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A systematic revision of Circassina from the western Caucasus region (Gastropoda: Hygromiidae)
Abstract
This paper presents a comprehensive revision of Circassina (Gastropoda: Hygromiidae). Circassina is divided into two subgenera. The nominotypical subgenus is characterized by an inflation forming a brim at the base of the dart apparatus, whereas Circassina (Abchasohela) is characterized by a granular shell sculpture. Circassina (Circassina) includes C. frutis (with the subspecies C. f. frutis, C. frutis circassica and C. frutis veselyi), which ranges from the northwestern slope of the Caucasus Mountains throughout most of Georgia to northernmost Armenia and the Eastern Pontic Mountains in Turkey, and C. lasistana new species, which is only known from the Vileyet Artvin in Turkey. Circassina (Abchasohela) includes C. christophori, C. pachnodes, C. pergranulata new species, C. septentrionalis new species and C. stephaniae. All species of Circassina (Abchasohela) are restricted to Abkhasia and the adjacent Russian regions in westernmost Caucasia. The shell and the genitalia of each taxon are described and illustrated. All synonyms and all locality records are listed. The distribution of the species is illustrated on a UTM-grid map.
... The family Hygromiidae Tryon, 1866 in the sense of Hausdorf & Bouchet (2005) is a highly diverse group of land snails with a distribution range stretching from Macaronesia in the west throughout Europe, the Mediterranean region, the temperate regions of Asia including the Caucasus region, southern Siberia, the Tian Shan and Altai Mountains to the Russian Far East and to East Africa (Schileyko, 1968(Schileyko, , 1970(Schileyko, , 1978a(Schileyko, , b, 2006bVerdcourt, 1969Verdcourt, , 1973Verdcourt, , 1974Verdcourt, , 1975Backhuys, 1975;Groh, 1984;Giusti & Manganelli, 1987;Neubert, 1998;Muñoz et al., 1999;Hausdorf, 2000a, b, 2001, 2003a, b, Seddon, 2008Sysoev & Schileyko, 2009;Welther-Schultes, 2012;Ganin & Striganova, 2012;Neubert & Bariche, 2013, Neiber & Hausdorf, 2015a. The morphology-based family-level taxonomy of the group largely rests upon a few characters relating to accessory organs of the genital system, i. e. the presence and development of the dart apparatus and parts belonging to these accessory copulatory structures (Nordsieck, 1987(Nordsieck, , 1993Schileyko, 2006b). ...
... However, Fruticocampylaea daghestana, the type species of the monotypic genus-level taxon Shileykoia, was nested within Fruticocampylaea. Circassina and Abchasohela as delimited by Hausdorf (2001) are polyphyletic. Fruticocampylaea christophori, which was tentatively classified with Abchasohela by Hausdorf (2001), although it differs from this group by the possession of a dart and an accessory sac, proved to be a Fruticocampylaea, whereas the true Abchasohela is neither closely related to Circassina (sensu stricto) nor to Fruticocampylaea as previously supposed, but belongs to the Monachaini (Chapter 8). ...
... Several studies identified the Greater Caucasus chain, which rises to 5,642 m above sea level, as a major barrier hampering colonisation of Ciscaucasia by mammals from Transcaucasian refugia (Orth et al., 2002;Seddon et al., 2002;Dubey et al., 2006). In contrast to this, the mainly passively dispersing snail Circassina, which is known from several areas north of the main ridge of the Greater Caucasus (Chapter 6; Hausdorf, 2001), and cold-adapted steppe vipers (Zinenko et al., 2015) crossed this barrier several times and lack deep phylogeographic splits corresponding to the main ridge of the Greater Caucasus. ...
... Circassina Hesse, 1921 (Gastropoda: Hygromiidae) is a group of land snails endemic to the western and central Caucasus region and the eastern Pontus. It includes only two species, C. frutis (Pfeiffer, 1859) and C. lasistana Hausdorf, 2001. Abchasohela Hudec & Lezhawa, 1971, which was formerly classified as a subgenus of Circassina (Hausdorf, 2001), proved to be more closely related to other hygromiid groups than to Circassina (Walther et al., 2015). ...
... It includes only two species, C. frutis (Pfeiffer, 1859) and C. lasistana Hausdorf, 2001. Abchasohela Hudec & Lezhawa, 1971, which was formerly classified as a subgenus of Circassina (Hausdorf, 2001), proved to be more closely related to other hygromiid groups than to Circassina (Walther et al., 2015). The Circassina frutis complex is exceptional among helicoid land snails because it is polymorphic with regard to the dart apparatus, i. e. the snails possess either a complete dart apparatus with a dart sac plus an accessory sac and mucus glands (C. ...
... Schileyko (1978) classified these morphotypes as subspecies, whereas Giusti & Mangenelli (1987) questioned whether they are related at all. Hausdorf (2001) followed Schileyko (1978), but questioned whether the morphotypes are actually genetic entities, because their ranges interdigitate in western Georgia. Furthermore, Hausdorf (2001) separated C. lasistana from C. frutis because of the much shorter spermatophoreforming flagellum. ...
