# Cepaea

> Mediated Wiki article. Canonical URL: https://mediated.wiki/source/Cepaea
> Markdown URL: https://mediated.wiki/source/Cepaea.md
> Source: https://en.wikipedia.org/wiki/Cepaea
> Source revision: 1313953732
> License: Creative Commons Attribution-ShareAlike 4.0 International (https://creativecommons.org/licenses/by-sa/4.0/)

Genus of gastropods

Not to be confused with the plant [Sedum cepaea](/source/Sedum_cepaea), the pink stonecrop.

Cepaea White-lipped snail Cepaea hortensis Scientific classification Kingdom: Animalia Phylum: Mollusca Class: Gastropoda Order: Stylommatophora Family: Helicidae Subfamily: Helicinae Tribe: Allognathini Genus: Cepaea Held, 1838[1] Type species Helix nemoralis Linnaeus, 1758 Synonyms Cepaea (Cepaea) Held, 1838 Helix (Tachea) W. Turton, 1831 (Invalid: junior homonym of Tachea Fleming, 1822 [Aves]) Hystrionica Scudder, 1882

***Cepaea*** is a [genus](/source/Genus) of large air-breathing [land snails](/source/Land_snail), [terrestrial](/source/Terrestrial_molluscs) [pulmonate](/source/Pulmonate) [gastropod](/source/Gastropod) [molluscs](/source/Mollusc) in the family [Helicidae](/source/Helicidae). The shells are often brightly coloured and patterned with brown stripes. The two living species in this genus, *C. nemoralis* and *C. hortensis*, are widespread and common in Western and Central Europe. In North America, *C. hortensis* is [native](/source/Native_species) on the northeast coast, but both species have been introduced elsewhere, and they are also spreading further east in Europe.[2] Both have been influential [model species](/source/Model_organisms) for ongoing studies of genetics and natural selection. Like many Helicidae, these snails use [love darts](/source/Love_dart) during mating.[3]

## Species

A comparison between *[Cepaea hortensis](/source/Cepaea_hortensis)* (left) and *[Caucasotachea vindobonensis](/source/Caucasotachea_vindobonensis)* (right)

For a long time, four living species were classified in the genus *Cepaea*. However, [molecular phylogenetic studies](/source/Molecular_phylogenetics) suggested that two of them should be placed in the genera *[Macularia](/source/Macularia)* and *[Caucasotachea](/source/Caucasotachea)*, which are not immediate relatives of either *Cepaea* or each other:[4][5]

- *[Cepaea hortensis](/source/White-lipped_snail)* (O. F. Müller, 1774) – white-lipped snail or garden banded snail

- *[Cepaea nemoralis](/source/Cepaea_nemoralis)* (Linnaeus, 1758) – brown-lipped snail or grove snail

- *Cepaea sylvatica* (Draparnaud, 1801), now *[Macularia sylvatica](/source/Macularia_sylvatica)*

- *Cepaea vindobonensis* (Férussac, 1821), now *[Caucasotachea vindobonensis](/source/Caucasotachea_vindobonensis)*

Several other fossil species have been described.[6]

## Interspecific relations

The range of *C. hortensis* extends further north than that of *C. nemoralis* in Scotland and Scandinavia and it is the only one of the two species in Iceland and the only one native to North America.[7][8][9] Likewise in the Swiss Alps *C. hortensis* is found as high as 2050 m, but *C. nemoralis* only up to 1600 m.[10] Conversely, the southern edge of the range lies further north in *C. hortensis*; unlike *C. nemoralis* it does not occur in Italy, and in Spain it has a more restricted distribution (in the north-east corner).[11][12]

Where the ranges overlap *C. hortensis* prefers cooler sites with longer and damper vegetation. But the two species often co-occur at a site, in which situation the densities of both affect each other's growth, fecundity and mortality. However, they differ somewhat in their behaviour: *C. hortensis* is more active at lower temperatures, aestivates higher on the vegetation and is more [diurnal](/source/Diurnality), although this appears to be independent of whether the other species is present or not.[13][14]

When given no choice of partner in the laboratory, the two *Cepaea* species can form hybrids, which will [backcross](/source/Backcross) with the parental species, but the fertility is very low.[15]

## Shell polymorphism

### Description and genetics

*Cepaea nemoralis*, showing colour and banding polymorphisms

The two *Cepaea* species share a [genetic polymorphism](/source/Genetic_polymorphism) for the colour and banding pattern of the shell.

