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ZooKeys 1010: 1-95 (2021)

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Numerous new records of tropical non-indigenous
species in the Eastern Mediterranean highlight
the challenges of their recognition and identification

Paolo G. Albano!”, Jan Steger'’, Piet A.J. Bakker”, Cesare Bogi?, Marija Bosnjak',
Tamar Guy-Haim°, Mehmet Fatih Huseyinoglu®,
Patrick I. LaFollette’, Hadas Lubinevsky°’, Martina Mulas>®,

Martina Stockinger', Michele Azzarone', Bruno Sabelli’

| Department of Palaeontology, University of Vienna, Althanstrasse 14, 1090, Vienna, Austria 2. Naturalis
Biodiversity Center, Darwinweg 2, 2333, CR Leiden, The Netherlands 3 Gruppo Matacologico Livornese, clo
Museo di Storia Naturale del Mediterraneo, via Roma 234, 57127, Livorno, Italy 4 Croatian Natural His-
tory Museum, Demetrova 1, Zagreb, Croatia § National Institute of Oceanography, Israel Oceanographic and
Limnological Research (IOLR), Haifa 3108001, Israel 6 Faculty of Maritime Studies, University of Kyrenia,
Karakum, Girne, Turkish Republic of Northern Cyprus 7 Malacology Section, Natural History Museum of Los
Angeles County, 900 Exposition Blvd., Los Angeles, CA 90007, USA 8 The Leon H. Charney School of Marine
Sciences, University of Haifa, 199 Aba Khoushy Ave., Mt. Carmel, Haifa 3498838, Israel 9 Museo di Zoologia
dell’ Universita di Bologna, via Selmi 3, 40126, Bologna, Italy

Corresponding author: Paolo G. Albano (pgalbano@gmail.com)

Academic editor: A.M. de Frias Martins | Received 17 September 2020 | Accepted 17 November 2020 | Published 13 January 2021
Attp://zoobank. org/45 DF30C9-AEB4-48AA-AC32-BBE77CB7191D

Citation: Albano PG, Steger J, Bakker PAJ, Bogi C, Bosnjak M, Guy-Haim T, Huseyinoglu ME, LaFollette PI,
Lubinevsky H, Mulas M, Stockinger M, Azzarone M, Sabelli B (2021) Numerous new records of tropical non-
indigenous species in the Eastern Mediterranean highlight the challenges of their recognition and identification.

ZooKeys 1010: 1-95. https://doi.org/10.3897/zookeys. 1010.58759

Abstract

New data on 52 non-indigenous mollusks in the Eastern Mediterranean Sea is reported. Fossarus sp.
(aff. aptus sensu Blatterer 2019), Coriophora lessepsiana Albano, Bakker & Sabelli, sp. nov., Cerithiopsis
sp. aff. pulvis, Joculator problematicus Albano & Steger, sp. nov., Cerithiopsis sp., Elachisina sp., Iravadia
aff. elongata, Vitrinella aff. Vitrinella sp. 1 (sensu Blatterer 2019), Melanella orientalis, Parvioris aft. di-
lecta, Odostomia cf. dalli, Oscilla virginiae, Parthenina cossmanni, Parthenina typica, Pyrgulina craticulata,
Turbonilla funiculata, Cylichna collyra, Musculus coenobitus, Musculus aft. viridulus, Chavania erythraea,

* These authors contributed equally to this work.

Copyright Paolo G. Albano et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

2 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Scintilla cf. violescens, lacra seychellarum and Corbula erythraeensis are new records for the Mediterranean.
An unidentified gastropod, Skeneidae indet., Triphora sp., Hypermastus sp., Sticteulima sp., Vitreolina cf.
philippi, Odostomia (s.1.) sp. 1, Henrya (?) sp., and Semelidae sp. are further potential new non-indigenous
species although their status should be confirmed upon final taxonomic assessment. Additionally, the sta-
tus of Dikoleps micalii, Hemiliostraca clandestina comb. nov. and H. athenamariae comb. nov. is changed
to non-indigenous, range extensions for nine species and the occurrence of living individuals for species
previously recorded from empty shells only are reported. Opimaphora blattereri Albano, Bakker & Sabelli,
sp. nov. is described from the Red Sea for comparison with the morphologically similar C. lessepsiana
Albano, Bakker & Sabelli, sp. nov. The taxonomic part is followed by a discussion on how intensive
fieldwork and cooperation among institutions and individuals enabled such a massive report, and how the
poor taxonomic knowledge of the Indo-Pacific fauna hampers non-indigenous species detection and iden-
tification. Finally, the hypothesis that the simultaneous analysis of quantitative benthic death assemblages

can support the assignment of non-indigenous status to taxonomically undetermined species is discussed.

Keywords

Cerithiopsidae, invasion biology, Lessepsian invasion, Mollusca, new species, Red Sea, taxonomy, Triphoridae

Introduction

The Eastern Mediterranean Sea is a hotspot of non-indigenous species introductions.
The opening of the Suez Canal in 1869 broke a long-standing biogeographic barrier
and enabled hundreds of Red Sea species to enter the basin and establish popula-
tions (Por 1978; Galil 2009; Zenetos et al. 2010, 2017; Zenetos and Galanidi 2020).
These so-called Lessepsian species are now recorded from all countries bordering
this basin west to Greece (Katsanevakis et al. 2009; Cinar et al. 2011; Ammar 2018;
Zenetos et al. 2018; Bariche and Fricke 2020; Crocetta et al. 2020) and some have
already reached the central Mediterranean, e.g., Tunisia (Ounifi-Ben Amor et al.
2015), Italy (Occhipinti-Ambrogi et al. 2011), and even France (Daniel et al. 2009;
Bodilis et al. 2011).

The introduction rate is an important metric to describe the invasion process.
Genuine variation in this rate can result from changes in vector efficacy, connectiv-
ity between the native and introduced range, and environmental conditions in the
recipient ecosystem. The introduction rate is often estimated from the discovery re-
cord (Solow and Costello 2004). However, even in well sampled and taxonomically
well-known groups like mollusks, multi-decadal time lags between introduction and
first detection have been quantified (Oliver 2015; Guy-Haim et al. 2017; Albano et
al. 2018), suggesting that the detection rate is a poor proxy of the introduction rate.
Indeed, although the discovery rate is increasing (Galil 2009; Raitsos et al. 2010),
estimates of the introduction rate corrected for temporal variation in sampling effort
for Lessepsian fishes showed that it was constant over ~ 1930-2010 (Belmaker et al.
2009). Still, the most recent enlargement of the Suez Canal has raised concerns that
the improved connectivity could increase the introduction rate of Lessepsian species

New non-indigenous species in the Mediterranean 3

(Galil et al. 2015). Additionally, rapid climate warming is particularly affecting the
Eastern Mediterranean (Ozer et al. 2017), causing, on the one hand, the decline of na-
tive species and, on the other hand, more favourable conditions for the establishment
of tropical species (Rilov 2016; Albano et al. 2021).

To monitor a dynamic process such as the Lessepsian invasion, intensive fieldwork
is mandatory. We indeed show that an intensive sampling effort coupled with identif-
cation at high taxonomic resolution and collaborative research among individuals and
institutions enabled the detection of 23 new Lessepsian mollusks, another nine species
which, upon further inspection, may prove to be new Lessepsian species, nine new re-
cords for Eastern Mediterranean countries, and new data for eleven already recognized
non-indigenous species. We here describe these new findings, providing detailed col-
lecting data, taxonomic comments, and comparisons with similar species.

Materials and methods

Origin of samples

The studied material comes from three main sources. First, sampling on the Israeli Med-
iterranean shelf performed in the context of the project “Historical ecology of Lessep-
sian migration” (HELM), in progress at the University of Vienna. Second, benthic as-
semblage monitoring by the Israel Oceanographic and Limnological Research (IOLR).
Third, smaller scale sampling by some of us, further detailed in the Results section.

Sampling in the framework of the HELM project was conducted on soft substrates
between 10 and 40 m depth with a van Veen grab, and on hard substrates between
5 and 30 m by diver-operated airlift suction sampling, using 0.5 mm mesh-size net
bags. Samples were sieved with a 0.5 mm mesh and the retained material fixed in 95%
ethanol. Both living individuals and empty shells were identified and counted.

IOLR conducts regular monitoring of Israeli soft bottom benthic assemblages in
the framework of the National Monitoring (NM) and focused sampling for environ-
mental assessment (APM DAN, Shafdan, Via Maris). The NM, APM DAN and Via
Maris projects sampled soft substrates with a 0.11 m* van Veen grab at depths between
6 and 12.5 m (NM), 22 and 26.5 m (APM DAN) and 18 and 26 m (Via Maris). Sam-
ples were sieved with a 250 um mesh. During the Shafdan project, three replicate sedi-
ment samples were taken at each station from a different 0.062 m? box-corer launch
(Ocean Instruments model 700 AL) twice a year in spring (May) and fall (October).
The samples were sieved on board with a 0.5 mm mesh. All samples were preserved
in 99% ethanol, stained with eosin solution (hence the pink hue that some specimens
bear) and picked for living individuals.

Finally, we included serendipitous findings by some of us or by colleagues within
our extended network, from multiple localities. For each species, we provide detailed
collecting data following the guidelines by Chester et al. (2019).

4 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Taxonomic assignment and non-indigenous status attribution

The depth of taxonomic assignment varies across taxa, mostly reflecting the available
knowledge on these groups in the Indo-Pacific province (the source pool of most non-
indigenous species in the Eastern Mediterranean). For families like the Triphoridae, some
of us (PGA, PAJB, and BS) have been conducting taxonomic research for a long time and
we have thus been able to describe new species as we have robust knowledge of inter- and
intraspecific variability and of type specimens (Albano et al. 2011, 2017, 2019; Albano
and Bakker 2016). For other families, like the Eulimidae, we focused our attention on
highlighting differences from native species and similarities with Indo-Pacific species,
because a more thorough coverage would have required revising the taxonomy of entire
Indo-Pacific species-groups, a task well beyond our objectives. In all cases, we strove to
provide detailed and high-quality images as a basis to foster further research and enable
the scientific community to refine our identifications. The use of qualifiers for species left
in open nomenclature follows the recommendations of Sigovini et al. (2016).

Acknowledging that an unsettled taxonomic status implies uncertainty in the as-
signment of non-indigenous status (Marchini et al. 2015), we here tagged as non-
indigenous only the species which: i) unequivocally belong to Indo-Pacific species;
ii) belong to clades (genera or families) that do not occur in the Mediterranean Sea,
even if left in open nomenclature; iii) belong to species whose diagnostic characters did
not enable a clear attribution to a non-Mediterranean clade but that were found alive
while not, or only very rarely, in the death assemblage (see Discussion). In contrast,
we tagged as “potential” non-indigenous species those whose morphological characters
did not allow for an unambiguous attribution to a non-Mediterranean clade and that
were found mostly, or exclusively, as empty shells.

Imaging and reporting

Small specimens were photographed with a Zeiss SteREO Discovery.V20 stereomicro-
scope, larger ones with a Nikon D7200 camera mounted on a stand, using a Nikon
Micro-Nikkor 60 mm lens. Photographs were stacked with Helicon Focus 6. Scanning
electron microscope (SEM) images were shot with a Fei Inspect S50 at low-vacuum
mode without coating. The internal shell morphology of Odostomia (s.1.) sp. 1, with a
particular focus on the intorted protoconch, was visualized using a Phoenix v|tome|x s
research edition computer tomographic (CT) scanner. The 3D-reconstruction and
virtual sections through the shell were produced with VGSTUDIO MAX 2.1 soft-
ware. Ihe X-ray image stack, mesh files, virtual sections, and a video showing the in-
terior of the shell are available from the Figshare repository (https://doi.org/10.6084/
m9.figshare.c.5215226). Plates were mounted with the image manipulation software
GIMP 2.

For each new non-indigenous species record, we report the size of at least one speci-
men (usually the one figured, unless otherwise stated). The systematic arrangement fol-
lows Bouchet et al. (2010, 2017). Table 1 summarizes the species treated in this work.

New non-indigenous species in the Mediterranean

Table |. List of the taxa treated in this paper, with indication of the novelty of the records.

Family | Taxon |—s—<‘itsSSSSCSsiCNvelty- — si | Taxon |—s—<‘itsSSSSCSsiCNvelty- — si Figure
a Uaieaia gags | __aaaeNS ore Malema 1-6 apie
Ceara — | _5 ceaadileadts «| —Tieor fling inane elieniaedsSe® (DTA IL =
Skeneidae Dikoleps micalii Declared NIS in the Mediterranean Sea. First record from Israel; Figure 2
first record of living individuals in the Mediterranean Sea
Figs
Planaxidae Fossarus sp. (aff. aptus sensu New NIS for the Mediterranean Sea Figure 4
Blatterer, 20 a
Naticidae Eunaticina papilla First record of living individuals in Israel Figure 5
Triphoridae Coriophora lessepsiana Albano, New NIS for the Mediterranean Sea Figure 6
Bakker & Sabelli, sp. nov.
Opimaphora blattereri Albano, | New species from the Red Sea, for comparison with the non- Figure 7
Bakker & Sabelli, sp. nov. indigenous Coriophora lessepsiana Albano, Bakker & Sabelli,
sp. nov.
Figure 8
Viriola cf. eles New reports of living individuals from Israel -
Cerithiopsidae Cerithiopsis sp. aff. pulvis New NIS for the Mediterranean Sea Figure 9
Cerithiopsis sp. New NIS for the Mediterranean Sea Figure 10
Joculator problematicus Albano New NIS for the Mediterranean Sea Figure 11
& Steger, sp. nov.
Elachisinidae Elachisina sp. New NIS for the Mediterranean Sea Figure 12
Iravadiidae Travadia aff. elongata New NIS for the Mediterranean Sea Figure 13
Vitrinellidae Vitrinella aff. Vitrinella sp. 1 New NIS for the Mediterranean Sea Figure 14
(sensu Blatterer 2019)
Fulinide Fie
Hemiliostraca clandestina Declared NIS in the Mediterranean Sea. First record from Israel; | 31 | Figure 16
comb. nov. first record of living individuals in the Mediterranean Sea
Figae
Sticteulima sp. Potential new NIS for the Mediterranean Sea 37 | Figure 19
Vitreolina cf. philippi Potential new NIS for the Mediterranean Sea 37 | Figure 16
Conidae Conus fumigatus First record from Israel 38 | Figure 20
Murchisonellidae Henrya (2) sp. Potental new NIS for the Mediterranean Sea 39 | Figure 21
Pyramidelide Figure
Odostomia (s.1.) sp. 1 Potential new NIS for the Mediterranean Sea Figure 23
Odostomia (s.l.) sp. 2 First record of a living individual in the Mediterranean Sea 44 | Figure 24
Oscilla virginiae New NIS for the Mediterranean Sea 45 | Figure 25
Parthenina cossmanni New NIS for the Mediterranean Sea 47 | Figure 26
Parthenina typica New NIS for the Mediterranean Sea 49 | Figure 27
:
Turbonilla funiculata New NIS for the Mediterranean Sea 53 | Figure 29
Cylichnidae Gjlichna collyra New NIS for the Mediterranean Sea 55 | Figure 30
Mnestiidae Mnestia girardi First record of living individuals in the Mediterranean Sea 57 -
Haminoeidae Atys angustatus First record from Greece 57 | Figure 31
Mild Figure
Figue3
Figure 35
Musculus | Musculus aff. viridulus | ——sNewNIS for the MediterraneanSea viridulus New NIS for the Mediterranean Sea Figure 36
Isognomonidae | Lsognomon ah a australica (sensu New record from Cyprus Figure 37
Angelidis and Polyzoulis 2018)
Lucinidae Pegophysema cf. philippiana First record of living individuals in the Mediterranean Sea Figure 38

Chavania erythraea

New NIS for the Mediterranean Sea

67 | Figure 39

Rugalucina angela

Additional records of living individuals from Israel

67

6 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)
Family Figure
Galeommatidae Nudiscintilla cf. glabra (sensu First record from Israel 68 | Figure 40
Mifsud and Ovalis 2012)

Galeommatidae Scintilla cf. violescens New NIS for the Mediterranean Sea 69 | Figure 41

Psammobiidae Gari pallida Additional records of living individuals from Israel 72 —
Semelidae Ervilia scaliola First record from Israel 72 | Figure 42
Lacra seychellarum New NIS for the Mediterranean Sea 72 | Figure 43
Figure

eed :
Cavbulida Figure

Abbreviations

BPBM

MZUB

Results

Bernice Pauahi Bishop Museum,

NHMW Natural

History Museum,

Landes-

Honolulu, Hawaii, USA; Vienna, Austria;

height; NM National Monitoring Israel;
“Historical ecology of Lessep-> OLML Oberésterreichisches

sian migration” project; museum Linz, Austria;

Israel Oceanographic and Lim- RMNH_ Rijksmuseum van Natuurlijke
nological Research; Historie (now Naturalis Bio-
length; diversity Center), Leiden, The
Natural History Museum of Los Netherlands;

Angeles County, Los Angeles, SMF Senckenberg Museum Frank-
California, United States; furt, Germany;

Museum national d'Histoire SMNH = Steinhardt Museum of Natural
naturelle, Paris, France; History, Tel Aviv, Israel;

Museo Civico di Storia Naturale sh/shs — empty shell/s;

di Genova “Giacomo Doria’,
Genova, Italy;

Museum of Zoology of the
University of Bologna, Italy;

Class Gastropoda Cuvier, 1795
Family unassigned (Caenogastropoda)

Unidentified gastropod

Figure 1

spcm/spcms live collected specimen/s;

vivv

W

valve/s;

width.

New records. IsRaEL * 1 spcm; Haifa Bay; 32.8211°N, 35.0196°E; depth 11 m; 2
Aug. 2015; soft substrate; grab; NM project (sample HM27(c)); size: H 2.5 mm, W

1.6 mm.

Remarks. We were not able to confidently assign this specimen to any family. The

general characters suggest that it is a caenogastropod. This specimen has apparently

New non-indigenous species in the Mediterranean 7

Figure |. Unidentified gastropod (Caenogastropoda), Haifa Bay, Israel (sample HM27(c)): front (A, B),
side (C) and back (D, E) views, protoconch (F, G). Scale bars: 1 mm (A=E); 0.3 mm (F-G).

traces of the animal inside and has thus been considered live collected. However, as it
was found in Haifa Bay, we cannot exclude that it comes from freshwater or transi-
tional ecosystems (the adjacent Kishon River and estuary) whose waters flow into the
bay. An anatomical study of the soft parts, should another living specimen become
available, will clarify the taxonomic placement of this intriguing species.

Family Conradiidae Golikov & Starobogatov, 1987

Conradia eutornisca (Melvill, 1918)

New records. IsraEL * 1 sh; Ashgelon; 31.7101°N, 34.5406°E; depth 31 m; 18
Sep. 2016; muddy-sand; grab; HELM project (sample SG30_5F) * 3 spcms;
Ashgelon; 31.6868°N, 34.5516°E; depth 11 m; 31 Oct. 2018; offshore rocky reef;
suction sampler; HELM project (samples $58_1M, S58_2F) * 3 spcms; Ashgelon;
31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef; suction

8 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

sampler; HELM project (sample S16_1M) © 1 spcm; same collecting data as for
preceding; depth 28 m; 31 Oct. 2018; HELM project (sample S59_3F) ¢ 1 spcm,
1 sh; west of Rosh HaNikra Islands; 33.0704°N, 35.0926°E; depth 12 m; 1 May
2018; rocky substrate; suction sampler; HELM project (samples $14_1F, $14_3F)
¢ 3 spcms; same collecting data as for preceding; 29 Oct. 2018; HELM project
(samples $52_1M, S52_3F, S52_3M) © 12 spcms, 4 shs; west of Rosh HaNikra
Islands; 33.0725°N, 35.0923°E; depth 20 m; 1 May 2018; rocky substrate; suction
sampler; HELM project (samples $13_1F, $13_1M, $13_2F, $13_3F, S13_3M) e
14 spcms; same collecting data as for preceding; depth 19 m; 29 Oct. 2018; HELM
projecte(samiplesS532 1h 'S5321M5:S5322F S53 2M, $5323FS53 3M); size:
2mm, W 1.7 mm.

Remarks. ‘The species has already been reported from Israel and Turkey (Bogi
and Galil 1999; Buzzurro and Cecalupo 2006), but only from empty shells. To the
best of our knowledge, this is the first record of living individuals from the Medi-
terranean Sea. Based on our observations, it occurs rather frequently on shallow
subtidal rocky substrates. The samples from Turkey led to the description of Par-
viturbo dibellai Buzzurro & Cecalupo, 2006, at that time supposed to be a native
species, but this name was later recognized to be a synonym of the Indo-Pacific Fos-
sarus eutorniscus (Rubio et al. 2015), attributed to Conradia by Janssen et al. (2011).
Janssen et al. (2011) highlighted, however, that the Red Sea specimens have seven
spiral cords instead of the five cited in the original description based on material
from Karachi (Pakistan). Specimens with five spiral cords occur also in the Persian
(Arabian) Gulf and rarely in the Red Sea (H. Dekker, pers. comm., November
2020). Further research is required to ascertain if these two morphologies belong to
two different taxa.

