Research Article |
Corresponding author: Paolo G. Albano ( pgalbano@gmail.com ) Academic editor: Antonio M. de Frias Martins
© 2021 Paolo G. Albano, Jan Steger, Piet A. J. Bakker, Cesare Bogi, Marija Bošnjak, Tamar Guy-Haim, Mehmet Fatih Huseyinoglu, Patrick I. LaFollette, Hadas Lubinevsky, Martina Mulas, Martina Stockinger, Michele Azzarone, Bruno Sabelli.
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.
Citation:
Albano PG, Steger J, Bakker PAJ, Bogi C, Bošnjak M, Guy-Haim T, Huseyinoglu MF, 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
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New data on 52 non-indigenous mollusks in the Eastern Mediterranean Sea is reported. Fossarus sp. (aff. aptus sensu
Cerithiopsidae, invasion biology, Lessepsian invasion, Mollusca, new species, Red Sea, taxonomy, Triphoridae
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 populations (
The introduction rate is an important metric to describe the invasion process. Genuine variation in this rate can result from changes in vector efficacy, connectivity between the native and introduced range, and environmental conditions in the recipient ecosystem. The introduction rate is often estimated from the discovery record (
To monitor a dynamic process such as the Lessepsian invasion, intensive fieldwork is mandatory. We indeed show that an intensive sampling effort coupled with identification 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 records for Eastern Mediterranean countries, and new data for eleven already recognized non-indigenous species. We here describe these new findings, providing detailed collecting data, taxonomic comments, and comparisons with similar species.
The studied material comes from three main sources. First, sampling on the Israeli Mediterranean shelf performed in the context of the project “Historical ecology of Lessepsian migration” (HELM), in progress at the University of Vienna. Second, benthic assemblage 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 environmental assessment (APM DAN, Shafdan, Via Maris). The NM, APM DAN and Via Maris projects sampled soft substrates with a 0.11 m2 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). Samples were sieved with a 250 μm mesh. During the Shafdan project, three replicate sediment samples were taken at each station from a different 0.062 m² 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
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 (
Acknowledging that an unsettled taxonomic status implies uncertainty in the assignment of non-indigenous status (
Small specimens were photographed with a Zeiss SteREO Discovery.V20 stereomicroscope, 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.l.) 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 software. The X-ray image stack, mesh files, virtual sections, and a video showing the interior 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 specimen (usually the one figured, unless otherwise stated). The systematic arrangement follows
List of the taxa treated in this paper, with indication of the novelty of the records.
Family | Taxon | Novelty | Page | Figure |
---|---|---|---|---|
– | Unidentified gastropod | Potential new NIS for the Mediterranean Sea | 6 | Figure |
Conradiidae | Conradia eutornisca | First record of living individuals in the Mediterranean Sea | 7 | – |
Skeneidae | Dikoleps micalii | Declared NIS in the Mediterranean Sea. First record from Israel; first record of living individuals in the Mediterranean Sea | 8 | Figure |
Skeneidae indet. | Potential new NIS for the Mediterranean Sea | 9 | Figure |
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Planaxidae | Fossarus sp. (aff. aptus sensu Blatterer, 2019) | New NIS for the Mediterranean Sea | 10 | Figure |
Epitoniidae | Cycloscala hyalina | First record of living individuals in the Mediterranean Sea | 11 | – |
Naticidae | Eunaticina papilla | First record of living individuals in Israel | 12 | Figure |
Triphoridae | Coriophora lessepsiana Albano, Bakker & Sabelli, sp. nov. | New NIS for the Mediterranean Sea | 12 | Figure |
Opimaphora blattereri Albano, Bakker & Sabelli, sp. nov. | New species from the Red Sea, for comparison with the non-indigenous Coriophora lessepsiana Albano, Bakker & Sabelli, sp. nov. | 15 | Figure |
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Triphora sp. | Potential new NIS for the Mediterranean Sea | 18 | Figure |
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Viriola cf. bayani | New reports of living individuals from Israel | 19 | – | |
Cerithiopsidae | Cerithiopsis sp. aff. pulvis | New NIS for the Mediterranean Sea | 20 | Figure |
Cerithiopsis sp. | New NIS for the Mediterranean Sea | 20 | Figure |
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Joculator problematicus Albano & Steger, sp. nov. | New NIS for the Mediterranean Sea | 23 | Figure |
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Elachisinidae | Elachisina sp. | New NIS for the Mediterranean Sea | 26 | Figure |
Iravadiidae | Iravadia aff. elongata | New NIS for the Mediterranean Sea | 26 | Figure |
Vitrinellidae |
Vitrinella aff. Vitrinella sp. 1 (sensu |
New NIS for the Mediterranean Sea | 29 | Figure |
