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Common periwinkle
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Common periwinkle
Periwinkle emerging from its shell, Sweden
L. littorea on the edge of a small sandy beach in Woods Hole, Massachusetts
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Subclass: Caenogastropoda
Order: Littorinimorpha
Family: Littorinidae
Genus: Littorina
Species:
L. littorea
Binomial name
Littorina littorea
Synonyms[2]
  • Littorina armoricana Locard, 1886
  • Littorina bartonensis Brown, 1843
  • Littorina communis Brown, 1843
  • Littorina litorea (Turton, 1819)
  • Littorina litorea var. brevicula Jeffreys, 1865
  • Littorina litorea var. paupercula Jeffreys, 1865
  • Littorina litorea var. sinistrorsa Jeffreys, 1865
  • Littorina litorea var. turrita Jeffreys, 1865
  • Littorina littorea f. intermedia Brøgger, 1901
  • Littorina littorea var. angulata Harmer, 1920
  • Littorina littorea var. antiqua Wood, 1848
  • Littorina littorea var. balteata Dautzenberg & Durouchoux, 1912
  • Littorina littorea var. complanata Harmer, 1920
  • Littorina littorea var. conica Harmer, 1920
  • Littorina littorea var. deformis Wood, 1848
  • Littorina littorea var. delphinula Wood, 1848
  • Littorina littorea var. distorta Harmer, 1920
  • Littorina littorea var. elegans Wood, 1848
  • Littorina littorea var. fuscofasciatus Middendorff, 1849
  • Littorina littorea var. major Dautzenberg & Durouchoux, 1913
  • Littorina littorea var. miniata Dautzenberg & Durouchoux, 1900
  • Littorina littorea var. pallida Dautzenberg & Durouchoux, 1900
  • Littorina littorea var. pallidefasciatus Middendorff, 1849
  • Littorina littorea var. parva Harmer, 1920
  • Littorina littorea var. picta Wood, 1848
  • Littorina littorea var. pyramidata Wood, 1848
  • Littorina littorea var. sanguinea Dautzenberg & Durouchoux, 1906
  • Littorina littorea var. truncata Harmer, 1920
  • Littorina littorea var. unicarinata Raeymaekers, 1889
  • Littorina parva Teilman-Friis, 1898
  • Littorina rudis var. aurantia Dautzenberg & P. Fischer, 1925
  • Littorina rudis var. reevei Harmer, 1921
  • Littorina sphaeroidalis Locard, 1886
  • Littorina vulgaris J. Sowerby, 1832
  • Turbo bicarinatus Woodward, 1833
  • Turbo carinatus Woodward, 1833
  • Turbo elongatus Woodward, 1833
  • Turbo litoreus Turton, 1819
  • Turbo littoreus Linnaeus, 1758 (basionym)
  • Turbo sulcatus Woodward, 1833
  • Turbo ustulatus Lamarck, 1822
  • Turbo ventricosus Woodward, 1833

The common periwinkle or winkle (Littorina littorea) is a species of small edible whelk or sea snail, a marine gastropod mollusc that has gills and an operculum, and is classified within the family Littorinidae, the periwinkles.[2]

This is a robust intertidal species with a dark and sometimes banded shell. It is native to the rocky shores of the northeastern, and introduced to the northwestern, Atlantic Ocean.

Description

[edit]
Shell of the common periwinkle

The shell is broadly ovate, thick, and sharply pointed except when eroded. The shell contains six to seven whorls with some fine threads and wrinkles. The color varies from grayish to gray-brown, often with dark spiral bands. The base of the columella is white.[3] The shell lacks an umbilicus. The white outer lip is sometimes checkered with brown patches. The inside of the shell is chocolate brown.

