The Radomaniola/Grossuana group from the Balkan Peninsula, with a description of Grossuana maceradica n. sp. and the designation of a neotype of Paludina hohenackeri Küster, 1853 (Caenogastropoda: Truncatelloidea: Hydrobiidae)

Abstract

Grossuana maceradica n. sp. from the Radika drainage, Macedonia is described. A lectotype is selected for Amnicola marginata Westerlund, 1881, and a neotype is designated for Paludina hohenackeri Küster, 1853. Photographs of type material of these and 2 other Balkan taxa, Amnicola filiola Westerlund, 1881 and Hydrobia haesitans Westerlund, 1881, are published here for the first time. © E. Schweizerbart'sche Verlagsbuchhandlung (Nägele u. Obermiller) and Senckenberg Gesellschaft für Naturforschung, 2017.

Full text

Archiv für Molluskenkunde 146 (2) 187–202 Frankfurt am Main, 20 Dec. 2017
© E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller) and Senckenberg Gesellschaft für Naturforschung, 2017, ISSN 1869-0963
DOI 10.1127/arch.moll/146/187-202
The Radomaniola/Grossuana group from the Balkan Peninsula,
with a description of Grossuana maceradica n. sp. and the
designation of a neotype of Paludina hohenackeri Küster, 1853
(Caenogastropoda: Truncatelloidea: Hydrobiidae)
H D. B, P G & V S S
1 Karneidstr. 8, 81545 München, Germany (boeters@t-online.de). 2 Biodiversity Research Laboratory, Schulstr. 3, 25491 Hetlingen, Germany
(gloeer@malaco.de). 3 Department of Invertebrates and Animal Ecology, Institute of Biology, Faculty of Natural Sciences and Mathematics,
SsCyril and Methodius University, Arhimedova 3, 1000 Skopje, Macedonia (vstamen@yahoo.com). • Corresponding author: H.D. Boeters.
Abstract. Grossuana maceradica n. sp. from the Radika drainage, Macedonia is described. A lectotype is
selected for Amnicola marginata Westerlund, 1881, and a neotype is designated for Paludina hohenackeri
Küster, 1853. Photographs of type material of these and 2 other Balkan taxa, Amnicola liola Westerlund,
1881 and Hydrobia haesitans
Key words. Paludina hohenackeri, Amnicola liola, Amnicola marginata, Hydrobia haesitans, ende-
mism in the Balkans, Radika river, Crni Drim river.
DOI. https://doi.org/10.1127/arch.moll/146/187-202
Introduction
Species of Belgrandiella A.J. Wagner, 1928 are distrib-
uted from southern France ( 1970, 2008), eastern
Austria ( 1994), northern Italy (&
1980,  1994), and through the Dinarides (-
 1975). They have also been reported from Bulgaria
( 1959, 1972, 1976,  2011, 2013,
& 2009,  1968,  [1928])
and Greece ( [1980]). Although 
described a Greek species as Orientalia delphica (now
Radomaniola delphica (Radoman, 1973)), 
[1980] insisted that it belongs to Belgrandiella. -
 (1985: 64) wrote, “We, however, have not found any
of its representatives either in the Hellenides, the south-
ern Balkan peninsula, Macedonia or Greece, nor in Asia
Minor”. Instead, he believed that Greece is inhabited by
representatives of Grossuana, such as G. serbica vurliana
Radoman, 1966, and of Radomaniola, such as R. curta
albanica Radoman, 1973 and R. delphica (Radoman,
1973) ( 1983). However, he did not consider
relevant earlier-named taxa. Recently  et al.
(2016) gave evidence, based on genetic analyses, that
Bulgaria is not inhabited by species of Belgrandiella, but
rather by Pontobelgrandiella Radoman, 1973.
To clarify the identity of a recently detected unknown
hydrobiid snail from Mavrovo National Park, Macedo-
nia, within the Crni Drim river drainage, we analysed
all known Balkan and Greek species of Belgrand iella,
Pontobelgrandiella, Radomaniola, and Grossuana.
Material and Methods
Abbreviations of collections.
BOE Boeters Collection (München)
CHARP Charpentier Collection (Lausanne)
GLÖER Glöer Collection (Hetlingen)
NMW Naturhistorisches Museum (Wien)
REI Peter & Alexander Reischütz Collection (Horn)
SMF Forschungsinstitut Senckenberg (Frankfurt am
Main)
WEST Westerlund Collection (Göteborg)
Examined material. Spring region of the Rosochka reka

level, Radika drainage, Nacionalen park mavrovo [Mav-
rovo National Park], about 12 km E of Debar, Macedonia;
    GLÖER/2
animals; BOE 3366/7 animals dissected + 2 mature + 1
juvenile animal; SMF 349125/1 animal (holotype).
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The snails were collected using a hand-net with a mesh
size of 500 µm and preserved in 75% ethanol. Whorls
were counted following  et al. (1970). An-
atomical investigations were done in accordance with
 (1999). Drawings of shells and animals were
made with a Zeiss 45° drawing prism. Length and diam-
eter of shells were measured with a 5 mm grid measure
      -
ments were rounded to the nearest 0.05 mm. Shells were
photographed using a Leica R8 camera with a Digital-
Modul-R.
Systematics. To show that the new species belongs to Gros -
suana, we have taken 13 genera with similar morphologi-
cal and anatomical characters into consideration (Table 1).
To justify the new species, we compare it to known
representatives of Belgrandiella A.J. Wagner, 1928, Pon-
tobelgrandiella Radoman, 1973, Grossuana Radoman,
1973, and Radomaniola Szarowska, 2006 because the
assignment of known species to one of these genera is
      
