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==Life history==
==Life history==
In ''H. pomatia'', sexual maturity is reached after 2–4 (–6) overwinterings, i.e. at the age of 2–4 (–6) years, but this differs between localities as well as between snails from the same clutch.<ref name=":3" /><ref name=":11" /><ref name=":16">{{Cite journal |last=Falkner |first=Gerhard |date=1984 |title=Das bayerische Weinbergschnecken-Projekt (Untersuchungen an <i>Helix pomatia</i> L.) |journal=Mitteilungen der deutschen malakozoologischen Gesellschaft |volume=37 |pages=182-197}}</ref> Adult snails cease to grow and form a thickened lip around the aperture. However, the snails sometimes mate already shortly before the lip is formed. The life span of ''H. pomatia'' may exceed 20 years in the wild.<ref name=":16" /> However, in the wild, they mostly live for just 5–9 years.<ref name=":11" /><ref name=":16" /> The age of ''H. pomatia'' individuals can be estimated by counting growth interruptions on the shell and the number of layers deposited on the aperture margin lip.<ref name=":11" /><ref>{{Cite journal |last=Pollard |first=E. |last2=Cooke |first2=A. S. |last3=Welch |first3=J. M. |date=1977 |title=The use of shell features in age determination of juvenile and adult Roman snails <i>Helix pomatia</i> |journal=Journal of Zoology, London |volume=183 |pages=269-279}}</ref><ref name=":16" /> The small species ''H. ceratina'' reaches maturity after 1.5–2 years and lives 4–5 years.<ref name=":12">{{Cite journal |last1=Camus |first1=Louise |last2=Poli |first2=Pedro |last3=Delaugerre |first3=Michel-Jean |last4=Dréano |first4=Stéphane |last5=Cucherat |first5=Xavier |last6=Natali |first6=Christine |last7=Guiller |first7=Annie |date=2023-05-22 |title=Unexpected and spatially structured genetic diversity of the relict population of the endangered corsican land snail Tyrrhenaria ceratina |url=https://link.springer.com/10.1007/s10592-023-01535-0 |journal=Conservation Genetics |language=en |doi=10.1007/s10592-023-01535-0 |s2cid=258855822 |issn=1566-0621}}</ref>
In ''H. pomatia'', sexual maturity is reached after 2–4 (–6) overwinterings, i.e. at the age of 2–4 (–6) years, but this differs between localities as well as between snails from the same clutch.<ref name=":3" /><ref name=":11" /><ref name=":16">{{Cite journal |last=Falkner |first=Gerhard |date=1984 |title=Das bayerische Weinbergschnecken-Projekt (Untersuchungen an <i>Helix pomatia</i> L.) |journal=Mitteilungen der deutschen malakozoologischen Gesellschaft |volume=37 |pages=182-197}}</ref> Adult snails cease to grow and form a thickened lip around the aperture. However, the snails sometimes mate already shortly before the lip is formed. The life span of ''H. pomatia'' may reach 30 years in the wild.<ref name=":16" /> However, in the wild, they mostly live for just 5–9 years.<ref name=":11" /><ref name=":16" /> The age of ''H. pomatia'' individuals can be estimated by counting growth interruptions on the shell and the number of layers deposited on the aperture margin lip.<ref name=":11" /><ref>{{Cite journal |last=Pollard |first=E. |last2=Cooke |first2=A. S. |last3=Welch |first3=J. M. |date=1977 |title=The use of shell features in age determination of juvenile and adult Roman snails <i>Helix pomatia</i> |journal=Journal of Zoology, London |volume=183 |pages=269-279}}</ref><ref name=":16" /> The small species ''H. ceratina'' reaches maturity after 1.5–2 years and lives 4–5 years.<ref name=":12">{{Cite journal |last1=Camus |first1=Louise |last2=Poli |first2=Pedro |last3=Delaugerre |first3=Michel-Jean |last4=Dréano |first4=Stéphane |last5=Cucherat |first5=Xavier |last6=Natali |first6=Christine |last7=Guiller |first7=Annie |date=2023-05-22 |title=Unexpected and spatially structured genetic diversity of the relict population of the endangered corsican land snail Tyrrhenaria ceratina |url=https://link.springer.com/10.1007/s10592-023-01535-0 |journal=Conservation Genetics |language=en |doi=10.1007/s10592-023-01535-0 |s2cid=258855822 |issn=1566-0621}}</ref>