The width of the shell ranges from 10 to 12 millimetres (38 to 12 in) at maturity,[4] with an average length of 16 to 38 mm (58 to 1+12 in). Shell height can reach up to 30 to 52 mm (1+18 to 2 in),[4][5][3] The length is measured from the end of the aperture to the apex. The height is measured by placing the shell with the aperture flat on a surface and measuring vertically.[6]

L. littorea can be highly variable in phenotype, with several different morphs known. Its phenotypic variations may be indicative of speciation, as opposed to phenotypic plasticity. This is of particular importance to evolutionary biology, as it may represent an opportunity to observe a transitional phase in the evolution of an organism.[7]

Life cycle

[edit]

L. littorea is oviparous, reproducing annually with internal fertilization of egg capsules that are then shed directly into the sea, leading to a planktotrophic larval development time of four to seven weeks.[8] Females lay 10,000 to 100,000 eggs contained in a corneous capsule from which pelagic larvae escape and eventually settle to the bottom. This species can breed year round depending on the local climate. Benson suggests that it reaches maturity at 10 mm and normally lives five to ten years.[9] while Moore suggests that maturity is reached in 18 months.[10] Some specimens have lived 20 years.

Female specimens have been observed to be ripe from February until end of May, when most are spawning. Male specimens are mainly ripe from January until the end of May and lose weight after copulation. The young seem to settle primarily from the end of May to the end of June,[citation needed] although other sources indicate earlier settlement.[10]

Growth rate

[edit]

A study in Plymouth Sound suggests an initial growth reaching up to 14 mm (12 in) in height December the first year, and 17.4 mm (58 in) by the end of the second year. Females seem to grow more rapidly than males, and in specimens above 25 mm (1 in) in height, females seem to dominate.[10] Another study undertaken in Blackwater Estuary, Essex showed growth reaching up to 8 mm (38 in) the first winter.[11]

Distribution

[edit]

Common periwinkles are native to the northeastern coasts of the Atlantic Ocean, including northern Spain, France, Great Britain, Ireland, Scandinavia, and Russia.[9]

There have been more than 14,000 observations made available as a dataset at the Global Biodiversity Information Facility - Littorina littorea,[12] which can be explored. More distribution information can also be found at Ocean Biographic Information System - Littorina littorea.[13] The NBN Gateway - Littorina littorea has a distribution map over the UK and Ireland.[14] These datasets may overlap.

Introductions to North America

[edit]

Common periwinkles were introduced to the Atlantic coast of North America, possibly by rock ballast in the mid-19th century.[15] This species is also found on the west coast of the United States, from Washington to California. The first recorded sighting in the East was in 1840 in the Gulf of St. Lawrence.[15] It is now abundant on rocky shores from New Jersey northward to Newfoundland.[9][8] In Canada, its range includes New Brunswick, Nova Scotia, Quebec, Newfoundland and Labrador.[3]

L. littorea is now the most common marine snail along the North Atlantic coast. It has changed North Atlantic intertidal ecosystems via grazing activities, altering the distribution and abundance of algae on rocky shores and converting soft-sediment habitats to hard substrates, as well as competitively displacing native species.[9][8]

Ecology

[edit]

Habitat

[edit]

The common periwinkle is mainly found on rocky shores in the higher and middle intertidal zone. It sometimes lives in small tide pools. It may also be found in muddy habitats such as estuaries and can reach depths of 55 metres (180 feet).[9] When exposed to either extreme cold or heat while climbing, a periwinkle will withdraw into its shell and start rolling, which may allow it to fall to the water.[16]

Zone

[edit]

Movement both horizontally and vertically in response to light and dark as well as temperatures have been observed, but over a short timespan the movement seems to be random.

Experiments seem to indicate that the snail responds to light and current, and moves accordingly.[17]

Feeding

[edit]

L. littorea is an omnivorous, grazing intertidal gastropod.[8] It is primarily an algae grazer, but it will feed on small invertebrates such as barnacle larvae. It uses its radula to scrape algae from rocks and, in the salt marsh community, pick up algae from cord grass or from the biofilm that covers the surface of mud in estuaries or bays. Macroalgae that are readily consumed include Ulva lactuca and Ulva intestinalis;[18] if provided, blue mussel can also be eaten.

The radula is taenioglossate, consisting of seven teeth per row: one middle tooth, flanked on each side by one lateral and two marginal teeth. The radula is used to scrape algae and detritus.

