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Revision of Helix LINNAEUS, 1758 in its eastern Mediterranean distribution area, and reassignment of Helix godetiana KOBELT, 1878 to Maltzanella HESSE, 1917 (Gastropoda, Pulmonata, Helicidae)
Abstract and Figures
The species of the genus Helix Linnaeus, 1758 from the eastern part of Europe and
Near East are revised. Based on the investigation of thousands of shells and more
than 100 preserved specimens, 29 species are recognised as valid for this area, for
which more than 250 validly described names are available. The type specimens for
almost all valid nominal taxa could be located and documented. The morphology of
the genital organs of most of the species is described and illustrated. Geo-referenced
maps are supplied to show the distribution of all species found in the investigated
area. For some species, lectotypes are designated in order to stabilise nomenclature
and to prevent further misidentifications. Helix godetiana Kobelt, 1878 is excluded
from Helix, and included in Maltzanella Hesse, 1917.
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... is a genus of large land snails including more than 30 species distributed in Europe, the Caucasus, the Middle East, and northern Africa. It has recently been the subject of extensive taxonomic, phylogenetic, and phylogeographic studies (Neubert 2014, Korábek et al. 2015, Psonis et al. 2015, Fiorentino et al. 2016, and its diversity is now fairly well known, often to the intraspecific level. Despite the recent revisions, the last species currently considered valid was described in 1939 (Neubert 2014), indicating that this is a well-studied group with complete species records. ...
... It has recently been the subject of extensive taxonomic, phylogenetic, and phylogeographic studies (Neubert 2014, Korábek et al. 2015, Psonis et al. 2015, Fiorentino et al. 2016, and its diversity is now fairly well known, often to the intraspecific level. Despite the recent revisions, the last species currently considered valid was described in 1939 (Neubert 2014), indicating that this is a well-studied group with complete species records. ...
... Their shells resemble Helix escherichi O. Boettger, 1898, a species that was placed in the Anatolian clade of H. (Helix) in the phylogenetic analyses of Korábek et al. (2015). However, Kayseri is more than 300 km away from the known range of this species on the western Anatolian plateau (Neubert 2014). The combination of colouration, shell sculpture, white aperture margins, and shape of the columellar region of the specimens from Kayseri also resemble some Maltzanella individuals (Schütt 1976). ...
Helix escherichi O. Boettger, 1898 from western Anatolia is a poorly known species that was placed in the Anatolian Helix (Helix) clade in previous phylogenetic analyses. We sequenced additional individuals, but these were found to belong to Maltzanella P. Hesse, 1917, the sister genus of Helix. Examination of the type materials and the largest set of shells to date, as well as anatomical studies,
revealed that 2 different species have been confused under the name H. escherichi in the past. While the true H. escherichi is actually a member of Maltzanella, the previously sequenced specimens represent a hitherto undescribed species, here named Helix (Helix) ankae sp. nov. We discuss the morphological differences and phylogenetic relationships between Maltzanella, Helix (Pelasga), Helix (Helix) and propose a new subgenus, Aegaeohelix subgen. nov. for Helix godetiana Kobelt, 1878.
... schiLeyKo 2006, neuBert 2014, KóraBeK et al. 2015, Psonis et al. 2015. According to neuBert (2014), the most recent and undoubtedly the most detailed review of the genus in the eastern Mediterranean, the large, dark-lipped shells on Rhodes belong to H. fathallae and not to H. cincta, which constitute two closely related taxa(neuBert 2014, KóraBeK et al. 2015. ...
... schiLeyKo 2006, neuBert 2014, KóraBeK et al. 2015, Psonis et al. 2015. According to neuBert (2014), the most recent and undoubtedly the most detailed review of the genus in the eastern Mediterranean, the large, dark-lipped shells on Rhodes belong to H. fathallae and not to H. cincta, which constitute two closely related taxa(neuBert 2014, KóraBeK et al. 2015. ...
