Helicidae
{{Short description|Family of gastropods}}
{{Automatic taxobox
| image = Helix pomatia 89a.jpg
| image_caption = Helix pomatia
| taxon = Helicidae
| authority = Rafinesque, 1815
| subdivision_ranks = Subfamilies
| subdivision = *Ariantinae
| type_genus = Helix Linnaeus, 1758
}}
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").M.P. Kerney & R.A.D. Cameron. 1979. A field guide to the land snails of Britain and northwestern Europe. Collins, London. The biologies of these two species in particular have been thoroughly studied and documented.
Shell description
File:Caucasotachea_atrolabiata.jpg]]
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.{{Cite journal |last=Nordsieck |first=Hartmut |date=1987 |title=Revision des Systems der Helicoidea (Gastropoda: Stylommatophora) |journal=Archiv für Molluskenkunde |volume=118 |pages=9–50}}
Genetics
In this family, the number of haploid chromosomes lies between 22 and 30.{{Cite journal |last=Thiriot-QuiéVreux |first=Catherine |title=Advances in Chromosomal Studies of Gastropod Molluscs |date=2003 |journal=Journal of Molluscan Studies |language=en |volume=69 |issue=3 |pages=187–202 |doi=10.1093/mollus/69.3.187 |issn=1464-3766|doi-access=free }}
In the {{Cite web |title=Darwin Tree of Life |url=https://www.darwintreeoflife.org/}}" project, four species (Cepaea nemoralis, Cepaea hortensis, Cornu aspersum, and Arianta arbustorum) are scheduled for whole genome sequencing and assembly ({{Cite web |title=Data portal |url=https://portal.darwintreeoflife.org/data}}).
Image:Snails-Italy.jpg; 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 (Eremina), Canary Islands (Theba, Hemicycla) and the Madeira Archipelago (Lampadia, Idiomela). Levantina extends far south in western Arabia, and Eremina desertella is distributed as south as Sudan, Eritrea and Puntland in Somalia.{{Cite book |last=Jickeli |first=Carl F. |url=https://www.biodiversitylibrary.org/item/46618#page/7/mode/1up |title=Fauna der Land- und Süsswasser-Mollusken Nord-Ost-Afrika's |publisher=E. Blochmann & Sohn |year=1874 |location=Dresden}}{{Cite journal |last=Neubert |first=Eike |date=1998 |title=Annotated checklist of the terrestrial and freshwater molluscs of the Arabian Peninsula with descriptions of new species |journal=Fauna of Arabia |volume=17 |pages=333–461}} Cepaea hortensis lives on Iceland{{Cite journal |last1=Arnason |first1=Einar |last2=Grant |first2=P. R. |date=1976 |title=Climatic Selection in Cepaea hortensis at the Northern Limit of Its Range in Iceland |url=https://www.jstor.org/stable/2407574 |journal=Evolution |volume=30 |issue=3 |pages=499–508 |doi=10.2307/2407574|jstor=2407574 |pmid=28563245 }} and in a small area in eastern Canada.{{Cite journal |last1=Pearce |first1=Timothy A. |last2=Olori |first2=Jennifer C. |last3=Kemezis |first3=Kathleen W. |date=2010 |title=Land Snails from St. Elzear Cave, Gaspé Peninsula, Quebec: Antiquity of Cepaea Hortensis in North America |url=http://www.bioone.org/doi/abs/10.2992/007.079.0105 |journal=Annals of Carnegie Museum |language=en |volume=79 |issue=1 |pages=65–78 |doi=10.2992/007.079.0105 |s2cid=84205240 |issn=0097-4463}} Some species, notably Cornu aspersum and Theba pisana have been introduced and become established in numerous different areas worldwide.
