Scaphopod species (Mollusca) of the Turkish Levantine and Aegean

B. ÖZTÜRK

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Turk J Zool

2011; 35(2): 199-211

© TÜBİTAK

doi:10.3906/zoo-0904-23

Scaphopod species (Mollusca) of the Turkish Levantine and

Aegean seas

Bilal ÖZTÜRK*

Ege University, Faculty of Fisheries, Department of Hydrobiology, 35100 Bornova, İzmir - TURKEY

Received: 28.04.2009

Abstract: An examination of benthic material collected from diff erent depths (0-875 m) and habitats along the Turk-

ish Levantine and Aegean seas between 1996 and 2008 has resulted in the identifi cation of 819 scaphopod specimens,

belonging to 10 species: Antalis dentalis (Linneaus, 1758), Antalis inaequicostata (Dautzenberg, 1891), Antalis panorma

(Chenu, 1843), Antalis rossati (Caprotti, 1966), Antalis vulgaris (da Costa, 1778), Fustiaria rubescens (Deshayes, 1825),

Entalina tetragona (Brocchi, 1814), Pulsellum lofotense (M. Sars, 1865), Cadulus jeff reysi (Monterosato, 1875), and Dis-

chides politus (S. Wood, 1842). Of these, P. lofotense and C. jeff reysi are new records for the Aegean molluscan fauna and

A. panorma and F. rubescens are new to the Levantine coast of Turkey. Of the identifi ed species, A. inaequicostata was

the most widely distributed along the Turkish coast, whereas C. jeff reysi was the rarest, and was encountered in only

2 places. Entalina tetragona was signifi cant as the deepest living species, sampled from depths between 96 and 875 m.

Some ecological characteristics and taxonomic remarks, with colour photographs of the identifi ed species, are provided.

Key words: Scaphopoda, Aegean Sea, Levantine Sea, ecology, distribution

Türkiye’nin Levantine ve Ege Denizi Scaphopoda (Mollusca) türleri

Özet: 1996-2008 yılları arasında Türkiye’nin Levantine ve Ege Denizi kıyılarının farklı derinlik (0-875 m) ve

biyotoplarından örneklenen bentik materyalin incelenmesi sonucu, 10 türe ait: Antalis dentalis (Linneaus, 1758), Anta-

lis inaequicostata (Dautzenberg, 1891), Antalis panorma (Chenu, 1843), Antalis rossati (Caprotti, 1966), Antalis vulgaris

(da Costa, 1778), Fustiaria rubescens (Deshayes, 1825), Entalina tetragona (Brocchi, 1814), Pulsellum lofotense (M. Sars

1865), Cadulus jeff reysi (Monterosato, 1875), ve Dischides politus (S. Wood, 1842) 819 skafopod bireyi saptanmıştır.

Bunlardan P. lofotense ve C. jeff reysi Ege Denizi Mollusca faunası için yeni kayıt olup, A. panorma ve F. rubescens ise

Türkiye’nin Levantine Denizi kıyılarından ilk defa rapor edilmiştir. Saptanan türlerden A. inaequicostata Türkiye

kıyılarında en geniş dağılıma sahip olmasına karşın, sadece iki istasyonda tespit edilen C. jeff reysi ise dağılımı en nadir

olan türdür. 96-875 m arasındaki derinliklerde bulunmuş olan E. tetragona, saptanan türler arasında en derin dağılımlı

tür olarak dikkat çekmektedir. Bu çalışmada, incelenen türlerin bazı ekolojik ve taksonomik özelliklerinin yanında,

bunların renkli fotoğrafl arına da yer verilmiştir.

Anahtar sözcükler: Scaphopoda, Ege Denizi, Levantine Denizi, ekoloji, dağılım

Research Article

* E-mail: [email protected]

Scaphopod species (Mollusca) of the Turkish Levantine and Aegean seas

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Introduction

Th e Levantine and Aegean seas are 2 distinct basins of the Mediterranean Sea, each with diff erent characteristics. Th e Aegean Sea is an area where the brackish waters of the Black Sea communicate with the more saline waters of the eastern Mediterranean Sea, whereas the Levantine Sea is the most oligotrophic basin in the Mediterranean. Th e conspicuous hydrological and ecological diff erences between the 2 basins greatly infl uence their biological diversity. According to Boudouresque (2004), the number of macroscopic marine species inhabiting the Mediterranean is estimated to be about 12,000, of which a total of almost 2000 are molluscs, and a few belong to the class Scaphopoda.

Scaphopods are bilaterally symmetrical marine molluscs, with an external tubular calcareous shell that is open at both ends, which is more or less markedly curved. Th ey are benthic animals, living partly embedded in the soft substrata between the shallow and abyssal realms to a depth of approximately 7000 m (Poppe and Goto, 1993), and feeding on foraminifers and small algae.

Scaphopods are relatively well-studied molluscs, both at the global and local scales. Faunistic investigations on this subject were undertaken in the 19th century, the monograph by Pilsbry and Sharp (1897-1898) being a particularly fundamental work in which the Scaphopoda was revised. More than half a century aft er Pilsbry and Sharp’s publication, Emerson (1962) revised and expanded their classifi cation. Emerson (1962), aft er reviewing the previous studies, gave detailed information on scaphopod classifi cation, with keys to the families, genera, and subgenera. He also added 8 new genera and 1 subgenus to the existing taxa. In addition to these major publications, the studies by Ludbrook (1960), Scarabino (1995), Steiner (1992, 1996), and Steiner and Kabat (2001, 2004) are the next important works on this subject. Scarabino (1995), in his study on tropical scaphopods, also described 3 new genera and 42 new species. Recently, Steiner and Kabat (2004) published a catalogue of species-group names of recent and fossil scaphopods, in which 517 species, belonging to 2 orders (Dentaliida da Costa, 1776 and Gadilida Starobogatov, 1974) and 45 genera (Steiner and Kabat, 2001), were listed as valid recent taxa in the world’s oceans.

