Cochlostoma revised: the subgenus Clessiniella Zallot et al., 2015 (Caenogastropoda, Cochlostomatidae)

Abstract

Five species of the subgenus Clessiniella of Cochlostoma (Cochlostomatidae) are recognized, viz. Cochlostoma (Clessiniella) villae (Strobel, 1851), Cochlostoma (Clessiniella) tergestinum (Westerlund, 1878), Cochlostoma (Clessiniella) waldemari (A.J. Wagner, 1897), Cochlostoma (Clessiniella) anomphale Boeckel, 1939 and Cochlostoma (Clessiniella) stelucarum sp. nov. The shells and the genitalia are described and the distributional data are summarized for all the species.

Full text

49
European Journal of Taxonomy 762: 49–95 ISSN 2118-9773
https://doi.org/10.5852/ejt.2021.762.1453 www.europeanjournaloftaxonomy.eu
2021 · Zallot E. et al.
This work is licensed under a Creative Commons Attribution License (CC BY 4.0).
Research article
urn:lsid:zoobank.org:pub:D1093CEB-7EBB-44A0-8078-8DC616B857F5
Cochlostoma revised: the subgenus Clessiniella Zallot et al., 2015
(Caenogastropoda, Cochlostomatidae)
Enrico ZALLOT 1,*, Willy DE MATTIA 2, Zoltán FEHÉR 3 & Edmund GITTENBERGER 4
1 Haagweg 29, 2681PA Monster, the Netherlands.
2 Central Research Laboratories of Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria.
3 Hungarian Natural History Museum, Baross 13, 1088 Budapest, Hungary.
4 Naturalis Biodiversity Center, P.O. Box 9517, NL-2300RA Leiden, the Netherlands.
* Corresponding author: ezallot@gmail.com
2 Email: wdemattia@gmail.com
3 Email: feher.zoltan@nhmus.hu
4 Email: egittenberger@yahoo.com
1 urn:lsid:zoobank.org:author:365E17AD-6938-4364-A526-F1BDA2663E10
2 urn:lsid:zoobank.org:author:C74A049D-9D7B-4A1F-878B-5BDFD104219F
3 urn:lsid:zoobank.org:author:E801EC76-8B1E-450B-993E-BBBE57C00EA9
4 urn:lsid:zoobank.org:author:D786C279-FC92-4D08-AF16-F79A9705E0AE
Abstract. Five species of the subgenus Clessiniella of Cochlostoma (Cochlostomatidae) are recognized,
viz. Cochlostoma (Clessiniella) villae (Strobel, 1851), Cochlostoma (Clessiniella) tergestinum
(Westerlund, 1878), Cochlostoma (Clessiniella) waldemari (A.J. Wagner, 1897), Cochlostoma
(Clessiniella) anomphale Boeckel, 1939 and Cochlostoma (Clessiniella) stelucarum sp. nov. The shells
and the genitalia are described and the distributional data are summarized for all the species.
Keywords. Taxonomy, morphology, Eastern Alps.
Zallot E., De Mattia W., Fehér Z. & Gittenberger E. 2021. Cochlostoma revised: the subgenus Clessiniella Zallot
et al., 2015 (Caenogastropoda, Cochlostomatidae). European Journal of Taxonomy 762: 49–95.
https://doi.org/10.5852/ejt.2021.762.1453
Introduction
The genus Cochlostoma Jan, 1830 includes predominantly rock-dwelling species distributed along the
mountain ranges surrounding the Mediterranean Sea. Until recently, the taxonomy of the genus was
based only on shell morphology. The lack of clear species-specic shell characters, however, makes
it extremely difcult in many cases to assign a population to a certain species. In our previous papers
(Zallot et al. 2015, 2018) we proved that the structure of the genitalia, together with the analyses of
some DNA markers, may facilitate identications while enabling a sound phylogenetic classication.
The results of this approach imply a revision of the shell-based taxonomy, which is still used in the
most recent European checklist (Welter-Schultes 2012). Whilst revising each of the subgenera of

European Journal of Taxonomy 762: 49–95 (2021)
50
Cochlostoma, we herewith analyze the subgenus Clessiniella Zallot, Groenenberg, De Mattia, Fehér &
Gittenberger, 2015, which includes a group of phylogenetically closely related species inhabiting the
eastern Alps and are characterized by a diagnostic structure of the female genitalia.
Material and methods
The sampling localities, mapped in Fig. 1, are also indicated for each species in the taxonomic section
of this paper. An approximate position is used for the pre-GPS samples. The availability of female
and male specimens from the different sampling localities was extremely various (dry or alcohol
preserved, number of male and female specimens) and therefore the level of analysis of each sample
was consequently different, as reported in Table 1. The methods for depicting the shells, obtaining
measurements and dissecting genitalia have been discussed previously in Zallot (2002: 94–97) and
Zallot et al. (2015: 64–67; 2018: 2–8). Here, we provide a description of the characters of the shell and
of the genitalia that are used for classication.
Abbreviations
Shell characters
Ap. incl. = inclination of the apertural edge in lateral view in respect to the main axis (no. 13 in
Fig. 2)
H 1st/4th = ratio between the height of the 1st and that of the 4th whorl. It is an index of the slenderness
of the shell
Ha/Ht = ratio between the height of the aperture measured along the main axis and the total height
Ht = total height: measured along the columella (no. 1 in Fig. 2)
Ht/W = ratio between the total height and the width of the basal whorl
Fig. 1. Map of the sampling localities. Blue dots = Cochlostoma (Clessiniella) villae (Strobel, 1851);
red dots = Co. (Cl.) stelucarum sp. nov.; green dots = Co. (Cl.) tergestinum (Westerlund, 1878); yellow
dots = Co. (Cl.) waldemari (A.J. Wagner, 1897); orange dots = Co. (Cl.) anomphale Boeckel, 1939.

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
51
Table 1. Number of specimens analyzed for each sampling locality. Abbreviations: FG = analysis of the female genitalia; FS = female shell measured;
MG = analysis of the male genitalia; MS = male shell measured; V = visual assessment of the species based on the shell.

European Journal of Taxonomy 762: 49–95 (2021)
52
LipW = the apertural lip, on the columellar side, enlarges and attens (no. 11 in Fig. 2). LipW
is the ratio between the projection in frontal view of the maximum width of the lip on
the columellar side and the maximum width of the aperture
LobeH = projection in frontal view of the height of the external lobe (no. 5 in Fig. 2)
Prt = number of whorls of the protoconch. The embryonic shell is recognizable by a change
of ribbing and often of the color (Fig. 3)
Prt/4th = ratio between the width of the basal whorl of the protoconch and the width of the shell
measured along the 4th suture. This ratio summarizes the relative size of the protoconch
in respect to the whole shell
Ribs/mm 1st = number of ribs per mm on the 1st whorl. The number of ribs per mm is calculated by
measuring the distance between the rst and the last of the most central ve ribs close
to the upper suture of the whorl (no. 15 in Fig. 2)
Ribs/mm 4th = number of ribs per mm on the 4th whorl. The number of ribs per mm is calculated by
measuring the distance between the rst and the last of the most central ve ribs of a
whorl, in frontal view, close to the basal suture of the whorl (no. 16 in Fig. 2)
Fig. 2. Measurements taken on the shell of Clessiniella Zallot et al. 2015. 1 = height (H); 2 = height
of the 1st whorl; 3 = height of the 4th whorl; 4 = height of the aperture (Ha); 5 = height of the external
lobe; 6 = width of the 1st whorl (W); 7 = width of the 1st suture; 8 = width of the 2nd whorl; 9 = width of
the 4th whorl; 10 = width of the aperture (Wa); 11 = maximum width of the lip on the columellar side;
12 = width of the basal whorl of the protoconch; 13 = inclination of the aperture; 14 = inclination of
the ribs; 15 = distance between the 5 central ribs of the body whorl; 16 = distance between the 5 central
ribs of the 4th whorl. The gure in the lower right corner shows how the whorls (green) and sutures
(red) are counted.

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
53
Ribs incl. = inclination of the ribs in respect to the plane orthogonal to the columellar axis (no. 14
in Fig. 2)
Smooth Prt = number of protoconch whorls, starting from the tip point that are smooth at 40×
magnication (no. 1 in Fig. 3)
W = width: measured at right angles to the columella (no. 6 in Fig. 2)
W 1st/4th = ratio between the width of the 1st and that of the 4th whorl. It is an index of the slenderness
of the shell
W round = ratio between the maximum width of the 2nd whorl and the width of the shell measured at
the 1st suture. This measure indicates the convexity of the whorl(s), a feature frequently
mentioned in the description of the taxa within Cochlostoma
Whorls = number of whorls: counted with the shell in front view. The whorls and the sutures are
counted bottom-up, so that the basal whorl is the 1st one and the suture above is the
1st suture as shown in Fig. 2.
Nominal or ordinal data
External lobe = the body whorl, approaching the aperture, widens in different way and, as
consequence, it changes the shape of the body whorl close to the aperture in
external view (Fig. 4A)
Ribs near ap. = approaching the aperture, the ribs often become weaker in comparison to their
strength on the front side of the body whorl. W = really weak ribs or smooth; I =
Fig. 3. Examples of protoconchs: the red dots mark the end of the protoconchs. A. Cochlostoma
(Clessiniella) anomphale Boeckel, 1939 (EZ0034). A1. Co. (Cl.) anomphale (EZ0034): 1 = end of the
rst smooth part of the protoconch; 2 = end of the protoconch. B. Co. (Cl.) tergestinum (Westerlund,
1878) (EZ0799). C. Co. (Cl.) villae (Strobel, 1851) (EZ0885). D. Co. (Cl.) waldemari (A.J. Wagner,
1897) (WdM6697). E. Co. (Cl.) stelucarum sp. nov. (EZ0104).