Rhodes, Symi and Chalki are three islands with different characteristics in the southeastern part of the Aegean Sea. In this work, we present their complete land snail fauna based on recent survey, samples from the collections of the Natural History Museum of Crete and a thorough bibliographic review. We report 67 land snail species from Rhodes, 34 species from Symi, and 32 from Chalki, with 4, 20 and 9 new records for each island, respectively. Also, one species has been added to the malacofauna of Greece, while many names, which were mentioned mainly in the 19th century, have been clarified, synonymised or abolished from the lists of the islands. Our results once again demonstrate the importance of the reproductive system as a tool for reliable assessment of the taxonomy of Aegean land snails, and that sampling during the rainy season should be the preferred methodological approach for collecting land snails in the Greek islands.
... La distinzione conchiologica tra queste due specie è, di solito, difficoltosa. Secondo Neubert (2014), tra le costole della teleoconca le conchiglie di H. melanostoma appaiono lisce mentre H. pronuba mostra piccoli granuli e nervature molto fini. Tuttavia, lo stesso autore (Neubert, 2014) ammette che nella Tunisia meridionale, Djerba in particolare, sono presenti popolazioni di difficile riconoscimento, con le due specie che potrebbero vivere in simpatria. ...
... Secondo Neubert (2014), tra le costole della teleoconca le conchiglie di H. melanostoma appaiono lisce mentre H. pronuba mostra piccoli granuli e nervature molto fini. Tuttavia, lo stesso autore (Neubert, 2014) ammette che nella Tunisia meridionale, Djerba in particolare, sono presenti popolazioni di difficile riconoscimento, con le due specie che potrebbero vivere in simpatria. ...
lleryana 37 (2) p. 103-111 luglio-dic 2019 novembre 2019 Riassunto Nel presente lavoro, gli Autori forniscono un loro origi-nale contributo alla conoscenza dell'Helix melanostoma Draparnaud, 1801 in Tunisia (Gastropoda: Stylommato-phora: Helicidae). In particolare, per questa specie si for-niscono ulteriori dati morfologici, anatomici, iconografi-ci, geonemici e bibliografici. Parole chiave Helicidae, Helix, Tunisia, morfologia, distribuzione. Abstract In the present work, the Authors show their original contribution to the knowledge of Helix melanostoma Draparnaud, 1801 in Tunisia (Gastropoda: Stylommato-phora: Helicidae). In particular, for this species further morphological, anatomical, iconographic, geonemic and bibliographic data are provided. Introduzione Il genere Helix Linnaeus, 1758 (Gastropoda: Stylomma-tophora: Helicidae) è uno dei più antichi taxon descritti comprendendo fin dall'inizio un enorme numero di spe-cie di molluschi terrestri anche molto diversi tra loro (co-me succedeva allora per quasi tutti i nuovi generi). Negli anni successivi, il perfezionamento degli studi si-stematici ha permesso di ridimensionare il numero di specie ad esso attribuite e di comprenderne meglio l'ori-ginario significato tassonomico anche in relazione alla specie tipo (H. pomatia Linnaeus, 1758). Attualmente, il genere Helix, suddiviso in diversi sotto-generi (Bank & Neubert, 2019), è diffuso in Europa, re-gione mediterranea e vicino Oriente, con diverse specie introdotte in altre regioni del mondo. Questo genere annovera, quindi, molluschi terrestri ca-ratterizzati morfologicamente dalla conchiglia quasi sempre globosa, liscia, con strie più o meno elevate con una densa granulazione sui giri superiori della teleocon-ca, fornita di cinque fasce di colore variabile di solito fuse tra loro in una banda sub e supraperiferica e columel-la bruno-nerastra o bianca. Negli organi genitali, il di-verticolo della borsa copulatrice è corto non superando mai il canale della borsa copulatrice, a volte può essere anche molto corto o assente, il muscolo retrattore del pe-ne è inserito sempre al terzo distale o al centro dell'epi-fallo, entrambe le papille peniali hanno una perforazio-ne centrale. Tra le specie del genere Helix, forniremo, nel presente la-voro, un contributo alla conoscenza delle popolazioni di H. melanostoma che vivono in Tunisia, dove questa specie è particolarmente diffusa. Materiali e metodi Conchiglie vuote ed esemplari vivi di H. melanostoma so-no stati raccolti a vista in diverse località della Tunisia e citate più avanti. Gli esemplari sono stati studiati per quanto riguarda dimensioni, morfologia della conchiglia e genitali. Le misure sono state effettuate con calibro di-gitale e lenti micrometriche. Le conchiglie sono state pu-*Via Principe di Paternò 3,