The length of the life cycle is dependent on environmental conditions. The time from hatching to first egg laying can be shortened to just 12–13 months under optimal conditions in the captivity, mainly by skipping hibernation.<ref name=":18" />
The length of the life cycle is dependent on environmental conditions. The time from hatching to first egg laying can be shortened to just 12–13 months under optimal conditions in the captivity, mainly by skipping hibernation.<ref name=":18" />
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Species of ''Helix'' live in a variety of habitats and under very different climatic regimes. Some species are exclusively found in open limestone rocky habitats (''H. secernenda''), while others tolerate acidic bedrock (e.g. ''H. pelagonesica'') or live predominantly in [[forest]]s (e.g. ''H. thessalica''). Members of the genus occur from [[temperate rainforest]]s (''H. buchii'') to [[Semi-arid climate|semi-arid]] regions (''Helix pronuba'').<ref>{{Cite journal |last=Korábek |first=Ondřej |date=2014 |title=Hlemýždi – stopařův průvodce nejen po Středomoří |url=https://ziva.avcr.cz/files/ziva/pdf/hlemyzdi-stoparuv-pruvodce-nejen-po-stredomori.pdf |journal=Živa |volume=2014 |issue=4 |pages=171–175}}</ref><ref name=":0" />
Species of ''Helix'' live in a variety of habitats and under very different climatic regimes. Some species are exclusively found in open limestone rocky habitats (''H. secernenda''), while others tolerate acidic bedrock (e.g. ''H. pelagonesica'') or live predominantly in [[forest]]s (e.g. ''H. thessalica''). Members of the genus occur from [[temperate rainforest]]s (''H. buchii'') to [[Semi-arid climate|semi-arid]] regions (''Helix pronuba'').<ref>{{Cite journal |last=Korábek |first=Ondřej |date=2014 |title=Hlemýždi – stopařův průvodce nejen po Středomoří |url=https://ziva.avcr.cz/files/ziva/pdf/hlemyzdi-stoparuv-pruvodce-nejen-po-stredomori.pdf |journal=Živa |volume=2014 |issue=4 |pages=171–175}}</ref><ref name=":0" />


''Helix pomatia'' feeds mainly on live plants and observations from England showed preference for somespecific plant species.<ref name=":11" />
''Helix pomatia'' feeds mainly on live plants.<ref name=":18" /><ref name=":11" /> Observations showed preference for somespecific plant species<ref name=":11" /> and avoidance of others.<ref name=":18" />


The activity of ''H. pomatia'' takes place mostly during the night, especially in juveniles.<ref name=":18" /> During winter, ''Helix'' snails hibernate in the soil.
A homing behaviour has been observed in ''H. pomatia.'' The snails disperse during the season, but tend to return to their hibernation grounds towards its end.<ref name=":11">{{Cite journal |last=Pollard |first=E. |date=1975 |title=Aspects of the Ecology of <i>Helix pomatia</i> L. |journal=[[The Journal of Animal Ecology]] |volume=44 |issue=1 |pages=305–329 |doi=10.2307/3865 |jstor=3865}}</ref> Young snails seem to be more dispersive than older ones.<ref name=":16" /> It was also observed that the snails may migrate to specific sites for egg laying.{{Citation needed|date=May 2024}}

A homing behaviour has been observed in ''H. pomatia.'' The snails disperse during the season, but tend to return to their hibernation grounds towards its end.<ref name=":20">{{Cite journal |last=Edelstam |first=Carl |last2=Palmer |first2=Carina |date=1950 |title=Homing behaviour in gastropodes |url=https://www.jstor.org/stable/3564796 |journal=Oikos |volume=2 |issue=2 |pages=259-270 |via=JSTOR}}</ref><ref name=":11">{{Cite journal |last=Pollard |first=E. |date=1975 |title=Aspects of the Ecology of <i>Helix pomatia</i> L. |journal=[[The Journal of Animal Ecology]] |volume=44 |issue=1 |pages=305–329 |doi=10.2307/3865 |jstor=3865}}</ref> It was also observed that the snails may migrate to specific sites for egg laying{{Citation needed|date=May 2024}} and aggregate for mating.<ref>{{Cite journal |last=Jarčuška |first=Benjamín |last2=Jarčušková Danková |first2=Lucia |date=2014 |title=Poznámky k zoskupovaniu slimáka záhradného <i>Helix pomatia</i> [Comments on the aggregation behaviour of the Roman snail (<i>Helix pomatia</i>)] |url=https://mollusca.sav.sk/pdf/13/13.Jarcuska.pdf |journal=Malacologica Bohemoslovaca |volume=13 |pages=114–115}}</ref> The snails are able to find their way back from distances of tens of meters.<ref name=":20" />