1–4, Appendices 1–4).
Thus, in addition to the newly decribed Grossuana
species of the Radomaniola/Grossuana group sensu -
et al. (2012), we summarize data on the types,
type localities, anatomy, and genetics of representatives
of Belgrandiella, Pontobelgrandiella, Radomaniola, and
Grossuana from the Balkan Peninsula (Table 1, Appen-
dices 1–4).
Despite having a penis with a simple outgrowth and
the presence of RS1 and RS2, as in Grossuana, Daph-
niola Radoman, 1973 is not mentioned in Table 1 because
D. graeca Radoman, 1973, the type species of this genus
and a junior synonym of Valvata exigua A. Schmidt,
1856, is characterised by a valvatiform shell (
 et al. 2001: 111).
Table 1. Comparison of Belgrandiella and related genera.
Genus Type species Intestine in roof
of mantle cavity
Caecum of
stomach Penis Receptaculum
Belgrandiella A.J. Wagner, 1928 aBelgrandia kusceri A.J. Wagner, 1914 Simply bent Missing Simple outgrowth RS1
Boleana Radoman, 1973
bBelgrandiella umbilicata Kuščer, 1932 Simply bent Missing Simple outgrowth RS1
Graecorientalia Radoman, 1973 cPseudamnicola vrissiana Radoman, 1966 ? ? Furcated outgrowth RS1+RS2
Grossuana Radoman, 1973 dGrossuana serbica Radoman, 1973 Simply bent ? Simple outgrowth RS1+RS2
Trichonia Radoman, 1973 eTrichonia kephalovrissonia Radoman, 1973 ? ? Furcated outgrowth RS1+RS2
Turkorientalia Radoman, 1973 fTurkorientalia anatolica Radoman, 1973 ? ? Weakly furcated outgrowth RS1+RS2
Graziana Radoman, 1975 gPaludina lacheineri Küster, 1853 ? Present Simple outgrowth RS1
Sarajana Radoman, 1975 hFrauenfeldia lacheineri apfelbecki Brancsik, 1888 ? ? Simple outgrowth RS1
Cavernisa Radoman, 1978 iBelgrandiella zaschevi Angelov, 1959 ? ? Simple outgrowth RS1
Pontobelgrandiella Radoman, 1978 jBelgraniella nitida Angelov 1972 Simply bent ? Two outgrowths RS1
Terranigra Radoman, 1978 kTerranigra kosovica Radoman, 1978 ? ? Simple RS1+RS2
Alzoniella Giusti & Bodon, 1984 lAlzoniella finalina Giusti & Bodon, 1984 Z-like loop Missing Basal and medial outgrowth RS1+RS2
Radomaniola Szarowska, 2006
pro Orientalina Radoman, 1978
pro Orientalia Radoman, 1972 m
Paludina curta Küster, 1853 ? ? Furcated outgrowth RS1+RS2
a Male genitalia of Belgrandia kusceri: B (1967: fig. 2A), R (1975: fig. 1, 1983: fig. 50). Female genitalia of B. kusceri: B
(1967: fig. 2A) and R (1975: fig. 1, 1983: fig. 50). In all species with investigated female genitalia (B. crucis, B. kusceri, B.
robusta, B. schleschi, B. substricta, B. superior, and B. umbilicata) the receptaculum (RS1) touches the bursa (B 1967: fig. 1B3, 82
fig. 2A3, R 1975: fig. 2, B 1967: fig. 3B3, fig. 5B3, fig. 2B3, fig. 2C3). Based on F & B (2015), B. krupensis
and B. zermanica can be included. Distribution in the Balkans: R (1975: fig. 11) and F & B (2015: fig. 1).
b According to F & B (2015), Boleana is a synonym of Belgrandiella. Male and female genitalia of B. umbilicata: B
(1967: fig. 2C) and R (1983: fig. 53).
c This genus might preoccupy Radomaniola Szarowska, 2006, but more study is needed; compare R’s (1983) figures 17 and 18
of R. curta and G. vrissiana, respectively.
d Intestine of Grossuana codreanui: S et al. (2007: figs 15, 16). Male and female genitalia of G. serbica: R (1983: fig. 24).
e F et al. (2012: fig. 14) referred to Radomaniola sensu stricto a genetially analysed sample of the type species, Trichonia
kephalovrissonia, but not from the type locality. Male and female genitalia of T. kephalovrissonia: R (1983: fig. 43).
f Male and female genitalia of Turkorientalia anatolica: R (1983: fig. 49).
g Caecum of Graziana acheineri at the proximal end of stomach: H (1994: fig. 7), but not confirmed by S (2006: figs
141–142, 153–157, 243–244). Male and female genitalia of G. lacheineri: B (1967: fig. 1A), R (1975: fig. 3, 1983: fig. 51),
and H (1994: figs 5B, 6E, F).
h A junior synonym of Belgrandiella. Male and female genitalia of Sarajana apfelbecki: R (1975: fig. 6, 1983: fig. 52).
i See Appendix 2, footnote 4. Male and female genitalia of Cavernisa zaschevi: R (1983: fig. 60).
j Intestine of Pontobelgrandiella pandurskii: G (2011a: fig. 1.1). Male and female genitalia of P. nitida: R (1983: fig. 59).
Male genitalia of P. tanevi: G (2013: figs 3, 4).
k Male and female genitalia: R (1978: fig. 1, 1983: fig. 30).
l Stomach and intestine of Alzoniella finalina: G & B ([1984]: fig. 2G, H). Male and female genitalia of A. finalina: G &
B ([1984]: figs 2A–F, 2G). The relationship between Alzoniella and Terranigra has not yet been discussed.
m Male and female genitalia of Radomaniola curta: R (1983: fig. 17).
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Figure 1. Type localities of species of Belgrandiella; red dot:
Grossuana maceradica n. sp. Localities: 1, B. apfelbecki
(Brancsik, 1888). 2, B. bozidarcurcici Glöer & Pešić, 2014. 3,
B. bumasta Schütt, 1960. 4, B. conica Radoman, 1975. 5, B.
croatica (Hirc, 1881). 6, B. crucis (Kuščer, 1928). 7, B. dabriana
Radoman, 1975. 8, B. driniana (Radoman, 1975). 9, B. erythro-
poma Schütt, 1959. 10, B. fontinalis (F.J. Schmidt, 1847). 11,
B. globulosa Bole, 1979. 12, B. hershleri Slapnik, 1997. 13, B.
hyalis Westerlund, 1886. 14, B. koprivnensis Radoman, 1975.
15, B. krupensis Radoman, 1973. 16, B. kusceri (A.J. Wag-
ner, 1914). 17, B. novoselensis Radoman, 1975. 18, B. pageti
Schütt, 1970. 19, B. robusta Radoman, 1975. 20, B. schleschi
Kuščer, 1932. 21, B. substricta (Kuščer, 1932). 22, B. superior
Kuščer, 1932. 23, B. travnicensis (Radoman, 1975). 24, B. um-
bilicata Kuščer, 1932. 25, B. zermanica Radoman, 1973.
Figure 4. Type localities of species of Radomaniola and of
Graecorientalia; red dot: Grossuana maceradica n. sp. Locali-
ties: 1, R. (?) aytosensis Georgiev, 2012. 2, R. bosniaca Rado-
man, 1973. 3, R. bulgarica Glöer & Georgiev, 2009. 4, R. curta
(Küster, 1853). 5, R. elongata Radoman, 1973. 6, R. (?) filiola
(Westerlund, 1881). 7, R. (?) haesitans (Westerlund, 1881). 8,
R.(?) hessei (Kobelt, 1891). 9, R. lacustris Radoman, 1983. 10,
R. montana Radoman, 1973. 11, R. radostinae Georgiev, 2012.
12, R. rhodopensis Glöer & Georgiev, 2009. 13, R. seminula
(Frauenfeld, 1863). 14, R. strandzhica Georgiev & Glöer, 2013.
15, R. tritonum Bourguignat, 1852. 16, Graecorientalia vris-
siana (Radoman, 1966).
Figure 3. Type localities of Grossuana species in the Balkans;
red dot: Grossuana maceradica n. sp. Localities: 1, G. angelt-
sekovi Glöer & Georgiev, 2009. 2, G. codreanui (Grossu, 1946).
3, G. delphica Radoman, 1973. 4, G. derventica Georgiev &
Glöer, 2013. 5, G. euxina (A.J. Wagner, 1928). 6, G. falniow-
ski Georgiev, Glöer, Dedov & Irikov, 2015. 7, 8, not shown,
see Appendix 3 for G. hohenackeri (Küster, 1853) and G. mar-
ginata (Westerlund, 1881). 9, G. serbica Radoman, 1973. 10,
G. slavyanica Georgiev & Glöer, 2013. 11, G. thracica Glöer &
Georgiev, 2009. 12, G. vurliana (Radoman, 1966).
Figure 2. Type localities of species of Pontobelgrandiella;
red dot: Grossuana maceradica n. sp. Localities: 1, P. angelovi
(Pintér, 1968). 2, P. bachkovoensis (Glöer & Georgiev, 2009).
3, P. bulgarica (Angelov, 1972). 4, P. bureschi (Angelov, 1976).
5, P. dobrostanica (Glöer & Georgiev, 2009). 6, P. hessei (A.J.
Wagner, 1928). 7, P. (?) maarensis (Georgiev, 2013). 8, P. nitida
Angelov, 1972. 9, P. pandurskii (Georgiev, 2011). 10, P. pusilla
(Angelov, 1959). 11, P. stanimirae (Georgiev, 2011). 12, P. tanevi
Georgiev, 2013. 13, P. zagoraensis (Glöer & Georgiev, 2009).
Systematics
Grossuana maceradica n. sp.
Figures 5A–C, 6
Holotype. SMF 349125.
Paratypes. GLÖER/2 specimens; BOE 3366/7 specimens.
Type locality. Spring region of the Rosochka reka [river]
  

nacionalen park Mavrovo [Mavrovo National Park], c.
12 km E of Debar, Macedonia.
The Rosochka reka is a river in the Radika drainage,
which is a right tributary of the Crni Drim [=Black Drim],