... Nei cesti adibiti alla vendita ripieni di questi elicidi vivi e mantenuti in attività da una costante umidità, in almeno tre occasioni (1980, 1989 e 2004) sono stati osservati anche alcuni esemplari vivi di Helix melanostoma Draparnaud, 1801. L'Helix melanostoma è un altro elicide edule diffuso in Francia, Marocco, Algeria, Tunisia e Malta (Neubert, 2014;Bank, 2017;Bank R.A. & Neubert E., 2020) ma introdotta anche in Spagna (Martínez-Ortí & Robles, 2003) e Italia (Hallgass & Vannozzi, 2010;Vuolo, 2010;Cianfanelli & Bodon, 2014). È specie termofila molto comune soprattutto nella vicina Tunisia dove si rinviene in quasi tutti gli ambienti naturali e antropizzati (Sparacio, 2018;Sparacio et al., 2019). ...
Alleryana 38 (2) p. 127-133 luglio-dic. 2020 novembre 2020 Riassunto La possibilità che le numerose attività dell'uomo possa-no facilitare l'introduzione di specie alloctone è un feno-meno noto e ha coinvolto, nei secoli, numerose specie animali e vegetali che si trovano, ai nostri giorni, lontano dal loro originario areale. Una di queste attività è il commercio dei molluschi terre-stri eduli praticato fin dall'antichità con lo spostamento di popolazioni viventi da una regione all'altra. Nel presente lavoro, sono riportate e documentate alcu-ne nuove e brevi osservazioni effettuate nei mercati sici-liani, in particolare della città di Palermo, sul commercio dei Molluschi terrestri eduli e le conseguenti implicazio-ni tassonomiche e faunistiche. Parole chiave Molluschi terrestri eduli, attività umane, specie allocto-ne, distribuzione geografica. Abstract [Short faunistic observations on the edible land snails trade in Sicily] The possibility that the numerous human activities can facilitate the introduction of alien species is known and has involved, over the centuries, numerous animal and plant species that are nowadays far from their original distribution area. One of these activities is the trade in edible terrestrial snails practiced since ancient times with the movement of living populations from one region to another. In the present work, some observations made in the Sicilian markets, and in the city of Palermo in particular , on the trade of edible terrestrial snails are reported and documented and brief considerations are made on their taxonomic and faunal implications. Introduzione Il problema delle specie alloctone, inserite in un ambien-te e in un areale diverso dall'originale, è stato da sempre facilitato e accentuato dalla presenza dell'uomo e delle sue attività. Il commercio, e quindi lo spostamento di merci da un paese all'altro, è stato, ed è ancora, una delle attività principalmente collegate a questo fenomeno. Per i molluschi terrestri, oltre allo spostamento passivo delle varie specie da un regione all'altra con le merci, più o meno casualmente, va considerato anche un trasporto attivo legato al commercio di alcune specie di esse per uso alimentare che avranno, quindi, maggiori possibilità di acclimatarsi nei paesi di destinazione, come già suc-cesso in diversi paesi del mondo. La facilità nella raccolta e/o nell'allevamento e le ottime proprietà nutritive sono le principali caratteristiche che hanno decretato il successo di alcune specie di molluschi terrestri nell'alimentazione dell'uomo fin dalla preisto-ria. Nell'area euro-mediterranea, per esempio, gli antichi Greci e i Romani le consumavano abitualmente come ci-bo e attribuivano a queste specie eduli anche proprietà mediche e afrodisiache. Citando qualche esempio, nel "De Re Coquinaria" di Marco Gavio Apicio (I secolo a.C.