Taxonomy
Image:Snail black on grass2.jpg]]
Image:Iberus gualterianus alonensis3.JPG alonensis]]
File:Snail-wiki-120-Zachi-Evenor.jpg]]
The family Helicidae contains 3 subfamilies (according to molecular phylogenetic analyses{{Cite journal|last1=Razkin|first1=Oihana|last2=Gómez-Moliner|first2=Benjamín Juán|last3=Prieto|first3=Carlos Enrique|last4=Martínez-Ortí|first4=Alberto|last5=Arrébola|first5=José Ramón|last6=Muñoz|first6=Benito|last7=Chueca|first7=Luis Javier|last8=Madeira|first8=María José|date=2015-02-01|title=Molecular phylogeny of the western Palaearctic Helicoidea (Gastropoda, Stylommatophora)|journal=Molecular Phylogenetics and Evolution|volume=83|pages=99–117|doi=10.1016/j.ympev.2014.11.014|pmid=25485783}}{{Cite journal|last1=Neiber|first1=Marco T.|last2=Hausdorf|first2=Bernhard|date=2015-12-01|title=Molecular phylogeny reveals the polyphyly of the snail genus Cepaea (Gastropoda: Helicidae)|journal=Molecular Phylogenetics and Evolution|volume=93|pages=143–149|doi=10.1016/j.ympev.2015.07.022|pmid=26256642}}{{Cite journal |last1=Neiber |first1=Marco T |last2=Korábek |first2=Ondřej |last3=Glaubrecht |first3=Matthias |last4=Hausdorf |first4=Bernhard |date=2022-04-11 |title=A misinterpreted disjunction: the phylogenetic relationships of the North African land snail Gyrostomella (Gastropoda: Stylommatophora: Helicidae) |url=https://academic.oup.com/zoolinnean/article/194/4/1236/6445973 |journal=Zoological Journal of the Linnean Society |language=en |volume=194 |issue=4 |pages=1236–1251 |doi=10.1093/zoolinnean/zlab059 |issn=0024-4082}}{{Cite journal |last1=Holyoak |first1=David T. |last2=Holyoak |first2=Geraldine A. |last3=Gómez Moliner |first3=Benjamín M. |last4=Chueca |first4=Luis J. |date=2020 |title=Phylogeny, species-limits and taxonomic revision of the Otalini (Helicidae) from north-west Africa |journal=Journal of Conchology |volume=43 |pages=551–611}}):
= 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|date=July 2023}}
== Tribe Allognathini Westerlund, 1903<ref name=":1">{{Cite journal |last1=Neiber |first1=Marco T. |last2=Chueca |first2=Luis J. |last3=Caro |first3=Amaia |last4=Teixeira |first4=Dinarte |last5=Schlegel |first5=Kevin A. |last6=Gómez-Moliner |first6=Benjamín J. |last7=Walther |first7=Frank |last8=Glaubrecht |first8=Matthias |last9=Hausdorf |first9=Bernhard |date=2021 |title=Incorporating palaeogeography into ancestral area estimation can explain the disjunct distribution of land snails in Macaronesia and the Balearic Islands (Helicidae: Allognathini) |url=https://linkinghub.elsevier.com/retrieve/pii/S1055790321001299 |journal=[[Molecular Phylogenetics and Evolution]] |language=en |volume=162 |pages=107196 |doi=10.1016/j.ympev.2021.107196|pmid=33965538 }}</ref> ==
== Tribe [[Helicini]] [[Rafinesque]], 1815<ref>{{Cite journal |last1=Korábek |first1=Ondřej |last2=Juřičková |first2=Lucie |last3=Petrusek |first3=Adam |date=2021-12-31 |title=Diversity of Land Snail Tribe Helicini (Gastropoda: Stylommatophora: Helicidae): Where Do We Stand after 20 Years of Sequencing Mitochondrial Markers? |journal=Diversity |language=en |volume=14 |issue=1 |pages=24 |doi=10.3390/d14010024 |issn=1424-2818 |doi-access=free }}</ref> ==
- 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<ref name="Neiber">{{cite journal |author1=Marco T Neiber |author2=Ondřej Korábek |author3=Matthias Glaubrecht |author4=Bernhard Hausdorf |year=2022 |title=A misinterpreted disjunction: the phylogenetic relationships of the North African land snail ''Gyrostomella'' (Gastropoda: Stylommatophora: Helicidae) |url=https://doi.org/10.1093/zoolinnean/zlab059 |journal=Zoological Journal of the Linnean Society |volume=194 |issue=4 |pages=1236–1251 |doi=10.1093/zoolinnean/zlab059}}</ref><ref name=":2" /> ==
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.
