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Ecologica Montenegrina 10: 26-30 (2017) This journal is available online at: www.biotaxa.org/em

https://zoobank.org/urn:lsid:zoobank.org:pub:476BC71C-E055-47A4-8280-205F37A6A981

Cotyorica nemethi n. gen. n. sp., a remarkable Tertiary relict of the subfamily Phaedusinae (: : ) from northern Turkey

JOZEF GREGO1 & MIKLÓS SZEKERES1

1 Horná Mičiná 219, SK-97401 Banská Bystrica, Slovakia, e-mail: [email protected] 2 Institute of Plant Biology, Biological Research Centre of the Hungarian Academy of Sciences, Temesvári krt. 62, H- 6726 Szeged, Hungary, e-mail: [email protected]

Received: 10 February 2017 │ Accepted by V. Pešić: 15 March 2017 │ Published online: 17 March 2017.

Abstract Cotyorica nemethi n. sp., one of the Tertiary relict western members of the geographically disjunct subfamily Phaedusinae, is described from Ordu Province of northern Turkey. It shows similarity to the species of the subterranean genera Graecophaedusa Rähle, 1982 and Nothoserrulina Németh & Szekeres, 1995 having high positioned lamellae inferior and subcolumellaris, but differs from these to an extent that justifies distinct generic status.

Key words: Phaedusinae, endemism, Tertiary relict, subterranean habitat.

Introduction

Climatically well balanced coastal regions of the Mediterranean, Black and Caspian Seas with mild winters harbour a remarkably rich and diverse fauna of the family Clausiliidae. Among its unique endemic elements are a few species that represent Tertiary relicts of the mainly Asiatic Phaedusinae subfamily (Nordsieck 1978). In addition to the favourable climate, the survival of most of these clausiliids was ensured by their subterranean habitats capable of providing frost-free and permanent high humidity environment. Due to the hidden life style of several members in this 'European' group (Nordsieck 1978) of the Phadusinae (also including taxa in the Caucasus and Hyrcanian areas), only recent studies (e.g., Rähle 1982; Neubert 1992; Majoros et al. 1994; Nordsieck 1994; Németh & Szekeres 1995; Gittenberger 2000; Reischütz et al. 2016) gave an idea of their previously underestimated diversity. Helped by novel collecting methods, in less then four decades these research efforts increased the number of described species substantially, from 10 (Nordsieck 1978) to 28. A recently discovered species of the subterranean Phaedusinae, representing a new endemic genus of the Black Sea coast, is a further addition to the diversity of these remarkable clausiliids.

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Material and Methods

The sample containing the holotype was taken from the upper 2 cm layer of soil at the bottom of an east- exposed cliff. Measurements and photographs were taken at the Naturhistorisches Museum (Vienna) with a Nikon DS-F2.5 camera mounted on a Nikon SMZ25 stereomicroscope, using NIS-Elements Microscope Imaging Software with an Extended Depth of Focus (EDF) patch. Line drawing of the aperture was done using a photographic template.

Systematics

Cotyorica n. gen.

Type species: Cotyorica nemethi n. sp.

Diagnosis: The diagnostic features of the new genus are the same as those of the type species.

Etymology: The new genus is named after the ancient name (Cotyora) of Ordu, a major town near the type locality.

Cotyorica nemethi n. sp. (Figs 1, 2)

Type material: Turkey, Ordu Province, coastal slope between Perşembe and Mersin (41°07' N 37°47' E), 20 m, leg. L. Németh 30.06.1991. Holotype, HNHM 100140, to be deposited at the Hungarian Natural History Museum (Budapest).

Figure 1. Cotyorica nemethi n. sp. Holotype, 11.5 mm.

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COTYORICA NEMETHI N. GEN. N. SP. FROM NORTHERN TURKEY

Diagnosis: Non-decollated small species of the Phaedusinae with strongly costate neck, all around serrate peristome, high positioned, deep divided lamella inferior ending in two parallel branches, uninterrupted lamella subcolumellaris, as well as short upper and longer lower plicae on the lateral side.

1 Description: The colourless shell with slightly bent spire consists of 9 /3 flat whorls. The dense ribs of the surface become stronger, sharper and widely spaced at the neck. The ovoid peristome is projected, at the basis not backward-bent. Its wide, reflexed margin has strong plicae all around, except at the sinulus. The lamella superior is moderately emerged, in front view not more conspicuous than the interlamellar plicae. Inward it is smoothly continuous with the lamella spiralis. Deeper than visible through the aperture the lamella inferior divides in two parallel branches, which descend along the lamella spiralis and terminate marginally as strong plicae above half height of the aperture. The uninterrupted lamella subcolumellaris also ends marginally, separated from the lower branch of the inferior by a strong plica. The plica principalis starts ventrolaterally and, gradually converging toward the suture, reaches close to the aperture. On the lateral side it is accompanied by a short upper and a farther forward reaching, twice longer lower palatal plica. The deep plate is not visible through the aperture. Dimensions of the holotype: shell height 11.5 mm, shell width 2.9 mm, aperture height 2.2 mm, aperture width 1.7 mm.

Figure 2. Lamella positions in the aperture of Cotyorica nemethi n. sp. The lamellae superior (Lsu), inferior (Lin) and subcolumellaris (Lsc) are marked.

Etymology: The new species is named after László Németh, the collector of the type specimen and an esteemed friend of the authors, who substantially contributed to the research of the Turkish malacofauna.

