
A Case History of Littorina littorea in California, USA | PLOS One
We characterized three newly discovered populations of an introduced gastropod, Littorina littorea (Linné, 1758), in California, USA, comparing them to ...
[36]
Littorina littorea - GISD
Mar 11, 2005 · Littorina littorea is a mollusc which is often distributed on rocky coasts, from the upper shore into the sub-littoral.
[37]
The Resolution of Littorina Littorea's North American Origin - PubMed
Based on mitochondrial sequences of 370 L. littorea and 196 C. lingua individuals, our results demonstrate a significant reduction in genetic diversity in North ...
[38]
Direct and indirect effects of Littorina littorea (L.) on barnacles ...
Aug 6, 2025 · For example, molluscan grazers can negatively affect the settlement and recruitment of barnacles through the detachment of newly settled larvae ...
[39]
δ 18 O-inferred salinity from Littorina littorea (L.) gastropods in a ...
Nov 8, 2019 · The lower range of salinity tolerance of L. littorea is unknown, but it can thrive in salinities >11.5–14 psu through to fully marine ...
[40]
Littornia and Fucus: Effects of Herbivores, Substratum Heterogeneity ...
Littorina snails speed up succession by eating early algae, releasing Fucus, but can prevent Fucus establishment on smooth surfaces. Substratum microtopography ...
[41]
Dietary preferences of the common periwinkle, Littorinalittorea (L.)
Littorina littorea (L.), was highly selective in favour of the foliose ephemeral green algae, Ulva lactuca L. and Enteromorpha intestinalis (L.) Link.Missing: microalgae diatoms
[42]
The susceptibility of benthic microalgae to periwinkle (Littorina ...
L. littorea is a moderately selective periphyton grazer. It fed on all algal species present, but the tube-dwelling diatoms Amphipleura rutilans and Haslea ...
[43]
Effects of feeding level and body size on food utilization of the ...
Feeding rate increased from 3.4 to 28.7, 2.0 to 19.1 and 0.2 to 7.2 mg/g/day in young, intermediate and old snails respectively. The corresponding increase in ...
[44]
Interactions of the circadian and circatidal rhythms of the littoral ...
Snails acclimated to non-tidal conditions were active at night. This rhythm persisted under constant conditions for 5–10 days, after which it too faded out.
[45]
https://www.tandfonline.com/doi/full/10.1080/15376516.2025.2581130
[46]
[PDF] Grazing on green algae by the periwinkle Littorina
Although Enteromorpha and Ulva are shown to be high quality food, Littorina littorea clearly prefers firm substrates to mats of green algae. As slow moving ...
[47]
Ocean acidification but not nutrient enrichment reduces grazing and ...
We tested whether a common marine herbivorous snail, Littorina littorea, would increase consumption rates of macroalgae (Ulva and Fucus) under ocean ...Snail Grazing (live Algal... · Results · Grazing (live Algal Thalli)Missing: Enteromorpha | Show results with:Enteromorpha
[48]
(PDF) Grazing on green algae by the periwinkle Littorina littorea in ...
Aug 7, 2025 · At normal snail densities, L. littorea grazing bulldozes sediments from hard substrate and precludes the presence of an algal canopy.
[49]
[PDF] I Ain't Afraid of No Crab: Intertidal Gastropod Littorina Littorea ...
Littorina littorea is an invasive species in Maine, genetic evidence suggests they were introduced to Nova Scotia from Europe. They were first found in New.Missing: taxonomic | Show results with:taxonomic
[50]
[PDF] Periwinkle (Littorina littorea) - à www.publications.gc.ca
Egg production increases directly with shell height. Females in the 14 mm size class are releasing less than. 1000 embryos compared to over 50,000 for. 19 mm ...
[51]
The influence of trematodes on the macroalgae consumption by the ...
Jul 29, 2008 · Littorina littorea serves as first intermediate host to a range of trematode species that also include benthic invertebrates, fish and ...Introduction · Results · DiscussionMissing: diet microalgae diatoms
[52]
The influence of Cryptocotyle lingua (Digenea:Platyhelminthes ...
Infection was found to retard growth and correlate with reduced shell height in some animals. A strong relationship was found between foot colour and infection.
[53]
[PDF] Habitat-specific size structure variations in periwinkle populations ...
Apr 30, 2010 · Wave exposure is considered one of the most important factors affecting gastropod shell size distribution. For example, Hylleberg and ...
