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Helicidae
	Helix pomatia
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Mollusca
Class:	Gastropoda
Order:	Stylommatophora
Superfamily:	Helicoidea
Family:	Helicidae; Rafinesque, 1815
Type genus
	Helix Linnaeus, 1758;
Subfamilies
	Ariantinae; Murellinae; Helicinae;

Helicidae is a large, diverse family of western Palaearctic, medium to large-sized, air-breathing land snails, sometimes called the "typical snails." It includes some of the largest European land snails, several species are common in anthropogenic habitats, and some became invasive on other continents. A number of species in this family are valued as food items, including Cornu aspersum (formerly Helix aspersa; "petit gris") the brown or garden snail, and Helix pomatia (the "escargot").^[1] The biologies of these two species in particular have been thoroughly studied and documented.

Shell description

The shells are usually flattened or depressed conical. Globular shells are found in the genera Helix, Maltzanella, Lindholmia, Cornu, Cantareus, Eremina, and Idiomella. One species, Cylindrus obtusus, has a cylindrical shell. In some genera, especially in Cepaea, the shells are brightly colored and patterned.

Anatomy

Helicidae typically have a ribbed jaw, bursa copulatrix with a diverticulum, and one dart sac accompanied by a pair of (usually) branched, tubular mucous glands inserting at the base of the dart sac.^[2]

Genetics

In this family, the number of haploid chromosomes lies between 22 and 30.^[3]

In the "Darwin Tree of Life"." project, four species (Cepaea nemoralis, Cepaea hortensis, Cornu aspersum, and Arianta arbustorum) are scheduled for whole genome sequencing and assembly ("Data portal".).

Helicid snails for sale as food in Italy; from the front *Eobania vermiculata*, *Cantareus apertus*, and *Helix* sp.

Distribution

The core of helicids is distributed in from the Caucasus through Turkey and Europe to North Africa. However, some genera or species live beyond these limits. Helicids occur on Cape Verde^[2] (Eremina), Canary Islands (Theba, Hemicycla) and the Madeira Archipelago (Lampadia, Idiomela).^[4] Levantina extends far south in western Arabia, and Eremina desertella is distributed as south as Sudan, Eritrea and Puntland in Somalia.^[5]^[6] Cepaea hortensis lives on Iceland^[7] and in a small area in eastern Canada.^[8] Some species, notably Cornu aspersum and Theba pisana have been introduced and become established in numerous different areas worldwide.

Taxonomy

The family Helicidae contains 3 subfamilies (according to molecular phylogenetic analyses^[9]^[10]^[11]^[12]):

Subfamily Helicinae Rafinesque, 1815

Genital system anatomy (does not apply on all species, as derived states are found in some of them): mucous glands divided into 2 or more branches, love dart with four blades (vanes) along its length, two penial papillae/verges.^{[citation needed]}

Tribe Allognathini Westerlund, 1903^[4]

Allognathus
Cepaea Held, 1838
Hemicycla
Iberus
Idiomela T. Cockerell, 1921
Lampadia

Tribe Helicini Rafinesque, 1815^[13]

Aristena Psonis, Vardinoyannis & Poulakakis, 2022
Amanica Nordsieck, 2017
Caucasotachea Boettger, 1909
Codringtonia Kobelt, 1898
Helix Linnaeus, 1758 - type genus
Isaurica Kobelt, 1901
Levantina Kobelt, 1871
Lindholmia Hesse, 1918
Maltzanella Hesse, 1917
Neocrassa Subai, 2005

Tribe Thebini Wenz, 1923^[14]^[12]

A 2022 phylogenetic analysis proposed that all groups of the Maghreb radiation belonged to a single tribe, Thebini, without support for a separate Otalini tribe. The same study proposed a new tribe, Maculariini trib. nov. containing the genus Macularia due to the wide geographic disjunction between the western Alpine Macularia and the primarily Maghrebian Thebini tribe.^[14]

Cantareus Risso, 1826
Cornu Born, 1778
Eobania P. Hesse, 1913
Eremina Pfeiffer, 1855
Gyrostomella P. Hesse, 1911
Loxana Pallary, 1899
Massylaea Möllendorff, 1898
Otala Schumacher, 1817
Rossmaessleria P. Hesse, 1907
Theba Risso, 1826

Tribe Maculariini Neiber, Korábek, Glaubrecht & Hausdorf, 2021

Macularia Albers, 1850

Subfamily Murellinae Hesse, 1918^[10]

