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The Genus Trochulus Chemnitz, 1786 (Gastropoda: Pulmonata: Hygromiidae) - A Taxonomic Revision

December 2009
Folia Malacologica 17(3):101-176

December 2009
17(3):101-176

Authors:

University of Wroclaw

The revision of the genus Trochulus Chemnitz, 1786 is based on conchological and anatomical characters of 4,377 adult specimens (3,190 dry shells and 1,187 alcohol-preserved snails) from 303 localities. Shell descriptions with the variation ranges, based on morphometric analysis, reproductive system descriptions with figures, identification key, synonymy, and distribution maps are provided. The most significant diagnostic characters are: shell shape and colour, umbilicus shape and size, durability of hairs, penis/epiphallus length ratio, flagellum/epiphallus length ratio, number and arrangement of mucous glands, length and location of dart sacs. Three names are synonymised: T. plebeius, T. concinnus and T. hispidus. Phylogenetic analysis yielded 96 equally parsimonious trees; the strict consensus procedure resulted in an incompletely resolved cladogram, with two distinct monophyletic groups of species: the villosus group including villosus, czarnohoricus, unidentatus and bielzi, and the striolatus group, with striolatus, graminicolus, montanus, caelatus, hispidus, suberectus, edentulus, bakowskii, luridus, filicinus and leucozonus. These two groups and erjaveci, biconicus, villosulus, clandestinus, lubomirskii and piccardi, form a phylogenetic bush. The monophyletic groups appearing in the cladogram are not compatible with any of the earlier proposed divisions. Most members of Trochulus inhabit the Alps and the Carpathians. The only widely distributed species is T. hispidus. The second most widespread species is T. striolatus. The remaining species are endemics with limited ranges. The number of species within particular parts of the range points to the Alpine-Carpathian area as the diversity centre of the genus.

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THE GENUS TROCHULUS CHEMNITZ, 1786

(GASTROPODA: PULMONATA: HYGROMIIDAE)

– A TAXONOMIC REVISION

MA£GORZATA PROÆKÓW

Museum of Natural History, Wroc³aw University, Sienkiewicza 21, 50-335 Wroc³aw, Poland

(e-mail: mprockow@biol.uni.wroc.pl)

BSTRACT: The revision of the genus Trochulus Chemnitz, 1786 is based on conchological and anatomical char

acters of 4,377 adult specimens (3,190 dry shells and 1,187 alcohol-preserved snails) from 303 localities. Shell

descriptions with the variation ranges, based on morphometric analysis, reproductive system descriptions with

figures, identification key, synonymy, and distribution maps are provided. The most significant diagnostic

characters are: shell shape and colour, umbilicus shape and size, durability of hairs, penis/epiphallus length

ratio, flagellum/epiphallus length ratio, number and arrangement of mucous glands, length and location of

dart sacs. Three names are synonymised: T. plebeius, T. concinnus and T. hispidus. Phylogenetic analysis yielded

96 equally parsimonious trees; the strict consensus procedure resulted in an incompletely resolved cladogram,

with two distinct monophyletic groups of species: the villosus group including villosus, czarnohoricus, unidentatus

and bielzi, and the striolatus group, with striolatus, graminicolus, montanus, caelatus, hispidus, suberectus, edentulus,

bakowskii, luridus, filicinus and leucozonus. These two groups and erjaveci, biconicus, villosulus, clandestinus,

lubomirskii and piccardi, form a phylogenetic bush. The monophyletic groups appearing in the cladogram are

not compatible with any of the earlier proposed divisions. Most members of Trochulus inhabit the Alps and the

Carpathians. The only widely distributed species is T. hispidus. The second most widespread species is T.

striolatus. The remaining species are endemics with limited ranges. The number of species within particular

parts of the range points to the Alpine-Carpathian area as the diversity centre of the genus.

EY WORDS: terrestrial snails, Hygromiidae, Trochulus, revision, phylogeny

INTRODUCTION

The genus Trochulus is commonly known as Trichia,

but the ruling by ICZN, Opinion 2079 (BZN 61 (3)

Sep 2004) resulted in replacing Trichia Hartmann,

1840, which turned out to be a junior homonym of

Trichia de Haan, 1839 (Crustacea: Brachyura), with

Trochulus Chemnitz, 1786.

Prior to this study the genus Trichia included 20

nominal species (K

ERNEY et al. 1983) inhabiting Eu

rope, especially the Alps and the Carpathians, some

of them with many subspecies and forms (W

AGNER

1915, KLOETI-HAUSER 1920, POLIÑSKI 1928, FORCART

1965, SHILEYKO 1978a). Recently, a new species Tro

chulus piccardi Pfenninger et Pfenninger

, 2005 was de

scribed from the Swiss Alps. Moreover, some species

from northern Africa were included in the genus:

Trichia roseotincta (Forbes), T. faidherbiana (Bour

guignat), T. baccueti (Bourguignat), T. mongrandiana

(Bourguignat) and T. zonitomaea (Letourneux). The

anatomical data provided by S

ZIGETHYI (1976) who

analysed only the penial papilla of these species, do

not solve the problem of their generic status. The

conchological features of the specimens deposited in

the collection of the Museum and Institute of Zool

ogy, Polish Academy of Science (Warsaw) and habitat

preferences of some species suggest that in all likeli

hood they represent another genus and, conse

quently, they are excluded from this publication.

None of the numerous previous papers dealt with

all the members of the genus (W

AGNER 1915, KLOETI-

HAUSER 1920, POLIÑSKI 1928, FORCART 1965,

HILEYKO 1978a, BANK 1995), their classification and

phylogenetic relationships among them were ob

FOLIA

MALACOLOGICA

ISSN 1506-7629

The Association of Polish Malacologists

& Faculty of Biology, Adam Mickiewicz University

Poznañ 2009

Vol. 17(3): 101–176

doi:10.2478/v10125-009-0013-0

scure, and there were no data on their conchological

and anatomical variation; likewise there was no infor

mation on the anatomy of some species.

Most species of Trochulus show a wide concho

logical variation, resulting in identification problems.

A conspicuous example is the fact that L

OCARD

(1895–1896) – a representative of the French Nou

velle École – distinguished 55 species of the genus in

France alone. As a result of the synonimisation done

by G

ERMAIN (1929) their number decreased to three!

An additional impediment is the fact that most of the

original descriptions are very laconic and it is unclear

which species they refer to, besides, the type speci

mens of most species have not been preserved.

Considering the identification difficulties, the lack

of data on the shell and genital variation ranges of

most species (except shell variation in T. hispidus, see

AGGS 1985, PROÆKÓW 1997, and some morpho

metric data on T. leucozonus, see B

ANK 1995) and in

completeness of the distribution data, a revision of

Trochulus seemed to be necessary.

HISTORICAL ACCOUNT

The Trochulus research history starts with

INNAEUS’s (1758) description – as Helix hispida – of

the first and most common member of the genus.

Subsequent species, like other members of the family

Hygromiidae, were for a long time placed in the ge-

nus Helix Linnaeus, 1758.

Two trends, resulting in two groups of papers, can

be distinguished in the previous studies on Trochulus:

some authors tried to solve classification problems

within the genus, while others focused on detailed in

vestigations of lower rank taxa, describing new spe

cies, subspecies and forms, or providing detailed data

on distribution, ecology and anatomy of some mem

bers of the genus.

The first attempts at revisions were made by

LESSIN (1874) and LOCARD (1888) at the end of the

19th c. Both studies were based only on conchological

characters and the latter author in his subsequent pa

pers (L

OCARD 1895–1896), like RISSO (1826), GRAS

(1840), BOURGUIGNAT (1864), MABILLE (1868,

1877), L

ETOURNEUX (1869), RAMBUR (1869),

ERVAIN (1880) and CAZIOT (1910), introduced an

excessive number of specific names, having no justifi

cation in reality.

AGNER (1915) published a revision of the Balkan

members of the genus, based on both conchological

and anatomical characters. He included the investi

gated species in the genus Fruticicola sensu W

AGNER

(1915), in which he distiguished two subgenera:

Fruticicola sensu stricto and Perforatella Schlüter, 1838.

However, in his study he did not specify the subgener

ic characters. Fruticicola was characterised as inhabit

ing “Westliche Balkanländer von der Save bis nach

Mazedonien” (W

AGNER 1915: 51 [479]), and

Perforatella as snails with a narrow shell and partly or

entirely covered umbilicus, 5.5–7 slowly increasing

whorls, a well developed lip where a basal tooth may

be present, shell surface with fine growth lines and

dense, short setae. He described their radulae and re

productive systems as typical (W

AGNER 1915: 58

[486]).

LOETI-HAUSER (1920) published a revision of

seven Swiss species of Trichia, describing their genital

characters. However, the author neither described

the shells nor preserved the specimens; he only gave

exact data on his collecting localities.

OLIÑSKI (1928) carried out a critical concho

logical and anatomical revision of the Alpine and

Carpathian species. He placed them within three sec

tions of the genus Fruticicola Held, 1837: Petasina

Beck, 1847, Filicinella Poliñski, 1928 and Edentiella

Poliñski, 1928, giving shell and reproductive system

characters, and type species as well. He also analysed

the distribution of the included species.

In 1965 F

ORCART clarified the taxonomic status of

some problematic species, based on conchological

and anatomical studies of materials collected, among

other localities, in the sites listed by K

LOETI-HAUSER

(1920).

HILEYKO (1978a, b) provided shell and reproduc-

tive system descriptions of Trichia species inhabiting

the area of the former Soviet Union and proposed

their cassification. Trichia sensu auct. (W

AGNER 1915,

OLIÑSKI 1928, FORCART 1965) and closely related

genera were treated as “Trichia sensu lato” and di-

vided into several genera. The members of

Trichia/Trochulus in its traditional sense were included

in three of them, i.e. Plicuteria Shileyko, 1978, Trichia

Hartmann, 1840, with subgenera Petasina Beck, 1847

and Trichia s. str., and Edentiella Poliñski, 1924.

ALKNER (1985) accepted POLIÑSKI’s (1928)

“groupe Perforatella auct.” as “eine echte monophy

letische Einheit und lassen sich – ebenfalls unter der

Einschränkung des jetzigen Kenntnisstands – mit

keiner anderen Gattung der Trichiinae zu einer

gesichert monophyletischen Gruppe vereinigen” and

thereby elevated the subgenus Petasina Beck, 1847 to

the generic rank, retaining its contents and system as

proposed by P

OLIÑSKI (1928).

Later N

ORDSIECK (1993), in his proposed classifi

cation system of Palaearctic Hygromiidae, distin

guished three genera within Trichia sensu K

ERNEY et

al. (1983), i.e. Trichia Hartmann, 1840, Petasina Beck,

1847 (+Edentiella Poliñski, 1928) and Plicuteria

Shileyko, 1978. Petasina was divided in two subgenera:

Petasina s. str. and Edentiella.

Recently, three molecular analyses of helicoid phy

logeny have been published (S

TEINKE et al. 2004,

OENE &SCHULENBURG 2005, MANGANELLI et al.

2005). However, they deal only with some species and

focus on phylogenetic analysis at the level of family or

selected genera.

102 Ma³gorzata Proæków

Among the researchers dealing with the systemat

ics of Trochulus at the specific or subspecific level it is

necessary to mention W

AGNER (1915) and POLIÑSKI

(1928), both of whom described a large number of

subspecies, and F

ORCART (1965), who tried to unravel

the systematic complexity among Trichia hispida sensu

lato. Only few studies dealt with morphometric analy

sis of Trichia from Great Britain (N

AGGS 1985) and

shell variation of T. hispida from Poland (P

ROÆKÓW

1997). KRAUSP’s (1952), MAHLER’s (1952), HUDEC’s

(1964, 1965, 1972) and K

AISER’s (1966) papers all

contributed to the knowledge of the genus, as well as

later F

ALKNER’s (1985) and BANK’s (1995) articles

aiming to solve the systematic position of some sub

species in the genus Petasina.

Trochulus plebeius and T. hispidus in the Czech Re

public have been subject to karyological analysis. All

the studied populations have identical numbers of

chromosomes (n=23) of a similar morphology. Se

quence analysis of 16S rDNA and COl genes showed

two divergent groups of T. hispidus populations, dif

fering in their geographic distribution (north-western

versus eastern) but with overlapping morphology

RABAKOVA et al. 2007).

The recent Fauna Europea project (B

ANK 2006)

lists 49 taxa of Trochulus and Petasina. They belong to

two (Trochulus with 26 taxa and Plicuteria with 1

taxon), and three (Petasina with 6 taxa, Edentiella with

7 taxa and Filicinella with 9 taxa) subgenera, respect

ively. The high number of subspecies (20) is striking.

Some of them seem to be dubious entities requiring

further studies.

MATERIAL AND METHODS

The material used in the study was obtained from

the following collections: BMNH – The Natural His-

tory Museum, London, England, MIZW – Museum

and Institute of Zoology, Polish Academy of Science,

Warsaw, MPW – Natural History Museum, Wroc³aw

University, NHMB – Naturhistorisches Museum,

Bern, NHMW – Naturhistorisches Museum, Wien,

UAM – Department of General Zoology, Adam

Mickiewicz University, Poznañ, U£ – Department of

Invertebrate Zoology and Hydrobiology, University of

£ódz, ZW – specimens collected especially for the

purpose of this study, now deposited at MPW.

A total of 4,377 adult specimens, including 3,190

dry shells and 1,187 alcohol-preserved snails were

examined. In the list of samples “alc.” denotes

alcohol-preserved material and “s.” – dry shells. The

shells were measured with the accuracy of 0.1 mm.

The measurements are shown in Fig. 1. Besides, the

following coefficients of shell proportions were calcu

lated: height/width ratio (H/W), relative height of

body whorl = body whorl height/shell height ratio

(bwH/H), umbilicus relative diameter = umbilicus

major diameter/shell diameter ratio (U/D), ratio of

umbilicus major to its minor diameter (u/U). Whorls

were counted according to E

HRMANN’s (1933)

method. Morphometric data were statistically ana

lysed with respect to the significance of differences

between populations (Duncan test). Statistical analy

sis was performed with Statistica 5. The specimens for

anatomical examination were drowned in water and

preserved in 70–75% ethyl alcohol. Prior to dissection

their shells were dissolved in 1N (10%) HCl (5–10

min), and the remaining periostracum removed.

All figures were drawn using the stereomicroscope

equipped with an eyepiece grid. Photographs of the

respective type specimens were obtained from the

Naturhistorisches Museum in Vienna. SEM photos of

gold-coated shells were taken at the laboratory of the

Institute of Materials Science and Applied Mechanics,

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 103

Fig. 1. Shell measurements: H – shell height, W – shell width,

D – shell diameter, bwH – body whorl height, h – aper

ture height, w – aperture width, U – umbilicus major di

ameter, u – umbilicus minor diameter

Wroc³aw University of Technology, using the micro

scope Jeol JSM–5800 LV.

The members of the Nouvelle École (such as

OCARD 1880, 1881, 1882, 1884–1889, 1888, 1894) de

scribed many species of the genus Trochulus, which

were later synonymised. In cases when references

and/or specimens of nominal species are not avail

able, the synonyms are marked with an asterisk and

cited after G

ERMAIN (1929).

GENERAL CHARACTERISTICS OF THE GENUS

Diagnosis

Shell from conical to flattened; shell diameter

3.5–14.3 mm; whorls 4.5–7.0; lip developed to various

extent; shell surface with radial striae and hairs at

least in juveniles. On internal vaginal walls longitudi

nal plicae; four groups of two mucous glands each, or

glands not grouped.

Shell

Shells of all members of Trochulus are dextral.

Their shape varies. Most of them are depressed coni-

cal, roundish-conical (T. striolatus) or strongly flat-

tened (T. villosus, T. caelatus). Shells of some species

vary widely in shape, for example T. hispidus varies

from much elevated to quite flattened. The variation

is mainly of interpopulation, but to some extent also

individual character.

The number of whorls in adult specimens ranges

from 4.7–5.0 in T. lubomirskii to 7.0 in T. unidentatus.

The individual variation is small: differences within

populations reach 0.5–1 whorl. The number of whorls

appears to show no or only slight differences among

the species. Most interspecific differences are usually

not greater than the interpopulation variation. Con

vexity of whorls varies greatly between species: from

very flat, especially in species with flattened shells, to

pretty convex in some species with conical shells. In

some members of the genus (T. hispidus, T. striolatus)

this character varies individually and between popula

tions.

The umbilicus, depending on the species, is round

or oval, open and wide so that the earlier whorls are

visible, and its major diameter is then up to 0.24 shell

diameter (T. erjaveci), partly covered by the colu

mellar aperture margin, or entirely covered (some

specimens of T. bielzi). In species with open, wide um

bilicus the character is constant, whereas in those with

partly covered umbilicus it varies inividually.

In all species the aperture is elliptical or crescen

tic, its margin usually regularly curved. It is not much

variable. In the majority of species it is equipped with

a white lip, sometimes weakly developed (T. lubo

mirskii, T. villosulus). The lip may be situated right on

the aperture margin (T. luridus, T. leucozonus)or

slightly removed from it (T. hispidus). In T. unidentatus

there is a white tooth on the basal aperture margin,

sometimes poorly visible. The remaining species do

not have any apertural barriers.

The greatest dimension of adult specimens in Tro

chulus (shell width) ranges from 3.5–5.5 mm (the

smallest specimens of T. edentulus) to 7.8–14.3 mm

(the largest specimens of T. erjaveci). The aperture

size ranges from 1.7 × 2.8 mm (T. edentulus)to4.2×

7.2 (big specimens of T. erjaveci). The intrapopulation

variation of shell size is wide.

104 Ma³gorzata Proæków

Figs 2–4. Shell surface sculpture. Undamaged shells of: 2 – T.

luridus,3–T. bakowskii,4–T. villosus

In all species juvenile shells and younger whorls of

adult specimens, except the protoconch, are covered

with hairs (Figs 2–6). Their length, thickness and den

sity vary between species but shows also some individ

ual variation. In some species (T. villosus, T. villosulus)

the hairs are rather permanent, while in others, soon

after forming each shell increment, they are lost

partly (T. bakowskii, T. hispidus) or entirely (T. lubo

mirskii, T. erjaveci), leaving characterisic scars on the

shell surface (Fig. 7).

The shell sculpture between the hairs (Figs 2–6)

consists of periostracal convexities which form a net

work of different density, with most components often

arranged radially. On shells devoid of periostracum

only slightly irregular growth lines are visible (Fig. 8).

Only the shell surface of T. lubomirskii is covered with

regular radial rows of elongated convexities (Fig. 9).

In all the studied species the surface of embryonic

whorls is entirely smooth (Fig. 10).

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 105

Figs 5–7. Shell surface sculpture. Undamaged shells of: 5 – T.

bielzi, 6–T. unidentatus, 7 – scars left by lost hairs on a

shell of T. hispidus devoid of periostracum

Figs 8–10. Shell surface sculpture. Shells of: 8 – T. clande

stinus, 9–T. lubomirskii devoid of periostracum, 10 – sur

face of embryonic shell of T. caelatus

The shell colour varies among species: from very

light, nearly transparent, horny-yellow, through light

brown, to dark brown or reddish-brown. There is also

some individual variation. In a part of individuals of

some species (T. hispidus, T. suberectus, T. striolatus,

T. montanus, T. caelatus, T. clandestinus, T. graminicolus,

T. erjaveci, T. unidentatus, T. edentulus, T. filicinus,

T. leucozonus, T. luridus, T. bakowskii, T. bielzi) a light,

spiral band runs along the body whorl, at approxi

mately half of its height. Shells of albino specimens

are white and almost completely transparent.

Reproductive system

The structure of the reproductive organs follows

the hygromiid pattern and their nomenclature is

shown in Figure 11. The gonad (glandula hermaph

roditica) consists of varied number (6–12) of light

lobes embedded in the digestive gland. Their collect

ing ducts merge to form the hermaphrodite duct

(ductus hermaphroditicus), which in its initial section

is thin and straight, the remaining section, closer to

the albumen gland, being thicker and coiled. Just

next to the albumen gland the hermaphrodite duct

106 Ma³gorzata Proæków

Fig. 11. Reproductive system: ag – albumen gland, ds – dart sacs, ep – epiphallus, fl – flagellum, ga – genital atrium, hd – her

maphroditic duct, hg – hermaphroditic gland, mg – mucous glands, o – oviduct, p – penis, pr – penial retractor, sc –

spermatheca, sd – spermatheca duct, so – spermoviduct, vd – vas deferens, vl – lower vagina, vu – upper vagina

forms a characteristic bend, called the fertilisation

chamber. The albumen gland (glandula albuminalis)

is large and elongate, whitish or cream-coloured. Its

size depends on the phase of reproductive activity: in

male-phase individuals it is considerably smaller than

in individuals producing and lying eggs. The spermo

viduct (spermoviductus) is incompletely divided into

female and male sections.

The free oviduct section (oviductus) is very short

and passes into the vagina, consisting of the upper va

gina (from the outlet of the mucous glands to the tips

of the inner dart sacs) and the lower vagina (from the

base of the dart sacs to the genital atrium). In a great

majority of species there are four pairs of mucous

glands (glandulae mucosae), but some reductions or

secondary branches are also possible, and then the to

tal number may vary from four to ten. The only spe

cies with constantly lower number of the mucous

glands are T. suberectus – six and T. biconicus – four.

The length and position of the glands vary: from very

long (2.5 mm) and situated just above the tips of the

dart sacs (T. edentulus), through equally long, but lo

cated in the upper section of the vagina (T. luridus, T.

leucozonus) to short and situated just above the dart

sacs (T. villosus).

The dart sacs are placed symmetrically in pairs, on

both sides of the vagina. The sacs lying near the va-

gina – internal – may be as long as the outer sacs

(T. hispidus, T. suberectus, T. bakowskii, T. edentulus,

T. luridus), slightly longer (T. villosulus, T. striolatus,

T. montanus, T. bielzi, T. villosus, T. biconicus, T. erjaveci,

T. caelatus, T. graminicolus, T. leucozonus, T. filicinus)or

cosiderably longer, reaching the outlet of the mucous

glands (T. unidentatus). The exception is T. piccardi,in

which no accessory dart sacs are visible. However,

since the main dart sacs are unusually wide, a fusion

of the sacs is possible (P

FENNINGER & PFENNINGER

2005). In all species (no data on T. piccardi) only the

outer dart sacs contain darts, one each. The

spermathecal duct (ductus spermathecae) opens to

the vagina. The spermathecal duct may be thin and

very long, like in T. villosulus, T. bakowskii, T. edentulus,

in which the spermatheca (receptaculum seminis)

reaches the albumen gland, or thick, straight and

short (T. lubomirskii), or else coiled (T. unidentatus),

reaching approximately 1/2 spermoviduct length (T.

hispidus, T. suberectus, T. villosus). The shape of the

spermatheca varies among species: from roundish (T.

striolatus) or oval (T. montanus, T caelatus, T. villosulus)

to irregularly club-like (T. lubomirskii) or hammer-like

(T. unidentatus). The lower vagina (below the dart

sacs) can be long and narrowing toward the genital

atrium (T. bielzi, T. luridus, T. villosus) or long and cy

lindrical (T. edentulus, T. bakowskii). On the internal

walls of the vagina there are four to six longitudinal

folds, depending on the species.

The spermiduct (vas deferens) passes into the

epiphallus which terminates with the flagellum. The

latter is as long as the epiphallus or slightly longer (T.

hispidus). The flagellum is always longer than the

epiphallus in T. bakowskii, T. bielzi, T. leucozonus, T.

filicinus, only slightly longer in T. hispidus,T. suberectus,

but considerably shorter in T. lubomirskii, T.

clandestinus, and in T. unidentatus the flagellum

reaches about 2/3 epiphallus length. Folds are visible

on the cross-section of the epiphallus, their size de

pending on the species. The penis is massive and con

ical (T. bakowskii), cylindrical (T. bielzi, T. villosus, T.

filicinus) or fusiform (T. edentulus, T. villosulus). Inside

the penis, in its section closer to the genital atrium,

there is a papilla with an apically situated pore. On its

cross-section, three or more cavities are visible devel

oped to various extent, depending on the species.

The insertion of the penial retractor muscle

(musculus retractor penis) is situated at the

epiphallus/penis junction. The genital atrium

(atrium genitale) is short.

Ecology and bionomics

A great majority of Trochulus species inhabit humid

and shaded habitats in forests or scrub along streams.

Some are found in montane (T. edentulus, T. villosus,

T. suberectus, T. unidentatus, T. bakowskii, T. villosulus)

or submontane forests (T. bielzi). Even species which

are widely distributed and regarded as euryoecious

(T. hispidus, T. striolatus) prefer not very dry places. All

the species like to spend humid summer days among

or on plants (Urtica dioica, Petasites sp., Caltha palustris,

Cirsium oleraceum), in other weather conditions and

seasons they are found on the ground, in leaf-litter,

under rotting timber, in moss and under stones. The

exceptions are three species. T. biconicus inhabiting

grassy limestone screes, T. graminicolus – known only

from the type locality – found in grassy screes in

sparse forests on steep mountain slopes, with pines,

spruces and junipers, and T. piccardi living in exten

sively used pastures on south-facing slopes. Except T.

clandestinus, which lives in both natural and synan

thropic habitats (gardens, parks and road margins) of

canton Bern (Switzerland), T. striolatus – a synan

thrope of northern France and the British Isles, and

T. hispidus found in anthropogenic sites in its entire

distribution area, the remaining members of the ge

nus are limited to natural or nearly-natural habitats.

The reproductive biology of Trochulus is complete

ly obscure. It is only known that eggs of T. hispidus are

calcified, white and spherical, their diameter is about

1.5 mm. Considering the fact that upon dissection no

eggs were found in the female ducts, probably all species

are strictly oviparous i.e. the eggs are laid immediately

after they have been formed.

Distribution

Disregarding the north African species with their

very doubtful placement in the genus Trochulus (see

Introduction), the majority of species inhabit the Alps

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 107

and the Carpathians (Fig. 12). The only widely distrib

uted species is T. hispidus, occuring in western, central

and eastern Europe, and a part of northern Europe.

The second largest distribution area is that of T.

striolatus which inhabits the British Isles, northern

France, Switzerland and Austria, the Netherlands,

southern Germany and south-western Slovakia. The

remaining species are endemics of limited range.

Thus, four groups may be distiguished: 1. Alpine

group, limited to the whole Alps or their small parts –

9 species (T. suberectus, T. villosus, T. biconicus, T.

caelatus, T. edentulus, T. clandestinus, T. graminicolus,

T. montanus, T. piccardi), 2. Carpathian group, distrib

uted in the Carpathians or their parts – 5 species

(T. villosulus, T. bakowskii, T. bielzi, T. lubomirskii, T.

czarnohoricus), 3. Alpine-Carpathian group – 1 species

(T. unidentatus) and 4. East-Alpine-Dinaric group, lim

ited to the Eastern Alps and/or the Dinaric Mts – 4

species (T. erjaveci, T. filicinus, T. leucozonus, T. luridus).

SYSTEMATIC INDEX OF SPECIES

Genus: Trochulus Chemnitz, 1786

1. Trochulus bakowskii (Poliñski, 1924)

2. Trochulus biconicus (Eder, 1917)

3. Trochulus bielzi (A. E. Bielz, 1860)

4. Trochulus caelatus (Studer, 1820)

5. Trochulus clandestinus (Hartmann, 1821)

108 Ma³gorzata Proæków

Fig. 12. Distribution of Trochulus (diversification centres indicated in dark)

6. Trochulus czarnohoricus (Poliñski, 1924)

7. Trochulus edentulus (Draparnaud, 1805)

8. Trochulus erjaveci (Brusina, 1870)

9. Trochulus filicinus (L. Pfeiffer, 1841)

10.Trochulus graminicolus (Falkner, 1973)

11.Trochulus hispidus (Linnaeus, 1758)

12.Trochulus leucozonus (C. Pfeiffer, 1828)

13.Trochulus lubomirskii (Œlósarski, 1881)

14.Trochulus luridus (C. Pfeiffer, 1828)

15.Trochulus montanus (Studer, 1820)

16.Trochulus piccardi Pfenninger et Pfenninger, 2005

17.Trochulus striolatus

(C. Pfeiffer, 1828)

18.Trochulus suberectus (Clessin, 1878)

19.Trochulus unidentatus (Draparnaud, 1805)

20.Trochulus villosulus (Rossmässler, 1838)

21.Trochulus villosus (Draparnaud, 1805)

22.Trochulus waldemari (Wagner, 1912) nomen dubium

KEY FOR SPECIES IDENTIFICATION

Note! The key serves to determine adults of all the taxa listed above, except T. czarnohoricus with no current

anatomical data and T. waldemari, specimens of which were unavailable.