European Journal of Taxonomy 762: 49–95 (2021)
54
Fig. 4. Features of the shell of Clessiniella Zallot et al., 2015. A. Shape of the body whorl approaching
the aperture. B. Strength of the spots on the shell (0 = spotless shell to 2 = really evident spots).
C. Strength of the ribs (1 = weak ribs to 3 = really prominent ribs). D. Strength of the ribs on the
external side of the body whorl approaching the aperture (W= smooth shell; I = intermediate; S =
prominent ribs).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
55
less strong than on the front side of the body whorl; S = as strong as in the front
side (Fig. 4D)
Rib prominence = from 0 = ribless shell to 3 = really strong/prominent ribs (Fig. 4C)
Spots noticeability = from 0 = spotless shell to 2 = really noticeable spots (Figs 4B, 12). The shell
spots are areas of the shell that are darker than the background. They are usually
reddish/brown.
Female genitalia (Figs 5–6)
Club-shaped seminal receptacle: seminal receptacle with narrow basis, gradually enlarging towards the
apex where it reaches the maximum width: SS = short, club-shaped seminal receptacle; SM = club-
shaped seminal receptacle of average length; SL = long, club-shaped seminal receptacle.
Oval-shaped seminal receptacle: seminal receptacle with recognizable basis and apex, with maximum
width close to the midline; BS = short, oval-shaped seminal receptacle; BM = oval-shaped seminal
receptacle of average length; BL = long, oval-shaped seminal receptacle.
Loops set up: the visceral oviduct, between the nal loop and the seminal receptacle, is more or less
tortuous and the resulting loops are more or less clustered together (Fig. 6B): S = supercial loops, quite
distinct from one another; O = loops overlapping; C = loops tightly clustered.
Loops main position: the loops formed by the visceral oviduct can be differently arranged in relation
to the apex of the seminal receptacle (Fig. 6C): O = loops positioned over (= dorsally) the apex of the
seminal receptacle; DH = loops positioned close to the apex, on the visceral side; DM = loops positioned
roughly halfway between the apex and the nal loop.
CBC position: the pedunculus of the bursa copulatrix can connect to it in different relative positions:
from 0 = apically to 3 = ventrally to 4 = distally (Fig. 6D).
SRwidth: Fig. 6E.
SR apex position: position of the apex of the seminal receptacle relative to the body: from 0 to 0.5 =
ventral side of the body; from 0.6 to 1 = dorsal side of the body (Fig. 6G)
SR curvature: the seminal receptacle can have a portion of its apical part (the one connected to the
visceral oviduct) curved downwards (apex towards the ventral side of the body) (Fig. 6F): from 0 =
apex straight, not curved to 0.5 = half seminal receptacle curved to 1 = inverted seminal receptacle
(connection with the distal oviduct on the dorsal side of the body and connection with the visceral
oviduct on the ventral side of the body).
JUG position: the junction of the uterus gland with the copulatory channel can be more or less far from
the connection between the pedunculus of the bursa and the distal oviduct: from 0 = really close to 3 =
really far (Fig. 6H).
Loops n.: the visceral oviduct can have a different number of loops. A rough count of them gives a
measure of the complexity of the loops path (and of the length of the visceral oviduct in between the
nal loop and the apex of the seminal receptacle).
DNA sequence analysis
For DNA extraction from small pieces of tissue of ethanol preserved specimens we used the DNeasy
Blood & Tissue Kit (Qiagen) following the manufacturer’s protocol. A 309 bp long gene fragment of
the nuclear Histone H3 gene was amplied by polymerase chain reaction (PCR) using the following

European Journal of Taxonomy 762: 49–95 (2021)
56
primers: H3ColF (Colgan et al. 2000) and H3PulR (Uit de Weerd & Gittenberger 2013). For PCR,
TopTaq DNA Polymerase (Qiagen) was used. The PCR prole started with an initial denaturation for
3 min at 94°C, followed by 40 cycles of 30 s at 94°C, annealing for 30 s at 57°C and elongation for 60 s
at 72°C, terminated by a nal elongation phase of 10 min at 72°C. Sequencing (both directions) was
performed by LGC Genomics (Berlin, Germany) using the PCR primers.
Fig. 5. Genitalia in Clessiniella Zallot et al., 2015. A. Female genitalia (Cochlostoma (Clessiniella)
tergestinum (Westerlund, 1878) (WdM6998)). 1 = ovary; 2 = visceral oviduct; 3 = proximal loop;
4 = loops of the visceral oviduct; 5 = seminal receptacle; 6 = distal oviduct; 7 = pedunculus of the
bursa copulatrix; 8 = bursa copulatrix; 9 = copulatory duct; 10 = channel of the uterine gland; 11 =
uterine gland. B. Male genitalia (Co. (Cl.) tergestinum (WdM7012)). 1 = anus; 2 = fold delimiting the
sperm groove; 3 = sperm pocket; 4 = penial funnel; 5 = body spermiduct; 6 = penis with the internal
spermiduct.

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
57
Institutional abbreviations
EZ = private collection of Enrico Zallot
MIZPAS = Muzeum i Instytut Zoologii PAN, Warsaw, Poland
NHMW = Naturhistorisches Museum, Vienna, Austria
PU = Museo di Storia Naturale dell’Universita’ di Parma, Italy
RMNH = Naturalis Biodiversity Center, Leiden, the Netherlands
SMF = Naturmuseum Senckenberg, Frankfurt am Main, Germany
WDM = private collection of Willy De Mattia
Results
Morphological characters
Five species of Cochlostoma, one of which is new to science and here described, show the female genital
characters of the subgenus Clessiniella as described by Zallot et al. (2015: 82). In Table 2 we present the
measurements of the shell characters of the types and the average, minimum, maximum and standard
Fig 6. Female genital features. A. SR shape. B. Loops set up. C. Loops main position. D. CBC position.
E. SR width. F. SR curvature. G. SR apex position. H. JUG position.

European Journal of Taxonomy 762: 49–95 (2021)
58
Table 2. Shell measurements. Abbreviation: N = number of female specimens analyzed.
# whorls (count)
H (mm)
W (mm)
H/W
Ha/H
H 1st/4th
W 1st/4th
w round.
Lip W
Lobe H
Ribs incl. ( °)
Ap. Incl. (°)
Ribs/mm 1st
Ribs/mm 4th
Prt (count)
Smooth Prt (count)
W Prt/4th
Clessiniella N214 215 215 215 215 215 215 215 215 215 215 198 204 215 141 135 196
Ave rage 8.4 7.9 3.0 2.6 0. 22 2.6 1.9 1.15 0. 14 0.12 66. 5 15.7 8 7 2.2 1.5 0.61
Min 7. 2 6.1 2.5 2.3 0.18 2. 0 1.6 1.02 0.06 0.03 59.3 5.4 4 4 1.8 1.1 0.42
Max 10.3 10.8 3.8 3.0 0.27 3. 4 2.2 1.20 0.21 0.23 76.5 26.5 17 14 2. 5 1.8 0.81
StdDev 0.6 0.7 0. 2 0.1 0.02 0.2 0. 1 0. 02 0. 03 0. 03 3.3 3.9 2 2 0.1 0.1 0.08
C. (C.) anomphale paratype 1 ♀7.5 6.2 2. 56 2.43 0. 23 2.75 2.01 0. 13 0.18 0. 11 65.4 18.6 10 10 0.67 0.69
paratype 2 ♀7.2 6.2 2. 63 2.35 0. 22 2.98 2.09 0. 15 0.14 0. 10 64.8 15.7 11 90. 71 0.73
N15 15 15 15 15 15 15 15 15 15 15 14 15 15 10 10 13
Ave rage 8.4 7.8 3.0 2.6 0. 21 2.6 1.9 1.15 0. 13 0.07 67. 3 16.4 11 92.2 1.4 0.56
Min 8. 0 6. 9 2. 8 2. 5 0.19 2.2 1.7 1. 11 0.10 0.03 64.2 9.3 7 7 2.0 1.3 0.49
Max 9.1 8.5 3.2 2.7 0.22 3.0 2.1 1. 17 0.17 0.11 69. 7 20. 3 17 14 2.3 1.6 0. 65
StdDev 0.4 0.4 0. 1 0.1 0.01 0.2 0. 1 0. 02 0. 02 0. 02 2.1 3.5 3 2 0.1 0.1 0.05
C. (C.) stelucarum holotype ♀10.0 8.0 3.20
paratype ♀10.3 8. 7 3.09 2. 81 0.19 2.47 1. 67 0.17 0. 14 0.07 69.3 17.7 10 60.64 0.43
paratype ♂9.1 7. 0 3.00
N14 14 14 14 14 14 14 14 14 14 14 11 14 14 8 7 14
Ave rage 9.6 8.4 3.1 2.7 0. 20 2.3 1.8 1.14 0. 12 0.08 70. 0 18.4 8 7 2.2 1.5 0. 48
Min 8. 4 7. 4 2. 9 2. 5 0.18 2.0 1.6 1. 11 0.08 0.04 66.5 15.2 5 5 2.1 1.3 0.42
Max 10.3 9.5 3.3 2.8 0.22 2.6 1.9 1. 17 0.14 0.12 75. 7 20. 2 11 82.3 1.8 0.58
StdDev 0.6 0.8 0. 2 0.1 0.01 0.2 0. 1 0. 02 0. 02 0. 02 2.2 1.7 2 1 0.1 0.2 0.06
C. (C.) tergestinum syntype ♀9.1 7. 8 2.83 2.75 0.22 2.70 1. 88 0.13 0.16 0.14 68.0 13.9 6 4 0.65 0. 54
syntype ♂8.2 7.4 2. 98 2.49 0.23 2.66 1.55 0. 13 0.12 0.12 66.8 12.6 6 5 0.77 0.52
N108 108 108 108 108 108 108 108 108 108 108 99 102 108 69 67 93
Ave rage 8.3 7.8 3.0 2.6 0.22 2.7 2.0 1.15 0.14 0.12 65.5 16.9 8 7 2.2 1.5 0. 63
Min 7. 3 6. 1 2. 5 2. 3 0.18 2.3 1.7 1. 02 0.06 0.05 59.3 5.9 5 4 1.8 1.1 0.51
Max 10.0 9.6 3.4 2.9 0.27 3.2 2.2 1. 20 0.20 0.23 70. 2 26. 5 16 13 2.5 1.7 0. 76
StdDev 0.4 0.7 0. 2 0.1 0.02 0.2 0. 1 0. 02 0. 03 0. 04 2.3 3.4 2 1 0.1 0.1 0.06
C. (C.) villae syntype ♀9.2 10. 1 3.43 2.94 0.19 1.8 1. 62 0.14 0.11 0. 09 75.6 18.18 5 4 - -
N23 24 24 24 24 24 24 24 24 24 24 22 23 24 16 15 24
Ave rage 8.8 8.8 3.3 2.7 0. 22 2.6 1.8 1.13 0. 13 0.10 72. 1 15.7 6 5 2.3 1.4 0. 53
Min 8. 2 7. 1 2. 9 2. 5 0.20 2.2 1.7 1. 09 0.08 0.06 64.6 6.5 4 4 2.0 1.3 0.43
Max 9.4 10.8 3. 8 3. 0 0.24 3.2 2.0 1.19 0.18 0. 15 76.5 21.7 9 8 2.4 1.7 0.63
StdDev 0.4 0.9 0. 3 0.1 0.01 0.2 0. 1 0. 02 0. 03 0. 02 2.6 3.2 1 1 0.1 0.2 0.05
C. (C.) waldemari syntype ♀7.5 7.3 2.72 2. 68 0.26 2. 93 1.93 0.15 0. 16 0.15 64. 7 4.9 811 0.72 0.70
N54 54 54 54 54 54 54 54 54 54 54 52 50 54 38 36 52
Ave rage 8.0 7.8 3.1 2.5 0. 23 2.7 2.0 1.14 0. 15 0.13 65. 0 12.9 9 9 2.2 1.4 0. 67
Min 7. 2 7. 0 2. 8 2. 4 0.21 2.3 1.7 1. 10 0.10 0.06 60.0 5.4 5 5 2.0 1.1 0.55
Max 8.6 8.8 3.4 2.7 0.27 3.4 2.2 1. 19 0.21 0.18 69. 7 20. 4 14 14 2.5 1.7 0. 81
StdDev 0.3 0.4 0. 1 0.1 0.01 0.2 0. 1 0. 02 0. 03 0. 02 2.4 3.9 2 2 0.1 0.1 0.05

