Journal of Biological Research-Thessaloniki 14: 161 – 179, 2010 J. Biol. Res.-Thessalon. is available online at http://www.jbr.gr Indexed in: WoS (Web of Science, ISI Thomson), SCOPUS, CAS (Chemical Abstracts Service) and DOAJ (Directory of Open Access Journals)

The of Thessaloniki & Thermaikos Gulfs (North Aegean Sea, ) with emphasis on new for Hellenic waters

Thanasis MANOUSIS1, George MPARDAKIS2, Constantinos PARASKEVOPOULOS2 and Sofia GALINOU-MITSOUDI2* 1 P.O. Box 48K, Epanomi 575 00, Greece 2 Alexander Technological Educational Institute of Thessaloniki, Department of Fisheries & Aquaculture Technology, 632 00 Nea Moudania, Chalkidiki, Greece

Received: 9 July 2009 Accepted after revision: 31 March 2010

The marine molluscan bivalve fauna of Thessaloniki Gulf (Thessaloniki Bay included) and Ther- maikos Gulf was investigated and collected by diving and trawling during the period from March 2005 to October 2008, after three decades from the completion of the previous relevant study. 188 species belonging to 48 families were identified and their biodiversity was compared with the current checklists of marine molluscs for Northern Aegean Sea and Hellenic seas based on pre- vious surveys. In this bivalve collection nine species are new for the Greek fauna, 18 are new for N Aegean Sea and six of them represent lessepsian migrants from the Red Sea (all of them new for the Hellenic waters). The occurrence of new species for the study area and Hellenic seas is discussed.

Key words: molluscs, marine bivalves, invasive species, Thermaikos Gulf, North Aegean Sea.

INTRODUCTION Thessaloniki Gulf (Thessaloniki Bay included) and Thermaikos Gulf (NW Aegean Sea) are referred The comprises less than 1% in generally as Thermaikos Gulf by numerous papers surface area and volume as compared with the world but yet includes a total of around 8500 species and comprise one of the most complicated and mul- of macroscopic organisms, which indicates a rich bio- ti-used ecosystems of the E Mediterranean Sea since diversity (Bianchi & Morri, 2000). This could be ex- i) it receives an input of nutrient-rich freshwater from plained by its geological history (Maldonado, 1985; five rivers and some protected wetlands (Ramsar Ruffel, 1997) and the conspicuous ecological and hy- convention and Natura 2000 network) resulting in drological differences in various localities (Sará, very high productivity and biodiversity of its coastal 1985). Moreover, biodiversity, mainly in ∂ Mediter- waters, ii) it is a main navigation line leading to the ranean Sea –close to the Suez Canal– and the Strait city of Thessaloniki and iii) it is affected by human of Gibraltar, has undergone modifications during re- activities in all its coastal zone (i.e. urbanization, agri- cent decades, following the introduction of non-in- culture, fisheries, mussel farming and recreation). digenous species, attributed to the expansion of ship- Currently, Thermaikos and Thessaloniki Gulfs are ping traffic, aquaculture activities and migration (Por, considered as a sensitive ecosystem (and rather is), 1978, 1990; Galil & Zenetos, 2002; Streftaris et al., according to the term “sensitive” that has been preci- 2005). sely defined by Hiscock & Tyler-Walters (2006) and Tyler-Walters et al. (2009). For such an ecosystem, al- * Corresponding author: tel.: +30 23730 26457, fax: +30 though there are numerous environmental investiga- 23730 26450, e-mail: [email protected] tions, only a few are referred to its fauna and even to

