Annotated checklist of the terrestrial and freshwater molluscs of the Arabian Peninsula with description of new species

Abstract

A b s t r a c t : In this work, all valid species and lists of primary synonyms of terrestrial and freshwater Gastropoda and Bivalvia known from the Arabian Peninsula are given. The type specimens of many of the species are illustrated. All species are described and anatomical details are given whenever possible. Zoogeographic relationships of the malacofauna of the area are discussed. A hypothesis explaining the time scales and pathways of colonisation of the Arabian Peninsula by molluscs is given. Ecological data are added, where known, to further the knowledge of autecology and distributional patterns. The following taxa are described as new to science: Stenothyra arabica n. sp., Arabivitrina jansseni n. sp., Cerastus schweinfurthi brunneus n. ssp., Cerastus schweinfurthi apicostatus n. ssp., Oxychilus (Costoxychilus) n. subgen., Oxychilus (Costoxychilus) profundus n. sp., Levantina (Laevihelix) n. subgen., Levantina (Laevihelix) symensi n. sp., Levantina (Laevihelix) asira n. sp., Levantina (Laevihelix) asagittata n. sp., Levantina (Laevihelix) semitecta n. sp. Additionally, several taxa are recorded as new to the malacofauna of the Arabian Peninsula: Assiminea nitida nitida (Pease, 1864), Gastrocopta klunzingeri (Jickeli, 1873), Boysia boysii (L. Pfeiffer, 1846), Lauria cylindracea (Da Costa, 1778), Toltecia pusilla (Lowe, 1831), Cecilioides aff. tumulorum (Bourguignat, 1859), Cecilioides acicula (O.F. Müller, 1774). New generic or subgeneric combinations are introduced: Homorus splendens (Thiele, 1910), Homorus arabica (Connolly, 1941), Streptostele (Raffraya) scotti (Connolly, 1941). Lectotypes for the following species are designated: Hydrobia lactea (Küster, 1852), Bithynia badiella (Küster, 1853).

Full text

FAUNA OF ARABIA 17: 333–461 Date of publication: 30.11.1998
Annotated checklist of the terrestrial and
freshwater molluscs of the Arabian Peninsula
with descriptions of new species
Eike Neubert
Abstract: In this work, all valid species and lists of primary synonyms of terrestrial and freshwater Gastropoda and Bivalvia
known from the Arabian Peninsula are given. The type specimens of many of the species are illustrated. All species are described
and anatomical details are given whenever possible. Zoogeographic relationships of the malacofauna of the area are discussed. A
hypothesis explaining the time scales and pathways of colonisation of the Arabian Peninsula by molluscs is given. Ecological data
are added, where known, to further the knowledge of autecology and distributional patterns.
The following taxa are described as new to science: Stenothyra arabica n. sp., Arabivitrina jansseni n. sp., Cerastus schweinfurthi
brunneus n. ssp., Cerastus schweinfurthi apicostatus n. ssp., Oxychilus (Costoxychilus) n. subgen., Oxychilus (Costoxychilus)
profundus n. sp., Levantina (Laevihelix) n. subgen., Levantina (Laevihelix) symensi n. sp., Levantina (Laevihelix) asira n. sp.,
Levantina (Laevihelix) asagittata n. sp., Levantina (Laevihelix) semitecta n. sp. Additionally, several taxa are recorded as new to
the malacofauna of the Arabian Peninsula: Assiminea nitida nitida (Pease, 1864), Gastrocopta klunzingeri (Jickeli, 1873), Boysia
boysii (L. Pfeiffer, 1846), Lauria cylindracea (Da Costa, 1778), Toltecia pusilla (Lowe, 1831), Cecilioides aff. tumulorum
(Bourguignat, 1859), Cecilioides acicula (O.F. Müller, 1774). New generic or subgeneric combinations are introduced: Homorus
splendens (Thiele, 1910), Homorus arabica (Connolly, 1941), Streptostele (Raffraya) scotti (Connolly, 1941). Lectotypes for the
following species are designated: Hydrobia lactea (Küster, 1852), Bithynia badiella (Küster, 1853).
INTRODUCTION
The malacofauna of the Arabian Peninsula and its importance for zoogeographical analysis has
largely been neglected throughout the history of malacological investigations. There are several

334 E. NEUBERT
reasons for our current lack of knowledge concerning a country which covers an area equivalent to
nearly 80 % of the Indian subcontinent. The large deserts do not support a diverse fauna and flora
compared to most of the remaining areas in the world. Such extreme habitats are usually not
inhabited by molluscs, but many species are able to survive under steppe conditions. In the desert
fringes, there is more molluscan life than one expects. In the area concerned, molluscs are
concentrated in more humid places such as oases, where they may be abundant. The moderate
climate of the mountainous areas at the western, southern and eastern rim of the Arabian plate
creates better “mollusc” conditions, which is reflected in the high abundance of some species and a
medium diversity of the fauna.
Another reason for the gap in our knowledge is the fact that many of the naturalists of the last
century did not visit the Arabian Peninsula because of its climate, diseases and other obstacles. This
is illustrated by the fact that there is no malacological information available concerning a mountain
stretch of more than 1000 km along the Arabian shores of the Red Sea.
Nevertheless, some scientists did record the wildlife, including molluscs, of the Arabian
Peninsula or neighbouring countries, for example Forskål, Ehrenberg, Jickeli and Bourguignat.
Additional data were given by PALADILHE (1872), V. MARTENS (1889), JOUSSEAUME (1889, 1890,
1899), PALLARY (1925, 1928) and others. More recently, a baseline study was published by
CONNOLLY (1941), who was the last to present a checklist of the Arabian terrestrial and freshwater
molluscs.
The aim of the work presented here is the continuation of Connolly’s baseline compilation of
the Arabian malacofauna in order to encourage more detailed research. Thus, this paper is not
basically a revision of the Arabian malacofauna but hopes to stimulate one. The geographical scope
covers the complete Arabian Peninsula. The north is delimited by the border of the Kingdom of
Saudi Arabia. In the south, the island of Socotra was excluded, although it is part of the Republic
of Yemen. The malacofauna of Socotra is highly endemic and differs profoundly to what is known
from the Arabian Peninsula. A revision of the Socotran malacofauna is in preparation.
There have only been two revisions of Arabian molluscs to date. The first was the revision of
the genus Revoilia of the Pomatiasidae by CROWLEY & PAIN (1978), the second the revision of the
Arabian Buliminidae and Cerastidae by MORDAN (1986). The results of Mordan are widely
accepted here, with some minor notes. No figures of the taxa concerned are given, because
excellent photographs are available in the publication mentioned. Descriptions and other informa-
tion were taken from this source and, where material was available, some additions were made.
MATERIAL AND METHODS
This investigation was basically stimulated by material which was collected by P. Symens and M.
Werner during an ornithological field trip in the south-western Arabian mountains. Although
quite poor in number, the species diversity was astonishing since it contained several new species
and several other taxa never recorded previously in the Arabian Peninsula. The stock for the south-
western molluscs was also enriched by the collections of W. Schneider and F. Krupp, S. Newton,
H. Dekker and F. Ceuninck van Capelle. Additional material was collected during a short field
trip by R. Janssen, E. Neubert, S. Newton and P. Symens in early June 1995 (this team of
collectors is referred to as “E. Neubert et al.” in the lists of material). As all material collected from
the provinces of Makkah, Baha, Asir and Jizan was found in the mountains or their foothills, a
geographical rather than a political region is adopted here; collectively the term “Asir mountain

Terrestrial and freshwater molluscs of the Arabian Peninsula 335
region” is used throughout. Shells from the Eastern Province were mainly collected by R. Kinzelbach
and R. Janssen & E. Neubert. Collections from Yemen and Oman by A. Liebegott, Frankfurt, S.
Neubert, Lautertal, J. Wittmann and I. and H. Pauscher, Darmstadt were included.
The aim of this publication was to include as much type material as possible. In many cases,
taxonomic and/or nomenclatural problems could only be discussed but not resolved, as they would
require much more material. Nonetheless, it was possible to check many nominal taxa which
enabled the author to clarify several questions and contribute results to some problems which are
still under discussion. Unfortunately, it was not possible to investigate the type material of
Paladilhe as access to his collection in Montpellier was denied by the authorities. Only an
unpublished manuscript on the content of Paladilhe’s collection was supplied, informing that
twelve lots from Aden are housed there. It is very likely that they contain the syntype specimens of
several taxa described by Paladilhe. The behaviour of this institution is regrettable since it prevents
scientific progress.
Generic diagnoses were derived partly from ZILCH (1960) and, where necessary, improved by
the results of more recent work and the author’s own observations.
All drawings were made using a camera lucida.
In the lists of material, the number of specimens/valves in the lot are indicated by a number in
parentheses following acronyms of (private) collections, e.g. “LIEB (9)”, unless they are encoded
in the collection number of official institutions.
Throughout the text, brackets contain annotations by the author.
Terminology
For the description of genital morphology, the germinative tissues are considered to be the
proximal parts of the genitalia. Thus, the atrium is distal, the penis boundary to the atrium is also
distal, its boundary to the epiphallus proximal. Unfortunately, the use of terms describing the
anatomy of the copulatory organs is still not uniform, although they have been discussed in detail
by several authors (cf. H. NORDSIECK 1966, 1985). This is important as the names of particular
structures should only be used for homologous organs; for example, a flagellum is by definition
the proximal terminal appendage of the epiphallus. Names like “penial flagellum” and others cause
complete confusion. For this reason, a definition of the terms used here and in forthcoming
publications is given to make absolutely clear which structure is discussed.
In the plesiomorphous state, the male copulatory organ of the Stylommatophora consists of a
penis and an epiphallus. They are differentiated by internal and external structures. The
boundaries may be indicated by either the presence of appendages, an epiphallial papilla or sudden
change in the structure of pilasters. They are no t given by the inserting point of the penial
retractor muscles as was automatically used by many authors (cf. publications of Hesse). The male
copulatory organ may completely or partially be ensheathed by muscular fascicles, the muscular
sheath. It may be connected to the retractor muscles or a simple envelope. The retractor system
may be split and should be named after the insertion point on the different parts of the copulatory
organ, i.e. penial retractor muscle, epiphallial retractor muscle, appendicular
retractor muscle etc.
The penis may have a penial appendix inserting in the median part of the penial tube. It
can be simple or subdivided into several parts and may project with the appendicular papilla
into the penis. The penial caecum is a proximal appendage branching off at the boundary of
the penis and epiphallus. The epiphallus may have a medium appendage, the epiphallial
caecum. As outlined before, the proximal terminal appendage is called a flagellum for

336 E. NEUBERT
historical reasons. Internally, the male copulatory organ may have none, one or two papillae. To be
correct, the papilla originating from the epiphallus and projecting into the penial lumen represents
the epiphallial papilla, the second papilla then is the penial papilla. In almost all cases only a single
papilla is present. In future, this single papilla should be called penial papilla regardless of the
fact that it might be a true penial or epiphallial papilla, as the homology is difficult to ascertain in
many taxa. A problem arises for taxa where a second papilla is present (for example many
Helicinae). In this case, the name epiphallial papilla for the proximal papilla is appropriate as there
is no doubt about the homology.
Acronyms of collections used for this study:
BM(NH) The Natural History Museum, London
DEKK private collection H. Dekker, Winkel
GIROD private collection A. Girod, Milano
HLMD Hessisches Landesmuseum Darmstadt
IBA private collection M. Ibañez, La Laguna, Teneriffa
IRSNB Institut Royal des Sciences Naturelles de Belgique, Bruxelles
I.a.e.n. Italian archaeological excavation number
IZPAN Instytut Zoologici Polska Akademia Nauk
KINZ private collection R. Kinzelbach, Rostock
LIEB private collection A. Liebegott, Frankfurt
MHNG Muséum d’Histoire Naturelle Genève
MNHNP Muséum National d’Histoire Naturelle, Paris
MZL Musée Zoologique, Lausanne
NEUB private collection E. Neubert
NHMB Naturhistorisches Museum Basel
NMW National Museum of Wales, Cardiff
NNM Nationaal Natuurhistorisch Museum, Leiden
ONHM Oman Natural History Museum, Muscat
SMF Senckenberg Museum, Frankfurt
SNMNH Saudi Arabian National Museum of Natural History, Riyadh
ZMHB Zoologisches Museum Humboldt Universität Berlin
ZMZ Zoologisches Museum der Universität Zürich
Abbreviations used for anatomical details:
BC Bursa copulatrix
MRA Musculus retractor appendiculae
MRP Musculus retractor penis
VD Vas deferens
All measurements are given in millimetres. Abbreviations used for conchological details:
D Maximum diameter of the shell
H Maximum height of the shell
PD Maximum diameter of the peristome
PH Maximum height of the peristome
R1Number of ribs on 1 mm of the body whorl
W Number of whorls

Terrestrial and freshwater molluscs of the Arabian Peninsula 337
RESULTS
Gastropoda
Prosobranchia
Family Pomatiasidae
Genus Revoilia Bourguignat, 1881
1881 Revoilia Bourguignat. — Mollusques terrestres et fluviatiles recueillis en Afrique dans les pays des Çomalis Med-
jourtin: 9.
Diagnosis: Low discoidal to conical shells with a slightly raised apex. The whorls are
sculptured by numerous spiral threads. The aperture is circular with a continuous peristome. A
columellar shield may close the umbilicus. A callosity spreads from the upper rim of the shell to
the body whorl. The operculum is calcareous.
Type species: Revoilia milneedwardsi Bourguignat, 1881.
The genus Revoilia was revised by CROWLEY & PAIN (1978). A subsequent investigation of the type
material of the Arabian Revoilia species revealed that their point of view cannot be adopted here.
Subgenus Socotora Pallary, 1925
1925 Socotora Pallary. — In: Little, O.H.: The geography and geology of Makalla, South Arabia. Geological survey of
Egypt: 231.
1925 Arabia Pallary. — In: Little, O.H.: The geography and geology of Makalla, South Arabia. Geological survey of Egypt:
232.
Diagnosis: Conical ventricose shells, the umbilicus may be closed by the columellar shield.
The peristome is not continuous, being offset at the point of insertion of the inner lip. The
operculum is convex on the outside with a depressed nucleus. It displays four whorls with the last
rapidly increasing.
Type species: Cyclostoma albicans Gray & Sowerby, 1839.
Revoilia (Socotora) clausa (Sowerby, 1847) Figs 1-3, 13
1847 Cyclostoma clausum Sowerby. — Thesaurus Conchyliorum I: 128, pl. 31, figs 266-267 (Yemen, Arabia).
1882 Georgia yemenica Bourguignat. — Mission de G. Révoil au pays Çomalis. Faune et Flore des Pays Çomalis: 70
(Yemen).
Material: Yemen: Jabal Sabar close to Ta’izz, 23.III.1988, A. Liebegott, LIEB (9); same locality, XII.1988, Rabin, LIEB
(9); at the road from al-Hodeida to Menaha, escarpment, 1200 m, 25.III.1988, A. Liebegott, LIEB (4).
Type material: not available, probably lost.
Description: The small shell is depressed, the last whorl increases rapidly in width. The
dome-shaped protoconch consists of two whorls. It is relatively big in diameter compared to the
size of the teleoconch and to the absolute size of the protoconch of other species. The teleoconch
whorls are rounded with a characteristically formed shallow suture. The uppermost part of each
whorl is “pasted” upwards to the wall of the preceding whorl closing the suture. Subsuturally, there
is a shallow depression or furrow without any spiral sculpture.
The teleoconch sculpture consists of very fine and densely spaced axial striae and coarser
overriding spirals. These spirals are not present on the base of the shell. In many specimens, a
bluish to red band colours the protoconch. In fresh specimens, the teleoconch colour varies from
pink to corneous brown with a prominent supramedian maroon-brown band. Often, a second
subsutural band is indicated by a darker brown coloration of the upper parts of the whorls.

338 E. NEUBERT
Figs 1-12: (original size × 3), 1-3: Revoilia (Socotora) clausa, specimen from Yemen, Jabal Sabar close to Ta’izz. 1: Frontal view. 2:
Apical view. 3: Ventral view. 4-9: Revoilia (Socotora) dhofarense. 4-6: Holotype BM(NH) 96.4.28.3. 4: Frontal view. 5: Apical
view. 6: Ventral view. 7-9: Holotype of Otopoma consimile, BM(NH) 96.4.28.2. 7: Frontal view. 8: Apical view. 9: Ventral view.
10-12: Revoilia (Socotora) hinduorum, holotype BM(NH) 19657. 10: Frontal view. 20: Apical view. 21: Ventral view.
The aperture is circular, the peristome only slightly thickened and reflected. The umbilicus is
closed by a thin callose plate in adult specimens (see remarks for R. (S.) dhofarense). The umbilical
furrow is straight and deeply excavated.
Measurements (illustrated specimen, Figs 1-3): H = 8.50; D = 10.60; PH = 6.20; PD = 5.60;
W = 4.25.
Remarks: Unfortunately, the type specimen of Cyclostoma clausum has been lost. The
description given by SOWERBY (1847) completely matches the shells of the populations presented
here. The unique characters of the suture, the depressed form and colour are clearly indicated by
him. The hitherto known range of this species seems to be a quite small area in the mountains of

Terrestrial and freshwater molluscs of the Arabian Peninsula 339
northern Yemen. Its congeners can be found from Aden along the coast eastwards to the Dhofar
area in Oman (Fig. 13).
Revoilia (Socotora) dhofarense (Melvill & Ponsonby, 1896) Figs 4-9, 13
1896 Otopoma dhofarense Melvill & Ponsonby. — Proc. Malac. Soc. London 2: 2, pl. 1, figs 9-11 (Dhofar).
1896 Otopoma consimile Melvill & Ponsonby. — Proc. Malac. Soc. London 2: 2, pl. 1, figs 5-7 (Dhofar).
Material: Yemen (all collected by H. Dekker & F. Ceuninck v. Capelle): al-Mahrah, Hawf, E side of village,
16°38'03"N 53°02'34"E, rocks with algae and sand patches, 1.X.1995, DEKK (17); al-Mahrah, Ras Sharwayn, 15°23'49"N
51°34'25"E, limestone rocks, 26.IX.1995, DEKK (38); al-Mahrah, between al-Fatk and Damquat, 16°33'N 52°47'E, limestone
rocks, 1.X.1995, DEKK (38). — O m a n (all Girod ded. 1997): Dhofar, NE slope of Qara mountains, Jibjat area (I.a.e.n.
KR 142), SMF 311671/19; same locality, (I.a.e.n. KR 143), SMF 311672/37 (+ fragments); same locality (I.a.e.n. KR 114),
SMF 311673/11; same locality (I.a.e.n. KR 117), SMF 311674/70 (+ fragments), GIROD (10); same locality (I.a.e.n. KR ?1?),
SMF 311675/15; same locality (I.a.e.n. KR 126B), SMF 311677/4.
Type material: Holotype Otopoma dhofarense, BM(NH) 96.4.28.3.; holotype Otopoma
consimile, BM(NH) 96.4.28.2.
Description: The small shell is high and conical. The prominent protoconch consists of
two whorls and is covered by dense rugose axial striae. The teleoconch whorls are rounded with a
deep suture. Their sculpture consists of prominent spirals and somewhat weaker axials. In some
specimens, these sculptural elements are similar in strength, causing a cancellate pattern of the
surface of the shell. The base of the shell is nearly smooth.
The aperture is circular, the peristome somewhat thickened and reflected. The umbilicus is
open to closed with a shallow u-shaped umbilical furrow.
The colour of the shell is white but there are specimens with a partially to completely red
protoconch. Usually, this suprasutural coloration can be observed until the fourth whorl where it
fades. A yellowish to red median band may be present, in some specimens a second subsutural
band can be found on the last whorl.
Measurements: Holotype Otopoma dhofarense (Figs 4-6): H = 11.10; D = 12.10; PH = 6.10;
PD = 5.20; W = 5.25. Holotype Otopoma consimile (Figs 7-9): H = 8.25; D = 10.80; PH = 5.30;
PD = 5.20; W = 5.
Remarks: According to the author’s observations, the presence or absence of the umbilical
plate, and thus a closed or open umbilicus, depends on the individual age of the specimen. It is
very likely that adult growth is limited to the umbilical region and causes a stepwise covering of the
umbilicus. Otopoma consimile is recognised to be a synonym as it differs only in its more broadened
shape of the teleoconch. At Hawf in Yemen as well as in the Jibjat area in Oman, R. (S.) dhofarense
and R. (S.) bentiana occur sympatrically thus proving their status as separate species (Fig. 13).
Revoilia (Socotora) bentiana (Melvill, 1895) Figs 14-22, 13
1895 Otopoma bentianum Melvill. — Proc. Malac. Soc. London 1: 224, pl. XIV, fig. 8 (Hadramaut, Arabia).
1896 Otopoma hadramauticum Melvill & Ponsonby. — Proc. Malac. Soc. London 2: 2, pl. 1, figs 1-3 (Dhofar).
1925 Arabia bentianum var. daliyana Pallary. — In: Little, O.H.: The geography and geology of Makalla, South Arabia.
Geological survey of Egypt: 226, pl. 35, figs 4-5 (Jabal el Da’liya).
1925 Arabia littlei Pallary. — In: Little, O.H.: The geography and geology of Makalla, South Arabia. Geological survey of
Egypt: 226, pl. 35, figs 1-3 (Makalla).
1925 Arabia hadramauticum var. minor Pallary. — In: Little, O.H.: The geography and geology of Makalla, South Arabia.
Geological survey of Egypt: 226, pl. 35, fig. 6 [Introduced only on the plate. Obviously a confusion with A. lucida
var. minor].
1925 Arabia lucida Pallary. — In: Little, O.H.: The geography and geology of Makalla, South Arabia. Geological survey
of Egypt: 226, pl. 35, figs 11-13 (Jabal al-Ma’âwis and Quarn el Ghail).
1925 Arabia lucida var. minor Pallary. — In: Little, O.H.: The geography and geology of Makalla, South Arabia.
Geological survey of Egypt: 227 (ditto).

340 E. NEUBERT
Fig. 13: Distribution of Revoilia (Socotora) spp. in Arabia.
1925 Georgia pulchella Pallary. — In: Little, O.H.: The geography and geology of Makalla, South Arabia. Geological
survey of Egypt: 232, pl. 35, fig. 15 (Socotra).
Material: Yemen: Hadramaut, Jabal Abdulla Kerim, 3.IV.1988, A. Liebegott, LIEB (2); al-Mahrah, between Damquat
and Jahib, 16°35.5'N 52°53'E, limestone rocks, 1.X.1995, H. Dekker & F. Ceuninck v. Capelle, DEKK (19); al-Mahrah, W of
Nishtun, 15°53'N 52°05'E, limestone rocks, 28.IX.1995, H. Dekker & F. Ceuninck v. Capelle, DEKK (24); al-Mahrah,
Nishtun, 15°48'54"N 52°11'26"E, sandy and rocky beach, 5-6.X.1995, H. Dekker & F. Ceuninck v. Capelle, DEKK (18); al-
Mahrah, Hawf, E side of village, 16°38'03"N 53°02'34"E, rocks with algae and sand patches, 1.X.1995, H. Dekker & F.
Ceuninck v. Capelle, DEKK (4). — Om a n : Dhofar, Taqah between Salalah and Mirbat, I.1993, H. Pauscher, NEUB (11);
Dhofar, NE slope of Qara mountains, Jibjat area (I.a.e.n. KR 31), Girod ded. 1997, SMF 311676/1, GIROD (1); Dhofar,
Khwar Rowri, 17°03'N 54°25'E, 16.VI.1980, A.B. Paltrinieri, ONHM 610.54 (3); Dhofar, Khwar Rowri, 17°03'N 54°25'E,
16. & 18.VI.1980, A.B. Paltrinieri, ONHM 610.53 (11).
Type material: Holotype Otopoma bentianum, BM(NH) 95.4.10.1.; holotype Otopoma
hadramauticum, BM(NH) 96.4.28.1; two syntypes Arabia lucida, IRSNB I.G. 10591.
Description: The basically white shell is broadly compressed, with the last whorl increasing
considerably in width. The submammillate and somewhat elevated protoconch is formed by
2-2.25 whorls with a finely wrinkled axial sculpture. The teleoconch whorls are well rounded with
a deep and simple suture. The fine and regular axial striae are crossed by coarser spiral threads
which disappear on the shell’s base.
Figs 14-22: (original size × 3), Revoilia (Socotora) bentiana. 14-16: Holotype BM(NH) 95.4.10.1. 14: Frontal view. 15: Apical
view. 16: Ventral view. 17-19: Holotype of Otopoma hadramauticum, BM(NH) 96.4.28.1. 17: Frontal view. 18: Apical view. 19:
Ventral view. 20-22: Syntype of Arabia lucida, IRSNB I.G. 10591. 20: Frontal view. 21: Apical view. 22: Ventral view.

Terrestrial and freshwater molluscs of the Arabian Peninsula 341

342 E. NEUBERT
The circular aperture displays a thickened lip, the peristomial rim is thickened and reflected.
The wide umbilicus is covered by a thin callose plate which is bordered by a u-shaped furrow of
medium depth.
Measurements: Holotype Otopoma bentianum (Figs 14-16): H = 15.0; D = 22.2; PH = 14.9;
PD = 9.9; W = 5.25. Holotype Otopoma hadramauticum (Figs 17-19): H = 14.0; D = 16.4;
PH = 8.7; PD = 7.9; W = 5. Illustrated syntype Arabia lucida (Figs 20-22): H = 15.3; D = 20.6;
PH = 10.5; PD = 10.2; W = 4.75.
Remarks: Otopoma hadramauticum (Figs 17-19) differs from R. (S.) bentiana in its yellow
shell with medium red band, its more conical shape and smaller dimensions. Since it is known
from several populations of Revoilia species that coloured and purely white shells may be present,
these characters seem to be of only minor taxonomic value. The taxa described by Pallary reflect
the variability of the species, although it has to be stressed that the holotype of Otopoma bentiana
(BM(NH) 95.4.10.1.) represents an extraordinarily broad specimen. Arabia lucida (Figs 20-22)
shows no conchological difference from R. (S.) bentiana except for a somewhat more conical shape
of the teleoconch.
In 1978, CROWLEY & PAIN reviewed the genus Revoilia. Concerning the Arabian species of this
genus, they relegated most of the taxa described to the synonymy of R. clausa. Clearly, this opinion
was without material from the western parts of Yemen, resulting in subsequent misinterpretation
of this species. As outlined before, R. clausa can easily be identified by its size and sutural
characters. These characters have only been recorded in shells from a very small area, while all other
Arabian shells investigated have a deep and simple suture. The problems encountered by CROWLEY
& PAIN (1978) in defining R. clausa clearly arose because they mixed up all three species present in
Arabia. The type locality was correctly given by SOWERBY (1847) as Yemen. For this reason, the
designation of a type locality “Dhofar, 800 miles east of Aden” for R. clausa has to be rejected. The
distribution of the Arabian species of Revoilia is shown in Fig. 13.
In their R. clausa mix-up, the authors also integrated Otopoma hinduorum Blanford, 1864, from
Kattawar, Western India. The holotype of this species (BM(NH) 19657) is illustrated herein
(Figs 10-12: H = 11.55; D = 11.7; PH = 6.5; PD = 6.15; W = 5.25) and can be compared with the
other species. It also has a deep suture which proves that it is not identical with R. clausa. In the
author’s opinion, this taxon has some superficial similarity with R. dhofarense, but it should be studied
in context not only with the Arabian but also with the Socotran species of this genus. It is not clear
whether R. hinduorum has been introduced to India by human activities or if it is an autochthonous
species of this subcontinent. These problems should be clarified with the help of biochemical
methods. Until this is done, R. hinduorum (Blanford, 1864) should be kept as a separate species.
Genus Lithidion Gray, 1850
1850 Lithidion Gray in Baird. — Nomen. Moll. Coll. Brit. Mus. I: 35.
Diagnosis: Flattened and depressed shells with a mammillate protoconch. The umbilicus
opens wide perspectively. The surface of the teleoconch is smooth or displays spiral ribs.
Type species: Cyclostoma lithidion Sowerby, 1847.
Lithidion lithidion (Sowerby, 1847) Figs 23-28
1847 Cyclostoma lithidion Sowerby. — Thesaurus Conchyliorum 1: 111, pl. 31, fig. 262 (Yemen, Arabia).
1850 Cyclostoma niveum Petit de la Sausaye. — J. Conchyl. 1: 52, pl. 3, fig. 7 (“Arabie, province d’Hyemen”).
1881 Lithidion marmorosum Godwin-Austen. — Proc. Zool. Soc.: 256, pl. 28, figs 6-6 c (Socotra: “very common
everywhere on the ground”).

Terrestrial and freshwater molluscs of the Arabian Peninsula 343
Figs 23-28: (original size × 4), Lithidion lithidion. 23-25: Possible type of Cyclostoma lithidion, BM(NH) 1843.10.2.122. 23:
Frontal view. 24: Apical view. 25: Ventral view. 26-28: Syntype of Cyclostoma niveum, MNHNP. 26: Frontal view. 27: Apical
view. 28: Ventral view.
Type material: Possible type Cyclostoma lithidion, BM(NH) 1843.10.2.122; two syntypes
Cyclostoma niveum, MNHNP.
Description: The small shell is discoidal depressed. It may be pure white to red-brown,
with regular axial zigzag flames which never extend to the umbilicus.
The brown to bluish mammillate protoconch is smooth and consists of two whorls. Its axis is
oblique causing the last protoconch whorl to become nearly completely submerged under the first
teleoconch whorl.
The teleoconch whorls are well rounded with a deep suture. They are sculptured by several
strong spirals which may become weaker on the body whorl. Often, the suprasutural spiral is more
pronounced forming a distinct keel. On the subsurface, the spirals disappear. The inner umbilical
walls are smooth. The umbilicus is wide and funnel-shaped.
The subcircular aperture is strengthened by a narrow lip, the upper peristomial rim is
somewhat reflected. It protrudes on the body whorl leaving a shallow sinusoid angulation. The
peristomial rims are connected by a thin to obsolete callus.
Measurements: Possible type Cyclostoma lithidion (Figs 23-25): H = 5.20; D = 9.60;
PH = 3.70; PD = 3.60; W = 4.5. Illustrated syntype Cyclostoma niveum (Figs 26-28): H = 4.30;
D = 10.20; PH = 3.40; PD = 3.45; W = 5.
Remarks: This species has not been recorded from the Arabian mainland again since the
description of the taxa C. lithidion and C. niveum. They differ in teleoconch sculpture, with less
pronounced spirals in C. niveum. Judging from several lots of L. lithidion from Socotra, presence
or absence of the spirals varies between individual shells. Intergrades are also known concerning
the coloration from pure white to marmorate.

344 E. NEUBERT
This genus is speciose on Socotra and the island of Abd el-Kuri and is still in need of revision.
As in other Socotran species, L. lithidion may have been introduced by human activity (cf.
Achatinelloides spp.).
Family Hydrobiidae
Genus Hydrobia Hartmann, 1821
1821 Hydrobia Hartmann. — Neue Alpina 1: 258.
Diagnosis: Small conical shells with an elevated spire. The operculum is thin, corneous and
spiral. The central radular tooth displays one small denticle on either side of the base. For
nomenclatural problems of the name Hydrobia refer to KABAT & HERSHLER (1993: 27).
Type species: Cyclostoma acutum Draparnaud, 1805.
Hydrobia lactea (Küster, 1852) Figs 29-30
1852 Paludina lactea Küster. — In: Martini & Chemnitz: Conchylien-Cabinet 1 (21): 50, Taf. 10, Fig. 5-6 (Mossul, Iraq).
Material: Saudi Arabia: Eastern Province, al-Hasa Oasis, al-Qarn, freshwater river to NW, 25°32'N 49°38'E,
27.I.1992, R. Kinzelbach & E. Neubert, SNMNH-MO 1/18, SMF 311521/17; al-Hasa Oasis, al-’Uyun palm plantation,
freshwater ditch with dense vegetation, 25°36'N 49°34'E, 27.I.1992, R. Kinzelbach, SNMNH-MO 2/100, SMF 311522/100,
KINZ (26); al-Qatif Oasis, freshwater course W of town, 26°36'N 49°58'E, 31.I.1992, R. Kinzelbach & I.A. Nader, SNMNH-
MO 3/200, SMF 311523/200, KINZ (60); al-Aba Oasis (Saadah, Dumayn), 9.VI.1992, R. Kinzelbach, SNMNH-MO 4/34,
SMF 311524/34; artificial pond east of main road Safwa-Jubail, 2 km N of exit Safwa, 9.VI.1992, R. Kinzelbach, SNMNH-
MO 7/120, SMF 311527/120; al-Qatif Oasis, palm plantations W of centre, irrigation channels, 14.VI.1992, R. Kinzelbach,
SNMNH-MO 6/72, SMF 311525/72; Tarut, plantations N of Tarut centre, 14.VI.1992, R. Kinzelbach, SNMNH-MO 5/7;
small pond E of the highway Khobar-Jubail, 3.4 km S of exit Juaymah, 25.V.1995, E. Neubert & R. Janssen, SNMNH-MO-
8/307, SMF 311526/307.
Type material: The original description by KÜSTER (1852) used material originating from
the collection of Charpentier, who obtained his material from Parreyss. The type lot (MZL)
contains two vials (six and two specimens) and five labels; two labels were handwritten by Parreyss,
one by Küster and two by Charpentier. Küster’s label reads “No. 69 Pal. lactea”, Parreyss’s read
“Paludina lactea” and “Palud. sp. England?”, Charpentier’s read “Pal. lactea Parr., Parreyss 1845,
Tigris prés Mossoul” and “Pro Bulimo missui potius Paludina”. The last label also contains a note
in an unidentified handwriting “Persia austr., Hohenacker 1890”. Small green labels reading “113”
on the vials and Charpentier’s first label, and numbers in red reading “210” and “211” are
interpreted as catalogue numbers. The note means that Charpentier received the specimens under
the generic name Bulimus (a frequently used generic name at this time) but he believed that it was
closer to Paludina. The whereabouts of the label with “England?” cannot be explained.
In the author’s opinion, the specimens mentioned have to be considered as syntypes because
they obviously originate from the collector Parreyss and are housed in Charpentier’s collection.
Küster’s label indicates that he definitely had these specimens at his disposal when describing
Paludina lactea. Moreover, the specimens match his description (text quoted below) exactly. For
this reason, a specimen which is close to Küster’s measurements and figure is herewith selected as
lectotype. The types are housed in the collection of the Musée Zoologique, Lausanne (MZL).
Original description: “Testa parva, anguste rimata, conica, tenuiuscula, diaphana,
nitida, subtilissime striata, alba; spira elata, obtusiuscula; anfractibus 6 planiusculis, superne
obsolete marginatis; apertura ovata; peristomate recto, acuto; margine columellari reflexo. Höhe
1.1/2 ''', Breite 0.3/5 '''. Aufenthalt: Mossul in Persien (Director v. Charpentiers Sammlung).” —
This translates to: Shell small with a narrow umbilical slit, conical, thin-walled, translucent and
glossy with very fine [axial] striations, white; the spire is elongated with a blunt apex; there are six

Terrestrial and freshwater molluscs of the Arabian Peninsula 345
flat whorls, subsuturally with a faint white line; the aperture is ovate with the peristome not
reflected, thin; the columellar part of the peristome is slightly reflected. Height c. 3 mm, width c.
1.2 mm. Locality: Mossul in Persia (in the collection of director v. Charpentier).
Additional descriptive notes: The protoconch is minutely granulated (magnification
> 60 ×) and consists of 1.5 whorls. The suture is of medium depth. Very fine and faint spiral lines
are present on the shell surface of some specimens. The umbilicus may be closed to open with all
intergrades.
Measurements (designated lectotype, Fig. 29): H = 2.95; D = 1.50; PH = 1.15; PD = 0.85;
W = 6.
Remarks: The description by KÜSTER (1852) generally matches the characters of the
specimens from the Arabian Peninsula. The shell’s outline varies from slender to broad conical. In
all populations investigated, a few very tall specimens occur with the last whorl increasing rapidly.
Since only dry shells were available, no information can be given about whether this phenomenon
represents a sexual dimorphism or is caused by parasites.
According to BROWN & WRIGHT (1980) and BROWN & GALLAGHER (1985), this species is
widespread in Eastern Saudi Arabia. The additional localities given here all originate from the same
area given by these authors. Unfortunately, there are no records from other freshwater habitats
north of Jubail towards Kuwait where Hydrobia lactea might be expected.
The classification of this species within Hydrobia s. lat. may be debatable and needs to be
proven by the anatomical investigation of topotypical specimens.
Hydrobia glaucovirens (Melvill & Ponsonby, 1896) Fig. 31
1896 Paludestrina glaucovirens Melvill & Ponsonby. — Proc. Malac. Soc. London 2: 3, pl. I, fig. 8 (Dhofar region, Oman).
Material: BM(NH) 1896.4.28.5-6, two syntypes.
Description: Both syntypes have five whorls. The only differences found from H. lactea
are the more globose whorls and, subsequently, the deeper suture of H. glaucovirens. All other
conchological characters are very similar to small specimens of H. lactea. Further detailed studies
on anatomical details of both taxa are necessary in order to decide on the specific status of H.
glaucovirens.
Measurements (illustrated syntype, Fig. 31): H = 2.45; D = 1.55; W = 5.
Remarks: This taxon is only known from the type area and has not been recorded again
since its description.
Family Bithyniidae
Genus Bithynia Leach, 1818
1818 Bithynia Leach. — Abel’s narrative of Journey into interior of China: 362.
Diagnosis: Small to medium-sized turreted shells with a calcareous and concentrically
ridged operculum.
Type species: Helix tentaculata Linnaeus, 1758.
Bithynia badiella (Küster, 1853) Fig. 32
1853 Paludina badiella Küster. — In: Martini & Chemnitz: Conchylien-Cabinet 1 (21): 62, Taf. 11, Fig. 25-28 (Beirut).
Type material: As in Hydrobia lactea, syntypes of Paludina badiella were investigated from
the collection of Charpentier (MZL). There are two vials, each containing five specimens. There
are three labels, two from Parreyss and one from Charpentier. The Parreyss labels read “1844,

346 E. NEUBERT
Paludina badiella, Barut” and “Nr. 2, Palud. kephissus”, the one of Charpentier reads “Pal. badiella
Parr., 56, nr. 2 Cephyssus, Parreyss 1844, Barut”. The catalogue number 56 (green label) is also
glued to both vials. The name “kephissus” seems to be an unpublished manuscript name of
Parreyss, and Charpentier obviously kept them conspecific. The specimens had been collected alive
as there are several individuals with an operculum. It is possible that Parreyss intended to dedicate
a different name to the few juvenile specimens.
KÜSTER (1853) states that the specimens used for his description originate from the Charpentier
collection ex Parreyss. His collection, or at least parts of the original lot because Küster’s label is
missing, is considered to contain the syntypes of Paludina badiella. To define the nomenclature of
this species, a lectotype (MZL) is herewith designated. As the specimen illustrated by Küster
cannot be traced with certainty, a specimen which provides most of the morphological information
given in the original description and in additional notes has been chosen.
There is a probable syntype in the collection of the SMF (SMF 143392/1) with an original
label of Parreyss reading “Paludina badiella”. It was provisionally selected and labelled “Neotype”
by Zilch, but never published by him. Later, this specimen was measured and illustrated by
SCHÜTT (1983: 34, Taf. 2, Fig. 31) as a neotype without any critical remark or discussion. In the
author’s opinion, this procedure was not correct as it is now proven that the original lot
containing nearly the full set of information still exists. As a consequence, the SMF specimen
has no “neotype” status, as the erroneous publication of a “neotype” by Schütt is not concordant
with the ICZN rules.
Original description: “Testa minuta, subperforato-rimata, globoso-conica, tenuis,
diaphana, subtiliter striata, nitidula, ferruginea; spira aperturam aequante, subconica, obtusiuscula,
anfractibus 5 convexis, superne planulatis, sed non angulatis; apertura ovata, marginibus continuis;
reflexiusculis. Höhe 1.3/4 ''', Breite 1.1/2 '''. Aufenthalt: bei Beirut in Syrien (Director von
Charpentiers Sammlung).” — This translates to: Shell minute, somewhat perforate to rimate,
globose conical, fragile, transparent, finely striate, somewhat glossy, dark coloured; spire of the
same dimensions as the aperture, subconical, somewhat blunt, with five convex whorls, flattened,
but not angulate above; the aperture is ovate with the rims connected and somewhat reflected.
Height c. 3.5 mm, width c. 3 mm. Locality: close to Beirut in Syria (in the collection of director v.
Charpentier).
Additional descriptive notes: The greenish yellowish shell is broad conical with a
very blunt protoconch. The whorls of the teleoconch are rounded but flat-topped below the suture.
The suture is very deep. The umbilicus is rimate and always open. The aperture is semi-ovate with
a sharp peristome. The peristomial rims may be free or connected to the body wall. The operculum
displays fine concentric striae with the nucleus shifted in a basolateral position.
Measurements (designated lectotype, Fig. 32): H = 3.90; D = 3.50; PH = 2.45; PD = 2.05;
W = 4.5.
Remarks: This species was recorded by BROWN & WRIGHT (1980) from the Safak Wells in
the Abu Washiyah area of the Khaibar Oasis. It was identified by Mandahl-Barth.
In his work on the freshwater molluscs of the Orontes river in Syria, SCHÜTT (1983) reduced
the number of Bithynia species described from the Levant area from 13 to two valid species. A re-
examination of the material stored in the Senckenberg Museum revealed several problems which
still have to be solved. From a conchological point of view, Schütt’s two-species concept is reliable.
One of the species can be characterised as described above, the other one differs by its more
elongate slender shell, the closed or narrow slit-like umbilicus, a shallow to medium deep suture,
evenly rounded whorl and an operculum with a more central nucleus. For the latter species, the
name Bithynia phialensis (Conrad, 1852) can be used.

Terrestrial and freshwater molluscs of the Arabian Peninsula 347
Figs 29-36: (original size × 10), 29-30: Hydrobia lactea. 29: Lectotype MZL, frontal view. 30: Specimen from small pond in
Eastern Province (SMF 311526/307), frontal view. 31: Hydrobia glaucovirens, syntype BM(NH) 1896.4.28.5-6, frontal view. 32:
Bithynia badiella, lectotype MZL, frontal view. 33-34: Stenothyra arabica n. sp., holotype SMF 311265. 33: Frontal view. 34:
Ventral view. 35: Iravadia quadrasi, lectotype SMF 225147, frontal view. 36: Assiminea nitida nitida, specimen from al-Qatif
Oasis (SMF 311529), frontal view.
Using these characters for species distinction, the results are partly contradictory to Schütt’s
synonymy and distribution map. In particular the taxa B. saulcyi and B. hawadieriana, both
described by Bourguignat in 1853, are not synonymous with B. phialensis but with B. badiella. To
prove this assumption, the most important parts of the original diagnosis of both taxa and of B.
badiella are quoted:
Bithynia saulcyi (BOURGUIGNAT 1853: 63, pl. II, figs 43-45), original description: “Testa
subconico-ventricosa, perforata […] anfractibus 5 convexis, sutura impressa […] operculo nucleo
subcentrali”. — This translates to: The shell is subconical ventricose, perforate […] with five
convex whorls, impressed suture […] the nucleus of the operculum is subcentral.
Bithynia hawadieriana (BOURGUIGNAT 1853: 63, pl. II, figs 46, 47), original description: “Testa
ovato-conoidea, perforata […] sutura valde impressa […] operculo striis concentricis, ac nucleo
subcentrali ornato”. — This translates to: The shell is ovate conical, perforate […] the suture
strongly impressed […] the operculum with concentric striae and a subcentral nucleus.
Bithynia badiella (KÜSTER loc. cit.), original description: “Das Gehäuse ist fast durchgehend
geritzt […] die fünf Windungen stark gewölbt, oben verflacht […] der Mittelpunkt (des Deckels)
unter der Mitte nach rechts liegend”. — This translates to: The shell with a slit-like umbilicus […]
the five whorls are strongly rounded and flat on top […] the nucleus (of the operculum) shifted to
the right below the centre.
Comparing the figures of both authors, the conspecificity of the taxa seems to be likely and
should be proven by investigating Bourguignat’s types. If this point of view can be supported, a
problem of priority arises, because the pages 57-80 of the “Conchylien-Cabinet 1 (21)” had been
published in 1853 and not in 1852 (as for example cited by BROWN & WRIGHT 1980: 345). Only
the exact publication dates then can decide about the priority of Küster’s vs. Bourguignat’s name.

348 E. NEUBERT
Additionally, this group should be compared very carefully to Bithynia “species” from Greece,
Asia Minor and Iran and to east-Mediterranean forms of Bithynia leachi (Sheppard, 1823). Since a
revision of this group is far beyond the scope of this work, the nomenclatural problems and the
task of synonymising the other taxa is left for subsequent investigations.
Family Stenothyridae
Genus Stenothyra Benson, 1854
1854 Stenothyra Benson. — Ann. Mag. Nat. Hist. 18: 496.
Diagnosis: Small ovoid to oval-elongate shells with an almost circular aperture. The shell is
often compressed dorsoventrally. The aperture is separated from the body whorl by a characteristi-
cally triangular and somewhat tumid area. Internally, a small parietal lamella can be found running
parallel to the peristome. The surface of the shell is smooth or ornamented with rows of small
bristles and spiral rows of minute punctuation. Internally, the corneous to calcareous operculum
has two parallel ridges.
Type species: Nematura deltae Benson, 1836.
Stenothyra arabica n. sp. Figs 33-34
Type material and locus typicus: holotype SMF 311265, paratypes SMF 311266/6, U.A.E., Ras al-
Khaimah, S of Rams, 25°52'N 56°00'E, material from excavations at the sea shore, 11.VII.1995, M. Apel.
Diagnosis: A minute Stenothyra species with a humped outline to the shell, a slit-like
opened umbilicus and red-brown maculation on the dorsal surface of the last whorl.
Description: The small oval and somewhat tumid shell reaches a maximum height of
2.2 mm. The protoconch consists of 1.5 smooth whorls and is quite blunt.
The teleoconch whorls are smooth, any punctuation is absent. The upper two whorls are
evenly rounded with a suture of medium depth. The following whorls increase considerably in size.
The suture is very shallow and, especially in the last whorl, adpressed to the upper whorl producing
a somewhat humped outline of the last two whorls. The body whorl is only slightly compressed
dorsoventrally, the outline is rounded.
The aperture is circular with a minute angle at the place where the suture of the descending
last whorls meets the peristome. The parietal triangular area is small and shows some faint growth
lines. On the dorsal surface of the body whorl, irregularly arranged red-brown spots are found. The
umbilicus is slit-like open.
Measurements (holotype, Figs 33-34): H = 1.80; D = 1.15; PH = 0.55; PD = 0.55,
W = 4.5.
Etymology: This species is called S. arabica as it is known only from its Arabian type locality.
Affinities: Compared to S. minima (Sowerby, 1836), the new species is smaller and differs
in the shape of the last whorls (SUBBA RAO 1989: 89, figs 151-154, 163-164). Stenothyra minima is
recorded from western India (Kathiawar, Little Rann of Cutch to Sri Lanka) and is close to S.
arabica n. sp. (ANNANDALE & PRASHAD 1921: 129). BRANDT (1974: 131) reports that in S. minima
the patches are often connected by obsolete bands while in S. moussoni v. Martens, 1897, there is
only one big patch of brown pigment on the dorsum. Stenothyra maculata Brandt, 1974 differs by
its size, the deeper sutures and the subangulate whorls and S. prasongi Brandt, 1974 by size,
punctuation and its basal keel. In S. hungerfordiana Nevill, 1880, the outline of the teleoconch is
evenly oval and the umbilicus is completely closed.
This is the first record for this genus for the Arabian Peninsula.

Terrestrial and freshwater molluscs of the Arabian Peninsula 349
Genus Gangetia Ancey, 1890
1890 Gangetia Ancey. — Bull. Soc. Malac. France 7: 163.
Diagnosis: Very small shells with a well-rounded body whorl. The aperture is somewhat
protruding, the thick operculum is paucispiral with two small transverse ridges on the inner
surface.
Type species: Hydrobia (Belgrandia) miliacea Nevill, 1880.
Gangetia miliacea (Nevill, 1880)
1880 Hydrobia (Belgrandia) miliacea Nevill. — J. Asiatic Soc. Bengal 49 (2): 161 (Port Canning, West Bengal).
Description: The minute shell is conical, the protoconch somewhat flattened. The teleo-
conch whorls are rounded with a suture of medium depth. The surface of the teleoconch whorls is
smooth. The aperture is somewhat elongate ovoid, there is no triangular area between the aperture
and the body whorl compared to species of Stenothyra.
Remarks: In the Arabian Peninsula, this species is known only from the Dhofar area, Oman,
where it lives in brackish waters or even in mangrove areas on algae and water plants (BROWN &
GALLAGHER 1985: 130-131). It is also known from the Ganges-delta in India (SUBBA RAO 1989).
Gangetia miliacea was probably introduced by human activities.
Family Iravadiidae
Genus Iravadia Blanford, 1867
1867 Iravadia Blanford. — J. Asiatic Soc. Bengal 36 (2): 56.
Diagnosis: The shell is elongated ovate-conical, an umbilicus is usually absent. The spiral
sculpture predominates, axial ribs may occur. The protoconch is depressed to sometimes dome-
shaped. The nucleus of the oval operculum is on the columellar edge, the operculum shows
concentric growth lines. The cephalic tentacles are often black-banded or spotted. For more
detailed information see PONDER (1984).
Type species: Iravadia ornata Blanford, 1867.
Iravadia quadrasi (O. Boettger, 1893) Fig. 35
1893 Alvania quadrasi O. Boettger. — Nachrbl. dtsch. malakozool. Ges. 25 (7/8): 101 (Philippines, Malabon close to
Manila).
Material: U.A.E.: Ras al-Khaimah, S of Rams, 25°52'N 56°00'E, material from excavations at the sea shore,
11.VII.1995, M. Apel, SMF 311528/6.
Type material: Lectotype Alvania quadrasi, SMF 225147.
Description: The shell is conical elongate with a short truncate protoconch. The teleo-
conch whorls are flat to somewhat rounded with a suture of medium depth. The surface is covered
by two (midbody whorls) to six (body whorl) spiral ridges crossed by strong overriding axial ribs
thus causing a reticulate pattern. The complete shell surface is covered by microscopically fine
spiral lines. The aperture is subcircular with a small varix behind the lip.
Measurements (lectotype SMF 225147, Fig. 35): H = 4.05; D = 2.10; PH = 1.80; PD = 1.40;
W = 4.5.
Remarks: This species was recorded by BROWN & GALLAGHER (1985) from several localities
along the Oman coast. According to PONDER (1984) it is widespread throughout brackish waters
in the Indopacific area.