1. Umbilicus entirely open and wide, previous whorls always visible. Umbilicus major

diameter/shell diameter ratio 0.1–0.3 . . . . . . . . . ........................2

– Umbilicus narrow, sometimes punctured, partly or entirely covered by columellar

aperture margin. Umbilicus major diameter/shell diameter ratio 0.02–0.13 .............13

2. Shell roundish-conical or nearly flattened; no hairs or hairs short and easily lost in adults.

Aperture with white lip inside . . . . . . . . . . . . . ........................3

– Shell flattened, low spire with blunt apex; no hairs or hairs long, curved and rather durable

but not dense. Aperture with white lip, sometimes lip poorly developed ................7

3. Shell width 5–10 mm (not more than 10 mm), with short, thin hairs, easily lost in adults.

Inner and outer dart sacs approximately equal in length .......................4

– Shell width usually more than 10 mm (9–14 mm), no hairs in adults. Inner dart sacs

reach slightly beyond outer ones . . . . . . . . . . . ........................5

4. Four pairs of mucous glands. Shell width 5.5–10 mm, shell height 3–6.7 mm, shape variable:

from roundish-conical to nearly flattened, sometimes with light band body whorl . . . T. hispidus,p.128

– Six mucous glands. Similar to the preceding species. Shell width 5–7.5 mm,

shell height 4–5.6 mm, roundish-conical. . . . . . .................T. suberectus, p. 153

5. Flagellum longer than epiphallus. Shell width 8–14 mm, shell height 4.5–9 mm.

Aperture with distinct lip inside . . . . . . . . . . . . ........................6

– Flagellum shorter than epiphallus. Shell width 9–12.5 mm, shell height 5.5–7 mm.

Body whorl inflated. Aperture with weakly developed lip inside ..........T. clandestinus, p. 118

6. Flagellum slightly longer than epiphallus; epiphallus as long as penis or longer.

Inner dart sacs reach only slightly beyond outer ones, considerably more massive.

Spermatheca roundish . . . . . . . . . . . . . . . . . . . . ............T. striolatus, p. 146

– Flagellum considerably longer than epiphallus; epiphallus shorter than penis.

Inner dart sacs reach far beyond outer ones. Spermatheca oval ...........T. montanus, p. 143

7. Shell height up to 6 mm, shell width up to 10 mm. Umbilicus diameter 0.8–3 mm. . . ........8

– Shell height more than 6 mm (6–8.5 mm), shell width more than 10 mm (10–14.7 mm).

Umbilicus diameter 1.9–3.5 mm . . . . . . . . . . . ........................12

8. Shell covered with long and sparse hairs in both juveniles and adults ........T. villosulus,p.162

– Shell devoid of hairs in adults . . . . . . . . . . . . . ........................9

9. Shell width up to 9.5 mm, shell height up to 5 mm. Mucous glands inserting directly

above tips of inner dart sacs .....................................10

– Shell width more than 9.5 mm, shell height usually more than 5 mm.

Outlet of mucous glands removed from tips of inner dart sacs ..........T. graminicolus, p. 127

10. Aperture narrow, crescentic. Inner and outer dart sacs distinctly visible.

Shell flattened or nearly disc-like with low spire . . . ........................11

– Aperture elliptical. No inner dart sacs visible. Shell depressed

but with elevated apex . . . . . ...........................T. piccardi, p. 146

11. Shell small: width up to 7 mm, height up to 3.5 mm. Whorls tightly coiled.Four long

mucous glands (ca. 2.5 mm) . . . . . . . . . . . . . . . ..............T. biconicus, p. 112

– Shell larger: width more than 7 mm, height more than 3.5 mm. Whorls moderately

coiled. Eight short mucous glands grouped in pairs ..................T. caelatus,p.116

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 109

12. Shell covered with long, up to 1 mm, hairs in both juveniles and adults. Lip poorly

developed. No light band on body whorl . . . . . . . ................T. villosus, p. 164

– Shell devoid of hairs in adults. Lip distinct. Sometimes light band on body whorl . . . T. erjaveci, p. 124

13. Aperture simple . . . . . . . . . . . . . . . . . . . ........................14

– Aperture with prominent white tooth on basal margin. Inner dart sacs up to 2× longer

than outer ones . . . . . . . . . . . . . . . . . . . . . . ............T. unidentatus, p. 155

14. Shell translucent, whitish-yellow to greenish-yellow. Aperture rounded with very weakly developed lip.

Flagellum conical, very short, considerably shorter than epiphallus .........T. lubomirskii, p. 140

– Shell horny-brown to reddish-brown, often with light band on body whorl. Aperture

with distinct lip inside. Flagellum and epiphallus approximately equal in length ...........15

15. Dart sacs situated just below outlet of mucous glands. Vagina long and cylindrical.

Shell small: width 5–8.5 mm, height 2.8–5.5 mm; roundish-conical with tightly coiled whorls .....16

– Outlet of mucous glands and tips of inner dart sacs ca. 1.5–4 mm apart. Vagina

expands at dart sacs region and narrows toward genital atrium. Shell larger:

width 7.5–12.6 mm, height 4.5–8.3 mm; roundish with moderately tightly coiled whorls .......17

16. Flagellum 1.5–2 × longer than epiphallus which is shorter than conical penis. Shell

roundish-conical. Hairs fine, easily lost in adults, scars hardly visible .........T. bakowskii, p. 110

– Flagellum slightly shorter than epiphallus or approximately as long; epiphallus as long as fusiform

penis or slightly longer. Shell very similar to T. bakowskii, shape variable from conical to flat. Juveniles

with dense hairs, often lost in adults, then shell slightly shiny and finely striated . . . T. edentulus, p. 121

17. Flagellum 1.5–2 × as long as epiphallus; penis cylindrical .......................18

– Flagellum only slightly longer than epiphallus; penis fusiform ....................19

18. Spermatheca does not reach albumen gland. Umbilicus very narrow, most often

entirely covered by columellar aperture margin. Hairs short, curved and easily lost,

leaving pronounced scars . . . . . . . . . . . . . . ..................T. bielzi, p. 114

– Spermatheca reaches albumen gland. Umbilicus very narrow, sometimes partly or nearly

entirely covered by columellar aperture margin; when open, most often round.

Both juveniles and adults with short and dense hairs .................T. filicinus, p. 125

19. Shell devoid of hairs in adults. Flagellum considerably longer than epiphallus.

Inner dart sacs slightly longer and narrower than outer ones ............T. leucozonus, p. 138

– Shell with short, fine and sparse hairs. Flagellum as long as epiphallus or slightly longer.

Inner and outer dart sacs approximately equal in size .................T. luridus, p. 141

SYSTEMATIC REVIEW

Trochulus bakowskii (Poliñski, 1924)

Fruticicola bielzi b¹kowskii POLIÑSKI 1924: 196, pl.

13, fig. 72–74. Locus typicus: Ukraine [formerly Po

land]: upper Prut valley: Dora and Tatarów; and East

Carpathians: E slopes of Czarnohora. Syntypes: Mu

seum of Natural History, Kraków.

Material examined

Poland: Bieszczady Mts: Bere¿ki near Ustrzyki

Górne, 10.08.1972, leg. Studenckie Ko³o Biologów

Wroc³aw, 1788, MPW, 5 alc.; Terebowiec Valley,

6.08.1972, leg. Studenckie Ko³o Biologów Wroc³aw,

1787, MPW, 32 alc.; Bystry Stream Valley, 6.08.1972

leg. Studenckie Ko³o Biologów Wroc³aw, 1876, MPW,

19 alc.; Ustrzyki Górne, 21.08.1996, ZW, 5 s.+38 alc.;

Cisna, 24.08.1996, ZW, 28 alc.; Wetlina, 25.08.1996,

ZW, 24 alc.; Moczarne, 25.08.1996, ZW, 49 alc.; s.

prec. loc., Bieszczady, 10.08.1972, leg. Studenckie

Ko³o Biologów Wroc³aw, 1790, MPW, 40 alc.

Shell (Figs 13–15)

Shell roundish-conical with 5.1–6.6 regularly and

slowly increasing whorls. Shell height 2.8–5.1 mm,

shell width 5.2–7.3 mm, height/width ratio 0.57–0.76,

body whorl height 2.5–3.6 mm, relative height of

body whorl 0.63–0.81, aperture height 1.6–2.6 mm,

aperture width 2.6–3.7 mm, umbilicus major diam

eter 0.2–0.7 mm, umbilicus minor diameter 0.2–0.6

mm, umbilicus major diameter/shell diameter ratio

0.03–0.1. Aperture transversely crescentic (wider than

high) with distinct white lip inside. Umbilicus narrow,

punctured, always open and most often round. Hairs

fine, easily lost in adults, scars hardly visible. Shell

horny-brown to reddish-brown, shiny with light band

on body whorl.

Reproductive system (Figs 16–20)

Most often four pairs of rather long mucous

glands; sometimes glands reduced to three pairs, in

other specimens with secondary branches. Inner dart

sacs as large as outer ones, well separated. Vagina very

110 Ma³gorzata Proæków

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 111

Figs 13–20. T. bakowskii. Specimen from Moczarne, Bieszczady, Poland, ZW. 13–15 – shell: 13 – apertural view, 14 – umbilical

view, 15 – apical view; 16–20 – reproductive system: 16 – general view, 17 – longitudinal section of vagina, 18 – penial pa

pilla, 19 – cross-section of penial papilla, 20 – cross-section of epiphallus. Scale bar 5 mm

long (ca. 5 mm) and cylindrical. Flagellum 1.5–2×

longer than epiphallus which is shorter than conical

penis. Spermatheca duct long and bent. Spermatheca

oval, reaching albumen gland. Longitudinal section

of vagina, penial papilla, cross-section of penial pa

pilla and epiphallus shown in Figs 17–20.

Ecology

T. bakowskii lives in very damp places in the herb

layer of montane forests between 700 and 1,500 m

a.s.l.; it is found in tussocks of vegetation (Petasites sp.,

Caltha palustris, Cirsium oleraceum) or leaf-litter. It pre

fers beech-fir forests (Fageto-Abietum) with Alnus

incana and Petasites albus; in spruce forests (Piceetum

altiherbosum) it is found in tussucks of Adenostyle sp.,

Doronicum sp. and Mulgedium sp. (H

UDEC 1965).

Distribution (Fig. 21)

A North-East-Carpathian species; it occurs in

Slovakia (Muransky-Kras, Pol’ana, eastern part of the

Low Tatra Mts) (H

UDEC 1965) Poland (Bieszczady

and an isolated locality on the Ropa River near Uœcie

Gorlickie) (R

IEDEL 1988) Ukraine (Svidovec;

Gorgany; in the east to Czarnohora) (P

OLIÑSKI 1924,

ERNEY et al. 1983) and Hungary (FALKNER et al.

2001).

Remarks

According to S

HILEYKO (1978b), there are 6–8 not

grouped mucous glands. All the adult specimens ana-

tomically examined had eight or six mucous glands,

always grouped in pairs.

UDEC (1965) noted that the flagellum was ap-

proximately as long as the epiphallus but most often

shorter, which was not confirmed by my studies. In all

the dissected adults the flagellum was considerably

longer than the epiphallus (see also P

OLIÑSKI 1928

and S

HILEYKO 1978b). Moreover, in the figured speci

men (H

UDEC 1965) the outlet of the mucous glands

was situated at a distance from the upper tips of the

inner dart sacs, while in the examined specimens they

were at the same level.

Trochulus biconicus (Eder, 1917)

Fruticicola biconica EDER 1917: 119, figs 1–4. Locus

typicus: Switzerland: Canton Nidwalden: Bannalpass,

steep SW-facing screes slope, 2,000 m a.s.l.

Material examined

Switzerland: Bannalppass, 2,100–2,200 m,

3.09.1967, coll. M. Wüthrich, Nr 1918, NHMB, 6 s.+2

alc.; Bannalppass, 2,150 m, leg. Böckel, coll. Klemm

50496, NHMW, 5 s.

Shell (Figs 22–24)

Shell flattened, with low spire. Whorls 5–6.5 tightly

coiled, slowly but regularly increasing, body whorl

only ca. twice wider than the first one. Shell height

2.5–3.5 mm, shell width 5.0–7.0 mm, height/width ra-

tio 0.48–0.57, body whorl height 2.5–2.8 mm, relative

height of body whorl 0.78–0.9, aperture height

1.7–2.2 mm, aperture width 2.5–3.1 mm, umbilicus

major diameter 0.9–1.3 mm, umbilicus minor diame-

ter 0.8–1.1 mm, umbilicus major diameter/shell di-

ameter ratio 0.16–0.19. Aperture narrow, crescentic,

with whitish lip. Basal and palatal aperture margins

slightly reflexed. Umbilicus deep and always open.

Adults hairless. Shell pale brown with irregular

growth lines distinct above, much weaker below.

Reproductive system (Figs 25–29)

Four long mucous glands. Inner dart sacs slightly

longer than outer ones. Vagina long and cylindrical.

112 Ma³gorzata Proæków

Fig. 21. Distribution of T. bakowskii. Black triangles – localities of origin of the examined material

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 113

Figs 22–29. T. biconicus. Specimen from Bannalppass (type locality), Switzerland, coll. M. Wüthrich, Nr 1918, NHMB. 22–24 –

shell: 22 – apertural view, 23 – umbilical view, 24 – apical view; 25–29 – reproductive system: 25 – general view, 26 – longitudi

nal section of vagina, 27 – penial papilla, 28 – cross-section of penial papilla, 29 – cross-section of epiphallus. Scale bar 5 mm

Flagellum longer than epiphallus, which is longer

than fusiform penis. Spermatheca duct straight.

Spermatheca oval, not reaching albumen gland. Lon

gitudinal section of vagina, penial papilla, cross-

section of penial papilla and epiphallus shown in Figs

26–29.

Ecology

T. biconicus lives in grassy limestone screes, on

mountain slopes with rock outcrops and sparse vege

tation, ca. 2,000 m a.s.l. and higher.

Distribution (Fig. 30)

A microendemic species; earlier known only from

its type locality in Bannalppass (canton Nidwalden) in

Switzerland. However, since 2005 it has been regularly

found in other localities: Isenthal (canton Uri);

Kerns, Engelberg, Ruchstock (canton Obwalden);

Wolfenschiessen, Beckenried (canton Nidwalden)

AGGENSTOS 2006).

Trochulus bielzi (A. E. Bielz, 1860)

Helix bielzi A. E. BIELZ 1859: 216. [nomen nudum]

[see also F

ALKNER 1995: 100]

Helix bielzi A. E. B

IELZ 1860: 151. Terra typica: Ro

mania: Transylvania.

Helix euconus W

ESTERLUND 1890: 121. Locus typi

cus: Poland: near Przemyœl

Fruticicola bielzi euconus var. globulina POLIÑSKI

1928: 187, pl. XXVIII, fig. 38. Locus typicus: Ukraine:

south of Ko³omyja: “Czarny Las” near Wierbi¹¿.

Material examined

Poland: Bieszczady Mts: Wetlina, Wierch

Muchanin, 9.07.1962, leg. C. Dziadosz et W. Starêga,

MIZW, 4 s.; Cisna, 24.08.1996, ZW, 16 s.+6 alc.;

Wetlina, 25.08.1996, ZW, 7 alc.; Dwernik, 26.08.1996,

ZW, 1 s.+3 alc.; Dwerniczek, 26.08.1996, ZW, 1 s.; s.

prec. loc., Bieszczady, 10.08.1972, leg. Studenckie

Ko³o Biologów Wroc³aw, Nr 1789, MPW, 14 alc.; Ro

mania: Transylvania: Attelsloch near Schässburg

[=Schäßburg=Sighiºoara], 1918, coll. Poliñski

228/37, MIZW, 9 s.; Mediasch [=Mediaº], coll.

Poliñski 228/37, MIZW, 2 s.

Shell (Figs 31–33)

Shell roundish-conical with 5.5–6.25 regularly in

creasing convex whorls. Shell height 5.2–7.3 mm,

shell width 7.5–10.1 mm, height/width ratio

0.61–0.78, body whorl height 4.0–5.1 mm, relative

height of body whorl 0.64–0.81, aperture height

2.7–3.6 mm, aperture width 4.1–5.3 mm, umbilicus

major and minor diameter 0.2–0.4 mm, umbilicus

major diameter/shell diameter ratio 0.02–0.13. Aper

ture transversely oval, with white lip inside. Umbilicus

very narrow, most often entirely covered by

columellar aperture margin. Hairs short, curved and

easily lost leaving pronounced scars. Shell horny-

yellow to reddish-brown, matt, often with light band

on body whorl.

Reproductive system (Figs 34–38)

Four pairs of short mucous glands situated around

upper vagina, ca. 1.5 mm from tips of inner dart sacs,

which are slightly longer than outer ones. Vagina long

(ca. 6.0 mm); its expanded dart sac region narrows to-

ward genital atrium. Flagellum up to 1.5× longer than

epiphallus, which is as long as cylindrical penis or

slightly shorter. Spermatheca duct thick and long.

Spermatheca oval, not reaching albumen gland. Lon-

gitudinal section of vagina, penial papilla, cross-section

of penial papilla and epiphallus shown in Figs 35–38.

Ecology

T. bielzi lives in damp herbal layer of forests; it shel

ters in leaf litter and rotting timber, but also often stays

on the underside of leaves of e.g. Petasites sp., Caltha

palustris, Cirsium oleraceum. It is found in the foothills

and lower mountain altitudes, up to 600–700 m.

Distribution (Fig. 39)

An Eastern-Carpathian species; known from

Slovakia (Carpathians), Poland (Bieszczady, eastern

Beskid Niski Mts, Pogórze Dynowskie foothills; in the

west to the Wis³ok River) (R

IEDEL 1988), Romania

(Transylvania) (K

ERNEY et al. 1983) and Ukraine

(Carpathians: Czeremosz-Tal) (P

OLIÑSKI 1924, 1928).

Remarks

Populations inhabiting the northern part of the

range, including Poland, were identified as a subspe

cies T. bielzi euconus (Westerlund

, 1890). Moreover,

among the specimens of euconus P

OLIÑSKI (1928) dis

tinguished smaller individuals with a very weakly de

veloped lip, which he designated as Fruticicola bielzi

euconus var. globulina. Besides the detailed shell de

scription, he gave figures and descriptions of the re

productive system, but without specifying any differ

ences between the two taxa.

114 Ma³gorzata Proæków

Fig. 30. Distribution of T. biconicus. Black triangle – locality

of origin of the examined material

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 115

Figs 31–38. T. bielzi. Specimen from Cisna, Bieszczady, Poland, ZW. 31–33 – shell: 31 – apertural view, 32 – umbilical view, 33 –

apical view; 34– 38 – reproductive system: 34 – general view, 35 – longitudinal section of vagina, 36 – penial papilla, 37 –

cross-section of penial papilla, 38 – cross-section of epiphallus. Scale bar 5 mm

Comparison of shells of T. bielzi (HUDEC 1972)

from the Slovak Carpathians and from various Roma-

nian populations showed that specimens correspond-

ing to the description of euconus and its forms, also oc

curred among Romanian populations. The variation

is probably of individual and to some degree

interpopulation characters and does not justify the

taxonomic distinctions.

UDEC (1972) remarked that juvenile T. bielzi had

the flagellum approximately as long as the epiphallus.

However, in his figure of the reproductive system, the

flagellum seems to be longer than the epiphallus

which may be caused by the fact that the figured

flagellum is twisted, or the figure shows an adult speci

men (which is not specified). In this study, in all the

anatomically examined specimens, the flagellum was

up to 1.5x longer than the epiphallus.

Trochulus caelatus (Studer, 1820)

Helix caelata Vallot, 1801 [nomen oblitum – see

ALKNER 1995: 99, TURNER et al. 1998: 343]

Glischrus (Helix) caelata S

TUDER 1820: 86 [12].

Terra typica: Switzerland: Jura, in forests, at humid

rocks. Lectotype: NHMB.

Helix corrugata Var. γ – coelata H

ARTMANN 1821:

236, pl II, fig. 12. Terra typica: Switzerland: mountains

in Bernese Oberland; Jura.

Helix glypta L

OCARD 1880–1881: 95. Locus typicus:

France: dep. Rhône-Alpes: environs of Lyon; dep.

Ain: Laumusse and Hauleville [according to F

ALKNER

(1995: 99) this name is also nomenclatorically cor

rect].

Helix coelomphala L

OCARD 1888: 352. Locus typicus:

Switzerland: Mt. Righi, Zurich, Lucerne and Jura

neuchâtelois; Germany: Dillingen near Saarlouis,

Günzburg, Dinkelsbühl; E France: dep. Savoie: envi

rons of Annecy, Chambéry, Aix-les-Bains, Mouxy,

Albertville; dep. Isère: vicinity of Genoble, Sassenage,

Saint-Martin-le-Vinoux, Allevard-les-Bains,

Grande-Chartreuse, Sappey, Voreppe; dep.

Hautes-Alpes: Barcelonnette; dep. Rhône: environs of

Lyon, alluvions of Rhône N Lyon, Brotteaux,

Sathonay, Pape; dep. Ain: Laumusse, Bugey, alluvions

of Suran; dep. Jura: Bief-du-Fourg, Saint-Claude,

Poligny; dep. Côte-d’Or: Châtillon-sur-Seine; envi

rons of Paris; Bionville near Metz; dep. Finistère:

Brest.* [see F

ALKNER 1990: 202, FALKNER et al. 2001:

57, F

ALKNER &FALKNER 2008 – Trochulus coelompha

la (Locard, 1888)]

Helix coelata C

LESSIN 1874: 187, pl. VIII, fig. 5.

Material examined

Switzerland: Mt. Wandfluh, Bettlach, Jura, 1,280

m, 11.06.1961, coll. M. Wüthrich Nr 2330, NHMB, 6

s.+2 alc.; Gorges de Moutier valley, Jura, 506 m,

27.06.1969, coll. M. Wüthrich Nr 228, NHMB, 6 s.+2

alc.; Gänsbrunnen, cant. Solothurn, MIZW, 5 s.

Shell (Figs 40–42)

Shell strongly flattened, nearly discoidal, convex

underneath. Whorls flat, 5.0–5.7. Shell height 3.7–5.0

mm, shell width 8.0–9.5 mm, height/width ratio

0.47–0.59, body whorl height 3.3–4.2 mm, relative

height of body whorl 0.84–0.95, aperture height

2.3–3.0 mm, aperture width 3.5–4.6 mm, umbilicus

major diameter 1.5–2.4 mm, umbilicus minor diam

eter 1.2–1.9 mm, umbilicus major diameter/shell di

ameter ratio 0.17–0.2. Aperture with white lip. Umbili

cus deep, open and wide. Hairs present only in very

young juveniles. Shell reddish-brown, shiny, promi

nently striated, sometimes with light band on body

whorl.

Reproductive system (Figs 43–47)

Four pairs of short mucous glands. Inner dart sacs

narrower but slightly longer than outer ones. Vagina

long and cylindrical. Flagellum longer than

epiphallus which is shorter or as long as fusiform pe

nis. Spermatheca duct short and straight.

Spermatheca oval, reaching ca. 2/3 spermoviduct

length. Longitudinal section of vagina, penial papilla,

cross-section of penial papilla and epiphallus shown

in Figs 44–47.

116 Ma³gorzata Proæków

Fig. 39. Distribution of T. bielzi. Black triangles – localities of

origin of the examined material

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 117

Figs 40–47. T. caelatus. Specimen from Bettlach, Wandfluh, Switzerland, coll. M. Wüthrich, Nr 2330, NHMB. 40–42 – shell: 40

– apertural view, 41 – umbilical view, 42 – apical view; 43–47 – reproductive system: 43 – general view, 44 – longitudinal sec

tion of vagina, 45 – penial papilla, 46 – cross-section of penial papilla, 47 – cross-section of epiphallus. Scale bar 5 mm

Ecology

T. caelatus lives in shaded limestone habitats, be

tween 500 and 1,300 m a.s.l.

Distribution (Fig. 48)

An endemic species; known only from Switzerland:

Jura, the middle Birs valley (T

URNER et al. 1998).

Remark

The reproductive system is very similar to those of

T. striolatus and T. montanus. It differs in its very short

dart sacs.

Trochulus clandestinus (Hartmann, 1821)

Helix corrugata Var. α H. clandestina HARTMANN

1821: 236. Locus typicus: Austria: Vienna and Switzer

land: near Zürich.

Helix Putonii C

LESSIN 1874: 314. Terra typica: Bel

gium and NE France: Vosges Mountains. [see

ALKNER et al. 2001: 58 – Trochulus clandestinus putonii

(Clessin, 1874)]

Material examined

Switzerland: Culm, cant. Aargau, 569a 44, MPW, 2

s.; Boll-Sinneringen, cant. Bern, 24.07.1977, coll. M.

Wüthrich Nr 3842/50, NHMB, 6 s.+2 alc.; Wattenwil,

Bellevue, cant. Bern, 27.05.1967, coll. M. Wüthrich Nr

3117/47, NHMB, 6 s.+2 alc.; Schleitheim, cant.

Schaffhausen, coll. Sterki 31800, NHMW, 5 s.

Shell (Figs 49–51)

Shell roundish-conical with 5.3–6.0 whorls. Shell

height 5.6–7.0 mm, shell width 9.3–12.3 mm,

height/width ratio 0.55–0.78, body whorl height

4.5–6.0 mm, relative height of body whorl 0.79–0.88,

aperture height 3.4–4.5 mm, aperture width 4.3–6.1

mm, umbilicus major diameter 1.3–2.2 mm, umbili

cus minor diameter 1.0–1.7 mm, umbilicus major di

ameter/shell diameter ratio 0.15–0.22. Aperture with

weakly developed lip. Umbilicus wide and open.

Adults hairless. Shell light horny-brown to yellow

ish-grey, shiny, body whorl inflated, sometimes with

light band.

Reproductive system (Figs 52–56)

Four pairs of short mucous glands situated around

upper vagina, ca. 1 mm from tips of inner dart sacs,

which are slightly longer than more massive outer

ones. Vagina long and cylindrical. Flagellum consider

ably shorter than epiphallus, which is longer than

conical penis. Spermatheca duct thin and straight.

Spermatheca oval, not reaching albumen gland. Lon

gitudinal section of vagina, penial papilla, cross-sec

tion of penial papilla and epiphallus, and shown in

Figs 53–56.

Ecology

T. clandestinus is a montane species, inhabiting the

herb layer in forests, scrubstreams, but is also found

in cultivated land (as a synanthrope in parks, gardens,

verges and in ruderal vegetation); between 400 and

1,000 m a.s.l., maximum up to 2,300 m a.s.l.

Distribution (Fig. 57)

Known from Switzerland (middle and northern

Jura; the northern Alps; canton Bern; Vierwaldstätter

See to the Thur valley), Liechtenstein and southern

Germany (Hegau) (F

ORCART 1965, KERNEY et al.

1983, T

URNER et al. 1998).

Remarks

The species is often confused with T. striolatus and

T. montanus, but it differs from them in more rapidly

increasing whorls, resulting in shells of the same size

having fewer whorls. Likewise, the very short

flagellum is distinctive (F

ORCART 1965).

