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''Littorina saxatilis'',
common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often con ...
the rough periwinkle, is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of small
sea snail Sea snails are slow-moving marine (ocean), marine gastropod Mollusca, molluscs, usually with visible external shells, such as whelk or abalone. They share the Taxonomic classification, taxonomic class Gastropoda with slugs, which are distinguishe ...
, a marine
gastropod Gastropods (), commonly known as slugs and snails, belong to a large Taxonomy (biology), taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and fro ...
mollusc Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
in the family
Littorinidae The Littorinidae are a taxonomic family of over 200 species of sea snails, marine gastropod molluscs in the clade Littorinimorpha, commonly known as periwinkles and found worldwide. Names In English-speaking countries, gastropod molluscs from ...
, the winkles or periwinkles. First identified in the 1700s, it has been misidentified as a new species 112 times.


Distribution

This species is native to the shores of the North
Atlantic Ocean The Atlantic Ocean is the second largest of the world's five borders of the oceans, oceanic divisions, with an area of about . It covers approximately 17% of Earth#Surface, Earth's surface and about 24% of its water surface area. During the ...
, including
Hudson Bay Hudson Bay, sometimes called Hudson's Bay (usually historically), is a large body of Saline water, saltwater in northeastern Canada with a surface area of . It is located north of Ontario, west of Quebec, northeast of Manitoba, and southeast o ...
,
Baffin Island Baffin Island (formerly Baffin Land), in the Canadian territory of Nunavut, is the largest island in Canada, the second-largest island in the Americas (behind Greenland), and the fifth-largest island in the world. Its area is (slightly smal ...
,
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
, and the
Barents Sea The Barents Sea ( , also ; , ; ) is a marginal sea of the Arctic Ocean, located off the northern coasts of Norway and Russia and divided between Norwegian and Russian territorial waters.World Wildlife Fund, 2008. It was known earlier among Russi ...
, south along the American East Coast to
Chesapeake Bay The Chesapeake Bay ( ) is the largest estuary in the United States. The bay is located in the Mid-Atlantic (United States), Mid-Atlantic region and is primarily separated from the Atlantic Ocean by the Delmarva Peninsula, including parts of the Ea ...
, and along the European coast to the
Straits of Gibraltar The Strait of Gibraltar is a narrow strait that connects the Atlantic Ocean to the Mediterranean Sea and separates Europe from Africa. The two continents are separated by 7.7 nautical miles (14.2 kilometers, 8.9 miles) at its narrowest point. Fe ...
. Only one population of this intertidal species occurred in the Mediterranean basin i.e., Venice lagoon. The Venice population represents the earliest confirmed introduction of an exotic species in the Mediterranean Sea. Abiotic environmental features such as salinity and water temperature have influenced the past and current distributions of this snail and limited its invasion of the Mediterranean Sea. This species has also been introduced to
San Francisco Bay San Francisco Bay (Chochenyo language, Chochenyo: 'ommu) is a large tidal estuary in the United States, U.S. state of California, and gives its name to the San Francisco Bay Area. It is dominated by the cities of San Francisco, California, San ...
, on the
West Coast of the United States The West Coast of the United States, also known as the Pacific Coast and the Western Seaboard, is the coastline along which the Western United States meets the North Pacific Ocean. The term typically refers to the Contiguous United States, contig ...
, where it was first observed in 1992.


Shell description

The shell in life often appears green with algae, but the shell itself can be white, red, or brown, sometimes with checkered lines. The
shell Shell may refer to: Architecture and design * Shell (structure), a thin structure ** Concrete shell, a thin shell of concrete, usually with no interior columns or exterior buttresses Science Biology * Seashell, a hard outer layer of a marine ani ...
has 4–5
whorls A whorl ( or ) is an individual circle, oval, volution or equivalent in a whorled pattern, which consists of a spiral or multiple concentric objects (including circles, ovals and arcs). In nature File:Photograph and axial plane floral diagram ...
. Maximum recorded shell length is 19 mm.''Littorina saxatilis'' (Olivi)
Malacolog 4.1.1. A Database of Western Atlantic Marine Mollusca.
Welch J. J. (2010). "The "Island Rule" and Deep-Sea Gastropods: Re-Examining the Evidence". '' PLoS ONE'' 5(1): e8776. . File:Littorina saxatilis shell.png, Apertural view File:Littorina saxatilis 003.jpg, Abapertural view File:Littorina saxatilis001.jpg, A series of shells of ''Littorina saxatilis''


