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| type_genus = [[Helix (gastropod)|Helix]] <small>[[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]</small>
| type_genus = [[Helix (gastropod)|Helix]] <small>[[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]</small>
}}
}}
'''Helicidae''' is a large, diverse [[family (biology)|family]] of small to large, air-breathing [[land snail]]s, sometimes called the "typical snails."


A number of species in this family are valued as food items, including ''[[Cornu aspersum]]'' (formerly ''Helix aspersa'') the brown or garden snail, and ''[[Helix pomatia]]'' the "[[escargot]]".<ref>M.P. Kerney & R.A.D. Cameron. 1979. ''A field guide to the land snails of Britain and northwestern Europe''. Collins, London.</ref> The biologies of these two species in particular have been thoroughly studied and documented.
'''Helicidae''' is a large, diverse [[family (biology)|family]] of western [[Palearctic realm|Palaearctic]], medium to large-sized, air-breathing [[land snail]]s, 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]]").<ref>M.P. Kerney & R.A.D. Cameron. 1979. ''A field guide to the land snails of Britain and northwestern Europe''. Collins, London.</ref> The biologies of these two species in particular have been thoroughly studied and documented.


== Shell description ==
== Shell description ==
The [[gastropod shell|shell]]s of these snails are mostly rather globular in shape. In some genera, such as ''[[Cepaea]]'', the shells are brightly colored and patterned.
The [[Mollusc shell|shells]] are usually flattened or depressed conical. Globular shells are found in the genera ''[[Helix (gastropod)|Helix]]'', ''[[Maltzanella]]'', ''[[Lindholmia]]'', ''[[Cornu aspersum|Cornu]]'', ''[[Cantareus]]'', ''[[Eremina vermiculosa|Eremina]]'', and ''[[Idiomela subplicata|Idiomella]]''. One species, ''[[Cylindrus obtusus]]'', has a cylindrical shell. In some genera, especially in ''[[Cepaea]]'', the shells are brightly colored and patterned.


== Anatomy ==
== 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.<ref name=":0">{{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}}</ref>
The animal is capable of complete retraction within the shell. The tail is without a mucous gland or projection.<ref name="manual 3">[[George Washington Tryon|Tryon G. W.]] 1887 ''Manual of conchology; structural and systematic. With illustrations of the species''. Second series: Pulmonata. [https://archive.org/details/manualconch03tryorich Volume 3]. Helicidae - Volume I. page [https://archive.org/stream/manualconch03tryorich#page/n7/mode/2up 3]–4.</ref>{{Unreliable source?|date=December 2008}}


== Genetics ==
The mouth is always provided with a jaw, which is striate, ribbed, sulcate or plicate, sometimes composed of several imbricating pieces. The [[radula]] is composed of many transverse horizontal series of teeth, the centrals tricuspidate, about the size of the laterals, laterals bicuspid, or tricuspid with the interior cusp obsolete, marginals usually wider than high, short with two or three small cusps.<ref name="manual 3"/>{{Unreliable source?|date=December 2008}}
In this family, the number of [[haploid]] [[chromosome]]s lies between 22 and 30.<ref>{{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 }}</ref>


In the {{Cite web |title=Darwin Tree of Life |url=https://www.darwintreeoflife.org/}}" project, four species (''[[Cepaea nemoralis]]'', [[White-lipped snail|''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}}).
In the soft parts the most obvious distinction is the lack of a [[caudal mucous pit]], and their possessing a sculptured jaw. Typically, there is a distinction in the dentition, although in some species the lateral teeth take on a pseudozonitoid appearance; even in such cases the extreme marginals in ''Helix'' remain short and very obtuse.<ref name="manual 3"/>{{Unreliable source?|date=December 2008}}

This family of snails is defined by the anatomical presence of a [[diverticulum]]. Some genera within this family create and use [[love dart]]s as part of their mating behavior. These snails have one dart apparatus. The dart sac has no accessory sac, and contains two tubular glands, inserted at the base of the dart sac.

In this family, the number of [[haploid]] [[chromosome]]s lies between 21 and 30 (according to the values in this table).<ref name="Barker">Barker G. M.: Gastropods on Land: ''Phylogeny, Diversity and Adaptive Morphology''. in Barker G. M. (ed.): ''[https://books.google.com/books?id=WlvX-9Wt0toC&hl The biology of terrestrial molluscs]''. CABI Publishing, Oxon, UK, 2001, {{ISBN|0-85199-318-4}}. 1-146, cited pages: 139 and 142.</ref>


[[Image:Snails-Italy.jpg|thumb|Helicid snails for sale as food in [[Italy]]; from the front ''[[Eobania vermiculata]]'', ''[[Cantareus apertus]]'', and ''Helix'' sp.]]
[[Image:Snails-Italy.jpg|thumb|Helicid snails for sale as food in [[Italy]]; from the front ''[[Eobania vermiculata]]'', ''[[Cantareus apertus]]'', and ''Helix'' sp.]]