In the past century, especially in its second half, several studies (Stork, 1934; Caprotti, 1965, 1968, 1979; Steiner, 1997) were carried out on the Mediterranean scaphopods. Stork (1934), in his study performed in the Adriatic Sea, reported 9 species from the area and gave detailed information on the identifi ed species, with a key for family, genus, and species determinations. More recently, Caprotti (1979) carried out an examination of Neogene and recent Mediterranean scaphopods, in particular discussing and comparing the living species with the fossilised ones. According to Sabelli et al. (1990), 16 scaphopod species have been recorded from the Mediterranean, of which 8 species were also recorded from the Levantine Sea (Cypriot coast) (Öztürk et al., 2003), and 10 species from the Aegean Sea (Koukouras and Kevrekidis, 1986). According to the studies performed along the Turkish coasts (Ostroumoff , 1896; Marion, 1898; Oberling, 1969-1971; Aartsen and Kinzelbach, 1990; Buzzurro and Greppi, 1996; Demir, 2003; Çevik and Sarıhan, 2004), a total of 8 species occur in the Levantine Sea, Aegean Sea, and the Sea of Marmara.

Th e present study focuses on some morphological features of the shells, along with some ecological and distributional information on the scaphopod species occurring along the Turkish Levantine and Aegean coasts.

Materials and methods

Scaphopod specimens examined in this study were collected during various cruises or research projects with diff erent purposes, carried out along the Turkish Levantine and Aegean coasts during the period from 1996 to 2008. Th e greater part of the Aegean Sea material was collected during research cruises in this area, undertaken by R/V Hippocampus in the summer and autumn of 2000, whereas the Levantine Sea material was sampled mostly within Project Number: 104 Y 065, supported by TÜBİTAK (Th e Scientifi c and Technological Research Council of Turkey). Th e remaining material was collected by R/V K. Piri Reis and Egesüf, or with the aid of other boats. An important portion of the material from Fethiye Bay (Levantine Sea) was obtained during a project funded by the Environmental Protection Agency for Special Areas (Republic of Turkey Ministry of Environment and Forestry).

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Benthic samples were collected from diff erent habitats at more than 500 stations and at depths reaching a maximum of 875 m. Scuba diving and snorkelling were used at shallow-water stations, whereas at deeper stations, the material was taken by using standard sampling gears such as an anchor dredge, a beam-trawl, and a Van Veen grab. Once on board, the material was sieved with a 0.5-mm mesh and the retained fauna was preserved in a 10% seawater-formalin solution.

In the laboratory, the sampled material was sorted into diff erent groups. Scaphopod specimens were separated from other benthic groups, identifi ed at the species level, and counted. Some shell features, such as length (= height) and aperture diameter (and apex in some species) of the specimens belonging to diff erent species, were measured using callipers or an ocular micrometer. Of the empty shells, only those with well-preserved characteristics were taken into consideration. Th e location of the stations (67) from which the scaphopod specimens were collected (Figure 1), and some ecological characteristics, such as biotope type and station depth, are also given (Table 1). Th e investigated specimens, with individual catalogue numbers, have been deposited in the museum collections of the Faculty of Fisheries (ESFM), Ege University (İzmir-Turkey).

Results

Th e examination of 664 scaphopod individuals and 155 empty shells (total 819), collected from the Levantine and Aegean coasts of Turkey, yielded 10 species: Antalis dentalis (Linneaus, 1758), Antalis inaequicostata (Dautzenberg, 1891), Antalis panorma (Chenu, 1843), Antalis rossati (Caprotti, 1966), Antalis vulgaris (da Costa, 1778), Fustiaria rubescens (Deshayes, 1825), Entalina tetragona (Brocchi, 1814), Pulsellum lofotense (M. Sars, 1865), Cadulus jeff reysi (Monterosato, 1875), and Dischides politus (S. Wood, 1842). Of these, 2 species (Pulsellum lofotense and Cadulus jeff reysi) are new records for the Aegean mollusc fauna and 2 species (Antalis panorma and Fustiaria rubescens) are new to the Levantine coast of Turkey. As for the families, Dentaliidae was represented by 5 species and 476 specimens (425 ind. + 51 sh.), all belonging to the genus Antalis Adams, H.&A., 1854 (A. dentalis, A. inaequicostata, A. panorma, A. rossati, and A. vulgaris); the Fustiariidae by 1 species (F. rubescens) and 74 specimens (61 ind. + 13 sh.); the Pulsellidae by 1 species (Pulsellum lofotense) and 6 individuals; the Gadilidae by 2 species (Cadulus jeff reysi and Dischides politus) and 125 specimens (104 ind., 21 sh.); and the Entalinidae by 1 species (Entalina tetragona) and 138 specimens (68 ind.+ 70 sh.) (Table 2).

40°00ʹΝ

N

S

W ET U R K E Y

35°00ʹΕ30°00ʹΕ26°00ʹΕ

36°00ʹΝ

Figure 1. Study area with location of the stations where scaphopod species were sampled.


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Table 1. Coordinates, sampling dates, depths, and biotope characterisations of the stations.