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
59
deviation values of the characters at the subgenus and species level. In Table 3 we report of each species
the features as explained in Fig. 3. A discriminant analysis based on the shell features is presented in
Fig. 8. In Figs 9–11 the relations between some of the most relevant shell characters are graphically
illustrated.
Female genitalia. The data regarding the female genital features are presented in Table 4.
Table 3. Percentage of the analyzed female specimens presenting the shell feature as explained in Fig. 3
within each species of Clessiniella Zallot et al., 2015. In bold the feature status as presented by the
types. Abbreviation: N = number of analyzed specimens.
External lobe
anomphale
stelucarum
tergestinum
villae
waldemari
spots
anomphale
stelucarum
tergestinum
villae
waldemari
N16 14 108 24 55 N17 14 111 25 56
FF 43% 25% 83% 2% 018% 4% 93%
AF 57% 15% 9% 129% 47% 28% 7%
II 60% 13% 89% 271% 100% 35% 68%
OB 100% 4%
rib strenght
anomphale
stelucarum
tergestinum
villae
waldemari
ribs near ap.
anomphale
stelucarum
tergestinum
villae
waldemari
N17 14 111 25 56 N16 14 109 24 24
0 W 50% 50% 90% 46% 98%
16% I38% 50% 10% 33% 2%
282% 21% 29% 84% S13% 21%
312% 79% 71% 100% 16%

European Journal of Taxonomy 762: 49–95 (2021)
60
Table 4. Number of female specimens presenting the genital features as explained in Fig. 5. Abbreviation:
N = number of analyzed specimens.
SR shape
SR linearity
Loops set up
Loops main position
CBC position
BC size
SR width
SR apex position
% of bent SR
JUG position
Loops number
Clessiniella N196 196 196 195 195 189 196 185 194 171 195
SM 19 S190 S63 O 6 0 190 1 6 1 18 ≤0.5 6 0 51 0114 113
SL 166 U 6 O 62 DH 69 1 5 2 43 2110 0.6 15 0.1 72 0.5 36 240
BM 1 T C 71 DM 120 2 3 114 368 0.7 45 0.2 48 121 357
BL 10 3 4 26 0.8 64 0.3 15 1.5 451
4≥0.9 55 ≥0.4 8 2 5 28
≥2.5 ≥6 6
C.(C.) anomphale N13 13 13 13 13 13 13 12 13 12 13
SM 7 S 13 S13 O 3 0 13 1 1 1 2 ≤0.5 3 0 13 0 6 1 2
SL 6 U O DH 10 1 2 12 212 0.6 40.1 0.5 6 2 9
BM T C DM 2 3 3 0.7 10.2 1 3 2
BL 3 4 0.8 30.3 1.5 4
4≥0.9 1≥0.4 2 5
≥2.5 ≥6
C. (C.) s teluca rum N11 11 11 11 11 11 11 10 11 911
SM 1 S 11 S11 O 3 0 9 1 1 ≤0.5 1 0 7 0 2 1 9
SL 5 U O DH 7 1 2 2 5 2 11 0.6 0.1 40.5 5 2 2
BM 1 T C DM 1 2 3 6 3 0.7 40.2 1 2 3
BL 4 3 4 0.8 50.3 1.5 4
4≥0.9 ≥0.4 2 5
≥2.5 ≥6
C. (C.) tergestinum N104 104 104 104 103 101 104 98 102 90 103
SM S99 S 5 O 0 102 1 2 1 2 ≤0.5 0 6 0 60 1 1
SL 98 U 5 O 46 DH 21 1 1 2 12 254 0.6 40.1 43 0.5 19 2 6
BM T C 53 DM 83 2 3 69 348 0.7 23 0.2 37 111 327
BL 6 3 4 18 0.8 34 0.3 11 1.5 441
4≥0.9 37 ≥0.4 5 2 5 22
≥2.5 ≥6 6
C. (C.) villae N18 18 18 18 18 18 18 18 18 17 18
SM S17 S 6 O 0 18 1 1 12 ≤0.5 0 3 0 4 1
SL 18 U 1 O 1 DH 7 1 2 2 2 6 0.6 0.1 80.5 5 2 1
BM T C 11 DM 11 2 3 14 30.7 10 0.2 7 1 8 3 8
BL 3 4 2 0.8 40.3 1.5 4 6
4≥0.9 4≥0.4 2 5 3
≥2.5 ≥6
C. (C.) waldemari N50 50 50 50 50 46 50 46 50 43 50
SM 11 S50 S28 O 0 48 1 3 1 2 ≤0.5 2 0 22 042 1 1
SL 39 U O 15 DH 24 1 2 2 11 228 0.6 70.1 18 0.5 1 2 22
BM T C 7 DM 26 2 3 26 320 0.7 70. 2 4 1 3 20
BL 3 4 6 0.8 18 0.3 41.5 4 4
4≥0.9 12 ≥0.4 2 2 5 3
≥2.5 ≥6

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
61
Table 5. GenBank voucher (H3).
Taxon Location Isolate GenBankVoucher (H3)
Cochl. (Clessiniella) anomphale Eisenkappel, Austria EZ0034 KP408278
Cochl. (Clessiniella) stelucarum Anduins, Italy EZ0102 KX120829
Cochl. (Clessiniella) stelucarum Cornino, Italy EZ0104 KX120830
Cochl. (Clessiniella) stelucarum Mt Bernadi a, Italy EZ0107 KX120831
Cochl. (Clessiniella) tergestinum Val Rosandra, Ital y EZ0840 KP408283
Cochl. (Clessiniella) tergestinum Rakow Skocj an, Sloveni a EZ0838 KP408282
Cochl. (Clessiniella) tergestinum Drivenik spring, Croatia WdM6714 KP408281
Cochl. (Clessiniella) tergestinum Jakačici, Croatia WdM6699 KP408280
Cochl. (Clessiniella) tergestinum Rasko Polje, Croatia EZ0813 KP408279
Cochl. (Clessiniella) villae P erarolo, Ital y EZ0891 KP408284
Cochl. (Clessiniella) waldemari Mt Klek, Croati a EZ0904 KP408289
Cochl. (Clessiniella) waldemari Benete, Slovenia WdM6805 KP408288
Cochl. (Clessiniella) waldemari Vrsic Pass, Sloveni a EZ0896 KP408287
Cochl. (Clessiniella) waldemari Tisovec, Slovenia WdM6749 KP408286
Cochl. (Clessiniella) waldemari Polog-E of Čepovan, Slovenia WdM6842 KP408285
Cochl. (Clessiniella) waldemari Mt Klek, Croati a EZ0905 KX120837
Cochl. (Cochlostoma) septemspirale Plitvice, Croatia WdM3031 KP408291
Fig. 7. Statistical parsimony network based on Histone H3 sequences of species in the subgenus
Clessiniella Zallot et al., 2015. Cochlostoma (Cochlostoma) septemspirale (Razoumowsky, 1789) was
used as an outgroup. For sample IDs we refer to Table 5.

European Journal of Taxonomy 762: 49–95 (2021)
62
Fig. 8. Discriminant analysis based on shell features of the species in Clessiniella Zallot et al., 2015.
Violet dots = Cochlostoma (Clessiniella) anomphale Boeckel, 1939; red squares = Co. (Cl.) stelucarum
sp. nov.; green triangles = Co. (Cl.) tergestinum (Westerlund, 1878); blue diamonds = Co. (Cl.) villae
(Strobel, 1851); black triangles = Co. (Cl.) waldemari (A.J. Wagner, 1897). Analysis performed with
the software ‛Past’ (https://palaeo-electronica.org).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
63
Fig. 9. Number of whorls vs height of the species in Clessiniella Zallot et al., 2015. Violet dots =
Cochlostoma (Clessiniella) anomphale Boeckel, 1939; red squares = Co. (Cl.) stelucarum sp. nov.;
green triangles = Co. (Cl.) tergestinum (Westerlund, 1878); blue diamonds = Co. (Cl.) villae (Strobel,
1851); black triangles = Co. (Cl.) waldemari (A.J. Wagner, 1897). Analysis performed with the software
‘Past’ (https://palaeo-electronica.org).
Fig. 10. Width of the 1st whorl vs height (bottom) of the species in Clessiniella Zallot et al., 2015.
Violet dots = Cochlostoma (Clessiniella) anomphale Boeckel, 1939; red squares = Co. (Cl.) stelucarum
sp. nov.; green triangles = Co. (Cl.) tergestinum (Westerlund, 1878); blue diamonds = Co. (Cl.) villae
(Strobel, 1851); black triangles = Co. (Cl.) waldemari (A.J. Wagner, 1897). Analysis performed with
the software ‘Past’ (https://palaeo-electronica.org).

European Journal of Taxonomy 762: 49–95 (2021)
64
DNA analysis
Female specimens of each species were used for DNA extraction. A Histone fragment (H3) was the
only DNA marker we could successfully amplify in the 5 species. New H3 sequences were deposited
in GenBank (KX120829–31 and KX120837, see Table 5). They were aligned with the sequences
of Clessiniella published by Zallot et al. (2015) (Fig. 3). As Zallot et al. (2015) have found that
Cochlostoma s. str. is the sister clade of Clessiniella in the H3 tree, a sequence of Co. (Co.) septemspirale
(Razoumowsky, 1789) was used as outgroup. Because of the low resolution, the phylogeny does not
show a tree-like structure. Therefore, we illustrated in Fig. 7 the relationships by a statistical parsimony
network (Clément et al. 2002) calculated by Popart ver. 1.7. (Leigh & Bryant 2015). It suggests that Co.
(Cl.) stelucarum sp. nov. might be the closest relative of Co. (Cl.) anomphale Boeckel, 1939 and that
Co. (Cl.) tergestinum (Westerlund, 1878) and Co. (Cl.) villae (Strobel, 1851) as closely related sister
species. Because the limited number of specimens analyzed (only 1 specimen in Co. (Cl.) anomphale
and Co. (Cl.) villae), however, further and more extensive molecular research would be needed.
Taxonomy
Subclass Caenogastropoda Cox, 1960
Order Architaenioglossa Haller, 1892
Superfamily Cyclophoroidea J.E. Gray, 1847
Family Cochlostomatidae Kobelt, 1902
Genus Cochlostoma Jan, 1830
Subgenus Clessiniella Zallot, Groenenberg, De Mattia, Fehér & Gittenberger, 2015
Clessiniella Zallot, Groenenberg, De Mattia, Fehér & Gittenberger, 2015: 80.
Type species
Pomatias maculatum var. villae Strobel, 1851, by original designation.
Remarks
The classication follows MolluscaBase (2021) except for the subgeneric name. Contrary to ICZN
Art. 10.4, Neglecta Wagner, 1897 is used in MolluscaBase as a senior synonym of Clessiniella. Wagner
(1897: 6 [570]) introduced Neglecta as “Formenkreis Neglecta”. It cannot be denied that the German
term Formenkreis is “.. a term such as “superspecies””. As a consequence, according to the ICZN ruling,
Neglecta “.. is not deemed to be a genus-group name” (ICZN Art. 10.4, see also Zallot et al. 2015).
Strobel (1851) and De Betta & Martinati (1855) named this entity Pomatias maculatum var. Villae – but
referring to Pomatias maculatus Draparnaud, 1805 (= Co. (Co.) septemspirale).
Diagnosis of the subgenus
Shell. Ribbed teleoconch with straight ribs, variable in strength and shape. The columellar lobe is
curved backwards, covering the umbilicus. The teleoconch spots (if present) form 2 rows on the whorls
(often, a 3rd row of spots is visible on the body whorl): the upper one close to the suture with rectangular
spots abapically elongated; the lower one roughly ⅓ up on the whorls (Fig. 12). In some populations
the 2 rows of spots merge and there is a continuous vertical darker band on the whorls. The protoconch
is dull and robust with ne riblets starting after 1.2 whorls or more from the apex; it is transparent and
fragile or the ribbing starts close to the apex in other subgenera.
Female genitalia. There is an apical connection of the pedunculus to the bursa copulatrix, a long
seminal receptacle (normally club-shaped), which runs parallel to the pedunculus. The junction of the
uterus gland is situated close to the connection between the pedunculus and the distal oviduct. This

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
65
Fig. 11. Ribs per mm on the 1st whorl vs ribs on the 4th whorls of the species in Clessiniella Zallot
et al., 2015. Violet dots = Cochlostoma (Clessiniella) anomphale Boeckel, 1939; red squares =
Co. (Cl.) stelucarum sp. nov.; green triangles = Co. (Cl.) tergestinum (Westerlund, 1878); blue diamonds
= Co. (Cl.) villae (Strobel, 1851); black triangles = Co. (Cl.) waldemari (A.J. Wagner, 1897). Analysis
performed with the software ‘Past’ (https://palaeo-electronica.org).