Eulimidae | Hypermastus sp. | Potential new NIS for the Mediterranean Sea | 31 | Figure |
Hemiliostraca clandestina comb. nov. | Declared NIS in the Mediterranean Sea. First record from Israel; first record of living individuals in the Mediterranean Sea | 31 | Figure |
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Melanella orientalis | New NIS for the Mediterranean. | 33 | Figure |
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Parvioris aff. dilecta | New NIS for the Mediterranean Sea | 35 | Figure |
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Sticteulima sp. | Potential new NIS for the Mediterranean Sea | 37 | Figure |
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Vitreolina cf. philippi | Potential new NIS for the Mediterranean Sea | 37 | Figure |
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Conidae | Conus fumigatus | First record from Israel | 38 | Figure |
Murchisonellidae | Henrya (?) sp. | Potental new NIS for the Mediterranean Sea | 39 | Figure |
Pyramidellidae | Odostomia cf. dalli | New NIS for the Mediterranean Sea | 41 | Figure |
Odostomia (s.l.) sp. 1 | Potential new NIS for the Mediterranean Sea | 41 | Figure |
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Odostomia (s.l.) sp. 2 | First record of a living individual in the Mediterranean Sea | 44 | Figure |
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Oscilla virginiae | New NIS for the Mediterranean Sea | 45 | Figure |
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Parthenina cossmanni | New NIS for the Mediterranean Sea | 47 | Figure |
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Parthenina typica | New NIS for the Mediterranean Sea | 49 | Figure |
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Pyrgulina craticulata | New NIS for the Mediterranean Sea | 49 | Figure |
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Pyrgulina nana | First record of living individuals in Israel | 53 | – | |
Turbonilla funiculata | New NIS for the Mediterranean Sea | 53 | Figure |
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Cylichnidae | Cylichna collyra | New NIS for the Mediterranean Sea | 55 | Figure |
Mnestiidae | Mnestia girardi | First record of living individuals in the Mediterranean Sea | 57 | – |
Haminoeidae | Atys angustatus | First record from Greece | 57 | Figure |
Mytilidae | Arcuatula perfragilis | Additional records of living individuals from Israel | 58 | Figure |
Lioberus ligneus | First record from Cyprus and Israel | 58 | Figure |
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Musculus coenobitus | New NIS for the Mediterranean Sea | 60 | Figure |
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Musculus aff. viridulus | New NIS for the Mediterranean Sea | 61 | Figure |
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Isognomonidae |
Isognomon aff. australica (sensu |
New record from Cyprus | 65 | Figure |
Lucinidae | Pegophysema cf. philippiana | First record of living individuals in the Mediterranean Sea | 66 | Figure |
Chavania erythraea | New NIS for the Mediterranean Sea | 67 | Figure |
|
Rugalucina angela | Additional records of living individuals from Israel | 67 | – | |
Galeommatidae |
Nudiscintilla cf. glabra (sensu |
First record from Israel | 68 | Figure |
Galeommatidae | Scintilla cf. violescens | New NIS for the Mediterranean Sea | 69 | Figure |
Psammobiidae | Gari pallida | Additional records of living individuals from Israel | 72 | – |
Semelidae | Ervilia scaliola | First record from Israel | 72 | Figure |
Iacra seychellarum | New NIS for the Mediterranean Sea | 72 | Figure |
|
Semelidae sp. | Potential new NIS for the Mediterranean Sea | 74 | Figure |
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Veneridae | Clementia papyracea | Additional records of living individuals from Israel | 76 | – |
Corbulidae | Corbula erythraeensis | New NIS for the Mediterranean Sea | 76 | Figure |
H height;
HELM “Historical ecology of Lessepsian migration” project;
IOLR Israel Oceanographic and Limnological Research;
L length;
MZUB Museum of Zoology of the University of Bologna, Italy;
NM National Monitoring Israel;
SMNH Steinhardt Museum of Natural History, Tel Aviv, Israel;
sh/shs empty shell/s;
spcm/spcms live collected specimen/s;
v/vv valve/s;
W width.
Unidentified gastropod
Figure
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 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 transitional 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.
Israel • 1 sh; Ashqelon; 31.7101°N, 34.5406°E; depth 31 m; 18 Sep. 2016; muddy-sand; grab; HELM project (sample SG30_5F) • 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_2F) • 3 spcms; Ashqelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef; suction 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 S14_1F, S14_3F) • 3 spcms; same collecting data as for preceding; 29 Oct. 2018; HELM project (samples S52_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 S13_1F, S13_1M, S13_2F, S13_3F, S13_3M) • 14 spcms; same collecting data as for preceding; depth 19 m; 29 Oct. 2018; HELM project (samples S53_1F, S53_1M, S53_2F, S53_2M, S53_3F, S53_3M); size: H 2 mm, W 1.7 mm.
The species has already been reported from Israel and Turkey (
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.