Coordinates

St Date Depth Biotope Location

Lat. Long. (m)

1 40°41ʹ40ʺ 25°58ʹ30ʺ 04.08.2000 15 Mud Saros Bay2 40°34ʹ45ʺ 26°09ʹ25ʺ 04.08.2000 8 P. oceanica + Sand Saros Bay3 40°33ʹ00ʺ 26°30ʹ20ʺ 03.08.2000 82 Sandy mud Saros Bay4 40°36ʹ30ʺ 26°28ʹ00ʺ 04.08.2000 21 Sand + Mud Saros Bay5 40°25ʹ38ʺ 26°25ʹ57ʺ 03.08.2000 8 P. oceanica + Sand Saros Bay6.1 40°27ʹ40ʺ 26°29ʹ57ʺ 03.08.2000 135 Mud Saros Bay6.2 40°23ʹ46ʺ 26°21ʹ46ʺ 03.08.2000 105 Sand + Mud Saros Bay7 40°26ʹ36ʺ 26°17ʹ47ʺ 14.05.2001 680 Mud Saros Bay8 40°17ʹ06ʺ 25°45ʹ05ʺ 15.05.2001 875 Mud Saros Bay9 40°13ʹ20ʺ 26°03ʹ00ʺ 02.08.2000 96 Mud North of Gökçeada10.1 40°04ʹ45ʺ 26°10ʹ50ʺ 13.08.2000 29 Sand Gelibolu Peninsula10.2 40°12ʹ15ʺ 26°03ʹ55ʺ 13.08.2000 28 Sand Gelibolu Peninsula11 40°05ʹ45ʺ 25°50ʹ45ʺ 13.08.2000 27 P. oceanica +Sand Gökçeada12 39°58ʹ50ʺ 26°03ʹ25ʺ 13.08.2000 30 Sand + Mud Ent. Çanakkale St.13 39°55ʹ30ʺ 25°50ʹ20ʺ 30.07.2000 77 Sand South of Gökçeada14 39°39ʹ15ʺ 26°02ʹ00ʺ 29.07.2000 70 Sand + Mud Bababurnu15 39°23ʹ10ʺ 26°45ʹ12ʺ 18.08.2000 38 Sand + Mud Ayvalık16 39°15ʹ00ʺ 26°32ʹ05ʺ 28.07.2000 53 Sand + Mud Ayvalık17 - - 25.03.1996 11.5 Mud Ayvalık18 39°00ʹ10ʺ 26°44ʹ28ʺ 28.07.2000 50 Mud Dikili19.1 38°54ʹ28ʺ 26°49ʹ57ʺ 17.03.2005 20 P. oceanica + Sand Kara Island (Çandarlı Bay)19.2 38°55ʹ29ʺ 26°49ʹ30ʺ 17.03.2003 46 Coralligenous sand Kızkulesi (Çandarlı Bay)19.3 38°51ʹ04ʺ 26°54ʹ56ʺ 07.03.2003 50 Coralligenous Tavşan Island (Çandarlı Bay)20 - - 19.10.1999 10-20 Mud Nemrut Bay (Aliağa)21 38°41ʹ25ʺ 26°58ʹ17ʺ 12.01.2001 24 Mud İzmir Bay22 38°44ʹ50ʺ 26°42ʹ00ʺ 11.04.2000 30 Sandy mud İzmir Bay23 38°43ʹ33ʺ 26°39ʹ00ʺ 08.03.2000 28 Sandy mud İzmir Bay24 38°38ʹ50ʺ 26°50ʹ50ʺ 13.10.1999 49 Sand İzmir Bay25 38°38ʹ33ʺ 26°58ʹ33ʺ 13.10.1998 68 Sand İzmir Bay26 38°41ʹ17ʺ 26°58ʹ17ʺ 11.01.2001 77 Sandy mud İzmir Bay27 38°44ʹ10ʺ 26°22ʹ00ʺ 12.09.2000 183 Mud Karaburun28 38°25ʹ16ʺ 26°21ʹ20ʺ 05.10.2000 5 Sand Karaburun Peninsula29.1 01.07.1999 6-61 Sand, Mud, Muddy sand; Sandy mud Çeşme 01.07.2000 Çeşme 14.09.2000 Çeşme 05.10.2000 Çeşme 22.02.2001 Çeşme 03.06.2005 Çeşme 09.05.2007 Çeşme 03.10.2007 Çeşme29.2 26.09.2002 10-65 Sand, Mud, Muddy sand; Sandy mud Çeşme 24.02.2003 Çeşme 01.05.2003 Çeşme 09.06.2003 Çeşme 27.04.2004 Çeşme 01.12.2004 Çeşme 22.05.2004 Çeşme 29.06.2004 Çeşme 07.03.2007 Çeşme 09.05.2007 Çeşme 11.09.2007 Çeşme 03.10.2007 Çeşme 05.02.2008 Çeşme 30 38°09ʹ30ʺ 26°17ʹ40ʺ 14.09.2000 113 Sand + Mud Çeşme31 38°08ʹ13ʺ 26°43ʹ00ʺ 30.09.2000 150 Mud Sığacık32 38°03ʹ10ʺ 26°56ʹ00ʺ 30.09.2000 41 Sand + Mud Kuşadası Bay33.1 37°55ʹ18ʺ 27°07ʹ41ʺ 14.09.2000 78 Mud Kuşadası Bay33.2 37°23ʹ55ʺ 27°06ʹ52ʺ 15.09.2000 71 Sand + Mud Kuşadası Bay33.3 37°50ʹ22ʺ 27°13ʹ11ʺ 08.10.2005 10-50 Sandy mud Kuşadası Bay

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Table 1. (Continued).

Coordinates

St Date Depth Biotope Location

Lat. Long. (m)

