Cepaea nemoralis: Difference between revisions

Lemon or Grove snail; Cepaea nemoralis
	Cepaea nemoralis
Conservation status
	; Least Concern (IUCN 3.1)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Mollusca
Class:	Gastropoda
Order:	Stylommatophora
Family:	Helicidae
Genus:	Cepaea
Species:	C. nemoralis
Binomial name
	Cepaea nemoralis; (Linnaeus, 1758)
Synonyms
	Cepaea (Cepaea) nemoralis (Linnaeus, 1758); Helix nemoralis Linnaeus, 1758;

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The grove snail, brown-lipped snail or lemon snail (Cepaea nemoralis) is a species of air-breathing land snail, a terrestrial pulmonate gastropod mollusc.^[3]

It is one of the most common large species of land snail in Europe, and has been introduced to North America.

Subspecies

Cepaea nemoralis etrusca (Rossmässler, 1835)^[4]
Cepaea nemoralis nemoralis (Linnaeus, 1758)

Cepaea nemoralis is the type species of the genus Cepaea.^[5] It is used as a model organism in ecological genetics, including in citizen science projects.^[6]^[7]

Description

Cepaea nemoralis is among the largest and, because of its bright colouration, one of the best-known snails in Western Europe.^[6] The colour of the shell is highly variable; it ranges from brown, through pink, to yellow or even whitish, with or without one to five dark-brown bands.^[8] Names for many colour variants were coined in the nineteenth century but this system has been replaced by an independent scoring of shell colour and the presence/absence and fusion of individual bands numbered 1 to 5.^[9]

The thickened and slightly out-turned apertural lip of adults is usually dark brown, but can be white in some regions.^[10] The umbilicus is closed in adults but narrowly open in juveniles.^[8] The shell surface is semi-glossy. An adult shell consists of 4½–5½ whorls, with a width of 18–25 mm and a height of 12–22 mm.^[8]

Identification

Cepaea nemoralis is closely related to Cepaea hortensis. They share much the same habitat and exhibit a similar range of shell colours and banding patterns. Cepaea nemoralis tends to grow larger, but usually the species can most easily be recognised by the colour of the lip of adult shells. In a high proportion of regions, C. nemoralis consistently has a dark-brown lip to its shell, whilst C. hortensis has a white lip.^[10]

Love darts of Cepaea, cross sections on left^[11]

C. nemoralis

C. hortensis

In areas where lip colour is variable, dissection is necessary. A cross-section of the love dart of C. nemoralis shows a cross with simple blades, whereas that of C. hortensis has bifurcated blades. The mucous gland has 3 or fewer branches in C. nemoralis, but 4 or more in C. hortensis.^[12]

Two superficially similar species Caucasotachea vindobonensis and Macularia sylvatica both have a lip that is brown near the columella becoming pale towards the suture, and they have fine growth ridges on the shell whereas in both Cepaea species it is smooth. Also, M. sylvatica is distinct in having a small blunt tooth in its aperture, whilst the lowest brown band on the shell of C. vindobonensis lies noticeably closer to the columella than in Cepaea.^[13]

Colour polymorphism

Cepaea nemoralis is highly polymorphic in shell colour and banding. The background colour of the shell varies along a continuum from brown through pink to yellow and sometimes almost white.^[14] Additionally the shells can be with or without dark bands. The bands vary in intensity of colour, in width and in number, from zero to five. The genetics underlying this variation is extensively understood and is shared with C. hortensis.^[10]

The polymorphism has also been intensely studied for its evolution and ecology. For instance, in stable habitats shells tend to be darker in woodland than in open habitats.^[15] The explanation might be camouflage or climatic selection: paler, more reflective colours in sunny environments reduce water loss and overheating. Climatic selection can also explain why yellow shells are more common in the south.^[6]

Another question is why the variation persists, usually even within a locality. Researchers have variously argued that the cause is random genetic drift and founder effects, different selection pressures in different areas with mixing by migration, and balanced polymorphism. Balanced polymorphism could arise when a predator like the song thrush develops a 'search image' for the commonest morph, so that the rarer morphs are less likely to be predated. Natural selection would then favour a diversity of colours and patterns as an antipredator adaptation. Most probably, the polymorphism has multiple causes.^[10]^[16]

Different coloration and banding of the shells of Cepaea nemoralis

A diversity of forms
Yellow unbanded
Yellow with several bands
Yellow mid-banded
Reddish-brown unbanded

Distribution

Snail traversing a leaf.

The native distribution of C. nemoralis is from Western and Northern Europe to Central Europe, but it has been spreading eastwards especially over the last few decades.^[17]^[18] Thus it is known from most of the Iberian Peninsula,^[19] France, Great Britain, Ireland, Belgium, the Netherlands, Germany, Denmark, southern Sweden and Norway, Switzerland, Austria, the Czech Republic,^[20]^[21] Hungary, Bosnia, Croatia, and the northern half of Italy.^[8]^[22] In Central and Eastern Europe it has spread particularly along the Baltic coast (e.g. in Poland,^[18] Latvia, Estonia, southern Finland,^[23] the east coast of Sweden)^[24] but also now elsewhere in Poland,^[18] and in Ukraine, Belarus and Russia.^[25] In Slovakia it was still known only from a single garden centre by 2020.^[26]

At the northern edge of its range,C. nemoralis is rare and scattered in northern Scotland, where it has been introduced.^[8] It is not found in the Hebrides, Orkney or Shetland.^[8] It seems to have been affected by air pollution and soil acidification in some parts of England.^[8]

Starting in 1857, there have been multiple introductions to North America, where it now occurs widely in Canada (from British Columbia to Newfoundland; in Ontario it is a pest of vineyards) and in the north-eastern part of the US, with further occurrences further south (e.g. California, Colorado, Texas and South Carolina).^[27]^[28]^[29]

Both in America and Eastern Europe it is known that some introductions have been deliberate.^[28]^[25]

The range of C. hortensis mostly overlaps that of C. nemoralis but extends further north and less far south.^[12]

Biology and ecology

This is a very common and widespread species in Western Europe, occupying a wide range of habitats from coastal dunes to woodlands with full canopy cover, including gardens and abandoned land.^[6] In Eastern Europe, where it is a new arrival, it has turned up particularly in urban areas and other disturbed habitats.^[18] It can be found up to an altitude of 1600 m in the Alps,^[30] 1800 m in the Pyrenees, 900 m in Wales, 600 m in Scotland.^[8] Density of adults is often of the order of 2 per square metre.^[31]

This species feeds mainly on dead or senescent plants.^[6]^[8] It prefers broad-leaved plants over grasses.^[32] Although mostly not a pest of crops,^[8] it can be a nuisance in vineyards because it is inadvertently picked with the grapes.^[33]

Like all pulmonate land snails, it is a hermaphrodite, and this species must mate to produce fertile eggs.^[6] Mating tends to be concentrated in late spring and early summer, though it can continue through the autumn.^[6] The snails may store the sperm they receive from their partner for some time, and individual broods can have mixed paternity.^[6] In Britain it lays clutches of 30–50 (in France 40–80) oval eggs between June and August (in France May–October, in W France until November).^[8] The size of the egg is 3.1 × 2.6 mm^[34] or egg diameter can be 2.3–3.0 mm.^[8] Juveniles hatch after 15–20 days.^[8]

This snail is comparatively slow-growing, taking 1 to 3 years to develop from an egg to a breeding adult.^[35] The life-span for this species is up to seven or eight years, with the annual survival rate of adults about 50% (= 3% over five years, older adults suffer higher mortalities).^[8] In winter, the snails may hibernate, but can become active again during warm spells.^[6]

Cepaea nemoralis is known experimentally to be a host for Angiostrongylus vasorum.^[36] Predators of Cepaea nemoralis include the song thrush (Turdus philomelos).