118 Ma³gorzata Proæków

Fig. 48. Distribution of T. caelatus. Black triangles – localities

of origin of the examined material

Fig. 57. Distribution of T. clandestinus. Black triangles – local

ities of origin of the examined material

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 119

Figs 49–56. T. clandestinus. Specimen from Boll-Sinneringen, Switzerland, coll. M. Wüthrich Nr 3842/50, NHMB. 49–51 –

shell: 49 – apertural view, 50 – apical view, 51 – umbilical view; 52–56 – reproductive system: 52 – general view, 53 –

longitudinal section of vagina, 54 – penial papilla, 55 – cross-section of penial papilla, 56 – cross-section of epiphallus.

Scale bar 5 mm

Trochulus czarnohoricus (Poliñski, 1924)

Fruticicola czarnohorica POLIÑSKI 1924: 193–195,

figs 75–77. Locus typicus: Ukraine: district Nadwórna:

south of Worochta and west of Ardzheludza: in the

valley between Kiczera and Kukul Ridges (“Kiczera

-Kukul-Rücken”).

Material examined

Ukraine: valley between Kiczera and Kukul Ridges,

12.04.1921, leg. Sz. Tenenbaum, MIZW, 16 s. [1

lectotype and 15 paralectotypes]

Shell (Figs 58–60)

Shell small, roundish with 4.5–4.75 whorls. Shell

height 4.1–4.5 mm, shell width 4.9–5.6 mm,

height/width ratio 0.73–0.89, body whorl height

3.0–3.2 mm, relative height of body whorl 0.66–0.78,

aperture height 2.0–2.2 mm, aperture width 2.7–2.9

mm, umbilicus diameter 0.8–0.9 mm, umbilicus ma

jor diameter/shell diameter ratio 0.16–0.18. Aperture

slightly broader than high, with very weakly devel

oped lip. Body whorl approximately 1.5× wider than

penultimate and not descending. Umbilicus narrow,

partly covered by columellar aperture margin. Juven-

iles with fine hairs, which in adults are lost, leaving

scars. Shell horny-brown, semi-translucent.

Ecology

The dry shells were found on a stream bank in a fir

forest at ca. 1,300 m a.s.l. (P

OLIÑSKI 1924).

Distribution (Fig. 61)

A species known only from the type locality in the

Ukrainian Eastern Carpathians.

Remarks

The species differs from its congeners in its smaller

size, roundish shell (the height/width ratio is the

highest in the genus: 0.73–0.89) as well as the rela

tively low body whorl (relative height 0.66–0.78), and

consequently, rather high spire. Further studies are

required to decide about its taxonomic status.

120 Ma³gorzata Proæków

Figs 58–60. T. czarnohoricus. Shell. Lectotype: 58 – apertural

view, 59 – umbilical view, 60 – apical view. Scale bar 5 mm

Fig. 61. Distribution of T. czarnohoricus. Black triangles – lo

calities of origin of the examined material

Trochulus edentulus (Draparnaud, 1805)

Helix edentula DRAPARNAUD 1805: 80, pl. VII, fig.

14. Terra typica: France: Bresse. Type specimen:

NHMW.

Helix depilata D

RAPARNAUD 1801: 72. Locus typicus:

France: environs of Lyon: Mont Pilat. [non Helix

depilata C. Pfeiffer] [see also F

ALKNER et al. 2002: 151]

Helix limnifera H

ELD 1836: 273. Terra typica: Ger

many: Bavaria. Locus typicus restrictus (according to

ALKNER 1985): Berchtesgadener Alpen:

Watzmann-Massiv.

Helix Cobresiana var. d?L.P

FEIFFER 1846: 190, pl.

99. figs 19–21. Locus typicus: Austria: Styria: Mürztal.

Helix unidentata var. subleucozona W

ESTERLUND

1889: 32, no 89. Locus typicus: Austria: Styria:

Mürztal.

Helix Lorteti L

OCARD 1894: 104. Terra typica:

France: mountainous regions of Dauphiné and

Savoie.*

Fruticicola edentula helvetica P

OLIÑSKI 1928: 195, pl.

XXVIII, figs 42–43. Locus typicus: Switzerland:

Füringen, Frutt, Zürich; Germany: Schwarzwald:

Schramberg; Baden-Württemberg: Aichtal valley near

Nürtingen.

Fruticicola edentula helvetica var. (natio) suevica

OLIÑSKI 1928: 199 (shell), 152 (genitalia). Locus

typicus: Germany: Schwäbische Alb: Seeburger Tal

near Urach.

Material examined

Austria: Golling an der Salzach near Salzburg,

6.11.1997, leg. A. Riedel et F. Seidl 10/97, MIZW, 31

s.+11 alc.; Tiefbrunnau, Salzkammergut, 4.07.1998,

ZW, 6 alc.; Vordersee, Salzkammergut, 4.07.1998, ZW,

2 alc.; Ostermiething, Upper Austria, 04.1926, coll.

Poliñski 228/37, MIZW, 4 s.; Rettengraben gorge, Mt.

Floning, Styria, ex coll. Wagner, MIZW, 13 s.

[subleucozona]; Thörlgraben gorge near Kapfenberg,

Styria, ex coll. Wagner, MIZW, 20 s. [subleucozona];

Schafferweg near Admont, Styria, ex coll. Wagner,

MIZW, 2 s.; Röthelstein castle near Admont, Styria, ex

coll. Wagner, MIZW, 1 s.; s. prec. loc., Carynthia, ex

coll. Gallenstein, MIZW, 1 s.; Germany: Urach,

Württemberg, coll. Poliñski 228/37, MIZW, 5 s.;

Seeburger Tal valley near Urach, Württemberg,

28.04.1928, leg. D. Geyer, MIZW, 4 s. [suevica];

Schamberg, Schwarzwald, Württemberg, coll. Poliñski

228/37, MIZW, 7 s.; Aichtal valley near Nürtingen, S

Stuttgart, ± 300 m, coll. Poliñski 228/37, MIZW, 6 s.;

Moosburg, Bavaria, 30.06.1998, ZW, 2 s.; Wim

bachklamm gorge near Reichenhall, Bavaria, coll.

Poliñski 228/37, MIZW, 6 s. [subleucozona]; Switzer

land: Aiguebelle near Veyrier, S Geneva, 450 m, 04.

1929, coll. Poliñski 228/37, MIZW, 13 s.; Pomier, Mt.

Salève, S Geneva, ± 900 m, 04.1929, leg J. Favre,

MIZW, 4 s.; Vaux forest, between Chasseron and

Fleurier, Jura Vaudois, ± 1,200 m, 09.1928, coll.

Poliñski 228/37, MIZW, 4 s.; Zürich, coll. Poliñski

228/37, MIZW, 3 s.; Frutt, cant. Obwalden, ± 1,700 m,

coll. Poliñski 228/37, MIZW, 6 s. [helvetica]; s. prec.

loc., Swiss Jura, 1928, coll. Poliñski 228/37, MIZW, 1 s.

Shell (Figs 62–67)

Shape variable, from conical to flat, with 5.25–7.0

tightly coiled whorls, body whorl angled at periphery.

Shell height 3.5–5.3, shell width 5.5–8.2,

height/width ratio 0.57–0.74, body whorl height

2.7–3.6 mm, relative height of body whorl 0.57–0.71,

aperture height 1.7–2.5 mm, aperture width 2.8–4.4

mm, umbilicus major diameter 0.2–0.6 mm, umbili

cus minor diameter 0.2–0.6 mm, umbilicus major di

ameter/shell diameter ratio 0.04–0.1. Aperture with

white lip inside, its basal wall slightly thickened but

without teeth; aperture margin slightly reflected. Um

bilicus very narrow and often round, sometimes partly

or entirely covered by columellar aperture margin. Ju

veniles with dense hairs often lost in adults, then shell

slightly shiny and finely striated. Shell horny to red

dish-brown with light band on body whorl.

Reproductive system (Figs 68–72)

Four pairs of rather long mucous glands. Both in-

ner and outer dart sacs approximately equal in size,

situated around upper vagina just below outlet of mu-

cous glands. Vagina relatively long and cylindrical.

Flagellum usually slightly shorter than epiphallus or

approximately equal in length. Epiphallus as long as

fusiform penis or somewhat longer. Spermatheca

duct long. Spermatheca large and elongate, reaching

albumen gland. Longitudinal section of vagina,

penial papilla, cross-section of penial papilla and

epiphallus shown in Figs 69–72.

Ecology

T. edentulus lives among herbaceous vegetation, in

leaf litter and under stones in damp montane forests;

between 300 and 2,000 m a.s.l. (usually not found

above the timberline).

Distribution (Fig. 73)

A West-Alpine species; known from France (Jura;

southern Vosges; in the west to the Rhone River and

the upper Loire River), Switzerland (Jura; from

Genfersee to Bodensee; Alpenrheintal; northern part

of reg. Waadt; Berner, Vierwaldstätter and Glarner

Alps; western Walliser Alps; Nordbünden), Germany

(pre-Alpine region in southern Bavaria: along the

Salzach and Inn Rivers; Bavarian Alps; Bavarian For

est) and Austria (widely distributed except the east

ern part) (P

OLIÑSKI 1928, KERNEY et al. 1983, TURNER

et al. 1998).

Remarks

Some populations from Switzerland and Germany

were described as Fruticicola edentula helvetica Poliñski,

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 121

122 Ma³gorzata Proæków

Figs 62–64. T. edentulus. Specimen from Golling, Salzburg,

Austria, MIZW. Shell: 62 – apertural view, 63 – umbilical

view, 64 – apical view. Scale bar 5 mm

Figs 65–67. T. edentulus. Specimen from Draparnaud's col

lection, NHMW. Shell: 65 – apertural view, 66 – umbilical

view, 67 – apical view. Scale bar 5 mm

1928. The subspecies differs from edentulus s. str. in its

smaller size and narrower umbilicus, sometimes en

tirely covered. The upper and underside of the shell

are similarly convex thus slightly resembling T.

biconicus. Another form, described from Schwäbische

Alb by P

OLIÑSKI (1928) – Fruticicola edentula helvetica

var. (natio) suevica – was distinguished from the other

froms of edentulus by its higher spire and much nar

rower aperture. Its reproductive system did not gener

ally differ from the nominate species (P

OLIÑSKI 1928:

152). The shell material from the respective regions

at my disposal was scanty and no alcohol-preserved

specimens were available which made it impossible to

solve the taxonomy of these entities.

The name Fruticicola edentula helvetica var. (natio)

suevica Poliñski, 1928 is infrasubspecific (Art. 45.6.4.1

ICZN), but was made available by J

AECKEL jun. (1962:

182).

According to F

ALKNER (1985, 1990) two main-

forms can be distiguished i.e. the western shells more

conical, flattened on underside, with a very narrow, al

most covered umbilicus and hairs present only in ju

veniles (including T. edentula s. str., T. edentula helvetica

and T. edentula suevica); the eastern shells are more

flattened, rounded underneath, with umbilicus open

or half open, and stronger and more permanent hairs

(including T. edentula limnifera and T. edentula

subleucozona). He also pointed out the possible transi

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 123

Figs 68–72. T. edentulus. Specimen from Golling, Salzburg, Austria, MIZW. Reproductive system: 68 – general view, 69 – longi

tudinal section of vagina, 70 – penial papilla, 71 – cross-section of penial papilla, 72 – cross-section of epiphallus. Scale bar

5mm

Fig. 73. Distribution of T. edentulus. Black triangles – local

ities of origin of the examined material

tion zone between limnifera and subleucozona, running

along the Salzach River and the lower Inn River. The

subspecies show no differences in the structure of

their genitalia.

Trochulus erjaveci (Brusina, 1870)

Helix (Fruticicola) Erjaveci BRUSINA 1870: 26. Locus

typicus: Croatia: reg. Varazdinska: Vidovec; Agram

[=Zagreb]; reg. Karlovaèka: Ozalj and Slunj; reg.

Lièko-senjska: Plitvice and Oštaria.

Helix Hirci C

LESSIN 1883: 198. Locus typicus:

Croatia: Schneeberg [=Snjeznik] and Veliki Risnjak.

Helix osoria B

RANCSIK 1889: 69, pl. II, fig. 4. Locus

typicus: Bosnia and Herzegovina: Miljacka valley.

Helix haueri B

RANCSIK 1889: 69, pl. II, fig. 3. Locus

typicus: Bosnia and Herzegovina: Mt. Trebeviæ.

Helix (Trichia) erjaveci blaui K

OBELT 1892: 6, pl. 122,

fig. 727

. Locus typicus: Bosnia and Herzegovina: near

Sarajevo.

Euomphalia (?) floerickei K

OBELT 1898: 162. Locus

typicus: Montenegro: Moraca.

Fruticicola erjaveci var. cincta S

OÓS 1904: 295. Locus

typicus: Croatia: Lika-Krbava [historic county, now in

south-western Croatia]: Goszpics [=Gospiæ].

Fruticicola erjaveci var. syrmiensis S

OÓS 1904: 295. Lo-

cus typicus: Serbia: Krušedol.

Fruticicola erjaveci leptolasia W

AGNER 1912: 250. Lo-

cus typicus: Bosnia and Herzegovina: Vlasiè near

Travnik and Jablanica.

Fruticicola erjaveci osoria oreinos W

AGNER 1915: 481,

pl. 18, figs. 154a-d (shell, genitalia). Locus typicus:

Bosnia and Herzegovina: Ljubièna and Radovina near

Celebiè.

Trichia erjaveci hajlensis Jaeckel in J

AECKEL &MEISE

1956: 29, pl. 1, fig. 4. Locus typicus: Serbia (Kosovo):

Hajla Planina 1,900 m a.l.s.

Material examined

Bosnia and Herzegovina: Vlašiènear Travnik,

MIZW, 5 s.; Croatia: Vorstadt Nova Ves near Agram

[=Zagreb], MIZW, 6 s.; Agram Tramvag, MIZW, 24 s.;

Agram, MIZW, 17 s.; Agram; leg. Èastek 1912, coll.

Poliñski 228/37, MIZW, 3 s.; Zagreb, MIZW, 12 s.;

Maksimir near Agram [now a part of Zagreb], MIZW,

10 s.; Brušane, reg. Lièko-senjska, MIZW, 1 s.; Kostaj

nica, reg. Sisaèko-moslavaèka, MIZW, 3 s.; Brodske

gory, MIZW, 3 s.; Serbia: Krušedol [monastery],

MIZW, 1 s.

Shell (Figs 74–76)

Shell cone-shaped, sometimes flattened, with

5.25–6.0 whorls. Shell height 5.6–8.5 mm, shell width

10.0–14.7 mm, height/width ratio 0.49–0.63, body

whorl height 5.2–7.3 mm, relative height of body

whorl 0,76–0,90, aperture height 3.0–4.3 mm, aper

ture width 5.4–7.0 mm, umbilicus major diameter

1.9–3.4 mm, umbilicus minor diameter 1.5–2.7 mm,

umbilicus major diameter/shell diameter ratio

0.16–0.24. Aperture transversely oval with distinct

white lip inside. Basal and palatal aperture margins

slightly reflexed. Umbilicus open, earlier whorls vis

ible. Adults hairless. Shell yellowish-white, sometimes

with lighter band on body whorl, matt and smooth,

sometimes translucent.

124 Ma³gorzata Proæków

Figs 74–76. T. erjaveci. Specimen from Zagreb – Croatia,

MIZW. Shell: 74 – apertural view, 75 – umbilical view, 76

– apical view. Scale bar 5 mm

Reproductive system

The description below is given according to W

NER

(1915) because material for anatomical examina

tion was unavailable.

Four pairs of short mucous glands, sometimes with

secondary branches. Inner dart sacs slightly longer

than outer ones. Vagina cylindrical. Flagellum shorter

than epiphallus, which is longer than fusiform penis.

Spermatheca duct short and thick. Spermatheca oval,

reaching ca. 1/2 spermoviduct length.

Ecology

T. erjaveci lives in damp, shaded forests, rarely in

open places; it is found on vegetation (preferably

shrubs). In the area of Batak Village it was collected at

1,100 m a.s.l. (I

RIKOV &MOLLOV 2006).

Distribution (Fig. 77)

A North-Western Balkan species; found in Croatia,

Bosnia and Herzegovina, Montenegro to Macedonia,

also in Hungary (environs of Budapest and south of

the country: Mecsek mountain range) (W

AGNER

1915, KERNEY et al. 1983), known from northern Alba

nia (D

HORA &WELTER-SCHULTES 1996) and Bulgaria

(Ropotamo National Park, Stara Planina, west of

Topolowgrad, southwest of Svilengrad, the Western

Rhodopes: only in the area of Batak Village) (D

JANOV

&LIKHAREV 1975, IRIKOV &MOLLOV 2006).

Trochulus filicinus (L. Pfeiffer, 1841)

Helix filicina L. PFEIFFER 1841: 39. Terra typica:

Slovenia: “Carniolia” [=Carniola].

Helix (Fruticicola) Bielzi var. bosnensis M

ÖLLENDORFF

1873: 35. Terra typica: Bosnia: Igmangebirge.

Fruticicola filicina filicina natio styriaca P

OLIÑSKI

1928: 181, pl. XXVII, fig. 32 [not 92]. Locus typicus:

Austria: Styria: Graz.

Material examined

Austria: Styria: Bärenschützklamm, NHMW, 5 s.;

Bärenschütz, Hochlantsch, MIZW, 1 s.; Hochlantsch,

coll. Poliñski 228/37, MIZW, 10 s.; Fronleiten, 1918,

leg. A. J. Wagner, coll. Poliñski 228/37, MIZW, 2 s.;

Ruine Peggau, coll. Edlauer 27407, NHMW, 12 s.;

Austria and Italy: Carnische Alpen [=Carnic Alps],

coll. Gallenstein 53532, NHMW, 12 s.; Slovenia: Mt.

Velika Gora, SW of Reifnitz (=Ribnica), Krain

[=Carniola], coll. Edlauer 20807, NHMW, 8 s.; Isonzo

[=Soèa] valley, Woltschach [=Volèe], W of Tolmin,

coll. Wagner, MIZW, 2 alc.; Mt. Grintovec, Kamnik

Alps, coll. Poliñski 228/37, MIZW, 1 s.

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 125

Fig. 77. Distribution of T. erjaveci. Black triangles – localities of origin of the examined material

Shell (Figs 78–80)

Shell roundish with 5.5–6.3 whorls. Shell height

5.3–7.6 mm, shell width 7.7–11.4 mm, height/width

ratio 0.56–0.73, body whorl height 4.3–5.8 mm, rela

tive height of body whorl 0.76–0.96, aperture height

2.7–4.1 mm, aperture width 4.3–6.2 mm, umbilicus

major diameter 0.2–0.9 mm, umbilicus minor diam

eter 0.2–0.9 mm, umbilicus major diameter/shell di

ameter ratio 0.02–0.11. Aperture oblique, basal aper

ture margin horizontally expanded and slightly re

flected, with distinct lip inside. Umbilicus very nar

126 Ma³gorzata Proæków

Figs 78–85. T. filicinus. Specimen from Hochlantsch, Styria, Austria, coll. Poliñski 228/37, MIZW. 78–80 – shell: 78 – apertural

view, 79 – apical view, 80 – umbilical view; 81–85 – reproductive system: 81 – general view, 82 – longitudinal section of va

gina, 83 – penial papilla, 84 – cross-section of penial papilla, 85 – cross-section of epiphallus. Scale bar 5 mm

row, sometimes partly or nearly entirely covered by

columellar aperture margin; when open, most often

round. Both juveniles and adults with short and dense

hairs. Shell horny-brown with prominent growth

lines, light band on body whorl.

Reproductive system (Figs 81–85)

Four pairs of short mucous glands situated around

upper vagina, ca. 3.5 mm from tips of inner dart sacs,

which are slightly longer than outer ones. Vagina very

long (ca. 9 mm); its expanded dart sac region narrows

toward genital atrium. Flagellum almost twice as long

as epiphallus, which is shorter than very long, cylin

drical penis. Spermatheca duct long and bent.

Spermatheca elongate, reaching albumen gland.

Longitudinal section of vagina, penial papilla,

cross-section of penial papilla and epiphallus shown

in Figs 82–85.

Ecology

T. filicinus lives in shaded, damp forests in the up

lands and lower mountain altitudes; it is quite often

found in river valleys.

Distribution (Fig. 86)

An East-Alpine and Carpathian species; known

from Austria (Carynthia, Styria), western Slovakia

(Inovec Mts), Hungary (Mecsek; Vértes; Örtilos;

Zalamerenge), Slovenia (Julian Alps: in the south to

Tolmin) and northern Italy (only in Alpi Giulie and

Carso) (P

OLIÑSKI 1928, KERNEY et al. 1983,

ANGANELLI et al. 1995, ÈEJKA et al. 2007); the

Dinaric Alps to southern Serbia (F

ALKNER 1990).

Remarks

The geographical “natio” styriaca is smaller and

more roundish than the nominate species: it has

rounded aperture, umbilicus almost covered, shorter

and sparser hairs (P

OLIÑSKI 1928).

Trochulus graminicolus (Falkner, 1973)

Trichia (Trichia) graminicola FALKNER 1973: 211, figs

1–4 (shell), fig. 6 (genitalia). Locus typicus: Germany:

Bayern: region of Donaueschingen: Eichberg near

Blumberg, W slopes at half distance between Achdorf

and Eschach. Holotype: Senckenberg-Museum Frank

furt SMF 228814.

The description of the species is given according

to F

ALKNER (1973) because I had neither dry shells

nor alcohol-preserved material at my disposal, and

failed to get access to the type material.

Shell

Shell strongly flattened, usually twice broader than

high, with nearly flat spire and 5.6–6.2 whorls (usually

5.7). Body whorl ca. 1.5 × wider than penultimate. Su-

ture deep, especially between the penultimate and

body whorl. Shell height 4.7–5.3 mm, shell major di-

ameter 9.6–10.7 mm, shell minor diameter 8.4–9.4

mm, aperture height 3.4–3.9 mm, aperture width

4.2–5.1 mm, umbilicus major diameter 2.1–3.0 mm,

umbilicus minor diameter 1.6–2.4 mm., umbilicus

major diameter/shell diameter ratio 0.22–0.29. Aper-

ture oblique, aperture margin reflected, with white

lip. Umbilicus wide. Adults hairless, in juveniles hairs

short and weak (0.1–0.2 mm long, near umbilicus

0.03–0.06 mm). Shell dark horny to reddish-brown, ir

regularly prominently striated above and smooth, dis

tinctly convex below; shiny, sometimes with light band

on body whorl.

Reproductive system

Four pairs of short mucous glands, sometimes with

secondary branches or reduced to four in total. Inner

dart sacs slightly longer than club-shaped outer ones.

Vagina very long and cylindrical (6.8–8.9 mm); dis

tance between outlet of mucous glands and tips of in

ner dart sacs (called by F

ALKNER Vaginahals = vaginal

neck, which I call upper vagina) equal to ca. 1/3 va

gina length. Flagellum usually longer than epiphallus,

which is longer than penis; but all these parts can be

approximately equal in length. Spermatheca duct

thin and long. Spermatheca elongatedly oval, not

reaching albumen gland.

Ecology

T. graminicolus lives in grassy screes in sparse

montane forests on steep slopes, with pine, spruce

and juniper (Calamagrostio variae-Pinetum commu

nity). Both juveniles and adults stay firmly attached to

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 127

Fig. 86. Distribution of T. filicinus. Black triangles – localities

of origin of the examined material

grass blades, especially to their underside. While feed

ing, they choose wider damaged blades. It is not

found directly on the ground or higher vegetation

ALKNER 1973).

Distribution (Fig. 87)

An endemic species; known only from south-west

ern Germany (Baden-Württemberg, Schwäbische Alb

near Geislingen and Blumberg) (F

ALKNER 1973).

Remarks

It differs from T. hispidus in a flatter shell and a

more oblique aperture. The vaginal neck is longer

than in the other members of Trochulus.

Trochulus hispidus (Linnaeus, 1758)

Helix hispida LINNAEUS 1758: 771. Terra typica:

Sweden.

Helix sericea

DRAPARNAUD 1801: 85. Terra typica:

not given in original description – probably France.

Helix plebeium D

RAPARNAUD 1805: 105, pl. VII, fig.

5. Terra typica: not given in original description –

probably France. Syn. nov.

Helix depilata C. P

FEIFFER 1821: 35, pl. II, fig. 18.

Locus typicus: Germany: near Cassel [=Kassel]:

“Bellevuegarten” [Palais Bellevue] and on

Kratzenberge, in grass and under fallen leaves.*

Helicella Prevostina R

ISSO 1826: 73. Terra typica:

France: dep. Alpes-Maritimes.*

Helix concinna J

EFFREYS 1830: 336. Locus typicus:

Wales: vicinity of Swansea, “under stones & c. in dry

places”; England: near Bristol, “among the rejecta

menta of the Avon river” [non Helix concinna Dupuy,

1848]. Syn nov.

Helix hispida Var. nana J

EFFREYS 1862: 199. Locus

typicus: England: Isle of Wight (Metcalfe), fresh

water.*

Helix Vendeana L

ETOURNEUX 1869: 60. Locus ty

picus: France: dep. Vendée: near Fontenay-le-

Comte.*

Helix Gratianopolitana R

AMBUR 1869: 267. Locus

typicus: France: environs of Grenoble.*

Helix liberta W

ESTERLUND 1871: 54. Locus typicus:

Sweden: prov. Skåne: near Lund, in garden.*

Helix Bellovacina M

ABILLE 1877: 305. Locus typicus:

France: dep. Oise:

“in humidis silvisque pagi

Compendiaci Galiae” [=probably Compiègne].*

Helix Steneligma Bourguignat in M

ABILLE 1877: 305.

Locus typicus: France: Pyrenees and dep. Oise:

“in silvis

pagi Compendiaci” [=probably Compiègne].*

Helix Elaverana Bourguignat in M

ABILLE 1877: 305.

Locus typicus: France: Pyrenees and environs of

Paris.*

Helix saporosa M

ABILLE 1877: 305. Locus typicus:

France: dep. Aisne:

“in silvis pagi Suessoniensis

Galiae” [=Soissons].*

Helix Axonana M

ABILLE 1877: 306. Locus typicus:

France: dep. Aisne: “in silvis ad Castrium-Theodorici

Galiae” [=Château-Thierry].*

Helix Matronica M

ABILLE 1877: 306. Locus typicus:

France: environs of Paris.*

Helix Goossensi M

ABILLE 1877: 306. Locus typicus:

France: dep. Essone: “in locis limpsis ad Firmitatem

Adelahidis Galiae” [=La Ferté-Alais].*

Helix hispidosa Letourneux in S

ERVAIN 1880: 60.

Locus typicus: France: dep. Loire-Atlantique: Vertou

near Nantes (coll. Letourneux); Spain: alluvions of

Arta [=Arga] near Pampelune (coll. Servain). [non

Helix hispidosa Bourguignat in Fagot, 1879]*

Helix badiella Ziegler in L

OCARD 1881: 107, fig. on

p. 107. Locus typicus: France: dep. Seine-et-Marne:

banks of Marne River towards drains of Pomponne

[lake], between Thorigny and Pomponne”.*

Helix urbana Coutagne in L

OCARD 1881: 107, fig.

on p. 108. Locus typicus: France: dep. Seine-et-Marne:

banks of Marne River, in environs of Lagny and

Pomponne.*

Helix Latiniacensis L

OCARD 1881: 108, fig. on p.

109. Locus typicus: France: dep. Seine-et-Marne: envi

rons of Paris: Lagny, Thorigny, Pomponne, Carnetin;

dep. Seine-et-Oise: Argenteuil, palace of Versailles;

Seine: Vincennes and Charenton (coll. Lo

card); Saint-Mandè forest and Paris: Arsenal street

near Bastille (coll. Coutagne).*

Helix subbadiella Bourguignat in L

OCARD 1882: 74,

317. Locus typicus: France: environs of Paris; dep.

Alpes-Maritimes: Monton (coll. Bourguignat).*

Helix Vendoperanensis Bourguignat in L

OCARD

1882: 76, 317. Locus typicus: France: dep. Aube: Ven

deuvre-sur Barse and Viélaines near Rosières; envi

rons of Lyon (coll. Bourguignat).*

128 Ma³gorzata Proæków

Fig. 87. Distribution of T. graminicolus

As FALKNER et al. (2002) point out, the name plebeius has been wrongly applied to the taxon Trochulus sericeus

(Draparnaud, 1801). F

ORCART (1965) originally proposed the replacement of the name sericeus by the younger name

plebeius to sort out a difficult taxonomic problem relating to the validity of the name sericeus (Draparnaud, 1801) in

contradistinction to sericeus (Müller, 1774). However, in so doing, he synonymised what have subsequently been regarded

as two distinct species, one widespread in north-west Europe, the other confined to the alpine region (A

NDERSON 2005).