34 37°23ʹ55ʺ 27°06ʹ52ʺ 15.09.2000 79 Mud Didim35 37°19ʹ30ʺ 27°29ʹ00ʺ 16.09.2000 19 Sand + Mud Güllük Bay36 37°16ʹ00ʺ 27°35ʹ30ʺ 17.09.2000 13 Mud Güllük Bay37.1 37°09ʹ00ʺ 27°29ʹ30ʺ 17.09.2000 44 Sand + Mud Güllük Bay37.2 37°13ʹ08ʺ 27°32ʹ15ʺ 16.09.2000 31 Mud Güllük Bay38 37°12ʹ43ʺ 27°12ʹ18ʺ 29.09.2000 85 Mud Güllük Bay39 37°03ʹ50ʺ 27°13ʹ30ʺ 17.09.2000 37 Caulerpa racemosa + P. oceanica + Bodrum Sandy mud40 36°56ʹ45ʺ 27°16ʹ32ʺ 17.09.2000 9.5-31 Sand + Algae Bodrum 21.10.2001 16.02.2002 05.09.2002 41 36°59ʹ00ʺ 27°32ʹ35ʺ 18.09.2000 47 Sand + Algae Bodrum42 36°59ʹ30ʺ 27°47ʹ56ʺ 18.09.2000 64 Sand Gökova Bay43 36°58ʹ30ʺ 27°57ʹ10ʺ 18.09.2000 109 Sand + Mud Gökova Bay44 36°59ʹ00ʺ 27°05ʹ35ʺ 18.09.2000 82 Mud Gökova Bay45 37°02ʹ39ʺ 28°19ʹ20ʺ 18.09.2000 7 Mud Gökova Bay46 36°54ʹ40ʺ 28°09ʹ57ʺ 19.09.2000 19 Sand + Mud Gökova Bay47.1 36°49ʹ07ʺ 27°52ʹ10ʺ 20.09.2000 54 Sand + Mud Gökova Bay47.2 36°47ʹ58ʺ 27°41ʹ25ʺ 20.09.2000 51 Sand + Algae Gökova Bay48 36°44ʹ26ʺ 27°24ʹ19ʺ 20.09.2000 12 Sand + Algae Gökova Bay49.1 36°43ʹ16ʺ 27°42ʹ10ʺ 21.09.2000 47 Sand + Mud Datça49.2 36°45ʹ08ʺ 27°47ʹ00ʺ 21.09.2000 26 Sand + Algae Datça49.3 36°42ʹ45ʺ 28°05ʹ58ʺ 14.09.2000 54 Mud Datça49.4 36°42ʹ30ʺ 28°00ʹ15ʺ 21.09.2000 57 Sand Datça50.1 36°40ʹ31ʺ 28°09ʹ51ʺ 22.09.2000 44 Sand + Algae Marmaris50.2 36°50ʹ40ʺ 28°16ʹ10ʺ 23.09.2000 19 Mud Marmaris51 36°44ʹ30ʺ 28°31ʹ30ʺ 23.09.2000 136 Mud Marmaris52.1 36°36ʹ25ʺ 28°52ʹ08ʺ 20.06.2008 5-30 Sand Fethiye Bay52.2 36°36ʹ05ʺ 28°51ʹ57ʺ 24.06.2008 5-30 Sand Fethiye Bay52.3 36°38ʹ27ʺ 28°53ʹ54ʺ 24.06.2008 15 H. stipulacea + Sand Fethiye Bay52.4 36°39ʹ55ʺ 28°51ʹ26ʺ 30.06.2008 5-30 Sand Fethiye Bay52.5 36°41ʹ48ʺ 28°52ʹ10ʺ 01.07.2008 5 Sand Fethiye Bay53.1 36°38ʹ28ʺ 29°02ʹ37ʺ 06.10.2005 200 Mud Fethiye Bay53.2 36°37ʹ56ʺ 29°04ʹ31ʺ 06.10.2005 50 Sandy mud Fethiye Bay53.3 36°39ʹ24ʺ 29°04ʹ44ʺ 06.10.2005 50 Sandy mud Fethiye Bay53.4 36°38ʹ23ʺ 29°06ʹ46ʺ 06.10.2005 10 Mud Fethiye Bay53.5 36°41ʹ39ʺ 28°55ʹ41ʺ 07.07.2008 30 Sand Fethiye Bay53.6 36°45ʹ03ʺ 28°55ʹ50ʺ 05.07.2008 0-5 Sand Fethiye Bay53.7 36°40ʹ05ʺ 28°53ʹ09ʺ 26.06.2008 5 Sand Fethiye Bay53.8 36°42ʹ16ʺ 28°54ʹ25ʺ 02.07.2008 5-30 Mud Fethiye Bay53.9 36°45ʹ03ʺ 28°55ʹ50ʺ 22.09.2008 30 Sand Fethiye Bay54 36°39ʹ29ʺ 29°06ʹ02ʺ 06.10.2005 25 Sandy mud Fethiye Bay55 36°38ʹ38ʺ 29°04ʹ36ʺ 05.10.2005 1.5 Sand Fethiye Bay56 36°12ʹ06ʺ 29°37ʹ30ʺ 03.10.2005 9 Sand Kaş57 36°17ʹ24ʺ 30°13ʹ01ʺ 30.09.2005 50 Mud Finike58.1 36°02ʹ37ʺ 32°54ʹ06ʺ 23.09.2005 75 Muddy sand Anamur58.2 36°02ʹ02ʺ 32°53ʹ59ʺ 23.09.2005 200 Sandy mud Anamur58.3 36°02ʹ31ʺ 32°54ʹ54ʺ 23.09.2005 200 Mud Anamur58.4 36°02ʹ21ʺ 32°54ʹ01ʺ 23.09.2005 100 Mud Anamur59 36°18ʹ51ʺ 33°51ʹ47ʺ 19.09.2005 1 Sandy mud near Taşucu 60 36°37ʹ50ʺ 34°37ʹ47ʺ 17.09.2005 75 Mud Mersin61 36°33ʹ22ʺ 35°34ʹ17ʺ 10.09.2005 10 Muddy sand İskenderun Bay62 36°46ʹ04ʺ 35°47ʹ45ʺ 10.09.2005 9 Muddy sand İskenderun Bay63 36°52ʹ21ʺ 35°55ʹ05ʺ 07.03.2007 30 Mud İskenderun Bay64.1 36°43ʹ32ʺ 36°10ʹ28ʺ 09.09.2005 25 Muddy sand İskenderun Bay64.2 36°43ʹ19ʺ 36°09ʹ30ʺ 09.09.2005 50 Sandy mud İskenderun Bay65 36°38ʹ11ʺ 36°06ʹ48ʺ 08.09.2005 72 Mud İskenderun Bay66 36°21ʹ15ʺ 35°44ʹ27ʺ 10.09.2005 75 Sandy mud İskenderun Bay67 36°23ʹ18ʺ 35°39ʹ26ʺ 10.09.2005 100 Mud İskenderun Bay


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Table 2. Stations where the scaphopod specimens were collected and some shell features of the identifi ed species (H: Mean height

(length) of the shell with ± standard errors, and maximum value (mm) in brackets; D: Mean diameter of the aperture with ±

standard errors, and maximum value (mm) in brackets; H/D: Th e ratio between the mean height and mean diameter with ±

standard errors).