References

This article includes public domain text from the reference^[8] and CC-BY-2.5 text from the reference^[6]

^ 2013 IUCN Red List of Threatened Species. <www.iucnredlist.org>. Cited 28 December 2013.
^ Linnaeus C. (1758). Systema naturae per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. pp. [1-4], 1-824. Holmiae. (Salvius).
^ "Cepaea nemoralis (Linnaeus, 1758)". MolluscaBase. Flanders Marine Institute. Retrieved 23 April 2023.
^ "Cepaea nemoralis etrusca (Rossmässler, 1835)". MolluscaBase. Flanders Marine Institute. Retrieved 23 April 2023.
^ "Cepaea Held, 1838". MolluscaBase. Flanders Marine Institute. Retrieved 23 April 2023.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Silvertown, J.; Cook, L.; Cameron, R.; Dodd, M.; McConway, K.; Worthington, J.; Skelton, P.; Anton, C.; Bossdorf, O.; Baur, B.; Schilthuizen, M.; Fontaine, B.; Sattmann, H.; Bertorelle, G.; Correia, M.; Oliveira, C.; Pokryszko, B.; Ożgo, M.; Stalažs, A.; Gill, E.; Rammul, Ü.; Sólymos, P.; Féher, Z.; Juan, X. (27 April 2011). "Citizen science reveals unexpected continental-scale evolutionary change in a model organism". PLOS ONE. 6 (4): e18927. doi:10.1371/journal.pone.0018927. PMC 3083392. PMID 21556137.
^ Kerstes, N.A.G.; Breeschoten, T.; Kalkman, V.J.; Schilthuizen, M. (19 July 2019). "Snail shell colour evolution in urban heat islands detected via citizen science". Communications Biology. 2 (1): 264. doi:10.1038/s42003-019-0511-6. PMC 6642149. PMID 31341963.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o "Species summary for Cepaea nemoralis". AnimalBase, last modified 6 February 2011, accessed 1 May 2011.
^ Cain, A.J. (1988). "The scoring of polymorphic colour and pattern variation and its genetic basis in molluscan shells". Malacologia. 28: 1–15.
^ ^a ^b ^c ^d Jones, J.S.; Leith, B.H.; Rawlings, P. (1977). "Polymorphism in Cepaea: a problem with too many solutions?". Annual Review of Ecology and Systematics. 8 (1): 109–143. doi:10.1146/annurev.es.08.110177.000545.
^ Koene, J.M.; Schulenburg, H. (30 March 2005). "Shooting darts: co-evolution and counter-adaptation in hermaphroditic snails". BMC Evolutionary Biology. 5 (1): 25. doi:10.1186/1471-2148-5-25. PMC 1080126. PMID 15799778.
^ ^a ^b Welter-Schultes, F.W. (2012). European non-marine molluscs, a guide for species identification = Bestimmungsbuch für europäische Land- und Süsswassermollusken. Göttingen: Planet Poster Editions. ISBN 9783933922755.
^ Kerney, M.P.; Cameron, R.A.D. (1979). A field guide to the land snails of Britain and north-west Europe. London: Collins. ISBN 000219676X.
^ Davison, A.; Jackson, H.J.; Murphy, E.W.; Reader, T. (August 2019). "Discrete or indiscrete? Redefining the colour polymorphism of the land snail Cepaea nemoralis". Heredity. 123 (2): 162–175. doi:10.1038/s41437-019-0189-z. PMC 6629550. PMID 30804571.
^ Cameron, R.A.D.; Cook, L.M. (May 2012). "Habitat and the shell polymorphism of Cepaea nemoralis (L.): interrogating the Evolution Megalab database". Journal of Molluscan Studies. 78 (2): 179–184. doi:10.1093/mollus/eyr052.
^ Cook, L.M. (29 October 1998). "A two–stage model for Cepaea polymorphism". Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences. 353 (1375): 1577–1593. doi:10.1098/rstb.1998.0311. PMC 1692378.
^ Dvořák L., Honěk A. & Martínková Z. (2003). "The spread of Cepaea nemoralis (L.) populations in the Czech Republic". 2003 BCPC Symposiumproceedings No. 80: Slugs & snails: agricultural, veterinary & environmental perspectives: 99-102.
^ ^a ^b ^c ^d Ożgo, M.; Cameron, R.A.D.; Horsák, M.; Pokryszko, B.; Chudaś, M.; Cichy, A.; Kaczmarek, S.; Kobak, J.; Marzec, M.; Mierzwa-Szymkowiak, D.; Parzonko, D.; Pyka, G.; Rosin, Z.; Skawina, A.; Soroka, M.; Sulikowska-Drozd, A.; Surowiec, T.; Szymanek, M.; Templin, J.; Urbańska, M.; Zając, K.; Zielska, J.; Żbikowska, E.; Żołądek, J. (17 April 2019). "Cepaea nemoralis (Gastropoda: Pulmonata) in Poland: patterns of variation in a range-expanding species". Biological Journal of the Linnean Society. 127 (1): 1–11. doi:10.1093/biolinnean/blz029.
^ Cadevall, J.; Orozco, A. (2016). Caracoles y babosas de la Península Ibérica y Baleares. Barcelona: Omega. ISBN 9788428215992.
^ Juřičková, L.; Horsák, M.; Beran, L. (2001). "Check-list of the molluscs (Mollusca) of the Czech Republic". Acta Soc. Zool. Bohem. 65: 25–40.
^ Peltanová, Alena; Dvořák, Libor; Juřičková, Lucie (1 April 2012). "The spread of non-native Cepaea nemoralis and Monacha cartusiana (Gastropoda: Pulmonata) in the Czech Republic with comments on other land snail immigrants". Biologia. 67 (2): 384–389. doi:10.2478/s11756-012-0020-2.
^ Ramos‐Gonzalez, D.; Saenko, S.V.; Davison, A. (August 2022). "Deep structure, long‐distance migration and admixture in the colour polymorphic land snail Cepaea nemoralis". Journal of Evolutionary Biology. 35 (8): 1110–1125. doi:10.1111/jeb.14060. PMC 9541890. PMID 35830483.
^ Koivunen, A.; Malinen, P.; Ormio, H.; Terhivuo, J.; Valovirta, I. (2014). Suomen kotilot ja etanat : opas maanilviäisten maailmaan. Helsinki: Tibiale. ISBN 9789526754468.
^ Ożgo, M.; Örstan, A.; Kirschenstein, M.; Cameron, R. (May 2016). "Dispersal of land snails by sea storms". Journal of Molluscan Studies. 82 (2): 341–343. doi:10.1093/mollus/eyv060.
^ ^a ^b Gural-Sverlova, N.; Egorov, R.; Kruglova, O.; Kovalevich, N.; Gural, R. (27 September 2021). "Introduced land snail Cepaea nemoralis (Gastropoda: Helicidae) in Eastern Europe: spreading history and the shell colouration variability". Malacologica Bohemoslovaca. 20: 75–91. doi:10.5817/MaB2021-20-75.
^ Čejka, T. (3 August 2020). "The terrestrial molluscan fauna in the Slovak part of the Danubian Lowland: an annotated checklist". Authorea. doi:10.22541/au.159620996.68925423/v2. S2CID 242206099.
^ Layton, K.K.S.; Warne, C.P.K.; Nicolai, A.; Ansart, A.; deWaard, J.R. (April 2019). "Molecular evidence for multiple introductions of the banded grove snail (Cepaea nemoralis) in North America" (PDF). Canadian Journal of Zoology. 97 (4): 392–398. doi:10.1139/cjz-2018-0084. S2CID 91897997.
^ ^a ^b Whitson, M. (September 2005). "Cepaea nemoralis (Gastropoda, Helicidae): The invited invader". Journal of the Kentucky Academy of Science. 66 (2): 82–88. doi:10.3101/1098-7096(2006)66[82:CNGHTI]2.0.CO;2. S2CID 86005518.
^ Burke, T.E. (2013). Land snails and slugs of the Pacific Northwest. Corvallis, Oregon: Oregon State University Press. ISBN 978-0-87071-685-0.
^ Turner, H.; Kuiper, J.G.J.; Thew, N.; Bernasconi, R.; Rüetschi, J.; Wüthrich, M.; Gosteli, M. (1998). Atlas der Mollusken der Schweiz und Liechtensteins. Neuchâtel: CentreSuisse de cartographie de la faune. ISBN 2884140131.
^ Williamson, P.; Cameron, R.A.D.; Carter, M.A. (February 1977). "Population dynamics of the landsnail Cepaea nemoralis L.: a six-year study". The Journal of Animal Ecology. 46 (1): 181. doi:10.2307/3955. JSTOR 3955.
^ Chang, H.-W.; Emlen, John M. (1 April 1993). "Seasonal variation of microhabitat distribution of the polymorphic land snail Cepaea nemoralis". Oecologia. 93 (4): 501–507. doi:10.1007/BF00328957. PMID 28313817. S2CID 22594544.
^ Giffard, B.; Winter, S.; Guidoni, S.; Nicolai, A.; Castaldini, M.; Cluzeau, D.; Coll, P.; Cortet, J.; Le Cadre, E.; d’Errico, G.; Forneck, A.; Gagnarli, E.; Griesser, M.; Guernion, M.; Lagomarsino, A.; Landi, S.; Le Bissonnais, Y.; Mania, E.; Mocali, S.; Preda, C.; Priori, S.; Reineke, A; Rusch, A.; Schroers, H.-J.; Simoni, S.; Steiner, M.; Temneanu, E.; Bacher, S.; Costantini, E.A.C.; Zaller, J.; Leyer, I. (2022). "Vineyard management and its impacts on soil biodiversity, functions, and ecosystem services". Frontiers in Ecology and Evolution. 10. doi:10.3389/fevo.2022.850272.
^ Heller J. (2001). Life History Strategies. In: Barker G. M. (ed.). The biology of terrestrial molluscs. CABI Publishing, Oxon, UK, ISBN 0-85199-318-4. 1-146, cited page: 428.
^ Oosterhoff, L. M. (1 January 1976). "Variation in growth rate as an ecological factor in the landsnail Cepaea nemoralis (L.)". Netherlands Journal of Zoology. 27 (1): 1–132. doi:10.1163/002829677X00072. S2CID 55537228.
^ Conboy G.A. (30 May 2000). "Canine Angiostrongylosis (French Heartworm)". In: Bowman D. D. (Ed.) Companion and Exotic Animal Parasitology. International Veterinary Information Service. Accessed 24 November 2009.