350 E. NEUBERT
Family Assimineidae
Genus Assiminea Fleming, 1828
1828 Assiminea Fleming. — Hist. Brit. Anim.: 275.
Diagnosis (cf. BROWN 1994: 95): Small to medium-sized globose to conical shells. The
operculum is paucispiral and entirely corneous. The radula has an accessory plate between lateral
and first marginal tooth. The central tooth lacks basal denticles.
Type species: Assiminea grayana Fleming, 1828.
Assiminea nitida nitida (Pease, 1864) Fig. 36
1865 Hydrocena nitida Pease in Charpentier. — Proc. Zool. Soc. London 1864: 674 (no type locality).
Material: Saudi Arabia: Eastern Province, al-Qatif Oasis, freshwater course W of town, 26°36'N 49°58'E,
31.I.1992, R. Kinzelbach & I.A. Nader, SNMNH-MO 9/15, SMF 311529/15, KINZ (3); al-Aba Oasis (Saadah, Dumayn),
9.VI.1992, R. Kinzelbach, SNMNH-MO 10/6, SMF 311530/6; al-Qatif Oasis, palm plantations W of centre, irrigation
channels, 14.VI.1992, R. Kinzelbach, SNMNH-MO 11/24, SMF 311531/24, NEUB (10); Tarut, plantations N of Tarut
centre, 14.VI.1992, R. Kinzelbach, SNMNH-MO 12/22, SMF 311532/22.
Description: The shells are small elongate conical reaching a height of 2.5 mm. They are
deep brown, fresh specimens are glossy with very fine growth lines. A fine spiral sculpture
(magnification c. 50 ×) may be present. Subsuturally, a spiral thread is visible in almost all
specimens. An additional spiral thread may occur surrounding the periomphalum.
The aperture is pear-shaped with a sharp peristome. The columella is thickened, the umbilicus
closed. A thin callus connects the peristome.
Measurements (illustrated specimen, Fig. 36): H = 2.70; D = 1.80; W = 5.5.
Remarks: This species is well known from many localities in the Indopacific area (ABBOTT
1958) but is recorded here for the first time from the Arabian Peninsula. It is divided into several
subspecies which differ in having more or less developed subsutural spiral and umbilical threads.
The nominate subspecies lives in estuarine habitats of the Indopacific shores. Unfortunately, there
is no precise information on the salinity of the water bodies where Assiminea nitida lives. Some of
them may be brackish, but others seem to be freshwater courses.
It should be noted that in the specimens from the Arabian Peninsula the protoconch is
somewhat broader than in specimens illustrated by BRANDT (1974: pl. 11, fig. 91) from Thailand.
Family Thiaridae
Genus Thiara Roeding, 1798
1798 Thiara Roeding. — Museum Boltenianum 2: 109.
Diagnosis: The shells are broadly conical to turreted elongate. The upper whorls may be
shouldered with projecting spines. The paucispiral operculum displays a basal nucleus. The females
are parthenogenetic, males are rare.
Type species: Helix amarula Linnaeus, 1758.
Thiara scabra (O.F. Müller, 1774)
1774 Buccinum scabrum O.F. Müller. — Vermium terrestrium et fluviatilium 2: 136 (“In paludosis littoris Coromandel”).
Material: Oman: Fanja W of Muscat, I.1993, H. Pauscher, NEUB (7); Oasis of Buraimi, I.1993, H. Pauscher, NEUB
(1); Wadi Manqal, effluent of Wadi Hilm (Qalhat), near Kibdiyah (= Kebdah), 22°41'N 59°18'E, 26.IV.1978, A.B. Paltrinieri,
ONHM 610.27 (1); Wadi Qalhat/Hilm, 22°41'N 59°22'E, 18.IV.1978, A.B. Paltrinieri, ONHM 610.23 (1).

Terrestrial and freshwater molluscs of the Arabian Peninsula 351
Description: Conical ovate shell with usually 8-12 whorls separated by a deep suture, the
apex is often eroded. The upper whorls are regularly ribbed, the ribs are often spinose at the
shoulder. On the last whorls, the lower part of the ribs is weak or the ribs are completely absent.
The lower part of the whorls displays numerous spiral threads which can be very strong on the
base of the last whorl. The shells are yellowish to olive-green with an irregular pattern of red-brown
spots. The aperture is acute oval, the base is subcanaliculated.
Remarks: Thiara scabra is known only from some localities in northern Oman and Wadi
Hadramaut in Yemen (BROWN 1994: 101).
Genus Melanoides Olivier, 1804
1804 Melanoides Olivier. — Voyage a l’Empire Ottoman 2: 40.
Diagnosis: The shells are turreted elongate with axial ribs often nodulose when crossed by
spiral sculptural elements. The paucispiral operculum displays a basal nucleus. The females are
parthenogenetic with a brood pouch which is separated from the oviduct.
Type species: Melanoides fasciolata Olivier, 1804 = Nerita tuberculata O.F. Müller, 1774.
Melanoides tuberculata (O.F. Müller, 1774)
1774 Nerita tuberculata O.F. Müller. — Vermium terrestrium et fluviatilium 2: 191 (“In littore Coromandel”).
Material: Saudi Arabia: Eastern province, Jubail, Sabkhat al-Fasl, saltwater canal near Royal Commission building,
22./28.I.1992, R. Kinzelbach, SNMNH-MO 144/136, SMF 311698/20, KINZ (20); Eastern province, Ayn ibn Umayr,
26°41'N 49°51'E, 16.II.1994, P. Symens, SNMNH-MO 145/12, SMF 311699/5; Jubail, Haii al-Bahr, 27°05'N 49°35'E,
8-31.I.1992, R. Kinzelbach & E. Neubert, SNMNH-MO 146/2; al-Hasa Oasis, al-’Uyun palm plantation, freshwater ditch with
dense vegetation, 25°36'N 49°34'E, 27.I.1992, R. Kinzelbach & E. Neubert, SNMNH-MO 147/100, SMF 311700/10, KINZ
(10); same locality and date, in loam excavated from ditches, R. Kinzelbach, W. Schneider & E. Neubert, SNMNH-MO 148/22,
SMF 311701/10, KINZ (10); same locality, 14.XII.1992, R. Kinzelbach & M. Werner, SNMNH-MO 149/17, SMF 311702/10,
KINZ (6); al-Qatif Oasis, freshwater course W of town, 26°36'N 49°58'E, 31.I.1992, R. Kinzelbach & I.A. Nader, SNMNH-
MO 150/19, SMF 311703/10; al-Aba Oasis (Saadah, Dumayn), 9.VI.1992, R. Kinzelbach, SNMNH-MO 151/67, SMF
311704/10, KINZ (10); artificial pond east of main road Safwa-Jubail, 2 km N of exit Safwa, 9.VI.1992, R. Kinzelbach,
SNMNH-MO 152/2; al-Qatif Oasis, palm plantations W of centre, irrigation channels, 14.VI.1992, R. Kinzelbach, SNMNH-
MO 153/100, SMF 311706/15, KINZ (15); Tarut, plantations N of Tarut centre, 14.VI.1992, R. Kinzelbach, SNMNH-
MO 154/150, SMF 311707/20, KINZ (20); small pond E of the highway Khobar-Jubail, 3.4 km S of exit Juaymah, 25.V.1995,
E. Neubert & R. Janssen, SNMNH-MO 155/5; Central province, al-Aflaj, Laila Lakes, 22°11'N 46°42'E, 550 m, 13.III.1990,
F. Krupp & W. Schneider, SNMNH-MO 157/10, SMF 311711/11; Central province, Wadi Dawasir, 20°30'N 44°46'E, 800 m,
13.III.1990, F. Krupp & W. Schneider, SNMNH-MO 158/5, SMF 311712/5; Asir mountain region, al-Mekwah, Marble
Village, 700 m, II.1994, P. Symens & M. Werner, SNMNH-MO 156/6, SMF 311708/6, SMF 311709/1 (preserved); Asir
mountain region, Wadi Ilyab, 20°04'N 40°53'E, 160 m, 17.III.1990, F. Krupp & W. Schneider, SNMNH-MO 159/7,
SMF 311713/7; Asir mountain region, Wadi Turabah, 20°32'N 41°17'E, 1450 m, 20.III.1990, F. Krupp & W. Schneider,
SNMNH-MO 160/18, SMF 311714/18; Southern Hijaz, Wadi Murwani, 22°12'N 39°39'E, 360 m, 22.III.1990, F. Krupp &
W. Schneider, SNMNH-MO 161/11, SMF 311715/10. — Yemen: Wadi near Kihab, 20.II.1990, M. Scholz, NEUB (5);
Hadramaut, Buraat, 6.IV.1988, A. Liebegott, LIEB (2). — O m a n : Fanja W of Muscat, I.1993, H. Pauscher, NEUB (17); Jabal
Akhdar, oasis between ar-Rustaq and al-Hazm, I.1993, H. Pauscher, NEUB (1); Eastern Hajar, Jabal Bani Jabir, Wadi Bani
Khalid east of al-Mintrib, I.1993, H. Pauscher, NEUB (12); Jabal Akhdar, irrigation channels at Nizwa, I.1993, H. Pauscher,
NEUB (15)¸ SMF 311687/3, Dhofar, NE slope of Qara mountains, Jibjat area (I.a.e.n. KR 53), Girod ded. 1997, GIROD (12),
SMF 311688/14 (+ fragments); same locality (I.a.e.n. KR 159), Girod ded. 1997, SMF 311689/3 (+ fragments); same locality
(I.a.e.n. KR 73), Girod ded. 1997, SMF 311692/5; same locality (I.a.e.n. KR 6B), Girod ded. 1997, GIROD; Dhofar, Wadi
Darbat, 17°06'N 54°27'E, 16.VI.1980, A.B. Paltrinieri, ONHM 610.29 (2); Dhofar, Wadi Darbat, 17°02'N 54°26'E,
16.VI.1980, A.B. Paltrinieri, ONHM 610.11 (4); Wahiba Sands, unknown locality, 19.IV.1978, A.B. Paltrinieri, ONHM 610.24
(2); Wadi Bani Khalid, 23°30'N 59°08'E, 16.IV.1978, A.B. Paltrinieri, ONHM 610.1 (1); Wadi Bani Khalid, c. 23°30'N
59°08'E, II.1978, A.B. Paltrinieri, ONHM 610.8 (3); Wadi Qalhat/Hilm, 22°41'N 59°22'E, 17.V.1978, A.B. Paltrinieri,
ONHM 610.22 (1); Dhofar, Ain Arzat (= Razat), 17°07'N 54°14'E, 15.VI.1978, A.B. Paltrinieri, ONHM 610.10 (9);
Den/Dann, Jabal Khawr, 23°10'N 57°20'E, 21.III.1977, A.B. Paltrinieri, ONHM 610.7 (7); Wadi Gholshin, above wadi,

352 E. NEUBERT
450 m, A.B. Paltrinieri, ONHM 610.4 (4); Wadi Manqal, effluent of Wadi Hilm (Qalhat), near Kibdiyah (= Kebdah), 22°41'N
59°18'E, 26.IV.1978, A.B. Paltrinieri, ONHM 610.27 (10); same data, ONHM 610.28 (19); Wadi Bani Awf, 23°17'N
57°28'E, 2.V.1978, A.B. Paltrinieri, ONHM 610.26 (1); Wadi al-Batha S of Bilad Bani bu Ali, 22°00'N 59°20'E, 27.IV.1978,
A.B. Paltrinieri, ONHM 610.19 (27); lower Wadi Sumayl, 23°25'N 58°08'E, 25.II.1986, A.B. Paltrinieri, ONHM 610.16 (9);
Wadi Qalhat/Hilm, lowest pool, 22°41'N 59°22'E, 18.IV.1978, A.B. Paltrinieri, ONHM 610.16 (9); same data, ONHM 610.23
(32); Wadi Bani Khalid, ad-Dawa, 23°36'N 59°05'E, 17.IV.1975, A.B. Paltrinieri, ONHM 610.13 (1); Buraimi Falaj, 24°15'N
55°45'E, 14.XII.1976, A.B. Paltrinieri, ONHM 610.17 (11).
Description: Slender conical shells reaching up to 40 mm length. The upper whorls are
ribbed, on the lower whorls the ribs become weak or are absent. The ribs are crossed by spiral
threads causing a reticulate pattern. Usually, the base of the last whorl is dominated by the spiral
sculpture. All sculptural elements may vary to a big extent. The basic colour of the shell is olive-
green to dark red-brown. Many shells display a pattern of red spiral dots or short stripes which
may be arranged irregularly or form oblique to zigzag-shaped flames.
Remarks: This species is by far the most common freshwater snail of the Arabian Peninsula.
Due to its parthenogenetic mode of reproduction, a single specimen is sufficient to build up a
complete population. Melanoides tuberculata is widespread all over the world owing to human
activities.
Melanoides sp. cf. plicaria (Born, 1780)
1780 Helix plicaria Born. — Test. mus. Caes. Vindob.: 389, Taf. 16, Fig. 14 (“Patria ignota”).
Remarks: This taxon, with the shallow sutures and flattened sides of the whorls, was
reported by BROWN & GALLAGHER (1985: 134-135, fig. 9) from the U.A.E. Comparing his
descriptions and figures to those of STAHRMÜHLNER (1976: 582-584, Taf. 16, Fig. 182-194) of
Melanoides plicaria, some remarkable differences can be found. The shells of the Arabian Mela-
noides species are stout and much smaller, the ratio of height/breadth is different. The ribs of the
Pacific shells are less pronounced while the spiral grooves seem to be stronger. They are obviously
not conspecific.
As BROWN states several times, a revision of the Thiaridae is urgently needed to help to
identify this taxon.
Family Melanopsidae
Genus Melanopsis Férussac, 1807
1807 Melanopsis Férussac. — Essai d’une methode conchyliologique appliquée aux Mollusques fluviatiles et terrestres …:
70.
Diagnosis: Medium to large ovate shells which may be smooth to coarsely ribbed. The
acute-oval aperture usually with a broad distinct parietal callus, often concentrated like an
“angularis”, and a distinct siphonal canal. The operculum is paucispiral with a basal nucleus.
Type species: Buccinum praemorsum Linnaeus, 1758.
Melanopsis praemorsa (Linnaeus, 1758)
1758 Buccinum praemorsum Linnaeus. — Systema Naturae, ed. 10: 740 (“In Europa australiore”).
Material: Saudi Arabia: Eastern province, al-Aba Oasis (Saadah, Dumayn), 9.VI.1992, R. Kinzelbach, SNMNH-
MO 13/18, SMF 311533/18, KINZ (4); al-Qatif Oasis, freshwater course W of town, 26°36'N 49°58'E, 31.I.1992, R.
Kinzelbach & I.A. Nader, SNMNH-MO 14/50, SMF 311534/50; al-Qatif Oasis, palm plantations W of centre, irrigation
channels, 14.VI.1992, R. Kinzelbach, SNMNH-MO 15/33, SMF 311535/33; Tarut, plantations N of Tarut centre, 14.VI.1992,
R. Kinzelbach, SNMNH-MO 16/10, SMF 311536/10.

Terrestrial and freshwater molluscs of the Arabian Peninsula 353
Description: Shell small to medium-sized, non-umbilicate, turreted, broad conical. In
adults, the protoconch and the initial whorls are usually eroded. The teleoconch whorls are flat
and smooth to heavily ribbed. The suture is shallow and somewhat indented in ribbed shells.
The aperture is oval with an acute sinulus. In fresh specimens, the interior of the aperture
glazes from white to deep red-brown. Parietal, there is a big angularis. The columella is obliquely
truncate, the siphonal canal is short and straight. The columellar and parietal area is connected by
a callose shield which is often bluish.
Uneroded shells are a deep olive-green colour with up to three red-brown spiral bands.
Remarks: GLAUBRECHT (1993) presented the geographical distribution and systematics of
Melanopsis; he stated that the circum-Mediterranean population of Melanopsis consists of one
superspecies which should be named praemorsa.
According to AMADON (1966: 245), a superspecies has to constitute a monophyletic entity and
has to be composed of taxa which occur entirely or essentially allopatrically and are morphologi-
cally different. There is no doubt about polymorphism in the M. praemorsa-group but there is no
attempt in Glaubrecht’s work to prove or even discuss their monophyletic origin. There is no
outgroup-comparison, no discussion about homologous or analogous characters, and no character
states are defined. Allopatry is also not proven. The fossil history of Melanopsis throughout the
Tertiary is not given. There is no final statement about the generic placement of the isolated
Indopacific species as Glaubrecht assigns them to Melanopsis or Zemelanopsis Finlay, 1927. The
challenging distributional pattern “circum-mediterranean/New Zealand/New Caledonia” (the only
allopatry to be found in the group) is not investigated nor sufficiently explained. Thus, as the main
conditions for the superspecies concept are not addressed, his proposal for the M. praemorsa-
complex is rejected.
The very careful use of this theory and its delimitation in ornithology is exemplified by FRY et
al. (1988) in their magnificent monographic work on African birds. In all cases, only allo- or
parapatric species are accepted to form a superspecies. Others, whose ranges are known to overlap,
were removed from superspecies to form independent species (as for the Ground Hornbills
Bucorvus abyssinicus and B. cafer, see FRY et al. 1988: XIII). In the introduction to Vol. IV (KEITH
et al. 1992), the authors discuss their view of the superspecies concept again in detail, leaving no
doubt about its theoretical framework.
Finally, the problem remains unresolved and for this reason the author retains the name
Melanopsis “praemorsa” for all taxa. However, it should be kept in mind that for the Levantinian
group the names M. buccinoidea Olivier, 1801 for the smooth form, and M. costata Olivier, 1804
and M. nodosa Férussac, 1823 for the ribbed shells exist, representing the oldest names available.
The shells reported here from the oases in the Eastern Province all belong to the ribbed forms.
The localities given here correspond with those by BROWN & WRIGHT (1980) and seem to
represent the southern distribution limits of Melanopsis.
Pulmonata
Family Lymnaeidae
Genus Radix Montfort, 1810
1810 Radix Montfort. — Conchyliologie systématique et classification méthodique des Coquilles 2: 266.
Diagnosis: The spire of the shell is short. In contrast, the body whorl is considerably
enlarged.
Type species: Helix auricularia Linnaeus, 1758.

354 E. NEUBERT
A revision of Radix, based on anatomical and shell characters, is urgently needed. It is
surprising that a taxon like the Lymnaeidae, which is well represented in Europe, is still in such a
confused taxonomical state. Reliable identification is difficult since almost all species display a high
plasticity in shell characters. The presence of ecomorphs is the result of many of the widespread
“species” living in habitats of quite differing ecological structure. Consequently, a plethora of
names was, and still is being, created by several authors. Unfortunately, the last (and only) global
revision of the family by HUBENDICK (1951) does not provide a reliable solution.
During an expedition in 1990 as part of the Zoological Survey in Saudi Arabia, several lots of
Lymnaeidae were collected (KRUPP et al. 1990). In particular, the lymnaeid specimens from the al-
Aflaj basin in the central part of the peninsula deserve a more detailed investigation. This has been
postponed until preserved specimens are available for dissection.
Radix auricularia (Linnaeus, 1758)
1758 Helix auricularia Linnaeus. — Systema Naturae, ed. 10: 774 (“in Europae fluviis, stagnis”).
Material: Oman: Jabal Akhdar, Wadi al-Yemen, lower dam, 23°04'N 57°41'E, V.1980, A.B. Paltrinieri, ONHM
610.9 (2).
Description: The medium-sized to large shell can be characterised by its very short and
pointed spire compared to the height of the extremely large aperture. The outline of the shell is
concave, the upper rim of the aperture is nearly horizontal.
Remarks: This species was recorded by BROWN & WRIGHT (1980: 347) from suitable
habitats all over the peninsula. The specimens in the lot from Oman recorded here have a shape
very similar to that of R. auricularia. There was a third shell in this lot which has to be assigned to
R. natalensis in the sense used here. Thus it is very likely that both species live sympatrically in the
area. To determine this, more detailed research on the anatomy is needed for both the genus Radix
in general and the species R. natalensis in particular.
Radix natalensis (Krauss, 1848)
1848 Limnaeus natalensis Krauss. — Die südafrikanischen Mollusken: 85, Tab. V, Fig. 15 (“Natal, Südafrika”).
1894 Lymnaea arabica Smith. — Proc. Malac. Soc. London 1: 142, textfig. 3 (Oman).
Material: Saudi Arabia: Eastern Province, al-Hasa Oasis, al-’Uyun palm plantation, freshwater ditch with dense
vegetation, 25°36'N 49°34'E, 27.I.1992, R. Kinzelbach & E. Neubert, SNMNH-MO 17/6, SMF 311537/7, SMF 311553/1
(preserved); same locality, loam excavated from ditches, 14.XII.1992, R. Kinzelbach & M. Werner, SNMNH-MO 18/14,
SMF 311538/14; same locality, loam excavated from ditches, 27.I.1992, R. Kinzelbach, E. Neubert & W. Schneider, SNMNH-
MO 19/6, SMF 311539/6; al-Qatif Oasis, palm plantations W of centre, irrigation channels, 14.VI.1992, R. Kinzelbach,
SNMNH-MO 20/2, SMF 311540/3, KINZ (5); al-Aba Oasis (Saadah, Dumayn), 9.VI.1992, R. Kinzelbach, SNMNH-
MO 21/14, SMF 311541/14; al-Qatif Oasis, freshwater course W of town, 26°36'N 49°58'E, 31.I.1992, R. Kinzelbach & I.A.
Nader, SNMNH-MO 22/4, SMF 311542/4; Ayn Ibn Umayr, 16.II.1994, P. Symens, SNMNH-MO 23/1, SMF 311543/1. —
Om a n : Eastern Hajar, Jabal Bani Jabir, Wadi Bani Khalid E of al-Mintrib, I.1993, H. Pauscher, NEUB; Wadi Bani Khalid, c.
23°30'N 59°08'E, II.1978, A.B. Paltrinieri, ONHM (12); Wadi Bani Khalid, 23°30'N 59°08'E, 16.IV.1978, A.B. Paltrinieri,
ONHM 610.1 (2); Wadi Jizzi, 24°23'N 56°41'E, 12.XII.1976, A.B. Paltrinieri, ONHM 610.2 (7); Dhofar, Ain Arzat (= Razat),
17°07'N 54°14'E, 15.VI.1978, A.B. Paltrinieri, ONHM 610.10 (1); Wadi Gholshin, above wadi, 450 m, A.B. Paltrinieri
ONHM 610.4; Wadi Manqal, effluent of Wadi Hilm (Qalhat), near Kibdiyah (= Kebdah), 22°41'N 59°18'E, 26.IV.1978, A.B.
Paltrinieri, ONHM 610.27 (9); same data, A.B. Paltrinieri, ONHM 610.28 (21); Wadi Bani Awf, 23°17'N 57°28'E, 2.V.1978,
A.B. Paltrinieri, ONHM 610.26 (18); lower Wadi Sumayl, 23°25'N 58°08'E, 25.II.1986, A.B. Paltrinieri, ONHM 610.16 (4);
Wadi Qalhat/Hilm, lowest pool, 22°41'N 59°22'E, 18.IV.1978, A.B. Paltrinieri, ONHM 610.16 (11); Wadi Bani Khalid, ad-
Dawa, 23°36'N 59°05'E, 17.IV.1975, A.B. Paltrinieri, ONHM 610.13 (4); Jabal Akhdar, Wadi al-Yemen, lower dam, 23°04'N
57°41'E, V.1980, A.B. Paltrinieri, ONHM 610.9 (1); Wadi Qalhat/Hilm, 22°41'N 59°22'E, 18.IV.1978, A.B. Paltrinieri,
ONHM 610.23 (1).
Description: The shell of this species is medium-sized, its spire is shorter than the aperture,
which is only moderately expanded. The protoconch is acute, the suture of both protoconch and

Terrestrial and freshwater molluscs of the Arabian Peninsula 355
teleoconch is very deep. The surface of the shell has spiral rows of transverse grooves. The
columella is somewhat twisted (cf. BROWN 1994: fig. 79 a).
Remarks: This species is known from several localities from the Arabian Peninsula and was
described from this area under the name Lymnaea arabica by SMITH (1894). BROWN (1994)
frequently suggested that this species intergrades with Radix auricularia. However, all the material
available is close to R. natalensis sensu auct., so the specimens are identified as this species here. No
shells in the various collections investigated showed any affinity to R. auricularia except for one lot
from Oman (cf. paragraph on Radix auricularia).
Genus Galba Schrank, 1803
1803 Galba Schrank. — Fauna Boica (2) 2: 262, 285.
Diagnosis: As for the species.
Type species: Galba pusilla Schrank, 1803 = Buccinum truncatulum O.F. Müller, 1774.
Galba truncatula (O.F. Müller, 1774)
1774 Buccinum truncatulum O.F. Müller. — Vermium terrestrium et fluviatilium 2: 130 (“In agro Thangelstedtiensis
Saxoniae”).
Material: Saudi Arabia: Asir mountain region, Raydah escarpment, 1800 m, 29.I.1994, P. Symens & M. Werner,
SMF 311544/3 (preserved); Raydah escarpment, al-Aqaba spring, 2310 m, 18°11'50.0"N 42°24'30.2"E, 5.V.1994, W.
Schneider & F. Krupp, SMF 311545/6, HLMD (6) (all preserved); Asir mountain region, Raydah escarpment, 2350 m,
29.I.1994, P. Symens & M. Werner, SNMNH-MO 24/1.
Description: Small turreted shells with a spire of nearly the same height as the aperture.
The protoconch is conical and smooth. The suture of the teleoconch is deeply impressed, its
whorls are well rounded. The columella is straight and reflected over the umbilicus. The umbilicus
is often open and slit-like.
Remarks: Galba truncatula was hitherto only known from Yemen (BROWN & WRIGHT 1980:
347). The material presented here provides the first records for this species from the mountainous
regions of south-west Saudi Arabia.
Genus Stagnicola Leach in Jeffreys, 1830
1830 Stagnicola Leach in Jeffreys. — Trans. Linn. Soc. London 16 (2): 376.
Diagnosis: Turreted shells with a spire of the same height or much longer than the aperture.
Type species: Buccinum palustre O.F. Müller, 1774.
Stagnicola palustris (O.F. Müller, 1774)
1774 Buccinum palustre O.F. Müller. — Vermium terrestrium et fluviatilium 2: 131 (“In paludosis”).
Description: The shell is turreted, the suture of the teleoconch whorls is deep. The height
of the spire always exceeds the height of the aperture. A coarse malleate sculpture is present on the
shell’s surface. An umbilicus is absent. The columellar margin is slightly broadened and somewhat
twisted.
Remarks: There is one record of this species from south-western Saudi Arabia from the
Sudah stream in the Khamis Mushayt-Abha area (3100 m) (BROWN & WRIGHT 1980: 347). This
locality is far from the hitherto known distribution of this species. Further anatomical studies are
required to confirm this identification.

356 E. NEUBERT
Family Planorbidae
Genus Planorbis O.F. Müller, 1774
1774 Planorbis O.F. Müller. — Vermium terrestrium et fluviatilium 2: 152.
Diagnosis: Medium-sized disciform shells with blunt or carinate whorls. Short penis
sheath, penis without sclerotisation or stylet.
Type species: Helix planorbis Linnaeus, 1758.
Planorbis planorbis (Linnaeus, 1758)
1758 Helix planorbis Linnaeus. — Systema Naturae, ed. 10: 769 (“In Europae stagnis”).
Description: The corneous shell may reach up to 15 mm in diameter and a height of
3 mm. The upper surface is nearly flat, the subsurface slightly concave. Usually, the diameter of
the last whorl reaches twice the width of the preceding one. There is a sharp to blunt keel on the
upper rim of the shell (the shell is sinistral).
Remarks: Planorbis planorbis was recorded by BROWN & WRIGHT (1980) from dry shells at
the archaeological excavation site in Gerrha south of Dhahran, Saudi Arabia. There are no
additional records from the Arabian Peninsula and it is doubtful whether this species actually lives
in the Arabian Peninsula.
Genus Gyraulus Charpentier, 1837
1837 Gyraulus Charpentier. — Mém. Soc. Helv. Sci. Nat. 1: 21.
Diagnosis: Small disciform shells with the whorls rapidly increasing in width. The shell
surface may be smooth or covered by a fine spiral to reticulate sculpture. The penis has a
subterminal opening and a sclerotised stylet.
Type species: Planorbis hispidus Draparnaud, 1805 (= Planorbis albus O.F. Müller, 1774).
Remarks: Although revised by MEIER-BROOK (1983: 86), the taxonomy of the Gyraulus
species of the east-Mediterranean region is still unclear. Most commonly, the names Gyraulus
ehrenbergi (Beck, 1837), G. piscinarum, G. hebraicum (both Bourguignat, 1852) and G. euphraticus
(Mousson, 1874) are used for the Turkish and Levantine species. In the author’s opinion, G.
ehrenbergi is restricted to the Nile area and does not occur in Syria or Turkey. This view is based on
a large number of Gyraulus specimens collected by Kinzelbach in Syria and Kinzelbach & Neubert
in Turkey (SMF and NEUB). All Gyraulus specimens from Egypt, the type area of G. ehrenbergi,
are smooth and they never display spiral sculptural patterns on the shell surface. This is in contrast
to almost all specimens investigated from the Levant, which show faint to very strong spiral
sculptural elements. Their presence or absence depends on the circumstances of the collection. Old
and eroded specimens often loose the periostracal sculpture, but under high magnification small
wrinkles may be observed indicating the presence of these structures. A reliable revision needs to
be undertaken to clarify the taxonomy of the G. piscinarum-complex.
Gyraulus piscinarum (Bourguignat, 1852)
1852 Planorbis piscinarum Bourguignat. — Testacea novissima quae Cl. de Saulcy in itinere per Orientem annis 1850 et
1851, collegit: 22 (“In aquis piscinarum ac stagnantibus prope Heliopolim Syriae” [= Baalbek]).
Description: This medium-sized Gyraulus species reaches a diameter of up to 6 mm. The
shell has three to four rapidly increasing whorls. The whorls are rounded to only slightly angulate
at the periphery. The shell’s subsurface is convex. The surface shows axial growth lines, which may
be somewhat pronounced. A faint reticulate sculpture is present.

Terrestrial and freshwater molluscs of the Arabian Peninsula 357
Remarks: MEIER-BROOK (1983: 52) dissected specimens from Diyarbakir, Turkey. As he
outlines, this species is characterised by its diffuse mantle pigmentation and the tadpole-type BC.
Topotypic material from Baalbek, Lebanon should be examined in detail to confirm these
characters.
In the Arabian Peninsula, this species was recorded by BROWN & GALLAGHER (1985: 138) from
Oman. In the author’s opinion, these specimens should be re-examined carefully and compared to
G. convexiusculus.
Gyraulus convexiusculus (Hutton, 1849)
1849 Planorbis convexiusculus Hutton. — J. Asiatic Soc. Bengal 18 (2): 657 (Candahar, Afghanistan).
Material: Saudi Arabia: Eastern Province, al-Qatif Oasis, palm plantations W of centre, irrigation channels,
14.VI.1992, R. Kinzelbach, SNMNH-MO 25/2; al-Hasa Oasis, al-’Uyun palm plantation, freshwater ditch with dense
vegetation, 25°36'N 49°34'E, 27.I.1992, R. Kinzelbach & E. Neubert, SNMNH-MO 26/1, SMF 311549/2, SMF 311552/2
(preserved), KINZ (1); same locality, loam excavated from ditches, 14.XII.1992, R. Kinzelbach & M. Werner, SMF 311546/1;
al-Qatif Oasis, freshwater course W of town, 26°36'N 49°58'E, 31.I.1992, R. Kinzelbach & I.A. Nader, SNMNH-MO 27/4,
SMF 311547/2; al-Aba Oasis (Saadah, Dumayn), 9.VI.1992, R. Kinzelbach, SNMNH-MO 28/16, SMF 311548/16. —
O m a n : Wadi Qalhat/Hilm, 22°41'N 59°22'E, 18.IV.1978, A.B. Paltrinieri, ONHM 610.23.
Description: A small Gyraulus species reaching up to 5 mm in diameter and about 1.8 mm
in height; the species is thus rather squat. The last whorl only deflects slightly, its periphery is
rounded to angled, often with a fringe of dark periostracum. The shell’s surface is glossy, but fine
growth lines are visible. A reticulate pattern is absent.
Remarks: Adult specimens of G. convexiusculus often show a somewhat thickened lip in the
aperture. It is not known whether this character is present in all populations. This species differs
from G. euphraticus (Mousson, 1874), which might also occur in this area, in that the latter species
is more flattened and its whorls increase more slowly. From the Arabian Peninsula, G. convexius-
culus is recorded from al-Qatif and al-Hasa oases and also from Bahrain, Oman (Trucial Coast)
and Yemen (BROWN & WRIGHT 1980: 349). Identification of specimens from the south-western
area of the Arabian Peninsula is doubtful and needs to be re-checked anatomically. Confusion with
Abyssinian taxa should also be taken into account.
Genus Biomphalaria Preston, 1910
1910 Biomphalaria Preston. — Ann. Mag. Nat. Hist. (8) 6: 535.
Diagnosis: Medium-sized discoid shells with convex, angular or carinate whorls. Tooth-like
lamella may occur on the inner wall of the shell near the aperture.
Type species: Biomphalaria smithi Preston, 1910.
Biomphalaria arabica (Melvill & Ponsonby, 1896)
1896 Planorbis arabicus Melvill & Ponsonby. — Proc. Malac. Soc. London 2: 3, pl. 1, figs 15-17 (Dhofar).
Material: Saudi Arabia: Asir mountain region, Wadi al-Sharan at the road from Taif to al-Baha, S of Bani Sa’ad,
1750 m, 2.VI.1995, E. Neubert et al., SNMNH-MO 29/1; Wadi Turabah, 1500 m, 20°28'N 41°15'E, 20.III.1990, W.
Schneider & F. Krupp, SNMNH-MO 40/2, SMF 311568/3 (preserved); Northern Hijaz, Khaibar Oasis, 750 m, 25°44'N
39°17'E, 28.III.1990, W. Schneider & F. Krupp, SNMNH-MO 41/2, SMF 311569/1 (preserved). — Yemen: Hadramaut,
area of Abiad, al-Achmar at the road from Aden to Makalla, c. 700 m, 1.IV.1988, A. Liebegott, LIEB (1). — O m a n : Dhofar,
NE slope of Qara mountains, Jibjat area (I.a.e.n. KR 73), Girod ded. 1997, SMF 311690/5; same locality, (I.a.e.n. KR 53),
Girod ded. 1997, SMF 311691/4, GIROD (1); same locality, (I.a.e.n. KR 159), Girod ded. 1997, SMF 311693/12; Dhofar,
Wadi Darbat, 17°06'N 54°27'E, 16.VI.1980, A.B. Paltrinieri; ONHM 610.29 (19); Dhofar, Ain Arzat (= Razat), 17°07'N
54°14'E, 15.VI.1978, A.B. Paltrinieri, ONHM 610.10 (20); Dhofar, Falaj Arzat, 17°03'N 54°13'E, 17.VI.1980, A.B.
Paltrinieri, ONHM 610.5 (1); Dhofar, Khwar Rowri, 17°03'N 54°25'E, 16.VI.1980, A.B. Paltrinieri, ONHM 610.54 (2);

358 E. NEUBERT
Dhofar, Khwar Rowri, 17°03'N 54°25'E, 16. & 18.VI.1980, A.B. Paltrinieri, ONHM 610.53 (4); Dhofar, Khwar Sawli,
17°02'N 54°20'E, 16.VI.1980, A.B. Paltrinieri, ONHM 610.51 (1); Dhofar, Wadi Sahalnaut, 17°08'N 54°11'E, 17.VI.1980,
A.B. Paltrinieri, ONHM 610.14 (4).
Description: The discoidal shell measures 5.2 mm in height and up to 17 mm in diameter.
The umbilicus is wide, reaching one-third of the shell diameter. The whorls are rounded to bluntly
angular.
Remarks: This species is widespread in all freshwater habitats in the western part of the
Arabian Peninsula. The taxonomic position (BROWN 1994: 195, fig. 122) has to be re-evaluated as
conspecificity with Biomphalaria pfeifferi (Krauss, 1848) from Africa might be possible.
Genus Bulinus Müller, 1781
1781 Bulinus Müller. — Der Naturforscher 15: 6.
Diagnosis: Sinistral spired shell, with gently curved whorls, angular or carinate. The penis
does not project freely into the penis sheath. It is a long, coiled, eversible tube attaching at the
upper and lower end of the sheath.
Type species: Bulinus senegalensis Müller, 1781.
Bulinus truncatus (Audouin, 1827)
1827 Physa truncata Audouin. — Descr. de l’Égypte II: 166; Atlas pl. 2, fig. 27 (Egypt).
Description: The shell of this species is pale and of medium size (reaching 10 mm in
height, 6 mm in diameter) and may vary considerably. In particular, the spire may be elongate to
truncate. The whorls are bluntly shouldered, the columellar margin is narrow and somewhat
twisted.
Remarks: This species is one of the best-known molluscan taxa because of its medical
importance. As with several of its congeners, it acts as an important intermediate host of Schisto-
soma haematobium. The species is widespread in tropical Africa and is found throughout the
western part of the Arabian Peninsula (BROWN & WRIGHT 1980). These authors indicate two
records from the eastern part of the peninsula. For more detailed information on this species,
BROWN (1994) should be consulted.
Bulinus beccari (Paladilhe, 1872)
1872 Physa beccari Paladilhe. — Ann. Mus. Stor. Nat. Genova 3: 23, tav. 1, figs 7-8 (Yemen: dry stream bed near Aden).
Material: Saudi Arabia: Southern Hijaz, Wadi Murwani, 360 m, 22°12'N 39°39'E, 22.III.1990, W. Schneider &
F. Krupp, SNMNH-MO 38/3, SMF 311565/2; Southern Hijaz, Wadi Ilyab, 160 m, 20°04'N 40°53'E, 17.III.1990, W.
Schneider & F. Krupp, SNMNH-MO 39/1, SMF 311566/1 (preserved), SMF 311567/2.
Description: This Bulinus species differs from B. truncatus in its smaller size (7 mm in
height, 3.4 mm in diameter) and more turreted shell. The whorls are evenly rounded and smooth
and thus differ from the Egyptian Bulinus forskalii (Ehrenberg, 1831), which is similar but always
ribbed.
Remarks: This species ranges throughout the south-western region of the Arabian Peninsula.
A detailed catalogue of localities is given in BROWN & WRIGHT (1980: 352).
Bulinus wrighti Mandahl-Barth, 1965
1965 Bulinus reticulatus wrighti Mandahl-Barth. — Bull. World Hlth. Org. 33: 41 (Aden).
Description: The small shell has a reticulate sculpture on the surface. It is globose and has
a large umbilicus. The spire is truncate with a deep suture. The whorls are somewhat flat-topped.

Terrestrial and freshwater molluscs of the Arabian Peninsula 359
Remarks: This species is known from several localities in Saudi Arabia, Yemen and Oman.
It seems to be restricted to the eastern part of the Arabian Peninsula. For a complete list see BROWN
& WRIGHT (1980: 352-353) and BROWN (1994: 247).
Genus Indoplanorbis Annandale & Prashad, 1920
1920 Indoplanorbis Annandale & Prashad. — Rec. Indian Mus. 22 (4): 578.
Diagnosis: Medium-sized discoid shells, whorls higher than wide. Anatomical characters
similar to those of Bulinus.
Type species: Planorbis exustus Deshayes, 1834.
Indoplanorbis exustus (Deshayes, 1834)
1834 Planorbis exustus Deshayes. — In: Belanger: Voy. Indes-Orientales: 417, pl. 1, figs 11-13 (“dans les lieux marécageux
de la côte du Malabar”).
Material: Saudi Arabia: Eastern Province, al-Hasa Oasis, al-’Uyun palm plantation, freshwater sewage channel,
25°36'N 49°34'E, 27.I.1992, R. Kinzelbach & E. Neubert, SNMNH-MO 30/30, SMF 311550/20 (preserved), SMF 311551/30,
KINZ (10). — O m a n (all collected by A.B. Paltrinieri, unless otherwise stated): Eastern Hajar, Jabal Bani Jabir, Wadi Bani
Khalid E of al-Mintrib, I.1993, H. Pauscher, NEUB (2); Wadi Bani Khalid, c. 23°30'N 59°08'E, II.1978, ONHM 610.8 (26);
same locality, 16.IV.1978, ONHM 610.1 (2); Wadi Qalhat/Hilm, 22°41'N 59°22'E, 17.V.1978, ONHM 610.22 (3); Wadi
Manqal, effluent of Wadi Hilm (Qalhat), near Kibdiyah (= Kebdah), 22°41'N 59°18'E, 26.IV.1978, ONHM 610.27 (40); Wadi
al-Batha S of Bilad Bani bu Ali, 22°00'N 59°20'E, 27.IV.1978, ONHM 610.19 (4); Wadi Hilm (Bani Ghabir) at Shofan, in
irrigation basin, 26.II.1985, ONHM 610.15 (10); Sayq, lower Wadi Bani Khalid, 23°30'N 59°07'E, 17.IV.1978, ONHM 610.18
(24); Wadi Qalhat/Hilm, lowest pool, 22°41'N 59°22'E, 18.IV.1978, ONHM 610.16 (5); same data, ONHM 610.23 (44).
Description: The left-coiled discoidal shell is greenish in fresh specimens. It reaches a
diameter of up to 20 mm and a height of nearly 10 mm. The spire and the umbilical areas are
deeply excavated. The teleoconch whorls are bluntly shouldered and somewhat inflated. Their
sculpture consists of coarse and regularly spaced ribs. The aperture is obliquely crescent-shaped.
Usually, the peristome is sharp but a thickened callose lip may be present in the interior of the
aperture.
Remarks: I. exustus is widespread in India and adjacent tropical areas of Asia. In addition to
southern Arabia, it also occurs on Socotra Island and is known to have been introduced by human
activities to a few localities in Africa (BROWN 1994: 208).
Family Physidae
Genus Physella Haldeman, 1843
1843 Physella Haldeman. — Monograph of the freshwater univalve Mollusca of the United States 6: 14.
Diagnosis: Medium-sized left-coiled shell (compare to Bulinus). The surface of the shell is
smooth. The mantle digitations only occur on the parietal side of the mantle.
Type species: Physa globosa Haldeman, 1841.
Physella acuta (Draparnaud, 1805)
1805 Physa acuta Draparnaud. — Hist. nat. Moll. terr. fluv. France: 55, pl. III, figs 10-11 (France, Garonne).
Material: Saudi Arabia: Northern Hijaz, Khaibar Oasis, 750 m, 25°44'N 39°17'E, 28.III.1990, W. Schneider & F.
Krupp, SNMNH-MO 37/5, SMF 311563/3 (preserved), SMF 311564/3. Eastern Province, al-Qatif Oasis, palm plantations W
of centre, irrigation channels, 14.VI.1992, R. Kinzelbach, SNMNH-MO 34/110, SMF 311560/30; Tarut, plantations N of
Tarut centre, 14.VI.1992, R. Kinzelbach, SNMNH-MO 35/51, SMF 311561/15, KINZ (5); al-Qatif Oasis, freshwater course
W of town, 26°36'N 49°58'E, 31.I.1992, R. Kinzelbach & I.A. Nader, SNMNH-MO 36/18, SMF 311562/15. — Yemen:
Dhi Bin N of Sana’a, from a cistern, 6.III.1990, M. Scholz, NEUB (2 preserved).

360 E. NEUBERT
Description: The small shell measures 15 mm in height and up to 9 mm in diameter. The
spire is elevated with evenly rounded whorls. The protoconch whorls are narrowly coiled and acute
compared to Bulinus species. The suture has a medium depth. The aperture is broadly excavated with
a sharp peristome. Often, there is a faint labial callus in the aperture, particularly in the basal part.
Remarks: This species is known from a few localities in the Arabian Peninsula (spring at
Khobar, Medina Province; small canals near Riyadh; cf. BROWN & WRIGHT 1980: 353). It is easily
dispersed by human activity. The author agrees with VAN DAMME (1984: 52) that Physella acuta is a
neozoon which was introduced to Europe long ago, maybe soon after the colonisation of the New
World, and not from Europe to northern America as indicated by BURCH (1989: 188). Almost all
species of the Physidae live in temperate areas of northern America. The fossil record during the
Tertiary is very vague because of confusion of the shells with species of the Bulininae. During the
Pliocene, the records are very scarce. Moreover, it is difficult to draw a line between Aplexa (which
seems to be holarctic, at least from fossil records) and Physa s. lat. based on shell morphological
grounds only. Obviously, the species started its colonisation in the western part of the continent,
spreading extremely fast over large areas of the Old World.
Family Ancylidae
Genus Ancylus O.F. Müller, 1774
1774 Ancylus O.F. Müller. — Vermium terrestrium et fluviatilium 2: 199.
Diagnosis: Small limpet-like shells with a blunt apex. The apex and often the shell show
coarse radial ridges.
Type species: Ancylus fluviatilis O.F. Müller, 1774.
Ancylus fluviatilis O.F. Müller, 1774
1774 Ancylus fluviatilis O.F. Müller. — Vermium terrestrium et fluviatilium 2: 201 (“In ripa sabulosa fluvii Ilm Saxoniae”).
1874 Ancylus compressus Jickeli. — Verh. Kaiserl. Leop.-Carol. Dtsch. Akad. Naturf. 37: 223, Taf. VII, Fig. 26 (“Hamaszen
bei Mekerka im Toqor”).
1874 Ancylus abyssinicus Jickeli. — Verh. Kaiserl. Leop.-Carol. Dtsch. Akad. Naturf. 37: 223, Taf. III, Fig. 5-6, Taf. VII,
Fig. 27-28 (“Hamaszen auf dem Wege von Genda nach Asmara, bei Mekarka im Toquor, in Zuflüssen des Anseba”
[= Eritrea]).
1941 Pseudancylus argenteus Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 37, pl. 3, figs 11-12
(Hadramaut: pools in Shab Samu’a, Hureidha).
Material: Saudi Arabia: Asir mountain region, Raydah escarpment, 1800 m, 29.I.1994, P. Symens & M. Werner,
SMF 311554/1 (preserved); Raydah escarpment, al-Aqaba spring, 2310 m, 18°11'50.0"N 42°24'30.2"E, 5.V.1994, W.
Schneider & F. Krupp, SNMNH-MO 31/8, SMF 311555/3 (preserved), SMF 311556/3, HLMD (2); Wadi Bani Mazin,
18°14'N 42°26'E, 14.III.1990, F. Krupp & W. Schneider, SNMNH-MO 32/16, SMF 311557/6 (preserved), SMF 311558/13;
Wadi al-Fajir, 18°14'N 42°13'E, 15.III.1990, F. Krupp & W. Schneider, SNMNH-MO 33/5, SMF 311559/5, HLMD (5). —
Yemen: Dhi Bin, N of Sana’a, from a cistern, 6.III.1990, M. Scholz, NEUB (1, preserved).
Description: The thin-walled limpet-like shell is oval to subcircular in its basal outline.
The protoconch is small and bluntly impressed. The apical part of the shell is always slightly
inclined to the right and is close to the end of the shell but never exceeds it. There are numerous
coarse riblets running from the apex to the rim of the shell. When crossed by growth-lines, a
reticulate pattern may be observed on the surface.
Remarks: The synonymy of Ancylus fluviatilis is very complex and has been compiled by
HUBENDICK (1970: 10-20). To avoid confusion here, only those names which were used for
specimens of A. fluviatilis from Arabia or originate from neighbouring areas are cited. HUBENDICK’s
map (1970: fig. 84) shows the distribution of Ancylus fluviatilis; only the northern area of Turkey
should be added.