Several species (e.g. ''H. pomatia'', ''H. figulina'') build a thick, calcified [[epiphragm]] that closes the shell's aperture during [[hibernation]] or [[aestivation]]. The epiphragm is followed inside the shell by a few additional membranes made of dried mucus.<ref>{{Cite book |last=Schileyko |first=A. A. |title=Mollyuski, Tom III, vyp. 6. Nazemnye mollyuski nadsemeystva Helicoidea |publisher=Nauka |year=1978 |series=Fauna SSSR |location=Leningrad}}</ref> When the animal emerges from the dormancy, it discards the calcareous epiphragm using the posterior part of the foot.{{Citation needed|date=July 2023}}
Several species (e.g. ''H. pomatia'', ''H. figulina'') build a thick, calcified [[epiphragm]] that closes the shell's aperture during [[hibernation]] or [[aestivation]]. The epiphragm is followed inside the shell by a few additional membranes made of dried mucus.<ref>{{Cite book |last=Schileyko |first=A. A. |title=Mollyuski, Tom III, vyp. 6. Nazemnye mollyuski nadsemeystva Helicoidea |publisher=Nauka |year=1978 |series=Fauna SSSR |location=Leningrad}}</ref> When the animal emerges from the dormancy, it discards the calcareous epiphragm using the posterior part of the foot.{{Citation needed|date=July 2023}}
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=== Predators and parasites ===
=== Predators and parasites ===
Several bird species prey on ''Helix'' species. This has been observed in ''[[Common pheasant|Phasianus colchicus]]'', ''[[Eurasian stone-curlew|Burhinus oedicemus]]'', [[European roller|''Coracias garrulus'']], ''[[Great grey shrike|Lanius excubitor]]'', [[Corvidae|corvids]], and probably other birds.<ref>{{Cite journal |last=Mienis |first=Henk |date=1993 |title=Welke vogels eten Wijngaardslakken? |url=https://natuurtijdschriften.nl/pub/600870/CNMV1993273001005.pdf |journal=Correspondentieblad van de Nederlandse Malacologische Vereniging |volume=273 |pages=92–93}}</ref><ref name=":13">{{Cite book |last=Allen |first=John A. |title=Natural Enemies of Terrestrial Molluscs |publisher=CABI Publishing |year=2004 |isbn=0-85199-319-2 |editor-last=Barker |editor-first=G.M. |location=Cambridge (USA) |pages=1–36 |chapter=Avian an mammalian predators of terrestrial gastropods}}</ref> [[Mammal]]s prey on ''Helix'', too. The known predators include [[hedgehog]], [[shrew]], [[rodent]]s (''[[Rattus]]'', ''[[Apodemus]]'') and the wild boar.<ref>{{Cite journal |last1=Schley |first1=Laurent |last2=Roper |first2=Timothy J. |date=2003 |title=Diet of wild boar Sus scrofa in Western Europe, with particular reference to consumption of agricultural crops: Diet of wild boar |url=http://doi.wiley.com/10.1046/j.1365-2907.2003.00010.x |journal=[[Mammal Review]] |language=en |volume=33 |issue=1 |pages=43–56 |doi=10.1046/j.1365-2907.2003.00010.x}}</ref><ref name=":13" /> The [[slow worm]] consumes molluscs, including ''Helix''.<ref>{{Cite book |last=Lapota-Ferreira |first=Iara Lúcia |title=Natural Enemies of Terrestrial Molluscs |publisher=CABI Publishing |year=2004 |isbn=0-85199-319-2 |editor-last=Barker |editor-first=G. M. |location=Cambridge (USA) |pages=427–482 |chapter=Reptilian predators of terrestrial gastropods}}</ref> ''Helix'' snails are also attacked by various [[beetle]]s and [[Fly|flies]]. The beetle predators belong to the families [[Ground beetle|Carabidae]] and [[Firefly|Lampyridae]].<ref name=":11" /><ref name=":16" /><ref>{{Cite book |last=Symondson |first=Wiliam O. C. |title=Natural Enemies of Terrestrial Molluscs |publisher=CABI Publishing |year=2004 |isbn=0-85199-319-2 |editor-last=Barker |editor-first=G.M. |location=Cambridge (USA) |pages=37–84 |chapter=Coleoptera (Carabidae, Staphylinidae, Lampyridae, Drilidae and Silphidae) as predators of terrestrial gastropods}}</ref> The predation by birds, small mammals, and beetles mostly affects juveniles.<ref name=":11" /> Larvae of flies from several families attack ''Helix'' snails and may kill even adults ([[Phoridae]], [[Muscidae]], [[Flesh fly|Sarcophagidae]], [[Sciomyzidae]]).<ref>{{Cite book |last1=Coupland |first1=James B. |last2=Barker |first2=Gary B. |title=Natural Enemies of Terrestrial Molluscs |publisher=CABI Publishing |year=2004 |isbn=0-85199-319-2 |editor-last=Barker |editor-first=G. M. |location=Cambridge (USA) |pages=85–158 |chapter=Diptera as predators and parasitoids of terrestrial gastropods, with emphasis on Phoridae, Calliphoridae, Sarcophagidae, Muscidae and Fanniidae}}</ref>
Several bird species prey on ''Helix'' species. This has been observed in ''[[Common pheasant|Phasianus colchicus]]'', ''[[Eurasian stone-curlew|Burhinus oedicemus]]'', [[European roller|''Coracias garrulus'']], ''[[Great grey shrike|Lanius excubitor]]'', [[Corvidae|corvids]], and probably other birds.<ref>{{Cite journal |last=Mienis |first=Henk |date=1993 |title=Welke vogels eten Wijngaardslakken? |url=https://natuurtijdschriften.nl/pub/600870/CNMV1993273001005.pdf |journal=Correspondentieblad van de Nederlandse Malacologische Vereniging |volume=273 |pages=92–93}}</ref><ref name=":13">{{Cite book |last=Allen |first=John A. |title=Natural Enemies of Terrestrial Molluscs |publisher=CABI Publishing |year=2004 |isbn=0-85199-319-2 |editor-last=Barker |editor-first=G.M. |location=Cambridge (USA) |pages=1–36 |chapter=Avian an mammalian predators of terrestrial gastropods}}</ref> [[Mammal]]s prey on ''Helix'', too. The known predators include [[hedgehog]], [[Mole (animal)|mole]], [[shrew]], [[rodent]]s (''[[Rattus]]'', ''[[Apodemus]]'') and the wild boar.<ref>{{Cite journal |last1=Schley |first1=Laurent |last2=Roper |first2=Timothy J. |date=2003 |title=Diet of wild boar Sus scrofa in Western Europe, with particular reference to consumption of agricultural crops: Diet of wild boar |url=http://doi.wiley.com/10.1046/j.1365-2907.2003.00010.x |journal=[[Mammal Review]] |language=en |volume=33 |issue=1 |pages=43–56 |doi=10.1046/j.1365-2907.2003.00010.x}}</ref><ref name=":13" /> The [[slow worm]] consumes molluscs, including ''Helix''.<ref>{{Cite book |last=Lapota-Ferreira |first=Iara Lúcia |title=Natural Enemies of Terrestrial Molluscs |publisher=CABI Publishing |year=2004 |isbn=0-85199-319-2 |editor-last=Barker |editor-first=G. M. |location=Cambridge (USA) |pages=427–482 |chapter=Reptilian predators of terrestrial gastropods}}</ref> ''Helix'' snails are also attacked by various [[beetle]]s and [[Fly|flies]]. The beetle predators belong to the families [[Ground beetle|Carabidae]] and [[Firefly|Lampyridae]].<ref name=":11" /><ref name=":16" /><ref>{{Cite book |last=Symondson |first=Wiliam O. C. |title=Natural Enemies of Terrestrial Molluscs |publisher=CABI Publishing |year=2004 |isbn=0-85199-319-2 |editor-last=Barker |editor-first=G.M. |location=Cambridge (USA) |pages=37–84 |chapter=Coleoptera (Carabidae, Staphylinidae, Lampyridae, Drilidae and Silphidae) as predators of terrestrial gastropods}}</ref> The predation by birds, small mammals, and beetles mostly affects juveniles.<ref name=":11" /> Larvae of flies from several families attack ''Helix'' snails and may kill even adults ([[Phoridae]], [[Muscidae]], [[Flesh fly|Sarcophagidae]], [[Sciomyzidae]]).<ref>{{Cite book |last1=Coupland |first1=James B. |last2=Barker |first2=Gary B. |title=Natural Enemies of Terrestrial Molluscs |publisher=CABI Publishing |year=2004 |isbn=0-85199-319-2 |editor-last=Barker |editor-first=G. M. |location=Cambridge (USA) |pages=85–158 |chapter=Diptera as predators and parasitoids of terrestrial gastropods, with emphasis on Phoridae, Calliphoridae, Sarcophagidae, Muscidae and Fanniidae}}</ref>