Etymology. The species’ name is a composite word and
refers to Macedonia and the Radica river.
Shell (Fig. 5A–C). Ovoid, transparent, smooth; whorls

neither ascending nor descending on shell wall; parietal
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Figure 5. A–C. Grossuana maceradica n. sp. Macedonia, Rosochka reka; V. Slavevska-Stamenković leg. 2015: (A, C) BOE 3366/2
paratypes; (B) SMF 349125/holotype). D–G. Grossuana (?) hohenackeri (Küster, 1853), shells: (D) neotype (CHARP); (E, F) 2 other
shells (CHARP); (G) shell [K 1853: pl. 10 fig. 20; height of shell 1.69 mm]). H–J. Grossuana (?) marginata (Westerlund, 1881).
“Graecia ad Avlochades”, Evia [= Euboea]; WEST (Göteborg)/4526: (H) lectotype.; (I, J) 2 paralectotypes. K, L. Radomaniola (?)
filiola (Westerlund, 1881). “Graecia, Patras, Fonte Salevale”; WEST (Göteborg)/4527 (2 syntypes?). M, N. Radomaniola (?) haesi-
tans (Westerlund, 1881), “Graecia, Santa Maura, Megali Vressi”; WEST (Göteborg)/4385: (M) syntype; (N) syntype?).
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border of aperture fused with shell wall; palatal and basal
border of aperture neither broadened nor thickened; colu-
mellar border slightly broadened; umbilicus closed by
columellar border or narrowly open as a slit.
Measurements: height 1.97–2.33 mm, mean 2.11 mm
(N = 4); diameter 1.44–1.58 mm, mean 1.51 mm (N = 4);
height:diameter = 1.40.
Operculum. Light brownish, transparent, showing an un -

Animal (Fig. 6). Head, mantle and body whorls pig-
N
simply bent in the roof of the mantle cavity (N = 4).
Males:females = 2:5 (N = 7).

top, formed like a long wedge provided with a hook-like
middle appendix. Penial duct runs laterally down to the
tip of the penis on the opposite side of the appendix. Base
of penis shows in its inactive state bellows-like folds (N =
2) in front of appendix.
Female genitalia: bursa spherical, provided with a
long duct, distal spherical receptaculum (RS1) with a
pronounced duct, which, however, is too short to allow re-
ceptaculum to touch bursa; proximal receptaculum (RS2)
       
front of its turn at bursa and accompanies renal oviduct to
the turn, where proximal receptaculum touches bursa (N
= 2). Proximal receptaculum iridescent, a muscular tube.
Habitat and distribution (Fig. 7). Known only from the
type locality. Here, the Rosochka reka is a small moun-
tain stream with clean, cold, fast running water. Living
animals were found on limestone substrates, mostly on
stones covered with aquatic mosses.
 Taxa of the Radomaniola/Gros-
suana group reported from neighbouring areas are R.
curta kicavica Radoman, 1973, R. curta albanica Rado-
man, 1973, G. serbica serbica macedonica Radoman,
1973, and G. serbica scupica Radoman, 1973.
According to 
subspecies of R. curta occur in the area of the Adriatic
drainage, and R. c. kicavica allegedly even in Crni Drim
river drainage ( 1983), the same drainage
as the new species. The shells of both subspecies of R.
Figure 6. Grossuana maceradica n. sp. Macedonia, Rosochka reka; V. Slavevska-Stamenković leg. 2015; BOE 3366/paratypes.
A, B. Shells, ♂. C. Shell, ♀. D. Same ♂ as A, head, mantle cavity opened to show penis and gill. E. Same ♂ as B, penis. F. Same
♀ as C, renal oviduct with RS1 and bursa, intestine partially broken away. G. ♀, renal oviduct with bursa, oviduct slightly turned
to show RS2, intestine partially broken away. Abbreviations: BC = bursa copulatrix; GI = gill; IN = intestine; OS = osphradium;
PE = penis; RO = renal oviduct; RS1 = distal receptaculum seminis; RS2 = proximal distal receptaculum seminis; VS = vas
derens. Scale bar: 2 mm in A–C, 1 mm in D–G.
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192
Archiv für Molluskenkunde · 146 (2) 2017
curta are more elongate than shells of the new species,
height : diameter 1.70 and 1.80 ( 1983: pl. 2

of R. curta  
top) outgrowth on the left side” (
41, G. maceradica
n. sp. the penis shows a simple hook-like outgrowth.
Finally, in females of the new species, the proximal
receptaculum (RS2) is tube-like and not, as in R. curta,
drop-like and borne by a short duct (
17).
Whereas 
penis of G. serbica as “long, with a very prolonged,
pointed top and a weak outgrowth on the left side, which
is hardly visible in many specimens”, in G. maceradica
n. sp. the penis shows a prominent hook-like outgrowth.
 (1983: 59) described the shell of G. s.
macedonica as “roundish, more shortened than that in all
up to now known subspecies of the genus, with weakly
     G.
maceradica n. sp. the shell is somewhat more elongate,
which is expressed by a height:diameter ratio of 1.40
instead of 1.35 (
In G. s. scupica the axis of the ovoid aperture is less
inclined to the axis of the shell than in G. macera dica n.
sp. (i.e., 18° vs 20° ).
The type localities of G. s. macedonica and G. s. scu-
pica are both within the Vardar drainage. This system
is part of the Aegean drainage area and not the Adriatic
drainage area where G. maceradica n. sp. occurs.
Discussion
Discrimination of spring snails
Geological and topographical aspects. Common
wisdom has it that species of spring snails are locally
restricted to small areas, as for example in Belgrandi-
ella. Recently, however,  &  (2015)
critically discussed this view with regards to 4 nominal
Balkan taxa. They argued that shells alone cannot be the
basis for species discrimination in the Truncatelloidea.
But they also cited  (1983) as claiming that
-
niowski’s and Beran’s view, based on their interpretation
of Radoman, should be adopted not just for Belgrandiella
but for other so-called spring snails.
-
ible, but that spring snails may indeed be geographically
restricted to very small areas has been corroborated, for
example, for Bythinella badensis (Boeters, 1981), based
on conchological, anatomical, genetic, and geographical
data. Although Bythinella dunkeri (Frauenfeld, 1857)
and B. badensis inhabit the same massif, B. badensis
only occurs in a small southern part of the Schwarzwald.
These species were isolated from each other in pre-
Quaternary times because the southern Schwarzwald was

the Rhine as today. Thus, Radoman’s approach of using
drainage systems as a guide to distribution patterns can
be useful.
Grossuana maceradica n. sp. is the only species of
Grossuana yet known from an Adriatic drainage. All
neighbouring localities reported by  
1985) and corroborated by 
1M) for Grossuana are within the Crna reka drainage,