Family Skeneidae W. Clark, 1851

Dikoleps micalii Agamennone, Sbrana, Nardi, Siragusa & Germana, 2020
Figure 2

New records. IsraEL * 4 spcms; Haifa Bay; 32.8211°N, 35.0196°E; depth 11 m; 2
Aug. 2015; soft substrate; grab; NM project (samples HM27(a) and HM27(c)); size of
largest specimen: H 0.7 mm, W 0.7 mm.

Remarks. This species has been recently described from sediment collected in
2016 at 33-45 m depth at Karpathos and Samos islands in the eastern Aegean Sea
(Agamennone et al. 2020b). The authors discussed but declined the possibility that
this is a Lessepsian species, but one of us (BS) observed non-distinguishable specimens
from the Red Sea and we received reports of further indistinguishable specimens from
the Persian (Arabian) Gulf (H. Dekker, pers. comm., November 2020). This is the first

record for Israel; all specimens were live collected.

New non-indigenous species in the Mediterranean 9

Figure 2. Dikoleps micalii Agamennone, Sbrana, Nardi, Siragusa & Germana, 2020, Haifa Bay, Is-

rael: front (A) and apical (B) views, umbilicus (C), microsculpture of body whorl (D), apical view
of protoconch (E) and microsculpture of the base (F). Photograph courtesy A. Bonfitto. Scale bars:
0.2 mm (A=C); 0.05 mm (D-F).

Skeneidae indet.
Figure 3

New records. IsraEt * 1 sh; Akko; 32.92°N, 35.07°E; depth 4 m; 22 Oct. 1998; shell
grit sample; size: H 0.6 mm, W 1.0 mm.

Remarks. ‘This tiny gastropod (largest diameter 1 mm) is characterized by a small
but solid shell, ~ 0.75 whorls of protoconch with axial costae visible near the proto-
teleoconch transition (more costae closer to the nucleus may be abraded), and two tel-
eoconch whorls with numerous regular spiral cords. ‘The shoulder is slightly angulated
near the lip. Umbilicus open, large. Shell white, slightly translucent. No native Medi-
terranean species shares these features. Only Skenea catenoides (Monterosato, 1877) has
a similarly solid shell with numerous regular spiral cords, but it can be distinguished
easily by the three nodulose thicker spiral cords on the base and the lack of angulation
at the shoulder. Both Mediterranean (e.g., Circulus striatus (Philippi, 1836) and Red
Sea Circulus (e.g., C. novemcarinatus (Melvill, 1906a)) and C. octoliratus (Carpenter,
1856)) can be distinguished by the multispiral protoconch and the much more promi-
nent spiral cords. It is most likely a new, probably still unnamed, Indo-Pacific species
in the Mediterranean.

10 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 3. Skeneidae indet., Akko, Israel: apical (A, B), front (C, F), side (D, E), umbilical (G, H) and
back (I) views, protoconch (J). Scale bars: 0.5 mm (A=); 0.1 mm (J).

Family Planaxidae Gray, 1850

Fossarus sp. (aff. aptus sensu Blatterer 2019)
Figure 4

New records. Israzt * 1 sh; Ashqelon; 31.6868°N, 34.5516°E; depth 11 m; 31 Oct.
2018; offshore rocky reef; suction sampler; HELM project (sample $58_3M); size: H
3.3 mm, W 2.7 mm.

New non-indigenous species in the Mediterranean iat

Figure 4. Fossarus sp. (aff. aptus sensu Blatterer, 2019), Ashgqelon, Israel, HELM project (sample
S58_3M): front (A), side (B) and back (C) views. Scale bar: 1 mm.

Remarks. We found a single empty shell of this Fossarus that can be readily distin-
guished from the Mediterranean F ambiguus (Linnaeus, 1758), which bears prominent
spiral ridges and has a depressed spire. In contrast, our shell bears numerous spiral cords
and has a high spire. This shell is extremely similar to “Fossarus aff. aptus Melvill, 1912”
illustrated by Blatterer (2019: plate 87, fig. 7a-f). Especially the largest specimen (plate 87,
fig. 7a, b) bears a sculpture of similarly depressed and closely arranged spiral cords, has a
very similar profile and a large elongated umbilical area. Blatterer’s specimen shows, how-
ever, more regularly alternated thicker and finer cords, whereas in our specimen this feature
is not so evident. Our specimen is almost the double in size and rather worn, which may
explain the observed differences in sculpture. The extreme similarity with Blatterer’s Red
Sea specimens suggests that this is a new non-indigenous species in the Mediterranean Sea.

The name aptus is problematic. Originally introduced by Melvill (1912) for a
species from the Persian (Arabian) Gulf, it is currently considered a synonym of the
Atlanto-Mediterranean F ambiguus (MolluscaBase 2020), but no revision of this genus
is available. However, this name fits neither Blatterer’s specimens nor ours because
E aptus is characterized by five strong spiral keels (indeed similar to F ambiguus).

Family Epitoniidae Berry, 1910 (1812)

Cycloscala hyalina (G.B. Sowerby II, 1844)

New records. IsRAEL * 2 spcms; west of Rosh HaNikra Islands; 33.0704°N, 35.0926°E;
depth 12 m; 29 Oct. 2018; rocky substrate; suction sampler; HELM project (samples
S52_1M, S52_2F); size of largest specimen: H 3.7 mm, W 2.0 mm.

Remarks. The species has been recently recorded for the Mediterranean Israeli
coastline based on empty shells collected off the Soreg desalination plant, ~ 15 km south
of Tel Aviv (Scaperrotta et al. 2019). The species is also known from Cyprus (Cecalupo
and Quadri 1994), Turkey (Giunchi et al. 2001), and Greece (Scaperrotta et al. 2019).
To our knowledge, this is the first record of living individuals in the Mediterranean Sea.

12 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Family Naticidae Guilding, 1834

Eunaticina papilla (Gmelin, 1791)
Figure 5

New records. IsRAEL * 1 spcm; Ashdod; 31.8758°N, 34.6465°E; depth 27 m; 17 May
2017; soft substrate; grab; APM DAN project (sample 8C); size: H 1.1 mm, W 1.4 mm
(illustrated specimen) * 1 sh; rocky reef off Sdot Yam; 32.5111°N, 34.8702°E; depth
28 m; 1 Nov. 2018; hard substrate; suction sampler; HELM project (sample S60_2M).
Remarks. We here report the finding of a living individual of Eunaticina papilla
from the Israeli Mediterranean shelf. This juvenile specimen can be assigned to E. papilla
because of its overall shape, the sculpture of fine spiral cords, the large umbilicus and the
morphology of the thin corneus operculum (Figure 5D). The species has already been re-
ported in the Mediterranean Sea from Iskenderun in eastern Turkey with a living individ-
ual (Oztiirk and Bitlis Bakir 2013). An empty shell was collected near Shiqmona, Israel,
in November 2019 and reported as Eunaticina linneana (Récluz, 1843) (Schechter and
Mienis 2020), a name considered a junior synonym of £. papilla by Beu et al. (2004).

Family Triphoridae Gray, 1847

Coriophora lessepsiana Albano, Bakker & Sabelli, sp. nov.
http://zoobank.org/B69 1 1508-20E6-43B2-A01C-694571ADG60FE
Figure 6

Type material. Holotype. Ecyrt ¢ sh; Sinai (Red Sea), Dahab, dive site “Blue Hole”;
28.572°N, 34.538°E; depth 4 m; 2017; H. Blatterer leg; NHMW-MO-113282.

Paratypes. Ecyrt ¢ sh; Sinai (Red Sea), Dahab, dive site “Tigerhouse”; 28.567°N,
34.533°E; depth 7 m; 2015; H. Blatterer leg.; OLML LIEV 2019/70/1 (paratype 1)
sh; Sinai (Red Sea), Dahab, dive site “Caves”; 28.416°N, 34.456°E; depth 20 m; 2017;
H. Blatterer leg.; MNHN-IM-2014-7546 (paratype 2)

SUDAN °* sh; Arous, ca 30 km N of Port Sudan; 19.90°N, 37.23°E; depth 25-30 m;
2-8 Apr. 1975; G. Spada leg.; MZUB 60254 (paratype 3).

Additional material examined. Ecypt ¢ 1 sh; Sinai (Red Sea), Dahab, dive site
“Caves”; 28.416°N, 34.456°E; depth unspecified; 2012; H. Blatterer leg. * 1 sh; same col-
lecting data as for preceding; depth 15 m; 2015 © 1 sh; same collecting data as for preced-
ing; depth 14 m; 2017 ¢ 1 sh; same collecting data as for preceding; depth 18 m; 2017 ¢ 1
sh (juv.); Sinai (Red Sea), Dahab, dive site “Blue Hole”; floor of cave in cliff face; 28.57°N,
34.54°E; depth 25 m; Oct. 1994; D. Korkos leg.; H. Dekker coll. reg. no. 22017.

Records from the Mediterranean Sea. IsRAEL * 1 sh; west of Rosh HaNikra Is-
lands; 33.0704°N, 35.0926°E; depth 12 m; 29 Oct. 2018; HELM project (sample
S52_ 2M); NHMW-MO-112930/LM/0169; size: H 3.1 mm, W 1.2 mm (illustrated
shell, Figure ON, O).

New non-indigenous species in the Mediterranean 3

Figure 5. Eunaticina papilla (Gmelin, 1791), juvenile, Ashdod, Israel, APM DAN project (sample 8C):
front (A), side (B) and back (C) views, operculum (D), base (E) and apical view (F). The pink hue is due

to staining with eosin solution. Scale bar: 0.5 mm.

Diagnosis. Shell cyrtoconoid of ~ 3 mm with 11 whorls and multispiral proto-
conch. Nucleus with hemispherical granules. Sculpture of three spiral cords of which
two with elevated tubercles larger than their interspaces; second cord appearing later.
Peristome apparently without bifurcating spiral cords.

Description. Color: protoconch light brown; first teleoconch whorls whitish, with
the first spiral cord becoming brown after one to three whorls. The second spiral cord
acquires this brown color only on the dorsal part of the last whorl. The fourth cord,
visible only on the last whorl, is brown. ‘The base is light brown.

Dimensions: H 2.6 mm, W 1.0 mm (holotype); H 2.4 mm, W 0.9 mm (paratype
1); H 2.7 mm, W 1.1 mm (paratype 2, without apex); H 3.1 mm, W 1.2 mm (Medi-
terranean specimen, without apex).

Protoconch: multispiral with five whorls, H 530 wm (holotype), 553 um (paratype 1).

Protoconch I: 1.5 whorls with hemispherical granules, nucleus height of 114 um
(holotype), 104 um (paratype 1) and a maximum diameter of 154 um (holotype), 145
um (paratype 1).

Protoconch II: 3.5 monocarinated whorls with axial orthogonal riblets with a
maximum diameter of 305 um (holotype), 311 um (paratype 1), 323 pm (paratype 2),
268 um (Mediterranean specimen).

Teleoconch: 6 (holotype, paratype 1 and 2), 7.5 (Mediterranean specimen) whorls,
height: 2.04 mm (holotype), 1.83 mm (paratype 1), 2.43 mm (paratype 2) and 2.95

mm (Mediterranean specimen).

14 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

eT

.
2
:

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is

Figure 6. Coriophora lessepsiana Albano, Bakker & Sabelli, sp. nov. A-J holotype, NHMW-MO-113282,
dive site “Blue Hole”, Dahab, Sinai, Egypt: front (A, B), side (C, D) and back (E) views, nucleus (F),
protoconch (G, H) and peristome (I, J) K, M paratype 1, OLML LIEV 2019/70/1, dive site “Tigerhouse”,
Dahab, Sinai, Egypt: front (IK), side (L) and back (M) views. N, O NHMW-MO-112930/LM/0169,
west of Rosh HaNikra Islands, Israel (Mediterranean), HELM project (sample $52_2M): front (IN) and
side (O) views. Scale bars: 0.5 mm (A=E, K-O); 0.1 mm (F); 0.2 mm (G, H); 0.3 mm (I, J).

New non-indigenous species in the Mediterranean 15

The tuberculate first and third spiral cords start simultaneously after the proto-
conch with the same size, the third later becomes progressively larger and more acute.
The second spiral cord appears only on the last whorl and is smaller than the others in
front view, becoming of similar size to the first dorsally. In the second half of the last
whorl, a very thin smooth suprasutural cord is visible. The base shows a fourth rather
smooth cord of the same color as the first, followed by a fifth and sixth cord that are
smooth and very pale in color. Anterior siphonal canal short, tubular, and oblique;
posterior siphonal canal a simple notch. Peristome without microsculpture and appar-
ently without bifurcating spiral cords.

The Mediterranean specimen is larger, has three white whorls after the protoconch
and the second spiral cord appears on the seventh whorl, remaining still smaller than
the others.

Etymology. Named after the Lessepsian invasion (Por 1978), because we first
found this Red Sea species on the Mediterranean Israeli shelf. The species epithet is an
adjective in nominative singular feminine.

Remarks. The Mediterranean specimen is larger and broader than the Red Sea
ones. Triphorids do show a morphological dimorphism characterized by smaller and
larger morphs and we think that we captured this dimorphism in our samples. See
under Opimaphora blattereri Albano, Bakker & Sabelli, sp. nov. for a comparison with
similar species.

Opimaphora blattereri Albano, Bakker & Sabelli, sp. nov.
http://zoobank.org/9133ECDF-5F41-4E68-80C2-D20A74549047
Figure 7

Type material. Holotype. Ecyrt * sh; Sinai (Red Sea), Dahab, dive site “Islands”;
28.476°N, 34.513°E; depth 10 m; 2015; H. Blatterer leg.; NHMW-MO-113283.

Paratypes. Ecyrt ¢ sh; same collecting data as for holotype; OLML LIEV 2019/70/2
(paratype 1) ¢ sh; Sinai (Red Sea), Dahab, dive site “Labyrinth”; 28.478°N, 34.514°E;
depth unspecified; 2011; H. Blatterer leg; MNHN-IM-2014-7547 (paratype 2).

Additional material examined. Ecyrt ¢ 3 shs (juv. and fragments); same collect-
ing data as for holotype ¢ 9 shs (juv. and fragments); same collecting data as paratype 2
¢ 1 sh; Sinai (Red Sea), Dahab, dive site “Rick’s Reef”; 28.557°N, 34.524°E; 2012; H.
Blatterer leg. ¢ 1 sh; Sinai (Red Sea), Dahab, dive site “Caves”; 28.416°N, 34.456°E;
depth 45 m; 2012; H. Blatterer leg. * 3 shs (juv.); Sinai (Red Sea), Dahab, dive site
“Canyon”; 28.553°N, 34.522°E; depth 29 m; 2012; H. Blatterer leg. * 3 shs (juv.);
Sinai (Red Sea), Dahab, Masbay Bay; 28.497°N, 34.518°E; depth 5 m; 2015; H. Blat-
terer leg. ¢ 2 shs; Marsa Abu Makhadiq (Makadi Bay), SW side of bay, station 03;
26.9889°N, 33.9036°E; beached shell grit; 24 Sep. — 4 Oct. 1999; H. Dekker leg.; H.
Dekker coll. reg. nos. 37192 and 37201).

Diagnosis. Shell cyrtoconoid of less than 3 mm with 11 (holotype) or 12 (paratype
2) whorls and multispiral protoconch. Nucleus with hemispherical granules. Sculpture

16

Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 7. Opimaphora blattereri Albano, Bakker & Sabelli, sp. nov., dive site “Islands”, Dahab, Sinai, Egypt
A-J holotype, NHMW-MO-113283: front (A, B), side (C, D) and back (E) views, nucleus (F), proto-
conch (G, H) and peristome (I, J) K-M paratype 1, LIEV 2019/70/2: front (KK), side (L) and back (M)
views. Scale bars: 0.5 mm (A=E, K—M), 0.1 mm (F); 0.2 mm (G, H); 0.3 mm (I, J).

New non-indigenous species in the Mediterranean is

of three spiral cords with round tubercles larger than their interspaces; the second cord
appears only on the fourth whorl, initially as a thin smooth thread. Microsculpture
absent on the teleoconch whorls, present on the peristome, which bears bifurcating
spiral cords.

Description. Color: protoconch brown; whitish first teleoconch whorls with the
first spiral cord becoming brown after two whorls. Light brown irregular patches are
randomly distributed on the teleoconch, usually covering one or, more frequently,
two tubercles. ‘The base background is white, with the color patches of the last whorl
extending onto it. The tip of the anterior siphon is brown.

Dimensions: H 2.7 mm, W 0.9 mm (holotype); H 2.6 mm, W 1.0 mm (paratype
1, without apex); H 3.0 mm, W 1.0 (paratype 2).

Protoconch: multispiral with 5.5 whorls (holotype), 5 (paratype 2, but first whorl
worn); height: 566 um (holotype), 644 um (paratype 2).

Protoconch I: 1.5 whorls with hemispherical granules, nucleus with a height of
109 um (holotype), 122 um (paratype 2), and a maximum diameter of 371 um (holo-
type), 380 um (paratype 2).

Protoconch II: 3.5 whorls with axial orthogonal riblets with a maximum dia-
meter of 371 um (holotype), 380 um (paratype 2). First two whorls monocarinated,
then bicarinated.

Teleoconch: 6 (holotype), 7.5 (paratype 1), 7 (paratype 2) whorls, height: 2.24
mm (holotype), 2.43 mm (paratype 1), 2.50 mm (paratype 2).

The tuberculate first and third spiral cords start simultaneously after the proto-
conch with the same size, whereas the second cord appears from the fourth to the sev-
enth teleoconch whorl, depending on shell size. This cord is initially thin and closer to
the first one, it progressively increases its size until reaching that of the other two cords
on the last whorl. On the second half of the shell, a very thin smooth suprasutural
cord is visible. The second cord bifurcates on the peristome. The base shows a fourth
rather smooth cord, and a fifth and sixth smooth ones; these cords become towards the
peristome more granulated. On the peristome, below the third spiral thread, microscu-
Ipture is visible as fine spiral lines. Anterior siphonal canal tubular, short and oblique;
posterior siphonal canal a simple notch.

Etymology. This species is named after Hubert Blatterer, Austrian conchologist, in
recognition of his work on Red Sea mollusks. Moreover, he contributed to our work
on Lessepsian species by granting us access to the material he collected in the Red Sea
and by donating the type series of O. blattereri and Coriophora lessepsiana. The species
epithet is a noun in the genitive case.

Remarks. We describe O. blattereri as new because of the similar color pattern to
C. lessepsiana Albano, Bakker & Sabelli, sp. nov., even if it has not been reported from
the Mediterranean Sea. The two species can be easily distinguished because C. lessep-
siana has an monocarinated protoconch while O. blattereri has a bicarinated one; the
second spiral cord of O. blattereri never becomes brownish as in C. lessepsiana; O. blat-
tereri has a white background on the base and a distinct brown end of the anterior
siphonal canal, whereas C. lessepsiana has a light brown base and the anterior siphon

18 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

has not a colored end; the teleoconch of O. blattereri has irregular light brown patches,
particularly evident on fresh specimens; this feature is totally absent in C. lessepsiana.

We have seen specimens very similar to O. blattereri collected in Madagascar, New
Caledonia, and French Polynesia. A revision of the group in the Indo-Pacific province
is beyond the scope of this paper; however, this species likely has a broad distribution.

Opimaphora blattereri and C. lessepsiana share their color pattern of brown to orange
spiral cords on a white background with other Indo-Pacific species. Litharium bilinea-
tum (Kosuge, 1962) (holotype illustrated by Higo et al. (2001)), Costatophora iniqua
(Jousseaume 1898) (= Notosinister kawamurai Kosuge, 1962, type material illustrated
by Higo et al. (2001) and Albano et al. (2019)) and Aclophora albozonata Laseron,
1958 can be easily distinguished by having three fully developed spiral cords since the
early teleoconch. /niforis formosula (Hervier, 1898) and Mastonia peanites Jousseaume,
1898 (= Mastonia squamosa Kosuge, 1962, type material again illustrated by Higo et
al. (2001) and Albano et al. (2019)) have only two spiral cords, but the former has
three or four dark brown lines on the last whorl, whereas the latter has a dark brown
last whorl with lighter tubercles. Triphora fulvescens Hervier, 1898 also has a similar
color pattern, but the second spiral cord remains a very fine thread even on the last
whorl and the tubercles are whitish even on the first cord (on an orange background).
Some species show a delayed appearance of the second spiral cord: Nototriphora regina
(Hedley, 1903) has a brown tip of the anterior siphonal canal similarly to O. blattereri,
but lacks the patches on the whorls and has an orange line on the third spiral cord on
the last whorl; Coriophora tigris Laseron, 1958 has a paucispiral protoconch; Cautor
similis (Pease, 1871) has larger and more densely arranged tubercles, a brown fourth
spiral cord and white base. Last, a few species have a similar color pattern, but with
an inverted pattern: the first spiral cord is white and the third orange to brown, like
Mastonia cingulifera (Pease, 1861), which also has a dark yellow teleoconch, Mastonia
funebris Jousseaume, 1884 and Mastonia tulipa Jousseaume, 1898 with a brown and
white base, respectively.

Triphora sp.
Figure 8

New records. IsrazL * 1 sh; north of Atlit; 32.7433°N, 34.9067°E; depth 40 m;
20 Sep. 2016; coarse biogenic sediment in a pool among rocks covered by corallig-
enous formations; grab; HELM project (sample NG40_2M); NHMW-MO-112930/
LM/0170; size: H 4.5 mm, W 1.4 mm.