A well known [[Facultative parasite|facultative]] [[Parasitism|parasite]] of land snails, including ''Helix'', is the [[nematode]] ''[[Phasmarhabditis hermaphrodita|Phasmarhabiditis hermaphrodita]]''.<ref>{{Cite book |last1=Morand |first1=Serge |last2=Wilson |first2=Michael J. |last3=Glen |first3=David M. |title=Natural Enemies of Terrestrial Molluscs |publisher=CABI Publishing |year=2004 |isbn=0-85199-319-2 |editor-last=Barker |editor-first=G. M. |location=Cambridge (USA) |pages=525–558 |chapter=Nematodes (Nematodes) parasitic in terrestrial gastropods}}</ref> The parasitic mite ''[[Riccardoella limacum]]'' is found on ''Helix'' species.<ref>{{Cite journal |last1=Turk |first1=Frank A. |last2=Phillips |first2=Stella‐Maris |date=1946 |title=A Monograph of the Slug Mite—Riccardoella limacum (Schrank.). |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1096-3642.1946.tb00102.x |journal=[[Proceedings of the Zoological Society of London]] |language=en |volume=115 |issue=3–4 |pages=448–472 |doi=10.1111/j.1096-3642.1946.tb00102.x |issn=0370-2774}}</ref>
A well known [[Facultative parasite|facultative]] [[Parasitism|parasite]] of land snails, including ''Helix'', is the [[nematode]] ''[[Phasmarhabditis hermaphrodita|Phasmarhabiditis hermaphrodita]]''.<ref>{{Cite book |last1=Morand |first1=Serge |last2=Wilson |first2=Michael J. |last3=Glen |first3=David M. |title=Natural Enemies of Terrestrial Molluscs |publisher=CABI Publishing |year=2004 |isbn=0-85199-319-2 |editor-last=Barker |editor-first=G. M. |location=Cambridge (USA) |pages=525–558 |chapter=Nematodes (Nematodes) parasitic in terrestrial gastropods}}</ref> The parasitic mite ''[[Riccardoella limacum]]'' is found on ''Helix'' species.<ref>{{Cite journal |last1=Turk |first1=Frank A. |last2=Phillips |first2=Stella‐Maris |date=1946 |title=A Monograph of the Slug Mite—Riccardoella limacum (Schrank.). |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1096-3642.1946.tb00102.x |journal=[[Proceedings of the Zoological Society of London]] |language=en |volume=115 |issue=3–4 |pages=448–472 |doi=10.1111/j.1096-3642.1946.tb00102.x |issn=0370-2774}}</ref>

Revision as of 09:12, 16 May 2024

Helix
Temporal range: Miocene–recent
Helix pomatia
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Order: Stylommatophora
Family: Helicidae
Subfamily: Helicinae
Tribe: Helicini
Genus: Helix
Linnaeus, 1758
Type species
Helix pomatia
Synonyms

see text

Helix is a genus of large, air-breathing land snails native to the western Palaearctic and characterized by a globular shell.[1][2]

It is the type genus of the family Helicidae, and one of the animal genera described by Carl Linnaeus[3] at the dawn of the zoological nomenclature.

