Mediterranean Marine Science

Vol. 9, 2008

Molluscan of minor commercial interest in Hellenic seas: Distribution, exploitation and conservation status

KATSANEVAKIS S. European Commission, Joint Research Centre, Institute for Environment and Sustainability, Ispra LEFKADITOU E. Hellenic Centre for Marine Research, Institute of Marine Biological Resources, Agios Kosmas, P.C. 16610, Elliniko, Athens GALINOU-MITSOUDI S. Fisheries & Aquaculture Technology, Alexander Technological Educational Institute of Thessaloniki, 63200, Nea Moudania KOUTSOUBAS D. University of the Aegean, Department of Marine Science, University Hill, 81100 Mytilini ZENETOS A. Hellenic Centre for Marine Research, Institute of Marine Biological Resources, Agios Kosmas, P.C. 16610, Elliniko, Athens https://doi.org/10.12681/mms.145

Copyright © 2008

To cite this article:

KATSANEVAKIS, S., LEFKADITOU, E., GALINOU-MITSOUDI, S., KOUTSOUBAS, D., & ZENETOS, A. (2008). Molluscan species of minor commercial interest in Hellenic seas: Distribution, exploitation and conservation status. Mediterranean Marine Science, 9(1), 77-118. doi:https://doi.org/10.12681/mms.145

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | Review Article

Mediterranean Marine Science Volume 9/1, 2008, 77-118

Molluscan species of minor commercial interest in Hellenic seas: Distribution, exploitation and conservation status

S. KATSANEVAKIS1, E. LEFKADITOU1, S. GALINOU-MITSOUDI2, D. KOUTSOUBAS3 and A. ZENETOS1

1 Hellenic Centre for Marine Research, Institute of Marine Biological Resources, P.O. Box 712, 19013, Anavissos, Attica, Hellas 2 Fisheries & Aquaculture Technology, Alexander Technological Educational Institute of Thessaloniki, 63200, Nea Moudania, Hellas 3 Department of Marine Sciences, School of Environment, University of the Aegean, 81100, Mytilini, Hellas e-mail: [email protected]

Abstract

Recognising the need for data gathering at taxon level besides the taxa of commercial interest and those listed as endangered in the Protocol of the Barcelona Convention, but still exploited by Man, the present study attempts to gather and review the available scientific information on molluscs of minor commercial importance in order to assist in the adequate management and protection of their popula- tions. Forty one species (18 gastropods, 13 bivalves, and 10 ) of minor commercial interest are treated in the present work with details on their biogeographic distribution, exploitation and conser- vation status in Hellas. Apart from a few species (e.g., Pinna nobilis, , Donacilla cornea), and these only at a local scale, there is no population assessment in Hellenic seas. The existing legislation for eleven of these species is not enforced in practice, and seems insufficient to guarantee their conservation. It is suggested that targeted collection data, networked nationally and internationally, should be promoted so this invaluable source of biodiversity information can be accessed for conserva- tion and planning purposes.

Keywords: exploitation; Conservation; Hellas.

Introduction the development of high density human populations near the coast has resulted in Exploitation of mollusca as food and as unprecedented and unsustainable levels of ornaments in Hellas has a history dating pressure nowadays. Unfortunately, sea- back to the Paleolithic age (SHACKLE- shells are collected in those areas least TON & VAN ANDEL, 2007). However, capable of withstanding exploitation. While

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | shelled mollusca are heavily exploited, some have been destructive and have caused a species are impacted by uncon- significant decline in the Mediterranean trolled and unreported fishing, which may populations of several species (e.g., Pinna have a significant effect on their populations. nobilis) and have brought others to near A total of 1160 mollusc species (1 extinction (e.g., ferruginea) Solenogastra, 2 Caudofoveata, 771 Gas- (BOUDOURESQUE, 2004). In addition, tropoda, 308 , 19 Polyplacophora, basic knowledge on the biology and ecolo- 12 Scaphopoda and 47 Cephalopoda) have gy of these species is either lacking or is been recorded so far in Hellenic seas scattered in the literature. (KOUTSOUBAS, 2004; CHINTIRO- The red list species approach to con- GLOU et al., 2005; ZENETOS et al., 2005; servation requires detailed knowledge LEFKADITOU, 2007; DELAMOTTE & concerning individual taxa (distribution, VARDALA-THEODOROU, 2007). A declines, threatening processes, biological total of 27 of these species (seventeen attributes, etc.). However, it addresses bivalves, four gastropods and six only the tip of the biodiversity iceberg. The cephalopods), comprising less than 2.5 % first step in knowledge acquisition of the molluscan fauna of the Hellenic required for the taxon-based approach to seas, have a commercial interest, particu- biodiversity and conservation is the study larly in fisheries and aquaculture since and documentation of at least the major they are collected and/or cultivated for components of the fauna. Hence, the pres- human consumption. The status of the ent study attempts to gather and review fishing of these commercially important the available scientific information on molluscs has been recently reviewed molluscs of minor commercial importance (KOUTSOUBAS et al., 2007; LEFKA- in order to assist in the adequate manage- DITOU, 2007). ment and protection of their populations. However, apart from those commer- cially important species, there are many Materials and Methods other molluscs, which, although not com- mercially exploited, are either systemati- Forty one species (18 gastropods, 13 cally or occasionally collected for human bivalves, and 10 cephalopods) of minor consumption and other uses (fishing bait, commercial interest were reviewed in the jewellery, decoration, private and public present work (Table 1). The distribution of collections). Furthermore, several species, the listed species in Hellenic seas was gen- for example some common small-sized erally based on ZENETOS (1986; 1996), cephalopods, constitute trawl fisheries by- KOUTSOUBAS (1992), KOUTSOUBAS catches which may have a marketable frac- et al. (1992, 1997, 1999, 2000a,b), tion. Bivalve and gastropod molluscs are ZENETOS et al. (2004, 2007), ANTO- also frequently used to biomonitor heavy NIADOU et al. (2005), DELAMOTTE & metal pollution in the marine environ- VARDALA-THEODOROU (2007), ment. In most cases, the distribution, the LEFKADITOU (2007), EVAGELO- degree of exploitation, and the state of POULOS & KOUTSOUBAS (2008), grey these populations have not been assessed literature such as academic dissertations and in Hellenic seas. Uncontrolled and unre- HCMR or EU technical reports, and/or ported fishing or other human activities on unpublished/personal observations.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | g), , sh ersity (continued) Table 1 109/2002 = Presidential Decree 109/2002) are also given. Petalida /Rayed Mediterranean limpetStiktopetalida /Ristic limpetAgriopetalida /Rough , China limpetSalingaraki, Arthrotos trochos /Articulate monodontSalingaraki, Trochos fraoula /Turbinate monodontSalingarakiStrovilos, Mati tis Panagias (the opperculum)/Rough turboSkaltsini, Striftari, Kerato /Mediterranean ceritheKonos tis Persias ED, FB, BIOM Gourounitsa / ED, FB, BIOM JWL, Gourounitsa COL /Cowrie ED, FB, BIOM ED, FB, BIOMGourounitsa /CowrieBouchona, Kochyla /Giant ton 109/2002 Agkathotritonas /Wandering triton, Olive trumpet 109/2002 FB ED, FB, BIOMTritonas /Knobed triton, lamp tritonTritonas, Bourou /Variegated tritonMitra /Zoned miter 109/2002 Konos /Cone shellPetrosolinas, Valanos,Chourmas /European date COLBC, PBC Pinna /Pen shell, Fan shell ED, FB COL, JWL ED ED, COL ED, COLBC, PBC, 67/1981 COL COL, JWL COL COL, JWL BC, PBC, 67/1981 BC, PBC, 67/1981 BC, PBC, 67/1981 92/43/EC, BC, PBC BC, PBC BC, PBC ED, COL COL ED, COL, BIOM BC, PBC 92/43/EC, PBC, 67/1981 in the Mediterranean of the Barcelona Convention, 92/43/EC = EC Habitats Directive 92/43, 67/1981 = Presidential Decree 67/1981 and the legislation for their protection (BC = Bern Convention, PBC = Protocol for Specially Protected Areas and Biological Div Mollusc species (, Bivalvia and Cephalopoda) of minor commercial interest in Hellenic seas. Common Hellenic and Engli names, exploitation modes (ED = Edible, FB = Fishing Bait, JWL = Jewelry, COL = Shell Collections, BIOM = used for Biomonitorin Scientific nameGASTROPODA Common Hellenic /English namesPatella ulyssiponensis Osilinus articulatus Osilinus turbinatus Phorcus mutabilis Bolma rugosa Cerithium vulgatum Strombus persicus Erosaria spurca lurida Zonaria pyrum Tonna galea Exploitation modeRanella olearium Charonia lampas Legislation Charonia variegata Mitra zonata Conus mediterraneus BIVALVIA Lithophaga lithophaga Pinna nobilis

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | (continued) Table 1 Margaritoforo stridi /Rayed pearl oysterFterostrido /European wing ysterKalogria, Chlamyda /Variegated scallopGrammoto chteni /Little bay scallopGaidouropodaro, Vassiliko stridi /European thorny ysterPortogaliko stridi /Giant (or Pacific) cupped oysterAmmokochylo, Mikri ahivada /Banded wedge shellSolinas, Ammosolinas /European razor clamSolinas, Ammosolinas /Giant razor ED, COL Samari, Zamponaki /Sandwich ED, JWL , BIOM Daktilo, Ftera aggelon, Folada /Common piddock ED, BIOM ED, COL Kalamaraki /Midsize ED, BIOM ED, COL Kalamari /Veined squid ED, COL Thrapsalo, Katamachi /European flying squidThrapsalo /Lesser flying ED, squidFB, BIOMElegant ED, FB Kokkinosoupia /Pink cuttlefish ED, FB 109/2002 Mikrosoupia /Stout bodtailSoupitsa /Lentil bodtailSoupitsa /Common bodtail ED, FB, Monokero COL109/2002 chtapodi /Unihorned BC, PBC ED, FB 109/2002 ED ED ED, FB ED ED ED ED ED ED Scientific nameBIVALVIA Pinctada radiata Pteria hirundo Common Hellenic /English namesChlamys varia Crassadoma multistriata Spondylus gaederopus gigas Donacilla cornea Solen marginatus minor strigilatus dactylus CEPHALOPODA Alloteuthis media Exploitation modeLoligo forbesi Todarodes sagittatus Legislation Todaropsis eblanae Sepia elegans Sepia orbignyana Rossia macrosoma Rondeletiola minor oweniana Scaeurgus unicirrhus