... circassica (= C. frutis (Pfeiffer, 1859)) without dart and accessory sacs as H. stephaniae and synonymized Abchasohela with Circassina, accordingly. Hausdorf (2001) reestablished Abchasohela as a subgenus of Circassina for species with a granular shell sculpture and tentatively included Helix christophori Rosen, 1911 in Abchasohela because it has a granular shell sculpture. However, in contrast to all other Abchasohela species, H. christophori possesses a dart and an accessory sac. ...
... The ML and BI analyses weakly support a sister group relationship between Fruticocampylaea and Circassina (sensu stricto) (BS: 46 %, PP: 0.83) that is also supported by the reduction of the dart apparatus from two to one set of dart and accessory sacs and the presence of a conical plug, via which the dart apparatus inserts into the lumen of the vagina (see Systematics). Circassina and Abchasohela as delimited by Hausdorf (2001) are polyphyletic. Fruticocampylaea christophori, which was tentatively classified with Abchasohela by Hausdorf (2001), although it differs from this group by the possession of a dart and an accessory sac, proved to be a Fruticocampylaea, whereas the true Abchasohela (classified as a synonym of Circassina by Schileyko (1978b) and as a subgenus of Circassina by Hausdorf (2001)), which is represented by A. pachnodes in our analyses, is neither closely related to Circassina (sensu stricto) nor to Fruticocampylaea (Fig. 45). ...
... Circassina and Abchasohela as delimited by Hausdorf (2001) are polyphyletic. Fruticocampylaea christophori, which was tentatively classified with Abchasohela by Hausdorf (2001), although it differs from this group by the possession of a dart and an accessory sac, proved to be a Fruticocampylaea, whereas the true Abchasohela (classified as a synonym of Circassina by Schileyko (1978b) and as a subgenus of Circassina by Hausdorf (2001)), which is represented by A. pachnodes in our analyses, is neither closely related to Circassina (sensu stricto) nor to Fruticocampylaea (Fig. 45). ...
... He misidentified the form of Circassina circassica (D Circassina frutis (Pfeiffer, 1859)) without dart and accessory sacs as Hela stephaniae and synonymized Abchasohela with Circassina, accordingly. Hausdorf (2001) re-established Abchasohela as a subgenus of Circassina for species with a granular shell sculpture and tentatively included Helix christophori Rosen, 1911 in Abchasohela because it has a granular shell sculpture. However, in contrast to all other Abchasohela species, H. christophori possesses a dart and an accessory sac. ...
... Remarks: For nomenclatorial remarks on this species see Hausdorf (2001). Distribution (Fig. 42). ...
... The ML and BI analyses weakly support a sister group relationship between Fruticocampylaea and Circassina (sensu stricto) (BS: 46%, PP: 0.83) that is also supported by the reduction of the dart apparatus from two to one set of dart and accessory sacs and the presence of a conical plug, via which the dart apparatus inserts into the lumen of the vagina (see Systematics). Circassina and Abchasohela as delimited by Hausdorf (2001) are polyphyletic. Fruticocampylaea christophori, which was tentatively classified with Abchasohela by Hausdorf (2001), although it differs from this group by the possession of a dart and an accessory sac, proved to be a Fruticocampylaea, whereas the true Abchasohela (classified as a synonym of Circassina by Schileyko (1978b) and as a subgenus of Circassina by Hausdorf (2001)), which is represented by A. pachnodes in our analyses, is neither closely related to Circassina (sensu stricto) nor to Fruticocampylaea (Fig. 45). ...
This paper presents a systematic revision and a molecular phylogenetic analysis of the Caucasian land snail genus Fruticocampylaea. The genus is newly delimited based on the reduction of the cavities adjoining the seminal duct in the penial papilla. Shell and genitalia of all five species (F. narzanensis, F. kobensis, F. tushetica sp. nov., F. christophori, F. daghestana) are described and figures provided. All synonyms and all locality records are listed. Maximum likelihood and Bayesian analyses of mitochondrial and nuclear DNA sequences (fragments of cox1, 16S rDNA, ITS2 and 28S rDNA) confirm the monophyly of Fruticocampylaea. The reduction of the dart apparatus and the conical plug, via which the dart apparatus inserts into the vagina, as well as the molecular phylogenetic analyses, suggests a sister group relationship between Fruticocampylaea and Circassina (without Abchasohela). Furthermore, the molecular phylogenetic analyses indicate that the Fruticocampylaea species originated in a rapid radiation. The uplift of the Greater Caucasus in the Late Miocene or Pliocene or climatic changes at the end of the Pliocene or in the early Pleistocene may have caused the radiation of Fruticocampylaea. Low intraspecific variability can be explained by population bottlenecks during Pleistocene glacial periods followed by postglacial population increase.http://zoobank.org/urn:lsid:zoobank.org:pub:AB15158D-21A3-4945-8D49-F7DE8E406E2B
... Circassina Hesse, 1921 (Gastropoda: Hygromiidae) is a group of land snails endemic to the western and central Caucasus region and the eastern Pontus. It includes only two species, C. frutis (Pfeiffer, 1859) and C. lasistana Hausdorf, 2001. Abchasohela Hudec & Lezhawa, 1971, which was formerly classified as a subgenus of Circassina (Hausdorf, 2001), proved to be more closely related to other hygromiid groups than to Circassina (Walther et al., in press). ...