In the Radika drainage only 2 Radomaniola/Gros-
suana taxa are known. These are G. maceradica n. sp.
and R. R. curta kicavica. According to Radoman (1983:
44), R. c. kikavica is “Spread … in springs in drainage
areas of the rivers Radika, and Crni Drim [entered by the
Radika at Debar], including some springs in the Ohrid
Basin … and in the Prespa Basin”. However, the type
locality of R. c. kicavica is “Izvor, a big spring about 16
km [west] from Kicevo town” ( 1983: 44), and
the Izvor spring supplies the Treska river, which belongs
to the Aegean drainage and not to the Adriatic drainage,
as does the Radika.
 (1985: 110) described the Drim as “mainly
a mountain river, about 300 km long, with many whirl-
pools, rapids and waterfalls ... naturally it contains no
marsh or spring forms except for Theodoxus uviatilis
... this river could not have played the role of an immi-
gration way ...”. This view might apply to species which
have not invaded interstitial habitats.
Concerning the alleged occurrences of R. curta kica-
vica in springs that supply Lake Ohrid and R. c. curta in
the Zeta river drainage ( 1853,  1983,
 et al. 2012),  (1985: 111) argued
that “Ohrid non-endemic forms”, such as R. c. kicavica,
“date in this area from the remote past of this basin [Lake
Ohrid], from the time of its broad intercommunication
with neighbouring waters at a very low altitude above sea
level”. However, with regard to an invasion of the Drim
drainage by the Radomaniola/Grossuana group, 
Figure 7. Type locality of Grossuana maceradica n. sp., Mavrovo
National Park, Macedonia. Photograph: D. Jovanovska.
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B et al. · Radomaniola/Grossuana of the Balkans and Greece
et al. (2004: map 10) showed that in the Middle–Late
Pliocene (3.4–1.8 Myr) the southernmost foothills of the
-
water lakes and marshes.
It is strange that  (1983: 44) thought that R.
c. kicavica occurs in 2 separate drainages. Even if passive
transport by birds or water beetles ( 1979, 1982,
1993) is not excluded, it is questionable whether his R.
c. kicavica from Radika and Crni Drim river drainages
is the same as that from its type locality in Treska river
drainage.
From ’s (1983) ideas on speciation in the
Lake Ohrid basin, it might be concluded that the ances-
tors of G. maceradica n. sp. immigrated to its cur rent
locality “in the remote past”, and that it is an endemic,
not non-endemic, species. This might explain the unique
characters seen in males of this species.
Anatomical aspects.
species of spring snails, such those of the Radomaniola/
Grossuana group, depends on which species are being
compared. In R. maceradica n. sp., the penis carries a pro-
nounced hook instead of a weak outgrowth, as described
for the type species, G. serbica ( 1983).
In contrast,   
8G19) have shown that in Grossuana the penis may not
have any outgrowth.  et al. (2012) investigated
females of G. serbica from the type locality, but their
     
 (1983: 57) described this species as having the
“Genital chamber of a characteristic pear-shaped form,
elongated and proportionally strongly developed”. How-
ever,  et al. (2012: table 1, row M3) depicted
a spherical bursa. Furthermore, 
shows that the distal receptaculum (RS1) nearly touches
the base of the bursa, but according to  et al.


they represent only physiological, stochastic variation, or
something else.
Excursus for some Greek taxa of the
Radomaniola/Grossuana group
 (1966) described from Greece Pseudamni cola
vrissiana and Pseudamnicola vurliana, and later, Orienta-
lina delphica, Trichonia kephalovrissonia, and Trichonia
trichonica ( 1973), but he did not consider taxa
described from that country earlier. After Hydrobia trito-
num Bourguignat, 1852 and Paludina curta Küster, 1853,
Paludina hohenackeri Küster, 1853, from the Balkans,
is the third of the 15 Radomaniola taxa to be described
(Appendix 4).  (1983) had overlooked that
lectotypes of Radomaniola seminula (Frauenfeld, 1863)
[Amnicola] and R. (?) hessei (Kobelt, 1891) [Pseudam-
nicola] were selected by  

 [1980] misleadingly thought that R. seminula
and R. (?) hessei, as well as R. (?) liola Westerlund,
1881 [Amnicola] and R. (?) haesitans Westerlund, 1881
[Hydrobia], belong to Belgrandiella.
 et al. (2012) studied R. tritonum (Bour-
guignat, 1852) [Hydrobia], Grossuana hohenackeri
(Küster, 1853) [Paludina], and G. marginata (Westerlund,
1881) [Amnicola] but without making any comparisons
with type specimens. However, among these 3 taxa, R.
tritonum      
only by topotypes. According to  (1853:
64), “Ce Mollusque habite sous les feuilles des plantes
aquatiques des eaux fangeuses du marais de Lerne, en
Grèce” [This mollusc lives under the leaves of aquatic
plants from the muddy waters of the marsh of Lerne in
Greece]. 
examined a sample from a spring at Mili (Lérni), Pelo-
ponnisos.
Here, we discuss the identity of Paludina hohenack-
eri Küster, 1853, Amnicola liola Westerlund, 1881, A.
marginata Westerlund, 1881, and Hydrobia haesitans
Westerlund, 1881.
Paludina hohenackeri Küster, 1853
Figures 5D–G, 9B, C
Material. Syntypes of Paludina hohenackeri are un-
known (cf.  2006: 70); we have not found any
syntypes at SMF and NHMW. Original material in the
Charpentier Collection: 31 shells + 1 juvenile shell in-
crusted within aperture. Three labels accompany the
syntypes: (1) label with green sticker “94”, “Pal. Ho-
henackeri Charp. Küst. Tab. 13 f[ig]. 18–19. Hohenacre
Grèce 1848”; (2) label: “208 [in red] Hohenackeri Charp.
Paludina brevis [“brevis” crossed out] Hohenaceri Grae-
Hohenackeri Charp.”
In Charpentier’s catalogue it is listed under Paludina as
“94 Hohenackeri Charp. - Grèce”; the number of speci-
mens is given as 25.
Neotype.  (1853: 77) received his material from
Charpentier (“Paludina hohenackeri, Charpentier in
litt”). To clarify the identity of B. hohenackeri, we hereby
select a neotype from the original material.
Measurements of neotype: height 1.68 mm, diameter
1.10 mm.
The shape and measurements of this neotype corre-
spond to      
        
    1853: 77) by taking 1
Paris line as 2.26 mm, the result is a height of 1.69 mm
and a diameter of 1.13 mm (cf. the conversion by -

measurements and those of the neotype are shown in
Table 2.
Paludina hohenackeri sensu  [1980] and 
(2006). The type locality of P. hohenackeri is Greece,
     [1980] treated P.
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hohenackeri as a species of Belgrandiella and gave its
distribution as “(restr. Thessalien) ... Insel Euböa ... Thra-
zien ...”.  (2006: 55), refering to  [1980],
gave its distribution as follows: “GR-AEG [= North
Aegean Islands] (Limnos), GR-GRC [= Greek mainland
(including Andikíthira, Evia, the Ionian Islands, Samo-
thráki, the Northern Sporades, and Thassos] (mainland;
Evia; Thassos). Endemic for Greece”. However, neither
P. hohenack-
eri. In comparison to specimens of the original material
(see below), the shell of B. hohenackeri sensu 
[1980] is less conical.
Paludina hohenackeri in the literature.  et
al. (2012, 2015) treated P. hohenackeri as a species of
Grossuana.
Operculum and pigmentation. The dried animal of the
neotype, seen through the shell wall, is black. Its opercu-
lum is chestnut-coloured.
 Concern-
ing the Radomaniola/Grossuana group in Greece, we
have compared the original material in the Charpentier
Collection to shells from 56 localities scattered through-
out the country, including 29 samples collected by P. and
A. Reischütz, [1980], 2 shells
        