Edited by Foxit Reader

For Evaluation Only.

Helix Vocoutiana Bourguignat in LOCARD 1882: 76,

317. Locus typicus: France: dep. Isère: La Salette near

Corps (coll. Bourguignat).*

Helix Hypsellina Pons in L

OCARD 1882: 78, 318. Lo

cus typicus: France: dep. Aveyron: Estaing (coll.

Bourguignat).*

Helix microgyra Bourguignat in L

OCARD 1882: 79,

319. Locus typicus: France: dep. Basses-Pyrènèes

[=Pyrénées-Atlantiques]: valley near Pic du Ger, above

Eaux-Bonnes (coll. Bourguignat).*

Helix Cularensis Bourguignat in L

OCARD 1882: 79,

319. Locus typicus: France: dep. Isère: Sassenage near

Grenoble (coll. Bourguignat).*

Helix hispidella Bourguignat in L

OCARD 1882: 79.

Locus typicus: France: dep. Loire-Inférieure

[=Loire-Atlantique]: Vertou near Nantes (coll.

Letourneux); dep. Aube: Othe forest,

vis-à-vis Montaigu near Bouilly (coll. Bourguignat).*

Helix Ataxiaca F

AGOT 1883: 220. Locus typicus:

France: dep. Aude: above Quillau, Fanges forest.

ALKNER et al. 2001: 58 – Trochulus ataxiacus (Fagot,

1884)].

Helix Beaudouini L

OCARD 1887: 165. Locus typicus:

France: dep. Côte-d’Or: environs of Châtillon-sur-

Seine.*

Helix Duesmensis L

OCARD 1887: 168. Locus typicus:

France: dep. Côte-d’Or: environs of Châtillon-sur-

Seine and Auxonne; dep. Aube: Courtenot and

Troyes; dep. Marne: Châlons-sur-Marne; dep.

Meurthe-et-Moselle: Bionville; dep. Moselle: Fouligny

and environs of Metz.*

Helix latiscensis L

OCARD 1887: 172. Locus typicus:

France: dep. Côte-d’Or: environs of Châtillon-sur-

Seine.*

Helix Sarinica L

OCARD 1887: 174. Locus typicus:

Switzerland: Sarine valley near Fribourg; France: allu

vions of Rhone, Lyon; dep. Ain: Chartreuse-de-Portes

and Dampierre; dep. Côte-d’Or: Châtillon-sur-Seine;

dep. Aube: environs of Troyes.*

Helix montigena L

OCARD 1894: 116. Locus typicus:

France: Ste Foy lès Lyon; dep. Ain: Chevry and Haute

ville; sources of the Doubs River.*

Helix Segusiana L

OCARD 1894: 116. Terra typica:

France: deps: Ain, Isère, Savoie and Haute-Saône.*

Helix Alixae Bourguignat in L

OCARD 1894: 120. Lo

cus typicus: France: dep. Hautes-Pyrènèes: Loudres.*

Helix Drunasiana L

OCARD 1894: 121. Locus typicus:

France: dep. Drôme: Die and Col du Rousset.*

Helix Barcelonnettensis Bourguignat in L

OCARD

1894: 125. Locus typicus: France: dep. Ain: Hauteville;

dep. Savoie: Albertville; dep. Hautes-Alpes: Barcelon

nette.*

Helix foeni L

OCARD 1894: 126. Terra typica: Fran

ce.*

Helix Pictavica Bourguignat in L

OCARD 1894: 127.

Locus typicus: France: Clain valley; dep. Vienne: be

tween Poitiers and Saint-Benolt.*

Helix Niverniaca L

OCARD 1894: 127. Locus typicus:

France: dep. Nièvre: environs of Nevers; dep.

Charente: Flèac.*

Helix Salinae Bourguignat in L

OCARD 1894: 131.

Locus typicus: France: dep. Jura: Salins.*

Helix subniverniaca C

AZIOT 1910: 115. Terra typica:

France: dep. Alpes-Maritimes.*

Helix Orzeszkoi C

AZIOT 1910: 117, pl. VI, fig. 36.

Terra typica: France: dep. Alpes-Maritimes.*

Material examined

Austria: Trins im Gschnitztal, Tyrol, MIZW, 2 s.

[plebeius]; Strasse von norders nach Martinsbrück,

1,300 m, Tyrol, MIZW, 5 s. [plebeius]; St. Jodok, Tyrol;

2.07.1998, ZW, 30 alc.; between Zirl and Innsbruck,

Tyrol, 2.07.1998, ZW, 15 alc.; Teichwiesen near Bruck

an der Leitha, Lower Austria, coll. Oberwimmer,

NHMW, 17 s.; loess Nußdorf ob der Traisen towards

Neusiedl, Lower Austria; coll. Klemm 63023, NHMW,

18 s.; Fischamend, Lower Austria, coll. Sterki 1,

NHMW, 21 s.; Schwöll near Pfongau, Upper Austria,

28.03.1946, coll. Klemm 23697, NHMW, 29 s.

[plebeius]; s. prec. loc., Carinthia, coll. Gallenstein

53569, NHMW, 22 s.; Britain: Box Hill near Dorking,

Surrey; 18–25.09.1963, leg. J. F. Peake & A. J. Knight,

BMNH, 10 s; Portholme Meadow, Huntingdon;

1.08.1988, leg. D. T. Bilton, BMNH, 3 alc. [plebeius];

France: Lyon; Jickeli, MIZW, 1 s. [plebeius]; Mt.

Ceindre near Lyon; coll. O. Retowski 41/39, MIZW, 3

s. [plebeius]; s. prec. loc. Frankreich; coll. Edlauer

48822, NHMW, 6 s. [plebeius]; Germany: Mauer an der

Elsenz near Heidelberg, Baden-Württemberg; coll.

Klemm 19416, NHMW, 11 s. [concinnus]; Freiburg,

MPW, 4 s. [concinnus]; Bonn, Breslauer Promenade,

MPW, 31 s.; Günzburg, Bavaria, coll. Edlauer 35288,

NHMW, 26 s.; Dinkelscherben, Bavaria, MIZW, 4 s.

[plebeius]; Netherlands: Amsterdam, NHMW, 15 s.;

Poland: Ziemia Lubuska: Zielona Góra, 03.1964, leg.

£u¿na, U£, 20 s.; Wielkopolska: Stêszew, 24.07.1979,

leg. E. Koralewska, UAM, 112 s.; Chyby, 18.06.1969,

leg. M. Stachowiak, UAM, 56 s.; Poznañ, 13.05.1991,

leg. E. Koralewska, UAM, 58 alc.; R¹bczyn, leg.

Strojkowska, UAM, 60 s.; Jarocin, 30.10.1976,

13.03.1977, leg. M. Micha³kiewicz, UAM, 179 s.;

Wêgierki, 2.07.1973, leg. E. Koralewska, UAM, 101 s.;

Lower Silesia: Zieleniec, Bystrzyca Dusznicka valley,

24.05.1997, ZW, 35 alc. [plebeius]; Wa³brzych, 07.1994,

leg. T. Maltz, MPW, 8 s.+58 alc.; Lubawka, 9.09.1994,

leg. M. Majkowska, MPW, 41 s.+27 alc.; Muszkowice,

26.08.1994, leg. M. Majkowska, E. Kuznik, MPW, 12

s.+59 alc.; O³awa, 30.07, 27.08.1973, leg. G. Notz,

MPW, 41 s.+203 alc.; Arnoldsmühle [=Jarno³tów in

Wroc³aw], MPW, 3 s. [concinnus]; Kapsdorf bei

Breslau [=Czerñczyce near Wroc³aw], MPW, 1 s.

[concinnus]; Greiffenstein [=Gryf castle in

Proszówka], MPW, 5 s.; Pomorze: Gdynia, 17.08.1993,

leg. T. Maltz, MPW, 51 s.; Mi³osna near Kwidzyñ, leg.

B. & A. Piechoccy, U£, 9 s.; Warmia and Mazury:

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 129

130 Ma³gorzata Proæków

Figs 88–95. T. hispidus. Shell. 88–91 – apertural view: 88 – specimen from Lubawka, Sudetes, Poland, ZW, 89 – specimen from

Muszkowice, Sudetes, Poland, ZW, 90 – specimen from Parzynów, Poland, ZW, 91 – specimen from Go³dap, Poland, ZW;

92–95 – umbilical view: 92 – specimen from Lubawka, Sudetes, Poland, ZW, 93 – specimen from Muszkowice, Sudety Foot

hills, Poland, ZW, 94 – specimen from Günzburg, Germany, NHMW, 95 – specimen from Zieleniec, Sudetes, Poland, ZW.

Scale bar 5 mm

Kudypy nature reserve, on the Pas³êka River,

5.09.1966, leg. A. Piechocki, U£, 3 s.; on the Pas³êka

River near Wapnik, 8.09.1966, leg. A. Piechocki, U£, 6

s.; Go³dap, on lake Go³dap, 25.08.1994, leg. M.

Majkowska, A. Leœniewska, MPW, 156 s.; B³¹dziszki,

26.08.1994, leg. T. Maltz, B. M. Pokryszko, MPW, 90

alc.; Mazowsze: Radom, 13–15.04.1919, leg. K. Gajl,

MIZW, 118 s.; Ziemia £ódzka: £ódz, 17.11.1960, leg.

A. Piechocki, U£, 2 s.; Œwiêtokrzyskie Mts: Œw. Krzy¿,

Polana Bielnik, 19, 24.06.1977, leg. A. Piechocki, U£,

3 s.; Œw. Krzy¿, near monastery, 20.06.1970, leg. A.

Piechocki, U£, 4 s.; Œw. Krzy¿, slope, 20.09.1973, leg.

A. Piechocki, U£, 10 s.; Kielce, Karczówka hill,

29.04.1978, leg. A. Piechocki, U£, 4 s.; S³owik near

Kielce, 30.04.1978, leg. A. Piechocki, U£, 1 s.; Wy¿yna

Ma³opolska: Z³oty Potok, Wa³y nature reserve,

7.07.1976, leg. E. Ruszkowska, U£, 2 s.; Bi³goraj,

17.07.1910, leg. Lorez, MIZW, 114 s.; Polesie: Che³m,

03.1923, leg. W. Wolberg, MIZW, 113 s.; Staw,

29.08.1994, leg. M. Majkowska, A. Leœniewska, MPW,

159 s.+46 alc.; Roztocze: Zwierzyniec, Roztoczañski

National Park, 23.07.1987, I. Kow. Jeziorko, U£, 14 s.;

¯urawnica, 1.08.1988, leg. A. Piechocki, U£, 11 s.;

Tartaczna Góra, 19.07.1989, leg. A. Piechocki, U£, 1

s.; Szewnia Dolna, 20.10.1989, leg. M. Puchyr, U£, 2 s.;

Turzyniec, 10.05.1989, leg. A. Piechocki, U£, 9 s.;

Ukraine: Taschlik [=Tashlyk], Podolien, 1873, coll. O.

Retowski 41/39, MIZW, 13 s. [plebeius];

Shell (Figs 88–95)

Shape variable: from roundish-conical to nearly

flattened, with 5–6 moderately convex whorls. Shell

height 3.0–6.7 mm, shell width 5.4–10.5 mm;

height/width ratio 0.47–0.78, body whorl height

2.9–5.2 mm, relative height of body whorl 0.73–0.93,

aperture height 1.9–3.8 mm, aperture width 2.6–5.0

mm, umbilicus major diameter 0.5–2.5 mm, umbili

cus minor diameter 0.6–2.3 mm, umbilicus major di

ameter/shell diameter ratio 0.1–0.31. Aperture with

thin white lip inside. Umbilicus entirely open and

wide, earlier whorls always visible. Hairs thin, short

and slightly curved, in adults often lost, leaving pro

nounced scars. Shell cream-coloured to dark brown,

some specimens have light band on body whorl.

Shell variation

Individual and inter-population variation of al

most all the studied morphometric characters is very

considerable (Figs 96–108). Each of the studied popu

lations differs statistically significantly from the re

maining ones in at least one character and the

intra-population variation ranges of most parameters

are very wide. Variation of some populations has been

discussed in P

ROÆKÓW (1997); a few more popula

tions are added here.

Almost every examined population differs statisti

cally significantly from the remaining ones in at least

one character. Most populations differ in their shell

size (Figs 96–103), and proportions (Figs 105–108),

and in the number of whorls (Fig. 104). Some pairs of

populations differ in all or nearly all the parameters

examined, e.g. those from Bonn, Muszkowice and

Zieleniec. Likewise, populations from Fischamend,

Jarocin and Go³dap differ from the remaining ones in

such characters as shell width and height/width ratio.

The least variable characters are relative height of

body whorl and umbilicus relative diameter.

Reproductive system (Figs 109–113)

Four pairs of short mucous glands, sometimes with

additional branches or some branches may be reduced

to 4–6 in total. Inner and outer dart sacs approximately

equal in length. Vagina cylindrical. Flagellum as long

as epiphallus or slightly longer. Epiphallus longer than

massive, fusiform and slightly bent penis. Spermatheca

duct straight. Spermatheca oval, reaching ca. 1/2

spermoviduct length. Longitudinal section of vagina,

penial papilla, cross-section of penial papilla and

epiphallus shown in Figs 110–113.

Ecology

T. hispidus lives in various damp habitats: in sparse

forests, semi-open (shrubland) and open (meadows,

cultivated areas) places, never in very dry sites; in sum-

mer it climbs plants, preferably stinging nettles

(Urtica dioica). It is usually found below 1,600 m a.s.l.,

but in the Alps and the Pyrenees it occurs up to

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 131

Fig. 96. Interpopulation variation of T. hispidus: shell height.

Statistically significant differences (Duncan test, p <

0.05) between the following populations: 1–2, 1–3, 1–5,

1–6, 1–7, 1–8, 1–9, 2–9, 2–10, 2–11, 3–10, 3–11, 4–9,

4–10, 5–10, 5–11, 6–9, 6–10, 6–11, 7–8, 7–9, 7–10, 7–11,

8–9, 8–10, 8–11, 9–10, 9–11

132 Ma³gorzata Proæków

Fig. 97. Interpopulation variation of T. hispidus: shell width.

Statistically significant differences (Duncan test, p <

0.05) between the following populations: 1–2, 1–3, 1–4,

1–5, 1–7, 1–8, 1–9, 1–10, 1–11, 2–5, 2–7, 2–9, 3–5, 3–7,

4–5, 4–7, 5–10, 6–7, 6–8, 7–8, 7–9, 7–10, 7–11, 8–9, 9–10

Fig. 98. Interpopulation variation of T. hispidus: aperture

height. Statistically significant differences (Duncan test,

p < 0.05) between the following populations: 1–2, 1–3,

1–6, 1–7, 1–9, 1–10, 2–9, 2–10, 3–9, 3–10, 4–9, 4–10, 5–6,

5–7, 5–9, 6–8, 6–9, 6–10, 6–11, 7–8, 7–9, 7–10, 7–11, 8–9,

8–10, 9–10, 9–11

Fig. 99. Interpopulation variation of T. hispidus: aperture

width. Statistically significant differences (Duncan test, p

< 0.05) between the following populations: 1–6, 1–7, 1–9,

2–6, 2–7, 2–9, 2–11, 3–9, 4–9, 5–7, 5–9, 6–8, 6–9, 6–10,

6–11, 7–8, 7–9, 7–10, 7–11, 8–9, 8–10, 9–10, 9–11

Fig. 100. Interpopulation variation of T. hispidus: body whorl

height. Statistically significant differences (Duncan test, p

< 0.05) between the following populations: 1–2, 1–6, 1–7,

1–8, 1–9, 1–10, 1–11, 2–9, 2–10, 2–11, 3–9, 3–10, 3–11,

4–9, 4–10,. 4–11, 5–7, 5–9, 5–10, 5–11, 6–7, 6–9, 6–10,

6–11, 7–8, 7–9, 7–10, 7–11, 8–9, 8–10, 8–11, 9–10, 9–11

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 133

Fig. 101. Interpopulation variation of T. hispidus: shell diam

eter. Statistically significant differences (Duncan test, p <

0.05) between the following populations: 1–2, 1–3, 1–4,

1–6, 1–7, 1–8, 1–9, 1–10, 1–11, 2–5, 2–7, 3–5, 3–7, 4–5,

4–7, 5–6, 5–7, 5–8, 5–9, 5–10, 5–11, 6–7, 6–8, 7–8, 7–9,

7–10, 7–11, 8–9, 8–10

Fig. 102. Interpopulation variation of T. hispidus: umbilicus

major diameter. Statistically significant differences

(Duncan test, p < 0.05) between the following popula

tions: 1–4, 1–5, 1–6, 1–8, 1–9, 1–10, 1–11, 2–4, 2–5, 2–6,

2–8, 2–9, 2–10, 2–11, 3–4, 3–5, 3–6, 3–8, 3–9, 3–10, 3–11,

4–5, 4–6, 4–7, 4–8, 4–9, 4–10, 4–11, 5–6, 5–7, 5–8, 5–9,

5–10, 5–11, 6–7, 6–9, 6–10, 6–11, 7–8, 7–9, 7–10, 7–11,

8–9, 8–10, 8–11, 9–10, 9–11

Fig. 103. Interpopulation variation of T. hispidus: umbilicus

minor diameter. Statistically significant differences

(Duncan test, p < 0.05) between the following popula-

tions: 1–4, 1–5, 1–6, 1–8, 1–9, 1–10, 1–11, 2–4, 2–5, 2–6,

2–8, 2–9, 2–10, 2–11, 3–4, 3–5, 3–8, 3–9, 3–10, 3–11, 4–5,

4–6, 4–7, 4–8, 4–9, 4–10, 4–11, 5–6, 5–7, 5–8, 5–10, 5–11,

6–7, 6–9, 6–10, 6–11, 7–8, 7–9, 7–10, 7–11, 8–9, 8–10,

8–11, 9–10, 9–11

Fig. 104. Interpopulation variation of T. hispidus: number of

whorls. Statistically significant differences (Duncan test,

p < 0.05) between the following populations: 1–2, 1–4,

1–6, 1–7, 1–9, 1–10, 1–11, 2–3, 2–5, 2–8, 2–9, 3–9, 3–11,

4–5, 4–9, 5–6, 5–7, 5–8, 5–10, 5–11, 6–8, 6–9, 7–8, 7–9,

8–9, 8–10, 8–11, 9–10, 9–11

134 Ma³gorzata Proæków

Fig. 105. Interpopulation variation of T. hispidus: shell

height/width ratio. Statistically significant differences

(Duncan test, p < 0.05) between the following popula-

tions: 1–2, 1–3, 1–5, 1–7, 1–8, 1–9, 1–10, 1–11, 2–4, 2–5,

2–6, 2–7, 2–10, 2–11, 3–4, 3–5, 3–6, 3–7, 3–10, 3–11, 4–5,

4–8, 4–9, 4–10, 5–6, 5–7, 5–8, 5–9, 5–10, 5–11, 6–7, 6–8,

6–9, 6–10, 6–11, 7–8, 7–9, 7–10, 8–9, 8–10, 8–11, 9–10,

9–11, 10–11

Fig. 106. Interpopulation variation of T. hispidus: umbilicus

major diameter/shell diameter. Statistically significant

differences (Duncan test, p < 0.05) between the follow

ing populations: 1–4, 1–5, 1–6, 1–7, 1–8, 1–9, 1–10, 1–11,

2–4, 2–5, 2–6, 2–7, 2–8, 2–9, 2–10, 2–11, 3–4, 3–5, 3–6,

3–8, 3–9, 3–10, 3–11, 4–5, 4–6, 4–7, 4–8, 4–9, 4–10, 4–11,

5–7, 5–8, 5–9, 5–10, 5–11, 6–8, 6–9, 6–10, 6–11, 7–9, 7–10,

7–11, 8–9, 8–10, 8–11, 9–10, 9–11

Fig. 107. Interpopulation variation of T. hispidus: minor/ma-

jor diameter of umbilicus. Statistically significant differ-

ences (Duncan test, p < 0.05) between the following

populations: 1–7, 2–7, 3–7, 4–7, 6–7, 7–8, 7–9, 7–10,

7–11, 8–9

Fig. 108. Interpopulation variation of T. hispidus: relative

height of body whorl. Statistically significant differences

(Duncan test, p < 0.05) between the following popula

tions: 1–2, 1–3, 1–5, 1–8, 1–10, 1–11, 2–4, 2–5, 2–7, 3–4,

3–6, 3–7, 3–8, 3–9, 4–5, 4–10, 4–11, 5–6, 5–7, 5–8, 5–9,

5–10, 6–11, 7–8, 7–9, 7–10, 7–11, 8–11, 9–11

2,300 m a.s.l. In Spain it usually lives in dry places,

only single sites are damp, e.g. stream banks, vicinity

of springs (A

LTONAGA et al. 1994).

Distribution (Fig. 114)

A widely distributed European species: in the

north, along the Atlantic coast of Scandinava, it

reaches the Arctic circle (S

HILEYKO 1978a, RIEDEL

1988), in the east – southern Ukraine, the Urals and

St. Petersburg (F

ORCART 1965), in the west – the Brit

ish Isles (K

ERNEY et al. 1983). Its southern distribu

tion border is not precisely known: it has been re

corded from north-eastern Spain (from Catalonia to

the Picos de Europa Mts; common in the Basque

Country; in the south to province de Teruel in Aragon

and Mt. Moncayo in Sistema Ibérico) (A

LTONAGA et

al. 1994), northern Italy (M

ANGANELLI et al. 1995); in

western Bulgaria it has only few records (near Tran,

not confirmed in the Western Rhodopes)

(DAMJANOV

&LIKHAREV 1975, IRIKOV &MOLLOV 2006, IRIKOV &

RÖSS 2008), northern Croatia (Bansko Brdo)

HERKOVICH et al. 2008). In Poland T. hispidus is

common in the whole country except the Carpa

thians, in the Sudetes recorded up to 800 m a.s.l.

IKTOR & WIKTOR 1968).

Remarks

Till recently T. hispidus was regarded as distinct

from the very similar T. plebeius (Draparnaud,

1805)/sericeus (Draparnaud, 1801) and T. concinnus

(Jeffreys, 1830), descibed from western and central Eu

rope (see synonyms). Delimitation of the three forms

has always been regarded as problematic. Analysis of

individual and inter-population shell variation of T.

hispidus from Poland revealed a very wide variation

range (P

ROÆKÓW 1997). With respect to shell propor

The genus Trochulus Cheminitz, 1786 – a taxonomic revision 135

Figs 109–113. T. hispidus. Specimen from Lubawka, Sudetes, Poland, ZW. Reproductive system: 109 – general view, 110 – lon

gitudinal section of vagina, 111 – penial papilla, 112 – cross-section of penial papilla, 113 – cross-section of epiphallus.

Scale bar 5 mm

tions, T. hispidus from Muszkowice and Zieleniec dis

tinctly departs from the remaining Polish populations,

thus resembling T. plebeius. Biometrical studies on an

extensive material (a total of 2,339 specimens) (Figs

115–122) showed that the shell variation ranges of T.

plebeius, T. concinnus and T. hispidus overlapped, and it

was impossible to indicate any diagnostic characters of

the three nominal taxa. Similar results, although based

on a poorer material, were obtained by N

AGGS (1985).

136 Ma³gorzata Proæków

Fig. 114. Distribution of T. hispidus. Black triangles – localities of origin of the examined material

Fig. 115. Morphometric comparison of shells of T. hispidus,

T. plebeius and T. concinnus. Shell height plotted against

height/width ratio

Therefore, in spite of the very laconic descriptions of

T. plebeius (D

RAPARNAUD 1805: 105) and T. concinnus

EFFREYS 1830: 336), it seems to be reasonable to

synonymise T. plebeius, T. concinnus and T. hispidus. The

single shell (subadult) labelled as a type of T. plebeius,

deposited in D

RAPARNAUD’s collection in Vienna, actu

ally is not a Trochulus, but a Monachoides species, and

was no doubt mislaid (W

INTER 1990), which was al

ready remarked by L

OCARD (1895).

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 137

Fig. 116. Morphometric comparison of shells of T. hispidus,

T. plebeius and T. concinnus. Shell width plotted against

shell height

Fig. 118. Morphometric comparison of shells of T. hispidus,

T. plebeius and T. concinnus. Ratio of umbilicus major to

shell diameter plotted against height/width ratio

Fig. 120. Morphometric comparison of shells of T. hispidus

and T. plebeius with populations from Zieleniec and

Muszkowice. Shell width plotted against shell height

Fig. 119. Morphometric comparison of shells of T. hispidus

and T. plebeius with populations from Zieleniec and Musz

kowice. Shell height plotted against height/width ratio

Fig. 117. Morphometric comparison of shells of T. hispidus,

T. plebeius and T. concinnus. Relative height of body whorl

plotted against hight/width ratio

Trochulus leucozonus (C. Pfeiffer, 1828)

Helix leucozona C. PFEIFFER 1828: 34, pl. VI, figs

19–20. Terra typica: “Illyrien”.

Helix dolopida C

RISTOFORI &JAN 1832: 1. Terra

typica: “Ital. bor.” [probably North Italy]

Helix leucozona var. ovirensis R

OSSMÄSSLER 1838: 4,

pl. XXXI, fig. 434. Locus typicus: Austria: Carinthia:

Ovir summit, under stones.

Helix leucozona var. heteromorpha W

ESTERLUND

1876–1878: 52. Terra typica: Austria: “Carinthia”.

Helix leucozona var. Erjaveci C

LESSIN 1887: 122. Lo

cus typicus: Slovenia: “Im Friaul, Eingang zur Grotte

Vodnik ober Tublje auf dem Karste”. [according to

ANK (1995) a synonym of leucozona heteromorpha]

Material examined

Austria: Vellach near Eisenkappel, Karawanken,

leg. V. Sturany, MIZW, 1 alc.; between Kozje [=Kos

siach] and ‘Ir’, ± 1650 m, Karawanken, 09.1913, coll.

Poliñski 228/37, MIZW, 19 s.; Bärntal, Karawanken,

MIZW, 1 s.; Velika Kotla (=Gerlouc=Gerloutz), Kara

wanken, 18.09.1913, leg. W. Poliñski, MIZW, 3 s.;

Loiblstrasse, Karawanken, 19.09.1913, coll. Poliñski

228/37, MIZW, 1 s.; s. prec. loc., Karawanken, ex coll.

H. Gallenstein, coll. Poliñski 228/37, MIZW, 2 s.;

Hochobir, 1,100–2,000 m, MIZW, 8 s. [ovirensis]; Mt.

Dobratsch, Carynthia, MIZW, 11 s. [ovirensis]; Villach,

Carynthia, 1918, MIZW, 3 s.; Croatia: between Rijeka

and Opatija, 09.1913, leg. W. Poliñski, coll. Poliñski

228/37, MIZW, 2 s.: between Rann and Agram [=Za

greb], MIZW, 4 s. [dolopida]; Italy: Sette Comuni,

prov. Veneto, MIZW, 5 alc.: Dosso dei Morti, Ada

mello Alps, 2,000 m, 27.07.1932, coll. Edlauer 10620,

NHMW, 15 s.: Matajur, Julian Alps, 1,000 m, coll.

Poliñski 228/37, MIZW, 3 s.: Slovenia: Gonobitz

[=Slovenske Konjice], Bachergebirge [=Pohorje],

coll. Poliñski 228/37, MIZW, 1 s.; Mt. Grintovec,

Kamnik Alps, 1,800 m, coll. Edlauer 10635, NHMW, 7

s.; Mt. Ursula [= Uršlja Gora], Karawanken, coll.