-I secolo d.C.) viene spiegato come bisogna ripulire le conchiglie (con latte e sale) prima di consumarle e vengono descritte quattro ricette per cucinarle. Nel Satyricon di Petronio (I secolo d.C.), il protagonista Encolpio, insieme a Gitone e Ascito, riceve l'invito a un banchetto organizzato nella casa del ricco Trimalcione e in un brano di quest'opera viene raccontato come il cuo-co servì "delle chiocciole su una graticola d'argento". Plinio il Vecchio, nella sua "Naturalis Historia" (77-78 d.C.), racconta di come i ricchi romani mangiassero le chiocciole provenienti da specifici allevamenti ("coclea-ri") realizzati su idea di Fulvio Lippino (nel 49 a.C.) che le importava anche da altre regioni. Anche Ateneo di Naucrati (II-III sec. d.C.) parla di que
... It is also present in the northern promontory of this mountain ridge (Bouaziz-Yahiatene et al., 2019). It is also noted that three species are present at all sampling stations, namely, Cornu aspersum which is a species of Mediterranean origin (Welter-Schultes, 2012), colonizing various environments, particularly entropized environments as well as agricultural environments (Neubert, 2014;Ballard et al., 2021), and Xerosecta cespitum and Cernuella virgata, which have a Mediterranean distribution and are considered the most abundant, colonizing open, xeric, and sunny habitats (Arrébola, 1995;Welter-Schultes, 2012). In conclusion, in the regions of Tizi-Ouzou and Bouira, 29 species of terrestrial mollusks belonging to 25 genera and 19 families were identified. ...
The snail and slug fauna of the Kabylia region (Bouira and Tizi-Ouzou) was studied during the years 2018/2019 and 2020/2021, focusing on seven stations belonging to three altitudinal levels (low, middle, and high altitude). A total of 5111 individuals were collected and 29 species were identified, with confirmation of the existence of Leiostyla anglica in the Kabylia region. Our results showed that altitude has a significant effect on the distribution and abundance of terrestrial gastropods in the study area – the low-altitude sites had the highest number of species (S = 21), while the high-altitude sites had the lowest species richness (S = 10).
... Cornu aspersum is present only as an alien species in North Macedonia (Cvetkovska-Gjorgievska et al. 2019). The native range of H. lucorum covers Balkan Peninsula, Asia Minor to Iran (Neubert 2014). However, its values as a bioindicator has not been assessed so far. ...
Heavy metal pollution of the environment is especially apparent within or around urban areas and around industrial facilities. This paper deals with the concentration of zinc, copper, manganese and cadmium in the soil, leaves of fodder plants and in the body parts of the Turkish snail (H. lucorum). The study included ten localities, arranged in four groups, situated in and around the city of Skopje. The first group consisted of five urban localities that were in the central city area. The second group included two suburban localities. The third group consisted of two rural sites. The fourth group represented one site that was located away from the metropolitan area (control site). Soils, leaves of four plant species (Cardaria draba, Lamium purpureum, Rumex crispus and Taraxacum officinale) and a sample of 10 adult snails were collected from each locality. Heavy metals content was determined by wet combustion and atomic absorption spectroscopy. Digestive gland was an organ with the highest amount of all heavy metals and with proven bioaccumulation for Zn and Cd; it showed a prominent correlation with soil concentrations for the four heavy metals but not with the examined plant species. The foot and shell of the snails had limited indicator values since they exhibited weak correlation with soil concentrations of Cu and Mn.
... The investigated species are the most widespread edible snails in Montenegro [14], and therefore important for human consumption. Regarding the growing interest in edible snails as a potential group of alternative foods, the determination of the factors that affect the content of metals is gaining in importance. ...