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | A general map of Hellas with the includes the Presidential Decree 67/1981, names of the gulfs and other locations the EC Habitats Directive 92/43, the Bern mentioned in the text is given in Figure 1. Convention, and the Protocol for Specially General information on the biology and Protected Areas and Biological Diversity in ecology of the species was gathered after a the Mediterranean of the Barcelona Con- thorough search of international and vention (hereafter: Protocol of the national literature. Nomenclature for gas- Barcelona Convention). For species tropods and bivalves followed the Check included in Annex IV of the Directive List of European Marine Mollusca 92/43/EC, all forms of deliberate capture or CLEMAM (http://www.somali.asso.fr/ killing of specimens in the wild, deliberate clemam/biotaxis.php), whereas that for disturbance, particularly during the period cephalopods followed the classification of of breeding, deliberate destruction or tak- living coleoid species of the Mediter- ing of eggs from the wild, and deterioration ranean Sea by MANGOLD & von or destruction of breeding sites or resting BOLETZKY (1987). places are strictly prohibited. For species Legislation concerning the conserva- included in Annex II of the Bern Conven- tion of each species is reported where tion, the collection, keeping in captivity, available (Table 1). The existing legislation killing, destruction or collecting of eggs, for the conservation of marine species disturbance especially during the reproduc-

Fig. 1: A general map of Hellas, with indications of the main locations mentioned in the text.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | tion period, possession or commerce of live treated together. or dead specimens are strictly prohibited. The selection of the forty-one species For species included in Annex II of the listed in this paper was based on data avail- Protocol of the Barcelona Convention, col- ability and the expert knowledge of the lection, capture, killing, commerce, trans- authors. Several other species, not includ- portation and disturbance especially during ed in this list might also be considered as reproduction should be prohibited or con- molluscs of minor commercial interest, as trolled. The Presidential Decree (PD) for example, Glycymeris glycymeris, Gly- 67/1981 prohibits killing, causing injury, cymeris bimaculata, Acanthocardia tubercu- capture, collection, commerce and trans- lata, Glossus humanus, Phalium granula- portation of all included species. Further- tum, Cassidaria echinophora, which are more, according to the PD 86/98 (regard- collected in great numbers by shell collec- ing shell-fishing) as reformed by the PD tors. In this review, we included those 227/2003, and the PD 109/2002 (regarding species that we subjectively considered as fishing baits), fishing, transportation and most important. commerce of any species of shelled mollus- ca not included in these PDs is prohibited. Results Thus, among the bivalves and gastropods of this study, exploitation is prohibited for 1. Gastropods all species except Solecurtus strigilatus, Ensis siliqua, Solen marginatus, Pholas Family : Patella caerulea (Lin- dactylus and Patella spp., which are includ- naeus, 1758), P. rustica (= lusitanica) (Lin- ed in PD 109/2002. naeus, 1758), P. ulyssiponensis (= aspera) The species listed below are sorted (Gmelin, 1791) first taxonomically (gastropods, bivalves and cephalopods) and then phyllogeneti- General: Three species have been cally. For each species, relevant informa- reported in Hellenic seas: Patella caerulea, tion is categorized into four sections, i.e. P. rustica (= lusitanica), and P. ulyssipo- General, with some diagnostic features of nensis (= aspera) (KOUTSOUBAS, 1992; the species and useful information on its DELAMOTTE & VARDALA-THEO- autecology, Distribution where the global DOROU, 2007). (Patella sp.) are and the Hellenic distribution of the species common inhabitants of the hard substrate is given, Exploitation, where the mode of communities in the intertidal and upper exploitation and the uses of the species in subtidal zones. They have a strong conical the past and at present are reported, and shell with radiating ridges, which may have Conservation Status, in which possible pop- epiphytes or epizooites growing on it. Lim- ulation assessments of the species and rel- pets attach themselves to the substratum evant legislation for its conservation are using their strong muscular foot, which presented. In Exploitation, the real mode enables them to withstand heavy wave of exploitation is presented and not that action. They are herbivorous and feed according to existing legislation, as in most mostly at night grazing on algae found on cases legislation is violated and species for rocks. They use a very hard-toothed radu- which fishing is prohibited may be exten- la to scrape rock surfaces, often limiting sively exploited. Con-generic species are algal growth on exposed and moderately

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | exposed shores. They have a homing General: Monodonts are common behaviour and return to the same spot inhabitants of the hard substrate commu- after eating, using sensitive chemorecep- nities in the upper subtidal zone. The tors to find their way back; sometimes they height of Osilinus turbinatus, O. articulatus, use the same ‘home’ for their whole life. and Phorcus mutabilis may reach 30 mm, P. caerulea may reach a length of 70 mm, 30 mm and 18 mm respectively. They are P. ulyssiponensis a length of 50 mm, while herbivorous and feed mostly at night graz- P. rustica is relatively smaller, attaining a ing on algae found on rocks. length of 45 mm. Distribution: All three species are Distribution: P. caerulea is present abundant in the Mediterranean basin, throughout the Mediterranean Sea and in while O. atriculatus and O. turbinatus are a few places in the eastern Atlantic (Bay of also found in the eastern Atlantic. In Hel- Biscay, Canary Islands, Moroccan coast). lenic seas, O. turbinatus seems to be com- P. ulyssiponensis has the widest distribu- mon everywhere, while O. articulatus has tion, being abundant on Atlantic shores been reported in the Saronikos, Evvoikos from Norway to Mauritania, in the and Pagasitikos Gulfs, along the Pelopon- Mediterranean and Black Sea, and in the nesian coasts, the Sporades, the N. Macaronesian Islands. P. rustica occurs in Aegean and Ionian Sea, and P. mutabilis the Mediterranean Sea and on the Atlantic in the Saronikos, Korinthiakos and coasts from Biarritz (France) to Maurita- Evvoikos Gulfs, the N. Aegean, along the nia, in , the Canary Islands, Peloponnesian coasts, Rodos and the Ion- Selvagens, Desertas, and Madeira. P. ian Sea. caerulea is the most common limpet in Exploitation: Small quantities of Mon- Hellenic seas. P. rustica, and P. ulyssipo- odonts (especially O. turbinatus) are col- nensis are also common in most Hellenic lected for human consumption and spo- coastal areas. radically offered in small taverns in fishing Exploitation: Limpets are widely col- villages especially in the Dodekanisos lected for human consumption and as fish- islands (S Aegean Sea) and the Chalkidiki ing bait. They are also used as bioindica- Peninsula (N Aegean Sea). They are also tors in heavy metal bioaccumulation stud- used as bioindicators in heavy metal bioac- ies (e.g., CONTI & CECCHETTI, 2003). cumulation studies (e.g., CONTI & Their fishing in Hellenic seas is regulated CECCHETTI, 2003). by the PD 109/2002. Conservation Status: Presumably, the Conservation Status: Although no three species are abundant in Hellenic population assessment has been conduct- seas. There is no relevant legislation. No ed, all three species seem to be rather population assessment has been conduct- common. ed in Hellenic seas.

Family Trochidae: Osilinus turbinatus Bolma (= Astraea) rugosa (Linnaeus, 1767) (= Monodonta turbinata) (Von Born, 1778), Osilinus articulatus (= Monodonta General: Bolma rugosa is an inhabitant articulata) (Lamarck, 1822), and Phorcus of photophilic algae or coralligenous mutabilis (= Monodonta mutabilis) (Philip- assemblages and Posidonia oceanica pi, 1846) meadows (KOUTSOUBAS, 1992;

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | ANTONIADOU et al., 2005; personal nating the archaeological record in the observations). Its shell may reach a height Franchthi Cave, an Upper Paleolithic find of 60 mm and it has a thick, glossy, cream in the southern Peloponnisos, Hellas. Per- to orange, calcified operculum, which in haps this small, gregarious was col- Hellas is called ‘Mati tis Panagias’ (i.e., lected in such numbers not for food but for ‘the Virgin Mary’s eye’). the purpose of conversion into body orna- Distribution: Bolma rugosa is found in ments, as the perforations in most shells the Mediterranean and E Atlantic suggest (SHACKLETON & VAN (around the Mediterranean basin), ANDEL, 2007). Nowadays, there is only including Madeira, the Azores and the limited exploitation of the species and they Canary Islands. B. rugosa is common and are used as fishing bait. may be found almost everywhere in Hel- Conservation Status: C. vulgatum is lenic seas. very common. However, there is no popu- Exploitation: In Hellenic seas, B. lation assessment in Hellenic seas. rugosa is exploited (especially on the islands of the Kyklades plateau) for its Strombus (= Conomurex) persicus (= de- beautifully coloured calcified operculum, corus raybaudii) (Swainson, 1821) which is utilized in jewellery making. Conservation Status: It is not protect- General: Strombus persicus may ed by legislation. No population assess- exceed a height of 75 mm, is gonochoristic ment has been conducted in Hellenic with sexual dimorphism in its radula seas. (MUTLU, 2004). Larvae are plank- totrophic, ensuring long-distance disper- Cerithium vulgatum (Bruguière, 1792) sal. All Strombus species are herbivorous, feeding on algae. In the Indo-Pacific General: Approximately 300 species regions, S. persicus lives on sandy bottoms belong to the family Cerithiidae, which and in coral sand (ABBOTT, 1960). In the are small and medium sized shells found Mediterranean, adult S. persicus inhabit in temperate and tropical seas. C. vulga- sandy or slightly gravelly bottoms and feed tum adults may reach a height of 5–7 cm. on seaweeds and detritus, while juveniles C. vulgatum may be found in a variety of (shell length <2 cm) were found either liv- biotopes, mainly on muddy substrate, on ing in colonies on small rocks covered by rocky and pebble bottoms, and in seagrass algae or on sandy bottoms with detritus beds. It prefers areas of low wave intensi- (MUTLU, 2004). ty, such as lagoons and embayments. It is Distribution: S. persicus is a subtropical a herbivore, feeding mainly on small species, initially restricted to the south algae. Arabian and part of the Iranian coast, that Distribution: C. vulgatum lives in the has entered the Mediterranean presum- Mediterranean Sea and E Atlantic, from ably via shipping through the Suez Canal the British Isles to the coast of Senegal. In and has become locally invasive in the Hellenic seas, C. vulgatum is abundant south-eastern Mediterranean (OLI- along most coastal areas. VERIO, 1995; STREFTARIS & Exploitation: Cerithium vulgatum, has ZENETOS, 2006). S. persicus was first been found in a range of habitats, domi- recorded in the Mediterranean Sea in 1978

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | in Iskenderum (southern Turkish coasts) exploited in Rodos Island over recent (NICOLAY, 1986) and was successively years and sporadically offered for human found in Israel, Hellas, Cyprus, Syria and consumption in fish restaurants. Lebanon (ZENETOS et al., 2004). In Hel- Conservation Status: It is a successful las, it has been established in Rodos, the invader in the Mediterranean Sea and its south Peloponnisos coasts, Kriti, the Kyk- spatial distribution keeps increasing lades, the Argolikos, Evvoikos, and (ZENETOS et al., 2004; YOUNG, 2007; Saronikos gulfs (ZENETOS et al., 2004; ZENETOS et al., 2007). There is no rele- YOUNG, 2007; ZENETOS et al., 2007; vant legislation for its conservation. personal observations); it has not yet been Although there are detailed records of its reported in the Ionian or the N Aegean invasion into the Mediterranean and Hel- Sea (Fig. 2). lenic seas, no population assessment has Exploitation: The species has been been conducted in Hellas.

