Corresponding author: Barna Páll-Gergely (
Academic editor: Eike Neubert
The genital anatomy of
The occurrence of only one
The anatomy of the Alpine and the Illyric
Recently,
Recently, living specimens of
Furthermore, we discuss the orculid species reported from Romania. Two species, namely
The comprehensive map (
Photographs of several focal planes were made with a Wild Makroskop M420 and a Nikon DS Camera Control Unit DS-L2. The different layers were combined with Helicon Focus 4.75 Pro to obtain one completely focused image.
Shells were directly observed without coating under a low vacuum SEM (Miniscope TM-1000, Hitachi High-Technologies, Tokyo). Teleoconch sculpture was noted on the dorsal or dorsolateral area of the penultimate whorl.
Shell yellowish–greenish to dark brown; cylindrical to conical and elongated; 8–10 weakly convex whorls; sculpture of first 0.5–1.0 protoconch whorl usually smooth, but may be of fine spiral lines, which may be extremely weak; teleoconch axial sculpture variable, ranging from irregular growth lines to equally spaced, conspicuous radial structure; apertural barriers: one parietal and 1–3 columellar lamellae; palatal side of the aperture smooth or with strong tooth or thickening parallel to the apertural lip; parietal callus weak, subangularis sometimes present; palatalis plicae missing.
Penis cylindrical, penial caecum of variable length and shape; penial appendix absent; interior of penis, epiphallus and caecum with longitudinal folds; retractor muscle attaches to the penis-epiphallus junction on the opposite side of the penial caecum; diverticulum absent; distal part of vas deferens sometimes slightly swollen, entering epiphallus terminally; bursa copulatrix long, club-like.
Detailed anatomical and conchological diagnoses were provided by
Some African genera, such as
In general, species of
Shells and a living specimens of
SEM of shells of various
SEM of the shell sculpture of various
Genital anatomy of
Genital anatomy of
Genital anatomy of
Schematic drawings of the penial caecum of
Spermatophores.
Distribution map of
Shell smoothish with irregular growth lines; apex somewhat conical, not blunt; aperture with 2–3 columellar lamellae; penial caecum simple and usually longer than half the length of the penis; its base often not conspicuously thickened.
The soft anatomy of various
The penial caecum of
A third columellar lamella is rarely present, but can occur in a small percentage of individuals within a population.
Most species have limited distributions in the Alps (mainly Austria).
Our knowledge of the distribution of
Shell usually with strong axial sculpture (irregular ribs), with two columellar lamellae, apex rather rounded, not attenuate. The penial caecum usually consists of two parts (‘’tubercles’’) and its length is less than half that of the penis.
The name of this new subgenus refers to its distribution in the Illyrian and Banatic biogeographical regions. It is feminine.
Montenegro, Albania, Northwestern Greece, Kosovo, Macedonia (
The reproductive anatomy of
RO, Jud. Bihor, Munţii Pădurea Craiului, Şuncuiuş, Valley of Crişul Repede, in front of Peştera Vantului (cave), limestone, leg.: Bata, Danyik, Deli, 11.04.2011. (anatomically examined); RO, Gyalui-havasok (Munţii Gilăului), Runki szakadék (gulch of Runk), leg. Papp, J., 22.07.1959, HNHM 73030/3; RO, Bihar Mts (Munţii Apuseni)., Felsőgirda (Gârda de Sus), Ordincus valley., leg. Kovács, Gy., 30.05.1985, HNHM 68284/2; RO, Muntii Bihorului, Baita, Piatra Graitoare, environment of the Crisu Baitei River, leg. Kovács, Gy., 23.08.1974, HNHM 68283/2; Černath. (Chernathal) bls Badern (?) v. Mehadia, leg. Jetschin 1882, NHMSK 4874/5; Forstgra (Forstgartens) bei im Černathal, Banat, leg. Jetschin 1882, NHMSK 4875/2; Klausenburg, Györgyfalvaer Wald, leg. Marzlof 1891, NHMSK 7470/8; Zalathna gegen den Judenberg, leg. Barth 1866–1906, NHMSK 7468/8; Banat, Herkulesbad, leg. Deubel 1895 May–Juni, NHMSK 7469/6; Steierdorf bis zur Höhle Panur, leg. Jetschin 1885, NHMSK 4876/4; Černathal b. Mehadia, leg. Jetschin 1885, NHMSK 4877/4 (‘’
Two specimens were anatomically examined. Penis slim, with the retractor muscle attached at its distal end; penial caecum very small, vestigial, consisting of two “tubercles”; epiphallus very long and cylindrical; there is clear distinction between the vas deferens and the epiphallus; vas deferens long and relatively thick; a slender retractor muscle is attached near the proximal end. Vagina short and thick, but pedunculus relatively long; bursa copulatrix extremely long, with the distal end slightly expanded. In one specimen an elongated, simple spermatophore was found with the apical portion slightly thickened.