European Journal of Taxonomy 762: 49–95 (2021)
66
combination of characters in the female genitalia is most diagnostic to distinguish Clessiniella from the
other subgenera.
male genitalia. Apart from its apical part, the penial spermiduct is tortuous. The body spermiduct is
protruding (often almost detached from the body) more than in the other subgenera and either straight or
more or less tortuous. The posterior side of the penis is wrinkled.
Identication key for the species
1. Body whorl inated shortly before the lip (Fig. 4A: OB). In the female genitalia, simplied loop path
with 1–2 loops ............................................................................Co. (Cl.) anomphale Boeckel, 1939
– Body whorl not inated before the lip (Fig. 4A: FF, AF, II) ............................................................. 2
2. In the female genitalia, the loops of the visceral oviduct are 1–2 ........Co. (Cl.) stelucarum sp. nov.
– In the female genitalia, the loops of the visceral oviduct are more than 2 ....................................... 3
3. In the male genitalia, the body spermiduct is twisted ............... Co. (Cl.) waldemari (Wagner, 1897)
– In the male genitalia, the body spermiduct is straight ...................................................................... 4
4. Ribs of the shell widely spaced and barely inclined, protoconch relatively small (Fig. 13). Slim,
club-shaped seminal receptacle in the female genitalia ..................... Co. (Cl.) villae (Strobel, 1851)
– Ribs less widely spaced and more inclined, relatively larger protoconch. Less slim seminal
receptacle .......................................................................... Co. (Cl.) tergestinum (Westerlund, 1878)
Fig. 12. Most common spot patterns on the shells of the species of Clessiniella Zallot et al., 2015.

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
67
Cochlostoma (Clessiniella) villae (Strobel, 1851)
Figs 1, 3C, 14D, 15D, 16D, 17–21
Pomatias maculatum var. villae Strobel, 1851: 17 (“Mizzole presso Verona”).
Pomatias maculatum var. villae De Betta & Martinati, 1855: 74 (“Prov. Verona, Mizzole”).
Cochlostoma (Auritus) villae – Alzona 1971: 22.
Cochlostoma villae − Welter-Schultes 2012: 102.
Diagnosis
The last whorl, approaching the aperture, hardly enlarges (Fig. 4A: FF). Very prominent and widely
spaced teleoconch ribs, barely inclined in respect to the vertical axis. Spots almost always visible. The
protoconch is relatively small. The apertural lip poorly developed. Visceral oviduct with convoluted
loops; thin seminal receptacle. Straight body spermiduct.
Type material
Syntypes
ITALY • 1 ♀; “Pomatias villae Spinel. Mizzole. Villa 49” [label of the original material] [Veneto, Verona,
Mizzole] [thus, it could be interpreted as collected in 1849 by Villa]; PU1610-C1 (Fig. 19) • 2 ♂♂; same
label data as for preceding; PU1610-C2, PU1610-C3 (Fig. 20).
Other material (fresh specimens from the type locality have also been analyzed)
ITALY • Altissimo; 45.61511° N, 11.25225° E; 2009; Zallot leg.; EZ0883 • Arqua’ Petrarca, surroundings;
45.26194° N, 11.69623° E; 1999; Zallot leg.; EZ0895 • Bolca; 45.59389° N, 11.20972° E; 2010; Zallot
leg.; EZ0882 • Ca’Gottolo; 45.58921° N, 10.90787° E; Jun. 2010; Scarlassara leg.; EZ1015 • Cornedo
Vicentino-Cereda; 45.60667° N, 11.36833° E; 1999; Zallot leg.; EZ0887 • Covoli di Velo; 45.60889° N,
11.12083° E; 2010; Zallot leg.; EZ0879 • Ferrazza; 45.62137° N, 11.19538° E; Apr. 2010; Zallot leg.;
Fig. 13. Shell characters (ribs inclination and relative size of the protoconch) which could be helpful to
distinguish Cochlostoma (Clessiniella) villae (Strobel, 1851) (blue diamonds) from Co. (Cl.) tergestinum
(Westerlund, 1878) (green triangles).

European Journal of Taxonomy 762: 49–95 (2021)
68
EZ0881 • Fumane-Molina; 45.56942° N, 10.89856° E; Jun. 2010; Scarlassara leg.; EZ0936 • Lago di
Fimon; 45.46972° N, 11.54° E; 2009; Zallot leg.; EZ0892 • Lumignano; 45.45607° N, 11.57914° E;
2010; Zallot leg.; EZ0893 • Mizzole; 45.48662° N, 11.05524° E; Apr. 2010; Zallot leg.; EZ0878 • Monte
delle Piume; 45.35477° N, 11.45641° E; May 2010; G. Zallot leg.; EZ0889 • Mt Rusta; 45.28735° N,
11.68563° E; 2002; Zallot leg.; EZ0894 • Novale-Valdagno; 45.66528° N, 11.29528° E; 2000; Zallot leg.;
EZ0884 • Perarolo dei Monti Berici, 45.47694° N, 11.50056° E; 1999, Zallot leg.; EZ0891 • Priabona;
45.63389° N, 11.37417° E; 2000; Zallot leg.; EZ0888 • San Giovanni di Zovencedo; 45.42833° N,
11.49417° E; 1999; Zallot leg.; EZ0890 • Tregnago-Finetti; 45.52866° N, 11.1806° E; 2005; Zallot leg.;
EZ0880 • Valdagno; 45.64991° N, 11.30079° E; 2000; Zallot leg.; EZ0885.
Fig. 14. The ve species of Clessiniella Zallot et al., 2015. A–D. Picture of a female (top) and male
(bottom) specimen from the type localities. A. Cochlostoma (Clessiniella) anomphale Boeckel,
1939. Vellach, Austria (EZ0991). B. Co. (Cl.) stelucarum sp. nov., Cornino, Italy (EZ0104).
C. Co. (Cl.) tergestinum (Westerlund, 1878), Val Rosandra, Italy (EZ0840). D. Co. (Cl.) villae (Strobel,
1851), Mizzole, Italy (EZ0878). E. Specimens from a locality close to the type one: Co. (Cl.) waldemari
(A.J.Wagner, 1897), Mt Klek, Croatia (EZ0905).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
69
Fig. 15. Comparison of female genitalia. A. Cochlostoma (Clessiniella) anomphale Boeckel, 1939. Vellach,
Austria (EZ0991). B. Co. (Cl.) stelucarum sp. nov., Cornino, Italy (EZ0104). C. Co. (Cl.) tergestinum
(Westerlund, 1878), Val Rosandra, Italy (EZ0840). D. Co. (Cl.) villae (Strobel, 1851), Mizzole, Italy
(EZ0878). E. Co. (Cl.) waldemari (A.J.Wagner, 1897), Mt Klek, Croatia (EZ0905).
Fig. 16. Comparison of male genitalia. A. Cochlostoma (Clessiniella) anomphale Boeckel, 1939. Vellach,
Austria (EZ0991). B. Co. (Cl.) stelucarum sp. nov., Cornino, Italy (EZ0104). C. Co. (Cl.) tergestinum
(Westerlund, 1878), Val Rosandra, Italy (EZ0840). D. Co. (Cl.) villae (Strobel, 1851), Mizzole, Italy
(EZ0878). E. Co. (Cl.) waldemari (A.J.Wagner, 1897), Tisovec, Croatia (WdM6750).

European Journal of Taxonomy 762: 49–95 (2021)
70
Description
Shell. The teleoconch whorls have whitish, very prominent, rather straight and widely spaced
ribs on the upper whorls (see Fig. 11), but smooth specimens are found occasionally (see
https://www.naturamediterraneo.com/forum/topic.asp?TOPIC_ID=43816).
lip. Moderately developed, and more or less attened by the internal callus of the aperture. It gently
curves backwards at the columellar side, starting rather far from the upper part of the body whorl.
On lateral view, the body whorl, approaching the aperture, in most of the specimens barely enlarging
(Fig. 4A: FF, Table 3).
Female genitalia. There is a long and relatively thin (as compared to the other species of Clessiniella;
see Table 5) seminal receptacle with a curved apex.
male genitalia. The penis is long and inated. The penial spermiduct is slightly twisted and occupies
only the central part of the penis. There is a clearly protruding but straight body spermiduct.
Fig. 17. Shell variability in Cochlostoma (Clessiniella) villae (Strobel, 1851). A. Mizzole, Italy
(EZ0878). B. Covoli di Velo, Italy (EZ0879). C. Ferrazza, Italy (EZ0881). D. Mt delle Piume, Italy
(EZ0889). E. Perarolo, Italy (EZ0891). F. Lumignano, Italy (EZ0893). G. Ca’ Gottolo, Italy (EZ1015).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
71
Distribution
The species inhabits a relatively small area at the foothills of the eastern Alps. It can also be found on
the hills stretching down the Venetian atland of the “pianura Veneta” (Colli Berici and Colli Euganei).
It has been found sympatric with Cochlostoma (Cochlostoma) septemspirale and Cochlostoma
(Eupomatias) philippianum in some localities of the province of Verona (Fernando Scarlassara, personal
communication).
Habitat preference
The snails are found on rocky cliffs as well as on smaller boulders and dry stone walls in woody places.
Remarks
The range of this species is far from the almost continuous area in the east where the other Clessiniella
species are known from. The species has been reported (and still is in the checklist of the Italian fauna
– www.faunaitalia.it) as Cochlostoma villae (De Betta & Martinati, 1855) but Bank (1988) found that
Strobel (1851) described it as Pomatias maculatum var. villae Spinelli, four years before De Betta &
Martinati (1855). The description provided by Strobel (1851) is poor: “Varieta’ generalmente maggiore,
coi giri della spira piu’ gon e colle coste trasversali piu’ regolari, e saglienti” = “variety generally
bigger, with more rounded whorls and more regular and strong ribs” and follows “P. Villae Spinelli in
Fig. 18. Variation of the female genitalia in Cochlostoma (Clessiniella) villae (Strobel, 1851).
A. Mizzole, Italy (EZ0878). B. Ferrazza, Italy (EZ0881). C. Mt delle Piume, Italy (EZ0889). D1–
D3. Perarolo, Italy (EZ0891). E. Mt Rusta, Italy (EZ0894). F. Ca’ Gottolo, Italy (EZ1015).