[54]
Growth inhibition of native Littorina saxatilis (Olivi) by introduced L ...
The presence of L. littorea depressed the growth rate of L. saxatilis, while L. littorea grew better in the presence of L. saxatilis than conspecifics.Missing: spread | Show results with:spread
[55]
[PDF] Distribution of epibionts and their effects on the marine snail Littorina ...
The snail's shell can be exploited by epibiotic organisms such as barnacles, bryozoans, and encrusting calcareous algae. However, an in- depth examination of ...
[56]
Periwinkles Fast Facts (U.S. National Park Service)
Aug 24, 2021 · Shell: The hard shell is part of the periwinkle's body and grows with it. Their shells can be up to 3 cm long. It provides protection from ...
[57]
Periwinkle (Littorina littorea) intrashell δ18O and δ13C records from ...
Aug 7, 2025 · (1) The relationship of dry body weight to shell height in Littorina littorea ... Stable isotopes, elemental distribution, and growth rings of ...
[58]
valuing and managing traditional shellfish gathering for Littorina ...
The lifecycle of L. littorea has a planktonic larval stage. Reproduction involves internal fertilization with egg capsules released into the plankton. Following ...1 Introduction · 3 Results · 4 Discussion<|control11|><|separator|>
[59]
Periwinkles - Serious Eats
To eat a periwinkle, you must bring the opening of the shell to your lips and suck: not too forcefully like a vacuum but not too gently, either. At first you'll ...
[60]
Periwinkle Escargot with Sweet & Sour Tamarind Sauce (Ốc Xào Me)
20 minMar 12, 2020 · A Vietnamese street food favorite. Periwinkle sea snails tossed in a tangy, savory-sweet tamarind sauce make. Easy to make and full of flavor.
[61]
Periwinkle, common, Norwegian - Matvaretabellen
100g of Norwegian periwinkle has 87 kcal, 1.4g fat, 15.2g protein, 3.3g carbohydrate, 80g water, and 367 kJ energy.
[62]
Littorina littorea, Common periwinkle : fisheries - SeaLifeBase
In shallow and deep subtidal mussel beds of Mytilus edulis. Occurs in both hard substrates and in soft bottoms, particularly on stones, boulders, on muddy and ...
[63]
[PDF] Final Report 2002 - Marine Institute Open Access Repository
A shell height of approximately 8-9mm is achieved by the end of the first year (Williams,. 1964). This increases to about 16mm by the end of the third year. A ...
[64]
Maine Seafood Guide - Periwinkles & Whelks
Periwinkles are collected by hand (“wrinkling”) or with a 6' or smaller drag. See the vessel and gear guide for more information.
[65]
Uptake and depuration of 131 I by the edible periwinkle Littorina ...
This study will help to make radiological assessments more realistic by introducing a verifiable concentration factor and biological half-life into the ...
[66]
Periwinkles an under-heralded seafood product - Island Institute
Mar 20, 2017 · A commercial fishing license with a periwinkle permit is required to gather and sell periwinkles; 2 quarts are allowed for personal use. The ...
[67]
https://www.islandinstitute.org/working-waterfront/periwinkles-an-under-heralded-seafood-product/
[68]
https://www.researchgate.net/publication/255602237_An_Assessment_of_the_Potential_for_the_Sustainable_Development_of_the_Edible_Periwinkle_Littorina_littorea_Industry_in_Ireland
[69]
Declines over the last two decades of five intertidal invertebrate ...
Oct 20, 2020 · ... Littorina littorea, and Nucella lapillus). The declines in mussels and the common periwinkle (L. littorea) are correlated with warming sea ...Missing: breeding | Show results with:breeding
[70]
Common periwinkle - Rating ID: 1319 | Good Fish Guide
Over the last five years, recorded landings of periwinkle (Littorina spp.) have more than halved from 21 tonnes in 2018 to 9 tonnes in 2022, whilst their ...Missing: yield | Show results with:yield
[71]
A Case History of Littorina littorea in California, USA - PubMed Central
Jan 10, 2011 · L. littorea is oviparous, reproducing annually with internal fertilization of egg capsules that are then shed directly into the sea, leading to ...
[72]
Responses of Littorina spp. Intertidal Snails to Thermal Extremes ...
Jan 29, 2025 · Here, we investigate this relationship with a study of the survival of Littorina littorea, L. ... larvae that undergo long-range dispersal, while ...