FMIB 48521 Two rows of the radula of Littorina littorea L

Phlorotannins in the brown algae Fucus vesiculosus and Ascophyllum nodosum act as chemical defenses against L. littorea.[19]

Parasites

[edit]

The common periwinkle can act as a host for various parasites, including Renicola roscovita, Cryptocotyle lingua, Microphallus pygmaeus and Himasthla sp. More studies are needed before any conclusions regarding the effect of parasites on growth can be reached. It seems that growth rate is primarily affected on available food and time available for feeding, rather than parasites.[20]

Polydora ciliata has also been found to excavate burrows in the shell of the common periwinkle when the snail is mature (above 10 mm long). The reason why this happens only to mature snails is not yet known, but one hypothesis is that a mature snail will excrete a signal substance which attracts the P. ciliata larvae. Another hypothesis is that a mature snail has a change in the shell surface that makes it suitable for P. ciliata larvae to settle. The infection by this parasite does not seem to alter the growth and proportions of the snail shell.[21]

Mortality

[edit]

A mortality rate of up to 94% per annum has been observed for the first two months, followed by up to 60% per annum for the rest of the first year:

...out of every 950 shells of all ages [collected] at that time, 850 are first year, and 100 are in their second or subsequent year.[22]

Older individuals above 15 months old seem to have a mortality of only 23% per annum. Cercaria emasculans is known to be fatal to the snail, but this does not account for the observed mortality.[10]

Human use

[edit]
Remains of an ancient meal. Winkle shells from Cantabrian Lower Magdalenian layer (15 000 before present) in the Altamira cave

This species appears in prehistoric shellfish middens throughout Europe, and is believed to have been an important source of food since at least 7500 B.C.E. in Scotland.[23][full citation needed] It is still collected in quantity in Scotland, mostly for export to the Continent and also for local consumption. The official landings figures for Scotland indicate over 2,000 tonnes of winkles are exported annually. This makes winkles the sixth most important shellfish harvested in Scotland in terms of tonnage, and seventh most important in terms of value. However, since actual harvests are probably twice reported levels, the species may actually be the fourth and sixth most important, respectively.[24][full citation needed]

Periwinkles are usually picked off the rocks by hand or caught in a drag from a boat. They are mostly eaten in the coastal areas of Scotland, England, Wales and Ireland, where they are commonly referred to as winkles or in some areas buckies, willicks, or wilks. In Belgium, they are called kreukels or caracoles.

They are commonly sold in paper bags near beaches in Ireland and Scotland, boiled in their local seawater, with a pin attached to the bag to enable the extraction of the soft parts from the shell.

Periwinkles are considered a delicacy in African and Asian cuisines. The meat is high in protein, omega-3 fatty acids and low in fat; according to the USDA National Nutrient Database for Standard Reference, raw snails in general are about 80% water, 15% protein, and 1.4% fat.

Periwinkles are also used as bait for catching small fish. The shell is usually crushed and the soft parts extracted and put on a hook.

In accordance with their history as an ancient food source in Atlantic Europe, they are harvested and consumed in the Azores Islands by the Portuguese people, where they are usually called búzios, the generic name for sea snails.

Supply chain

[edit]

As for seafood supply chains in general, the supply chain consists of a collector, processor, distributor and finally the retailer. The true nature of the supply chain is usually more complex and opaque, with the potential for records of harvesting areas and date of catch to be falsified.

Collection

[edit]

Commonly harvested in buckets by workers walking in the intertidal zone on low tide; other methods have been tried.

In Maine, the snails are commonly collected by a dredge towed from a vessel.

In Norway, snorkeling has also been used.

A report on the state of the periwinkle industry in Ireland suggests a maximum catch size in order to preserve the population[citation needed].

External videos
video icon Dredging for periwinkles, YouTube video
video icon Snorkeling for periwinkles, YouTube video

Processor

[edit]

The processor buys in bulk from the collector, involving a possibly long transport route by land in a refrigerator truck or airplane, taking care to avoid temperatures below 0 °C.

If fresh seawater is readily available, the periwinkles are first graded if possible, using a machine custom built for the purpose. The method used for grading differs, but two proven methods include a Trommel screen with horizontal bars instead of a mesh, and a circle-throw vibrating machine also using bars. The price to purchase a complete sorting machine can be €10,000 or more.

Periwinkles are graded by number of snails per kilogram. The following table displays some common grades in France. The actual value depends upon supply and demand, with seasonal variations. The actual ranges may also differ from each establishment.