... It includes only two species, C. frutis (Pfeiffer, 1859) and C. lasistana Hausdorf, 2001. Abchasohela Hudec & Lezhawa, 1971, which was formerly classified as a subgenus of Circassina (Hausdorf, 2001), proved to be more closely related to other hygromiid groups than to Circassina (Walther et al., in press). The Circassina frutis complex is exceptional among helicoid land snails because it is polymorphic with regard to the dart apparatus, i.e. the snails possess either a complete dart apparatus with a dart sac plus an accessory sac and mucus glands (C. ...
... Schileyko (1978) classified these morphotypes as subspecies, whereas Giusti and Manganelli (1987) questioned whether they are related at all. Hausdorf (2001) followed Schileyko (1978), but questioned whether the morphotypes are actually genetic entities, because their ranges interdigitate in western Georgia. Furthermore, Hausdorf (2001) separated C. lasistana from C. frutis because of the much shorter spermatophore-forming flagellum. ...
Snails in the genus Cepaea are important model organisms in ecogenetic studies because of their colour and banding polymorphism. The monophyly of this group has been almost unanimously assumed based on superficial similarities in shell form and colouration. However, molecular phylogenetic analyses based on mitochondrial and nuclear sequences of 20 genera of Helicidae unequivocally demonstrated that Cepaea as currently understood is a polyphyletic assemblage. Only C. nemoralis and C. hortensis are retained in Cepaea, whereas C. vindobonensis is referred to Caucasotachea and C. sylvatica to Macularia based on our molecular phylogeny. Cepaea and Macularia belong to the western clade of the Helicinae, whereas Caucasotachea is nested in the eastern clade which probably diverged in the late Eocene. Because of the large phylogenetic distances between Cepaea, Macularia and Caucasotachea, it has to be shown whether the genetic mechanism underlying the simpler banding polymorphism in C. vindobonensis and M. sylvatica is a simpler version of the supergene that determines the polymorphism in Cepaea in the strict sense. This case illustrates the importance of sound phylogenetic analyses as a basis for any predictions in comparative biology.
Copyright © 2015. Published by Elsevier Inc.
... Circassina Hesse, 1921 (Gastropoda: Hygromiidae) is a group of land snails endemic to the western and central Caucasus region and the eastern Pontus. It includes only two species, C. frutis (Pfeiffer, 1859) and C. lasistana Hausdorf, 2001. Abchasohela Hudec & Lezhawa, 1971, which was formerly classified as a subgenus of Circassina (Hausdorf, 2001), proved to be more closely related to other hygromiid groups than to Circassina (Walther et al., in press). ...
... It includes only two species, C. frutis (Pfeiffer, 1859) and C. lasistana Hausdorf, 2001. Abchasohela Hudec & Lezhawa, 1971, which was formerly classified as a subgenus of Circassina (Hausdorf, 2001), proved to be more closely related to other hygromiid groups than to Circassina (Walther et al., in press). The Circassina frutis complex is exceptional among helicoid land snails because it is polymorphic with regard to the dart apparatus, i.e. the snails possess either a complete dart apparatus with a dart sac plus an accessory sac and mucus glands (C. ...
... Schileyko (1978) classified these morphotypes as subspecies, whereas Giusti and Manganelli (1987) questioned whether they are related at all. Hausdorf (2001) followed Schileyko (1978), but questioned whether the morphotypes are actually genetic entities, because their ranges interdigitate in western Georgia. Furthermore, Hausdorf (2001) separated C. lasistana from C. frutis because of the much shorter spermatophore-forming flagellum. ...
The phylogeny and historical biogeography of the Caucasian land snail genus Circassina was reconstructed using multilocus amplified fragment length polymorphism (AFLP) data and mitochondrial DNA sequences. Diversification within the group started with a divergence of populations from the western Lesser Caucasus from those of the Greater Caucasus during the late Miocene. Distinct AFLP clusters and major mitochondrial clades separated by long internal branches lend evidence to the hypothesis of separate glacial refuges in the Lesser and Greater Caucasus during the Pleistocene. High genetic distances across low geographic distances and admixture analysis revealed a phylogeographic boundary running through the Colchis lowlands, which may have been established and maintained in part by repeated transgressions of the Black Sea during the Pleistocene and Holocene. Localities in Ciscaucasia were probably colonised through long-distance dispersal across the main ridge of the Greater Caucasus. The phylogeny implies multiple independent losses of accessory genital organs, i.e. dart sac and mucus glands, within Circassina. None of the anatomically defined (sub-) species distinguished so far is monophyletic and there is gene flow between the two main population groups across the Colchis lowlands. Thus, we propose to classify these population groups as subspecies of a single species.