== Distribution ==
== Distribution ==
The core of helicids is distributed in from the [[Caucasus Mountains|Caucasus]] through [[Turkey]] and [[Europe]] to [[North Africa]]. However, some genera or species live beyond these limits. Helicids occur on [[Cape Verde]]<ref name=":0" /> (''[[Eremina]]''), Canary Islands (''[[Theba]]'', ''[[Hemicycla]]'') and the [[Madeira Archipelago]] (''[[Lampadia]]'', ''[[Idiomela subplicata|Idiomela]]'').<ref name=":1" /> ''[[Levantina]]'' extends far south in western [[Arabian Peninsula|Arabia]], and [[Eremina|''Eremina desertella'']] is distributed as south as [[Sudan]], [[Eritrea]] and [[Puntland]] in [[Somalia]].<ref>{{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}}</ref><ref>{{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}}</ref> ''[[White-lipped snail|Cepaea hortensis]]'' lives on [[Iceland]]<ref>{{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 (journal)|Evolution]] |volume=30 |issue=3 |pages=499–508 |doi=10.2307/2407574|jstor=2407574 |pmid=28563245 }}</ref> and in a small area in eastern [[Canada]].<ref>{{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}}</ref> Some species, notably ''[[Cornu aspersum]]'' and ''[[Theba pisana]]'' have been introduced and become established in numerous different areas worldwide.
The family is native to [[Eurasia]] and belongs to the Western [[Palearctic]] group.

However some species, notably ''[[Cornu aspersum]]'', have been introduced and become established in numerous different areas worldwide.

Many accidental introductions happen because the eggs of most Helicidae are laid in the soil, and they may thus easily travel unnoticed along with landscaping plants.


== Genera ==
== Taxonomy ==
[[Image:Snail black on grass2.jpg|thumb|''[[Arianta arbustorum]]'']]
[[Image:Snail black on grass2.jpg|thumb|''[[Arianta arbustorum]]'']]
[[Image:Iberus gualterianus alonensis3.JPG|thumb|''[[Iberus gualtieranus]] alonensis'']]
[[Image:Iberus gualterianus alonensis3.JPG|thumb|''[[Iberus gualtieranus]] alonensis'']]
[[File:Snail-wiki-120-Zachi-Evenor.jpg|thumb|''[[Cornu aspersum]]'']]
[[File:Snail-wiki-120-Zachi-Evenor.jpg|thumb|''[[Cornu aspersum]]'']]


The family Helicidae contains 3 subfamilies (according to molecular phylogenetic analyses<ref>{{Cite journal|last1=Korábek|first1=Ondřej|last2=Petrusek|first2=Adam|last3=Neubert|first3=Eike|last4=Juřičková|first4=Lucie|date=2015-05-01|title=Molecular phylogeny of the genus Helix (Pulmonata: Helicidae)|journal=Zoologica Scripta|language=en|volume=44|issue=3|pages=263–280|doi=10.1111/zsc.12101|s2cid=84673484 |issn=1463-6409}}</ref><ref>{{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}}</ref><ref>{{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}}</ref>):
The family Helicidae contains 3 subfamilies (according to molecular phylogenetic analyses<ref>{{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}}</ref><ref name=":3">{{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}}</ref><ref>{{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}}</ref><ref name=":2">{{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}}</ref>):