External links

Cepaea nemoralis at Animalbase taxonomy,short description, distribution, biology,status (threats), images
Cepaea nemoralis images at Encyclopedia of Life
Evolution MegaLab was an online survey of Cepaea polymorphism (no longer online) Archived 2019-07-06 at the Wayback Machine
Life & Environment, University of the West of Scotland

[1] 2013 IUCN Red List of Threatened Species. <www.iucnredlist.org>. Cited 28 December 2013.

[2] Linnaeus C. (1758). Systema naturae per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. pp. [1-4], 1-824. Holmiae. (Salvius).

[3] "Cepaea nemoralis (Linnaeus, 1758)". MolluscaBase. Flanders Marine Institute. Retrieved 23 April 2023.

[4] "Cepaea nemoralis etrusca (Rossmässler, 1835)". MolluscaBase. Flanders Marine Institute. Retrieved 23 April 2023.

[5] "Cepaea Held, 1838". MolluscaBase. Flanders Marine Institute. Retrieved 23 April 2023.

[Silvertown2011-6] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Silvertown, J.; Cook, L.; Cameron, R.; Dodd, M.; McConway, K.; Worthington, J.; Skelton, P.; Anton, C.; Bossdorf, O.; Baur, B.; Schilthuizen, M.; Fontaine, B.; Sattmann, H.; Bertorelle, G.; Correia, M.; Oliveira, C.; Pokryszko, B.; Ożgo, M.; Stalažs, A.; Gill, E.; Rammul, Ü.; Sólymos, P.; Féher, Z.; Juan, X. (27 April 2011). "Citizen science reveals unexpected continental-scale evolutionary change in a model organism". PLOS ONE. 6 (4): e18927. doi:10.1371/journal.pone.0018927. PMC 3083392. PMID 21556137.

[7] Kerstes, N.A.G.; Breeschoten, T.; Kalkman, V.J.; Schilthuizen, M. (19 July 2019). "Snail shell colour evolution in urban heat islands detected via citizen science". Communications Biology. 2 (1): 264. doi:10.1038/s42003-019-0511-6. PMC 6642149. PMID 31341963.

[AB-8] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o "Species summary for Cepaea nemoralis". AnimalBase, last modified 6 February 2011, accessed 1 May 2011.

[9] Cain, A.J. (1988). "The scoring of polymorphic colour and pattern variation and its genetic basis in molluscan shells". Malacologia. 28: 1–15.

[Jones1977-10] Jones, J.S.; Leith, B.H.; Rawlings, P. (1977). "Polymorphism in Cepaea: a problem with too many solutions?". Annual Review of Ecology and Systematics. 8 (1): 109–143. doi:10.1146/annurev.es.08.110177.000545.

[11] Koene, J.M.; Schulenburg, H. (30 March 2005). "Shooting darts: co-evolution and counter-adaptation in hermaphroditic snails". BMC Evolutionary Biology. 5 (1): 25. doi:10.1186/1471-2148-5-25. PMC 1080126. PMID 15799778.

[Welter-Schultes2012-12] Welter-Schultes, F.W. (2012). European non-marine molluscs, a guide for species identification = Bestimmungsbuch für europäische Land- und Süsswassermollusken. Göttingen: Planet Poster Editions. ISBN 9783933922755.

[Kerney1979-13] Kerney, M.P.; Cameron, R.A.D. (1979). A field guide to the land snails of Britain and north-west Europe. London: Collins. ISBN 000219676X.

[Davison2019-14] Davison, A.; Jackson, H.J.; Murphy, E.W.; Reader, T. (August 2019). "Discrete or indiscrete? Redefining the colour polymorphism of the land snail Cepaea nemoralis". Heredity. 123 (2): 162–175. doi:10.1038/s41437-019-0189-z. PMC 6629550. PMID 30804571.

[Cameron2012-15] Cameron, R.A.D.; Cook, L.M. (May 2012). "Habitat and the shell polymorphism of Cepaea nemoralis (L.): interrogating the Evolution Megalab database". Journal of Molluscan Studies. 78 (2): 179–184. doi:10.1093/mollus/eyr052.

[Cook1998-16] Cook, L.M. (29 October 1998). "A two–stage model for Cepaea polymorphism". Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences. 353 (1375): 1577–1593. doi:10.1098/rstb.1998.0311. PMC 1692378.

[17] Dvořák L., Honěk A. & Martínková Z. (2003). "The spread of Cepaea nemoralis (L.) populations in the Czech Republic". 2003 BCPC Symposiumproceedings No. 80: Slugs & snails: agricultural, veterinary & environmental perspectives: 99-102.

[Ozgo2019-18] Ożgo, M.; Cameron, R.A.D.; Horsák, M.; Pokryszko, B.; Chudaś, M.; Cichy, A.; Kaczmarek, S.; Kobak, J.; Marzec, M.; Mierzwa-Szymkowiak, D.; Parzonko, D.; Pyka, G.; Rosin, Z.; Skawina, A.; Soroka, M.; Sulikowska-Drozd, A.; Surowiec, T.; Szymanek, M.; Templin, J.; Urbańska, M.; Zając, K.; Zielska, J.; Żbikowska, E.; Żołądek, J. (17 April 2019). "Cepaea nemoralis (Gastropoda: Pulmonata) in Poland: patterns of variation in a range-expanding species". Biological Journal of the Linnean Society. 127 (1): 1–11. doi:10.1093/biolinnean/blz029.

[19] Cadevall, J.; Orozco, A. (2016). Caracoles y babosas de la Península Ibérica y Baleares. Barcelona: Omega. ISBN 9788428215992.

[20] Juřičková, L.; Horsák, M.; Beran, L. (2001). "Check-list of the molluscs (Mollusca) of the Czech Republic". Acta Soc. Zool. Bohem. 65: 25–40.

[21] Peltanová, Alena; Dvořák, Libor; Juřičková, Lucie (1 April 2012). "The spread of non-native Cepaea nemoralis and Monacha cartusiana (Gastropoda: Pulmonata) in the Czech Republic with comments on other land snail immigrants". Biologia. 67 (2): 384–389. doi:10.2478/s11756-012-0020-2.

[22] Ramos‐Gonzalez, D.; Saenko, S.V.; Davison, A. (August 2022). "Deep structure, long‐distance migration and admixture in the colour polymorphic land snail Cepaea nemoralis". Journal of Evolutionary Biology. 35 (8): 1110–1125. doi:10.1111/jeb.14060. PMC 9541890. PMID 35830483.

[23] Koivunen, A.; Malinen, P.; Ormio, H.; Terhivuo, J.; Valovirta, I. (2014). Suomen kotilot ja etanat : opas maanilviäisten maailmaan. Helsinki: Tibiale. ISBN 9789526754468.

[24] Ożgo, M.; Örstan, A.; Kirschenstein, M.; Cameron, R. (May 2016). "Dispersal of land snails by sea storms". Journal of Molluscan Studies. 82 (2): 341–343. doi:10.1093/mollus/eyv060.

[Gural-Sverlova2021-25] Gural-Sverlova, N.; Egorov, R.; Kruglova, O.; Kovalevich, N.; Gural, R. (27 September 2021). "Introduced land snail Cepaea nemoralis (Gastropoda: Helicidae) in Eastern Europe: spreading history and the shell colouration variability". Malacologica Bohemoslovaca. 20: 75–91. doi:10.5817/MaB2021-20-75.