Oberwimmer, NHMW, 4 s. [dolopida]; Mt. Ursula

[=Uršlja Gora], Karawanken, coll. Klemm 69381,

NHMW, 15 ok. [dolopida]; Mt. Stol, Krain [=Carniola],

22.09.1923, leg. W. Poliñski, coll. Poliñski 228/37,

MIZW, 8 s.; Mt. Razor, between Isonzo [=Soèa] valley

and Woheinersee [=Bohinjsko jezero], MIZW, 4 s.;

Isonzo [=Soèa] valley, environs of Tolmin, coll. Wag

ner, MIZW, 1 s.+2 alc.; Mt. Èaven, “Tarnovaner Wald”,

NW Haidenschaft [=Ajdovšèina], MIZW, 3 s.

Shell (Figs 123–125)

Shell conical to roundish with 5.5–6.5 regularly

coiled, convex whorls and dome-shaped spire. Shell

height 4.5–7.6 mm, shell width 7.7–11.4 mm,

height/width ratio 0.54–0.76, body whorl height

4.0–4.9 mm (one specimen 5.2 mm), relative height

of body whorl 0.7–0.79, aperture height 2.6–3.6 mm,

aperture width 3.6–5.3 mm, umbilicus major diam

eter 0.2–0.8 mm, umbilicus minor diameter 0.2–0.8

mm, umbilicus major diameter/shell diameter ratio

0.06–0.1. Aperture slightly oblique, crescentic; aper

ture margin sharp, its short columellar wall passes

into horizontal or almost horizontal basal wall with

distinct shiny lip inside. Umbilicus narrow, usually

open but sometimes partly or, very rarely, entirely cov

ered by columellar aperture margin. Adults hairless.

Shell pale to horny brown, with fine growth lines,

light band on body whorl.

138 Ma³gorzata Proæków

Fig. 121. Morphometric comparison of shells of T. hispidus

and T. plebeius with populations from Zieleniec and

Muszkowice. Relative height of body whorl plotted

against hight/width ratio

Fig. 122. Morphometric comparison of shells of T. hispidus

and T. plebeius with populations from Zieleniec and

Muszkowice. Ratio of umbilicus major diameter to shell

diameter plotted against height/width ratio

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 139

Figs 123–130. T. leucozonus. Specimen from Villach, Carynthia, Austria, 1918, MIZW. 123–125 – shell: 123 – apertural view,

124 – apical view, 125 – umbilical view; 126–130 – reproductive system: 126 – general view, 127 – longitudinal section of va

gina, 128 – penial papilla, 129 – cross-section of penial papilla, 130 – cross-section of epiphallus. Scale bar 5 mm

Reproductive system (Figs 126–130)

Four pairs of short mucous glands situated around

upper vagina, ca. 4 mm upper tips of inner dart sacs,

which are slightly longer and narrower than outer

ones. Vagina long (6.0–8.7 mm); its expanded dart

sac region narrows toward genital atrium. Flagellum

considerably longer than epiphallus, which is shorter

than fusiform penis. Spermatheca duct straight.

Spermatheca elongate, reaching ca. 1/2 spermovi

duct length. Longitudinal section of vagina, penial

papilla, cross-section of penial papilla and epiphallus

shown in Figs 127–130.

Ecology

T. leucozonus lives in damp shaded forests, under

stones, in leaf litter and in mosses; it is found up to the

alpine zone. According to B

ANK (1995) its various

forms reach the following altitudes: T. leucozonus s. str.

275–1,900 m a.s.l.; T. leucozonus ovirensis 1,700–2,150 m

a.s.l. and T. leucozonus heteromorpha 220–2,200 m a.s.l.

Distribution (Fig. 131)

A South-East-Alpine species; found in Austria

(Carynthia), Slovenia (Pohorje; Kamnik Alps;

Carniola; Julian Alps: source of Soèa; western

Slovenia: the environs of Sava sources and Èaven sum-

mit), northern Italy and Croatia (P

OLIÑSKI 1928,

ANK 1995, MANGANELLI et al. 1995).

Remarks

Based on the umbilicus diameter, B

ANK (1995) dis-

tinguished two geographical subspecies: leucozonus s.

str. (umbilicus open or only partly covered, 1/13–1/7

of shell diameter) and heteromorpha (umbilicus cov

ered to a great extent, not more than 1/15 of shell di

ameter). According to that autor subspecies ovirensis

represents an Alpine form (living between 1,700 and

2,150 m a.s.l.) derived from leucozonus s. str. Besides,

he ragards subspecies erjaveci as a synonym of

heteromorpha.

The examined specimens of ovirensis had distinctly

smaller shells (height 4.1–6.3 mm, width 6.5–9.4 mm)

compared to leucozonus s. str. (height 4.5–7.6 mm,

width 7.7–11.4 mm) and fewer whorls (ovirensis

5.1–6.0, mean 5.45; leucozonus s. str. 5.25–6.3, mean

5.8).

The examined shells of specimens from Mt. Ursula

and from “between Rann and Agram”, representing

dolopida, were much larger: height 6.5–8.2 mm, width

9.2–12.2 mm, body whorl height 4.4–6.1 mm, aper

ture height 4.4–6.1 mm and aperture width 4.8–6.1

mm. On the other hand, their proportions were simi

lar to those of leucozonus s. str.: height/width ratio

0.59–0.74, relative height of body whorl 0.68–0.81,

umbilicus major diameter/shell diameter ratio

0.04–0.09. The number of whorls (5.8–6.7) and umbi

licus diameter (major 0.4–0.9 mm, minor 0.4–0.8

mm) were also very similar. Because of the unavail

ability of specimens from the locality and alcohol-

preserved material, the taxonomic status of this taxon

has to remain uncertain. However, B

ANK (1995) placed

dolopida among the synonyms of leucozonus s. str.

Trochulus lubomirskii (Œlósarski, 1881)

Helix (Fruticicola) Lubomirski ŒLÓSARSKI 1881: 319,

pl. X, figs A-D. Locus typicus: Poland: Œwiêty Krzy¿

(£ysa Góra) and Ogrodzieniec.

Helix Clessini U

LIÈNY 1884: 7–8, figs on p. 3. Locus

typicus: Czech Republic: Moravia: near Olomouc and

Brno.

Material examined

Austria: Wien, MIZW, 1 s.; Korneuburg near Wien,

coll. Poliñski 228/37, MIZW, 1 s.; Poland: Lower

Silesia: Mt. Œlê¿a, 27.04.1955, leg. A. Wiktor, Nr 98,

MPW, 5 s.; Cracow-Wieluñ Jura: Mirów castle ruins, leg.

E. Poliñska 1920, MIZW, 7 s.; Œwiêtokrzyskie Mts: Œw.

Krzy¿ [=£ysa Góra], 22.10.1971, leg. A. Piechocki, U£,

8 s.; Beskid Wyspowy: Limanowa, 15.08.1999, leg. E.

Su³kowski, ZW, 4 alc.; Pogórze Przemyskie: Przemyœl,

coll. O. Retowski 41/39, MIZW, 5 s.; Bieszczady Mts:

Ustrzyki Górne, 21.08.1996, ZW, 5 alc.; Slovakia: Plaveè

castle ruins, reg. Spiš, 540–547 m, 12.08.1922, coll.

Poliñski 228/37, MIZW, 1 s.; Skalky near Uják [=Údol],

reg. Prešov, 12.08.1922, coll. Poliñski 228/37, MIZW, 1

s.; Árvaváralja [=Oravský Podzámok], reg. Zilina, coll.

Poliñski 228/37, MIZW, 1 s.; Ukraine: Z³oczów

[=Zo³ocziw], MIZW, 7 s.

Shell (Figs 132–134)

Shell conical, dome-shaped spire with acute apex;

4.7–5 moderately tightly coiled whorls. Dimensions:

shell height 5.0–7.0 mm, shell width 7.0–10.0 mm,

height/width ratio 0.62–0.77, body whorl height

4.2–5.7 mm, relative height of body whorl 0.82–0.88,

aperture height 3.0–3,7 mm, aperture width 4.0–5.3

mm, umbilicus major diameter 0.3–0.9 mm, umbili

cus minor diameter 0.3–0.7 mm, umbilicus major di

ameter/shell diameter ratio 0.06–0.11. Aperture

rounded, aperture margin thin with very weakly de

140 Ma³gorzata Proæków

Fig. 131. Distribution of T. leucozonus. Black triangles – local

ities of origin of the examined material

veloped lip. Umbilicus round and very narrow but al

ways open. Few, very short (0.1 mm) and fine hairs,

usually lost in adults. Shell translucent, whitish-yellow

to greenish-yellow, weakly shiny.

Reproductive system (Figs 135–139)

Four pairs of short mucous glands situated around

upper vagina, ca. 1.5 mm from tips of inner dart sacs

which reach beyond outer ones and are much more

massive. Vagina rather long and cylindrical. Very

short and conical flagellum passes into thick cylindri

cal epiphallus, which is longer than fusiform penis.

Spermoviduct bent just near outlet of free oviduct.

Spermatheca duct rather thick, straight and short.

Spermatheca big and irregularly club-shaped, not

reaching albumen gland. Longitudinal section of va

gina, penial papilla, cross-section of penial papilla

and epiphallus shown in Figs 136–139.

Ecology

T. lubomirskii usually lives in dense herbal vegetation

(Petasites sp.) in relatively damp forests (river plains,

mountains up to 1,600 m a.s.l.); it is found under

stones, sometimes on limestone rocks and meadows.

Distribution (Fig. 140)

A Carpathian species; known from Austria (only in

Lower Austria: Danube valley), the Czech Republic and

Slovakia (scattered records in northern Bohemia,

northern and eastern Moravia, the Beskidy Mts, the Slo-

vak Carpathians); Poland (Carpathians – localities from

the Tatra Mts not confirmed, Carpathian Foothills, to

the Eastern Sudetes, single localities in the Œwiêto-

krzyskie Mts, Mt. Œlê¿a – probably absent, Trzebnickie

Hills and environs of Krotoszyn) (R

IEDEL 1988, WIKTOR

2004), Hungary (Somoskoujfalu, the Zemplén Mts,

Bárna) (K

ERNEY et al. 1983) and western Ukraine

(Podolia: Worochta, Transcarpathian region: Jasina

[=Yasinya=Yasina=Jasinja=Körösmezö]) (P

OLIÑSKI

1924).

Remarks

The internal structure of the vagina presented by

HILEYKO (1978a, b) does not correspond with the

structure of the examined specimens (fig. 136).

Trochulus luridus (C. Pfeiffer, 1828)

Helix lurida C. PFEIFFER 1828: 33, pl. VI, figs 14–15.

Terra typica: “Illyrien”.

Helix lurida var. separanda R

OSSMÄSSLER 1835: 35,

Nr 360. Terra typica: Slovenia: Krain.

Material examined

Austria: Petzen, Karawanken, leg. Gredler, 1927,

coll. Poliñski, MIZW, 2 s.; Bosnia and Herzegovina:

Vlašiæ mountain range near Travnik, coll. Poliñski

228/37, MIZW, 5 s.; Mt. Trebeviè near Sarajevo, coll.

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 141

Figs 132–134. T. lubomirskii. Specimen from Ustrzyki Górne,

Bieszczady, Poland, ZW. Shell: 132 – apertural view, 133 –

umbilical view, 134 – apical view. Scale bar 5 mm

Poliñski 228/37, MIZW, 2 s.; Mt. Dragoljub, N

Sarajevo, coll. Poliñski 228/37, MIZW, 2 s.; Croatia:

Fiume [=Rijeka], MIZW, 1 s.; Šestine near Agram

[=Zagreb], coll. Poliñski 228/37, MIZW, 7 s.; Agram

[=Zagreb], coll. Poliñski 228/37, MIZW, 1 s.; Zagreb,

1918, leg. Wagner, MIZW, 2 s.; Lasinjsko, S Zagreb,

coll. Poliñski 228/37, MIZW, 2 s.; Tuskanec near

Zagreb, MIZW, 5 s.; Virovitica, reg. Virovitièk-Pod

ravska, MIZW, 2 s.; Jankovac, reg. Brodsko-Posavska,

coll. Poliñski 228/37, MIZW, 4 s.; Kapela near

Jezerane, reg. Lièko-Senjska, coll. Poliñski 228/37,

MIZW, 5 s.; Ogulin, reg. Karlovaèka, coll. Poliñski

228/37, MIZW, 3 s.; s. prec. loc., coll. Poliñski 228/37,

MIZW, 5 s.; Italy: Pazanella, Tyrol, 1,500 m, 1916, coll.

Edlauer 6442, NHMW, 3 s.; Cormons, prov. Gorizia,

MIZW, 2 s.; Serbia: Mt. Stolac, near Bajina Bašta, coll.

Poliñski 228/37, MIZW, 1 s.; Slovenia: Kankertal

[=Kokra valley], Kamnik Alps, coll. Poliñski 228/37,

MIZW, 1 s.; environs of Kranj (=Krainburg),03.1928,

leg. L. Kušcer, MIZW, 2 alc.; Rothweinklamm

[=Radovna valley] near Bled, Julian Alps, 06.1934,

leg. Klemm, coll. Klemm 2407, NHMW, 7 s.;

142 Ma³gorzata Proæków

Figs 135–139. T. lubomirskii. Specimen from Ustrzyki Górne, Bieszczady, Poland, ZW. Reproductive system: 135 – general view,

136 – longitudinal section of vagina, 137 – penial papilla, 138 – cross-section of penial papilla, 139 – cross-section of

epiphallus. Scale bar 5 mm

Fig. 140. Distribution of T. lubomirskii. Black triangles – localities of origin of the examined material

Ulrichsberg [=Šenturška gora], Krain=Carniola, coll.

Poliñski 228/37, MIZW, 3 s.; s. prec. loc., leg. Kušèer,

03.1928, MIZW, 2 alc.

Shell (Figs 141–143)

Shell roundish with 5.5–6.6 whorls. Shell height

5.0–8.3 mm, shell width 7.8–12.6 mm, height/width

ratio 0.51–0.71, body whorl height 3.8–6.0 mm, rela

tive height of body whorl 0.75–0.88, aperture height

2.3–4.0 mm, aperture width 4.0–6.2 mm, umbilicus

major diameter 0.2–1.1 mm, umbilicus minor diame

ter 0.2–1.1 mm, umbilicus major diameter/shell di

ameter ratio 0.02–0.1. Aperture oblique, aperture

margin sharp, slightly reflected with reddish lip in

side. Umbilicus very narrow, sometimes partly cov

ered by columellar aperture margin. Short, fine and

rather sparse. Shell horny-brown to reddish-brown,

slightly shiny, sometimes with light band on body

whorl.

Reproductive system (Figs 144–148)

Four pairs of rather long (ca. 4 mm) mucous

glands situated around upper vagina, ca. 2 mm from

tips of inner dart sacs. Inner and outer dart sacs ap-

proximately equal in size. Vagina very long; its very ex-

panded dart sac region narrows toward genital

atrium. Flagellum slightly longer the epiphallus

which is shorter than fusiform penis, or all three sec-

tions of approximately equal length. Spermatheca

duct thin and straight. Spermatheca elongate, reach-

ing ca. spermoviduct 2/3 length. Longitudinal sec-

tion of vagina, penial papilla, cross-section of penial

papilla and epiphallus shown in Figs 145–148.

Ecology

T. luridus lives in damp, shaded forests, among leaf

litter.

Distribution (Fig. 149)

A South-Alpine species; known from Austria

(Carinthia: S slopes of Karawanken), Slovenia

(Kamnik Alps, Julian Alps), Croatia (in the east from

Zagreb, Korana valley to Velika and Mala Kapela),

western Bosnia and Herzegowina, northern Italy (en

virons of Gorizia, Sette Comuni, S. Tirol) (P

OLIÑSKI

1928, MANGANELLI et al. 1995).

Trochulus montanus (Studer, 1820)

Glischrus (Helix) montana STUDER 1820: 86 [12].

Locus typicus: Switzerland: Berner Jura, in forests and

on damp rocks. Based on juveniles [see also F

ORCART

1957: 194]. Lectotype: NHMB.

Helix submontana M

ABILLE 1868: 22. Locus typicus:

France: dep. Jura: Saint-Amour; dep. Ain: Bellegarde.

= Helix Pascali Mabille

, 1868: 22. [in synonymy]*

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 143

Figs 141–143. T. luridus. Specimen from Agram, Croatia, coll.

Poliñski 228/37, MIZW. Shell: 141 – apertural view, 142 –

apical view, 143 – umbilical view. Scale bar 5 mm

Helix Dubisiana Coutagne in LOCARD 1882: 77,

318. Locus typicus: France: Haut-Doubs Mountains:

Mont Dore; dep. Jura (coll. Coutagne).*

Helix plebicola L

OCARD 1888: 366. Locus typicus:

France: dep. Ain: Tenay and Colombier; N of Lyon:

alluvions of Rhône; dep. Haute-Savoie: Évian; dep.

Jura: Bief-du-Fourg; dep. Aube (coll. Bourguignat).*

Material examined

Switzerland: Taubenlochschlucht near Biel, cant.

Berne, 18.07.1935, coll. Edlauer 22122, NHMW, 5 s.;

Taubenlochschlucht near Biel, cant. Berne, coll.

Klemm 54283, NHMW, 4 s.; Les Roudez near

Delémont, cant. Jura, 19.04.1969, leg. Falkner, 85033,

NHMW, 5 alc.

Shell (Figs 150–152)

Shell roundish-conical, 5.25–6.0 convex whorls with

moderately deep suture. Shell height 4.5–7.0 mm, shell

width 9.3–12.0 mm; height/width ratio 0.47–0.78,

body whorl height 4.5–5.3 mm, relative height of body

whorl 0.77–0.92, aperture height 3.0–4.1 mm, aperture

width 4.5–5.7 mm, umbilicus major diameter 1.7–2.0

mm, umbilicus minor diameter 1.1–1.8 mm, umbilicus

major diameter/shell diameter ratio 0.11–0.26. Aper

ture with white lip. Umbilicus open and moderately

wide, earlier whorls visible. Few thin hairs present only

in juveniles. Colour variable: from horny-yellow to red

dish-brown, shell shiny, finely striated, usually with

light band on body whorl.

Reproductive system (Fig. 153)

Four pairs of short mucous glands. Inner dart sacs

reach beyond outer ones. Vagina long and cylindrical.

Flagellum cosiderably longer than epiphallus, which

is shorter than massive and fusiform penis. Sper

matheca duct long. Spermatheca oval, reaching ca.

2/3 spermoviduct length.

Ecology

T. montanus lives in forests on steep mountain

slopes, on shaded rocks; between 400 and 1,600 m a.s.l.

Distribution (Fig. 154)

A North-Alpine species; known from France (Alps

of Savoy: Grand Chartereuse), Switzerland (only in

144 Ma³gorzata Proæków

Figs 144–148. T. luridus. Specimen from Kranj, Slovenia,

coll. Poliñski 228/37, MIZW. Reproductive system: 144 –

general view, 145 – longitudinal section of vagina, 146 –

penial papilla, 147 – cross-section of penial papilla, 148 –

cross-section of epiphallus. Scale bar 5 mm

Fig. 149. Distribution of T. luridus. Black triangles – localities

of origin of the examined material

Jura) and Italy (Western Alps) (FORCART 1965,

ANGANELLI et al. 1995, TURNER et al. 1998).

Remarks

It is not always possible to distinguish this species

from T. striolatus based on conchological characters

alone. The two species show only slight differences in

the structure of their reproductive system. T.

montanus has the flagellum considerably longer than

the epiphallus, and the inner dart sacs reach far be

yond the outer ones. T. striolatus has the flagellum as

long as the epiphallus or slightly longer, and the inner

dart sacs are only slightly longer than the outer ones.

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 145

Figs 150–153. T. montanus. Specimen from Les Roudez near Delémont, Switzerland, 85033, NHMW. 150–152 – shell: 150 –

apertural view, 151 – umbilical view, 152 – apical view; 153 – reproductive system. Scale bar 5 mm

Fig. 154. Distribution of T. montanus. Black triangles – local

ities of origin of the examined material

Trochulus piccardi Pfenninger et Pfenninger, 2005

Trichia piccardi HAUSSER 2005: 153. [nomen

nudum]

Trochulus piccardi P

FENNINGER &PFENNINGER

2005: 264, figs 4–5 (shell and genitalia). Locus typi

cus: Switzerland: canton Vaud: Commune de Chateau

d’Oex, meadow at the foot of the church-hill,

(46°28’37’’N, 07°07’83’’E, 970 m a.s.l.). Holotype:

Naturmuseum Senckenberg, Frankfurt am Main –

SMF 328194; paratypes: Bayerische Staatssammlung,

München and Naturhistorische Museum, Bern – SMF

328195/11.

Since the species was described only recently, the

original description is detailed and supported with

molecular data, and its separate status raises no

doubts, the description below is given after

FENNINGER &PFENNINGER (2005).

Shell and animal

Shell rather thick-walled, glossy, opaque, light

brown, depressed with elevated apex. Mean height

3.9±0.4 (range 3.3–5.0), width 6.4±0.7 (5.6–8.7).

Mean number of whorls 4.7±0.3 (4.3–5.3); whorls de-

scending with pronounced suture. Elliptical aperture

broader than high, downwards slanted, often with

whitish rim inside. Shell increment reflected in

slanted striation on upperside. Periphery round and

not shouldered. Umbilicus ca. 15% of shell width.

Shell hairless; no hair scars visible on first whorls.

Body light, with wedge-shaped dark grey pigment spot

on head behind ommatophores.

Reproductive system

The reproductive system departs in some respects

from the typical Trochulus structure. Neither male nor

female parts are pigmented. The three sections of the

male ducts have a length ratio of ca. 1:1.6:1.2 (19.5

mm total length), with a strongly coiled flagellum.

The penial retractor muscle attaches to the epiphallus

directly behind the penis. No accessory dart sacs are

visible in the female tract. However, since the main

dart sacs are unusually broad, fusion of the sacs is

possible. Eight mucous glands present, inserting di

rectly above dart sacs.

Habitat

The snail lives on extensively used pastures on

south-facing slopes; it is found on the ground, among

the vegetation composed of typical alpine herbs and

grasses.

Distribution (Fig. 155)

Plain d’Afflon, near Enney. Fresh shells were

found in various, ecologically similar places in the

lower Saanen-valley.

Trochulus striolatus (C. Pfeiffer, 1828)

Cochlea rufescens DA COSTA 1778: 80, pl. IV, fig. 6.

Locus typicus: England: Cornwall and Hampshire;

Wales: Flintshire: Leeswood and others. [see also

ALKNER 1995: 99]

Helix striolata C. P

FEIFFER 1828: 28, pl. VI, fig. 8. Lo

cus typicus: Germany: near Heidelberg, in gardens.

Helix montana C. P

FEIFFER, 1828: 33, pl. VI, fig. 9.

Locus typicus: Germany: near Heidelberg, the castle

hill; Austria: forests near Vienna.

Helix danubialis C

LESSIN 1874: 184, pl. VIII, fig. 4.

Locus typicus: Germany: Bavaria: near Dillingen,

woods on the border of the Bavarian Danube.

Helix altenana Klees in L

OCARD 1882: 79. Locus

typicus: France: dep. Nord: vicinity of Lille; dep.

Pas-de-Calais: Boulogne-sur-Mer; dep. Aube:

Bar-sur-Seine (coll. Bourguignat) and Courtenot.*

Helix abludens L

OCARD 1888: 334. Locus typicus:

Jersey; Ireland: environs of Dublin; France: dep.

Pas-de-Calais: Boulogne-sur-Mer. [see Falkner et al.

2001: 57 – Trochulus striolatus abludens (Locard, 1888)]

Helix rufescentella L

OCARD 1894: 128–129. Locus

typicus: France: dep. Nord: Valenciennes and Lille.*

Helix (Trichia) britannica W

ESTERLUND 1894: 164.

Locus typicus: England: London.*

Helix (Hygromia, Fruticicola) montana var. juvavensis

EYER 1914: 276. Locus typicus: Austria: north slopes

of Schafberg. Paratypes: NHMW.

Material examined

Austria: Hollerberg, north of See am Mondsee,

Upper Austria, 1945, coll. Klemm 56657, NHMW, 19 s.;

Scharflingerpass, Salzkammergut, 600 m, 8.09.1997,

leg. A. Riedel & F. Seidl, MIZW, 19 s.; Schafberg,

Salzkammergut, ± 1,700 m, 8.09.1997, leg. A. Riedel

& F. Seidl, MIZW, 7 alc. [juvavensis]; Schafberg,

Salzkammergut, coll. Rusnov R/105/5, NHMW, 5 s.

[paratypes T. striolatus juvavensis]; Höllengebirge,

Upper Austria, 21.09.1968, coll. Klemm 86624,

NHMW, 4 alc. [juvavensis]; Höllengebirge, cable car

station, Upper Austria, NHMW, 10 s. [juvavensis];

146 Ma³gorzata Proæków

Fig. 155. Distribution of T. piccardi

Höllengebirge, cable car station, Upper Austria, coll.

Klemm 80844: NHMW, 19 s.; Mt. Hochlecken,

Höllengebirge plateau, 1,600 m, Upper Austria,

NHMW, 30 s.; Mt. Feuerkogel, Höllengebirge near

Ebensee, Upper Austria, 1927, coll. Edlauer 6873, 11 s.;

Fischamend, Lower Austria, coll. Rusnov, NHMW, 22 s.

[danubialis]; Grimsing, Wachau valley, Lower Austria,

coll. Klemm 63024, NHMW, 12 s. [danubialis]; Britain:

Box Hill near Dorking, county Surrey, 18–25.09.1963,

leg. J. F. Peake & A. J. Knight, BMNH, 18 alc.; Epsom,

county Surrey, V 1939, leg. A. E. Ellis, Nr 2136, BMNH,

16 s.; Durrington near Salisbury, county Wiltshire, leg.

A. E. Ellis, Nr 2136, BMNH, 9 s.; Porton near Salisbury,

county Wiltshire, 08.1917, leg. Boycott, Nr 1218,

BMNH, 32 s.; Cheltenham, county Gloucestershire,

2.06.1954, leg. I. S. Hawkins, Nr 1392–1396, BMNH, 29

s.; Germany: Ulm, Württemberg, leg. Prinzing, MIZW,

19 s.; Geislingen an der Steige, Württemberg, coll.

Edlauer 9103, NHMW, 16 s.; Mt. Hohenstausen near

Göppingen, Schwäbische Alb, leg. Regius, coll. Geyer

47033, NHMW, 13 s.; Nürburg castle ruins near

Adenau, Rhineland-Palatinate, coll. O. Retowski

41/39, MIZW, 8 s.; Günzburg, Bavaria, coll. Edlauer

36332, NHMW, 26 s.; Sonthofen, Bavarian Alps, leg.

Gutten, MIZW, 10 s.

Shell (Figs 156–158)

Shell roundish-conical with 5.3–6.5 moderately con-

vex whorls, body whorl slightly expanded and often an-

gled. Shell height 5.2–8.8 mm, shell width 8.5–14.0 mm;

height/width ratio 0.51–0.7, body whorl height 4.3–7.1

mm, relative height of body whorl 0.75–0.94, aperture

height 3.0–5.5 mm, aperture width 3.9–7.2 mm, umbili-

cus major diameter 1.5–2.7 mm, umbilicus minor diame

ter 0.9–2.4 mm, umbilicus major diameter/shell diame

ter ratio 0.1–0.24. Aperture with white lip, aperture mar

gin very slightly reflected. Umbilicus open, earlier whorls

visible. Adults usually hairless. Shell dark reddish-brown

to creamy-yellow, usually with light band on body whorl.