Terrestrial and freshwater molluscs of the Arabian Peninsula 361
Fig. 37: Male genital
organs of Laevicaulis
alte, specimen from
Jubail, Haii al-Bahr,
research station.
Family Veronicellidae
Genus Laevicaulis Simroth, 1913
1913 Laevicaulis Simroth. — In: Voeltzkow, A.: Reise in Ostafrika in den Jahren 1903-1905 3: 147.
Diagnosis: A terrestrial slug with the upper body surface (= notum) covered by minute
warts. The sole is distinctly differentiated from the neighbouring hyponotum and characterised by
minute transverse lamellae. The female genital opening is situated in the posterior half of the
hyponotum. The digitiform glands of the penial gland open directly into the papilla. Anteriorly,
the intestines are delimited by a fold of the intestinum (FORCART 1953).
Type species: Vaginula comorensis Fischer, 1883.
Laevicaulis alte (Férussac, 1823) Fig. 37
1823 Vaginulus alte Férussac. — Tabl. syst. Limaces: 14, pl. VIIIA, fig. 8, pl. VIIIB, fig. 5 (“Les environs de Pondichéry”
[= India, Pondicherry S of Madras]).
Material: Saudi Arabia: Hijaz, Jeddah, 21°29'N 39°11'E, 29.X.1983, J. Grainger, NHMB (1); same locality,
7.XI.1984, W. Büttiker, NHMB (1); same locality, 6.VI.1985, W. Büttiker, NHMB (4); same locality, 31.III.1985, J. Grainger,
NHMB (1); Hakimah, 85 m, 15.X.1985, W. Büttiker, NHMB (1); Riyadh, park of Central Hospital, 17.X.1990, H. Tayeb,
SNMNH-MO 42/3, SMF 311570/2; Dammam, gardens and parks in the city, 30.IX.1989, H. Tayeb, SMF 311571/1; Jubail,
Haii al-Bahr, research station, 27.XI.1993, M. Almarri, SNMNH-MO 42/3, SMF 311572/2. — Om a n : Hawiyah, pool,
280 m, 22°23'N 58°51'E, 15.I.1986, J. Gallagher, NHMB (1).
Description: The slugs may reach up to 50 mm in length in preserved specimens (Saudi
Arabia), but FORCART (1953: 64) also reports a body length of 80 mm in specimens from Africa.
In fresh specimens, the body is dark grey to black with a longitudinal bright stripe.
Anatomical details: One of the specimens collected in Jubail was dissected (Fig. 37). The
organs match exactly the descriptions given by GRIMPE & HOFFMANN (1925). The pedal gland is
coiled several times and opens with a funnel-shaped tube below the buccal mass. The male genital
system opens on the left side above the snout. Its distal parts consist of a heavily coiled VD which
is sheathed by the MRP and projects into the copulatory organ. This consists of a massive
muscular sheath and a slender “papilla”. The VD enters this “papilla” centrally, its duct opens on

362 E. NEUBERT
top of the organ. Basally, a ring-shaped structure can be observed. Parallel to the penial sheath, an
accessory organ is found consisting of a bundle of glandular tubes with an annulated surface and a
sheath enveloping the common duct of this organ which is also formed like a papilla.
Remarks: This species is widespread in the Indopacific area due to human activities. Its
natural origin is doubtful, but FORCART (1953: 68) states “From the fact, that the genus Laevicaulis
has its main development in East- and Central-Africa, may be concluded that Laevicaulis alte was
primarily an African species”. The records in Arabia are all from areas in the vicinity of humans
(gardens). It was never found in natural habitats in humid mountainous areas, which supports
Forcart’s view of dispersal by humans outside tropical Africa.
Family Vertiginidae
Genus Gastrocopta Wollaston, 1878
1878 Gastrocopta Wollaston. — Testacea Atlantica: 515.
Diagnosis: Minute oval to oval-conical shells with a rimate umbilicus. Angularis and
parietalis are connected to form a sometimes bifid lamella, columellar and palatal lamellae are
present.
Type species: Pupa acarus Benson, 1856.
Gastrocopta antinorii (Paladilhe, 1872) Figs 38, 52
1872 Pupa antinorii Paladilhe. — Ann. Mus. Stor. Nat. Genova 3: 21, tav. 1, figs 11-12 (“Environs d’Aden”).
Type material: not available.
Original description: “Testa late ac profunde umbilicata, cylindracea, vix subovata,
angustissime striatula, parum nitida, albido-lutescens; spira ad apicem minutum, obtusulum,
convexo-conica; anfractibus 7.5 parum convexis, rapide, a tertio praesertim, accrescentibus, sutura
parum profunda separatis; ultimo ad aperturam valde ascendente, postice biscrobiculato, ad
aperturam substrangulato, umbilicum carina evanescente circumdante; margine libero convexo-
subsinuoso. Apertura subovato, superne latiore; pariete aperturali plica elongatula dentiformi,
medium versus, munito; peristomate disjuncto, expanso, reflexo; margine externo arcuatulo, plica
minuta, extremitati liberae dentis parietis aperturali sat approximata, instructo; plica palatali 1,
scrobiculum leve extus formante; plica columellari sat immersa; columella vix subarcuata, margini-
bus, ob incurvationem superam labri, valde approximatis”. — This translates to: The shell is
broadly and deeply umbilicate, cylindrical, subovate, very finely striate, not glossy, of a yellowish
white colour; the spire is convex subconical towards the minute apex; 7.5 subconvex whorls
enlarging very fast until the third whorl, separated by a shallow suture; the last one ascends
strongly towards the aperture and is somewhat constricted with two minute furrows backwards,
the umbilicus surrounded by a faint keel; the free margin is somewhat convex sinusoid. The
aperture is subovate and broadened at the top; on the parietal side of the aperture, an elongate
tooth-like lamella is present; the peristome is disconnected, inflated and reflected; the outer margin
is somewhat curved and shows an elongate callosity reaching its maximum extension opposite the
parietal tooth; there is one palatal lamella, forming a shallow furrow outside; a columellar tooth of
medium strength, the columella only somewhat arcuate, its margin somewhat thickened at the top.
PALADILHE (loc. cit.) gives the measurements of this shell (Fig. 38) as H = 4 mm, D = 2 mm.
Remarks: It is difficult to comment on this species as it has never been found again and
thus its generic position is provisional. To the author’s knowledge, there is no species recorded from
the African coastal mountains opposite Arabia which might be conspecific (Fig. 52).

Terrestrial and freshwater molluscs of the Arabian Peninsula 363
Gastrocopta klunzingeri (Jickeli, 1873) Figs 39, 52
1873 Pupa klunzingeri Jickeli. — Malakozool. Bl. 20: 106 (“Abyssinien, Prov. Hamaszen”, “auf dem Weg von Genda nach
Asmara, nicht selten auf der Hochebene Rara-Beit-Andu und Bagos auf dem Berge von Keren” [= Eritrea]).
1874 Pupa klunzingeri. — Jickeli; Verh. Kaiserl. Leop.-Carol. Dtsch. Akad. Naturf. 37: 116, Taf. V, Fig. 8.
Material: Yemen: Red Sea, ad-Durayhimi, 14°40'N 42°59'E, sand beach, 16.IV.1993, H. Dekker & F. Ceuninck v.
Capelle, DEKK (2).
Type material: Syntypes SMF 4725/3, coll. O. Boettger ex Jickeli labelled “Hamaszen,
Rora Beit Andu”. A second lot from the same locality, SMF 10641/2, coll. Reinhardt ex Jickeli.
Description: The minute shell is turreted and brown in fresh specimens. The blunt and
considerably thick protoconch consists of two whorls which are sculptured by fine warts.
The teleoconch whorls are rounded to bluntly shouldered subsuturally with a deep suture.
The surface of the shell is sculptured by coarse and oblique sharp ribs. Interaxially, the shell is
finely granulated.
The aperture is rounded to subquadrate. The lip is flared but not recurved and very fragile at
the rim. The dentition consists of five teeth. The angularis is very prominent and somewhat
recurved, producing an oval sinulus. It is connected to the parietalis by an oblique lamella. The
parietalis is truncate in the interior of the shell. The columellaris is a broad perpendicular plate.
Basally, there is only a minute basalis left reaching the same size as the suprapalatalis. The palatalis
itself is a strong elongate lamella reaching far into the interior of the aperture. In some specimens
it is visible by a shallow furrow on the last whorl.
The peristomial rims are connected to each other by a weak callose shield which may spread
over the lower part of the preceding whorl.
The last whorl is somewhat compressed at the base. The umbilicus is open and surrounded by
a funnel-shaped periomphalum.
Measurements (illustrated syntype ex Jickeli SMF 4725, Fig. 39): H = 2.35; D = 1.20;
W = 5.
Remarks: This species was hitherto known only from the Ethiopian region and is recorded
here for the first time from the Arabian Peninsula. A syntype specimen ex Jickeli (SMF 4725) from
“Hamaszen, Rora Beit Andu”, Eritrea, is shown in Fig. 39.
Genus Boysia L. Pfeiffer, 1849
1849 Boysia L. Pfeiffer. — Z. Malakozool. 6: 105.
Diagnosis: As for the species.
Type species: Tomogeres boysii L. Pfeiffer, 1846.
Boysia boysii (L. Pfeiffer, 1846) Figs 40, 52
1846 Tomogeres boysii L. Pfeiffer. — Symbolae ad Historiam Heliceorum III: 82 (“Bengalia”).
Material: Saudi Arabia: Asir mountain region, Abha escarpment, 2500 m, I.1994, P. Symens & M. Werner,
SNMNH-MO 44/2, SMF 311573/1; Asir mountain region, Raydah escarpment, 2350 m, 29.I.1994, P. Symens & M. Werner,
SMF 311574/1 (protoconch).
Description: The minute shell is corneous brown and has a conical spire. The protoconch
is smooth. The whorls of the teleoconch are bordered by a deep suture. The surface of the
teleoconch displays very fine and dense axial striations. The last whorl is bent upwards and ascends
to reach at least half the height of the spire.
The peristome has the form of an oblique triangle. Its rim is reflected and slightly callose
internally. It is not connected to the teleoconch. The umbilicus is open and formed like a small

364 E. NEUBERT
elongated slit. In a creeping animal, the spire presumably points in the direction of movement.
This is in contrast to species with a typically situated aperture, where the spire always points in the
opposite direction.
Measurements (illustrated specimen, Fig. 40): H = 3.20; D = 3.20; PH = 1.30; PD = 1.40;
W = 6.
Remarks: This species was originally described from India. GUDE (1914) reports on five
localities in north-western India where Boysia boysii is found. ZILCH (1960: 147, Abb. 494)
illustrates a specimen from the Nerbudda valley. The specimens from the Arabian Peninsula were
compared to this specimen (SMF 51683). No conchological differences were found, so the
specimens from Saudi Arabia are assigned to this species (Fig. 52).
Family Pupillidae
Genus Pupoides L. Pfeiffer, 1854
1854 Pupoides L. Pfeiffer. — Malakozool. Bl. 1: 192.
Diagnosis: Minute turreted oval-elongate shells. The protoconch is blunt, the teleoconch
almost smooth. The ovate aperture with a small to medium-sized angularis, the peristomial rims
reflected, a labial callus is often present.
Type species: Bulimus nitidulus L. Pfeiffer, 1848.
Pupoides coenopictus (Hutton, 1834) Figs 41-48, 53
1834 Pupa coenopicta Hutton. — J. Asiatic Soc. Bengal 3: 93.
1872 Bulimus samavaensis Mousson in Paladilhe. — Ann. Mus. Stor. Nat. Genova 3: 14, tav. 1, figs 20-21 (“Samava sur
les bords de l’Euphrate”).
1872 Bulimus vermiformis Paladilhe. — Ann. Mus. Stor. Nat. Genova 3: 15, tav. 1, figs 24-25 (“Environs d’Aden”).
1872 Bulimus cerealis Paladilhe. — Ann. Mus. Stor. Nat. Genova 3: 16, tav. 1, figs 22-23 (“Environs d’Aden”).
1875 Buliminus fabianus Gredler. — Nachrbl. dtsch. malakozool. Ges. 7: 87 (“Aus dem Land der Schilluck-Neger”).
1876 Bulimus mahariscus Bourguignat. — Species novissimae molluscorum in Europaeo systemati detectae, notis diag-
nosticis succinctis breviter descriptae. Premiére centurie: 21 (“Kursi près d’Aden”).
1876 Bulimus euphraticus Bourguignat. — Species novissimae molluscorum in Europaeo systemati detectae, notis diag-
nosticis succinctis breviter descriptae. Premiére centurie: 22 (“Sur les bords de l’Euphrate aux environs de Samava”).
1876 Bulimus marebiensis Bourguignat. — Species novissimae molluscorum in Europaeo systemati detectae, notis
diagnosticis succinctis breviter descriptae. Premiére centurie: 23 (“Mareb, près des ruines de l’antique Saba”).
1876 Bulimus kursiensis Bourguignat. — Species novissimae molluscorum in Europaeo systemati detectae, notis diag-
nosticis succinctis breviter descriptae. Premiére centurie: 23 (“Kursi près d’Aden”).
1889 Bulimus ragius Jousseaume. — Bull. Soc. Malac. France 6: 347 (“aux environs d’Aden […] aux alentours de
Massaouah”).
Material: Saudi Arabia: Eastern Province, beach debris at Haii al-Bahr, N of Jubail, I.1992, E. Neubert, SNMNH-
MO 45/3, SMF 311575/2; Eastern Province, Jubail, gardens at Dauhat ad-Dafi, 7.I.1994, P. Symens, SNMNH-MO 46/8,
SMF 311576/7, NEUB (7) (all preserved); Eastern Province, Phragmites vegetation on a pond at the highway Khobar-Jubail,
3.4 rkm S of exit to Juaymah, 25.V.1995, E. Neubert & R. Janssen (2). — Yemen, sub B. samavaensis: Aden, SMF 51510/2,
coll. Kobelt ex Rolle; same locality, SMF 247062/3, coll. Bosch ex Rolle; same locality, SMF 247061/3, coll. Bosch ex Rolle;
same locality, SMF 51509/2, coll. O. Boettger ex Schweinfurth; same locality, SMF 200569/9, coll. Jaeckel. — sub B.
vermiformis: Aden, SMF 51511/3, coll. Kobelt ex Rolle. — sub B. mahariscus: Aden, SMF 51507/2, coll. O. Boettger ex
Ponsonby. — sub B. cerealis: Aden, SMF 51546/3, coll. Kobelt ex Rolle; Hadramaut, Wadi bin Ali, SMF 50275 ex v. Wissmann;
Hadramaut, Wadi Djirdan, SMF 50273 ex v. Wissmann; Hadramaut, Wadi Amd close to Shamikh at Horeida, SMF 50274 ex
v. Wissmann; Midi, Marsa Baqlah, 16°21'N 42°47'E, beach, mangroves, 24.IV.1993, H. Dekker & F. Ceuninck v. Capelle,
DEKK (3); Midi, Oreste Point, 16°22'N 42°46'E, sand beach, rocks, 24.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK
(2); Red Sea, al-Murk Island, 15°38'N 42°38'E, beach, 11-12.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (2); Red
Sea, ad-Durayhimi, 14°40'N 42°59'E, sand beach, 16.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (9); al-Mahrah,
Hawf, E side of village, 16°38'03"N 53°02'34"E, rocks with algae and sand patches, 1.X.1995, H. Dekker & F. Ceuninck v.

Terrestrial and freshwater molluscs of the Arabian Peninsula 365
Figs 38-51: (original size × 10), all in frontal view. 38: Gastrocopta antinorii ex PALADILHE (1872: tav. 1, fig. 12). 39:
Gastrocopta klunzingeri, syntype SMF 4725. 40: Boysia boysii, specimen from Asir mountain region, Abha escarpment. 41-48:
Pupoides coenopictus. 41: Bulimus vermiformis ex PALADILHE (1872: tav 1, fig. 25). 42: Bulimus cerealis ex PALADILHE (1872:
tav. 1, fig. 22). 43: Syntype of Buliminus fabianus, SMF 4740. 44: Syntype of Bulimus mahariscus, MHNG. 45: Syntype of
Bulimus euphraticus, MHNG. 46: Holotype of Bulimus marebiensis, MHNG. 47: Holotype of Bulimus kursiensis, MHNG.
48: Syntype of Bulimus ragius, MNHNP. 49-50: Lauria cylindracea. 49: Specimen from Asir mountain region, Abha
escarpment (SMF 311578). 50: Syntype of Pupa bruguierei, SMF 51781 a. 51: Granopupa granum, specimen from Asir
mountain region, W of al-Atawah.

366 E. NEUBERT
Capelle, DEKK (13). — O m a n : Jabal Akhdar, Kahf Hoti (Hoti carstic cave system), 990 m, 23°06'14"N 57°21'55"E (= N of
Tanuf, E of al-Hambra) (many fragments).
Type material: Syntypes Buliminus fabianus, SMF 4740/8 ex coll. O. Boettger ex Gredler
ex L. Pfeiffer; 6 syntypes Bulimus mahariscus, MHNG; 2 syntypes Bulimus euphraticus, MHNG;
holotype Bulimus marebiensis, MHNG; holotype Bulimus kursiensis, MHNG; syntype Bulimus
ragius, MNHNP.
Description: The small brown shells are conical, elongated. The smooth and blunt
protoconch consists of 1.5 whorls.
The teleoconch whorls are rounded with a suture of medium depth. They are covered by fine
axial growth lines. In some specimens stronger growth lines may occur, but they are irregular and
may represent growth stops. Microscopically, a dense punctuation can be detected on the shell’s
surface (magnification > 40 ×).
The aperture is semi-circular. The lip is marked by a thin callus, the peristomial rims are flared
and sometimes reflected. A weak to strong angularis is always present. The umbilicus is elongate
and open.
Remarks: The list of synonyms only contains names which were given to specimens from
the Arabian Peninsula and some adjacent areas. For a more complete compilation of synonyms of
this widespread species refer to SEDDON (1992). Pupoides coenopictus is also known from the
Dahlak archipelago in the Red Sea as well as from Sudan, Eritrea, Djibouti and adjacent areas of
Ethiopia and from several localities in Oman (Fig. 53).
Genus Lauria Gray in Turton, 1840
1840 Lauria Gray in Turton. — A manual of the land and freshwater shells of the British Islands 2: 193.
Diagnosis: Small oval to subcylindrical, turreted shells, teleoconch with faint to strong axial
ribs. The aperture often with an angularis, parietalis and columellaris, these lamellae may be
reduced.
Type species: Turbo cylindraceus Da Costa, 1778.
Lauria cylindracea (Da Costa, 1778) Figs 49-50, 52
1778 Turbo cylindraceus Da Costa. — Test. Brit.: 89, pl. V, fig. 16.
Material: Saudi Arabia: Asir mountain region, Abha escarpment, Juniperus forest, 2500 m, II.1994, P. Symens &
M. Werner, SNMNH-MO 48/5, SMF 311578/5; same data but 2300 m, SMF 311579/1; Asir mountain region, Raydah
escarpment, 2350 m, 29.I.1994, P. Symens & M. Werner, SNMNH-MO 49/2.
Type material: Syntypes Pupa bruguièrei, SMF 517812 a and b, southern Ethiopia, Habab
mountains Enjelal and Bagla, c. 7995 ft, coll. Reinhardt ex Jickeli; SMF 4624/1, same locality, coll.
O. Boettger ex Jickeli; SMF 62025/2, same locality, coll. Kobelt ex Jickeli.
Description: The small brown shell is oval in outline. The smooth protoconch consists of
two whorls and is only slightly rounded. The teleoconch whorls are covered by fine, dense and
obliquely arranged axial riblets. The suture is shallow.
The aperture is subquadrate with a reflected peristome strengthened by a small white lip. The
umbilicus is open. The dentition consists of a weak columellaris and an almost strong angularis
which continues into the interior of the shell. It can also be found in juvenile specimens which also
display several small perpendicular palatal callosities in their shells (cf. KERNEY et al. 1983: 121).
Measurements (illustrated specimen, Fig. 49): H = 4.30; D = 2.00; W = 8.
Remarks: This species is widespread in Europe. It lives in temperate semi-humid environ-
ments. Thus, it may be expected to occur in high altitude mountains exposed to upwelling

Terrestrial and freshwater molluscs of the Arabian Peninsula 367
Fig. 52: Distribution of several Pupilloidea and one Succineidae species on the Arabian Peninsula.
clouds in the Arabian Peninsula. This is the first record for this species from the Arabian
Peninsula (Fig. 52).
VAN BRUGGEN (1991) discusses the distribution of Lauria cylindracea and draws attention to
PILSBRY’s note (1922: 61) on the close relationship of Pupa bruguierei Jickeli, 1874, to L. cylindra-
cea. In the author’s opinion, this taxon indeed represents the latter species, but a final decision
should be postponed until a revision of all taxa of Lauria is conducted. As the SMF houses
syntypic material of P. bruguierei (cf. material), a well-preserved specimen (SMF 517812 a) is also
illustrated here (Fig. 50). Compared to the Arabian specimens, P. bruguierei is somewhat smaller,
its whorls are more compressed and the suture is deeper. Bearing in mind the variation of shell
characters in L. cylindracea it seems doubtful whether these differences are sufficient to define a
separate species.
Family Chondrinidae
Genus Granopupa O. Boettger, 1889
1889 Granopupa O. Boettger. — Jahrb. Nassau. Ver. Naturk. Wiesbaden 42: 249.
Diagnosis: Small cylindrical shells with up to nine whorls reaching a length of approxi-
mately 6-7 mm. The whorls are covered with fine evenly spaced ribs. There are four palatal, one
parietal and two columellar lamellae. In the male genital system, a penial caecum is missing.
Internally, there are coarse papillae in the distal epiphallus lumen present (GITTENBERGER 1973:
36-41, textfigs 6, 7 a-b, 8).
Type species: Pupa granum Draparnaud, 1801.

368 E. NEUBERT
Granopupa granum (Draparnaud, 1801) Fig. 51-52
1801 Pupa granum Draparnaud. — Tab. Moll. France: 59.
Material: Saudi Arabia: Asir mountain region, at the road from al-Baha to Taif, 2.8 rkm W of al-Atawah,
3.VI.1995, E. Neubert et al., SNMNH-MO 50/6, SMF 311580/5, SMF 311581/1 (preserved).
Description: Since this species is the smallest member of the Chondrinidae, it is not
usually confused with other species of the family. The following additional characters are given: in
the peristome, the palatalis inferior is the strongest lamella. The columellaris is often stronger than
the infracolumellaris.
Remarks: This species is widespread in the western Palaearctic region (Fig. 52).
Genus Granaria Held, 1837
1837 Granaria Held. — Isis (Oken) 1837 (12): 918.
Diagnosis: Small to medium-sized cylindrical shells. There are four palatal, one parietal and
two columellar lamellae. A penial caecum is present, the MRP inserts in a medium position on the
epiphallus.
Type species: Pupa frumentum Draparnaud, 1801.
Granaria arabica (Dohrn, 1859) Fig. 52
1859 Pupa arabica Dohrn. — Malakozool. Bl. 6: 203 (Arabia).
Description (cf. GITTENBERGER 1973: 41): The shell is cylindrical fusiform and is formed
by 6.5-7 whorls. The subconvex whorls are irregularly striped. The peristome is only weakly
thickened, the peristomial rims are not interconnected. The parietal wall displays a strong
parietalis, the angularis is weak. There are two short columellar lamellae and three palatal lamellae
with a very strong palatalis inferior.
Measurements: H = 5.1-6.1, D = 2.3-2.4.
Remarks: The description by GITTENBERGER (loc. cit.) is based on material collected by H.
Scott in 1938 at the pass over Jabal Sumara in Yemen between Ibb and Yarim. There are no
additional records known for this species (Fig. 52).
Family Punctidae
Genus Toltecia Pilsbry, 1926
1926 Toltecia Pilsbry. — Proc. Acad. Nat. Sci. Philadelphia 78: 116.
Diagnosis: Minute discoidal shell with a subdepressed to somewhat elevated spire. Teleo-
conch whorls with widely spaced riblets and a very fine impressed spiral sculpture.
Type species: Thysanophora (Toltecia) jaliscoensis Pilsbry, 1926.
Toltecia pusilla (Lowe, 1831) Figs 53-62
1831 Helix (Helicella) pusilla Lowe. — Trans. Cambr. Phil. Soc. 4 (1): 46, pl. 5, fig. 17.
1865 Helix (Patula) cryophila v. Martens. — Malakozool. Bl. 12: 182 (Ethiopia, Simien mountains near Bayeta).
1881 Patula lederi O. Boettger. — Jahrb. dtsch. malakozool. Ges. 7: 380 (Lenkoran).
1905 Punctum lederi var. meridionalis O. Boettger. — Nachrbl. dtsch. malakozool. Ges. 37: 102 (“Sarus Genist bei Adana”
[= Turkey, debris of the Sarus river close to Adana]).
Material: Saudi Arabia: Asir mountain region, King Khalid descent near Baha, 2215 m, 3.VI.1995, E. Neubert et
al., SNMNH-MO 51/3, SMF 311582/2. — sub H. cryophilum: Eritrea, Mekarka Prov. Hamaszen, SMF 197475/1, coll. O.
Boettger ex Jickeli.
Type material: Lectotype Patula lederi, SMF 225643, coll. O. Boettger; paralectotypes
SMF 225644/2, coll. O. Boettger; paralectotypes SMF 225645/2, coll. v. Moellendorff. Lectotype

Terrestrial and freshwater molluscs of the Arabian Peninsula 369
Fig. 53: Distribution of Pupoides coenopictus and Toltecia pusilla on the Arabian Peninsula.
Punctum lederi var. meridionalis, SMF 225174/1, coll. O. Boettger ex Nägele; paralectotypes
SMF 225175/6, coll. O. Boettger ex Nägele; paralectotypes SMF 120361/8, coll. C.R. Boettger
ex O. Boettger ex Nägele; paralectotypes SMF 225176/2, ex Nägele. The lectotypes of both, Patula
lederi and Punctum lederi var. meridionalis, were designated by Zilch (unpublished) and are
validated here.
Description: The minute shells are brown, depressed conical with the spire only slightly
elevated. The protoconch is smooth apart from some very faint spiral ridges. The whorls are
subcircular and form a deep suture. The body whorl is not completely rounded but somewhat
angular in cross-section. It descends under the preceding whorl; the peristome is circular, its upper
rims are not connected. The umbilicus is wide and measures approximately one third of the major
diameter.
The shell surface displays two kinds of sculptural elements. The first are the wide-spaced ribs
which are thin and sharply pointed in juvenile or fresh shells and may be eroded in dead shells.
The second element is the fine spiral sculpture which consists of densely packed and finely beaded
threads. Both elements evenly cover surface and subsurface of the shell and are easily visible in the
umbilicus.

370 E. NEUBERT
Measurements: Illustrated specimen (Figs 54-56): H = 1.25; D = 1.90; W = c. 4. Lectotype
Punctum lederi H = 1.4; D = 2.5. Lectotype Punctum lederi var. meridionalis H = 1.3; D = 2.15.
Remarks: VERDCOURT (1991: 354) reports that the holotype of Helix cryophila v. Martens,
1865, in the ZMHB is heavily damaged. Nevertheless, he found traces of spiral sculpture on the
base of the type specimen. The figures given by JICKELI (1874: Taf. IV, Fig. 17 a-d) are very similar
to the specimen kept in the SMF except that the ribs are more pronounced in the figures. The
SMF specimen is an adult and, as Jickeli points out, the ribs may become eroded in adult shells.
The taxa Patula lederi and Punctum lederi var. meridionalis are so similar to Helix cryophilum
that it is impossible to keep them separate. The sculpture of widely spaced ribs with the interaxial
spiral threads is very characteristic of this species. Both shell measurements and width of the
umbilicus are similar. The lectotype of P. lederi has a somewhat more elevated spire, but the
paratype lots contain specimens which are nearly identical with those from the Asir mountains.
Patula lederi was already mentioned by GITTENBERGER et al. (1980: 12) as a possible synonym
taxon of Toltecia pusilla. The identity of many similar minute endodontid taxa is insufficiently
known. It is possible that there is only one species involved, with a nearly global distribution,
which then for reasons of priority has to bear the name of the New Zealand taxon Paralaoma
caputspinulae (Reeve, 1852) (Gittenberger, pers. comm. 1996). These relationships are not proven
exactly and are far beyond the scope of this publication. Therefore, Lowe’s name for this taxon is
used here until nomenclature is settled.
Family Succineidae
Genus Quickia Odhner, 1950
1950 Quickia Odhner. — Proc. Malacozool. Soc. 28: 206.
Diagnosis: Small succineid shell with deep sutures. The spire is short and mammillate. The
penial sheath is absent. Usually, the penial retractor passes underneath the right optic tentacle. The
epiphallus is strongly reduced or missing, the penis a simple tube. Quickia s. str. differs from its
subgenus Burchella Patterson, 1970 by having a tissue sheet covering the distal penis (combined
after ODHNER 1950 and PATTERSON 1971).
Type species: Succinea concisa Morelet, 1848.
Quickia (Quickia) concisa (Morelet, 1848) Figs 52, 63
1848 Succinea concisa Morelet. — Rev. Zool. (Soc. Cuv.) 11: 351.
Material: Saudi Arabia: Asir mountain region, Wadi Juwa, 150 m, 1.II.1994, P. Symens & M. Werner, SNMNH-
MO 83/1, SMF 311615/1.
Description: The small species reaches about 5 mm in length. The blunt and oblique
protoconch consists of about 1.5 whorls. Its surface is sculptured by minute pits (magnifica-
tion > 40 ×).
The suture of both teleoconch and protoconch is deep. The surface of the teleoconch whorls
displays a dull glistening and is sculptured by fine densely packed irregular riblets. The aperture is
oval acute and reaches approximately 1.5 times the length of the spire. The columella is straight
and diverging from the palatal peristome.
Remarks: This species lives in tropical Africa and is also known from several islands in
the Indian Ocean. In Arabia, it was first recorded by MORDAN (1988: 398) from Oman in the
Wahiba Sands region at Hawiyah near Mintrib, 22°23'N 58°51'E, under decaying banana shoots
in a date garden. The material from Wadi Juwa represents the first record of this species for Saudi
Arabia (Fig. 52).

Terrestrial and freshwater molluscs of the Arabian Peninsula 371
Figs 54-63: (original size × 10), 54-62: Toltecia pusilla. 54-56: Specimen from Asir mountain region, King Khalid descent near
Baha (SMF 311582). 54: Apical view. 55: Subsurface. 56: Detail of surface. 57-59: Lectotype of Patula lederi, SMF 225643. 57:
Frontal view. 58: Apical view. 59: Subsurface. 60-62: Lectotype of Punctum lederi var. meridionalis, SMF 225176. 60: Frontal
view. 61: Apical view. 62: Subsurface. 63: Quickia concisa, specimen from Asir mountain region, Wadi Juwa.
Family Subulinidae
No te : The generic placement of many species within the Subulinidae has not yet been deter-
mined exactly; many genera are ill defined and anatomical data are lacking for most of the taxa. A
phylogenetic revision of the Subulinidae based on conchological and anatomical details is urgently
needed. The definition of genera on conchological grounds only is doubtful.
As an example of the practice of adding names whilst disregarding the taxonomic problem
within the family, the description of the subfamily Glessulinae by Schileyko in SCHILEYKO &
KUZNETSOV (1996: 158-160) may be cited. The author states: “Current subdivision of the family
Subulinidae into three recent subfamilies (Subulininae, Rumininae and Obeliscinae) seems to be
artificial, because none of them has a clear differential diagnosis (compare diagnoses given by Zilch
1959).” This is a clear description of the present situation, but it does not prevent him from
introducing an additional subfamily. His taxon is described without a differential diagnosis, nor
are the relationships within the family discussed. His arguments are based on the dissection of one
species of Glessula v. Martens, 1860, i.e. Glessula serena (Benson, 1860), and refers to the figure of
the sexual organs of Glessula orophila (Benson, 1849) given by SEMPER (1870: pl. XII, fig. 15), a
figure which was reproduced by PILSBRY (1909: pl. 15, figs 2-3) and is regarded to be representative
for Glessula.
In the author’s opinion, this is a poor reason to introduce a subfamily containing the speciose
genus Glessula, even without having dissected the type species of Glessula (Achatina gemma Benson,
1849). Moreover, the structure called “flagellum” by Schileyko (SCHILEYKO & KUZNETSOV 1996:

372 E. NEUBERT
160, fig. 2 B; Glessula serena) is obviously an epiphallial caecum, as proved by its branching
position and the structure of the proximal epiphallus. The figure of SEMPER (loc. cit.) is much too
small to judge the true position of this “appendix” (which is elongately stalked). It is possibly a
true flagellum, as its branches are more proximal than in G. serena. This reinforces Schileyko’s
statement of the “artificial groups” within the Subulinidae.
More attention should be paid to the exact structure of the genital organs. Important charac-
ters within the Subulinidae seem to be presence or absence of an epiphallial papilla, flagellum,
epiphallial and/or penial caecum, penial muscular sheath and the attachment site of the MRP, and
others. In the last character, the origin of the retractor muscles may also play an important role.
Adherence at both the epiphallus and the penial caecum seems to be a plesiomorphous state, as it
can be observed in many taxa. Reduction of the epiphallus leads to a strengthening of the caecal
muscle which then acts as MRP (cf. H.B. BAKER 1927: 2 ff, 22 ff, pl. XX, figs 99-100 for Subulina
and Leptinaria).
Genus Zootecus Westerlund, 1887
1887 Zootecus Westerlund. — Fauna der in der paläarctischen Region lebenden Binnenconchylien III: 3.
Diagnosis: Cylindrical to pupoid shell with a conical apex. The teleoconch surface is
sculptured by wavy growth line. The aperture is considerably small with a thickened peristome.
Type species: Pupa insularis Ehrenberg, 1831.
Zootecus insularis (Ehrenberg, 1831) Figs 64-66
1831 Pupa insularis Ehrenberg. — Symbolae Physicae: unpaginated [13th page from beginning of chapter Animalia
Mollusca] (“Habitat in insula Cameran, quae prope Maris rubrum ostium australe inter Loheiam et Moccham iuxta
Arabiae felicis littus sita est” [= Yemen, Island Kameran N of al-Hodeida]).
1851 Pupa adenensis L. Pfeiffer. — Z. Malakozool. 8: 27 (“Ad rupes vulcanicas prope Aden Arabiae”).
1876 Buliminus ducoureti Bourguignat. — Species novissimae molluscorum in Europaeo systemati detectae, notis diag-
nosticis succinctis breviter descriptae. Premiére centurie: 20.
1943 Imparietula wissmanni Wenz. — Arch. Molluskenk. 75 (2/3): 150, textfig. (“Wadi Mahredun, Nord Djol, Hadra-
maut”).
Material: Saudi Arabia: Bukairat al-Hakimah, storage lake of Wadi Jizan, E of Jizan, 28.IV.1994, W. Schneider &
F. Krupp, SNMNH-MO 52/30, SMF 311583/30, HLMD (4); Asir mountain region, Wadi Jawah, SE of al-Ahrida, 29.IV.1994,
W. Schneider & F. Krupp, SNMNH-MO 53/2, SMF 311584/2; Asir mountain region, Wadi Juwa, 150 m, 1.II.1994, P. Symens
& M. Werner, SNMNH-MO 54/9, SMF 311585/9; Asir mountain region, Wadi Marabah, 1300 m, 29.I.1994, P. Symens &
M. Werner, SNMNH-MO 55/1, SMF 311586/1. — Ye m e n : Aden, SMF 157246 a, 157246/5, coll. Kobelt ex Schweinfurth;
same locality, SMF 157247/4, coll. K.L. Pfeiffer ex Mus. Berlin; same locality, SMF 296617/1, coll. Nägele ex v. Martens ex
Schweinfurth; same locality, SMF 211491/2, coll. Jaeckel; same locality, SMF 296627/3, coll. C.R. Boettger; Badjil, SMF
296621/6, coll. Reinhardt ex Schweinfurth; same locality, Febana lowlands, SMF 296622/7, coll Reinhardt ex Schweinfurth. —
sub Zootecus insularis adenensis: Aden, SMF 296631/2, coll. Bosch ex Rolle; same locality, SMF 296630/6, coll. v. Moellendorff;
Hadramaut, Wadi Djirdan, SMF 50264/4 ex v. Wissmann; Hadramaut, Wadi bin Ali, SMF 50269/7 ex v. Wissmann; Hadra-
maut, northern Djol, Dawadanu-Wadi Djari, SMF 50267/2 ex v. Wissmann; Hadramaut, Wadi Amd, loess wall at Shamikh,
close to Horeida, SMF 50266/5 ex v. Wissmann; Wadi Amd, loess wall at Amd, close to Hebre, SMF 50265/6 ex v. Wissmann;
southern Tihama, Chocha, 24.III.1988, A. Liebegott, LIEB (1); at the road from al-Hodeida to Menaha, foothills of the
escarpment, 800 m, 25.III.1988, A. Liebegott, LIEB (10); Hadramaut, Buraat, 6.IV.1988, A. Liebegott, LIEB (1); Red Sea, as-
Salif, 15°18'N 42°40'E, beach and rocks, 13.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (1); Red Sea, Ukban Island,
15°31'N 42°22'E, sand beach and rocks, 12.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (3); Red Sea, Midi, 16°20'N
42°47'E, mudflats, 24.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (> 50); Midi, Oreste Point, 16°22'N 42°46'E, sand
beach, rocks, 24.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (6); Red Sea, al-Murk Island, 15°38'N 42°38'E, beach,
11-12.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (32); Sharas, Wadi Sharas, NE of Hajjah, old river sediments,
800 m, 9.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (1); Red Sea, Abu Zahr, at al-Khawkhah Tourist Village,
13°51'N 43°14'E, sand beach, 19.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (2); Red Sea, al-Urj, 15°06'N 42°52'E,

Terrestrial and freshwater molluscs of the Arabian Peninsula 373
beach, mangroves, 15.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (1); Red Sea, ad-Durayhimi, 14°40'N 42°59'E,
sand beach, 16.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (6); al-Mahrah, Hawf, E side of village, 16°38'03"N
53°02'34"E, rocks with algae and sand patches, 1.X.1995, H. Dekker & F. Ceuninck v. Capelle, DEKK (14); al-Mahrah,
between Damquat and al-Fatk, 16°32'24"N 52°46'00"E, sand beach, patches of rock, 4.X.1995, H. Dekker & F. Ceuninck v.
Capelle, DEKK (1). — O m a n: Fanja, W of Muscat, I.1993, H. Pauscher, NEUB (37); Jabal Akhdar, oasis between ar-Rustaq
and al-Hazm, I.1993, H. Pauscher, NEUB (2); Oasis N of Nizwa, palm plantations, 13.III.1995, J. Wittmann, NEUB (13);
Oasis of Buraimi, I.1993, H. Pauscher, NEUB (2); Wahiba Sands, unknown locality, 19.IV.1978, A.B. Paltrinieri, ONHM 610.24
(20); Den/Dann, Jabal Khawr, 23°10'N 57°20'E, 21.III.1977, A.B. Paltrinieri, ONHM 610.7 (2); Wadi Manqal, effluent of
Wadi Hilm (Qalhat), near Kibdiyah (= Kebdah, 22°41'N 59°18'E, 26.IV.1978, A.B. Paltrinieri, ONHM 610.27 (3); Wadi al-
Batha S of Bilad Bani bu Ali, 22°00'N 59°20'E, 27.IV.1978, A.B. Paltrinieri, ONHM 610.19 (28); Sint, in Wadi, 23°07'N
57°05'E, 18.XII.1976, A.B. Paltrinieri, ONHM 610.20 (10); Wadi Bani Khalid, 23°30'N 59°08'E, 16.IV.1978, A.B. Paltrinieri,
ONHM 610.1 (1); Sayq, lower Wadi Bani Khalid, 23°30'N 59°07'E, 17.IV.1978, A.B. Paltrinieri, ONHM 610.18 (1); Wadi
Qalhat/Hilm, lowest pool, 22°41'N 59°22'E, 18.IV.1978, A.B. Paltrinieri, ONHM 610.16 (1); Wadi Qalhat/Hilm, 22°41'N
59°22'E, 18.IV.1978, A.B. Paltrinieri, ONHM 610.23 (9); Jabal Misfeh, various sites, 23°13'N 57°07'E, 14.III.1986, A.B.
Paltrinieri, ONHM 610.12 (1); Jabal Akhdar, Wadi al-Yemen, lower dam, 23°04'N 57°41'E, V.1980, A.B. Paltrinieri,
ONHM 610.9 (1); Wadi Selma (in Wadi Bani Awf = Salmah), 23°12'N 57°22'E, 2.V.1978, A.B. Paltrinieri, ONHM 610.50
(27, + juveniles). — The SMF houses numerous lots of Z. insularis from the Dahlak archipelago and Suakin (Sudan) mainly
originating from Rüppell and Jickeli.
Type material: Holotype Imparietula wissmanni, SMF 50242, paratypes SMF 50268/18.
WENZ (1943 b: 241) later identified I. wissmanni with Zootecus insularis adenensis. One specimen
ex coll. Reinhardt from the island of Kameran (SMF 296623/1) is present in the collection of the
Senckenberg Museum. The original label reads “Cylindrus insularis Ehrenberg, Insel Cameran”. As
Reinhardt exchanged material from the Museum in Berlin with v. Martens (often proven in coll.
SMF), it is likely that this specimen was taken from the original lot of Ehrenberg and thus it
probably represents a syntype and is illustrated here.
Description: The thick white shell is turreted with the apical whorls broad and conical in
shape. The teleoconch whorls are nearly cylindrical. The protoconch shows some irregular
wrinkles. The teleoconch whorls display very fine axial riblets which are often arched subsuturally.
Irregular spiral grooves may interrupt the axial sculpture.
The aperture is subquadrate with a thickened peristome. The umbilicus is open although
sometimes nearly covered by the reflection of the columellar rim of the peristome. In many
specimens, the peristomial rims are connected by a thick parietal callosity. Specimens from the
Arabian Peninsula may reach a height of up to 15 mm and a diameter of 5-6 mm.
Measurements (illustrated holotype Imparietula wissmanni, SMF 50242, Fig. 66): H = 12.0,
D = 3.7.
Remarks: This species is widespread in the Saharo-Sindian region. Consequently, a large
number of names was introduced from several places within its distribution. The synonymy given
here only refers to taxa within the Arabian region. A revision of the “species” usually assigned to
Zootecus is urgently needed.
On the Arabian Peninsula, Zootecus insularis seems to be restricted to the semi-arid habitats.
There are no records from the humid mountains in the south-west nor from desert areas in the
interior of the peninsula (Fig. 64). There is also a distribution gap in Oman east of Dhofar where
the species is obviously missing (MORDAN 1980 a: 110, 1988: 399).
Genus Allopeas H.B. Baker, 1935
1935 Allopeas H.B. Baker. — Nautilus 48 (3): 84.
Diagnosis: The shells are slender turreted with a blunt protoconch and a simple straight
columella. The distal penis is enveloped by a heavy muscular sheath. The epiphallus is short and

374 E. NEUBERT
Fig. 64: Distribution of Zootecus insularis and Allopeas gracilis on the Arabian Peninsula.
projects into the penial lumen with a conical papilla with a central perforation. A penial caecum
alongside the epiphallus with a retractor muscle uniting with the epiphallial MRP (PILSBRY 1946:
177, figs 84, 8-10).
Type species: Bulimus gracilis Hutton, 1834.
Allopeas was created by H.B. Baker “to include species of Lamellaxis with elongate radular
marginals which retain tricuspid fascies, with better developed accessory penial caecum than in
Leptopeas and Lamellaxis s. str., with relatively simple straight columella, and without distinctly
colored shell cuticule”. In the author’s opinion, the value of radular characters for generic
definition is questionable in many groups of terrestrial molluscs. The differentiation of the male
genital organ supplies much more reliable criteria, although it only can be used when sufficient
anatomical data are present. This is not the case, in particular for Lamellaxis Strebel & Pfeffer,
1882 (type species Spiraxis mexicanicus L. Pfeiffer, 1866). Allopeas is retained as a separate genus
here because its type species has a blunt protoconch and a crenulate suture, characters which are
not known from species of Lamellaxis.
Allopeas gracilis (Hutton, 1834) Figs 64, 67
1834 Bulimus gracilis Hutton. — J. Asiatic Soc. Bengal 3: 93.
1872 Limicolaria bourguignati Paladilhe. — Ann. Mus. Stor. Nat. Genova 3: 18, tav. 1, figs 13-14 (“Environs d’Aden”).

Terrestrial and freshwater molluscs of the Arabian Peninsula 375
Material: Saudi Arabia: Eastern Province, al-Qatif Oasis, freshwater course W of town, 26°36'N 49°58'E,
31.I.1992, R. Kinzelbach & I.A. Nader, SNMNH-MO 56/1; Eastern Province, al-Aba Oasis (Saadah, Dumayn), 9.VI.1992, R.
Kinzelbach, SNMNH-MO 57/1, SMF 311705/1; Eastern Province, Jubail, Dauhat ad-Dafi, in gardens, 11.XII.1993, P. Symens
& M. Werner, SMF 311587/2; Eastern Province, Ayn ibn Umayr, 26°41'N 49°51'E, 16.II.1994, P. Symens, SNMNH-
MO 58/1. — Yemen: Red Sea, ad-Durayhimi, 14°40'N 42°59'E, sand beach, 16.IV.1993, H. Dekker & F. Ceuninck v.
Capelle, DEKK (3). — O m an : Oasis N of Nizwa, palm plantations, 13.III.1995, J. Wittmann, NEUB (2); Wadi Qalhat/Hilm,
22°41'N 59°22'E, 18.IV.1978, A.B. Paltrinieri, ONHM 610.23.
Description: The opaque slender shell is conical elongate. The protoconch is dome-shaped
and smooth within the first whorl, but sutural crenulation starts with the second protoconch whorl.
The teleoconch whorls are evenly rounded with a deep suture which is crenulated by minute
papillae. The surface of the whorls is covered by fine and dense axial striae which are curved
suprasuturally.
The aperture is oval and lacks any dentition. The columella is straight and somewhat thickened.
The umbilicus is closed.
Remarks: The neotropical A. gracilis is widespread in the Indopacific area and seems to be
distributed easily by human activity. The records given here extend the range along the coast of the
Arabian Gulf considerably. More attention should be paid to inland oases as they are potential
dwelling grounds for a species with this autecology (Fig. 64).
Genus Obeliscella Jousseaume, 1889
1889 Obeliscella Jousseaume. — Bull. Soc. Malac. France 6: 359.
Diagnosis: High spired shells, slender conical. The whorls are smooth and shiny. The
columellar margin is straight, the columellar rim projects over the umbilicus, closing it completely.
Type species: Bulimus lucidissima Paladilhe, 1872.
Obeliscella lucidissima (Paladilhe, 1872) Figs 68-69, 73
1872 Bulimus lucidissimus Paladilhe. — Ann. Mus. Stor. Nat. Genova 3: 17, tav. 1, figs 18-19 (Aden).
1889 Ennea ? lucidissima v. Martens. — Nachrbl. dtsch. malakozool. Ges. 21 (9/10): 152 (“Vorhügel bei Badjil und am
Gebel Bura bei Chalifa”).
1890 Obeliscella ? Martensi Jousseaume. — Bull. Soc. Malac. France 7: 99 (“Badjil, Gebel Bura bei Chalifa”).
1896 Stenogyra bentiae Melvill & Ponsonby. — Proc. Malac. Soc. London 2: 1, pl. 1, fig. 4 (Dhofar).
Material: Yemen: Badjil, SMF 157245/10, coll. Reinhardt ex Schweinfurth; same locality, SMF 296678/3, coll. O.
Boettger ex Schweinfurth; same locality, SMF 296677/2, coll. Kobelt ex v. Martens; between Amran and Hajjah, 27.III.1988,
A. Liebegott, LIEB (1); at the road from al-Hodeida to Menaha, foothills of the escarpment, 800 m, 25.III.1988, A. Liebegott,
LIEB (7); same data but escarpment, 1200 m, LIEB (7); Sharas, Wadi Sharas, NE of Hajjah, 800 m, old river sediments,
9.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (1); al-Mahrah, Hawf, E side of village, 16°38'03"N 53°02'34"E, rocks
with algae and sand patches, 1.X.1995, H. Dekker & F. Ceuninck v. Capelle, DEKK (17); al-Mahrah, between Damquat and
Jahib, 16°35.5'N 52°53'E, limestone rocks, 1.X.1995, H. Dekker & F. Ceuninck v. Capelle, DEKK (13). — O m a n :
SMF 311686/1, Dhofar, NE slope of Qara mountains, Jibjat area (I.a.e.n. KR 31), Girod ded. 1997.
Type material: Two syntypes Stenogyra bentiae, BM(NH) 1895.7.10.5 and 1896.4.28.4.
De s cri p ti o n : The turreted shell is light corneous with a broad, dome-shaped and smooth
protoconch. The teleoconch whorls are nearly flat, the suture is very shallow. Subsuturally, a faint
white borderline is present. The whorls are glossy and smooth apart from very weak oblique axial
striae with irregular interspaces. There is no umbilicus.
The aperture is pyriform, the peristome is blunt without any callosities. The columellar
margin is almost straight, a thin to moderately thick callus connects the columellar area and the
upper peristomial rim.
Measurements (illustrated syntype S. bentiae BM(NH), Fig. 69): H = 14.50; D = 3.80;
PH = 4.20; PD = 2.80; W = 10.5.