The objective of the present study is to determine the differences between the mineral content of various organs of Helix vladika and H. secernenda, the two most abundant edible snail species in Montenegro. The bioaccumulation of 12 examined elements (zinc, manganese, copper, aluminum, cadmium, lead, nickel, iron, chromium, lithium, selenium and mercury) was determined in the hepatopancreas, albumen gland, digestive tract, reproductive system, mantle, foot and shell from three sampling sites (Biogradska Gora, Nikšić and Malesija). The examined populations of H. vladika and H. secernenda showed a significant difference in their lithium and selenium contents. The levels of the most examined metals (Zn, Mn, Cu, Al, Cd, Pb, Se and Hg) varied significantly among organs. The digestive tract and hepatopancreas tend to bioaccumulate selenium and cadmium. The general mean concentration of cadmium in the examined snail tissues exceeded the maximum allowable level at the Biogradska Gora and Malesija sites. Therefore, the use of the Montenegrin edible snails collected from the wilderness for human consumption seems to be limited by their higher bioaccumulation capacity for toxic elements such as cadmium.
... The species, for which a neotype was recently fixed (Giusti et al., 2015), is native to the northern Levant (Korábek et al., 2021), but was introduced to the northern Adriatic basin in ancient times (cf. Welter-Schultes, 2012;Neubert, 2014). Linnaeus, 1758Mienis, 1988cBeckmann, 1992 sub Helix lucorum; Giusti et al., 1995sub Helix lucorum. ...
The first record in Malta, and the second record in Europe, of the tropical terrestrial gastropod Paropeas achatinaceum (L. Pfeiffer, 1846) is presented, with an annotated bibliography of the allochthonous (and non-marine) gastropod fauna hitherto recorded from the Maltese archipelago.
In the west of Ukraine, the Turkish snail Helix lucorum was first discovered in 2020 at one of the household plots in Chortkiv, Ternopil region. It was found that the owner of the house specially brought snails from France for his children. Until the beginning of the 21st century, H. lucorum was known in Ukraine only from the mountainous Crimea, where it was presumably brought by Greek colonists. The distribution area of this species in Crimea gradually expanded, in particular, due to targeted introductions. Since 2010, single population of H. lucorum have been recorded in Ukraine outside of Crimea. Judging by the shell colouration, only some of them could have been founded by individuals from the Crimea, with almost unnoticeable spiral banding and distinct radial streaks. Other populations are of unclear origin, and their founders may have been brought from different parts of the present range of H. lucorum. In particular, shells with wide dark, often fused bands and a distinct light gap between them on the shell periphery are characteristic of most known introduced European populations of H. lucorum, and are also common in the Krasnodar region of Russia, from where they could have been introduced to the east of Crimea (Kerch Peninsula).
A b s t r a c t : In this work, all valid species and lists of primary synonyms of terrestrial and freshwater Gastropoda and Bivalvia known from the Arabian Peninsula are given. The type specimens of many of the species are illustrated. All species are described and anatomical details are given whenever possible. Zoogeographic relationships of the malacofauna of the area are discussed. A hypothesis explaining the time scales and pathways of colonisation of the Arabian Peninsula by molluscs is given. Ecological data are added, where known, to further the knowledge of autecology and distributional patterns. The following taxa are described as new to science: Stenothyra arabica n. sp., Arabivitrina jansseni n. sp., Cerastus schweinfurthi brunneus n. ssp., Cerastus schweinfurthi apicostatus n. ssp., Oxychilus (Costoxychilus) n. subgen., Oxychilus (Costoxychilus) profundus n. sp., Levantina (Laevihelix) n. subgen., Levantina (Laevihelix) symensi n. sp., Levantina (Laevihelix) asira n. sp., Levantina (Laevihelix) asagittata n. sp., Levantina (Laevihelix) semitecta n. sp. Additionally, several taxa are recorded as new to the malacofauna of the Arabian Peninsula: Assiminea nitida nitida (Pease, 1864), Gastrocopta klunzingeri (Jickeli, 1873), Boysia boysii (L. Pfeiffer, 1846), Lauria cylindracea (Da Costa, 1778), Toltecia pusilla (Lowe, 1831), Cecilioides aff. tumulorum (Bourguignat, 1859), Cecilioides acicula (O.F. Müller, 1774). New generic or subgeneric combinations are introduced: Homorus splendens (Thiele, 1910), Homorus arabica (Connolly, 1941), Streptostele (Raffraya) scotti (Connolly, 1941). Lectotypes for the following species are designated: Hydrobia lactea (Küster, 1852), Bithynia badiella (Küster, 1853).