Ecology


Habitat

This species frequently lives in
salt marsh A salt marsh, saltmarsh or salting, also known as a coastal salt marsh or a tidal marsh, is a coastal ecosystem in the upper coastal intertidal zone between land and open saltwater or brackish water that is regularly flooded by the tides. I ...
es. it can also be found in crevices of intertidal
bedrock In geology, bedrock is solid rock that lies under loose material ( regolith) within the crust of Earth or another terrestrial planet. Definition Bedrock is the solid rock that underlies looser surface material. An exposed portion of bed ...
, in empty
barnacle Barnacles are arthropods of the subclass (taxonomy), subclass Cirripedia in the subphylum Crustacean, Crustacea. They are related to crabs and lobsters, with similar Nauplius (larva), nauplius larvae. Barnacles are exclusively marine invertebra ...
shells, and under rocks. Like many other periwinkles, this species can survive long exposures out of the water. The species has been recorded alive from depth range 0 – 46 m or up to 183 m (for shells only). In the exposed Galician coast in the Northern Spain, two well differentiated
ecotype Ecotypes are organisms which belong to the same species but possess different phenotypical features as a result of environmental factors such as elevation, climate and predation. Ecotypes can be seen in wide geographical distributions and may event ...
s are adapted to different shore levels and habitats. The RB ecotype (Ridged and Banded) lives on
barnacle Barnacles are arthropods of the subclass (taxonomy), subclass Cirripedia in the subphylum Crustacean, Crustacea. They are related to crabs and lobsters, with similar Nauplius (larva), nauplius larvae. Barnacles are exclusively marine invertebra ...
s in the upper shore. This ecotype displays a larger and more robust shell to resist the attack from predators such as
crab Crabs are decapod crustaceans of the infraorder Brachyura (meaning "short tailed" in Greek language, Greek), which typically have a very short projecting tail-like abdomen#Arthropoda, abdomen, usually hidden entirely under the Thorax (arthropo ...
s, and a smaller shell aperture in order to reduce the desiccation due to high sunshine exposure. The SU ecotype (Smooth and Unbanded) is found at the lower shore living on mussels. This ecotype shows a smaller and thinner shell with a wider shell aperture to allocate a relatively larger muscular foot providing a higher ability to avoid the dislodgment caused by the heavy wave action. Both ecotypes coexist in an intermediate habitat at the middle shore.


Genetics

''Littorina saxatilis'' has been shown to be an excellent model system for speciation genetics. In the seminal 2001 paper, Wilding ''et al'' demonstrated, using
amplified fragment length polymorphism Amplified fragment length polymorphism (AFLP-PCR or AFLP) is a PCR-based tool used in genetics research, DNA fingerprinting, and in the practice of genetic engineering. Developed in the early 1990s by Pieter Vos, AFLP uses restriction enzymes t ...
, that the low shore M form of the species were divergent from the high shore H form at number of loci despite gene flow between the forms.


Feeding habits

This snail is a
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
which grazes on the surface of rocks and mud flats.


Life cycle

The marine snail ''Littorina saxatilis'' has separate sexes,
internal fertilization Internal fertilization is the union of an egg and sperm cell during sexual reproduction inside the female body. Internal fertilization, unlike its counterpart, external fertilization, brings more control to the female with reproduction. For inte ...
, and a brood pouch with non-planktonic shelled embryos.