Habitat: Cotyorica nemethi n. sp. was recovered as an empty shell from a litter-soil sample collected at a cliff. As its closest relatives Graecophaedusa sperrlei Rähle, 1982 and Nothoserrulina subterranea Németh & Szekeres, 1995 were found in narrow underground passages between rocks (Németh & Szekeres 1995; Reischütz et al. 2008), it is assumed that this species may have similar habitat preference. This is also suggested by the highly local occurrence of the new species in a zoogeographically well researched region. The holotype may have been washed out from crevices of the cliff by rain water.

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Remarks Among non-decollated western Phaedusinae C. nemethi n. sp. shows superficial similarity to Serrulina serrulata (Pfeiffer, 1847) that differs markedly by its steeply descending and low ending lamella inferior, a feature shared between members of the genera Serrulina Mousson, 1873, Dobatia Nordsieck, 1973, Sciocochlea Boettger, 1935, and Tsoukatosia Gittenberger, 2000. By contrast, the high positioned lamellae inferior and subcolumellaris of the new species (Fig. 2) resemble those seen in the monotypic genera Graecophaedusa Rähle, 1982 and Nothoserrulina Németh & Szekeres, 1995. Nevertheless, C. nemethi n. sp. is well distinguishable from Graecophaedusa sperrlei by the narrow apex and less retracted palatal plicae, from Nothoserrulina subterranea by the spindle-shaped shell and the absence of minor folds between the upper and lower plicae, and from both species by the projected aperture with non-recessed basal rim, and the strongly serrate parietal peristome margin. Considering these differences relative to the morphological coherence of species in the non-monotypic genera (e.g., Serrulina, Sciocochlea, Tsoukatosa), they justify the classification of the new species in a distinct genus. Ample all year round rainfall and the narrow, within 15°C range of monthly average temperatures of the southern and eastern Black Sea coast provide ideal climatic conditions for the survival of Phaedusinae species, which were abundant elements of the European Clausiliidae fauna before the Pleistocene period (Nordsieck 2000; Reischütz et al. 2016). This temperate region is home to four highly endemic species (Fig. 3), all of which live in shallow subterranean habitats (Culver & Pipan 2014) and, in accordance with their relict status, belong to monotypic genera.

Figure 3. Occurrence of highly endemic Phaedusinae species along the northeastern Aegean and the Black Sea coast. Graecophaedusa sperrlei Rähle, 1982 (1); Nothoserrulina subterranea Németh & Szekeres, 1995 (2); Cotyorica nemethi n. sp. (3); Pontophaedusella ofensis Nordsieck, 1994 (4); Truncatophaedusa evae Majoros, Németh & Szili- Kovács, 1994 (5).

Acknowledgements The authors are thankful to Anita Eschner and Ronald Janssen for granting access to the public collections at their care, to László Németh, as well as Alexander and Peter L. Reischütz, for providing valuable material and discussions, as well as to Zoltán Fehér for his helpful comments and for preparing the photographic images.

References

Culver, D.C. & Pipan, T. (2014) Shallow subterranean habitats: Ecology, evolution and conservation. Oxford University Press, 258 pp. Gittenberger, E. (2000) Serrulininae in Greece, there may be more (Gastropoda, Pulmonata, Clausiliidae). Basteria, 64, 81–87.

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Majoros, G., Németh, L. & Szili-Kovács, T. (1994) Truncatophaedusa evae n. gen. n. sp.: A new representative of Serrulininae from the western Caucasus (Pulmonata: Clausiliidae). Archiv für Molluskenkunde, 123, 123–126. Németh, L. & Szekeres, M. (1995) New and little known species of the Serrulina group from northern Turkey (Gastropoda: Pulmonata: Clausiliidae). Archiv für Molluskenkunde, 124, 93–96. Neubert, E. (1992) Descriptions of new taxa of the Clausiliidae from Turkey (: ). Zoology in the Middle East, 7, 65–86. Nordsieck, H. (1978) Zur Anatomie und Systematik der Clausilien, 20. Die rezenten Arten der Serrulininae und der Gattung Caspiophaedusa. Archiv für Molluskenkunde, 109, 91–101. Nordsieck, H. (1994) Türkische Clausiliidae, 2: Neue Taxa der Unterfamilien Serrulininae und Mentissoideinae in Anatolien (Gastropoda: Stylommatophora). Stuttgarter Beiträge zur Naturkunde, Serie A, 513, 1–36. Nordsieck, H. (2000) Annotated check-list of the fossil (pre-Pleistocene) Clausiliidae (Gastropoda: Stylommatophora) from central and western Europe. Mitteilungen der Deutschen Malakozoologischen Gesellschaft, 65, 1–15. Rähle, W. (1982) Graecophaedusa sperrlei n. gen., n. sp.: ein bemerkenswertes Tertiärrelikt aus Griechisch- Mazedonien. Archiv für Molluskenkunde, 112, 45–48. Reischütz, A., Reischütz, P.L. & Fischer, W. (2008) Helleniká pantoía, 15: Kein Ende abzusehen - Zur Kenntnis der Gattung Balcanodiscus Riedel & Urbanski 1964 (Gastropoda: Pulmonata: Zonitidae). Nachrichtenblatt der Ersten Vorarlberger Malakologischen Gesellschaft, 15, 11–13. Reischütz, A., Reischütz, P.L. & Szekeres, M. (2016) The Clausiliidae subfamily Phaedusinae (Gastropoda, Pulmonata) in the Balkans. Nachrichtenblatt der Ersten Vorarlberger Malakologischen Gesellschaft, 23, 93–117.

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