European Journal of Taxonomy 762: 49–95 (2021)
72
Fig. 19. Cochlostoma (Clessiniella) villae (Strobel, 1851), syntype, ♀, at the Museo di Storia Naturale
of Parma University, Italy (PU1610-C1).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
73
esemplari raccolti a Mizzole presso Verona.” = “P. villae Spinelli on specimens collected in Mizzole
near Verona.”
A few years later, De Betta & Martinati (1855) gave almost the same poor description: “c. elongatior,
costulis anfract. super. distinctioribus” = “taller shell, ribs in the upper whorls spaced” and follows
“Prov. Verona, Mizzole. Spin. ex spec.” = “Verona Province, Mizzole, specimens provided by Spinelli.”
It could be, as was common at the time, that the specimens used to describe the species by both Strobel
(1851) and De Betta & Martinati (1855) have had a tortuous path, after being collected by Villa in
1849 (as it appears from Strobel’s label), given to Spinelli later on, who gave the collector’s name to
the species (as far as is known, however, Giovanni Battista Spinelli never published the specic epithet
Fig. 20. Cochlostoma (Clessiniella) villae (Strobel, 1851), syntypes, ♂♂, at the Museo di Storia
Naturale of Parma University, Italy (1610-C2, left; 1610-C3, right).

European Journal of Taxonomy 762: 49–95 (2021)
74
Fig. 21. Shell of Cochlostoma (Clessiniella) villae (Strobel, 1851). Collezione De Betta, Museo di
Storia Naturale di Verona (photo Folco Giusti).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
75
villae, probably using the name only in private correspondence). Spinelli eventually gave part of the
material to Strobel and part to De Betta (the De Betta material is currently kept at the Museo di Storia
Naturale di Verona, see Fig. 21).
Cochlostoma (Clessiniella) anomphale Boeckel, 1939
Figs 1, 3A, 3A1, 14A, 15A, 16A, 22–24
Cochlostoma septemspirale anomphale Boeckel, 1939: 41–50 (“Karawanken, Trögern. Weg nach
Vellach”).
Cochlostoma septemspirale anomphale – Zilch 1958: 57.
Cochlostoma (Turritus) anomphale – Gittenberger 1971: 115–118.
Cochlostoma anomphale – Welter-Schultes 2012: 91.
Diagnosis
Shell quite densely ribbed, spots on the teleoconch generally clearly visible, body whorl inated shortly
before the lip. Visceral oviduct simplied with 1–2 supercial loops at most; seminal receptacle always
club-shaped.
Fig. 22. Shell variability in Cochlostoma (Clessiniella) anomphale Boeckel, 1939. A. Savinjska Dolina,
Slovenia (WdM1579). B. Eisenkappel, Austria (EZ0034). C. Stahovika, Slovenia (EZ0035). D. Vellach,
Austria (EZ0991). E. Solčava, Slovenia (WdM7010). F. Logarska Dolina, Slovenia (EZ0994).

European Journal of Taxonomy 762: 49–95 (2021)
76
Type material
Paratypes
AUSTRIA • 2 specs; Trögern, way to Vellach; SMF 42967/2 (Fig. 24).
Other material (we analyzed fresh material from the type locality)
AUSTRIA • EisenKappel; 46.48583° N, 14.59111° E; 2010, Mildner leg.; EZ0034 • Vellach; 46.42332° N,
14.53639° E; Aug. 2010; De Mattia leg.; EZ0991.
SLOVENIA • 210 road, N of Kokra; 46.34318° N, 14.48336° E; Aug. 2010; De Mattia leg.; EZ0993
• Gornji Grad; 46.2934° N, 14.82796° E; Aug. 2010; De Mattia leg.; EZ0996 • Kamniska Bistrika;
46.32722° N, 14.59° E; 2010; Zallot leg.; EZ0033 • Kokra; 46.30356° N, 14.4771° E; Aug. 2010; De
Mattia leg.; EZ0995 • Logarska Dolina; 46.41959° N, 14.65074° E; Aug. 2010; De Mattia leg.; EZ0994
• Logarska Dolina; 46.42334° N, 14.66301° E; Aug. 2010; De Mattia leg.; EZ0992 • Podvolovliek-
Stahovica; 46.28787° N, 14.69765° E; Aug. 2005; Zallot leg.; EZ0036 • Savinjska Dolina; 46.36596° N,
14.7318° E; 2010; De Mattia leg.; WdM1579 • Solčava; 46.42002° N, 14.19172° E; Jul. 2010; De
Mattia leg.; WdM7010 • Stahovika; 46.26528° N, 14.60361° E; 2005; Zallot leg.; EZ0035 • SV Anton;
46.28787° N, 14.69765° E; 2010; Zallot leg.; EZ0032.
Remarks
The holotype and 23 paratypes should be in the Boeckel collection at the Naturkundemuseum Erfurt
(Hartmann et al. 2018: 26). However, the shells could not be traced when this article was prepared
(C. Unger in mail October 14, 2020).Type locality - Austria, Trögern, way to Vellach.
Fig. 23. Variation of the female genitalia in Cochlostoma (Clesssiniella) anomphale Boeckel, 1939.
A. Eisenkappel, Austria (EZ0034). B. Savinjska Dolina, Slovenia (WdM1579). C. Vellach, Austria
(EZ0991). D. Logarska Dolina, Slovenia (EZ0994). E1–E2. Solčava, Slovenia (WdM7010).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
77
Fig. 24. Cochlostoma (Clessiniella) anomphale Boeckel, 1939. Paratypes at the Senckenberg Museum
Frankfurt (SMF42967/2).

European Journal of Taxonomy 762: 49–95 (2021)
78
Description
Shell. The outside of the body whorl is inated shortly before the apertural lip (Fig. 4A: OB). The lip
is poorly developed; it is almost vertical at the palatal side, and curved backwards and covering the
umbilicus at the columellar side. The shells are more densely sculptured and the ribs are weaker than in
the other species of Clessiniella. Most of the shells have spots.
Female genitalia. There is a spindle-like seminal receptacle. The visceral oviduct has only 1–2,
supercial, loops.
male genitalia. The swollen penis is longer than the body and has a central spermiduct. There is a
protruding, straight, body spermiduct.
Distribution
Cochlostoma (Clessiniella) anomphale inhabits the Karawanken in south-eastern Austria and northern
Slovenia.
Habitat preference
The animals can be found under and on stones and boulders in shady habitats in forests.
Remarks
This species was initially described as a subspecies of Cochlostoma (Cochlostoma) septemspirale.
Gittenberger (1971) considered it a separate species because of the sympatry with Co. (Co.)
semptemspirale heydenianum and the lack of conchologically intermediate specimens. This is an
epitome of quite common Cochlostoma taxonomical issues: based on shell morphology it was initially
described as subspecies of a taxon which is only distantly related.
Cochlostoma (Clessiniella) stelucarum sp. nov.
urn:lsid:zoobank.org:act:4609B41C-3611-4F8A-9048-5F97EA0446E6
Figs 1, 3E, 14B, 15B, 16B, 15–27
Diagnosis
Slender shell with 8½–10½ rather widely ribbed whorls. Spots always visible. Poorly developed lip with
FF or AF external lobe. Visceral oviduct simplied and seminal receptacle sometimes oval.
Etymology
The epithet ‘steluca’ is an acronym noun formed from the initial syllables of the names of the wife and
two daughters (Stefania, Ludovica and Camilla Zallot) of the senior author.
Type material (Fig. 27)
Holotype
ITALY • ♀ (in ethanol); Friuli Venezia Giulia, Udine, Cornino; 46.2297° N, 13.0250° E; 180 m a.s.l.;
5 Jun. 2002; Zallot leg.; rocks surrounding the lake; RMNH.MOL.347617.
Paratypes
ITALY • 1 ♂ (in ethanol); same collection data as for holotype; RMNH.MOL.347618 • 1 ♀ (empty
shell); same collection data as for holotype; GenBank voucher H3: KX120830; RMNH.MOL.347619.

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
79
Other material
ITALY • Anduins-cliffs; 46.23755° N, 12.97106° E; 1997; Zallot leg.; EZ0102 • Campone; 46.26194° N,
12.83333° E; 2010; Zallot leg.; EZ0098 • Castelmonte; 46.094° N, 13.52095° E; Oct. 2010; Margelli leg.;
EZ1020 • Cjanet-Peonis; 46.25917° N, 13.04111° E; 2002; Zallot leg.; EZ0105 • Cornino; 46.22972° N,
13.025° E; 2002; Zallot leg.; EZ0104 • F. Piccola Mt Valinins; 46.245° N, 12.80111° E; 2000; Zallot
leg.; EZ0095 • Mt Bernadia; 46.23361° N, 13.25556° E; 2005; Zallot leg.; EZ0107 • Mt Ciaurlec;
46.21226° N, 12.88018° E; 2005; Zallot leg.; EZ0099 • Mt Prat; 46.24577° N, 12.9907° E; 2002; Zallot
leg.; EZ0103 • Mt Valinis; 46.23417° N, 12.8075° E; 2002; Zallot leg.; EZ0096 • San Vito D’asio;
46.2337° N, 12.94531° E; 1997; Zallot leg.; EZ0101 • Tramonti-Campone; 46.27833° N, 12.81917° E;
2000; Zallot leg.; EZ0097 • Val di Torre-all’imbocco; 46.23861° N, 13.23111° E; 1998; Zallot leg.;
EZ0106 • Valle D’Arzino; 46.30278° N, 12.92972° E; 2000; Zallot leg.; EZ0100.
Fig 25. Shell variability in Cochlostoma (Clessiniella) stelucarum sp. nov. A. Cornino, Italy (EZ0104).
B. Mt Valinis, Italy (EZ0096). C. Anduins, Italy (EZ0102). D. Mt Prat, Italy (EZ0103). E. Val di Torre,
Italy (EZ0106).