Members of the genus first appeared in the fossil record during the Miocene.[4]

Well-known species include Helix pomatia (Roman snail, Burgundy snail, or edible snail) and Helix lucorum (Turkish snail). Cornu aspersum (garden snail), though externally similar and long classified as a member of Helix (as "Helix aspersa"), is not closely related to Helix[5][6] and belongs to a different tribe of Helicinae.[7]

Taxonomy

In Linnaeus' 10th edition of Systema Naturae, which marks the beginning of the zoological nomenclature, the generic name Helix had been used for a variety of terrestrial (e.g. Zonites algirus), freshwater (e.g. Lymnaea stagnalis), and marine (e.g. Fossarus ambiguus) gastropods. Later authors restricted the name's use to stylommatophoran species with flattened to globular shells, including zonitids and other groups. In the course of the 1800s, several thousand species of Europe and abroad have been described in Helix.[8][9] By the early 1900s, the genus was split into many separate genera, leaving only species closely related to its type species Helix pomatia in the genus. However, due to the previously broad concept of the genus, Helix is part of the original combination (basionym) of many gastropod names and there still are many nominal taxa described in Helix whose generic placement remains unresolved (taxa inquirenda),[10] although they clearly do not refer to any species of Helix in its present sense.

In the 2000s, Helix has been subject to extensive molecular phylogenetic studies and taxonomic revisions.[1][11][2][12][13] These led to the exclusion of several species, most notably the garden snail, and inclusion of others (H. ceratina, H. nicaeensis). Maltzanella, for long considered a subgenus of Helix, was also formally removed from the genus,[14] but is the sister group of Helix.

Two subgenera are currently recognized:[2][15]

  • Helix (Helix) Linnaeus, 1758
  • Helix (Pelasga) Hesse, 1908
  • Helix (Aegaeohelix) Korábek & Hausdorf, 2023

Description

Helix comprises large land snails species, with shell diameter of 2–6 cm. The shell is globular to conical, with five darker bands that may be variably reduced or fuse together. The globular shell distinguishes Helix from most of the related genera (tribe Helicini), except for Maltzanella and Lindholmia. The surface has a structure of fine transversal ribs, developed to a varying degree, and there may be very fine spiral grooves as well. The shell is never malleated. Colour of the foot varies. It may be grey, brown, black or pink; the back of the foot is dark in several species.

The shells of Helix species are dextral. Sinistral individuals are very rare, but are occasionally found (e.g. H. pomatia,[16] H. thessalica[17] and H. lutescens[18]).

Characters on the genital system have been used to define the genus and its subgenera. Unlike Cornu, the penis of Helix contains two papillae with a central opening. There appears to be a tendency for a shortening of the diverticulum of bursa copulatrix and of the eppiphallus, but there is an overlap with related genera in these characters. Mucous glands adjoining the dart sac are usually richly branched.

Distribution

Helix is a western Palaearctic genus. The species diversity is concentrated to the Balkans and Anatolia, with the greatest phylogenetic diversity in Greece. The natural western distribution limits run through mainland France (Helix pomatia), Corsica (Helix ceratina), and Algeria (Helix melanostoma). In the north, the natural distribution of H. pomatia reaches central Germany and the southern margins of the North European plain. The southernmost species live in North Africa (H. melanostoma, H. pronuba) and the southern Levant (H. engaddensis). The eastern limits are reached in western Iran and Iraqi Kurdistan (H. salomonica) and in the Caucasus (H. lucorum); H. thessalica reaches through Ukraine at least to the western Russian frontier.[1][2]

Genetics

Haploid genome size was estimated to be nearly 4 Gbp (C-value 4 pg) with a GC-content of ~42%,[19][20] but it is unclear which species was studied due to a discrepancy between the stated species and sample origin. The haploid number of chromosomes is 27 (studied species were H. lucorum, H. buchii, H. pomatia, H. gussoneana and H. straminea).[21][22][23][24] In H. pomatia, all chromosomes have median or sub-median centromeres.[25] Small supernumerary chromosomes were reported from H. pomatia from England.[25]

The mitochondrial genome of H. pomatia is available (ca. 14 070 bp long).[26][27]

Genital system

Scheme of the genital system in Helix and related Helicidae.[28] D - love-dart, S - stylophore or dart sac, MG - mucus glands, P - penis, EP - epiphallus, FL - flagellum, BTD - bursa tract diverticulum, BT - bursa tract, BC - bursa copulatrix, SRO - spermatophore-receiving organ, SP - spermathecae, sperm storage organ, FP - fertilization pouch, AG - albumen gland, G - genital opening, HD - hermaphroditic duct, OT - ovotestis, PRM - penis retractor muscle, SO - spermoviduct, V - vaginal duct, VD - vas deferens

The structure of the genital system corresponds in most aspects to that of other Helicidae. Its anatomy and function have been studied in detail in H. pomatia.[29][30][31]

As all stylommatophorans, Helix snails are hermaphrodites. Sperms and egg cells are produced in a common gonad, the ovotestis (hermaphroditic gland), which is embedded in the hepatopancreas (digestive gland) near the apex of the shell. Gametes are transported through a hermaphroditic duct (ovotestis duct) to the fertilization pouch–spermatheca complex (carrefour) embedded at the base of the albumen gland.[32] In this organ, the foreign sperm (sperm from the other individual) is stored in spermathecal sacs (receptacula seminis) and egg fertilization takes place in the fertilisation pouch. The albumen gland provides nourishment for the developing fertilized eggs, and its size greatly varies with the stage of the reproductive cycle. The snail's own sperm and fertilized eggs are transported by specialised regions of the spermoviduct (sperm groove and uterus), which distally separate into a male (vas deferens) and female (free oviduct) parts of the genital system.[32]

The male genitalia consist of a tube that serves the formation of a spermatophore and its transfer into the female parts of the mating partner. The penis is the most distal and muscular part. A spermatophore is formed in the epiphallus (between vas deferens and penis) and the flagellum (continuation of the epiphallus proximally from the vas deferens opening); the latter forms the tail of the spermatophore. During copulation, the penis everts (like a sleeve turned inside out)[29] and is in this process inserted in the vagina. A retractor muscle attaches on the epiphallus and retracts the male genitalia after copulation.

