identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
16294ED587C75DD4B93A9695E52F01FC.text	16294ED587C75DD4B93A9695E52F01FC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helicidae Rafinesque 1815	<div><p>Family Helicidae Rafinesque, 1815</p><p>References.</p><p>Schileyko 1978; Nordsieck 1987; Razkin et al. 2015.</p><p>Type genus.</p><p>Helix Linnaeus, 1758</p></div>	https://treatment.plazi.org/id/16294ED587C75DD4B93A9695E52F01FC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
E8F214919FE65BA79929E5A2E592E57D.text	E8F214919FE65BA79929E5A2E592E57D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helicinae Rafinesque 1815	<div><p>Subfamily Helicinae Rafinesque, 1815</p><p>References.</p><p>Hesse 1918; Schileyko 1978; Razkin et al. 2015.</p></div>	https://treatment.plazi.org/id/E8F214919FE65BA79929E5A2E592E57D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
2F68F9EC1CB55F17843C7750C5221C0B.text	2F68F9EC1CB55F17843C7750C5221C0B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helicini Rafinesque 1815	<div><p>Tribe Helicini Rafinesque, 1815</p><p>References.</p><p>Razkin et al. 2015; Neiber et al. 2022.</p></div>	https://treatment.plazi.org/id/2F68F9EC1CB55F17843C7750C5221C0B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
10EF5060A5435B47834E2B5E80D3F675.text	10EF5060A5435B47834E2B5E80D3F675.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix (Helix) borealis Mousson 1859	<div><p>Helix (Helix) borealis Mousson, 1859</p><p>Figs 28, 29</p><p>References.</p><p>Neubert 2014; Korábek et al. 2021.</p><p>Description.</p><p>Shell (Fig. 28) mid-sized, globular to slightly conical; aperture rounded and in the more globular populations spacious; umbilicus missing; protoconch small; shell surface rather smooth, with fine growth lines; basal colour ranging from nearly white to pale brown; banding pattern varies from completely reduced bands to well-developed and contrasting; the upper three bands may be separated only on the upper whorls or well separated up to the last whorl; aperture margins and the parietal region brown and much darker than the rest of the shell, the colour ranges from vivid pale brown with orange tones (Ionian Islands, Epirus) to dark with purple tones (Peloponnese, Evia); mantle margins pale. Animal (Fig. 29) pale grey or pale brown, often but not always with a dark back of the foot.</p><p>Distribution and habitat.</p><p>Typical Helix borealis is distributed over much of Peloponnese, the east of central Greece (Aetolia, Acarnania, Phocis, Evrytania) and in western Epirus (Fig. 30). It also occurs on the Ionian Islands. The inhabited environments include as disparate types as rocky phrygana, pine forests on sand dunes at the sea level, and open fir forests in the mountains. Geophilous. Besides typical H. borealis, there are two other, rather divergent lineages in Greece. One occurs on northern Evia and Northern Sporades. It is found in rocky habitats on limestones. The second has a small range in the mountains of western Crete.</p><p>Remarks.</p><p>Some H. borealis populations have a globular shell with shape resembling that of H. figulina, but with a smoother, not regularly ribbed surface. There is also always some brown colouration of the apertural margins. In colour, H. borealis is often very similar to H. pelagonesica, but the two differ in shell shape and in the small protoconch of H. borealis . The globular-shelled form of H. pelagonesica from the tip of Sithonia (see below) is similar to H. borealis also in the shell shape but has a larger protoconch. In Epirus, H. borealis occurs syntopically with H. schlaeflii . Although the colouration may be similar there, H. schlaeflii is much larger and has a large protoconch.</p><p>In the mitochondrial phylogeny (Fig. 2), H. borealis appears as possibly polyphyletic. However, the relevant parts of the tree are unresolved, so this is likely just an issue of low phylogenetic signal.</p></div>	https://treatment.plazi.org/id/10EF5060A5435B47834E2B5E80D3F675	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
285A409F94F15D17A9B989E5D43AC2AD.text	285A409F94F15D17A9B989E5D43AC2AD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix (Helix) Linnaeus 1758	<div><p>Subgenus Helix Linnaeus, 1758</p><p>References.</p><p>Hesse 1908; Neubert 2014; Korábek and Hausdorf 2023.</p></div>	https://treatment.plazi.org/id/285A409F94F15D17A9B989E5D43AC2AD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