Grade name Number per kilogram
Small Unknown
Medium Unknown
Large 180–240
Jumbo 140–180
Super Jumbo 90–140

After grading, the periwinkles are "climbed" close to the consumer, which involves checking whether they are still alive. This can take anywhere from a few hours to several days, depending on how healthy the periwinkles are and the temperature of the water they climb in. Any periwinkles left immobile at the bottom are considered dead and are discarded. It is not uncommon to have up to 8% waste in a shipment.

Hereafter, the winkles are commonly packed in smaller quantities before being distributed to customers. Mesh bags from 3 to 10 kg are common.

Distributor

[edit]

To sell large quantities, distributors are commonly used to move the periwinkles to the retailer. These have networks of transport available both internationally, regionally and locally inside a city. Several distributors are usually involved in the complete journey, each focusing on its own part of the transport network.

Retail

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The common periwinkle is sold by fishmongers at seafood markets in large cities around the world, and is also commonly found in seafood restaurants as an appetizer or as a part of a seafood platter. In some countries, pubs may serve periwinkles as a snack.

Most of the volume fished is consumed by France, Belgium, Spain and the Netherlands.

Methods to increase commercial value

[edit]

Ongrowing has been investigated as a potential way of increasing commercial value, but no documented pilot facilities have been established. By harvesting the periwinkle during the summer and storing them with feed until December, not only should the grade have been increased, but the market value should be higher since supply is lower in the cold winter months.[25]

Aquaculture

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Raising the common periwinkle has not been a focus due to its abundance in nature and relatively low price; however, there are potential benefits from aquaculture of this species, including a more controlled environment, easier harvesting, less damages from predators, as well as saving the natural population from commercial harvesting.

Packaging

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Commonly packed in 3 kg boxes by the processor, the box is usually polystyrene foam or thin wood, depending on the market demands. Holes in the box ensures that any water lost by the snails drains out, so that they remain in better condition for longer. A label indicates the fishing zone, packaging date, and any other information required by law.

Storing

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In a refrigerator, the common periwinkle can usually be stored for up to a week, but this may vary depending on how long they have been stored prior to sale, and how they have been kept since the moment they are fished. As long as they are kept moist and cold, they can survive well for a longer period of time. It is not recommended to store at temperatures below 0 degrees Celsius, even if research has shown a Median Lower Lethal Temperature of -13.0 degree Celsius.[26] Even if the common periwinkle survives when put back into seawater, they seem to be unable to move and climb.

See also

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References

[edit]

Further reading

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[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
The common periwinkle (Littorina littorea), also known as the edible periwinkle, is a marine gastropod mollusc in the family , characterized by its sharply conical shell reaching a maximum height of 52 mm, typically dark grey-brown with spiral darker lines and a white . Native to the northeastern from northern to the White Sea in , 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 and species. This species is gonochoristic, with separate sexes, reaching 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. It forms dense aggregations on hard substrates and tolerates brackish conditions, playing a key role as a in coastal ecosystems by controlling algal growth and serving as prey for birds, , and . Introduced to around the 19th century—likely via transatlantic shipping—it has become invasive along the eastern seaboard from to , competing with native snails like Littorina saxatilis and altering marsh habitats through intensified herbivory on grasses such as . 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 often boiled and served with . Additionally, L. littorea acts as a for marine pollutants due to its sensitivity to contaminants; its was sequenced in 2025, aiding on its biology and invasiveness.

Taxonomy

Classification

The common periwinkle, Littorina littorea, belongs to the kingdom Animalia, phylum , class , order Littorinimorpha, family , genus Littorina, and species L. littorea. The Littorinidae family, to which L. littorea belongs, underwent significant diversification during the period (approximately 23–2.6 million years ago), with fossil records from the 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. Estimates based on these fossils and 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. 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. However, phylogenetic analyses indicate that L. littorea occupies a distinct position in the Littorina, forming a separate from the more closely related L. saxatilis and L. obtusata, which belong to the Neritrema. studies, including sequences of the oxidase subunit I (COI) , further confirm these genetic distinctions, revealing unique haplotypes for L. littorea that underscore its monophyletic status relative to its congeners.