The background colour of the shell ranges from dark brown, through pink to yellow or even approaching white. This variation is continuous, but there are peaks in the distribution corresponding to brown, pink and yellow morphs.[16] The colour is mainly determined by [alleles](/source/Alleles) at a single [locus](/source/Locus_(genetics)) with brown [dominant](/source/Dominance_(genetics)) to pink, which is dominant to yellow.[14]

Up to five bands (very rarely more) run spirally around the shell, numbered 1 to 5 with the larger numbers further from the shell apex. The conventional scoring annotation is to write 12345 if all bands are present and separated, but to replace a number with 0 if a band is absent from its usual position and to enclose numbers in parentheses if bands are fused with their neighbours. Thus 003(45) would mean that the top two bands are absent and the lower two fused.[17]

A dominant allele at one locus causes the absence of all bands, a dominant allele at another locus causes the loss of all bands except band 3, and a dominant allele at a third locus causes the loss of just bands 1 and 2. The first of these three loci is closely [linked](/source/Genetic_linkage) to the locus determining shell colour, to another influencing the spread of the band pigment, and to one determining the colour of the lip and bands. This collection of linked loci are part of a [supergene](/source/Supergene). A consequence of this arrangement is that the shells of different background colours within a population often exhibit different ratios of banded to unbanded shells: this is an example of [linkage disequilibrium](/source/Linkage_disequilibrium).[14]

The bands are usually dark brown, but this is affected by genes influencing intensity and colouration (e.g. black or orange). Another locus (part of the supergene) determines whether the band is continuous or forms a sequence of spots. The genetics underlying the fusion of adjacent bands is not well understood.[14]

### Evolutionary explanations

Proportion of *C. nemoralis* shells that are yellow

In both species, most populations exhibit polymorphism in one or more of these shell characters. Nevertheless, statistically we can detect systematic variation at continental scales, and also between habitats, and at various scales down to a few tens of metres. There is also statistical evidence of change with time, based both on comparisons between sub-fossil and modern shells,[18] and on resampling the same sites some decades apart, although the latter has more often found little change over the period ([stasis](/source/Stasis_(biology))).[19] Very much research in [ecological genetics](/source/Ecological_genetics) has addressed the reasons for both the variation and the systematic trends.[14][20]

The two selection pressures that might most feasibly act on the appearance of shells are [climatic selection](https://en.wikipedia.org/w/index.php?title=Climatic_selection&action=edit&redlink=1) and predation. Darker shells heat up more quickly in the sun, which might well be advantageous for cold-blooded animals in shaded woodland but risks causing overheating and death in open habitats.[21][22] This trade-off is also presumed to be responsible for the greater proportion of yellow *C. nemoralis* to the south, but it is curious why the trend is not present in *C. hortensis*.[14] Contrary to predictions, recent global warming has not led to a detectable increase in yellow morphs on a continental scale.[23] The use of photosensitive paint has shown that paler morphs spend more time exposed to the sun, which may imply that the shell polymorphism allows different morphs to coexist at a site by occupying different microhabitats.[24]

Song thrush anvil with broken *Cepaea* shells

Both temperature regulation and predation make the same prediction of pale shells in open habitats and dark shells in woodland, so—although the prediction has often been confirmed[25]—it is difficult to test which is the more important explanation. However, [song thrushes](/source/Song_thrush) (*Turdus philomelos*) break open *Cepaea* shells on stones ("anvils"), allowing a comparison of those they predate with those present in the local environment. Besides the directional selection favouring camouflaged individuals,[26] visually searching predators might cause [apostatic selection](/source/Apostatic_selection). The hypothesis is that they form a search image for the commonest morphs, favouring whichever morphs are locally rare, thus promoting diversity.[27] As well as its visual effect, the shell pigments are associated with differences in shell strength, so may affect predation by predators searching non-visually, for instance at night.[28]

Several studies have demonstrated a predicted evolutionary response of shell appearance to a change of habitat.[29] However, the association of shell appearance and habitat is not always consistent, especially in more disturbed environments,[25] so it is believed that random effects are also influential, particularly [founder effects](/source/Founder_effects). The two *Cepaea* species colonised much of Europe only within the last 4000 generations,[20] so the time available for selection to act has been limited, and local anthropogenetic disturbances must often have reversed which morphs are optimal. Moreover, snails disperse more slowly than many other animals, so the most suitable genes may be locally absent.