57FAD28A002056138CAE28A454E647CE.text	57FAD28A002056138CAE28A454E647CE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix (Helix) lucorum Linnaeus 1758	<div><p>Helix (Helix) lucorum Linnaeus, 1758</p><p>Figs 31, 32</p><p>References.</p><p>ICZN 2002; Neubert 2014; Korábek et al. 2018, 2023 a; Korábek 2020.</p><p>Description.</p><p>Shell (Fig. 31) mid-sized to large, broadly conical to depressed conical, with relatively narrow body whorl and low aperture; umbilicus present in juveniles but usually fully closed in adults; protoconch small relative to shell size; shell surface smooth; basal colour of the shell whitish, but usually largely covered by dark brown fused bands; characteristic are transverse dark bands marking growth interruptions; conspicuous whitish band along the shell periphery; aperture margins straight at the upper insertion but becoming reflected towards the lower lip and columella, brown inside; columellar margin oblique and often with an internal ridge. Animal (Fig. 32) uniformly brown including the mantle margins; diaphragm paper-like.</p><p>Distribution and habitat.</p><p>Found commonly in northern Greece except for west of the Pindos Mts. (Fig. 33) but may also be found synanthropically elsewhere (e. g. Peloponnese). Broadly distributed and somewhat invasive species, currently extending its range in Europe. In the northeastern Aegean (Samothraki, Lesvos) a different morphotype with globular shells occurs. Helix lucorum lives in various shrubs and herbs and in deciduous forests, but avoids dry Mediterranean types of habitats (phrygana, maquis, exposed rocks). Commonly synanthropic, in some areas exclusively so. Often climbs on vegetation.</p><p>Remarks.</p><p>Helix lucorum is very variably coloured and also details of the shell and aperture shape vary. The forms present in mainland Greece are typically very darkly coloured, with a white band on the periphery. The animal is also rather darkly coloured. It has a similar shell shape as H. straminea and H. pelagonesica, but besides the colour it differs in smooth shell surface and presence of strong transverse banding, visible at least at the bottom of the shell. We found H. lucorum syntopic with H. figulina, H. thessalica and H. philibinensis .</p></div>	https://treatment.plazi.org/id/57FAD28A002056138CAE28A454E647CE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
393122E946E45C2F8F51AFF7F91040D8.text	393122E946E45C2F8F51AFF7F91040D8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix (Helix) pelagonesica subsp. pelagonesica (Rolle 1898)	<div><p>Helix (Helix) pelagonesica pelagonesica (Rolle, 1898)</p><p>Figs 4, 5, 6, 7, 8, 9</p><p>References.</p><p>Neubert 2014; Korábek et al. 2022.</p><p>Description.</p><p>Shell (Fig. 4) middle-sized, conical with relatively flat base; individual whorls low and tightly coiled; deep suture giving the shell a stepped look; umbilicus fully covered or there is very narrow slit between the shell bottom and the reflected columellar margin of the aperture; protoconch large, making the apex blunt; aperture small and low, with oblique columella; aperture margins and the parietal area dark brown with purple hue; shell surface usually with fine ribs, pale brown, but often corroded even in live individuals; four bands typically present (because 2 + 3 fuse), brown with reddish or purple tones; bands interrupted by fine growth lines and lightly coloured ribs; aperture margins straight, columellar margin thickened, margins and the parietal area dark purple-brown. Animal grey (Fig. 5).</p><p>Distribution and habitat.</p><p>It has a small range (Fig. 10), within which it occurs sporadically, even though it can be locally abundant. In the north, it was found in the hills on both sides of the valley of Vardar / Axios (Kilkis Regional Unit), e. g. west of Polykastro or near the Dojran Lake. The northernmost record is from northeast of Negotino in North Macedonia. To the south, H. p. pelagonesica is distributed over Chalkidiki, additional populations are known from foothills of Mount Olympus and from Pelion near Volos (in Makrinitsa). The type locality is the island of Kyra Panagia (Pelagonisi) in the Northern Sporades. The characteristic habitat is a loose shrub with Quercus ( Q. coccifera or similar species) and Paliurus spina-christi . It seems to rest relatively frequently on branches of shrubs.</p><p>Remarks.