=== Subfamily Helicinae Rafinesque, 1815 ===
=== Subfamily Helicinae Rafinesque, 1815 ===
In this subfamily the glands are divided. The love dart has four blades or vanes. There are usually two penial papillae.
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>{{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 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> ====
*''[[Allognathus]]''
*''[[Allognathus]]''
*''[[Cepaea]]'' <small>Held, 1838</small>
*''[[Cepaea]]'' <small>Held, 1838</small>
*''[[Hemicycla]]''
*''[[Hemicycla]]''
*''[[Iberus]]''
*''[[Iberus]]''
*''[[Pseudotachea]]''
*''[[Idiomela]]'' <small>T. Cockerell, 1921</small>
*''[[Idiomela]]'' <small>T. Cockerell, 1921</small>
*''[[Lampadia]]''
*''[[Lampadia]]''
Line 59: Line 50:
==== 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> ====
==== 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> ====
* ''[[Levantina rechingeri|Aristena]]'' <small>Psonis, Vardinoyannis & Poulakakis, 2022</small>
* ''[[Levantina rechingeri|Aristena]]'' <small>Psonis, Vardinoyannis & Poulakakis, 2022</small>
* ''[[Amanica]]'' <small>Nordsieck, 2017</small>
* ''[[Amanica praecellens|Amanica]]'' <small>Nordsieck, 2017</small>
* ''[[Caucasotachea]]'' <small>[[Caesar Rudolf Boettger|Boettger]], 1909</small>
* ''[[Caucasotachea]]'' <small>[[Caesar Rudolf Boettger|Boettger]], 1909</small>
* ''[[Codringtonia]]'' <small>[[Wilhelm Kobelt|Kobelt]], 1898</small>
* ''[[Codringtonia]]'' <small>[[Wilhelm Kobelt|Kobelt]], 1898</small>
Line 67: Line 58:
* ''[[Lindholmia]]'' <small>Hesse, 1918</small>
* ''[[Lindholmia]]'' <small>Hesse, 1918</small>
* ''[[Maltzanella]]'' <small>Hesse, 1917</small>
* ''[[Maltzanella]]'' <small>Hesse, 1917</small>
* ''[[Neocrassa]]'' <small>Subai, 2005</small>
* ''[[Neocrassa neocrassa|Neocrassa]]'' <small>Subai, 2005</small>


⚫
==== 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" /> ====
==== Tribe Otalini ====
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.<ref name="Neiber" />
* ''[[Cantareus]]'' <small>[[Antoine Risso|Risso]], 1826</small>
* ''[[Cantareus]]'' <small>[[Antoine Risso|Risso]], 1826</small>
* ''[[Cornu (gastropod)|Cornu]]'' <small>Born, 1778</small>
* ''[[Cornu (gastropod)|Cornu]]'' <small>Born, 1778</small>
* ''[[Eobania]]'' <small>P. Hesse, 1913</small>
* ''[[Eobania]]'' <small>P. Hesse, 1913</small>
⚫
* ''[[Eremina]]'' <small>Pfeiffer, 1855</small>
⚫
* ''[[Gyrostomella]]'' <small>P. Hesse, 1911</small>
⚫
* ''[[Loxana]]'' <small>Pallary, 1899</small>
* ''[[Massylaea]]'' <small>Möllendorff, 1898</small>
* ''[[Massylaea]]'' <small>Möllendorff, 1898</small>
* ''[[Otala]]'' <small>Schumacher, 1817</small>
* ''[[Otala]]'' <small>Schumacher, 1817</small>
* ''[[Rossmaessleria]]''
* ''[[Rossmaessleria]]'' <small>P. Hesse, 1907</small>

==== Tribe Thebini====
* ''[[Theba]]'' <small>[[Antoine Risso|Risso]], 1826</small>
* ''[[Theba]]'' <small>[[Antoine Risso|Risso]], 1826</small>


==== Tribe Maculariini Neiber, Korábek, Glaubrecht & Hausdorf, 2021 ====
⚫
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.<ref name="Neiber">{{cite journal|author1=Marco T Neiber|author2=Ondřej Korábek|author3=Matthias Glaubrecht|author4=Bernhard Hausdorf|title=A misinterpreted disjunction: the phylogenetic relationships of the North African land snail ''Gyrostomella'' (Gastropoda: Stylommatophora: Helicidae)|journal=Zoological Journal of the Linnean Society| volume = 194| issue= 4|year= 2022| pages= 1236–1251|url= https://doi.org/10.1093/zoolinnean/zlab059|doi=10.1093/zoolinnean/zlab059}}</ref> 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.<ref name = "Neiber"/>
⚫
* ''[[Macularia]]'' <small>Albers, 1850</small>


⚫
=== Subfamily Murellinae Hesse, 1918<ref name=":3" /> ===
⚫
* ''[[Macularia]]'' <small>Albers, 1850</small>
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.<ref name=":4">{{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}}</ref>