The female part consists of a vagina (sometimes called the copulatory canal[33]) and the bursa copulatrix (gametolytic gland) with its stalk and usually a diverticulum of the stalk. The vagina serves the transport of the foreign spermatophore and of eggs. The bursa is attached by a thin stalk to the vagina (marking the boundary between vagina and the free oviduct). The stalk in most cases bears a diverticulum, a blind tube that receives the front part of the spermatophore if present. The diverticulum has been proposed to be a remnant of a seminal duct that originally transported foreign sperm into the fertilization pouch.[34] Sperm leave the tail of the spermatophore and migrate into the oviduct and then to the fertilization pouch; the vast majority of the sperm does not escape in this way and is digested in the bursa.[31][32] In Helix, there is tendency for a reduction of the diverticulum, and it can be missing in several species.[2]

Dart apparatus, although positioned on the vagina, is functionally also part of the male genitalia. It is composed of a single muscular dart sac and two mucous glands (digitiform or accessory glands) on its sides. The mucous glands are branched at their base; the number of branches varies between Helix species. The single dart is aragonitic, straight or only weakly curved, with four blades (vanes) along its length and a corona at its base.[35][5] The dart apparatus is missing in Helix salomonica.[2] The mucous glands produce mucus that covers the dart during shooting and is thereby injected into the body of the partner, where it induces shortening of the diverticulum and peristaltic movements of the bursa stalk that help the foreign sperm to escape lysis in the bursa.[33]

The male and female parts open into a common atrium and a genital pore positioned ventrally behind the right optic tentacle.

A rare teratological individual with paired male genitalia (penis, epiphallus, flagellum) has been reported from Germany.[36]

Reproduction

Helix pomatia in the initial phase of the mating sequence.

The aspects of reproduction have been studied primarily in H. pomatia, with limited information from other species.

Mating behaviour has been described several times for H. pomatia.[29][37] In the initial phase, the two snails raise their feet and press the soles against each other and touch each other's tentacles and mouthparts. This takes 15–30 min. Some time later, the dart shooting takes place, although many matings progress without a love dart being employed. The mucous glands produce a whitish secretion just before the shooting, that contains hormones promoting the compound that improves preservation of foreign sperm in the receiving individual.[38] Then, again after a pause, comes the copulation, usually preceded by several unsuccessful attempts in which the reciprocal insertion of penes is not achieved and the genital organs are partially retracted back into the body. Finally, both individuals simultaneously insert the penes into each other's female opening. Within ca. 4–7 min the spermatophore is formed and transferred, after which the snail disengage and retract the everted genital organs. However, the complete reception of the spermatophore takes another 2–3 hours, during which the snails remain partially retracted and inactive.[39][40]

It has been reported that only one spermatophore is usually transferred during copulation in H. pomatia, so one animal functions as a male and one as a female in each mating. According to that report it is mostly the older snail who lay eggs, while younger function as males.[39]

In H. pomatia, mating takes place mostly from May to June, but often continues more sporadically up to the autumn[30][41]. The snails copulate usually with multiple mates[30][41]. Received foreign sperm may be stored for more than a year before fertilization.[31] Eggs are laid into a chamber dug in the soil by the parent 4–6 days after mating[39]. The eggs are formed only as the nest is built.[42] As in other pulmonates, the eggs are rich in galactogen produced by the albumen gland. The eggshell is partially mineralized, with crystals of calcium cabonate in a flexible membrane.[32] Clutch size is given in the literature within the range 3–93.[43] Hatching follows roughly 25–26 (range 18–31[41]) days after egg laying, but the snails remain additional 8–10 days in the nest.[39] An individual may lay more than one clutch per season.[41] The clutch size may be different in other species. The range reported for H. albescens (smaller than H. pomatia, with larger eggs) is only 7–22 eggs per clutch.[43]

The sperm morphology follows the basic pattern known from "pulmonates". Mitochondria are fused and form a continuous sheath around the flagellum. Large part of mitochondrial derivative is made up by a proteinaceous paracrystalline structure, in which there is a glycogen-filled canal. The canal runs helically along the flagellum, forming a so-called glycogen helix. There is only a single, loosely coiled glycogen helix in Helix.[44][45]

Life history

In H. pomatia, sexual maturity is reached after 2–4 (–6) overwinterings, i.e. at the age of 2–4 (–6) years, but this differs between localities as well as between snails from the same clutch.[39][46][47] Adult snails cease to grow and form a thickened lip around the aperture. However, the snails sometimes mate already shortly before the lip is formed. The life span of H. pomatia may reach 30 years in the wild.[47] However, in the wild, they mostly live for just 5–9 years.[46][47] The age of H. pomatia individuals can be estimated by counting growth interruptions on the shell and the number of layers deposited on the aperture margin lip.[46][48][47] The small species H. ceratina reaches maturity after 1.5–2 years and lives 4–5 years.[49]