For instance, biologists were at one time puzzled by the phenomenon of "[area effects](https://en.wikipedia.org/w/index.php?title=Area_effects&action=edit&redlink=1)"; the same morph of *Cepaea* may be found consistently over a wide area but in adjacent areas of similar habitat a different set of morphs predominate instead, with a sharp transition between. The explanation accepted nowadays is that relatively recently a change of habitat allowed the rapid colonisation of vacant areas by descendants of a few founder individuals until the colony had expanded out to areas occupied by other populations; subsequently intraspecific competition slowed the dispersal of genes into the neighbouring, occupied areas.[30][31][32] Nevertheless, occasional transfer of genes between areas of different habitat is proposed to be important in maintaining the local diversity of phenotypes.[20]

## References

1. **[^](#cite_ref-1)** [Held, F.](/source/Friedrich_Held) (1838). "Notizen über die Weichthiere Bayerns. (Fortsetzung)". *Isis (Leipzig)*. **1837** (12): 902–921.

1. **[^](#cite_ref-Gural-Sverlova2021_2-0)** 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"](https://doi.org/10.5817%2FMaB2021-20-75). *Malacologica Bohemoslovaca*. **20**: 75–91. [doi](/source/Doi_(identifier)):[10.5817/MaB2021-20-75](https://doi.org/10.5817%2FMaB2021-20-75).

1. **[^](#cite_ref-Nordsieck_3-0)** Robert Nordsieck. ["Banded Snails (*Cepaea* Held 1838)"](http://www.molluscs.at/gastropoda/terrestrial.html?/gastropoda/terrestrial/banded_snails.html). *The Living World of Molluscs*. Retrieved 19 August 2015.

1. **[^](#cite_ref-4)** Neiber, M.T.; Hausdorf, B. (2015). "Molecular phylogeny reveals the polyphyly of the snail genus *Cepaea* (Gastropoda: Helicidae)". *Molecular Phylogenetics and Evolution*. **93**: 143–149. [doi](/source/Doi_(identifier)):[10.1016/j.ympev.2015.07.022](https://doi.org/10.1016%2Fj.ympev.2015.07.022). [PMID](/source/PMID_(identifier)) [26256642](https://pubmed.ncbi.nlm.nih.gov/26256642).

1. **[^](#cite_ref-5)** Neiber, M.T.; Sagorny, C.; Hausdorf, B. (2016). ["Increasing the number of molecular markers resolves the phylogenetic relationship of "*Cepaea*" *vindobonensis* (Pfeiffer 1828) with *Caucasotachea* Boettger 1909 (Gastropoda: Pulmonata: Helicidae)"](https://doi.org/10.1111%2Fjzs.12116). *Journal of Zoological Systematics and Evolutionary Research*. **54** (1): 40–45. [doi](/source/Doi_(identifier)):[10.1111/jzs.12116](https://doi.org/10.1111%2Fjzs.12116).

1. **[^](#cite_ref-MolluscaBase_6-0)** MolluscaBase Eds. ["*Cepaea* Held, 1838"](https://www.molluscabase.org/aphia.php?p=taxdetails&id=235791). *MolluscaBase*. Flanders Marine Institute. Retrieved 14 June 2025.

1. **[^](#cite_ref-7)** Bengtson, S.-A.; Nilsson, A.; Nordström, S.; Rrundgren, S. (February 1976). "Polymorphism in relation to habitat in the snail *Cepaea hortensis* in Iceland". *Journal of Zoology*. **178** (2): 173–188. [doi](/source/Doi_(identifier)):[10.1111/j.1469-7998.1976.tb06006.x](https://doi.org/10.1111%2Fj.1469-7998.1976.tb06006.x).