European Journal of Taxonomy 762: 49–95 (2021)
80
Description
Shell. The shell has prominent spots and strong and rounded ribs, which are regularly and widely
spaced and coloured like the background. The ribs become less prominent on the body whorl. The body
whorl slightly and gradually enlarges while approaching the aperture (Fig. 4A: FF, AF). In comparison
with the other species of the subgenus, the whorls increase more slowly in size so that, given the same
shell height, there is a higher number of whorls (Fig. 9).
Female genitalia. The visceral oviduct has only 1–2 supercial loops, what makes it one of the simplest
in the genus Cochlostoma. The seminal receptacle is thin and may be oval, thus not always club-shaped
as in the other species.
male genitalia. The penis is longer than the body and inated, although less so than in the other species
of Clessiniella. The penial spermiduct is gently twisted. The body spermiduct is straight. There is a
long groove, ending in a well-formed sperm pocket, delimited at the frontal side by an ascending sperm
funnel.
Distribution
The species inhabits a restricted area in the eastern Southern Pre-Alps. Our sampling localities are
located between Val Tramontina to the west and Val di Torre to the east (Friuli region, Italy).
Habitat preference
Like in most species of Cochlostoma, these are obligate rock-dwelling snails but, rather than on the
surface of limestone cliffs, their main habitat is on smaller surfaces of stones and boulders which are
normally found in large numbers at the foot of the cliffs.
Fig 26. Variation of the female genitalia in Cochlostoma (Clessiniella) stelucarum sp. nov. A. Mt Valinis,
Italy (EZ0096). B. Campone, Italy (EZ0098). C. Anduins, Italy (EZ0102). D1–D3. Cornino, Italy
(EZ0104). E. Cjanet-Peonis, Italy (EZ0105). F. Mt Bernadia, Italy (EZ0107).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
81
Remarks
Cochlostoma (Clessiniella) stelucarum sp. nov. was introduced as Cochlostoma sp. by Zallot (2002:
98). At most of the localities, it lives syntopically with either Co. (Eupomatias) philippianum (Gredler,
1853) or Co. (Eupomatias) henricae (Strobel, 1851). Near the village of Anduins it has been found a
few centimeters apart from both Co. (Eu.) henricae and Co. (Cochlostoma) septemspirale on stones and
boulders. This is one of the rare cases where 3 species of Cochlostoma, belonging to different subgenera,
live syntopically. The range of the species is situated in between the range of Co. (Cl.) villae in the west
and that of the other species of Clessiniella in the east.
Fig. 27. Cochlostoma (Clessiniella) stelucarum sp. nov. Top left: holotype, ♀ (RMNH.MOL.347617),
H = 8 mm. Top right: paratype, ♂ (RMNH.MOL.347618), H = 7 mm. Bottom: paratype, ♀ (RMNH.
MOL.347619), empty shell, H = 8.7 mm.

European Journal of Taxonomy 762: 49–95 (2021)
82
Cochlostoma (Clessiniella) tergestinum (Westerlund, 1878)
Figs 1, 3B, 5, 14C, 15C, 16C, 28–30, 34 (bottom), 35 (bottom), 36 (top)
Pomatias tergestinus Westerlund, 1878: 108–109 (“Triest”).
Pomatias tortivus Westerlund, 1885: 124 (“Fiume, San Daniel u. Karstgebirge”, which should be Rijeka
in Croatia and Štanjel in Slovenia).
Pomatias tergestinus var. grahovana A.J. Wagner, 1897: 616, pl. 9 g. 89 (“Grahovo bei Fiume”, a
locality ca 10 km north of Rijeka in Croatia).
Pomatias tergestinus var. tortiva – A.J. Wagner 1897: 616, pl. 9 g. 88.
Cochlostoma tergestinum − Welter-Schultes 2012: 102.
Diagnosis
Shell very variable, lip curved backwards more or less abruptly at the columellar side. Strong ribs on the
teleoconch, in most of the populations widely spaced and, on the upper whorls, whitish and contrasting
with the corneous, dark brown to light yellow, shell background. Visceral oviduct with convoluted loops
path; body spermiduct straight.
Type material
Syntypes
ITALY • “Triest” [Friuli Venezia Giulia, Trieste]; Westerlund leg.; SMF160677/1 • same label data as for
preceding; SMF 160676/7 (Fig. 30).
Other material
AUSTRIA • Dobratsch; 46.59222° N, 13.6825° E; 2009; Mildner and Giovannelli leg.; EZ0799 •
Kranzwand; 46.588799° N, 13.75002° E; Jul. 2015; AL leg.; NHMW109000.
CROATIA • Sveta Nedelja near Labin; 45.13741° N, 14.14488° E; 2000; De Mattia leg.; WdM6961
• Brest, Cicaria; 45.45444° N, 14.00625° E; Apr. 2008; De Mattia leg.; WdM6715 • Drivenik spring;
45.24147° N, 14.64828° E; Mar. 2009; De Mattia leg.; WdM6714 • from A6 to 501 down Bakar;
45.30543° N, 14.58579° E; Jul. 2009; De Mattia and Zallot leg.; EZ0938 • Jakačici; 45.179494° N,
14.004384° E; 2009; De Mattia leg.; WdM6699 • Kremenje-Buje; 45.43558° N, 13.68499° E; Jan.
2008; De Mattia leg.; WdM6400 • Kukurini; 45.19369° N, 14.07216° E; Feb. 2009; De Mattia leg.;
WdM6698 • Limski Kanal; 45.12957° N, 13.74039° E; 2010; De Mattia leg.; WdM6693 • Limski
Kanal; 45.12957° N, 13.74039° E; Jun. 2010; De Mattia leg.; WdM6998 • Mala Ucka; 45.27605° N,
14.19715° E; 2009; De Mattia leg.; WdM6768 • Meja, Bakarac; 45.3037° N, 14.58348° E; 2009; De
Mattia leg.; WdM6951 • Momjan; 45.442° N, 13.71384° E; 2009; De Mattia leg.; WdM6854 • Mt Učka,
beech forest; 45.30636° N, 14.20865° E; 2009; De Mattia leg.; WdM6794 • Mt Učka, top; 45.28967° N,
14.20341° E; 2009; De Mattia leg.; WdM6794 • Pazin; 45.24134° N, 13.93099° E; Jun. 1986; Drimmer
leg.; HNHM97227 • Permani; 45.43742° N, 14.28905° E; 2009; De Mattia leg.; WdM6708 • Raša;
45.0872° N, 14.08854° E; 2010; De Mattia leg.; WdM6968 • Rasko Polje; 45.15638° N, 14.0538° E;
2010; De Mattia leg.• EZ0813 • Rasko Polje; 45.15638° N, 14.0538° E; 1900; De Mattia leg.; EZ1027
• Zminj; 45.13991° N, 13.88873° E; 2000; De Mattia leg.; WdM6700.
ITALY • Val Rosandra; 45.62° N, 13.87139° E; 2000; Zallot leg.; EZ0840 • Torri di Monrupino;
45.71878° N, 13.80594° E; 1998; Zallot leg.; EZ0839.
SLOVENIA • Abram-Nanos Mt; 45.82055° N, 14.01433° E; 2008; De Mattia leg.; WdM6731
• Brestovica pri Komnu; 45.80996° N, 13.65016° E; 2009; De Mattia leg.; WdM6948 • Čepno;
45.67071° N, 14.09732° E; Jun. 2009; De Mattia leg.; WdM6866 • Črnotiče-Črni Kal; 45.55276° N,

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
83
13.88633° E; Mar. 2009; De Mattia leg.; EZ1028 • Dolina Nanos-Abran-Podkraj; 45.79813° N,
14.06699° E; Apr. 2009; De Mattia leg.; WdM6733 • Farjev Vrh; 45.92429° N, 13.95442° E; 2009; De
Mattia leg.; WdM6740 • Golac, Vela Vrata; 45.51315° N, 14.02969° E; 2009; De Mattia leg.; WdM6762
• Gorjansko; 45.78485° N, 13.70068° E; 2000; De Mattia leg.; WdM6950 • Hruščica; 45.85832° N,
14.1117° E; 2000; De Mattia leg.; WdM2409 • Ilirska Bistrika; 45.56944° N, 14.23917° E; 2009;
Zallot leg.; EZ0821 • Kovk; 45.94614° N, 13.95058° E; 2015 • De Mattia leg.; WdM6737 • Križna
Jama; 45.82003° N, 14.24579° E; Jun. 2009; Jochum leg.; HNHM97142 • Markovščina-Dimnice cave;
45.58357° N, 14.03994° E; 2009; De Mattia leg.; WdM6765 • Mt Kobilica; 46.06528° N, 13.80907° E;
2005; De Mattia leg.; WdM6849 • Mt Nanos, top, loc. Plesa; 45.79813° N, 14.06699° E; Jun. 2005;
De Mattia leg.; WdM0000 • Mt Snežnik 1161; 45.56816° N, 14.37049° E; Jun. 2010; De Mattia
leg.; WdM7012 • Mt Snežnik 1390; 45.57179° N, 14.41044° E; Jun. 2010; De Mattia leg.; EZ0965 •
Mt Snežnik 1447; 45.57877° N, 14.43839° E; Jun. 2010; De Mattia leg.; EZ0964 • Mt Snežnik 1450;
45.58195° N, 14.43129° E; Jun. 2010; De Mattia leg.; EZ0962 • Mt Snežnik car parking; 45.58235° N,
14.43117° E; Jun. 2002; De Mattia leg.; WdM7006 • Mt Vremščica; 45.69898° N, 14.03388° E; 2009;
De Mattia leg.; WdM6758 • near Gorenje; 45.82976° N, 14.10755° E; 2009; Zallot leg.; EZ0817 •
near Logatec; 45.88076° N, 14.27448° E; 2009; Zallot leg.; EZ0828 • Osp, 45.57071° N, 13.86168° E;
2010; De Mattia leg.; WdM6756 • Ospo, small cliff; 45.57119° N, 13.8582° E; May 2010; De Mattia
leg.; WdM6993 • Otlica; 45.92822° N, 13.90918° E; 1986; De Mattia leg.; WdM6752 • Planina;
45.82405° N, 14.24708° E; 2000; Zallot leg.; EZ0823 • Planina; 45.82972° N, 14.25056° E; 2000;
Zallot leg.; EZ0824 • Planina, Postojna; 45.83° N, 14.26056° E; 2009; Zallot leg.; EZ0826 • Podgorje;
45.53872° N, 13.9449° E; 2000; De Mattia leg.; WdM6735 • Podgorje; 45.5364° N, 13.9463° E; 2009;
De Mattia leg.; WdM6735 • Podlešče-Bainsizza; 46.05738° N, 13.72709° E; May 2009; De Mattia leg.;
WdM6814 • Prešnica; 45.56587° N, 13.94191° E; Mar 2009; De Mattia leg.; WdM6734 • Pri Cerkvi-
Bainsizza; 46.04133° N, 13.77584° E; 2009; De Mattia leg.; WdM6834 • Rakow Skocjan; 45.79467° N,
14.28948° E; 2010; Zallot leg.; EZ0838 • San Canzian (Škocjan); 45.66532° N, 13.99356° E; 2009; De
Mattia leg.; WdM6754 • Senožeče; 45.71421° N, 14.07856° E; 2009; De Mattia leg.; WdM6865 • 10 km
S of Vrhnika; 45.88549° N, 14.26091° E; 2000; Zallot leg.; EZ0827 • Vrhnika, Močilnik; 45.95591° N,
14.29268° E; 2009; Zallot leg.; EZ0829.
Remarks
The original indication of the type locality is rather vague, since only a city in the Friuli-Venezia Giulia
region, in the easternmost part of Italy is indicated. The closest to Trieste where we found the species is
at “Val Rosandra”, 8 km north-east of the city. The fresh specimens collected there have shell features
similar to those of the syntypes. We therefore consider them as topotypical.
Description
Shell. The size of the shells is extremely variable (height 6.1–9.6 mm in the females) as well as their
general morphology (see Fig. 28). Several populations are spotless whereas others have clear spots on
the teleoconch. Depending on the population, the lip can be either poorly or even hugely developed. In
lateral view, the body whorl, while approaching the aperture, may widen in various ways, depending on
the population (Fig. 4A: FF, AF, II). The H/W ratio is very variable and the slenderness of the shells in
different populations varies accordingly.
Female genitalia. There is a long, club-shaped seminal receptacle. In some cases there is a change of
colour at its base, presumably indicating the starting point of the rather long distal oviduct. As shown in
Table 5, there is a considerable variability in the part of the seminal receptacle that is curved down and
in the number of loops.
male genitalia. The inated penis is longer than the body. The penial spermiduct is conned to the
central part of the penis and gently twisted. The body spermiduct is straight and well dened.