 et al. (2012). All localities of samples col-
          
by Schütt, Radoman and Falniowski et al. are shown in
Figure 9. At a glance, 2 samples only, collected by P. and
A. Reischütz (Figs 9A, 9C) might be P. hohenackeri.
These are:
Locality 4 (Fig. 8): Thermo, a spring at Trihonida lake
(Fig. 9A).
Locality 9 (Fig. 8): a spring in Kefalari at Lerna (Fig. 9C).
A detailed comparison of the neotype (Fig. 9B) with the
shells from these 2 localities (Figs 9A, 9C) shows, how-
ever, that only the shell from Kefalari at Lerna represents
G. (?) hohenackeri. The shell from Thermo (in front view
of Fig. 9A) has a narrower second whorl and a less-vaulted
last whorl; the angle of the aperture, as formed by the
palatal and parietal border, is not lifted upwards, and the
columellar border is much less turned over the umbilicus.
Because of its wealth of water, Lerna was an impor-
tant ancient place rich in myths and has always attracted
travellers. One such traveller, Louis Félicien Joseph
Caignart de Saulcy, a globetrotter, botanist, paleontologist,
entomologist, and malacologist, collected a sample of
snails in Lerna; these were used by Bourguignat for his
decription of Hydrobia tritonum, a species of Grossuana.
It is our view that Charpentier’s material originated
from Lerna. Charpentier was a well-known collector who
 -
ter receiving this material, he forwarded it to Küster for
description. It is not a surprise that the Charpentier Col-
lection also contains a sample of Bourguignat’s Hydrobia
tritonum collected by Saulcy (Charpentier catalogue: 20,
no. 63). Therefore, we cannot accept ’s ([1980]:

locality to Thessaly. Schütt had seen neither syntypes nor
the original material in the Charpentier Collection, and
his photograph of P. hohenackeri ( 1980]: pl.

(Table 2).
Further investigations are needed to determine if P.
hohenackeri Küster, 1853 is a junior synonym of Hydro-
bia tritonum Bourguignat, 1852. Its type locality is
number 42 in Figure 8.
Amnicola marginata Westerlund, 1881
Figure 5H–J
Material. Westerlund Collection (Göteborg), 4526: (1)
numerous shells + 1 shell Bythinella sp.; label: “Amn.
marginata n. sp. Euboea ad Arlachades. Thiesse”; (2) c.
50 shells + 3 shells Bythinella sp.; label: “Amn. margi-
nata n. sp. Euboea, Arlachlades [sic]. Blanc”; (3) 3 shells;
label: “2. Amnicola marginata West. Eubée”.
We have not seen the material in the Westerlund Col-
lection (Stockholm), RM 14.20.
Lectotype.        
sample mentioning “Thiesse”. Measurements of the
shell: height 1.73 mm, diameter 1.20 mm. For compari-
son, ’s (1881: 68) measurements are “Long.
1¾, diam. 1 mm”.
Amnicola marginata in the literature.  [1980],
followed by  (2006), treated this species as B.
hohenackeri marginata, but according to  et
al. (2012, 2015), A. marginata is a species of Grossuana.
Operculum and pigmentation. The dried animal of the
lectotype, seen through the shell wall, is black. Its oper-
culum is chestnut-coloured.
Type locality. We have not been able to identify the type
locality “Graecia ad Avlochades”, even by taking the
Table 2. Measurements of Paludina hohenackeri Küster, 1853.
Taxonomic reference /collection Height (mm) Diameter (mm) Height:diameter
Paludina hohenackeri Küster, 1853: 77 1.69 [¾ ]1.13 [½ ]1.50:1
Neotype of Paludina hohenackeri (Charpentier Collection) 1.68 1.10 1.53:1
Grossuana hohenackeri sensu F et al. 2012: 25 g. 3 G19 1.63 1.07 1.52:1
Grossuana hohenackeri sensu S 1980: pl. 10 g. 20; SMF 262 332 1.83 1.27 1.44:1
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Figure 8. Localities of species of the Radomaniola/Grossuana
group in Greece: 1, Drama, municipal pool, P. & A. Reischütz
leg.. vii.2006; BOE 3369; 2, spring at Monolithio [at Kalen-
dini?], P. & A. Reischütz leg.; BOE 3370; 3, Louros, valley at
Ag. Georgios, P. & A. Reischütz leg.; BOE 3371; 4, Thermo,
spring at Trihonida lake, P. & A. Reischütz leg.; BOE 3372; 5,
Quelle at Amphitea at Ioannina, P. & A. Reischütz leg.; BOE
3373; 6, spring at Olimbias–Arnea road at crossroads Varvara,
P. & A. Reischütz leg.; BOE 3374; 7, spring in Kristallopigi
W of Oliki [at Paramithia?], P. & A. Reischütz leg. viii.2003;
BOE 3375; 8, Peloponisos, ancient spring of Phigalia E of
Za[c]haro, viii. 2003 P. & A. Reischütz leg. viii.2003; BOE 3376;
9, Peloponisos, spring in Kefalari at Lerna, P. & A. Reischütz
leg. viii.2008; BOE 3377; 10, Peloponisos, spring S of Vrisari
W of Kalavrita, P. & A. Reischütz leg. viii.2005; BOE 3378; 11,
Peloponisos, Altomira S of Kambos, P. & A. Reischütz leg.
viii.2005; BOE 3379; 12, Peloponisos, spring in Aetos at Dorio,
P. & A. Reischütz leg. viii.2005; BOE 3380; 13, Evvia, spring
in Steni Dirfos, P. & A. Reischütz leg. vii.2006; BOE 3381; 14,
spring at Kastro Livadia, P. & A. Reischütz leg. vii.2007; BOE
3382; 15, Evvia, spring 3 km SW of Agriopotamo [Agriovota-
no?], P. & A. Reischütz leg. vii.2007; BOE 3383; 16, Maked,
spring in Paradeissos [Paradisos] W of Nestos [?], P. & A. Reis-
chütz leg. vii.2007; BOE 3384; 17, Anatoliko Frangista NW of
Karpenision [Karpenisi], spring at open-air swimming pool, P.
& A. Reischütz leg. vii. 2008; BOE 3385; 18, spring in Eptahori
W of Konitsa, P. & A. Reischütz leg. vii.2010; BOE 3386; 19,
Ag. Sophia, at Trihonida lake, spring, P. & A. Reischütz leg.
vii.2010; BOE 3387; 20, captured spring downhill Anavra S of
Thermopiles, P. & A. Reischütz leg. vii.2010; BOE 3388; 21 Pe -
loponisos, Likeo, spring at church in direction of Neda, P. & A.
Reischütz leg. vii.2010; BOE 3389; 22, Ag. Oros, Athos, spring
Agiasma at Ag. Athanasio, P. & A. Reischütz leg. ix.2013; BOE
3390; 23, Ag. Oros, Athos, spring S of Megistis Lavras, P. &
A. Reischütz leg. ix. 2013; BOE 3391; 24, Nagia N of Parga,
spring, P. & A. Reischütz leg. iv.2014; BOE 3392; 25, Pelo-
ponisos, Killini, Roman baths, P. & A. Reischütz leg. iv.2014;
BOE 3393; 26, spring W of Vonitsa, at road at Pla Panagia, P.
& A. Reischütz leg. iv.2014; BOE 3394; 27, Evvia, spring bet-
ween Kokkinohori and Karistos, P. & A. Reischütz leg. iv.2016;
BOE 3395; 28, Piges Vellas S of Konitsa, P. & A. Reischütz
leg. v.2016; BOE 3396; 29, spring in Vrisoula in direction of
Kranea N [W?] of Amvrakikos Kolpos, P. & A. Reischütz leg.
v. 2016; BOE 3397; 30, Levkas, Kaligoni; S 1980: pl. 9
fig. 16; 31, Zante; loc. cit.: pl. 9 fig. 17; 32, Moules at Arta;
loc. cit.: pl. 9 fig. 18; 33, Delphi; loc. cit.: pl. 9 fig. 19; 34, Ve-
lestinon; loc. cit.: pl. 10 fig. 20; 35, Kavalla; loc. cit.: pl. 10 fig.
20; 36, Delfi; R 1983: 46, pl. 2 fig. 31; 37, Vrissia; loc
cit.: 85, pl. 6 fig. 88; 38–52, F et al. 2012: 21 fig. 1
G1-G15, 24 fig. 2 G1–G15; 53–56, loc. cit.: 21 fig. 1 G17–G20,
24 fig. 2 G17–G20.
labels of the syntypes into consideration. 
&  (1879: 141) mentioned “Nord de l’Eubée à
Arlachades [sic]”, but in another context (i.e., the label