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 (
Dikoleps micalii Agamennone, Sbrana, Nardi, Siragusa & Germanà, 2020, Haifa Bay, Israel: 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).
Israel • 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.
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 teleoconch whorls with numerous regular spiral cords. The shoulder is slightly angulated near the lip. Umbilicus open, large. Shell white, slightly translucent. No native Mediterranean 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 prominent spiral cords. It is most likely a new, probably still unnamed, Indo-Pacific species in the Mediterranean.
Israel • 1 sh; Ashqelon; 31.6868°N, 34.5516°E; depth 11 m; 31 Oct. 2018; offshore rocky reef; suction sampler; HELM project (sample S58_3M); size: H 3.3 mm, W 2.7 mm.
We found a single empty shell of this Fossarus that can be readily distinguished 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
The name aptus is problematic. Originally introduced by
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.
The species has been recently recorded for the Mediterranean Israeli coastline based on empty shells collected off the Soreq desalination plant, ~ 15 km south of Tel Aviv (
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).
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
Holotype. Egypt • sh; Sinai (Red Sea), Dahab, dive site “Blue Hole”; 28.572°N, 34.538°E; depth 4 m; 2017; H. Blatterer leg.;
Paratypes. Egypt • sh; Sinai (Red Sea), Dahab, dive site “Tigerhouse”; 28.567°N, 34.533°E; depth 7 m; 2015; H. Blatterer leg.;
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).
Egypt • 1 sh; Sinai (Red Sea), Dahab, dive site “Caves”; 28.416°N, 34.456°E; depth unspecified; 2012; H. Blatterer leg. • 1 sh; same collecting data as for preceding; depth 15 m; 2015 • 1 sh; same collecting data as for preceding; 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.
Israel • 1 sh; west of Rosh HaNikra Islands; 33.0704°N, 35.0926°E; depth 12 m; 29 Oct. 2018; HELM project (sample S52_2M);
Coriophora lessepsiana Albano, Bakker & Sabelli, sp. nov. A–J holotype,
Shell cyrtoconoid of ~ 3 mm with 11 whorls and multispiral protoconch. 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.
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 (Mediterranean specimen, without apex).
Protoconch : multispiral with five whorls, H 530 µm (holotype), 553 µm (paratype 1).
Protoconch I: 1.5 whorls with hemispherical granules, nucleus height of 114 µm (holotype), 104 µm (paratype 1) and a maximum diameter of 154 µm (holotype), 145 µm (paratype 1).
Protoconch II: 3.5 monocarinated whorls with axial orthogonal riblets with a maximum diameter of 305 µm (holotype), 311 µm (paratype 1), 323 µm (paratype 2), 268 µm (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).
The tuberculate first and third spiral cords start simultaneously after the protoconch 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 apparently 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.
Named after the Lessepsian invasion (
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.
Holotype. Egypt • sh; Sinai (Red Sea), Dahab, dive site “Islands”; 28.476°N, 34.513°E; depth 10 m; 2015; H. Blatterer leg.;
Paratypes. Egypt • sh; same collecting data as for holotype;
Egypt • 3 shs (juv. and fragments); same collecting 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. Blatterer 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).
Opimaphora blattereri Albano, Bakker & Sabelli, sp. nov., dive site “Islands”, Dahab, Sinai, Egypt A–J holotype,
Shell cyrtoconoid of less than 3 mm with 11 (holotype) or 12 (paratype 2) whorls and multispiral protoconch. Nucleus with hemispherical granules. Sculpture 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.
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 µm (holotype), 644 µm (paratype 2).
Protoconch I: 1.5 whorls with hemispherical granules, nucleus with a height of 109 µm (holotype), 122 µm (paratype 2), and a maximum diameter of 371 µm (holotype), 380 µm (paratype 2).
Protoconch II: 3.5 whorls with axial orthogonal riblets with a maximum diameter of 371 µm (holotype), 380 µm (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 protoconch with the same size, whereas the second cord appears from the fourth to the seventh 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, microsculpture is visible as fine spiral lines. Anterior siphonal canal tubular, short and oblique; posterior siphonal canal a simple notch.
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.
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. lessepsiana 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. blattereri 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 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 bilineatum (Kosuge, 1962) (holotype illustrated by
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);
We found a single, adult, empty shell. It likely possesses a large paucispiral 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.
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_2F, 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 S52_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 S13_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_3F, S53_3M); size of largest specimen: H 11.6 mm, W 2.7 mm.