Fig. 2: Known distribution of Strombus persicus in Hellenic seas.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | Family : Luria (= ) (Spirastrella cunctatrix, Cacospongia sp.), lurida (Linnaeus, 1758), Erosaria (= and colonial hydrozoans (Eudendrium sp.) Cypraea) spurca (Linnaeus, 1758), Zonar- (LIPPARINI & NEGRA, 2002). Zonaria ia (= Cypraea) pyrum (Gmelin, 1791) pyrum may exceed 50 mm in height. Speci- mens from the N Aegean were found on General: The shells are extremely rocky substrate with many Axinella spp. smooth and shiny and exhibit a great vari- sponges, at a depth of 25 m (KOU- ety of colouration, size, and sculpture. The TSOUBAS, 1992). French call them ‘porcelains’ because of Distribution: The family Cypraeidae their texture. The of the () includes approximately 190 usually covers most of the shell surface species, mostly distributed in tropical and keeping it in excellent condition through- subtropical waters. is the most out its life and without epibionts. Cowries common cowry of the Hellenic seas. Its have no operculum and a long, very nar- distribution extends to the Mediterranean row, aperture which is lined with ‘teeth’. Sea and E and W , while Luria lurida may exceed 65 mm in height. after the opening of the Suez Canal it has It lives in rocky areas of the subtidal zone also invaded the Red Sea and is consid- and is a nocturnal animal. During daylight ered as an anti-lessepsian migrant it usually remains hidden in crevices or in (KOUTSOUBAS, 1992; DELAMOTTE sponge colonies, while at night it is active & VARDALA-THEODOROU, 2007). searching for its prey. L. lurida selectively L. lurida has been reported in the feeds on sponges having a strong prefer- Saronikos, Evvoikos, Pagasitikos, ence for Aplysina aerophoba. It has also Lakonikos, and Korinthiakos gulfs, the been observed to prey on the sponges Kyklades, the Dodekanissa, Kriti, the N Chondrosia reniformis, Chondrilla nucula, Aegean, and the Ionian Sea. Erosaria spur- and Suberites domuncula, while the ascidi- ca is found in the Mediterranean, the Red an Ciona intestinalis is also preyed on occa- Sea and the Atlantic Ocean. It is less com- sionally, when of small size (DONEDU & mon in Hellenic seas than L. lurida and it MANUNZA, 1995). In the N Aegean, L. has been reported in the Saronikos, lurida specimens have been found in Evvoikos, Messiniakos and Korinthiakos colonies of A. aerophoba and Petrosia fici- gulfs, the Kyklades, the Dodekanisos, formis (KOUTSOUBAS, 1992). Although Kriti, Rodos, Samos, and the N Aegean. its longevity is not accurately known, L. Zonaria pyrum appears to be the least lurida specimens have been kept alive in abundant of the three species. It is found aquariums for more than 11 years. in the Mediterranean Sea and E Atlantic. Erosaria spurca may reach 39 mm in In Hellenic seas it has been reported in the height. It is a nocturnal animal, remaining Evvoikos Gulf, along the Peloponnisos hidden during daylight hours, and lives in coasts, the Kyklades, the Dodekanisos, rocky subtidal areas and in seagrass beds. Samos, the N Aegean, Kriti, and the Ion- In Messina (Sicily) it is normally found on ian Sea. substrates such as shaded walls covered by Exploitation: All three species are chlorophyceans (Cladophora prolifera), among the most beautiful shells in Hel- pheophyceans (Padina pavonica, Colpo- lenic seas and are very popular among menia sinuosa), encrusting demosponges shell collectors. They are not edible and

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | are exploited to be used in jewellery or to Distribution: The family Tonnidae be sold to shell collectors. includes approximately 30 species living in Conservation Status: All three species temperate and tropical seas. T. galea is a are under strict protection according to the cosmopolitan species. It has a wide distri- Bern Convention (Annex II), the Protocol bution in Hellenic seas and has been of the Barcelona Convention (Annex II), recorded in the Saronikos, Korinthiakos, and the Presidential Decree 67/1981. and Evvoikos gulfs, the N Aegean Sea (SE However, they are still illegally collected Thermaikos Gulf, Chalkidiki - SE Toron- and sold, and the relevant legislation has in aios Gulf), the Kyklades, the Ionian Sea practice never been implemented. No pop- and the Cretan Sea. ulation assessment has been conducted in Exploitation: T. galea is edible and is Hellenic seas for any of the three species. sporadically collected in many Hellenic areas (Evvoikos Gulf, Gulf of Kavala, Tonna (= Dolium) galea (Linnaeus, 1758) Thracian Sea) by divers for its flesh and also for its shell or as a by-catch by fishing General: T. galea is one of the largest gear (e.g. trawlers, nets). In a fisheries’ sur- Mediterranean gastropods with a height vey using bottom nets in the SE Ther- that may exceed 29 cm. It is gonochoristic maikos Gulf during summer, 1 individual and has no operculum. It lives in per 1500 m nets per day was caught in 25 % sandy/muddy bottoms and seagrass beds, of the cases (GALINOU MITSOUDI, at depths from a few meters to 120 m. It is unpublished data). an active carnivore, mostly predating on Conservation Status: Although, no holothurians but also on sea urchins and population assessment has been conduct- starfish. T. galea is equipped with a partial- ed in Hellenic seas, it is believed that its ly invaginable and highly extensible pro- populations have greatly declined in the boscis, which is capable of engulfing prey last decades. It is a protected species that rarely manages to escape. T. galea has according to Annex II of the Bern Con- enlarged salivary glands whose secretion vention, the Protocol of the Barcelona contains 2.20–4.88 % sulfuric acid in addi- Convention (Annex II), and the Presiden- tion to hydrochloric acid (FÄNGE & tial Decree 67/1981; however, it is still LIDMAN, 1976) that can mediate the dis- exploited. solving of the prey integuments. The acquisition of enlarged salivary glands Family Ranellidae: Charonia lampas (Lin- capable of secreting acid was first discov- naeus, 1758), Charonia variegata (= trito- ered in 1853 by Troschel who found that nis variegata) (Lamarck, 1816) after the mollusc shell was crushed it pro- truded the proboscis over a distance of 13- General: Charonia lampas and C. varie- 16 cm and released a jet of clear fluid gata (tritons) are among the largest gas- inducing active dissolution of a marble tropods of the Mediterranean Sea with a floor (FÄNGE & LIDMAN, 1976). T. height that may exceed 35 cm. They are galea is a nocturnal species and is usually gonochoristic gastropods and have a cor- buried in the sand during daylight hours neous operculum. Adult tritons are active for protection. The is its predators and mainly feed on starfish, sea main natural predator. urchins, sea cucumbers, and molluscs. Tri-

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | ton locates its prey by chemoreception of (Annex II) and the Protocol of the current-born prey substances. It grips its Barcelona Convention (Annex II). No prey with its muscular foot and uses its population assessment has been conduct- radula to saw through the prey’s integu- ed in Hellenic seas. ments. Once it has penetrated, its paralyz- ing saliva subdues the prey and the snail Ranella (= Argobuccinum) olearium (Lin- feeds on it. naeus, 1758) Distribution: C. variegata lives in rocky shores of temperate and tropical waters in General: Ranella olearium is an outer the Atlantic Ocean and the Mediterranean shelf–upper bathyal species of the family Sea. It occurs mainly in the warm waters of Ranellidae. It is a large probosciferous gas- the western Atlantic, in the Azores, the tropod with a height that may exceed 22 cm. Canary and Cape Verde Islands (eastern Ranellinae are generalist predators, mostly Atlantic) and in the eastern Mediter- feeding on echinoderms, ascidians, and ranean; in the western Mediterranean it polychaetes; however, little is known about has been observed only sporadically the specific feeding habits of R. olearium. (TEMPLADO, 1991). In Hellenic seas, C. Distribution: R. olearium is a cosmo- variegata has been reported in the politan species and is mostly found in the Saronikos, Korinthiakos, Lakonikos and NE Atlantic (SW British Isles to Maureta- Evvoikos gulfs, the Kyklades, Kriti, nia, including the Azores, Canary Islands, Dodekanisos, the N Aegean, and the Ion- Lusitanian and Meteor group seamounts), ian Sea. C. lampas is found in the E the Mediterranean Sea, S Africa (Indian Atlantic Ocean and the Mediterranean Ocean) and New Zealand, while it has also Sea. In Hellenic seas, it has been reported been recorded occasionally in the W and in the N Aegean Sea and the coasts of the SE Atlantic, the Caribbean Sea, the S Indi- Peloponnisos, Samos, and Kriti. an Ocean, and the SW Pacific Exploitation: From ancient times, peo- (SCARABINO, 2003). In Hellenic seas, it ple of many different cultures, mainly fish- has been reported around the Sporades ermen, have removed the tip of the shell Islands, in the Evvoikos, on the Kymi of the triton or drilled a hole in the tip and coasts, and Chalkidiki. then utilized the shell as a trumpet to Exploitation: It is a by-catch species of communicate over long distances. Many trawl fishery. It is popular among shell col- people find triton shells striking and lectors. attractive and so they are collected and Conservation Status: The species is sold as part of the international shell strictly protected according to the Bern trade. This has contributed to the over- Convention (Annex II) and the Protocol of fishing and depletion of many triton pop- the Barcelona Convention (Annex II). ulations worldwide. However, its populations are probably Conservation Status: Nowadays, tri- greatly affected by trawling. In a study of tons are protected in many areas of the the effects of commercial trawling on the world. In the Mediterranean, both C. lam- malacological communities on the conti- pas and C. variegata have been declared as nental shelf and slope of Portugal endangered species and are strictly pro- (MALAQUIAS et al., 2006), R. olearium tected according to the Bern Convention accounted for 7.3 % of the total number of