Copyright © 2015 Elsevier Inc. All rights reserved.
... The land mollusc's fauna of the Caucasus (318 species of Gastropoda, according to Walther et al. 2014) has been intensively studied over the last decades (Hausdorf 2000(Hausdorf , 2001Neiber et al. 2015;2022 etc.). At the same time, the nematode fauna associated with gastropods of this region has remained largely understudied. ...
... Species radiation has been investigated for several elements of the biota, including terrestrial gastropods, within the Caucasus hotspots Neiber et al. 2015Neiber et al. , 2017, but no data were available for the gastropod-associated nematodes prior to our study. However, the gastropod fauna of the Caucasus region has been extensively studied, and its phylogenetic relationships with gastropods of neighbouring regions and the evolutionary history of many taxa have been discussed (Hausdorf, 2000(Hausdorf, , 2001Neiber et al. 2022). The richness of the gastropod fauna in the Caucasus (318 species, 66% of which are endemic; Walther et al. 2014) suggests the same for the nematode fauna. ...
Caucasus is known as one of the few biodiversity hotspots in Europe and is characterised by rich gastropod fauna while the nematode fauna in association with gastropods has remained largely understudied. Surveys conducted in 2019 and 2021 in the North Caucasus of the Russian Federation (Stavropol Upland and western and central parts of Krasnodar Krai) has revealed the presence of three new species of Pellioditis , a facultative parasite of land gastropods, and two species of obligate parasites, the intestinal parasite Angiostoma kimmeriense and a new, still undescribed species of a larval ectoparasite Alloionema sp. The new associations of Cruznema sp. and Rhabditophanes sp. with land gastropods were recorded for the first time in the Russian Federation. The new species of Pellioditis Dougherty, 1953 described here is based on the analysis of morphology and molecular studies of two distant and morphologically distinct strains, thermalis and sindicae. Pellioditis thermalis n. sp. was characterised by females possessing a tail of about 95–100 μm long, broadly conical in shape in the thermalis and sindicae strains, with a rounded anterior part and a subulate terminal part as long as the former; prominent phasmids located at the mid-tail, equatorial vulva position, a lateral field of three ridges (four incisions), males with spicules featuring a hole at the distal tip, ensheathed infective juveniles with average length 717 μm in the thermals strain and 771 μm in the sindicae strain, and exsheathed ones 644 μm and 682 μm, respectively. ITS-based phylogenetic analyses revealed that all Pellioditis species found in Ciscaucasia and Transcaucasia probably belong to two separate clades, with independent evolutionary histories of colonisation of this area. The entire Caucasus range area appears to serve as a biodiversity hotspot for the genus Pellioditis , presumably due to its complicated geological history and repeated isolation events for its terrestrial mollusc hosts.
... As a result, large amount of taxonomic works usually directed by Russian scientists were published in Russian language and to lesser extent in Georgian language journals. Furthermore, a number of papers are still appearing regularly based on old preserved material kept outside Georgia (see for instance Hausdorf (2001) and Evsyukov et al. (2016)). Later on, much of the pre-modern work as well as data directly derived from museum collections was included in multiauthored series of books such as "Keys to Animals in the Fauna of the USSR" ("Фауна СССР" in Russian, published by the Zoological Institute of the Academy of Sciences of Russia). ...
... Further, biodiversity studies done by former Soviet scientists or, on rare occasions, by foreign researchers were usually based on existing literature sources and museum collections kept outside Georgia, without updating or obtaining new field data (e.g. Hausdorf 2001). ...
We evaluated progress towards animal biodiversity research in Georgia, a key area in the Caucasus biodiversity hotspot. By reviewing recently (1990–2018) published articles in all areas of animal diversity research, we unmasked the trends in biodiversity inventory, ecological and biogeographical studies, and conservation issues in Georgia. We concluded that species inventory and biodiversity research in Georgia has significantly increased during the last ten years, however the rate and extent of investigation is far from satisfactory. Major gaps remain in all branches of animal diversity research in Georgia, and consequently existing knowledge is inadequate to address modern challenges related to species and ecosystem conservation. We urge local governmental authorities and international scientific societies to support development of stronger research facilities and cultivate interest in biodiversity inventory and research in Georgia as an important step towards maintaining globally important biodiversity in the Caucasus.
... As a result, large amount of taxonomic works usually directed by Russian scientists were published in Russian language and to lesser extent in Georgian language journals. Furthermore, a number of papers are still appearing regularly based on old preserved material kept outside Georgia (see for instance Hausdorf (2001) and Evsyukov et al. (2016)). ...