[26] Čejka, T. (3 August 2020). "The terrestrial molluscan fauna in the Slovak part of the Danubian Lowland: an annotated checklist". Authorea. doi:10.22541/au.159620996.68925423/v2. S2CID 242206099.

[27] Layton, K.K.S.; Warne, C.P.K.; Nicolai, A.; Ansart, A.; deWaard, J.R. (April 2019). "Molecular evidence for multiple introductions of the banded grove snail (Cepaea nemoralis) in North America" (PDF). Canadian Journal of Zoology. 97 (4): 392–398. doi:10.1139/cjz-2018-0084. S2CID 91897997.

[Whitson2005-28] Whitson, M. (September 2005). "Cepaea nemoralis (Gastropoda, Helicidae): The invited invader". Journal of the Kentucky Academy of Science. 66 (2): 82–88. doi:10.3101/1098-7096(2006)66[82:CNGHTI]2.0.CO;2. S2CID 86005518.

[29] Burke, T.E. (2013). Land snails and slugs of the Pacific Northwest. Corvallis, Oregon: Oregon State University Press. ISBN 978-0-87071-685-0.

[30] Turner, H.; Kuiper, J.G.J.; Thew, N.; Bernasconi, R.; Rüetschi, J.; Wüthrich, M.; Gosteli, M. (1998). Atlas der Mollusken der Schweiz und Liechtensteins. Neuchâtel: CentreSuisse de cartographie de la faune. ISBN 2884140131.

[Williamson1977-31] Williamson, P.; Cameron, R.A.D.; Carter, M.A. (February 1977). "Population dynamics of the landsnail Cepaea nemoralis L.: a six-year study". The Journal of Animal Ecology. 46 (1): 181. doi:10.2307/3955. JSTOR 3955.

[32] Chang, H.-W.; Emlen, John M. (1 April 1993). "Seasonal variation of microhabitat distribution of the polymorphic land snail Cepaea nemoralis". Oecologia. 93 (4): 501–507. doi:10.1007/BF00328957. PMID 28313817. S2CID 22594544.

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[Barker-34] Heller J. (2001). Life History Strategies. In: Barker G. M. (ed.). The biology of terrestrial molluscs. CABI Publishing, Oxon, UK, ISBN 0-85199-318-4. 1-146, cited page: 428.

[35] Oosterhoff, L. M. (1 January 1976). "Variation in growth rate as an ecological factor in the landsnail Cepaea nemoralis (L.)". Netherlands Journal of Zoology. 27 (1): 1–132. doi:10.1163/002829677X00072. S2CID 55537228.