At the type locality, A. marginata apparently co-exists
with a species of Bythinella (shell height 2.95–3.35 mm,
N = 2). This may be useful for the rediscovery of the type
locality.
Amnicola filiola Westerlund, 1881
Figure 5K, L
Material. In Westerlund Collection (Göteborg) 4527: (1)
4 shells, label (printed: “Collectio Molluscorum Typica
C. A. Westerlund”): “Amn. liola West. Graecia, Patras”;
(2) 4 shells; label: “Pal. liola W. Gr., Patras.”
Measurements of the largest shell: height 1.75 mm,
diameter 1.35 mm, height:diameter = 1.30 [instead
       1881: 68),
height:diameter = 1.50].

the original description and the syntypes, and because we
have not seen material in Westerlund Collection (Stock-
holm) RM 14.21, we hesitate to select a lectotype.
Amnicola liola in the literature. According to 
([1980]: 128), A. liola is a junior synonym of Amnicola
seminula Frauenfeld, 1863, which he regards to be a spe-
cies of Belgrandiella.  (2006) accepted this synony-
my.  et al. (2012) examined a specimen from
near Patras, although they did not provide it with a name.
Hydrobia haesitans Westerlund, 1881
Figure 5M, N
Material. Westerlund Collection (Göteborg) 4385: (1)
4 shells; label (printed: “Collectio Molluscorum Typica
C.A. Westerlund”): “Hydr. haesitans W. Gr., St. Maura
Blanc”; (2) 3 shells; label: “Pal. haesitans W. Graecia,
Santa Maura.”

height 1.925 mm, diameter 1.175 mm [instead of “Long.
2¾ diam. 1½ mm” ( 1881: 69)]. The heights
of the 6 mature shells, belonging to another species, are

We hesitate to select this single shell as the lectotype,
because its aperture is not fully developed and because
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 
entire second sample, belong to another species; we have
also not seen the material in the Westerlund Collection

exist at the type locality needs to be determined.
H. haesitans in the literature.   
in terpreted this species as belonging to Belgrandiella
and wrote “penis slim with appendix; one receptaculum

no indication of the loca lity from where the dissected ma-
terial came.  (2006) accepted Schütt and placed this
species in Belgrandiella.
Operculum and pigmentation. The dried animal of the
specimen with the largest shell, as seen through the shell
wall, is black. The operculum is horn-coloured and not
chestnut-coloured.
Acknowledgements
We thank Anita Eschner (Wien), Gerhard Falkner (Hörl-
kofen), Edmund Gittenberger (Leiden), and Ronald
Janssen (Frankfurt am Main) for helpful information,
Andrzej Falniowski (Krakow) as highly valued referee,
Anne Freitag (Lausanne), Ted von Proschwitz (Göte-
borg), and Peter and Alexander Reischütz (Horn) for
supporting us in an extraordinarily generous manner with
material for comparative purposes. In addition we like to
thank David Walker, who polished our English.
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cologique de la Grèce inclus l’Epire et la Thessalie. Coquilles
extramarines. Naples: Frères Tornese.
, M.L. (2008) Two records of the regional endemic hydro-
biid snail Grossuana codreanui (Grossu, 1946) in Bulgaria (Do-
bruja) and some nomenclatorial notes. Mollusca 26: 163–167.
Manuscript submitted 7 May 2017
Revised manuscript accetped 18 July 2017
Appendix 1
List of nominal taxa of Belgrandiella,
or regarded as belonging to this genus,
on the Balkan Peninsula1
For type localities see Figure 1.
B. apfelbecki (Brancsik, 1888) [Frauenfeldia lachei-
neri]. Type locality: “im Bosnaursprung (Vrelo Bosne)”
[at Ilitza, c. 10 km SW of Sarajevo, Bosnia-Herzegovina].
Lectotype: Male and female
genitalia: 
B. bozidarcurcici     Type locality:
“Bosnia and Herzegovina, Republic of Srpska, Banjaluka,

 &

B. bumasta Schütt, 1960. Type locality: “Spaltenge-
wässer der Rugovska Klisure bei Pec, autonomer Bezirk
Kosmet”. Holotype (?): 
B. conica Radoman, 1975. Type locality: “Mrzlik, a
 
of Obrh” [Slovenia]. Lectotype:  (1991: pl. 6


B. croatica (Hirc, 1881) [Bythinella]. Type locality: “in
einem Bächlein bei Brod [na Kupi] gegen das Dorfe
Lesnica zu” [Croatia]. Topotype:  (1970: pl. 8

B. crucis [Frauenfeldia (?)]. Type local-
       
Syntype: 
32. Male and female genitalia: 
B. dabriana Radoman, 1975. Type locality: “source of
the river Dabar, about 6–8 km southern of Sanski most,
by Dabarska pecina (Dabar cave)” [Bosnia-Hercegovina].
Lectotype: 
to 
B. driniana (Radoman, 1975) [Sarajana]. Type locality:
“spring in the village Kaostica, about 2 km above Drina
river, above the road Visegrad–Ustipraca” [Bosnia-
Hercegovina]. Lectotype:     
62, corresponding to 

B. erythropoma Schütt, 1959. Type locality: “in einer
Quelle unterhalb Vrelo Knijeginjac und oberhalb des
Wasserreservoir [sic] Maratin am Abhange des Berges
Trebovic [Trebevic] nach Sarajewo hin” [Bosnia-Herce-
govina]. Paratype: 
B. fontinalis (F. J. Schmidt, 1847) [Paludinella]. Type
locality: “in einer Quelle bei Lustthal [dol c. 11 km NE
of Ljubljana]” [Slovenia]. Lectotype:  1970: pl.

B. globulosa Bole, 1979. Type locality: “Mrzla jama pri
Blocicah, 3,5 km NNW od Loza”; “Kleine Wasserhöhle
Mrzla jama bei Blocice, 3,5 km NNW von Loz.” Holo-
type: 
B. hershleri Slapnik, 1997. Type locality: “Spring by
Prusnik, Sava hills, south of the main Sava valley road

east of Ljubljana, Slovenia.” Paratypes:  1997:
        
