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{{Short description|Family of gastropods}}
{{Taxobox
{{Automatic taxobox
| color = pink
| name = Helicidae
| image = Helix pomatia 89a.jpg
| image = Helix pomatia 89a.jpg
| image_caption = ''[[Helix pomatia]]''
| image_width = 225px
| regnum = [[Animal]]ia
| taxon = Helicidae
| authority = [[Constantine Samuel Rafinesque|Rafinesque]], 1815
| phylum = [[Mollusca]]
| classis = [[Gastropoda]]
| subclassis = [[Orthogastropoda]]
| superordo = [[Heterobranchia]]
| ordo = [[Pulmonata]]
| superfamilia = [[Helicoidea]]
| familia = '''Helicidae'''
| familia_authority = [[Rafinesque]], [[1815]]
| subdivision_ranks = Subfamilies
| subdivision_ranks = Subfamilies
| subdivision =
| subdivision = *[[Ariantinae]]
[[Ariantinae]]<br>
*[[Murellinae]]
[[Helicinae]]
*[[Helicinae]]
| 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 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.
The '''Helicidae''', sometimes known as the typical snails, are a [[taxonomic]] [[family (biology)|family]] of medium sized to large, air-breathing, land [[snail]]s. In other words, they are small to large [[terrestrial]] [[pulmonate]] [[gastropod]] [[mollusk]]s.


== Shell description ==
Well-known species in this family include ''[[Helix aspersa]]'' the garden snail, and ''[[Helix pomatia]]'' the "[[escargot]]".
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.


==Distribution==
== 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 family is native to Europe.


== Genetics ==
However some of the species, such as for example ''[[Helix aspersa]]'', have been introduced and become established in numerous different places worldwide.
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}}).
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.


[[Image:Snails-Italy.jpg|thumb|Helicid snails for sale as food in [[Italy]]; from the front ''[[Eobania vermiculata]]'', ''[[Cantareus apertus]]'', and ''Helix'' sp.]]
==Shell description==
The shells of these snails are mostly rather globular in shape. In some genera, such as ''[[Cepaea]]'', the shells are brightly colored and patterned.


== Distribution ==
==Subfamilies and genera within the family Helicidae==
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.
* subfamily [[Ariantinae]] Mörch, 1864
** ''[[Arianta]]'' Turton, 1831
** ''[[Chilostoma]]'' Fitzinger, 1833
** ''[[Cylindrus]]'' Fitzinger, 1833
** ''[[Helicigona]]'' A. Férussac, 1821
** ''[[Isognomostoma]]'' Fitzinger, 1833


== Taxonomy ==
* subfamily [[Helicinae]] Rafinesque, 1815
[[Image:Snail black on grass2.jpg|thumb|''[[Arianta arbustorum]]'']]
** ''[[Cantareus]]'' Risso, 1826
[[Image:Iberus gualterianus alonensis3.JPG|thumb|''[[Iberus gualtieranus]] alonensis'']]
** ''[[Cepaea]]'' Held, 1838
[[File:Snail-wiki-120-Zachi-Evenor.jpg|thumb|''[[Cornu aspersum]]'']]
** ''[[Cornu (genus)|Cornu]]'' Born, 1778
** ''[[Eobania]]'' P. Hesse, 1913
** ''[[Helix (genus)|Helix]]'' Linnaeus, 1758
** ''[[Idiomela]]'' T. Cockerell, 1921
** ''[[Theba]]'' Risso, 1826 - ''[[Theba pisana]]'' (O. F. Müller, 1774)
** ''[[Otala]]'' Schumacher, 1817


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>):
==References==
<references/>
* M.P. Kerney & R.A.D. Cameron, 1979, a field guide to the land snails of Britain and northwestern Europe, Collins, London


=== Subfamily Helicinae Rafinesque, 1815 ===
{{commonscat|Helicidae}}
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> ====
[[Category:Helicidae| ]]
*''[[Allognathus]]''
*''[[Cepaea]]'' <small>Held, 1838</small>
*''[[Hemicycla]]''
*''[[Iberus]]''
*''[[Idiomela]]'' <small>T. Cockerell, 1921</small>
*''[[Lampadia]]''