</p><p>Neubert (2014: fig. 224) figured a shell of a snail from the tip of the Sithonia peninsula, Chalkidiki (NMBE 524743), which was quite similar to the brightly coloured form of H. schlaeflii from Corfu. He discussed this population with the first author in 2011 and back then both eventually identified it as H. schlaeflii, speculating it was possibly an anthropogenic occurrence. Following the examination of new material and phylogenetic analysis (Fig. 1), we currently consider the form living at the tip of Sithonia to be a variety of H. p. pelagonesica . Its globular shell shape is a clear difference to typical H. p. pelagonesica, but this is a question of the height of individual whorls driven probably by the angle of the columella, so a change in a single parameter of the shell geometry is responsible for the difference. In some shells, the columella is not vertical and the difference in shell shape is much weaker (whorls are lower, suture deeper). The colouration of the shell and animal is identical to typical H. p. pelagonesica, whose sample was previously sequenced from a locality on the northern foot of a limestone hill near Porto Koufo (just over a kilometre from where we collected). The strongest argument for a separate taxon for the snails from the tip of Sithonia is the mitochondrial lineage found there, which does not fall within or sister to the clade comprising typical H. p. pelagonesica samples. However, the conchological similarities as well as geographic proximity suggest that there is a gene flow; the divergent mtDNA might be a relic of a period of separation that was then followed by merging back with H. p. pelagonesica . We found this form (Figs 7, 8) on the limestone hills on both sides of the Maratia Beach at the southernmost tip of the peninsula. This habitat contrasts with geological conditions on the rest of Sithonia, which is covered by substrates generally unfavourable for snails, in large part by granite and other plutonic rocks (Christofides et al. 2007). However, a strict dependence on limestone is unlikely, because Neubert (2014) also found a locality of the same form north of the limestone occurrences (17 road km south of Sarti, ~ 39.993°N, 23.959°E; NMBE 524745). Typical H. p. pelagonesica tolerates non-limestone substrates.</p></div>	https://treatment.plazi.org/id/393122E946E45C2F8F51AFF7F91040D8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
4B4C8C92A9255699BF32CDACC2C95420.text	4B4C8C92A9255699BF32CDACC2C95420.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix (Helix) pelagonesica subsp. thembones Korabek, Jurickova & Hausdorf 2025	<div><p>Helix (Helix) pelagonesica thembones Korábek, Juřičková &amp; Hausdorf subsp. nov.</p><p>Figs 11, 12, 13</p><p>Type material.</p><p>Holotype: (Fig. 11 A) Greece • diameter 36 mm, height 37 mm; Thessaly, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=21.7468&amp;materialsCitation.latitude=39.3574" title="Search Plazi for locations around (long 21.7468/lat 39.3574)">Morfovouni</a>, hill on the NW outskirts of the village; 39.3574°N, 21.7468°E; 15 Apr. 2023; O. Korábek et al. leg.; NMP P 6 M 44038. The type locality is a northern side of the top of a small hill at the northwestern margin of Morfovouni (Μορφοβούνι; formerly Βουνέσι), Karditsa regional unit, Thessaly (Θεσσαλία) . Paratypes: Greece • 15 shells, 9 bodies in ethanol; Thessaly, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=21.7468&amp;materialsCitation.latitude=39.3574" title="Search Plazi for locations around (long 21.7468/lat 39.3574)">Morfovouni</a>, hill on the NW outskirts of the village; 39.3574°N, 21.7468°E; 15 Apr. 2023; O. Korábek et al. leg.; NMP P 6 M 42943 • 2 shells; <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=21.7468&amp;materialsCitation.latitude=39.3574" title="Search Plazi for locations around (long 21.7468/lat 39.3574)">Morfovouni</a>, hill on the NW outskirts of the village; 39.3574°N, 21.7468°E; 15 Apr. 2023; O. Korábek et al. leg.; ZMH 141525 • 12 shells; south of Mouzaki, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=21.6642&amp;materialsCitation.latitude=39.4135" title="Search Plazi for locations around (long 21.6642/lat 39.4135)">by a road to Porti</a>; 39.4135°N, 21.6642°E; 15 Apr. 2023; O. Korábek et al. leg.; NMP P 6 M 42942 • 6 shells; <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=21.7318&amp;materialsCitation.latitude=39.3747" title="Search Plazi for locations around (long 21.7318/lat 39.3747)">between Morfovouni and Ellinopyrgos</a>, slope above a forest road; 39.3747°N, 21.7318°E; 16 Apr. 2023; O. Korábek et al. leg.; NMP P 6 M 42944 • 1 shell; rocks northwest of Morfovouni; 18 Jun. 1985; B. Hausdorf leg.; ZMH 137861 • 1 shell; Morfovouni, crystallic rocks; 18 May 1995; P. Subai leg.; NMBE 524759 • 3 shells; Morfovouni, northern fringes, limestone rocks; 12 May 1997; P. Subai &amp; M. Szekeres leg.; NMBE 524756 • 1 complete individual in alcohol; <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=21.7464&amp;materialsCitation.latitude=39.3567" title="Search