== Tribe Maculariini Neiber, Korábek, Glaubrecht & Hausdorf, 2021 ==
- Macularia Albers, 1850
= Subfamily Murellinae Hesse, 1918<ref name=":3" /> =
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.{{Cite book |last=Hesse |first=Paul |url=https://www.biodiversitylibrary.org/item/55071#page/287/mode/1up |title=Iconographie der Land- & Süsswasser-Mollusken mit vorzüglicher Berücksichtigung der Europäischen noch nicht abgebildeten Arten. Neu Folge. Vierzehnter Band. |publisher=C. W. Kreidel |year=1908 |location=Wiesbaden}}
Distributed in Sardinia, Corsica, the Apennine Peninsula and Sicily.{{Cite journal |last1=Fiorentino |first1=Viviana |last2=Salomone |first2=Nicola |last3=Manganelli |first3=Giuseppe |last4=Giusti |first4=Folco |date=2010 |title=Historical biogeography of Tyrrhenian land snails: The Marmorana–Tyrrheniberus radiation (Pulmonata, Helicidae) |url=https://linkinghub.elsevier.com/retrieve/pii/S105579030900493X |journal=Molecular Phylogenetics and Evolution |language=en |volume=55 |issue=1 |pages=26–37 |doi=10.1016/j.ympev.2009.11.024|pmid=19995610 }}
- Marmorana W. Hartmann, 1844
- Tacheocampylaea
- Tyrrheniberus
= Subfamily Ariantinae Mörch, 1864<ref>{{Cite journal |last1=Groenenberg |first1=Dick S.J. |last2=Subai |first2=Peter |last3=Gittenberger |first3=Edmund |date=2016-01-28 |editor-first= |title=Systematics of Ariantinae (Gastropoda, Pulmonata, Helicidae), a new approach to an old problem |journal=Contributions to Zoology |volume=85 |issue=1 |pages=37–65l |doi=10.1163/18759866-08501003 |issn=1383-4517|doi-access=free }}</ref> =
Genital system anatomy: mucous glands divided into 2 branches or undivided, love dart with two blades on the tip, one penial papilla.{{Cite journal |last=Nordsieck |first=Hartmut |date=1987 |title=Revision des Systems der Helicoidea (Gastropoda: Stylommatophora) |journal=Archiv für Molluskenkunde |volume=118 |pages=9–50}}
- 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
- {{extinct}}Pseudochloritis C. R. Boettger, 1909{{cite journal | last1 = Höltke | first1 = Olaf | last2 = Rasser | first2 = Michael W | year = 2015 | title = Pseudochloritis insignis–a peculiar large land-snail from the Miocene of SW Germany: taxonomic status and census of morphologically related forms | journal = Journal of Conchology | volume = 42 | issue = 1| page = 1 }}
- {{extinct}}Mesodontopsis Pilsbry, 1895
- {{extinct}}Metacampylaea Pilsbry, 1895
- {{extinct}}Paradrobacia H. Nordsieck, 2014
- {{extinct}}Pseudoklikia H. Nordsieck, 2018
= ''Incertae sedis'' =
- †Megalotachea Pfeffer, 1930
References
{{Reflist}}
External links
{{Commons category|Helicidae}}
{{Taxonbar|from=Q1418257}}
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