row, right.
1The objective of this appendix is not to determine whether taxa that have not yet been anatomically or genetically studied belong to Belgrandi-
ella or Graziana. It is also not our objective to clarify whether the Graziana species described by  (1975), G. adriolitoralis, G. glin-
ensis, G. papukensis, G. slavonica, and G. vrbasensis, actually belong to Graziana, or rather, to Belgrandiella. Based on genetic analyses by
et al. (2016), Belgrandiella is not present in Bulgaria. Thus, the following 7 species, described as Belgrandiella, belong to Ponto-
belgrandiella: angelovi, dobro stanica, nitida, pandurskii, stanimirae, and zagoraensis. According to  (1994: 223) B. kuesteri Boeters,
1970, pro Paludina minu tissima Küster, 1853 non Grateloup, 1838, belongs to Graziana. Type locality: “in Krain [Slovenia] am Grosskahlen-
berg [Smarna Gora c. 8 km NW of Ljubljana] in Quellen”. Topotype of original sample in F.J. Schmidt Collection (
For Belgrandiella zaschevi Angelov, 1959, see Appendix 2, footnote 4. We could not determine whether Belgrandiella kropae, described by
 (1994) in his dissertation, is valid according to Chapter 3 of the Code (ICZN 1999).
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2& (2015), concluded that B. krupensis together with 3 other nominal species (i.e., B. kusceri, B. umbilicata and B.
zermanica), constitute a single biological species with slight genetic variation. However, Falniowski and Beran may not have investigated
animals from the type locality; shells of B. krupensis
&
3B. kusceri and B. umbilicata. According to & (2015), the COI sequences of both were identical.
4The type locality of Belgrandiella zaschevi Angelov, 1959 is “Die große Karstquelle bei der Höhle ‘Duschnik’, Dorf Iskrez [Iskretz N of
Syntype: -
Cavernisa Radoman,
1978, or if it belongs to Pontobelgrandiella because of its Bulgarian type locality, or if it belongs to Belgrandiella, as perhaps does P. (?)
maarensis 
B. hyalis Westerlund, 1886 [Paludinella (Bythinella)].
Type locality: “Croatien bei Skrad” [between Rijeka and
Karlovac, Croatia]. Lectotype: 
B. koprivnensis Radoman, 1975. Type locality: “spring
Krcana, about 2 km from the road Cazin-Buzim, near the
village Donja Koprivna” [Bosnia-Hercegovina]. Lec-
totype:        

B. krupensis Radoman, 1973 2 Type local-
ity: “spring of small river Krupa, the right tributary of
Zrmanja river” [Croatia]. Lectotype:  1991: pl.


B. kusceri (A.J. Wagner, 1914) [Belgrandia].3 Type
locality: “im Schlamme des Rakbaches (Rakowski potok)
bei Rakek [c. 5 km SE of Planina] in Krain” [Slovenia].
Syntype:       Male and female
genitalia:        

B. novoselensis Radoman, 1975. Type locality: “small
spring in Nova sela, by the road Kocevje-[Brod-na-Kupi-]
Delnice, on the left side of the river Kupa” [Slovenia].
Lectotype: 
to 
B. pageti Schütt, 1970. Type locality: “Tounjcica-Höhle
    -
nica bei Tounj nahe Ogulin, Kroatien.” Holotype: 
1970
B. robusta Radoman, 1975. Type locality: “Obrh, the big
    
-
        

genitalia: 
B. schleschi[Pseudamnicola]. Type local-
ity: “Höhle Krizna jama [at Loz, Slovenia]. Topotype?:
   

B. substricta   [Microsalpinx]. Type lo-
cality: “Bistraquellen” [in Bistra c. 3 km SE of Vrhnika,
Slovenia]. Syntype:       
ing). Male and female genitalia: 
B. superior   [Belgrandiella kusceri] [not
       
[Cerknica basin] … Originalfundort die Speiquelle Jezer-
ski obrh” [Slovenia]. Topotype?:  1975: 63 pl.


B. travnicensis (Radoman, 1975) [Sarajana]. Type local-
ity: “Plava voda in Travnik town” [Bosnia-Hercegovina].
Lectotypes:        -
sponding to 

B. umbilicata   Type locality: “Mocilnik
(die Hauptquelle der Ljubljanica)” [at S border of Vrh-
nika, Slovenia]. Topotype:    
Male and female genitalia: 
B. zermanicaType locality:
“the middle course of Zrmanja river, Dalmatia” [Croatia].
Lectotype: 
to 
Appendix 2
List of nominal taxa of Pontobelgrandiella,
or regarded as belonging to this genus,
from the Balkan Peninsula
For type localities, see Figure 2.4
P. angelovi (Pintér, 1968) [Belgrandiella]. Type local -
ity: “Bulgarien, Balkan-Gebirge (Stara Planina), eine
    
vom gleich namigen Dorf.” Holotype:   
1. Paratype:  &-
 et al. 2016:
table 1.
P. bachkovoensis (Glöer & Georgiev, 2009) [Belgrandi-
ella]. Type locality: “Bachkovo village, West Rhodopes.
   
[Bulgaria]. Holotype: & 
Male genitalia: &
simple).
P. bulgarica (Angelov, 1972) [Belgrandiella]. Type
locality: “Karstquelle beim Höhlenausgang bei dem Dorf

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Archiv für Molluskenkunde · 146 (2) 2017
[Bulgaria]. Holotype:     
Male genitalia: 
P. bureschi (Angelov, 1976) [Belgrandiella]. Type loca-
lity: “Karstquelle, die das Bad beim Dorf Bankja (Bezirk
      
versorgt.” Syntype: 
P. dobrostanica (Glöer & Georgiev, 2009) [Belgrandi-
ella]. Type locality: “Gargina Dupka cave, about 20 m
      
    Holotype:  &
       &
 et
al. 2016: table 1.
P. hessei (A.J. Wagner, 1928) [Belgrandia (Belgran-
diella)]. Type locality: “die Höhle Temnata Dupka bei
     
in N. Bulgarien”. Syntype:     
74–77); topotype: 
P. (?) maarensis (Georgiev, 2013) [Belgrandiella].5 Type
locality: “Urushka Maara cave, near village of Krushuna,
   
      2013:
   
2B (“penis … with … a small lobe in the middle on its
left side.”).
P. nitida Angelov, 1972 [Belgrandiella]. Type locality:
“Karstquelle beim Höhlenausgang bei dem Dorf Pola-
      
[Bulgaria]. Holotype:     
topotype: 
genitalia: 
P. pandurskii (Georgiev, 2011) [Belgrandiella]. Type
     
village of Devetaki, Lovech town area, Devetashko pla-
teau, northern Bulgaria”. Holotype:  2011a: 9
 
Molecular data:  et al. 2016: table 1.
P. pusilla (Angelov, 1959) [Belgrandiella]. Type locality:
“Die Karstquelle ‘Petreski Isvor’ in der Umgebung der
    -
kangebirge” [Bulgaria]. Syntype: 
(drawing); topotype: 
P. stanimirae (Georgiev, 2011) [Belgrandiella]. Type
locality: “Zmeyova Dupka cave, near Tryavna town, Stara

512 m alt. Holotype: -
talia: 
et al. 2016: 3 table 1.
P. tanevi Georgiev, 2013. Type locality: “Parnitsite cave,
near village of Bezhanovo, Pre-Balkan area, northern Bul-


Molecular data:  et al. 2016: table 1).
P. zagoraensis (Glöer & Georgiev, 2009) [Belgrandi-
ella]. Type locality: “spring near Bedechka River, park
‘Krairechen’, town of Stara Zagora” [Bulgaria]. Syntype:
 &      
 &      
 et al. 2016: 3 table 1.
Appendix 3
List of nominal taxa of Grossuana,
or regarded as belonging to this genus,
from the Balkan Peninsula
For type localities, see Figure 3.6
G. angeltsekovi Glöer & Georgiev, 2009. Type local-
ity: “Bachkovo village, W Rhodopes, spring in sand”,
      
paratype:  & 
 & 

2 outgrowths). Molecular data:  et al. 2015:
table 1.
G. codreanui (Grossu, 1946) [Paladilhiopsis].7 Type
locality: “dans la grande source captée et murée d’Ac-

[Bulgaria]. Syntype:     
shell from Balcic:  et al.