Species StationDepth

(m)H D H/D

Antalis dentalis

(Linneaus, 1758)

St. 24: 1 sp.; St. 29.2: 6 sp.; St. 33.3: 2 sp.; St. 6.1: 1 sp.; St. 46:

1 sp. 25-49

19.14 ± 10.23

(31.5)

2.38 ± 0.65

(3.30)8.08 ± 3.29

Antalis inaequicostata

(Dautzenberg, 1891)

St. 1: 4 sp.; St.4: 2 sp.; St. 6.1: 2 sp.; St. 14: 1 sp.; St. 15: 6 sp., 2

sh.; St. 16: 3 sp.; St. 18: 3 sp.; St. 19.2: 1 sp.; St. 19.3: 1 sp.; St. 20:

3 sp.; St. 21: 2 sp.; St. 22: 2 sp.; St. 23: 2 sp.; St. 24: 6 sp.; St. 25:

1 sp.; St. 26: 6 sp.; St. 29.1: 4 sp.; St. 29.2: 88 sp., 4 sh.; St. 32: 6

sp.; St. 33.1: 2 sp., 2 sh.; St. 33.3: 2 sp.; St. 35: 1 sp., 5 sh.; St. 37:

9 sp.; St. 38: 9 sp.; St. 39: 4 sp., 1 sh.; St. 40: 3 sp.; St. 45: 1 sp.; St.

46: 1 sp, 1 sh.; St. 47.1: 3 sp.; St. 49:1: 1 sp., 2 sh.; St.49.3: 2 sp.,

2 sh.; St.50.2:4 sp., 2 sh.; St. 43.9: 1 sp.; St. 53.4:1 sp.; St. 54.1: 2

sp.; St. 57: 5 sp.; St.58: 3 sp.; St. 60: 4 sp.; St. 61: 5 sp.; St. 62: 3

sp.; St. 64.2: 5 sp., 1 sh.; St. 65: 3 sp.; St. 66: 2 sp.; St. 67: 6 sp.

12-10025.55 ± 1.51

(41.6)

2.72 ± 0.14

(3.79)8.14 ± 0.44

Antalis panorma

(Chenu, 1843)

St. 6.2: 1 sp.; St. 25: 1 sp.; St. 27: 12 sp., 4 sh.; St. 37.2: 4 sp.; St.

43: 1 sp.; St. 51: 31 sp., 8 sh.; St. 53.1: 1 sp.; St. 58.3: 19 sp.; St.

58.4: 18 sp.

68-20028.04 ± 1.70

(40.4)

2.35 ± 0.11

(3.20)11.40 ± 0.44

Antalis rossati

(Caprotti, 1966)St. 17: 5 sp.; St.21: 1 sp.; St. 24: 1 sp.; St. 36: 6 sh. 11.5-49

24.94 ± 2.53

(41.2)

2.56 ± 0.17

(3.48)9.46 ± 0.40

Antalis vulgaris

(da Costa, 1778)

St. 2: 2 sp.; St. 10.1: 5 sp.; St. 19.1: 1 sp.; St. 19.2: 4 sp.; St. 28: 1

sp.; St. 29.1: 18 sp., 5 sh., St. 29.2: 6 sp.; St. 40: 26 sp., 3 sh.,; St.

48: 1 sp.; St.49.2: 3 sp.; St. 52.1: 2 sp.; St. 52.2: 3 sp., 1 sh.; St.

52.4: 1 sp.; St. 52.5: 1; St. 53.5: 2 sp.; St. 53.6: 2 sp.; St. 53.7: 3

sp.; St. 53.8: 2 sp.; St. 54.2: 3 sp.; St. 54.3: 1 sp.; St. 55: 3 sp., St.

56: 1 sp.; St. 58: 2 sp.; St. 59: 1; St. 64.1: 2 sp., 2 sh.

4-75 21.60 ± 1.59

(48.3)

2.50 ± 0.15

(4.60)7.04 ± 0.36

Fustiaria rubescens

(Deshayes, 1825)

St. 29.1: 7 sp.; St. 29.2: 8 sp.; St.33.3: 2 sp., 7 sh.; St. 35: 8 sp.; St.

37.1: 5 sp.; St. 40:7 sp.; St. 50.2: 1 sp.; St. 52.3: 1 sp.; St.53.2: 2

sp.; St. 61: 17 sp., 6 sh.; St. 63: 3 sp.

5-5213.24 ± 1.49

(30.2)

1.31 ± 0.14

(2.90)7.65 ± 0.75

Entalina tetragona

(Brocchi, 1814)

St.6.1: 12 sp., 4 sh.; St. 6.2: 1 sp.; St. 7: 3 sp., 23 sh.; St. 8: 5 sp.;

St.9: 3 sp., 2 sh.; St. 27: 10 sp.; St.31: 27 sp.; St. 48: 1 sp.; 29

sh., St. 31: 1 sp.; St. 53.1: 5 sp., 12 sh.

96-8757.64 ± 0.38

(11.1)

0.92 ± 0.02

(1.16)8.19 ± 0.28

Pulsellum lofotense

(M. Sars, 1865)

St.13: 2 sp.; St. 27: 1 sp.; St. 29.2: 1 sp.; St. 33.2: 1 sp.; St.43:

1 sp.65-183

2.88 ± 0.39

(4.0)

0.36 ± 0.02

(0.39)7.75 ± 0.76

Cadulus jeff reysi

(Monterosato, 1875)St. 27: 4 sp.; St. 31: 2 sp. 150-183

2.94 ± 0.60

(3.0)- -

Dischides politus

(S. Wood, 1842)

St. 3: 2 sp.; St. 10.1: 19 sp., St. 10.2: 1 sp.; 7 sh.; St. 11: 1 sp.; St.