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | individuals caught in the fish trawls. Conus mediterraneus (= ventricosus) Although most of the individuals caught by (Hwass in Bruguière, 1792) trawlers are discarded, mortality may occur as a consequence of aggressive han- General: Conus mediterraneus has a dling by fishers, thermal stress, or from set- distinctively conical shape and can grow up tling on unsuitable substratum. The status to 7 cm. It has a brown, elliptical, corneous of R. olearium populations has not been operculum of length about 30 % of the assessed in Hellenic seas or anywhere else. aperture length. C. mediterraneus is a predatory, vermivorous species, being Mitra zonata (Marryat, 1818) active mostly at night. It catches its prey using singular, venomous, barbed, har- General: Mitra zonata belongs to the poon type radular teeth that are thrust into family Mitridae and is well differentiated their prey with the use of their long and from the other congeneric species due to its highly maneuverable proboscis. In a study larger size. It seems to be rather rare, and of its feeding ecology in Tunisia, C. lives in sandy, coralligenous, or muddy bot- mediterraneus was found to eat fourteen toms of deep waters. KOUTSOUBAS species of polychaete which together with (1992) has reported the collection of three a gastropod of the Nereididae species samples that were on a red coral (Coralli- were the most common food items um rubrum) colony at a depth of 75 m. It (TAYLOR, 1987). Cone possess a may exceed a height of 85 mm. Species of large, coiled venom gland, and use their the family Mitridae are carnivores usually composite venom to envenomate their feeding on sipunculans and small gas- prey and to defend themselves from pred- tropods; however, the specific diet of M. ators and competitors. This venom con- zonata is unknown. tains a wide array of cysteine-rich, neu- Distribution: M. zonata occurs in a few roactive oligopeptides (named conopep- localities of the E Atlantic Ocean (e.g. the tides), which interfere with neuromuscular Canary Islands, the Azores, Madeira, transmission (MASSILIA et al., 2001). C. Angola) and the Mediterranean Sea. In mediterraneus crude venom extracts have a Hellenic seas, it has been reported in the paralytic effect on polychaete worms and Saronikos and Evvoikos gulfs, Chalkidiki, on molluscs (FAINZILBER & Kymi (Evvoia), the Ionian Sea, and N Spo- KLEIMAN, 1989). rades Islands. Distribution: The family Conidae has Exploitation: The startling contrast of many representatives in tropical and sub- its beautiful colours and its rarity make M. tropical seas (approximately 400 species) zonata very appreciated and popular but C. mediterraneus is the single Conus among shell collectors. It is edible and is species living in the Mediterranean Sea; it eaten in Croatia; it is not eaten in Hellas. is also found on the coasts of western Conservation Status: M. zonata is under Africa. It is a very common species of the strict protection according to the Bern rocky upper subtidal zone in Hellenic seas Convention (Annex II) and the Protocol of and may be found everywhere, from the N the Barcelona Convention (Annex II). No Aegean to the Libyan Sea, and from the population assessment has been conduct- Ionian Sea to the Dodekanisos. ed in Hellenic seas. Exploitation: Its exploitation is very

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | limited, for private or public collections. It Gulf, the Ionian islands, the Ipeiros coast, is not edible. the Korinthiakos Gulf, the Sea of Kythira, Conservation Status: Presumably, the the N Sporades Islands, the Dodekanisos, species is abundant. There is no relevant Kriti, Lesvos island., Maliakos, Argolikos legislation. No population assessment has and Chalkidiki (Fig. 3) (ZENETOS 1986; been conducted in Hellenic seas. 1996; GALINOU-MITSOUDI & SINIS, 1994, 1997; GEROVASILEIOU et al., 2. Bivalves 2007; personal observations). Exploitation: In the past, L. lithophaga Lithophaga lithophaga (Linnaeus, 1758) was extensively exploited in Hellenic seas and it was found in markets and General: The European date mussel fish restaurants in many locations. The Lithophaga lithophaga is an endolithic species is still exploited by divers (illegally) species and an inhabitant of hard substrata as a delicious food resource, especially in (calcified) communities in the midlittoral Evvoia. and upper sublittoral zones (GALINOU- Conservation Status: The species is MITSOUDI & SINIS, 1994). The autoe- under strict protection according to the cology (ecology, population dynamic, biol- 92/43/EC Directive (Annex IV), the Bern ogy) of L. lithophaga has been extensively Convention (Annex II), and the Protocol studied in the Evvoikos Gulf (GALINOU- of the Barcelona Convention (Annex II). MITSOUDI & SINIS, 1994, 1995). The The population of Evvoikos Gulf has been population density of this species, which assessed in the past (GALINOU- seems to be dependent on the substratum MITSOUDI & SINIS, 1994). type (limestones), the presence of other organisms (e.g. endolithic), predation and Pinna nobilis (Linnaeus, 1758) environmental conditions, is higher in the N. Evvoikos Gulf (36 inds/dm3). The age of General: The fan mussel Pinna nobilis first reproduction is 2+ years and the occurs at depths of between 0.5 and 60 m, reproductive activity is annual and contin- typically in soft-bottom areas overgrown uous. The main reproductive period is dur- by seagrass (mostly Posidonia oceanica ing the summer and autumn at water tem- and Cymodocea nodosa meadows) but peratures > 14ÆC. L. lithophaga is a long- also in unvegetated sandy bottoms lived species (> 54 years) and its growth (KATSANEVAKIS, 2006; 2007a). It is rate is one of the slowest among bivalves. the largest Mediterranean bivalve and one The highest growth rate is observed at the of the largest in the world, attaining end of spring and early summer. In the lengths of up to 120 cm. P. nobilis is long- Evvoikos Gulf, the annual productivity of lived and may live over 20 years L. lithophaga was 1659.6 g/m2 and the (GALINOU-MITSOUDI et al., 2006). annual turnover ratio (P/B) was 0.845, Fan live partially buried (usually which is among the lowest values reported >35 % of their length) by the anterior in the literature. portion of the shell and attached by their Distribution: L. lithophaga is endemic byssus threads to the substratum. Repro- in the Mediterranean Sea. In Hellenic duction of P. nobilis takes place by means seas, the species is found in the Evvoikos of external fertilization and its success

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | Fig. 3: Population density hotspots of Lithophaga lithophaga. depends on the proximity of other individ- causes quickly diminishes as the fan mus- uals spawning synchronously. When a sels grow in size (KATSANEVAKIS, population becomes sparse (as is the case 2007b). Growth rates have a seasonal pat- for most fan mussel populations in the tern, with an extended period of very slow Mediterranean) failure of fertilization growth between late autumn and early becomes a critical issue for the survival of spring and a peak in growth rates during the species. The planktonic period is late spring - early summer, probably relat- short, hence populations (especially in ed to an optimum combination of temper- closed embayments), which are relatively ature and food availability (KATSA- isolated are not easily recolonised if NEVAKIS, 2007b). depleted. Natural mortality is strikingly Distribution: P. nobilis is endemic to size dependent and P. nobilis suffers high the Mediterranean Sea. The global popu- natural mortality during the first year of lation of P. nobilis has been greatly life; the probability of death by natural reduced over the past few decades as a

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | result of recreational and commercial fish- GEROVASILEIOU et al., 2007; personal ing for food, the use of its shell for decora- observations). tive purposes, and incidental mortality by Exploitation: From ancient times until trawlers, bottom nets, or anchoring. early in the 20th century, P. nobilis was Although P. nobilis has become rare in exploited for its byssus, from which an many parts of the Mediterranean, impor- extremely fine and valuable fabric was pro- tant local populations still exist in Hellenic duced, called ‘sea silk’. P. nobilis has also seas (Fig. 4), especially in the Korinthi- been extensively exploited for human con- akos, Evvoikos, and Thermaikos gulfs, the sumption and use of its shell for decorative islands of Chios and Lesvos (NE Aegean), purposes. ‘Semi-quality’ pearls may be NW Kriti, N Karpathos, as well as the Ion- found in some aged fan mussels. Nowadays ian Sea (ZENETOS, 1986; 1996; HAMES it has been declared an endangered species et al., 2001; GALINOU-MITSOUDI et al., and its exploitation has been banned. How- 2006; KATSANEVAKIS, 2006; 2007a; ever, it is still illegally exploited and in cer-

Fig. 4: Population density hotspots of Pinna nobilis.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | tain areas fishing mortality greatly exceeds natural mortality and is a critical determi- nant of the spatial distribution of the species. In Lake Vouliagmeni, mortality ., 2006 due to illegal fishing by free divers has caused a very low population density of the et al species in shallow areas (KATSA- NEVAKIS, 2007b). P. nobilis has been used as a bioindicator in heavy metal bioaccumu- ., 2001 lation studies. In Hellenic seas, it was select- et al ed as an indicator species to study chromi- um contamination in a gulf receiving tan- nery wastes (CATSIKI et al., 1994). Conservation Status: The species is under strict protection according to the 92/43/EC Directive (Annex IV), the Proto- in Hellenic sites. col of the Barcelona Convention (Annex II), and the Presidential Decree 67/1981. A few populations of P. nobilis have been assessed in Hellenic seas, and some synop- Pinna nobilis tic results of reported population density or ) Abundance (inds) Reference abundance are given in Table 2. In two of 2 these areas (Vouliagmeni Lake and Souda Bay) total abundance of P. nobilis has been Table 2 estimated using Density Surface Modelling with Distance Sampling (KATSA- NEVAKIS, 2007a), while in another three areas population densities have been esti- mated (Table 2). Specific mtDNA regions were genetically characterized in pinnids from the Thessaloniki and Thermaikos gulfs, Vouliagmeni Lake and NE Chios. These results could contribute to screening genetic variability and population dynam- (hectares) mean max estimate 95%CI ics, as well as for conservation plans for Estimated population density or abundance of species protection and/or aquaculture investigations (KATSARES et al., 2008).