==== 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> ====
[[fr:Helicidae]]
* ''[[Levantina rechingeri|Aristena]]'' <small>Psonis, Vardinoyannis & Poulakakis, 2022</small>
[[lt:Helicidai]]
* ''[[Amanica praecellens|Amanica]]'' <small>Nordsieck, 2017</small>
[[pl:Ślimakowate]]
* ''[[Caucasotachea]]'' <small>[[Caesar Rudolf Boettger|Boettger]], 1909</small>
[[sk:Slimákovité]]
* ''[[Codringtonia]]'' <small>[[Wilhelm Kobelt|Kobelt]], 1898</small>
[[fi:Tarhakotilot]]
* ''[[Helix (gastropod)|Helix]]'' <small>[[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]</small> - [[type genus]]
* ''[[Isaurica (gastropod)|Isaurica]]'' <small>Kobelt, 1901</small>
* ''[[Levantina]]'' <small>Kobelt, 1871</small>
* ''[[Lindholmia]]'' <small>Hesse, 1918</small>
* ''[[Maltzanella]]'' <small>Hesse, 1917</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" /> ====
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>
* ''[[Cornu (gastropod)|Cornu]]'' <small>Born, 1778</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>
* ''[[Otala]]'' <small>Schumacher, 1817</small>
* ''[[Rossmaessleria]]'' <small>P. Hesse, 1907</small>
* ''[[Theba]]'' <small>[[Antoine Risso|Risso]], 1826</small>

==== Tribe Maculariini Neiber, Korábek, Glaubrecht & Hausdorf, 2021 ====
* ''[[Macularia]]'' <small>Albers, 1850</small>

=== 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.<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>
*''[[Marmorana]]'' <small>W. Hartmann, 1844</small>
*''[[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 |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> ===
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>
*''[[Campylaea]]'' <small>H. Beck, 1837</small>
*''[[Campylaeopsis]]'' <small>A.J. Wagner, 1914</small>
* ''[[Cattania]]'' <small>Brusina, 1904</small>
*''[[Causa]]'' <small>Schileyko, 1971</small>
*''[[Chilostoma]]'' <small>Fitzinger, 1833</small>
*''[[Corneola]]'' <small>Held, 1838</small>
*''[[Cylindrus]]'' <small>Fitzinger, 1833</small>
*''[[Delphinatia]]'' <small>P. Hesse, 1931</small>
*''[[Dinarica]]'' <small>Kobelt, 1902</small>
*''[[Drobacia]]'' <small>Brusina, 1904</small>
*''[[Faustina (gastropod)|Faustina]]'' <small>Kobelt, 1904</small>
*''[[Helicigona]]'' <small>A. Férussac, 1821</small>
*''[[Isognomostoma]]'' <small>Fitzinger, 1833</small>
*''[[Josephinella]]'' <small>F. Haas, 1936</small>
*''[[Kollarix]]'' <small>Groenenberg, Subai & E. Gittenberger, 2016</small>
*''[[Kosicia]]'' <small>Brusina, 1904</small>
*''[[Liburnica]]'' <small>Kobelt, 1904</small>
*''[[Pseudotrizona]]'' <small>Groenenberg, Subai & E. Gittenberger, 2016</small>
*''[[Thiessea]]'' <small>Kobelt, 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}}''[[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'' ===
*†''[[Megalotachea]]'' <small>Pfeffer, 1930</small>

== References ==
{{Reflist}}

== External links ==
{{Commons category|Helicidae}}

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

[[Category:Helicidae| ]]
[[Category:Helicoidea]]
[[Category:Gastropod families]]
[[Category:Taxa named by Constantine Samuel Rafinesque]]

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

[edit]

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

[edit]

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

[edit]

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

[edit]

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

[edit]

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]

[edit]

Tribe Helicini Rafinesque, 1815[13]

[edit]

Tribe Thebini Wenz, 1923[14][12]

[edit]

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

[edit]

Subfamily Murellinae Hesse, 1918[10]

[edit]

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

[edit]
  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.
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