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@@ Line 16: / Line 16: @@
 The '''grove snail''', '''brown-lipped snail''' or '''lemon snail''' ('''''Cepaea nemoralis''''') is a [[species]] of air-breathing [[land snail]], a [[terrestrial molluscs|terrestrial]] [[pulmonate]] [[gastropod]] [[mollusc]].<ref>{{cite web |title=''Cepaea nemoralis'' (Linnaeus, 1758) |url=https://molluscabase.org/aphia.php?p=taxdetails&id=235792 |website=MolluscaBase |publisher=Flanders Marine Institute |access-date=23 April 2023}}</ref>
-It is one of the commonest large species of land snail in [[Europe]], and has been introduced to [[North America]].
+It is one of the most common large species of land snail in [[Europe]], and has been introduced to [[North America]].
 ;Subspecies:
@@ Line 22: / Line 22: @@
 * ''Cepaea nemoralis nemoralis'' <small>(Linnaeus, 1758)</small>
-''Cepaea nemoralis'' is the type species of the genus ''Cepaea''.<ref>{{cite web |title=''Cepaea'' Held, 1838 |url=https://molluscabase.org/aphia.php?p=taxdetails&id=235791 |website=MolluscaBase |publisher=Flanders Marine Institute |access-date=23 April 2023}}</ref> It is used as a [[model organism]] in [[ecological genetics]], including in [[citizen science]] projects.<ref name="Silvertown2011"/><ref>{{cite journal |last1=Kerstes |first1=N.A.G. |last2=Breeschoten |first2=T. |last3=Kalkman |first3=V.J. |last4=Schilthuizen |first4=M. |title=Snail shell colour evolution in urban heat islands detected via citizen science |journal=Communications Biology |date=19 July 2019 |volume=2 |issue=1 |pages=264 |doi=10.1038/s42003-019-0511-6}}</ref>
+''Cepaea nemoralis'' is the type species of the genus ''Cepaea''.<ref>{{cite web |title=''Cepaea'' Held, 1838 |url=https://molluscabase.org/aphia.php?p=taxdetails&id=235791 |website=MolluscaBase |publisher=Flanders Marine Institute |access-date=23 April 2023}}</ref> It is used as a [[model organism]] in [[ecological genetics]], including in [[citizen science]] projects.<ref name="Silvertown2011"/><ref>{{cite journal |last1=Kerstes |first1=N.A.G. |last2=Breeschoten |first2=T. |last3=Kalkman |first3=V.J. |last4=Schilthuizen |first4=M. |title=Snail shell colour evolution in urban heat islands detected via citizen science |journal=Communications Biology |date=19 July 2019 |volume=2 |issue=1 |pages=264 |doi=10.1038/s42003-019-0511-6|pmid=31341963 |pmc=6642149 }}</ref>
 ==Description==
@@ Line 38: / Line 38: @@
  | direction = vertical
  | width = 200
- | header = Love darts of ''Cepaea'', cross sections on left<ref>{{cite journal |last1=Koene |first1=J.M. |last2=Schulenburg |first2=H. |title=Shooting darts: co-evolution and counter-adaptation in hermaphroditic snails |journal=BMC Evolutionary Biology |date=30 March 2005 |volume=5 |issue=1 |pages=25 |doi=10.1186/1471-2148-5-25}}</ref>
+ | header = Love darts of ''Cepaea'', cross sections on left<ref>{{cite journal |last1=Koene |first1=J.M. |last2=Schulenburg |first2=H. |title=Shooting darts: co-evolution and counter-adaptation in hermaphroditic snails |journal=BMC Evolutionary Biology |date=30 March 2005 |volume=5 |issue=1 |pages=25 |doi=10.1186/1471-2148-5-25|pmid=15799778 |pmc=1080126 |doi-access=free }}</ref>
  | image1 = Cepaea nemoralis dart.jpg
  | alt1 = sketch of dart
@@ Line 47: / Line 47: @@
  | caption2 = ''C. hortensis''
-}}In areas where lip colour is variable, dissection is necessary. A cross-section of the [[love dart]] of ''C. nemoralis'' shows a cross with simple blades, whereas that of ''C. hortensis'' has bifurcated blades. The mucus gland has 3 or fewer branches in ''C. nemoralis'', but 4 or more in ''C. hortensis''.<ref name=Welter-Schultes2012>{{cite book |last1=Welter-Schultes |first1=F.W. |title=European non-marine molluscs, a guide for species identification = Bestimmungsbuch für europäische Land- und Süsswassermollusken |date=2012 |publisher=Planet Poster Editions |location=Göttingen |isbn=9783933922755}}</ref>
+}}In areas where lip colour is variable, dissection is necessary. A cross-section of the [[love dart]] of ''C. nemoralis'' shows a cross with simple blades, whereas that of ''C. hortensis'' has bifurcated blades. The mucous gland has 3 or fewer branches in ''C. nemoralis'', but 4 or more in ''C. hortensis''.<ref name=Welter-Schultes2012>{{cite book |last1=Welter-Schultes |first1=F.W. |title=European non-marine molluscs, a guide for species identification = Bestimmungsbuch für europäische Land- und Süsswassermollusken |date=2012 |publisher=Planet Poster Editions |location=Göttingen |isbn=9783933922755}}</ref>
 Two superficially similar species ''[[Caucasotachea vindobonensis]]'' and ''[[Macularia sylvatica]]'' both have a lip that is brown near the [[Gastropod shell#Parts of the shell|columella]] becoming pale towards the [[Gastropod shell#Parts of the shell|suture]], and they have fine growth ridges on the shell whereas in both ''Cepaea'' species it is smooth. Also, ''M. sylvatica'' is distinct in having a small blunt tooth in its [[Gastropod shell#Parts of the shell|aperture]], whilst the lowest brown band on the shell of ''C. vindobonensis'' lies noticeably closer to the columella than in ''Cepaea''.<ref name="Kerney1979">{{cite book |last1=Kerney |first1=M.P. |last2=Cameron |first2=R.A.D. |title=A field guide to the land snails of Britain and north-west Europe |date=1979 |publisher=Collins |location=London |isbn=000219676X}}</ref>
 ==Colour polymorphism==
-{{Main|Cepaea#The_shell_polymorphism|l1=''Cepaea'': the shell polymorphism}}
+{{Main|Cepaea#Shell polymorphism}}
-''Cepaea nemoralis'' is highly [[Polymorphism (biology)|polymorphic]] in shell colour and banding. The background colour of the shell various along a continuum from brown through pink to yellow and sometimes almost white.<ref name="Davison2019">{{cite journal |last1=Davison |first1=A. |last2=Jackson |first2=H.J. |last3=Murphy |first3=E.W. |last4=Reader |first4=T. |title=Discrete or indiscrete? Redefining the colour polymorphism of the land snail ''Cepaea nemoralis'' |journal=Heredity |date=August 2019 |volume=123 |issue=2 |pages=162–175 |doi=10.1038/s41437-019-0189-z}}</ref> Additionally the shells can be with or without dark bands. The bands vary in intensity of colour, in width and in number, from zero to five. The genetics underlying this variation is extensively understood and is shared with ''C. hortensis''.<ref name=Jones1977/>
+''Cepaea nemoralis'' is highly [[Polymorphism (biology)|polymorphic]] in shell colour and banding. The background colour of the shell varies along a continuum from brown through pink to yellow and sometimes almost white.<ref name="Davison2019">{{cite journal |last1=Davison |first1=A. |last2=Jackson |first2=H.J. |last3=Murphy |first3=E.W. |last4=Reader |first4=T. |title=Discrete or indiscrete? Redefining the colour polymorphism of the land snail ''Cepaea nemoralis'' |journal=Heredity |date=August 2019 |volume=123 |issue=2 |pages=162–175 |doi=10.1038/s41437-019-0189-z|pmid=30804571 |pmc=6629550 |doi-access=free }}</ref> Additionally the shells can be with or without dark bands. The bands vary in intensity of colour, in width and in number, from zero to five. The genetics underlying this variation is extensively understood and is shared with ''C. hortensis''.<ref name=Jones1977/>
-The polymorphism has also been intensely studied for its [[evolution]] and [[ecology]]. For instance, in stable habitats shells tend to be darker in woodland than in open habitats.<ref name="Cameron2012">{{cite journal |last1=Cameron |first1=R.A.D. |last2=Cook |first2=L.M. |title=Habitat and the shell polymorphism of ''Cepaea nemoralis'' (L.): interrogating the Evolution Megalab database |journal=Journal of Molluscan Studies |date=May 2012 |volume=78 |issue=2 |pages=179–184 |doi=10.1093/mollus/eyr052}}</ref> The explanation might be [[camouflage]] or climatic selection: paler, more reflective colours in sunny environments reduce water loss and overheating. Climatic selection can also explain why yellow shells are commoner in the south.<ref name=Silvertown2011>{{cite journal |last1=Silvertown |first1=J. |last2=Cook |first2=L. |last3=Cameron |first3=R. |last4=Dodd |first4=M. |last5=McConway |first5=K. |last6=Worthington |first6=J. |last7=Skelton |first7=P. |last8=Anton |first8=C. |last9=Bossdorf |first9=O. |last10=Baur |first10=B. |last11=Schilthuizen |first11=M. |last12=Fontaine |first12=B. |last13=Sattmann |first13=H. |last14=Bertorelle |first14=G. |last15=Correia |first15=M. |last16=Oliveira |first16=C. |last17=Pokryszko |first17=B. |last18=Ożgo |first18=M. |last19=Stalažs |first19=A. |last20=Gill |first20=E. |last21=Rammul |first21=Ü. |last22=Sólymos |first22=P. |last23=Féher |first23=Z. |last24=Juan |first24=X. |title=Citizen science reveals unexpected continental-scale evolutionary change in a model organism |journal=PLoS ONE |date=27 April 2011 |volume=6 |issue=4 |pages=e18927 |doi=10.1371/journal.pone.0018927}}</ref>