Plazi for locations around (long 21.7464/lat 39.3567)">Morfovouni, northern fringes</a>, east-oriented limestone rocks; 39.3567°N, 21.7464°E; 23 Apr. 2003; P. Subai leg.; NMBE 524760 • 1 shell; Mouzaki, rocks 1.5 km towards Kryonéri; 19 Jun. 1985; B. Hausdorf leg.; ZMH 137863 • 1 shell; 800 south of Mouzaki, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=21.6659&amp;materialsCitation.latitude=39.4153" title="Search Plazi for locations around (long 21.6659/lat 39.4153)">junction to Porti</a>; 39.4153°N 21.6659°E; 10 Apr. 1988; P. Subai leg.; NMBE 524757 • 1 shell; 1.6 km south of Mouzaki, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=21.663&amp;materialsCitation.latitude=39.4113" title="Search Plazi for locations around (long 21.663/lat 39.4113)">junction to Porti</a>; 39.4113°N 21.6630°E; 23 Apr. 2003; P. Subai leg.; NMBE 524746 • 1 shell; Elati, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=21.5179&amp;materialsCitation.latitude=39.571" title="Search Plazi for locations around (long 21.5179/lat 39.571)">7.3 km in direction to Pertouli and 2.9 km east on field road</a>, coniferous forest, on limestone rocks; 39.5710°N, 21.5179°E; 23 Jul. 1990; P. Subai leg.; NMBE 524754 • 1 shell; Elati, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=21.5179&amp;materialsCitation.latitude=39.571" title="Search Plazi for locations around (long 21.5179/lat 39.571)">7.3 km in direction to Pertouli and 2.9 km east on field road</a>, coniferous forest, on limestone rocks; 39.5710°N, 21.5179°E; 17 May 1991; P. Subai leg.; NMBE 524755 • 1 shell; Elati, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=21.5179&amp;materialsCitation.latitude=39.571" title="Search Plazi for locations around (long 21.5179/lat 39.571)">7.3 km in direction to Pertouli and 2.9 km east on field road</a>, coniferous forest, on limestone rocks; 39.5710°N, 21.5179°E; 16 May 1995; P. Subai leg.; NMBE 524754 .</p><p>Diagnosis.</p><p>Helix pelagonesica thembones differs from H. p. pelagonesica in the pale brownish shell with inconspicuous bands and paler apertural margins. Helix pelagonesica thembones has globular shells, whereas most populations of H. p. pelagonesica have more conical shells with a relatively smaller aperture.</p><p>Description.</p><p>Middle-sized globular shell (diameter 32–42 mm, height 33–43 mm) resembling in shape Helix pomatia; no umbilicus; protoconch large (~ 5–6 mm in diameter at 1 whorl); aperture semicircular; aperture margins and especially columella may be darker than the rest of the shell, with columella often meat-coloured to brown; shell surface relatively smooth, with only fine irregular riblets, pale brownish, sometimes with pinkish hue when alive; bands are inconspicuous, not much darker than the background, but present (2 + 3 and 4, sometimes weakly also 5, positioned close to the shell axis); apertural margins straight, slightly reflected only towards columella; animal pale brown to greyish brown, mantle margins pale.</p><p>Genital system (Fig. 13) with proximal epiphallus (sensu Korábek and Hausdorf 2023) much shorter than distal epiphallus and penis combined (4–5 vs 12–15 mm); flagellum well-developed (38–49 mm), mucous gland with many branches and of similar length as or longer than the dart sac; distal pedunculus of bursa copulatrix considerably thicker than the proximal part and leading to a well-developed diverticulum; diverticulum shorter than the proximal pedunculus (12–17 vs 21–32 mm), thick and flattened, narrowing towards tip (heavily swollen in the individual with a spermatophore in the pedunculus); distal genitalia (penis, epiphallus, vagina, the distal-most pedunculus) white.</p><p>Etymology.</p><p>Named after the opening track of the Alice in Chains’ 1992 album “ Dirt ” as a little reminder to all in power that they are also “ gonna end up a big ol’ pile of them bones ”. Noun in apposition.</p><p>Distribution.</p><p>The species was found on the band of hills that directly adjoin the Plain of Thessaly from the west (Fig. 14). The southern-most known locality is Morfovouni, the northernmost is ca 7 km north of Elati, so the known range extent is only slightly more than 30 km.</p><p>Ecology.</p><p>At the type locality, the greatest concentration of individuals was in an area overgrown by Phlomis fruticosa L. (Fig. 15). The same plant dominated vegetation in a small clearing in an oak forest northwest of Morfovouni, where we also found the species. The biology of the subspecies has not been studied. A spermatophore found in the bursa pedunculus of one of the dissected individuals indicates that mating takes place (at least partially) in April.