Biodiversity Journal, 2021,12 (2): 467–474 https://doi.org/10.31396/Biodiv.Jour.2021.12.2.467.474

Rediscovery of Assyriella rechingeri (Fuchs et Käufel, 1936) ( ) in Karpathos Island (Dodecanese, Greece)

Mauro Grano1 & Cristina Cattaneo2

¹Via Valcenischia 24, 00141 Rome, Italy; e-mail: [email protected] ²Via Eleonora d’Arborea 12, 00162 Rome, Italy; e-mail: [email protected]

ABSTRACT The aim of this work is to confirm the occurrence of several living populations of Assyriella rechingeri (Fuchs et Käufel, 1936) (Gastropoda Helicidae) in Karpathos Island (Dodecanese, Greece) since the species was believed to be extinct in the past by some authors due to the dis- covery of only subfossils specimens. Its systematic status and the sympatry with Levantina spiriplana (Glaubrecht, 1993) and L. malziana (L. Pfeiffer, 1861) are discussed.

KEY WORDS Assyriella rechingeri; endemism; Kali Limni.

Received 26.02.2021; accepted 30.04.2021; published online 30.06.2021

INTRODUCTION tomella (instead of Gyrostoma). As knowledge of species, localities, distribution areas and system- Assyriella rechingeri (Fuchs et Käufel, 1936) atic position of the supraspecific taxa mentioned (Helicidae family, Helicinae subfamily, Helicini above increased, Assyriella was treated as a sepa- tribe) is part of a group of conchologically similar rate genus (Glaubrecht 1993a, 1993b; Subai, genera found between the central Mediterranean to 1994). Molecular data are in Korabek et al. (2014). Middle East and the Caucasus, which are close rel- Currently, Assyriella is considered para- ative of the genus Helix Linnaeus, 1758. These gen- phyletic with respect to Levantina (Korábek et al., era are: Maltzanella P. Hesse, 1917; Lindholmiola 2014). The supraspecific taxa Codringtonia, P. Hesse, 1977; Levantina Kobelt, 1871; Assyriella Isaurica, Assyriella and Levantina evidently de- Hesse, 1908; Gyrostomella Hesse, 1911, Isaurica veloped from a common root, just like the North Kobelt, 1901; Codringtonia Kobelt, 1898 (Hesse, African Gyrostomella. Schütt stated that South- 1920; Boettger & Wenz, 1921; Subai, 1994). eastern Anatolia was the radiation centre of the Hesse (1908) had proposed the taxon As- genus Assyriella (Schütt & Subai, 1996). From syriella as a section of the Levantina for the here it may have spread in the Middle East and species “guttata Olivier, 1804”. Levantina had westward in some islands as Cyprus and been split into the subgenera Codringtonia, Karpathos (Akbayin et al., 2001). The spreading Isaurica and Levantina s. str., and the latter was of genus Assyriella towards west and south took split into Assyriella, Levantina and Gyrostoma place following the isolation and separation of sections for shell anatomy reasons. Subsequently, Codringtonia, Isaurica and Levantina genera Hesse (1918) elevated Assyriella to the rank of (Schütt & Subai, 1996). Eustatic fluctuations in subgenus close to Levantina s. str. and Gyros- sea level, especially during the Messinian period 468 MAURO GRANO & CRISTINA CATTANEO