Helix is a genus of large Western Palaearctic land snails, particularly diverse in the Mediterranean region. Despite size and attractiveness of its members, it has an unsettled taxonomy, and no data are available on its intrageneric phylogenetic relationships. One of the problematic Helix taxa is the widely distributed, economically important, and conchologically very variable H. lucorum. Two distinct forms may be encountered under this name in the Apennine Peninsula: a typical one in the north, and a form originally described as Helix straminea in central and southern Italy. To evaluate the status of H. straminea and its relationships to Italian and Balkan Helix fauna, we combined shell morphology, geometric morphometrics, and phylogenetic analysis based on two mitochondrial genes. Distribution data were improved by drawing information from unambiguously identifiable photographs posted online. Based on our results, Helix straminea is redescribed as a separate species, and we find it to have a disjunctive trans-Adriatic range with closest relatives in the western Balkans. We provide an insight into relationships and intraspecific variability within Helix, a first step towards a comprehensive revision of the genus. On the example of Italian Helix fauna, we demonstrate how understanding of snail zoogeography may change with improving taxonomy.
Two species of genus Helix Linnaeus, 1758 (Mollusca: Gastropoda: Helicidae) endemic to the Caucasus region are known from
Georgia and northeastern Turkey: Helix buchii Dubois de Montpereux, 1839 and the recently-described but disputed Helix goderdziana
Mumladze, Tarkhnishvili and Pokryszko, 2008. The latter species is the largest land snail throughout non-tropical Eurasia. We compared
shell shapes and genital morphology of the two species. We analyzed mitochondrial COI and nuclear 18S ribosomal RNA and ITS1 gene
fragments in 39 specimens of H. buchii and H. goderdziana from ten locations from the entire distribution range of these species, together
with 13 specimens of the widespread H. lucorum Linnaeus, 1758 and H. pomatia Linnaeus, 1758. Based on shell morphology alone, most
of the individuals of the two species can be discriminated using multivariate approaches. The species have different fl agellum/diverticulum
ratios, and the foot coloration is a fully diagnostic morphological character. Molecular genetic analysis revealed little variation in 18S+ITS1
fragment, and eleven COI haplotypes. Phylogenetic analyses support reciprocal monophyly of H. buchii and H. goderdziana. The genetic
distances signifi cantly correlate with the geographic and morphological distances; correlation of morphological distances with geography is
insignifi cant. The basal lineages of both species are found within two distinct glacial refugia, a result which matches the separation of eastern
and western evolutionary lineages of other relicts of the Western Caucasus. The present distribution of H. goderdziana coincides with the
expected refugial borders, whereas H. buchii is likely to have extended its geographical range since the last glaciation.
Types of land and freshwater mollusc species described from material collected on the expedition are listed and figured. -from Authors
Phylogenetic relationships of the Aegean Helix cincta and Helix nucula with congeneric species found in Greece were inferred using mitochondrial DNA sequence data. Twenty-three specimens from mainland Greece, Aegean Islands, Cyprus and North Africa were analysed, revealing that (1) H. nucula is monophyletic, (2) H. cincta from Greece and Cyprus is paraphyletic, and so questions arise regarding the taxonomy of this species, and (3) H. cf. cincta from Tunisia might be considered as a distinct evolutionary lineage. Moreover Helix valentini, an endemic species of Kalymnos Island group, is clustered within the lineage of H. c. anatolica, so supporting the synonymy of the two ?species?, and the elevation of H. c. anatolica to species level. Hence, our results stress the need for a taxonomic reconsideration of H. cincta in the Aegean Sea, indicating that sequence data can prove useful in overcoming taxonomic issues at both species and subspecies level.