The species inhabits deciduous forests. It is found most commonly between small stones and leaf litter on the forest floor or under hazelnut (
Least concern (LC) according to IUCN criteria (
All living specimens found were covered in mud, causing them to appear like tiny grains of soil. The ribbed shell is possibly an adaptation for camouflaging. The photographs herein are of cleaned shells.
Albania, Bjeshkët e Nemuna (Prokletije Mts), above village Okol, near pass Qafa e Pejës, W slope of Mt. Maja e Popluks, at a spring on limestone, 1660 m,
One specimen was dissected. Penis cylindrical and slim, with a short, but thick penial caecum, the proximal portion broader than the short and slimmer distal portion; retractor muscle attaches at the penis–epiphallus transition; epiphallus more than twice as long as the penis and much thicker, its transition to the vas deferens is gradual, barely discernable; there is a slim retractor muscle attached to the proximal portion of the epiphallus; proxim\al portion of the vas deferens thicker than the distal part. Vagina and free pedunculus extremely short; bursa copulatrix almost twice as long as the combined length of the penis–epiphallus complex.
Our observations on the genitalia agree with that of
Shell with conical apex and strong axial sculpture (irregular axial growth lines), with three columellar lamellae (columellar, supracolumellar and one short lamellae above), palatalis reaches its maximum height on the dorsolateral side. Penial caecum very long with thickened base, canal connecting the proximal end of the epiphallus to the penial caecum.
The new subgenus is named in honour of Dr Bernhard Hausdorf (University of Hamburg), who first noted the unusual shell characters of
See under
According to
Bulgaria, Strandzha Mts., Kondolovo village,
Two specimens were dissected. Penis cylindrical, relatively long; retractor muscle short, attaches on the proximal portion; penial caecum very long, with a thickened base and a cylindrical distal portion; an additional canal (?) connects the proximal end of the epiphallus with the penial caecum; epiphallus long, with a thickened distal part; the separation between the vas deferens and epiphallus is distinct; vas deferens relatively thick. Vagina cylindrical and relatively short; bursa copulatrix extremely long with a pointed end.
A developing egg covered with small calcareous crystals was found in the uterus of the figured specimen. In the other specimen, an elongated, simple bursa copulatrix was found with a slightly thickened apical part.
South-Eastern Bulgaria and North-Western Turkey.
The type series (12 shells) of
We were able to find
The species is very rare wherever it has been encountered yet, especially in Turkey. On two occasions, in 2007 and 2010, Barna Páll-Gergely spent about 4–5 hours at the locality in vil. Bursa, but found only one specimen in 2007. The other locality (Vil. Bolu) was visited in 2005 and 2006 for similar lengths of time and only one specimen was found in 2006. Atanas Irikov collected 9 living specimens and about 10 empty shells in an hour near Kondolovo in Bulgaria.
Listed as Vulnerable (V) under IUCN criteria (
Two of four living specimens had beetle (possibly drilid beetle) larvae in the body whorl.
The dissected specimens were collected about 23 km south-southwest of the type locality. The Strandzha Mountains (incl. the collecting site) belongs to the drainage of the Ropotamo River. It is reasonable to suppose that Urbański’s population was “washed down” from somewhere in the Strandzha Mts. and settled a temporary subpopulation in the Ropotamo floodplain. This might be a reasonable explanation why A. Irikov could not find this species in the type locality.
Based on available information we suggest using
Two Romanian samples of Grossu in the collection of MNINGA are labelled as
In this paper we describe the genitalia of the Eastern European
The genus can be subdivided into three subgenera (
Three species included herein have some shell and anatomical characters which differ from characters used to the features mentioned in the diagnoses of certain subgenera: (1) the shell sculpture of many populations of
Based on available literature, the occurrence of
We are very grateful to those colleagues who provided access to their museum collections: A. Eschner (NHMW), E. Neubert (NMBE), R. Janssen (SMF), A. Mesaroş (NHMS), Z. Fehér (HNHM), to O. Merkl (HNHM) for determining the beetle larvae found in