Synonyms

* ''Litorina groenlandica'' Menke, 1830 * ''Litorina incarnata'' Philippi, 1846 * ''Litorina marmorata'' L. Pfeiffer, 1839 * ''Litorina sulcata'' Menke, 1830 * ''Littorina castanea'' Deshayes in Deshayes & Milne Edwards, 1843 * ''Littorina danieli'' Locard, 1886 * ''Littorina groenlandica'' (Menke, 1830) * ''Littorina neglecta'' Bean, 1844 * ''Littorina nervillei'' Dautzenberg, 1893 * ''Littorina nervillei'' var. ''major'' Pallary in Seurat, 1924 * ''Littorina nigrolineata'' Gray, 1839 * ''Littorina palliata'' var. ''turritella'' Schlesch, 1916 * ''Littorina rudis'' (Maton, 1797) (synonym) * ''Littorina rudis'' f. ''elatior'' Middendorff, 1849 * ''Littorina rudis'' var. ''albida'' Dautzenberg, 1887 * ''Littorina rudis'' var. ''alticola'' Dacie, 1917 * ''Littorina rudis'' var. ''aurantia'' Dautzenberg, 1887 * ''Littorina rudis'' var. ''brevis'' Dautzenberg, 1887 *'' Littorina rudis'' var. ''conoidea'' Schlesch, 1916 * ''Littorina rudis'' var. ''fasciata'' Dautzenberg, 1887 * ''Littorina rudis'' var. ''finmarchia'' Herzenstein, 1885 * ''Littorina rudis'' var. ''globosa'' Jeffreys, 1865 * ''Littorina rudis'' var. ''globosa'' Martel, 1901 * ''Littorina rudis'' var. ''laevis'' Jeffreys, 1865 * ''Littorina rudis'' var. ''major'' Dautzenberg & P. Fisher, 1912 * ''Littorina rudis'' var. ''rubescens'' Monterosato, 1878 * ''Littorina rudis'' var. ''scotia'' S.M. Smith, 1979 * ''Littorina rudis'' var. ''similis'' Jeffreys, 1865 * ''Littorina rudis'' var. ''sulcata'' Martel, 1901 * ''Littorina rudis'' var. ''tenebrosapallida'' L.E. Adams, 1896 * ''Littorina rudis'' var. ''tessellata'' Dautzenberg, 1893 * ''Littorina saxatile'' La Roque, 1953 * ''Littorina saxatile saxatile'' La Roque, 1953 * ''Littorina saxatilis'' Johnston, 1842 * ''Littorina saxatilis'' f. ''abbreviata'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis'' f. ''conoidea'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis'' f. ''elongata'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis'' f. ''minor'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis groenlandica'' (Menke, 1830) * ''Littorina saxatilis groenlandica'' var. ''sculpta'' Schlesch, 1931 * ''Littorina saxatilis jugosa'' Montagu, 1803 * ''Littorina saxatilis jugosa'' var. ''bynei'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis jugosa'' var. ''tenuis'' James, 1968 * ''Littorina saxatilis nigrolineata'' Gray, 1839 * ''Littorina saxatilis rudis'' (Maton, 1797) * ''Littorina saxatilis rudis'' var. ''rudissimoides'' James, 1968 * ''Littorina saxatilis scotia'' Graham, 1988 * ''Littorina saxatilis tenebrosa'' ( Montagu, 1803) * ''Littorina saxatilis tenebrosa'' var. ''biinterrupta'' Fischer-Piette & Gaillard, 1963 * ''Littorina saxatilis tenebrosa'' var. ''bizonaria'' James, 1963 * ''Littorina saxatilis tenebrosa'' var. ''elata'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis tenebrosa'' var. ''maculata'' Fischer-Piette & Gaillard, 1963 * ''Littorina saxatilis'' var. ''clarilineata'' Fischer-Piette & Gaillard, 1971 * ''Littorina saxatilis'' var. ''flammulata'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis'' var. ''fulva'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis'' var. ''fusca'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis'' var. ''gascae'' Fischer-Piette & Gaillard, 1971 * ''Littorina saxatilis'' var. ''groenlandica'' (Menke, 1830) * ''Littorina saxatilis'' var. ''hieroglyphica'' Fischer-Piette, Gaillard & Jouin, 1961 * ''Littorina saxatilis'' var. ''interrupta'' Fischer-Piette, Gaillard & Jouin, 1961 * ''Littorina saxatilis'' var. ''lagunae'' Barnes, 1993 * ''Littorina saxatilis'' var. ''lineata'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis'' var. ''lugubris'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis'' var. ''nigra'' Fischer-Piette & Gaillard, 1971 * ''Littorina saxatilis'' var. ''nojensis'' Fischer-Piette & Gaillard, 1964 * ''Littorina saxatilis'' var. ''rubra'' Fischer-Piette & Gaillard, 1971 * ''Littorina saxatilis'' var. ''rubrolineata'' Fischer-Piette, Gaillard & Delmas, 1967 * ''Littorina saxatilis'' var. ''salvati'' Fischer-Piette, Gaillard & Delmas, 1967 * ''Littorina saxatilis'' var. ''sanguinea'' Coen, 1933 * ''Littorina saxatilis'' var. ''sellensis'' Fischer-Piette & Gaillard, 1964 * ''Littorina saxatilis'' var. ''tractibus'' Fischer-Piette, Gaillard & Jouin, 1961 * ''Littorina saxatilis'' var. ''trifasciata'' Dautzenberg & P. Fisher, 1912 * ''Littorina saxatilis zonata'' Daniel, 1883 * ''Littorina saxoides'' Nardo, 1847 * ''Littorina simplex'' Reeve, 1857 * ''Littorina tenebrosa'' ( Montagu, 1803) * ''Littorina tenebrosa'' f. ''elatior'' Middendorff, 1849 * ''Littorina tenebrosa'' var. ''costulata'' Middendorff, 1849 * ''Littorina tenebrosa'' var. ''densecostulata'' Middendorff, 1849 * ''Littorina tenebrosa'' var. ''grisolacteus'' Middendorff, 1849 * ''Littorina tenebrosa'' var. ''intermedia'' Forbes & Hanley, 1850 * ''Littorina tenebrosa'' var. ''rubidus'' Middendorff, 1849 * ''Littorina tenebrosa'' var. ''tessellatus'' Middendorff, 1849 * ''Littorina tenebrosa'' var. ''zonatus'' Middendorff, 1849 * ''Littorina zonaria'' Bean, 1844 * ''Nerita rustica'' Nardo, 1847 * ''Turbo obligatus'' Say, 1822 * ''Turbo rudis'' Maton, 1797 * ''Turbo rudissimus'' Johnston, 1842