Distributed in Sardinia, Corsica, the Apennine Peninsula and Sicily.<ref name=":4" /><ref>{{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 }}</ref>
⚫
=== Subfamily Murellinae ===
⚫
*''[[Marmorana]]'' <small>W. Hartmann, 1844</small>
==== Tribe [[Murellini]] Hesse, 1918 ====
*''[[Marmorana]]'' <small>W. Hartmann, 1844</small> - This genus might actually belong to the subfamily Ariantinae and not to the subfamily Helicinae, as has been assumed until now.<ref>Nordsieck H. (2006). ''Higher classification of Helicoidea and the molecular analyses of their phylogeny''. [http://www.hnords.de/5356429d6b117f602/index.html hnords.de] {{Webarchive|url=https://web.archive.org/web/20120314225859/http://www.hnords.de/5356429d6b117f602/index.html |date=2012-03-14 }}, Accessed 20/06/12</ref>
*''[[Tacheocampylaea]]''
*''[[Tacheocampylaea]]''
*''[[Tyrrheniberus]]''
*''[[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-last=Arntzen |editor-first=J.W. |title=Systematics of Ariantinae (Gastropoda, Pulmonata, Helicidae), a new approach to an old problem |url=https://brill.com/view/journals/ctoz/85/1/article-p37_3.xml |journal=Contributions to Zoology |volume=85 |issue=1 |pages=37–65l |doi=10.1163/18759866-08501003 |issn=1383-4517}}</ref> ===
=== 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 |url=https://brill.com/view/journals/ctoz/85/1/article-p37_3.xml |journal=Contributions to Zoology |volume=85 |issue=1 |pages=37–65l |doi=10.1163/18759866-08501003 |issn=1383-4517|doi-access=free }}</ref> ===
In this subfamily the glands are divided or undivided. The love darts contain two to four blades. There is only one penial papilla.
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|date=July 2023}}
*''[[Arianta]]'' <small>[[William Turton|Turton]], 1831</small>
*''[[Arianta]]'' <small>[[William Turton|Turton]], 1831</small>
*''[[Campylaea]]'' <small>H. Beck, 1837</small>
*''[[Campylaea]]'' <small>H. Beck, 1837</small>
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*''[[Kosicia]]'' <small>Brusina, 1904</small>
*''[[Kosicia]]'' <small>Brusina, 1904</small>
*''[[Liburnica]]'' <small>Kobelt, 1904</small>
*''[[Liburnica]]'' <small>Kobelt, 1904</small>
⚫
*''[[Mesodontopsis]]'' <small>Pilsbry, 1895</small> †
⚫
*''[[Metacampylaea]]'' <small>Pilsbry, 1895</small> †
⚫
*''[[Paradrobacia]]'' <small>H. Nordsieck, 2014</small> †
⚫
*''[[Pseudoklikia]]'' <small>H. Nordsieck, 2018</small> †
*''[[Pseudotrizona]]'' <small>Groenenberg, Subai & E. Gittenberger, 2016</small>
*''[[Pseudotrizona]]'' <small>Groenenberg, Subai & E. Gittenberger, 2016</small>
*''[[Thiessea]]'' <small>Kobelt, 1904</small>
*''[[Thiessea]]'' <small>Kobelt, 1904</small>
*''[[Vidovicia]]'' <small>Brusina, 1904</small>
*''[[Vidovicia]]'' <small>Brusina, 1904</small>
*{{extinct}}''[[Pseudochloritis]]'' C. R. Boettger, 1909<ref name=hotrasser>{{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 }}</ref>
*{{extinct}}''[[Pseudochloritis]]'' C. R. Boettger, 1909<ref name=hotrasser>{{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 }}</ref>
*{{extinct}}''[[Mesodontopsis]]'' <small>Pilsbry, 1895</small>
*{{extinct}}''[[Metacampylaea]]'' <small>Pilsbry, 1895</small>
*{{extinct}}''[[Paradrobacia]]'' <small>H. Nordsieck, 2014</small>
*{{extinct}}''[[Pseudoklikia]]'' <small>H. Nordsieck, 2018</small>


=== ''Incertae sedis'' ===
=== ''Incertae sedis'' ===
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{{Commons category|Helicidae}}
{{Commons category|Helicidae}}


{{taxonbar|from=Q1418257}}
{{Taxonbar|from=Q1418257}}
{{Authority control}}


[[Category:Helicidae| ]]
[[Category:Helicidae| ]]

Latest revision as of 07:01, 17 May 2024

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

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

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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

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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

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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

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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

[edit]
Arianta arbustorum
Iberus gualtieranus alonensis
Cornu aspersum

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

Subfamily Helicinae Rafinesque, 1815

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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]

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Tribe Helicini Rafinesque, 1815[13]

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Tribe Thebini Wenz, 1923[14][12]

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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]

Tribe Maculariini Neiber, Korábek, Glaubrecht & Hausdorf, 2021

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Subfamily Murellinae Hesse, 1918[10]

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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]

Subfamily Ariantinae Mörch, 1864[17]

[edit]

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]

Incertae sedis

[edit]

References

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  1. ^ M.P. Kerney & R.A.D. Cameron. 1979. A field guide to the land snails of Britain and northwestern Europe. Collins, London.
  2. ^ a b 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.
  4. ^ 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.
  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.
  10. ^ 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.
  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.
  12. ^ 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.
  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.
  14. ^ 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.
  15. ^ 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.
  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.
  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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