European Journal of Taxonomy 762: 49–95 (2021)
84
Fig. 28. Shell variability in Cochlostoma (Clessiniella) tergestinum (Westerlund, 1878). A. Mt Kobilica,
Slovenia (WdM6849). B. Limski Kanal, Croatia (WdM6693). C. Dobratsch, Austria (EZ0799).
D. Farjev Vrh, Slovenia (WdM6740). E. Rasko Polje, Croatia (EZ0813). F. Raša, Croatia (WdM6968).
G. Val Rosandra, Italy (EZ840). H. Mala Učka, top, Croatia (WdM6794). I. Ilirska Bistrika, Slovenia
(EZ0921). L. Permani, Croatia (WdM6708). M. Vrhnika, Slovenia (EZ0827). N. Near Bakar, Croatia
(EZ0938). O. Mala Učka, Croatia (WdM6768). P. Mt Snežnik 1161, Slovenia (WdM7012).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
85
Distribution
This species inhabits the eastern side of the province of Trieste in Italy, the southwestern part of Slovenia
and the Istrian peninsula. It has also been found in several Croatian localities in the surroundings of
Bakar and, quite far from its main range, on the Villacher Alps, in Austria (see Mildner 1997). The
species has been erroneously reported for Albania (see Fehér & Erőss 2009).
Habitat preference
It is an obligate rock-dwelling species. Co. (Cl.) tergestinum is found in a variety of habitats, from open,
exposed limestone cliffs to shady spots in deciduous and evergreen forests.
Remarks
According to Zilch (1958) and Alzona (1971) there are 3 subspecies of Co. (Cl.) tergestinum, viz. the
nominate subspecies Co. (Clessiniella) tergestinum tergestinum (Westerlund, 1878), Co. (Clessiniella)
tergestinum grahovanum (A.J. Wagner, 1897) and Co. (Clessiniella) tergestinum tortivum (Westerlund,
1885).
Fig. 29. Variation of the female genitalia in Cochlostoma (Clessiniella) tergestinum (Westerlund, 1878).
A. Pri Cerkvi, Slovenia (WdM6834). B. Brestovica, Slovenia (WdM6948). C. Dobratsch, Austria
(EZ0799). D. Goriansko, Slovenia (WdM6950). E. Prešnica, Slovenia (WdM6734). F. Farjev Vrh,
Slovenia (WdM6740). G. Rasko Polje, Croatia (EZ0813). H. Rasa, Croatia (WdM6968). I. Permani,
Croatia (WdM6708). L. Meja-Bakarac, Croatia (WdM6951). M. Jakačici, Croatia (WdM6699).
N. Drivenik, Croatia (WdM6714).

European Journal of Taxonomy 762: 49–95 (2021)
86
Fig. 30. Cochlostoma (Clessiniella) tergestinum (Westerlund, 1878), syntype at the Senckenberg
Museum Frankfurt (SMF160676/7).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
87
For the time being, we here synonymize these nominal taxa because the conchological features which
supposedly characterize them, like measures, density and prominence of the ribs, and the structure of the
lip, are population features without an obvious geographical pattern. There is no signicant correlation
between latitude and longitude of the locality of the sample and the character states that are mentioned
in the original description of the so-called subspecies. Moreover, in the surroundings of Trieste, the type
locality of the nominate subspecies, only populations with some ‘tortivum’ shell features, like the thin
lip, can be found (De Mattia & Prodan 2006).
Some of the populations have a peculiar shell morphology (see for instance Fig. 28E–F); their
taxonomical status should be investigated more thoroughly especially since the available molecular
H3 analysis indicates that population “EZ0813” from Rasko Polje, corresponding to the specimen in
Fig. 28E, stands apart from the others.
Cochlostoma (Clessiniella) waldemari (Wagner, 1897)
Figs 1, 3D, 14E, 15E, 16E, 31–33, 34 (top), 35 (top), 36 (bottom)
Pomatias (Auritus) waldemari A.J. Wagner, 1897: 616, pl. 9, 90 (“Ogulin”).
Cochlostoma waldemari – Welter-Schultes 2012: 103.
Diagnosis
Spotless shell. Moderately strong, more or less widely spaced ribs. Columellar lobe gently curved
backwards. External lobe of type “II” in almost all the populations. The body spermiduct is huge and
twisted instead of straight as in all the other species of Clessiniella.
Type material
Syntypes
CROATIA • ♀; Ogulin; A.J. Wagner leg.; MIZPAS-W-6979a (Fig. 33) • 2 ♀♀, 3 ♂♂; same collection
data as for preceding; MIZPAS-W-6979/5.
Other material
CROATIA • Kapela pass, Ogulin; 45.07534° N, 15.20837° E; 2009; De Mattia leg.; EZ0909 •
Klek Mt, close to the top; 45.25904° N, 15.14044° E; 2009; Zallot leg.; EZ0904 • Klek Mt, cliffs
halfway; 45.25438° N, 15.14561° E; 1996; Zallot leg.; EZ0905 • Mt Risnjak; 45.43095° N, 14.6188°
E; 1400 m a.s.l.; 2009; De Mattia leg.; EZ0845 • Mt Risnjak; 45.39072° N, 14.61748° E; 1000 m a.s.l.;
2010; Zallot leg.; EZ0907 • Ročko Polje; 45.3607° N, 14.09534° E; 2009; De Mattia leg.; WdM6697
• Rudnica; 45.23402° N, 15.33827° E; 2009; De Mattia leg.; EZ0906 • Spilja Vrelo; 45.31777° N,
14.71065° E; 2009; Zallot leg.; EZ0899 • Vitunj-Ogulin; 45.29114° N, 15.14041° E; 2005; Zallot leg.;
EZ0903.
SLOVENIA • Benete, 45.76192° N, 14.56752° E; 2009; De Mattia leg.; WdM6805 • Bresovica; 45.95607° N,
14.43533° E; 2006; De Mattia leg.; EZ0897 • Bresovica pri Predgradu; 45.55164° N, 15.0437° E; 2009;
De Mattia leg.; WdM6612 • Divje Jezero; 45.9827° N, 14.0277° E; 2009; De Mattia leg.; WdM6840 •
Dol-E of Čepovan; 46.06445° N, 13.81189° E; 2009; De Mattia leg.; WdM6837 • Jelenja Vas; 45.5115° N,
15.05346° E; 2009; De Mattia leg.; WdM6804 • Krempa Kolpa; 45.53417° N, 15.09528° E; 1996; Mildner
leg.; EZ0902 • Marija Ceplje Kolpa; 45.53444° N, 15.09389° E; 2006; Mildner leg.; EZ0901 • Mt Snežnik
1150; 45.56913° N, 14.321° E; Jun. 2010; De Mattia leg.; WdM7014 • Mt Snežnik 1620; 45.58845° N,
14.44209° E; Jun. 2010; De Mattia leg.; EZ0963 • Mt Snežnik car parking; 45.58235° N, 14.43117° E; Jun.
2010; De Mattia leg.; WdM7006 • Polog-E of Čepovan; 46.06387° N, 13.81275° E; 2009; De Mattia leg.;
WdM6842 • SW Pescena Glava; 46.05156° N, 13.73844° E; 843 m a.s.l.; 2009; De Mattia leg.; WdM6829

European Journal of Taxonomy 762: 49–95 (2021)
88
• Tisovec; 45.81809° N, 14.76859° E; 2009; De Mattia leg.; WdM6749 • same collection data as for
preceding; WdM6750 • Vrsic Pass; 46.44278° N, 13.76417° E; 1985; Zallot leg.; EZ0896 • Zagradec-
Polyane; 45.86028° N, 14.83417° E; Bodon leg.; EZ0900.
Description
Shell. The shell is more or less light corneous brown and spotless. The size of the aperture varies
considerably among the populations; shells with a large aperture and a correspondingly large lip occur
next to shells with a poorly developed lip and a relatively small aperture. The lip is gently curved
backwards on the columellar side.
Female genitalia. The female genitalia are hardly distinguishable from those of Co. (Cl.) tergestinum
if not for the more relaxed loops.
male genitalia. There is a long, inated penis. The penial spermiduct is slightly twisted and occupies
only the central part of the penis. The body spermiduct is twisted and conspicuously protruding (see
Figs 34–35).
Fig. 31. Shell variability in Cochlostoma (Clessiniella) waldemari (A.J. Wagner, 1897). A. Polog-E of
Čepovan, Slovenia (WdM6842). B. Vrsic Pass, Slovenia (EZ0896). C. Benete, Slovenia (WdM6805).
D. Mt Klek, Croatia (EZ0904). E. Bresovica pri Predgradu, Slovenia (WdM6612).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
89
Distribution
The species inhabits the north-western and south-eastern part of Slovenia and an adjacent area in Croatia.
It is reported from Austria for the Karawanken and the Gailtaler Alpen (Edlinger & Mildner 1979). Its
range largely surrounds that of Co. (Cl.) tergestinum (see Fig. 1).
Habitat preference
The snails are found on rocky cliffs as well as on boulders in woody areas.
Remarks
To assign a given population to Co. (Cl.) waldemari rather than to Co. (Cl.) tergestinum on the basis
of the morphology of the shell and the female genitalia alone could be challenging if not impossible
(see Figs 34–35). Only the structure of the male genitalia allows a convincing identication of the
species. The H3 analysis (Fig. 7) conrms that these sibling species belong to separate clades within
Clessiniella.
Boeckel (1939) considered Co. (Cl.) waldemari a subspecies of Co. (Cl.) tergestinum. This view was
contradicted by Bole (1994). In some areas, like on Mount Snežnik in southern Slovenia, the two species
live not far from each other and here they can be distinguished also conchologically while differing in
the presence or absence of the spots on the teleoconch (with Co. (Cl.) waldemari having a spotless shell)
and in the shape of the body whorl approaching the aperture, in lateral view (Fig. 36).
Fig. 32. Variation of the female genitalia in (A.J. Wagner, 1897). A. Polog-E of Čepovan, Slovenia
(WdM6842). B. Dol-E of Čepovan, Slovenia (WdM6837). C. Divje Jezero, Slovenia (WdM6840).
D. Tisovec, Slovenia (WdM6749). E. Vrsic Pass, Slovenia (EZ0896). F. Benete, Slovenia (WdM6805).
G. Vitunin spring, Croatia (EZ0903). H. Mt Klek, Croatia (EZ0905).