161 162 Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs its bivalve molluscs exclusively, i.e. Sakellariou (1957), MATERIALS AND METHODS Zenetos (1996) and Zenetos et al. (2005). Research Sampling of live, mainly, specimens was conducted in over the last 50 years on the benthic fauna of Ther- the coastal areas of Thessaloniki Bay, Thessaloniki maikos gulf has revealed the presence of several bi- Gulf and Thermaikos Gulf (NW Aegean sea, Gree- valve species (Zarkanellas, 1980; Antoniadou et al., ce) (Fig. 1) during the period from 2005 to 2008. The 2004). samples were collected by a) a van Veen type grab of The main goals of this study are: a) to contribute a 1000 cm2 sampled surface (all stations), b) a profes- to the knowledge of bivalve biodiversity with an up- sional (traditional) shell fisheries dredge (130 cm wi- dated catalogue of bivalve species of Thermaikos de, nylon drag net of 2.2 cm mesh) (stations C, D, E, Gulf, b) to compare the present biological diversity of H, I), c) a professional trawl (station area S) and d) the area with that recorded in publications of the past snorkeling (stations K, L, M, N). Species were colle- and c) to improve our knowledge on the distribution cted by sieving the soft substrate through a 5 mm, a 2 and expansion of invasive bivalve species. This pur- mm and a 0.5 mm mesh sieve. After rinsing with fresh pose was facilitated by the use of i) several books, at- water, shells were treated with 25% glycerin in etha- lases and on-line services, such as Oliver (1992), Pop- nol and stored at room temperature. All species col- pe & Goto (1993), Giannuzzi-Savelli et al. (2001), Ca- lected were accompanied by data regarding synonyms, chia et al. (2004), Doneddu & Trainito (2005), Repet- shell sizes, habitat characters and collection depths and to et al. (2005), Delamotte & Vardala-Theodorou dates. The nomenclature followed in this study was (2008), ELNAIS (https://services.ath.hcmr.gr/spec) and that of European Register of Marine Species (ERMS) Natuurhistorisch Museum Rotterdam (www.nmr-pics.nl), (www.marbef.org) and the species recognition was based ii) faunistic and review articles (i.e. Salas, 1996; De- on systematic guides and atlases listed in References. mir, 2003; Gofas & Salas, 2008) and iii) relevant stud- For the species nomenclature update (30 November ies of the Mollusca fauna in Hellenic areas (i.e. Zene- 2009) beside the ERMS, the CLEMAM on-line Da- tos et al., 2005, 2007, 2009a; Ovalis & Zenetos, 2007). tabase was used. The specimens are deposited in the

FIG. 1. Map of the study area: 1. Thessa- loniki Bay (stations F and G); 2. Thessa- loniki Gulf (stations D, E, H and I); 3. Thermaikos Gulf (stations A-C, J-R and S). Collection stations: A. Scala Litocho- rou; B. Kitros; C. Axios River Estuaries; D. Chalastra; E. Naziki; F. Paliomana; G. Micro Emvolo; H. MACEDONIA Airport; I. Ajia Triada; J. Aggelohori; K. Palioura; L. Paralia; M. Cape of Epanomi; N. Pota- mos; O. Nea Kallikratia; P. Nea Mouda- nia; Q. Sani; R. Poseidi; S. Trawled area. w. Wetlands. Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs 163

S. (M) 30-40

R. (S) 4-10 . Naziki; E Q. (SM) 5-10 . Nea Mouda- P

P. (SM) 4-10 + . Chalastra;

D O. (SM) 6-11

N. (S) 2-20 . Nea Kallikratia; O S + + M. (SM) 2-10 +++ and + L. (SM) 3-8 ++ + R - J , . Potamos; K. (SM) 3-10 + + ++ + C N - A . Axios River Estuaries; C

J. (SM) 8-12 ++ + + + ++ I. (SM) 5-10 ++ + . Kitros; B ++++ + +++ ++ + H. (M) 5-13 ++++ . Cape of Epanomi;

G. (M) 3-10 +++ + + ++ M

F. (M) 8-10 +++ + ++ + + ++++ ): sandy and muddy. *indicates new species for N Aegean Sea **new ; Thermaikos Gulf: stations I SM . Paralia; E. (M) 4-8 + +++++ + L . Scala Litochorou; and A +++ ++ +

H D. (SM) 5-12 , E ): Sandy; ( ,

S C. (SM) 5-10 D . Palioura; K B. (SM) 8-15

A. (S) 6-12 ): Muddy; ( M . Aggelohori; J STATION Depth (m) ; Thessaloniki Gulf: stations . Ajia Triada; G I and F (SUBSTRATE TYPE) (Lamarck 1819) (Reeve 1858) (Poli 1795) (Linné 1758) (Philippi 1844) (Linné 1758) (Cartaine 1835) (Scacchi 1832) (Lamarck 1819) (Müler 1776) (Say 1822) (Defrance 1816) (Linné 1767) (Linné 1758) Carrozza 1987 Winckworth 1930 (Linné 1758) (Poli 1795) Bronn 1831 Mayer 1868) Poli 1795 (Linné 1758) ( Linné 1758 . MACEDONIA Airport; H . Trawled area. The substrate type is indicated by ( Nuculana pella S Nucula nucleus Nucula sulcata Lithophaga lithophaga Modiolarca subpicta Modiolula phaseolina Solemya togata Nucula nitidosa ** Anadara polii Striarca lactea Glycymeris bimaculata Gregariella petagnae Modiolus adriaticus Nuculana illirica Anadara transversa Arca noae Arca tetragona Barbatia clathrata Glycymeris glycymeris Gregariella semigranata Asperarca nodulosa Barbatia barbata Glycymeris violacescens . Micro Emvolo; . Poseidi; G R . Sani; Q . Paliomana; nia; F species for Hellenic waters. Thessaloniki Bay: stations TABLE 1. Bivalvia species in a taxonomic order and their occurrence the collection stations: FAMILYSOLEMYIDAE NUCULIDAE Species NUCULANIDAE ARCIDAE NOETIIDAE GLYCYMERIDIDAE 164 Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs

S. (M) 30-40 +

R. (S) 4-10

Q. (SM) 5-10 + + P. (SM) 4-10 +

O. (SM) 6-11

N. (S) 2-20 + M. (SM) 2-10 +

L. (SM) 3-8 +

K. (SM) 3-10 ++ ++ + + + +

J. (SM) 8-12 ++ + ++ + I. (SM) 5-10 ++

H. (M) 5-13 +++

G. (M) 3-10 ++ + +

F. (M) 8-10

E. (M) 4-8 ++++++ + ++ ++++ + ++ +++++ + +++ ++ D. (SM) 5-12 ++++++++++ +

C. (SM) 5-10

B. (SM) 8-15 ++++++ ++++++++++

A. (S) 6-12 ++ +++++++++++ + + STATION Depth (m) (SUBSTRATE TYPE) (Poli 1795) (Risso 1826) (Poli 1795) Lamarck 1819 (Linné 1758) (Gmelin 1791) (Poli 1795) (Risso 1826) (Locard 1889) (Gmelin 1791) (Laskey 1811) (Poli 1795) (Linné 1758) (Poli 1795) (Linné 1767) (Leach 1814) (Dillwyn 1817) (Linné 1758) (Linné 1767) (Linné 1758) (Linné 1758) (Linné 1758) (Linné 1758) Linné 1758 Linné 1758 lineatus Mytilaster marioni Lissopecten hyalinus Pecten jacobeus Pseudamussium clavatum Palliolum incomparabile Delectopecten vitreus Mytilus galloprovincialis Pinna nobilis Crassadoma multistriata Chlamys varia Mytilus edulis Atrina pectinata Pinctada radiata Pteria hirundo Musculus discors Musculus costulatus * * Mytilaster minimus Modiolus barbatus Similipecten similis Chlamys proteus Aequipecten opercularis Chlamys glabra Chlamys pesfelis Chlamys flexuosa PINNIDAE PTERIIDAE PECTINIDAE TABLE 1. Continued FAMILY Species Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs 165

S. (M) 30-40

R. (S) 4-10

Q. (SM) 5-10

P. (SM) 4-10

O. (SM) 6-11 +

N. (S) 2-20

M. (SM) 2-10 + + L. (SM) 3-8 + +

K. (SM) 3-10 ++ + + +

J. (SM) 8-12

I. (SM) 5-10 ++ + +++ + + ++

H. (M) 5-13 ++ ++ ++ ++ G. (M) 3-10 ++

F. (M) 8-10 +++ +++ + ++++ ++ E. (M) 4-8 ++++++ ++++ + + +

D. (SM) 5-12 ++ +++++++ + + ++ ++++

C. (SM) 5-10

B. (SM) 8-15

A. (S) 6-12 STATION Depth (m) l 1775) å (SUBSTRATE TYPE) (Linné 1761) (Poli 1795) (Linné 1758) (Müller 1776) Linné 1758 (Forssk (Montagu 1803) (Deshayes 1850) (Olivi 1792) (Linné 1758) Linné 1758 (Linné 1767) (Jeffreys 1881) (Philippi 1836) van Aartsen 2004 (Payraudeau 1826) (Sowerby G.B. I 1823) (Costa O.G. 1829) (Montagu 1803) (Linné 1758) (Gmelin 1791) Linné 1758 (Linné 1758) Diplodonta globosa Diplodonta bogii Pododesmus patelliformis Lima lima Diplodonta rotundata Anomia ephippium Spondylus gaederopus Heteranomia squamula Pododesmus aculeatus Limaria hians Limaria tuberculata Limea loscombii Ostrea edulis Ostreola stentina Neopycnodonte cochlear Anodontia fragilis Loripes lacteus Myrtea spinifera Ctena decussata Lucinella divaricata Lucinoma borealis Thyasira subovata Diplodonta brocchii ** LIMIDAE TABLE 1. Continued FAMILYSPONDYLIDAE Species OSTREIDAE GRYPHAEIDAE LUCINIDAE THYASIRIDAE UNGULINIDAE ** 166 Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs

S. (M) 30-40

R. (S) 4-10

Q. (SM) 5-10

P. (SM) 4-10

O. (SM) 6-11

N. (S) 2-20 + + + M. (SM) 2-10 + ++ L. (SM) 3-8 +

K. (SM) 3-10 ++ ++ + +

J. (SM) 8-12 ++ ++ +++ + + I. (SM) 5-10 +++ ++++

H. (M) 5-13 +++ +++++ + + + G. (M) 3-10 + ++

F. (M) 8-10 ++ + + ++ ++ + E. (M) 4-8 +++ + +++ + +

D. (SM) 5-12

C. (SM) 5-10

B. (SM) 8-15

A. (S) 6-12 STATION Depth (m) (SUBSTRATE TYPE) (Jonas 1846) (Lamarck 1819) (Linné 1758) (Montagu 1803) (Montagu 1808) (Turton 1822) (Sturany 1899) Sowerby in Turton 1825 Linné 1758 (Montagu 1803) Broderip 1834 (Linné 1758) (Clark W. 1852) (Philippi 1836) (Linné 1758) Lamarck 1819 Reeve 1846 (Lovén 1846) (Poli 1795) (Linné 1767) (Costa O.G. 1829) pacifica Cardites akabana corbieri Chama aspersa * ** Chama asperella Digitaria digitaria * Hemilepton nitidum Glans aculeata Galeomma turtoni Kellia suborbicularis Epilepton clarkiae Kurtiella bidentata Tellimya ferruginosa Bornia sebetia Chama gryphoides Scacchia oblonga Glans trapezia ** Cardita calyculata Tellimya tenella Venericardia antiquata * GALEOMMATIDAE MONTACUTIDAE ASTARTIDAE KELLIIDAE CARDITIDAE TABLE 1. Continued FAMILY Species Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs 167

S. (M) 30-40

R. (S) 4-10

Q. (SM) 5-10

P. (SM) 4-10

O. (SM) 6-11 +

N. (S) 2-20 ++

M. (SM) 2-10 ++ ++ + L. (SM) 3-8 +

K. (SM) 3-10 + ++++ + + + +++ ++ +++++ + ++ ++ + ++ J. (SM) 8-12 +++++

I. (SM) 5-10 ++++ + ++ ++ +++++++++++++ + + ++ + ++

H. (M) 5-13 ++++++++ ++

G. (M) 3-10

F. (M) 8-10

E. (M) 4-8 ++++

D. (SM) 5-12

C. (SM) 5-10 +

B. (SM) 8-15

A. (S) 6-12 STATION Depth (m) (Sowerby G.B.II 1841) (SUBSTRATE TYPE) (Linné 1758) (Poli 1795) Cecalupo & Quadri 1996 (Gmelin 1791) (Philippi 1836) (Bucquoy, Dautzenberg (Linné 1758) (Linné 1758) (Locard 1899) (Poiret 1789) (Lightfoot in Solander 1786) (Gmelin 1791) (Philippi 1844) (Gmelin 1791) (Linné 1758) (da Costa 1778) (Sowerby G.B. II 1840) (Pennant 1777) (Linné 1758) Pulteney 1799 (Poli 1795) Von Born 1778 (Chenu 1843) (Linné 1758) Monterosatus primus Parvicardium scriptum Parvicardium trapezium Ensis minor glauca Spisula subtruncata * & Dollfus 1892) * Plagiocardium papillosum Solen marginatus Ensis ensis * ** Parvicardium ovale Parvicardium scabrum Parvicardium exiguum Parvicardium minimum Laevicardium oblongum Acanthocardia aculeata Laevicardium crassum Donacilla cornea Phaxas pellucidus Acanthocardia echinata Acanthocardia spinosa Cerastoderma edule Cerastoderma glaucum Acanthocardia paucicostata Acanthocardia tuberculata SOLENIDAE PHARIDAE TABLE 1. Continued FAMILYCARDIIDAE Species MESODESMATIDAE 168 Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs

S. (M) 30-40

R. (S) 4-10

Q. (SM) 5-10

P. (SM) 4-10

O. (SM) 6-11

N. (S) 2-20 + + + + M. (SM) 2-10 +

L. (SM) 3-8

K. (SM) 3-10 ++++ +++ + ++++ + + + + + ++ +++++ +++ + + ++ +++ + ++ J. (SM) 8-12 + +++++++++++++++ + + + +++ + + I. (SM) 5-10 +++++++ +