Remarks. We found a single, adult, empty shell. It likely possesses a large paucispi-
ral protoconch, but it is incomplete in our shell. The second spiral cord starts at mid-
shell height, the fourth and fifth spiral cords are smooth, and the posterior siphonal
canal is shallow. It is brown in color with darker spiral cords. We have not been able to
assign it to a species so far, but it is distinctly different from all known Mediterranean
species and most likely belongs to the Indo-Pacific fauna.

New non-indigenous species in the Mediterranean 19

Figure 8. Ziiphora sp., NHMW-MO-112930/LM/0170, north of Atlit, Israel, HELM project (sample
NG40_2M): front (A), side (B) and back (C) views, protoconch (D). Scale bars: 1 mm (A=C); 0.2 mm (D).

Viriola cf. bayani Jousseaume, 1884

New records. IsRAEL * 3 spcms; Ashqelon; 31.6868°N, 34.5516°E; depth 11 m; 31
Oct. 2018; offshore rocky reef; suction sampler; HELM project (samples S58_1M,
S58_2E S58_2M) ¢ 3 spcms; off Tel Aviv Marina; 32.0871°N, 34.7635°E; depth 7 m;
8 Nov. 2018; rocky reef, suction sampler; HELM project (sample S67_1M) * 7 spcms;
west of Rosh HaNikra Islands; 33.0704°N, 35.0926°E; depth 12 m; rocky substrate;
suction sampler; 29 Oct. 2018; HELM project (samples $52_1M, S52_3F) ¢ 1 spcm;
west of Rosh HaNikra Islands; 33.0725°N, 35.0923°E; depth 20 m; 1 May 2018;
rocky substrate; suction sampler; HELM project (sample $13_3M) * 9 spcms; same
collecting data as for preceding; depth 19 m; 29 Oct. 2018; HELM project (samples
S53_1M, S53_2F S53_2M, S53_3K, S53_3M); size of largest specimen: H 11.6 mm,
W 2.7 mm.

Remarks. ‘This species was first recorded from Israel by Steger et al. (2018) based on
three living individuals from Palmachim, southern Israel. We here report multiple living
individuals all along the Israeli coast, confirming its establishment. The species shows a
broad distribution in the Eastern Mediterranean ranging from Greece to Turkey and Cy-
prus (Micali et al. 2017; Stamouli et al. 2017; Angelidis and Polyzoulis 2018; Chartosia et

20 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

al. 2018). Its final taxonomic assignment requires the clarification of the relation between
several other Viriola such as V. corrugata (Hinds, 1843), V. senafirensis (Sturany, 1903),
and V. tricincta (Dunker, 1882) (Albano and Bakker 2016; Albano et al. 2017, 2019).

Family Cerithiopsidae H. Adams & A. Adams, 1853

Cerithiopsis sp. aff. pulvis (Issel, 1869)
Figure 9D—F

New records. Israet * 1 sh; Ashgelon; 31.6868°N, 34.5516°E; depth 11 m; 31 Oct.
2018; offshore rocky reef; suction sampler; HELM project (sample S58_1F) ¢ 1 spcm;
Ashgqelon; 31.6891°N, 34.5257°E; depth 28 m; 31 Oct. 2018; offshore rocky reef;
suction sampler; HELM project (sample S59_3F); NHMW-MO-112930/LM/0174;
size: H 1.8 mm, W 0.7 mm (illustrated specimen).

Additional material examined. Cerithiopsis pulvis (Issel, 1869): ISRAEL * 2 spcms,
1 sh; Ashgelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr. 2018; offshore rocky reef;
suction sampler; HELM project (sample $12_1F) * 5 spcms; same collecting data as
for preceding; depth 11 m; 31 Oct. 2018; HELM project (samples $58_1F S58_1M,
S58_2F) * 5 spcms, 2 shs; Ashgelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018;
offshore rocky reef; suction sampler; HELM project (samples S16_1F, S16_2E S16_2M)
¢ 19 spcms; same collecting data as for preceding; depth 28 m; 31 Oct. 2018; HELM
project (samples S59_1F S59_1M, S59_2F, S59_3F) © 8 spcms; west of Rosh HaNikra
Islands; 33.0704°N, 35.0926°E; depth 12 m; 29 Oct. 2018; rocky substrate; suction
sampler; HELM project (samples $52_2F, $52_2M, S52_3M) © 1 spcm; west of Rosh
HaNikra Islands; 33.0725°N, 35.0923°E; depth 20 m; 1 May 2018; rocky substrate; suc-
tion sampler; HELM project (sample S13_1M) * 6 spcms; same collecting data as for pre-
ceding; depth 19 m; 29 Oct. 2018; HELM project (samples $53_1F, $53_1M, S53_3F).

Remarks. ‘This species superficially resembles the Lessepsian Cerithiopsis pulvis but
has a more cyrtoconoid shape and a greater ratio between the height of the last whorl
and that of the shell. The base is not concave as in C. pulvis, bears a fourth spiral cord
which is more prominently tuberculate, and an additional fifth tuberculate cord that
is not present in typical C. pulvis. Additionally, the siphonal canal bears numerous fine
cords. The color pattern is similar to C. pulvis which has orange bands on white back-
ground; in contrast, in C. aff. pulvis these are brown and yellowish, respectively. It is
distinct from any native Mediterranean species and clearly belongs to an Indo-Pacific
clade. It is here considered a new non-indigenous species.

Cerithiopsis sp.
Figure 10

New records. Israet * 1 sh; Shiqmona Beach; 32.8259°N, 34.9555°E; beached; 4 Jan.
2008; size: H 3.5 mm, W 1.2 mm.

New non-indigenous species in the Mediterranean 21

Figure 9. Comparison between Cerithiopsis pulvis (Issel, 1869) and Cerithiopsis sp. aff. pulvis. A—C Ce-
rithiopsis pulvis, Ashqelon, Israel, HELM project (sample $16_1F): front (A), side (B) and back (C)
views. D=F Cerithiopsis sp. aff. pulvis, NHMW-MO-112930/LM/0174, Ashgelon, Israel, HELM project
(sample S59_3F): front (D), side (E) and back (F) views. Scale bars: 0.5 mm.

Remarks. This beautiful species has almost eight teleoconch whorls bearing two
strong spiral cords with oblong tubercles at the intersection with prosocline axial ribs.
Interspaces between spiral cords are approximately as large as the cords themselves, and
interspaces between the axial ribs are double the size of the ribs. A third smooth thick
cord delimits the rather flat base and is visible above the suture throughout most of the
teleoconch. The protoconch is smooth with very fine and extremely short axial riblets

22 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 10. Cerithiopsis sp., Shiqmona Beach, Israel: front (A, B), side (C) and back (D) views, pro-
toconch oriented to highlight the transition to the teleoconch (E, F), microsculpture (G). Scale bars:
1 mm (A=D); 0.2 mm (E, F); 0.4 mm (G).

just below the suture; it is multispiral but broken in our specimen in which only the
last two whorls are preserved. The slender shape, the two strong spiral cords and the
smooth flat base distinguish it at once from all native Mediterranean species suggesting
it is a new non-indigenous species in the basin.

Among Indo-Pacific cerithiopsids, Synthopsis lauta Cecalupo & Perugia, 2013, de-
scribed from Vanuatu, is among the few similar species we were able to trace. However,
the interspace between the spiral cords is broader, the tubercles on the first spiral cord
of the last whorl are larger than those on the second cord, and the teleoconch is shorter
with just six whorls. Additionally, the color pattern with white tubercles, yellowish in-
terspaces, deep brown suture and violet protoconch is strikingly different from the one
of our shell. We have some reservations that S. /auta, as well as our specimen, belong
to the genus Synthopsis Laseron, 1956 that was described as bearing three tuberculate
spiral cords on the whole teleoconch (Laseron 1956). Pending a molecular phylogeny
of the family, we consider this feature important at the genus level. ‘Therefore, we as-
sign our specimen to the nominotypical genus Cerithiopsis, in the wait of a better un-
derstanding of cerithiopsid systematics. The specimen identified as Horologica gregaria

New non-indigenous species in the Mediterranean 23

Cecalupo & Perugia, 2012 and illustrated in the recent revision of Cerithiopsidae from
South Madagascar (Cecalupo and Perugia 2014b: fig. 8G) is also similar to ours; that
specimen, however, has a distinct basal spiral cord which is absent in our specimen.
The latter character, the prominence of the tuberculate spiral cords and the evident but
rather flat third cord also raise some doubts that the specimen from South Madagascar
is conspecific with the H. gregaria originally described from the Central Philippines
(Cecalupo and Perugia 2011). Last, the Sudanese specimen of Horologica cf. taenia-
ta Cecalupo & Perugia, 2013 illustrated by Cecalupo and Perugia (2016: fig. 1P—S)
shares the general features of our shell but can be distinguished by the first spiral cord
that tends to split into two separate cords, and by the color pattern of white teleoconch
and orange base.

Joculator problematicus Albano & Steger, sp. nov.
http://zoobank.org/ 1 D9DDB2C-99D0-40A6-824A-93F4149B3550
Figure 11

Type material. Holotype. Isrart * spcm; Ashqelon; 31.6868°N, 34.5516°E; depth
11 m; 31 Oct. 2018; offshore rocky reef; suction sampler; HELM project (sample
S58_3F); NHMW-MO-113580.

Paratypes. \sRAEL * spcm; west of Rosh HaNikra Islands; 33.0704°N, 35.0926°E;
depth 12 m; 1 May 2018; rocky substrate; suction sampler; HELM project (sample
S14_2F); MNHN-IM-2012-25505 (paratype 1) * spcm; same collecting data as for
paratype 1; HELM project (sample $14_4F); MZUB 60400 (paratype 2) * spcm; same
collecting data as for paratype 1; 29 Oct. 2018; HELM project (sample $52_3F); SMF
360591 (paratype 3) * spcm; same collecting data as for paratype 1; HELM project
(sample $14_2F); SMNH MO 99705 (paratype 4).

Additional material examined. IsrazEL * 5 spcms; Ashqelon; 31.6868°N,
34.5516°E; depth 12 m; 30 Apr. 2018; offshore rocky reef; suction sampler; HELM
project (samples $12_1E $12_2E S12_3F) ¢ 6 spcms; same collecting data as for pre-
ceding; depth 11 m; 31 Oct. 2018; HELM project (samples $58_1F, $58_2E, S58_3F)
¢ 1 spcm; Ashgelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky
reef; suction sampler; HELM project (sample S16_2F) * 2 spcms; same collecting
data as for preceding; depth 28 m; 31 Oct. 2018; HELM project (samples S59_3F
S59_3M) © 1 sh; Sdot Yam; 32.5299°N, 34.8599°E; depth 24 m; 3 May 2018; rocky
substrate; suction sampler; HELM project (sample S17_1F) * 3 spcms; west of Rosh
HaNikra Islands; 33.0704°N, 35.0926°E; depth 12 m; 1 May 2018; rocky substrate;
suction sampler; HELM project (samples $14_2F, S14 _4F) * 16 spcms; same collect-
ing data as for preceding; 29 Oct. 2018; HELM project (samples $52_1F S$52_1M,
S52_2EF, S52_3F) © 2 spcms; west of Rosh HaNikra Islands; 33.0725°N, 35.0923°E;
depth 20 m; 1 May 2018; rocky substrate; suction sampler; HELM project (sample
S13_1F) * 9 spcms; same collecting data as for preceding; depth 19 m; 29 Oct. 2018;
HELM project (samples $53_1F, S53_2F S53_3E S53_3M).

24 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure | |. Joculator problematicus Albano & Steger, sp. nov., holotype, NHMW-MO-113580, Ashqelon,
Israel, HELM project (sample $58_3F): front (A, B), left side (C), right side (D, E) and back (F) views,
protoconch (G, H), microsculpture (I) and base and siphonal canal (J). Scale bars: 0.5 mm (A-+F);
0.2 mm (G, H); 0.3 mm (I); 0.4 mm (J).

Diagnosis. Very small bulbous brown shell, of - 1.5 mm in height and < 1 mm in
width, with a relatively short, almost smooth protoconch.

Description. Color: Protoconch white, teleoconch brown with white outer lip margin.

Dimensions: H 1.6 mm, W 0.7 mm (holotype), H 1.4 mm, W 0.7 mm (paratype
1), H 1.5 mm, W 0.7 mm (paratype 2), H 1.5 mm, W 0.7 mm (paratype 3), H 1.6
mm, W 0.8 mm (paratype 4).

Protoconch: composed of 3.5 whorls with no clear demarcation between proto-
conchs | and I], height: ~ 300 um, width ~ 200 um (holotype), but accurate measure-
ment hampered by the last protoconch whorl being covered by the first teleoconch
whorl. It appears smooth except for growth lines and fine pustules covering the lower
half of the first whorl and sparsely present apically and abapically on the following
whorls (only visible with scanning electron microscopy at high magnification).

New non-indigenous species in the Mediterranean 25

Teleoconch: 4 whorls (holotype), height: 1.4 mm (holotype). It bears three spiral
cords of equal size, with tubercles at the intersection of orthocline axial ribs. The base
is contracted and has two additional tuberculate spiral cords. Tubercles become oblong
near the lip. Anterior siphonal canal short, reverted upwards, formed by a prong-like
protrusion of the anterior outer lip (Figure 11A); posterior siphonal canal notch-like.

Etymology. The name problematicus refers to the difficult task of recognizing and
identifying non-indigenous species belonging to groups whose taxonomy in the tropi-
cal seas is poorly known (see Discussion). The species epithet is an adjective in nomina-
tive singular masculine.

Remarks. This species is characterized by its bulbous contour and constricted last
whorl which justify its inclusion in the genus /Joculator Hedley, 1909 (Hedley 1909;
Marshall 1978).

The Cerithiopsidae of the Indo-Pacific have been subject to numerous in-depth
studies (Cecalupo and Perugia 2011, 2013, 2014a, b, 2016, 2017a, b, 2018, 2019a,
b, c). Still, this species does not fit any of the known species. Among the most similar
species in terms of shell shape and ornamentation, /oculator itiensis Cecalupo & Peru-
gia, 2014 has one teleoconch whorl more and a different color pattern characterized
by light brown first whorl and base, /. olivoideus Cecalupo & Perugia, 2018 can be
distinguished by its clearly prosocline axial ribs and greyish tubercles, and /. sekensis
Cecalupo & Perugia, 2018 has only two spiral cords and blunter axial ribs on the first
teleoconch whorl, in addition to a blunter siphonal canal.

There are several more species of small brown bulbous /ocu/ator often distinguish-
able only by subtle character differences. Joculator priorai Cecalupo & Perugia, 2012 is
corneous in color and has a pointed protoconch with one additional whorl; moreover,
in our specimens the interspaces between the spiral cords are smaller. Joculator pupiformis
Cecalupo & Perugia, 2012 has one protoconch and one teleoconch whorl more, the
tubercles are oblong, and the base lacks a clearly visible fifth tuberculate spiral cord.
Joculator fuscus Cecalupo & Perugia, 2012 has much broader interspaces between cords
and a wide subquadrangular aperture which is, in contrast, quite small in our speci-
mens. Joculator furvus Cecalupo & Perugia, 2012 has a neat abapical smooth cord on
the protoconch, one teleoconch whorl less and a broader aperture. Joculator carpatinus
Cecalupo & Perugia, 2012 has one protoconch whorl more, one teleoconch whorl less,
a broader aperture and a fine abapical thread on the protoconch. Joculator caliginosus Ce-
calupo & Perugia, 2012 has one protoconch whorl more and one teleoconch whorl less,
the basal fourth and fifth cords are only weakly tuberculate whereas they are neatly tu-
berculate in our specimens. Joculator coffeus and_J. subglobosus, both Cecalupo & Perugia,
2013, have one clear abapical thread on the protoconch, one teleoconch whorl less, the
shell has a more roundish shape and the lip does not reach anteriorly the siphonal canal,
almost covering it, like in our specimens. The other representatives of Joculator include
also other more elongated species that can be easily distinguished from our specimens.

This species is superficially similar to the native Mediterranean Cerithiopsis ladae Prki¢
& Buzzurro, 2007, which, however, can be distinguished at once for not having the last
protoconch whorl partially covered by the first teleoconch whorl and lacking the prong-

26 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

like process of the anterior outer lip. Additionally, tubercles in C. dadae on the last whorl
are more elongated, subrectangular, and the shell profile is less bulbous. Cerithiopsis grep-
pii Buzzurro and Cecalupo, 2005, described from Turkey, has a rather oval profile, but
not as bulbous as in our species; additionally, it has a paucispiral protoconch. Cerithiopsis
micalii (Cecalupo and Villari, 1997), which also has a somewhat oval shell profile, can
be quickly distinguished by its protoconch whose last two whorls bear strong axial ribs.

Unfortunately, a revision of Red Sea Cerithiopsidae is lacking, but given that Jocu-
lator is a broadly distributed genus in the Indo-Pacific province, we consider J. prob-
lematicus another previously undescribed Indo-Pacific species recently introduced to the
Mediterranean Sea.

Family Elachisinidae Ponder, 1985

Elachisina sp.
Figure 12A—-E

New records. IsraEL * 1 spcm; north of Atlit; 32.7417°N, 34.9177°E; depth 31 m; 25
Apr. 2017; sand; grab; HELM project (sample NG30_8M) * 14 spcms, 1 sh; west of
Rosh HaNikra Islands; 33.0725°N, 35.0923°E; depth 20 m; 1 May 2018; rocky sub-
strate; suction sampler; HELM project (samples $13_1E $13_1M, S13_2E S13_3L);
size of largest specimen: H 1.6 mm, W 1.3 mm ® 9 spcms; same collecting data as for pre-
ceding; depth 19 m; 29 Oct. 2018; HELM project (samples $53_1F, $53_2E S53_3F).

Additional material examined. Elachisina robertsoni Kay, 1979: UNITED STATEs ®
1 sh; Hawaii, Oahu, Maunalua Bay; BPBM 9754 (holotype).

Remarks. ‘The morphology of this species is unique among the native mollusks of
the Mediterranean, which does not host any shallow water Elachisinidae. Therefore, we
consider it a new non-indigenous species in the basin.

The only Indo-Pacific Elachisina we are aware of is E. robertsoni Kay, 1979, which
indeed shares the general characters of our species. However, it can be readily distin-
guished by the thicker and fewer spiral cords, less rounded whorls and sigmoid, rather
than strongly prosocline, aperture profile. Elachisina sp. is more similar to the West-
African E. tenuisculpta (Rolan and Gofas 2003), but the Israeli shells have more rounded
whorls, a greater height/width ratio and smaller ratio between aperture and shell height.

Family Iravadiidae Thiele, 1928

Travadia aff. elongata (Hornung & Mermod, 1928)
Figure 13

New records. Israzt * 1 spcm; Ashqelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr.
2018; offshore rocky reef; suction sampler; HELM project (sample $12_1F) ¢ 5 shs;
same collecting data as for preceding; depth 11 m; 31 Oct 2018; HELM project (sam-
ple S58_2F); size: H 2.8 mm, W 1.1 mm (largest (illustrated) shell).

New non-indigenous species in the Mediterranean 27

Figure 12. Comparison between Elachisina sp. and Elachisina robertsoni Kay, 1979 A=-E Elachisina
sp., west of Rosh HaNikra Islands, Israel, HELM project (sample $13_3L): front (A, B), side (C) and
back (D, E) views FH Elachisina robertsoni, BPBM 9754 (holotype), Maunalua Bay, Oahu, Hawaii:
front (F), side (G) and back (H) views (photograph courtesy N. Young). Scale bars: 0.5 mm.

Additional material examined. /ravadia aff. elongata: SUDAN * 2 shs; Arusa (near
Port Sudan); 19.90°N, 37.23°E; shallow water; 1975; shell-grit; G. Spada leg.; MZUB.

Travadia elongata (Hornung & Mermod, 1928): Ertrrea * 1 sh; Massawa; depth
30 m; 1870; A. Issel leg.; syntype in MSNG; size: H 3.9 mm, W 1.4 mm.

28 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 13. lravadia aff. elongata (Hornung & Mermod, 1928) A=-F Ashqelon, Israel, HELM project
(sample $58_2F): front (A, B) and back (C, D) views, protoconch (E) and detail of sculpture on the body
whorl (F) G, H Arusa (near Port Sudan), Sudan: front view of an adult (G) and juvenile (H) shell. Scale
bars: 1 mm (A=D, G, H); 0.2 mm (E); 0.3 mm (F).

Remarks. This species is characterized by a turriform shell with up to five convex
whorls, separated by a marked suture and a blunt, flat, and smooth protoconch. The
sculpture consists of flat spiral ridges (12-14 on the penultimate whorl) that become
more raised at both the adapical and abapical parts of the whorls, and which overlie
numerous axial lines (Figure 13F), resulting in a reticulate surface.

New non-indigenous species in the Mediterranean 29

The closest match to our specimens is [ravadia elongata (Hornung & Mermod,
1928) which was described from material collected by Arturo Issel in the Red Sea off
Massawa, Eritrea, at 30 m depth (Hornung and Mermod 1928). Compared to our
material, however, the syntype of /. elongata is larger (height 3.9 mm vs. 2.8 mm in
our largest shell) and has seven less convex whorls. Further, the apical part of its spire
has a slightly concave profile and thus appears more tapered. According to Issel’s de-
scription, the sculpture of 1. elongata consists of spiral ridges (12 on the penultimate
and 22 on the last whorl) as well as growth lines, although the latter are not indicated
in the accompanying line drawing. This suggests that the axial component might be
less evident in /. elongata than in our specimens, however, the poor preservation of the
shell surface of the syntype of J. elongata did not allow a reliable comparison with our
material. Slightly eroded shells very similar to our specimens have been collected from
the Sudanese Red Sea (Figure 13G, H), confirming that the material from Israel indeed
represents an Indo-Pacific species rather than an undescribed Mediterranean taxon.