Nomenclature

The binomial name of the common periwinkle is Littorina littorea (Linnaeus, 1758). The name Littorina derives from the Latin littor-, a stem of littus meaning "seashore," combined with the -ina, reflecting the shore-dwelling of the snails in this . The specific littorea is the feminine form of littoreus, meaning "of the shore" or "shore-related," emphasizing the species' intertidal lifestyle. The species was originally described by in the 10th edition of Systema Naturae per regna tria naturae, published in 1758, where it was initially named Turbo littoreus on page 761. 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. These synonyms were resolved under the principles of the (ICZN), which prioritizes the original Linnaean name and stabilizes Littorina littorea as the valid binomial within the family . In English, the species is commonly known as the common periwinkle, winkle, edible winkle, or wrinkled winkle. Equivalent common names in other languages include littorine commune or bigorneau commun in French.

Description

Shell characteristics

The shell of the common periwinkle, Littorina littorea, is solid and heavy, exhibiting an ovate-conical or shape with a prominent onion-like . It is dextrally coiled, typically featuring 6 to 7 whorls that are flat-sided with shallow sutures, and lacks an umbilicus. The is ovate, with a thin and sharp outer lip; the interior is often brown, while the is white. Mature shells commonly reach heights of 10 to 52 mm, providing robust protection in intertidal environments. The shell surface is sculptured with fine growth lines, low narrow axial , 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. width measures approximately 10 to 12 mm in adults. L. littorea demonstrates 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.

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. This structure, composed primarily of , fits snugly against the shell's opening, preventing and predation during exposure. The foot is a large, organ enabling locomotion across rocky substrates and to surfaces via a thin layer of pedal , which provides both for movement and stickiness for temporary attachment against wave dislodgement. The , a ribbon-like in the , features approximately 280 rows of teeth arranged in a taenioglossate pattern with seven teeth per transverse row, facilitating the grazing of and from substrates. Respiration occurs primarily through a ctenidium, or , located within the mantle cavity, where water is drawn in for via across the gill filaments into the hemocyanin-containing blood. The incorporates a well-developed in the mantle cavity, a chemosensory organ that detects waterborne chemical cues to guide and avoid predators. This radula-mediated grazing process, as detailed in the feeding section, underscores the periwinkle's role as an algal consumer in intertidal communities.

Reproduction and life history

Reproductive strategies

The common periwinkle, Littorina littorea, is gonochoristic, with separate sexes determined genetically and maintaining an approximately 1:1 in natural populations. Males are distinguished by the presence of a during the breeding season, while females possess a capsule for production. Reproduction involves internal fertilization achieved through copulation, during which the male everts its to transfer spermatophores directly into the female's reproductive tract. Males exhibit mate preference for larger females, which are typically more fecund, influencing success in this promiscuous . Breeding occurs primarily from to in the native North Atlantic range, triggered when temperatures exceed 10°C, though the exact timing varies with , local , and (e.g., earlier maximum spawning in January in estuaries). 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. True simultaneous hermaphroditism is absent, but conditions—where females develop male characteristics in the —have been reported at low frequencies in stressed populations exposed to pollutants like , potentially reducing fertility. These anomalies are rare in unimpacted habitats and do not represent a standard reproductive strategy.

Larval stages

The common periwinkle, Littorina littorea, is oviparous and releases egg capsules containing numerous embryos into the , where they develop into planktotrophic veliger larvae that rely on for nutrition during their free-swimming phase. The eggs typically hatch after 5–10 days, depending on temperature and , emerging as veliger larvae measuring about 180 μm in . These larvae spend 3–6 weeks in the , 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. 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. 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. 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. The extended planktonic duration provides significant dispersal potential, with larvae capable of traveling up to 100 km via ocean currents, promoting and recolonization of suitable habitats across the North Atlantic. 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 , , and other planktivores, as well as environmental stressors like temperature fluctuations.

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. Growth rates vary by , with females typically growing faster than males. This 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 and availability. 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. Abundant food resources, particularly and macroalgae, accelerate size attainment to maturation thresholds, while scarcity delays it. Adult L. littorea have a lifespan ranging from 5 to 20 years, with most individuals living 5-10 years in natural populations, though exceptional up to 20 years has been observed in . 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. In later years, is evident through reduced mobility and slower growth increments, leading to increased vulnerability to environmental stressors and predation.