Pinctada radiata (Leach, 1814)

General: Pinctada radiata is an epifau- nal suspension feeder of the subtidal zone and a fouling species, living attached by its byssus to hard substrata. Usually it attains a Lake Vouliagmeni (Korinthiakos)Lake Vouliagmeni (Korinthiakos) 150.4Souda Bay (Kriti) 150.4NE Chios Isl.(Aegean) 2004NE Kefallonia Isl.(Ionian) 2006 0.0057 0.0045 0.17 0.24 0.088 1471.5 8500 6770 4200 –12900 2006 2007 KATSANEVAKIS, 2006 5450 0.0089–8400 0.06 2000 KATSANEVAKIS, 2007a 0.53 0.07 130900 100600 –170400 KATSANEVAKIS, unpublished data 0.08 GALINOU-MITSOUDI, unpublished data HAMES Epanomi (Thermaikos Gulf) 0.007 2004 1.04 1.3 GALINOU-MITSOUDI Location Study Area Year Density (inds/m

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | length of 50-65 mm, but may exceed 100 as a bioindicator of heavy metal pollution mm in length. It is a protandric hermaphro- (e.g., AL-MADFA, et al., 1998; GÖKSU et dite species with sex inversion occurring in al., 2005). shells of 32-57 mm length. Gonad maturity Conservation Status: It is a successful is controlled by temperature (TLIG invader into the Mediterranean Sea and its ZOUARI & ZAOUALI, 1994). spatial distribution keeps increasing Distribution: Pinctada radiata origi- (ZENETOS et al., 2004; ZENETOS et al., nates from the Indo-Pacific and the Red 2007). There is no relevant legislation for Sea. Following the opening of the Suez its conservation and there is no need for Canal, P. radiata has entered the far east- protective measures. Although there are ern Mediterranean. It was also intention- detailed records of its invasion in the ally imported for mariculture in many Mediterranean and Hellenic seas, no pop- areas of Hellas and Italy during the last ulation assessment has been conducted in century. Up to date, it has been recorded Hellas. as common in the eastern Mediterranean with sporadic occurrences in the western Pteria hirundo (Linnaeus, 1758) basin (ZENETOS et al., 2004). Besides its adaptation to the subtropical environ- General: Pteria hirundo may reach 10 ment of the south-eastern Mediter- cm in length. It has an aggregated distribu- ranean, its tolerance to chemical contam- tion and usually many individuals are ination has enhanced its expansion in found attached by their robust byssus to enclosed polluted ecosystems. In Hellenic gorgonians, on rocky bottoms, or in seas (Fig. 5), it is mostly reported close to clumps of many individuals on muddy bot- areas where it was originally introduced toms in the sublittoral and bathyal zones. for aquaculture (mainly in the Distribution: P. hirundo is found in the Dodekanisos islands, Argolikos, the Mediterranean Sea and the E Atlantic Saronikos, Evvoia, and the island of from the British Isles south to the coasts Lesvos in the N Aegean Sea). Recently, of Portugal. In Hellenic seas, it has been the species was also recorded in the Kyk- reported in the Saronikos, Patraikos, Pag- lades Isles complex (in 2006) and in Kriti asitikos, and N Evvoikos gulfs, Astakos, (in 2003) (ZENETOS et al., 2007). the Kyklades, the N Sporades Islands, and Exploitation: Pearl shells (mother-of- Chalkidiki (Toronaios and S Sithonia). pearl) have been used by the people of Exploitation: Its exploitation is very the Pacific and other regions as utensils, limited, for private or public collections. implements and ornamentation, while the Conservation Status: No population itself has been a basic food item. assessment has been conducted in Hel- Pearl oyster meat is a delicacy in many lenic seas. western cultures. However, mother-of- pearl shells are of little commercial value Family Pectinidae: Chlamys varia (Lin- in the Mediterranean. Because of its toler- naeus, 1758), Crassadoma (Chlamys) mul- ance to chemical contamination and the tistriata (Poli, 1795) ability of to accumulate metals to several orders of magnitude higher than General: Chlamys varia is a the background medium, it has been used with a thin, narrow, elongated oval shell

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | Fig. 5: Known distribution of Pinctada radiata in Hellenic seas. that may reach a length of 80 mm. Its during late spring and summer (PEN~A et colour is very variable and may be white, al., 1996). yellow, orange, pink, red, purple, or Distribution: C. varia is found in the brown. Crassadoma multistriata looks like Mediterranean Sea and the E Atlantic C. varia but is shorter and with distinctive Ocean from the British Isles south to Sene- decoration. It usually attains a length of gal, while C. multistriata is found in the 15–30 mm. Both species live on sandy, Mediterranean Sea and the E Atlantic muddy, or rocky bottoms of the sublittoral Ocean from the Faroe Islands to the coasts zone. Young individuals are either free-liv- near the Mediterranean. In Hellenic seas, ing or attached by byssus threads, while C. varia is quite common and seems to be adults are usually permanently cemented present everywhere on the Hellenic coasts. to hard surfaces by their right valve. Both C. multistriata has been reported in the Sa- species are commonly found in Posidonia ronikos, Patraikos, Korinthiakos, Evvoikos, oceanica beds. The spat settlement occurs and Messiniakos gulfs, the Sporades, the

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | Kyklades, the Dodekanisos, Kymi (Evvoia), Exploitation: Archaeological evidence Kavo Doro, and Chalkidiki. shows that people in Neolithic Europe Exploitation: Both species are edible traded the shells of S. gaederopus to make and in a few European locations (e.g., on bangles, bracelets, belt-buckles and other the French Atlantic coasts) C. varia is tar- ornaments as long as 5000 years ago. The geted by a commercial fishery. In Hellenic shells were mostly harvested from the seas C. varia is locally exploited for human Aegean Sea and then transported into consumption. C. varia has been reported continental Europe. S. gaederopus orna- to highly concentrate trace metals and ments from the Neolithic age have been radionuclides and has been used as a bio- found in many caves and archeological monitor of marine pollution (BUSTA- sites in Hellas (IFANTIDIS, 2006). It is MANTE et al., 2002; BUSTAMANTE & edible and has been collected for human MIRAMAND, 2005). Because of their consumption since antiquity. In some beautiful and variable colouration both C. areas it is considered a delicacy and for this varia and C. multistriata are popular reason was named the ‘royal oyster’. It is among shell collectors. also popular among shell collectors. Conservation Status: No population Conservation Status: No population assessment has been conducted in Hel- assessment has been conducted in Hel- lenic seas. lenic seas. POPPE & GOTTO (1993) pointed out that the species was rather Spondylus gaederopus (Linnaeus, 1758) common till the beginning of the 1980s, when (for unknown reasons) its popula- General: Spondylous gaederopus may tions markedly declined or even disap- exceed a height of 15 cm. It lives on rocky peared in some areas. bottoms of the sublittoral zone from 1 m to about 50 m depth, permanently cemented Crassostrea gigas (= angulata) (Thun- to hard surfaces by its right valve. It prefers berg, 1793) exposed positions on vertical rocks with slow but steadily moving currents (POPPE General: Crassostrea gigas is a large & GOTO, 1993). The species is often oyster whose size may exceed 30 cm (and accompanied by the sponge Crambe may reach 40 cm in exceptional cases). The crambe that covers the upper valve and shell surface is extremely rough, coarsely camouflages its shell. sculptured with concentric ridges and 6–7 Distribution: S. gaederopus is found in thick and bold ribs that commonly give the the Mediterranean and the Atlantic Ocean shell edges an undulating appearance. The near Gibraltar, including Madeira, the shape of the shell varies depending on the Azores, the Canaries, and the Cape Verde environmental conditions. It prefers firm Islands. In Hellenic seas, it has been substrates of the sublittoral zone usually at recorded in the Saronikos, Lakonikos, depths between 5 and 40 m and occasion- Messiniakos, Thermaikos, Evvoikos, ally to depths of about 80 m. It may also be Patraikos, and Korinthiakos gulfs, the Pelo- found on mud or mud-sand bottoms ponnisos, the Kyklades, the Dodekanisos, attached to debris, rocks, or other bivalve Kriti, Astakos, Kymi (Evvoia), Kefallonia, shells. It is an estuarine species and opti- and the Chalkidiki peninsula. mal salinity range is between 20 and 25ò.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | It is a protandrous hermaphrodite and the et al., 2003). It is not a target species in timing of sex change is erratic and season- Hellenic seas, as it has low population den- al. Spawning usually depends on a rise in sities and narrow distribution. water temperatures above 20ÆC, giving Conservation Status: It is classified as approximately 50-100 million eggs in a sin- one of the Worst Invasive Species in gle spawning; larvae are planktonic (FAO, Europe (EEA, 2007). Its dense aggrega- 2008). The biology, ecology, and especially tions limit food and space for other benthic the aquaculture of the species have been species (competition), it destroys habitats extensively studied (~1000 related pub- and causes eutrophication of water. It also lished papers in the Science Citation Index causes causes serious decline in macrofau- between 1970 and 2007). na and zooplankton, but enhances bacte- Distribution: C. gigas originates in the ria, microfauna and meiofauna. No popu- coastal and marine waters of Japan, North lation assessment has been conducted in West Pacific ocean. It has been introduced Hellenic seas. for mariculture into North America, and is now found from SE Alaska to Baja Cali- Donacilla (= Mactra) cornea (Poli, 1791) fornia. In Europe, it was first introduced in the Atlantic (Portugal, Spain, France, Ire- General: Donacilla cornea is an inhabi- land, U.K.), and may be found from tant of the midlittoral soft substratum and Morocco to Norway. It is also widespread is found mostly in moderately exposed in the Mediterranean (France, Adriatic beaches with medium to very coarse sand lagoons, Malta, Spain, Tunisia, Italy, Hel- (MAVIDIS et al., 2006). It is a relatively las, Turkey). In Hellenic seas, it has been small bivalve and its maximum length may reported in the Patraikos and Korinthi- reach 25 mm. The individuals of D. cornea akos gulfs (Zenetos et al, 2004). There is are distinctively patterned and coloured, also unconfirmed information that the exhibiting one of the most striking exam- species was imported for aquaculture into ples of massive polymorphism in nature the Thermaikos Gulf in 2007. (OWEN & WHITELEY, 1988). D. cornea Exploitation: It is both fished and culti- is gonochoristic, its life expectancy is 3–4 vated for human consumption in many years, has an annual reproductive cycle, is countries. Through its potential for rapid mature at a length of approximately 10 growth and its wide ranging tolerance to mm, and spawning (in the Thermaikos environmental conditions, it has become Gulf) occurs between March and July with the oyster of choice for cultivation in many a peak in June (MAVIDIS & KOUKOU- regions of the world. It is the dominant RAS, 2001a). In a study of the midlittoral in a growing United States aqua- soft substratum assemblages of the North culture industry along the Pacific Coast. Aegean Sea (KOUKOURAS & RUSSO, By 2003, global production of this species 1991), among the five different assem- had expanded to 4.38 million tonnes, more blages that were described, one assem- than any other species of fish, mollusc or blage was characterized by the very high crustacean. Nearly 84 % of global C. gigas dominance of D. cornea and the poly- production was in China (FAO, 2008). C. chaete Ophelia bicornis. This assemblage gigas has also been used as a bioindicator was dominant in the greatest part of the of heavy metal pollution (e.g., SHULKIN Thermaikos and Strymonikos gulfs and