Among Mediterranean iravadiids, our specimens superficially resemble only Cera-
tia proxima (Forbes and Hanley, 1850). This species, however, lacks axial sculpture.
Interestingly, Hornung and Mermod (1928) also mention the presence of this latter
species at Assab (Eritrea) and “tle Saldadin” (Zeila, northern Somalia). While obviously
based on a misidentification — C. proxima has an Eastern Atlantic-Mediterranean distri-
bution (Bouchet and Warén 1993; Hoiseter 2009) — one might speculate that this re-
cord could be the result of a confusion of C. proxima with the [ravadia presented here.

Family Vitrinellidae Bush, 1897

Vitrinella aff. Vitrinella sp. 1 (sensu Blatterer 2019)
Figure 14

New records. Isrart * 1 sh; Ashdod; 31.8697°N, 34.6473°E; depth 24 m; Sep. 2019;
soft substrate; grab; APM DAN project (sample 10B); size: H 0.4 mm, W 0.7 mm.
Remarks. This tiny gastropod defeated all our attempts to identify it. It consists
of a protoconch and a teleoconch of ~ 1.5 whorls each. Sculpture is absent, except
for two spiral ridges that run on the shoulder and on the base. A third ridge runs
periumbilically (Figure 14E). Broad umbilicus, roundish aperture. Our shell closely
resembles the Vitrinella sp. 1 illustrated by Blatterer (2019: plate 127, fig. 12a—j) from
the Dahab region in the northern Red Sea, which, however, apparently bears fine spi-
ral threads in the umbilicus (fig. 12e, and unpublished figures). SEM images of our
shell show that its surface is taphonomically altered; additionally, Blatterer’s specimens
look slightly more mature, reaching 2 teleoconch whorls. The significance of these
features should be re-assessed upon a satisfying revision of these tiny gastropods from
the Indo-Pacific province. Another similar shell is illustrated by Janssen et al. (2011,
plate 19, figs. 3a—b), which apparently has less conspicuous or absent spiral ridges as
long as can be judged from the optical illustrations provided. It is worth mentioning

that gastropods belonging to the family Clenchiellidae D.W. Taylor, 1966 share the

30 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 14. Vitrinella aff. Vitrinella sp. 1 (sensu Blatterer 2019), Ashdod, Israel, APM DAN project (sam-
ple 10B): apical (A, B), umbilical (C, D), front (E) and side (F) views, protoconch (G) and back view
(H). Scale bars: 0.3 mm (A=F, H); 0.1 mm (G).

small size, low spire, wide umbilicus and presence of strong spiral keels we observed in
our specimen (Ponder et al. 2014); the latter, however, lacks the numerous finer spiral
cords that characterize clenchiellids. Additionally, these gastropods occur in mangrove

New non-indigenous species in the Mediterranean 31

swamps or adjacent habitats in tropical estuaries, a kind of habitat that does not occur
in Israel. The shell shape and sculpture (in particular the strong spiral keels) distinguish
it at once from native Mediterranean species. The extreme similarity with the shell il-
lustrated in Blatterer’s book suggests that the species belongs to a Red Sea clade and is
here considered a new non-indigenous species in the Mediterranean Sea.

Family Eulimidae Philippi, 1853

Hypermastus sp.
Figure 15

New records. IsraxEt * 4 shs; off Tel Aviv Marina; 32.0871°N, 34.7635°E; depth 7 m;
8 Nov. 2018; rocky reef; suction sampler; HELM project (sample S67_3F); size: H 1.9
mm, L 0.5 mm (illustrated shell).

Remarks. This slender eulimid is characterized by a constriction at the transition
between the protoconch and the teleoconch (Figure 15A, B, F). This feature distin-
guishes it at once from any native Mediterranean species. The protoconch is ~ 2.5
whorls, apparently without any ornamentation. The teleoconch is very slender, made
of 6 translucent-white whorls, with flat sides, inconspicuous suture; the lip profile is
arched. These characters fit the genus Hypermastus Pilsbry, 1899 to which we tentative-
ly assign the species (Warén 1991b). We were not able to assign it to any Indo-Pacific
species, but the family is among the most diverse and least known in that province
(Bouchet et al. 2002), thus it could be another still undescribed species recently intro-
duced into the Mediterranean Sea.

Hemiliostraca clandestina (Mifsud & Ovalis, 2019), comb. nov.
Figure 16A—C

New records. IsraEL * 1 spcm; Ashgelon; 31.6868°N, 34.5516°E; depth 12 m; 30
Apr. 2018; offshore rocky reef; suction sampler; HELM project (sample $12_1F) ¢ 38
spcms; Ashgelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky
reef; suction sampler; HELM project (samples S16_1K S16_2F S16_2M); size: H
2.7 mm, L 0.9 mm (illustrated shell) * 16 spcms; same collecting data as for preced-
ing; depth 28 m; 31 Oct. 2018; HELM project (samples $59_1F, $59_2F, S59_3F) e
1 spcm; west of Rosh HaNikra Islands; 33.0725°N, 35.0923°E; depth 20 m; 1 May
2018; rocky substrate; suction sampler; HELM project (sample S13_3F).

Remarks. Sticteulima clandestina and S. athenamariae, both Mifsud & Ovalis, 2019,
were described on specimens collected in Turkey (Mifsud and Ovalis 2019). However,
both belong to species present in the Red Sea and were illustrated by Blatterer (2019)
for the Gulf of Aqaba on plate 131, fig. 8a—d and plate 131, fig. 9a—h, respectively.
Sticteulima clandestina appears rather variable but our specimens clearly match Mifsud
and Ovalis (2019: fig. 1B). Both S. clandestina and S. athenamariae look closely related
to Hemiliostraca and thus we propose the new combinations Hemiliostraca clandestina

32 Paolo G. Albano etal. / ZooKeys 1010: 1-95 (2021)

Figure 15. Aypermastus sp., Tel Aviv, Israel, HELM project (sample S67_3F): front (A, B), side (C, D) and
back (E) views, apex with optical microscope (F) and SEM (G) showing the protoconch-teleoconch transition

(the shell had a different orientation in images F and G). The arrow marks the constriction between proto-

conch and teleoconch that is a diagnostic character for this species. Scale bars: 0.5 mm (A=E); 0.2 mm (F,G).

and Hemiliostraca athenamariae. This is the first record of H. clandestina in Israel, but
the species has been recorded for Lebanon based on empty shells collected in 1999
(Crocetta et al. 2020). Consequently, it is likely present here since at least 1999, with a
~ 20 year time-lag in first detection as quantified also for other non-indigenous species

New non-indigenous species in the Mediterranean 33

Figure 16. Hemiliostraca clandestina (Mifsud & Ovalis, 2019), and comparison between Vitreolina

philippi (de Rayneval & Ponzi, 1854) and Vitreolina sp. A-C Hemiliostraca clandestina, Ashqelon, Israel,
HELM project (sample S16_1F): front (A), side (B) and back (C) views. D Vitreolina philippi (de Ray-
neval & Ponzi, 1854), Plakias, Crete, Greece (sample Rh.05_5M): front view. E, F Vitreolina cf. philippi,
Ashgelon, Israel, HELM project (S16_2F): front (E) and side (F) views. Scale bars: 0.5 mm.

in the Mediterranean Sea (Crooks 2005; Albano et al. 2018). This is also the first record
of living individuals from the Mediterranean Sea. Despite the relatively large number of
living individuals, we did not find any attached to an echinoderm host; this is consist-
ent with the fact that some eulimids actively leave the host if disturbed (Warén 1984).

Melanella orientalis Agamennone, Micali & Siragusa, 2020
Figure 17

New records. IsRAEL * 5 spcms; Ashgelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr.
2018; offshore rocky reef; suction sampler; HELM project (samples $12_1F $12_1M,
S12_3F); size: H 2.7 mm, W 1.0 (illustrated specimen) * 1 spcm; same collecting data
as for preceding; depth 11 m; 31 Oct. 2018; HELM project (sample S$58_2F) ¢ 3 spc-
ms; Ashgelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef;
suction sampler; HELM project (samples $16_1F S16_2F) * 1 spcm; same collecting
data as for preceding; depth 28 m; 31 Oct. 2018; HELM project (sample $59_1F).
Remarks. ‘This species can be distinguished from Mediterranean Melanella by its
gently curved whorls, straight spire with fewer whorls and thinner shell than most
species. It superficially resembles the Red Sea “Eulima” orthophyes Sturany, 1903 (type
illustrated by Albano et al. (2017)), which can be distinguished because of its slightly
bent apical whorls and the unusual pustulous sculpture of the protoconch. ‘The species
presented here is apparently already widespread in the Eastern Mediterranean (Aga-
mennone et al. 2020a). We found only living individuals and no empty shells. Because
of this, and the low likelihood that a so widespread species in shallow depths in the

34 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure | 7. Melanella orientalis Agamennone, Micali & Siragusa, 2020, Ashqelon, Israel, HELM project (sam-
ple S12_1M): front (A, B), side (C, D) and back (E) views, apex (F). Scale bars: 1 mm (A=E); 0.2 mm (F).

Eastern Mediterranean would have escaped detection for long, we consider it a new
non-indigenous species in the basin. The specimens reported as Melanella sp. by Al-
bano et al. (2020) from mesophotic reefs off northern Israel belong to this species.

New non-indigenous species in the Mediterranean EB)

Parvioris aff. dilecta (E.A. Smith, 1899)
Figure 18E—M

New records. IsrAEL * 1 sh; north of Atlit; 32.7433°N, 34.9067°E; depth 40 m; 20
Sep. 2016; coarse biogenic sediment in a pool among rocks covered by coralligenous
formations; grab; HELM project (sample NG40_2M); size: H 4.9 mm, W 2.2 mm (il-
lustrated shell, Figure 18E—H) ¢ 3 spcms; west of Rosh HaNikra Islands; 33.0704°N,
35.0926°E; depth 12 m; 29 Oct. 2018; rocky substrate; suction sampler; HELM pro-
ject (samples $52_1K $52_1M, S52_2F).

Additional material examined. Parvioris ibizenca (FE Nordsieck, 1968): IsRAEL ® 1
sh; west of Rosh HaNikra Islands; 33.0704°N, 35.0926°E; depth 12 m; 29 Oct. 2018;
rocky substrate; suction sampler; HELM project (sample $52_2F).

Remarks. The genus Parvioris Warén, 1981 was erected for a group of numer-
ous conchologically very similar species of which many are still undescribed (Warén
1981). The species here reported is very similar in general shape and size to P dilecta
(Warén 1981: 146), especially the morphs illustrated here in Figure 18I-M. However,
it has a multispiral protoconch of ~ 4.5 whorls (Figure 18H), whereas P dilecta has a
paucispiral protoconch of ~ 1.5 whorls. The type of protoconch is considered to be
related to the developmental mode, which was regarded a diagnostic character at the
species level for most molluscan lineages (Hoagland and Robertson 1988; Bouchet
1989). Because Warén (1984) suggested that the number of protoconch whorls is
rather constant within species in Eulimidae, we currently do not consider our ma-
terial conspecific with P dilecta, but only closely related (thus the “aff.” notation).
However, there is increasing evidence that poecilogony, the intraspecific variation in
developmental mode, occurs in Caenogastropoda (McDonald et al. 2014), Neogas-
tropoda (Russini et al. 2020), and Sacoglossa (Krug 1998; Ellingson and Krug 2006;
Vendetti et al. 2012).

Parvioris aff. dilecta can be easily distinguished from the native P ibizenca because
of a more arched apical part and because of the protoconch morphology: both have
multispiral protoconchs, but P ibizenca has shorter whorls and a distinct profile which
inflates at the third whorl, in contrast with the more slender and regular profile of P
aff. dilecta. Our specimens are likely conspecific with those identified as Melanella
sp. 1 by Blatterer (2019) from Dahab, Red Sea, suggesting that it is indeed a new
Lessepsian species. An additional issue is whether the animal color is diagnostic at
the species level like in other groups whose shells offer few diagnostic morphological
characters, e.g., Mediterranean Granulina (Neogastropoda: Granulinidae) and Gib-
berula (Neogastropoda: Cystiscidae) (Gofas 1990, 1992). Some of our live collected
specimens show a light yellow-white color (e.g., Figure 18I-K) whereas others have a
brownish animal (e.g., Figure 18K—M). The final attribution of our findings to a spe-
cies requires a thorough revision of Parvioris, which is beyond the scope of this paper.
The specimens reported as Parvioris sp. by Albano et al. (2020) from mesophotic reefs
off northern Israel belong to this species.

36 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 18. Comparison between Parvioris aff. dilecta (E.A. Smith, 1899) and Parvioris ibizenca (F. Nor-
dsieck, 1968) A=-D Parvioris ibizenca, west of Rosh HaNikra Islands, Israel, HELM project (sample
S52_2F): front (A), side (B) and back (C) views, protoconch (D) E=H Parvioris aff. dilecta, north of
Atlit, Israel, HELM project (sample NG40_2M): front (E), side (F) and back (G) views, protoconch (H)
I Parvioris aff. dilecta, west of Rosh HaNikra Islands, Israel, HELM project (sample $52_1F): front view
J-K Parvioris aff. dilecta, west of Rosh HaNikra Islands, Israel, HELM project (sample $52_1M): front (J)
and side (KK) views L, M Parvioris aff. dilecta, same collecting data as for preceding, HELM project (sample
S52_2F): front (L) and side (M) views. Scale bars: 0.5 mm (A=C, I=-M); 0.2 mm (D, H); 1 mm (E=G).

New non-indigenous species in the Mediterranean 37

Sticteulima sp.
Figure 19

New records. IsrazL * 1 sh; north of Atlit; 32.7820° N, 34.9466° E; depth 10 m;
21 Sep. 2016; sand; grab; HELM project (sample NG10_1F); size: H 1.4 mm, W
0.6 mm (illustrated shell, Figure 19A—F) * 1 spcm; Ashgelon; 31.6891°N, 34.5257°E;
depth 28 m; 31 Oct. 2018; offshore rocky reef; suction sampler; HELM project (sam-
ple S59_1F).

Remarks. We place this species in Sticteulima due to its small size, slender profile
with high and rather flat whorls (Warén 1984). In contrast to the native S. jeffreysiana
(Brusina, 1869) and the Lessepsian S. /entiginosa (A. Adams, 1861), it is colorless, also
in live-collected specimens, and stouter. Further, this species does not match any of
the known small-sized Mediterranean eulimids. It can be readily distinguished from
Vitreolina curva (Monterosato, 1874) and Melanella levantina (Oliverio, Buzzurro &
Villa, 1994) by the lack of the strongly arched apical whorls. This feature differentiates
it at once also form other Red Sea small-sized eulimids (Blatterer 2019). Melanella
petitiana (Brusina, 1869) is larger, has more numerous whorls (our Sticteulima has a
fully thickened lip suggesting that it is an adult) and has a less prominent lip profile.
Nanobalcis nana (Monterosato, 1878) (type illustrated by Appolloni et al. (2018)) has
shorter whorls, especially the last one, which is also much broader than in this species.
It can also be easily distinguished from Hemiliostraca athenamariae (Mifsud & Ovalis,
2019) by the lack of any color pattern, and the more inflated lip profile with a deeper
posterior sinus. Sticteulima sp. may be a new non-indigenous species in the Mediter-
ranean Sea.

Vitreolina cf. philippi (de Rayneval & Ponzi, 1854)
Figure 16E, F

New records. IsRAEL * 4 spcms; Ashqelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May
2018; offshore rocky reef; suction sampler; HELM project (samples $16_1F, $16_2F);
size: H 2.6 mm, W 0.9 mm.

Additional material examined. Vitreolina philippi (de Rayneval & Ponzi, 1854):
GREECE ® Crete, Plakias; 35.1796°N, 24.3957°E; depth 5 m; 24 Sep. 2017; Posido-
nia oceanica rhizomes; suction sampler (sample Rh.05_5M).

Remarks. This Vitreolina is extremely similar to the native V. philippi, but the
animal is whitish with a yellowish digestive gland (Figure 16E, F), in contrast to
the peculiar color pattern of typical V. philippi with a white background and red
dots (Figure 16D). Vitreolina is known to be gonochorous (Warén 1984) but it is
unclear if this different color pattern, never reported from the Mediterranean, can
be related to sex. We suspect that this could be another new Lessepsian species for
the Mediterranean Sea, because we observed several Mediterranean-Red Sea species
pairs that are morphologically extremely similar. If we are correct, the occurrence

38 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 19. Sticteulima sp. A-F north of Atlit, Israel, HELM project (sample NG10_1F): front (A, B),
side (C, D) and back (E) views, protoconch (F) G=I Ashgelon, Israel, HELM project (sample S59_1F):
front (G), side (H) and back (I) views. Scale bars: 0.3 mm (A=E, G=I); 0.1 mm (F).

and distribution of this species in the Mediterranean may be difficult to trace, be-
cause empty shells, the most easily collected, are virtually indistinguishable from the

native V. philippi.

Family Conidae J. Fleming, 1822

Conus fumigatus Hwass in Bruguiére, 1792
Figure 20

New records. IsraEL * 3 shs; north of Atlit; 32.7433°N, 34.9067°E; depth 40 m; 20
Sep. 2016; coarse biogenic sediment in a pool among rocks covered by coralligenous for-
mations; grab; HELM project (sample NG40_2M); NHMW-MO-112930/LM/0175;
size of the largest shell (illustrated shell 2, Figure 20D-F): H 7.2 mm, W 4.0 mm.

New non-indigenous species in the Mediterranean 39

Figure 20. Conus fumigatus Hwass in Bruguiére, 1792, NHMW-MO-112930/LM/0175, north of
Atlit, Israel, HELM project (sample NG40_2M) A=C Shell 1: front (A), side (B) and back (C) views
D-F Shell 2: front (D), side (E) and back (F) views. Scale bars: 1 mm.

Remarks. Conus fumigatus was first recorded from the Mediterranean Sea in Libya
(Réckel 1986) but not recorded again for three decades until a recent report from Syria
(Ammar 2018). This is the first finding in Israel, filling the distributional gap from the
Suez Canal northward; only shells of juveniles have been found so far.

Family Murchisonellidae T.L. Casey, 1904

Henrya (?) sp.
Figure 21

New records. IsRaEL * 1 spcm; Palmachim; 31.9574°N, 34.6645°E; depth 36.2 m; 24 May
2017; soft substrate; box-corer; Shafdan project (sample 24(B)); size: H 1.6 mm, W 0.8 mm.

40 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 21. Henrya (?) sp., Palmachim, Israel, Shafdan project (sample 24(B)): front (A, B), side (C, D)
and back (E, F) views, apex (G) and apical view of the protoconch (H). Scale bars: 0.5 mm (A-F);
0.2 mm (G, H).

Remarks. We were unable to assign this species to any Mediterranean or Indo-
Pacific species, despite its conspicuous combination of shell characters. Our single
specimen has an elongated, pupoid shell with convex whorls, a narrow but deeply
incised suture, and a heterostrophic protoconch of type B (diameter: 250 um). The
surface is glossy and smooth except for densely spaced, very fine growth lines. The
latter are straight, slightly prosocline on the spire, becoming orthocline near the aper-
ture. [he aperture is drop-shaped with a simple, thin lip that is slightly reflected at the
columella. An umbilical chink is present. The shell is translucid-white, ornamented
with a single, broad, light brown spiral color band. The shell morphology is similar
to species of the murchisonellid genus Henrya Bartsch, 1947. However, the three cur-
rently known species of that genus were described form the tropical West Atlantic
(Florida, Bahamas, and Yucatan) (Bartsch 1947), and none of them has a brown color
band. For these reasons, the lack of anatomical and molecular data, and the fact that
only a single specimen was available for study, we refrained from a definitive generic

New non-indigenous species in the Mediterranean 4]

assignment. This species is potentially another non-indigenous one originating from
the Indo-Pacific.

Among Mediterranean gastropods, the shell shape somewhat resembles the ira-
vadiid Hyala vitrea (Montagu, 1803), however, the semi-immersed protoconch and
brown color band of Henrya (?) sp. immediately set it apart. The heterobranch Cima
minima (Jeffreys, 1858) is smaller, has a more concial shape, flexuous growth lines,
and also lacks the brown band (van Aartsen 1981; Gofas et al. 2011; Scaperrotta et al.
2012; Giannuzzi-Savelli et al. 2014).

Family Pyramidellidae Gray, 1840

Odostomia cf. dalli (Hornung & Mermod, 1925)
Figure 22

New records. IsRAEL * 4 spcms; west of Rosh HaNikra Islands; 33.0704°N, 35.0926°E;
depth 12 m; 29 Oct. 2018; rocky substrate; suction sampler; HELM project (samples
S52_1KF S52_2F); size: H 2.1 mm, W 1.0 mm (illustrated specimen) * 1 spcm; west
of Rosh HaNikra Islands; 33.0725°N, 35.0923°E; depth 19 m; 29 Oct. 2018; rocky
substrate; suction sampler; HELM project (sample $53_1F).