Distribution

Native range

The common periwinkle (Littorina littorea) is native to the rocky intertidal shores of the northeastern , with its original geographic distribution spanning from the in northern northward to the in northern . This range encompasses the coastal regions of France, the , , and the , where the species occupies a variety of subtidal and intertidal habitats on moderately exposed to sheltered rocky shores. Prior to significant human-mediated dispersal in the , populations maintained relative stability across this extent, reflecting long-term adaptation to the region's post-glacial environmental dynamics. 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 . 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 at higher extremes. 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. Genetic studies reveal a cline in diversity across the native range, with the highest levels observed in southern European populations, such as those in , where gene diversity reaches up to 0.436. This southern richness contrasts with lower values in northern regions (e.g., 0.181 in ), a pattern driven by post-glacial recolonization from Iberian refugia following the , leading to successive founder effects and reduced variability northward. Such phylogeographic structure underscores the species' evolutionary history tied to Pleistocene climate fluctuations within its native Atlantic domain.

Introduced ranges

The common periwinkle, Littorina littorea, was introduced to the northwestern Atlantic coast of by the mid-19th century, with the first documented record occurring in the 1840s at , . This introduction likely originated from European populations, facilitated by transatlantic shipping vectors such as ballast rock discharge and hull fouling from vessels arriving from and . High propagule pressure from frequent ship arrivals—over 800 European vessels entering 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. By 1873, populations had reached , extending to by 1875, New Jersey by 1892, and further south to by 1970 and / between 1959 and 1989; today, established populations span from Red Bay, Labrador, to . Beyond the northwestern Atlantic, L. littorea has seen sporadic introductions elsewhere, though most have failed to establish self-sustaining populations. On the of , individuals were first noted in 1937, with larger groups appearing in (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 to Washington. In the Mediterranean, introductions occurred in and in 1978, but populations became extinct by 1988. Introductions to the remain unconfirmed and unestablished, with no verified records of persistence in regions like or . The species is classified as invasive in by the IUCN Global Invasive Species Database, reflecting its widespread establishment and displacement of native intertidal communities. Genetic analyses of introduced populations reveal a history of multiple events, with East Coast North American populations exhibiting relatively high mitochondrial diversity—suggesting repeated introductions possibly dating back to the late 18th or early , or even earlier Viking-era translocations—though overall is reduced compared to native European stocks, indicative of founder effects and bottlenecks. In contrast, Pacific Coast introductions show unexpectedly high without significant bottlenecks but lack the needed for persistence.

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. 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. 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. 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. 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. 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. To endure prolonged aerial exposure during low , L. littorea seals its shell operculum with , minimizing water loss and maintaining internal hydration. 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. These higher densities in protected areas reflect the ' preference for microhabitats with reduced wave action and greater moisture retention.

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. While predominantly herbivorous, L. littorea displays omnivorous tendencies by incidentally ingesting , , and small , such as juvenile or polychaetes, during radula scraping. These supplementary 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 . Feeding rates vary with age, size, and conditions, but young individuals can consume up to approximately 28 mg of per gram of body weight per day under optimal circumstances, supporting rapid growth and metabolic demands. Foraging behavior in L. littorea is closely synchronized with tidal cycles, with most occurring during submersion when movement is facilitated and desiccation risk is low; activity often persists nocturnally under constant conditions, reflecting an underlying . On exposed shores, individuals tend to forage more nocturnally to minimize daytime heat and 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. 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.

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, , is a primary predator, exerting strong lethal and sublethal effects by crushing shells and inducing behavioral changes such as reduced foraging in periwinkles. Shorebirds, including (Haematopus ostralegus and Haematopus palliatus), actively forage on periwinkles by hammering or stabbing their shells, particularly targeting exposed individuals in the . Various species, such as those in coastal waters, also consume periwinkles, contributing to overall mortality, especially among juveniles post-settlement. Predation collectively accounts for substantial adult mortality, with rates estimated at 20-30% annually in some populations, influencing periwinkle distribution and density. 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. 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. 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. 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 in overlapping zones. Symbiotic relationships include mutualism with epibiotic algae, such as encrusting species that provide against visual predators while gaining a stable substrate on the periwinkle's shell. As an in , 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.