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | was located on coarse sandy substrates. Solen marginatus (Pulteney, 1799) Mean densities of D. cornea (± SE) were 1993 (± 350) inds/m2 and 1255 (± 205) General: Solen marginatus is a large, inds/m2 in the D. cornea–O. bicornis assem- shallow-water, elongate clam, which may blages of the Thermaikos and Strymonikos reach a length of 13 cm. It is an inhabitant gulfs, respectively. On Korinos beach of the midlittoral and upper sublittoral (Thermaikos Gulf) the abundance of D. zone, living burried in the sand. It is a filter cornea had two peaks, one at the mean sea feeder and lives in a vertical position in the level (with a density that exceeded 12000 sediment in more or less permanent tubes inds/m2) and a second below the low tide in which they can ascend or descend. With level (MAVIDIS & KOUKOURAS, its specially adopted piston-like foot, 2001b). which can be inflated with blood, it is able Distribution: It is found in the Mediter- to burrow rapidly in its tube or even dig a ranean, the Black Sea, and the E Atlantic new tube. in areas close to the Mediterranean basin. Distribution: S. marginatus is found in In Hellenic seas, D. cornea is very abun- the eastern Atlantic from Mauretania to dant in the N Aegean and is also found in the North Sea, including the Mediter- the Evvoikos, Saronikos, Lakonikos, ranean and the Black Sea. In Hellenic Amvrakikos, and Patraikos gulfs, the Kyk- seas, S. marginatus has been reported from lades, the Ionian and the Cretan Seas. the coasts of the Peloponnisos (both in the Exploitation: The species is collected Ionian Sea and the Aegean Sea) and from by hand in very small quantities and used the Amvrakikos, Thermaikos, Pagasitikos occasionally as fishing bait and rather and Saronikos gulfs. rarely for human consumption. There is Exploitation: In Hellenic seas, it is ample evidence that Donacilla cornea exploited locally both for human consump- along with Tapes decussatus and, Donax tion and as fishing bait. Its fishing is regu- trunculus were among the marine lated by the PD 109/2002. S. marginatus molluscan resources available to has been used as a bioindicator of marine Franchthi's inhabitants around 13,000 BC pollution by organochlorine compounds (SHACKLETON & VAN ANDEL, 2007). such as PCBs and DDTs (THOMPSON et D. cornea has been used as a bioindicator al., 1999). of heavy metal pollution (e.g., REGOLI et Conservation Status: No population al., 1991). assessment has been conducted in Hel- Conservation Status: It is very abun- lenic seas. dant in many midlittoral soft substratum areas. The populations of the Thermaikos Ensis minor (Chenu, 1843) and Strymonikos gulfs have been assessed in the past (KOUKOURAS & RUSSO, General: Ensis minor is a large burrow- 1991; MAVIDIS & KOUKOURAS, ing, shallow-water, elongate clam, which 2001b). There is no relevant legislation. may reach a length of 20 cm. It is an inhab- The regional IUCN status of Critically itant of the midlittoral and upper sublit- Endangered species was proposed for D. toral zone, living buried in the sand mostly cornea in the Black Sea (MICU & MICU, at depths shallower than 4 m, but in the 2006). Mediterranean may be found down to

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | about 25 m (VAN URK, 1964). It is a fil- all of its siphons, which contract rhythmi- ter feeder and lives in a vertical position in cally as they float away in the water col- the sediment in more or less permanent umn; this is probably part of the escape tubes. In the Gulf of Trieste, gametogene- strategy of the animal, as the aborted parts sis begins in September and the maximum will attract the predator and the bivalve reproductive activity occurs between Jan- will have the chance to escape, while its uary and March (VALLI et al., 1983). siphons will soon regenerate (BROMLEY Distribution: E. minor is distributed in & ASGAARD, 1990). the eastern Atlantic, from Norway to Por- Distribution: S. strigilatus is found in tugal, including the Mediterranean Sea. In the Mediterranean Sea and on neighbour- Hellenic seas, E. minor has been reported ing E Atlantic coasts. In Hellenic seas, it from Kriti and the Saronikos, Pagasitikos has been reported in the Saronikos, Ther- and Thermaikos gulfs. maikos, Evvoikos, Patraikos, Korinthi- Exploitation: In Hellenic seas, it is akos, and Kavala gulfs, and the coasts of exploited locally both for human consump- the Peloponnisos, Kefallonia, and Kriti. tion and as fishing bait. In Italy it is exten- Exploitation: Its exploitation is very sively fished and is one of the most impor- limited, for human consumption and for tant fishery resources of the Gulf of Tri- private or public collections. It is also used este (DEL PIERO & DACAPRILE, as fishing bait, especially in the North 1998). Its fishing in Hellenic seas is regu- Evvoikos Gulf. Its fishing in Hellenic seas lated by the PD 109/2002. is regulated by the PD 109/2002. Conservation Status: No population Conservation Status: No population assessment has been conducted in Hel- assessment has been conducted in Hel- lenic seas. lenic seas.

Solecurtus strigilatus (Linnaeus, 1758) (Linnaeus, 1758)

General: Solecurtus strigilatus may General: Pholas dactylus grows up to exceed 10 cm in length. It lives buried in 15 cm long. It lives in midlittoral and shal- the soft substrata of the sublittoral and, low sublittoral areas bored into soft rocks rarely, of the bathyal zones. It burrows rap- such as, shale, peat, chalk, sandstone, stiff idly and to relatively great depths. Adults clay, or wood. It creates a conical burrow, may live 50 cm below the sea floor and which has a narrow entrance and a larger attempts to excavate them from their bur- rounded chamber. Where abundant, its rows release rapid escape reactions where- borings can severely compromise the by the descend obliquely still fur- structural stability of the shore, and can ther into the sediment (BROMLEY & result in increased rates of coastal erosion ASGAARD, 1990). The location of S. (PINN et al., 2005). P. dactylus gives off a strigilatus individuals on the seafloor is blue, luminous secretion when irritated. Its marked by two conspicuous round orifices luminescence has been known since about 1 cm in diameter and 2 cm apart, ancient times (e.g., Pliny the Elder docu- with siphons never observed to project mented the luminescence of P. dactylus from these openings. When in danger of even whilst being eaten) and has a historic predation S. strigilatus will abort parts or place in the biochemistry of biolumines-

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | cence, since it was the first organism used (ALIDROMITI, 2007). A. media is among to describe the oxygen-requiring luciferin- the main cephalopod species of the conti- luciferase reaction that is universal in bio- nental shelf community (GONZALEZ & luminescence. It is a gonochoristic species S NCHEZ, 2002; LEFKADITOU et al., with a spawning period during summer 2004) and is mostly found on sandy and and a life span of 14 years (HILL, 2006). muddy grounds and also in brackish waters Distribution: It is found in the Mediter- (UNSAL et al., 1999). It inhabits a wide ranean, the Black Sea, the Red Sea, and in depth range from the surface down to 600 the E Atlantic from the British Isles south m, but is mostly found at depths <150 m to the coasts of Morocco. In Hellenic seas, (LEFKADITOU, 2007). It performs sea- it has been reported in the Thermaikos, sonal migrations between offshore (in win- Kalloni, Maliakos, Evvoikos, Saronikos, ter) and inshore (in summer) areas Amvrakikos, and Argolikos gulfs, on the (ROPER et al., 1984), as well as diel verti- Ionian coasts of the Peloponnisos, and cal migrations (ZUEV & NESIS, 1971). Kefallonia. Maturation takes place over a wide range Exploitation: It has been extensively of mantle lengths. Spawning in the Me- fished for human consumption and as fish- diterranean occurs at depths of 10-100 m ing bait. mostly in sandy areas or seagrass mead- Conservation Status: P. dactylus was ows. Spawning period lasts from March to once prevalent across the entire Mediter- October in the western part of the ranean and on the Atlantic coast of Mediterranean and potential fecundity Europe, but it has disappeared from most (PF) has been estimated to be about 950- sites due to human collection for food and 1400 eggs (MANGOLD-WIRZ, 1963), bait and/or as a result of marine pollution. whereas in the eastern part spawning The species is under strict protection occurs all year round and PF is some 1000- according to the Bern Convention (Annex 4000 eggs, most of which are released dur- II), and the Protocol of the Barcelona ing multiple spawning with female growth Convention (Annex II). Surprisingly, it is continuing between spawning events included as legally collected fishing bait in (LAPTIKHOVSKY et al., 2002). The diet PD 109/2002 but the Bern Convention of A. media consists of larvae and juveniles (adopted by the Hellenic Law 1335/1983) of fishes, copepods and euphausiids and the Barcelona Convention prevail (ZUEV & NESIS, 1971). A. media is over the PD. No population assessment preyed on by demersal fish of medium and has been conducted in Hellenic seas. large size, mostly by gadids and some elas- mobranchs (ZUEV & NESIS, 1971; 3. Cephalopods BELLO, 1997; VELASCO et al., 2001). Distribution: A. media is common in Alloteuthis media (Linnaeus, 1758) the Mediterranean Sea including the Sea of Marmara. In the E Atlantic its range General: A. media is a small-sized extends from 60Æ to 21Æ N, but it is mainly member of the family Loliginidae, with found in the southern part of the North females attaining a mantle length up to Sea, the Irish Sea, the English Channel, 12 cm and males up to 9 cm. The duration the Bay of Biscay and off the south coast of of its life cycle is about one year the Iberian peninsula. It is widespread in