(6 Ma), allowed the ancestors of Assyriella to geri must have reached the island by this time. cross the Mid Aegean Trench (MAT) in the Karpathos was reintegrated with the mainland dur- Aegean and to colonize the Peloponnese, whose ing the Messinian salinity crisis (5–6 Ma) and was further development isolated the Codringtonia joined with Rhodes and Anatolia in Lower genus. Subsequently, sea level rise and tectonic Pliocene (Daams & Van der Weerd, 1980), but pre- uplift isolated the Isaurica genus. The very dis- sumably remained isolated at the end of the middle joint distribution of the Levantina genus on some Pliocene and during the whole Pleistocene (Der- Aegean islands and on the Syrian-Israeli mainland mitzakis, 1991). This isolation would have led the dates back to a subsequent lowering of the sea Assyriella population of Karpathos and Cyprus to level during a Pleistocene glaciation (Pfannen- an evolutionary change and to an adaptation to dif- stiel, 1944). The lowering of the coastline up to ferent environmental conditions. Hence, A. rechin- 200 m allowed migrations of freshwater and ter- geri on Karpathos and bellardii on Cyprus can be restrial molluscs along the Mediterranean coast construed as vicarious taxa of the Assyriella favouring the spreading of the genus Assyriella. species group. A similar disjoint distribution in the Furthermore, volcanic eruptions in the southern eastern Mediterranean can be observed in Lev- part of the Caucasus favoured the isolation of antina spiriplana (Glaubrecht, 1993). The Eastern population with the emergence of geographically Mediterranean land Levantina displays a dis- isolated species toward east (Schütt & Subai, junct range spanning the Middle East (Levant), 1996). Cyprus, few locations along the Aegean Turkish The genus Assyriella Hesse, 1909 includes 13 coast between Bodrum and Datça, the islands of species: A. bellardii (Mousson, 1854), A. cer- Rhodes and Karpathos and a few surrounding islets atomma (Pfeiffer, 1856), A. cilicica (Kobelt, 1895), (Dodecanese). In the Levant, it is represented by A. djulfensis (Dubois de Montpéreux, 1840), A. es- two species parapatrically distributed: L. caesare- cheriana (Bourguignat, 1864), A. guttata, A. kur- ana (Mousson, 1854) in the north and L. hi- distana (Pfeiffer, 1862), A. mardinensis (Kobelt, erosolyma (Mousson, 1854) in the south. In the 1900), A. naegelei (Kobelt, 1901), A. ninivita (Gal- Dodecanese L. spiriplana and L. malziana (L. land, 1885), A. rechingeri (Fuchs & Käufel, 1936), Pfeiffer, 1861) occur together (Ketmaier & A. thospitis Schütt et Subai, 1996 and A. vanensis Glaubrecht, 2015). In Karpathos, both species are Schütt et Subai, 1996. Assyriella guttata (from the parapatrically distributed, and leave in sympatry Gulf of Iskenderun to Mersin), A. bellardii (Cyprus) with A. rechingeri sharing area up to 500 m a.s.l. and A. rechingeri are the westernmost representa- tives of the genus. Assyriella rechingeri is an exclusive endemic species of Karpathos Island in the Dodecanese MATERIALS AND METHODS Archipelago (Greece) and perhaps is a relic popu- lation of the Assyriella group. The relic occurrence Study area of A. rechingeri on Karpathos and A. bellardii on Cyprus suggests that Assyriella was once more Karpathos is the second largest Dodecanese is- widely distributed in the eastern Mediterranean as land after Rhodes and lies in the SE Aegean Sea, it is even today (Schütt & Subai, 1996) and can halfway between Crete and Rhodes (39 km from also be interpreted as allospecies of a widely dis- Crete and 25 km from Rhodes). The island is 49 tributed A. guttata superspecies (Glaubrecht, km long and 15 km wide and covers an area of 1994). It could be argued that the Assyriella popu- 302.15 square kilometres. Together with Kasos lations were separated from the Assyriella distri- and Armathia in the southwest, the nearby Saria in butional area (Eastern Anatolia) after Miocene due the north and several offshore islets, Karpathos to climatic and geological changes. Palaeogeo- forms the Karpathos Archipelago and is a con- graphical data reveal that Karpathos was detached stituent part of the South Aegean Island Arc from the island chain of Rhodes and thus from the (Greuter et al., 1983). The three larger islands con- mainland of Asia Minor as early as in the late tain widespread mountainous areas with some Miocene (5–6 Ma). Thus, the ancestors of rechin- higher peaks, culminating on Karpathos in the Kali Rediscovery of Assyriella rechingeri (Gastropoda Helicidae) in Karpathos Island (Dodecanese, Greece) 469