1. **[^](#cite_ref-8)** Valovirta, I.; Halkka, O. (12 February 2009). ["Colour polymorphism in northern peripheral populations of *Cepaea hortensis*"](https://doi.org/10.1111%2Fj.1601-5223.1976.tb01576.x). *Hereditas*. **83** (1): 123–126. [doi](/source/Doi_(identifier)):[10.1111/j.1601-5223.1976.tb01576.x](https://doi.org/10.1111%2Fj.1601-5223.1976.tb01576.x). [PMID](/source/PMID_(identifier)) [965238](https://pubmed.ncbi.nlm.nih.gov/965238).

1. **[^](#cite_ref-9)** Pearce, T.A.; Olori, J.C.; Kemezis, K.W. (10 August 2010). "Land snails from St. Elzear cave, Gaspé Peninsula, Quebec: antiquity of *Cepaea hortensis* in North America". *Annals of Carnegie Museum*. **79** (1): 65–78. [doi](/source/Doi_(identifier)):[10.2992/007.079.0105](https://doi.org/10.2992%2F007.079.0105).

1. **[^](#cite_ref-10)** 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](/source/ISBN_(identifier)) [2-88414-013-1](https://en.wikipedia.org/wiki/Special:BookSources/2-88414-013-1).

1. **[^](#cite_ref-11)** Cadevall, J.; Orozco, A. (2016). *Caracoles y babosas de la Península Ibérica y Baleares*. Barcelona: Omega. [ISBN](/source/ISBN_(identifier)) [978-84-282-1599-2](https://en.wikipedia.org/wiki/Special:BookSources/978-84-282-1599-2).

1. **[^](#cite_ref-12)** Stoch, F. ["Checklist of the Italian Fauna On-line version 2.1"](https://www.faunaitalia.it/checklist/index.html). *www.faunaitalia.it*. Ministry of Environment, Directorate-General for Nature and Sea Protection (PNM). Retrieved 22 April 2023.

1. **[^](#cite_ref-13)** Cowie, R.H.; Jones, J. S. (1987). "Ecological interactions between *Cepaea nemoralis* and *Cepaea hortensis*: competition, invasion but no niche displacement". *Functional Ecology*. **1** (2): 91–97. [doi](/source/Doi_(identifier)):[10.2307/2389710](https://doi.org/10.2307%2F2389710). [JSTOR](/source/JSTOR_(identifier)) [2389710](https://www.jstor.org/stable/2389710).

1. ^ [***a***](#cite_ref-Jones1977_14-0) [***b***](#cite_ref-Jones1977_14-1) [***c***](#cite_ref-Jones1977_14-2) [***d***](#cite_ref-Jones1977_14-3) [***e***](#cite_ref-Jones1977_14-4) [***f***](#cite_ref-Jones1977_14-5) Jones, J.S.; Leith, B.H.; Rawlings, P. (November 1977). "Polymorphism in *Cepaea*: a problem with too many solutions?". *Annual Review of Ecology and Systematics*. **8** (1): 109–143. [doi](/source/Doi_(identifier)):[10.1146/annurev.es.08.110177.000545](https://doi.org/10.1146%2Fannurev.es.08.110177.000545).

1. **[^](#cite_ref-15)** Lang, A. (1908). *Über die Bastarde von*Helix hortensis*Müller und*Helix nemoralis*L. Eine Untersuchung zur experimentellen Vererbungslehre*. Jena: Fischer.

1. **[^](#cite_ref-Davison2019_16-0)** 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*"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6629550). *Heredity*. **123** (2): 162–175. [doi](/source/Doi_(identifier)):[10.1038/s41437-019-0189-z](https://doi.org/10.1038%2Fs41437-019-0189-z). [PMC](/source/PMC_(identifier)) [6629550](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6629550). [PMID](/source/PMID_(identifier)) [30804571](https://pubmed.ncbi.nlm.nih.gov/30804571).

1. **[^](#cite_ref-Cain1988_17-0)** Cain, A.J. (1988). ["The scoring of polymorphic colour and pattern variation and its genetic basis in molluscan shells"](https://www.biodiversitylibrary.org/page/13030417). *Malacologia*. **28**: 1–15.

1. **[^](#cite_ref-18)** Cain, A.J. (1971). "Colour and banding morphs in subfossil samples of the snail *Cepaea*". In Creed, E.R. (ed.). *Ecological genetics and evolution*. Oxford: Blackwell. pp. 65–92.