H. (M) 5-13 ++

G. (M) 3-10

F. (M) 8-10

E. (M) 4-8 ++ ++++ + + ++

D. (SM) 5-12

C. (SM) 5-10 ++

B. (SM) 8-15 +++++++++

A. (S) 6-12 + ++++++ STATION Depth (m) (SUBSTRATE TYPE) (Payraudeau 1826) (Linné 1758) Poli 1795 (Gmelin 1791) (Costa O.G. 1829) Linné 1758 Linné 1758 (Pennant 1777) Lamarck 1818 Linné 1758 (Linné 1758) Lovén 1846 (Scacchi 1834) (Montagu 1808) Poli 1795 Linné 1758 Poli 1791 Brocchi 1814 (Turton 1822) Gmelin 1791 (Pennant 1777) (Lamarck 1818) (Gmelin 1791) da Costa 1778 Poli 1791 (Müller O.F. 1776) (Wood W. 1802) Tellina donacina Tellina distorta Tellina fabula Tellina incarnata Tellina nitida Tellina planata Scrobicularia cottardi Abra prismatica Abra alba Macoma cumana Arcopagia crassa Gastrana fragilis Tellina pulchella Tellina pygmaea Arcopagia balaustina Abra longicallus Donax venustus Tellina serrata Tellina tenuis Capsella variegata Donax semistriatus Donax trunculus Abra nitida Gari costulata Gari depressa Gari fervensis Gari tellinella SEMELIDAE TELLINIDAE SCROBICULARIIDAE DONACIDAE PSAMMOBIIDAE FAMILY Species TABLE 1. Continued Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs 169

S. (M) 30-40 +

R. (S) 4-10 +

Q. (SM) 5-10

P. (SM) 4-10

O. (SM) 6-11

N. (S) 2-20 + M. (SM) 2-10 +

L. (SM) 3-8

K. (SM) 3-10 ++ ++ ++ + ++ +++ + ++ + + ++ +++ + ++ + + + J. (SM) 8-12 ++ ++ ++

I. (SM) 5-10 ++++ ++++++++ + + + +++++ +++ +++++ + ++ + +

H. (M) 5-13 ++++++ ++ ++ G. (M) 3-10 +

F. (M) 8-10 + +++++ + +++++++ E. (M) 4-8 +++++++++++

D. (SM) 5-12

C. (SM) 5-10

B. (SM) 8-15

A. (S) 6-12 STATION Depth (m) (SUBSTRATE TYPE) (Dautzenberg (Gmelin 1791) (da Costa 1778) (Linné 1758) (Linné 1758) (Born 1778) Roemer 1860 (da Costa 1778) (Pennant 1777) (da Costa 1778) (Forbes 1844) (Turton 1822) (Tiberi 1855) (Linné 1758) (Linné 1758) (Montagu 1803) Linné 1758 (Linné 1758) (Pennant 1777) (Linné 1758) (Linné 1758) (Locard 1886) (Gmelin 1791) Linné 1758 (Poli 1795) (Linné 1758) Dosinia erythraea punctigera Circenita calipyga Pitar rudis Ruditapes decussatus Timoclea ovata casina Venerupis senegalensis Paphia lucens Paphia rhomboides Pitar mediterranea Gouldia minima Paphia aurea & Fischer H. 1906) ** Dosinia exoleta Dosinia lupinus Clausinella fasciata ** Irus irus Chamelea striatula ** Chamelea gallina Solecurtus strigilatus humanus Callista chione Solecurtus scopula Azorinus chamasolen Venus verrucosa Kelliella abyssicola TABLE 1. Continued FAMILYSOLECURTIDAE Species KELLIELLIDAE 170 Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs

S. (M) 30-40

R. (S) 4-10 +

Q. (SM) 5-10

P. (SM) 4-10

O. (SM) 6-11

N. (S) 2-20 + + +++ + M. (SM) 2-10 +

L. (SM) 3-8 +

K. (SM) 3-10 ++ + + J. (SM) 8-12 ++ + ++ + +

I. (SM) 5-10 +++

H. (M) 5-13 +++

G. (M) 3-10

F. (M) 8-10 +++ + + +

E. (M) 4-8 ++ +++++ +++++++ + +

D. (SM) 5-12

C. (SM) 5-10

B. (SM) 8-15

A. (S) 6-12 +++++++++ STATION Depth (m) (Costa O.G. 1829) Spengler 1783 (Linné 1758) (SUBSTRATE TYPE) (Olivi 1792) (MacGillivray 1827) (Pennant 1777) (Philippson 1788) Deshayes 1830 (Payraudeau 1826) (Poli 1791) (Pulteney 1799) (Montagu 1803) (Linné 1758) Linné 1758 (Linné 1767) (Linné 1767) Linné 1758 (Olivi 1792) Gastrochaena cymbium Barnea candida Pholas dactylus Teredo navalis corbuloides Thracia distorta Thracia papyracea Petricola lajonkairii Hiatella rugosa Cuspidaria cuspidata Hiatella arctica Thracia pubescens Petricola lithophaga Gastrochaena dubia Thracia villosiuscula Corbula gibba Lentidium mediterraneum * Pandora inaequivalvis PHOLADIDAE TEREDINIDAE PETRICOLIDAE CUSPIDARIIDAE HIATELLIDAE GASTROCHAENIDAE CORBULIDAE PANDORIDAE TABLE 1. Continued FAMILY Species Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs 171 premises of the Alexander Technological Educational collected from sandy-muddy bottoms of the sub- Institute of Thessaloniki and in the premises of Dr. T. littoral zone of S Thessaloniki Gulf and E Ther- Manousis. Scientists are welcome to have access to maikos Gulf. A total of seven live specimens and the biological material at will. two shells were collected with their lengths rang- ing from 1.6 mm to 7.6 mm (some bearing their RESULTS ) (Fig. 2E). 5. MESODESMATIDAE. Two live specimens of As a result of this investigation, approximately 10000 the very characteristic species Monterosatus pri- specimens were collected from variable substrates mus (Locard 1899) (Fig. 2F) was collected from such as sandy (stations N and R), muddy (station S) sandy-muddy bottom of the littoral zone of E and sandy and muddy in all the rest of the stations Thermaikos (3.2 mm and 4.4 mm in length, respe- (Fig. 1). A total of 188 species belonging to 48 fami- ctively). lies were identified (Table 1). Among the 48 families, 6. VENERIDAE. Three species of this family were the most species-rich was VENERIDAE with 21 spe- collected from sandy-muddy bottoms of the sub- cies followed by CARDIIDAE (16 species), TELLI- NIDAE and MYTILIDAE (14 species each) and PE- littoral zone from the east coasts of the study a- CTINIDAE (13 species). The number of bivalve spe- rea: Circenita calipyga (Born 1778) (one shell 8.6 cies that are reported for the first time from the study mm in length) (Fig. 3A), Clausinella punctigera area is approximately 100, increasing the number of (Dautzenberg & Fischer H. 1906) (Fig. 3B) (two species in a percentage close to 100%. Among the live specimens, 7.3 mm and 10 mm in length, res- identified species, nine are referred for the first time pectively), and Dosinia erythraea Roemer 1860 (Fig. to the Hellenic fauna while these nine species together 3C) (two live specimens with length of 35 mm and with nine more species are referred for the first time 44 mm, respectively). to the N Aegean Sea. In the study area, six invasive species originated from the Red Sea and three of W New records for N Aegean Sea Mediterranean Sea or Atlantic origin are recognized. Apart from the nine new species for the Greek waters, The majority of the new species for the Hellenic wa- nine more bivalve species are referred for the first ti- ters is endobenthic of soft substrates and one, Gastro- me for N Aegean. chaena cymbium, of calcareous substrates like empty 1. MYTILIDAE. Two species of Mytilidae were col- bivalve or gastropod shells. lected live from sandy-muddy bottoms of the sub- The new species (for Hellenic fauna and N Aegean littoral zone of E Thermaikos: Mytilaster lineatus Sea), their origin and mode of life are presented in (Gmelin 1791) (Fig. 3D) (10 live specimens with Table 2. lengths ranging from 3.3 mm to 5.0 mm) and Myti- laster marioni (Locard 1889) (Fig. 3E) (one live New records for Hellenic waters specimen, 4.9 mm in length). Basic eco-geographical information for the new re- 2. CHAMIDAE. Two species of this family were cords per family in the Hellenic seas is given below: collected from rocky bottoms (live specimens): 1. NUCULANIDAE. Only one shell of the species Chama asperella Lamarck 1819 (Fig. 4A) (four li- Nuculana illirica Carrozza 1987 was found in the ve specimens with length ranging from 8.2 mm to littoral zone of E Thermaikos in sandy-muddy 17.6 mm), Chama aspersa Reeve 1846 (Fig. 4B) bottom, 6.8 mm in length (Fig. 2A). (two live specimens 15 mm and 26 mm in length, 2. UNGULINIDAE. Two species of this family were respectively) and Pseudochama corbieri (Jonas collected: Diplodonta bogii van Aartsen 2004 (Fig. 1846) (Fig. 4C) (two live specimens of 13.1 mm 2B) (one live specimen of 7 mm) and Diplodonta and 19.1 mm, respectively). globosa (Forsskål 1775) (Fig. 2C) (one live speci- 3. CARDIIDAE. Two small species of this family men of 14 mm) in sandy-muddy bottom. were collected: Parvicardium scriptum (Bucquoy, 3. CHAMIDAE. One species of this family was col- Dautzenberg & Dollfus 1892) (Fig. 4D) (three live lected live from rocky bottom: Chama pacifica specimens ranging from 2.0 to 3.9 mm in length), Broderip 1834 (Fig. 2D) (one specimen of 6.7 mm and Parvicardium trapezium Cecalupo & Quadri in length). 1996 (Fig. 4E) (four live specimens with lengths 4. CARDITIDAE. Cardites akabana (Sturany 1899), ranging from 2.0 mm to 6.2 mm). 172 Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs . ; L G and F . Palioura; K . Aggelohori; J . Ajia Triada; I endobenthic epibenthic epibenthic endobenthic feeder - endobenthic epibenthic epibenthic endobenthic epibenthic epibenthic Mode of life (Todd, 2001) Found Origin ). Sandy and muddy. Thessaloniki Bay: station