</p><p>Remarks.</p><p>The majority of the Helix pelagonesica populations differ in a conical shell shape from the plesiomorphic more globular shell shape in Helix . This conical shell shape characterises not only the populations in the contiguous main range of the species in Greek Macedonia and North Macedonia, for which the name Helix pelagonesica vardarica Knipper, 1939 has been proposed, but also isolates of the species in Thessalia, for which the name Helix volensis Kobelt, 1906 was proposed, and the population from the island Kyra Panagia (= Pelagonisi) in the Northern Sporades, for which the species name was originally proposed. The populations with conical shell shape are monophyletic in the mitochondrial tree (Fig. 2). They are hardly differentiated. Therefore, the names given to these populations were synonymised (Neubert 2014). In contrast, there are populations from the surroundings of Morfovouni in western Thessaly and from the tip of Sithonia, the middle “ finger ” of the Chalkidiki peninsula in Macedonia, which differ from other populations of Helix pelagonesica so strongly in a globular shell shape that Neubert (2014) misidentified them as H. schlaeflii . It turned out that these globular populations are not related to H. schlaeflii (Fig. 2) but are the next relatives of H. pelagonesica . They were obviously isolated from the main lineage of H. p. pelagonesica before its shell became conical. However, the globular populations had different fates. There is probably no gene flow between the populations in the isolate in western Thessaly and those in the main range of the species because of the large geographic distance. This is supported by the distinctive pale shell of these populations (compare Figs 4, 5 to Figs 11, 12). Therefore, we suggest separating the populations from western Thessaly as a distinct subspecies Helix pelagonesica thembones from the other populations of Helix pelagonesica .</p><p>In contrast, the population from the tip of Sithonia came probably in secondary contact with the conical populations when they colonised the rest of Sithonia. We suppose that the globular population from the tip of Sithonia is connected by gene flow with the neighbouring H. pelagonesica populations, which are only about a kilometre apart, and that the differentiated populations are in the process of merging. Therefore, we currently classify this population as H. p. pelagonesica despite the still recognisable difference in shell shape and the deep split between the mitochondrial lineages of these populations (Fig. 2). The relationships between the H. pelagonesica populations should be re-examined by multi-locus markers. If it would turn out that gene flow between the globular population from the tip of Sithonia and the neighbouring populations remained restricted despite their proximity and that their genetic differentiation based on the multi-locus data reflects still the deep split between their mitochondrial lineages, it should be re-considered to classify also the population from the tip of Sithonia as a distinct subspecies.</p><p>We did not find any noteworthy differences between the genital system of H. pelagonesica thembones, typical H. pelagonesica and the form from the tip of Sithonia. The latter might have a somewhat shorter flagellum (35–37 mm, n = 2) compared to H. pelagonesica thembones (38–49 mm, n = 5), but both overlap with the typical H. pelagonesica (35–45 mm, n = 3).</p><p>Helix pelagonesica thembones is usually smaller than H. schlaeflii and has a smoother shell surface without the irregular whitish patterns typical for the latter. It differs from individuals of H. borealis with reduced banding in much paler colouration of the aperture margins and a much larger protoconch. Furthermore, the mucous glands in H. borealis are shorter, reaching only to the half of the dart sac, and diverticulum of bursa copulatrix may be much shorter in some individuals. Compared to H. philibinensis, which is usually distinctly banded, H. pelagonesica thembones has a higher aperture.</p></div>	https://treatment.plazi.org/id/4B4C8C92A9255699BF32CDACC2C95420	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