Human interactions

Culinary and historical uses

The common periwinkle (Littorina littorea) has been exploited by humans since , with its shells frequently appearing in archaeological shell middens along the Atlantic coasts of , including sites in , , , and . These middens indicate early collection for food, dating back to the mid-Holocene period in regions like Denmark's , around 5000 BCE, where periwinkle shells served as a dietary staple in coastal settlements. In , periwinkle shells have been found in Norse settlements in Newfoundland dating to approximately 1000 CE and in Indigenous sites in around 1300 CE, reflecting consumption following its introduction from . During medieval and early modern periods in , periwinkles were a common source, particularly in coastal communities where they were harvested seasonally, often in early spring as a "starvation " to supplement dwindling winter supplies. Archaeological from sites like Skaill Farm in , , dated to the 14th–15th centuries and 17th–19th centuries, shows concentrated collection during , suggesting their role as an accessible protein source amid scarcity. While Roman records primarily document consumption of land snails, marine periwinkles like L. littorea likely contributed to broader diets in ancient , as inferred from similar assemblages. In culinary traditions, common periwinkles are typically prepared by or for 5–10 minutes in salted to loosen the , which is then extracted from the shell using a pin, , or by sucking directly from the . 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. Nutritionally, periwinkle 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. Beyond direct consumption, periwinkles have held cultural significance as since at least the , particularly in European and North American fisheries where they were collected to attract like and . Recent 2025 archaeological research utilizing periwinkle shells from middens in the broader European Atlantic, including sites in , has confirmed their prehistoric and medieval exploitation patterns, employing sclerochronology to reconstruct seasonal harvesting and environmental conditions.

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. In some regions, such as parts of the , 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 beds. Harvesting is largely unregulated in many areas, including , with no quotas or closed seasons enforced, though participants must comply with hygiene standards under Regulations (EC) No 852/2004 and 853/2004, requiring registration and record-keeping. Historically, annual yields from and exceeded 2,000 tonnes combined, with alone exporting around 3,651 tonnes in the late , though recent figures show a mean of 808 tonnes per year in from 2004 to 2015 due to fluctuating landings and reduced collector participation. As of 2024, periwinkle landings are not separately reported in national stock reviews, reflecting limited monitoring of this . The supply chain begins with collectors, often numbering around 500 part-time pickers , who sort periwinkles by size on-site or at collection points, discarding undersized individuals (typically below 13-16 mm, depending on local guidelines). Larger specimens (>16 mm ) fetch higher prices and are prioritized for markets. Collected periwinkles are transported to approximately 26 wholesalers or processors , where they undergo cleaning, purging in clean seawater to expel sand and reduce microbial loads, and sometimes depuration processes to minimize contaminants like or radionuclides. Distributors then package them live in ventilated containers for , primarily as fresh product, with some frozen options for longer transport. Major markets are export-oriented, with (especially , , the , and ) receiving the bulk for human consumption as a in seafood dishes. Shipments from northeastern , including and , also reach and other European countries via hubs like , supporting both food and emerging bait uses in the and for . 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 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.

Conservation and impacts

In , the common periwinkle (Littorina littorea) is considered an , particularly along the northeastern Atlantic coast, where it has significantly altered intertidal community structures by grazing on ephemeral and facilitating the dominance of slower-growing like Chondrus crispus over native macroalgae such as Fucus vesiculosus. 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 like Littorina saxatilis and Littorina obtusata, through resource competition and predation on eggs, thereby reshaping food webs and parasite communities. The 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 to . Globally, L. littorea has no assessment and is not evaluated for , reflecting its widespread abundance in native European ranges where it is considered of least concern. 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 , linked to ocean warming and reduced recruitment. is prompting northward range shifts in native populations, with fossil and subfossil evidence suggesting historical expansions into cooler waters like under warmer conditions, a pattern that may accelerate with ongoing global temperature rises. Management efforts focus on regulatory measures and monitoring rather than widespread eradication, as the species is well-established in many areas. In the , harvesting is regulated with minimum size limits—typically 13–16 mm shell height in regions like the —to protect juveniles and sustain stocks, though no quotas exist for commercial quantities. In the United States, eradication trials on the , including in , 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.

References

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