Limni (1215 m), on Kasos in the Megalo Prionas RESULTS (601 m), and on Saria in the Pachy Vouno (629 m) (Grano & Cattaneo, 2019). On Karpathos the only Assyriella rechingeri (Fuchs et Käufel, 1936) flat areas are near Afiartis in the south, near Lastos in the centre, and Avlona in the north. Karpathos is 1936. Levantina (Codringtonia) rechingeri Fuchs built up of limestone and dolomitic limestone & Käufel, Arch. Naturgesch., (NF) 5 (4): 658, (Jurassic-Eocene) in the northernmost part, flysch Abb. XI, F. 35 A–C, locus typicus: Insel (Eocene) in most of the northern and the southern Karpathos: Gipfelstock des Kalolimni, 800– part, Neogene deposit in the southernmost area. 1000 m. Holotypus: NMW 535 (lost). The central part of the island corresponding to the 1939. Codringtonia (Isaurica) rechingeri, - K. L. Kali Limni Mountain complex, the area near Pfeiffer & Wächtler, Arch. Molluskenkunde, 71 Menetes (south) and Pigadia (south-east), are con- (2/3): 57, Abb. 10c (as Isaurica rechingeri). stituted by the Kalilimni Unit, a parautochthonous 1987. Codringtonia intusplicata rechingeri, - Zilch, carbonate platform (Jurassic-Eocene) overlaid by Arch. Molluskenkunde, 117 (1986) (4/6): 245. the Xindothio Unit (Triassic-Cretaceous) almost 1994. Assyriella rechingeri, - Glaubrecht, Verh. built up of ophiolites (Cordey & Quillévéré, 2019). naturwiss. Ver. Hamburg, (NF) 34: 373, F. 1 (Karte), F. 2 (shell). Methods Shell dirty white with yellowish hue, with five Our naturalistic research has been carried out on brown colour bands, often interrupted by white Karpathos twice, the first from 28 July to 17 August zigzag lines, but blurrier than in L. spiriplana. 2019 and the second from 10 August to 17 August Smooth, white aperture with a short and distinct 2020. During the second session (2020) A. rechin- callus at the columellar base, peristome well re- geri was found in four different areas. For an opti- flected, sharp, with margins connected by a thin mal research, the entire island has been only visited parietal layer, umbilicus narrow and deep, partly on foot along roads, paths, and streams. Only empty covered by the reflected columellar margin. The shells were found in several sites - which are men- size is 15.5–21 x 31–38 mm. Named in honour of tioned below - therefore only a conchological study the Austrian botanist and phytogeografer Karl of the specimens was possible to perform. The Heinz Rechingher (Wien 16 October 1906 - Wien shells were cleaned with water and a brush after 30 December 1998). being soaked for about two minutes in a solution Based on a single heavily corroded specimen with 60% water (H2O) and 40% NaClO. The spec- collected in 1935 by Rechinger on the Kali Limni imens are kept in the private collection of one of the massif, the species was initially described as Lev- authors (MG, Rome, Italy). For the of antina (Codringtonia) rechingeri Fuchs et Käufel, this species the Fauna Europaea Project (Bank, 1936, type locality “Karpathos, Kalolimni moun- 2017) and MolluscaBase (2021) has been followed. tain peak region, 800–1000 m” (Fuchs & Käufel, The collected material is of 12 specimens of 1936). The holotype is deposited in the Naturhis- which 5 examined (Table 1). The shell variables torisches Museum of Wien MNW 535 (lost, Schütt shown in Fig. 1 were measured on the specimens in & Subai 1996). Subsequently, Pfeiffer found other millimeters with a calliper of 1/100 mm precision. specimens (Pfeiffer & Wächtler, 1939) and based A–B: Shell Height (ShH); A–C: Spire Height on the shell morphology and on biogeographical (SpH); D–G: Shell Diameter (ShD); E–F: Trunca- reasons, suggested to assign the taxon to Isaurica tion Width (TrW); E–G: Aperture Width (ApW); Kobelt, 1901 rather than to the Greek Codringtonia H–I: Aperture Height (ApH). Kobelt, 1898. Later, Glaubrecht (1994) assigned The specimens of the Figs. 2–4 are from Profitis rechingeri from Karpathos to the Assyriella genus, Ilias (35°37’20’’N - 27°08’09’’E 500 m), the spec- as already supposed by Subai (1994), highlighting imen of the Fig. 5 is from Profitis Ilias the differences with Isaurica. The latter differs (35°37’20’’N - 27°08’09’’E 500 m) (Fig. 2), and the from rechingeri by several features such as the specimen of the Fig. 6 is from Kali Limni larger and spherical shell, the diameter-to-callus (35°35’42’’N - 27°07’25’’E 1100 m). length ratio, and the umbilical region always com- 470 MAURO GRANO & CRISTINA CATTANEO