... This trend has continued into the present day to some extent. A noteworthy condition of the first decade of independence was an almost complete absence of young researchers.Further, biodiversity studies done by former Soviet scientists or, on rare occasions, by foreign researchers were usually based on existing literature sources and museum collections kept outside Georgia, without updating or obtaining new field data (e.g.Hausdorf 2001).Hence, the first fifteen years of independence (period of 1990-2005) is represented by a marked gap in taxonomic and systematics research for Georgian fauna, with <10 indexed articles per year published. Instead, publication in local periodicals, most notably the Proceedings of the Institute of Zoology, was the main pathway for information sharing about Georgia in this time, even though almost all articles published there were non-peer reviewed, outdated, literature-based check lists, or systematic contributions. ...
We evaluated progress towards animal biodiversity research in Georgia, a key area in the Caucasus biodiversity hotspot. By reviewing recently (1990-2018) published articles in all areas of animal diversity research, we unmasked the trends in biodiversity inventory, ecological and biogeographical studies, and conservation issues in Georgia. We concluded that species inventory and biodiversity research in Georgia has significantly increased during the last ten years, however the rate and extent of investigation is far from satisfactory. Major gaps remain in all branches of animal diversity research in Georgia, and consequently existing knowledge is inadequate to address modern challenges related to species and ecosystem conservation. We urge local governmental authorities and international scientific societies to support development of stronger research facilities and cultivate interest in biodiversity inventory and research in Georgia as an important step towards maintaining globally important biodiversity in the Caucasus.
... Using this system, in this paper we revise the delimitation of the species of the Caucasigenini and discuss their geographical variation and distribution based on recently collected specimens as well as material from all major research collections. We describe all species of the Caucasigenini except those of the genera Circassina Hesse, 1921 and Fruticocampy laea Kobelt, 1871, which have been revised previously (Hausdorf 2001, Neiber & Hausdorf 2015, Walther et al. 2016a Figure 1. Bayesian 50% majority consensus tree based on the combined analysis of cox1, 16S rDNA and 5.8S rDNA + ITS2 + 28S rDNA sequences. ...
... Circassina and Fruticocampylaea have been revised previously (Hausdorf 2001, Neiber & Hausdorf 2015, Walther et al. 2016a) and will not be discussed here in detail. In the other genera, the 2 dart and 2 accessory sacs are about equally broad and the apical portions of the accessory sacs surmount the dart sacs. ...
The species systematics of the tribe Caucasigenini (Gastropoda: Hygromiidae) from the Caucasus region is revised based on recently collected specimens as well as material from all major research collections. Shells and genitalia of the 14 species assigned to the genera Caucasigena, Anoplitella, Hygrohelicopsis, Diodontella, Dioscuria, Teberdinia, and Lazicana are described and figured. Caucasigena armeniaca, C. rengarteni, C. schileykoi and the form from the Fiagdon valley labelled erroneously as C. tschetschenica, are classified as subspecies of C. eichwaldi because transitional forms among these taxa exist. Anoplitella antonwagneri is separated from A. schaposchnikovi as a distinct species. Diodontella nubigena is placed in the synonymy of D. stschukini, and Kokotschashvilia eberhardi as well as K. tanta are placed in the synonymy of Lazicana suanetica. Lazicana makvalae, Dioscuria thalestris, Dioscuria abchasica, Dioscuria caucasicola, Dioscuria prometheus, Teberdinia flavolimbata, and T. phaeolaema are confirmed as distinct. The anatomy of Dioscuria prometheus is described for the first time. The name Helix thalestris is fixed by the designation of a neotype for the taxon for which it is in current usage. The geographic ranges of the species are summarized based on a comprehensive compilation of locality records and illustrated in distribution maps. The morphological variability of the Caucasigenini species corresponds to their phylogenetic patterns. The Fruticocampylaea species are separated by long branches, probably caused by bottlenecks as a result of the Pleistocene ice ages, and diversified only recently. This is reflected in little intraspecific morphological variation. In contrast, Circassina, Caucasigena, and Lazicana show a continuous branching pattern, and the greater diversity of genetic lineages in these groups is reflected by more extensive intraspecific morphological variability. The contrast between the patterns found in Fruticocampylaea versus Circassina, Caucasigena and Lazicana demonstrates the importance of extinction in the speciation process. The differentiation of Fruticocampylaea into sharply delimited species was not an adaptive radiation into different niches, but was initiated by the extinction of intermediate populations. Such non-adaptive radiations are more likely in organisms with limited dispersal abilities, whereas ecological differentiation might play a more important role in the speciation of organisms with better dispersal abilities. © E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller) and Senckenberg Gesellschaft für Naturforschung, 2018.
... The possession of one or several dart sacs is usually a species-specific character. The only species of the Helicoidea which is known to be polymorphic with regard to the presence or absence of a dart sac is Circassina frutis (Pfeiffer, 1859) from the Caucasus region (Hausdorf, 2001;Neiber & Hausdorf, 2015;Schileyko, 1978). Schileyko (1996) also reported a lack of the dart sac in one of six examined Cylindrus obtusus (Draparnaud, 1805) from the eastern Alps. ...