Genital system anatomy (does not apply on all species, as derived states are found in some of them): mucous glands weakly branched or undivided, love dart with four blades along its length, one penial papilla.^[15]

Distributed in Sardinia, Corsica, the Apennine Peninsula and Sicily.^[15]^[16]

Marmorana W. Hartmann, 1844
Tacheocampylaea
Tyrrheniberus

Subfamily Ariantinae Mörch, 1864^[17]

Genital system anatomy (does not apply on all species, as derived states are found in some of them): mucous glands divided into 2 branches or undivided, love dart with two blades on the tip, one penial papilla.^{[citation needed]}

Arianta Turton, 1831
Campylaea H. Beck, 1837
Campylaeopsis A.J. Wagner, 1914
Cattania Brusina, 1904
Causa Schileyko, 1971
Chilostoma Fitzinger, 1833
Corneola Held, 1838
Cylindrus Fitzinger, 1833
Delphinatia P. Hesse, 1931
Dinarica Kobelt, 1902
Drobacia Brusina, 1904
Faustina Kobelt, 1904
Helicigona A. Férussac, 1821
Isognomostoma Fitzinger, 1833
Josephinella F. Haas, 1936
Kollarix Groenenberg, Subai & E. Gittenberger, 2016
Kosicia Brusina, 1904
Liburnica Kobelt, 1904
Pseudotrizona Groenenberg, Subai & E. Gittenberger, 2016
Thiessea Kobelt, 1904
Vidovicia Brusina, 1904
†Pseudochloritis C. R. Boettger, 1909^[18]
†Mesodontopsis Pilsbry, 1895
†Metacampylaea Pilsbry, 1895
†Paradrobacia H. Nordsieck, 2014
†Pseudoklikia H. Nordsieck, 2018