5F606D81DAF05B4CBD92444959C5BC59.text	5F606D81DAF05B4CBD92444959C5BC59.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix (Helix) philibinensis Rossmassler 1839	<div><p>Helix (Helix) philibinensis Rossmässler, 1839</p><p>Figs 25, 26</p><p>References.</p><p>Neubert 2014; Korábek et al. 2022.</p><p>Description.</p><p>Shell (Fig. 25) small, globular or slightly conical; body whorl expanded but bent downwards, so the aperture is relatively small; no umbilicus; protoconch middle-sized, but relatively large to shell size, making the apex blunt; shell surface with some growth lines but quite smooth; shell colour is whitish to pale brown; bands variously developed, brown on brown shells and with red or purple hue on whitish shells; all bands often separate on older whorls but may fuse towards the aperture; aperture margins straight, usually brown to some degree at least at the columella which may be very dark. Foot (Fig. 26) pale grey with dark brown back.</p><p>Distribution and habitat.</p><p>Distributed from Lake Prespa (northwestern Greece) to the island of Thasos (northeastern Greece) (Fig. 27). The southernmost populations occur on Chalkidiki (the base of the peninsula) and in the south of Western Macedonia (by Servia). In the north, it reaches almost to Skopje in North Macedonia and to Plovdiv in Bulgaria. The core of the range is in Central Macedonia, elsewhere the distribution is highly fragmented. Typically living in open, shrubby habitats, often on rocky slopes, but sometimes found even on forested valley bottoms. It appears to be indifferent to bedrock, as we found it on limestones, but also on granite and gneiss. Though typically resting on the ground, we saw it climbing onto vegetation during rain.</p></div>	https://treatment.plazi.org/id/5F606D81DAF05B4CBD92444959C5BC59	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
263DCD36E7BA5EBBB576733FE6049051.text	263DCD36E7BA5EBBB576733FE6049051.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix (Helix) schlaeflii Mousson 1859	<div><p>Helix (Helix) schlaeflii Mousson, 1859</p><p>Figs 16, 17</p><p>References.</p><p>Neubert 2014; Korábek et al. 2022.</p><p>Description.</p><p>Shell large, globular to conical; umbilicus sometimes slit-like but usually completely covered; protoconch large; shell surface with irregular ribs; basal colour whitish or very pale brown, more rarely the whole shell is brown; banding often reduced with upper bands 2 and 3 fused or partially fused and the lower two fuzzy and faint, but some populations have well developed, contrasting, reddish brown bands; bands with irregular whitish interruptions; aperture margins straight, only slightly reflected towards the columella; apertural lips and in particular the columellar triangle orange- or meat-brown to violet-brown, but the colouration is sometimes only faint and is missing or only weakly developed in the parietal area; mantle margins pale; animal very pale brown or grey to yellowish.</p><p>Distribution and habitat.</p><p>Common in Epirus and adjacent Western Macedonia (eastern limits uncertain, but apparently west of Kastoria, Neapoli and Grevena); occurs also on Kerkyra (Corfu). The range of the species as currently accepted extends to central Albania and up to the Galičica Mountains between the lakes Ochrid and Prespa (Fig. 18). In Greece, it occurs in a range of habitats, from Mediterranean-type shrubby vegetation on limestones in low altitudes in the west to more temperate landscapes on sandstones in the east, but occurs also in pine forests as well as on the margins of beech forests in altitudes over 1000 m. Geophilous.</p><p>Remarks.</p><p>Differences to H. thessalica and H. borealis are described under the respective species. Helix schlaeflii differs from H. straminea in globular shape with an expanded last whorl and a larger aperture; the Greek populations often differ in a pale colouration, because H. straminea has typically more vivid colours. Helix straminea also lacks the whitish pattern on the bands characteristic for H. schlaeflii and often has a darker, brown foot.</p><p>Helix schlaeflii is not monophyletic in the mitochondrial tree (Fig. 2). This is due to three issues. First, there are two mitochondrial clades in this species that are deeply divergent and whose mutual relationships remain unresolved. Second, the population from Krujë (central Albania, isolate SH 5 in the tree) possesses a mitochondrial lineage introgressed from Helix secernenda Rossmässler, 1847 . Third, a sample of H. straminea yielded a mitochondrial lineage typical for H. schlaeflii, apparently also due to introgression. Mitochondrial introgressions are not rare in the western-Balkan radiation of Helix (Korábek et al. 2022) and an ancient introgression may in fact account also for the presence of two divergent lineages within H. schlaeflii . Introgressions are generally a major problem for species identification using mitochondrial data (Funk and Omland 2003).</p></div>	https://treatment.plazi.org/id/263DCD36E7BA5EBBB576733FE6049051	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