pletely covered. Instead, rechingeri shows a rein- enough to justify different species assignments forced, everted aperture lip and the umbilicus al- (Glaubrecht, 1994; Ketmaier & Glaubrecht, 2015). most entirely open. Furthermore, all rechingeri Assyriella rechingeri is considered CR, Criti- shells show a tooth-like protuberance on the lower cally Endangered, in the EU Red List. The - lip that is less pronounced in the Isaurica forms Base database www.animalbase.uni-goettingen.de (Glaubrecht, 1994). This tooth like projection and report: “Rare, usually only subfossil shells are broadly everted aperture lip, as in rechingeri, are found. It is not known if live are still present features only for the genus Assyriella, especially somewhere, and if yes, by which factors they are for its type species A. guttata and for A. bellardii. threatened. Glaubrecht 1994 suspected that the In addition, both the latter two species have a con- species was extinct”. spicuous, at most half-covered, umbilicus. It is worth noting that Assyriella like Levantina and the northern African Gyrostomella includes snails dis- DISCUSSION playing either a wide-open umbilicus or an almost covered umbilical region. In Assyriella as in Lev- The specimens from Kali Limni are signifi- antina the discrepancy between umbilicate and cantly smaller than those of Profitis Ilias (Figs. 2– non-umbilicate forms is still regarded relevant 6). The specimens have been found at different

Figure 1. Measurements of shell variables (modified by Akbayin et al., 2001).

Table 1. Measured variables of five specimens of Assyriella rechingeri (see text). Rediscovery of Assyriella rechingeri (Gastropoda Helicidae) in Karpathos Island (Dodecanese, Greece) 471

Figures 2–6. Assyriella rechingeri from Karpatos island. Figs. 2–4: Profitis Ilias. Fig. 5: Profitis Ilias. Fig. 6: Kali Limni. altitudes (from 500 to 1100 m a.s.l.) in several sites area is occupied only by A. rechingeri. Previous all included in the Kali Limni massif: Mt. Kali competition with Levantina genus may have caused Limni, Mt. Kollas and Mt. Profitis Ilias, Also the the limited distribution of A. rechingeri in the Kali specimens collected by Pfeiffer (1937) and by Limni massif (Glaubrecht, 1994). Competition be- Glaubrecht (1989), come all from Mt. Kali Limni tween these related species, with a probably similar and from the mountainous reliefs in the nearby as ecological niche, would have involved a specializa- Mt. Melloura (south-east of Lastos), Mt. Kollas and tion of A. rechingeri to occupy a niche characterized Mt. Volada (Glaubrecht, 1994). We confirm as by specific climatic (higher altitude) and edaphic stated by Glaubrecht (1994), that it seems to be no conditions. It is noteworthy that on Cyprus we have evidence of the presence of the species in other the same situation, as both the aforementioned parts of the island. Assyriella rechingeri lives in species of Levantina occur in sympatry with A. bel- sympatry in Karpathos with L. spiriplana (Olivier, lardii Mousson, 1854 (Glaubrecht, 1994). 1801) and L. malziana, which show a parapatric distribution. Both the latter species are abundantly present throughout the island with the exception for CONCLUSIONS the Kali Limni from 500 to maximum height. Our discovery of some specimens of A. rechingeri to- The summer period during which our research gether with L. spiriplana on Mt. Kollas at an alti- took place did not allow us to find live specimens tude of 500 m would suggest an overlap between of A. rechingeri. However, numerous specimens the distribution range of both species. However, were found in excellent condition and therefore re- from 500 m up to the summit of Mt. Kali Limni, the cently died. In addition, some specimens were also 472 MAURO GRANO & CRISTINA CATTANEO

Figure 7. Entrance of the burrow of Rattus rattus predator of molluscs (Karpatos island).

Figure 8. Limestone quarries and roads on the eastern slope of the Mt. Kollas (Karpatos island). Rediscovery of Assyriella rechingeri (Gastropoda Helicidae) in Karpathos Island (Dodecanese, Greece) 473