Incertae sedis

†Megalotachea Pfeffer, 1930

References

^ M.P. Kerney & R.A.D. Cameron. 1979. A field guide to the land snails of Britain and northwestern Europe. Collins, London.
^ ^a ^b Nordsieck, Hartmut (1987). "Revision des Systems der Helicoidea (Gastropoda: Stylommatophora)". Archiv für Molluskenkunde. 118: 9–50.
^ Thiriot-QuiéVreux, Catherine (2003). "Advances in Chromosomal Studies of Gastropod Molluscs". Journal of Molluscan Studies. 69 (3): 187–202. doi:10.1093/mollus/69.3.187. ISSN 1464-3766.
^ ^a ^b Neiber, Marco T.; Chueca, Luis J.; Caro, Amaia; Teixeira, Dinarte; Schlegel, Kevin A.; Gómez-Moliner, Benjamín J.; Walther, Frank; Glaubrecht, Matthias; Hausdorf, Bernhard (2021). "Incorporating palaeogeography into ancestral area estimation can explain the disjunct distribution of land snails in Macaronesia and the Balearic Islands (Helicidae: Allognathini)". Molecular Phylogenetics and Evolution. 162: 107196. doi:10.1016/j.ympev.2021.107196. PMID 33965538.
^ Jickeli, Carl F. (1874). Fauna der Land- und Süsswasser-Mollusken Nord-Ost-Afrika's. Dresden: E. Blochmann & Sohn.
^ Neubert, Eike (1998). "Annotated checklist of the terrestrial and freshwater molluscs of the Arabian Peninsula with descriptions of new species". Fauna of Arabia. 17: 333–461.
^ Arnason, Einar; Grant, P. R. (1976). "Climatic Selection in Cepaea hortensis at the Northern Limit of Its Range in Iceland". Evolution. 30 (3): 499–508. doi:10.2307/2407574. JSTOR 2407574. PMID 28563245.
^ Pearce, Timothy A.; Olori, Jennifer C.; Kemezis, Kathleen W. (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:10.2992/007.079.0105. ISSN 0097-4463. S2CID 84205240.
^ Razkin, Oihana; Gómez-Moliner, Benjamín Juán; Prieto, Carlos Enrique; Martínez-Ortí, Alberto; Arrébola, José Ramón; Muñoz, Benito; Chueca, Luis Javier; Madeira, María José (2015-02-01). "Molecular phylogeny of the western Palaearctic Helicoidea (Gastropoda, Stylommatophora)". Molecular Phylogenetics and Evolution. 83: 99–117. doi:10.1016/j.ympev.2014.11.014. PMID 25485783.
^ ^a ^b Neiber, Marco T.; Hausdorf, Bernhard (2015-12-01). "Molecular phylogeny reveals the polyphyly of the snail genus Cepaea (Gastropoda: Helicidae)". Molecular Phylogenetics and Evolution. 93: 143–149. doi:10.1016/j.ympev.2015.07.022. PMID 26256642.
^ Neiber, Marco T; Korábek, Ondřej; Glaubrecht, Matthias; Hausdorf, Bernhard (2022-04-11). "A misinterpreted disjunction: the phylogenetic relationships of the North African land snail Gyrostomella (Gastropoda: Stylommatophora: Helicidae)". Zoological Journal of the Linnean Society. 194 (4): 1236–1251. doi:10.1093/zoolinnean/zlab059. ISSN 0024-4082.
^ ^a ^b Holyoak, David T.; Holyoak, Geraldine A.; Gómez Moliner, Benjamín M.; Chueca, Luis J. (2020). "Phylogeny, species-limits and taxonomic revision of the Otalini (Helicidae) from north-west Africa". Journal of Conchology. 43: 551–611.
^ Korábek, Ondřej; Juřičková, Lucie; Petrusek, Adam (2021-12-31). "Diversity of Land Snail Tribe Helicini (Gastropoda: Stylommatophora: Helicidae): Where Do We Stand after 20 Years of Sequencing Mitochondrial Markers?". Diversity. 14 (1): 24. doi:10.3390/d14010024. ISSN 1424-2818.
^ ^a ^b Marco T Neiber; Ondřej Korábek; Matthias Glaubrecht; Bernhard Hausdorf (2022). "A misinterpreted disjunction: the phylogenetic relationships of the North African land snail Gyrostomella (Gastropoda: Stylommatophora: Helicidae)". Zoological Journal of the Linnean Society. 194 (4): 1236–1251. doi:10.1093/zoolinnean/zlab059.
^ ^a ^b Hesse, Paul (1908). Iconographie der Land- & Süsswasser-Mollusken mit vorzüglicher Berücksichtigung der Europäischen noch nicht abgebildeten Arten. Neu Folge. Vierzehnter Band. Wiesbaden: C. W. Kreidel.
^ Fiorentino, Viviana; Salomone, Nicola; Manganelli, Giuseppe; Giusti, Folco (2010). "Historical biogeography of Tyrrhenian land snails: The Marmorana–Tyrrheniberus radiation (Pulmonata, Helicidae)". Molecular Phylogenetics and Evolution. 55 (1): 26–37. doi:10.1016/j.ympev.2009.11.024. PMID 19995610.
^ Groenenberg, Dick S.J.; Subai, Peter; Gittenberger, Edmund (2016-01-28). "Systematics of Ariantinae (Gastropoda, Pulmonata, Helicidae), a new approach to an old problem". Contributions to Zoology. 85 (1): 37–65l. doi:10.1163/18759866-08501003. ISSN 1383-4517.
^ Höltke, Olaf; Rasser, Michael W (2015). "Pseudochloritis insignis–a peculiar large land-snail from the Miocene of SW Germany: taxonomic status and census of morphologically related forms". Journal of Conchology. 42 (1): 1.

External links

[1] M.P. Kerney & R.A.D. Cameron. 1979. A field guide to the land snails of Britain and northwestern Europe. Collins, London.

[:0-2] Nordsieck, Hartmut (1987). "Revision des Systems der Helicoidea (Gastropoda: Stylommatophora)". Archiv für Molluskenkunde. 118: 9–50.

[3] Thiriot-QuiéVreux, Catherine (2003). "Advances in Chromosomal Studies of Gastropod Molluscs". Journal of Molluscan Studies. 69 (3): 187–202. doi:10.1093/mollus/69.3.187. ISSN 1464-3766.

[:1-4] Neiber, Marco T.; Chueca, Luis J.; Caro, Amaia; Teixeira, Dinarte; Schlegel, Kevin A.; Gómez-Moliner, Benjamín J.; Walther, Frank; Glaubrecht, Matthias; Hausdorf, Bernhard (2021). "Incorporating palaeogeography into ancestral area estimation can explain the disjunct distribution of land snails in Macaronesia and the Balearic Islands (Helicidae: Allognathini)". Molecular Phylogenetics and Evolution. 162: 107196. doi:10.1016/j.ympev.2021.107196. PMID 33965538.

[5] Jickeli, Carl F. (1874). Fauna der Land- und Süsswasser-Mollusken Nord-Ost-Afrika's. Dresden: E. Blochmann & Sohn.