This species was first recorded from Israel by
Israel • 1 sh; Ashqelon; 31.6868°N, 34.5516°E; depth 11 m; 31 Oct. 2018; offshore rocky reef; suction sampler; HELM project (sample S58_1F) • 1 spcm; Ashqelon; 31.6891°N, 34.5257°E; depth 28 m; 31 Oct. 2018; offshore rocky reef; suction sampler; HELM project (sample S59_3F);
Cerithiopsis pulvis (Issel, 1869): Israel • 2 spcms, 1 sh; Ashqelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr. 2018; offshore rocky reef; suction sampler; HELM project (sample S12_1F) • 5 spcms; same collecting data as for preceding; depth 11 m; 31 Oct. 2018; HELM project (samples S58_1F, S58_1M, S58_2F) • 5 spcms, 2 shs; Ashqelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef; suction sampler; HELM project (samples S16_1F, S16_2F, 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 S52_2F, S52_2M, S52_3M) • 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_1M) • 6 spcms; same collecting data as for preceding; depth 19 m; 29 Oct. 2018; HELM project (samples S53_1F, S53_1M, S53_3F).
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 background; 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.
Comparison between Cerithiopsis pulvis (Issel, 1869) and Cerithiopsis sp. aff. pulvis. A–C Cerithiopsis pulvis, Ashqelon, Israel, HELM project (sample S16_1F): front (A), side (B) and back (C) views. D–F Cerithiopsis sp. aff. pulvis,
Israel • 1 sh; Shiqmona Beach; 32.8259°N, 34.9555°E; beached; 4 Jan. 2008; size: H 3.5 mm, W 1.2 mm.
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 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, described 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 interspaces, deep brown suture and violet protoconch is strikingly different from the one of our shell. We have some reservations that S. lauta, 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 (
Holotype. Israel • spcm; Ashqelon; 31.6868°N, 34.5516°E; depth 11 m; 31 Oct. 2018; offshore rocky reef; suction sampler; HELM project (sample S58_3F);
Paratypes. Israel • 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);
Israel • 5 spcms; Ashqelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr. 2018; offshore rocky reef; suction sampler; HELM project (samples S12_1F, S12_2F, S12_3F) • 6 spcms; same collecting data as for preceding; depth 11 m; 31 Oct. 2018; HELM project (samples S58_1F, S58_2F, S58_3F) • 1 spcm; Ashqelon; 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 S14_2F, S14_4F) • 16 spcms; same collecting data as for preceding; 29 Oct. 2018; HELM project (samples S52_1F, S52_1M, S52_2F, 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 S53_1F, S53_2F, S53_3F, S53_3M).
Joculator problematicus Albano & Steger, sp. nov., holotype,
Very small bulbous brown shell, of ~ 1.5 mm in height and < 1 mm in width, with a relatively short, almost smooth protoconch.
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 protoconchs I and II, height: ~ 300 µm, width ~ 200 µm (holotype), but accurate measurement 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).
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
The name problematicus refers to the difficult task of recognizing and identifying non-indigenous species belonging to groups whose taxonomy in the tropical seas is poorly known (see Discussion). The species epithet is an adjective in nominative singular masculine.
This species is characterized by its bulbous contour and constricted last whorl which justify its inclusion in the genus Joculator Hedley, 1909 (
The Cerithiopsidae of the Indo-Pacific have been subject to numerous in-depth studies (
There are several more species of small brown bulbous Joculator often distinguishable 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 specimens. 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 Cecalupo & 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 tuberculate 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 Prkić & 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-like process of the anterior outer lip. Additionally, tubercles in C. ladae on the last whorl are more elongated, subrectangular, and the shell profile is less bulbous. Cerithiopsis greppii 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 Joculator is a broadly distributed genus in the Indo-Pacific province, we consider J. problematicus another previously undescribed Indo-Pacific species recently introduced to the Mediterranean Sea.
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 substrate; suction sampler; HELM project (samples S13_1F, S13_1M, S13_2F, S13_3L); size of largest specimen: H 1.6 mm, W 1.3 mm • 9 spcms; same collecting data as for preceding; depth 19 m; 29 Oct. 2018; HELM project (samples S53_1F, S53_2F, S53_3F).
Elachisina robertsoni Kay, 1979: United States • 1 sh; Hawaii, Oahu, Maunalua Bay;
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 distinguished 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 (
Comparison between Elachisina sp. and Elachisina robertsoni Kay, 1979 A–E Elachisina sp., west of Rosh HaNikra Islands, Israel, HELM project (sample S13_3L): front (A, B), side (C) and back (D, E) views F–H Elachisina robertsoni,
Israel • 1 spcm; Ashqelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr. 2018; offshore rocky reef; suction sampler; HELM project (sample S12_1F) • 5 shs; same collecting data as for preceding; depth 11 m; 31 Oct 2018; HELM project (sample S58_2F); size: H 2.8 mm, W 1.1 mm (largest (illustrated) shell).
Iravadia aff. elongata: Sudan • 2 shs; Arusa (near Port Sudan); 19.90°N, 37.23°E; shallow water; 1975; shell-grit; G. Spada leg.; MZUB.