1. **[^](#cite_ref-19)** Cameron, R.A.D. (22 May 1992). "Change and stability in *Cepaea* populations over 25 years: a case of climatic selection". *Proceedings of the Royal Society of London. Series B: Biological Sciences*. **248** (1322): 181–187. [doi](/source/Doi_(identifier)):[10.1098/rspb.1992.0060](https://doi.org/10.1098%2Frspb.1992.0060).

1. ^ [***a***](#cite_ref-Cook1998_20-0) [***b***](#cite_ref-Cook1998_20-1) [***c***](#cite_ref-Cook1998_20-2) Cook, L.M. (29 October 1998). ["A two–stage model for *Cepaea* polymorphism"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1692378). *Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences*. **353** (1375): 1577–1593. [doi](/source/Doi_(identifier)):[10.1098/rstb.1998.0311](https://doi.org/10.1098%2Frstb.1998.0311). [PMC](/source/PMC_(identifier)) [1692378](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1692378).

1. **[^](#cite_ref-21)** Richardson, A.M.M. (February 1974). "Differential climatic selection in natural population of land snail *Cepaea nemoralis*". *Nature*. **247** (5442): 572–573. [doi](/source/Doi_(identifier)):[10.1038/247572a0](https://doi.org/10.1038%2F247572a0).

1. **[^](#cite_ref-22)** Chang, H.-W. (October 1991). "Activity and weight loss in relation to solar radiation in the polymorphic land snail *Cepaea nemoralis*". *Journal of Zoology*. **225** (2): 213–225. [doi](/source/Doi_(identifier)):[10.1111/j.1469-7998.1991.tb03812.x](https://doi.org/10.1111%2Fj.1469-7998.1991.tb03812.x).

1. **[^](#cite_ref-23)** 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"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3083392). *PLOS ONE*. **6** (4) e18927. [doi](/source/Doi_(identifier)):[10.1371/journal.pone.0018927](https://doi.org/10.1371%2Fjournal.pone.0018927). [PMC](/source/PMC_(identifier)) [3083392](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3083392). [PMID](/source/PMID_(identifier)) [21556137](https://pubmed.ncbi.nlm.nih.gov/21556137).

1. **[^](#cite_ref-24)** Jones, J.S. (August 1982). "Genetic differences in individual behaviour associated with shell polymorphism in the snail *Cepaea nemoralis*". *Nature*. **298** (5876): 749–750. [doi](/source/Doi_(identifier)):[10.1038/298749a0](https://doi.org/10.1038%2F298749a0).

1. ^ [***a***](#cite_ref-Cameron2012_25-0) [***b***](#cite_ref-Cameron2012_25-1) Cameron, R.A.D.; Cook, L.M. (May 2012). ["Habitat and the shell polymorphism of *Cepaea nemoralis* (L.): interrogating the Evolution Megalab database"](https://doi.org/10.1093%2Fmollus%2Feyr052). *Journal of Molluscan Studies*. **78** (2): 179–184. [doi](/source/Doi_(identifier)):[10.1093/mollus/eyr052](https://doi.org/10.1093%2Fmollus%2Feyr052).

1. **[^](#cite_ref-26)** Sheppard, P.M. (April 1951). ["Fluctuations in the selective value of certain phenotypes in the polymorphic land snail *Cepæa nemoralis* (L.)"](https://doi.org/10.1038%2Fhdy.1951.8). *Heredity*. **5** (1): 125–134. [doi](/source/Doi_(identifier)):[10.1038/hdy.1951.8](https://doi.org/10.1038%2Fhdy.1951.8). [PMID](/source/PMID_(identifier)) [14840763](https://pubmed.ncbi.nlm.nih.gov/14840763).

1. **[^](#cite_ref-27)** Tucker, G.M. (June 1991). "Apostatic selection by song thrushes (*Turdus philomelos*) feeding on the snail *Cepaea hortensis*". *Biological Journal of the Linnean Society*. **43** (2): 149–156. [doi](/source/Doi_(identifier)):[10.1111/j.1095-8312.1991.tb00590.x](https://doi.org/10.1111%2Fj.1095-8312.1991.tb00590.x).