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | Hellenic seas and has been reported as the females and have higher growth rates most frequently caught cephalopod (PIERCE et al., 1994a,b). Adult body size species on the shelf of the N. Aegean usually reaches 100–650 mm ML in males (LEFKADITOU, 2006), S. Aegean (weight range 155–3700 g) and 175–350 (LEFKADITOU et al., 2003b), and the mm ML in females (weight range Ionian Sea (Table 3). 200–1150 g), while maximum values of 937 Exploitation: A. media is a by-catch of mm ML and 8308 g for males, and 462 mm bottom trawlers and beach-seiners that is ML and 2184 g for females have been mostly discarded by Hellenic trawlers reported from the Azores (MARTINS, (MACHIAS et al., 2001), whereas its dis- 1982). It has an annual life-cycle and is card percentage by Italian and Spanish semelparous, exhibiting ‘intermittent ter- trawlers fishing in the western Mediter- minal spawning’, in which the females lay ranean ranges from 10 to 50 % and is con- 1000 to 23000 eggs in batches and die sidered accidental due to the fast sorting shortly after spawning (PIERCE et al., operations on deck (SARTOR et al., 1994b). L. forbesi migrates to shallower 1998). waters to spawn. A large number of prey Conservation Status: No population species, including various polychaetes, assessment has been conducted in Hel- molluscs, crustacean and fish have been lenic seas. identified in L. forbesi stomachs, with crus- taceans being relatively more important in Loligo forbesi (Steenstrup, 1856) the diet of small squid while larger squid prey predominantly on fish (MARTINS, General: Loligo forbesi is a large-sized 1982; PIERCE et al., 1994b). Among loliginid; males generally grow larger than cephalopod remains, those of L. forbesi

Table 3 Indicative population density (in number of individuals per square km)of cephalopod species of minor importance,estimated during the MEDITS survey of 2005, on shelf (10-200 m)and upper slope (200-500)of the Ionian Sea and the region of Argosaronikos.

Ionian Sea Argosaronikos Species 10-200 m 200-500 m 10-200 m 200-500 m Alloteuthis media 2948 82 453 95 Loligo forbesi 2 38 2 112 Todarodes sagittatus 013014 Todaropsis eblanae 26 124 0 17 Sepia elegans 259 24 231 163 Sepia orbignyana 46 5 777 238 40 428 35 53 Rondeletiola minor 7 296 8 34 Rossia macrosoma 5 17 0 277* Scaeurgus unicirrhus 59 3 232 126 *exceptionally high value observed only in 1995

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | have also been occasionally identified. in sea surface temperature after 1993 and Large demersal fish and some marine subsequent high levels (CHEN et al., mammals have been reported to prey on 2004). No population assessment has been L. forbesi (DALY et al., 2001; SANTOS et conducted in Hellenic seas. al., 2001). Distribution: L. forbesi occurs in the Todarodes sagittatus (Lamarck, 1798) Mediterranean Sea and the NE Atlantic from 20 ÆN to 63 ÆN, where it is more General: T. sagittatus is a large squid of abundant in the northern part of its range. the family Ommastrephidae. Its mantle In the temperate part of its distributional length attains 75 cm, but usually in trawl range it is mainly found over the shelf, but catches it does not exceed 35 cm, as imma- in subtropical areas and in the Mediter- ture individuals make up the vast majority ranean in deeper waters, reaching depths of specimens (BORGES & WALLACE, of 720 m in the Ionian Sea 1993; QUETGLAS et al., 1998; LORDAN (LEFKADITOU et al., 2003a). It is wide- et al., 2001; LEFKADITOU, unpublished spread in Hellenic seas, being more com- data). This neritic and oceanic squid mon in the neritic zone of regions with occurs from surface to near-bottom water abrupt waters like the Ionian Sea and the layers over the continental shelf and slope area of the Kyklades, Kriti and the up to 2500 m. It carries out vertical move- Dodekanisos islands, while it is scarce ments between the surface at night and north of 40Æ ¡ in the Aegean Sea near the sea bottom during the day (KOUTSOUBAS et al., 1999; LEFKA- (MANGOLD-WIRZ, 1963; KORZUM et DITOU, 2006). al., 1979). Estimates of its life span range Exploitation: L. forbesi is largely fished from 9 to 14 months. Its life span was as a by-catch from bottom trawl fisheries. found longer in higher latitudes and about Fishery statistics do not distinguish L. a month longer in females than in earlier forbesi from L. vulgaris, since they are of maturing males. T. sagittatus is a fast grow- similar appearance and equal commercial ing and early maturing squid, its matura- value. However, landings in Scottish tion found earlier and subsequently its waters can generally be assumed to be L. growth rate lower in NW African waters forbesi (PIERCE et al., 1994a), while those (ARKHIPKIN et al., 1999) compared with in the Mediterranean mainly consist of L. those of northernmost areas (LORDAN et vulgaris (GUERRA et al., 1994). There is al., 2001; QUETGLAS & MORALES- no estimation of the marketable propor- NIN, 2004). Mature individuals are pres- tion of L. forbesi in the landings of lolig- ent all year round but spawning appears to inids in Hellenic fisheries. peak in autumn-winter (QUETGLAS et Conservation Status: Since the early al., 1998; ARKHIPKIN et al., 1999). 1990s, L. forbesi has apparently disap- Potential fecundity estimates range from peared from much of the southern part of 200000 to 950000 eggs (LAPTI- its range in the eastern Atlantic (France KHOVSKY & NIGMATULLIN, 1999; and Portugal), while at the same time its LORDAN et al., 2001). Its diet consists of abundance in northern waters (Scotland) fish, crustaceans and cephalopods (in has increased. This shift in its distribution decreasing order of importance), with can- range may be associated with the increase nibalism also identified (QUETGLAS et

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | al., 1999; LORDAN et al., 2001). Younger high commercial value. It is marketed T. sagittatus in some areas have shown fresh, frozen, salted or dried and is also preference for small pelagic prey in com- used as bait in finfish fisheries. parison to specimens in more advanced Conservation Status: No population adult stages (PIATKOWSKI et al., 1998), assessment has been conducted in Hel- a fact that indicates their preying mostly in lenic seas. No official statistics are avail- upper water layers. T. sagittatus is the most able for T. sagittatus in any ICES area or abundant prey in the stomachs of sword- the Mediterranean Sea, because landings fish from the eastern Mediterranean sug- of ommastrephid species (Illex coindetii, gesting a high abundance of the species Todaropsis eblanae, Todarodes sagittatus, particularly during summer (PERISTE- and Ommastrephes bartramii) are all RAKI et al., 2005). Apart from swordfish, pooled together (PEREIRA et al., 1998; other large pelagic fishes, sharks and JEREB, 2000). Thus, no population cetaceans are important predators of this assessment is possible based on commer- species (ROPER et al., 1984). cial catches. Distribution: T. sagittatus is broadly dis- tributed throughout the north and north- Todaropsis eblanae (Ball, 1841) east Atlantic, from the Arctic Ocean to about 13ÆS and to approximately 40ÆW, General: T. eblanae is a demersal mid- including the North Sea and the Mediter- sized ommastrephid squid, with a maxi- ranean Sea, where it is more abundant off mum recorded mantle length 29 cm for the Tunisian and Libyan coasts, in the females and 23 cm for males (ROBIN et straits of Sicily and Messina, in the north- al., 2002). It is associated with sandy and ern Tyrrhenian Sea and in the central muddy bottoms and it mostly occurs on the Adriatic (BELLO, 1985; JEREB & shelf break and upper slope, not ascending RAGONESE, 1994; RELINI et al.,1999). to the surface or approaching the shore. Its In Hellenic seas it has been reported in populations have been considered as the Aegean (N< 40Æ 30’) and the Ionian indicative of the cephalopod community at Seas, being caught at depths between 120 these depths, in the south-eastern Atlantic and 800 m, more frequently during sum- (ROELEVELD et al., 1992), the Balearic mer and autumn, but never exceeding 10 Sea (QUETGLAS et al., 2000), the north individuals per hour of trawling Tyrrhenian (SANCHEZ et al., 1998), the (LEFKADITOU et al., 2003a). south-eastern Adriatic (KRSTULOVIC- Exploitation: A directed fishery by jig- SIFNER et al., 2005) and the eastern Ion- ging machines exists for this species off ian Sea (LEFKADITOU et al., 2001). Norway, with highly variable annual yields, Mature individuals are found all year which peaked in the early 1980s reaching round, with maximum incidence of mature 10,000 metric tons. It usually constitutes of females from mid-spring to early autumn a by-catch of trawl fisheries and occasion- (GONZALEZ et al., 1994; RELINI et al., ally of long-line and pelagic net fisheries, 1999; ROBIN et al., 2002). T. eblanae fol- while it is also fished with hand-jigs by arti- lows the multiple pattern of spawning, its sanal and sports fishermen, mainly during PF decreasing with latitude and reaching summer over deep waters. In some 275 000 for larger females off West Africa Mediterranean countries it has a relatively (LAPTIKHOVSKY & NIGMATULLIN,