Iravadia elongata (Hornung & Mermod, 1928): Eritrea • 1 sh; Massawa; depth 30 m; 1870; A. Issel leg.; syntype in
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
The closest match to our specimens is Iravadia elongata (Hornung & Mermod, 1928) which was described from material collected by Arturo Issel in the Red Sea off Massawa, Eritrea, at 30 m depth (
Among Mediterranean iravadiids, our specimens superficially resemble only Ceratia proxima (Forbes and Hanley, 1850). This species, however, lacks axial sculpture. Interestingly,
Iravadia aff. elongata (Hornung & Mermod, 1928) A–F Ashqelon, Israel, HELM project (sample S58_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).
Israel • 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.
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
Israel • 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).
This slender eulimid is characterized by a constriction at the transition between the protoconch and the teleoconch (Figure
Hypermastus 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 protoconch and teleoconch that is a diagnostic character for this species. Scale bars: 0.5 mm (A–E); 0.2 mm (F, G).
Israel • 1 spcm; Ashqelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr. 2018; offshore rocky reef; suction sampler; HELM project (sample S12_1F) • 38 spcms; Ashqelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef; suction sampler; HELM project (samples S16_1F, S16_2F, S16_2M); size: H 2.7 mm, L 0.9 mm (illustrated shell) • 16 spcms; same collecting data as for preceding; depth 28 m; 31 Oct. 2018; HELM project (samples S59_1F, S59_2F, S59_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).
Sticteulima clandestina and S. athenamariae, both Mifsud & Ovalis, 2019, were described on specimens collected in Turkey (
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 Rayneval & Ponzi, 1854), Plakias, Crete, Greece (sample Rh.05_5M): front view. E, F Vitreolina cf. philippi, Ashqelon, Israel, HELM project (S16_2F): front (E) and side (F) views. Scale bars: 0.5 mm.
Israel • 5 spcms; Ashqelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr. 2018; offshore rocky reef; suction sampler; HELM project (samples S12_1F, S12_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 S58_2F) • 3 spcms; Ashqelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef; suction sampler; HELM project (samples S16_1F, S16_2F) • 1 spcm; same collecting data as for preceding; depth 28 m; 31 Oct. 2018; HELM project (sample S59_1F).
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
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 (illustrated shell, Figure
Parvioris ibizenca (F. 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 S52_2F).
The genus Parvioris Warén, 1981 was erected for a group of numerous conchologically very similar species of which many are still undescribed (
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
Comparison between Parvioris aff. dilecta (E.A. Smith, 1899) and Parvioris ibizenca (F. Nordsieck, 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 S52_1F): front view J–K Parvioris aff. dilecta, west of Rosh HaNikra Islands, Israel, HELM project (sample S52_1M): front (J) and side (K) 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).
Israel • 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
We place this species in Sticteulima due to its small size, slender profile with high and rather flat whorls (
Israel • 4 spcms; Ashqelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef; suction sampler; HELM project (samples S16_1F, S16_2F); size: H 2.6 mm, W 0.9 mm.
Vitreolina philippi (de Rayneval & Ponzi, 1854): GREECE • Crete, Plakias; 35.1796°N, 24.3957°E; depth 5 m; 24 Sep. 2017; Posidonia oceanica rhizomes; suction sampler (sample Rh.05_5M).
This Vitreolina is extremely similar to the native V. philippi, but the animal is whitish with a yellowish digestive gland (Figure
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 formations; grab; HELM project (sample NG40_2M);
Conus fumigatus was first recorded from the Mediterranean Sea in Libya (
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.
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 µm). 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 aperture. The 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 currently known species of that genus were described form the tropical West Atlantic (Florida, Bahamas, and Yucatan) (
Among Mediterranean gastropods, the shell shape somewhat resembles the iravadiid 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
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_1F, 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 S53_1F).
The shell of this species is white and rather solid, with convex, unkeeled 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
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 (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).
This species is characterized by a translucid-white, cylindrical shell with ~ 3 whorls, and an intorted protoconch of type C (Figure
Odostomia sp. 1 does not resemble any known Mediterranean pyramidellid; considering 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.,
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).
Israel • 1 spcm; Soreq desalination plant; 31.9420°N, 34.6896°E; depth 17.4 m; 19 May 2015; soft substrate; grab; Soreq project (sample S027).
Israel • 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; previously figured by
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 (
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 unaware 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.
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).
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.;
Indonesia • 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.;
Taiwan • 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 & P.F. Liu leg.;
Miralda sp. (Oscilla jocosa sensu van
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
This species has just been described from the infralittoral of Jordan and also occurs in the Egyptian Red Sea (
Within the Mediterranean, O. virginiae is superficially similar only to two other non-indigenous pyramidellids, Cingulina isseli (Tryon, 1886) and Miralda sp. (Figure
Comparison between Oscilla virginiae Peñas, Rolán & 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
Israel • 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
Parthenina indistincta (Montagu, 1808): Israel • 2 shs; Ashqelon; 31.7002°N, 34.5498°E; depth 21 m; 18 Sep. 2016; sand; grab; HELM project (sample SG20_2F) • 1 sh; Ashqelon; 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).