376 E. NEUBERT
Figs 65-71: (original size × 3), all in frontal view. 65-66: Zootecus insularis. 65: Specimen from Island of Kameran, SMF 296623.
66: Holotype of Imparietula wissmanni, SMF 50242. 67: Allopeas gracilis, specimen from Oman, Oasis N of Nizwa. 68-69:
Obeliscella lucidissima. 68: Bulimus lucidissimus ex PALADILHE (1872: tav. 1, fig. 18). 69: Syntype of Stenogyra bentiae,
BM(NH) 1895.7.10.5. 70: Homorus splendens, specimen from Yemen, Menaha (SMF 145498). 71: Homorus arabica, holotype
of Subulina arabica, BM(NH) 1939.4.19.220.
Remarks: In 1889, V. MARTENS reported upon specimens of O. lucidissima from Badjil,
Yemen which he transferred, with some doubts, to the genus Ennea.
Parts of his description read: “… Apertura paululum obliqua, ¼ longitudinis superans,
subovata, superne acutangula, peristomate recto obtuso crassiusculo, margine columellari sat
obliquo, basi subangulato, superne dilatato et appresso, in callum parietalem distinctum abeunte.”
— This translates to: The aperture is only weakly oblique, reaching a quarter of the total length. It
is subovate, acutely angulate above. The peristome is straight and bluntly thickened, the columellar
margin is somewhat oblique, the base subangulate, extended above and pressed to [the body
whorl], converting in a distinct parietal callosity.
This excerpt indicates that he discussed O. lucidissima (parietal callus, form of the aperture and
columella).
In 1890, JOUSSEAUME introduces the name O. martensi, referring to the record of lucidissima
published by v. MARTENS (1889). Jousseaume gives no plausible reason why he considered his O.
martensi to be different from lucidissima. A note, that he had checked the material, is missing. In
the author’s opinion, it was a misunderstanding of Jousseaume, who erroneously believed that the
question mark of v. Martens meant that he doubted his identification. For this reason, Jousseaume
introduced this substitute name.
The Senckenberg Museum houses parts (?) of the questionable lot from Badjil described by v.
Martens. The original label of one of these lots (SMF 296677/2) reads “Ennea ? lucidissima” in the
handwriting of v. Martens. It contains two specimens of O. lucidissima. These specimens are here
considered to represent syntypes of Obeliscella ? Martensi Jousseaume, 1890. Comparing these type
specimens to the syntypes of Stenogyra bentiae and the figures of Bulimus lucidissima given by
PALADILHE (1872) reveal that all these nominal taxa can be considered as synonyms since there is
no conchological difference.
THIELE (1910) notes that he checked the original lot of v. Martens which contained specimens
looking different, leading him to his description of Subulina splendens. It is presumed that he failed
to find the original lot of v. Martens (missing or lost?) in the ZMHB. These findings should be
checked in the ZMHB. If this lot is definitely lost, one of the syntypic specimens from the SMF

Terrestrial and freshwater molluscs of the Arabian Peninsula 377
may be used for a lectotype designation of O. martensi (if necessary, as O. martensi is simply a
synonym!).
Judging from the material presented here, it seems highly probable that both species, O.
lucidissima and H. splendens, live sympatrically at Badjil (Fig. 73).
Genus Homorus Albers, 1850
1850 Homorus Albers. — Die Heliceen nach natürlicher Verwandtschaft systematisch geordnet, 2. Aufl.: 196.
Diagnosis: Slender elongate turreted shells with a truncate and concave columella, non-
umbilicate. The protoconch is subcylindrical and blunt.
Type species: Achatina cyanostoma L. Pfeiffer, 1842.
Usually, the two Arabian species discussed here are classified within Subulina Beck, 1837 (type
species Bulimus octona Bruguière, 1789). This species was investigated anatomically by H.B. BAKER
(1927: 2, pl. XX, fig. 99). It is characterised by the presence of a strong muscular penial sheath
with a swollen and internally folded distal penis, an extremely elongate penial tube, a reduced or
missing epiphallus and a retractor muscle inserting at the penial caecum. The male part of the
genitalia differs profoundly from what is described here for “Subulina” splendens.
Unfortunately, there are insufficient data available on the genital anatomy of the Homorus
species, although this genus is well represented in the Ethiopian area and Central Africa. PILSBRY
(1919: 114, figs 41 a, b) depicted the genital organs of H. (Subulona) clarus Pilsbry, 1919, and H.
(Oreohomorus) bequaerti Pilsbry, 1919. In the latter species, only a juvenile specimen was dissected
showing a simple penis. In H. clarus, however, the penis is swollen distally (papilla inside?) with a
slightly longer epiphallus. Clearly, there are no appendices to be found on the male organs. This
morphology is close to what is found in the Arabian species. Thus, a congenerity of both Arabian
species seems to be very likely, as the shells also share the same conchological characteristics. A
definition of Homorus and its phylogenetic relationship to Subulona v. Martens, 1889, Itiopiana
Preston, 1910, Oreohomorus Pilsbry, 1919, and Striosubulina Thiele, 1933 is completely un-
resolved. For this reason, a subgeneric classification should be postponed until more anatomical
data on African members of the family Subulinidae in general are available.
Homorus splendens (Thiele, 1910) Figs 70, 72-73
1910 Subulina splendens Thiele. — Sitzungsber. Ges. naturf. Fr. Berlin 1910: 283, Taf. 10, Fig. 5 (Badjil [Yemen]).
Material: Saudi Arabia: Asir mountain region, Raydah escarpment, 2350 m, 29.I.1994, P. Symens & M. Werner,
SNMNH-MO 59/10, SMF 311588/9; Asir mountain region, Abha escarpment, Juniperus forest, 2300 m, II.1994, P. Symens
& M. Werner, SNMNH-MO 60/3, SMF 311589/4 (all preserved). — Yemen: Menaha, 2300 m, SMF 145498/1, coll.
Reinhardt ex Schweinfurth; Bani al-Harith, Wadi Dhar (= Zahr), Qaryat al-Qabil, NE of Sana’a, 2200 m, between leaves, wood
and soil, 8.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (11).
Description: The corneous to olivaceous shell is turreted and glossy. The protoconch
consists of two smooth whorls. The second protoconch whorl is somewhat higher than the
preceding first teleoconch whorl. These whorls are flat and have a shallow suture. The surface of
the teleoconch is covered by fine axial striae.
The aperture is elongate pyriform. The peristome is acute, a parietal callosity is lacking. The
columellar margin is concave, the columella obliquely truncate.
Genital morphology (Fig. 72): The short penis is bipartite with a globular proximal
part. Internally, this part is filled by a huge penial papilla with a central perforation. In the distal
penis tube, two small longitudinal pilasters are present fusing with two faint vaginal pilasters. The
epiphallus is a tube as long as the penis. Internally, there are two main longitudinal pilasters which

378 E. NEUBERT
Fig. 72:
Genital organs of
Homorus splendens.
are folded transversely. The VD enters the epiphallus with a minute papilla. The MRP inserts at
the boundary of epiphallus and VD and attaches at the diaphragm.
The vagina is extremely long. Internally, only the distal lumen shows two pilasters, the
proximal tube is smooth. The stalk of the BC branches off basally. A diverticulum is missing, and
the pedunculus is somewhat swollen in its basal part. The BC is a small rounded vesicle.
Measurements (illustrated specimen SMF 145498/1, Fig. 70): H = 10.30; D = 2.95;
PH = 2.80; PD = 1.40; W = 9.
Homorus arabica (Connolly, 1941) Figs 71, 73
1941 Subulina arabica Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 24, pl. 3, fig. 2 (Wadi
Dareija SW of Dhala).
Type material: Holotype Subulina arabica, BM(NH) 1939.4.19.220, paratype BM(NH)
1939.4.19.221.
Description: The corneous shell is slender and turreted. The protoconch consists of three
subcylindrical whorls. The teleoconch whorls increase rapidly in diameter. They are glossy and
evenly rounded with a suture of medium depth. Fine axial striae are present.
The aperture is obliquely pyriform with a concave columellar margin. The columella is
truncate above the base of the shell.
Measurements (holotype, Fig. 71): H = 11.95; D = 3.15; PH = 2.20; PD = 1.60; W = 8.5.
Remarks: This species is close to H. splendens but is kept separate here as its suture is deeper,
the whorls are more rounded and the protoconch is slightly constricted which, in contrast to
CONNOLLY (1941: 25), is characteristic of his H. arabica (Fig. 71) and not of H. splendens (Fig. 70).
Family Streptaxidae
Genus Streptostele Dohrn, 1866
1866 Streptostele Dohrn. — Malakozool. Bl. 13: 128.
Diagnosis: The fragile shell is elongate conical. The whorls are sculptured by oblique ribs
or become smooth on the lower whorls. The suture is somewhat crenulate in many of the ribbed

Terrestrial and freshwater molluscs of the Arabian Peninsula 379
species. The aperture is ovate and sometimes displays a weak denticle. The columella is somewhat
oblique and broadened.
Type species: Bulimus fastigiatus Morelet, 1848.
Subgenus Raffraya Bourguignat, 1883
1883 Raffraya Bourguignat. — Histoire Malacologique de l’Abyssinie: 66.
Diagnosis: The shells are completely ribbed. The oblique aperture is characterised by an
excavated sinulus with a palatal callosity or denticle on the upper peristomial rim.
Type species: Raffraya Milne-Edwardsi Bourguignat, 1883.
Streptostele (Raffraya) scotti Connolly, 1941 Figs 73-74
1941 Streptostele scotti Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 18, pl. 3, fig. 1 (Jabal
Jihaf, 7700 ft, cliffs near spring [Yemen]).
Type material: Holotype Streptostele scotti, BM(NH) 1939.4.19.190, five paratypes
BM(NH) 1939.4.19.191-195.
Description: The small corneous shell is turreted and slender. The smooth and glossy
protoconch consists of 2.5 whorls and nearly reaches the diameter of the teleoconch. The upper
teleoconch whorls are rounded, the lower whorls are more flat. The suture is of moderate depth.
The surface of the whorls is covered by evenly spaced and densely packed ribs.
The aperture is considerably small, obliquely pyriform and somewhat squarish at the base.
The peristome is slightly thickened with a small recurved sinulus. Below the sinulus, the
peristome may protrude in a small callose triangle. The columellar margin is straight and
somewhat enlarged covering the umbilicus. The parietal callus is thin with a medium to strong
angularis.
Measurements (holotype, Fig. 74): H = 6.90; D = 1.80; PH = 1.55; PD = 1.40; W = 8.5.
Remarks: This species is only known from the type locality. It is here assigned to the
subgenus Raffraya because of the obvious conchological similarities to S. (R.) milneedwardsi.
This Ethiopian species differs by the shape of shell, aperture and its rimate umbilicus.
Genus Gulella L. Pfeiffer, 1856
1856 Gulella L. Pfeiffer. — Malakozool. Bl. 2: 173.
Diagnosis: Small ovoid to turreted shells, smooth to coarsely ribbed. The aperture is
rounded to nearly quadrate with a flared peristome. The almost heavy dentition is composed of
the parietalis, which may be connected to an angularis. The aperture is often constricted palatally,
displaying a strong palatalis. One to two columellar lamellae may be present.
Type species: Pupa menkeana L. Pfeiffer, 1853.
Gulella bicolor (Hutton, 1834)
1834 Pupa bicolor Hutton. — J. Asiatic Soc. Bengal 3: 93.
Description: The glossy shells are turreted and slightly fusiform. The smooth protoconch
consists of 2.5 whorls and is somewhat bulbous. The following teleoconch whorls are smooth and
somewhat rounded, the suture is of medium depth. Minute subsutural riblets characterise the
upper parts of the whorls and disappear subsequently.

380 E. NEUBERT
Fig. 73: Distribution of some species of Subulinidae, Streptaxidae and Urocyclidae in the south-eastern part of the Arabian
Peninsula.
The subquadrate aperture is very small. On the upper edge, the sinulus is delimited by the
strong lamella formed by the angularis and parietalis and the strong palatal fold situated perpen-
dicular on the inner apertural wall. A minute basalis is present. The single columellaris is situated
deep in the aperture and is truncate basally.
The last whorl is coarsely ribbed on the cervix, a deep squarish furrow corresponds to the
palatalis. The peristome partially protrudes at the palatal insertion, forming a subacute triangle in
the peristomial outline. The peristome is slightly thickened and somewhat reflected. A thin parietal
callus connects the peristomial rims. The umbilicus opens with a minute perforation.
Measurements (according to MORDAN 1988: 399, fig. 2): H = 5.30; D = 1.86; PH = 1.50;
PD = 1.36; W = 7.
Remarks: Gulella bicolor is widespread in the Asian area. There is no additional record apart
from that published by MORDAN (1988: 399) from Hawiyah in Oman.
This species is usually assigned to the subgenus Huttonella L. Pfeiffer, 1855. Since a subgeneric
assignment of the other species described herein has to be postponed until more data are available,
a subgeneric placement is rejected. Arabia is an area of marginal importance for the taxonomic
revision of this genus, which contains a plethora of unrevised generic and specific names.
Gulella isseli (Paladilhe, 1872) Figs 73, 75-76
1872 Ennea isseli Paladilhe. — Ann. Mus. Stor. Nat. Genova 3: 19, tav. 1, figs 5-6 (“Environs d’Aden”).
Material: Yemen: Sharas, Wadi Sharas, NE of Hajjah, 800 m, old river sediments, 9.IV.1993, H. Dekker & F.
Ceuninck v. Capelle, DEKK (1).

Terrestrial and freshwater molluscs of the Arabian Peninsula 381
De s cri p ti o n : The small fragile shell is turreted and nearly cylindrical. Its surface is glossy,
the shell described here is white, but it is not known whether this represents the natural coloration
of this species.
The protoconch consists of two smooth and dome-shaped whorls. The teleoconch whorls are
smooth apart from some irregularly spaced oblique axial stripes. The whorls are flattened, the
suture is shallow and displays characteristic minute papillae.
The aperture is semi-quadrate, the peristome thickened, its rims are not connected. The strong
angularis connects to the short parietalis which is indicated by a small depression. The sinulus is
deep and somewhat recurved, the peristomial rim is connected to the angularis. There are two
palatal denticles situated on an oblique palatal callosity. The small basalis is acute. In PALADILHE’s
original figure (1872: tav. 1, fig. 6), the basalis is absent although it is mentioned in the text
(“… margine basali tuberculo dentiformi munito”). There are two somewhat oblique columellar
lamellae. The umbilicus is slit-like.
Measurements (illustrated specimen ex coll. Dekker, Fig. 76): H = 4.50; D = 1.80; W = 6.
Remarks: This species seems to be very rare. The original locality “Aden” is vague and may
be better understood as “southern Yemen” (Fig. 73).
Gulella schweinfurthi (Thiele, 1910) Figs 73, 77, Plate 1
1910 Ennea schweinfurthi Thiele. — Sitzungsber. Ges. naturf. Fr. Berlin 1910: 283, Taf. X, Fig. 6 (Menaha).
Material: Saudi Arabia: Asir mountain region, Raydah escarpment, 2350 m, 29.I.1994, P. Symens & M. Werner,
SNMNH-MO 61/3, SMF 311590/2; same data but 1800 m, SMF 311591/1. — Yemen: Bani al-Harith, Wadi Dhar (= Zahr),
Qaryat al-Qabil, NE of Sana’a, 2200 m, between leaves, wood and soil, 8.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK
(4 adults, 2 fragments).
Description: The turreted shell is elongate cylindrical, the last whorl is somewhat nar-
rowed in some specimens. The colour of fresh shells is bright corneous with a greenish glaze while
eroded shells are purely white.
The protoconch consists of 2.5 whorls which display a finely beaded spiral sculpture. The
teleoconch whorls are rather flat with a suture of medium depth. The surface is covered with
regularly spaced oblique ribs. Between the ribs, minute spiral striae are present. Apically, the ribs
are somewhat broadened to form small papillae.
The subquadrate aperture has a shallow sinulus. The peristome is slightly thickened and
strongly reflected. The peristomial rims are connected by a thin callus. The dentition is reduced,
only a weak angularis remains. The umbilicus is slit-like and always open.
The animals are orange to deep red in colour (Plate 1).
Measurements (illustrated specimen ex coll. Dekker, Fig. 77): H = 7.55; D = 2.55;
PH = 2.10; PD = 1.70; W = 10.5.
Remarks: This species seems to be confined to the higher altitudes in the south-western
mountain belt (Fig. 73).
Gulella protruda Neubert & Frank, 1996 Figs 78-80
1996 Gulella protruda Neubert & Frank. — Arch. Molluskenk. 126 (1/2): 125, Fig. 1-3 (Oman: Jabal Akhdar, Hoti cave
system, 990 m).
Material: Oman: Jabal Sira, Jabal Akhdar, 15.V.1972, M.D. Gallagher, BM(NH) (78), SMF 311720/3; Wadi
Qalhat/Hilm, 22°41'N 59°22'E, 18.IV.1978, A.B. Paltrinieri, ONHM 610.23.
Description: The dome-shaped protoconch consists of two whorls, its diameter reaches
more than half of the diameter of the shell. Its surface is finely granulated, the sutures of the
protoconch are less deep than those of the teleoconch.
The small shell is turreted, cylindrical and may consist of five to eight whorls. The surface of
the teleoconch whorls is densely covered by minute but strong oblique ribs. The whorls are narrow

382 E. NEUBERT
and evenly rounded, the subcanaliculate suture is deep. The last whorl is flat with a blunt keel
surrounding the periomphalum. The umbilicus is narrowly open.
The last whorl is characterised by two furrows, an upper shallow one and a deep and broad
one situated basolaterally. These furrows correspond to two palatal denticles. The last whorl
protrudes from the teleoconch to form a subtriangular aperture. The peristome is only slightly
thickened and somewhat reflected.
The dentition consists of four tooth-like lamellae which are all situated deep in the aperture.
The upper palatalis is slightly elongated, the lower palatalis is thin in the interior of the shell but
expands considerably towards the aperture to form a nodule. The columellaris, situated basally on
the columella, is a thick bifid callus. There is no trace of this denticle on the columella in the
interior of the shell. The parietalis is a strong and broad lamella and arched frontally. A weak
angularis may be observed left of the shallow sinus.
Measurements (illustrated specimen SMF 311720 a, Figs 78-80): H = 2.75; D = 1.45.
Figs 74-85: (original size × 6), 74: Streptostele (Raffraya) scotti, holotype BM(NH) 1939.4.19.190, frontal view. 75-76: Gulella
isseli. 75: Ennea isseli ex PALADILHE (1872: tav. 1, fig. 6), frontal view. 76: Specimen from Yemen, Wadi Sharas, frontal view. 77:
Gulella schweinfurthi, specimen from Yemen, Bani al-Harith, Wadi Dhar, frontal view. 78-80: Gulella protruda, specimen from
Oman, Jabal Akhdar (SMF 311720 a). 78: Frontal view. 79: Dorsal view. 80: Lateral view. 81-83: Gudeella rufocincta, holotype
BM(NH) 139.4.19.196-207. 81: Frontal view. 82: Apical view. 83: Subsurface. 84-85: Gudeella eremias, syntype BM(NH)
96.4.28.4-8. 84: Frontal view. 85: Apical view.

Terrestrial and freshwater molluscs of the Arabian Peninsula 383
Figs 86-87: Gudeella rufocincta, preserved animal. 86: Apical view with mantle collar and the two digitiform mantle lobes. Note
the pointed upright “tail” with porus of mucous gland. 80: Lateral view.
Family Urocyclidae
Genus Gudeella Preston, 1913
1913 Gudeella Preston. — Proc. Malac. Soc. London 10: 285 [nom. nov. pro Thapsiella Gude, 1911 non Fischer, 1885].
Diagnosis: Small, fragile, depressed to low conical shells. They are umbilicate, the peri-
stome is simple and the columellar margin somewhat reflected.
Type species: Thapsia masukuensis E.A. Smith, 1899.
Gudeella rufocincta Connolly, 1941 Figs 73, 81-83, 86-87
1941 Gudeëlla rufocincta Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 19, pl. 3, figs 7-9 (Foot
of Jabal Harir, dead leaves under wild fig tree, 5000' [Yemen]).
Material: Saudi Arabia: Asir mountain region, Raydah escarpment, 2100 m, 10.VII.1995, S. Newton, SNMNH-
MO 62/1, SMF 311592/1; Raydah escarpment, 2300 m, VII.1995, S. Newton, SMF 311593/1 (preserved); Asir mountain
region, Abha escarpment, Juniperus forest, 2300 m, II.1994, P. Symens & M. Werner, SNMNH-MO 63/1. — Yemen: Bani
al-Harith, Wadi Dhar (= Zahr), Qaryat al-Qabil, NE of Sana’a, 2200 m, between leaves, wood and soil, 8.IV.1993, H. Dekker
& F. Ceuninck v. Capelle, DEKK.
Type material: Holotype and paratype Gudeella rufocincta, BM(NH) 139.4.19.196-207
(holotype marked, no separate catalogue number).
Description: The shell is depressed with a low spire. It is milky opaque with a red-brown
spiral band running slightly above the periphery and may be seen suprasuturally. The protoconch
and teleoconch are smooth and glossy. The complete surface is covered by very fine spiral grooves
(magnification > 30 ×) which become wavy on the last whorl. The teleoconch whorls are narrowly
coiled and rather flat with a shallow suture.
The last whorl is bluntly angulate above the periphery, the subsurface is flattened. The nearly
circular umbilicus is narrow and deep with steep walls. The aperture is obliquely crescent-shaped.
The peristome is sharp with a weak labial callus. The columellar margin is straight with a small
callus reflecting over the umbilicus.
The (preserved) animal is bluish coloured. The foot is elongated and very narrow, a medium
zone is marked. The “tail” is acute with a triangular lobe pointing upwards. In the centre of this lobe,
a bright field marks the pore of the mucous gland. The mantle collar is subdivided into three parts.
The basic one covers the neck and supports the right mantle lobe. The medium one is a thin bilobed
shield above the first one but reaches also to the left side of the neck. The last one represents the
upper wall of the mantle and consists of a thin and recurved lamella supporting the left mantle
lobe. Both mantle lobes are elongate and acute reaching at least over the entire shell (Figs 86-87).

384 E. NEUBERT
The outer lung wall is irregularly patterned by dark pigments.
Measurements (holotype, Figs 81-83): H = 3.25; D = 6.65; PH = 1.80; PD = 2.20;
W = 4.5.
Remarks: The preserved specimen illustrated was collected at Raydah escarpment at 2300 m
(Fig. 73). Its shell measured 5.8 mm in diameter. Unfortunately, the reproductive organs were not
mature and only a rudimentary female part was present.
Gudeella eremias (Melvill & Ponsonby, 1896) Figs 73, 84-85
1896 Hyalinia (Arnouldia) eremias Melvill & Ponsonby. — Proc. Malac. Soc. London 2: 1, pl. 1, figs 12-14 (Dhofar).
Type material: Two syntypes Gudeella eremias, BM(NH) 96.4.28.4-8, one of them glued
to a piece of paper. The unglued syntype is illustrated.
Description: The brown shell is small with an elevated to submammillate spire. The
surface of the teleoconch is smooth and glossy and covered by very fine spiral furrows. The suture
is shallow and bordered by a subopaque line. The subsurface of the shell is well rounded, the
interior walls of the umbilicus bear strong spiral ridges.
The last whorl of the teleoconch is bluntly angulate. The subquadrate aperture has a sharp
peristome and is not callose.
Measurements (illustrated syntype, Figs 84-85): H = 3.15; D = 4.60; PH = 2.60; PD = 2.50;
W = 4.
Remarks: This species was hitherto only known from a restricted area in Oman at Dhofar
(Khadrafi, Jabal Qamr, 16°42'N 53°09'E) close to the Yemen border, where it occurs sympatrically
with G. rufocincta (MORDAN 1980 a: 107 and Fig. 73 herein).
Family Vitrinidae
Genus Arabivitrina Thiele, 1931
1931 Arabivitrina Thiele. — Handbuch der systematischen Weichtierkunde I (2): 600.
Diagnosis: Vagina with a voluminous glandula amatoria, the proximal vagina part is coated
by glandular tissue. Penis with a longitudinal pilaster consisting at least of an apical glandular part
followed by a narrow to broadened part with transverse lamellae.
Type species: Vitrina arabica Thiele, 1910.
Species of Arabivitrina are known from a vast area in Ethiopia and from the south-western
Arabian Peninsula. Its distribution patterns are more or less the same as exhibited in the genus
Lejeania. A general revision comprising all taxa involved in Arabivitrina is still pending.
Arabivitrina arabica (Thiele, 1910) Figs 88-90, 94-100, 107
1910 Vitrina arabica Thiele. — Sitzungsber. Ges. naturf. Fr. Berlin 1910: 281, Taf. X, Fig. 1, 1 a (Menaha).
Material: Saudi Arabia: Asir mountain region, Raydah escarpment, 2350 m, 29.I.1994, P. Symens & M. Werner,
SNMNH-MO 64/8, SMF 311594/7; Raydah escarpment, Juniperus forest, 2130 m, 18°11'41.6"N 42°24'10.8"E, 5.V.1994, W.
Schneider & F. Krupp, SNMNH-MO 76/3, SMF 311606/3, HLMD (3) (all preserved); Raydah escarpment, 1800 m,
29.I.1994, P. Symens & M. Werner, SNMNH-MO 65/3, SMF 311595/3 (all preserved); Raydah escarpment, 2050 m,
25.III.1994, W. Schneider & F. Krupp, SNMNH-MO 66/3, SMF 311596/2; Raydah escarpment, al-Aqaba spring, 2310 m,
18°11'50.0"N 42°24'30.2"E, 5.V.1994, W. Schneider & F. Krupp, SNMNH-MO 67/4, SNMNH-MO 68/3 (preserved),
SMF 311597/2, SMF 311598/3 (preserved), HLMD (2); Raydah escarpment, 2200 m, I.1995, S. Newton, SNMNH-
MO 69/1, SMF 311599/1; Raydah escarpment, 2100 m, 10.VII.1995, S. Newton, SNMNH-MO 70/9, SMF 311600/8,
SMF 311601/2 (preserved); Raydah escarpment, 2300 m, 10.VII.1995, S. Newton, SNMNH-MO 71/5, SNMNH-MO 72/5
(preserved), SMF 311602/3, SMF 311603/3 (preserved); Asir mountain region, Jabal Qahar, 17°39'N 42°53'E, II.1996, S.
Newton, SNMNH-MO 73/7, SMF 311604/6; Asir mountain region, Abha escarpment, Juniperus forest, 2500 m, II.1994, P.

Terrestrial and freshwater molluscs of the Arabian Peninsula 385
Figs 88-93: (original size × 3), 88-90: Arabivitrina arabica, syntype SMF 158370 a. 88: Frontal view. 89: Apical view. 90:
Subsurface. 91-93: Arabivitrina jansseni n. sp., holotype SMF 311267. 91: Frontal view. 92: Apical view. 93: Subsurface.
Symens & M. Werner, SNMNH-MO 74/2; Abha escarpment, Juniperus forest, 2300 m, II.1994, P. Symens & M. Werner,
SNMNH-MO 75/2, SMF 311605/1. — Yemen: Pass over the Sumara mountains N of Sana’a, XII.1992, S. Neubert, NEUB
(8); Dschibbla, between Sana’a and Ta’izz, 22.III.1988, A. Liebegott, LIEB (1); Jabal Sabar close to Ta’izz, 23.III.1988, A.
Liebegott, LIEB (3); on top of the Sumara pass, 22.III.1988, A. Liebegott, LIEB (9); Bani Matar, Jabal an-Nabi Shu’ayb, W of
Sana’a, 3660 m, under stones, 27.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK.
Type material: Syntype SMF 158370 a, coll. Reinhardt (= ZILCH 1960: Abb. 842); syn-
types SMF 158370/14, coll. Reinhardt; syntypes SMF 158371/2, coll. v. Moellendorff; syntypes
SMF 158372/3, coll. O. Boettger. All lots originate from: “S-Arabien: Menaha (7000 Fuß)
Schweinfurth” thus representing parts of the type lot.
Description: The fragile shell is medium to large in size. Shells of well-preserved specimens
are covered by a greenish periostracum. This may erode easily leaving a white shell.
The protoconch is smooth and consists of one whorl. Under high magnifications (50 ×), a
dense pattern of spiral punctuation can be observed. The teleoconch increases quickly in diameter,
producing an ear-shaped shell. Very often, blunt axial riblets occur running from the suture to the
periphery. The subsurface is smooth. The spire is always elevated with an impressed suture.

386 E. NEUBERT
Figs 94-100: Genital organs of Arabivitrina arabica. 94-95: Specimen from Asir mountain region, Raydah escarpment, 1800 m.
94: Genital apparatus. 95: Proximal vagina dissected, showing vaginal papilla. 96-97: Specimen from Asir mountain region,
Raydah escarpment, 2100 m. 96: Genital apparatus. 97: Penis dissected, showing penial gland, lamellate pilaster and two main
pilasters. 98-100: Specimen from Yemen, Pass over the Sumara mountains N of Sana’a. 98: Genital apparatus. 99: Proximal
vagina dissected, showing vaginal papilla. 100: Penis dissected, showing penial gland.

Terrestrial and freshwater molluscs of the Arabian Peninsula 387
The aperture is depressed elliptical and bordered by a small fringe of periostracum. The
columella is weakly callose and sometimes white.
Measurements: Illustrated syntype SMF 158370 a (Figs 88-90): H = 11.2; D = 16.9;
PH = 11.0; PD = 11.9; W = 3.5. The maximum measurements were taken from a specimen from
Raydah escarpment with H = 15.6; D = 21.4; PH = 15.0; PD = 15.0; W = 3.25.
Genital morphology (Figs 94-100): The penis is a broad tube, the MRP inserts apically
on a short and blunt protrusion. The VD enters the penis sublaterally where the penis is coated by
a glandular tissue, the penial gland. Internally, three remarkable structures are present. The first is
a longitudinal pilaster subdivided by several deep constrictions. The second is an organ with a
complicated structure. It starts apically at the attachment site of the MRP. This part is constituted
by a pilaster with a coating of glandular tissue which is soft and can easily be stripped off. It is
followed by a second triangular part which is covered by lamellae perpendicular to the longitudinal
axis of the penis. Distally, a short and narrow stretch of undifferentiated pilaster occurs before it
broadens again and has a lateral row of several thick and hard nodules or warts. They are connected
to the third structure, a small pilaster, which runs parallel to the nodular part of the main pilaster on
the inner surface of the penis and exhibits the same warts. Distally, the main pilaster narrows and
unites with atrial pilasters. In adult specimens, the vagina is shorter than, or as long as, the penis.
The vagina is a broad tube subdivided into a proximal and distal part. The proximal part is
formed by a huge glandula amatoria projecting with a curved papilla of medium size into the
vaginal lumen. The papilla is covered by minute wrinkled crests of tissue, the pore lies centrally on
the papilla’s apex. The glandula amatoria is covered by a dense glandular tissue with racemose
follicles. Thin stretches of connective tissue cover the gland longitudinally. The pedunculus is thin,
the BC a well-rounded vesicle close to the free oviduct. The distal part of the vagina is thin-walled
and lacks any remarkable internal structures. The atrium is very long.
Remarks: The specimens from Yemen (pass over Sumara mountains) have a strongly
elevated spire. The shells are more conical than the syntypes studied but can still be assigned to A.
arabica (Fig. 107).
Arabivitrina jansseni n. sp. Figs 91-93, 101-107, Plate 2
Type material and locus typicus: holotype SMF 311267, paratypes SNMNH-MO 81/67, SNMNH-MO 82/10
(preserved), SMF 311268/43, SMF 311613/10 a and b (a = preserved animals, b = shells), SMF 311614/5 (preserved), NNM
(5), BM(NH) 1996282 (5), IBA (3 shells, 2 preserved specimens), Saudi Arabia, Asir mountain region, King Khalid descent near
Baha, 2215 m, 3.VI.1995, E. Neubert et al. — Non-type material: Saudi Arabia: Asir mountain region, 24 rkm N
of Baha, 2120 m, 28.I.1994, P. Symens & M. Werner, SNMNH-MO 89/1; Asir mountain region, 15 rkm S of Bani Sa’ad S of
Taif, 2300 m, Juniperus forest, 2.VI.1995, E. Neubert et al., SMF 311621/3, SMF 311607/1 (preserved); Asir mountain region,
small wadi at the right side of the road from Taif to al-Baha, S of Gharie, 1700 m, 2.VI.1995, E. Neubert et al., SNMNH-
MO 77/5, SMF 311608/4, SMF 311609/1 (preserved); Asir mountain region, rocky area 50 rkm S of al-Baha, 2000 m,
2./3.VI.1995, E. Neubert et al., SNMNH-MO 78/17, SMF 311610/17; Asir mountain region, at the road from Taif to al-Baha,
18 rkm N of al-Mandaq, 2050 m, 2.VI.1995, E. Neubert et al., SNMNH-MO 79/2, SMF 311611/2; Asir mountain region, al-
Abna descent near Baha, 2000 m, 3.VI.1995, E. Neubert et al., SNMNH-MO 80/13, SMF 311612/13.
Diagnosis: A species of Arabivitrina with a subglobose shell. The spire is dome-shaped, the
suture very shallow.
Description: The medium-sized shell of well-preserved specimens is opaque-greenish. The
protoconch is small, consists of one whorl and shows micropunctuations arranged in spiral rows.
The teleoconch is subglobose, the body whorl is inflated and covered by very blunt axial riblets.
The spire is low conical dome-shaped and never exceeds the outline of the shell. The suture of
the first teleoconch whorl is very shallow and bordered by a broad whitish line. The aperture is
subcircular and bordered by a narrow fringe of periostracum. The umbilical area is closed by a
small, often white, callosity, a slight callus extends to the upper rim of the peristome.

388 E. NEUBERT
The animal is not able to withdraw completely into its shell (Fig. 101).
Measurements (holotype, Figs 91-93): H = 14.1; D = 18.0; PH = 12.8; PD = 11.5;
W = 3.5.
Genital morphology (holotype, Figs 102-106): There are no major differences from A.
arabica with respect to the basic structure of the genital organs of this species. The vagina is twice
Figs 101-106: Arabivitrina jansseni n. sp. 101: Preserved animal. Note mantle collar and small right mantle lobe. 102-104:
Paratype. 102: Genital apparatus. 103: Proximal vagina dissected, showing vaginal papilla. 104: Penis dissected, showing penial
gland. 105-106: Specimen from Asir mountain region, S of Bani Sa’ad S of Taif. 105: Genital apparatus. 106: Penis dissected,
showing penial gland, lamellate pilaster and two main pilasters.

Terrestrial and freshwater molluscs of the Arabian Peninsula 389
Fig. 107: Distribution of Arabivitrina spp. on the Arabian Peninsula.
as long as the penis in adults, but most of the specimens investigated were juveniles. This character
state should investigated in detail in adults.
Etymology: This species is named in honour of Dr. R. Janssen, curator of the malacological
department of the Senckenberg Museum, who collected this new species, together with the author,
during a field trip in Saudi Arabia in 1995.
Affinities: This species is close to A. arabica. It differs from this taxon by its more globose
shell, the dome-shaped spire, the subcircular aperture and the narrower mode of coiling. Although
specimens from the Abha and Raydah escarpments look quite similar, the spire of these shells is
always elevated superseding the body whorl. The general shape of the shell is remarkably constant
in both species.
Remarks: There is insufficient material present to define the species’ distribution, but it is
evident that A. arabica lives in Yemen and the southern mountains of Asir while A. jansseni n. sp.
is only found in the northern part of the Asir mountain region. Both species are clearly confined
to humid places and are very abundant in habitats with malacophyllic herbs and shrubs. The more
arid Juniperus forests are also inhabited by both Arabivitrina species but the population density is
significantly lower (Fig. 107).
“Vitrina” gruneri L. Pfeiffer, 1846
1846 Vitrina gruneri L. Pfeiffer. — Symbolae ad Historiam Heliceorum III: 81 (Arabia).
Type material: The collection of L. Pfeiffer was lost during World War II. There are also
no specimens in Gruner’s collection, which is housed in the Löbbecke Museum in Düsseldorf,

390 E. NEUBERT
Germany (J. Boscheinen, pers. comm.). To give an impression of what gruneri might be, the text
of the original description of L. Pfeiffer is quoted here:
“T. imperforata, globoso-depressa, glaberrima, quarum nitens, olivaceo-cornea; spira vix
elevata; sutura albo-marginata; anfr. 3 1/2 convexiusculi, ultimus subdepressus; apertura perobli-
qua, lunato-ovalis; perist. simplex marginibus conniventibus, columellari arcuato, subinflexo.–
Diam. 8, alt. vix 5 mill.– Arabia. (Gruner).” — This translates to: Shell imperforate, globose-
depressed, very smooth, brilliant, olive-green-horny; the spire is scarcely elevated, the suture with
a white rim; 3½ hardly convex whorls, the last somewhat depressed; the aperture is somewhat
oblique, oval to crescent-shaped; the peristome is simple with the margins inclining, the columella
is arched and somewhat curved.
Remarks: On the one hand, this description matches characters of many species of Vitri-
nidae. On the other hand, the number of whorls indicates that the specimen(s) on which L.
Pfeiffer based his description were adult. The ratio between shell diameter and height makes it very
probable that this is indeed a separate species.
Remarks on the position of Arabivitrina
In the author’s opinion, the internal structures of the penis have not been investigated adequately
in the Phenacolimax-group, although their importance for taxonomic research (as well as for the
mating behaviour of the animals) is evident. This makes it necessary to review briefly some data
on other genera which are presumed to be closely related. The Plutoniinae Cockerell, 1893 are
characterised by the presence of a specialised vaginal organ, the glandula amatoria. The subfamily
contains several genera with an endemic distribution on the Macronesian Islands.
Phenacolimax major (A. Férussac, 1807) is known from a widespread area in the temperate
zones in the west of the western Palaearctic. Its genital organs are characterised briefly here
(Figs 108-110). The penis is composed of two parts. The distal part is a simple tube, while the
proximal part is covered by a muscular sheath and, close to the branching of the MRP, an external
penial gland. The VD submerges under the muscular sheath in the middle of the penis and
connects to the penial gland. Internally, there is only one narrow elongate pilaster in the distal
penial tube. The conical proximal part is filled by thick glandular tissues. In between the penial
pilaster and the gland, a subtrigonal massive pilaster with perpendicular ridges can be found.
The vagina is elongate with a massive conical glandula amatoria. The proximal part of the
glandula amatoria is covered by a thick layer of glandular tissue of a bilobed heart shape. The BC
is small, its stalk short, the vesicle itself somewhat elongate. Internally, the vaginal tube is simple
and smooth. The glandula amatoria projects into the vaginal lumen by an acute papilla with a central
perforation. The surface of the papilla is characterised by scale-like ridges comparable to a fir cone.
In Arabivitrina, length ratios of penis/vagina may vary to a great extent, depending on the
ontogenetic stage of the animal considered. In juvenile specimens, the vagina is always consider-
ably longer than the penis. Thus, for species definition only adult specimens should be used.
FORCART (1956) described anatomical details of several Abyssinian taxa of Arabivitrina which are
close to what is presented here for Arabivitrina arabica. In particular, the “torus” of Forcart matches
exactly the structure described here under the name major pilaster. It is much more differentiated
than Forcart describes, as he believes that the complete torus is built up of glandular tissue and
shows, for example, “peculiar transverse folds” (FORCART 1956: 119, fig. 1; sub P. (A.) abyssinicus
(L. Pfeiffer, 1848)).
Insulivitrina lamarcki (Férussac, 1821), the type species of Insulivitrina, differs in several
respects. The big right mantle lobe covers more than half of the shell. In the genitalia, a glandular

Terrestrial and freshwater molluscs of the Arabian Peninsula 391
Figs 108-116: Genital organs of Vitrinidae species related to Arabivitrina. 108-110: Phenacolimax major, specimen from France,
Le Puy. 108: Genital apparatus. 109: Penis dissected, showing penial gland, sublamellate pilaster and main pilaster. 110:
Proximal vagina dissected, showing vaginal papilla. 111-113: Insulivitrina lamarcki, specimen from Spain, Teneriffa, Agua Garcia.
111: Genital apparatus. 112: Penis dissected, showing penial gland and main pilasters. 113: Proximal vagina dissected, showing
vaginal papilla. 114-116: cf. Madeirovitrina sp., specimen from Portugal, Madeira, Ribeiro Frio. 114: Genital apparatus. 115:
Penis dissected, showing main pilastral structures. 116: Proximal vagina dissected, showing vaginal papilla and vaginal conus.

392 E. NEUBERT
tissue sheath covers the glandula amatoria. This result differs from that of IBAÑEZ et al. (1987: 126),
who stressed the absence of this tissue as a diagnostic character of Insulivitrina (Figs 111-113). The
penis is bipartite with a somewhat globular proximal and a tubiform distal part. Internally, the
proximal part harbours an oviform penial gland, its tissue also expands to the surface of the penis
at the boundary with the VD. The penial gland is not connected to the pilaster. There is only one
pilaster present, although it is deeply cleft distally and proximally. Transverse lamellae are very
weak, warts and similar structures are missing completely. In the author’s opinion, the term “penial
papilla” as used by IBAÑEZ et al. (1987) is not completely correct, because a papilla is a structure
serving as an outlet for a duct, which is not the case here. It is very likely that this “papilla peneana”
represents the uppermost pilastral part plus glandula since it is situated in the same place as it is in
Arabivitrina. In the female part, the construction of the vaginal papilla is different. The papilla is
elongate (short conical with central perforation in Arabivitrina) and deeply cleft, leaving two lobes.
The inner surface of these lobes displays a reticulate structure like honey-combs. Obviously, this
papilla is everted and defolded during copulation forming the “semilunar fold” as illustrated by
IBAÑEZ et al. (1987: 129, fig. 11). For all the other species described in this genus, the inner
structure of the penis and vagina has not been investigated sufficiently (cf. VALIDO et al. 1993).
The morphology of the genital organs indicates that the generic definition of Insulivitrina needs to
be re-evaluated. It possibly consists of several lineages (cf. length of atrium, digitiform “appendix”
in I. machadoi, MRP cone in I. mascaensis etc.), deserving a phylogenetically focused investigation.
A revision of this group, discussing the anatomical features of Insulivitrina and related taxa, is in
preparation (Ibañez, pers. comm.).
For anatomical research, the author received two specimens of Vitrinidae from K. Groh
labelled as Madeirovitrina nitidus (no type material). The differences from the figures of M. nitidus
as given by GROH & HEMMEN (1986: 189, fig. 2A) are striking. It is probable that the specimen
investigated represents a new species or genus. As it is beyond the scope of this publication, the
problem is postponed, and the taxon is dealt with here as cf. Madeirovitrina sp. In this species, the
penis is bipartite, the VD inserts at the base of the proximal penial part (Figs 114-116). This part
is filled by a heavily folded pilaster which is completely covered by the penial gland tissue (partly
removed in Fig. 115 to show pilaster). The distal part of the penis is also characterised by thick
pilasters fusing basally with the single atrial pilaster. Their surface as well as the inner surface of the
penial wall is covered by a minute honey-comb sculpture (indicated by longitudinal stripes in
Fig. 115). This structure can also be seen without opening the distal part of the penis. The vagina
is short, the glandula amatoria enlarged and covered by glandular tissue. Internally, a conical and
curved vaginal papilla is present with a central perforation. Distal to the papilla is a conical
(vaginal) organ which is striated by transverse lamellae. Only in Madeirovitrina behnii (Lowe,
1851) does an elongate pilaster seem to be present, sculptured with small warts in transverse rows
(for discussion of this species see GROH & HEMMEN 1986: 196). The results from ODHNER (1937)
differ from what is described for M. nitidus and still have to be reconfirmed.
The inner structures of the genital organs of Guerrina Odhner, 1954 are still to be described.
Apart from its shell, this genus seems to be well characterised by its short glandula amatoria and
the small-lobed vaginal gland (IBAÑEZ et al. 1987, VALIDO et al. 1993). In Gallandia Bourguignat,
1880, a glandular roll is present, fusing with two pilasters originating in the vagina (HAUSDORF
1995: 67, 69).
From the data given here it is evident that Arabivitrina is close to Insulivitrina, not only in
conchological characters but also in the structure of their genital organs. Both genera display
elongate major penial pilasters subdivided into at least three functional parts. The lamellate part of
the major penial pilaster and the penial warts as well as the structure of the vaginal papilla may be

Terrestrial and freshwater molluscs of the Arabian Peninsula 393
used as intergenerically differentiating characters. Both differ from cf. Madeirovitrina sp., as this
taxon shows a single multi-lobed penial pilaster with a honey-comb structure and a lamellate
vaginal cone. An epiphallus, as mentioned by GROH & HEMMEN (1986: 186) for Insulivitrina, is
not present in the family Vitrinidae as the sperm generally is not transferred in a spermatophore.
All species dissected have a penial gland which often expands to the outer surface of the penis. At
present it is not possible to assess the reliability of the characters discussed. A reconstruction of the
phylogenetic relationships within the Phenacolimax stock has to be postponed until all species of
this group are investigated. Nevertheless, this compilation shows the importance of the character
set “inner penial and vaginal structure”. As shown by FORCART (1949) for Phenacolimax, vagina
and penis evaginate completely during copulation. The function of vaginal and penial glands are
unknown but it is evident that the inner part of the latter is also exposed to the sexual partner.
Thus, most of the structures formerly inside the organs are presented to the partner and certainly
play an important role in partner-partner recognition. Missing or differing structures thus produce
and maintain reproductive barriers, so knowledge of them is absolutely necessary for both
taxonomic and phylogenetic analysis.
Plates 1-4: 1: Gulella schweinfurthi, specimen from Yemen, Bani al-Harith, Wadi Dhar. Photo H. Dekker. 2: Arabivitrina jansseni
n. sp., specimen from Asir mountain region, S of Bani Sa’ad S of Taif. Photo E. Neubert. 3: Cerastus schweinfurthi schweinfurthi,
specimen from Yemen, Bani al-Harith, Wadi Dhar. Photo H. Dekker. 4: Araboxychilus sabaeus, specimen from Yemen, Bani al-
Harith, Wadi Dhar. Photo H. Dekker.

394 E. NEUBERT
Family Buliminidae
Genus Mordania Bank & Neubert, 1998
1998 Mordania Bank & Neubert. — Basteria 61 (4/6): 80.
Diagnosis: Medium-sized pupiform shells with a rounded and expanded aperture. A well-
developed penial papilla is present. The penis is bipartite with an elongate proximal part. The
penial caecum is located close to the insertion of the epiphallus. The lumen of the epiphallus is
septated. The epiphallus passes without a flagellum into the VD.
Type species: Bulimus omanensis E.A. Smith, 1894.
No te : The taxon Bulimus omanensis E.A. Smith, 1894 was transferred to Mastus by MORDAN
(1986: 210). This generic assignment is incorrect for several reasons, as discussed by BANK &
NEUBERT (1998). There is no Mastus species present on the Arabian Peninsula, although it might be
possible that species of this genus occur in the unexplored northern areas close to Jordan or Syria.
Mordania omanensis (E.A. Smith, 1894) Figs 117-118
1894 Bulimus omanensis E.A. Smith. — Proc. Malac. Soc. London 1: 141, textfig. 1 (Green mountain, Oman).
Material: Oman: Oasis of Buraimi, I.1993, H. Pauscher, NEUB (1); Wadi Manqal, effluent of Wadi Hilm (Qalhat),
near Kibdiyah (= Kebdah), 22°41'N 59°18'E, 26.IV.1978, A.B. Paltrinieri, ONHM 610.27 (1); Jabal Misfeh, various sites,
23°13'N 57°07'E, 14.III.1986, A.B. Paltrinieri, ONHM 610.12 (3); Jabal Akhdar, Wadi al-Yemen, lower dam, 23°04'N
57°41'E, V.1980, A.B. Paltrinieri, ONHM 610.9 (3).
Type material: Lectotype Bulimus omanensis, BM(NH) 1894.3.22.5, four paralectotypes
BM(NH) 1894.3.22.6-9.
Description: The turreted shell is broad conical and white. The broad and blunt proto-
conch consists of 2.5 whorls with an extremely fine granulated surface. The teleoconch whorls are
flat with a very shallow suture. The surface of the whorls is glossy with a sculpture of wavy spiral
lines (magnification 50 ×). The only axial sculptural elements visible are fine and very oblique
growth lines.
The aperture is dominant, its height reaches more than half of the total shell height. The
peristome is broadly reflected but not recurved. Its rims are connected by a faint callus, a weak
angularis is present. The shell is slit-like umbilicated with a funnel-shaped periomphalum.
Measurements: Lectotype (Fig. 117): H = 23.9; D = 12.2. Specimen from Buraimi
(Fig. 118): H = 17.0; D = 9.7; PH = 9.2; PD = 7.4; W = 6.5. The shell diameter is measured
including the reflected peristome. The diameter behind the peristome is 8.2 mm.
Genus Pseudonapaeus Westerlund, 1887
1887 Pseudonapaeus Westerlund. — Fauna der in der paläarctischen Region lebenden Binnenconchylien III: 66.
1887 Subzebrinus Westerlund. — Fauna der in der paläarctischen Region lebenden Binnenconchylien III: 66.
Diagnosis: Shells pupiform to elongate, medium-sized, unicoloured brown or white, often
with brown oblique axial flames. Some species may be ribbed. For more information see MATEKIN
(1959), SCHILEYKO (1984), and UVALIEVA (1990).
Type species: Buliminus asiaticus v. Martens, 1881.
No te : The species Bulimus jousseaumi E.A. Smith, 1894 was transferred to Imparietula
Lindholm, 1925, by MORDAN (1986: 216). This genus is restricted to the eastern parts of Turkey
and some areas in neighbouring countries. For the use of the generic name Pseudonapaeus see BANK
& NEUBERT 1998.