... Sexual selection should result in a rapid fixation of morphs with dart sac or appendicula in helicoid species, because a higher proportion of the sperm of mutants without dart sac (or appendicula) will be digested by the copulation partner (Chase, 2007). Actually, Monacha kuznetsovi is the only species of the Helicoidea besides Circassina frutis (Hausdorf, 2001;Neiber & Hausdorf, 2015;Schileyko, 1978) that is known to be polymorphic with regard to the possession of a dart sac or its homologue, the appendicula. ...
During mating the dart apparatus of the land snail group Helicoidea transfers an allohormone that increases paternity success. Thus, this organ is under sexual selection. Despite this selective advantage, the dart sac has independently been lost in many lineages in the Helicoidea, presumably because natural selection counteracts sexual selection. However, the selective pressures that result in the loss of the organ have not been examined experimentally because most species show no variation in the presence or absence of the dart sac. Using nuclear multilocus data and mitochondrial sequences, we show that a so far misidentified morph of the genus Monacha without appendicula, a homologue of the dart sac, is conspecific with M. kuznetsovi with appendicula. This is only the second case of polymorphy with regard to the presence or absence of the dart sac or its homologue in the Helicoidea. Monacha kuznetsovi might therefore be a suitable model organism to study how the interplay between sexual selection and natural selection may affect the evolution of the genital organs.
... It can be speculated that the divergence between these two strains occurred in the Caucasus region, with its well-known isolation events (Nikishin et al., 2010) and reputation as a biodiversity hotspot (Mumladze et al., 2014;Neiber et al., 2015Neiber et al., , 2017. Rapid species radiation in the Caucasus biodiversity hotspot has been demonstrated for terrestrial gastropods (Hausdorf, 2000(Hausdorf, , 2001Neiber et al., 2022) and, more recently, for gastropod-associated nematodes (Ivanova & Spiridonov, 2023). ...
A member of the genus Pellioditis Dougherty, 1953 (syn. Phasmarhabditis Andrássy, 1976) was detected in three introduced gastropod species in Moscow for the first time. Both the nematode and its gastropod hosts were found to have originated from the Caucasus region. No native gastropod species as well as the alien invasive slug Arion vulgaris were infected by P. akhaldaba . The Moscow strain of P. akhaldaba is very close to P. akhaldaba from Georgia in the ITS rDNA (7 bp difference) but they differ significantly in the sequences of COI mtDNA (50 bp difference). However, phylogenetic analysis of both loci showed that these two strains were the closest relative to all other species and strains of the genus Pellioditis studied.
... The genera in question are Caucasigena Lindholm, 1927(including Anoplitella Lindholm, 1929and Dioscuria Lindholm, 1927Nordsieck, 1993 proposed to classify Dioscuria as a distinct genus), Diodontella Lindholm, 1929, Hygrohelicopsis Schileyko, 1978, Kokotschashvilia Hudec & Lezhawa, 1969, and Teberdinia Schileyko, 1978. However, Schileyko (1978b, 1991 did not recognize that also Fruticocampylaea Kobelt, 1871, Shileykoia Hudec, 1969(synonymized with Fruticocampylaea by Walther, Neiber, & Hausdorf, 2016a, and Circassina Hesse, 1921, with one dart apparatus (which is even lost in some Circassina frutis (Pfeiffer, 1859) populations; see Schileyko, 1978b;Hausdorf, 2001;Neiber & Hausdorf, 2015) were also descendants of this radiation, but classified them in the Hygromiinae or Hygromiini. Nordsieck (1993) classified the groups with two dart apparatus also in the Trichiini, whereas he included the groups with one dart apparatus together with Kalitinaia Hudec & Lezhawa, 1967(now Geomitridae), Caucasocressa Hesse, 1921(now Trochulinae, Monachaini), Metafruticicola Ihering, 1892, and Hiltrudia Nordsieck, 1993, in the Metafruticicolini Schileyko, 1972. ...
The Caucasigenini is an endemic radiation of hygromiid land snails from the Caucasus region. A phylogenetic analysis of morphological characters of the genitalia and the shell showed that the morphological characters are insufficient for resolving the relationships within the Caucasigenini. Convergences of the few parsimony informative characters in other groups of the Hygromiidae demonstrate that these characters are not reliable indicators of phylogenetic relationships. Phylogenetic analyses of sequences of cox1, 16S rDNA, 5.8S rDNA, ITS2 and 28S rDNA revealed several well-supported groups. The relationships among these groups could not be resolved. It is likely that these groups originated in a rapid radiation during the uplift of the Caucasus. Based on the molecular phylogeny, we propose a new classification of the species of the Caucasigenini and establish a new genus, Lazicana gen. n.