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 profile, 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
The type material of P. cossmanni was collected from the Red Sea of Massawa (Eritrea) at a depth of 30 m (
Among native Mediterranean species, P. cossmanni superficially resembles Parthenina 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 (
We suspect the two shells illustrated as P. indistincta in
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, Ashqelon, 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).
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; Palmachim; 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).
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 columellar 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 μm, which is slightly smaller than 270–290 μm stated by
Parthenina typica (Laseron, 1959) was described from eastern Australia (
Israel • 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 S12_2F, S12_3F) • 17 spcms, 1 sh; same collecting data as for preceding; depth 11 m; 31 Oct. 2018; HELM project (samples S58_1F, 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 sampler; HELM project (sample S16_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 S52_2F, 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 S13_3F) • 2 spcms; same collecting data as for preceding; depth 19 m; 29 Oct. 2018; HELM project (sample S53_1F).
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.;
Madagascar • 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 gorgonians; J.H. McLean leg.;
Maldives • 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;
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.;
Spiralinella incerta (Milaschewitsch, 1916): Israel • 1 sh; north of Atlit; 32.7820°N, 34.9466°E; depth 10 m; 21 Sep. 2016; sand; grab; HELM project (sample NG10_1F) • 1sh; north of Atlit; 32.7422°N, 34.9181°E; depth 30 m; 20 Sep. 2016; sand; grab; HELM project (sample NG30_2F) • 2 shs; Ashqelon; 31.7002°N, 34.5498°E; depth 21 m; 18 Sep. 2016; sand; grab; HELM project (sample SG20_2F) • 1 sh; Ashqelon; 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 S14_4F); size: H 1.5 mm, W 0.7 mm (illustrated shell).
Pyrgulina craticulata has been reported only from the Red Sea so far, however, a search of previously uncatalogued lots of pyramidellid shells in the
Here, we provide the first records of P. craticulata for the Mediterranean Sea. Several 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 ornamentation, this species closely resembles the native Spiralinella incerta (Figure
Comparison between Pyrgulina craticulata (Issel, 1869) and Spiralinella incerta (Milaschewitsch, 1916) A–I Pyrgulina craticulata, Ashqelon, Israel, HELM project (sample S58_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 S14_4F): front (J, K) and back (L, M) views, detail of the sculpture of the body whorl (N). Scale bars: 0.5 mm (A–D, J–M); 0.2 mm (E, F, I, N); 0.1 mm (G, H).
Israel • 2 spcms; Ashqelon; 31.6891°N, 34.5257°E; depth 28 m; 31 Oct 2018; offshore rocky reef; suction sampler; HELM project (sample S59_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 S52_1F, S52_2F).
Until now, the first record of this species from the Mediterranean was considered to be by Öztürk and van Aartsen (2006), who reported on material obtained from shallow-water sediment samples collected along the Turkish Levantine (Viransehir, Mersin Bay) and Aegean coasts (Güllük Bay) in 1997 and 2000, respectively. However, already
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 S52_1F, S52_2F, S52_3F); size: H 1.8 mm, W 0.6 mm (illustrated specimen).
China • 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.;
Pakistan • 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.;
Sri 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.;
Vietnam • 1 sh; station 1328 of the Nha-Trang Oceanography Institute: “Entre les îles des Pêcheurs, Hon-Mung, le banc du Castleragh et l’isobathe 50. (En dehors et au S. de la baie de Nha-Trang)” [between Îles des Pêcheurs, Hon-Mung, Banc du Castleragh (= Castlereagh) and isobath 50. (outside of and to the south of the Bay of Nha-Trang)]; sediment sample; dredge;
Turbonilla funiculata de Folin, 1868, west of Rosh HaNikra Islands, Israel, HELM project (sample S52_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 (H) and side (I) views of the protoconch. Scale bars: 0.5 mm (A–D); 0.1 mm (E, H, I); 0.2 mm (F, G).
Shells of Turbonilla funiculata are polymorphic with respect to their shape (
Turbonilla funiculata has been previously reported from Fiji, Hong Kong, Indonesia, New Caledonia, Thailand, the Solomon Islands and Vietnam from shore to 396 m depth (
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, 34.6551°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; Palmachim; 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 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)).
Oman • 13 shs; off Muscat; 24°58'N, 56°54'E; 156 fathoms (285 m) depth; NMW.1955.158.00578 (F.W. Townsend coll.).
We record here for the first time in the Mediterranean 13 living individuals of Cylichna collyra, a cephalaspidean originally described from the Gulf of Oman (
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, S52_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 S13_2M, S13_3F) • 12 spcms; same collecting data as for preceding; depth 19 m; 29 Oct. 2018; HELM project (samples S53_1F, S53_2F, S53_2M, S53_3F).