[6] Neubert, Eike (1998). "Annotated checklist of the terrestrial and freshwater molluscs of the Arabian Peninsula with descriptions of new species". Fauna of Arabia. 17: 333–461.

[7] Arnason, Einar; Grant, P. R. (1976). "Climatic Selection in Cepaea hortensis at the Northern Limit of Its Range in Iceland". Evolution. 30 (3): 499–508. doi:10.2307/2407574. JSTOR 2407574. PMID 28563245.

[8] Pearce, Timothy A.; Olori, Jennifer C.; Kemezis, Kathleen W. (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:10.2992/007.079.0105. ISSN 0097-4463. S2CID 84205240.

[9] Razkin, Oihana; Gómez-Moliner, Benjamín Juán; Prieto, Carlos Enrique; Martínez-Ortí, Alberto; Arrébola, José Ramón; Muñoz, Benito; Chueca, Luis Javier; Madeira, María José (2015-02-01). "Molecular phylogeny of the western Palaearctic Helicoidea (Gastropoda, Stylommatophora)". Molecular Phylogenetics and Evolution. 83: 99–117. doi:10.1016/j.ympev.2014.11.014. PMID 25485783.

[:3-10] Neiber, Marco T.; Hausdorf, Bernhard (2015-12-01). "Molecular phylogeny reveals the polyphyly of the snail genus Cepaea (Gastropoda: Helicidae)". Molecular Phylogenetics and Evolution. 93: 143–149. doi:10.1016/j.ympev.2015.07.022. PMID 26256642.

[11] Neiber, Marco T; Korábek, Ondřej; Glaubrecht, Matthias; Hausdorf, Bernhard (2022-04-11). "A misinterpreted disjunction: the phylogenetic relationships of the North African land snail Gyrostomella (Gastropoda: Stylommatophora: Helicidae)". Zoological Journal of the Linnean Society. 194 (4): 1236–1251. doi:10.1093/zoolinnean/zlab059. ISSN 0024-4082.

[:2-12] Holyoak, David T.; Holyoak, Geraldine A.; Gómez Moliner, Benjamín M.; Chueca, Luis J. (2020). "Phylogeny, species-limits and taxonomic revision of the Otalini (Helicidae) from north-west Africa". Journal of Conchology. 43: 551–611.

[13] Korábek, Ondřej; Juřičková, Lucie; Petrusek, Adam (2021-12-31). "Diversity of Land Snail Tribe Helicini (Gastropoda: Stylommatophora: Helicidae): Where Do We Stand after 20 Years of Sequencing Mitochondrial Markers?". Diversity. 14 (1): 24. doi:10.3390/d14010024. ISSN 1424-2818.

[Neiber-14] Marco T Neiber; Ondřej Korábek; Matthias Glaubrecht; Bernhard Hausdorf (2022). "A misinterpreted disjunction: the phylogenetic relationships of the North African land snail Gyrostomella (Gastropoda: Stylommatophora: Helicidae)". Zoological Journal of the Linnean Society. 194 (4): 1236–1251. doi:10.1093/zoolinnean/zlab059.

[:4-15] Hesse, Paul (1908). Iconographie der Land- & Süsswasser-Mollusken mit vorzüglicher Berücksichtigung der Europäischen noch nicht abgebildeten Arten. Neu Folge. Vierzehnter Band. Wiesbaden: C. W. Kreidel.

[16] Fiorentino, Viviana; Salomone, Nicola; Manganelli, Giuseppe; Giusti, Folco (2010). "Historical biogeography of Tyrrhenian land snails: The Marmorana–Tyrrheniberus radiation (Pulmonata, Helicidae)". Molecular Phylogenetics and Evolution. 55 (1): 26–37. doi:10.1016/j.ympev.2009.11.024. PMID 19995610.

[17] Groenenberg, Dick S.J.; Subai, Peter; Gittenberger, Edmund (2016-01-28). "Systematics of Ariantinae (Gastropoda, Pulmonata, Helicidae), a new approach to an old problem". Contributions to Zoology. 85 (1): 37–65l. doi:10.1163/18759866-08501003. ISSN 1383-4517.

[hotrasser-18] Höltke, Olaf; Rasser, Michael W (2015). "Pseudochloritis insignis–a peculiar large land-snail from the Miocene of SW Germany: taxonomic status and census of morphologically related forms". Journal of Conchology. 42 (1): 1.

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Helicidae

Shell description

Anatomy

Genetics

Distribution