+The polymorphism has also been intensely studied for its [[evolution]] and [[ecology]]. For instance, in stable habitats shells tend to be darker in woodland than in open habitats.<ref name="Cameron2012">{{cite journal |last1=Cameron |first1=R.A.D. |last2=Cook |first2=L.M. |title=Habitat and the shell polymorphism of ''Cepaea nemoralis'' (L.): interrogating the Evolution Megalab database |journal=Journal of Molluscan Studies |date=May 2012 |volume=78 |issue=2 |pages=179–184 |doi=10.1093/mollus/eyr052|doi-access=free }}</ref> The explanation might be [[camouflage]] or climatic selection: paler, more reflective colours in sunny environments reduce water loss and overheating. Climatic selection can also explain why yellow shells are more common in the south.<ref name=Silvertown2011>{{cite journal |last1=Silvertown |first1=J. |last2=Cook |first2=L. |last3=Cameron |first3=R. |last4=Dodd |first4=M. |last5=McConway |first5=K. |last6=Worthington |first6=J. |last7=Skelton |first7=P. |last8=Anton |first8=C. |last9=Bossdorf |first9=O. |last10=Baur |first10=B. |last11=Schilthuizen |first11=M. |last12=Fontaine |first12=B. |last13=Sattmann |first13=H. |last14=Bertorelle |first14=G. |last15=Correia |first15=M. |last16=Oliveira |first16=C. |last17=Pokryszko |first17=B. |last18=Ożgo |first18=M. |last19=Stalažs |first19=A. |last20=Gill |first20=E. |last21=Rammul |first21=Ü. |last22=Sólymos |first22=P. |last23=Féher |first23=Z. |last24=Juan |first24=X. |title=Citizen science reveals unexpected continental-scale evolutionary change in a model organism |journal=PLOS ONE |date=27 April 2011 |volume=6 |issue=4 |pages=e18927 |doi=10.1371/journal.pone.0018927|pmid=21556137 |pmc=3083392 |doi-access=free }}</ref>
-Another question is why the variation persists, usually even within a locality. Researchers have variously argued that the cause is random [[genetic drift]] and [[founder effects]], different [[natural selection|selection]] pressures in different areas with mixing by migration, and balanced polymorphism. Balanced polymorphism could arise when a predator like the [[song thrush]] develops a 'search image' for the commonest morph, so that the rarer morphs are less likely to be predated. Natural selection would then favour a diversity of [[animal coloration|colours and patterns]] as an [[antipredator adaptation]]. Most probably, the polymorphism has multiple causes.<ref name="Jones1977" /><ref name="Cook1998">{{cite journal |last1=Cook |first1=L.M. |title=A two–stage model for ''Cepaea'' polymorphism |journal=Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences |date=29 October 1998 |volume=353 |issue=1375 |pages=1577–1593 |doi=10.1098/rstb.1998.0311}}</ref>
+Another question is why the variation persists, usually even within a locality. Researchers have variously argued that the cause is random [[genetic drift]] and [[founder effects]], different [[natural selection|selection]] pressures in different areas with mixing by migration, and balanced polymorphism. Balanced polymorphism could arise when a predator like the [[song thrush]] develops a 'search image' for the commonest morph, so that the rarer morphs are less likely to be predated. Natural selection would then favour a diversity of [[animal coloration|colours and patterns]] as an [[antipredator adaptation]]. Most probably, the polymorphism has multiple causes.<ref name="Jones1977" /><ref name="Cook1998">{{cite journal |last1=Cook |first1=L.M. |title=A two–stage model for ''Cepaea'' polymorphism |journal=Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences |date=29 October 1998 |volume=353 |issue=1375 |pages=1577–1593 |doi=10.1098/rstb.1998.0311|pmc=1692378 }}</ref>
 {{gallery |title=Different coloration and banding of the shells of ''Cepaea nemoralis'' |height=135 |mode=packed
@@ Line 70: / Line 70: @@
 ==Distribution==
 [[File:Snail traversing a leaf - 20190801.webm|thumb|Snail traversing a leaf.]]
-The native distribution of ''C. nemoralis'' is from Western and Northern Europe to Central Europe, but it has been spreading eastwards especially over the last few decades.<ref>Dvořák L., Honěk A. & Martínková Z. (2003). [https://web.archive.org/web/20060525044725/http://www.mollusca.cz/malakologie/Cepaea-nemoralis.jpg "The spread of ''Cepaea nemoralis'' (L.) populations in the Czech Republic"]. ''2003 BCPC Symposiumproceedings No. 80: Slugs & snails: agricultural, veterinary & environmental perspectives'':  99-102.</ref><ref name=Ozgo2019/> Thus it is known from most of the Iberian Peninsula,<ref>{{cite book |last1=Cadevall |first1=J. |last2=Orozco |first2=A. |title=Caracoles y babosas de la Península Ibérica y Baleares |date=2016 |publisher=Omega |location=Barcelona |isbn=9788428215992}}</ref> [[List of non-marine molluscs of France|France]], [[List of non-marine molluscs of Great Britain|Great Britain]], [[List of non-marine molluscs of Ireland|Ireland]], Belgium, [[List of non-marine molluscs of the Netherlands|the Netherlands]], [[List of non-marine molluscs of Germany|Germany]], Denmark, southern Sweden and Norway, [[List of non-marine molluscs of Switzerland|Switzerland]], [[List of non-marine molluscs of Austria|Austria]], the [[List of molluscs recorded in the Czech Republic|Czech Republic]],<ref>{{cite journal |last1=Juřičková |first1=L. |last2=Horsák |first2=M. |last3=Beran |first3=L. | year = 2001 | title = Check-list of the molluscs (Mollusca) of the Czech Republic | journal = Acta Soc. Zool. Bohem. | volume = 65 | pages = 25–40 }}</ref><ref>{{cite journal |last1=Peltanová |first1=Alena |last2=Dvořák |first2=Libor |last3=Juřičková |first3=Lucie |title=The spread of non-native ''Cepaea nemoralis'' and ''Monacha cartusiana'' (Gastropoda: Pulmonata) in the Czech Republic with comments on other land snail immigrants |journal=Biologia |date=1 April 2012 |volume=67 |issue=2 |pages=384–389 |doi=10.2478/s11756-012-0020-2}}</ref> Hungary, Bosnia, Croatia, and the northern half of Italy.<ref name="AB"/><ref>{{cite journal |last1=Ramos‐Gonzalez |first1=D. |last2=Saenko |first2=S.V. |last3=Davison |first3=A. |title=Deep structure, long‐distance migration and admixture in the colour polymorphic land snail ''Cepaea nemoralis'' |journal=Journal of Evolutionary Biology |date=August 2022 |volume=35 |issue=8 |pages=1110–1125 |doi=10.1111/jeb.14060}}</ref> In Central and Eastern Europe it has spread particularly along the Baltic coast (e.g. in [[List of non-marine molluscs of Poland|Poland]],<ref name=Ozgo2019>{{cite journal |last1=Ożgo |first1=M. |last2=Cameron |first2=R.A.D. |last3=Horsák |first3=M. |last4=Pokryszko |first4=B. |last5=Chudaś |first5=M. |last6=Cichy |first6=A. |last7=Kaczmarek |first7=S. |last8=Kobak |first8=J. |last9=Marzec |first9=M. |last10=Mierzwa-Szymkowiak |first10=D. |last11=Parzonko |first11=D. |last12=Pyka |first12=G. |last13=Rosin |first13=Z. |last14=Skawina |first14=A. |last15=Soroka |first15=M. |last16=Sulikowska-Drozd |first16=A. |last17=Surowiec |first17=T. |last18=Szymanek |first18=M. |last19=Templin |first19=J. |last20=Urbańska |first20=M. |last21=Zając |first21=K. |last22=Zielska |first22=J. |last23=Żbikowska |first23=E. |last24=Żołądek |first24=J. |title=Cepaea nemoralis (Gastropoda: Pulmonata) in Poland: patterns of variation in a range-expanding species |journal=Biological Journal of the Linnean Society |date=17 April 2019 |volume=127 |issue=1 |pages=1–11 |doi=10.1093/biolinnean/blz029}}</ref> [[List of non-marine molluscs of Latvia|Latvia]], Estonia, southern Finland,<ref>{{cite book |last1=Koivunen |first1=A. |last2=Malinen |first2=P. |last3=Ormio |first3=H. |last4=Terhivuo |first4=J. |last5=Valovirta |first5=I. |title=Suomen kotilot ja etanat : opas maanilviäisten maailmaan |date=2014 |publisher=Tibiale |location=Helsinki |isbn=9789526754468}}</ref> the east coast of Sweden)<ref>{{cite journal |last1=Ożgo |first1=M. |last2=Örstan |first2=A. |last3=Kirschenstein |first3=M. |last4=Cameron |first4=R. |title=Dispersal of land snails by sea storms |journal=Journal of Molluscan Studies |date=May 2016 |volume=82 |issue=2 |pages=341–343 |doi=10.1093/mollus/eyv060}}</ref> but also now elsewhere in Poland,<ref name=Ozgo2019/> and in [[List of non-marine molluscs of Ukraine|Ukraine]], Belarus and Russia.<ref name=Gural-Sverlova2021>{{cite journal |last1=Gural-Sverlova |first1=N. |last2=Egorov |first2=R. |last3=Kruglova |first3=O. |last4=Kovalevich |first4=N. |last5=Gural |first5=R. |title=Introduced land snail ''Cepaea nemoralis'' (Gastropoda: Helicidae) in Eastern Europe: spreading history and the shell colouration variability |journal=Malacologica Bohemoslovaca |date=27 September 2021 |volume=20 |pages=75–91 |doi=10.5817/MaB2021-20-75}}</ref> In Slovakia it was still known only from a single garden centre by 2020.<ref>{{cite journal |last1=Čejka |first1=T. |title=The terrestrial molluscan fauna in the Slovak part of the Danubian Lowland: an annotated checklist |journal=Authorea |date=3 August 2020 |doi=10.22541/au.159620996.68925423/v2}}</ref>