SM R. (S) 4-10 O. (SM) 6-11 M. (SM) 2-10 ). Sandy, ( . MACEDONIA Airport; S L. (SM) 3-8 H K. (SM) 3-10 J. (SM) 8-12 ). Muddy, (

I. (SM) 5-10 + + Suspension feeder - alive Rare M

Micro Emvolo; H. (M) 5-13

G. G. (M) 3-10 F. (M) 8-10 . Paliomana; F Station R Depth (m) and O (Subtrate type) , M - J l 1775) + Suspension feeder - alive Red Sea å (Bucquoy, + + Suspension feeder - alive South Aegean (Jonas 1846) + + Suspension feeder - alive South Aegean . Poseidi. The substrate type is indicated by ( (Forssk R (Sturany 1899) + + Suspension feeder - alive Red Sea (Locard 1889) + Suspension feeder - alive Red Sea (Gmelin 1791) + Suspension feeder - alive South Aegean Carrozza 1987 + Subsurface deposit shell Adriatic Sea van Aartsen 2004 + Suspension feeder - alive Red Sea Broderip 1834 + Suspension feeder - alive Red Sea Lamarck 1819 + + + + Suspension feeder - alive Red Sea Reeve 1846 + Suspension feeder - alive Red Sea ; Thermaikos Gulf: stations I . Nea Kallikratia; , O Nuculana illirica Diplodonta bogii Cardites akabana Diplodonta globosa Chama pacifica H Pseudochama corbieri Mytilaster lineatus Chama asperella Parvicardium scriptum Parvicardium trapezium Mytilaster marioni Chama aspersa * ** * ** * ** * Dautzenberg & Dollfus 1892)* Cecalupo & Quadri 1996 endobenthic endobenthic * ** * ** . Cape of Epanomi; M Paralia; TABLE 2. Bivalvia species new for Greek waters** and N Aegean Sea*. Gulf of Thessaloniki: stations NUCULANIDAE MYTILIDAE UNGULINIDAE CHAMIDAE CARDITIDAE CARDIIDAE FAMILY Species Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs 173 endobenthic endobenthic terranean endobenthic endolithic in calcareous Mode of life (Todd, 2001) Found Origin substrates endobenthic

R. (S) 4-10 O. (SM) 6-11 M. (SM) 2-10 L. (SM) 3-8 K. (SM) 3-10 J. (SM) 8-12 I. (SM) 5-10 H. (M) 5-13

G. (M) 3-10 + + Suspension feeder - alive Atlantic F. (M) 8-10 Station Depth (m) (Subtrate type) Spengler 1783 + + + + + + Suspension feeder - alive South Aegean (Locard 1899) + Suspension feeder - alive West Medi- (Born 1778) + Suspension feeder - shell Red Sea Roemer 1860 + Suspension feeder - shell Red Sea (Chenu 1843) + Suspension Feeder - alive South Aegean Dosinia erythraea Clausinella punctigera Monterosatus primus Circenita calipyga Ensis minor Gastrochaena cymbium (Dautzenberg & Fischer H. 1906)** endobenthic ** ** * ** * TABLE 2. Continued MESODESMATIDAE PHARIDAE VENERIDAE GASTROCHAENIDAE FAMILY Species 174 Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs

FIG. 2. New records for Hellenic waters: A. Nuculana illirica.B.Diplodonta bogii.C. Diplodonta globosa.D. Chama pacifica. E. Cardites akabana. F. Monterosatus primus. Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs 175

FIG. 3. New records for Hellenic waters (A-C) and N Aegean Sea (D-E): A. Circenita calipyga. B. Clausinella punctigera.C. Dosinia erythraea. D. Mytilaster lineatus. E. Mytilaster marioni. 176 Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs

FIG. 4. New records for N Aegean Sea: A. Chama asperella. B. Chama aspersa. C. Pseudochama corbieri. D. Parvicardium scriptum. E. Parvicardium trapezium. F. Ensis minor. G. Gastrochaena cymbium. Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs 177