12: 1 sp.; St. 14: 2 sh.; St. 15: 2 sp.; St. 16: 7 sp., 5 sh.; St. 18: 1

sp.; St. 22: 1 sp.; St. 29.1: 33 sp., 1 sh.,; St. 29.2: 7 sp.; St. 30: 1

sp.; St. 32: 1 sp.; St. 33.1: 3 sp.; St. 33.2: 1 sp.; St. 34: 2 sp.; St.

35: 1 sp., 1 sh.; St. 38: 1 sp.; St. 39: 1 sp.; St. 40: 1 sp.; St. 41: 4

sp., 1 sh.; St. 42: 1 sp.; St. 43: 1 sp.; St. 47: 2 sh., St. 49.3: 2 sp.;

St. 49.4: 2 sp.; St. 50.1: 1 sp., 1 sh.; St. 50.2: 1 sh.

19-1135.11 ± 0.28

(8.8)

0.56 ± 0.02

(0.92)8. 09 ± 0.40

B. ÖZTÜRK

205

Of the identifi ed species, Antalis inaequicostata was the most common in the investigated area and comprised 30% of the total specimens collected, whereas both Pulsellum lofotense and Cadulus jeff reysi were represented by only 6 individuals (Table 2). All of the determined species were found in soft substrata (Table 1). Antalis antalis, A. inaequicostata, A. rossati, A. vulgaris, and Fustiaria rubescens were found at depths up to 100 m. Antalis panorma, P. lofotense, and C. jeff reysi dominated the depths over 100 m (68-200 m). Entalina tetragona had the deepest distribution, being found at depths up to 875 m (Tables 1 and 2).

Family DENTALIIDAE Gray, 1847

Antalis dentalis (Linneaus, 1758)

(Figure 2)

Dentalium dentalis Linnaeus, 1758

Material examined: 11 individuals from 5 stations (Table 2).

Remarks: Th e shell of this species has a longitudinal primary sculpture consisting of about 10 primary ribs (sometimes more: up to 15) in the posterior part. On the shell of some specimens, toward the aperture, between the primary longitudinal ribs, there are secondary ribs, which are mostly in the same thickness range as the primary ones. When secondary ribs are present, they generally disappear before reaching the middle of the shell, which diff erentiates

it from A. inaequicostata. Th ere are also specimens with fi ne striations between the ribs, which are fewer in number (4-6) than in related species. In addition, the shell is more slender and generally more fragile, with a circular aperture and without a truncate apex and calcareous pipe in it (the other diff erence from A. inaequicostata).

Th e mean shell length of the investigated specimens was 19.1 mm, and the mean diameter of the aperture was 2.4 mm (Table 2).

Distribution: Eastern Atlantic and Mediterranean Sea. Depth range: 0-300 m (Steiner and Kabat, 2004: 583).

Antalis inaequicostata (Dautzenberg, 1891)

(Figure 3)

Dentalium inaequicostatum Dautzenberg, 1891

Material examined: 224 individuals and 22 empty shells from 44 stations (Table 2).

Remarks: Th e shell is solid, and the posterior aperture appears more or less truncated, polygonal, and generally has an oval short central pipe in adult specimens (another diff erence between this species and the preceding one). Th ere are 8-11 strong primary longitudinal ribs towards the posterior aperture, which increase in number towards the anterior end, with the intercalation of the secondary and the tertiary ribs, which are of unequal thickness. In the shell of some specimens, all the ribs can disappear

a

B

A

Figure 2. Antalis dentalis: general view of 2 specimens (A = St.

24, 27.2 mm, 49 m; B = St. 29.2, 19.5 mm, 45 m) and

aperture of the shell A (a = 0.7 mm).

Figure 3. Antalis inaequicostata: general view of diff erent shells

(A = St. 39, 33.2 mm, 37 m; B = St. 19.2, 27.8 mm, 46

m; C = St. 19.2, 36.2 mm, 46 m; D = St. 34, 17.9 mm, 79

m) and the apertures of the shells A (a = 3.2 mm) and

B (b = 3.4 mm).

a

b

A

B

D

C


206

towards the aperture. Th e secondary ribs can reach to the posterior half of the shell, but they rarely reach near the apex (Figure 3C). Th ere are also fi ne lines (9-12) between the ribs.

Th e mean length and diameter of the aperture were calculated as 25.6 mm and 2.7 mm, respectively.


Antalis panorma (Chenu, 1843)

(Figure 4)

Dentalium panormum Chenu, 1843

Material: 78 individuals and 22 empty shells from 9 stations (Table 2).

Remarks: Th e sculpture of the shell consists of 10-24 longitudinal ribs in the posterior part near the apex, and then they increase in number with another 10-20 ribs, which become weaker. All of these ribs disappear in the anterior part near the aperture, or transform into thin lines. Th e shell is whitish or yellow.

Th e mean length of the investigated specimens was 28.0 mm, and the mean diameter of the aperture was 2.4 mm. Th e ratio of H/D was 11.4 (Table 2).

Distribution: Eastern Atlantic and the

Mediterranean Sea. Depth range: 54-350 m (Steiner

and Kabat, 2004: 626).

Antalis rossati (Caprotti, 1966)

(Figure 5)

Dentalium rossati Caprotti, 1966

Material: 7 individuals and 6 empty shells from 4

stations (Table 2).

Remarks: Among the dentaliid species occurring

along the Turkish coasts, this species is similar in

general aspect to A. dentalis, A. inaequicostata,

and A. vulgaris. It diff ers from A. dentalis and A.

inaequicostata by its longitudinal ribs barely reaching

the middle part of the shell and clear concentric growth

lines towards the anterior end. As for the diff erence

from A. vulgaris, the well-marked longitudinal ribs

in the posterior end of A. rossati are lower in number

(approximately 10) in contrast to the longitudinal

ribs (28-33) of A. vulgaris. Th e pinkish colour of the

shell of A. rossati is also characteristic.