Shell variation

Almost every studied population differs statistically

significantly from the remaining ones in at least one

character (Figs 159–171). Some pairs of populations

differ in all or nearly all the parameters examined,

e.g. those from Höllengebirge, Schafberg and

Scharflingerpass. Some characters, such as shell

height and width (Figs 159, 160), aperture height and

width (Figs 161, 162), body whorl height (Fig. 163),

and umbilicus major and minor diameter (Figs 165,

166) are very much variable. The least variable char

acters are those describing the shell proportions i.e.

height/width ratio (Fig. 168), relative height of body

whorl (Fig. 171) and umbilicus relative diameter (Fig.

169). The shell diameter (Fig. 164) and umbilicus mi

nor/major diameter ratio (Fig. 170) show no statisti

cally significant differences between the examined

populations.

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 147

Figs 156–158. T. striolatus. Specimen from Ulm, Württem

berg, Germany, MIZW. Shell: 156 – apertural view, 157 –

umbilical view, 158 – apical view. Scale bar 5 mm

148 Ma³gorzata Proæków

Fig. 159. Interpopulation variation of T. striolatus: shell

height. Statistically significant differences (Duncan test,

p < 0.05) between the following populations: 1–6, 1–8,

1–9, 1–10, 2–6, 2–8, 2–9, 3–4, 3–6, 3–7, 3–8, 3–9, 3–10,

4–6, 4–9, 4–10, 5–6, 5–8, 5–9, 5–10, 6–7, 6–8, 6–10, 7–8,

7–9, 7–10, 8–9, 8–10

Fig. 160. Interpopulation variation of T. striolatus: shell

width. Statistically significant differences (Duncan test, p

< 0.05) between the following populations: 1–2, 1–4, 1–5,

1–6, 1–7, 1–8, 1–9, 1–10, 2–3, 2–6, 2–8, 2–9, 2–10, 3–4,

3–6, 3–7, 3–8, 3–9, 3–10, 4–6, 4–8, 4–9, 4–10, 5–6, 5–8,

5–9, 5–10, 6–7, 6–8, 6–9, 6–10, 7–8, 7–9, 7–10, 8–9, 8–10

Fig. 161. Interpopulation variation of T. striolatus: aperture

height. Statistically significant differences (Duncan test,

p < 0.05) between the following populations: 1–2, 1–4,

1–5, 1–6, 1–7, 1–8, 1–9, 1–10, 2–4, 2–5, 2–6, 2–7, 2–8, 2–9,

2–10, 3–4, 3–5, 3–6, 3–7, 3–8, 3–9, 3–10, 4–5, 4–6, 4–9,

4–10, 5–6, 5–8, 5–9, 5–10, 6–7, 6–8, 6–9, 6–10, 7–9, 7–10,

8–9, 8–10

Fig. 162. Interpopulation variation of T. striolatus: aperture

width. Statistical significance of differences (Duncan

test, p < 0.05) are between the following populations:

1–3, 1–4, 1–5, 1–6, 1–8, 1–9, 1–10, 2–3, 2–4, 2–5, 2–6, 2–8,

2–9, 2–10, 3–4, 3–5, 3–6, 3–7, 3–8, 3–9, 3–10, 4–5, 4–6,

4–7, 4–9, 4–10, 5–6, 5–8, 5–9, 5–10, 6–7, 6–8, 6–10, 7–8,

7–9, 7–10, 8–9, 8–10

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 149

Fig. 163. Interpopulation variation of T. striolatus: body

whorl height. Statistically significant differences

(Duncan test, p < 0.05) between the following popula-

tions: 1–4, 1–6, 1–8, 1–9, 1–10, 2–3, 2–4, 2–6, 2–8, 2–9,

2–10, 3–4, 3–6, 3–7, 3–8, 3–9, 3–10, 4–5, 4–6, 4–9, 4–10,

5–6, 5–8, 5–9, 5–10, 6–7, 6–8, 6–9, 6–10, 7–8, 7–9, 7–10,

8–9, 8–10

Fig. 164. Interpopulation variation of T. striolatus: shell di

ameter. No statistically significant differences between

the studied populations

Fig. 165. Interpopulation variation of T. striolatus: umbilicus

major diameter. Statistically significant differences

(Duncan test, p < 0,05) between the following popula-

tions: 1–2, 1–5, 1–6, 1–8, 1–9, 1–10, 2–3, 2–4, 2–9, 2–10,

3–5, 3–6, 3–7, 3–8, 3–9, 3–10, 4–5, 4–6, 4–8, 4–9, 4–10,

5–9, 5–10, 6–9, 6–10, 7–10, 8–9, 8–10

Fig. 166. Interpopulation variation of T. striolatus: umbilicus

minor diameter. Statistically significant differences

(Duncan test, p < 0.05) between the following popula

tions: 1–2, 1–5, 1–6, 1–7, 1–8, 1–9, 1–10, 2–3, 2–5, 2–6,

2–9, 2–10, 3–5, 3–6, 3–7, 3–8, 3–9, 3–10, 4–5, 4–6, 4–7,

4–8, 4–9, 4–10, 5–9, 5–10, 6–9, 6–10, 7–10, 8–9, 8–10

150 Ma³gorzata Proæków

Fig. 167. Interpopulation variation of T. striolatus: number of

whorls. Statistically significant differences (Duncan test,

p < 0.05) between the following populations: 1–5, 2–5,

3–9, 3–10, 4–9, 4–10, 5–6, 5–7, 5–9, 5–10, 6–10, 7–8, 8–9,

8–10

Fig. 168. Interpopulation variation of T. striolatus: shell

height/width ratio. Statistically significant differences

(Duncan test, p < 0.05) between the following popula

tions: 1–2, 1–5, 2–3, 2–6, 2–7, 3–5, 5–6, 5–7

Fig. 169. Interpopulation variation of T. striolatus: umbilicus

major diameter/shell diameter. Statistically significant

differences (Duncan test, p < 0.05) between the follow-

ing populations: 2–4, 2–6, 4–5, 4–7, 5–6, 6–7, 6–8, 6–10

Fig. 170. Interpopulation variation of T. striolatus: umbilicus

minor/major diameter ratio. No statistically significant

differences between the studied populations

Reproductive system (Figs 172–176)

Four pairs of short mucous glands. Inner dart sacs

reach only slightly beyond outer ones, which are con-

siderably more massive. Vagina long, its dart sac re

gion narrows toward genital atrium. Flagellum longer

than epiphallus. Epiphallus as long as massive and

fusiform penis, or longer. Spermatheca duct thick.

Spermatheca roundish, not reaching albumen gland.

Longitudinal section of vagina, penial papilla,

cross-section of penial papilla and epiphallus shown

in Figs 172–176.

Ecology

T. striolatus lives in humid and shaded habitats in

forests, scrub and roadsides; also in gardens (espe

cially in northern France and the British Isles where it

is an introduced synanthrope); in Austria found be

tween 150 and 2,000 m a.l.s. (K

LEMM 1974), in north

ern Switzerland at 400–800 m a.s.l.

Distribution (Fig. 177)

A North-West-European species; known from the

British Isles, northern France (to the Seine River),

southern part of the Netherlands, southern Germany

(to the middle Rhine River; mainly in river valleys and

on lowlands), northern Switzerland (canton

Schaffhausen), Austria (except the southern part)

and Slovakia (Danube floodplain, southern part of

Little Carpathians and Zilinská Basin) and Hungary

ORCART 1965, KERNEY et al. 1983, ÈEJKA 2000).

Remarks

The specimens from Fischamend were deter-

mined as Trochulus striolatus danubialis (Clessin, 1874).

However, the shell variation ranges of this population

and those of T. striolatus from the remainig localities

overlap (Figs 159–171). Besides, the differences be-

tween this population and the remaining ones are as

statistically significant as those among the popula

tions of T. striolatus s. str. Moreover, the reproductive

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 151

Fig. 171. Interpopulation variation of T. striolatus: relative

height of body whorl. Statistically significant differences

(Duncan test, p < 0.05) between the following popula-

tions: 1–4, 2–4, 2–5, 2–8, 2–9, 2–10, 3–4, 3–5, 3–6, 3–8,

3–9, 3–10, 4–7, 5–7, 7–8, 7–10

Figs 172–176. T. striolatus. Specimen from Box Hill near Dorking, Great Britain, BMNH. Reproductive system: 172 – general

view, 173 – longitudinal section of vagina, 174 – penial papilla, 175 – cross-section of penial papilla, 176 – cross-section of

epiphallus. Scale bar 5 mm

system of the examined specimens from Great Britain

displays characters which are present in both striolatus

and danubialis described in S

HILEYKO (1978b), i.e. the

expanded dart sac region narrows toward the genital

atrium and the flagellum is longer than the

epiphallus (like in striolatus;S

HILEYKO 1978b). How-

ever, the spermatheca duct is short and thick, and the

spermatheca does not reach the albumen gland (like

in danubialis;S

HILEYKO 1978b). Thus, the above data

support F

ORCART’s (1965) statement that “we cannot

recognize geographical subspecies, but that the area

of distribution of T. striolata is split up into colonies

with local variations”. In this context, it is likely that

the subspecies danubialis is conspecific with Trochulus

striolatus (

C. Pfeiffer, 1828), despite the fact that

HILEYKO (1978b) regards it as a distinct species.

The pecularities of the external morphology of the

reproductive system of T. striolatus danubialis were pre

viously noted by H

UDEC (1964). He did not compare

its structure to T. striolatus s. str., but to other mem

bers of Trochulus (T. hispidus, T. villosulus, T. sericeus

and T. lubomirskii) inhabiting the Czech Republic.

Two charactes were recognized as subspecific i.e. the

inner dart sacs considerably thinner than the outer

ones and the relatively thick spermathecal duct, com

pared to free oviduct. The features given by H

UDEC

(1964) are, in fact, identical with those of T. striolatus

s. str. from Great Britain (specimens examined cf. Figs

172–176).

Specimens from Schafberg and Höllengebirge rep

resent subspecies T. striolatus juvavensis (Geyer, 1914)

(Figs 178–180). They differ most of all in smaller shell

height and width (mean values 5.61 mm and 9.23 mm,

respectively). The mean aperture height and width are

also small (3.22 mm and 4.61 mm, respectively). In T.

striolatus s. str. the values are in the following ranges:

shell height 5.7–7.2 mm (mean 6.82 mm) (Fig. 159),

shell width 9.8–12.4 mm (mean 11.59 mm) (Fig. 160),

aperture height 3.3–4.6 mm (mean 4.27 mm) (Fig.

161) and aperture width 4.7–6.4 mm (mean 5.91 mm)

(Fig. 162). The coefficients describing the shell pro

portions are similar in all the studied populations, and

their variation ranges overlap (Figs 168–171). The

152 Ma³gorzata Proæków

Fig. 177. Distribution of T. striolatus. Black triangles – local

ities of origin of the examined material

Figs 178–180. T striolatus juvavensis. Paratype. Specimen

from Schafberg, Salzkammergut, Austria, coll. Rusnov

R/105/5, NHMW. Shell: 178 – apertural view, 179 – um

bilical view, 180 – apical view. Scale bar 5 mm

mean number of whorls is 5.53 in T. striolatus juvavensis

and a slightly higher in T. striolatus s. str. (5.65–5.89,

mean of all populations 5.76); but also their variation

ranges overlap (Fig. 167).

The reproductive systems in T. striolatus s. str. (Figs

172–185) and T. striolatus juvavensis (Figs 181–185)

display considerable similarities both in shape and

proportions of their component parts. Thus,

ORCART’s (1965) suggestion that juvavensis Geyer,

1914 from Schafberg is probably conspecific with T.

suberecta seems ungrounded.

Trochulus suberectus (Clessin, 1878)

Helix suberecta CLESSIN 1878b: 46. Locus typicus:

Germany: Bavaria: Dreifaltigkeitsberg above

Kleinprüfening near Regensburg (higher fossil loess

layers), near Passau; Austria: Linz; Elbe and Rhine

valleys.

Material examined

Austria: Mühl near St. Jodok, Tyrol, 2.07.1998, ZW,

4 alc.

Shell (Figs 186–188)

Shell roundish-conical with 5–5.2 convex whorls.

Shell height 4.0–5.6 mm, shell width 5.0–7.5 mm,

height/width ratio 0,63–0,71, body whorl height

3.5–3.9 mm, relative height of body whorl 0.79–0.83,

aperture height 2.7–3.0 mm, aperture width 3.7–3.9

mm, umbilicus major diameter 0.7–1.1 mm, umbili

cus minor diameter 0.6–1.0 mm, umbilicus major di

ameter/shell diameter ratio 0.14–0.15. Aperture with

thin white lip. Umbilicus entirely open and wide, ear-

lier whorls visible. Hairs thin, short and slightly

curved, often lost in adults, leaving pronounced scars.

Shell light to dark brown, some specimens with light

band on body whorl.

Reproductive system (Figs 189–193)

Six short mucous glands. Inner and outer dart sacs

approximately equal in size. Vagina long and cylindri

cal. Flagellum longer than epiphallus, which is

shorter than conical penis. Spermatheca duct rather

thick. Spermatheca oval, reaching ca. 1/2

spermoviduct length. Longitudinal section of vagina,

penial papilla, cross-section of penial papilla and

epiphallus shown in Figs 190–193.

Ecology

T. suberectus lives in montane habitats between

1,000 and 2,000 m a.s.l., in shaded scrub along

streams; it shelters in tussocks of vegetation or under

fallen leaves.

Distribution (Fig. 194)

A species known from some localities in southern

Germany (Schwäbische Alb), western Austria (Tirol:

Trins, Mühl near St. Jodok, Lech valley), north-west

ern Italy and eastern Switzerland (Unter Engadin:

Swiss National Park, Münster valley) (F

ORCART 1965,

ERNEY et al. 1983).

Remarks

Conchologically T. suberectus is very similar to T.

hispidus. It differs from other members of Trochulus in

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 153

Figs 181–185. T. striolatus juvavensis. Specimen from Schafberg – Salzkammergut, Austria, MIZW. Reproductive system: 181 –

general view, 182 – longitudinal section of vagina, 183 – penial papilla, 184 – cross-section of penial papilla, 185 –

cross-section of epiphallus. Scale bar 5 mm

154 Ma³gorzata Proæków

Figs 186–193. T. suberectus. Specimen from Mühl near St. Jodok, Tirol, Austria, ZW. 186–188 – shell: 186 – apertural view, 187

– umbilical view, 188 – apical view; 189–193 – reproductive system: 189 – general view, 190 – longitudinal section of va

gina, 191 – penial papilla, 192 – cross-section of penial papilla, 193 – cross-section of epiphallus. Scale bar 5 mm

having 6 branches of the mucous glands. However, dis

section of eastern Swiss specimens showed that the

form was conspecific with Trochulus sericeus sensu Dra

parnaud, 1801, where occasionally some of the eight

mucous glands can be missing (T

URNER et al. 1998).

Trochulus unidentatus (Draparnaud, 1805)

Helix unidentata DRAPARNAUD 1805: 81, pl. VII, fig.

15. Terra typica: France: Bresse. Type specimen:

NHMW.

Helix Cobresiana A

LTEN 1812: 79, pl. IX, fig. 18. Lo-

cus typicus: Germany: vicinity of Lechhausen [now a

part of Augsburg] and outside Ublaß.

Helix monodon F

ÉRUSSAC 1821–1822: 35 [quarto

edition], no 122. Locus typicus: France: La

Franche-Comté; Germany: Augsburg; E Switzerland.

Helix ventricosa C

RISTOFORI &JAN 1832: 1, no 6–54.

Terra typica: Amer. Jamaica (labels probably mislead)

[non Helix ventricosa Müller].*

Helix unidens Ziegler in

L. PFEIFFER 1848: 151. [in

synonymy] Terra typica: Germany, France, Italy: Al

pine summits”.*

Helix (Fruticicola) unidentata var. alpestris C

LESSIN

1878a: 84, pl. III, fig. 12. Locus typicus: Austria: Tyrol:

Lafatsch and Styria [not Kärnten]: Schneealpe (noti

fied by Tschapeck).

Helix unidentata var. anodonta T

SCHAPECK 1880:

190, pl. VI, fig. 4. Locus typicus: Austria: Styria:

Prebichel-Passe.

Fruticicola unidentata var. (natio) norica P

OLIÑSKI

1928: 161–163, pl. XXVII, fig. 20. Locus typicus: Aus

tria: Styria: Präbühel-Pass [=Prebichl]; Lower Austria:

Schönbühel near Melk; Kuchl near Salzburg;

Salzburg Alterbachgenist; Ach near Burghausen

[near Salzburg].

Fruticicola unidentata var. subalpestris P

OLIÑSKI 1928:

163–165, tab, XXVII, figs 22–23. Locus typicus: Swit

zerland:

“Basler Jura” and Valzeina (canton Grau

bünden); Italy: Tyrol: Glisenklamm near Sterzing [=

Vipiteno]; Germany: Bavaria: Burgkirchen an der Alz;

Austria: Styria: Seegraben near Aflenz.

Fruticicola unidentata var. (natio) subtecta P

OLIÑSKI

1928: 167, pl. XXVII, figs 24–26. Locus typicus: Aus

tria: various localities near Vienna: Markenstein ruins

near Vöslau; Gutenstein in Piestingtal; gorges near

Fischau; Hoche Wand – Dürnbachklamm; Hoche

Wand near Stollhof; Hoche Wand above Leiter

grabens; Sirningtal (between Puchberg and Stixen

stein); vicinity of Payerbach; Klause near Mariensee in

Wechselgebiet; Pittental valley (Scheiblingkirchen).

[see F

ALKNER 1990: 206, FALKNER et al. 2001: 58 –

Petasina subtecta]

Fruticicola unidentata var. carpatica P

OLIÑSKI 1928:

170–172, pl. XXVII, figs 27–28. Locus typicus: Poland:

Carpathians: Zamki Orawskie (S slopes of Carpa

thians between Babia Góra and Tatra Mts); Zaryte

(gm. Limanowa), Mszana Dolna; Homole gorge near

Jaworki; £omnica (commune Nowy S¹cz); Mt.

Cisownice; Trzy Korony; Tatra Mts (N slopes); Kopa

Magóry (Tatra Mts); “Jatki Bialskie” (Tatry Bialskie).

Material examined

Austria: Tirol: Trins, 1.07.1998, ZW, 5 s.; St.

Jodok, 2.07.1998, ZW, 5 alc.; Vals, 2.07.1998, ZW, 5

alc.; Mühl near St. Jodok, 2.07.1998, ZW, 5 alc.;

Putzalpe near Reutte, coll. O. Retowski 41/39,

MIZW, 5 s. [alpestris]; Austria: Rottenbach near

Haag am Hausruck, 9.09.1997, leg. A. Riedel et F.

Seidl, MIZW, 3 s.; Pupping near Eferding, 9.09.1997,

leg. A. Riedel et F. Seidl, MIZW, 1 s.; Überackern,

Braunau am Inn, 7.09.1997, leg. A. Riedel et F. Seidl,

MIZW, 1 s.; Wildshut, on Salzbach River near St.

Pantaleon, 7.09.1997, leg. A. Riedel et F. Seidl,

MIZW, 7 s.+7 alc.; Sengsengebirg Wald, 1380 m,

08.1958, NHMW, 28 s.; Ramsau, Salzkammergut,

4.07.1998, ZW, 5 alc.; Salzburg: Salzach-Klamm

Gorge near Pass Lueg, 6.09.1997, leg. A. Riedel et F.

Seidl, MIZW, 5 s.+3 alc.; Golling near Salzburg,

6.09.1997, leg. A. Riedel et F. Seidl, MIZW, 23 s.;

Scharflingerpass, Salzkammergut, 8.09.1997, leg. A.

Riedel et F. Seidl, MIZW, 1 s.; Mt. Schafberg,

Salzkammergut, 8.09.1997, leg. A. Riedel et F. Seidl,

MIZW, 4 s.+6 alc.; Kuchl, Salzburg, leg. A. Wagner

12, MIZW, 10 s.; Alterbachgeniste, Salzburg, coll.

Rusnov, NHMW, 68 s.; Hinterseegenist near

Faistenau, Salzburg, coll. Klemm 64768, NHMW, 25

s. [norica]; Tiefbrunnau, Salzkammergut, 4.07.1998,

ZW, 10 alc.; Ebenau, Salzkammergut, 4.07.1998, ZW,

7 alc.; between Fuschl and St. Gilgen,

Salzkammergut, 5.07.1998, ZW, 5 alc.; environs of Vi

enna and Lower Austria: Dornbacher Park near

Wien, coll. Poliñski 228/37, MIZW, 6 s.; Augarten

Wien, coll. Poliñski 228/37, MIZW, 3 s.; Schneeberg

near Wien, 1,900 m, 1918, coll. Poliñski 228/37,

MIZW, 3 s. [alpestris]; Hinterbrühl near Mödling,

coll. Poliñski 228/37, MIZW, 1 s.; Maleiten near

Fischau, coll. Poliñski 228/37, MIZW, 10 s.;

Gutenstein in Piestingtal, coll. Poliñski 228/37,

MIZW, 1 s.; Dürnbachklamm, coll. Poliñski 228/37,

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 155

Fig. 194. Distribution of T. suberectus. Black triangle – locality

of origin of the examined material

MIZW, 6 s.; Styria: Mt. Leobner Mauer, Eisenerzer

Alpen, 1,800–2,000 m, coll. Edlauer 43925, NHMW,

30 s. [subalpestris]; Prebichl, ex coll. A. J. Wagner, coll.

Poliñski 228/37, MIZW, 4 s.; Präbuhel, cotypes, ex

coll. Tschapeck, coll. Poliñski 228/37, MIZW, 2 s.

[anodonta]; Gleisdorf, on Raba River, coll. Poliñski

228/37, MIZW, 4 s.; Mürzenist near Kapfenberg, coll.

Poliñski 228/37, MIZW, 3 s.; between Aussee and

Grundlsee, 15.08.1912, leg. A. J. Wagner, coll. Poliñski

228/37, MIZW, 3 s.; Klachau near Mitterndorf [=Bad

Mitterndorf], coll. Rusnov, NHMW, 13 s. [anodonta];

Carynthia: Villach, coll. Poliñski 228/37, MIZW, 3 s.;

Magdalensberg, coll. Edlauer 50642, NHMW, 31 s.; s.

prec. loc., Catynthia, coll. Poliñski 228/37, MIZW, 2 s.;

Croatia: s. prec. loc., ex coll. Jickeli, coll. Poliñski

228/37, MIZW, 2 s.; Czech Republic: Srbsko near

Karlštejn, Bøeznice, 25.09.1960, leg. A. Riedel, 66/60,

MIZW, 3 alc.; Ruine Landsberg b. Geiersberg

[=Lanšperk castle ruins near Letohrad, reg.

Pardubice], coll. O. Retowski 41/39, MIZW, 6 s.;

Landsberg [=Lanšperk], Böhmen, coll. Poliñski

228/37, MIZW, 5 s.; Germany: Bavaria: Ulm, leg.

Prinzing, coll. Poliñski 228/37, MIZW, 1 s.; München,

on Isar River, coll. Poliñski 228/37, MIZW, 1 s.;

Moosburg, 30.06.1998, ZW, 39 alc.; Raitenhaslach

near Burghausen, 7.09.1997, leg. A. Riedel et F. Seidl,

MIZW, 1 s.; Grafussing near Simbach, 10.09.1997, leg.

A. Riedel et F. Seidl, MIZW, 9 s.; Gemeinde Ering,

5.09.1997, MIZW, 11 alc.; Italy: Ugovizza, Friuli Val

Canale valley (between Drava and Tagliamento

Rivers), leg. Alzona, 09.1925, MIZW, 5 s.; Poland:

Sudetes: Kletno, Kleœnica stream valley, 17.08.1996,

ZW, 35 s.+12 alc.; Beskid Œl¹ski: Cisownica near

Ustroñ, 1918, leg. Wagner, MIZW, 4 s. [carpatica];

Beskid ¯ywiecki: Ciapków Stream Valley near Rycerka,

24.09.1976, leg. A. Wiktor, Nr 1920, MPW, 3 alc.;

Abrahamów Stream Valley near Rycerka, 26.09.1976,

leg. A. Wiktor, Nr 1921, MPW, 2 alc.; stream valley on

Przegibek, 24.09.1976, leg. A. Wiktor, Nr 1917, MPW,

2 alc.; nature reserve Œrubita, 13.08.1970, leg. A.

Wiktor, Nr 1759, MPW, 2 alc.; Mt. Babia Góra,

4.08.1970, leg. A. Wiktor, Nr 1705, MPW, 5 alc.; below

Palice, Mt. Babia Góra, 25.05.1964, leg. A. Wiktor, Nr

1172, MPW, 2 alc.; above Wid³y, Mt. Babia Góra,

23.08.1964, leg. A. Wiktor, Nr 1164, MPW, 2 alc.;

Gorce: Rzeki, 16.08.1975, leg. A. Wiktor, Nr 1897,

MPW, 6 alc.; Mt. Marsza³ek, 7.07.1962, leg. A. Wiktor,

Nr 846, MPW, 5 alc.; Mt. Luboñ, 27.08.1962, leg. A.

Wiktor, Nr 874, MPW, 1 alc.; Beskid S¹decki: Valley of

Szczawny Potok, 19.08.1961, leg. A. Wiktor, Nr 822a,

MPW, 1 alc.; Sopotnicki Stream Valley, 4.07.1962, leg.

A. Wiktor, Nr 843, MPW, 2 alc.; Pieniny Mts:

Czorsztyn, 28.08.1963, leg. A. Wiktor, Nr 1063, MPW,

4 alc.; Valley of Pieniñski Potok, 8.08.1972, leg. A.

Riedel, 10/72, MIZW, 9 s.; Mt. Sokolica, 14.08.1961,

leg. A. Wiktor, Nr 821, MPW, 2 alc.; Trzy Ska³ki near

Jarmuta massif, 9.07.1962, leg. A. Wiktor, Nr 850,

MPW, 1 alc.; Mt. Góra Zamkowa, 22.08.1961, leg. A.

Wiktor, Nr 827, MPW, 1 alc.; Valley of Ociemny Potok,

19.08.1961, leg. A. Wiktor, Nr 822b, MPW, 3 alc.;

Ligarka Glade, 8.08.1962, leg. A. Wiktor, Nr 881,

MPW, 4 alc.; Tatra Mts: Zakopane, Pod Capkami

quarry, 1914, leg. W. Poliñski, coll. Poliñski 228/37,

MIZW, 8 s.; Ma³a £¹ka Valley, 7.07.1997, ZW, 11 s.+15

alc.

Shell (Figs 195–200)

Shell dome-shaped with 5–7 tightly coiled convex

whorls, rounded at periphery. Shell height 3.4–6.6

mm, shell width 5.1–9.5 mm, height/width ratio

0.54–0.82, body whorl height 2.9–4.9 mm, relative

height of body whorl 0.62–0.89, aperture height

1.7–3.2 mm, aperture width 3.0–5.2 mm, umbilicus di

ameter 0.2–0.8 mm, umbilicus major diameter/shell

diameter ratio 0.03–0.11. Aperture with whitish or

pinkish lip inside and prominent white tooth on basal

margin. Umbilicus very narrow, punctured, sometimes

partly or entirely covered by columellar aperture mar

gin. Juveniles with dense and fine hairs, in adults hairs

easily lost. Shell horny to reddish-brown, finely stri

ated, sometimes with light band on body whorl.

Shell variation

Almost every examined population differs statisti-

cally significantly from the remaining ones in at least

one character (Figs 201–211). Some pairs of popula-

tions differ in all or nearly all the parameters exam-

ined, e.g. those from Kletno in the Sudetes, Dolina

Ma³ej £¹ki in the Tatra Mts., Alterbachgeniste in Aus-

tria. The shell height (Fig. 201), aperture width (Fig.

202) and body whorl height (Fig. 205) vary very

widely. The least variable characters are those describ

ing the shell proportions i.e. height/width ratio (Fig.

210), relative height of body whorl (Fig. 211).

Reproductive system (Figs 212–216)

Four pairs of long (ca. 2 mm) mucous glands. In

ner pair of long, well-separated dart sacs reaches out

let of mucous glands, outer pair considerably shorter.

Vagina long and cylindrical. Flagellum reaches ca.