4. PHARIDAE. One specimen of Ensis minor (Che- The presence of the non-native Cardites akabana nu 1843), 121 mm in length (Fig. 4F) was collected in the Mediterranean waters was first reported by C¸ e- live from the littoral zone of E Thermaikos. viker & Albaryak (2006), while that of Pseudochama 5. GASTROCHAENIDAE. From six stations of the corbieri in the S Aegean Sea has been reported by Ze- sublittoral zone of E Thermaikos Gulf, Thessalo- netos et al. (2003). niki Gulf and Thessaloniki Bay, more than 30 live The occurrence of Claucinella punctigera (two live specimens of Gastrochaena cymbium Spengler 1783, specimens in Thessaloniki Bay and Thermaikos Gulf) 5.4-5.7 mm in length, were dissected out of their is reported for the first time from E Mediterranean calcareous capsules (Fig. 4G). and has already been referred from W Mediterra- nean Sea (Repetto et al., 2005). The gap of this spe- DISCUSSION cies distribution between W and E Mediterranean The last list of bivalves included 96 species from Ther- could be attributed either to its rather recent transfer maikos Gulf (Zenetos, 1996). The present survey has with ships ballasts (for similar cases, see also Zenetos almost doubled the number of the known species et al., 2008, 2009b) or due to its rarity. from that area and presents new data on the biodiver- The invasion progress of an alien species may be sity of Thermaikos Gulf. At the same time, it reveals attributed to its life strategy and history, hydrodyna- the presence of 18 new species from the N Aegean mics (currents), geo-climatic status and changes and Sea and, among them, nine new species for Greece anthropogenic activities, although certain mediated (Table 2). This means that approximately 3% of the parameters for invasion success of bivalves, such as registered species in this work is not included in the the body size, the growth rate and fecundity (Roy et latest checklists for Hellenic bivalves (Zenetos et al., al., 2002), may have contributed to their expansion. 2005, 2007). Six new species for Greece, namely Di- For eight out of the nine new records for Hellenic plodonta bogii van Aartsen 2004, Diplodonta globosa seas, two species have previously been found in W (Forsskål 1775), Chama pacifica Broderip 1834, Car- Mediterranean Sea or E Atlantic Ocean and six in dites akabana (Sturany 1899), Circenita calipyga (Born the Red Sea; these seas represent the major pathways 1778) and Dosinia erythraea Roemer 1860, are new for the alien bivalves to the Greek waters due to a) lessepsian migrants in Hellenic waters raising the num- ships ballasts, b) northwards currents in Suez Gulf ber of marine alien species from 14 (ELNAIS, Zene- (Rady et al., 1998) and c) water entrance from the At- tos et al., 2009a) to 20. Except of D. globosa, the other lantic Ocean in the Mediterranean Sea, distributing five species from the Red Sea have been already re- larvae to the E and W Mediterranean, respectively corded from the coasts of Levantine Sea (Zenetos et (Shefer et al., 2004). Besides the ships ballasts, Pan- al., 2003; C¸ inar et al., 2005; C¸ eviker & Albaryak, 2006; cucci-Papadopoulou et al. (2005) referred the Black Galil, 2007). Sea as the main source of alien species for Thessalo- Gastrochaena cymbium was firstly referred from S niki Bay, although this could not be supported any Aegean by Tenekidis (1989) and was characterized longer according to the recent information presented and listed as “casual” alien species (ELNAIS, Zene- in the research effort of Zenetos et al. (2009a). tos et al., 2009a). After the frequent occurence of G. The distribution of Pinctada radiata has expanded cymbium along the north-east coasts of the study a- north to the E Thermaikos along the navigation line rea, its characterization as “casual” should be rather to the harbour of Thessaloniki. This may be attrib- changed to “established”. uted to the same reasons, namely shipping and mari- The four Chamidae species collected in the study culture activities as well as to epibiosis on the sea tur- area and originated from the Red Sea seem to be well tle Caretta caretta that is believed to be responsible expanded and established, sinse large size shells be- for its distribution to the Hellenic seas (Oliverio et al., longing to each one of them have been found (be- 1992; Zenetos et al., 2003). sides of the mentioned alive specimens). Thus, the ex- We feel that further research emphasizing on pansion of Chama aspersa and C. asperella from the small in size species (most of the new records of this Red Sea to the E Mediterranean in 2004 and to S Ae- study were <5 mm), symbionts, epibionts and rare gean Sea in 2007 presented by Ovalis & Zenetos (2007) species needs to be contucted in order to further ex- continuous to the N Aegean Sea. plore the present biodiversity of bivalves in the area. 178 Thanasis Manousis et al. — The Bivalvia Mollusca of Thessaloniki & Thermaikos Gulfs

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