See also

*
Common periwinkle 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. This is a ...


References

This article incorporates CC-BY-2.0 text from the reference Martínez-Fernández M., Bernatchez L., Rolán-Alvarez E. & Quesada H. (2010). "Insights into the role of differential gene expression on the ecological adaptation of the snail ''Littorina saxatilis''". ''
BMC Evolutionary Biology ''BMC Ecology and Evolution'' (since January 2021), previously ''BMC Evolutionary Biology'' (2001–2020), is a peer-reviewed open access scientific journal covering all fields of evolutionary biology, including phylogenetics and palaeontology ...
'' 10: 356. .
Wilding C. S., Butlin R. K. & Grahame J. (2001). "Differential gene expression between parapatric morphs of ''Littorina saxatilis'' detected using RFLP markers". ''
Journal of Evolutionary Biology The ''Journal of Evolutionary Biology'' is a peer-reviewed scientific journal published monthly covering the field of evolutionary biology. It is owned by the European Society for Evolutionary Biology. The founding editor-in-chief was Stephen C. ...
'' 14: 4. .


Further reading

* Carvajal-Rodríguez, A.; Conde-Padín, P. & Rolán-Alvarez, E. (2005). "Decomposing shell form into size and shape by geometric morphometric methods in two sympatric ecotypes of ''Littorina saxatilis''. ''
Journal of Molluscan Studies The ''Journal of Molluscan Studies'' is the peer-reviewed scientific journal of the Malacological Society of London, covering research in malacology.
'' 71: 313–318. * Galindo, J.; Morán, P. & Rolán-Alvarez, E. (2009). "Comparing geographical genetic differentiation between candidate and noncandidate loci for adaptation strengthens support for parallel ecological divergence in the marine snail ''Littorina saxatilis''". ''Molecular Ecology'' 18: 919–930. * Hayward, P. J.; Ryland, J. S. (Ed.) (1990). ''The marine fauna of the British Isles and North-West Europe: 1. Introduction and protozoans to arthropods''. Clarendon Press: Oxford, UK. . 627 pp. * Johannesson, B. (1986). "Shell morphology of ''Littorina saxatilis'' Olivi the relative importance of physical factors and predation". ''Journal of Experimental Marine Biology and Ecology'' 102: 183–195. * Johannesson, K.; Johannesson, B. & Rolán-Alvarez, E. (1993). "Morphological differentiation and genetic cohesiveness over a micro-environmental gradient in the marine snail ''Littorina saxatilis''". ''Evolution'' 47: 1770–1787. * Reid, D. G. (1989a) "The comparative morphology, phylogeny and evolution of the gastropod family Littorinidae". ''Philosophical Transactions of the Royal Society of London, Series B'' 324: 1–110 * Reid, D. G. (1996). ''Systematics and evolution of Littorina''. The Ray Society 463p. * Rolán-Alvarez, E. (2007). "Sympatric speciation as a by-product of ecological adaptation in the Galicia ''Littorina saxatilis'' hybrid zone". ''
Journal of Molluscan Studies The ''Journal of Molluscan Studies'' is the peer-reviewed scientific journal of the Malacological Society of London, covering research in malacology.
'' 73: 1–10. * Rolán-Alvarez, E.; Johannesson, K. & Erlandsson, J. (1997). "The maintenance of a cline in the marine snail ''Littorina saxatilis'': the role of home site advantage and hybrid fitness in ecotype formation". ''Evolution'' 51: 1838–1847.


External links


In the British Isles


{{DEFAULTSORT:Littorina Saxatilis Littorinidae Gastropods described in 1792 Taxa named by Giuseppe Olivi