European Journal of Taxonomy 762: 49–95 (2021)
90
Discussion
The lack of clear diagnostic features in shell morphology, with intraspecic sometimes overlapping
interspecic variability as in Pyrenean Cochlostomatidae (Gofas 2001), is one of the reasons of
the problematic taxonomy at the species level within the genus Cochlostoma. Based on only shell
morphology, several populations of 4 of the 5 species of Clessiniella cannot be identied with certainty.
The lack of clear distinguishing shell features may explain why a species like Co. (Cl.) stelucarum
sp. nov. went unnoticed in an area that may be considered well explored. The multivariate analyses of
Fig. 33. Cochlostoma (Clessiniella) waldemari (A.J. Wagner, 1897), syntype, ♀, at the Muzeum i
Instytut Zoologii PAN, Warsaw, Poland (MIZPAS-W-6979a).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
91
Fig. 34. Example of two populations of species of Clessiniella Zallot et al., 2015 which are
impossible to distinguish without analyzing the male genitalia. Female shells. Top: Cochlostoma
(Clessiniella) waldemari (A.J. Wagner, 1897), Polog-E of Čepovan, Slovenia (WdM6842). Bottom:
Co. (Cl.) tergestinum (Westerlund, 1878), Jakačici, Croatia (WdM6699).

European Journal of Taxonomy 762: 49–95 (2021)
92
the shell measurements also fail to unequivocally diagnose the species. The shell features that cannot
be quantied may be helpful for identication, but are hardly decisive, except for the identication of
Co. (Cl.) anomphale. There is no general, always satisfying, single morphological dataset that enables
indisputable identications. The analysis of the female genital morphology sets apart Co. (Cl.) anomphale
and Co. (Cl.) stelucarum sp. nov. from the other species but it does not clarify everything in Cochlostoma
taxonomy. Similarly, the investigation of the male genitalia reveals a taxonomical relevant feature for
the diagnosis of Co. (Cl.) waldemari but not of the other species.
A combined analysis of shell and genitalia is needed to clarify the taxonomic relationships within the
genus in a satisfactory way. It should not be overlooked, however, that the results of our approach do not
Fig. 35. Example of two populations of species of Clessiniella Zallot et al., 2015 which are impossible
to distinguish without analyzing the male genitals. Male shells. Top: Cochlostoma (Clessiniella)
waldemari (A.J. Wagner, 1897), Mt Klek, Croatia (EZ0904). Bottom: Co. (Cl.) tergestinum (Westerlund,
1878), Val Rosandra, Italy (EZ0840).

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
93
fundamentally differ from the views of our predecessors except for the phylogenetic relations amongst
taxa. Three species of Cochlostoma (Clessiniella) got their names in the 19th century, one in the 20th
century, and one species was discovered only in modern times.
Acknowledgements
The authors are grateful to Dr Daniele Giannetti and Dr Cristina Menta of the Universita’ degli studi di
Parma (Italy) for the kind collaboration in nding and taking pictures of Strobel’s material. We thank
Roberta Salmaso of the Museo civico di Storia Naturale di Verona (Italy) for all the information and
pictures of the De Betta material. We thank Sigrid Hof of the Senckenberg Forschungsinstitut for the
photos of the types of Co. (Cl.) anomphale and Co. (Cl.) tergestinum. We are grateful to Dr Fernando
Scarlassara for the precious information about Co. (Cl.) villae.
Fig. 36. Top: Cochlostoma (Clessiniella) tergestinum (Westerlund, 1878), Mt Snežnik 1161 m a.s.l.,
Slovenia (WdM7012). Bottom: Co. (Cl.) waldemari (A.J. Wagner, 1897), Mt Snežnik 1150 m a.s.l.,
Slovenia (WdM7014).

European Journal of Taxonomy 762: 49–95 (2021)
94
References
Alzona C. 1971. Malacofauna Italica. Catalogo e bibliograa dei molluschi viventi, terrestri e d’acqua
dolce. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
111: 1–433.
Bank R.A. 1988. Revision der nordostitalienischen Arten und Unterarten der Gattung Cochlostoma Jan,
1830 (Gastropoda Prosobranchia: Cyclophoridae). Basteria 52 (4/6): 151–170.
Boeckel W. 1939. Cochlostoma-Arten aus den Karawanken. Archiv für Molluskenkunde 71: 41–50.
Bole J. 1994. Rod Cochlostoma Jan 1830 (Gastropoda, Prosobranchia, Cochlostomatidae) v Sloveniji.
Razprave [Dissertationes], Slovenska Akademija Znanosti in Umetnosti [Academia Scientiarum et
Artium Slovenica], Razred za Naravoslovne Vede [Classis 4] 35 (11): 187–217.
Clement M., Snell Q., Walker P., Posada D. & Crandall K. 2000. TCS: estimating gene genealogies.
Proceedings 16th International Parallel and Distributed Processing Symposium 2002: 7.
https://doi.org/10.1109/IPDPS.2002.1016585
Colgan D.J., Ponder W.F. & Eggler P.E. 2000. Gastropod evolutionary rates and phylogenetic relationships
assessed using partial 28S rRNA and histone H3 sequences. Zoologica Scripta 29: 29–63.
https://doi.org/10.1046/j.1463-6409.2000.00021.x
De Betta E. & Martinati P. 1855. Catalogo dei Molluschi Terrestri e Fluviatili Viventi nelle Provincie
Venete di Edoardo de Betta e Pietropaolo Martinati. G. Antonelli, Verona.
https://doi.org/10.5962/bhl.title.13113
De Mattia W. & Prodan M. 2006. Nuovi dati sulla distribuzione di Cochlostoma (Turritus) tergestinum
tortivum (Westerlund, 1885) e Cochlostoma (Cochlostoma) scalarinum scalarinum (Villa & Villa, 1841)
in Italia (Gastropoda: Prosobranchia: Cochlostomatidae). Atti del Museo Civico di Storia Naturale di
Trieste 52: 205–213.
Edlinger K. & Mildner P. 1979. Monographie der in Kärnten lebenden Prosobranchiergattung.
Cochlostoma. Carinthia II 169: 281–304.
Fehér Z. & Erõss Z. 2009. Checklist of the Albanian mollusc fauna. Schriften zur Malakozoologie 25:
22–38.
Gittenberger E. 1971. Zur näheren Kenntnis von Cochlostoma (Turritus) anomphale Boeckel
(Prosobranchia-Cyclophoridae). Basteria 35 (6): 115–118.
Giusti F. & Selmi M.G. 1985. The seminal receptacle and sperm storage in Cochlostoma montanum
(Issel) (Gastropoda: Prosobranchia). Journal of Morphology 184: 121–133.
https://doi.org/10.1002/jmor.1051840204
Gofas S. 2001. The systematics of Pyrenean and Cantabrian Cochlostoma (Gastropoda, Cyclophoroidea)
revisited. Journal of Natural History 35 (9): 1277–1369. https://doi.org/10.1080/002229301750384301
Hartmann M., Grimm H. & Scheidt U. 2018. ERFURT: the Naturkundemuseum Erfurt. In: Beck L. (ed.)
Zoological Collections of Germany. Natural History Collections: 295–309. Springer, Cham.
https://doi.org/10.1007/978-3-319-44321-8_26
Leigh J.W. & Bryant D. 2015. POPART: full-feature software for haplotype network construction.
Methods in Ecology and Evolution 6: 1110–1116. https://doi.org/10.1111/2041-210X.12410
MolluscaBase eds. 2021. MolluscaBase. Available from http://www.molluscabase.org [accessed 24 Apr.
2021]. https://doi.org/10.14284/448
Strobel P. 1851. Notizie Malacostatiche sul Trentino. Fusi, Pavia. Available from
https://www.biodiversitylibrary.org/page/46194631 [accessed 12 Jul. 2021].

ZALLOT E. et al., Revision of Clessiniella (Caenogastropoda)
95
Uit de Weerd D.R. & Gittenberger E. 2013. Phylogeny of the land snail family Clausiliidae (Gastropoda:
Pulmonata). Molecular Phylogenetics and Evolution 67: 201–216.
https://doi.org/10.1016/j.ympev.2013.01.011
Wagner A.J. 1897. Monographie der Gattung Pomatias Studer. Denkschriften der kaiserlichen Akademie
der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe 64: 565–632.
https://doi.org/10.5962/bhl.title.10792
Welter-Schultes F.W. 2012. European non-marine Molluscs, a Guide for Species Identication. Planet
Poster Editions, Göttingen.
Westerlund C.A. 1878. Diagnosen neuer Mollusken. Nachrichtsblatt der Deutschen Malakozoologischen
Gesellschaft 10: 108–109.
Westerlund C.A. 1885. Fauna der in der paläarctischen Region (Europa, Kaukasien, Sibirien, Turan,
Persien, Kurdistan, Armenien, Mesopotamien, Kleinasien, Syrien, Arabien, Egypten, Tripolis, Tunesien,
Algerien und Marocco) lebenden Binnenconchylien. V. Fam. Succinidæ, Auriculidæ, Limnæidæ,
Cyclostomidæ & Hydrocenidæ. Lund.
Zallot E. 2002. Alcune note sul genere Cochlostoma Jan, 1830 (Gastropoda, Prosobranchia) in Friuli
(Italia nord-orientale). Gortania 24: 93–113.
Zallot E., Groenenberg D.S.J., De Mattia W., Fehér Z. & Gittenberger E. 2015. Genera, subgenera and
species of the Cochlostomatidae (Gastropoda, Caenogastropoda, Cochlostomatidae). Basteria 78 (4–6):
63–88.
Zallot E., Fehér Z., Bamberger S. & Gittenberger E. 2018. Cochlostoma revised: the subgenus Lovcenia
Zallot et al., 2015 (Caenogastropoda, Cochlostomatidae). European Journal of Taxonomy 464: 1–25.
https://doi.org/10.5852/ejt.2018.464
Zilch A. 1958. Die Typen und Typoide des Natur-Museums Senckenberg, 21: Mollusca, Cyclophoridae,
Craspedopominae-Cochlostominae. Archiv für Molluskenkunde 87 (1/3): 53–76.
Manuscript received: 10 December 2020
Manuscript accepted: 10 June 2021
Published on: 6 August 2021
Topic editor: Rudy Jocqué
Section editor: Thierry Backeljau
Desk editor: Pepe Fernández
Printed versions of all papers are also deposited in the libraries of the institutes that are members of the
EJT consortium: Muséum national d’histoire naturelle, Paris, France; Meise Botanic Garden, Belgium;
Royal Museum for Central Africa, Tervuren, Belgium; Royal Belgian Institute of Natural Sciences,
Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark; Naturalis Biodiversity
Center, Leiden, the Netherlands; Museo Nacional de Ciencias Naturales-CSIC, Madrid, Spain; Real
Jardín Botánico de Madrid CSIC, Spain; Zoological Research Museum Alexander Koenig, Bonn,
Germany; National Museum, Prague, Czech Republic.