Remarks. The shell of this species is white and rather solid, with convex, un-
keeled whorls and a deep, narrow suture. The columellar tooth is visible in frontal
view; there are no lirae inside the aperture. The outer surface appears smooth at
first sight but bears numerous very fine spiral lines. The protoconch is of type A2,
tending to type B. In ethanol-preserved specimens, the soft body is yellowish-white,
with the eyes well visible through the shell (Figure 22A). Odostomia cf. dalli differs
in its shell morphology from all known Mediterranean Odostomiinae, but bears
close resemblance to the illustration of the type specimen of Odostomia dalli from
Sarad Island (“He de Sarato”), Dahlak Archipelago, Eritrean Red Sea (Hornung and
Mermod 1925). In contrast to our material, however, O. dalli was described as lack-
ing both, spiral sculpture and a visible columellar fold, although a columellar tooth
seems to be indicated in the line drawing accompanying the original description.
A rigorous assessment of potential conspecificity between O. dalli and our material
therefore awaits a thorough study of the type material of the former, but the close
similarity suggests that this is a new non-indigenous species in the Mediterranean
Sea. This interpretation is also supported by the lack of empty shells in death assem-
blages (see Discussion).

Odostomia (s.1.) sp. 1
Figure 23

New records. IsRaEL * 1 spcm; Haifa Bay; 32.8211°N, 35.0196°E; depth 11 m; 2 Aug.
2015; soft substrate; grab; NM project (sample HM27(c)); size: H 1.4 mm, W 0.7 mm

42 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 22. Odostomia cf. dalli (Hornung & Mermod, 1925), west of Rosh HaNikra Islands, Israel,
HELM project (sample $52_2F): front (A) and back (B) views. Scale bar: 0.5 mm.

(illustrated specimen) * 8 spcms; 31.9364°N, 34.6846°E; depth 20.2 m; 11 Oct. 2012;
sandy substrate; grab; Via Maris project (sample VM40).

Remarks. This species is characterized by a translucid-white, cylindrical shell with ~
3 whorls, and an intorted protoconch of type C (Figure 231) whose columella is oriented
at an angle of ~ 160° relative to the teleoconch axis (revealed by wCT-imaging, Figure
23H and additional scans available at https://doi.org/10.6084/m9. figshare.c.5215226).
The growth lines are slightly prosocline on the spire while becoming almost orthocline
on the body whorl; an extremely faint spiral microsculpture is present on the apical part
of the whorls, but only visible in high-magnification SEM images (Figure 23G). This
species differs from Odostomia cf. dalli by its smaller size (height up to 1.4 mm), the
more cylindrical shape, shallower suture, and the absence of a visible columellar tooth.
Although this species, in terms of size and overall shape, somewhat resembles representa-
tives of the fresh- and brackish water-dwelling family Hydrobiidae, the fact that numer-
ous living specimens were found in a fully marine environment and its heterostrophic
protoconch unambiguously identify it as member of the family Pyramidellidae.

Odostomia sp. 1 does not resemble any known Mediterranean pyramidellid; consider-
ing the great number of confirmed introductions of Indo-Pacific microgastropods to the
eastern Mediterranean Sea, we therefore suspect that also this taxon might be a Lessepsian
species. Among Indo-Pacific Odostomiinae, O. bullula Gould, 1861 (e.g., Johnson 1964;
Robba et al. 2004) is similar to our specimens, but differs by its more conical shape and
larger size (height to 2 mm, width to 1 mm). Another similar species, O. decouxi Saurin,
1959, was suggested to be a junior synonym of O, bullula (Robba et al. 2004).

New non-indigenous species in the Mediterranean 43

Figure 23. Odostomia (s.l.) sp. 1, Haifa Bay, Israel, NM project (sample HM27(c)): front (A, B), side
(C, D) and back (E, F) views, detail of the adapical part of the body whorl showing the extremely faint

spiral microsculpture (G), virtual section through the apical spire showing the columella of the intorted
protoconch (H) and apical view of the protoconch (I; surface partly corroded). The pink hue is due to
staining with eosin solution. Scale bars: 0.5 mm (A=F); 0.2 mm (G-I).

44 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 24. Odostomia (s.1.) sp. 2, Neve Yam, Atlit, Israel: front (A, B), side (C, D) and back (E, F) views,
apex (G) and apical view of the protoconch (H). Scale bars: 0.5 mm (A=F); 0.2 mm (G, H).

Odostomia (s.1.) sp. 2
Figure 24

New records. IsraEL * 1 spcm; Soreq desalination plant; 31.9420°N, 34.6896°E;
depth 17.4 m; 19 May 2015; soft substrate; grab; Soreg project (sample S027).

Additional material examined. Isrart * 1 sh; Neve Yam, Atlit; 32.6785°N,
34.9289°E; 6 Feb. 2006; beached; size H 1.3 mm, W 0.7 mm (illustrated shell; previ-
ously figured by Bogi and Galil (2006)).

Remarks. ‘The first record of this species is based on five well-preserved shells found
in a shell grit sample taken in 1995 on the beach of Yumurtalik, Adana, Turkey (Giunchi
et al. 2001). In 2006, another beached shell was found at Neve Yam, northern Israel (Bogi
and Galil 2006, re-illustrated in Figure 24 herein) and, according to these authors, the
species was also found in Israel by J.J. van Aartsen. A specimen of Odostomia sp. 2, from
the original lot from Yumurtalik, was recently figured by Giannuzzi-Savelli et al. (2014).
Here, we report the first finding of a living individual of Odostomia sp. 2 which was re-
covered from a sediment sample taken at the Soreq desalination plant, southern Israel.

New non-indigenous species in the Mediterranean 45

Since the first finding in Turkey 25 years ago, the identity of this most likely non-
indigenous species has remained unresolved. It differs from all known Mediterranean
Odostomiinae at first glance by the presence of two brown spiral bands. We are una-
ware of any Indo-Pacific pyramidellid resembling this taxon, and it may well represent
an undescribed species. To aid the further study of this taxon and raise awareness of
its presence and apparent spread in the Mediterranean, we here re-illustrate the well-
preserved shell from Neve Yam using light and scanning electron microscopy.

Oscilla virginiae Penas, Rolan & Sabelli, 2020
Figure 25A, B

New records. IsRAEL * 1 spcm; west of Rosh HaNikra Islands; 33.0704°N, 35.0926°E;
depth 12 m; 29 Oct. 2018; rocky substrate; suction sampler; HELM project (sample
S52_1F) * 1 spcm; west of Rosh HaNikra Islands; 33.0725°N, 35.0923°E; depth 19
m; 29 Oct. 2018; rocky substrate; suction sampler; HELM project (sample S53_1F);
size: H 1.6 mm, W 0.9 mm (illustrated specimen).

Additional material examined. Oscilla appeliusi (Hornung & Mermod, 1925):
EGYPT ¢ 1 sh; Sinai (Red Sea), south of Sharm-el-Sheik, Na’ama Bay; 27.8500°N,
34.2833°E; depth 0-6 m; 28 Sep. -4 Oct. 1978; rocky substrate; A.J. Ferreira leg; LACM
1978-79.4 © 1 sh; Sinai (Red Sea), Gulf of Aqaba, Strait of Tiran, Jackson Reef; 28.0167°N,
34.4667°E; depth 2—3 m; 31 Oct. 1985; sand and coral substrate; T. Bratcher leg.; LACM
1985-111.5 © 2 shs; Sinai (Red Sea), Gulf of Aqaba, Strait of Tiran, east side Jackson Reef;
28.000°N, 34.4667°E; depth 10 m; 9 Jul. 1988; coral rubble; J. H. Golden leg.; LACM
1988-118.1 © 4 shs; Sinai (Red Sea), off Ras Umm Sid, “Amphoras” dive site; 27.8667°N,
34.333°E; depth 18 m; 24 Jul. 1988; coral rubble; J.H. Golden leg.; LACM 1988-119.2.

InponEsiA * 7 shs; Papua Province, south-east of Biak Island, east side of Auki
Islet; 1.2300° S, 136.3367°E; depth 0 m; 5 Apr. 1988; rock; J.H. McLean & E. Abbott
leg.; LACM 1988-48.12.

Tarwan ° 7 shs; Tai-Pei County, east of Chi-lung (= Keelung), south-east side of
Pitou Chiao (= Pitou Nonkow); 25.1333°N, 121.9167°E; depth 0-3 m; 10 May 1988;
rocky tide pool; C.C. Coney & PE Liu leg.; LACM 1988-80.13.

Miralda sp. (Oscilla jocosa sensu van Aartsen et al. (1989)): IsraEL * 1 spcm; west
of Rosh HaNikra Islands; 33.0704°N, 35.0926°E; depth 12 m; 29 Oct. 2018; rocky
substrate; suction sampler; HELM project (sample $52_1F); size: H 2.0 mm, W 1.0
mm (illustrated specimen).

Remarks. Oscilla virginiae is characterized by a small-sized, white, conical shell
with a type A protoconch. The sculpture consists of thick, smooth spiral cords: the
first and second whorl bear two cords; the upper cord is broadest and bifurcates on
the third whorl, forming three cords on the last whorl, with the newly formed pair
remaining positioned very close one to each other (Figure 25A, B; Pefas et al., 2020).

This species has just been described from the infralittoral of Jordan and also occurs in
the Egyptian Red Sea (Pefias et al. 2020). It superficially resembles the Indo-Pacific O. ap-
peliusi (Hornung & Mermod, 1925), and indeed, a juvenile shell from Dahab (Egypt) was

46 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 25. Comparison between Oscilla virginiae Pefas, Rolan & Sabelli, 2020 and Miralda sp.
A, B Oscilla virginiae, west of Rosh HaNikra Islands, Israel, HELM project (sample S53_1F): front (A)
and back (B) views C, D Miralda sp. (Oscilla jocosa sensu van Aartsen et al. (1989)), west of Rosh HaNikra
Islands, Israel, HELM project (sample S52_1F): front (C) and back (D) views. Scale bars: 0.5 mm.

recently figured by Blatterer (2019: plate 212, fig. 17c, d) under this name. In contrast to O.
virginiae, however, O. appeliusi bears spiral cords more similar in thickness which are spaced
more equidistantly and closer to each other. Already on the second whorl, three cords are
present, and the uppermost cord does not evidently bifurcate (Pefas et al. 2020). Lastly, the
illustration of Hornung and Mermod (1925) suggests a greater number of spiral cords on
the last whorl. ‘To date, O. appeliusi has not been recorded from the Mediterranean Sea.
Within the Mediterranean, O. virginiae is superficially similar only to two other non-
indigenous pyramidellids, Cingulina isseli (Iryon, 1886) and Miralda sp. (Figure 25C, D).
The latter taxon has previously been reported under the name Oscilla jocosa Melvill, 1904,
despite recent evidence by Pefas & Rolan (2017) that it is not conspecific with Melvill’s
(1904) type material. Oscilla virginiae differs from C. isseli by its broader, more conical
shell, fewer whorls, smaller size (C. isseli reaches a height of ~ 3 mm), and the much less
pronounced axial sculpture between the spiral cords. Compared to Miralda sp., O. virgini-
ae differs by its smaller size (up to ~ 3 mm in Miralda sp.), the absence of beads on the two
upper spiral cords (Figure 25C, D), and stronger spiral cords on the base of the shell. An-
other Oscilla present in the Mediterranean Sea is O. galilae Bogi, Kharan & Yokes, 2012,
which can be easily distinguished from O. virginiae by the smaller size, the oval profile,
the oblique spiral sculpture, the more prominent axial sculpture and a type C protoconch.
Penas et al. (2020) illustrated O. galilae (their Fig. 383A—E) under the name O. cylindrica
(de Folin, 1879) suggesting synonymy between the two names. However, O. cylindrica
can be easily distinguished from O. galilae because it bears spiral cords without interspaces,
because it does not bear the fine axial lamellae typical of O. galilae and because of its more
cylindrical profile. The Red Sea records of O. virginiae and O. galilae by Pefias et al. enable

to assess that both species are non-indigenous in the Mediterranean Sea.

New non-indigenous species in the Mediterranean 47

Parthenina cossmanni (Hornung & Mermod, 1924)
Figure 26A—H

New records. Israe * 1 sh; north of Atlit; 32.7422°N, 34.9181°E; depth 30 m; 20
Sep. 2016; sand; grab; HELM project (sample NG30_2F); size: H 1.9 mm, W 0.9 mm
(illustrated shell, Figure 26A—F) * 9 spcms, 2 shs; Ashqelon; 31.7002°N, 34.5498°E;
depth 21 m; 18 Sep. 2016; sand; grab; HELM project (samples SG20_1F, SG20_2F
SG20_4F, SG20_5F); size: H 2.0 mm, W 0.8 mm (illustrated specimen, Figure 26G,
H) ¢ 1 spcm; Ashgelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr. 2018; offshore
rocky reef; suction sampler; HELM project (sample $12_1F).

Additional material examined. Parthenina indistincta (Montagu, 1808): IsRAEL
e 2 shs; Ashqelon; 31.7002°N, 34.5498°E; depth 21 m; 18 Sep. 2016; sand; grab;
HELM project (sample SG20_2F) ¢ 1 sh; Ashgelon; 31.7101°N, 34.5406°E; depth
31 m; silty sand, 15 cm below sediment surface; gravity corer; HELM project (sample
SC30_1_15L); size: H 1.7 mm, W 0.7 mm (illustrated shell).

Remarks. Parthenina cossmanni has an elongated-conical shell with flat-convex
whorls and a protoconch of type C. The whorls of the spire have a subangular pro-
file, whereas the body whorl in adult specimens is more convex and evenly rounded.
The axial sculpture is made of strong orthocline ribs that become slightly flexuous on
the body whorl in some specimens (Figure 26A—E). The spiral sculpture on the spire
consists of a single, thin, suprasutural cord; a second cord emerges on the penultimate
whorl, and three cords are present on the last whorl. The columellar tooth is weak
and deeply inset, and in some of the studied specimens hardly visible inside the aper-
ture. The soft body of ethanol-preserved specimens is yellowish, with the eyes visible
through the shell (Figure 26G, H).

The type material of P cossmanni was collected from the Red Sea of Massawa
(Eritrea) at a depth of 30 m (Hornung and Mermod 1924); the species was recently
recorded from Dahab (Gulf of Aqaba, northern Egypt) by Blatterer (2019), and from
Jordan by Pefas et al. (2020). Outside the Red Sea, it is known from Vietnam (Saurin
1959) and Thailand (Robba et al. 2004).

Among native Mediterranean species, P cossmanni superficially resembles Partheni-
na interstincta (J. Adams, 1797). The latter species, however, has only two spiral cords
on the last whorl and a more developed columellar tooth. P cossmanni is further similar
to P indistincta (Montagu 1808) (Figure 26I—L) which has a very weak, internal colu-
mellar fold (Warén 199 1a: 96, fig. 29f) and three (rarely four) spiral cords on the last
whorl. Compared to P cossmanni, however, the shell of P indistincta is more elongated
and has two spiral cords on the spire whorls.

We suspect the two shells illustrated as P indistincta in Oztiirk et al. (2011: fig. 10A,
B) might also be P cossmanni, considering their broad shape and overall morphology.
Oztiirk et al.’s material was collected in 2009 from a mud bottom at 9 m depth in Mersin
Bay (stn. 46, 36.7167°N, 34.8667°E), south-eastern Turkey; should our hypothesis be
confirmed upon re-examination of these shells, this would suggest that P cossmanni likely
has a wider distribution in the southeastern Mediterranean Sea. Our finding of several

48 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 26. Comparison between Parthenina cossmanni (Hornung & Mermod, 1924) and Parthenina
indistincta (Montagu, 1808) A=F Parthenina cossmanni, north of Atlit, Israel, HELM project (sample
NG30_2F): front (A, C) and back (B, D) views, detail of the spiral sculpture on the body whorl (E) and
apical view of the protoconch (F) G, H Parthenina cossmanni, Ashgelon, Israel, HELM project (sample
SG20_4F): front (G) and back (H) views I=L Parthenina indistincta, Ashqelon, Israel, HELM project
(sample SC30_1_15L): front (I, K) and back (J, L) views. Scale bars: 0.5 mm (A-E, G-L); 0.1 mm (F).

New non-indigenous species in the Mediterranean 49

living specimens on the Israeli shelf, together with the relative rarity of empty shells in
the samples, suggests that this species might have established locally only rather recently.

Parthenina typica (Laseron, 1959)
Figure 27

New records. IsRAEL * 2 spcms; Palmachim; 31.9737°N, 34.6767°E; depth 35.4 m;
2 Sep. 2015; soft substrate; box-corer; Shafdan project (sample 29(C)) * 4 spcms; Pal-
machim; 31.9685°N, 34.6732°E; depth 35.6 m; 2 Sep. 2015; soft substrate; box-corer;
Shafdan project (sample 26(B)); size: H 2.0 mm, W 0.8 mm (illustrated specimen) ¢ 1
sh; north of Atlit; 32.7433°N, 34.9067°E; depth 40 m; 20 Sep. 2016; coarse biogenic
sediment in a pool among rocks covered by coralligenous formations; grab; HELM
project (sample NG40_2M) ¢ 1 spcm; Ashqelon; 31.7487°N, 34.4960°E; depth 41 m;
18 Sep. 2016; sandy mud; grab; HELM project (sample SG40_1F).

Remarks. ‘This species is characterized by a straight, conical profile with flat whorls,
separated by a deep, canaliculate suture; the abapical part of the whorl is angulated.
The sculpture consists of straight axial ribs and a prominent suprasutural spiral cord;
the base is smooth except for faint continuations of the axial ribs; an internal columel-
lar fold is present and visible inside the aperture when slightly turning the shell to the
left side. The protoconch is of type C and in the illustrated specimen it has a diameter
of ~ 240 um, which is slightly smaller than 270-290 um stated by Pefias and Rolan
(2017) for this species.

Parthenina typica (Laseron, 1959) was described from eastern Australia (Laseron
1959) and subsequently recorded from the Solomon Islands, Fiji and the Philippines
at infralittoral to bathyal depths (Pefas and Roldan 2017). To our knowledge, it has
not been reported from the Indian Ocean nor from the Red Sea, however, its absence
could well represent an artifact of the limited knowledge of the micromollusk fauna of
these regions. Among native species, the conchologically highly variable Parthenina in-
terstincta (J. Adams, 1797) and P monozona (Brusina, 1869) are most similar, however,
they differ by having more rounded whorls and a greater number of axial ribs.

Pyrgulina craticulata (Issel, 1869)
Figure 28A-I

New records. Israzx ® 1 sh; north of Atlit; 32.7820°N, 34.9466°E; depth 10 m; 21 Sep.
2016; sand; grab; HELM project (sample NG10_1F) * 3 spcms; Ashqelon; 31.6868°N,
34.5516°E; depth 12 m; 30 Apr. 2018; offshore rocky reef; suction sampler; HELM
project (samples $12_2F, $12_3F) ¢ 17 spcms, 1 sh; same collecting data as for preced-
ing; depth 11 m; 31 Oct. 2018; HELM project (samples S58_1E S58_1M, S58_2F
S58_3F); size: H 1.5 mm, W 0.7 mm (illustrated specimen) * 2 spcms; Ashqelon;
31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef; suction sam-

50 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 27. Parthenina typica (Laseron, 1959), Palmachim, Israel, Shafdan project: front (A, B) and back
(C, D) views, apex (E) and apical view of the protoconch (F). Scale bars: 0.4 mm (A=D); 0.1 mm (E, F).

pler; HELM project (sample S$16_2F) ¢ 1 spcm; same collecting data as for preceding;
depth 28 m; 31 Oct. 2018; HELM project (sample S59_2F) * 2 spcms; west of Rosh
HaNikra Islands; 33.0704°N, 35.0926°E; depth 12 m; 29 Oct. 2018; rocky substrate;
suction sampler; HELM project (samples $52_2F, $52_3F) © 1 spcm; west of Rosh
HaNikra Islands; 33.0725°N, 35.0923°E; depth 20 m; 1 May 2018; rocky substrate;
suction sampler; HELM project (sample S13_3F) ¢ 2 spcms; same collecting data as for
preceding; depth 19 m; 29 Oct. 2018; HELM project (sample S53_1F).

Additional material examined. Pyrgulina craticulata: EGYPT ° 1 sh; Sinai (Red
Sea), Gulf of Aqaba, Strait of Tiran, Jackson Reef; 28.0167°N, 34.4667°E; depth
2-3 m; 31 Oct. 1985; sand and coral substrate; T. Bratcher leg.; LACM 1985-111.4.

Mapacascar ® 3 shs; Antsiranana Province, south of Nosy Be, out from Hellville
(= Andoany); 13.4500° S, 48.2500°E; depth 14 m; 5 Apr. 1989; coral heads and gor-
gonians; J.H. McLean leg.; LACM 1989-55.1.

New non-indigenous species in the Mediterranean 51

Figure 28. Comparison between Pyrgulina craticulata (Issel, 1869) and Spiralinella incerta (Milaschewitsch,
1916) A=l Pyrgulina craticulata, Ashqelon, Israel, HELM project (sample $58_3F): front (A, B) and back
(C, D) views, aperture and details of the sculpture on the base (E), top view (F), detail of the protoconch-
teleoconch transition (G), apical view of the protoconch (H) and detail of the sculpture of the body whorl (I)
J-N Spiralinella incerta, west of Rosh HaNikra Islands, Israel, HELM project (sample $14_4F): front J, K)
and back (L, M) views, detail of the sculpture of the body whorl (IN). Scale bars: 0.5 mm (A-D, J—-M);
0.2 mm (E, FI, N); 0.1 mm (G, H).