Th e mean length of the shells and the mean

diameter of the aperture were 24.9 mm and 2.6 mm,

respectively.

a

A

B

a

A

B

Cc

b

Figure 4. Antalis panorma: general view of 2 specimens (A = St.

58.2, 35 mm, 200 m; B = St. 58.3, 40.3 mm, 200 m) and

the aperture of the shell A (a = 2.8 mm).

Figure 5. Antalis rossati: general view of 3 specimens (A = St.17,

32.6 mm, 11.5 m; B = St. 17, 36.0 mm, 11.5 m; C = St.

22, 41.1 mm, 24 m), posterior (a) and anterior (b) parts

under magnifi cation, and aperture of the shell C (c =

3.4 mm).

B. ÖZTÜRK

207

Distribution: Mediterranean Sea. Depth range: 3-49 m (Steiner and Kabat, 2004: 639 and the present study).

Antalis vulgaris (da Costa, 1778)

(Figure 6)

Dentalium vulgare da Costa, 1778


Remarks: Th e shell is striated only posteriorly near the apex, and the number of thin ribs varies from 28 to 33. Th e anterior half of the shell is generally smooth. Some specimens have a central pipe at the apex. Th e shell ranges in colour from whitish, yellowish, to pinkish. Sometimes only the posterior part of the shell, near the apex, is pinkish.

Th e mean height of the studied shells and the mean diameter of the aperture were 21.6 mm and 2.5 mm, respectively (Table 2).

Distribution: Eastern Atlantic and the Mediterranean Sea. Depth range: 5-1000 m (Steiner and Kabat, 2004: 662).

Family FUSTIARIIDAE Steiner, 1991

Fustiaria rubescens (Deshayes, 1825)

(Figure 7)

Dentalium rubescens Deshayes, 1825


Remarks: Among the scaphopod species occurring along the Turkish coast, this species is characterised by its smooth, glossy shell with a short split at the apex (especially in young specimens) (Figure 7a). Th ey range in colour from brownish to greyish-white.

Of the measured specimens, the maximum length was 30.2 mm (Table 2).


Family ENTALINIDAE Chistikov, 1979

Entalina tetragona (Brocchi, 1814)

(Figure 8)

Dentalium tetragonum Brocchi, 1814


Remarks: Th is species is characterised by its angled cross-section. It has 5 primary ribs near the posterior end. Th e shell is whitish.

Th e mean length of the studied shells was 7.6 mm, with a maximum length of 11.1 mm, and the mean of the diameter of the aperture was 0.9 mm (Table 2)

a

A

B

C

D

cb

a

A

B

Figure 6. Antalis vulgaris: general view of diff erent specimens (A

= St. 29.1, 44.8 mm, 6 m; B = St. 55, 37.1 mm, 25 m; C

= St.40, 39.5 mm, 13.5 m; D = St. 53.5, 35.8 mm, 30 m),

posterior (a) and middle (b) parts under magnifi cation,

and aperture of the shell D (c = 3.8 mm).

Figure 7. Fustiaria rubescens: general view of 2 specimens (A = St.

53.2, 27.9 mm, 50 m); B = St. 29.2, 17.6 mm, 30 m) and

the slit at the apex (a) under magnifi cation.


208

Distribution: Eastern Atlantic (European coasts) and the Mediterranean. Depth range: 68-2664 m (Steiner and Kabat, 2004: 655).

Family PULSELLIDAE Scarabino in Boss, 1992

Pulsellum lofotense (M. Sars, 1865)

(Figure 9)

Siphonodentalium lofotense M. Sars, 1865

Material: 6 individuals from 5 stations (Table 2).

Remarks: Th e shell is thin, semitransparent, smooth, and rather fragile. Oblique growth lines are evident.

Th e mean length of the shells and the mean diameter of the aperture were 2.8 mm and 0.4 mm, respectively.

Distribution: Eastern Atlantic (Norway to Spain) and the Mediterranean. Depth range: 26-3500 m (Steiner and Kabat, 2004: 611).

Family GADILIDAE Stoliczka, 1868

Cadulus jeff reysi (Monterosato, 1875)

(Figure 10)

Helonyx jeff reysi Monterosato, 1875

Material: 6 individuals from 2 stations (Table 2).

Remarks: Th is species can be easily recognised by its shell when viewed from diff erent aspects. Th e shell is smooth and shiny with a less evident swelling in the middle of the dorsal side. Th e aperture is slightly laterally compressed, while the apex is slightly dorsoventrally depressed (Figures 10a and 10b).

Th e mean length of the studied 6 shells was 2.9 mm (Table 2).

Distribution: Eastern Atlantic and Mediterranean

Sea. Depth range: 90-2200 m (Steiner and Kabat,

2004: 605).

Dischides politus (S. Wood, 1842)

(Figure 11)

Ditrupa polita S. Wood, 1842

Material: 98 individuals and 21 empty shells from

29 stations (Table 2).

Remarks: Th e species is characterised by its

thin, smooth, and subcylindrical shell, in which the

posterior end is cleft . Th e ventral lobe is somewhat

shorter than the dorsal lobe. Growth lines are the

only shell sculpture. In adults, the anterior part of the

shell narrows at the end.

Th e mean length of the studied shells was 5.1 mm,

with a maximum length of 8.8 mm (Table 2).

Distribution: Eastern Atlantic and the

Mediterranean. Depth range: 9-324 m (Steiner and

Kabat, 2004: 632).

Discussion

In the benthic material sampled from the Turkish

Levantine and Aegean coasts, 10 scaphopod species

were identifi ed versus 16 nominal ones distributed

throughout the Mediterranean. All of the identifi ed

species were distributed along the Turkish Aegean

coast, whereas only 4 species (Antalis inaequicostata,

A. panorma, A. vulgaris, and Fustiaria rubescens)

were encountered along the Turkish Levantine coast.