2/3 length of straight, cylindrical epiphallus, which is

longer than straight or bent cylindrical penis.

Spermatheca duct slightly coiled. Spermatheca mas

sive, hammer-shaped, not reaching albumen gland.

Longitudinal section of vagina, penial papilla, cross-

section of penial papilla and epiphallus shown in Figs

213–216.

Ecology

T. unidentatus lives in damp montane forest habi

tats between 500 and 1,500 m a.s.l.; it is also quite fre

quent above the timberline in various habitats, up to

2,000 m a.s.l.; it is found along streams in leaf-litter,

rotting wood and, rarely, on vegetation.

156 Ma³gorzata Proæków

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 157

Figs 195–197. T. unidentatus. Specimen from Kletno,

Sudetes, Poland, ZW. Shell: 195 – apertural view, 196 –

apical view, 197 – umbilical view. Scale bar 5 mm

Figs 198–200. T. unidentatus. Specimen from Draparnaud's

collection, NHMW. Shell: 198 – apertural view, 199 – api

cal view, 200 – umbilical view. Scale bar 5 mm

158 Ma³gorzata Proæków

Fig. 201. Interpopulation variation of T. unidentaus: shell

height. Statistically significant differences (Duncan test,

p < 0.05) between the following populations: 1–2, 1–3,

1–4, 1–5, 1–6, 1–8, 2–3, 2–4, 2–5, 2–6, 2–7, 2–8, 3–4, 3–5,

3–6, 3–7, 4–7, 4–8, 5–7, 5–8, 6–7, 6–8, 7–8

Fig. 202. Interpopulation variation of T. unidentaus: shell

width. Statistically significant differences (Duncan test, p

< 0.05) between the following populations: 1–2, 1–3, 1–4,

1–5, 1–6, 1–7, 1–8, 2–3, 2–4, 2–5, 2–6, 2–7, 2–8, 3–4, 3–5,

3–6, 3–7, 3–8, 6–8, 7–8

Fig. 203. Interpopulation variation of T. unidentatus: aper-

ture height. Statistically significant differences (Duncan

test, p < 0.05) between the following populations: 1–2,

1–3, 1–4, 1–5, 1–6, 1–7, 1–8, 2–3, 2–4, 2–5, 2–6, 2–7, 2–8,

3–4, 3–5, 3–6, 3–7, 3–8

Fig. 204. Interpopulation variation of T. unidentatus: aper

ture width. Statistically significant differences (Duncan

test, p < 0.05) between the following populations: 1–2,

1–3, 1–4, 1–5, 1–6, 1–7, 1–8, 2–3, 2–4, 2–5, 2–6, 2–7, 2–8,

3–4, 3–5, 3–6, 3–7, 3–7, 3–8, 4–7, 4–8, 5–7, 6–7, 6–8, 7–8

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 159

Fig. 205. Interpopulation variation of T. unidentatus: body

whorl height. Statistically significant differences

(Duncan test, p < 0.05) between the following popula-

tions: 1–2, 1–3, 1–4, 1–5, 1–6, 1–7, 1–8, 2–3, 2–4, 2–5,

2–6, 2–7, 2–8, 3–4, 3–5, 3–6, 3–7, 4–7, 4–8, 5–8, 6–8, 7–8

Fig. 206. Interpopulation variation of T. unidentatus: shell di

ameter. Statistically significant differences (Duncan test,

p < 0.05) between the following populations: 1–2, 1–3,

1–4, 1–5, 1–6, 1–7, 1–8, 2–3, 2–4, 2–5, 2–6, 2–7, 2–8, 3–4,

3–5, 3–6, 3–7, 2–8, 7–8

Fig. 207. Interpopulation variation of T. unidentatus: umbili-

cus major diameter. Statistically significant differences

(Duncan test, p < 0.05) between the following popula-

tions: 1–3, 1–4, 1–5, 1–6, 1–7, 3–7

Fig. 208. Interpopulation variation of T. unidentatus: umbili

cus minor diameter. Statistically significant differences

(Duncan test, p < 0.05) between the following popula

tions: 1–3, 1–4, 1–5, 1–6, 1–7, 1–8, 2–7, 3–4, 3–5, 3–7,

4–5, 4–6, 4–7, 4–8, 5–7, 6–7, 7–8

Distribution (Fig. 217)

An East-Alpine and Carpathian species; known

from eastern and north-eastern Switzerland (two dis-

tribution centres: 1. Samnaun, Engadin and Val

Müstair; 2. Alpenrheintal and Nordbünden; to

Walensee, near Zürich, Thurtal, near Bodensee, the

High Rhine), Germany (Alpine regions, single locali

ties in the north to the Main River, Erzgebirge), Aus

tria (widespread), the Czech Republic and Slovakia

(Danube valley, Carpathians,

“Böhmische Masse”)

OLIÑSKI 1928, KERNEY et al. 1983, TURNER et al.

1998)

, also from Poland (Carpathians except

Bieszczady Mts, Cracow-Wieluñ Jura, Sudetes: in the

north to Srebrna Góra) (R

IEDEL 1988) and from

northern Hungary. It is also found in northern Italy

(MANGANELLI et al. 1995). Although, the species was

described from France its distribution there is doubt

ful (F

ALKNER 1990).

Remarks

Six local forms, varieties, races or subspecies were

described within unidentatus. Their identification is

sometimes very difficult. Five of the six were described

from Austria but only two of them occupy separate

range fragments and their populations are fairly easily

distinguished. The populations of the remaining three

forms are heterogeneous, which means that the race-

specific specimens are found among “typical” shells.

160 Ma³gorzata Proæków

Fig. 209. Interpopulation variation of T. unidentatus: num-

ber of whorls. Statistically significant differences

(Duncan test, p < 0.05) between the following popula-

tions: 1–3, 1–4, 1–5, 1–6, 1–7, 1–8, 2–3, 2–4, 2–5, 2–6,

2–7, 2–8, 3–6, 3–7, 4–7, 5–7, 6–7, 7–8

Fig. 210. Interpopulation variation of T. unidentatus: shell

height/width ratio. Statistically significant differences

(Duncan test, p < 0.05) between the following popula

tions: 1–6, 1–7, 2–5, 2–6, 2–7, 3–6, 3–7, 4–6, 4–7, 5–6,

5–7, 6–7, 6–8, 7–8

Fig. 211. Interpopulation variation of T. unidentatus: relative

height of body whorl. Statistically significant differences

(Duncan test, p < 0.05) between the following popula-

tions: 7–1, 7–2, 7–3, 7–4, 7–5, 7–6, 8–7

The first group includes alpestris – a very small

roundish form (shell height 3–4.5 mm, shell width

5–7 mm) from the high Alps (800–2,300 m a.s.l.), with

open umbilicus and the tooth more or less reduced,

and subtecta

– also small (shell width 5–6 mm) but

with punctured or covered umbilicus, lower spire and

convex underside; with fine, very short and regular

hairs (P

OLIÑSKI 1928, KLEMM 1974).

The second group comprises subalpestris,de

scribed from various localities in the Alps as charac

terised by a lower spire, narrower, partly covered um

bilicus, often less distinct tooth, and dense but easily

lost hairs (P

OLIÑSKI 1928), and norica – generally dis

tinguished by its larger size (shell diameter 8–10.2

mm) and the tooth developed to various extent

OLIÑSKI 1928). Shells, of approximately the same

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 161

Figs 212–216. T. unidentatus. Specimen from Kletno – Sudetes, Poland, ZW. Reproductive system: 212 – general view, 213 –

longitudinal section of vagina, 214 – penial papilla, 215 – cross-section of penial papilla, 216 – cross-section of epiphallus.

Scale bar 5 mm

Fig. 217. Distribution of T. unidentatus. Black triangles – localities of origin of the examined material

KLEMM (1954) adopted Trichia (Petasina) unidentata subtecta (Poliñski) as the valid name of a subspecies (Art. 45.6.4.1

ICZN). F

ALKNER (1990: 106, 1991) regarded it as a separate species “Rasse des Ostalpenrandes” (from “südlicher

Wienerwald bis ins Grazer Bergland”). It prefers drier and warmer places than unidentata s. str.

size as norica but with no tooth were named Fruticicola

unidentata f. anodonta, and often misidentified with

edentulus or leucozonus (K

LEMM 1974).

Finally, the shells of “natio” carpatica are more con

ical, spire dome-shaped, with the umbilicus almost

covered, the tooth variously developed, and hairs

more durable, very dense and short. The form is not

easily distinguishable from unidentatus s. str. (P

OLIÑ

SKI

1928).

In order to solve the taxonomic status of these taxa

further investigations are required, including de

tailed conchological and anatomical analyses, and

their distribution ranges.

In this study the specimens from Leobner Mauer

were determined as Trochulus unidentatus subalpestris

(Poliñski, 1928), and from Hinterseegenist as

Trochulus unidentatus norica (Poliñski, 1928). However,

the shell variation ranges of these populations overlap

with that of T. unidentatus s. str. from the remainig lo

calities (figs 201–211). Besides, the differences be

tween those populations and the remaining ones, and

among the populations of T. unidentatus s. str. are

equally statistically significant. Likewise, it was imposs-

ible to indicate any characters which would allow to

distinguish the subspecies unequivocally. Conse-

quently, it is assumed here that both names are syn-

onyms of Trochulus unidentatus (Draparnaud, 1805).

Trochulus villosulus (Rossmässler, 1838)

Helix villosula ROSSMÄSSLER 1838: 1. Terra typica:

Hungary.

Helix Pietruskyana L. P

FEIFFER, 1853: 124, no 615.

Terra typica: Ukraine: Podolia. [In Schnirkel

schnecken, Dritter Theil: 418, pl. 148, figs 11, 12.

Terra typica: Ukraine: Podolia. No title page with ref

erence on publication year, but according to W

ELTER-

SCHULTES (1999) it is 1854.]

Fruticicola (Trichia) pietruskiana f. elatior P

OLIÑSKI

1914: 28. Locus typicus: Poland: vicinity of Ojców.

Material examined

Poland: Cracow-Wieluñ Jura: Ojców National Park,

near church, 16.08.1970, leg. A. Wiktor, Nr 1722,

MPW, 5 alc.; Ojców National Park, Pr¹dnik Valley,

23.08.1973, leg. A. Wiktor, Nr 1846, MPW, 19 alc.;

Pieniny Mts.: Mt. Sokolica, 14.08.1961, leg. A. Wiktor,

Nr 821, MPW, 4 alc.; Czorsztyn, 31.08.1963, leg. A.

Wiktor, Nr 1069, MPW, 1 alc.; Œrutówka, 6.08.1961,

leg. A. Wiktor, Nr 809, MPW, 1 alc.; Valley of Pieniñski

Potok, 26.08.1961, leg. A. Wiktor, Nr 829, MPW, 12

alc.; between Rabsztyn and Homole Gorge, 9.07.1962,

leg. A. Wiktor, Nr 854, MPW, 4 alc.; Kroœcienko on

Dunajec, 11.08.1961, leg. A. Wiktor, Nr 808, MPW, 5

alc.; Bieszczady Mts: Ustrzyki Górne, 21.08.1996, ZW,

10 s.+25 alc.; Slovakia: Rosenberg [=Ruzomberok],

Èebra} a/d Waag, [=Mt. Èebra} on Váh River], coll. O.

Retowski, 41/39, MIZW, 11 s.; Vratnathal [=Vratna val

ley], Comitatus Trencsin [presently in W. Slovakia],

dr C. Brancsik, coll. O. Retowski, 41/39, MIZW, 5 s.

Shell (Figs 218–220)

Shell roundish-conical, sometimes nearly flat

tened, with 5–5.75 moderately tightly coiled whorls.

Shell height 4.5–6.0 mm, width 7.0–9.0 (10) mm;

height/width ratio 0.52–0.68, body whorl height

3.3–4.8 mm, relative height of body whorl 0.73–0.96,

aperture height 2.6–3.6 mm, aperture width 3.2–4.7

mm, umbilicus major diameter 0.9–2.0 mm, umbili

cus minor diameter 0.9–1.8 mm, umbilicus major di

ameter/shell diameter ratio 0.13–0.22. Aperture with

weakly developed, white lip. Umbilicus entirely open.

Hairs long, curved and rather durable. Shell dark

brown, with no band on body whorl.

Reproductive system (Figs 221–225)

Four pairs of short mucous glands. Dart sacs long

and massive. Inner dart sacs always reach beyond

outer ones. Vagina long, its dart sac region narrows to-

ward cylindrical lower vagina. Flagellum shorter than

epiphallus, which is longer than fusiform penis.

Spermatheca duct long and thin. Spermatheca oval,

almost reaching albumen gland. Longitudinal section

of vagina, penial papilla, cross-section of penial pa-

pilla and epiphallus shown in Figs 222–225.

Ecology

T. villosulus lives in mountains up to 1,600 m a.s.l.,

also in foothills in humid habitats: scrub along

streams, wet alder forests, preferably in tussocks of

vegetation (Petasites sp.) or rarely on the ground,

among the vegetation and rotting leaves (R

IEDEL

1988).

Distribution (Fig. 226)

A West-Carpathian species; found in the Czech

Repulic and Slovakia (in the Carpathians to the Mora

vian Gate) (K

ERNEY et al. 1983, JUØIÈKOVÁ et al. 2001)

and Poland (Carpathians with their foothills: in the

north to the Cracow-Wieluñ Jura and the Sando

mierska Lowland; in the west probaly to Upper Silesia

and along the Skawa River) (R

IEDEL 1988). Isolated

localities in the Polish Eastern Sudetes have not been

confirmed (R

IEDEL 1988).

Remarks

According to P

OLIÑSKI (1924) the flagellum is as

long as the epiphallus or slightly longer, and shorter

than the penis. In all the adult specimens anatomi

cally examined, both the flagellum and the penis were

shorter than the epiphallus.

162 Ma³gorzata Proæków

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 163

Figs 218–225. T. villosulus. Specimen from Ustrzyki Górne, Bieszczady, Poland, ZW. 218–220 – shell: 218 – apertural view, 219

– umbilical view, 220 – apical view; 221–225 – reproductive system: 221 – general wiev, 222 – longitudinal section of va

gina, 223 – penial papilla, 224 – cross-section of penial papilla, 225 – cross-section of epiphallus. Scale bar 5 mm

Trochulus villosus (Draparnaud, 1805)

Helix Villosa Studer in COXE 1789: 385. [nomen

nudum]

Helix villosa D

RAPARNAUD 1805: 104, pl. VII, fig. 18.

Locus typicus: France: montagnes de Savoie.

Helix pilosa A

LTEN 1812: 46, pl. IV: fig. 7. Locus

typicus: Germany: Siebenbrunnenfeld [near Augsburg],

“an den Lechkanalen” and environs of Stätzling.*

Helix phorochaetia B

OURGUIGNAT 1864: 52, pl. VI:

figs 9–14. Locus typicus: France: dep. Isère: Grande-

Chartreuse 3 km NW Saint-Pierre-de-Chartreuse, be-

tween Saint-Bruno and Chartreusette. [see W

INTER

1990 and FALKNER et al. 2001 – Trochulus phorochaetius

(Bourguignat, 1864)]

Fruticicola villosa var. alpicola E

DER 1921: 228. Locus

typicus: Switzerland: canton Nidwalden: Bannalppass.

[see G

ITTENBERGER &NEUTEBOOM 1991, FALKNER et

al. 2001 – Trochulus alpicola (Eder, 1921)]

Material examined

Austria: Vorarlberg: Bregenz, MIZW; 7 s.; Bregenz,

coll. Klemm 69410, NHMW, 6 s.+1 alc.; Bregenzer Wald,

74368, NHMW, 5 alc.; France: Besançon, 07.1924, leg.

M. Dyrdowska, coll. Poliñski 228/37, MIZW, 3 s.; Ger

many: Donaugehölze near Ulm, Württemberg, leg.

Prinzing, MIZW, 20 s.; Ulm, Württemberg, MIZW, 6 s.;

Sigmaringen a/Donau, Württemberg, coll. Geyer

47014, NHMW, 8 s.; Güzburg, Bavaria, coll. O. Retowski

41/39, MIZW, 3 s.; Switzerland: Schindellegi, cant.

Schwyz, coll. Edlauer 15611, NHMW, 8 s.; Ragatz [=Bad

Ragaz], cant. St. Gallen, MPW, 23 s.

Shell (Figs 227–229)

Shell flattened, with low spire and blunt apex,

5.5–6.0 convex whorls with deep suture. Shell height

6.0–8.0 mm (most often 7.0–7.5 mm), shell width

11.0–14.0 mm, height/width ratio 0.5–0.65, body

whorl height 5.6–6.8 mm, relative height of body

whorl 0.83–0.94, aperture height 3.6–4.8 mm, aper-

ture width 5.0–7.0 mm, umbilicus major diameter

2.1–3.2 mm, umbilicus minor diameter 1.9–2.7 mm,

umbilicus major diameter/shell diameter ratio

0.16–0.23. Aperture rounded with weakly developed lip.

Umbilicus wide and deep. Hairs long, up to 1.5 mm,

and rather sparce. Shell pale yellowish to reddish-brown,

body whorl slightly descending, without band.

Reproductive system (Fig. 230)

Four pairs of short mucous glands. Tips of inner

dart sacs reach beyond outer ones. Lower vagina ex

panded in dart sac region and narrowing toward long

genital atrium. Flagellum slightly shorter than

epiphallus, which is shorter than cylindical penis.

Spermatheca duct straight. Spermatheca small, oval,

reaching ca. 1/2 spermoviduct length.

Ecology

T. villosus lives in shaded and humid montane habi

tats, usually in forests, also alpine pastures; its occur

rence depends on high air humidity (in the Alps usu

ally found between 500 and 2,000 m a.s.l.).

Distribution (Fig. 231)

A North-West-Alpine species; known from eastern

France (Vosges Mts, Jura, Alps), Switzerland (Jura,

northern Alps, Mittelland) and southern Germany

(from the upper Rhine River valley to south-eastern

Bavaria; an isolated locality in Zippendorf near

Schwerin (Mecklenburg)) (K

ERNEY et al. 1983,

RAUSP 1952); found also in western Austria (Vo

rarlberg and northern Tirol) (K

ERNEY et al. 1983).

164 Ma³gorzata Proæków

Fig. 226. Distribution of T. villosulus. Black triangles – localities of origin of the examined material

Remarks

The taxonomic status of alpicola Eder, 1921 has re

mained controversial. Some authors (G

ITTENBERGER

&NEUTEBOOM 1991, FALKNER et al. 2001) regard it as

a distinct species, whereas others (T

URNER et al. 1998)

consider it to be a small alpine form of T. villosus. Its re

productive system does not differ from that of typical

T. villosus, which was already shown by E

DER (1921) in

his original work. Moreover, P

FENNINGER et al. (2005)

found a comparatively small genetic distance between

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 165

Figs 227–229. T. villosus. Specimen from Ragatz, Switzerland,

MPW. Shell: 227 – apertural view, 228 – apical view, 229 –

umbilical view. Scale bar 5 mm

Fig. 230. T. villosus. Specimen from Bregenz, Vorarlberg, Aus-

tria, coll. Klemm 69410, NHMW. Reproductive system.

Scale bar 5 mm

Fig. 231. Distribution of T. villosus. Black triangles – localities

of origin of the examined material

T. villosus and T. alpicolus, indicating a questionable

specific distinction between these two taxa.

Another taxon requiring taxonomic clarification is

phorochaetia Bourguignat, 1864, which was regarded as

a distinct species by W

INTER (1990) and FALKNER et

al. (2001). According to W

INTER (1990) its shell close

ly resembles either T. sericeus (Draparnaud, 1801) or

T. plebeius (Draparnaud, 1805), whereas its genitalia

are similar to the figure and measurements of T.

‘sericea’ given by K

LOETI-HAUSER (1920).

Nomen dubium

Trochulus waldemari (Wagner, 1912)

Fruticicola waldemari WAGNER 1912: 250. Locus

typcus: Bosnia and Herzegovina: near Sarajevo, Jajce

and Boèac near Banja Luka.

No specimens were available; no original material

was found in the collections of the two institutions

that keep Wagner’s material: Naturhistorisches Mu

seum Wien or Museum and Institute of Zoology, Pol

ish Academy of Science in Warsaw. According to W

NER’

s (1912) description and later paper of MAASSEN

(1985), it is impossible to establish which species they

are related to. The taxonomic status of T. waldemari

requires further studies.

PHYLOGENETIC ANALYSIS

CHARACTER ANALYSIS

Since it is difficult to specify the sister group of

Trochulus, the following genera of the Trichiinae

sensu Shileyko 1978b, were adopted as out-groups:

Nanaja Shileyko, 1978, Odontotrema Lindholm, 1927,

Leucozonella Lindholm, 1927, Hygrohelicopsis Shileyko,

1977, Teberdinia Shileyko, 1977, Kokotschashvilia Hu-

dec et Lezhava, 1969, Caucasigena Lindholm, 1927

and Anoplitella Lindholm, 1929. The Helicoidea seem

to have undergone much parallel evolution, both in

their shell characters and anatomy. Within numerous

genera, otherwise well-defined by their synapomor

phies, the same or similar structures have been ac

quired or reduced, so that intermediate stages of the

evolution are similar as well (for references and char

acter review see: S

HILEYKO 1978a, b). Consequently,

only some of the analysed characters could be polar

ised unequivocally.

The following shell and reproductive system char

acters were used in the phylogenetic analysis (num

bers correspond with the numbers in the character

matrix – Table 1).

1. Shell colour: body whorl banded (0) or mono

chrome (1). In the outgroups and other members

of the Hygromiidae banded shells are common.

Thus, this state, even when expressed only in

some individuals of the species, was regarded as

plesiomorphic.

2. Lip: distinct, well developed (0) or weakly devel

oped (1). In the outgroups the lip is mostly well

developed, often stronger on the basal aperture

margin, and sometimes provided with callus.

3. Hairs: absent in adults (0) or present in adults

(1). In outgroups short hairs, easily lost in adults

are common. Only exceptionally, in phylogeneti

cally remote genera, such as Helicodonta Férussac

1821, Causa Shileyko, 1971 and Isognomostoma

Fitzinger, 1833, hairs are preserved in adults.

4. Length of hairs: short or absent (0); long (1) or

very long (up to 1.5 mm) (2). Hairs in the

outgroups are short. The character was treated as

ordered and additively coded.

5. Umbilicus: wide open (0); or partly or entirely

covered, punctured (1). In the outgroups the um-

bilicus is wide or narrow, but at least a part of the

previous whorl is always visible; the umbilicus is

never punctured.

6. Penis shape: fusiform (00); cylindrical (10) or con

ical (01). The penis in the outgroups is fusiform.

7. Penis/epiphallus length ratio: penis as long as or

slightly shorter than epiphallus (0) or penis lon

ger than epiphallus (1). In the outgroups the pe

nis is as long as epiphallus or slightly shorter.

8. Flagellum/epiphallus length ratio: flagellum

shorter than epiphallus (00); flagellum slightly lon

ger than or as long as epiphallus (10) or flagellum

1.5–2× longer than epihallus (01). In the

outgroups the flagellum is shorter than epiphallus.

9. Penial papilla lumen (cross-section): simple (0);

complicated as a result of division in 2–3 parts (1)

or very complicated, divided in more than 3 parts

(2). All three character states are present in the

outgroups. Initially two alternative polarizations

were applied, one of which was chosen because it

yielded a more parsimonious cladogram. The

character was regarded as ordered character and

coded additively.

10. Epiphallus lumen: low folds (00); few folds of me

dium height + two high folds (10) or high folds

(01). In the outgroups the situation with some

low and two higer folds is common.

166 Ma³gorzata Proæków

11. Number and position of mucous glands: eight

glands grouped in pairs (1); four glands not

grouped in pairs (0) or six glands not grouped in

pairs (0). In the outgroups the mucous glands are

clearly grouped (two groups of 2–3) and tend to

branch secondarily to various extent. This state

may have given origin to the other two states

through division or reduction of some compo

nents; 4 or 6 mucous glands are autapomorphies

in biconicus and suberectus, respectively, and they

were excluded from the phylogenetic analysis.

12. Length of mucous glands: short (0) or long (1).

The mucous glands in the outgroups are long.

13. Dart sacs: inner and outer dart sacs of approxi

mately equal length (00); inner dart sacs slightly

longer than outer ones (10); inner dart sacs much

longer than outer ones (01); dart sacs fused (02).

In the outgroups the inner dart sacs are longer

than the outer ones. The character was treated as

partly ordered.

14. Section from outlet of mucous glands to tips of in

ner dart sacs: short or absent (0) or long (1). In

the outgroups this section is short or absent.

15. Shape of vagina: cylindrical (0) or narrowing to-

ward genital atrium (1). In the outgroups the va-

gina is cylindrical or slightly narrowing towards

the genital atrium.

16. Internal structure of vagina: two plicae inside

(01); three plicae inside (10) or four plicae in-

side, arranged in pairs (00). In the outgroups in

which plicae are present, they are arranged in

pairs i.e. the outlet of the dart sacs, on both sides,

is flanked by plicae.

17. Shape of spermatheca: oval (0); roundish (10) or

different shape (01). In the outgroups the sper

matheca is oval.

18. Length of spermatheca duct: short (0) or long (1).

In the outgroups the spermatheca duct is short.

RELATIONSHIPS AND CLASSIFICATION

PAUP programme (version 4.0b10 for Windows)

yielded 96 equally parsimonious trees; the strict con

sensus procedure resulted in the cladogram (Fig. 232)

81 steps long, CI = 0.33, RI = 0.49, RC = 0.16. The

cladogram shows numerous polytomies. Both this and

the relatively low values of the CI, RI and RC indices re

sult from the characters being distributed in such a way

that only few of them characterise consistent species

groups, and from many instances when character states

for the species were unknown (cf. Table 1). Con

firming or rejecting the phylogeny tentatively pro

posed here would require complete knowledge of cha

racter states for all taxa and support from molecular

studies. Nevertheless the cladogram makes it possible

to question some classifications proposed in the past.

None of the previous authors included in their

classifications all or even most of the species then

known. The subgenera in which W

AGNER (1915)

placed members of Trochulus in its present sense were:

Fruticicola s. str. and Perforatella Schlüter, 1838, includ

ing erjaveci, villosa, villosula, hispida, caelata, lubomirskii,

and unidentata, edentula, leucozona, lurida and bielzi,re

spectively. In the cladogram species of none of these

groups form monophyletic units; they do not seem to

be closer related within the group than with the spe

cies from outside the group. Some members of W

NER

’s Fruticicola s. str. are close to the common ances

tor, some are interspersed with species included in his

Perforatella.P

OLIÑSKI’s (1928) sections, correspond

ing to subgenera, comprised the following species:

Petasina Beck, 1847: unidentata, Filicinella Poliñski,

1928: filicina, leucozona and bielzi, Edentiella Poliñski,

1928: edentula and bakowskii. In the cladogram in Fig.

232 unidentata, representing the monotypic Petasina,

is the sister group to bielzi which was included in

Filicinella, the other two members of which form the

sister group to lurida (not considered by P

OLIÑSKI

1928) and then to edentula and bakowskii. Only the last

two species, which P

OLIÑSKI classified within

Edentiella, are closely related in the cladogram (sister

species). From phylogenetic point of view the section

Filicinella would not be monophyletic. According to

HILEYKO (1978a, b) members of Trochulus and

Petasina should be placed in the following genera:

Plicuteria Shileyko, 1978 (lubomirskii), Trichia

Hartmann, 1840, with subgenera Petasina Beck, 1847

(unidentata)andTrichia s. str. (hispida, villosula,

striolata), and Edentiella Poliñski, 1924 (bakowskii).

These genera and subgenera were defined on the ba-

sis of certain conchological and genital characters,

but their significance was not specified. The species of

the only non-monotypic taxon given by the author do

not form a monophyletic group in the cladogram.