The length of the life cycle is dependent on environmental conditions. The time from hatching to first egg laying can be shortened to just 12–13 months under optimal conditions in the captivity, mainly by skipping hibernation.[41]

Ecology and behaviour

Species of Helix live in a variety of habitats and under very different climatic regimes. Some species are exclusively found in open limestone rocky habitats (H. secernenda), while others tolerate acidic bedrock (e.g. H. pelagonesica) or live predominantly in forests (e.g. H. thessalica). Members of the genus occur from temperate rainforests (H. buchii) to semi-arid regions (Helix pronuba).[50][1]

Helix pomatia feeds mainly on live plants.[41][46] Observations showed preference for somespecific plant species[46] and avoidance of others.[41]

The activity of H. pomatia takes place mostly during the night, especially in juveniles.[41] During winter, Helix snails hibernate in the soil.

A homing behaviour has been observed in H. pomatia. The snails disperse during the season, but tend to return to their hibernation grounds towards its end.[51][46] It was also observed that the snails may migrate to specific sites for egg laying[citation needed] and aggregate for mating.[52] The snails are able to find their way back from distances of tens of meters.[51]

Several species (e.g. H. pomatia, H. figulina) build a thick, calcified epiphragm that closes the shell's aperture during hibernation or aestivation. The epiphragm is followed inside the shell by a few additional membranes made of dried mucus.[53] When the animal emerges from the dormancy, it discards the calcareous epiphragm using the posterior part of the foot.[citation needed]

Biotic interactions

Predators and parasites

Several bird species prey on Helix species. This has been observed in Phasianus colchicus, Burhinus oedicemus, Coracias garrulus, Lanius excubitor, corvids, and probably other birds.[54][55] Mammals prey on Helix, too. The known predators include hedgehog, mole, shrew, rodents (Rattus, Apodemus) and the wild boar.[56][55] The slow worm consumes molluscs, including Helix.[57] Helix snails are also attacked by various beetles and flies. The beetle predators belong to the families Carabidae and Lampyridae.[46][47][58] The predation by birds, small mammals, and beetles mostly affects juveniles.[46] Larvae of flies from several families attack Helix snails and may kill even adults (Phoridae, Muscidae, Sarcophagidae, Sciomyzidae).[59]

A well known facultative parasite of land snails, including Helix, is the nematode Phasmarhabiditis hermaphrodita.[60] The parasitic mite Riccardoella limacum is found on Helix species.[61] [62]

The kinetoplastid Cryptobia helicis lives in the bursa copulatrix of Helix pomatia.[31] The ciliate Tetrahymena limacis was also reported from Helix.[63]

Bacterial diseases of gastropods including Helix are known, but this field is not well researched.[64]

Influence on other species and the environment

Some bee species build their nests inside empty Helix shells (e.g. Rhodanthidium semptemdentatum).[citation needed]

Human use

A poster advertising a buy up of Helix snails for food processing (Czechia, 2008).

Some species, above all H. pomatia and H. lucorum, are collected for human consumption.[citation needed] The culinary use dates back several millennia and has been evidenced for several species across the genus' range. Mesolithic shell midden dated to 9370 ± 80 and 8110 ± 90 uncalibrated C-14 years bp and providing evidence of collecting was documented for H. pomatella in Abruzzo, Italy.[65]

Conservation

Most of the species included in the IUCN Red List are classified as Least Concern.[66] One of the species, H. ceratina, is critically endangered and the present known distribution is limited to a very small area near the Ajaccio airport.[67][49]

In the past, collection of wild H. pomatia for food led to fears of over-exploitation and the introduction of protection by law in several countries.[68]

List of extant species

Scientific name IUCN Red List
status 		Distribution Picture
Helix albescens
Rossmässler, 1839 		LC IUCN Azerbaijan, Armenia, Georgia, Russia, Ukraine[1]
Helix anctostoma
Martens, 1874 		Turkey (Nur Dagi and surroundings), Syria (northwestern)
Helix ankae
Korábek & Hausdorf, 2023 		Turkey (nortwestern Anatolia)[15]
Helix asemnis
Bourguignat, 1860 		LC IUCN
Helix borealis
Mousson, 1859 		DD IUCN Greece
Helix buchii
(Dubois de Montpéreux, 1840) 		Turkey (northeastern), Georgia, Armenia (northern)
Helix calabrica
Westerlund, 1876
Helix ceratina
Shuttleworth, 1843 		CR IUCN
Helix cincta
O. F. Müller, 1774 		LC IUCN Syria, Turkey (partly probably introduced), Cyprus (probably introduced), Greece (east Aegean, probably introduced), Croatia (introduced), Slovenia (introduced), Italy (introduced)[1][69]
Helix dormitoris
Kobelt, 1898 		LC IUCN
Helix engaddensis
Bourguignat, 1852 		Israel, West Bank, Jordan, Lebanon[70]
Helix fathallae
Nägele, 1901
Helix figulina
Rossmässler, 1839 		LC IUCN
Helix godetiana
Kobelt, 1878 		EN IUCN
Helix gussoneana
L. Pfeiffer, 1848
Helix kazouiniana
Pallary, 1939
Helix ligata
O. F. Müller, 1774 		DD IUCN
Helix lucorum
Linnaeus, 1758
Helix lutescens
Rossmässler, 1837 		LC IUCN
Helix melanostoma
Draparnaud, 1801 		Tunisia, Algeria, France (introduced), Spain (introduced)
Helix mileti
Kobelt, 1906
Helix nicaeensis
A. Férussac, 1821 		Turkey
Helix nucula
Mousson, 1854 		LC IUCN Lebanon, Syria, Cyprus,