Terrestrial and freshwater molluscs of the Arabian Peninsula 395
Pseudonapaeus jousseaumi (E.A. Smith, 1894) Fig. 119
1894 Bulimus jousseaumi E.A. Smith. — Proc. Malac. Soc. London 1: 142, textfig. 2 (Oman).
1905 Buliminus (Subzebrinus) Dautzenbergi Ancey. — J. Conchyl. 53: 262 [nom. nov. pro Bulimus jousseaumi E.A. Smith,
1894 non Ovella jousseaumei Jousseaume, 1890 (= Buliminus jousseaumi of authors)].
Material: Oman: Fanja, W of Muscat, I.1993, H. Pauscher, NEUB (7); Jabal Akhdar, Kahf Hoti (Hoti carstic cave
system), 990 m, 23°06'14"N 57°21'55"E (= N of Tanuf, E of al-Hambra), (many fragments); Wadi Bani Khalid, ad-Dawa,
23°36'N 59°05'E, 17.IV.1975, A.B. Paltrinieri, ONHM 610.13 (1); Jabal Akhdar, Wadi al-Yemen, lower dam, 23°04'N
57°41'E, V.1980, A.B. Paltrinieri, ONHM 610.9 (2); Wadi Selma, in Wadi Bani Awf (= Salmah), 23°12'N 57°22'E, 2.V.1978,
A.B. Paltrinieri, ONHM 610.50 (1).
Type material: Lectotype Bulimus jousseaumi, BM(NH) 1894.3.22.10, six paralectotypes
BM(NH) 1894.3.22.11 and 1900.6.8.69-73.
De s cri p tio n : The small turreted shells are broadly fusiform to conical. The protoconch is
very blunt and consists of two extremely fine granulated whorls. The basic colour of the teleoconch
is white with axial corneous streaks of the same width as the remaining white ones causing a
“zebra” pattern. Irregularly arranged blunt riblets indicating growth intervals may be observed.
The teleoconch whorls are glossy and flat. The suture is shallow. The oval aperture is some-
what thickened by a faint labial callus. The peristome is reflected and forms a narrow fringe. Its
rims are connected by a thick callus which ends by forming a prominent angularis. The umbilicus
is open and formed by an elongated slit, the periomphalum is funnel-shaped.
Measurements (lectotype Bulimus jousseaumi, Fig. 119): H = 12.1; D = 5.3; PH = 4.8;
PD = 3.9; W = 7.
Genus Paramastus Hesse, 1933
1933 Paramastus Hesse. — Arch. Naturgesch. 2 (2): 181.
Diagnosis: Subcylindrical to broadly fusiform turreted shells. Protoconch mammillate, the
initial teleoconch whorls are conical. The surface is characterised by a rugose axial sculpture with
spiral incisions. Peristome broadly flared, internally with a thickened lip, a small angularis is
present. The male copulatory organs with or without a penial appendix, a penial caecum and a
flagellum is always present.
Type species: Buliminus episomus Bourguignat, 1857 sensu HESSE 1933 (= Paramastus
cyprius Zilch, 1959).
Note: Paramastus consists of two lineages which are usually considered as subgenera, i.e.
Paramastus s. str. with a penial appendix and Cyrenaeus Heller, 1971 without a penial appendix.
Both Arabian taxa have never been collected alive, thus an assignment to one of the subgenera is
impossible. The author agrees with MORDAN (1986: 222), placing them provisionally in Para-
mastus because of the similarity of conchological characters.
Paramastus sabaeanus (Bourguignat, 1876) Fig. 120
1876 Buliminus sabaeanus Bourguignat. — Species novissimae molluscorum in Europaeo systemati detectae, notis
diagnosticis succinctis breviter descriptae. Premiére centurie: 19 (“Sabéens, pres de Mareb”).
Type material: Lectotype Buliminus sabaeanus, MHNG, one paralectotype.
Description: The shell is elongate pupiform. The smooth mammillate protoconch consists
of two whorls. Upper teleoconch whorls with a shallow suture. The aperture is ovoid with a faint
lip inside. The peristome is somewhat flared, the lip inconspicuous. The sculpture of the teleo-
conch consists of oblique axial striae. The umbilicus is closed.
Remarks: It is difficult to find conchological differences between the Arabian species and P.
episomus (Fig. 121). The latter species is ventricose, the upper whorls are flatter and the angularis

396 E. NEUBERT

Terrestrial and freshwater molluscs of the Arabian Peninsula 397
is stronger. In P. sabaeanus, the spiral sculpture is missing. Bearing in mind the conchological
variability of P. episomus, it is an act of caution to keep the Arabian species separate. There is a gap
of nearly 1000 km between the Palestinian and the Arabian distribution areas. It is not clear
whether this gap really exists or represents the absence of records, and thus it is not known whether
the Arabian populations are isolated from the Mediterranean ones or not. Moreover, the fact that
only two specimens are hitherto known of P. sabaeanus and one of P. hedjazicus gives rise to doubts
about the autochthonous origin of these taxa. The author collected intensively in the mountainous
area south of Taif, the type area of P. hedjazicus, and failed to find any specimens. For reasons of
comparison, figures of the type specimens of all three taxa are given here.
Measurements: Lectotype Bulimus sabaeanus (Fig. 120): H = 23.10; D = 10.50; PH = 9.60;
PD = 7.00; W = 8. Holotype (?) Paramastus episomus (Fig. 121): H = 17.45; D = 8.75; PH = 7.35;
PD = 5.40; W = 7.25.
Paramastus hedjazicus (Bourguignat, 1882) Fig. 122
1882 Buliminus hedjazicus Bourguignat. — Mission de G. Révoil au pays Çomalis. Faune et Flore des Pays Çomalis: 24,
pl. 1, fig. 12 (“Montagnes entre Djeddah et la Mecque”).
Type material: Holotype Buliminus hedjazicus, MHNG.
Description: The shell is small elongate turreted. The protoconch has the same structure
as in other species. There is a faint spiral sculpture on the top whorls, the axial elements are faint.
The aperture is narrow, a weak columellaris is present.
Remarks: For general remarks see under P. sabaeanus. The taxon P. hedjazicus is kept separate
from P. episomus until more material becomes available to supply sufficient information to
determine whether there are two distinct Arabian species.
Measurements (holotype, Fig. 122): H = 15.25; D = 6.45; PH = 6.00; PD = 4.35; W = 7.5.
Family Cerastidae
Genus Cerastus Albers, 1860
1860 Cerastus Albers. — Die Heliceen nach natürlicher Verwandtschaft systematisch geordnet, 2. Aufl.: 232.
Description: The medium to large-sized shells are elongate conical, the lip is undifferen-
tiated or weakly flared. The penis has a short, pointed caecum (see comments on this detail in C.
schweinfurthi). The MRP inserts on the epiphallus. The penial appendix has a moderately deve-
loped papilla. The MRA inserts basally at the central stalk, both muscles unite prior to attachment
to lung wall.
Type species: Bulimus distans L. Pfeiffer, 1856.
Figs 117-132: (original size × 2), all in frontal view. 117-118: Mordania omanensis. 117: Lectotype of Bulimus omanensis,
BM(NH) 1894.3.22.521. 118: Specimen from Oman, Oasis of Buraimi. 119: Pseudonapaeus jousseaumi, lectotype of Bulimus
jousseaumi, BM(NH) 1894.3.22.10. 120: Paramastus sabaeanus, lectotype of Buliminus sabaeanus, MHNG. 121: Paramastus
episomus, holotype (?) of Buliminus episomus, MHNG. 122: Paramastus hedjazicus, holotype of Buliminus hedjazicus, MHNG.
123-125: Cerastus schweinfurthi schweinfurthi. 123: Syntype of Buliminus schweinfurthi, SMF 7403 a. 124: Same, enlarged
protoconch and upper teleoconch whorls. 125: Holotype of Buliminus (Cerastus) schweinfurthi var. menahensis, SMF 7403 b.
126-127: Cerastus schweinfurthi brunneus n. ssp. 126: Holotype SMF 311347/1. 127: Same, enlarged protoconch and upper
teleoconch whorls. 128-129: Cerastus schweinfurthi apicostatus n. ssp. 128: Holotype SMF 311349 a. 129: Same, enlarged
protoconch and upper teleoconch whorls. 130: Euryptyxis labiosa, syntype of Buliminus (Petraeus) Pilsbryanus, NMW 1955.158.
131: Achatinelloides sebasmia, syntype MNHNP. 132: Achatinelloides jousseaumei, syntype MNHNP.

398 E. NEUBERT
Cerastus schweinfurthi schweinfurthi (v. Martens, 1895) Figs 123-125, 133-135, Plate 3
1895 Buliminus schweinfurthi v. Martens. — Sitzungsber. Ges. naturf. Fr. Berlin 1895 (6): 129 (“Menaha, im südlichen
Arabien, 7000 Fuß”).
1895 Buliminus schweinfurthi var. gracilior v. Martens. — Sitzungsber. Ges. naturf. Fr. Berlin 1895 (6): 129 (ditto).
1902 Buliminus (Cerastus) schweinfurthi var. menahensis Kobelt. — In: Martini & Chemnitz: Conchylien-Cabinet,
2. Auflage, I 13: 894, pl. 127, figs 22-23 (Menaha).
1941 Cerastus schweinfurthi var. maxima Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 26
(Jabal Harir, 7500 ft).
Material: Yemen: Bani al-Harith, Wadi Dhar (= Zahr), Qaryat al-Qabil, NE of Sana’a, 2200 m, between leaves, wood
and soil, 8.IV.1993, H. Dekker & F. Ceuninck v. Capelle, SMF 311619/1 a, SMF 311619/1 b (preserved), DEKK (8).
Type material: Syntypes Buliminus schweinfurthi, SMF 7403 a-d; holotype Buliminus
(Cerastus) schweinfurthi var. menahensis, SMF 7403 b/1.
Description: The big turreted shell is broad conical. In fresh specimens, the colour is
brown; in faded specimens it is light corneous.
The protoconch consists of two smooth whorls with a moderately deep suture. The teleoconch
whorls are slightly rounded with a shallow suture. The upper teleoconch whorls are densely ribbed
(Fig. 124). The ribs become weaker towards the aperture. They are crossed by deep spiral furrows,
thus producing a pattern of elongated beads on the last whorl.
The aperture is elongate oval, the peristome is slightly thickened to form a white lip. On the
columellar side, the peristome is reflected over the narrow and slit-like umbilicus producing a small
white triangular area.
Measurements: Illustrated syntype (Figs 123-124): H = 30.8; D = 17.9; PH = 17.8;
PD = 12.1; W = 7. Holotype of Buliminus (Cerastus) schweinfurthi var. menahensis (Fig. 125):
H = 29.6; D = 15.8; PH = 15.2; PD = 10.2; W = 7.5.
Genital morphology (SMF 311619/1 b, Figs 133-135): The distal penial tube is simple
and cylindrically shaped. The proximal part is wrapped in a muscular penis sheath. Proximal to
the sheath, there is a structure which, prior to now, was always interpreted as a caecum (HESSE
1933: 220, fig. 43; MORDAN 1986: 227, figs 15A, B). In fact, this “caecum” consists of a penial
and epiphallial part tightened by very dense connective tissue. This was proven by opening the
penis which contains coarse erected papillae at that part. The caecum is formed by a very small
vesical inflation of the penis. Here, the papillae are small and bear no sculptures.
The epiphallus is subdivided into a distal tube and a swollen proximal part. Its lumen is filled
with two main pilasters with interdigitating pits. The pilasters are spirally coiled. In the holotype
specimen of C. s. apicostatus n. ssp., a twice-coiled autospermatophore was present filling the space
between the pilasters. It matches exactly the figures given by HESSE (loc. cit.: fig. 43E). Towards
the VD, its diameter decreases continuously, the exact borders of both organs are not visible.
The penial appendix is more than three times the penis length. It is subdivided into three
parts. The first stretches distally from the branching point of the penis distad to the insertion of
the MRA. It is followed by a short and narrow, but thick-walled, part. The third part is thin-walled
and somewhat bulbous elongate. The inner walls are finely septate (not as coarse as indicated by
HESSE [loc. cit.: fig. 43C]). The appendicular papilla is simple and projects into the basal lumen of
the appendicula.
The retractor muscle is split into the MRA and the MRP. Both arms unite before attaching at
the diaphragm. The MRP inserts at the proximal third of the epiphallus, the MRA basally to the
medium part of the appendix.
The vagina is a stout voluminous structure enveloped by a soft tissue. Internally, it is com-
pletely smooth and has no pilasters. The pedunculus is short and opens after an inconspicuous
constriction into the BC. The inner wall of the pedunculus is sculptured by thick and short ob-

Terrestrial and freshwater molluscs of the Arabian Peninsula 399
Figs 133-135: Genital organs of Cerastus schweinfurthi schweinfurthi. 133: Genital apparatus. 134: Penis/epiphallus and penial
caecum. 135: Penial appendix.
liquely arranged pilasters. The pedunculus of the holotype specimen of C. s. apicostatus n. ssp.
harboured an autospermatophore which was folded into its lumen. Nevertheless, the spermato-
phore was still coiled. As shown by HESSE (loc. cit.), it is sculptured by a single row of digitated
scales which exactly match the pilasters of the pedunculus wall. Obviously, they act as an anchoring
structure for the spermatophore.
Remarks: The specimens of the type lot are quite similar in shape, mode of ribbing and
shape of the whorls. The nominate subspecies inhabits the southern part of Yemen.
Cerastus schweinfurthi brunneus n. ssp. Figs 126-127
Type material and locus typicus: holotype SMF 311347/1, paratypes SNMNH-MO 84/3, SMF 311348/3,
Saudi Arabia, Asir mountain region, Raydah escarpment, 2050 m, 25.III.1994, W. Schneider & F. Krupp. The type lot consists
of three adults and four apical fragments. — Non-type material: Saudi Arabia: Asir mountain region, Raydah
escarpment, 1850 m, 25.III.1994, W. Schneider & F. Krupp, SNMNH-MO 85/1, SMF 311616/1; Raydah escarpment,
2350 m, 29.I.1994, P. Symens & M. Werner, SNMNH-MO 86/3, SMF 311617/3; Raydah escarpment, 2100 m, 10.VII.1995,
S. Newton, SNMNH-MO 87/2, SMF 311618/1.
Diagnosis: A subspecies of C. schweinfurthi differing by its big elongate shell of dark
chestnut colour.
Description: The shell is elongate conical. The protoconch and the initial whorls of the
teleoconch are similar to that found in the nominal subspecies. The spiral sculpture is somewhat
denser and finer. The aperture is elongate, the columellar peristomial rim is excavated and
completely attached to the shell, thus closing the umbilicus.
Measurements (holotype, Figs 126-127): H = 41.8; D = 21.0; PH: 20.5; PD = 14.3;
W = 7.5.

400 E. NEUBERT
Etymology: This new subspecies is called C. s. brunneus because of its brown-coloured shell.
Remarks: This subspecies is less ventricose than the nominal subspecies and displays a
concave columella. These characters are stable in the few adult specimens available.
Cerastus schweinfurthi apicostatus n. ssp. Figs 128-129
Type material and locus typicus: holotype SMF 311349 a, dissected animal of holotype SMF 311349 b; three
paratypes in coll. DEKK, Yemen, al-Ashmur, between Amran and Hajjah, below and on limestone, 2700 m, 9.IV.1993, H.
Dekker & F. Ceuninck v. Capelle. The type lot consists of three adults (one collected alive) and one basal fragment. — No n -
type material: Yemen: between Amran and Hajjah, 27.III.1988, A. Liebegott, LIEB (2).
Diagnosis: A subspecies of C. schweinfurthi, which differs in the shape of its shell and the
sculpture of the upper whorls.
Description: This subspecies is much more slender than all forms known from the
nominate subspecies. It differs considerably in the sculpture of the upper teleoconch whorls. The
surface is covered by coarse, widely spaced ribs. In the interspaces, smaller ribs are nearly lacking.
The holotype specimen was collected alive and thus could be investigated anatomically. It
showed no difference to the nominal subspecies in the structure or length ratios of the genitalia.
Measurements (holotype, Figs 128-129): H = 35.9; D = 17.7; PH = 18.4; PD = 11.4;
W = 6.5.
Etymology: The name C. s. apicostatus is composed of the Latin words apex (= top) and
costa (= rib).
Remarks: Cerastus schweinfurthi apicostatus n. ssp. is known only from two localities in small
numbers, but the differences described seem to be stable. The material from coll. Liebegott can be
considered to be topotypic, since it is very likely that it originates from nearly the same locality as
the type lot does. Specimens from both lots are identical.
Cerastus scotti Connolly, 1941
1941 Cerastus scotti Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 26, pl. 3, fig. 14, textfig. 7
(Jabal Jihaf, 7100 ft.; Jabal Harir, 7500 ft).
Description (no material available, description according to MORDAN 1986: 228): The
shell is elongate ovoid. The fine ribs are initially strong and regularly spaced, becoming much
weaker and less regular [on the last whorls]. Weak spiral striae give a glossy appearance. The apex
is sharp, the protoconch initially smooth. The sutures are not deeply impressed. The umbilicus is
narrow and shallow, the aperture sharp and thin. The shell is colourless to very pale brown and
semi-transparent. The stronger early ribs may be translucent white.
Measurements (lectotype): H = 19.3; D = 10.6; PH = 9.6; PD = 6.2; W = 7.
Remarks: This species differs from C. schweinfurthi in its smaller size for an equivalent
number of whorls. This is also true for C. schweinfurthi apicostatus n. ssp. which is taller and seems
to have a thicker shell, although it also displays strong apical ribs which disappear later. It occurs
in the southernmost parts of the western Arabian mountain chains in Yemen.
Cerastus girwanensis Connolly, 1941
1941 Cerastus girwanensis Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 27, pl. 3, fig. 13 (Jabal
Girwan, near Ghaiman, 9 miles SE of Sana’a, 8700 ft).
Material: Saudi Arabia: Asir mountain region, Wadi al-Sharan, S of Bani Sa’ad, 1750 m, 2.VI.1995, E. Neubert &
R. Janssen, SNMNH-MO 88/1, SMF 311620/1.
Description: The turreted shell is broadly conical and unicoloured brown in fresh speci-
mens. The protoconch whorls are small and somewhat compressed apically with a very deep
suture. It is sculptured by axial wrinkles which subsequently become coarser to form the teleo-

Terrestrial and freshwater molluscs of the Arabian Peninsula 401
conch ribs. The teleoconch whorls are rounded with a suture of medium depth. The white and
sometimes translucent ribs are coarse and evenly spaced on the whole shell.
The oval aperture has a sharp rim, an internal lip is missing. The columellaris is somewhat
weaker compared to that of C. albonotatus. The columellar rim of the peristome is reflected over
the umbilicus which is rounded and opened.
Remarks: See Cerastus albonotatus.
Cerastus albonotatus Verdcourt, 1974
1974 Cerastua albonotata Verdcourt. — J. Conchyl. 111: 5, figs 2 a-c (Hillside 5 miles out of Ta’izz on airport road).
Material: Yemen: Jabal Sabar close to Ta’izz, 23.III.1988, A. Liebegott, LIEB (2); same locality, XII.1989, Rabin, LIEB (2).
Description: The medium-sized conical shell is brown with characteristically axial white
flames and dashes. The minute protoconch is finely ribbed axially. These ribs continuously enlarge
during growth of the whorls and constitute the teleoconch ribs. These are oblique and sharp,
irregularly spaced and more dense on the last whorl, particularly on the cervix. The suture is of
medium depth.
The oval aperture has a sharp and simple peristome. The columella bears an obliquely truncate
lamella. The columellar peristomial rim is somewhat reflected over the open umbilicus.
Measurements (holotype, according to MORDAN 1986): H = 18.2; D = 8.8; PH = 7.7;
PD = 5.5; W = 9.
Remarks: This species seems to be endemic to the area around Ta’izz, Yemen. As MORDAN
(1986: 228) outlines, it is close to C. girwanensis from which it differs in coloration, mode of
ribbing and its slightly narrower umbilicus. It is questionable whether it should be ranked as a
species or simply constitutes a local subspecies of C. girwanensis.
Genus Polychordia Connolly, 1941
1941 Polychordia Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 28.
Diagnosis: Slender elongate shells with deep sutures. The ribs are thin, raised, regularly and
widely spaced.
Type species: Polychordia pulcherrima Connolly, 1941.
Polychordia pulcherrima Connolly, 1941
1941 Polychordia pulcherrima Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 28, pl. 3, fig. 10
(Wadi Tahabad, north slope of Jabal Sabar, near Ta’izz, 5900 ft).
Description: The small shell (holotype measurements 13.8/5.2) is elongate conical. The
teleoconch whorls are somewhat shouldered in profile. The sculpture consists of very prominent
and sharp ribs which are regularly arranged and widely spaced. The shell is narrowly umbilicated,
the aperture has a sharp peristome without any labial callosities. The basic colour is pale brown
with the ribs white opaque.
Remarks: The author agrees completely with MORDAN (1986: 229), who stresses the
conchological similarity of P. pulcherrima with Cerastus girwanensis. The shell of C. albonotatus is
also close in conchological characters and distribution patterns. The specimens of C. albonotatus
studied display less intensive axial streaks and flames than the holotype (MORDAN 1986: fig. 13 e)
and only differ from P. pulcherrima in their mode of ribbing. This observation supports the
suggestion that the species actually belongs to Cerastus and that, following anatomical investiga-
tion, Polychordia will turn out to be a synonym of this genus.

402 E. NEUBERT
Genus Euryptyxis Fischer, 1883
1883 Euryptyxis Fischer. — Manual de Conchyliologie: 479.
Diagnosis: Medium to large-sized pupiform shells with an oblique aperture and a reflected
lip. Internally, a columellar fold may be present and visible on the columella. The penial caecum is
short and blunt, the penial appendix has a short, blunt papilla. The appendicular retractor inserts
at the junction of the basal and central portions of the appendix.
Type species: Pupa candida Lamarck, 1822.
Euryptyxis candida (Lamarck, 1822)
1775 ? Helix arabica Forskål. — Descriptiones animalium: 127 [nom. dub.] (“Lohajae”).
1822 Pupa candida Lamarck. — Anim. sans vertèbres II: 106.
1837 Buliminus forskalii Beck. — Index molluscorum: 68.
1843 Buliminus arata Recluz. — Rev. Zool. (Soc. Cuv.) VI: 4 (Socotra).
1882 Buliminus micraulaxus Bourguignat. — Mission de G. Révoil au pays Çomalis. Faune et Flore des Pays Çomalis: 17,
fig. 20 (“Sud de l’Arabie”).
1887 Buliminus eryx Westerlund. — Fauna der in der paläarctischen Region lebenden Binnenconchylien III: 64
(“Arabien”).
Material: Saudi Arabia: Asir mountain region, Bukairat al-Hakimah, storage lake of Wadi Jizan, E of Jizan,
28.IV.1994, W. Schneider & F. Krupp, SNMNH-MO 90/12, SMF 311622/12, HLMD (3); Asir mountain region, Wadi Jawah,
SE of al-Ahrida, 29.IV.1994, W. Schneider & F. Krupp, SNMNH-MO 91/4, SMF 311623/3; Asir mountain region, Wadi Juwa,
1.II.1994, P. Symens & M. Werner, SNMNH-MO 92/7, SMF 311624/7, NEUB (1). — Yemen: 10 rkm W of Menaha,
I.1990, S. Neubert, NEUB (2); Rihab, 21.II.1990, M. Scholz, NEUB (1); between Amran and Hajjah, 27.III.1988, A.
Liebegott, LIEB (2); at the road from al-Hodeida to Menaha, foothills of the escarpment, 800 m, 25.III.1988, A. Liebegott,
LIEB (2); at the road from al-Hodeida to Menaha, escarpment, 1200 m, 25.III.1988, A. Liebegott, LIEB (1); “Northern part of
Yemen”, XII.1989, Rabin, LIEB (3); Midi, Oreste Point, 16°22'N 42°46'E, sand beach, rocks, 24.IV.1993, H. Dekker & F.
Ceuninck v. Capelle, DEKK (1); Sharas, Wadi Sharas, NE of Hajjah, 800 m, old river sediments, 9.IV.1993, H. Dekker & F.
Ceuninck v. Capelle, DEKK (15); Red Sea, al-Murk Island, 15°38'N 42°38'E, beach, 11./12.IV.1993, H. Dekker & F. Ceuninck
v. Capelle, DEKK (1); Red Sea, al-Salif Peninsula, south coast, 15°11'N 42°49'E, sand beach, 13.IV.1993, H. Dekker & F.
Ceuninck v. Capelle, DEKK (1); Red Sea, Ukban Island, 15°31'N 42°22'E, sand beach and rocks, 12.IV.1993, H. Dekker & F.
Ceuninck v. Capelle, DEKK (12).
Description: The pupiform shell is stout with somewhat inflated median teleoconch
whorls. The surface of the teleoconch is covered by regular oblique and coarse axial ribs crossed by
fine spiral striae. The umbilicus is closed. The shell is opaque to brown-coloured.
The aperture has a flared, but never recurved, lip with a weak labial callus; the lip may be
deeply pigmented. Internally, there is a strong fold on the columella, which is not clearly visible
from the aperture.
Measurements (according to measurements taken from type material by MORDAN 1986:
230): H = 16.5-40.1; D = 10.0-19.9; PH = 9.8-18.5; PD = 6.2-14.0.
Remarks: Considerable variation in shell dimensions was also observed in the population
found at Wadi Juwa, where the dimensions varied between 26.2/14.2/14/11.1 and 17.8/9.5/8.2/6.6
in adult specimens.
Euryptyxis fragosa (L. Pfeiffer, 1842)
1821 Cochlogena fragosa Férussac. — Tabl. syst. Limaçon: 55, No. 421 [nomen nudum] (“Les grandes Indes?”).
1842 Bulimus fragosus L. Pfeiffer. — Symbolae ad Historiam Heliceorum II: 45 (Yemen, Arabia).
Material: Yemen: Dschibbla, between Sana’a and Ta’izz, 22.III.1988, A. Liebegott, LIEB (2); Jabal Sabar close to
Ta’izz, 23.III.1988, A. Liebegott, LIEB (16); same locality, XII.1989, Rabin, LIEB (1 adult, 3 juv.).
Description: The shell is elongate conical. The axial ribs are fine and regularly spaced on
the upper whorls becoming less evenly spaced and weaker on the lower whorls. A fine wavy spiral
sculpture is present.

Terrestrial and freshwater molluscs of the Arabian Peninsula 403
The aperture is elongate, straight to somewhat oblique. The peristome is flared but never
recurved. The columellaris is completely absent or very weak.
Measurements (specimen from Jabal Sabar close to Ta’izz): H = 27.7-35.7; D = 14.1-18.7;
PH = 13.3-18.0; PD = 9.2-12.0; W = 8-9 (smallest vs. tallest specimen, n = 16).
Euryptyxis labiosa (O.F. Müller, 1774) Fig. 130
1774 Helix labiosa O.F. Müller. — Vermium terrestrium et fluviatilium 2: 96 (“In India”).
1775 Helix arabica Forskål. — Descriptiones animalium: 127 (“Lohajae”).
1843 Pupa jehennei Recluz. — Rev. Zool. (Soc. Cuv.) VI: 4 (Socotra).
1882 ? Bulimus bruguieri Bourguignat. — Mission de G. Révoil au pays Çomalis. Faune et Flore des Pays Çomalis: 25
(“On ne connaît pas la patrie […] vraisemblablement de l’Yemen”).
1897 Bulimulus hypodon Pilsbry. — Nautilus 10: 102 (Lower California, exact locality unknown; err.!).
1900 Buliminus (Petraeus) Pilsbryanus Ancey. — Nautilus 14: 43 (Mountains of Yemen above Aden, Southern Arabia (Dr.
Jousseaume)).
Material: Saudi Arabia: Asir mountain region, al-Mekwah, Marble Village, 700 m, II.1994, P. Symens & M.
Werner, SNMNH-MO 93/2, SMF 311625/2; Asir mountain region, Jabal Qahar, 17°39'N 42°53'E, II.1996, S. Newton,
SNMNH-MO 94/2, SMF 311626/1. — Yemen: Dschibbla, between Sana’a and Ta’izz, 22.III.1988, A. Liebegott, LIEB (1);
al-Mahrah, between Damquat and al-Fatk, 16°32'24"N 52°46'00"E, sand beach, patches of rock, 4.X.1995, H. Dekker & F.
Ceuninck v. Capelle, DEKK (1); al-Mahrah, between al-Fatk and Damquat, 16°33'N 52°47'E, limestone rocks, 1.X.1995, H.
Dekker & F. Ceuninck v. Capelle, DEKK (6); al-Mahrah, Hawf, E side of village, 16°38'03"N 53°02'34"E, rocks with algae and
sand patches, 1.X.1995, H. Dekker & F. Ceuninck v. Capelle, DEKK (5).
Type material: Six syntypes Buliminus pilsbryanus, NMW 1955.158.
Description: The shell is elongate pupiform. The surface is covered by thin ribs, which are
closely set in the upper whorls. On the lower teleoconch whorls, the ribs become faint and the
interspace between the ribs is enlarged. In many specimens, the surface of the lower part of the
shell seems to be smooth. There is a fine and dense spiral sculpture consisting of small beaded
threads. The shell is coloured opaque white to brown with the aperture often more deeply
pigmented.
The aperture is somewhat oblique with a recurved peristome. A strong columellar tooth is
present in the aperture and can also be found internally on the columella.
Measurements: According to MORDAN (1986: 249, 39, specimens from four localities):
H = 25.0-28.0; D = 13.5-17.0; PH = 13.4-16.4; PD = 10.5-13.7. Illustrated syntype of Buliminus
pilsbryanus (Fig. 130): H = 21.8; D = 11.7; PH = 10.9; PD = 8.8; W = 7.5.
Remarks: This species differs from E. candida in its more elongate shape and the mode of
ribbing. The Melvill-Tomlin collection in Cardiff houses a single lot of Buliminus pilsbryanus
Ancey, 1900, containing six specimens from Aden. Investigation of this material revealed that it is
identical with E. labiosa. The specimens are shaped like the shells known from Yemen. On the last
whorls, the ribs become weak and disappear nearly completely, which is characteristic of this
species as described by ANCEY (1900: “Anfractus 7.25 convexiusculi […] superiores confertim
oblique costulati, sculptura in sequentibus debiliore, in inferioribus obsolescenti …”). Obviously,
the specimens were communicated to Ancey by Jousseaume, but this is not clear from the original
label since this only reads “Buliminus Pilsbryanus Ancey, Aden”. In the author’s opinion, the
designation of a lectotype should be postponed until the whereabouts of more original material is
traced (in coll. Jousseaume). However, the specimens match Ancey’s description so well that they
may be considered to be syntypes.
Euryptyxis latireflexa (Reeve, 1849)
1849 Bulimus latireflexus Reeve. — Conchologica Iconica 5: species 568, pl. LXXVIII, (Muscat?).
1882 Bulimus micraulax Bourguignat. — Mission de G. Révoil au pays Çomalis. Faune et Flore des Pays Çomalis: 17,
fig. 20 (“Sud de l’Arabie”).

404 E. NEUBERT
1894 Bulimus deflersi Jousseaume. — Bull. Soc. Philomath. Paris (3) 6 (3): 100 (Jabal el Areys).
1895 Buliminus lunti Melvill. — Proc. Malac. Soc. London 1: 224, pl. XIV, fig. 7 (Plateau 400 miles east of Aden,
Hadramaut).
1899 Petraeus socialis Jousseaume. — Le Naturaliste 2 (13): 8 (Schoukra).
1899 Petraeus schoukraensis Jousseaume. — Le Naturaliste 2 (13): 8 (Schoukra).
1903 Cerastus dinshawi Sykes. — Proc. Malac. Soc. London 5: 338 (Senna).
1925 Euryptyxis lunti var. makallensis Pallary. — In: Little, O.H.: The geography and geology of Makalla, South Arabia.
Geological survey of Egypt: 224, pl. XXXV, fig. 22 (Makalla and Jabal el Da’liya).
1925 Euryptyxis littlei Pallary. — In: Little, O.H.: The geography and geology of Makalla, South Arabia. Geological survey
of Egypt: 224, pl. XXXV, fig. 21 (Jabal el Da’liya, Jabal Mihta (Asfal el Ain), and Qam el Ghail).
1925 Euryptyxis littlei var. minor Pallary. — In: Little, O.H.: The geography and geology of Makalla, South Arabia.
Geological survey of Egypt: 224 (ditto).
1928 Euryptyxis leesi Pallary. — Proc. Malac. Soc. London 18: 41, pl. 1, figs a, b (Dhofar).
Material: Saudi Arabia: Asir mountain region, Raydah escarpment, 1850 m, 25.III.1994, W. Schneider & F.
Krupp, SNMNH-MO 95/1, SMF 311627/1. — Yem en : Mountains between Makalla and Wadi Hadramaut, IV.1988, A.
Liebegott, LIEB (2); al-Mahrah, between Damquat and Jahib, 16°35.5'N 52°53'E, limestone rocks, 1.X.1995, H. Dekker & F.
Ceuninck v. Capelle, DEKK (5). — Om a n : Dhofar, NE slope of Qara mountains, Jibjat area (I.a.e.n. KR ?1?), Girod ded.
1997, SMF 311678/4 (+ fragments), GIROD (2); same locality, (I.a.e.n. KR 117), Girod ded. 1997, SMF 311679/1; same
locality, (I.a.e.n. KR 143), Girod ded. 1997, SMF 311680/1; same locality, (I.a.e.n. KR 126B), Girod ded. 1997, SMF 311681/3;
same locality, (I.a.e.n. KR 31), Girod ded. 1997, SMF 311682/2 (+ fragments); same locality, (I.a.e.n. KR 127), Girod ded.
1997, SMF 311683 (only fragments); same locality, (I.a.e.n. KR 73), Girod ded. 1997, SMF 311694 (fragments); Dhofar, Wadi
Darbat, 17°06'N 54°27'E, 16.VI.1980, A.B. Paltrinieri; ONHM 610.29 (3); Dhofar, Ain Arzat (= Razat), 17°07'N 54°14'E,
15.VI.1978, A.B. Paltrinieri, ONHM 610.10 (1); Dhofar, Khwar Rowri, 17°03'N 54°25'E, 16.VI.1980, A.B. Paltrinieri,
ONHM 610.54 (2, + 4 fragments); Dhofar, Khwar Rowri, 17°03'N 54°25'E, 16. & 18.VI.1980, A.B. Paltrinieri, ONHM
610.53 (3).
Description: Apart from the aperture, the shell is similar to that of E. candida. The
shape varies considerably from elongate pupiform to inflated or subglobose. The surface sculp-
ture consists of dense axial ribs crossed by a fine spiral sculpture. The ribs become fainter on the
last whorl.
The aperture is elongated and, in typical specimens, characterised by the broadly reflected
peristome. The lip may be callose and is often tinted chocolate-brown. A columellaris can be seen
in the aperture; internally, a strong lamella can be observed on the columella.
Measurements (lectotype of Bulimus latireflexus, according to MORDAN 1986: 247):
H = 31.8; D = 15.1; PH = 14.9, PD = 11.1; W = 8.2.
Genus Achatinelloides Nevill, 1878
1878 Achatinelloides Nevill. — Handlist of Mollusca in the Indian Museum I: 131.
Diagnosis: Shells broad conical to fusiform. Aperture with a simple lip, the last whorl is
somewhat adpressed to the preceding whorl. MRP and MRA originate separately, the MRP inserts
at the epiphallus, the MRA at the top of the basal stalk of the penial appendix.
Type species: Bulimus socotorensis L. Pfeiffer, 1845.
Achatinelloides sebasmia (Jousseaume, 1889) Fig. 131
1889 Ovella sebasmia Jousseaume. — Bull. Soc. Malac. France 6: 350 (“environs d’Aden”).
Type material: Two syntypes Ovella sebasmia, coll. Jousseaume, MNHNP.
Description: The medium-sized shell is fusiform. It is white with brown axial flames which
sometimes form oblique zigzag patterns.
The protoconch consists of 2.5 smooth whorls. The teleoconch whorls are flat, the suture is
very shallow. The surface of the teleoconch is sculptured by blunt and densely packed ribs which
are arched subsuturally.

Terrestrial and freshwater molluscs of the Arabian Peninsula 405
The aperture is elongate elliptical with a weak lip internally. The peristome is simple but
somewhat thickened at the columellar insertion. A blunt angularis is present. The peristomial rims
are connected by a weak callus. The umbilicus is open and narrowly elongate. The last whorl is
somewhat compressed at the base forming a very blunt keel around the excavated periomphalum.
Measurements (illustrated syntype MNHNP, Fig. 131): H = 17.8; D = 8.8; PH = 9.2;
PD = 5.3; W = 7.
Remarks: see under Achatinelloides jousseaumei.
Achatinelloides jousseaumei (Jousseaume, 1890) Fig. 132
1890 Ovella jousseaumei Jousseaume. — Bull. Soc. Malac. France 7: 93, pl. 3, figs 7-8 (Mahala).
Type material: Syntype Ovella jousseaumei, coll. Bourguignat, MNHNP; six syntypes,
coll. Jousseaume, MNHNP.
Description: As already outlined by MORDAN (1986: 253) this taxon is conchologically
very close to A. sebasmia. It differs by its more inflated conical shell, the stronger colour patterns
and the somewhat stronger columellaris.
Remarks: MORDAN (1986) states that A. sebasmia is close to A. balfouri (Godwin-Austen,
1881) and A. jousseaumei to both A. dahamisensis (Smith, 1898) and A. homhilensis (Smith, 1898).
The author agrees completely with this opinion. Only a taxonomic revision of the endemic
Socotran malacofauna will clarify the status of the Arabian species which have very likely been
introduced to this area by humans.
Measurements (illustrated specimen, Fig. 132): H = 20.15; D = 12.10; PH = 11.05;
PD = 6.25; W = 7.5.
Genus Zebrinops Thiele, 1931
1931 Zebrinops Thiele. — Handbuch der systematischen Weichtierkunde II: 525.
Diagnosis: Turreted conical shells without callose lip. Penis elongate with a muscular sheath
and a well-developed caecum. The MRP inserts on the epiphallus, the MRA at the base of the
central stalk. Both arms originate separately. The appendix inserts on the penis close to the atrium
(cf. MORDAN 1986: 254-259, figs 36, 37; BOURGUIGNAT 1882: 100, fig. 13).
Type species: Limicolaria revoili Bourguignat, 1882.
Zebrinops albata (Férussac, 1827)
1827 Helix (Cochlogena) albata Férussac. — Bull. Sci. 10: 305 (“L’Arabie hereuse”).
1843 Bulimus bicinctus Recluz. — Rev. Zool. (Soc. Cuv.) VI: 4 (Socotra).
1858 Bulimus candidissimus L. Pfeiffer. — Malakozool. Bl. 5: 239 (Socotra).
1941 Zebrinops ventricosa Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 29, pl. 3, fig. 15,
textfig. 8 (Dhala).
Material: Yemen: Dschibbla, between Sana’a and Ta’izz, 22.III.1988, A. Liebegott, LIEB (3); Ta’izz, 23.III.1988, A.
Liebegott, LIEB (1); Ta’izz, 30 km NE of town, c. 2000 m, XII.1989, Rabin, LIEB (3).
Description: The white shell is turreted and broad conical in shape. In the early teleoconch
whorls, a brown spiral band starts suprasuturally. In the beginning, it consists of single dots, but
on the body whorl the dashes verge and form a compact band. A second spiral band may occur on
the last whorls. In addition, brown axial streaks and flames colour the last whorl. The coloration
may vary from pure white to the pattern described.
The protoconch consists of 2.5 smooth and evenly rounded whorls which are light corneous
in fresh shells. The teleoconch whorls are rounded with a suture of medium depth. The early

406 E. NEUBERT
Figs 136-141: (original size × 3), 136-138: Araboxychilus sabaeus, specimen from Asir mountain region, Raydah escarpment.
136: Frontal view. 137: Apical view. 138: Subsurface. 139-141: Oxychilus (Costoxychilus) profundus n. sp., holotype SMF 311269.
135: Frontal view. 136: Apical view. 137: Subsurface.
teleoconch whorls are covered by dense and regularly spaced fine ribs which later become less
prominent and are obsolete on the last whorls.
The aperture is oval elongate with a sharp and straight peristome. Internally, a weak callose lip
may be present. The columellar peristomial rim is straight and somewhat reflected over the open
umbilicus. The umbilicus is almost closed in juveniles.
Measurements (lectotype of Helix albata, according to MORDAN 1986: 255): H = 26.3;
D = 11.4; PH = 10.1; PD = 6.6; W = 8.9.
Family Zonitidae
Genus Araboxychilus Riedel, 1977
1977 Araboxychilus Riedel. — Ann. Zool. Warszawa 33: 509.
Diagnosis: Medium-sized shells with a conical spire. Juvenile whorls are sharply keeled, the
body whorl is bluntly angulated. The shell surface is covered by a dense reticulate pattern.
Type species: Trochomorpha sabaea v. Martens, 1889.

Terrestrial and freshwater molluscs of the Arabian Peninsula 407
Araboxychilus sabaeus (v. Martens, 1889) Figs 136-138, Plate 4
1889 Trochomorpha sabaea v. Martens. — Nachrbl. dtsch. malakozool. Ges. 21 (9/10): 146 (“Menaha, an berieselten
Felsen zwischen Moos”).
Material: Saudi Arabia: Asir mountain region, Abha escarpment, Juniperus forest, 2300 m, II.1994, P. Symens &
M. Werner, SNMNH-MO 96/1, SMF 311628/1; same data but 2500 m, SNMNH-MO 97/2, SMF 311629/2; Asir mountain
region, Raydah escarpment, 2350 m, 29.I.1994, P. Symens & M. Werner, SNMNH-MO 98/6; SMF 311630/6, NEUB (1);
same locality, 2050 m, 25.III.1994, W. Schneider & F. Krupp, SNMNH-MO 99/1; same locality, 2200 m, I.1995, S. Newton,
SMF 311631/1; same locality, 2100 m, 10.VII.1995, S. Newton, SNMNH-MO 100/1, SMF 311632/4; same data but 2300 m,
SNMNH-MO 101/1; Asir mountain region, King Khalid descent near Baha, 2215 m, 3.VI.1995, E. Neubert et al., SNMNH-
MO 102/4, SMF 311633/4, SMF 311641/1 (preserved). — Yemen: between Amran and Hajjah, 27.III.1988, A. Liebegott,
LIEB (1); Bani al-Harith, Wadi Dhar (= Zahr), Qaryat al-Qabil, NE of Sana’a, 2200 m, between leaves, wood and soil,
8.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (1).
Description: The fragile brown shells are medium-sized. The spire is subdepressed to conical.
In adult specimens, the body whorl is angulated whereas it is sharply keeled in juveniles. Sometimes,
these keeled whorls may be seen in injured adult shells when the peristome sinks under the periphery.
Usually, the upper rim of the peristome inserts exactly at the periphery, producing a shallow suture.
The subsurface of the shell is somewhat flattened. Compared to the diameter of the teleo-
conch, the diameter of the umbilicus is small. The body whorl is very broad, reaching twice the
width of the preceding one. The peristome is broadly oblique and sharp, its upper rim inserts
above the periphery. It is only slightly reflected over the umbilicus.
The surface of the shell is densely covered by minute elongated and comma-like angulated
beads arranged in numerous spiral rows. Under low magnification, this sculpture shows a reticulate
pattern and has a silky appearance to the naked eye. On the subsurface, the sculpture fades
completely leaving a smooth periumbilical region. Since this species is rare and not well known,
the measurements of a few adult specimens are given below.
The two preserved specimens were juveniles with immature genital organs.
Table 1. Measurements of five adult specimens of Araboxychilus sabaeus from Saudi Arabia, Raydah escarpment, 2350 m.
Height of
shell
Diameter of
shell
Height of
peristome
Diameter of
peristome
Number of
whorls
10.2 20.5 8.1 10.8 7.5
8.6 19.6 8.2 10.2 7
9.2 22.2 7.9 11.4 7
8 19.2 7.3 9.4 7
8.6 20.4 7.8 9.9 7
Genus Oxychilus Fitzinger, 1833
1833 Oxychilus Fitzinger. — Systematisches Verzeichnis der im Erzherzogthume Oesterreich vorkommenden Weichthiere: 100.
Diagnosis: Shells small to medium-sized, fragile, nearly flat to low conical. The umbilicus
may be narrow to wide, only a few species are without umbilicus. Usually, the shells are smooth
and glossy, some with fine spiral or reticulate patterns. For more detailed information refer to
RIEDEL (1980).
Type species: Helix cellaria O.F. Müller, 1774.
The genus Oxychilus contains more than 20 subgenera, which usually are identified precisely
by the anatomy of the reproductive organs. Unfortunately, no fertile specimens of the new species
were available. The outstanding conchological characters of this species make it necessary to
separate it at the subgeneric level from other congeneric taxa.

408 E. NEUBERT
Subgenus Costoxychilus n. subgen.
Description: A subgenus of Oxychilus with a narrowly coiled small shell. The last whorl is only
inconspicuously enlarged. The protoconch is sculptured by spiral rows of small wrinkles, the
teleoconch with fine and beaded axial riblets.
Type species (by present designation): Oxychilus (Costoxychilus) profundus n. sp.
Remarks: Costoxychilus n. subgen. is here tentatively placed in Oxychilus. Anatomical details
determine the taxonomic level and the phylogenetic relationships.
Oxychilus (Costoxychilus) profundus n. sp. Figs 139-141
Type material and locus typicus: holotype SMF 311269, paratypes SMF 311270/10, SNMNH-MO 103/10,
IZPAN (2), NEUB (2), Saudi Arabia, Asir mountain region, Raydah escarpment, 2350 m, 29.I.1994, P. Symens & M. Werner.
The type lot contained 25 specimens. — N on-type material: Saudi Arabia: Asir mountain region, Abha escarpment,
2500 m, II.1994, P. Symens & M. Werner, SNMNH-MO 104/5, SMF 311634/4; same locality, 2100 m, 10.VII.1995, S.
Newton, SMF 311635/1; same locality, 2310 m, 18°11'50.0"N 42°24'30.3"E, 5.V.1994, W. Schneider & F. Krupp, SNMNH-
MO 105/1, SMF 311651/1 (preserved).
Diagnosis: A small Oxychilus species with narrow whorls and a very dense beaded sculpture.
The umbilicus is nearly cylindrical and deep.
Description: The yellowish to brown shell is somewhat depressed with a flat to low conical
spire. The first part of the protoconch is smooth. The sculpture of the teleoconch starts within the
first protoconch whorl. It consists of fine and densely packed ribs which are intersected by up to
20 spiral rows. The ribs thus are composed of small comma-like beads. This structure may fade on
the body whorl. Although less prominent, it also covers the subsurface of the shell. The inner walls
of the umbilicus are smooth.
The narrowly coiled whorls are bordered by a suture of medium depth. They are bluntly
angulated with the upper rim of the peristome always attaching above the periphery. The peri-
stome is sharp, its diameter always exceeds the peristome’s height, thus producing an obliquely
crescent-shaped outline. The umbilicus is of medium width and nearly cylindrical and resembles a
bore hole (the diameter of the umbilicus does not increase substantially during growth of the shell).
Close to the type locality, one living specimen was collected. Unfortunately, the genital organs
of the juvenile had not developed yet. Even in its preserved state, the animal was deeply blue-
coloured with a yellowish foot sole. The foot was extremely narrow and elongate.
Measurements (holotype, Figs 139-141): H = 4.8; D = 10.0; PH = 3.5; PD = 4.7; W = 7.
Etymology: This species is called O. (C.) profundus for the deep umbilicus of the shell.
Affinities: Similar to Araboxychilus sabaeus, this species is isolated in the south-western area
of Arabia. Any affinities to the few Oxychilus species with ribbed or just sculptured shells are
superficial. Relationships to other Zonitidae can only be discussed when details of the genital
organs are investigated.
Family Ferussaciidae
Genus Cecilioides Férussac, 1814
1814 Cecilioides Férussac. — Mém. Géol.: 48.
Diagnosis: Minute needle-shaped shells with a blunt protoconch. The columella is arched
and obliquely truncate. The animals live subterraneously and have no eyes.
Type species: Buccinum acicula O.F. Müller, 1774.