+The native distribution of ''C. nemoralis'' is from Western and Northern Europe to Central Europe, but it has been spreading eastwards especially over the last few decades.<ref>Dvořák L., Honěk A. & Martínková Z. (2003). [https://web.archive.org/web/20060525044725/http://www.mollusca.cz/malakologie/Cepaea-nemoralis.jpg "The spread of ''Cepaea nemoralis'' (L.) populations in the Czech Republic"]. ''2003 BCPC Symposiumproceedings No. 80: Slugs & snails: agricultural, veterinary & environmental perspectives'':  99-102.</ref><ref name=Ozgo2019/> Thus it is known from most of the Iberian Peninsula,<ref>{{cite book |last1=Cadevall |first1=J. |last2=Orozco |first2=A. |title=Caracoles y babosas de la Península Ibérica y Baleares |date=2016 |publisher=Omega |location=Barcelona |isbn=9788428215992}}</ref> [[List of non-marine molluscs of France|France]], [[List of non-marine molluscs of Great Britain|Great Britain]], [[List of non-marine molluscs of Ireland|Ireland]], Belgium, [[List of non-marine molluscs of the Netherlands|the Netherlands]], [[List of non-marine molluscs of Germany|Germany]], Denmark, southern Sweden and Norway, [[List of non-marine molluscs of Switzerland|Switzerland]], [[List of non-marine molluscs of Austria|Austria]], the [[List of molluscs recorded in the Czech Republic|Czech Republic]],<ref>{{cite journal |last1=Juřičková |first1=L. |last2=Horsák |first2=M. |last3=Beran |first3=L. | year = 2001 | title = Check-list of the molluscs (Mollusca) of the Czech Republic | journal = Acta Soc. Zool. Bohem. | volume = 65 | pages = 25–40 }}</ref><ref>{{cite journal |last1=Peltanová |first1=Alena |last2=Dvořák |first2=Libor |last3=Juřičková |first3=Lucie |title=The spread of non-native ''Cepaea nemoralis'' and ''Monacha cartusiana'' (Gastropoda: Pulmonata) in the Czech Republic with comments on other land snail immigrants |journal=Biologia |date=1 April 2012 |volume=67 |issue=2 |pages=384–389 |doi=10.2478/s11756-012-0020-2|doi-access=free }}</ref> Hungary, Bosnia, Croatia, and the northern half of Italy.<ref name="AB"/><ref>{{cite journal |last1=Ramos‐Gonzalez |first1=D. |last2=Saenko |first2=S.V. |last3=Davison |first3=A. |title=Deep structure, long‐distance migration and admixture in the colour polymorphic land snail ''Cepaea nemoralis'' |journal=Journal of Evolutionary Biology |date=August 2022 |volume=35 |issue=8 |pages=1110–1125 |doi=10.1111/jeb.14060|pmid=35830483 |pmc=9541890 }}</ref> In Central and Eastern Europe it has spread particularly along the Baltic coast (e.g. in [[List of non-marine molluscs of Poland|Poland]],<ref name=Ozgo2019>{{cite journal |last1=Ożgo |first1=M. |last2=Cameron |first2=R.A.D. |last3=Horsák |first3=M. |last4=Pokryszko |first4=B. |last5=Chudaś |first5=M. |last6=Cichy |first6=A. |last7=Kaczmarek |first7=S. |last8=Kobak |first8=J. |last9=Marzec |first9=M. |last10=Mierzwa-Szymkowiak |first10=D. |last11=Parzonko |first11=D. |last12=Pyka |first12=G. |last13=Rosin |first13=Z. |last14=Skawina |first14=A. |last15=Soroka |first15=M. |last16=Sulikowska-Drozd |first16=A. |last17=Surowiec |first17=T. |last18=Szymanek |first18=M. |last19=Templin |first19=J. |last20=Urbańska |first20=M. |last21=Zając |first21=K. |last22=Zielska |first22=J. |last23=Żbikowska |first23=E. |last24=Żołądek |first24=J. |title=Cepaea nemoralis (Gastropoda: Pulmonata) in Poland: patterns of variation in a range-expanding species |journal=Biological Journal of the Linnean Society |date=17 April 2019 |volume=127 |issue=1 |pages=1–11 |doi=10.1093/biolinnean/blz029}}</ref> [[List of non-marine molluscs of Latvia|Latvia]], Estonia, southern Finland,<ref>{{cite book |last1=Koivunen |first1=A. |last2=Malinen |first2=P. |last3=Ormio |first3=H. |last4=Terhivuo |first4=J. |last5=Valovirta |first5=I. |title=Suomen kotilot ja etanat : opas maanilviäisten maailmaan |date=2014 |publisher=Tibiale |location=Helsinki |isbn=9789526754468}}</ref> the east coast of Sweden)<ref>{{cite journal |last1=Ożgo |first1=M. |last2=Örstan |first2=A. |last3=Kirschenstein |first3=M. |last4=Cameron |first4=R. |title=Dispersal of land snails by sea storms |journal=Journal of Molluscan Studies |date=May 2016 |volume=82 |issue=2 |pages=341–343 |doi=10.1093/mollus/eyv060|doi-access=free }}</ref> but also now elsewhere in Poland,<ref name=Ozgo2019/> and in [[List of non-marine molluscs of Ukraine|Ukraine]], Belarus and Russia.<ref name=Gural-Sverlova2021>{{cite journal |last1=Gural-Sverlova |first1=N. |last2=Egorov |first2=R. |last3=Kruglova |first3=O. |last4=Kovalevich |first4=N. |last5=Gural |first5=R. |title=Introduced land snail ''Cepaea nemoralis'' (Gastropoda: Helicidae) in Eastern Europe: spreading history and the shell colouration variability |journal=Malacologica Bohemoslovaca |date=27 September 2021 |volume=20 |pages=75–91 |doi=10.5817/MaB2021-20-75|doi-access=free }}</ref> In Slovakia it was still known only from a single garden centre by 2020.<ref>{{cite journal |last1=Čejka |first1=T. |title=The terrestrial molluscan fauna in the Slovak part of the Danubian Lowland: an annotated checklist |journal=Authorea |date=3 August 2020 |doi=10.22541/au.159620996.68925423/v2|s2cid=242206099 }}</ref>
 At the northern edge of its range,''C. nemoralis'' is rare and scattered in northern Scotland, where it has been introduced.<ref name="AB" /> It is not found in the [[Hebrides]], [[Orkney]] or [[Shetland]].<ref name="AB" /> It seems to have been affected by [[air pollution]] and [[soil acidification]] in some parts of England.<ref name="AB" />
-Starting in 1857, there have been multiple introductions to North America, where it now occurs widely in Canada (from British Columbia to Newfoundland; in Ontario it is a pest of vineyards) and in the north-eastern part of the US, with further occurrences further south (e.g. California, Colorado, Texas and South Carolina).<ref>{{cite journal |last1=Layton |first1=K.K.S. |last2=Warne |first2=C.P.K. |last3=Nicolai |first3=A. |last4=Ansart |first4=A. |last5=deWaard |first5=J.R. |title=Molecular evidence for multiple introductions of the banded grove snail (''Cepaea nemoralis'') in North America |journal=Canadian Journal of Zoology |date=April 2019 |volume=97 |issue=4 |pages=392–398 |doi=10.1139/cjz-2018-0084}}</ref><ref name=Whitson2005>{{cite journal |last1=Whitson |first1=M. |title=''Cepaea nemoralis'' (Gastropoda, Helicidae): The invited invader |journal=Journal of the Kentucky Academy of Science |date=September 2005 |volume=66 |issue=2 |pages=82–88 |doi=10.3101/1098-7096(2006)66[82:CNGHTI]2.0.CO;2}}</ref><ref>{{cite book |last1=Burke |first1=T.E. |title=Land snails and slugs of the Pacific Northwest |date=2013 |publisher=Oregon State University Press |location=Corvallis, Oregon |isbn=978-0-87071-685-0}}</ref>
+Starting in 1857, there have been multiple introductions to North America, where it now occurs widely in Canada (from British Columbia to Newfoundland; in Ontario it is a pest of vineyards) and in the north-eastern part of the US, with further occurrences further south (e.g. California, Colorado, Texas and South Carolina).<ref>{{cite journal |last1=Layton |first1=K.K.S. |last2=Warne |first2=C.P.K. |last3=Nicolai |first3=A. |last4=Ansart |first4=A. |last5=deWaard |first5=J.R. |title=Molecular evidence for multiple introductions of the banded grove snail (''Cepaea nemoralis'') in North America |journal=Canadian Journal of Zoology |date=April 2019 |volume=97 |issue=4 |pages=392–398 |doi=10.1139/cjz-2018-0084|s2cid=91897997 |url=https://hal-univ-rennes1.archives-ouvertes.fr/hal-01970952/file/Layton%20et%20al-2019-Molecular%20evidence%20for%20multiple%20introductions%20of%20the%20banded%20grove%20snail%20%281%29.pdf }}</ref><ref name=Whitson2005>{{cite journal |last1=Whitson |first1=M. |title=''Cepaea nemoralis'' (Gastropoda, Helicidae): The invited invader |journal=Journal of the Kentucky Academy of Science |date=September 2005 |volume=66 |issue=2 |pages=82–88 |doi=10.3101/1098-7096(2006)66[82:CNGHTI]2.0.CO;2|s2cid=86005518 |url=https://www.biodiversitylibrary.org/part/335690 }}</ref><ref>{{cite book |last1=Burke |first1=T.E. |title=Land snails and slugs of the Pacific Northwest |date=2013 |publisher=Oregon State University Press |location=Corvallis, Oregon |isbn=978-0-87071-685-0}}</ref>
 Both in America and Eastern Europe it is known that some introductions have been deliberate.<ref name=Whitson2005/><ref name=Gural-Sverlova2021/>
@@ Line 84: / Line 84: @@