52 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Matpives * 1 sh; North Male Atoll, 2 km north of Baros Island; 4.3167°N,
73.4167°E; depth 25-35 m; 10 Feb. 1983; gravel; leg. A.J. Ferreira; LACM 1983-9.7.

THAILAND ® 1 sh; Phuket Province, east side of Kaew Yai Island; 7.7450°N,
98.3083°E; depth 0-3 m; 24—26 Mar. 1985; coral; J.H. McLean leg.; LACM 1985-5.17
¢ 1 sh; same collecting data as for preceding; LACM 1985-5.18 6 shs; Phuket Province,
southern tip of Phromthep Cape; 7.7583°N, 98.3217°E; depth 0 m; 24 Mar. 1985;
boulder beach with coral; J.H. McLean leg.; LACM 1985-6.4 © 4 shs; Phuket Prov-
ince, east of Phuket, near Pee-Pee (= Phi Phi) Island, Hin Hmusang Rock; 7.7917°N,
98.5500°E; depth 15 m; 15 Feb. 1985; sand; A.J. Ferreira leg; LACM 1985-14.2.

Spiralinella incerta (Milaschewitsch, 1916): IsrazL® 1 sh; north of Atlit; 32.7820°N,
34.9466°E; depth 10 m; 21 Sep. 2016; sand; grab; HELM project (sample NG10_1F)
¢ Ish; north of Atlit; 32.7422°N, 34.9181°E; depth 30 m; 20 Sep. 2016; sand; grab;
HELM project (sample NG30_2F) ¢ 2 shs; Ashgelon; 31.7002°N, 34.5498°E; depth
21 m; 18 Sep. 2016; sand; grab; HELM project (sample SG20_2F) ¢ 1 sh; Ashgelon;
31.7101°N, 34.5406°E; depth 31 m; silty sand, 90 cm below sediment surface; gravity
corer; HELM project (sample SC30_1_90L) © 1 sh; west of Rosh HaNikra Islands;
33.0704°N, 35.0926°E; depth 12 m; 1 May 2018; rocky substrate; suction sampler;
HELM project (sample $14_4F); size: H 1.5 mm, W 0.7 mm (illustrated shell).

Remarks. Issel (1869) described Odontostomia craticulata based on an illustration
of a shell from the Red Sea by Savigny (1817: pl. 3, fig. 39); the syntype from the
Savigny collection (MNHN-IM-2000-34095) was later figured by Bouchet & Danri-
gal (1982) and is shown on the MNHN website (https://science.mnhn.fr/institution/
mnhn/collection/im/item/2000-34095?listIndex=158listCount=44; last accessed 26
July 2020). As Odontostomia is considered an unjustified emendation of Odostomia
(MolluscaBase 2020), a genus with very different conchological characters from P crat-
iculata, we follow the opinion of Pefias et al. (2020) and use the combination Pyrgulina
craticulata (Issel, 1869).

Pyrgulina craticulata has been reported only from the Red Sea so far, however, a
search of previously uncatalogued lots of pyramidellid shells in the LACM collection
by one of us (PILF) yielded specimens from Madagascar, the Maldives and Thailand,
confirming that this species has a much wider distribution in the Indian Ocean. Shells
of P craticulata seem indistinguishable from the illustration of Chrysallida tribulationis
(Hedley, 1909), a taxon recorded from Australia and the western Japan Sea (Hedley
1909; Higo et al. 2001). In the light of a possible distribution of P craticulata also in
the West Pacific, we recommend an assessment of potential synonymy between C.
tribulationis and P craticulata.

Here, we provide the first records of P craticulata for the Mediterranean Sea. Sev-
eral living specimens were found on hard substrates off southern and northern Israel,
suggesting it is established in the region. In terms of shell size, shape and type of orna-
mentation, this species closely resembles the native Spiralinella incerta (Figure 28J—N),
but has pronounced spiral cords in the interspaces of the axial ribs (Figure 28I vs. 28N)
which enable a reliable segregation of these two species. In addition, the axial ribs are
spaced more closely in S. incerta than in P craticulata.

New non-indigenous species in the Mediterranean 55

Pyrgulina nana Hornung & Mermod, 1924

New records. IsraEL * 2 spcms; Ashqelon; 31.6891°N, 34.5257°E; depth 28 m; 31 Oct
2018; offshore rocky reef; suction sampler; HELM project (sample $59_2F) ¢ 1 spcm;
west of Rosh HaNikra Islands; 33.0704°N, 35.0926°E; depth 12 m; 1 May 2018; rocky
substrate; suction sampler; HELM project (sample S14_4F) * 5 spcms; same collecting
data as for preceding; 29 Oct. 2018; HELM project (samples $52_1E S52_2F).
Remarks. Until now, the first record of this species from the Mediterranean was
considered to be by Oztiirk and van Aartsen (2006), who reported on material obtained
from shallow-water sediment samples collected along the Turkish Levantine (Viranse-
hir, Mersin Bay) and Aegean coasts (Giilliik Bay) in 1997 and 2000, respectively. How-
ever, already van der Linden and Eikenboom (1992) described and illustrated P nana
from the Levantine Sea (page 60, Figure 41), referring to it as Chrysallida spec. C in
the lack of a species-level identification. Their material consisted of a single individual,
likely an empty shell, but not specified by the authors, from Mersin (south-eastern
Turkey) with unknown collecting date, housed in the collection of J. van der Linden
(The Hague, The Netherlands). Although we were unable to examine this material, the
excellent and detailed line drawing provided enabled an unambiguous assignment of
Chrysallida spec. C to P nana; thus, van der Linden and Eikenboom (1992) should be
regarded the first Mediterranean record. Today, the known Mediterranean distribution
of P nana includes Turkey, Lebanon, and Israel (Bogi and Galil 2006; Giannuzzi-
Savelli et al. 2014). To our knowledge, this is the first record of living individuals of
P. nana from Israel; here, the species occurs along both the southern (Ashqelon) and
northern coasts (west of Rosh HaNikra Islands) on rocky bottoms at 12—28 m depth.

Turbonilla funiculata de Folin, 1868
Figure 29

New records. IsRaEL * 2 spcms; west of Rosh HaNikra Islands; 33.0704°N,
35.0926°E; depth 12 m; 1 May 2018; rocky substrate; suction sampler; HELM
project (sample S14_4F) * 15 spcms; same collecting data as for preceding; 29
Oct. 2018; HELM project (samples $52_1F, $52_2F, $52_3F); size: H 1.8 mm, W
0.6 mm (illustrated specimen).

Additional material examined. Curna ¢ 10 shs; Hong Kong, Tolo Channel, cove
at Hoi Sing Wan; 22.430°N, 114.2467°E; depth 0 m; 6 Apr. 1985; sand, rock and
oysters; J.H. McLean leg.; LACM 1985-12.2.

PaxisTAN ° 4 shs; Sind Province, small cove 4.8 km west of atomic power plant and
7 km west-northwest of Bulegi Point; 24.8000°N, 66.7250°E; depth 0-4 m; 19 Jan.
1979; rock and clay; C.C. Swift leg.; LACM 1979-1.3.

Srt Lanka * 1 sh; Southern Province, Tangalla, cove at Tangalla Bay Hotel;
6.0250°N, 80.8000°E; depth 0-1 m; 30-31 Jan. 1983; rock and clay; A.J. Ferreira
leg.; LACM 1983-5.2.

Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 29. Turbonilla funiculata de Folin, 1868, west of Rosh HaNikra Islands, Israel, HELM project
(sample $52_2F): front (A, B) and back (C, D) views, detail of the sculpture on the base (E), aperture (F),
detail of the sculpture on the back of the third whorl (G), apical (IH) and side (I) views of the protoconch.
Scale bars: 0.5 mm (A=D); 0.1 mm (E, H, 1); 0.2 mm (F, G).

VIETNAM ® 1 sh; station 1328 of the Nha-Trang Oceanography Institute: “Entre
les tles des Pécheurs, Hon-Mung, le banc du Castleragh et Pisobathe 50. (En dehors
et au S. de la baie de Nha-Trang)” [between Iles des Pécheurs, Hon-Mung, Banc du

New non-indigenous species in the Mediterranean 55

Castleragh (= Castlereagh) and isobath 50. (outside of and to the south of the Bay of
Nha-Trang)]; sediment sample; dredge; MNHN-IM-2000-21843 (holotype of Pyrgis-
cus mirandus Saurin, 1959).

Remarks. Shells of Turbonilla funiculata are polymorphic with respect to their
shape (Pefias and Roldan 2010 and our own observations) but the species can be readily
distinguished from all pyramidellids in the Mediterranean — the most similar being the
non-indigenous Turbonilla edgarii (Melvill, 1896), Turbonilla flaianoi Mazziotti, Aga-
mennone, Micali & Tisselli, 2006 and Pyrgulina fischeri Hornung & Mermod, 1925
— by the presence of a very marked subsutural constriction running along the whorls.
This constriction separates the pronounced, almost orthocline axial ribs into a larger
lower and a narrow upper, crown-like, portion. The ribs extend adapically beyond the
suture of the preceding whorl, giving the transition zone between subsequent whorls
a wavy appearance. The interspaces of the lower portion of the ribs bear several thin
spiral lines, while those between the narrow upper parts of the ribs are smooth. ‘The
protoconch is helicoid and of type A.

Turbonilla funiculata has been previously reported from Fiji, Hong Kong, Indone-
sia, New Caledonia, Thailand, the Solomon Islands and Vietnam from shore to 396 m
depth (Robba et al. 2004; Pefas and Rolan 2010). As Robba et al. (2004) already point-
ed out, the shell figured as Pyrgiscus microscopica (Laseron, 1959) by Okutani (2000:
712, fig. 68) is most likely 7’ funiculata, confirming that the species also occurs in Ja-
pan. This interpretation is re-afirmed by another illustration of the very same Japanese
specimen in Mazziotti et al. (2005: 81, fig. 1m, n), showing the subsutural constriction
characteristic of 7! funiculata. Shells of T’ funiculata were found by one of us (PILF) also
among hitherto unidentified lots of shells from Pakistan and Sri Lanka housed in the
LACM collection, demonstrating that this species also lives in the Indian Ocean. Here,
we report the first records of 7’ funiculata for the Mediterranean, where several living
specimens were collected in northern Israel on hard substrates at 12 m depth.

Family Cylichnidae H. Adams & A. Adams, 1854

Cylichna collyra Melvill, 1906
Figure 30

New records. IsraEL * 1 spcm; Palmachim; 31.9477°N, 34.6562°E; depth 37.5 m; 18
Oct. 2017; soft substrate; box-corer; Shafdan project (sample 22(A)); size: H 4.2 mm, W
1.5 mm (illustrated specimen) * 1 spcm; Palmachim; 31.9424°N, 34.655 1°E; depth 36.3
m; 2 May 2018; soft substrate; box-corer; Shafdan project (sample 21(A)); size: H 7.2
mm, W 4.0 mm ¢ 2 spcms; Palmachim; 31.9376°N, 34.6515°E; depth 36.7 m; 2 May
2018; soft substrate; box-corer; Shafdan project (sample 5(B)) * 1 spcm; Palmachim;
31.9685°N, 34.6732°E; depth 35.6 m; 2 May 2018; soft substrate; box-corer; Shafdan
project (sample 26(A)) * 1 spcm; Palmachim; 31.9477°N, 34.6562°E; depth 37.5 m;
2 May 2018; soft substrate; box-corer; Shafdan project (sample 22(B)) * 3 spcms; Pal-
machim; 31.9424°N, 34.6551°E; depth 36.3 m; 2 May 2018; soft substrate; box-corer;
Shafdan project (sample 21(C)) * 2 spcms; Palmachim; 31.9327°N, 34.6495°E; depth

56 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

~
Ze &

; oe pee * ee x §
c PEE op Wier St er,

<} pee aS eS
a eh We > *

ae

Figure 30. Cylichna collyra Melvill, 1906, Palmachim, Israel, Shafdan project (sample 22(A)): front (A, B),
side (C) and back (D) views, and apical (E) and anterior (F) microsculpture. Scale bars: 1 mm (A=D);
0.2 mm (E, F).

36.2 m; 2 May 2018; soft substrate; box-corer; Shafdan project (sample 4(A)) * 1 spcm;
Palmachim; 31.9574°N, 34.6645°E; depth 36.2 m; 2 May 2018; soft substrate; box-
corer; Shafdan project (sample 24(C)) * 1 spcm; Palmachim; 31.9477°N, 34.6562°E;
depth 37.5 m; 20 Oct. 2018; soft substrate; box-corer; Shafdan project (sample 22(A)).
Additional material examined. Oman °¢ 13 shs; off Muscat; 24°58'N, 56°54'E;
156 fathoms (285 m) depth; NMW.1955.158.00578 (EW. Townsend coll.).
Remarks. We record here for the first time in the Mediterranean 13 living in-
dividuals of Cylichna collyra, a cephalaspidean originally described from the Gulf of
Oman (Melvill 1906b). Cylichna collyra can be distinguished from the native Medi-
terranean C. cylindracea (Pennant, 1777) by its more elongated and slender shell, the
more tapering apical part, the color pattern characterized by fine brown spiral lines
apically and abapically, and the smaller size (C. cylindracea commonly reaches 1 cm

New non-indigenous species in the Mediterranean +f

in height whereas C. collyra attains approximately half that size). Cylichna villersii
(Audouin, 1826), another non-indigenous species of Red Sea origin recorded from
the Mediterranean coast of Israel (Bogi and Galil 2013a), is smaller (less than 2 mm),
less slender, has a more rounded base and stronger growth marks (not visible in C.
collyra), and bears two brown bands apically and abapically instead of the fine brown
lines. Cylichna biplicata (A. Adams in Sowerby, 1850), a species occurring on the con-
tinental platform in the Indo-West Pacific, shares with our specimens the cylindrical
shape and the color pattern of reddish-brown spiral bands apically and abapically
(Valdés 2008), but is larger, more elongated anteriorly, with a stronger columellar
tooth, and the colored spiral bands become a compact larger band apically. Cylichna
collyra has not been recorded from the Red Sea yet (Dekker and Orlin 2000).

Family Mnestiidae Oskars, Bouchet & Malaquias, 2015

Mnuestia girardi (Audouin, 1826)

New records. IsRAEL * 5 spcms; west of Rosh HaNikra Islands; 33.0704°N, 35.0926°E;
depth 12 m; 29 Oct. 2018; HELM project (samples S52_1F S$52_2F, S52_3F)
1 spcm, 2 shs; west of Rosh HaNikra Islands; 33.0725°N, 35.0923°E; depth 20 m;
1 May 2018; rocky substrate; suction sampler; HELM project (samples $13_2M,
S13_3F) ¢ 12 spcms; same collecting data as for preceding; depth 19 m; 29 Oct. 2018;
HELM project (samples $53_1F, $53_2F S53_2M, S53_3F).

Remarks. This species was first recorded in the Mediterranean Sea in 1974 with
the finding of few empty shells in the Bardawil Lagoons in Egypt (Mienis 1976). The
species has since been recorded also in Greece (Crocetta et al. 2017), Turkey (Cinar et
al. 2011), Cyprus (Katsanevakis et al. 2009), and Lebanon (Crocetta et al. 2020). A
single record is available from Israel based on shells collected in 2004 off Palmachim
(Bogi and Galil 2006). We here report living individuals from Israel for the first time.
This is also the first finding of living individuals in the Mediterranean Sea.

Family Haminoeidae Pilsbry, 1895

Atys angustatus E.A. Smith, 1872
Figure 31

New records. GREECE * | sh; Crete, Plakias; 35.1796°N, 24.3957°E; depth 15 m;
21 Sep. 2017; Posidonia oceanica rhizomes; suction sampler (sample Rh.15_5M); size:
H 2.6 mm, W not available because of broken aperture.

Remarks. Atys angustatus was first recorded in the Mediterranean Sea in 1974,
based on specimens collected at Haifa, Israel (Aartsen and Goud 2006). It has been
reported from Mersin, Turkey, since 1986 (Aartsen and Goud 2006) and multiple
records from Israel followed from several locations along its coast (Micali et al. 2016).
To the best of our knowledge, this is the first record from Greece.

58 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 31. Atys angustatus E.A. Smith, 1872, Plakias, Crete, Greece (sample Rh.15_5M): front view.
Scale bar: 0.5 mm.

Class Bivalvia Linnaeus, 1758
Family Mytilidae Rafinesque, 1815

Arcuatula perfragilis (Dunker, 1857)
Figure 32

New records. IsraeEL * 1 spcm; Palmachim; 31.9574°N, 34.6645°E; depth 36.2 m; 2
May 2018; soft substrate; box-corer; Shafdan project (sample 24(C)); size: L 13 mm,
H 5.3 mm ¢ | spcm; Palmachim; 31.9737°N, 34.6767°E; depth 35.4 m; 2 May 2018;
soft substrate; box-corer; Shafdan project (sample 29(A)).

Remarks. ‘This species was first recorded in Israel in 1960 from Bat Yam with living
individuals (Barash and Danin 1973). Further living individuals were recorded in the
1960s and early 1970s from Israel and Egypt (Bardawil Lagoons) (Barash and Danin
1971, 1973, 1977). Since then, however, no further living individuals were found, ques-
tioning the persistence of its populations in the Mediterranean. We report here the recent
finding of two living individuals, suggesting that the species indeed still occurs in Israel.

Lioberus ligneus (Reeve, 1858)
Figure 33A—D

New records. Cyprus * 2 shs; Turkish Republic of Northern Cyprus, southern tip
of Karpaz Peninsula; 35.6809°N, 34.5785°E; depth 1 m; 29 Jul. 2019; mixed sub-

New non-indigenous species in the Mediterranean 59

Figure 32. Arcuatula perfragilis (Dunker, 1857), Palmachim, Israel, Shafdan project. Left valve outer view.
The pink hue is due to staining with eosin solution. Photograph courtesy S. Bartolini. Scale bar: 2 mm.

strate with sand, rocks and Cystoseira; P.G. Albano leg. (sample NCY7H); NHMW-
MO-112930/LM/0173; size of the largest shell: L 30.2 mm, H 15.0 mm.

IsraEL * 1 spcm; Carmel Head; 32.8232°N, 34.9431°E; depth 12 m; 21 Apr.
2020; in a small patch of Galaxaura rugosa and Cystoseira sp.; M. Mulas leg. (sample
SK2); SMNH MO83605; size: L 17.6 mm, H 10.0 mm.

Additional material examined. Lioberus ligneus: Ecypt * 19 wv; Red Sea,
Suez; Jousseaume coll.; MNHN « 1 sh; Sinai (Red Sea), Dahab, dive site “Islands”;
28.4789°N, 34.5126°E; beached; Nov. 2007; H. Blatterer leg. ¢ 1 v; same collecting
data as for preceding; 2012 © 1 v; Sinai (Red Sea), Dahab, dive site “Lagoon”; depth
unspecified; 2008; H. Blatterer leg. * 1 v; same collecting data as for preceding; 2009.

Lioberus agglutinans (Cantraine, 1835): Porrucat * 1 v; Algarve, Canal of Olhao;
depth 3-7m, MNHN ® 2 wy; Algarve, Tavira, Pedra do Barril; depth 25 m; MNHN ° 1
sh, 1 v; Algarve, Sagres, Ponta da Baleeira; depth 17-23 m; MNHN ® 1 spcm; Algarve,
Tavira, off Cabanas; depth 14 m; MNHN «® 2 shs; Algarve, Olhao, Ilha da Barretta;
depth unspecified; MNHN.

Tunisia ® 1 sh, 8 vv; Gulf of Gabés, NW Boughrara Gulf; depth 10-15 m; MNHN
¢ 11 spcms, 42 wv; Gulf of Gabés, Djerba, Ajim Canal; depth 10-32 m; MNHN.

Remarks. We here report Lioberus ligneus for the first time from Israel and Cy-
prus. The Israeli live-collected specimen comes from a patch of Galaxaura rugosa
and Cystoseira sp. whereas the two empty but fresh shells from Cyprus were found
attached vertically to Cystoseira shoots. These findings suggest that this species in-
deed prefers vegetated habitats in the shallow subtidal. This species has been previ-
ously reported from Lebanon based on shells collected in 1999-2000 (Crocetta et al.
2013), suggesting that it has likely occurred undetected throughout the Levantine
Basin for long. ‘The distinction of L. igneus from the native Mediterranean Lioberus
agelutinans (Cantraine, 1835) is not straightforward because both species share the
elongated appearance, sculpture limited to concentric striae, and brown color. More-
over, they are morphologically variable. A fairly consistent character of L. ligneus in
the samples we inspected from the Red Sea is the darker internal color, often shad-
ing into violet. The group would deserve a taxonomic revision to unambiguously
distinguish its species.

60 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 33. Comparison between Lioberus ligneus (Reeve, 1858) and Lioberus agglutinans (Cantraine,
1835). A, B Lioberus ligneus, NHMW-MO-112930/LM/0173, Karpaz Peninsula, Turkish Republic of
Northern Cyprus, Cyprus: right valve outer (A) and left valve inner (B) views C, D L. ligneus, Dahab, Egypt

(H. Blatterer coll.): right valve outer (C) and left valve inner (D) views E, F Lioberus agelutinans, Olhao,
Ilha da Barretta, Algarve, Portugal: right valve outer (E) and left valve inner (F) views G, H L. agelutinans,
Tavira, off Cabanas, Algarve, Portugal: right valve outer (G) and left valve inner (IH) views. Scale bars: 5 mm.