Terrestrial and freshwater molluscs of the Arabian Peninsula 409
Cecilioides isseli (Paladilhe, 1872) Figs 142-143
1872 Caecilianella isseli Paladilhe. — Ann. Mus. Stor. Nat. Genova 3: 22, tav. 1, figs 9-10 (Aden).
Material: Yemen: Red Sea, ad-Durayhimi, 14°40'N 42°59'E, sand beach, 16.IV.1993, H. Dekker & F. Ceuninck v.
Capelle, SMF 3116395/5, DEKK (39).
Description: The blunt and broadened protoconch consists of 1.5-2.0 whorls. The
teleoconch whorls are smooth, gently rounded and somewhat constricted suprasuturally. The
suture is deep and bordered by a thin opaque line. The teleoconch whorls increase rapidly in size,
the body whorl is considerably elongated. In adult specimens, the body whorl (including the
aperture) reaches nearly twice the height of the spire (ratio 1:1.8-1:2.0).
Compared to other species, the aperture is short and broadens basally. The columella is
obliquely truncate. Parietal and columellar sides of the aperture are connected by a very thin
callus.
Remarks: This species is only known from its type area and the locality recorded here.
Measurements (illustrated specimen, Fig. 143): H = 4.30; D = 1.36, PH = 1.41; PD = 0.75,
W = 5.
Cecilioides tumulorum (Bourguignat, 1856) Fig. 144
1856 Caecilianella tumulorum Bourguignat. — Aménités Malacologiques I: 219 (Greece, Megara).
Material: Saudi Arabia: Asir mountain region, Wadi Juwa, 150 m, 1.II.1994, P. Symens & M. Werner, SNMNH-
MO 106/1, SMF 311636/1. — Yemen: Red Sea, ad-Durayhimi, 14°40'N 42°59'E, sand beach, 16.IV.1993, H. Dekker & F.
Ceuninck v. Capelle, DEKK (4). — Oma n : Dhofar, NE slope of Qara mountains, Jibjat area (I.a.e.n. ?1?), Girod ded. 1997,
SMF 311695/2; same locality, (I.a.e.n. KR 126B), Girod ded. 1997, SMF 311696/2; same locality, (I.a.e.n. KR 159), Girod
ded. 1997, SMF 311697/1.
Description: This species is characterised by its broadly ovate shell when compared to the
other species. The protoconch is dome-shaped and blunt. The teleoconch whorls are slightly
rounded with a shallow suture. The ratio of body whorl to spire ranges from 1:3 to 1:4. The
aperture is elongate and reaches twice the length of the spire. The parietal outline is gently curved,
the columella is very short and obliquely truncate.
Remarks: The Arabian shells presented here are only tentatively identified as C. tumulorum.
Fragments of shells which were found by the Italian archaeological excavation in Oman are here
also provisionally placed in C. tumulorum.
A plethora of names are created for shells belonging to this genus and a revision is pending.
Forms of Cecilioides with tumid whorls from the eastern Mediterranean and Caucasian area are
widespread in the collections and are usually identified as several different species. For this reason,
the identification of the Arabian specimens may turn out to be wrong. This is the first record of
this species from the Arabian Peninsula.
Measurements (illustrated specimen, Fig. 144): H = 5.05; D = 1.80, PH = 2.71; PD = 1.05,
W = 5.
Cecilioides acicula (O.F. Müller, 1774) Fig. 145
1774 Buccinum acicula O.F. Müller. — Vermium terrestrium et fluviatilium 2: 150 (Germany, Thangelstedt).
Material: Saudi Arabia: Asir mountain region, Abha escarpment, Juniperus forest, 2500 m, II.1994, P. Symens &
M. Werner, SNMNH-MO 107/2, SMF 311637/1; Asir mountain region, Raydah escarpment, 2350 m, 29.I.1994, P. Symens
& M. Werner, SNMNH-MO 108/1, SMF 311638/2. — Yemen: al-Mahrah, between Damquat and Jahib, 16°35.5'N
52°53'E, limestone rocks, 1.X.1995, H. Dekker & F. Ceuninck v. Capelle, DEKK (1).
Description: The broad protoconch is slightly bulbous and reaches or even exceeds the
diameter of the first teleoconch whorl. The teleoconch whorls are evenly rounded with a shallow
to medium deep suture, a border line is always absent. The body whorl is elongated (ratio of body

410 E. NEUBERT
Figs 142-148: (original size × 10), all in frontal view. 142-143: Cecilioides isseli. 142: Caecilianella isseli ex PALADILHE (1872:
tav. 1, fig. 10). 143: Specimen from Yemen, ad-Durayhimi. 144: Cecilioides aff. tumulorum, specimen from Yemen, ad-
Durayhimi. 145: Cecilioides acicula, specimen from Asir mountain region, Abha escarpment. 146-148: Coilostele paladilhiana.
146: Francesia scalaris ex PALADILHE (1872: tav. 1, figs 1-2, 4). 147: Lectotype of Coelostele isseli, MHNG. 148: Holotype of
Coelestele aegyptiaca, MHNG.
whorl to spire ranges from 1:1.5 to 1:2). The aperture is oval and narrow at the base. The
columella is short and obliquely truncate.
Remarks: This widespread European species differs from C. isseli in the size of its teleoconch
whorls, its smaller protoconch and the relatively shorter body whorl. Cecilioides tumulorum is
broader and shows a different ratio of body whorl to spire length. This is the first record for the
Arabian Peninsula.
Measurements (illustrated specimen, Fig. 145): H = 6.45; D = 1.81, PH = 2.33; PD = 0.82,
W = 6.
Genus Digoniaxis Jousseaume, 1889
1889 Digoniaxis Jousseaume. — Bull. Soc. Malac. France 6: 348.
This genus was described by Jousseaume for the species D. bourguignati Jousseaume, 1889,
which was found in Yemen “Sur la plage de Mahala, entre Aden et Steamer-Point”. JOUSSEAUME
(1889) discusses the position of his new genus and places it in the Ferussaciidae because of
characters of the spire and the fragility of the shell. The species was never recorded again.
For this investigation, two original lots were available from the MNHNP. One of them
contained a considerably damaged syntype from the collection of Jousseaume. The figure given by
JOUSSEAUME (1890: pl. III, figs 1-3) is much too broad and the whorls appear less shouldered. The
other lot consists of one complete shell and several fragments originating from the collection of de
Morgan from “Aden, Mer Rouge, 1861”.
In one of the fragments ex coll. de Morgan, the protoconch was still intact, displaying a
heterostrophic mode of coiling. This proves that Digoniaxis bourguignati is not a member of the
Ferussaciidae, nor a terrestrial species at all but belongs to the marine Pyramidellidae (Neubert,
in prep.).

Terrestrial and freshwater molluscs of the Arabian Peninsula 411
Genus Coilostele Benson, 1864
1864 Coilostele Benson. — Ann. Mag. Nat. Hist. 13: 136.
1872 Francesia Paladilhe. — Ann. Mus. Stor. Nat. Genova 3: 5.
Diagnosis: The turreted cylindrical shell is translucent and thin with a blunt protoconch of
usually three whorls. The teleoconch whorls with a straight outline, smooth to ribbed with a deep
suture. In the upper teleoconch whorls, the inner parietal walls and the columella are completely
resorbed. In the last two whorls, these structures are still present, the columella is indented with an
oblique lamella.
The phenomenon of resorbed inner shell walls was recently discussed by NAGGS (1997), who
stressed the striking similarity of this character state with what is known from Carychium spp. As a
consequence, Coilostele should be reclassified within the Ellobiidae. Such a dramatic change in
taxonomic position should be postponed until the status is confirmed by either anatomical or
biochemical methods.
Type species: Coilostele scalaris Benson, 1864.
Coilostele paladilhiana Nevill, 1878 Figs 146-148
1872 Francesia scalaris Paladilhe. — Ann. Mus. Stor. Nat. Genova 3: 10, tav. 1, figs 1-4. [non Coilostele scalaris Benson,
1864] (“Kursi près d’Aden”).
1878 Coilostele paladilhiana Nevill. — Handlist of Mollusca in the Indian Museum I: 162 (Aden, ex C. Arturo Issel)
[nom. nov. pro Francesia scalaris Paladilhe non Coilostele scalaris Benson, 1864].
1880 Coelestele aegyptiaca Bourguignat. — Description de diverses espèces de Coelestele et de Paladilhia découvertes en
Espagne par le Dr G. Servain: 12 (“Alluvions du Nil pres Damiette”).
1880 Coelostele isseli Bourguignat. — Description de diverses espèces de Coelestele et de Paladilhia découvertes en Espagne
par le Dr G. Servain: 15 (“Alluvions d’un torrent á Kursi, pres d’Aden, en Arabie”).
1880 Coelestelle arabica Bourguignat. — Description de diverses espèces de Coelestele et de Paladilhia découvertes en
Espagne par le Dr G. Servain: 15. [nom. nov. pro Francesia scalaris Paladilhe non Coilostele scalaris Benson, 1864].
1890 Coelostele bourguignati Jousseaume. — Bull. Soc. Malac. France 7: 95, pl. 3, figs 16-18 (“Alluvions du torrent de la
plaine de Mahala, près d’Aden”).
1890 Coelostele stenostoma Jousseaume. — Bull. Soc. Malac. France 7: 95, pl. 3, figs 19-21 (“Alluvions du torrent de la
plaine de Mahala, près d’Aden”).
Material: Yemen: Aden, SMF 230194/2, coll. Bosch ex Rolle; Red Sea, ad-Durayhimi, 14°40'N 42°59'E, sand beach,
16.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (33), SMF 311640/5; al-Mahrah, Nishtun, 15°48'54"N 52°11'26"E,
sandy and rocky beach, 5./6.X.1995, H. Dekker & F. Ceuninck v. Capelle, DEKK (2). — sub Coelestele aegyptiaca: unknown
locality, coll. Bourguignat, MHNG (15). — Om a n : Dhofar, NE slope of Qara mountains, Jibjat area (I.a.e.n. 126B), Girod
ded. 1997, SMF 311670/1 (protoconch and upper whorl).
Type material: PALADILHE (1872) identified the specimens from “Kursi prés d’Aden” as
the Indian Francesia scalaris, which is a different species. This mistake was corrected by NEVILL in
1878 who subsequently created the substitute name Coelestele paladilhiana for the Arabian species.
For the same reason, disregarding Nevill, BOURGUIGNAT (1880: 7) created the second substitute
name Coelestele arabica.
Lectotype Coelostele isseli, MHNG; six paralectotypes MHNG. The lectotype of C. isseli was
designated by Waldén in 1973 but (to the knowledge of the author) has not been published yet.
Holotype Coelostele aegyptiaca, MHNG (Fig. 148). This specimen was also designated by Waldén
in 1973 as “lectotype”, but as the type lot contains only one specimen it has to be regarded as the
holotype.
Coelostele aegyptiaca is here regarded as a synonym of Coilostele paladilhiana, because the type
specimen shows no differences from the other taxa recorded from the area. In particular, the most
important character, the “smooth” shell, turned out to be a complete misinterpretation. The type
specimen of C. aegyptiaca is ribbed, although the ribs are quite smooth. Shells from Massaua, as

412 E. NEUBERT
well as from Arabia, show some variation in the strength of the ribs. More importantly, the
protoconch and teleoconch form and the small columellar callus are so similar that no differen-
tiating characters could be found.
The types of C. bourguignati and C. stenostoma are probably lost (Bouchet, pers. comm. 1996).
Both taxa originate from the same lot from Mahala close to Aden and were found in the debris of
a river together with C. paladilhiana and C. isseli. Comparing the descriptions, no essential
differences exist. In both taxa, as well as in C. paladilhiana, the protoconch is blunt and exceeds
the diameter of the first teleoconch whorl. The aperture is obliquely elongate and the shape of the
specimens investigated varies to some extent from cylindrical to slightly conical. For this reason,
both taxa are believed to be conspecific with C. paladilhiana.
Description: The minute and fragile shells are cylindrical turreted and non-umbilicate.
They are white with a corneous glaze in fresh specimens. The protoconch consists of 2.5-3 smooth
and glossy whorls. They are blunt and quickly reach the width of the teleoconch whorls. The
teleoconch whorls are sculptured by ribs. The whorls are flat and shouldered subsuturally. The
suture is rather deep.
The oblique aperture is broadened basally. In adult specimens, a weak and narrow labial callus
may be present. The columella is twisted and bears a weak lamella which is much stronger
internally. The peristomial rims are connected by a thin callus which also spreads over the
umbilical area.
Measurements: Holotype of Coelestele aegyptiaca (Fig. 148): H = 2.40; D = 0.50, W = 7.
Lectotype of Coelestele isseli (Fig. 147): H = 3.20; D = 0.60; W=8.
Remarks: JOUSSEAUME (1890: 88, 99) records Coelestele aegyptiaca Bourguignat, 1880 from
“l’ile de Seed-Sayd, petite ile dans le voisinage de Massaouah” and “dans les alluvions du torrent de
la plaine de Mahala” which is in Yemen. Material from Massaua (Eritrea) still exists in the MHNG.
It is identical with that collected and recorded by ISSEL (1873: 522 ff.) at “Sceck-Sayd”. At that
time, the distinction between the smooth C. scalaris and the ribbed C. paladilhiana was not clear,
and Issel might have confused C. scalaris with what was later described as C. aegyptiaca. The
Sudanese specimens are indeed close to the lectotype specimen of C. aegyptiaca and are here
considered to be conspecific.
As a result, this species is hitherto only known from the few places in the lowlands of the
south-western area of Yemen, and also occurs in the coastal areas of Sudan. The type locality of C.
aegyptiaca at Damiette in the Nile delta has urgently to be reconfirmed.
This species is here recorded for the first time from Oman. The specimen originates from an
archaeological excavation and is badly damaged. Only the protoconch and parts of the first
teleoconch whorls are left. Nevertheless, there is no doubt about its identification. This record
extends the known range of C. paladilhiana considerably to the east.
Family Clausiliidae
Genus Macroptychia O. Boettger, 1877
1877 Macroptychia O. Boettger. — Clausilienstudien: 108.
Diagnosis: Small to medium-sized left-coiled shells. The upper lamella and the spiralis are
connected, a lunella is present. The cervix displays a blunt basal and dorsal keel. The protoconch
is flat and densely granulated.
Type species: Clausilia senaariensis L. Pfeiffer, 1855.

Terrestrial and freshwater molluscs of the Arabian Peninsula 413
Figs 149-152: (shells: original size × 4, cervices: original size × 6), 149-150: Macroptychia (Sabaeola) schweinfurthi, lectotype
ZMHB 41344 a. 149: Frontal view. 150: Cervix. 151-152: Macroptychia (Sabaeola) sumarana, syntype BM(NH) 1939.4.19.91.
151: Frontal view. 152: Cervix.
Subgenus Sabaeola Lindholm, 1925
1925 Sabaeola Lindholm. — Proc. Malac. Soc. London 16: 266.
Diagnosis: The protoconch is acute and of conic shape. The principalis and the upper
palatalis diverge, a subclaustralis is present, sometimes a very small basalis may be present.
Penis without a caecum, a penial papilla is present. The MRP has two arms with the epiphal-
lial part attaching proximally on the epiphallus. The penial arm inserts in a median position on
the distal part of the penis. The diverticulum is present but extremely weak.
Type species: Clausilia Schweinfurthi v. Martens, 1889.
Remarks: Lindholm based Sabaeola on the absence of the lunella, a character state which he
himself obviously never checked. He relied on the statement of v. MARTENS (1889), who wrote in
the description of C. schweinfurthi : “lunella nulla”. Investigation of the lectotype of C. schwein-
furthi (ZMHB 41344 a) revealed that the lunella is definitely present, thus eliminating the major
difference from Macroptychia s. str. Unfortunately, the genital morphology of the African Macro-
ptychia species is still unknown (Neubert, in prep.) and it is uncertain whether the conchological
differences described may be important enough to justify the status of Sabaeola. Thus it seems to
be reasonable to keep Sabaeola separated from Macroptychia until more anatomical details may
support either lumping or splitting.
Macroptychia (Sabaeola) schweinfurthi (v. Martens, 1889) Figs 149-150, 153-154
1889 Clausilia Schweinfurthi v. Martens. — Nachrbl. dtsch. malakozool. Ges. 21 (9/10): 152 (Yemen, Menaha, in rock
crevices of steep slopes between moss, 7500 ft).
Material: Saudi Arabia: Asir mountain region, Raydah escarpment, 2300 m, IV.1994, S. Newton, SNMNH-
MO 109/1, SMF 311642/1 (preserved). — Yemen: an-Nabi Shu’ayb mountains, area A, M.M. Al-Safadi, NEUB (11,
preserved); an-Nabi Shu’ayb mountains, area B, M.M. Al-Safadi, NEUB (3); Aiban mountains; Hajjah area NW of Sana’a, M.M.
Al-Safadi, NEUB (1, preserved).
Type material: Lectotype Clausilia schweinfurthi, ZMHB 41344 a.
Description: The shells are brown, slender fusiform with an acute protoconch consisting
of 2.5-3 granulated whorls. The whorls of the teleoconch are nearly flat with a flat suture bordered

414 E. NEUBERT
by a fine white line. They are finely ribbed, in some specimens groups of white ribs may occur. In
the cervical area, the ribs are somewhat coarser. The last whorl displays two blunt keels.
The peristome is subquadrate with two shallow basal furrows. The peristomial rim is reflected
and strengthened by a weak callosity.
The lunellar lies dorsally and consists of a principal lamella, the upper palatalis and a thick
evenly rounded lunella. The upper palatalis is often somewhat knobbly. A small subclaustralis is
present in almost all specimens, a faint basalis was observed in some specimens.
Upper lamella and spiralis are connected. The columellaris lies deep in the peristome and
reaches the same height internally as the spiralis. The subcolumellaris is arched and may be seen in
perpendicular view on the peristome. The clausilium plate is slender linguiform, apically callose
with the rims bent upwards.
Measurements (lectotype, Figs 149-150): H = 15.0; D = 3.3; PH = 3.3; PD = 2.5; W = 11;
R1 = 7.
Genital morphology (Figs 153-154): The elongated penis is subdivided into a distal and
a proximal part, with the lumen of the distal part filled with longitudinal pilasters. The proximal
part is somewhat swollen, in particular in the place where the penial papilla is situated. The
epiphallus is shorter than the penis, its lumen also displays pilasters. The MRP is divided into two
arms. The penial arm attaches in the middle of the distal part of the penis, the epiphallial muscle
close to the boundary of epiphallus and VD. A flagellum is not visible.
The vagina is shorter than the penis, the BC is elongated with a very weak diverticulum.
Macroptychia (Sabaeola) sumarana (Connolly, 1941) Figs 151-152, 155
1941 Clausilia sumarana Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 23, pl. 3, fig. 3,
textfig. 6 (Highest point of pass over Jabal Sumara, between Ibb and Yarim, 9750 ft).
Material: Yemen: Sumara mountains, M.M. Al-Safadi, NEUB (4 preserved); Aiban mountains, M.M. Al-Safadi,
NEUB (2 preserved).
Type material: Syntypes BM(NH) 1939.4.19.91-102.
Description: The brown fusiform shell is somewhat ventricose, the protoconch consists of
two whorls with a finely granulated surface. The whorls of the teleoconch are flat with a shallow
suture. A fine white border line is present in some specimens. The whorls are evenly and finely
ribbed. The last whorl has a blunt basal keel, the dorsal keel is very faint.
The peristome is subquadrate to oblique with a very shallow basal furrow. The peristomial rim
is reflected with a small palatal callosity.
The lunellar lies dorsally with a long principal lamella, the upper palatalis is short, not
connected to the lunella. The lunella is straight, a short subclaustralis is present.
The upper lamella is connected to the spiralis; the columellaris is situated deep in the aperture.
Internally, columellaris and spiralis end at the same height. The subcolumellaris is arched and ends
above the basal keel. It is visible in perpendicular view on the peristome. The clausilium plate is
linguiform and very slightly thickened at the apex.
Genital morphology (Fig. 155): The long penis is subdivided into two parts, the
epiphallus is shorter than the penis. A penial papilla is present. The MRP is split into a penial and
an epiphallial arm with the first attaching at the border of epiphallus and VD, the second in the
middle of the distal part of the penis.
The vagina reaches the length of the penis, the diverticulum is weak and branches off in the
basal part of the pedunculus.
Measurements (syntype, Figs 151-152): H = 13.90; D = 3.20; PH = 2.90; PD = 2.33;
W = 10.5; R1 = 9.

Terrestrial and freshwater molluscs of the Arabian Peninsula 415
Figs 153-155: Genital organs of Macroptychia (Sabaeola) species. 153-154: Macroptychia (Sabaeola) schweinfurthi, specimen from
Yemen, Hajjah area, NW of Sana’a. 153: Genital apparatus. 154: Penis and epiphallus dissected, showing longitudinal pilasters
and penial papilla. 155: Macroptychia (Sabaeola) sumarana, genital apparatus, specimen from Yemen, Sumara mountains.
Remarks: Anatomically, there are no major differences between M. (S.) schweinfurthi and
M. (S.) sumarana. On average, M. (S.) sumarana is smaller, the suture is deeper, the teleoconch is
more finely ribbed and the dorsal keel is very shallow, a character which may serve as a good
conchological criterion to discriminate between both taxa.
Family Sphincterochilidae
Genus Sphincterochila Ancey, 1887
1887 Sphincterochila Ancey. — Conch. Exchange II (2): 23 [genus XXVI].
Diagnosis: Heavy thick-walled spheroidal shells with a typical chalky surface. The shell may
be sharply keeled to rounded, its surface smooth to heavily granulated. The last whorl always
descends under the periphery of the preceding one. The genitalia are mainly characterised by the
presence of a penial stimulatory organ.
Type species: Helix boissieri Charpentier, 1847.
Bibliographic note: In the years 1886 and 1887, Ancey published an important contri-
bution in “The Conchologist’s Exchange” describing generic and subgeneric taxa of what he
believed to be Helicidae. This paper was split into six parts which are cited correctly under
“References” at the end of this work.

416 E. NEUBERT
Subgenus Sphincterochila s. str.
Diagnosis: The subdivision of the genus Sphincterochila follows the basic work of FORCART
(1972), who accepted three subgenera according to the fine anatomy of the penial stimulatory
organ. Sphincterochila s. str. is characterised by a heavy sheath with longitudinal pilasters filling its
lumen and is situated distally to the stimulatory gland. Both the ducts of the gland and the
“appendicula” apically enter the stimulator sheath.
Sphincterochila (Sphincterochila) prophetarum (Bourguignat, 1852)
1852 Helix prophetarum Bourguignat. — Testacea novissima quae Cl. de Saulcy in itinere per Orientem annis 1850 et
1851, collegit: 12 (“Palaestinam, in locis aridis circa Hierosolymam urbem”).
1901 Leucochroa arabica Pallary. — Bull. Inst. Egypt. 1901: 4, pl. 1, figs 1-2 (Ouadi Naouk [= Egypt, northern slope of
Jabal Galala el Baharieh S of Suez]).
1923 Albea prophetarum var. arabica sous-var. gracilis Pallary. — J. Conchyl. 68: 188, pl. X, fig. 3 (“Ouadi Raha et massif
du Moghara” [Sinai]).
1923 Albea prophetarum var. arabica sous-var. minor Pallary. — J. Conchyl. 68: 188, pl. X, fig. 2 (Ouadi Naouk).
Material: Saudi Arabia: Wadi Madaqarq, XII.1943, Waterston, BM(NH) 1945.8.23.186.
Description: The shell of this Sphincterochila species is small to medium-sized compared
to its congeners. The protoconch is smooth and somewhat elevated. Characteristically, the surface
of the teleoconch is smooth apart from some faint irregularly spaced axial riblets. The suture is of
medium depth, the whorls are rounded. The body whorl bears a faint blunt keel situated on the
periphery. The aperture is crescent-shaped with a weak angularis. The umbilicus is closed, the
periomphalum excavated.
Measurements: H = 11.0; D = 16.0; PH = 5.3; PD = 8.7, W = 5.
Remarks: The specimen from Wadi Madaqarq was mentioned by MORDAN (1980 b: 362)
under the name Sphincterochila (Albea) candidissima (Draparnaud, 1801). This species from the
western Mediterranean differs in both shell and anatomical characters from S. prophetarum and is
not known in the Palestinian area. Sphincterochila prophetarum inhabits the north-eastern moun-
tains of Egypt, the Sinai, the lower parts of Palestine, Jordan and the coastal mountains of Saudi
Arabia. There is no information about the southern limits of its range in Arabia.
Family Polygyridae
Genus Polygyra Say, 1818
1818 Polygyra Say. — J. Acad. Nat. Sci. Philadelphia 1 (10): 276.
Diagnosis: Shells depressed to discoidal. The whorls are narrowly coiled and often bluntly
angulate. The aperture is small, the parietal callus bears a short tooth.
Type species: Polygyra septemvolva Say, 1818.
Polygyra cereolus (Megerle von Mühlfeldt, 1816) Figs 156-160
1816 Helix cereolus Megerle von Mühlfeldt. — Ges. Naturf. Fr. Berlin, Mag. VIII (1): 11.
Material: Saudi Arabia: Eastern Province, beach debris at Haii al-Bahr, N of Jubail, I.1992, E. Neubert, SNMNH-
MO 110/1, SMF 311643/1; Eastern Province, in gardens in Jubail, N of Dhahran, XI.1993, P. Symens, SNMNH-MO 111/3,
SMF 311644/3; Eastern Province, Jubail, Dauhat ad-Dafi, in gardens, 11.XII.1993, P. Symens & M. Werner, SNMNH-
MO 112/5, SMF 311645/5, NEUB (2), SNMNH-MO 113/16 (preserved), SMF 311646/16 (preserved).
Description: The brown discoidal shell has a low conical spire. The protoconch is smooth
and is formed by two whorls.

Terrestrial and freshwater molluscs of the Arabian Peninsula 417
Fig. 156: Genital
organs of Pol ygyra
cereolus, specimen
from the Eastern
Province, Jubail,
Dauhat ad-Dafi.
The teleoconch whorls are narrowly coiled and shed by a deep suture. The surface is covered
by coarse and regularly spaced ribs which become weak on the subsurface of the shell. The body
whorl is characterised by a blunt subsutural shoulder and a shallow depression opposite the
aperture.
The aperture is somewhat trigonal with a callose lip. On the upper edge, the peristome is
nearly rectangular to attach to the body whorl. The peristomial rim is strongly recurved. There is a
thick parietal callus situated in between the peristomial rims. The lamella is bifid in several
specimens. Deep in the interior of the shell close to the shallow depression, another elongated
lamella occurs on the inner parietal wall of the shell.
The umbilicus is open and shaped like a bore hole. The body whorl suddenly opens to form a
wide and shallow periomphalum.
Genital morphology (SMF 311646, Fig. 156): Penis and epiphallus are of approxi-
mately the same length. The penis is characterised by an elongated glandular tissue which covers
half of its surface. Internally, there are two slender pilasters running from the atrium to the
beginning of the epiphallus. They are smooth in the penis, producing an open tube. In the
epiphallus, however, their rims are heavily folded and interdigitate, forming a closed tube.
The vagina is remarkably short, the pedunculus slender with an elongated pointed BC.
Remarks: This species is native to the south-eastern United States, in particular to Florida.
It is known to be transported by human activities. Recent findings indicate that the species is well
adapted to the artificial environments of gardens in the industrial city north of Jubail. The gardens
are irrigated regularly and, combined with the hot climate, a permanent tropical microclimate is
produced which can support a population of this species. The numbers of well-established garden
populations and the fact that the species even occurs in the debris of a waste water channel indicate
that it might have been introduced earlier than suspected (NEUBERT 1995). It is likely that it was
introduced by American employees of companies in the industrial city of Jubail.
In its natural habitats, the species varies considerably in respect to shell morphological
characters such as size, shape and dentition. However, the specimens found here show a very
limited variation in these characters. The genital organs differ in the length of the vagina. In

418 E. NEUBERT
specimens of P. cereolus from Florida, Key West as well as in P. septemvolva volvoxis (L. Pfeiffer,
1846) from Jacksonville, Florida (PILSBRY 1940: 581, fig. 378: 1, 4), the vagina is extraordinarily
elongate.
Family Hygromiidae
Hygromiidae gen. sp. indet. Fig. 161
Material: Yemen: al-Ashmur between Amran and Hajjah, below and on limestone, 2700 m, 9.IV.1993, H. Dekker &
F. Ceuninck van Capelle, DEKK (70).
One lot of presumably Hygromiidae from Yemen represents an unidentified species. A typical
fresh specimen (SMF 311647/15) is illustrated here. The lot contains numerous eroded white
shells with several brown spiral bands (a subperipheral is always present). The shell is subglobular
with a semi-lunate aperture. The umbilicus is open, the columellar rim reflects over the peri-
omphalum. Unfortunately, no preserved specimens are available to clarify the status of this taxon.
Genus Xerocrassa Monterosato, 1893
1893 Xerocrassa Monterosato. — Atti Accad. Palermo (3) 2: 23.
Diagnosis: Globular, trochiform or nearly flat shells of medium size. In the male genital
system, the flagellum is not longer than the epiphallus. A penial appendix is missing. The two dart
sacs and the glandulae mucosae are reduced.
Type species: Helix seetzeni L. Pfeiffer, 1847.
Xerocrassa seetzeni (L. Pfeiffer, 1847)
1847 Helix seetzeni L. Pfeiffer. — Z. Malakozool. 4: 14.
Description: The shell is globular and thick-walled. The protoconch is smooth, white with
a deep suture. The initial teleoconch whorls are flat, in some specimens bluntly keeled with a
shallow suture. The last whorls are rounded, with a deeper suture. The sculpture of the teleo-
conch consists of fine axial riblets on the first whorls which subsequently become rugose and
irregularly spaced.
The aperture is subglobular with a sharp peristome. Internally, a weak lip is present. The
columellar rim of the peristome extends over the periomphalum and nearly covers the umbilicus.
The basic colour is chalky white. Very often, obscure brown spiral bands occur. The genital organs
are described and depicted by HESSE (1934: 8, Taf. 2, Fig. 12 a-c). FORCART (1976: 145) reports
that the sole of the (living?) animal is coloured violet.
Remarks: This species was reported upon by MORDAN (1980 b: 362) from the northern part
of Saudi Arabia “N of Tabuk, approx. 100 km S of the Jordan border”. It is likely that it inhabits
the semi-desert habitats similar to those in Palestine and Syria.
Figs 157-173: (original size × 4), 157-160: Polygyra cereolus, specimen from Eastern Province, Jubail, Dauhat ad-Dafi. 157:
Frontal view. 158: Apical view. 159: Subsurface. 160: Inner parietal lamella. 161: Hygromiidae indet., specimen from Yemen,
al-Ashmur between Amran and Hajjah. 162-170: Xeropicta aff. mesopotamica. 162-164: Syntype of Helix (Xerophila) meso-
potamica, ZMZ 529959. 162: Frontal view. 163: Apical view. 164: Subsurface. 165-167: Syntype of Helix (Xerophila)
mesopotamica var. ghaesiana, ZMZ 504562. 165: Frontal view. 166: Apical view. 167: Subsurface. 168-170: Specimen from
Oman, Fanja, W of Muscat. 168: Frontal view. 169: Apical view. 170: Subsurface. 171-173: Xeropicta parableta. 171: Frontal
view. 172: Apical view. 173: Subsurface.

Terrestrial and freshwater molluscs of the Arabian Peninsula 419

420 E. NEUBERT
Genus Xeropicta Monterosato, 1893
1893 Xeropicta Monterosato. — Atti Accad. Palermo (3) 2: 24.
Diagnosis: Medium-sized to small shells with a subdepressed to subconoidal spire. The
umbilicus is open, the shell is always white with brown spiral bands. The vagina with two dart sacs
and two accessory sacs, which are always longer than the dart sacs. The penis with a penial
appendix. Distributed throughout the eastern Mediterranean area and the Middle East eastwards
to Afghanistan.
Type species: Helix krynickii Krynicki, 1833.
Xeropicta krynickii (Krynicki, 1833)
1833 Helix krynickii Krynicki. — Bull. Soc. Imp. Nat. Moscou 6: 434 (“Habitat inter montes calcareos Tauriae”).
1841 Helix vestalis L. Pfeiffer. — Symbolae ad Historiam Heliceorum I: 40 (“Aegyptus”).
1855 Helix joppensis A. Schmidt. — Abh. naturw. Ver. Sachsen und Thüringen 1: 29, Taf. 6, Fig. 34 (Jaffa).
Description: The shell is small to medium in size and very variable. It may have a
considerably flat to elevated spire. The protoconch consists of two smooth and often dark brown
whorls. The suture of the teleoconch whorls is shallow to moderately deep. The initial whorls of
the teleoconch are covered by fine and regularly spaced riblets. This sculpture is more coarse and
irregular towards the last whorl. The genital organs have often been described and depicted (cf.
GITTENBERGER 1991: 106, fig. 24-26).
Remarks: In the synonymy, the names of the taxa are given which may be of importance for
the area considered. The author here follows the view of several colleagues (Hausdorf, Nordsieck,
pers. comm., Bank, in litt.). There are no essential differences in either genital or conchological
characters. In general, the taxonomy of Xeropicta krynickii is still vague as minor conchological
differences have led to a plethora of descriptions thus complicating the situation. A revision is
urgently needed.
From the Arabian Peninsula, MORDAN (1980 b: 363) records this species under the name
Helicopsis (X.) vestalis from Jeddah. This record has to be re-confirmed by dissection.
Xeropicta mesopotamica (Mousson, 1874) Figs 162-170, 174
1874 Helix (Xerophila) mesopotamica Mousson. — J. Conchyl. 22: 22 (“dans toute la vallée de la Mesopotamie”).
1874 Helix (Xerophila) mesopotamica var. ghaesiana Mousson. — J. Conchyl. 22: 37 (“dans l’île de Ghaes, dans le golfe
Persique”).
1892 Helix (Xerophila) mesopotamica var. alepina Westerlund. — Verh. zool.-bot. Ges. Wien 42: 29 (“Syrien, bei Halep”).
Material: Saudi Arabia: Eastern Province, Tarut, plantations N of Tarut centre, 14.VI.1992, R. Kinzelbach,
SNMNH-MO 114/1; al-Qatif Oasis, palm plantations W of centre, irrigation channels, 14.VI.1992, R. Kinzelbach, SMF
311648/1; Jubail, in gardens at Dauhat ad-Dafi, 11.XII.1993, P. Symens & M. Werner, SMF 311649/1 (preserved); same
locality, 7.I.1994, P. Symens, SNMNH-MO 115/1 (preserved), SMF 311650/1 (preserved). — O m a n : Fanja, W of Muscat,
I.1993, I. & H. Pauscher, NEUB (7); Oasis of Buraimi, I.1993, I. & H. Pauscher, NEUB (1).
Type material: Although there is sufficient material present in the ZMZ, the designation
of a lectotype has been postponed until preserved specimens from Iraq can be studied. There are
seven lots labelled as Xeropicta mesopotamica, six of them with the original labels still present. In
the author’s opinion, only three of these lots represent the taxon X. mesopotamica and a lectotype
should be designated from one of these. They originate from “v.[on] Haleb n.[ach] Beredschek,
Schlaeffli 62” ZMZ 504565/19, “Diwanich am Euphrat (Wüste), Schlaeffli 62” ZMZ 504566/35
and “Samava” ZMZ 529959/4. Describing his H. (X.) mesopotamica var. ghaesiana, Mousson
himself records the nominate form as abundant at “Ninive, Tekrit, Bagdad, et Samava”, while the
variety was found on the Island Ghaes in the Arabian Gulf (syntypes ZMZ 504562) [= Keis Island,
Queys, Gazire-ye-Kis E of Bandar-e Lenge].

Terrestrial and freshwater molluscs of the Arabian Peninsula 421
The remaining lots represent different species of Xeropicta. In the author’s opinion, ZMZ
504561/10 (“v.[om] Küstengebirge nach Haleb”) represents Xeropicta krynickii, while the lots
ZMZ 504567 (“Syrien, nahe Biredschik”) and ZMZ 504364 (“Syrien, von Biredschek nach
Siverek”) contain specimens of Xeropicta derbentina (Krynicki, 1836).
Description: The shell is small to medium-sized with an often elevated spire. It is basically
cream-coloured with a somewhat glossy surface. Brown spirally arranged spots may occur, in some
specimens rudiments of subsutural spiral bands may be visible. Only the narrow peripheral brown
spiral band is always present.
The protoconch consists of two smooth and brown whorls. The teleoconch whorls are
rounded to often bluntly shouldered. The subsurface of the shell is flat. The surface is covered by
densely packed fine ribs. The ribbing of the last whorl is not essentially different from that of the
initial ones. The suture is of medium depth and only slightly indented. There are short and
irregularly arranged spiral furrows while the spiral sculpture of very fine and parallel grooves (as
seen in other Xeropicta species) is less abundant.
The aperture is subcircular, the lip in the interior. The peristome is simple and sharp. The
umbilicus is open and somewhat eccentric. It reaches about one sixth of the shell diameter. The
H. (X.) mesopotamica var. ghaesiana differs in having a somewhat flattened shell. The last whorl
is somewhat broader, but as this falls into the variation seen in so many Xeropicta shells, it
cannot be used as a differentiating character. The variety is thus relegated into the synonymy of
the nominate form.
Genital morphology (SMF 311649/1, Fig. 174): The short penis is characterised by a
voluminous conical distal part. The penial appendix branches off basally from the penis surmount-
ing it somewhat. Proximally, the MRP inserts from a small sheath and attaches at the diaphragm.
The epiphallus reaches more than twice the length of the penis. The flagellum is short, reaching
the length of the penis. Internally, a conical penial papilla is found filling the lumen of the penis
nearly completely.
The vagina reaches the same length as the penis. There are two dart sacs with two longer
accessory sacs. The four glandulae mucosae split in several branches. They insert at the vagina in
the same height as the BC does. The pedunculus is slender, the BC an elongate vesicle.
Measurements: Illustrated syntype of Xeropicta mesopotamica ex ZMZ 529959, Samava
(Figs 162-164): H = 6.00; D = 8.75; PH = 3.80; PD = 3.30; W = 5.5. Syntype of Helix (Xerophila)
mesopotamica var. ghaesiana ZMZ 504562 (Figs 165-167): H = 7.10; D = 11.10; PH = 4.50;
PD = 5.60; W = 6. Illustrated specimen from Fanja, Oman (Figs 168-170): H = 4.80; D = 8.10;
PH = 3.60; PD = 4.00; W = 5.
Remarks: This species is characterised by the very short penis and an appendage, which
branches off very close to the atrium. This is very similar to that recorded for X. acrotirica
(GITTENBERGER 1991: fig. 26). In that taxon, the penial appendix has the same basal position as in
the specimens described from Saudi Arabia. The penis is also swollen and the flagellum short, but
the shells differ considerably as in X. acrotirica, the mode of ribbing is much coarser, the shells are
more tumid and the umbilicus is less eccentric. As there is no topotypic material available for
anatomical investigation, the identification of the Arabian specimens as X. mesopotamica remains
provisional.
Another species which could be confused with the Arabian species is Helix (Xerophila)
parableta O. Boettger, 1881 (“Zahlreich in den Anschwemmungen des Araxes bei Nachitsche-
wan”). It was assigned to Xeropicta by SCHILEYKO (1978: 223), who investigated preserved animals
from “mountainous steppe areas of the Talysch close to the village Kelia” and stated “despite
considerable differences to other species of this genus, this species completely matches the

422 E. NEUBERT
diagnosis of the genus concerning characters of the genital organs. Differences to other members
of this genus particularly concern the following characters: Flagellum very short and conical, the
bursa copulatrix with a thin stalk and a big reservoir. Characteristic internal structures of the penial
appendix and in the distal part of the penial tube are less pronounced.” (translation by Hohorst).
He illustrates the investigated shell (SCHILEYKO 1978: Tab. XI, fig. 101), which seems to be
identified correctly. For reasons of comparison, the lectotype of X. parableta is illustrated here
(Figs 171-173). It is characterised by the prominent nodulose keel on the periphery of the body
whorl, the non-eccentric umbilicus, the coarse ribs and the considerably elevated spire. Both
figures make it evident that X. parableta represents a species separate from the Arabian ones.
Measurements (lectotype of Helix (X.) parableta, SMF 10348, Figs 171-173): H = 5.4;
D = 7.6; PH = 3.4; PD = 3.7; W = 5.5.
On the Iranian plateau and in the fringing inner-Asiatic steppe areas, another Xeropicta species
is very common. This species is usually called X. candaharica (L. Pfeiffer, 1846). It differs from X.
mesopotamica by its big depressed shell, the nearly smooth surface of the last whorl, the wider
umbilicus and the coarser mode of ribbing on the initial whorls. There are no major conchological
differences from X. derbentina, and the relative length of the flagellum is also similar. The base of
the penial papilla is somewhat swollen in X. derbentina, but the value of such an observation is
doubtful in respect to species delimitation (intraspecific variation?). Specimens from lots from
Kandahar and some other places in Afghanistan are considerably smaller with a more elevated spire.
To sum up, the conchological differences between most of the species of Xeropicta are small,
intergrading forms are abundant. It is still questionable whether anatomical details of the genital
organs are constant enough to serve as a reliable criterion for species differentiation. The complete
complex urgently needs to be investigated in more detail.
Genus Monacha Fitzinger, 1833
1833 Monacha Fitzinger. — Beitr. Landeskunde Oesterr. Enns (= Verh. Vaterl. Ges. Wien) 3: 95.
Diagnosis: Small to medium-sized depressed to subglobose and somewhat fragile shells.
The surface is smooth or bears small bristles and may exhibit some faint spiral grooves. The
subcircular aperture is often thickened by a lip which may be reddish-coloured. The teleoconch
whorls are milky white, an opaque peripheral band may occur.
Type species: Helix cartusiana O.F. Müller, 1774.
Monacha obstructa (L. Pfeiffer, 1842) Fig. 175
1842 Helix obstructa L. Pfeiffer. — Symbolae ad Historiam Heliceorum II: 35 (Tripoli).
Description: The shell is small (D = 10-12 mm), the spire is somewhat elevated and
broadly conical. The protoconch consists of 1.5 smooth whorls. The teleoconch is opaque, its
whorls are evenly rounded. They are sculptured by faint and irregularly arranged riblets, the
complete surface is covered by shallow pits. To the naked eye, the shell seems to be smooth.
The last whorl descends under the periphery of the preceding whorl. The peristome is sharp
and marked by a strong porcelaineous white lip which can also be seen from the outside as a thick
white callosity. The peristomial rim is red. The umbilicus is closed, but the last whorl forms a
characteristic funnel-shaped “pseudo-umbilicus”. The animal is cream-white, the outer lung wall is
coloured by a few dark brown blotches.
Genital morphology (Fig. 175): The penis is short conical, the epiphallus reaches twice
the length of the penis. A MRP is lacking. The flagellum is slender and very long, reaching the

Terrestrial and freshwater molluscs of the Arabian Peninsula 423
Figs 174-175: Genital organs. 174: Xeropicta aff. mesopotamica, specimen from Eastern province, Jubail. 175: Monacha obstructa,
specimen from Syria, Tall Shaik Hamad, Nahr al-Habur.
length of the epiphallus. The penial appendix branches off at the base of the penis. It is very long
reaching more than twice the length of penis and epiphallus combined. In the penis, a papilla is
present which adheres to the inner wall of the penis. The epiphallial lumen is filled by two thick
and several slender longitudinal pilasters.
The vagina is somewhat longer than the penis. There is only one stem of glandulae mucosae
which split into several short tubes.
Remarks: This species is known from a few localities (particularly oases) in Saudi Arabia and
seems to have been introduced by human activities (MORDAN 1980 b: 362). It is very abundant in
southern Turkey, Syria and Palestine. Since no shells or preserved specimens from Arabia were
available, anatomical details were described from specimens from northern Syria.
Genus Lejeania Ancey, 1887
1887 Lejeania Ancey. — Conch. Exchange I (12): 75 [genus XIX].
Diagnosis: Medium-sized depressed conical to subspheroidal shells. The umbilicus is
always open to perspectively enlarged. The female genital organs with two slender appendiculae.
The penial papilla consists of a conical velum and a bilobed inner part.
Type species: Helix darnaudi L. Pfeiffer, 1854.
The first to describe anatomical details of Lejeania was POLLONERA (1888: 27, tav. III, fig. 17),
who depicted the sexual organs of Helix lejeaniana Bourguignat, 1883. Later, PFEFFER (1931) and
VERDCOURT (1969) presented anatomical data on Ethiopian species of this genus.

424 E. NEUBERT
The most striking feature is the fact that in Ethiopian species the MRP is lacking (Pfeffer
investigated L. darnaudi and Verdcourt L. isseli (Morelet, 1872)). It is unlikely that three authors
overlooked a character of this importance. Additionally, VERDCOURT (1969: 178) reports that the
“right hand ocular retractor passes to the left of the genitalia”. A missing MRP means that the
complete male system has lost its attachment on the diaphragm. When retracting the protruding
genitalia after copulation, there are at least three possibilities for the resting position of the genitalia
in relation to the ocular retractor in the case of a missing MRP. The first would be the usual case,
with the ocular retractor passing between the male and female system with the penis below. The
second is the same position with the penis situated above the ocular retractor. Thirdly, the penis is
on the right side and the ocular retractor passes freely left to the genitalia to unite with the
columellar muscle.
The character state “missing MRP” was used by Hesse to define his Helicellinae. More recent
investigations have revealed that this character is present in several independently evolved groups
(SCHILEYKO 1978, NORDSIECK 1987). Following the ideas given here, the value of this character for
phylogenetic investigations seems to be vague.
The form of the appendiculae and the velum around the penial papilla support the view of
Pfeffer and Nordsieck, who placed Lejeania, with some reservations, in the Euomphaliini (NORD-
SIECK 1987: 31). Its exact position within the tribus should be clarified by revising the genus. Much
more anatomical research needs to be conducted in the Ethiopian species to determine the
characters and the phylogenetic relationships.
Lejeania is restricted to the humid mountainous regions of Ethiopia and the south-west of the
Arabian Peninsula. The northern distribution limits of the genus are unknown for Arabia. In
Africa, as far as is known, it only reaches the mountains in the north of Eritrea.
Lejeania darnaudi (L. Pfeiffer, 1856) Figs 176-178
1856 Helix darnaudi L. Pfeiffer. — Proc. Zool. Soc. London: 327 (Sennaar, interior of Africa).
The type species of Lejeania has never been described adequately. For this reason, the author is
taking this opportunity to present a description and figure of the holotype, even though L.
darnaudi is an Eritrean species which does not occur in Arabia.
Type material: One syntype Helix darnaudi, BM(NH) 1996024.
Description: The syntype specimen is subadult. The small shell is depressed globular. The
protoconch is smooth consisting of two whorls with a deep suture. The teleoconch whorls are
rounded with a suture of medium depth. The surface is sculptured by oblique riblets with irregular
interspaces. There are no scars of periostracal bristles, the space between the faint ribs is smooth
except for some faint and short spiral grooves. The aperture is rounded with a labial callus lacking
(juvenile character). The umbilicus is open but narrow, in adults the columellar rim may be
somewhat reflected.
The colour is light cream-brown and mottled with white. There is a broad white spiral band
subsuturally followed by a brown band. The periphery is marked by a significant white spiral
followed by several narrow white and brown bands.
Measurements (holotype, Figs 176-178): H = 5.7; D = 7.7; PH = 4.0; PD = 4.1; W = 5.
Remarks: In general, there are no major differences from the Arabian species of this
genus. On average, the Arabian L. leucosticta is larger, but globular to nearly flattened specimens
occur. A difference may be found in the granulated surface which is always displayed in Arabian
specimens. To prove the reliability of this character, more material from Ethiopia needs to be
studied.

Terrestrial and freshwater molluscs of the Arabian Peninsula 425
Figs 176-184: (original size × 4), 176-178: Lejeania darnaudi, syntype BM(NH) 1996024. 176: Frontal view. 177: Apical view.
178: Subsurface. 179-181: Lejeania leucosticta, syntype SMF 3516 b. 179: Frontal view. 180: Apical view. 181: Subsurface.
182-184: Lejeania aperta, syntype BM(NH) 1939.4.19.127. 182: Frontal view. 183: Apical view. 184: Subsurface.