 This is a very common and widespread species in Western Europe, occupying a wide range of habitats from coastal dunes to woodlands with full canopy cover, including gardens and abandoned land.<ref name="Silvertown2011"/> In Eastern Europe, where it is a new arrival, it has turned up particularly in urban areas and other disturbed habitats.<ref name=Ozgo2019/> It can be found up to an altitude of 1600 m in the Alps,<ref>{{cite book |last1=Turner |first1=H. |last2=Kuiper |first2=J.G.J. |last3=Thew |first3=N. |last4=Bernasconi |first4=R. |last5=Rüetschi |first5=J. |last6=Wüthrich |first6=M. |last7=Gosteli |first7=M. |title=Atlas der Mollusken der Schweiz und Liechtensteins |date=1998 |publisher=CentreSuisse de cartographie de la faune |location=Neuchâtel |isbn=2884140131}}</ref> 1800&nbsp;m in the Pyrenees, 900 m in Wales, 600 m in Scotland.<ref name="AB"/> Density of adults is often of the order of 2 per square metre.<ref name="Williamson1977">{{cite journal |last1=Williamson |first1=P. |last2=Cameron |first2=R.A.D. |last3=Carter |first3=M.A. |title=Population dynamics of the landsnail ''Cepaea nemoralis'' L.: a six-year study |journal=The Journal of Animal Ecology |date=February 1977 |volume=46 |issue=1 |pages=181 |doi=10.2307/3955 |jstor=3955}}</ref>
-This species feeds mainly on dead or senescent plants.<ref name="Silvertown2011"/><ref name="AB"/> It prefers broad-leaved plants over grasses.<ref>{{cite journal |last1=Chang |first1=H.-W. |last2=Emlen |first2=John M. |title=Seasonal variation of microhabitat distribution of the polymorphic land snail ''Cepaea nemoralis'' |journal=Oecologia |date=1 April 1993 |volume=93 |issue=4 |pages=501–507 |doi=10.1007/BF00328957}}</ref> Although mostly not a pest of crops,<ref name="AB"/> it can be a nuisance in vineyards because it is inadvertently picked with the grapes.<ref>{{cite journal |last1=Giffard |first1=B. |last2=Winter |first2=S. |last3=Guidoni |first3=S. |last4=Nicolai |first4=A. |last5=Castaldini |first5=M. |last6=Cluzeau |first6=D. |last7=Coll |first7=P. |last8=Cortet |first8=J. |last9=Le Cadre |first9=E. |last10=d’Errico |first10=G. |last11=Forneck |first11=A. |last12=Gagnarli |first12=E. |last13=Griesser |first13=M. |last14=Guernion |first14=M. |last15=Lagomarsino |first15=A. |last16=Landi |first16=S. |last17=Le Bissonnais |first17=Y. |last18=Mania |first18=E. |last19=Mocali |first19=S. |last20=Preda |first20=C. |last21=Priori |first21=S. |last22=Reineke |first22=A |last23=Rusch |first23=A. |last24=Schroers |first24=H.-J. |last25=Simoni |first25=S. |last26=Steiner |first26=M. |last27=Temneanu |first27=E. |last28=Bacher |first28=S. |last29=Costantini |first29=E.A.C. |last30=Zaller |first30=J. |last31=Leyer |first31=I. |title=Vineyard management and its impacts on soil biodiversity, functions, and ecosystem services |journal=Frontiers in Ecology and Evolution |date=2022 |volume=10 |doi=10.3389/fevo.2022.850272}}</ref>
+This species feeds mainly on dead or senescent plants.<ref name="Silvertown2011"/><ref name="AB"/> It prefers broad-leaved plants over grasses.<ref>{{cite journal |last1=Chang |first1=H.-W. |last2=Emlen |first2=John M. |title=Seasonal variation of microhabitat distribution of the polymorphic land snail ''Cepaea nemoralis'' |journal=Oecologia |date=1 April 1993 |volume=93 |issue=4 |pages=501–507 |doi=10.1007/BF00328957|pmid=28313817 |s2cid=22594544 }}</ref> Although mostly not a pest of crops,<ref name="AB"/> it can be a nuisance in vineyards because it is inadvertently picked with the grapes.<ref>{{cite journal |last1=Giffard |first1=B. |last2=Winter |first2=S. |last3=Guidoni |first3=S. |last4=Nicolai |first4=A. |last5=Castaldini |first5=M. |last6=Cluzeau |first6=D. |last7=Coll |first7=P. |last8=Cortet |first8=J. |last9=Le Cadre |first9=E. |last10=d’Errico |first10=G. |last11=Forneck |first11=A. |last12=Gagnarli |first12=E. |last13=Griesser |first13=M. |last14=Guernion |first14=M. |last15=Lagomarsino |first15=A. |last16=Landi |first16=S. |last17=Le Bissonnais |first17=Y. |last18=Mania |first18=E. |last19=Mocali |first19=S. |last20=Preda |first20=C. |last21=Priori |first21=S. |last22=Reineke |first22=A |last23=Rusch |first23=A. |last24=Schroers |first24=H.-J. |last25=Simoni |first25=S. |last26=Steiner |first26=M. |last27=Temneanu |first27=E. |last28=Bacher |first28=S. |last29=Costantini |first29=E.A.C. |last30=Zaller |first30=J. |last31=Leyer |first31=I. |title=Vineyard management and its impacts on soil biodiversity, functions, and ecosystem services |journal=Frontiers in Ecology and Evolution |date=2022 |volume=10 |doi=10.3389/fevo.2022.850272|doi-access=free }}</ref>
-Like all [[pulmonate]] land snails, it is [[hermaphrodite]], and this species must mate to produce fertile eggs.<ref name="Silvertown2011"/> Mating tends to be concentrated in late spring and early summer, though it can continue through the autumn.<ref name="Silvertown2011"/> The snails may store the sperm they receive from their partner for some time, and individual broods can have mixed paternity.<ref name="Silvertown2011"/> In Britain it lays clutches of 30–50 (in France 40–80) oval eggs between June and August (in France May–October, in W France until November).<ref name="AB"/> The size of the egg is 3.1 × 2.6&nbsp;mm<ref name="Barker">Heller J. (2001). ''Life History Strategies''. In: Barker G. M. (ed.). ''[https://books.google.com/books?id=WlvX-9Wt0toC&hl The biology of terrestrial molluscs]''. CABI Publishing, Oxon, UK, {{ISBN|0-85199-318-4}}. 1-146, cited page: 428.</ref> or egg diameter can be 2.3–3.0&nbsp;mm.<ref name="AB"/> Juveniles hatch after 15–20 days.<ref name="AB"/>
+Like all [[pulmonate]] land snails, it is a [[hermaphrodite]], and this species must mate to produce fertile eggs.<ref name="Silvertown2011"/> Mating tends to be concentrated in late spring and early summer, though it can continue through the autumn.<ref name="Silvertown2011"/> The snails may store the sperm they receive from their partner for some time, and individual broods can have mixed paternity.<ref name="Silvertown2011"/> In Britain it lays clutches of 30–50 (in France 40–80) oval eggs between June and August (in France May–October, in W France until November).<ref name="AB"/> The size of the egg is 3.1 × 2.6&nbsp;mm<ref name="Barker">Heller J. (2001). ''Life History Strategies''. In: Barker G. M. (ed.). ''[https://books.google.com/books?id=WlvX-9Wt0toC&hl The biology of terrestrial molluscs]''. CABI Publishing, Oxon, UK, {{ISBN|0-85199-318-4}}. 1-146, cited page: 428.</ref> or egg diameter can be 2.3–3.0&nbsp;mm.<ref name="AB"/> Juveniles hatch after 15–20 days.<ref name="AB"/>
-This snail is comparatively slow-growing, taking 1 to 3 years to develop from an egg to a breeding adult.<ref>{{cite journal |last1=Oosterhoff |first1=L. M. |title=Variation in growth rate as an ecological factor in the landsnail ''Cepaea nemoralis'' (L.) |journal=Netherlands Journal of Zoology |date=1 January 1976 |volume=27 |issue=1 |pages=1–132 |doi=10.1163/002829677X00072}}</ref> The life-span for this species is up to seven or eight years, with the annual survival rate of adults about 50% (= 3% over five years, older adults suffer higher mortalities).<ref name="AB"/> In winter, the snails may [[hibernate]], but can become active again during warm spells.<ref name="Silvertown2011"/>
+This snail is comparatively slow-growing, taking 1 to 3 years to develop from an egg to a breeding adult.<ref>{{cite journal |last1=Oosterhoff |first1=L. M. |title=Variation in growth rate as an ecological factor in the landsnail ''Cepaea nemoralis'' (L.) |journal=Netherlands Journal of Zoology |date=1 January 1976 |volume=27 |issue=1 |pages=1–132 |doi=10.1163/002829677X00072|s2cid=55537228 |url=https://research.rug.nl/en/publications/472621dd-1a18-4bed-8884-b144af81ee5d }}</ref> The life-span for this species is up to seven or eight years, with the annual survival rate of adults about 50% (= 3% over five years, older adults suffer higher mortalities).<ref name="AB"/> In winter, the snails may [[hibernate]], but can become active again during warm spells.<ref name="Silvertown2011"/>
 ''Cepaea nemoralis'' is known experimentally to be a host for ''[[Angiostrongylus vasorum]]''.<ref>Conboy G.A. (30 May 2000). [http://www.ivis.org/advances/Parasit_Bowman/conboy_angiostrongylosis/ivis.pdf "Canine Angiostrongylosis (French Heartworm)"]. In: Bowman D. D. (Ed.) ''Companion and Exotic Animal Parasitology''. International Veterinary Information Service. Accessed 24 November 2009.</ref> Predators of ''Cepaea nemoralis'' include the [[song thrush]] (''Turdus philomelos'').
@@ Line 108: / Line 108: @@
 [[Category:Helicidae]]
 [[Category:Gastropods described in 1758]]
-[[Category:Molluscs of Europe]]
+[[Category:Gastropods of Europe]]
 [[Category:Taxa named by Carl Linnaeus]]