Turkey (partly probably introduced)[1]

Helix pachya
Bourguignat, 1860
Helix pathetica
Mousson, 1854
Helix pelagonesica
(Rolle, 1898)
Helix philibinensis
Rossmässler, 1839 		LC IUCN Greece, North Macedonia, Bulgaria (southwestern), Albania (by Lake Prespa)
Helix pomacella
Mousson, 1854 		LC IUCN
Helix pomatella
Kobelt, 1876
Helix pomatia
Linnaeus, 1758 		LC IUCN
Helix pronuba
Westerlund & Blanc, 1879 		Egypt, Libya, Tunisia (southern), Greece (Crete and other islands, introduced)
Helix salomonica
Nägele, 1899 		Turkey (southeastern), Iran (western), Iraq (Kurdistan)
Helix schlaeflii
Mousson, 1859 		Albania (central and southern), Greece, North Macedonia (Galičica)[2][1]
Helix secernenda
Rossmässler, 1847 		LC IUCN Croatia, Bosnia and Herzegovina, Montenegro, Albania (northern)[2][1] Helix secernenda (Montenegro)
Helix straminea
Briganti, 1825 		LC IUCN Albania, North Macedonia (eastern), Italy Helix straminea (Italy)
Helix thessalica
O. Boettger, 1886 		LC IUCN
Helix valentini
Kobelt, 1891 		EN IUCN
Helix vladika
(Kobelt, 1898) 		LC IUCN Serbia, Montenegro

Fossil record

Several extinct taxa of Helix have been described:

  • Helix jasonis Mayer, 1856 (Ukraine: Sevastopol. Miocene: Tortonian[71])
  • Helix pseudoligata Sinzov, 1897 (Ukraine. Miocene: Sarmatian[71])
  • Helix kadolskyi Neubauer & Harzhauser, 2023 (nom. nov. for Helix toulai Kojumdgieva, 1969) (Bulgaria: Balchik. Miocene: middle Tortonian[72])
  • Helix barbeyana De Stefani in De Stefani et al., 1891
  • Helix krejcii Wenz in Krejci-Graf & Wenz, 1926
  • Helix mrazeci Sevastos, 1922
  • Helix sublutescens Wenz in Krejci & Wenz, 1926
  • Helix maeotica Steklov, 1966 (Russia: Chechnya: river Gums. Miocene: Maeotian[73]=?Tortonian[74])
  • Helix varnensis Toula, 1892 (Bulgaria: Varna. Miocene: Sarmatian)
  • Helix lucorum supralevantina Wenz, 1942 (Romania. Pliocene)

Some extant species are known from Quaternary deposits. The most studied species in this respect is H. pomatia, where the fossils have been used to document the earliest postglacial occurrences in Central Europe.[75] The earliest record in Czechia was dated directly by radiocarbon to 10,120-9,690 BP (but is likely a few hundred years younger);[76] fossils presumably older than 9,402–9,027 BP[77] or 9,403–9,003 BP[78] were found in Baden-Württemberg, Germany. Such records document the speed at which Helix species may extends their ranges by natural means of dispersal.

The quaternary land snail fossil record in more southern parts of Europe is scarce, but some records of Helix do exist. Helix figulina dated ~16,000 BP was recorded from the greek island Antikythera.[79] Helix borealis shells dated to 8,000–27,000 BP were reported from another island, Gavdos.[80]

Other records come from archaeological contexts.

Phylogeny

The phylogenetic relationships between Helix and related genera as well as the internal relationships within the genus have been so far studied only using partial sequences of mitochondrial genes and of the nuclear rRNA gene cluster.[81][1]

The cladogram shown is based on phylogenetic analyses of mitochondrial sequence data.[82][11][69][1]

Maltzanella
Helix

Synonyms

The following genus-level taxa are considered synonyms of Helix:

  • Callunea Scudder, 1882
  • Cochlea Da Costa, 1778
  • Coenatoria Held, 1838
  • Cunula Pallary, 1936
  • Glischrus S. Studer, 1820
  • Helicites W. Martin, 1809 (Established for fossils of Helix to distinguish them from extant members of that taxon. Invalid, available only for the purposes of the Principle of Homonymy (Art. 20))
  • Helicogena A. Férussac, 1821
  • Megastoma Scudder, 1882
  • Naegelea P. Hesse, 1918
  • Pachyphallus P. Hesse, 1918
  • Pentataenia A. Schmidt, 1855 (junior objective synonym)
  • Physospira Boettger, 1914
  • Pomatia Beck, 1837
  • Pomatiana Fagot, 1903
  • Pomatiella Pallary, 1909
  • Pseudofigulina P. Hesse, 1917
  • Rhododerma P. Hesse, 1918
  • Tacheopsis Boettger, 1909
  • Tammouzia Pallary, 1939
  • Tyrrhenaria P. Hesse, 1918

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