426 E. NEUBERT
Lejeania leucosticta (v. Martens, 1889) Figs 179-181, 185-186
1889 Helix leucosticta v. Martens. — Nachrbl. dtsch. malakozool. Ges. 21 (9/10): 147 (“Yemen: Menaha, an berieselten
Felsen; Uossil am Westabhang des Gebel Harasa, 7000 Fuß”).
Material: Saudi Arabia: Asir mountain region, Raydah escarpment, 2050 m, 25.III.1994, W. Schneider & F. Krupp,
SNMNH-MO 116/1; same data but 1850 m, SNMNH-MO 117/3, SMF 311652/2; same locality, 1800 m, 29.I.1994, P. Symens
& M. Werner, SNMNH-MO 118/1, SMF 311659/4 (preserved); same data but 2350 m, SNMNH-MO 119/1, SMF 311653/1;
same locality, 2100 m, 10.VII.1995, S. Newton, SNMNH-MO 120/1, SMF 311654/1, SMF 311655/2 (preserved); same data but
2300 m, SNMNH-MO 127/1 (preserved); Asir mountain region, Jabal Qahar, 17°39'N 42°53'E, II.1996, S. Newton, SNMNH-
MO 121/4; Asir mountain region, Abha escarpment, 2500 m, Juniperus forest, II.1994, P. Symens & M. Werner, SNMNH-
MO 122/1, SNMNH-MO 123/1 (preserved), SMF 311656/1; Asir mountain region, Raydah escarpment, 2310 m, 18°11'50.0"N
42°24'30.2"E, 5.V.1994, W. Schneider & F. Krupp, SNMNH-MO 124/3, SMF 311657/1; Asir mountain region, rocky area
50 rkm S of al-Baha, 2000 m, 2./3.VI.1995, E. Neubert et al., SNMNH-MO 125/2; Asir mountain region, small wadi at the right
side of the road from Taif to al-Baha, S of Gharie, 1700 m, 2.VI.1995, E. Neubert et al., SNMNH-MO 126/2, SMF 311658/1,
SMF 311562/1 (preserved); Asir mountain region, King Khalid descent near Baha, 2215 m, 3.VI.1995, E. Neubert et al.,
SNMNH-MO 128/5, SMF 311660/4, SNMNH-MO 129/2 (preserved), SMF 311661/3 (preserved); Asir mountain region, at the
road from Taif to al-Baha, 18 rkm N of al-Mandaq, 2050 m, 2.VI.1995, E. Neubert et al., SNMNH-MO 130/1, SMF 311663/1;
Asir mountain region, al-Abna descent near Baha, 2000 m, 3.VI.1995, E. Neubert et al., SNMNH-MO 131/8, SMF 311664/8,
SMF 311665/1 (preserved), NEUB (1). — Yemen: At the road from Kaukabam to Shibam, 2.III.1990, S. Neubert, NEUB (1);
Haqa close to Menaha, 26.III.1988, A. Liebegott, LIEB (2); on top of the Sumara pass, 22.III.1988, A. Liebegott, LIEB (5);
between Amran and Hajjah, 27.III.1988, A. Liebegott, LIEB (5); Jabal Sabar close to Ta’izz, 23.III.1988, A. Liebegott, LIEB (1);
same locality, XII.1989, Rabin, LIEB(1); Bani Matar, Jabal an-Nabi Shu’ayb (W of Sana’a), 3660 m, under stones, 27.IV.1993, H.
Dekker & F. Ceuninck v. Capelle, DEKK (7); Sharas, Wadi Sharas, NE of Hajjah, 800 m, old river sediments, 9.IV.1993, H.
Dekker & F. Ceuninck v. Capelle, DEKK (1); Bani al-Harith, Wadi Dhar (= Zahr), Qaryat al-Qabil, NE of Sana’a, 2200 m,
between leaves, wood and soil, 8.IV.1993, H. Dekker & F. Ceuninck v. Capelle, DEKK (2).
Type material (all specimens listed here are syntypes as they originate from the type lot):
Ye m e n: Menaha, SMF 3516B/5, coll. v. Moellendorff ex v. Martens; same locality, SMF 3516A/2,
coll. Kobelt ex v. Martens; same locality, SMF 138265/2, coll. O. Boettger ex v. Martens ex Schwein-
furth; same locality, SMF 138266/18, coll. Reinhardt ex O. Boettger ex v. Martens ex Schweinfurth;
Uossil, 4000', SMF 138268/8, coll. Reinhardt ex O. Boettger ex v. Martens ex Schweinfurth; same
locality, SMF 209011/2, ex Jaeckel ex Mus. Berlin; same locality, SMF 286756/1, coll. Bosch ex
Rolle ex Mus. Berlin.
Description: The thin brown shell is medium-sized with an irregular pattern of white
streaks and brown spiral bands. The shell coloration varies from nearly white to dark brown, where
the white maculation is strongly reduced. The protoconch is nearly smooth. The spire is elevated
subglobose with a suture of medium depth. The surface of the complete shell is covered by a dense
and prominent sculpture of small granules serving as a basic structure for short bristles. These
bristles are lost in adults. There are only faint riblets on the teleoconch. Several specimens from
the Raydah escarpment show strong spiral grooves on the surface.
The teleoconch whorls are rounded, juvenile shells display a blunt keel at the periphery of the shell.
The aperture is semi-ovate with a very thin lip in the interior. The peristome is sharp. A small
reflection of the columellar part of the peristome partly covers the umbilicus. The umbilicus is
narrow but always open.
The animal and the mantle are cream to grey. The lung cavity is yellowish to cream and
mottled by broad and dispersed black spots of pigment.
Measurements: Illustrated syntype SMF 3516 B (Figs 179-181): H = 11.60; D = 16.15;
PH = 8.60; PD = 8.80; W = 6.5. The largest specimen investigated (from the Raydah escarpment)
measures H = 15.1; D = 18.6; PH = 10.9; PD = 11.2; W = 6.25. Much smaller, but adult, speci-
mens are frequent.
Genital morphology (SMF 311655, Figs 185-186): The penis is a short tube lacking
any pilasters. Together with the epiphallus, it is enwrapped in a sheath of connective tissue which
tightens both organs to a compact mass. The proximal penial lumen is filled with the papilla. This

Terrestrial and freshwater molluscs of the Arabian Peninsula 427
Figs 185-186: Genital organs of Lejeania leucosticta, specimen from Asir mountain region, Raydah escarpment. 185: Genital
apparatus. 186: Penis and epiphallus dissected, showing pilasters and penial papilla.
organ is composed of a cup-shaped velum and a bilobed inner part. The inner surface of the velum
is sculptured by longitudinal pilasters. The two lobes are apically connected to the velum,
producing a small inner lumen with the pore to the epiphallus in between.
The epiphallus reaches twice the length of the penis and consists of thick muscular tissue.
Proximally, a pointed caecum reaching the same length as the epiphallus branches off. The VD is
very thick at its epiphallial boundary. The MRP attaches at the distal third of the epiphallus and
inserts at the diaphragm.
The vagina is short and bears two tubular appendiculae which are connected to each other by
a swollen perpendicular base. They are broadened at the base with a slender stalk and a small
vesicle apically. The lumen of the tubes is filled by simple elongate pilasters which are thickened
and rolled up in the small vesicle. Proximally, two bundles of glandulae mucosae occur. There are
three to six simple or bifid tubes branching off at the same level. The BC is elongate ovoid.
Lejeania aperta Connolly, 1941 Figs 182-184
1941 Lejeania aperta Connolly. — Brit. Mus. (Nat. Hist.) Exped. S.-W. Arabia 1937-8, Vol. 1: 22, pl. 3, figs 4-6, textfig. 4
(North slope of Jabal Jalal, above Naqil Isla pass, c. 9900 ft).
Type material: Three syntypes Lejeania aperta, two of them adult shells, BM(NH)
1939.4.19.127-129.
Description: The thin shell is depressed with an only scarcely elevated spire. The proto-
conch consists of 2.5 whorls and is sculptured by axial riblets. The teleoconch whorls are fine and
irregularly ribbed. The complete surface of the shell is covered by a dense sculpture of rounded
granules which serve as a basic structure for the periostracal bristles as seen in the juvenile syntype.

428 E. NEUBERT
The basic colour of the shell is white with a broad medium white spiral band on the periphery.
Between the suture and this medium band up to three additional bands may occur with the two
sutural bands always interrupted by white streaks. In the periomphal region, several brown bands
may occur with intercalated white flames. Overall, the shells have a mottled appearance.
The suture is deep, the last whorl of the teleoconch is rounded and bluntly keeled in juveniles.
The aperture is wide and subelliptical to nearly rounded. The peristome is only slightly
reflected and sharp. The umbilicus is wide and funnel-shaped.
Measurements (illustrated syntype, Figs 182-184): H = 9.5; D = 19.4; PH = 8.6; PD = 9.8;
W = 5.75.
Remarks: The differentiation of both Lejeania species recorded from the Arabian Peninsula
still poses problems. Several specimens of L. leucosticta have more flattened shells. Also the
diameter of the umbilicus varies considerably. Both taxa are treated here as separate species because
in L. aperta the umbilicus is extremely wide and the spire nearly completely flattened.
To date, L. aperta is only known from its type locality and is surrounded by populations of L.
leucosticta. It is possible that this taxon only represents a geographical race or an eco-phenotype of
L. leucosticta.
Family Helicidae
Genus Eremina L. Pfeiffer, 1855
1855 Eremina L. Pfeiffer. — Malakozool. Bl. 2: 139.
Diagnosis: Medium-sized to large depressed to subglobose shells. Usually, the shell walls
are thick, as an adaptation to life in the steppe. The surface is smooth to heavily ribbed. The genus
is widespread in arid areas from Morocco to Somalia.
Type species: Helix desertorum Forskål, 1775.
Eremina desertella (Jickeli, 1872) Figs 187-192
1872 Helix desertella Jickeli. — Nachrbl. dtsch. malakozool. Ges. 4: 62 (“Habitat litora maris rubri ad Habab”).
1881 Helix pisaniformis Bourguignat. — Mollusques terrestres et fluviatiles recueillis en Afrique dans les pays des Çomalis
Medjourtin: 3 (“Çomalis Medjourtin”).
1882 Helix çomaliana Bourguignat. — Mission de G. Révoil au pays Çomalis. Faune et Flore des Pays Çomalis: 8, pl. IV,
figs 67-69 (“au pied des broussailles entre la lacune de Tohen et le cap Gardafui”).
1882 Helix tiani Bourguignat. — Mission de G. Révoil au pays Çomalis. Faune et Flore des Pays Çomalis: 9, pl. IV,
figs 70-71 (“dans les sables, sous les pierres, non loin de la vallée de Tohen”).
1882 Helix tohenica Bourguignat. — Mission de G. Révoil au pays Çomalis. Faune et Flore des Pays Çomalis: 11, pl. IV,
figs 74-76 (“au pieds des roches arides du Cap Guardafui”).
Type material: Three syntypes Helix desertella, SMF 5363 a-c, ex coll. Jickeli; syntype
Helix tohenica, MNHNP.
Description: The shell is depressed conical and very thin compared to its congeners. The
smooth protoconch consists of 1.5 submammillate whorls. The teleoconch whorls are well
rounded with a suture of medium depth. Their surface is sculptured by fine and closely packed
riblets, but appears smooth to the naked eye. Some irregularly scattered small grooves can be found
predominantly on the subsurface of the shell. The last whorl is rounded to bluntly depressed and
descends towards the aperture. The aperture is nearly circular with a fine labial callus inside. The
peristome is only very slightly reflected, but a columellar reflection nearly covers the umbilicus
leaving a rimate elongate hole.
The surface of the teleoconch is basically white with up to four brown spiral bands. The
uppermost subsutural band usually consists of small patches, the remaining bands are solid brown.

Terrestrial and freshwater molluscs of the Arabian Peninsula 429
Figs 187-192: (original size × 2), Eremina desertella. 187-189: Syntype of Helix desertella, SMF 5363 a. 187: Frontal view. 188:
Apical view. 189: Subsurface. 190-192: Syntype of Helix tohenica, MNHNP. 190: Frontal view. 191: Apical view. 192:
Subsurface.
Measurements: Syntype of Helix desertella (Figs 187-189): H = 12.7; D = 19.4; PH = 9.8;
PD = 11.5; W = 4.5. Holotype of Helix tohenica, MNHNP (Figs 190-192): H = 15.0; D = 21.3;
PH = 11.0; PD = 12.2; W = 5.
Remarks: VERDCOURT (1960: 240) synonymises Helix comaliana, Helix tiani, Helix tohenica
and Helix pisaniformis, all described by Bourguignat in 1881 and 1882 with this species. Although
Verdcourt did not investigate the type material, his argument: “From the original figures I can find
no reasons for keeping any of these species distinct” is accepted here and H. tohenica is depicted.
The distribution of Eremina desertella is not well known. It has only been recorded from a
narrow stretch along the coast of the Red Sea and the Gulf of Aden southwards to Cape Guardafui
in Somalia. There are several records of this species by JOUSSEAUME (1889: 353) from Arabia. Two
are published under the name H. desertella and come from Jeddah and near the port of Loheiyah.
Accepting Verdcourt’s point of view, Jousseaume’s records for Helix tohenica from Aden and
Makalla may be added. It is remarkable that this species has never been recorded since. Introduc-
tion by human activities is likely and the coastal plains of the south-western Arabian Peninsula
should be investigated more thoroughly. In general, the habitat structure should support a
population of E. desertella.
Genus Eobania Hesse, 1913
1913 Eobania Hesse. — Nachrbl. dtsch. malakozool. Ges. 45: 13.
Diagnosis: Big helicoid shells with a depressed spire. The BC with an elongate diverticu-
lum. One very small dart sac. The flagellum is shorter than penis and epiphallus.
Type species: Helix vermiculata O.F. Müller, 1774.

430 E. NEUBERT
Eobania vermiculata (O.F. Müller, 1774)
1774 Helix vermiculata O.F. Müller. — Vermium terrestrium et fluviatilium 2: 20.
Material: Saudi Arabia: Asir mountain region, Taif, residential compound gardens, VII/VIII.1995, S. Newton,
SNMNH-MO 132/11, SMF 311666/3.
Description: The thick-walled shell of this species reaches a diameter of up to 30 mm. It is
depressed to low spherical. The whorls of the teleoconch are rounded with a shallow suture. The
last whorl descends abruptly below the periphery of the shell. The aperture is obliquely ovoid, the
peristome is somewhat reflected. Inside the aperture, a lip is present, which is strengthened at the
columellar to basal margin. The umbilicus is covered by a white porcelaineous callus.
Generally, the shell is creamy white to faint yellow. There are up to five brown spiral bands,
which are often fused. Nearly maroon brown specimens may occur. Usually, white axial flames
interrupt the bands thus producing an alternating pattern of bright and dark colours on the surface
of the shell.
Remarks: Eobania vermiculata is a well-known circum-Mediterranean species. It was
hitherto only recorded from Wadi Hanifa in central Saudi Arabia (MORDAN 1980 b: 364). This
additional record is not surprising since this species is well known for its survival capability and
has been introduced to many countries by human activity.
Genus Levantina Kobelt, 1871
1871 Levantina Kobelt. — Cat. Europ. Binnenconch.: 19.
Diagnosis: The medium-sized to large shells of this genus have a depressed to low conical
spire. Usually, juvenile shells display a sharp to blunt keel which becomes obsolete on the last third
of the body whorl of an adult shell. The flagellum is heavily coiled, the glandulae mucosae are very
small, branching of the glandulae is rare.
Type species: Helix spiriplana Olivier, 1801.
Subgenus Laevihelix n. subgen.
Diagnosis: A subgenus of Levantina with thin-walled shells, which are densely covered by a fine
granulation.
Description: This subgenus is characterised by its thin, light and corneous shells. Usually,
the surface of the protoconch and teleoconch is covered by a densely packed fine granulation. On
the protoconch, the granules are usually arranged in oblique axial rows. Juvenile shells are sharply
keeled, the suture of the teleoconch whorls is moderately deep.
The shells are smaller than in species of Levantina s. str., which usually have thicker and
“chalky” shells. If present, the granules are coarser and more irregularly arranged. Juveniles of
Levantina s. str. have flat and sharply keeled whorls, the suture of the upper whorls is always very
shallow and often the keels surmount the preceding whorl. Levantina (Laevihelix) differs from
Levantina (Gyrostomella) Hesse, 1908 by its colour, the lack of any radial sculptural elements and
the length of the flagellum.
Type species (by present designation): Levantina (Laevihelix) symensi n. sp.
Etymology: This subgenus is called Laevihelix as an illustration of the characteristically
thin-walled shells.
Remarks: Laevihelix is classified with Levantina as anatomical details of the genital organs
prove their close relationship. The coiled flagellum and the reduced glandulae mucosae are

Terrestrial and freshwater molluscs of the Arabian Peninsula 431
apomorphic character states of the genera Levantina and Assyriella Hesse, 1908. Apart from
conchological characters, Assyriella differs in having a diverticulum which is considerably longer
than in Levantina, and in the wider coiling of the flagellum.
It has to be stressed that the adaptive value of most of the character states used for supraspecific
definitions in the Helicinae is insufficiently known. Although SUBAI (1994) and SCHÜTT & SUBAI
(1996) discussed anatomical characters of Levantina, Assyriella, Codringtonia Kobelt, 1898, and
Isaurica Kobelt, 1901, many of the more than 20 genera of the Helicinae are still in need of
intensive investigation. These are necessary to sort out plesiomorphic and apomorphic character
states to establish a more phylogenetically based system for the whole group.
The first to mention the genus Levantina from Arabia was MORDAN (1980 b). He found two
species of this genus living in the south-west of Saudi Arabia, Levantina cf. caesareana and
Levantina sp. New collections from this area revealed that there are four species involved, all of
which are new to science. It is likely that there are more species to be found, in particular in the
northern mountainous belt of the Red Sea coast.
The classification of these species within the genus Levantina is based on conchological and
genital morphological characters. Juvenile shells of these species are keeled, as is typical for the
Levantina group. The genital morphology of three of the species was investigated. The flagellum is
extremely long and heavily coiled. The pedunculus with the BC is always shorter than the
diverticulum, but does not reach the relative length that can be seen in Assyriella.
These characters agree very well with the diagnosis of Levantina given by SUBAI (1994: 59).
The penial and epiphallial papillae and pilasters in the atrium and vagina were also investigated,
but did not reveal any suitable information. These characters vary intraspecifically (in the three
anatomically investigated Arabian species) to such an extent that they are of no use for systematic
purposes (for this problem cf. SZIGETHY 1976).
Levantina (Laevihelix) symensi n. sp. Figs 193-195, 205-206, 211
Type material and locus typicus: holotype SMF 311271, paratypes SNMNH-MO 133/1 a and b (a = shell,
b = preserved animal), SMF 311272/3 a and b (a = shell, b = preserved animal), SNMNH-MO 134/2 (and many fragments),
NEUB (1), Saudi Arabia, Asir mountain region, Jabal Dhaka SW of Taif, 2340 m, Juniperus forest, 4.VI.1995, E. Neubert et al.
— Non-type material: Saudi Arabia: Hijaz, Barrad, about 40 miles east of Mecca at about 4200 ft, II.1948,
Meinertzhagen, BM(NH) 1948.11.12.11-18 (8); Asir mountain region, al-Shafa, SW of Taif, 2040 m, 4.VI.1995, E. Neubert et
al., SNMNH-MO 135/12, SMF 311667/12; Asir mountain region, al-Hada escarpment, 2000 m, VII.1994, S. Newton,
SNMNH-MO 136/2; Asir mountain region, al-Hada escarpment, 2000 m, VII.1995, S. Newton, SNMNH-MO 137/8
(+ fragments), SMF 311668/8.
Diagnosis: A member of the genus Levantina (Laevihelix) with a comparatively small shell.
Its umbilicus is closed, the peristomial rims are not connected.
Description: The low conical subglobose shells are medium-sized with five whorls. The
protoconch consists of 1.5 whorls. Its surface is finely granulated, the granules are often arranged
in axial rows. This granulation continues over the rest of the whorls, but is less pronounced on the
surface of the last whorl. It nearly disappears on the subsurface. The suture of the protoconch is
deep, but the preceding suture of the juvenile shell is very shallow reflecting the somewhat
nodulose keel. The suture of the body whorl is deeper again, restricting the keel to the upper three
whorls of the shell. The last whorl is well rounded and bent downwards towards the peristome.
There is no additional surface sculpture apart from the granules, but the growth lines
sometimes may be slightly pronounced. The basic colour of the shells is light brown with five
maroon brown bands. The bands are visible in all shells and always well separated. On the upper
whorls, the bands are interrupted by small yellow zigzag flames, a pattern which is restricted to the
subsutural area on the last whorl.

432 E. NEUBERT

Terrestrial and freshwater molluscs of the Arabian Peninsula 433
The aperture of the shell is oblique elongated, the upper rim of the peristome is not connected
to the columellar reflection. The umbilicus is closed by an expansion of the peristomial shield. The
peristome is reflected and bears a slightly thickened white lip.
Genital morphology (five specimens investigated, holotype SMF 311271 illustrated,
Figs 205-206): The penis is conical with a slight constriction where the penial papilla is situated.
The epiphallial papilla is as long as the penial papilla. The proximal part of the penis is slender, the
penial retractor inserts at the boundary of the penis and the epiphallus. The flagellum is more than
twice as long as the penis and epiphallus. It is spirally coiled.
The vagina is shorter than the penis. The glandulae mucosae consist of two pairs of slender
tubes which have only one secondary branch. The diverticulum is longer than the BC and the
pedunculus.
The dart sac nearly reaches the length of the glandulae mucosae. The dart is slender and nearly
straight. Proximally, the four blades are equally sized reaching their maximum width in the middle
of the dart. Distally, the height of two opposite blades decreases.
Measurements (holotype, Figs 193-195): H = 14.2; D = 28.0; PH = 14.2; PD = 16.5;
W = 5.
Etymology: This new species is named in honour of the Belgian ornithologist Peter
Symens, who was the first to find living specimens (L. asagittata n. sp.) of this subgenus.
Note: Very probably, this species had already been mentioned by MORDAN (1980 b: 364,
fig. 2A) under the name Levantina cf. caesareana (Mousson, 1854) from Barrad, 40 miles E of
Mecca and from between Jeddah and Taif.
Levantina (Laevihelix) asagittata n. sp. Figs 196-198, 207-208, 211
Type material and locus typicus: holotype SMF 311273, paratypes SNMNH-MO 138/3 a and b (a = shell,
b = preserved animals) SMF 311274/8 a and b (a = shell, b = preserved animals), Saudi Arabia, Asir mountain region, 15 rkm S
of Bani Sa’ad, S of Taif, 2300 m, Juniperus forest, 2.VI.1995, E. Neubert et al. The specimens from the following lot are also
considered to be paratypes: Saudi Arabia, Asir mountain region, 15 rkm S of Bani Sa’ad, S of Taif, 2300 m, Juniperus forest,
28.I.1994, P. Symens & M. Werner, SNMNH-MO 139/1, SNMNH-MO 140/2 (preserved), SMF 311669/1 (preserved animal
of SNMNH-MO 139/1), NEUB (1 shell, 1 preserved animal).
Diagnosis: A species of the genus Levantina (Laevihelix) which resembles L. (L.) symensi
n. sp. conchologically but the complete dart system (i.e. dart sac and glandulae mucosae) is
missing.
Description: The shell is subglobose, the spire more elevated than in L. (L.) symensi n. sp.
The shells have five whorls, the last whorl is well rounded and bent downwards to the aperture.
The protoconch consists of 1.5-2 whorls and is covered by granules, often arranged in small riblets.
The surface of the preceding whorls is also covered by granules which in almost all shells disappear
on the last two thirds of the last whorl. The surface of the last whorl is almost smooth besides the
growth lines. In some shells, faint subsutural spiral grooves can be seen on the last whorl. The
subsurface is smooth in adult shells; in juveniles, the granulation may cover the complete shell.
The suture is moderately deep, juvenile shells are keeled.
Figs 193-204: (original size × 2), 193-195: Levantina (Laevihelix) symensi n. sp., holotype SMF 311271. 193: Frontal view. 194:
Apical view. 195: Subsurface. 196-198: Levantina (Laevihelix) asagittata n. sp., holotype SMF 311273. 196: Frontal view. 197:
Apical view. 198: Subsurface. 199-201: Levantina (Laevihelix) asira n. sp., holotype SMF 311275. 199: Frontal view. 200: Apical
view. 201: Subsurface. 202-204: Levantina (Laevihelix) semitecta n. sp., holotype BM(NH) 198064. 202: Frontal view. 203:
Apical view. 204: Subsurface.

434 E. NEUBERT
The aperture is oblique to subcircular, the peristome is slightly reflected. The umbilical region
of some shells is slightly excavated, the umbilicus is completely covered by a columellar shield. The
lip is thin and shows only a weak callosity. The rims of the peristome are not connected.
The basic colour of the shell is light brown, the deep brown spiral bands are always well
visible. The zigzag pattern of yellow flames is present and extends in many shells to the subsurface,
interrupting the lowest spiral band.
Genital morphology (13 specimens investigated, holotype SMF 311273 illustrated,
Figs 207-208): The penis is conical, widening distally. The boundary between the proximal and
distal parts of the penis is not visible. The penial retractor inserts at the boundary of penis and
epiphallus. The flagellum is very long, reaching more than twice the length of the penis and
epiphallus. The penial papilla is cone-shaped with transverse lamella as the small epiphallial papilla.
The vagina is short, dart sac and glandulae mucosae are missing completely. This character
was constant in all specimens investigated (two specimens were collected in 1994, 11 specimens
in 1995).
Measurements (holotype, Figs 196-198): H = 13.2; D = 26.0; PH = 14.3; PD = 14.4;
W = 5.
Etymology: This species is named for the missing dart apparatus.
Remarks: All specimens were found in a bushy Juniperus forest under crystalline boulders.
All specimens were actively creeping.
On the one hand, the conchological differences between L. (L.) symensi n. sp. and L. (L.)
asagittata n. sp. are slight and thus separation based on conchological characters is not possible.
On the other hand, the absence of the complete dart apparatus seems to be a constant feature in L.
(L.) asagittata n. sp. The dart sac and the glandulae mucosae play an important role in the
reproductive behaviour of the Helicoidea. It is one of the outstanding autapomorphies of this
group. It can be hypothesised that the complete loss of such a substantial anatomical detail will
cause a completely different mating behaviour. This is a very good example of a reproductive
barrier justifying the specific status of a taxon.
Levantina (Laevihelix) asira n. sp. Figs 199-201, 209-211
Type material and locus typicus: holotype SMF 311275 (with preserved animal), paratypes SNMNH-
MO 141/6, SMF 311276/1, NEUB (1 shell and 1 preserved animal), Saudi Arabia, Asir mountain region, small wadi at the
right side of the road from Taif to al-Baha, S of Gharie, in dense vegetation and under the bark of shrubs and small trees,
1700 m, 2.VI.1995, E. Neubert et al. — N on-type material: Saudi Arabia: Asir mountain region, al-Abna descent
near Baha, 2000 m, 3.VI.1995, E. Neubert et al., SNMNH-MO 142/2 (2 fragments).
Diagnosis: A member of the genus Levantina (Laevihelix) with a flattened spire. The
surface is granulose, the umbilicus open and only partly covered by the reflected peristomial rim.
The peristome is callose, interconnected and detached from the last whorl.
Description: The shell is medium-sized with a depressed, nearly flattened spire. The shell
consists of 4.5-5 whorls. The protoconch has 1.5-2 whorls, its surface is covered by numerous
small granules. On the first protoconch whorl, the granulose sculpture is less pronounced, small
axial riblets are more prominent. On the subsequent whorls, the granulose sculpture increases in
strength, covers the surface and subsurface of the shell and is visible even in the umbilicus. The
granules are prominent and can be seen by the naked eye.
The suture of the complete shell is moderately deep. Juvenile shells are keeled, the keel is
subnodulose. The last whorl descends towards the peristome. The aperture is enlarged and very
oblique. The peristome is reflected and thickened by a callose porcelaineous strong lip. The
peristomial callus is white. Upper and lower part of the peristome are connected and detached

Terrestrial and freshwater molluscs of the Arabian Peninsula 435
Figs 205-210: Genital organs. 205-206: Levantina (Laevihelix) symensi n. sp., paratype. 205: Genital apparatus. 206: Penis
dissected, showing penial papillae. 207-208: Levantina (Laevihelix) asagittata n. sp., paratype. 207: Genital apparatus. 208: Penis
dissected, showing penial papillae. 209-210: Levantina (Laevihelix) asira n. sp., paratype. 209: Genital apparatus. 210: Penis
dissected, showing penial papillae.

436 E. NEUBERT
from the last whorl. On the columellar side, the peristomial shield is partly reflected over the
umbilicus. The umbilicus is always open.
The basic colour of the shell is brown, the five spiral bands are not as obvious as in L. (L.)
symensi n. sp. or L. (L.) asagittata n. sp. The zigzag pattern of yellow flames is present on the upper
whorls; on the surface of the last whorl, it reaches the periphery of the shell.
Genital morphology (one paratype specimen investigated, Figs 209-210): Penis sub-
cylindrical, the boundary between the proximal and distal part of the penis not easily visible. The
flagellum is spirally coiled and reaches more than twice the length of the penis and epiphallus. The
penial retractor inserts at the boundary of the epiphallus and penis. The penial papilla is short and
somewhat rugose similar to the epiphallial papilla. Two Y-shaped pilasters surround the osculum
of the epiphallus (as in the two other species investigated).
The vagina is shorter than the penis. The glandulae mucosae are small with only very few
secondary branches. The diverticulum is very long. Unfortunately, the pedunculus was lost during
preparation. The dart is very close to that described in Levantina (Laevihelix) symensi n. sp.
Measurements (holotype, Figs 199-201): H = 11.2; D = 27.8; PH = 14.8; PD = 18.9;
W = 5.
Etymology: The species name is derived from the name of the typical area, the Asir
mountain region.
Remarks: Two specimens were found alive, clinging under the bark of a tree. They were
fixed to the wood by an extraordinarily strong diaphragm.
Levantina (Laevihelix) semitecta n. sp. Figs 202-204, 211
Type material and locus typicus: The type lot contained two specimens BM(NH) 198064. The original text
of the label reads: “Midian. Collected and presented by Capt. Burton”. A second label reads: “Label in E.A. Smith’s writing. If
this is Sir Richard Burton then prob. c. 1877 when he was surveying in area N.E. of Red Sea”.
Diagnosis: A small species of Levantina (Laevihelix) with a partly covered umbilicus.
Description: The shell is low-spired and depressed. Compared to the teleoconch, the
protoconch is quite large. It is finely rugose and bears faint granules. The teleoconch whorls are
rounded, the suture is deep. The last whorl is bluntly angulate and abruptly descends under the
periphery of the body whorl. The surface of the teleoconch bears fine axial riblets, suprasuturally
traces of granules are discernible. Both shells are greyish white and show the typical zigzag axial
flames. Presumably, the shells once had a light brown coloration.
The aperture is subcircular, the peristomial rim is reflected. On the columellar margin, the
reflection widens to form a shield covering half of the umbilicus and the umbilical area. The
umbilicus is open and narrow.
Measurements (holotype, Figs 202-204): H = 9.9; D = 17.9; PH = 9.1; PD = 11.1;
W = 4.5.
Etymology: This species is called L. (L.) semitecta for the characteristic shield covering half
of the umbilicus.
Remarks: Midian is the name of a landscape in the Northern Hijaz in the north-west of
Saudi Arabia which stretches from the area south of Jabal al-Lawz to al-Wagh at the Red Sea shore.
The boundaries of the area are not precisely defined, but they enclose a chain of mountains of
more than 200 km length and altitudes of up to 2350 m (Jabal al-Dubbagh). The substratum is
formed by basaltic rocks or sandstone. The environment seems to be similar to what is known
from habitat characteristics for the other Levantina (Laevihelix) species and is in contrast to the
habitat of Levantina s. str., which seems to be calciphilic. This species was already mentioned and
illustrated by MORDAN (1980 b: 364, fig. 2C).

Terrestrial and freshwater molluscs of the Arabian Peninsula 437
Fig. 211: Distribution of Levantina (Laevihelix) spp. on the Arabian Peninsula.
It would be very interesting to study the area and the animals in more detail. The question
arises as to whether L. (L.) asagittata n. sp. and L. (L.) symensi n. sp. live allopatrically or whether
there are sympatric populations. An investigation of the mating behaviour is urgently needed to
obtain more information about how a asagittate species of a usually sagittate group will mate.
A map displaying the distribution of Levantina in Arabia is supplied for a better understanding
of the remarkable expansion of this genus towards the south (Fig. 211).
Genus Cantareus Risso, 1826
1826 Cantareus Risso. — Histoire naturelle de l’Europe méridionale 4: 64.
Description: Medium-sized to large globose shells with the last whorl enlarged. Sculpture
of the teleoconch whorls malleate. There is only one penial papilla projecting into the proximal
lumen of the distal part of the penis. This part is characterised by an annular pad and an additional
“false papilla” which inserts laterally on the penial wall (SCHILEYKO 1978: 324, GIUSTI et al. 1995:
491, figs 616-622).
Type species: Helix naticoides Draparnaud, 1801 (= Helix aperta Born, 1778).

438 E. NEUBERT
Cantareus aspersus (O.F. Müller, 1774)
1774 Helix aspersa O.F. Müller. — Vermium terrestrium et fluviatilium 2: 59 (“In Italia”).
Material: Saudi Arabia: Asir mountain region, Taif, residential compound gardens, VII./VIII.1995, S. Newton,
SNMNH-MO 143/1.
Description: The shell of this species is depressed globose with an inflated voluminous last
whorl. Compared to most of the Helix species, the shell of C. aspersa is thin and more fragile. The
basic colour is yellow to light brown, but this coloration is nearly indiscernible as it is dominated
by dark spiral bands. The subsutural band is narrow and followed by the second and third which
are usually fused, forming a broad dark and compact spiral above the periphery. The last two
spirals are usually narrow and separate. This coloration pattern varies, completely dark shells or
shells lacking any spiral frequently occur. Irregular yellow zigzag flames may also be observed.
The smooth protoconch is very small and consists of almost two whorls. The teleoconch suture
is somewhat impressed. The teleoconch whorls are sculptured by rugose axial ribs which are
irregularly spaced. Most characteristic is the malleate to reticulate pattern on the surface of the shell.
The aperture is wide oval to rounded with a somewhat flared white peristome. A weak labial
callus is present. In adult specimens, the umbilicus is closed by a triangular reflection of the colu-
mellar peristomial rim. Internally, the colour of the aperture varies from white to opaque bluish.
Remarks: As in E. vermiculata, this species which is native to the western Mediterranean
area has been dispersed by human activities all over the world. Nonetheless, this is the first record
for this species from the Arabian Peninsula.
Bivalvia
Family Corbiculidae
Genus Corbicula Megerle von Mühlfeldt, 1811
1811 Corbicula Megerle von Mühlfeldt. — Ges. Naturf. Fr. Berlin, Mag. V (1): 56.
Diagnosis: Small trigonal heterodont freshwater mussels. Slightly anisolateral with three
cardinals in each valve. Inner margin of laterals finely denticulate. Ligament external, nymph
coarsely ridged. A pallial sinus is absent. Surface sculptured by fine to heavy concentric ribs.
Type species: Tellina fluminalis O.F. Müller, 1774.
Corbicula purpurea Prime, 1863
1863 Corbicula purpurea Prime. — Ann. Lyc. Nat. Hist. New York 8: 77 (“In flumine Tigris, Asiae”).
1874 ? Cyrena (Corbicula) Tigridis Mousson. — J. Conchyl. 22: 55 (“à Tekrit sur le Tigre”).
Material: Saudi Arabia: Eastern Province, al-Qatif Oasis, palm plantations W of centre, irrigation channels,
14.VI.1992, R. Kinzelbach, (7); al-Qatif Oasis, freshwater course W of town, 26°36'N 49°58'E, 31.I.1992, R. Kinzelbach &
I.A. Nader, (27); al-Hasa Oasis, al-’Uyun palm plantation, freshwater ditch with dense vegetation, 25°36'N 49°34'E, 27.I.1992,
R. Kinzelbach & M. Werner, (1); al-Hasa Oasis, al-’Uyun palm plantation, loam excavated from ditches, 25°36'N 49°34'E,
21.I.1992, R. Kinzelbach, E. Neubert & W. Schneider, (4).
Type material: The type designation by JOHNSON (1959) for this taxon is incorrect, a new
lectotype designation will be published by Neubert & Zajonz (in prep.).
Description: The valves are broadly trigonal and somewhat depressed. The umbos are
small and tinged violet with a dark triangle fading at its base. The ribs are fine and dense compared
to other species, in particular to the often syntopic C. fluminalis (O.F. Müller, 1774).
Remarks: Corbicula tigridis is included here provisionally in the synonymy of C. purpurea.
Very probably, the type material of this taxon is lost (T. Meier, pers. comm. 1997). The description
given by Mousson is very clear as he states “late rotundato-trigona […] extus transverse inaequaliter

Terrestrial and freshwater molluscs of the Arabian Peninsula 439
costulata, fulvescens, intus albo-caerulescens. Umbones parvuli, minus producti, nec inflati, antice
parum deflexi …”. This description matches the material and the syntypes of C. purpurea very well.
Under the name Corbicula fluminalis, BROWN & WRIGHT (1980: 354) had previously recorded
a Corbicula species for the Arabian Peninsula (al-Qatif and al-Hasa Oases). This material was not
checked, but its conspecificity is very likely as the material investigated here principally originates
from the same area.
Family Sphaeriidae
Genus Pisidium C. Pfeiffer, 1821
1821 Pisidium C. Pfeiffer. — Naturg. dtsch. Land-Moll. (1) 17: 123.
Diagnosis: Small lentil-shaped freshwater bivalves. The usually thin-walled shells are
anisolateral.
Type species: Tellina amnica O.F. Müller, 1774.
Pisidium casertanum (Poli, 1791)
1791 Cardium casertanum Poli. — Test. Sicil. II: 65.
Description: This species reaches a length of up to 4 mm. It is ovoid to subtrigonal, an
umbonal sculpture is absent. The surface is smooth, apart from concentric growth lines, and glossy
in fresh specimens. Very often, red-brown encrustations may be present. The cardinals of the right
valve are fused, those of the left valve small and close together.
Remarks: This ubiquitous species is widespread and seems to be introduced to suitable
freshwater courses easily. It was recorded for the Arabian Peninsula from the south-western area at
“Sadah stream at over 3.100 m in the Khamis Mushayi-Abha area” (BROWN & WRIGHT 1980: 354)
and “Yemen, Birkat Ghail Masnah, a pond south-west of Ma’abar, 8.400 ft” (CONNOLLY 1941: 35).
DISCUSSION
The checklist presented here contains 70 species of land molluscs and 27 species of freshwater
molluscs. Bearing in mind that an area of more than 3 million square kilometres (approximately
80 % of the surface of the Indian subcontinent) is discussed, this number is remarkably low. This
is caused by geological and climatological factors as well as insufficient knowledge of many parts
of the Arabian Peninsula from a malacological point of view. In particular, the mountain belts of
western Arabia are awaiting further malacological exploration. More information is available
concerning the southern regions of Yemen, the Dhofar and Hadramaut region, and the very
important Oman Mountains.
Distribution patterns of terrestrial and freshwater organisms generally reflect the geological
and climatological history of the area concerned. The interpretation of these patterns is one of the
most interesting conclusions to be drawn from data now available. It may serve as confirmation
and a reconstruction of geological events, climatic changes and, last but not least, even human
activities. In this context, molluscs play a prominent but widely neglected role as they are slow
moving, non-vagile animals. Their dispersion velocity equals geological time scales much more
than that of mobile animals like birds, and the analysis of distributional patterns gives an indirect
view of geological processes. For example, the distribution of the Acavidae, a family of landsnails
known from eastern South America, southern Africa, Madagascar, the Seychelles and Sri Lanka,

440 E. NEUBERT
reflects a typical Gondwana type distribution and provides information for both the fields of
geology and evolution. Moreover, the time scales of evolution within systematic entities become
visible and can be calibrated by geological processes. To prevent misinterpretation, the taxonomy
of the species concerned, their phyletic relationships and true distribution must be clarified. Often,
this is hampered by the existence of a over-abundance of available unrevised names for only a few
“biological” species, a problem which led to the present intensive systematic study of most of the
taxa living in Arabia.
In the following paragraphs, an analysis of the distribution patterns of the terrestrial and
freshwater malacofauna of the Arabian Peninsula from a zoogeographical point of view is pre-
sented. To facilitate the understanding of the results, brief outlines of the geological background
and the recent climatological structure are given.
A brief outline of the geological history of the Arabian Peninsula
The Arabian Peninsula forms part of the African plate from which it is separated by the Red Sea.
It is characterised by Precambrian volcanic rocks with large areas covered by younger sediments.
Plutonic intrusions have broken through the crust in several areas. Starting at the Cambrian, the
Tethys covered large areas of the eastern peninsula, accumulating sediments of several thousand
metres thickness. These conditions persisted throughout the Palaeozoic and Mesozoic times. At the
beginning of the Cenozoic, the development of the two basic geological provinces of the Arabian
Peninsula, the Arabian Shield and the Arabian Shelf, must be considered in parallel, as one is
substantially influenced by the formation of the Red Sea, the other by the regression of the Tethys
(Chapman in AL-SAYAD & ZÖTL 1978: 5).
In the late Cretaceous or early Eocene (60-50 million years b.p.), the Red Sea trough, the
northern elongation of the east African rift valley, opened for the first time caused by the rotation
of the Arabian plate. The basin was filled from the north by Tethyian waters and ended at a land
bridge between Africa and Arabia in the south. The first breakthrough connecting the Red Sea to
the ancient Indian Ocean took place in the late Oligocene 30 million years b.p. and lasted to the
early Miocene approximately 15 million years later. After this period, sea-floor spreading ceased
and evaporites indicate a partial or complete regression of the sea. At least in the shallow southern
parts, the former land bridge re-established. The sea-floor spreading started again in the Pliocene
about 5 million years b.p. lasting to Recent times (FRISCH & LOESCHKE 1993: 40). During the last
glaciation, the global sea level dropped approximately 100 m below the Recent level. Water fluxes
may have been severely restrained at this time, causing a salinity crisis in the interior basin of the
Red Sea. It is not clear whether a shallow water body or hypersaline swamps were present at Bab
el-Mandeb then, but the existence of a land bridge to be used by land molluscs is highly improbable.
The Arabian Shield represents the basic Precambrian African craton and stretches parallel to
the Red Sea coast extending eastwards into the interior of the peninsula. It consists of gneiss and
metamorphosed sedimentary and volcanic rocks. The western Arabian mountain belt is the result
of several stepwise elevations and reaches its maximum height in Yemen, at an altitude of 3760 m
contrasting to the low coastal plain, the Tihama. To the north, the altitude decreases, but never
drops much below 1000 m.
The Arabian Shelf comprises two thirds of the surface of the peninsula. Its foundation is
formed by the same Precambrian plate as in the Arabian Shield, but it is covered by a sequence of
continental and shallow-water marine sedimentary rocks. The thickness of this layer increases
eastwards and reaches more than 10,000 m beneath the Arabian Gulf. The opening of the Red Sea
was paralleled by an uplift of the coastal mountain belt, a gentle dipping of the complete Arabian

Terrestrial and freshwater molluscs of the Arabian Peninsula 441
plate, its anti-clockwise rotation and subduction under the Iranian subplate. This process was
accompanied by a stepwise regression of the Tethys disconnecting the ancient Indian Ocean from
what is now the Mediterranean Sea. In the middle Miocene, a marine transgression affected the
coastal plains of the peninsula but the interior remained terrestrial until the Present (POWERS et al.
1966: 20). During the second stage of alpine orogeny at the transition from Miocene to Pliocene,
the strata described were uplifted and folded, forming the mountains of the Zagros (opposite the
Arabian coast) and the Oman Mountains.
The nature of the Arabian Gulf as a subduction zone, where the Arabian plate gently dips
under the Iranian subplate of the Eurasian plate, is displayed by typical features such as a folded
mountain belt and a small island arc (FRISCH & LOESCHKE 1993: 94, figs 7-1). The sediment
volumes transported by the freshwater tributaries are insufficient to balance this process and fill
the basin. During the late Pliocene, several intervals of transgression are recorded up to 70 m above
recent sea level, as witnessed by huge inland Sabkhas in the Eastern Province of Saudi Arabia (Zötl
in AL-SAYAD & ZÖTL 1978: 290). In general, sea level fluctuations persisted until the Recent
period. The consequences are dramatic, as during glaciation periods amplitudes of 100 m (which
are said to have occurred several times) caused a nearly complete withdrawal of the sea from the
Arabian Gulf (Felber et al. in AL-SAYAD & ZÖTL 1978: 50 f.). Calculating the late Pleistocene
amplitudes at between 50 and 100 m, the shoreline and the delta of the Shatt al-Arab would have
been situated between Jubail (N of Dhahran) and the Strait of Hormuz.
Climatological factors
The major climatological factors reflect both the global position of the area and its particular
geomorphologic structure. The peninsula is situated in one of the major desert belts of the world,
reaching from the Cape Verde Islands in the west to the Thar Desert in India. It stretches within
latitude 32° to 12°N and longitude 36° to 60°E. Within this huge area of land, continental climate
can be expected. Concerning temperature, a gradient from north to south is observed. Thus, the
mean temperature values increase from 8 °C to approximately 20 °C in winter and from 27 °C to
approximately 36 °C during summer. These values may be exceeded considerably, with maximum
temperatures of up to 48 °C in summer and minimum temperatures below 0 °C in winter. In
general, the mountainous areas experience lower temperatures.
Rainfall variability is extremely high, as in all desert regions. Sporadic heavy rains may occur
in unpredictable intervals, as reflected in annual precipitation measured in Dhahran with 5.3 mm
in 1946 and 186.9 mm in 1974. On average, a mean precipitation exceeding 1000 mm per year
can only be found in the south-western mountains as these are influenced by the Indian Ocean
monsoons. Dhofar and Oman Mountains also receive a considerable amount of precipitation
(> 200 mm annually). The Tihama, as well as most other parts of the Arabian Peninsula, remain
dry with values of 100-200 mm per year. Only in the area of the Rub al-Khali, rainfall may be less
than 50 mm per year.
Origin of the Arabian malacofauna
It has to be stressed that Arabia is not an important centre of dispersal – unlike for example the
Alpine region, the Caucasus or the Maghreb region – as shown by its low degree of endemism at a
generic level. Its malacofauna is composed of several groups which invaded the peninsula at
different time levels and originated from different zoogeographical units. The groups are still
represented by species which show a similar distribution pattern. These are the Palaearctic, the

442 E. NEUBERT
Afrotropical and the Saharo-Sindian group. As a fourth group, species introduced by human
activities during historical times may be added. To facilitate access to finer structures of the Arabian
malacofauna, a table is given which puts most of the species in a zoogeographical context (Table 2).
The Palaearctic group
This group comprises species and genera which are well-known representatives of the western
Palaearctic. Within this group, several faunal lineages may be observed.
The first is the Mesopotamian fauna, which invaded the peninsula via the Euphrates/Tigris
valley. It is still to be found in lower Mesopotamia (Iraq), its delimitation in the north is not well
defined but may be found in the arc of the eastern Anatolian mountains. A few remnant species
are left on the Arabian coast and presumably in the valleys of the Iranian Zagros mountains too.
The second subgroup contains two endemic species of the Buliminidae living exclusively in
the Oman Mountains. They are closely related to the fauna found in the inner-Asiatic steppe areas
and can be characterised as Irano-Turanian elements.
A third group, the Levantinian fauna, spreads from Syria and Jordan to the south. At a specific
level, only Xeropicta krynickii is present throughout the Red Sea mountain belt. The others are
typical members of the Syrian malacofauna and are restricted to the northern region of Saudi
Arabia. The two species of Paramastus are endemic to southern Arabia, while in Levantina, a
separate diversification at a subgeneric level took place in the Arabian mountains. The distri-
butional gap shown for all Levantinian taxa in the northern and middle Arabian mountains
presumably represents an artefact caused by missing information.
The fourth subgroup comprises phylogenetically “old” species which are widespread in the
western Palaearctic and are known to represent, or to be close to, stem groups of more recently
evolved taxa. This is true for the freshwater species which are sporadically recorded from Arabia. It
is questionable whether they are allochthonous in the area and, owing to the general deserti-
fication, they now form a secondary isolated group or whether they have been introduced
accidentally by roosting birds during migration periods. The terrestrial snails, however, certainly
represent species who invaded the area by natural means. In particular, Lauria cylindracea and
Granopupa granum are widespread in the southern part of the western Palaearctic and are supposed
to be close to the hypothetical stem forms of their groups. This is similar to Cecilioides, where C.
acicula and C. tumulorum are widespread taxa often occurring sympatrically in the eastern
Mediterranean area. Only C. isseli is an endemic species for the south-western Arabian Peninsula.
Another subgroup of ancient Palaearctic taxa is formed by Vitrinidae and Clausiliidae. The
genera Arabivitrina and Macroptychia are speciose in the adjacent areas of the southern Red Sea. As
proven by anatomical and shell morphological details, their nearest relatives still survive on the
Macronesian Islands (Insulivitrina, Madeirovitrina and Boettgeria). Lejeania is close to Euomphalia
and thus shows Caucasian influence, as the diversification centre of this group is the Caucasian
mountains. The present status of Eremina desertella in Arabia remains unsolved as its records from
the southern Tihama need to be reconfirmed. This species ranges at least from Eritrea to Cape
Guardafui in Somalia.
The Afrotropical group
The Afrotropical fauna is well represented in Arabia. It can roughly be subdivided into two
subgroups, one which stems from the Ethiopian Highlands, the other is of tropical origin. The
function of the Ethiopian Highlands for diversification processes is not clear yet, as this area has
not been investigated sufficiently. From the data available, it is evident that it harbours a speciose

Terrestrial and freshwater molluscs of the Arabian Peninsula 443
and in parts endemic malacofauna. About 15 % of the total Arabian malacofauna has close
relationships to this area. In contrast to the much smaller contribution of the Afrotropical fauna,
only terrestrial gastropods are involved.
Afrotropical species were represented by five freshwater and three terrestrial snails only. This
group contains the well-known vector species of schistosomiasis (Bulinus spp.). The distribution
pattern of Quickia is interesting, as it is not only known from tropical Africa but also from several
Indopacific islands and the Indian subcontinent.
The Saharo-Sindian group
This group comprises species which are known from a vast area from northern Africa, the Arabian
Peninsula to the Middle East and northern India. It is composed of the holo-Saharo-Sindian group
containing Pupoides coenopictus and Zootecus insularis. Both species are known from the entire
range, but their distribution pattern is somewhat patchy, and they are absent from many areas.
Coilostele paladilhiana is known from both coastal plains at the southern Red Sea. Its congeners
range in Spain (NW-Africa?) and India. Obviously, the ancient distribution area is restricted to a
few relic localities.
Boysia boysii is the only true representative of the east-Saharo-Sindian subgroup. Now it is
known from two places separated by a distance of more than 4000 km. This is similar to the
distribution pattern of Cerastus, which has two centres, one in India and one in the Arabian/Ethio-
pian area (MORDAN 1992).
Endemism
Endemism at the species level is quite common in the Arabian malacofauna, as 47 of the species
mentioned (50 %) are restricted to the peninsula. At the supraspecific level, however, endemism is
only represented in four (sub)genera which are definitely restricted to the area concerned. Within
those, Obeliscella has African affinities, while Polychordia is close to the Saharo-Sindian Cerastus.
Its status as a separate genus and even subgenus is questionable but is retained until investigation
of the genital organs can clarify its position.
Araboxychilus sabaeus and Oxychilus (Costoxychilus) profundus n. sp. are true endemic elements
in the strict sense. They originate from the Palaearctic stock of the Zonitidae, and thus a relation
to the speciose eastern Mediterranean and Caucasian fauna is likely. Unfortunately, no anatomical
data are available, thus preventing a more precise analysis, but the data on the radula of Araboxy-
chilus indicate a relationship to the Oxychilini (RIEDEL 1980: 117). In general, the Zonitidae are of
Laurasian origin and the two Arabian taxa are the only species of this family inhabiting Gondwana
(NORDSIECK 1986: 100).
Introduced and ubiquitous species
The Indian Indoplanorbis exustus as well as Thiara scabra were presumably introduced, as both are
known from Oman, the adjacent northern territories and Socotra (!) which traditionally were bases
of trade with the Indian subcontinent. Dispersal by drinkable or ballast water is often observed in
freshwater molluscs. The same is true for Melanoides tuberculata, a widespread ubiquitous taxon
which may originate from the Indopacific area. Physella acuta also is spread widely by human activity,
its origin is doubtful (northern America?). Polygyra cereolus and Allopeas gracilis both were introduced
from the New World, with the first from Florida (NEUBERT 1995), the latter from Middle America.
The freshwater clam Pisidium casertanum is found in nearly all suitable habitats around the globe.