Special attention should be paid to the monotypic ge

nus Plicuteria. In the original version of his classifica

tion (S

HILEYKO 1978a) the author defined it as having

“vaginal plicae divided into rows of folds, flagellum

considerably shorter than epiphallus, spermatheca

duct very short and spermatheca very massive”. In the

English edition of the treatise (S

HILEYKO 1978b) he

pointed out that “regular transverse prismatic folds

that form a dense pattern on the internal wall of the

vagina” were unique to the genus Plicuteria, and not

found in any other species (cf. S

HILEYKO 1978a: 204,

fig. 215, 1978b: 31, fig. 107). Examining nine speci

mens of T. lubomirskii, in none of them did I find such

a pattern of folds on the vaginal walls; their structure

did not differ from the corresponding structures in

other members of Trochulus (cf. Fig. 136). Reasons for

describing a non-existent character by S

HILEYKO

(1978b) are uknown; perhaps it was a mistake (mis

placed labels, figures, etc.). T. lubomirskii, like T. uni

dentatus and bakowskii, included by S

HILEYKO (1978a,

b) in other, monotypic, genera and subgenera do not

depart from the remaining members of Trochulus.

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 167

In the cladogram (Fig. 232), despite it being not

completely resolved, two distinct monophyletic

groups of species can be distinguished, the villosus

group including villosus, czarnohoricus, unidentatus and

bielzi, and the striolatus group, with striolatus, graminico

lus, montanus, caelatus, hispidus, suberectus, edentulus,

bakowskii, luridus, filicinus and leucozonus. These two

groups and erjaveci, biconicus, villosulus, clandestinus,

168 Ma³gorzata Proæków

Fig. 232. Cladogram of Trochulus

lubomirskii and piccardi, form a phylogenetic bush.

The monophyletic groups appearing in the

cladogram (Fig. 232) are not compatible with any of

the proposed divisions. At present it is impossible to

attempt any subgeneric classification; any such at

tempt would result in two (supposedly) monophyletic

groups, viz. villosus group and striolatus group being

distinguished as subgenera, plus either a paraphyletic

subgenus including all the remaining species, or six

monotypic subgenera. Since anyway the genus in

cludes only about 20 species of rather small diversity

of characters, distinguishing any subgenera does not

seem justified.

DISTRIBUTION AND EVOLUTION

Discounting the North African species of very

doubtful relationship with the genus (see Introduc

tion), most members of Trochulus inhabit the Alps and

the Carpathians (Fig. 12). The only widely distributed

species is T. hispidus, widespread in Western, Central

and Eastern Europe, and partly Northern Europe. T.

striolatus is the second most widespread species. It in

habits the British Isles, northern France, Switzerland

and Austria, the Netherlands, southern Germany and

south-western Slovakia. The remaining species are

endemics with limited ranges and can be divided into

four grous: 1. Alpine group, with distribution re

stricted to the Alps or their parts – 9 species

(T. suberecus, T. villosus, T. biconicus, T. caelatus, T.

edentulus, T. clandestinus, T. graminicolus, T. montanus,

T. piccardi), 2. Carpathian group, distributed in the

Carpathians or their parts – 5 species (T. villosulus, T.

bakowskii, T. bielzi, T. lubomirskii, T. czarnohoricus), 3. Al

pine-Carpathian group – 1 species (T. unidentatus)

and 4. East-Alpine-Dinaric group, comprising species

widespread in the Eastern Alps and/or in the Dinaric

Alps – 4 species (T. erjaveci, T. filicinus, T. leucozonus, T.

luridus). Like the previously proposed classifications

within the genus, the distributional groups do not

match the phylogenetic lineages in the cladogram (cf.

Fig. 232).

The number of species within particular parts of

the range points to the Alpine-Carpathian area as the

diversity centre of the genus. Superimposing the dis

tribution ranges on the cladogram (Fig. 232) reveals

that often distribution changes must have been ac

companied by speciation events, since the members

of the two main monophyletic groups are encoun

tered in a few (2–3) distribution areas. In this context

the position of sister-species pairs is noteworthy. In

the cladogram (Fig. 232) there are four such pairs:

unidentatus/bielzi, hispidus/suberectus, edentulus/ba

kowskii, and filicinus/leucozonus. Only one of such

pairs (edentulus/bakowskii) is vicariating.

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 169

Table 1. Genus Trochulus – character matrix. Hyphens represent missing data

Taxon/character 123456789101112131415161718

erjaveci

0000000000–––00000–000

biconicus

100000001000001000001000

czarnohoricus

11101–––––––––––––

hispidus

001000001000110100001000

suberectus

001000111000100100001000

villosulus

111100000000110000000001

villosus

1112010100––100001–000

striolatus

000000001000110000110100

montanus

0000000110––100000–000

caelatus

000000011000010000010000

clandestinus

010000100020110000010000

graminicolus

0000010010––100010–000

lubomirskii

110000000001010000010010

unidentatus

001011000001010010001010

edentulus

001010000021011100010001

filicinus

001011010110111001110000

leucozonus

000011010110111001101000

luridus

001010011010111101110000

bakowskii

001010110111011100010001

bielzi

001011000121010000101000

piccardi

1000000000––000200–000

From the point of view evolution and distribu

tional changes, phylogenetic position of the two most

widespread species hispidus and striolatus is relevant.

The plesiomorphic nature of their characters could

point to their previously wider distribution, with later

reduction of the distribution ranges and the origin of

endemic species, whereas their position in the termi

nal parts of the cladogram could suggest the evol

ution within a limited area, with later spreading of few

species. Meanwhile in the cladogram (Fig. 232) the

two species belong to the striolatus group, of which

striolatus (with graminicolus) is the most plesiomorphic

member, whereas hispidus is a member of a more de

rived cluster in the same group. Their position and

the fact that the evolution of the genus most likely

started in the Alpine area, suggests an independent

expansion of their ranges.

ACKNOWLEDGEMENTS

I wish to express my sincere gratitude to Prof. dr.

hab. B

EATA M. POKRYSZKO, the supervisor of my the

sis, for her comments and suggestions during the

preparation of the manuscript, and improving the

English text. For the loan of material I am grateful to

Prof. dr. hab. A

DOLF RIEDEL (MIZW), Prof. dr. hab.

NDRZEJ PIECHOCKI (U£), Dr. hab. EL¯BIETA

KORALEWSKA-BATURA (UAM), Dr. MARGRET GOSTELI

(NHMB) and MSc. ANITA ESCHNER (NHMW).

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INDEX OF TAXONOMIC NAMES

abludens ....................146

Alixae .....................129

alpestris ..................155, 161

Alpicola ..................164, 165

altenana ....................146

anodonta ...............155, 156, 162

Ataxiaca ....................129

Axonana ....................128

badiella ....................128

bakowskii . . . . . 105, 106, 107, 108, 110–112, 167,

168, 169

Barcelonnettensis ................129

Beaudouini ...................129

Bellovacina ...................128

biconica ....................112

biconicus ........107, 108, 109, 112–114, 168

bielzi . . . . 104, 106, 107, 108, 110, 114–116, 125,

167, 168, 169

blaui .....................124

bosnensis ....................125

britannica ...................146

caelata ..................116, 167

caelatus . 104, 106, 107, 108, 109, 116–118, 168, 169

carpatica .................155, 156

cincta .....................124

clandestina ...................118

clandestinus ......106, 108, 109, 118–119, 168

Clessini ....................140

Cobresiana ................121, 155

Cochlea ....................146

coelata .....................116

coelomphala ..................116

concinna ....................128

concinnus ..............129, 135–137

corrugata .................116, 118

Cularensis ...................129

czarnohorica ..................120

czarnohoricus ........108, 109, 120, 168, 169

danubialis ............146, 147, 151, 152

depilata ..................121, 128

dolopida .................138, 140

Drunasiana ..................129

Dubisiana ...................144

Duesmensis ...................129

Edentiella ..............102, 103, 167

edentula ..................121, 167

edentulus .....104, 106, 107, 108, 110, 121–124,

162, 168, 169

elatior .....................162

Elaverana ...................128

erjaveci . . . 104, 105, 106, 107, 108, 110, 124–125,

138, 140, 167, 168

euconus ..................114, 116

Euomphalia ..................124

filicina .....................125

Filicinella ..............102, 103, 167

filicinus .....106, 107, 108, 110, 125–127, 167,

168, 169

floerickei ....................124

foeni ......................129

Glischrus .................116, 143

globulina ....................114

glypta .....................116

Goossensi ....................128

graminicola ..................127

graminicolus ......106, 107, 108, 109, 127–128,

168, 169, 170

Gratianopolitana ................128

Hajlensis ....................124

haueri .....................124

Helicella ....................128

helvetica ..................121, 123

heteromorpha ...............138, 140

Hirci .....................124

174 Ma³gorzata Proæków

hispida . . . . . . . . . . . . . 102, 103, 128, 167

hispidella ...................129

hispidosa ....................128

hispidus .....103, 104, 105, 106, 107, 108, 109,

128–137, 152, 153, 168, 169

Hygromia ...................146

Hypsellina ...................129

juvavensis ..............146, 152, 153

Latiniacensis ..................128

latiscensis ...................129

leptolasia ....................124

leucozona .................136, 167

leucozonus . . . . 104, 106, 107, 108, 110, 138–140,

162, 167, 168, 169

liberta .....................128

limnifera .................121, 123

Lorteti .....................121

lubomirskii . . . . 104, 105, 107, 108, 110, 140–141,

152, 167, 169

lurida ...................141, 167

luridus . . . . 104, 106, 107, 108, 110, 141–143, 168

Matronica ...................128

microgyra ...................129

monodon ....................155

montana .................143, 146

montanus . . . . 106, 107, 108, 109, 118, 143–145,

168, 169

montigena ...................129

nana .....................128

Niverniaca ...................129

norica ................155, 161, 162

oreinos .....................124

Orzeszkoi ....................129

osoria .....................124

ovirensis .................138, 140

Pascali .....................143

Petasina . . . . . . . . . . . . . 102, 103, 155, 167

Perforatella ................102, 167

phorochaetia ..................164

piccardi . . . . . . . . 101, 107, 108, 109, 146, 169

Pictavica ....................129

Pietruskyana ..................162

pilosa .....................164

plebeium ....................128

plebeius ........128, 129, 130, 135–137, 166

plebicola ....................143

Plicuteria ...............102, 103, 167

Prevostina ...................128

Putonii ....................118

rufescens ....................146

rufescentella ..................146

Salinae ....................129

saporosa ....................128

Sarinica ....................129

Segusiana ...................129

separanda ...................141

sericea .....................128

sericeus ...........128, 135, 152, 155, 166

Steneligma ...................128

striolata ...............146, 152, 167

striolatus .....104, 106, 107, 108, 109, 118, 145,

146–153, 168, 169, 170

styriaca ..................125, 127

subalpestris ...........155, 156, 161, 162

subbadiella ...................128

suberecta .................153, 161

suberectus .......106, 107, 108, 109, 153–155,

168, 169

subleucozona ................121, 123

submontana ..................143

subniverniaca .................129

subtecta ....................155

suevica .....................121

syrmiensis ...................124

unidens ....................155

unidentata ..............121, 155, 167

unidentatus . . . 104, 106, 107, 108, 110, 155–162,

168, 169

urbana ....................128

Vendeana ...................128

Vendoperanensis ................128

ventricosa ...................155

villosa ..................164, 167

villosula .................162, 167

villosulus .......104, 105, 107, 108, 109, 152,

162–163, 168, 169

villosus ..........104, 105, 107, 108, 110,

164–166, 168

Vocoutiana ...................129

waldemari .................109, 166

Received: May 15th, 2009

Accepted: July 20th, 2009

The genus Trochulus Chemnitz, 1786 – a taxonomic revision 175

On the verge of extinction - revision of a highly endangered Swiss alpine snail with description of a new genus, Raeticella gen. nov. (Gastropoda, Eupulmonata, Hygromiidae)

Article

Full-text available

Jun 2022

The phylogenetic status of the alpine land snail Fruticicola biconica has remained questionable since it was described by Eder in 1917. Considered a microendemic species from mountain tops in Central Switzerland, the shell is specially adapted for life under stones. Herein, we show via molecular and anatomical investigations that F. biconica neither belongs to the land snail genus Trochulus, nor to any other genus within Trochulini, but rather warrants placement within the newly established genus Raeticella Kneubühler, Baggenstos & Neubert, 2022. Phylogenetic analyses reveal that R. biconica is clearly separated from Trochulus. These findings are supported by morphological investigations of the shell and genitalia.

Speciation in sympatric species of land snails from the genus Trochulus (Gastropoda, Hygromiidae)

Article

Full-text available

Jan 2021
ZOOL SCR

The identification and designation of land snail species in the genus Trochulus on the basis of shell characteristics are problematic because of their great phenotypic plasticity. Some genetic analyses have proved inconclusive, with much variation within populations and apparent gene flow among them. We examined this issue by morphometric and molecular approaches on the morphologically similar species T. coelomphala, T. hispidus and T. striolatus, co-occurring in the Alpenvorland of Germany. While these species differed in shell and reproductive system morphology, there were forms that turned out intermediate in shell characters between T. coelomphala and T. hispidus but had genital morphology similar to T. coelomphala. Phylogenetic analysis, however, showed that these forms clustered neither with T. coelomphala nor T. hispidus but are sister to T. striolatus from the same region, which suggests that they evolved by way of sympatric speciation. Further, these analyses suggest that T. coelomphala diverged within T. hispidus; a crossing experiment indicated that they were interfertile. Expanding the study to include all available Trochulus sequences enabled us to infer evolutionary relationships between them and showed that T. hispidus is polyphyletic. Some Trochulus samples of one nominal species were grouped within others. The combination of phenotypic plasticity and possible mitochondrial DNA introgression illustrates the complex nature of evolutionary processes and the need for caution in the application of traditional taxonomic practice.

Temporal variation in climatic factors influences phenotypic diversity of Trochulus land snails

Article

Full-text available

Jul 2022

Organisms with limited dispersal capabilities should show phenotypic plasticity in situ to keep pace with environmental changes. Therefore, to study the influence of environmental variation on the phenotypic diversity, we chose land snails, Trochulus hispidus and T. sericeus, characterized by high population variability. We performed long-term field studies as well as laboratory and common garden experiments, which revealed that temporal environmental changes generate visible variation in shell size and shape of these snails. Many shell measurements of T. hispidus varied significantly with temperature and humidity in individual years. According to this, the first generation of T. hispidus, bred in controlled laboratory conditions, became significantly different in higher spire and narrower umbilicus from its wild parents. Interestingly, offspring produced by this generation and transplanted to wild conditions returned to the ‘wild’ flat and wide-umbilicated shell shape. Moreover, initially different species T. hispidus and T. sericeus transferred into common environment conditions revealed rapid and convergent shell modifications within one generation. Such morphological flexibility and high genetic variation can be evolutionarily favored, when the environment is heterogeneous in time. The impact of climate change on the shell morphometry can lead to incorrect taxonomic classification or delimitation of artificial taxa in land snails. These findings have also important implications in the context of changing climate and environment.

ANNOTATED LIST OF THE NON-MARINE MOLLUSCA OF BRITAIN AND IRELAND, 2020

Preprint

Full-text available

Sep 2020

A revised and updated list of non-marine Mollusca in Britain and Ireland based on Anderson (2005). Changes to Anderson (2005) are predicated upon recently published molecular studies.

Genome‐wide nuclear data confirm two species in the Alpine endemic land snail Noricella oreinos s.l. (Gastropoda, Hygromiidae)

Article

Full-text available

Jan 2020

The Austrian endemic land snail species Noricella oreinos (formerly Trochulus oreinos) occurs in the Northeastern Calcareous Alps at high elevations. Two morphologically highly similar subspecies N. o. oreinos and N. o. scheerpeltzi have been described. First analyses of mitochondrial and nuclear marker sequences indicated a high genetic divergence between them. In the present study, we aimed to assess gene flow between the two subspecies which should help to re-evaluate their taxonomic status. Sequence data and amplified fragment length polymorphism (AFLP) markers of 255 Noricella specimens covering the whole distribution range were analyzed. A clear geographic separation was found within the potential contact zone, the Haller Mauern mountain range. Samples of all western sites were part of the clade representing N. o. scheerpeltzi and almost all samples from the eastern sites clustered with N. o. oreinos. However, within two sampling sites of the eastern Haller Mauern, a few individuals possessed a COI sequence matching the N. o. oreinos clade whereas at the ITS2 locus they were heterozygous possessing the alleles of both taxa. Contrary to the ITS2 results indicating historical and/or ongoing hybridization, AFLP analyses of 202 individuals confirmed a clear separation of the two taxa congruent with the mitochondrial data. Although they occur on the same mountain range without any physical barrier, no indication of ongoing gene flow between the two taxa was found. Thus, we conclude that the two taxa are separate species N. oreinos and N. scheerpeltzi.

Case 3839 – Helix unidentata Draparnaud, 1805 (currently Petasina unidentata; Gastropoda, Hygromiidae): proposed conservation of the specific name and proposed fixation of the type species of the genus-group name Petasina Beck, 1847

Article

Dec 2021

Bernhard Hausdorf

This application has two purposes. The first, under Article 23.9.5 of the Code, is to conserve the specific name of the binomen Helix unidentata Draparnaud, 1805, a junior primary homonym of Helix unidentata Holten, 1802, by ruling that the former is not thereby invalid. These names are currently in use for two different species of stylommatophoran snails that have not been treated as congeneric since 1898. Helix unidentata Draparnaud, 1805 has been considered the type species of Petasina Beck, 1847 (Helicoidea, Hygromiidae,) by a majority of authors, whereas Helix unidentata Holten, 1802 is the valid name of Helix ventricosa De Cristofori & Jan, 1832, the type species of Stylodonta De Cristofori & Jan, 1832 (Acavidae, Rhytidoidea). The second purpose of this application, under Art. 81.1 of the Code, is to clarify the status of Petasina Beck, 1847 by (a) fixing Helix unidentata Draparnaud, 1805 as its type species and (b) confirming that Petasina Beck, 1847 is an available name independent of Petasia Beck, 1837. This is necessary to maintain current usage of not only Petasina Beck, 1847 but also another frequently used genus-group name, Euconulus Reinhardt, 1883 (Euconulidae, Trochomorphoidea). Finally, lectotypes are designated for Helix unidentata Draparnaud, 1805 and Helix edentula Draparnaud, 1805.

Lectotype designation with type locality restriction for Helix abludens, Locard 1888, with notes on the taxonomy of Trochulus striolatus (C. Pfeiffer, 1828) (Gastropoda: Hygromiidae)

Article

Full-text available

Oct 2021
J CONCHOL

Trochulus is a genus of European land snail with a complex and disputed taxonomy and an intricate evolutionary history. Many species and subspecies were described only on the basis of shell morphology, which is highly variable, such that the shell morphology of many species and subordinate taxa appear to overlap that of others. In resolving the phylogeny and nomenclature of the genus, it is crucial to study type material, which is often not available. Examination of Locard's material from the Muséum national d'Histoire naturelle (MNHN) in Paris (France) led to the discovery of two specimens of Helix abludens, Locard 1888. This material is considered to belong to the original lot used by Locard to define his species, which was regarded as Trochulus striolatus subsp. abludens but recently discarded. In order to definitively clarify its identity and to stabilise the name, the best-preserved specimen is selected and designated herein as a lectotype. It is fully described and illustrated. The other specimen from the same lot becomes a paralectotype. This designation allows the restriction of the type locality to Boulogne-sur-Mer in the department Pas-de-Calais in France.

Mollusques terrestres et d'eau douce des Pyrénées-Atlantiques : catalogue commenté des espèces, espèces patrimoniales, enjeux de connaissances et de conservation, bibliographie

Article

Full-text available

Jun 2020

Alain Bertrand

Mollusques continentaux Pyrénées-Atlantiques 10 juin 2020.doc

Article

Full-text available

Jun 2020

Alain Bertrand

Avec au moins 230 espèces de mollusques le département des Pyrénées-Atlantiques apparaît comme un département très riche et il semble être le département le plus riche de France métropolitaine . Soit plus du 1/3 des espèces de la faune de France continentale sur 1,4 % du territoire. Il est à noter que toutes les sous-espèces, ainsi que des « espèces » au statut taxonomique douteux ont été exclues. Dans un contexte biogéographie et une superficie comparables, la Cantabrie, située à l’ouest des Pyrénées dans le nord de l’Espagne n’en compte que 130 (RUIZ COBO ET VAZQUEZ TORO, 2019). Cette richesse spécifique trouve son origine dans plusieurs facteurs : 1. la position géographie et le contexte ; 2. biogéographique qui en découle aux confins du contact des plaques européenne et ibérique ; 3. la très grande diversité des climats, atlantique à alpin en versant nord, sub méditerranéen à alpin en versant sud ; 4. la position latitudinale et l’orientation qui a probablement contribué à créer des situations de refuge lors des glaciations du Quaternaire ; 5. pour les Hydrobiidae et Moitessieriidae décrits notamment par Boeters, on ne peut pas exclure un splittage créateur d’espèces, qui devra être confirmé/infirmé par la mise en œuvre de méthodes moléculaires, mais on ne peut pas non plus exclure la présence d’espèces inédites. Cent quatre vingt dix neuf publications de 1822 à 2020 ont été rassemblées. Les deux premières publications, qui font état d’espèces de mollusques des Pyrénées-Atlantiques datent du début du XIX -ème siècle ; FERUSSAC, 1821-1822 et LAMARCK (1822) mentionnent chacun une espèce ! Quatre malacologues, Fernand BERILLON, Paul FAGOT, Léopold De FOLIN et Alexandre MERMET on posés les bases de la connaissance de la malacologie du département au XIX siècle. La première monographie sur la malacofaune par MERMET date de 1843.L’activité des malacologues au cours de ces 200 années est marquée par deux périodes très productives, la première de 1870 à 1890, la seconde de 1980 2020. La longue période « stérile » de 1900 à 1970 trouve au moins en partie son origine dans les dégâts causés à la malacologie par les auteurs de la « Nouvelle Ecole » qui sous la « direction de Bourguignat » ont appliqué une conception très « originale » de la notion d’espèce. Dix huit pour cent des espèces sont des endémiques régionaux, pyrénéens, pyrénéo-cantabriques, etc. Ce chiffre est inférieur celui à mis en avant par l’INPN notamment. Ce dernier a été calculé sur des bases très différentes incluant des taxons non évalués avec des méthodes rigoureuses et notamment des sous-espèces qui s’avèrent n’être que des formes individuelles, des formes clinales, écologiques, etc. Onze pour cent des espèces ont été décrites depuis 1973 et d’autres sont en cours de description. Treize pour cent des espèces ont été mentionnées pour la première fois dans le département depuis 2010, 6 dans la littérature et vingt-cinq dans ce travail. Dix huit espèces mentionnées dans la littérature n’ont pas fait l’objet d’observations depuis le début du XIXème siècle. Trois espèces, au moins mentionnées dans la littérature ancienne l’ont très probablement été par erreur. Ce sont Cochlostoma septemspirale (Razoumowsky 1789), Chondrina farinesii (Des Moulins 1835), Cernuella neglecta (Draparnaud 1805). Malgré cette richesse spécifique et cette littérature abondante la malacofaune des Pyrénées-Atlantiques semble rester encore bien mal connue ! En effet : 1. moins d’un 1/3 seulement du territoire a été prospecté de manière significative ; ainsi le nord-est, bien que très anthropisé peut abriter des espèces présentes dans le Gers et qui n’ont pas été retrouvées comme Helicodonta obvoluta, Trochoidea elegans ou Chilostoma squamatinum ; 2. en altitude de vastes massifs calcaires restent très mal connus voir inconnus ; 3. les zones humides, que ce soit dans les basses vallées comme celle de l’Adour, ou bien les tourbières et bas marais sont quasiment inconnues ; 4. les riches faunes des sources et des eaux souterraines, un peu mieux connues, abritent probablement encore des espèces à décrire ; 5. les micro espèces terrestres sont largement sous prospectées ; 6. le genre Zospeum, reste à revoir car probablement étudié trop superficiellement ; 7. les limaces n’ont probablement pas encore été étudiées avec toute l’attention nécessaire à une bonne connaissance ; 8. les micro-bivalves, Sphaeriidae, n’ont pas fait l’objet de recherches spécifiques adaptées ; 9. les Cochlostomatidae et les Chondrinidae, doivent être revus avec la prospection des massifs (et pas seulement des bords de route !) et la mise en œuvre de méthodes moléculaires notamment. Huit espèces seulement ont été introduites plus ou moins récemment sont connues; 3 sont aquatiques et 5 terrestres. Potamopyrgus antipodarum se comporte comme une espèce invasive ; il peuple beaucoup des sources abritant des Bythinella et des Alzoniella. Il a également été noté dans le sous-écoulement des cours d’eau affluents du Saison en Soule où sont également présentes des espèces stygobies endémiques. Son impact sur les espèces de mollusques et d’invertébrés n’est pas documenté dans le département. En terme d’enjeux de conservation, une première analyse montre l’incohérence-inadéquation pour les espèces à statut légal (protection nationale, Protection Union Européenne) ainsi que celles avec des évaluations UICN et les connaissances disponibles. A l’opposé de nombreuses espèces qui nécessiteraient des mesures de conservation sont oubliées.

Liste de référence fonctionnelle et annotée des Mollusques continentaux (Mollusca, Gastropoda, Bivalvia) du Grand-Est (France)

Article

Full-text available

Dec 2019

Les plateformes collaboratives d’observation de la biodiversité, couplées à des référentiels taxonomiques numériques, permettent désormais de générer automatiquement des listes d’espèces et/ou atlas de répartition à diverses échelles tant temporelles que géographiques. Nous questionnons ici la fiabilité d’une telle liste d’espèces en prenant comme point de départ la pré-liste taxonomique des Mollusques de la région du Grand-Est générée à partir des données d’occurrences disponibles dans le Système d’Information sur la Nature et les Paysages (SINP) et du réferentiel associé TAXREF. Nous montrons que le taux de recouvrement entre cette pré-liste qui cite 233 taxons terminaux et celle élaborée par les auteurs de cet article est de l’ordre de 90 %. Les différences sont liées à 60,7 % à de fausses-absences, à 32,1 % à de fausses-présences et à 7,2 % à des traitements taxonomiques différents. Au final, nous documentons la présence de 241 taxons terminaux dont 156 Gastéropodes terrestres, 46 Gastéropodes aquatiques et 39 Bivalves. Nous montrons que plus de 40 % des espèces de Mollusque nécessitent une approche conjointe morpho-anatomique pour une identification fiable. C’est pourquoi, cette liste de référence dite fonctionnelle a mis l’accent, pour certains taxons, sur les critères spécifiques estimés comme stables et sur le niveau de difficulté lors de l’identification, et ce, afin de tenter d’homogénéiser l’implémentation des données d’occurrence et/ou de leur évaluation a posteriori. Nous recommandons que les données primaires enregistrées dans les bases de données collaboratives fassent référence à des spécimens déposés dans des collections publiques, aux caractères spécifiques observés et par la transmission de photographie(s) de bonnes qualités.

First find Of pomatias rivularis (Eichwald, 1829) (mollusca: pomatiidae) in croatia

Article

Full-text available

Sep 2008

Pomatias rivularis (Eichwald, 1829) specimens were collected on Bansko Brdo (or Bansko Hill, Baranya, Croatia) during spring 2007. This species proved to be new for the Croatian fauna. Populations of Pomatias elegans (O. F. Müller, 1774) were also found on Bansko Brdo, which is the only known area in Croatia where both these two species occur. The list of 40 mollusc species (35 snails, 5 bivalves) identified in the study area is presented. Among them, Ena montana (Draparnaud, 1801) was found for the first time in the NE part of the country.

Check list of the molluscs (Mollusca) of the Slovak Republic

Article

Full-text available

Mar 2009

The checklist of 245 mollusc species known so far from the Slovak Republic is presented, plus 11 species limited to greenhouses or thermal waters. Critical comments on species erroneously mentioned in recent publications from Slovakia are included.

System der Erd- und Flußschnecken der Schweiz. Mit vergleichender Aufzählung aller auch in den benachbarten Ländern, Deutschland, Frankreich und Italien sich vorfindenden Arten

Article