1. **[^](#cite_ref-28)** Rosin, Z.M.; Kobak, J.; Lesicki, A.; Tryjanowski, P. (September 2013). ["Differential shell strength of *Cepaea nemoralis* colour morphs—implications for their anti-predator defence"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3753478). *Naturwissenschaften*. **100** (9): 843–851. [doi](/source/Doi_(identifier)):[10.1007/s00114-013-1084-8](https://doi.org/10.1007%2Fs00114-013-1084-8). [PMC](/source/PMC_(identifier)) [3753478](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3753478). [PMID](/source/PMID_(identifier)) [23921905](https://pubmed.ncbi.nlm.nih.gov/23921905).

1. **[^](#cite_ref-29)** Ożgo, M. (February 2011). ["Rapid evolution in unstable habitats: a success story of the polymorphic land snail *Cepaea nemoralis* (Gastropoda: Pulmonata)"](https://doi.org/10.1111%2Fj.1095-8312.2010.01585.x). *Biological Journal of the Linnean Society*. **102** (2): 251–262. [doi](/source/Doi_(identifier)):[10.1111/j.1095-8312.2010.01585.x](https://doi.org/10.1111%2Fj.1095-8312.2010.01585.x).

1. **[^](#cite_ref-30)** Cameron, R.A.D.; Carter, M.A.; Palles-Clark, M.A. (November 1980). "*Cepaea* on Salisbury Plain: patterns of variation, landscape history and habitat stability". *Biological Journal of the Linnean Society*. **14** (3–4): 335–358. [doi](/source/Doi_(identifier)):[10.1111/j.1095-8312.1980.tb00113.x](https://doi.org/10.1111%2Fj.1095-8312.1980.tb00113.x).

1. **[^](#cite_ref-31)** Davison, A.; Clarke, B. (22 July 2000). ["History or current selection? A molecular analysis of 'area effects' in the land snail *Cepaea nemoralis*"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1690693). *Proceedings of the Royal Society of London. Series B: Biological Sciences*. **267** (1451): 1399–1405. [doi](/source/Doi_(identifier)):[10.1098/rspb.2000.1156](https://doi.org/10.1098%2Frspb.2000.1156). [PMC](/source/PMC_(identifier)) [1690693](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1690693). [PMID](/source/PMID_(identifier)) [10983823](https://pubmed.ncbi.nlm.nih.gov/10983823).

1. **[^](#cite_ref-32)** Cameron, R.A.D.; Dillon, P.J. (1984). ["Habitat stability, population histories and patterns of variation in *Cepaea*"](https://www.biodiversitylibrary.org/page/13145394). *Malacologia*. **25**: 271–291.

## Further reading

Wikimedia Commons has media related to [Cepaea](https://commons.wikimedia.org/wiki/Category:Cepaea).

- Yamazaki N.; Ueshima R.; Terrett J. A.; Yokobori S.; Kaifu M.; Segawa R.; Kobayashi T.; Numachi K.; Ueda T.; Nishikawa K.; Watanabe K.; Thomas R. H. (1997). ["Evolution of pulmonate gastropod mitochondrial genomes: comparisons of complete gene organizations of *Euhadra*, *Cepaea* and *Albinaria* and implications of unusual tRNA secondary structures"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1207859). *Genetics*. **145** (3): 749–758. [doi](/source/Doi_(identifier)):[10.1093/genetics/145.3.749](https://doi.org/10.1093%2Fgenetics%2F145.3.749). [PMC](/source/PMC_(identifier)) [1207859](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1207859). [PMID](/source/PMID_(identifier)) [9055084](https://pubmed.ncbi.nlm.nih.gov/9055084).

Taxon identifiers Cepaea Wikidata: Q747535 Wikispecies: Cepaea ADW: Cepaea BioLib: 2974 BOLD: 6001 CoL: 7NS7V EoL: 57692 EPPO: 1CEPAG Fauna Europaea: 426292 Fauna Europaea (new): fae4fba9-d099-40a2-8fec-6ca9545fb731 GBIF: 2294278 iNaturalist: 48587 IRMNG: 1354834 ITIS: 77908 NBN: NHMSYS0001701946 NCBI: 28834 Paleobiology Database: 114488 Plazi: D047E409-B02A-131D-FF02-0416FE90BFF2 WoRMS: 235791

---
Adapted from the Wikipedia article [Cepaea](https://en.wikipedia.org/wiki/Cepaea) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Cepaea?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.