19, 20,       
       et
  et al.
2015: table 1.
5Because the type locality of Belgrandiella maarensis Georgiev, 2013 lies within the distribution of Pontobelgrandiella, which is clearly
separate from Belgrandiella because of its distinctive male genitalia, this species is here listed with reservation as P. (?) maarensis.
6This list does not include the nominal taxa treated by  (1983) as subspecies of Grossuana serbica or of Radomaniola curta,
except for G. codreanui and G. vurliana, which  et al. (2007) and  et al. (2015) recognized as valid species.
According to  (1983) the following taxa are synonyms of R. curta: Amnicola miliaria Frauenfeld, 1863, Amnicola montenigrana
Frauenfeld, 1865, and Hydrobia consociellaGrossuana aytosensis Georgiev,
2012, see Appendix 4.
7According to  (2008), Paladilhiopsis codreanui Grossu, 1946 may be a junior synonym of Pseudamnicola consociella euxina
A.J. Wagner, 1928.
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G. delphica Radoman, 1973 [Orientalia8
Type locality: “spring in Delphi, by the main road Ath-
ens–Mesolongion, Greece”. Lectotype:  1991:

31. Molecular data:  et al. 2007: table 3,

2016: table 1.
G. derventica Georgiev & Glöer, 2013. Type locality:
“Dratchi Dupka cave, near village of Melnitsa, Der-
 
 &
5A,    -
 &  
single outgrowth on its left side”) and  et al.

G. euxina (A.J. Wagner, 1928) [Pseudamnicola con-
sociella]. Original localities: “Quelle Reka Devna bei
      
      
Bulgarien”. Syntypes:      
21–24 (drawings).
G. falniowski Georgiev, Glöer, Dedov & Irikov, 2015.
      
River, Krayrechen Park, city of Stara Zagora, Bulgaria,
    
     
    -

bearing small single lobe on its left side.”). Molecular
data:  et al. 2015: table 1.
G. hohenackeri (Küster, 1853) [Paludina]. Type local-
ity: “in Griechenland”. Neotype: herein, Figure 9B.
G. marginata (Westerlund, 1881) [Amnicola -
ured]. Type locality: “Graecia ad Avlochades” [“Nord
de l’Eubée à Arlachades [sic]” ( &
1879: 141); “Euboa, Arlachlades [sic]” (Blanc label in
Westerlund Collection)].
G. serbica Radoman, 1973 

    
 1983: pl.

… with … a weak outgrowth”); female genitalia: -

Molecular data:  et al. 2007: table 3, -

G. slavyanica Georgiev & Glöer, 2013. Type local-
ity: “Stream below water source at Goleshovo village,
 
-
 &   et al. 2015:
  &
(“penis bears a small lobe on its left side”). Molecular
data:  et al. 2015: table 1.
G. thracica Glöer & Georgiev, 2009. Type locality:
“where the water emerges from the spring of Chirpan
 
village of Bolyarino, Upper Thracian Lowland, southern

and paratypes:  & 

& 

G. vurliana (Radoman, 1966) [Pseudamnicola]. Type
locality: “Zivi u izvoru Kamena Vurla, blizu mesta Sv.
Konstantin, kod glavnog druma Larisa-Atina, iznad
ceste (Grcka)” [“the spring Kamena Vurla near the place
Ag. Konstantinos, above the main road Larisa–Athens,
Greece”;  1983: 60]. Syntype (?): 
 1991: pl.
  
Molecular date:  et al. 2007: table 3, 
 et al. 2016: table 1.
Appendix 4
List of nominal taxa of Radomaniola and
Graecorientalia, or regarded as belonging to
these genera, from the Balkan Peninsula9
For type localities, see Figure 4.
R. (?) aytosensis Georgiev, 2012 [Grossuana]. Type
locality: “Water source in a park forest north of Aytos
town, East Stara Planina Mts, southern Bulgaria”, 42°
-

genitalia:       
bearing a small single or bi-lobed [!] outhgrowth on its left
side”). Molecular data:  et al. 2015: table 1.
R. bosniaca Radoman, 1973 
“spring in the village Miljevci, between Bosanska Krupa
and Sanski Most, Bosnia”. Lectotype:  1991:

R. bulgarica Glöer & Georgiev, 2009. Type locality:
“Thermal spring south of the village of Ostra Mogila,
southern slope of Sarnena Sredna Gora Mts, Southern
     
Holotype and paratype:  &   
14.1, 14.2. Male genitalia:  &  2009:
       
Molecular data:  et al. 2015: table 1.
8 et al. (2007) and  et al. (2012, 2016) treated Orientalia delphica Radoman, 1973 as a species of Grossuana.
9For Graecorientalia, see also Table 1.
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Archiv für Molluskenkunde · 146 (2) 2017
R. curta (Küster, 1853) [Paludina].10 Type locality: “in

         
20,      
 1972: 196 (“Penis mit einem Appendix an der

  
        

R. elongata Radoman, 1973  -
ity: “spring near monastery Vranjina, Virpazar, Crna
Gora” [Vranjina island, lake Skutary;  1983].
Lectotype:       
to      -
      R. curta  
14M17.
R. (?) liola (Westerlund, 1881) [Amnicola  -
ured]. Type locality: “Graecia, Patras [Patrai], Fonte
Salevale”. Possible syntypes: herein, Figure 5K, L.
R. (?) haesitans (Westerlund, 1881) [Hydrobia] [not
      
Levkas = Levkada], Megali Vressi [Megali Vrisi at Kali-
goni]”. Topotype: 
R. (?) hessei (Kobelt, 1891) [Pseudamnicola]. Type local-
ity: “auf Zante [Zakinthos, Greece]”. Lectotype: 

R. lacustris Radoman, 1983. Type locality: “Lake
Skutary, along the sandy beach on the south lake bank,
below Murici village” [Montenegro]. Syntype (?): -
       

R. montana Radoman, 1973. Type locality: “spring
near stream Lukavac, about 15 km (above the main
road) north-west from Budva, Crna Gora.” Lectotype:


R. radostinae Georgiev, 2012 [Grossuana]. Type locality:
“small stream near the beginning of the path to ‘Madar-
ski Konnik’ monument at Shumensko Plateau area, near
village of Madara, 20 km from Shumen town, northern
     
   et al.
  2012: 116, 118
-
growth on the left side”). Molecular data:  et
al. 2015: table 1.
R. rhodopensis Glöer & Georgiev, 2009. Type locality:
“a small spring (water source), tributary of the Pavelsko
       
       
 & 
cf.  &  2009: 132 (“penis bears a double
lobe on its left edge”).
R. seminula (Frauenfeld, 1863) [Amnicola  -
ured]. Type locality: “aus Arcadien” [Greece]. Lectotype:

R. strandzhica Georgiev & Glöer, 2013. Type local-
ity: “in the spring emerging from the Izvora cave, west
     
-
 &
&
on its left side”).
R. tritonum Bourguignat, 1852 [Hydrobia
Type locality: “Graeciam, in aquis paludosis lacus Ler-
nae, habitat” [“Ce Mollusque habite sous les feuilles des
plantes aquatiques des eaux fangeuses du marais de Lerne
[Lerni at Mili], en Grèce”;  1853: 64]. Syn-
type:       
topotypes: 


Graecorientalia vrissiana (Radoman, 1966) [Pseudam-
nicola]. Type locality: “Zivi u malom izvoru kod mesta
Vrisija, pored glavnog druma Larisa–Atina, ispod ceste
(Grcka)” [“a small spring by the place Vrissia [SW of
Farsala], near the main road Larisa–Athens (below the
road), Greece”;  1983: 85]. Syntype: 
      1991:

88. Male and female genitalia:  
(“Penis … with … a ‘double’ outgrowth on the left side”).
10The list does not comprise nominal taxa treated by  (1983) as subspecies of Radomaniola curta. According to  (1983:
41) the following taxa are synonyms of Paludina curta Küster, 1853: Amnicola miliaria Frauenfeld, 1863, Amnicola montenigrana Frauen-
feld, 1865, and Hydrobia consociella
20171219-095537
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