In the studies previously carried out in the Aegean

Sea (Forbes, 1844; Sturany, 1895; Koukouras and

aA

a b

A

Figure 8. Entalina tetragona: general view of a specimen (A = St.

31, 7.9 mm, 150 m) and cross-section of a shell in the

middle part (a = 0.81 mm).

Figure 9. Pulsellum lofotense: general view of a specimen (A =

St. 13, 3.9 mm, 77 m), aperture (a = 396 μ × 264 μ) and

apex (b = 165 μ).

B. ÖZTÜRK

209

Kevrekidis, 1986; Demir, 2003), 10 species were reported from the area: Antalis agilis (Sars, G. O., 1872), A. dentalis, A. inaequicostata, A. panorma, A. rossati, A. vulgaris, Fustiaria rubescens, Entalina tetragona, Dischides politus, and Episiphon fi lum (Sowerby, G. B. II, 1860). With the exception of A. agilis and E. fi lum, all these species were also found during this study. In addition to those previously recorded from the Aegean Sea, Pulsellum lofotense

and Cadulus jeff reysi were new records for the Aegean molluscan fauna.

Antalis agilis, which was reported from several localities in the Aegean Sea (Koukoras and Kevrekidis, 1986: 190), was not found in this study, which may be due to its wide depth range. Although the known depth range for this species is from 20 to 5000 m (Koukoras and Kevrekidis, 1986: 190; Steiner and Kabat, 2004: 560), it seems that its lower distribution limit is in even deeper waters. Caprotti (1966a) indicated that this species is generally distributed at depths ranging from 200 to 600 m. Th e same reason may account for the absence of E. fi lum, known from depths of between 20 and 4784 m.

Pulsellum lofotense and C. jeff reysi, the 2 new records from the investigated region, were both found at stations located in the Aegean Sea. Pulsellum lofotense was obtained from 5 stations at depths between 77 and 183 m, whereas C. jeff reysi was found only at 2 stations at depths 150 and 183 m, respectively (Table 2). Pulsellum lofotense was previously reported by Koutsoubas et al. (2000) in the Cretan Sea from depths between 500 and 700 m, but as for C. jeff reysii, according to the relevant literature, this record seems to be the fi rst from the eastern Mediterranean. Th e specimens of both species were encountered in soft substrata consisting of sand, mud, or a mixture of both (Table 1).

In the studies carried out along the Turkish Levantine coast (Buzzurro and Greppi, 1996: 19; Çevik and Sarıhan, 2004: 95), 4 scaphopod species (Antalis dentalis, A. inaequicostata, A. vulgaris, and A. rossati) were reported from the area. Of these, A. dentalis, previously reported from this coastline by Çevik and Sarıhan (2004: 95), was not found in this study. Antalis panorma and F. rubescens, which were obtained from various soft substrata down to depths of 200 m, are newly added to the molluscan list of the Turkish Levantine coast, increasing the number of known species to 6.

Dischides politus, occurring in the Mediterranean and eastern Atlantic Ocean (Caprotti, 1979; Steiner and Kabat, 2004), was not encountered within the samples collected along the Levantine coast, although it is widespread along the Aegean coast. Its southern distribution limit was station 50, close to the border between the Aegean and Levantine coasts.

a

b

A

A

C

B

Figure 10. Cadulus jeff reysi: general view of a specimen (A = St.

31, 2.9 mm, 150 m), aperture (a = 350 μ × 325 μ) and

apex (b = 300 μ × 275 μ).

Figure 11. Dischides politus: general view of 3 specimens (St.

49.3, A = 6.6 mm, B = 5.3 mm, 54 m; C = St. 11, 3.7

mm, 27 m).


210

Th is species was recently reported from a nearby area (İztuzu), by Aartsen and Kinzelbach (1990), providing the fi rst, and so far only, record of the species from the Turkish coast.

Antalis rossati, one of the rarely encountered scaphopod species of the Turkish coast, and previously reported from İskenderun Bay and the Cypriot coast (Levantine Sea) (Öztürk et al., 2003; Çevik and Sarıhan, 2004), was found only at 4 Aegean stations, mostly in muddy bottoms at depths of between 11.5 and 49 m (Tables 1 and 2). Antalis rossati, which was described by Caprotti (1966b) from the Israeli coast and was previously considered to be an eastern Mediterranean species, was also recorded recently from the Spanish coast by Alzuria (1986).

According to Sturany (1895), Marion (1898), and Demir (2003), Antalis dentalis, A. inaequicostata, A. panorma, and A. vulgaris are also present in the Sea of Marmara. Marion (1898: 169) also reported F. rubescens from the Sea of Marmara, which is the only record of this species to date. However, the reports of Antalis entalis (Linneaus, 1758) from the Sea of Marmara (Tortonese, 1959) and the entrance to the Dardanelles (Colombo, 1885) are doubtful records, considering that the known distribution of this

species is the Atlantic Ocean (Caprotti, 1979: 232; Steiner and Kabat, 2004: 588). No scaphopod species has ever been reported from the Turkish coast of the Black Sea. Th is might be due to the great change in salinity (18‰) and some other features of Black Sea waters, compared to the Aegean Sea and the Sea of Marmara.

In conclusion, the Aegean Sea contained the highest number of scaphopod species compared with the adjacent seas, a fact which might be related to great diff erences in the ecological features such as temperature, salinity, habitats, and nutrients in the Aegean Sea.

Acknowledgements

Th anks are due to Sigurd Boletzky for polishing the English text. Th e constructive comments of the anonymous referees are much appreciated.

Th is work has been partially supported by TÜBİTAK (Project Number: 104 Y 065) and the Environmental Protection Agency for Special Areas (Republic of Turkey Ministry of Environment and Forestry).

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Scaphopod species (Mollusca) of the Turkish Levantine and Aegean