444 E. NEUBERT
The remaining species Laevicaulis alte (tropical Africa?), Gulella bicolor (Indopacific area) and
Eobania vermiculata and Cantareus aspersus (both of circum-Mediterranean origin) are frequently
dispersed by soil adhering to the roots of plants. Introduction of allochthonous vegetation, in
particular in private gardens, recreation areas or simply in plantings alongside roads supports
successful settlement of alien faunal elements. The impact of these artificial habitats to the
autochthonous malacofauna has not been studied yet.
Iravadia quadrasi is a species living in brackish to completely marine habitats. It is included
here because it is a tradition to consider brackish water species together with terrestrial molluscs.
As with the freshwater-dwelling Assiminea nitida, it is known from the entire Indopacific region.
In Stenothyra, all habitats from marine to freshwater are colonised. The specific salt tolerance is
striking, although no known species lives in all types of habitats. This group is of Laurasian origin
and is also represented by fossil taxa in the Tertiary of Europe.
Due to the rigid arid environment, the interior parts of the peninsula as well as the coastal
plains of eastern Arabia may serve as a dwelling ground for a few autochthonous species only.
Oases, the only suitable habitats in these areas, have been subject to high densities of humans,
cattle and trade supporting anthropochorous dispersal of snails during historical times. Nowadays,
these environments have been substantially enlarged by huge artificially irrigated areas, producing
ecological niches usually not present since the last pluvial period. These structures are stepping
stones for allochthonous elements which often displace the autochthonous fauna. This does not
pose a severe threat in irrigated desert areas as there is no natural malacofauna existing, but as
shown by the list given above, several of the so-called “pest snails” have already reached the
peninsula. In particular, the sensitive mountain areas in the south-western part of Arabia, with
their unique fauna, should be protected from a snail such as Cantareus aspersus, which has
colonised all continents at the expense of the autochthonous malacofauna.
Species Saharo-Sindian Palaearctic Afrotropical Intro- AP en- Indo-
holo east west Lev. Mes. Ir.-Tu. west Et.H. trop. duced demic pacific
Hydrobia lactea ●
Hydrobia glaucovirens ●?
Corbicula purpurea ●
Xeropicta aff. mesopotamica ●
Mordania omanensis ●
Pseudonapaeus jousseaumi ●
Bithynia badiella ●
Sphincterochila prophetarum ●
Xerocrassa seetzeni ●
Xeropicta krynickii ●
Monacha obstructa ●
Paramastus sabaeanus ●
Paramastus hedjazicus ●
Levantina (Laevihelix) symensi ●
Levantina (Laevihelix) asagittata ●
Levantina (Laevihelix) asira ●
Levantina (Laevihelix) semitecta ●
Table 2. Zoogeographical groups within the Arabian malacofauna.
 = species also present outside the Arabian Peninsula;  = with generic affinities or present generically. Abbreviations:
Lev. = Levantinian, Mes. = Mesopotamian, Ir.-Tu. = Irano-Turanian, Et.H. = Ethiopian Highlands, trop. = tropical, AP en-
demic = species endemic for the Arabian Peninsula, Indopacific = Indopacific origin, n = note (see below table), g = global.

Terrestrial and freshwater molluscs of the Arabian Peninsula 445
Species Saharo-Sindian Palaearctic Afrotropical Intro- AP en- Indo-
holo east west Lev. Mes. Ir.-Tu. west Et.H. trop. duced demic pacific
Melanopsis praemorsa ●
Radix auricularia ●?
Galba truncatula ●?
Stagnicola palustris ●?
Planorbis planorbis ●?
Lauria cylindracea ●
Granopupa granum ●
Granaria arabica ●
Cecilioides aff. tumulorum ●
Cecilioides acicula ●?
Cecilioides isseli ●
Ancylus fluviatilis ●●
Arabivitrina arabica 
●
Arabivitrina jansseni 
●
Macroptychia (Sabaeola)
schweinfurthi

●
Macroptychia (Sabaeola) sumarana 
●
Lejeania leucosticta  ●
Lejeania aperta  ●
Eremina desertella 
Revoilia (Socotora) clausa ●
Revoilia (Socotora) dhofarense ●
Revoilia (Socotora) bentiana ●
Gulella protruda n1 ●
Gulella isseli  ●
Gulella schweinfurthi  ●
Gastrocopta antinorii ●
Gastrocopta klunzingeri ●
Streptostele (Raffraya) scotti ●
Homorus splendens ●
Homorus arabica ●
Euryptyxis candida ●
Euryptyxis fragosa ●
Euryptyxis labiosa ●
Euryptyxis latireflexa ●
Zebrinops albata ●
Biomphalaria arabica ●
Bulinus truncatus ●●?
Bulinus beccari ●
Bulinus wrighti ●
Radix natalensis ●●?
Quickia concisa ●
Gudeella rufocincta ●
Gudeella eremias ●
Gangetia miliacea ●
Gyraulus convexiusculus ●
Boysia boysii ●
Pupoides coenopictus ●●
Zootecus insularis ●●

Coilostele paladilhiana ●●
Cerastus schweinfurthi schweinfurthi 
●
Cerastus schweinfurthi apicostatus 
●
Cerastus schweinfurthi brunneus 
●

446 E. NEUBERT
Species Saharo-Sindian Palaearctic Afrotropical Intro- AP en- Indo-
holo east west Lev. Mes. Ir.-Tu. west Et.H. trop. duced demic pacific
Obeliscella lucidissima ●
Polychordia pulcherrima ●
Araboxychilus sabaeus ●
Oxychilus (Costoxychilus) profundus ●
Thiara scabra ●●
Melanoides tuberculata ●●
Melanoides sp. cf. plicaria ●●
Indoplanorbis exustus ●●
Laevicaulis alte ●●
Gulella bicolor ●●
Physella acuta ●
Eobania vermiculata ●●
Cantareus aspersus ●●
Allopeas gracilis ●
Polygyra cereolus ●
Iravadia quadrasi ●
Stenothyra arabica ●
Assiminea nitida nitida ●
Toltecia pusilla ● g?
Achatinelloides sebasmia ● n2
Achatinelloides jousseaumei ● n2
Lithidion lithidion ● n2
Pisidium casertanum ● g?
Notes:
n1: The generic placement of this species may be incorrect as there are also shell morphological
affinities to Gibbulinopsis Germain, 1919, of the Pupillidae. This would radically change the
relationships of this taxon to the Irano-Turanian fauna as is indicated by its distribution in the
Oman mountains. This problem can only be solved by investigation of the anatomy of the
genital organs.
n2: Achatinelloides and Lithidion are two endemic genera of Socotra Island. Probably, the so-called
species A. sebasmia and A. jousseaumei were introduced and can probably be relegated to the
synonymy of true Socotran Achatinelloides species. Lithidion lithidion appears to have been
introduced accidentally.
Synthesis
As shown in the previous paragraphs, the Arabian terrestrial and freshwater malacofauna is
composed of a mosaic of several faunas. The question of the origin of this “melting pot” at the
boundaries of three major zoogeographical units and two ancient continents arises. The time
schedule and pathways of colonisation, invasion and eradication of this fauna can be reconstructed
by using major geological events.
The most important area for such an analysis is the western and south-western area of the
peninsula, as it is deeply influenced by the geological developments during the Tertiary. The
following features are striking:

Terrestrial and freshwater molluscs of the Arabian Peninsula 447
– There is a fauna common to both Arabia and the Ethiopian Highlands. This fauna is already
differentiated on specific level, with only very few species known from both coasts of the
isthmus.
– Within the Palaearctic group, there are two genera with affinities to the western Saharo-
Sindian group (Arabivitrina, Macroptychia), another part consists of phylogenetically ancient
and widespread western Palaearctic taxa (Lauria, Granopupa, Granaria, Cecilioides, Ancylus).
– There is another group of Levantinian origin (Paramastus, Levantina), which is not represented
in Ethiopia.
– All Arabian taxa of African origin still have congeneric representatives on the main continent.
– There are a few taxa with the Saharo-Sindian type of distribution left, which have a relic
character in almost all cases.
– The malacofaunas of the Dhofar region and the Oman Mountains differ considerably in their
composition and origin.
The following hypothesis is developed to explain the several colonisation events to the Arabian
Peninsula. During the Palaeocene and until the early Eocene, the northern part of the area was
covered by a southwards pointing lagoon of the Tethys. This trough was narrowed and deepened
during the Eocene, leaving a land bridge in the area of what is now Yemen, Eritrea and Djibouti.
In parallel, the regression of the Tethys from the northern parts of Africa opened a connection
reaching from the Atlantic to the area of Ethiopia/Arabia. During the Oligocene, this land mass
was already inhabited by a stem fauna of the Palaearctic invading to all directions possible. As the
Arabian Shelf was covered by the Tethys until the Miocene, this first invasion from Africa
(Palaearctic and Afrotropical!) towards Arabia could only reach the area which was later uplifted to
form the Red Sea mountain belt. This explains the considerable differences in the faunal structure
between the Oman Mountains and the Red Sea coast. This first invasion was stopped by the first
opening of the Red Sea to the Indian Ocean in the early Miocene.
During the salinity crisis at the transition from Miocene to Pliocene, a second land bridge was
available. In the author’s opinion, another invasion from the African continent during that time is
unlikely as the huge evaporite series known make it evident that the “bridge” was a salt-pan
forming a barrier impossible to cross for land molluscs. Moreover, the mountain belts had been
formed and uplifted and most of the species were already adapted to humid mountainous climates
as they are still today. The only species known to dwell in both high altitudes in the mountain belt
and the lowland Tihama to Red Sea islands is Euryptyxis candida. It is evident that the highland
malacofauna is composed of two basic elements, the Oligocene Palaearctic and Afrotropical stock.
On the Arabian Peninsula, the northern border of this fauna reaches the Asir south of Taif, whereas
the Hijaz and Midian region seems to be completely influenced by Levantine faunal elements.
The second considerably younger invasion originated in the Levant and was directed south-
wards. Tethyian sediments are known from the northern part of Saudi Arabia, Jordan and Syria
dating from the middle Miocene approximately 20 million years b.p. Starting at this time, an
establishment of the Levantine fauna via Asia Minor took place. This process itself will have lasted
several million years, establishing for example the ancestral Helicidae. As a consequence, it can be
assumed that the second invasion reached southern Arabia during late the Miocene to early
Pliocene approximately 8-5 million years b.p. As outlined previously, there was no possibility of
crossing the sea, and consequently there are no Buliminidae, Zonitidae and Helicidae known from
the neighbouring Ethiopian Highland malacofauna.
The malacofauna of the eastern parts of the Arabian Peninsula is also characterised by two
groups. The relic Irano-Turanian species inhabit the Oman Mountains and Musandam Peninsula.
The time of invasion can not be traced exactly, it might have been before the middle Miocene

448 E. NEUBERT
transgression or after this event parallel to the orogenic activity of the area. In agreement with
MORDAN (1984: 131), the latter alternative is more probable as both species of the Buliminidae
are congeneric, with taxa already present in Iran. As displayed by Paramastus and Cyrenaeus, a
(sub)generic differentiation within this family can roughly be estimated to last approximately
5-8 million years.
The second eastern Arabian group is considered here to be represented by typical lowland
species which accompanied the two major freshwater tributaries of the area, Euphrates and Tigris.
This faunal element was nearly completely eradicated by the Pliocene transgressions and increasing
aridity. It might have had its climax during the more humid phase at the transition from late
Pliocene to early Pleistocene (3.5-1.2 million years b.p.) as characterised by Hötzl & Zötl (in AL-
SAYAD & ZÖTL 1978: 310). The freshwater species Hydrobia lactea and Corbicula purpurea are not
present south of the al-Hufuf Oasis system (Hydrobia glaucovirens (= H. lactea?) may have been
introduced to Dhofar by human activity). Although not typical Mesopotamian faunal elements,
Melanopsis praemorsa invaded the eastern Arabian freshwater systems from Syria and Asia Minor,
Gyraulus convexiusculus from the Oriental direction via the Shatt al-Arab.
The Afrotropical elements are restricted to western Arabia extending to the Dhofar region in
the south. There are only two species crossing the border: Bulinus wrighti and Radix natalensis. In
the author’s opinion, these two species should not be used for zoogeographical analysis as they are
insufficiently delimited from a systematic point of view. The specific status of B. wrighti has not
been reconfirmed, and the genus Radix is in a confused state in Europe and a hazardous one in
Africa. Introduction of both can not be excluded from our present state of knowledge. A (sub)fossil
record is known for four freshwater species from Wadi al-Luhy south-west of Riyadh (Hötzl et al.
in AL-SAYAD & ZÖTL 1978: 205). The age of this fauna was determined by carbon-14 measure-
ment as 8400 ± 140 b.p. Of Afrotropical origin, Biomphalaria pfeifferi rueppellii (here considered
to be B. arabica), Bulinus truncatus and R. natalensis have been found. Melanoides tuberculata is of
minor value for zoogeographical analysis, but its presence shows that the species had already been
introduced in historical times. Thus, the border of the Ethiopian and Palaearctic malacofauna is to
be found east of the Dhofar area (MORDAN 1980 a: 110).
In the Saharo-Sindian group, Pupoides coenopictus, Zootecus insularis and Coilostele paladilhiana
are here considered to represent part of the first invasive phase. The first two species are widespread
taxa, the genus Coilostele has three relic distribution centres in Spain, Arabia/Ethiopia and India.
Obviously, Pupoides is an ancient taxon, several congeneric species are known from America, Africa
(?) and Asia. Its outset from the hypothesised Oligocene Palaearctic malacofauna is supported by
this distributional pattern. In Z. insularis, the relationship to an Oligocene African fauna is very
likely. As a species, its distribution is similar to that of P. coenopictus, but the genus is restricted to
the Saharo-Sindian range. It is surprising in this context that only a few species of the first invasion
had been able to leave the Arabian Shield and spread over the Arabian Shelf.
The affinities of Cerastus with both the Afrotropical and Oriental fauna were considered by
MORDAN (1992: 3). The type locality of the type species C. distans (Kharg Island, Arabian Gulf) is
surprising as the species was never recorded again from this place. Moreover, there is no species of
this genus known from the area between Yemen and northern India, and a question about the
correctness of Pfeiffer’s type locality (coll. Cuming) arises. Assuming, that C. distans is conspecific
with an Indian species of the genus, only two spreading centres are left for Cerastus, an African and
an Indian one. At this point, ancient Gondwana distributional types may play a role. There are
only a few species and genera concerned, namely the Succineidae Quickia with its Indian subgenus
Burchella and Hyalimax H. & A. Adams, 1855. Ancestral forms of the Bulininae had been
separated by the splitting of Africa and India, leading to the evolutionary lineage of Indoplanorbis

Terrestrial and freshwater molluscs of the Arabian Peninsula 449
exustus in India and Bulinus in Africa. There are more examples for ancient Gondwana taxa, but
this exceeds the scope of a discussion of the Arabian malacofauna.
Conclusions
The invasion pathways as hypothesised here offer the opportunity for some speculation on
phylogenetic relationships within the western Palaearctic malacofauna.
Vitrinidae: The stem form of the Phenacolimax stock was already present in the Oligocene,
inhabiting both the northern and southern borders of the Tethys. The southern lineages developed
throughout the entire range and may be characterised by their exceptionally enlarged shells. In this
context, the almost completely unknown “Vitrina” libanica Germain, 1912 from a restricted area
in the Lebanese mountains should be mentioned (TOHMÉ & TOHMÉ 1988: 70). This species also
displays a huge shell, but differs from all species of Arabivitrina by having shallow interior radially
arranged lamellae, a unique character within the family (material examined: two specimens,
MNHNP ex Pallary, “Beyrouth?”). It is likely that a hitherto undescribed species from eastern
Anatolia is closely related. In the author’s opinion, a separate genus should be introduced for this
species, but it should be postponed until preserved specimens are available for anatomical research.
A relationship between “Vitrina” libanica and the Oligocene fauna is hypothetical, but there is no
argument with the idea that the Levantine pathway may also have been used by Palaearctic
elements extending northwards via Arabia.
Clausiliidae: As for the Phenacolimax stock, Boettgeria and Macroptychia may have a very
similar history. Compared to most of the recent Clausiliidae of Europe (excluding the Serrulinae),
these genera display a considerable number of plesiomorphic character states, but a few autapo-
morphies prove their monophyletic origin and are shared with the few relic taxa spreading to
South Africa. The remaining plesiomorphic “Mentissoid” genera found in the northern part of the
eastern Mediterranean should be confined to the European Oligocene Mentissoideinae stock
(Neubert, in prep.).
Hygromiidae: The presence of Lejeania on both highland areas bordering the southern Red
Sea supports the view of NORDSIECK (1987: 31) that the Euomphalia stock is a basic group within
the Hygromiidae.
Helicidae: In 1995, GLAUBRECHT published a paper dealing with zoogeographical ideas
concerning Levantina. Again, the author introduces the superspecies concept to malacological
science. Apart from the fact that he obviously overlooked Mordan’s record of Levantina from the
Asir, his conclusion is the separation of two generic complexes as it was already indicated by SUBAI
in 1994. Unfortunately, these results are not discussed in a framework of middle and lower Tertiary
geological processes. The isolation of populations in mountain chains and islands in the eastern
Mediterranean area is clear to all malacologists working in this region.
Levantina as a generic group, together with Assyriella, is at least of late Miocene origin.
Presumably, it evolved from an ancestral stock in the Syrian and/or south Anatolian area spreading
from there in several directions. The southern branch split up into two directions. The Libyan one
gave rise to Gyrostomella, the Arabian to Laevihelix. This is concordant to Paramastus, where the
Libyan branch is separated as subgenus Cyrenaeus Heller, 1971.
Deduced from major geological events which formed the Arabian Peninsula since the end of
the Mesozoic, a hypothesis concerning the invasion of this landmass by terrestrial and freshwater
molluscs is given (Fig. 212).
Summarising the data given by previous authors and the work presented, the following ideas
concerning the stepwise invasion of the Arabian Peninsula by molluscs can be hypothesised:

450 E. NEUBERT
Fig. 212: Pathways of colonisation of the Arabian Peninsula by terrestrial molluscs. 1 = the Oligocene introgression from Africa.
2 = the Miocene Levantine introgression. 3 = the Miocene Irano-Turanian introgression. 4 = the Pliocene/Pleistocene Euphrates
introgression
– The first phase started during the Oligocene, where a Palaearctic fauna in northern Africa and
an Afrotropical fauna from the south entered the Arabian plate via a land bridge in the area of
Eritrea/Djibouti and Yemen.
– The second phase started in the lower Miocene but was supported by the Levantine branch of
the western Palaearctic fauna.
– A third Miocene invasion of Irano-Turanian character took place in the eastern parts of the
peninsula via the Zagros, Musandam to the Oman Mountains.
– The remnants of the youngest Pliocene/Pleistocene fauna can be found in the realm of the
former Shatt al-Arab valley.
ACKNOWLEDGEMENTS
In the first place I would like to thank Pascale and Peter Symens, and Drs. Anne and Steven
Newton for their friendship and help during my stays in Saudi Arabia and for collecting molluscs

Terrestrial and freshwater molluscs of the Arabian Peninsula 451
used for this study. I am very much indebted to Prof. Dr. R. Kinzelbach (Rostock) for the
possibility to include his material in this study and Dr. F. Krupp (Frankfurt), whose interest and
help greatly supported this study. Help and discussion about Vitrinidae with Dr. Alonso and Prof.
Dr. Ibañez, Teneriffa, is greatly acknowledged. Many thanks go to Dipl.-Biol. E. Feltkamp for
valuable help in editing and preparing maps and for critical remarks on inconsistencies of the text.
I am very grateful to private collectors and all custodians of the museums for loaning type material
and additional specimens from the Arabian Peninsula, i.e. Dr. P. Bouchet (Paris), Prof. Dr. W.
Büttiker (Basel), H. Dekker (Winkel), Dipl.-Biol. G. Falkner (Wörth-Hörlkofen), Dr. J. van
Goethem (Bruxelles), Dipl.-Biol. K. Groh (Hackenheim), Dr. A. Hänggi (Basel), Prof. Dr. W.
Hohorst (Frankfurt), Prof. Dr. R. Kilias (Berlin), Mrs. A. Liebegott (Frankfurt), T. Meier (Zürich),
Dr. P. Mordan (London), Prof. Dr. I.A. Nader (Riyadh), S. Neubert (Lautertal), I. & H. Pauscher
(Griesheim), Prof. Dr. A. Riedel (Warzawa), Prof. Dr. M. Al-Safadi (Sana’a), Dr. M. Sartori
(Lausanne), Dr. W. Schneider (Darmstadt), Dr. M. Seddon (Cardiff) and Dr. M. Werner. For
many discussions and critical remarks on the manuscript I would like to express my thanks to
OStr. H. Nordsieck (Villingen-Schwenningen), Dr. R. Janssen (Frankfurt), and Dr. A. Zilch
(Wächtersbach).
This work was supported by the European Commission, Brussels, the NCWCD (National
Commission for Wildlife Conservation and Development), Riyadh, and the Senckenberg Museum,
Frankfurt.
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Manuscript accepted: 30 January 1998
Author’s address:
Dr. Eike Neubert, Forschungsinstitut Senckenberg, Sektion Malakologie, Senckenberganlage 25, D – 60325 Frankfurt a.M.,
Germany.

456 E. NEUBERT
APPENDIX 1
Systematic list of valid taxa of the Arabian Peninsula
Family Pomatiasidae
Revoilia (Socotora) clausa (Sowerby, 1847)
Revoilia (Socotora) dhofarense (Melvill & Ponsonby, 1896)
Revoilia (Socotora) bentiana (Melvill, 1895)
Lithidion lithidion (Sowerby, 1847)
Family Hydrobiidae
Hydrobia lactea (Küster, 1852)
Hydrobia glaucovirens (Melvill & Ponsonby, 1896)
Family Bithyniidae
Bithynia badiella (Küster, 1852)
Family Stenothyridae
Stenothyra arabica n. sp.
Gangetia miliacea (Nevill, 1880)
Family Iravadiidae
Iravadia quadrasi (O. Boettger, 1893)
Family Assimineidae
Assiminea nitida nitida (Pease, 1865)
Family Thiaridae
Thiara scabra (O.F. Müller, 1774)
Melanoides tuberculata (O.F. Müller, 1774)
Melanoides sp. cf. plicaria (Born, 1780)
Family Melanopsidae
Melanopsis praemorsa (Linnaeus, 1758)
Family Lymnaeidae
Radix auricularia (Linnaeus, 1758)
Radix natalensis (Krauss, 1848)
Galba truncatula (O.F. Müller, 1774)
Stagnicola palustris (O.F. Müller, 1774)
Family Planorbidae
Planorbis planorbis (Linnaeus, 1758)
Gyraulus piscinarum (Bourguignat, 1852)
Gyraulus convexiusculus (Hutton, 1849)
Biomphalaria arabica (Melvill & Ponsonby, 1896)
Bulinus truncatus (Audouin, 1827)
Bulinus beccari (Paladilhe, 1872)
Bulinus wrighti Mandahl-Barth, 1965
Indoplanorbis exustus (Deshayes, 1834)
Family Physidae
Physella acuta (Draparnaud, 1805)
Family Ancylidae
Ancylus fluviatilis O.F. Müller, 1774
Family Veronicellidae
Laevicaulis alte (Férussac, 1823)
Family Vertiginidae
Gastrocopta antinorii (Paladilhe, 1872)
Gastrocopta klunzingeri (Jickeli, 1873)
Boysia boysii (L. Pfeiffer, 1846)
Family Pupillidae
Pupoides coenopictus (Hutton, 1834)
Lauria cylindracea (Da Costa, 1778)
Family Chondrinidae
Granopupa granum (Draparnaud, 1801)
Granaria arabica (Dohrn, 1859)
Family Punctidae
Toltecia pusilla (Lowe, 1831)
Family Succineidae
Quickia concisa (Morelet, 1848)
Family Subulinidae
Zootecus insularis (Ehrenberg, 1831)
Allopeas gracilis (Hutton, 1834)
Obeliscella lucidissima (Paladilhe, 1872)
Homorus splendens (Thiele, 1910)
Homorus arabica (Connolly, 1941)
Family Streptaxidae
Streptostele (Raffraya) scotti Connolly, 1941
Gulella bicolor (Hutton, 1834)
Gulella isseli (Paladilhe, 1872)
Gulella schweinfurthi (Thiele, 1910)
Gulella protruda Neubert & Frank, 1996
Family Urocyclidae
Gudeella rufocincta Connolly, 1941
Gudeella eremias (Melvill & Ponsonby, 1896)
Family Vitrinidae
Arabivitrina arabica (Thiele, 1910)
Arabivitrina jansseni n. sp.
“Vitrina” gruneri L. Pfeiffer, 1846
Family Buliminidae
Mordania omanensis (E.A. Smith, 1894)
Pseudonapaeus jousseaumi (Smith, 1894)
Paramastus sabaeanus (Bourguignat, 1876)
Paramastus hedjazicus (Bourguignat, 1882)
Family Cerastidae
Cerastus schweinfurthi schweinfurthi (v. Martens, 1895)
Cerastus schweinfurthi brunneus n. ssp.
Cerastus schweinfurthi apicostatus n. ssp.
Cerastus scotti Connolly, 1941
Cerastus girwanensis Connolly, 1941
Cerastus albonotatus Verdcourt, 1974
Polychordia pulcherrima Connolly, 1941
Euryptyxis candida (Lamarck, 1822)
Euryptyxis fragosa (L. Pfeiffer, 1842)
Euryptyxis labiosa O.F. Müller, 1774)
Euryptyxis latireflexa (Reeve, 1849)
Achatinelloides sebasmia (Jousseaume, 1889)
Achatinelloides jousseaumei (Jousseaume, 1890)
Zebrinops albata (Férussac, 1827)
Family Zonitidae
Araboxychilus sabaeus (v. Martens, 1889)
Oxychilus (Costoxychilus) profundus n. sp.

Terrestrial and freshwater molluscs of the Arabian Peninsula 457
Family Ferussaciidae
Cecilioides isseli (Paladilhe, 1872)
Cecilioides aff. tumulorum Bourguignat, 1859
Cecilioides acicula (O.F. Müller, 1774)
Coilostele paladilhiana Nevill, 1880
Family Clausiliidae
Macroptychia (Sabaeola) schweinfurthi (v. Martens, 1889)
Macroptychia (Sabaeola) sumarana (Connolly, 1941)
Family Sphincterochilidae
Sphincterochila prophetarum (Bourguignat, 1852)
Family Polygyridae
Polygyra cereolus (Megerle von Mühlfeldt, 1816)
Family Hygromiidae
Xerocrassa seetzeni (L. Pfeiffer, 1847)
Xeropicta krynickii (Krynicki, 1833)
Xeropicta aff. mesopotamica (Mousson, 1874)
Monacha obstructa (L. Pfeiffer, 1842)
Lejeania leucosticta (v. Martens, 1889)
Lejeania aperta Connolly, 1941
Hygromiidae gen. sp. indet.
Family Helicidae
Eremina desertella (Jickeli, 1872)
Eobania vermiculata (O.F. Müller, 1774)
Levantina (Laevihelix) symensi n. sp.
Levantina (Laevihelix) asagittata n. sp.
Levantina (Laevihelix) asira n. sp.
Levantina (Laevihelix) semitecta n. sp.
Cantareus aspersus (O.F. Müller, 1774)
Family Corbiculidae
Corbicula purpurea Prime, 1863
Family Sphaeriidae
Pisidium casertanum (Poli, 1791)

458 E. NEUBERT
APPENDIX 2
Reference list of names used by various authors for terrestrial and freshwater molluscs
of the Arabian Peninsula
abyssinicus, Ancylus Jickeli, 1874
= Ancylus fluviatilis ...................................................... 360
Achatinelloides Nevill, 1878 ............................................... 404
acicula, Buccinum O.F. Müller, 1774
= Cecilioides acicula ..................................................... 409
acuta, Physa Draparnaud, 1805
= Physella acuta ........................................................... 359
adenensis, Pupa L. Pfeiffer, 1851
= Zootecus insularis ...................................................... 372
aegyptiaca, Coelestele Bourguignat, 1880
= Coilostele paladilhiana .............................................. 411
albata, Helix (Cochlogena) Férussac, 1827
= Zebrinops albata ....................................................... 405
albonotata, Cerastua Verdcourt, 1974
= Cerastus albonotatus ................................................. 401
alepina, Helix (Xerophila) mesopotamica var.
Westerlund, 1892 = Xeropicta mesopotamica............... 420
Allopeas H.B. Baker, 1935 ................................................. 373
alte, Vaginulus Férussac, 1823
= Laevicaulis alte ......................................................... 361
Ancylus O.F. Müller, 1774 ................................................. 360
antinorii, Pupa Paladilhe, 1872
= Gastrocopta antinorii ................................................ 362
aperta, Lejeania Connolly, 1941
= Lejeania aperta ......................................................... 427
apicostatus, Cerastus schweinfurthi ................................... 400
Arabia Pallary, 1925............................................................ 337
arabica, Coelestelle Bourguignat, 1880
= Coilostele paladilhiana .............................................. 411
arabica, Helix Forskål, 1775
= Euryptyxis candida .................................................... 402
arabica, Helix Forskål, 1775
= Euryptyxis labiosa ..................................................... 403
arabica, Leucochroa Pallary, 1901
= Sphincterochila prophetarum .................................... 416
arabica, Lymnaea Smith, 1894
= Radix natalensis ........................................................ 354
arabica, Pupa Dohrn, 1859
= Granaria arabica ...................................................... 368
arabica, Stenothyra ........................................................... 348
arabica, Subulina Connolly, 1941
= Homorus arabica ...................................................... 378
arabica, Vitrina Thiele, 1910
= Arabivitrina arabica ................................................. 384
arabicus, Planorbis Melvill & Ponsonby, 1896
= Biomphalaria arabica ............................................... 357
Arabivitrina Thiele, 1931 .................................................. 3 84
Araboxychilus Riedel, 1977 ................................................ 406
arata, Buliminus Recluz, 1843
= Euryptyxis candida .................................................... 402
argenteus, Pseudancylus Connolly, 1941
= Ancylus fluviatilis ...................................................... 360
asagittata, Levantina (Laevihelix) .................................... 433
asira, Levantina (Laevihelix) ............................................ 434
aspersa, Helix O.F. Müller, 1774
= Cantareus aspersus .................................................... 438
Assiminea Fleming, 1828 ................................................... 350
auricularia, Helix Linnaeus, 1758
= Radix auricularia ...................................................... 354
badiella, Paludina Küster, 1853
= Bithynia badiella ...................................................... 345
beccari, Physa Paladilhe, 1872
= Bulinus beccari .......................................................... 358
bentiae, Stenogyra Melvill & Ponsonby, 1896
= Obeliscella lucidissima .............................................. 375
bentianum, Otopoma Melvill, 1895
= Revoilia (Socotora) bentiana ..................................... 339
bicinctus, Bulimus Recluz, 1843
= Zebrinops albata ....................................................... 405
bicolor, Pupa Hutton, 1834
= Gulella bicolor ........................................................... 379
Biomphalaria Preston, 1910 .............................................. 357
Bithynia Leach, 1818 ......................................................... 345
bourguignati, Coelostele Jousseaume, 1890
= Coilostele paladilhiana .............................................. 411
bourguignati, Limicolaria Paladilhe, 1872
= Allopeas gracilis ......................................................... 374
Boysia L. Pfeiffer, 1849 ...................................................... 363
boysii, Tomogeres L. Pfeiffer, 1846
= Boysia boysii .............................................................. 363
bruguieri, Bulimus Bourguignat, 1882
= Euryptyxis labiosa ..................................................... 403
brunneus, Cerastus schweinfurthi ..................................... 399
Bulinus Müller, 1781 ......................................................... 358
candida, Pupa Lamarck, 1822
= Euryptyxis candida .................................................... 402
candidissimus, Bulimus L. Pfeiffer, 1858
= Zebrinops albata ....................................................... 405
Cantareus Risso, 1826........................................................ 437
casertanum, Cardium Poli, 1791
= Pisidium casertanum ................................................ 439
Cecilioides Férussac, 1814 .................................................. 40 8
Cerastus Albers, 1860 ......................................................... 397
cerealis, Bulimus Paladilhe, 1872
= Pupoides coenopictus ................................................. 364
cereolus, Helix Megerle von Mühlfeldt, 1816
= Polygyra cereolus ....................................................... 416
clausum, Cyclostoma Sowerby, 1847
= Revoilia (Socotora) clausa .........................................337

Terrestrial and freshwater molluscs of the Arabian Peninsula 459
coenopicta, Pupa Hutton, 1834
= Pupoides coenopictus ................................................. 364
Coilostele Benson, 1864 ..................................................... 41 1
çomaliana, Helix Bourguignat, 1882
= Eremina desertella ..................................................... 428
compressus, Ancylus Jickeli, 1874
= Ancylus fluviatilis ...................................................... 360
concisa, Succinea Morelet, 1848
= Quickia (Quickia) concisa ........................................ 370
consimile, Otopoma Melvill & Ponsonby, 1896
= Revoilia (Socotora) dhofarense .................................. 339
convexiusculus, Planorbis Hutton, 1849
= Gyraulus convexiusculus ............................................ 357
Corbicula Megerle von Mühlfeldt, 1811 ........................... 438
Costoxychilus ..................................................................... 408
cryophila, Helix (Patula) v. Martens, 1865
= Toltecia pusilla .......................................................... 368
cylindraceus, Turbo Da Costa, 1778
= Lauria cylindracea .................................................... 366
daliyana, Arabia bentianum var. Pallary, 1925
= Revoilia (Socotora) bentiana ..................................... 339
darnaudi, Helix L. Pfeiffer, 1854
= Lejeania darnaudi .................................................... 424
Dautzenbergi, Buliminus (Subzebrinus) Ancey, 1905
= Pseudonapaeus jousseaumi ........................................ 395
deflersi, Bulimus Jousseaume, 1894
= Euryptyxis latireflexa ................................................ 404
desertella, Helix Jickeli, 1872
= Eremina desertella ..................................................... 428
dhofarense, Otopoma Melvill & Ponsonby, 1896
= Revoilia (Socotora) dhofarense .................................. 339
Digoniaxis Jousseaume, 1889 ............................................. 410
dinshawi, Cerastus Sykes, 1903
= Euryptyxis latireflexa ................................................ 404
ducoureti, Buliminus Bourguignat, 1876
= Zootecus insularis ...................................................... 372
Eobania Hesse, 1913 .......................................................... 429
eremias, Hyalinia (Arnouldia) Melvill & Ponsonby, 1896
= Gudeella eremias ....................................................... 384
Eremina L. Pfeiffer, 1855 ................................................... 428
eryx, Buliminus Westerlund, 1887
= Euryptyxis candida .................................................... 402
euphraticus, Bulimus Bourguignat, 1876
= Pupoides coenopictus ................................................. 364
Euryptyxis Fischer, 1883 .................................................... 402
exustus, Planorbis Deshayes, 1834
= Indoplanorbis exustus ................................................ 359
fabianus, Buliminus Gredler, 1875
= Pupoides coenopictus ................................................. 364
fluviatilis, Ancylus O.F. Müller, 1774
= Ancylus fluviatilis ...................................................... 360
forskalii, Buliminus Beck, 1837
= Euryptyxis candida .................................................... 402
fragosa, Cochlogena Férussac, 1821
= Euryptyxis fragosa ..................................................... 402
fragosus, Bulimus L. Pfeiffer, 1842
= Euryptyxis fragosa ..................................................... 402
Francesia Paladilhe, 1872
= Coilostele ................................................................... 411
Galba Schrank, 1803.......................................................... 355
Gangetia Ancey, 1890 ........................................................ 34 9
Gastrocopta Wollaston, 1878 ............................................. 362
ghaesiana, Helix (Xerophila) mesopotamica var.
Mousson, 1874 = Xeropicta mesopotamica .................. 420
girwanensis, Cerastus Connolly, 1941
= Cerastus girwanensis ................................................. 400
glaucovirens, Paludestrina Melvill & Ponsonby, 1896
= Hydrobia glaucovirens ............................................... 345
gracilior, Buliminus schweinfurthi var. v. Martens, 1895
= Cerastus schweinfurthi schweinfurthi ........................ 398
gracilis, Albea prophetarum var. arabica sous-var.
Pallary, 1923 = Sphincterochila prophetarum .............. 416
gracilis, Bulimus Hutton, 1834
= Allopeas gracilis ......................................................... 374
Granaria Held, 1837 ......................................................... 368
Granopupa O. Boettger, 1889 ........................................... 3 67
granum, Pupa Draparnaud, 1801
= Granopupa granum .................................................. 368
gruneri, Vitrina L. Pfeiffer, 1846
= ”Vitrina” gruneri ...................................................... 389
Gudeella Preston, 1913 ...................................................... 383
Gulella L. Pfeiffer, 1856 ..................................................... 3 79
Gyraulus Charpentier, 1837 ............................................... 356
hadramauticum, Otopoma Melvill & Ponsonby, 1896
= Revoilia (Socotora) bentiana ..................................... 339
hedjazicus, Buliminus Bourguignat, 1882
= Paramastus hedjazicus .............................................. 397
Homorus Albers, 1850 ........................................................ 377
Hyalinia (Arnouldia) eremias Melvill & Ponsonby, 1896
= Gudeella eremias ....................................................... 384
Hydrobia Hartmann, 1821 ................................................ 344
hypodon, Bulimulus Pilsbry, 1897
= Euryptyxis labiosa ..................................................... 403
Indoplanorbis Annandale & Prashad, 1920 ....................... 35 9
insularis, Pupa Ehrenberg, 1831
= Zootecus insularis ...................................................... 372
Iravadia Blanford, 1867 ..................................................... 349
isseli, Caecilianella Paladilhe, 1872
= Cecilioides isseli ......................................................... 409
isseli, Coelostele Bourguignat, 1880
= Coilostele paladilhiana .............................................. 411
isseli, Ennea Paladilhe, 1872
= Gulella isseli .............................................................. 380
jansseni, Arabivitrina ........................................................ 387
jehennei, Pupa Recluz, 1843
= Euryptyxis labiosa ..................................................... 403
joppensis, Helix A. Schmidt, 1855
= Xeropicta krynickii .................................................... 420
jousseaumei, Ovella Jousseaume, 1890
= Achatinelloides jousseaumei ...................................... 405
jousseaumi, Bulimus E.A. Smith, 1894
= Pseudonapaeus jousseaumi ........................................ 395
klunzingeri, Pupa Jickeli, 1873
= Gastrocopta klunzingeri ............................................ 363

460 E. NEUBERT
krynickii, Helix Krynicki, 1833
= Xeropicta krynickii .................................................... 420
kursiensis, Bulimus Bourguignat, 1876
= Pupoides coenopictus ................................................. 364
labiosa, Helix O.F. Müller, 1774
= Euryptyxis labiosa ..................................................... 403
lactea, Paludina Küster, 1852
= Hydrobia lactea ......................................................... 344
Laevicaulis Simroth, 1913 ................................................. 36 1
Laevihelix .......................................................................... 430
latireflexus, Bulimus Reeve, 1849
= Euryptyxis latireflexa ................................................ 403
Lauria Gray in Turton, 1840 ............................................. 366
lederi, Patula O. Boettger, 1881
= Toltecia pusilla .......................................................... 368
leesi, Euryptyxis Pallary, 1928
= Euryptyxis latireflexa ................................................ 404
Lejeania Ancey, 1887 ......................................................... 424
leucosticta, Helix v. Martens, 1889
= Lejeania leucosticta ...................................................426
Levantina Kobelt, 1871 ..................................................... 430
Lithidion Gray in Baird, 1850 ........................................... 342
lithidion, Cyclostoma Sowerby, 1847
= Lithidion lithidion .................................................... 342
littlei, Arabia Pallary, 1925
= Revoilia (Socotora) bentiana ..................................... 339
littlei, Euryptyxis Pallary, 1925
= Euryptyxis latireflexa ................................................ 404
lucida, Arabia Pallary, 1925
= Revoilia (Socotora) bentiana ..................................... 339
lucidissima, Ennea? v. Martens, 1889
= Obeliscella lucidissima .............................................. 375
lucidissimus, Bulimus Paladilhe, 1872
= Obeliscella lucidissima .............................................. 375
lunti, Buliminus Melvill, 1895
= Euryptyxis latireflexa ................................................ 404
Macroptychia O. Boettger, 1877 ........................................ 412
mahariscus, Bulimus Bourguignat, 1876
= Pupoides coenopictus ................................................. 364
makallensis, Euryptyxis lunti var. Pallary, 1925
= Euryptyxis latireflexa ................................................ 404
marebiensis, Bulimus Bourguignat, 1876
= Pupoides coenopictus ................................................. 364
marmorosum, Lithidion Godwin-Austen, 1881
= Lithidion lithidion .................................................... 342
Martensi, Obeliscella? Jousseaume, 1890
= Obeliscella lucidissima .............................................. 375
maxima, Cerastus schweinfurthi var. Connolly, 1941
= Cerastus schweinfurthi schweinfurthi ........................ 398
Melanoides Olivier, 1804 ................................................... 3 51
Melanopsis Férussac, 1807 ................................................. 352
menahensis, Buliminus (Cerastus) schweinfurthi var.
Kobelt, 1902 = Cerastus schweinfurthi schweinfurthi .. 398
meridionalis, Punctum lederi var. O. Boettger, 1905
= Toltecia pusilla .......................................................... 368
mesopotamica, Helix (Xerophila) Mousson, 1874
= Xeropicta mesopotamica ............................................ 420
micraulax, Bulimus Bourguignat, 1882
= Euryptyxis latireflexa ................................................ 403
micraulaxus, Buliminus Bourguignat, 1882
= Euryptyxis candida .................................................... 402
miliacea, Hydrobia (Belgrandia) Nevill, 1880
= Gangetia miliacea ..................................................... 349
minor, Albea prophetarum var. arabica sous-var.
Pallary, 1923 = Sphincterochila prophetarum .............. 416
minor, Arabia hadramauticum var. 1925
= Revoilia (Socotora) bentiana ..................................... 339
minor, Arabia lucida var. Pallary, 1925
= Revoilia (Socotora) bentiana ..................................... 339
minor, Euryptyxis littlei var. Pallary, 1925
= Euryptyxis latireflexa ................................................ 404
Monacha Fitzinger, 1833.................................................... 422
Mordania Bank & Neubert, 1998 ..................................... 394
natalensis, Limnaeus Krauss, 1848
= Radix natalensis ........................................................ 354
nitida, Hydrocena Pease in Charpentier, 1865
= Assiminea nitida nitida ............................................. 350
niveum, Cyclostoma Petit de la Sausaye, 1850
= Lithidion lithidion .................................................... 342
Obeliscella Jousseaume, 1889............................................. 375
obstructa, Helix L. Pfeiffer, 1842
= Monacha obstructa .................................................... 422
omanensis, Bulimus E.A. Smith, 1894
= Mordania omanensis ................................................. 394
Oxychilus Fitzinger, 1833 ................................................... 407
paladilhiana, Coilostele Nevill, 1878
= Coilostele paladilhiana .............................................. 411
palustre, Buccinum O.F. Müller, 1774
= Stagnicola palustris ................................................... 355
Paramastus Hesse, 1933..................................................... 395
Physella Haldeman, 1843 ................................................... 359
Pilsbryanus, Buliminus (Petraeus) Ancey, 1900
= Euryptyxis labiosa ..................................................... 403
pisaniformis, Helix Bourguignat, 1881
= Eremina desertella ..................................................... 428
piscinarum, Planorbis Bourguignat, 1852
= Gyraulus piscinarum ................................................. 356
Pisidium C. Pfeiffer, 1821.................................................. 43 9
Planorbis O.F. Müller, 1774 ..............................................356
planorbis, Helix Linnaeus, 1758
= Planorbis planorbis ................................................... 356
plicaria, Helix Born, 1780
= Melanoides sp. cf. plicaria ......................................... 352
Polychordia Connolly, 1941 ............................................... 40 1
Polygyra Say, 1818.............................................................. 416
praemorsum, Buccinum Linnaeus, 1758
= Melanopsis praemorsa ............................................... 352
profundus, Oxychilus (Costoxychilus) ................................ 408
prophetarum, Helix Bourguignat, 1852
= Sphincterochila prophetarum .................................... 416
protruda, Gulella Neubert & Frank, 1996
= Gulella protruda ....................................................... 381
Pseudonapaeus Westerlund, 1887 ...................................... 394

Terrestrial and freshwater molluscs of the Arabian Peninsula 461
pulchella, Georgia Pallary, 1925
= Revoilia (Socotora) bentiana ..................................... 339
pulcherrima, Polychordia Connolly, 1941
= Polychordia pulcherrima ........................................... 401
Pupoides L. Pfeiffer, 1854 .................................................. 364
purpurea, Corbicula Prime, 1863
= Corbicula purpurea ................................................... 438
pusilla, Helix (Helicella) Lowe, 1831
= Toltecia pusilla .......................................................... 368
quadrasi, Alvania O. Boettger, 1893
= Iravadia quadrasi ..................................................... 349
Quickia Odhner, 1950 ....................................................... 370
Radix Montfort, 1810 ........................................................ 3 53
Raffraya Bourguignat, 1883 .............................................. 379
ragius, Bulimus Jousseaume, 1889
= Pupoides coenopictus ................................................. 364
Revoilia Bourguignat, 1881 ............................................... 337
rufocincta, Gudeëlla Connolly, 1841
= Gudeella rufocincta ................................................... 383
sabaea, Trochomorpha v. Martens, 1889
= Araboxychilus sabaeus ............................................... 407
sabaeanus, Buliminus Bourguignat, 1876
= Paramastus sabaeanus ............................................... 395
Sabaeola Lindholm, 1925 .................................................. 41 3
samavaensis, Bulimus Mousson, 1873
= Pupoides coenopictus ................................................. 364
scabrum, Buccinum O.F. Müller, 1774
= Thiara scabra ............................................................ 350
scalaris, Francesia Paladilhe, 1872
= Coilostele paladilhiana .............................................. 411
schoukraensis, Petraeus Jousseaume, 1899
= Euryptyxis latireflexa ................................................ 404
schweinfurthi, Buliminus v. Martens, 1895
= Cerastus schweinfurthi schweinfurthi ........................ 398
Schweinfurthi, Clausilia v. Martens, 1889
= Macroptychia (Sabaeola) schweinfurthi .................... 413
schweinfurthi, Ennea Thiele, 1910
= Gulella schweinfurthi ................................................ 381
scotti, Cerastus Connolly, 1941
= Cerastus scotti ........................................................... 400
scotti, Streptostele Connolly, 1941
= Streptostele (Raffraya) scotti ...................................... 379
sebasmia, Ovella Jousseaume, 1889
= Achatinelloides sebasmia ........................................... 404
seetzeni, Helix L. Pfeiffer, 1847
= Xerocrassa seetzeni .................................................... 418
semitecta, Levantina (Laevihelix) ...................................... 436
socialis, Petraeus Jousseaume, 1899
= Euryptyxis latireflexa ................................................ 404
Socotora Pallary, 1925 ........................................................ 337
Sphincterochila Ancey, 1887 .............................................. 415
splendens, Subulina Thiele, 1910
= Homorus splendens .................................................... 377
Stagnicola Leach, 1830 ...................................................... 355
stenostoma, Coelostele Jousseaume, 1890
= Coilostele paladilhiana .............................................. 411
Stenothyra Benson, 1854 ................................................... 348
Streptostele Dohrn, 1866 .................................................... 378
sumarana, Clausilia Connolly, 1941
= Macroptychia (Sabaeola) sumarana .......................... 414
symensi, Levantina (Laevihelix) ........................................ 431
Thiara Roeding, 1798 ........................................................ 350
tiani, Helix Bourguignat, 1882
= Eremina desertella ..................................................... 428
Tigridis, Cyrena (Corbicula) Mousson, 1874
= Corbicula purpurea ................................................... 438
tohenica, Helix Bourguignat, 1882
= Eremina desertella ..................................................... 428
Toltecia Pilsbry, 1926 ......................................................... 368
truncata, Physa Audouin, 1827
= Bulinus truncatus ...................................................... 358
truncatulum, Buccinum O.F. Müller, 1774
= Galba truncatula ...................................................... 355
tuberculata, Nerita O.F. Müller, 1774
= Melanoides tuberculata ............................................. 351
tumulorum, Caecilianella Bourguignat, 1856
= Cecilioides tumulorum .............................................. 409
ventricosa, Zebrinops Connolly, 1941
= Zebrinops albata .......................................................405
vermiculata, Helix O.F. Müller, 1774
= Eobania vermiculata ................................................. 430
vermiformis, Bulimus Paladilhe, 1872
= Pupoides coenopictus ................................................. 364
vestalis, Helix L. Pfeiffer, 1841
= Xeropicta krynickii .................................................... 420
wissmanni, Imparietula Wenz, 1943
= Zootecus insularis ...................................................... 372
wrighti, Bulinus reticulatus Mandahl-Barth, 1965
= Bulinus wrighti ......................................................... 358
Xerocrassa Monterosato, 1893 ........................................... 418
Xeropicta Monterosato, 1893 ............................................. 42 0
yemenica, Georgia Bourguignat, 1882
= Revoilia (Socotora) clausa .........................................337
Zebrinops Thiele, 1931 ...................................................... 405
Zootecus Westerlund, 1887 ................................................ 3 72