Musculus coenobitus (Vaillant, 1865)
Figures 34A—D, 35A—F

New records. IsRAEL * 2 spcms; west of Rosh HaNikra Islands; 33.0704°N, 35.0926°E;
depth 12 m; 1 May 2018; rocky substrate; suction sampler; HELM project (sample
S14_4M) ¢ 4 spcms; same collecting data as for preceding; 29 Oct. 2018; HELM pro-
ject (samples $52_1M, S52_2L, $52_3M, S52_3L) © 1 sh; north of Nahariyya port;

New non-indigenous species in the Mediterranean 61

33.0127°N, 35.0896°E; beached; 29 Oct. 2018; HELM project (sample H3); size: L
10.2 mm, H 6.0 mm (illustrated shell, Figure 34A—D).

Additional material examined. Musculus subpictus (Cantraine, 1835): CROATIA ®
14 spcms; Istria, off Rovinj, western shore of Sveta Katarina; 45.0760°N, 13.6276°E; 3
Jul. 2008; dried-out fouling community collected from the surface of buoy-like float-
ing objects dumped on the shore (J. Steger coll.).

IsRAEL * 10 spcms, 3 vv; Ashgelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr.
2018; offshore rocky reef; suction sampler; HELM project (samples $12_1F $12_1M,
S12_1L, $12_2F $12_2M) ¢ 111 spcms, 2 wv; Ashqelon; 31.6891°N, 34.5257°E;
depth 25 m; 2 May 2018; offshore rocky reef; suction sampler; HELM project (sam-
ples S16_1E S16_1M, S16_1L, S16_2F S16_2M, S16_2L) 13 spcms; west of Rosh
HaNikra Islands; 33.0704°N, 35.0926°E; depth 12 m; 1 May 2018; rocky substrate;
suction sampler; HELM project (samples $14_2F, $14_2M, S14_3F S14_4F) « 1 v;
west of Rosh HaNikra Islands; 33.0725°N, 35.0923°E; depth 20 m; 1 May 2018; rocky
substrate; suction sampler; HELM project (sample $13_3M) * 1 spcm; same collecting
data as for preceding; depth 19 m; 29 Oct. 2018; HELM project (sample $53_2L).

Matta * 9 spcms; off Filfla Island; on a rafting fisherman's net; 1981 (PG. Albano
coll. ID:557).

Remarks. We found several living individuals of Musculus coenobitus in northern
Israel. It is the first record of this Red Sea species for the Mediterranean Sea. Musculus
coenobitus is very similar to the native M. subpictus, which can occur sympatrically, but
can be distinguished at once because of its reddish rather than greenish hue (Figure 34).
It is also more elongated with a more pointed posterior margin. Juvenile specimens are
bright red (Figure 35C), have interspaces between the radial riblets in the posterior part
of the shell almost as large as the riblets themselves, while in MZ. subpictus the riblets are
closely arranged with very narrow interspaces (Figure 35E, J). Musculus coenobitus also
shows a fine but distinct lamellar concentric sculpture in such interspaces, whereas .
subpictus shows more spatse fine concentric ridges (Figure 35E and J, respectively). The
relation of M. coenobitus with other Indo-Pacific species like MZ. cumingianus (Reeve,
1857) and M. cuneatus (Gould, 1861) deserves a revision (Oliver 1992).

Musculus aff. viridulus (H. Adams, 1871)
Figure 36A—E

New records. IsraEL * 1 spcm; Ashqelon; 31.6891°N, 34.5257°E; depth 25 m; 2
May 2018; offshore rocky reef; suction sampler; HELM project (sample $16_2F) * 15
spcms; same collecting data as for preceding; depth 28 m; 31 Oct. 2018; HELM pro-
ject (samples S59_1E S59_1M, S59_2F S59_2M) ¢ 2 spcms; west of Rosh HaNikra
Islands; 33.0704°N, 35.0926°E; depth 12 m; 1 May 2018; rocky substrate; suction
sampler; HELM project (samples $14_3E S14_4M) * 25 spcms; same collecting data
as for preceding; 29 Oct. 2018; HELM project (samples $52_1F, $52_2F S52_2M,
$52_21, $522 3K7S52_3M,S52"31); size: 3:6 mm; A 2.1, ‘mm (illustrated: -speci-
men, Figure 36A—E) * 1 spcm; west of Rosh HaNikra Islands; 33.0725°N, 35.0923°E;

Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 34. Comparison between Musculus coenobitus (Vaillant, 1865) and Musculus subpictus (Cantraine,
1835) A=D Musculus coenobitus, north of Nahariyya port, Israel, HELM project (sample H3): right valve
outer (A) and inner (D) views, left valve inner (B) and outer (C) views E-H Musculus subpictus, off Filfla,

Malta: right valve outer (E) and inner (H) views, left valve inner (F) and outer (G) views. Scale bars: 5 mm.

depth 20 m; 1 May 2018; rocky substrate; suction sampler; HELM project (sample
S13_3M) * 9 spcms; same collecting data as for preceding; depth 19 m; 29 Oct. 2018;
HELM project (samples.S53_16'S53:2F S53_2M, 55338 553,33):

New non-indigenous species in the Mediterranean 63

Figure 35. Comparison between Musculus coenobitus (Vaillant, 1865) and Musculus subpictus (Can-
traine, 1835) A=F Musculus coenobitus, west of Rosh HaNikra Islands, Israel, HELM project (sample
S52_2L): right valve outer (A, D) and inner (B) views, living specimen with byssus (C), sculpture of the
posterior (E) and anterior (F) area of the right valve G-K Musculus subpictus, Ashqelon, Israel, HELM

project (sample $12_1L): right valve outer (G, I) and inner (H) views, sculpture of the posterior (J) and
anterior (K) area. Scale bars: 1 mm (A, B, D, G-I), 2.5 mm (C); 0.5 mm (E, F, J, K).

Additional material examined. Musculus aff. viridulus. Eyer ¢ 1 v; Sinai (Red Sea),
Dahab, dive site “Caves”; 28.416°N, 34.456°E; depth 15 m; 2015; H. Blatterer leg. * 4 wv;
same collecting data as for preceding; depth 17-19 m; 2016; H. Blatterer leg. * 2 vv; same
collecting data as for preceding; depth 18 m; 2017; H. Blatterer leg. * 1 v; Sinai (Red Sea),
Dahab, dive site “Golden Blocks”; 28.436°N, 34.459°E; depth 18 m; 2018; H. Blatterer leg.

64 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 36. Comparison between Musculus aff. viridulus (H. Adams, 1871) and Musculus costulatus (Risso,
1826) A-E Musculus aff. viridulus, west of Rosh HaNikra Islands, Israel, HELM project (sample $52_3L):
left valve outer (A, B) and inner (D) views, right valve inner (C) and outer (E) views FAH Musculus aff.

viridulus, dive site “Caves”, Dahab, Sinai, Egypt: left valve outer (F, G) and inner (H) views I-M Musculus
costulatus, west of Rosh HaNikra Islands, Israel, HELM project (sample $53_2L): left valve outer (I, J) and

inner (L) views, right valve inner (KK) and outer (M) views. Scale bars: 1 mm.

Musculus costulatus (Risso, 1826): IsraEL * 9 spcms, 4 vv; Ashgelon; 31.6868°N,
34.5516°E; depth 12 m; 30 Apr. 2018; offshore rocky reef; suction sampler; HELM
project (samples $12_1E S12_1M, S12_1L, $12_2F $12_2M) ¢ 3 spcms; Ashqelon;
31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef; suction sampler;
HELM project (samples $16_1F S16_2F) ¢ 1 spcm; same collecting data as for preceding;

New non-indigenous species in the Mediterranean 65

depth 28 m; 31 Oct. 2018; HELM project (sample S59_1M) * 5 spcms; west of Rosh
HaNikra Islands; 33.0704°N, 35.0926°E; depth 12 m; 1 May 2018; rocky substrate; suc-
tion sampler; HELM project (samples $14_1L, $14_3E S14_3L, $14_4F) ¢ 56 spcms;
same collecting data as for preceding; 29 Oct. 2018; HELM project (samples $52_1F
$520°1M,S5211, 852° 2E.S52)2M,S52.2L, $523 :S52.3M, S5203L)'* L-viwest:of
Rosh HaNikra Islands; 33.0725°N, 35.0923°E; depth 20 m; 1 May 2018; rocky substrate;
suction sampler; HELM project (sample S13_3F) * 20 spcms; same collecting data as for
preceding; depth 19 m; 29 Oct. 2018; HELM project (sample S$53_2L).

Remarks. We report numerous living individuals of Musculus aff. viridulus from the
Mediterranean Israeli coastline. It is the first record of this Indo-Pacific species in the
Mediterranean Sea. This species can be readily distinguished from the native M. costu-
latus because the latter has a more oval outline and a much smaller number of riblets at
the same overall shell size (Figure 36). These riblets are also much larger compared to M.
aff. viridulus. The Atlanto-Mediterranean M. discors (Linnaeus, 1767) has a similarly fine
posterior sculpture but at the same size is much higher and at all sizes bears much more
prominent riblets anteriorly. The taxonomy of Musculus in the Indo-Pacific province is
not settled and the available images of Red Sea M. viridulus (Oliver 1992; Zuschin and
Oliver 2003) show a more oval species, hence our dubitative identification. Still, we are
confident that this is a Red Sea species because we examined indistinguishable speci-
mens from the northern Red Sea (Figure 36F—H). Blatterer (2019) illustrated these and
other similar specimens (plate 10, fig. 18a, b) as Gregariella ehrenbergi (Issel, 1869). We
recorded a morphologically distinct species as G. ehrenbergi from a buoy stranded on the
Israeli coastline (Steger et al. 2018; Ivki¢ et al. 2019). Gregariella ehrenbergi type material
is corroded by Byne’s disease and the original description likely refers to a juvenile speci-
men; the identity of this species deserves further scrutiny.

Family Isognomonidae Woodring, 1925 (1828)

Isognomon aff. australica (Reeve, 1858) (sensu Angelidis and Polyzoulis 2018)
Figure 37

New records. Cyprus * 1 sh; Turkish Republic of Northern Cyprus, Esentepe;
35.3589°N, 33.6078°E; depth 2 m; 28 Jul. 2019; rocky substrate; PG. Albano leg.
(sample NCY3H); size: L 11.8 mm, H 10.1 mm (Figure 37A, B, D, E).

GREECE * | spcm; Crete, Plakias; 35.1796°N, 24.3957°E; depth 5 m; 24 Sep. 2017;
Posidonia oceanica rhizomes; suction sampler (sample Rh.05_4M); size: L 4.4 mm,
H 4.4 mm (Figure 37C, F).

Remarks. ‘The discrimination and identification of the species of lsognomon Light-
foot, 1786 is difficult due to their morphological plasticity that is related to their cryp-
tic way of life. Still, the specimens recently reported from Astypalaia, in the Eastern
Aegean Sea (Lipej et al. 2017; Angelidis and Polyzoulis 2018), show clear morphologi-
cal differences from J. legumen (Gmelin, 1791), the established non-indigenous species
in the Mediterranean Sea: the main features of the sculpture are radial rather than
concentric and the shape of the shell can be very elongated rather than subquadrate.

66 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 37. lsognomon aff. australica (Reeve, 1858) (sensu Angelidis and Polyzoulis 2018) A, B, D, E Esentepe,
Cyprus (sample NCY3H): left valve outer (A) and inner (B) views, right valve inner (D) and outer (E) views
C, F Plakias, Crete (sample Rh.05_4M): left (C) and right (F) valve outer views. Scale bars: 2 mm.

We here report a juvenile living individual and an empty shell from Greece and Cy-
prus (for which this is a new record), respectively, which are not distinguishable from
those previously reported from Astypalaia. Juvenile shells (up to a size of - 5 mm) bear
a sculpture of radial ribs adorned by tubular spines. We do not comment upon the
choice of the australica name for this taxonomic entity by previous authors, lamenting
the lack of a thorough revision of this genus in the Indo-Pacific province.

Family Lucinidae J. Fleming, 1828

Pegophysema cf. philippiana (Reeve, 1850)
Figure 38

New records. IsraeEL * 1 spcm; Palmachim; 31.9574°N, 34.6645°E; depth 36.2 m; 2
May 2018; soft substrate; box-corer; Shafdan project (sample 24(B)); size: L 15 mm,
Fal,5 “aim.

Remarks. Pegophysema philippiana was first found in the Mediterranean Sea in
2018 as a single valve from south of Tel Aviv (Mienis 2019). The specimen here report-
ed is much smaller, and we acknowledge that the identification of juvenile individuals
of this genus can only be tentative. We do not consider it conspecific to native species
such as Loripinus fragilis (Philippi, 1836) because this latter species is much more in-
flated at this size, nor Loripes orbiculatus Poli, 1795 which has a different valve profile.
If the identification is confirmed, this would be the first live collected specimen of P
philippiana in the Mediterranean Sea.

New non-indigenous species in the Mediterranean 67

Figure 38. Pegophysema cf. philippiana (Reeve, 1850), Palmachim, Israel, Shafdan project (sample
24(B)): right valve outer (AX) and dorsal (B) views. The pink hue is due to staining with eosin solution.
Photograph courtesy S. Bartolini. Scale bar: 5 mm.

Chavania erythraea (Issel, 1869)
Figure 39

New records. IsRAEL * 1 spcm; Palmachim; 31.9292°N, 34.6405°E; depth 36.9 m; 29
May 2004; soft substrate; grab; NM project (station 19); size: L 3.2 mm, H 3.2 mm.

Remarks. We report a juvenile but living individual of Chavania erythraea, a luci-
nid occurring in the Red Sea, the Persian (Arabian) Gulf and the Arabian Sea (Glover
and ‘Taylor 2001). This is the first record of this species from the Mediterranean Sea.
Adults develop a commarginal lamellar sculpture.

Rugalucina angela (Melvill, 1899)
New records. IsRAEL * 1 spcm, 1 v; Ashgelon; 31.6868°N, 34.5516°E; depth 11 m;

31 Oct. 2018; offshore rocky reef; suction sampler; HELM project (samples $58_2M,
S58_3M).

68 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 39. Chavania erythraea (Issel, 1869), Palmachim, Israel, NM project (station 19): right valve in-
ner (A, D) and left valve outer (B, E) views, hinge of right (C) and left (F) valve. The pink hue is due to
staining with eosin solution. Scale bars: 1 mm (A, B, D, E); 0.5 mm (C, F).

Remarks. This species was first recorded as Pillucina vietnamica Zorina, 1978
(now in Rugalucina too) from the Mediterranean coast of Israel by Steger et al. (2018),
who also illustrated it. A recent molecular phylogeny showed that R. vietnamica is dis-
tinct from R. angela and that the non-indigenous species in the Mediterranean belongs
to this latter species, which occurs in the Red Sea and northwest Indian Ocean (Taylor

and Glover 2019).

Family Galeommatidae Gray, 1840

Nudiscintilla cf. glabra Littzen & Nielsen, 2005 (sensu Mifsud and Ovalis 2012)
Figure 40

New records. IsraEL * 1 v; Nahariyya, 200 m north of the entrance to the marina;
33.0149°N, 35.0890°E; depth 3-4 m; 6 Nov. 2018; pools with bioclastic sand in rocky
bottom; snorkelled; J. Steger leg.; HELM project (sample D7); size: L 10.0 mm, H
6.4 mm (Figure 40A—E) ¢ 1 spcm; Palmachim; 31.9285°N, 34.6947°E; depth 3 m; 7
Nov. 2018; attached to the lower valve of a living Lessepsian Spondylus; scuba diving;
hand-picked; J. Steger & A. Ivkié leg.; HELM project (sample H17); size: L 4.2 mm,
H 2.8 mm (Figure 40F, G).

New non-indigenous species in the Mediterranean 69

Remarks. ‘This non-indigenous species has first been recorded in the Mediterranean
Sea by Mifsud and Ovalis (2012) as Nudiscintilla cf. glabra Liitzen and Nielsen, 2005,
based on five living specimens collected at Yumurtalik, Adana (Turkey) in shallow wa-
ter. Their tentative identification was primarily guided by the external morphology of
the living animals, which had a smooth mantle surface. This feature is characteristic for
the monotypic genus Nudiscintilla (hence the genus name), but unusual among scin-
tilloid galeommatids in general. Although no observations on living individuals could
be made by us, the shell morphology of our material well matches that of the specimen
illustrated in Mifsud and Ovalis (2012: fig. 1), suggesting conspecificity. Our findings
represent the first records of this species from Israel. However, the dentition of the
right valve as seen in SEM images (Figure 40C-—E) clearly differs from that described
by Liitzen and Nielsen (2005) for Nudiscintilla: the latter has a single cardinal tooth
in each valve and no lateral teeth. However, the studied right valve — the hinge of the
single live-collected specimen was not examined to avoid damage — bears what appears
to be two cardinal teeth (Figure 40C) that are fused at their base (Figure 40D, E), as
well as a ridge posterior to the internal ligament which most likely is a lateral tooth.
This ridge seems to correspond to the left of the two swellings indicated by a pair of ar-
rows on the right hand side of Mifsud and Ovalis (2012: fig. le), while the right swell-
ing might correspond to a narrow ridge visible also on the dorsal margin of our valve.
Mifsud and Ovalis (2012) interpreted these features as aberrant shell growth, however,
the presence of such ridges also in our right valve (Figure 40D) speaks against this hy-
pothesis. Furthermore, their living individuals had a small tentacle situated above the
widely gaping anterior inhalant region (cf. Mifsud and Ovalis (2012: 8, fig. 2a), how-
ever, the illustration of N. glabra in Liitzen and Nielsen (2005: 292, fig. 38a) shows a
small tentacle in the posterior exhalant region of the reflected mantle. In the light of
the poorly developed taxonomy and great species diversity of galeommatid bivalves
in the Indo-Pacific, further observations on living specimens, thorough comparisons
with the type material from Thailand and molecular analyses are required to definitely
clarify the relationship of Mediterranean specimens with NV. glabra.

Scintilla cf. violescens Kuroda & Taki, 1961
Figure 41

New records. IsRAEL * 1 spcm; Haifa; depth 15 m; May 1999; biogenic sediment; B.S.
Galil leg.; size: L6.9 mm, H 5.2 mm.

Remarks. ‘The single shell found is trapezoid-oval (L:H ratio = 1.33), slightly high-
er posteriorly than anteriorly, translucid-white, and has a glossy external surface. The
valves are narrowly gaping, more widely in their posterior part. The umbones are pro-
sogyrate, pointed and submedian. The commarginal sculpture consists of fine growth
lines that are slightly wavy posteriorly, as well as irregular growth marks. Flat radial ribs
are present in the posterior part of the shell; they are visible, upon close examination,
also on the inside of the valves in the form of shallow markings. The inner surface,

70 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 40. Nudiscintilla cf. glabra Liitzen & Nielsen, 2005 (sensu Mifsud and Ovalis 2012) A—E Naha-
riyya, Israel, HELM project (sample D7): outer (A) and inner (B) views, detail of hinge (C) and hinge
in dorsal view (D, E) of right valve F, G Palmachim, Israel, HELM project (sample H17): right (F) and
left (G) valve outer views. Scale bars: 3 mm (A, B); 0.5 mm (C, D); 0.2 mm (E); 1 mm (F, G).

particularly of the right valve, is spotted by blister-like markings (Figure 41C, D). The
hinge of the right valve bears a single cardinal tooth, bent towards the anterior, and an
elongated posterior lateral tooth. The left valve has two cardinals, but the anterior one
is broken off (Figure 41G); a posterior lateral is present.

Lacking further material and observations on living individuals, which are of great
diagnostic importance in galeommatids, we refrained from assigning a definitive spe-
cific name to our shell. However, the overall shape, hinge dentition and the presence
of radial sculpture match well descriptions of Scintilla violescens Kuroda & Taki, 1961
(Arakawa 1961; Kuroda and Taki 1961), a species recorded from the intertidal and

New non-indigenous species in the Mediterranean jaa

Figure 41. Scintilla cf. violescens Kuroda & Taki, 1961, Haifa, Israel: right (A) and left (B) valve outer
views, left (C) and right (D) valve inner views, hinge of left (E, G) and right (F, H) valve. Scale bars:
2 mm (A=D); 0.5 mm (E=H).

shallow subtidal of Thailand and Japan (Huber 2015). In contrast to our shell, howev-
er, Kuroda and Taki (1961) mention the presence of radial sculpture on the entire sur-
face of the valves of their type material. While our Mediterranean shell is less elongated
than the specimens of S. violescens illustrated by Okutani (2000), Liitzen and Nielsen
(2005), and Huber (2015), it is very similar in outline to the shell shown by Arakawa
(1961: fig. 5B) (which was identified by T. Kuroda and I. Taki). Scintilla violescens
appears to be variable also with respect to shell size and coloration: specimens from
Thailand (maximum length = 10.5 mm, n = 12 spcms) all were considerably smaller

than the > 15 mm-long Japanese type (Kuroda and Taki 1961) and had a whitish in-

fig Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

stead of pale violet color (Liitzen and Nielsen 2005), like the Israeli shell. Considering
this great plasticity in shell characters, and the differences in living animal morphology
observed for Thai vs. Japanese specimens of S. violescens by Liitzen and Nielsen (2005),
the question arises whether more than one biological entity might be involved.