This species was first recorded in the Mediterranean Sea in 1974 with the finding of few empty shells in the Bardawil Lagoons in Egypt (
Greece • 1 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.
Atys angustatus was first recorded in the Mediterranean Sea in 1974, based on specimens collected at Haifa, Israel (
Israel • 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 • 1 spcm; Palmachim; 31.9737°N, 34.6767°E; depth 35.4 m; 2 May 2018; soft substrate; box-corer; Shafdan project (sample 29(A)).
This species was first recorded in Israel in 1960 from Bat Yam with living individuals (
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 substrate with sand, rocks and Cystoseira; P.G. Albano leg. (sample NCY7H);
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.
Lioberus ligneus: Egypt • 19 vv; Red Sea, Suez; Jousseaume coll.;
Lioberus agglutinans (Cantraine, 1835): Portugal • 1 v; Algarve, Canal of Olhão; depth 3–7m,
Tunisia • 1 sh, 8 vv; Gulf of Gabès, NW Boughrara Gulf; depth 10–15 m;
We here report Lioberus ligneus for the first time from Israel and Cyprus. 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 indeed prefers vegetated habitats in the shallow subtidal. This species has been previously reported from Lebanon based on shells collected in 1999–2000 (
Comparison between Lioberus ligneus (Reeve, 1858) and Lioberus agglutinans (Cantraine, 1835). A, B Lioberus ligneus,
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 project (samples S52_1M, S52_2L, S52_3M, S52_3L) • 1 sh; north of Nahariyya port; 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
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 floating objects dumped on the shore (J. Steger coll.).
Israel • 10 spcms, 3 vv; Ashqelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr. 2018; offshore rocky reef; suction sampler; HELM project (samples S12_1F, S12_1M, S12_1L, S12_2F, S12_2M) • 111 spcms, 2 vv; Ashqelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef; suction sampler; HELM project (samples S16_1F, 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 S14_2F, S14_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 S13_3M) • 1 spcm; same collecting data as for preceding; depth 19 m; 29 Oct. 2018; HELM project (sample S53_2L).
Malta • 9 spcms; off Filfla Island; on a rafting fisherman’s net; 1981 (P.G. Albano coll. ID 557).
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
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.
Comparison between Musculus coenobitus (Vaillant, 1865) and Musculus subpictus (Cantraine, 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 S12_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).
Israel • 1 spcm; Ashqelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef; suction sampler; HELM project (sample S16_2F) • 15 spcms; same collecting data as for preceding; depth 28 m; 31 Oct. 2018; HELM project (samples S59_1F, 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 S14_3F, S14_4M) • 25 spcms; same collecting data as for preceding; 29 Oct. 2018; HELM project (samples S52_1F, S52_2F, S52_2M, S52_2L, S52_3F, S52_3M, S52_3L); size: L 3.6 mm, H 2.1 mm (illustrated specimen, Figure
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 S52_3L): left valve outer (A, B) and inner (D) views, right valve inner (C) and outer (E) views F–H 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 S53_2L): left valve outer (I, J) and inner (L) views, right valve inner (K) and outer (M) views. Scale bars: 1 mm.
Musculus aff. viridulus: Egypt • 1 v; Sinai (Red Sea), Dahab, dive site “Caves”; 28.416°N, 34.456°E; depth 15 m; 2015; H. Blatterer leg. • 4 vv; 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.
Musculus costulatus (Risso, 1826): Israel • 9 spcms, 4 vv; Ashqelon; 31.6868°N, 34.5516°E; depth 12 m; 30 Apr. 2018; offshore rocky reef; suction sampler; HELM project (samples S12_1F, S12_1M, S12_1L, S12_2F, S12_2M) • 3 spcms; Ashqelon; 31.6891°N, 34.5257°E; depth 25 m; 2 May 2018; offshore rocky reef; suction sampler; HELM project (samples S16_1F, S16_2F) • 1 spcm; same collecting data as for preceding; 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; suction sampler; HELM project (samples S14_1L, S14_3F, S14_3L, S14_4F) • 56 spcms; same collecting data as for preceding; 29 Oct. 2018; HELM project (samples S52_1F, S52_1M, S52_1L, S52_2F, S52_2M, S52_2L, S52_3F, S52_3M, S52_3L) • 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 S13_3F) • 20 spcms; same collecting data as for preceding; depth 19 m; 29 Oct. 2018; HELM project (sample S53_2L).