encountered at the entrance of the numerous bur- Dermitzakis M.D., 1991. Paleogeography, geodynamic rows of Rattus rattus (Linnaeus, 1758) predators process and event stratigraphy during the Late Ceno- of molluscs in arid environments (Fig. 7). zoic of the Aegean area. In: Azzaroli A. (Ed.), Bio- Glaubrecht (1994) hypothesized that A. rechingeri geographical aspects of insularity. Atti dei Convegni may be extinct, fortunately, we can claim that this Lincei, 85: 263–288. Rome (Accademia Nazionale dei Lincei). species is not extinct even if its presence is rare Glaubrecht M., 1993a. Die Landschnecke Levantine with a very localized distribution. It could be a spiriplana im Ostmediterran: Johanniter-Kreuzritter- relic species and deserves to be overlooked like These oder Paleogeographie? Natur und Museum, many other species of land snails, animals and rare Frankfurt a.M., 123: 97–114. and endemic plants that occur in Karpathos and in Glaubrecht M., 1993b. Entdeckungsgeschichte von particular on the Kali Limni massif. In the last year Landschnecken der Gattung Levantina (, (pers. obs.) limestone quarries and roads have Helicinae). Mitteilungen der Deutschen Malakozool- been built on the eastern flank of the Mt. Kollas ogischen Gesellschaft, 50/51: 33–40. (Fig. 8). The tourism sector is increasingly devel- Glaubrecht M., 1994. Systematic and biogeography of oping in Karpathos and this involves the extraction Assyriella rechingeri (Gastropoda, Pulmonata, Heli- cidae) on Karpathos, Greece. Verh. Naturwiss. Ver. of large quantities of limestone for the construc- Hamburg, 34: 373–384. tion of new buildings and hotels. This represents Grano M. & Cattaneo C., 2019. First record of Rough- a serious damage for the environment since pecu- tailed Agama Stellagama stellio (Linnaeus, 1758) liar habitats for the fauna and for endemic plants (Reptilia, Agamidae) from Karpathos island (Dode- have already been eradicated. canese, Greece). Parnassiana Archives, 7: 51–54. Greuter W., Pleger R. & Raus T., 1983. The vascular flora of the Karpathos island group (Dodecanesos, ACKNOWLEDGEMENTS Greece). A preliminary checklist. Willdenowia, 13: 43–78. We are grateful to Fabio Liberto (Cefalù, Italy) and Hesse P., 1908. Die systematische Stellung von Helix leachii Fér. und gyrostoma Fér. Zoologische Joop Eikenboom (Melissant, The Netherlands) for Jahrbücher, 27: 313–320. their valuable help and advice. Also, a special Hesse P., 1918. Das Genus Levantina Kob. Nachrichten- thanks to Roberto Nistri for the “still life” photos. blatt der Bayerischen Bayerischen Malakozoologis- chen Gesellschaft, 50: 40–47. Hesse P., 1920. Iconographie der Land & Süsswasser- REFERENCES Mollusken mit vorzüglicher Berücksichtigung der eu- ropäischen noch nicht abgebildeten Arten von E.A. Akbayin H., Hevedanli M. & Şeşen R., 2001. Some As- Rossmässler. N.F. 23, 1–262, pl. 631–660. Wies- syriella species in South East Anatolia in Turkey dif- baden: Kreidel. ferentiated by multivariate statistical analysis Ketmaier V. & Glaubrecht M., 2015. The legacy of the (Gastropoda: Pulmonata, Helicidae). Turkey Journal Crusaders: Complex history of colonization and an- of Zoology, 25: 1–9. thropochory in the land snails Levantina (Gas- Bank R.A., 2017. Fauna Europaea Project. Checklist of tropoda, Pulmonata) in the Eastern Mediterranean. the land and freshwater Gastropoda of Europe, 176 pp. Zoosystematics and Evolution, 91: 81–89. Boettger C.R. & Wenz W., 1921. Zur Systematik der zu https://doi.org/10.3897/zse.91.4693 den Heliciden subfamilien Campylaeinae und Helic- Korábek O., Jurickova L. & Petrusek A., 2014. Resur- inae gehörigen tertiären Landschnecken. Archiv für recting Helix straminea, a forgotten escargot with Molluskenkunde, 53: 6–55. trans-Adriatic distribution: first insights into the ge- Cordey F. & Quillévéré F., 2019. Reassessing the age of netic variation within the genus Helix (Gastropoda: Karpathos ophiolite (Dodecanese, Greece): conse- Pulmonata). Zoological Journal of the Linnaean So- quences for Aegean correlations and Neotethys evo- ciety, 171: 72–91. lution. Geological Magazine, 157: 1–12. https://doi.org/10.1111/zoj.12122 https://doi.org/10.1017/S0016756819000657 MolluscaBase, 2021. MolluscaBase. Accessed at Daams R. & Weerd A. Van de, 1980. Early Pliocene http://www.molluscabase.org on 2021–01–05. small mammals from the Aegean Island of Karpathos https://doi.org/10.14284/448 (Greece) and their palaeogeographic significance. Pfannenstiel M., 1944. Die diluvialen entvicklungssta- Geologie en Mijnbouw, 59: 327–331. dien und die Urgeschiccthe von Dardanellen. Mar- 474 MAURO GRANO & CRISTINA CATTANEO

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