Irrespective of its unresolved specific affinity, the shell presented here clearly differs
from all native Mediterranean galeommatids and thus cannot be confused; it can be
easily distinguished from the non-indigneous Nudiscintilla cf. glabra (see above) by its
less elongated shell, smaller L:H-ratio and, most notably, the presence of radial sculp-
ture on the valves. Apart from the present shell, which was found in 1999 in Haifa Bay,
we know of no other material.

Family Psammobiidae J. Fleming, 1828
Gari pallida (Deshayes, 1855)

New records. IsRAEL * 6 spcms; Ashqelon; 31.7002° N, 34.5498° E; depth 21 m; 18
Sep. 2016; sand; grab; HELM project (samples SG20_1E SG20_3K SG20_3M).

Remarks. A single living individual of this species has been recently reported from
Palmachim, along the southern Mediterranean coast of Israel, representing the first record
from the Mediterranean Sea (Lubinevsky et al. 2018). We here confirm the establishment
of this species by reporting six further living individuals, again from southern Israel.

Family Semelidae Stoliczka, 1870 (1825)

Ervilia scaliola Issel, 1869
Figure 42

New records. IsraxL * 2 vv; Ashqelon; 31.7002° N, 34.5498° E; depth 21 m; 18 Sep.
2016; sand; grab; HELM project (sample SG20_2F) * 1 sh; Ashqelon; 31.6868°N,
34.5516°E; depth 11 m; 31 Oct. 2018; offshore rocky reef; suction sampler; HELM
project (sample $58_3M); NHMW-MO-112930/LM/0176; size: L 3.1 mm, H 2.2
mm (illustrated shell).

Remarks. FErvilia scaliola was first recorded from Turkey based on material col-
lected in 2013 by Zenetos and Ovalis (2014) who correctly described the complex
taxonomic status of this genus in the Indo-Pacific. We here record the species from
Israel for the first time. The complete shell from Ashqelon (Figure 42) is very fresh, and
thus probably originates from an extant population.

Tacra seychellarum (A. Adams, 1856)
Figure 43

New records. GREECE * 3 wy; Kos Island; 2005-2010; A. Storm leg.; RMNH.
MOL.342632; size: L 12.1 mm, H 9.8 mm.

New non-indigenous species in the Mediterranean ie:

Figure 42. Evrvilia scaliola Issel, 1869, NHMW-MO-112930/LM/0176, Ashgelon, Israel, HELM pro-
ject (sample S58_3M): right (A, C) and left (B, D) valve outer views, left (E, G) and right (F, H) valve

inner views. Scale bar: 1 mm.

74 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Figure 43. Jacra seychellarum (A. Adams, 1856), Kos Island, Greece, RMNH.MOL.342632: left valve outer (A)
and inner (B) views, hinge (C). Photograph courtesy T: van Haaren. Scale bar: 2 mm (A, B); 1 mm (C).

Remarks. We here report the finding of three beached valves of Jacra seychellarum
in Kos, Greece. lacra seychellarum can be readily distinguished from any Mediterranean
semelid by its thick valves, large chondrophore and different sculpture in three zones:
anterior with fine incised concentric lirations, median to posterior slope of fine incised
oblique lines becoming strongly divaricate over the posterior slope, posterior area with
closely spaced concentric incised lirations (Oliver 1992; Zuschin and Oliver 2003). It
has been recorded from the northern Red Sea by Blatterer (2019) but has a broader
distribution in the Indian Ocean (Oliver 1992). Because to our knowledge the find-
ing has not been followed by others, we suggest that living individuals are needed to
confirm the introduction and establishment of this species in the Mediterranean Sea.

Semelidae sp.

Figure 44

New records. IsRAEL * 1 spcm; Palmachim; 31.9737°N, 34.6767°E; depth 35.4 m; 24
May 2017; soft substrate; box-corer; Shafdan project (sample 29(A)); size: L 2.4 mm,
H 2.0 mm.

Additional material examined. Abra alba (W. Wood, 1802): IsRaet * 1 v; north
of Atlit; 32.7422°N, 34.9181°E; depth 30 m; 20 Sep. 2016; sand; grab; HELM project
(sample NG30_2M) ¢ 2 vv; Ashqelon; 31.7101°N, 34.5406°E; depth 31 m; 18 Sep.
2016; sand; grab; HELM project (sample SG30_5M).

Remarks. We were unable to identify this peculiar bivalve beyond family level, de-
spite a thorough search in the literature on Mediterranean and Indo-Pacific mollusks.
The shell of the single specimen found is roundly subtrigonal, fragile, with a rounded,
steeply sloping anterior and a subacute to subtruncate posterior part; the ventral mar-
gin is slightly concave, the umbo submedian. The outer surface is smooth, glossy, and
only sculptured by very fine growth lines. Although the small size and outline are
reminiscent of certain galeommatoid genera such as Bornia Philippi, 1836, which is
also represented in the Mediterranean Sea, the presence of both an external and an
internal ligament, the latter situated on a well-developed resilifer, is typical for the fam-
ily Semelidae (Oliver 1992; Beesley and Ross 1998; Huber 2010). The hinge of both

valves bears two cardinals; the posterior cardinal of the left valve is becoming obsolete

New non-indigenous species in the Mediterranean Fis)

Figure 44. Semelidae sp., Palmachim, Israel, Shafdan project (sample 29(A)): left valve outer view (A, B),
right valve inner view (D, E), hinge (C, F) and outer view (G, H), left valve inner view (J-K) and hinge (I, L).
The pink hue is due to staining with eosin solution. Scale bars: 1 mm (A, B, D, E, G, H, J, K); 0.5 mm (FL);
0.3 mm (C, I).

by encroachment of the internal ligament portion which also extends vertically beyond
the hinge line. Two well-developed laterals are present in the right valve, while in the
left valve, only a weak tooth-like ridge is present anteriorly, formed by the dorsal shell
margin. Due to the extremely smooth and glossy interior of the valves that renders
hardly visible even the adductor muscle scars, it remains unclear whether a deep pallial
sinus, another feature typical of semelids, is present in the studied specimen.

Lonoa katoi Habe, 1976, a semelid from Japan, shares with our species the small
size and irregular outline with an often concave ventral margin (related to its attached

76 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

lifestyle); however, the outer shell surface of L. katoi bears rough lamellae and fine
radial threads (Habe 1976; Okutani 2017), while that of the Israeli specimen is al-
most smooth. The most similar confamilial species from the Red Sea probably is Abra
aegyptiaca Oliver and Zuschin, 2000, however, it differs from the specimen described
here in shell shape, sculpture, the prosogyrate umbo, and features of the hinge such
as the shape of the anterior lateral tooth of the right valve. Similar-sized juveniles of
Mediterranean Abva spp., including A. a/ba, the most common species on the shallow
Israeli shelf, have a hinge morphology comparable to our specimen, but differ in their
outline and by having more protruding umbos (Scaperrotta et al. 2013). Abra tenuis is
most similar in shape, and a teratological specimen might approach the outline of our
shell; however, such a specimen would still differ from Semelidae sp. by the presence of
commarginal lines on the early dissoconch (Scaperrotta et al. 2013; Oliver et al. 2020).

Until the finding of further specimens, it remains open whether the present indi-
vidual is a juvenile or an adult of a small-sized species. Considering that only a single
specimen was found so far, the lack of known native Mediterranean species with a
similar morphology, and the geographical proximity of the Israeli coast to the Suez
Canal, we suspect that this species might be another Indo-Pacific taxon introduced to
the southeastern Mediterranean.

Family Veneridae Rafinesque, 1815

Clementia papyracea (Gmelin, 1791)

New records. IsraEL * 1 spcm; Haifa Bay; 32.8211°N, 35.0196°E; depth 11 m; 2 Aug.
2015; soft substrate; grab; NM project (station HM27).

Remarks. Clementia papyracea has been recorded in the Mediterranean Sea since
1937, but findings of living individuals are very scarce and limited to samples collected
in 1968 in El Arish, Egypt (Barash and Danin 1973), in 1975 in Haifa, Israel (Barash
and Danin 1977) and in 2012 in Ashgelon, Israel (Crocetta et al. 2016). We here re-
port a further living juvenile individual.

Family Corbulidae Lamarck, 1818

Corbula erythraeensis H. Adams, 1871
Figure 45A—K

New records. IsraxEL * 1 spcm; Ashgelon; 31.7002°N, 34.5498°E; depth 21 m; 18 Sep.
2016; sand; grab; HELM project (sample SG20_4F) ¢ 3 spcms; Ashgelon; 31.7487°N,
34.4960°E; depth 41 m; 27 Apr. 2017; sandy mud; grab; HELM project (sample
SG40_8M); sizes: L 2.9 mm, H 2.5 mm (illustrated specimen 1, Figure 45A—C); L 3.8
mm, H 3.1 mm (illustrated specimen 2, Figure 45D, E) * 2 spcms; Soreq desalination
plant; 31.94°N, 34.69°E; depth 17-22 m; 10 May 2009; soft substrate; grab; Soreq
project; size: L 1.9 mm, H 1.5 mm (Figure 45J, K).

New non-indigenous species in the Mediterranean 77

Figure 45. Comparison between Corbula erythraeensis H. Adams, 1871 and Corbula gibba (Olivi, 1792).
A-E Corbula erythraeensis, Ashgelon, Israel, HELM project (sample SG40_8M): A=-C Specimen 1: left
(A) and right (B) valve outer views, sculpture of the anterior left valve (C) D, E Specimen 2: left (D) and
right (E) valve outer views. F—H Corbula erythraeensis, Bay of Safaga, Egypt, NHMW-MO-106107: left
valve outer (F), ventral (G) and dorsal (H) views. | Corbula erythraeensis, same collecting data as for preced-
ing, NHMW-MO-106100: left valve outer view J, K Corbula erythraeensis, Soreq desalination plant, Israel,
Soreq project: left (J) and right (KK) valve outer views L,M Corbula gibba, same collecting data as for A-E:
left (L) and right (M) valve outer views N, O Corbula gibba, same collecting data as for J—K: left (N) and
right (O) valve outer views. Scale bars: 1 mm (A, B, D, E, F-I, L, M); 0.5 mm (C, J, K, N, O).

78 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

Additional material examined. Corbula erythraeensis: Ecypt ° 1 sh; Bay of Safaga;
26.8142°N, 33.9653°E; depth 39 m; 31 Oct. 1994; mud; scuba diving; M. Zuschin
leg.; ref. Oliver and Zuschin (2000), sample 94/4/b; NHMW-MO-106107; size: L
3.5 mm, H 2.5 mm (illustrated shell, Figure 45F—H) © 1 v; same collecting data as
for preceding; NHMW-MO-106100; size: L: 2.3 mm, H: 1.6 mm (illustrated valve,
Figure 45]).

Corbula gibba (Olivi, 1792): IsraEL * 271 spcms; Ashgelon; 31.7492°N, 34.4964°E;
depth 41 m; 27 Apr. 2017; sandy mud; grab; HELM project (sample SG40_8M)
300 spcms, 11 shs, 10 vv (5 right, 5 left); Soreq desalination plant; 31.94°N, 34.69°E;
depth 17—22 m; 10 May 2009; soft substrate; grab; Soreq project.

Remarks. Corbula erythraeensis is widespread in the northern Red Sea where it
has been recorded from the gulfs of Suez (MacAndrew 1870; Oliver 1992) and Aqaba
at Eilat (Edelman-Furstenberg and Faershtein 2010), as well as the northern Bay of
Safaga (Egypt) (Zuschin and Oliver 2003); outside the Red Sea, its distribution ranges
eastward to Pakistan (Huber 2010). Here we report the first findings of this species
from the Mediterranean. Corbula erythraeensis was found sympatrically with the com-
mon native Mediterranean Corbula gibba but was always present in very low numbers.
No empty shells have been found so far. While being similar in appearance to the
morphologically variable C. gibba, C. erythraeensis has a convex anterior dorsal margin
of the right valve (usually concave in C. gibba, particularly in smaller specimens), a
more inflated umbonal region, a regular concentric sculpture on the left valve (Oliver
1992), and its color always is whitish-yellowish (C. gibba frequently has a rosy pattern).
Juvenile individuals are more wedge-shaped than those of C. gibba.

Discussion

Massive reporting follows intensive fieldwork and broad cooperation

We have covered 52 species, reporting the finding of 23 new Lessepsian mollusks,
nine additional species that, upon final identification, may turn out to be further new
Lessepsian species, nine new records for Eastern Mediterranean countries and new
data for eleven already recognized non-indigenous species. Such a massive report is
derived from three characteristics of this study which translate into recommendations
for an effective approach to non-indigenous species detection and monitoring. First,
the intensive sampling effort and the effective sampling techniques of the “Historical
ecology of Lessepsian migration” (HELM) project and of the IOLR monitoring pro-
grams. The HELM project in particular targeted also hard substrates, poorly explored
at this taxonomic resolution in the Eastern Mediterranean, by suction sampling. ‘This
technique has repeatedly proved to be a very effective method on compact (e.g., coral
rubble, pebbles (Bouchet et al. 2002; Linnane et al. 2003; Ringvold et al. 2015; Evans
et al. 2018)), seagrass (Bonfitto et al. 1998; Albano and Sabelli 2012; Albano and
Stockinger 2019) or hard substrates (Templado et al. 2010) and has enabled here the

New non-indigenous species in the Mediterranean 79

collection of vast amounts of living micromollusks and their shells. Indeed, 26 out of
the 52 (50%) species treated here and 17 out of 32 (53%) new and potentially new
NIS came from suction samples on hard substrates. The IOLR surveys covered soft
substrates along the whole Israeli Mediterranean coastline with a dense station network
and multi-seasonal sampling that led to the detection of 16 out of 52 (31%) treated
species, notwithstanding several findings had already been published (e.g., Bogi and
Galil 2013b, Aartsen et al. 2015; Lubinevsky et al. 2018). Second, the use of fine mesh
sizes. The HELM samples were sieved with a 0.5 mm mesh and those of the IOLR
monitoring programs were sieved with either a 0.5 mm or even a 250 um mesh. Such
small sizes enabled retaining the large majority of invertebrates, including mollusks,
even those with very small and elongated shells, such as many Pyramidellidae. How-
ever, this approach requires an enormous effort when picking and sorting the samples
and the availability of high-level taxonomic expertise, since most small-sized species
belong to taxonomically challenging groups or represent juvenile individuals. Third,
and importantly, cooperation among institutions and individuals. Despite new records
of non-indigenous species are often scattered into short papers in the literature and
may become difficult to trace in the long term, recent efforts have demonstrated the
value of cooperation to build up large datasets (e.g., Katsanevakis et al. 2020). It is also
important to highlight the role that citizen scientists had in the detection of the new
non-indigenous species treated here by contributing to sample sorting, species identi-
fication and their taxonomic study.

The challenge of recognizing and identifying tropical non-indigenous species

Taxonomic uncertainty is recognized as a major impediment to the reliable inventory-
ing of non-indigenous species (McGeoch et al. 2012; Marchini et al. 2015; Katsane-
vakis and Moustakas 2018). Species whose identification is uncertain or whose taxo-
nomic status is unresolved were suggested to be excluded from inventories (Marchini et
al. 2015). Taxonomic uncertainty may also imply an uncertain non-indigenous status:
a species morphologically distinct from the native species pool can either be a yet
undescribed native species or a newly introduced species (which may be undescribed
too). If the new species belongs to a clade not occurring in the sampled range, then
the attribution of the non-indigenous status is well supported. Still, only the finding of
clearly conspecific individuals from the source pool would provide final evidence of the
non-indigenous status. We exemplified these cases with two new taxa described here:
Coriophora lessepsiana Albano, Bakker & Sabelli, sp. nov. clearly belongs to a tropical
clade of Triphoridae and the availability of material from the Red Sea enabled the
unambiguous attribution of the species to the Red Sea pool (hence the species name
lessepsiana). Joculator problematicus Albano & Steger, sp. nov. belongs to a genus absent
from the Mediterranean Sea. Despite multiple similar species have been recorded in
the Indo-Pacific province, we have not been able to find conspecific Red Sea material
(hence the species name problematicus, to highlight the uncertainty in attributing the
non-indigenous status when solid taxonomic and faunistic knowledge is lacking).

80 Paolo G. Albano et al. / ZooKeys 1010: 1-95 (2021)

A more complex case is represented by species that have few diagnostic characters,
hampering the unequivocal attribution to a tropical clade (e.g., within Eulimidae and
Pyramidellidae). We here propose to exploit the properties of death assemblages to
deliver a solid hypothesis of non-indigenous status. Death assemblages, the taxonomi-
cally identifiable, dead or discarded organic remains encountered in a landscape or
seabed such as molluscan shells, accumulate species richness over time (Kidwell 2013).
This property implies that death assemblages are good archives of local species diversity
and that species which have long occurred in a study site (like native ones) are likely
to be found in a death assemblage even if they are too rare to be regularly detected in
living censuses. Indeed, the inclusion of empty shells in field surveys increased the es-
timations of occupancy and detectability of native land snails (Albano et al. 2015). In
contrast, a species which has recently established a population in a new area would be
expected to be poorly detectable in the death assemblage because not enough time has
elapsed to contribute a significant number of skeletal parts to it. Consequently, newly
reported non-indigenous species found alive but not, or very rarely, in the death as-
semblage may represent newly introduced species. This approach can be applied when
living and death assemblages are sampled simultaneously with quantitative methods.
The main limitation of this approach is that it is applicable only to organisms with hard
skeletal parts like foraminiferans, ostracods and fishes (e.g. Agiadi and Albano 2020).

With biological invasions constituting a major element of global change, there is
understandable concern on the accuracy of non-indigenous species inventories (Mar-
chini et al. 2015; Katsanevakis and Moustakas 2018). However, also the risk of under-
estimating the magnitude of biological invasions must be considered. In this respect,
the Lessepsian invasion is a special case. The salinity reduction of the Bitter Lakes (Gal-
il 2006), the containment by dams of the annual Nile flood which used to modulate
salinity off its delta (Rilov and Galil 2009), and the multiple enlargements of the Suez
Canal (Galil et al. 2015) have certainly enhanced connectivity between the Mediter-
ranean Sea and the Indo-Pacific province during the 20" century. Additionally, in the
most recent decades, rising seawater temperatures in the Mediterranean Sea (Ozer et
al. 2017) make it increasingly suitable for the establishment of inherently thermophilic
Red Sea species. These factors combined render very likely that an increasing number
of Red Sea species manages to cross the Canal and to settle in the Mediterranean Sea.
The proper understanding of this phenomenon and of its drivers requires a timely and
detailed census of assemblages as well as a broad toolkit to overcome the limitations of
the taxonomic challenges associated with tropical species.

Authors’ contributions

PGA and JS conceived the study. PGA, JS, CB, MB, TGH, MFH, HL, MM, and MS
contributed to fieldwork, sample sorting and data acquisition. PGA, JS, PAJB, CB,
PILE and BS identified the specimens and contributed to the taxonomic discussion.

PGA, JS, and TGH contributed to the discussion. PGA, JS, PAJB, and MA prepared

New non-indigenous species in the Mediterranean 81

the figures. PGA and JS wrote the first draft of the manuscript, which then received
contributions by all co-authors.

Funding

This research has been conducted in the context of the “Historical ecology of Lessep-
sian migration” project funded by the Austrian Science Fund (FWF) P28983-B29 (PI:
PG. Albano). Sampling in Crete was supported by the “Kurzfristige wissenschaftli-
che Auslandsstipendien” program of the University of Vienna to M. Stockinger, the
non-profit organization Mare Mundi, and the diving school Dive2gether. Sampling in
northeastern Cyprus was conducted in the framework of the project “Classification of
coastal habitats of Northern Cyprus according to EUNIS protocol” (PI: EF Huseyno-
glu). The Faculty of Earth Sciences of the University of Vienna funded a citizen science
project conducted in October 2019. MB was supported by an Ernst Mach fellowship
of the OeAD (Osterreichischer Austauschdienst).

Acknowledgements

We thank Jonathan Belmaker, Bella S. Galil, Shahar Malamud, and Martin Zuschin for
their support at various stages of this study. Alberto Cecalupo, Italo Nofroni, P. Graham
Oliver, John Taylor, and Manuel Tenorio gave useful suggestions for the identification
of some species. Bruno Amati, Stefano Bartolini, Beata Dunne, Verena Gareis, Alex-
ander Heidenbauer, Anna Hinterplattner, Nadja Loferer, Denny Morchner, and Maria
Scaperrotta helped sorting the samples. Hubert Blatterer, Henk Dekker, Serge Gofas,
Virginie Héros, Anna Holmes, Ben Rowson, and Amy Storm put at our disposal mate-
rial for comparison. Franco Agamennone and Pasquale Micali shared information about
some species. Stefano Bartolini, Antonio Bonfitto, Ton van Haaren, Philippe Maestrati,
Norine Young, and Maria Tavano allowed us to use photographs they took. Emilia Ja-
rochowska and Christian Schulbert provided the wCT-scan and 3D-reconstruction of
Odostomia (s.1.) sp. 1. Henk Dekker commented on the manuscript. Sampling in Israel
was conducted under permit 41928 by the Israel Nature and Parks Authority.

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