... Evidence from the hygromiid genus Circassina Hesse, 1921(Hausdorf, 2001Neiber & Hausdorf, 2015a) and other helicoid families (e. g., Mejía & Zúñiga, 2007;Hirano et al., 2014;Köhler & Criscione, 2015) suggests that high levels of homoplasy in the genital system may also be more widespread in the Hygromiidae than previously expected. In the present contribution we will reconstruct the phylogeny of the Hygromiidae in the sense of Razkin et al. (2015) on the basis of a broad taxonomic sampling covering almost the complete generic diversity from the entire range of the family using mitochondrial as well as nuclear DNA sequences. ...
The family Hygromiidae is a highly diverse group of land snails with a distribution
range stretching throughout the Palearctic region from the Macaronesian Islands
in the west to the Russian Far East in the east and reaching southwards to the
northeastern Ethiopian region. The classification of the family largely rested on few
characters of the dart apparatus of the genitalia so far. We used nuclear and mitochondrial
DNA sequences of almost all genera to reconstruct the phylogeny of the
Hygromiidae. The molecular phylogenetic trees indicate widespread homoplasies
with regard to the characters of the dart apparatus so that many of the currently
accepted subdivisions of the family turned out to be polyphyletic. We newly delimit
three subfamilies within the family Hygromiidae, Hygromiinae (with three tribes),
Leptaxinae (with three tribes) and Trochulinae (with nine tribes), on the basis of our
phylogenetic analyses and describe Pseudotrichiini n. trib., Perforatellini n. trib.,
Cryptosaccini n. trib., Ganulini n. trib., Urticicolini n. trib., Caucasigenini n. trib.
and Ashfordini n. trib. as new tribes and Coronarchaica n. gen. and Noricella
n. gen. as new genera. The reconstruction of the biogeographical history of the
Hygromiidae indicated an origin of the family in the western Palaearctic, from
where the Tian Shan and Altai Mountain regions in Central Asia, the Macaronesian
Islands, the Caucasus region and sub-Saharan East Africa were colonized.
... More recently, new material was collected by A. N. Suvorov, P. V. Kijashko and L. Mumladze, who described several new species (Suvorov & Schileyko, 1991;Schileyko & Kijasko, 1999;Suvorov, 2002Suvorov, , 2003Suvorov, , 2006Kijashko, 2005Kijashko, , 2006Mumladze et al., 2008). B. Hausdorf revised some, at that time, poorly understood genera (Monacha, Circassina, Caucasocressa) that all belong to the family Hygromiidae (Hausdorf, 2000b(Hausdorf, , 2001(Hausdorf, , 2003 and the Orculidae from the Caucasus region (Hausdorf, 1996). Egorov (2008) and Sysoyev & Schileyko (2009) published synoptical works on XI. 1849, † 29. ...
... Also in other hygromiids polymorphism with regard to the presence of a dart (and accessory) sac, the homologue of the appendicula is very rare. The only known case is that of Circassina frutis (Schileyko 1978;Hausdorf 2001;Neiber & Hausdorf 2015a). The mucus glands, which are part of the dart apparatus of helicoids, produce an allohormone-like substance that is usually transmitted into the mating partner during courtship by the calcareous dart. ...
Monacha is the most species-rich genus of the family Hygromiidae with a centre of diversity in Anatolia. On the basis of the presence or absence of accessory genital appendages, the group was subdivided into three subgenera, Monacha s. str., Paratheba and Metatheba, in the past. We used mitochondrial and nuclear DNA sequences of a representative sample of species 1) to reconstruct the phylogeny of the major lineages of Monacha, 2) to reconstruct the evolution of the accessory genital appendages, and 3) to reconstruct the biogeography of the group. Our results show that the accessory genital appendages upon which the classification of Monacha into subgenera rested so far, that is the appendicula, which is homologous to the dart sac and/or its accessory sac in other helicoid land snails, and the penis retractor muscle, were lost several times independently in different lineages of Monacha. Even among sister species, these characters were found to be variable. Thus, the typologically defined subgenera are para- or polyphyletic assemblages. The reconstruction of the biogeographical history indicated an origin of Monacha in Anatolia and the adjacent Caucasus region which is consistent with the observation that most other genera currently included in the subfamily Monachainae are also endemic to this region. Most major Monacha lineages remained restricted to northern Anatolia. Southern Europe was colonized by Monacha s. str., the Aegean region by Aegaeotheba subgen. n., the Crimean Peninsula by Paratheba and the Levant by Platytheba. On the basis of our phylogenetic analyses and testing of alternative hypotheses, we propose to divide Monacha into eight subgenera. We newly delimit the previously proposed subgenera Monacha s. str., Metatheba, Paratheba and Platytheba, and describe Pontotheba subgen. n., Aegaeotheba subgen. n., Trichotheba subgen. n. and Rhytidotheba subgen. n. as new subgenera.