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. costulatus because the latter has a more oval outline and a much smaller number of riblets at the same overall shell size (Figure
Cyprus • 1 sh; Turkish Republic of Northern Cyprus, Esentepe; 35.3589°N, 33.6078°E; depth 2 m; 28 Jul. 2019; rocky substrate; P.G. Albano leg. (sample NCY3H); size: L 11.8 mm, H 10.1 mm (Figure
Greece • 1 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
The discrimination and identification of the species of Isognomon Lightfoot, 1786 is difficult due to their morphological plasticity that is related to their cryptic way of life. Still, the specimens recently reported from Astypalaia, in the Eastern Aegean Sea (
Isognomon aff. australica (Reeve, 1858) (sensu
Israel • 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, H 13.5 mm.
Pegophysema philippiana was first found in the Mediterranean Sea in 2018 as a single valve from south of Tel Aviv (
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.
We report a juvenile but living individual of Chavania erythraea, a lucinid occurring in the Red Sea, the Persian (Arabian) Gulf and the Arabian Sea (
Israel • 1 spcm, 1 v; Ashqelon; 31.6868°N, 34.5516°E; depth 11 m; 31 Oct. 2018; offshore rocky reef; suction sampler; HELM project (samples S58_2M, S58_3M).
This species was first recorded as Pillucina vietnamica Zorina, 1978 (now in Rugalucina too) from the Mediterranean coast of Israel by
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
Nudiscintilla cf. glabra Lützen & Nielsen, 2005 (sensu
This non-indigenous species has first been recorded in the Mediterranean Sea by
Israel • 1 spcm; Haifa; depth 15 m; May 1999; biogenic sediment; B.S. Galil leg.; size: L 6.9 mm, H 5.2 mm.
The single shell found is trapezoid-oval (L:H ratio = 1.33), slightly higher 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 prosogyrate, 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, particularly of the right valve, is spotted by blister-like markings (Figure
Lacking further material and observations on living individuals, which are of great diagnostic importance in galeommatids, we refrained from assigning a definitive specific 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 (
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 sculpture on the valves. Apart from the present shell, which was found in 1999 in Haifa Bay, we know of no other material.
Israel • 6 spcms; Ashqelon; 31.7002°N, 34.5498°E; depth 21 m; 18 Sep. 2016; sand; grab; HELM project (samples SG20_1F, SG20_3F, SG20_3M).
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 (
Israel • 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 S58_3M);
Ervilia scaliola was first recorded from Turkey based on material collected in 2013 by
GREECE • 3 vv; Kos Island; 2005–2010; A. Storm leg.;
We here report the finding of three beached valves of Iacra seychellarum in Kos, Greece. Iacra 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 (
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.
Abra alba (W. Wood, 1802): Israel • 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).
We were unable to identify this peculiar bivalve beyond family level, despite 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 margin 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 family Semelidae (
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 lifestyle); however, the outer shell surface of L. katoi bears rough lamellae and fine radial threads (
Until the finding of further specimens, it remains open whether the present individual 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.
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 (F, L); 0.3 mm (C, I).
Israel • 1 spcm; Haifa Bay; 32.8211°N, 35.0196°E; depth 11 m; 2 Aug. 2015; soft substrate; grab; NM project (station HM27).
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 (
Israel • 1 spcm; Ashqelon; 31.7002°N, 34.5498°E; depth 21 m; 18 Sep. 2016; sand; grab; HELM project (sample SG20_4F) • 3 spcms; Ashqelon; 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
Corbula erythraeensis: Egypt • 1 sh; Bay of Safaga; 26.8142°N, 33.9653°E; depth 39 m; 31 Oct. 1994; mud; scuba diving; M. Zuschin leg.; ref.
Corbula gibba (Olivi, 1792): Israel • 271 spcms; Ashqelon; 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.
Corbula erythraeensis is widespread in the northern Red Sea where it has been recorded from the gulfs of Suez (
Comparison between Corbula erythraeensis H. Adams, 1871 and Corbula gibba (Olivi, 1792). A–E Corbula erythraeensis, Ashqelon, 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,
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 programs. 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 (
Taxonomic uncertainty is recognized as a major impediment to the reliable inventorying of non-indigenous species (
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 taxonomically identifiable, dead or discarded organic remains encountered in a landscape or seabed such as molluscan shells, accumulate species richness over time (
With biological invasions constituting a major element of global change, there is understandable concern on the accuracy of non-indigenous species inventories (
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, PILF, 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 the figures. PGA and JS wrote the first draft of the manuscript, which then received contributions by all co-authors.
This research has been conducted in the context of the “Historical ecology of Lessepsian migration” project funded by the Austrian Science Fund (FWF) P28983-B29 (PI: P.G. Albano). Sampling in Crete was supported by the “Kurzfristige wissenschaftliche 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: F. Huseynoglu). 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 (Österreichischer Austauschdienst).
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, Alexander 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 material 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 Jarochowska and Christian Schulbert provided the µCT-scan and 3D-reconstruction of Odostomia (s.l.) sp. 1. Henk Dekker commented on the manuscript. Sampling in Israel was conducted under permit 41928 by the Israel Nature and Parks Authority.