5FD85ABDF58E502281D637369BCE86E1.text	5FD85ABDF58E502281D637369BCE86E1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix (Helix) straminea Briganti 1825	<div><p>Helix (Helix) straminea Briganti, 1825</p><p>Figs 19, 20</p><p>References.</p><p>Korábek et al. 2014, 2022; Petraccioli et al. 2021.</p><p>Description.</p><p>Shell (Fig. 19) large, conical, with varying relative height; umbilicus fully covered or more rarely slit-like; individual whorls low and tightly coiled; suture deep; last whorl not expanded; aperture small and low; no umbilicus; protoconch rather large; shell surface irregular with occasional lightly coloured ribs; basic colour pale grey, usually with four well-developed dark brown bands of which the middle pair is much thicker, but bands sometimes only faint; aperture margins brown, about as dark as the bands. Animal (Fig. 20) brown including mantle margins, rarely pale brown.</p><p>Distribution and habitat.</p><p>Helix straminea occurs in the northwest of North Macedonia and in central Albania (Fig. 21); besides that, it is widely distributed in the Apennines (Petraccioli et al. 2021; Korábek et al. 2022). This species is reported here from Greece for the first time; it was found in a floodplain of a small stream north of Kalambaka (Thessaly, central Greece). Two live individuals and several empty shells were collected in a gallery forest dominated by Platanus, but with a rich herb understorey. While the mitochondrial haplotype of these specimens belongs to one of the mitochondrial lineages of H. schlaeflii, they can be identified as H. straminea based on the more conical shell with a narrower last whorl and a smaller aperture (cf. Korábek et al. 2014: fig. 4 B). A closely related H. schlaeflii haplotype was already recorded in H. straminea before in central Albania, where two other sequenced individuals from the same population had H. straminea haplotypes as expected (Korábek et al. 2022). However, that population originated from central Albania and all the H. schlaeflii haplotypes related to the haplotype from Greece obtained here also originate from central Albania. Although H. straminea (as Helix vladika (Kobelt, 1898)) was reported from an unspecified locality in the valley of Vjosa at the border between Albania and Greece (Dhora and Welter-Schultes 1996), the current record is far from the range of the species and is perhaps explainable by an introduction. However, there may exist a second population in northern Greece on the east bank of the Ioannina Lake, as documented by one convincing and two possible records posted on iNaturalist (https://www.inaturalist.org/observations/180903383, https://www.inaturalist.org/observations/189208381, https://www.inaturalist.org/observations/121853021). Furthermore, the distribution of H. straminea in the Balkan part of its range is very scattered, so it is conceivable that it extends naturally also to northern Greece.</p><p>Helix straminea was in the Balkans found in different habitats, from beech forests to shrubs. There is no clear species boundary to the closely related H. vladika in the north (northern Albania to central Serbia).</p></div>	https://treatment.plazi.org/id/5FD85ABDF58E502281D637369BCE86E1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
18FD6CC02B7257329C6187FA98C808A1.text	18FD6CC02B7257329C6187FA98C808A1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix (Helix) thessalica Boettger 1886	<div><p>Helix (Helix) thessalica Boettger, 1886</p><p>Figs 22, 23</p><p>References.</p><p>Korábek et al. 2016 a, 2016 b, 2020, 2023 b; Korábek and Hausdorf 2024.</p><p>Description.</p><p>Shell (Fig. 22) large, globular; body whorl large; very spacious aperture; umbilicus narrow, completely covered or slit-like; shell surface with irregular fine ribs; strongly developed fine spiral grooves (well visible above the aperture); shell covered with a thick periostracum, yellowish brown; bands largely missing in Greek populations; aperture margins white in Greek populations; mantle pale. Animal (Fig. 23) pale brown or yellowish.</p><p>Distribution and habitat.</p><p>In Greece it lives in higher altitudes. It is relatively broadly distributed in the Rhodopes (Fig. 24), there are isolated occurrences in Thessaly (Ossa / Kissavos, Pelion), and we report it here from the Paiko Mts. (SW of Gevgelija). Its typical habitat in Greece is beech forests, where it can be found in places enriched in nutrients (e. g. along streams, in nettles). Geophilous, but juveniles climb on herbs and can be often found on their leaves.