The vast diversity of land snail forms is insufficiently understood even in seemingly familiar taxa. This holds for Helix Linnaeus, 1758, a genus with several common edible species which comprises the largest Western Palearctic snails. The taxonomy of this genus, which has a centre of diversity in the eastern Mediterranean, has recently undergone significant changes, in both the delimitation of the genus itself and its species‐level systematics. Here, we compare the lineage diversity of Helix, as revealed by two mitochondrial markers, with the conclusions of the recently published morphology‐based taxonomic revision. For the molecular analysis, we assembled a representative data set covering almost all species of the genus as recognized by the mentioned revision. We obtained sequences not only from fresh and preserved soft tissues but also from dried tissue remains (some of them decades old) from shell collections. Our results show that the genus Helix, in the narrow sense proposed by recent studies, is paraphyletic because the genus Tacheopsis was unambiguously revealed as one of the tip branches of Helix. The monophyly of several species, as presently recognized, was not supported; partly, this may be attributed to a lineage diversity overlooked so far. This holds also for the type species of the genus, H. pomatia, which comprises at least one additional lineage. Greece, the Aegean and western Turkey is the core area for the diversity of Helix and its relatives, and the region is probably a major long‐term refuge for large Helicidae. The highest species diversity is found along the Alpide belt from the western Balkans to southern Turkey. The diversity of Helix in Europe, north of Greece and the Apennines, is a result of a single European radiation. Our data also suggest that past human activities likely influenced the present‐day distributions of some species.
Helix is a genus of large Western Palaearctic land snails, particularly diverse in the Mediterranean region. Despite the large 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 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.
Exact locations of glacial refugia are relevant for the study of contemporary biodiversity, not only as places less disturbed during the climatic changes but also as sources of rapid expansion of the biota after the Last Glacial cycle. If continuously inhabited over several of the Quaternary glacial cycles, the refugia are readily identifiable by the accumulated genetic diversity. However, the sources of the Holocene range expansion, particularly important for the emergence of present‐day bio‐ and phylogeographic patterns and for realistic estimation of species’ expansion rates, might have been located at the fringes of the glacial species ranges and lack unique lineages. This problem is pertinent when the variation is explored at slowly evolving genetic markers. We suggest that the location of such source refugia may be approximated by reconstructing the geographic location as a continuous trait evolving along the branches of a phylogenetic tree. We applied this approach, using the BEAST software, on two large southeast European land snail species: Caucasotachea vindobonensis and Helix thessalica. We found evidence for C. vindobonensis refugia in the western Balkans; notable is an apparently old refugium in Bosnia and Herzegovina. The plausible sources of the species’ Holocene range expansion, however, were located around the south‐western end of the Carpathians. Although the source areas were likely similar in H. thessalica, some expansion sources suggested by the analyses (e.g., Podolia, Ukraine) appeared implausible and driven by sampling clustered in that area. The applied approach allows for additional exploitation of the mitochondrial data gathered during the past two decades of animal phylogeography studies.
Aim: Were postglacial recolonizations facilitated by persistence close to the colonized areas rather than by dispersal ability allowing for colonization from distant sources? This question is particularly relevant for organisms with low active dispersal abilities and lacking specialized propagules. Here we identified glacial refugia of four Central European land snail species, including one for which a northern glacial refugium in the region was indicated by indirectly dated fossils.Location: Central Europe and the Balkans.Taxon: Pulmonate land snails: Monachoides incarnatus (Hygromiidae) and Helix pomatia, Helix thessalica and Caucasotachea vindobonensis (Helicidae). Methods:We used continuous phylogeographic analysis of mitochondrial sequence data to trace the origin of postglacially expanding lineages. We assessed the credibility of results from our focal species, M. incarnatus, by comparison to three other broadly sympatric land snail species. In addition, we performed direct radiocarbon dating of fossil shells to verify the presumed glacial presence of M. incarnatus on the territories of Czechia and Slovakia and to test the reliability of the available fossil data. Results:In three of the four species, the molecular data supported refugia located south of the Pannonian Basin, from the south-eastern Alps to the south-western Carpathians, but not more northerly. The direct radiocarbon dating resulted in younger dates than previously assumed. However, the molecular data also revealed unexpected, yet substantial postglacial southward expansions of M. incarnatus and C. vindobonensis into the Balkans.Main Conclusions: Neither the phylogeographic analyses nor the direct radiocarbon dating provided evidence for the glacial survival of studied land snail species in Central Europe. The refugia located adjacent to the Pannonian Basin were the most important source of postglacial expansions to Central Europe, but were also the source of the expansion southwards. Both climatic factors and biological interactions might explain why such southward expansions seem rather rare in Europe.
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