... However, many species listed by him have subsequently been eliminated by synonymy, while others have been newly described or found in further localities. These data have therefore been supplemented by personal collection by L.M. and by information from Riedel (1966), Hausdorf (2000Hausdorf ( , 2001Hausdorf ( , 2003, Egorov & Greke (2005), Sysoev & Schileyko (2005), Schu¨tt (2005), Neubert & Bank (2006) and Pokryszko et al. (2011). Many of these studies also draw upon earlier museum records. ...
The globally significant Caucasus hotspot of biological diversity holds a rich and largely endemic fauna of land molluscs.
Georgia holds the majority of these regional endemics. Land molluscs are particularly sensitive indicators of habitat quality
and faunal diversity. In this study, we examine the extent to which the existing network of protected areas (PAs) within Georgia
captures the hotspots of endemic molluscan diversity. We collected and mapped the records of Georgian and Caucasian endemic
species onto a 20 × 20 km2 UTM grid to identify the most important endemic areas in Georgia. We related these to the existing network of PAs. Less than
half of the richest grid cells included significant PAs. Although those endemics with the smallest known ranges were better
protected than the remainder, the incomplete state of knowledge means that our estimates of existing protection are surely
optimistic. To date, the designation of PAs in Georgia has not used distributional data for invertebrates, although elsewhere
they have been shown to be an effective aid to planning and management for conservation. Further surveys of molluscs and their
monitoring in existing PAs can and should inform a systematic conservation strategy in Georgia.
У каталозі в систематичному порядку описано матеріали по наземних молюсках, які зберігаються в основному фонді Державного природознавчого музею НАН України. Вони репрезентують 250 видів, зібраних на території України та інших країн. Наведено короткі відомості щодо історії формування малакологічного фонду музею та, зокрема, фондової колекції наземних молюсків, її сучасного стану, а також деякі зауваження фауністичного і систематичного характеру. Каталог буде корисним науковцям-зоологам, аспірантам, студентам біологічних спеціальностей вищих навчальних закладів.
У каталозі в систематичному порядку описано матеріали по наземних молюсках, які зберігаються в основному фонді Державного природознавчого музею НАН України. Вони репрезентують 250 видів, зібраних на території України та інших країн. Наведено короткі відомості щодо історії формування малакологічного фонду музею та, зокрема, фондової колекції наземних молюсків, її сучасного стану, а також деякі зауваження фауністичного і систематичного характеру. Каталог буде корисним науковцям-зоологам, аспірантам, студентам біологічних спеціальностей вищих навчальних закладів.
Sistematic inventory and monitoring of the mollusc fauna present in the "Bosque Vereador Pedro Medeiros" (total of 15 native and 9 synanthropic exotic gastropod species), a urban park with preserved remanescent Atlantic forest in the continental Florianópolis territory, Santa Catarina State/ SC, Central Southern Brazil region, is available (pp. 16-17) ...
Over the last 30 years, a considerable amount of Metafruticicola material has been collected. Here, a monographic revision is provided of the genus, which is mainly based on conchological characters, but descriptions of the genital system of eight species (and in addition two subspecies) are given as well. The distribution area of the genus, namely the eastern Mediterranean, ranges from Albania to Isreal.In total,24Species are recognized,Which are Classified into four subgenera Two species are polytypic, accounting for eight subspecies. All synonyms and localities are listed, the distribution is illustrated on maps, and scanning electron micrographs are given to provide an insight of the shell microsculpture (protoconch and teleoconch). New for science are: one subgenus (Rothifruticicola), six species (M. crassicosta, M. monticola, M. ottmari, M. pieperi, M. rugosissima, M. uluborluensis) and one subspecies (M. nicosiana maasseni).©2013 E. Schweizerbartsche Verlagsbuchhandlung (Nagele u. Obermiller).
The Caucasus region is among the most important Eurasian biodiversity hotspots (Zazanashvili et al. 2004, Walther et al. 2014). Its mollusc fauna is far from being well-explored and, as indicated by a number of recent studies, one could reasonably expect further taxonomical discoveries in this region. The diversity and distribution of the mollusc fauna are especially little known, as only few precise locality records have been published for most of the species (Walther et al. 2014, see also http://www.caucasus-snails.uni-hamburg.de/CaucasianLandSnails-Dateien/Checklist.html). This paper has three objectives, namely (i) clarifying the question of possible genital polymorphism in Pontophaedusa funiculum (Mousson) (Fig. 1), (ii) presenting georeferenced distribution records (Appendix 1) and shell photographs (Figs 2–15) of clausiliid species collected during a field trip in Georgia in June, 2012 in order to contribute the better knowledge of the region's biodiversity, and (iii) providing a comprehensive list of Caucasian door-snail literature.
The genus Caucasocressa is revised. Ten Caucasocressa species are known from north-east Turkey and south-west Georgia. Five new species are described. For every taxon the shell is described and illustrated and the variability of the genitalia (as far as known) is shown. All synonyms are listed. The distribution is summarized and all locality records are listed. The distribution of the species is illustrated on a UTM grid-map.
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