</p><p>Remarks.</p><p>The most similar species in Greece is H. schlaeflii . In Helix thessalica, the shell is in most cases much more darkly coloured with a distinctly developed periostracum. Greek populations lack any brown colour on the columella, which in turn is characteristic for H. schlaeflii . Helix thessalica has a dark grey penis, epiphallus and vagina. The description as provided fits the Greek populations, but in other parts of the range the species may be distinctively banded, with brown apertural margins and darker foot.</p></div>	https://treatment.plazi.org/id/18FD6CC02B7257329C6187FA98C808A1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
B9F5F964764C536E944AC994E8C54847.text	B9F5F964764C536E944AC994E8C54847.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix (Pelasga) figulina Rossmassler 1839	<div><p>Helix (Pelasga) figulina Rossmässler, 1839</p><p>Figs 34, 35</p><p>References.</p><p>Neubert 2014; Korábek et al. 2022.</p><p>Description.</p><p>Shell (Fig. 34) very small (the smallest Helix species in Greece), globular, with large body whorl and spacious aperture; no umbilicus; protoconch very small; shell surface with regular, rounded ribs and lacking spiral sculpture; shell pale greyish or brownish, with the lower two bands narrow and the upper three faint and most often partly fused; aperture margins straight and white; white columella rounded and smoothly transitioning into the palatal area. Animal (Fig. 35) pale brownish with darker, brown or reddish-brown, back, mantle margins pale grey; calcareous diaphragm conspicuously convex and attached to the very margins of the aperture.</p><p>Distribution and habitat.</p><p>Very common species distributed over large part of mainland Greece and the Peloponnese (Fig. 36), but completely missing from the west (&lt;21.5– 22.0°E). It is broadly distributed from the southeast of North Macedonia (valleys of Strumica and Vardar, isolated occurrences reported up to Kumanovo) and southeastern Bulgaria (Thrace) to the Aegean islands (Cyclades, Northern Sporades, Lesvos, Samothraki, etc.). It also lives in a small area of western Anatolia (e. g. the ancient Pergamon and Troy). Fossils were found on Crete (Kotsakiozi et al. 2012). It lives in open, often exposed habitats with low vegetation. May be difficult to find alive when inactive because it buries itself into the soil. Geophilous.</p><p>Remarks.</p><p>Helix figulina is easily recognisable due to small size, a very small protoconch, globular shell shape with large aperture, rounded columella smoothly transitioning to the bottom of the previous whorl, completely white aperture margins, and regularly ribbed surface. It may be found syntopically or nearly so with H. lucorum, H. borealis, H. philibinensis and H. pelagonesica . Helix philibinensis is the most similar species overlapping in size, but it has a blunter apex, smoother shell surface, smaller aperture which usually has at least partially coloured margins.</p></div>	https://treatment.plazi.org/id/B9F5F964764C536E944AC994E8C54847	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
99A7F357DC735A90950FB4CFFD2CA00C.text	99A7F357DC735A90950FB4CFFD2CA00C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix (Pelasga) Hesse 1908	<div><p>Subgenus Pelasga Hesse, 1908</p><p>References.</p><p>Hesse 1908; Neubert 2014; Korábek and Hausdorf 2023.</p><p>Type species.</p><p>Helix (Helicogena) pelasgica Kobelt 1904 = Helix figulina Rossmässler, 1839, by subsequent designation (Hesse 1918: 38).</p></div>	https://treatment.plazi.org/id/99A7F357DC735A90950FB4CFFD2CA00C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
3663380B95865A6094409C18574CBF31.text	3663380B95865A6094409C18574CBF31.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Helix Linnaeus 1758	<div><p>Genus Helix Linnaeus, 1758</p><p>References.</p><p>Hesse 1908; Giusti et al. 1995; Neubert and Bank 2006; Korábek and Hausdorf 2023).</p><p>Type species.</p><p>Helix pomatia Linnaeus, 1758, by subsequent designation (de Montfort 1810: 231).</p></div>	https://treatment.plazi.org/id/3663380B95865A6094409C18574CBF31	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Korábek, Ondřej;Dolejš, Petr;Coufal, Radovan;Juřičková, Lucie;Kubíková, Kateřina;Hausdorf, Bernhard	Korábek, Ondřej, Dolejš, Petr, Coufal, Radovan, Juřičková, Lucie, Kubíková, Kateřina, Hausdorf, Bernhard (2025): A new distinctive lineage of Helix (Gastropoda, Stylommatophora, Helicidae), with a guide to Helix species from mainland Greece. ZooKeys 1249: 93-135, DOI: 10.3897/zookeys.1249.143635