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | 1999) but not exceeding 28 000 in Scottish been conducted during the summer over waters (HASTIE et al., 1994). Teleostean the continental shelf and slope since 1994. fish have been found to compose the major part of its stomach contents, with Sepia elegans (Blainville, 1827) crustacean and cephalopod (including its own species) percentages varying by geo- General: Sepia elegans is a small cuttle- graphic area and specimen size (RASERO fish; males attain a mantle length of 72 mm et al., 1996; LORDAN et al., 1998). Its and females up to 89 mm, their maximum most important predators are teleost fish, weight not exceeding 60 g (REID et al. toothed whales and selachians. 2005). S. elegans is a sublittoral species of Distribution: T. eblanae is distributed in sandy and sandy-muddy bottoms of the the Mediterranean Sea and E Atlantic, continental shelf and the upper slope, from 61Æ 15' N to 40Æ S, as well as in the being among the dominant cephalopod SW Pacific and the SW Indian off species on the upper shelf (SANCHEZ et Australia. It is widespread in Hellenic seas, al. 1998; KRSTULOVIC-SIFNER et al., where it has been recorded from 100 to 850 2005; LEFKADITOU, 2007). In the Sea m of depth, being more abundant on the of Marmara, S. elegans has also been upper slope of regions with steep waters, recorded in brackish waters (18-25 psu) like the Toroneos Gulf (LEFKADITOU, (UNSAL et al., 1999), which indicates a 2006) and the Ionian Sea (Table 3). high degree of salinity tolerance. Mature Exploitation: T. eblanae is a by-catch in individuals are present all year round, with bottom trawl fisheries that is either dis- maximum incidence during summer and carded or landed with Illex coindetii. No autumn in the north Aegean Sea (D’ official statistics or research estimates exist ONGHIA et al., 1996), while during spring on the landed catch of T. eblanae from and summer in the western Mediterranean Hellenic waters. Studies of trawl fishery (MANGOLD-WIRZ, 1963). A seasonal landings have shown a marked seasonality migration has been reported in some areas of the species catches, with a peak during with individuals spending their winter in summer in the northern Tyrhenian Sea deep waters (200–400 m) and then migrat- (BELCARI et al., 1998) and almost equal ing into shallower waters in spring and levels of T. eblanae and I. coindetii landings summer, which however, has not been from northern Spanish Atlantic waters observed in other areas (RELINI et al., (GONZALEZ et al., 1994). 1999). Feeding studies conducted in the E. Conservation Status: Due to its pooling Atlantic have shown the preference of S. with other ommastrephid species in fishery elegans for small crustaceans, fish and statistics, it is not possible to assess catch polychaetes (REID & JEREB, 2005). Its levels as an index of stock biomass. Indica- remains have been frequently found in the tive trends of density and biomass indices stomach contents of Scyliorhinus canicula for this species on the fishing grounds and Raya clavata from the Aegean Sea across the north coasts of the Mediter- (KABASAKAL, 2002). ranean Sea might probably be feasible Distribution: S. elegans is found in the through the database of the MEDITS eastern Atlantic from Ireland and western (International Bottom Trawl Surrvey in Scotland (~50ÆN) to the Sahara Bank and the Mediteranean) surveys, which have Namibia (21ÆS) and throughout the

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | Mediterranean Sea including the brackish Mediterranean Sea, existing also in the waters of the Sea of Marmara (UNSAL et brackish waters of the Sea of Marmara al., 1999) It is common in Hellenic seas (REID et al., 2005). Its bathymetric range where it has been reported at depths of extends from a few meters down to middle between 20 and 600 m. slope, with its deepest record from a haul Exploitation: S. elegans is a by-catch at 700-770 m in the southern Aegean Sea species of trawl fisheries; its catches (MEDITS – International bottom trawl exhibiting a marketable proportion in Por- survey in the Mediterranean, unpublished tugal, Spain, Italy and Hellas (SARTOR et data). However, S. orbignyana is more fre- al., 1998; MACHIAS et al., 2001; quently caught on lower continental shelf, SENDAO et al., 2002) depending on the being an indicative species of cephalopod landing port and sale perspectives of the assemblages at these depths (SANCHEZ species, which is usually marketed mixed et al., 1998; QUETGLAS et al., 2002; with S. orbignyana and small S. officinalis. LEFKADITOU, 2006). Its remains have In some areas S. elegans represents a sig- been noted among prey species of Raya nificant percentage of the catch and con- clavata from the Aegean Sea stitutes a valuable resource locally (REID (KABASAKAL, 2002) and the demersal et al., 2005). fish community in the Bay of Biscay Conservation Status: No population (VELASCO et al., 2001). assessment has been conducted in Hel- Exploitation: Sepia orbignyana repre- lenic seas or other geographical areas. sents a common trawl by-catch, regularly present in Italian and Spanish markets Sepia orbignyana (Ferussac, 1826) (RELINI et al., 1999), being marketable also in Hellas (MACHIAS et al., 2001). It General: Sepia orbignyana is a cuttle- is marketed frozen and fresh, usually fish of medium size, with maximum mantle mixed with S. elegans and small individuals length recorded in the Mediterranean Sea of the common cuttlefish Sepia officinalis, 9.3 cm for males and 9.6 cm for females so that there are no separate statistical (REID et al., 2005). Mature females are data for this species’ landings. caught from spring to autumn while Conservation Status: Density and bio- mature males are collected during all sea- mass data have been collected systemati- sons (RELINI et al., 1999; cally for Sepia orbignyana during the trawl LEFKADITOU, 2006), a fact that indi- surveys conducted in the Mediterranean cates earlier maturation of the males. waters of European countries since the Mature females carry about 400 oocytes summer of 1994, but are not sufficient for and males about 100 spermatophores stock assessment of this short-lived (REID et al., 2005). According to a study species. of its feeding in the Tuscan waters, crus- taceans constitute its preferred prey, fol- Rossia macrosoma (Delle Chiaje, 1830) lowed by fish, cephalopods and echino- derms (RELINI et al., 1999). General: Rossia macrosoma is the Distribution: Sepia orbignyana extends largest sepiolid species in the Mediter- from Irish waters south to Angola (17Æ S) ranean Sea; its mantle length attaining 5.4 in the eastern Atlantic and throughout the cm for males and 8.4 cm for females

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | (MANGOLD-WIRTZ, 1963; the Mediterranean (MEDITS), conducted LEFKADITOU, 2006). Mature speci- during the summer over the continental mens of both sexes are found during all shelf and slope of European countries seasons, indicating an extended spawning since 1994. period that probably peaks in spring and autumn (REID & JEREB, 2005). Poten- Rondeletiola minor (Naef, 1912) tial fecundity reaches 208 eggs, while counts of spermatophores inside a male’s General: Rondeletiola minor is one of Sac of Needham do not exceed 150 the smaller members of the family Sepioli- (LEFKADITOU, unpublished data). No dae, attaining a mantle length up to 2.3 cm information is available on its feeding (REID & JEREB 2005) and recognized by habits. Remains of R. macrosoma have the roundish pair of light organs present been identified among the stomach con- inside the mantle cavity on the ventral side tents of demersal fishes and chon- of the ink sac, easily seen from the ventral drichthyans (BELLO, 1997; VELASCO et mantle opening. Mature individuals are al., 2001). found throughout the year, with females ma- Distribution: It is widespread in the turing at 1.3-1.7 cm and males at 1.1-1.3 cm Mediterranean Sea and across the north- (SALMAN & KATAGAN, 1996; LEFKA- eastern Atlantic, extending also into DITOU & KASPIRIS, 2004), a fact waters around Greenland (REID & explaining the almost exclusive presence of JEREB, 2005). It has been reported at mature individuals among males collected depths ranging between 32 and 900 m, by selective bottom trawl nets. Potential caught more frequently on upper slopes. It fecundity of females reaches 160 eggs has been found in the Aegean Sea and the (LEFKADITOU unpublished data) while Ionian Sea, at depths between 72 and 700 in males 280 spermatophores have been m. Mature individuals of both sexes are counted on average (SALMAN & distributed mainly on upper slopes where- KATAGAN, 1996). R. minor shows an as younger individuals are mainly found on aggregation forming behaviour, being the lower shelf (LEFKADITOU & among the cephalopod species character- KASPIRIS, 2004; ROSA et al., 2006). izing shelf-break areas (GONZALEZ & Exploitation: It consistently exhibits SANCHEZ, 2002; LEFKADITOU, 2006). both a marketable and discarded amount R. minor remains have been reported in in Hellenic, Italian and Spanish trawl fish- the stomach content of Raya clavata eries (SARTOR et al., 1998; MACHIAS (BELLO, 1997). et al., 2001). There are no official statistics Distribution: Rondeletiola minor is for R. macrosoma commercial catches, widespread in the Mediterranean Sea which are usually landed mixed with other including the brackish waters of the Sea of sepiolids and small eledonids. Marmara (UNSAL et al., 1999) and in the Conservation Status: There are no eastern Atlantic from the north coast of studies targeting assessment of this the Iberian Peninsula up to the waters off species’ stocks. Data on the number of Namibia (30oS) (REID & JEREB 2005). individuals and their total weight by haul It has been reported at depths ranging have been collected in the framework of from 17 to 900 m, with its deepest records the International bottom trawl survey in reported from the northeastern Ionian Sea

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | (LEFKADITOU et al., 2003a). In the Aristaeomorpha foliacea (BELLO & Aegean Sea it has been caught down to PIPITONE, 2002). 770 m, with the greatest part of its popula- Distribution: Sepietta oweniana is wide- tion distributed over the continental slope spread in the Mediterranean Sea including (LEFKADITOU et al., 2003b, LEFKADI- the southwestern part of the Sea of Mar- TOU & KASPIRIS, 2004). mara (UNSAL et al., 1999) and in the Exploitation: Rondeletiola minor is a northeastern Atlantic from Norway to by-catch species of trawl fisheries, known Mauritania (15o N), while its occurrence in to be consumed only in Italy (SARTOR et the Indian Ocean is doubtful (REID & al., 1998). JEREB, 2005). It is present on soft bot- Conservation Status: No population tom, of continental shelf and slope, but assessment has been conducted for this more abundant on the upper slope, where species across its distributional range. it has usually been found among the dom- inant species of cephalopod assemblages Sepietta oweniana (Pfeffer, 1908) (SANCHEZ et al., 1998; BIAGI et al., 2002; GONZALEZ & SANCHEZ, 2002; General: Sepietta oweniana is the most LEFKADITOU, 2006). It is common in abundant species of the family. Hellenic seas, where it has been recorded Its mantle length reaches 5 cm in the at depths of between 20 and 820 m, being North Sea but does not exceed 4 cm in the the most abundant cephalopod species on Mediterranean Sea, with females general- the eastern Ionian slope (Table 3). ly maturing at around 2.5 cm and males at Exploitation: Sepietta oweniana consti- around 2.0 cm. Mature individuals are tutes the major part of landed sepiolids, present throughout the year with maxi- particularly appreciated in some markets mum incidence of mature females of the western Mediterranean, represent- observed from spring to autumn (RELINI ing a significant by-catch of trawl fisheries, et al., 1999; LEFKADITOU & especially in Norwegian lobster fishing KASPIRIS, 2004). Fecunded eggs are grounds (RELINI et al., 1999). According released in one or several spawning events to the results of a study of Hellenic trawl and potential fecundity ranges from 50 to fisheries’ discards for 1995-1998, by-catch- 240 eggs, while counts of male’s sper- es of this species were only discarded matophores range between 50 and 824 (MACHIAS et al., 2001). (SALMAN, 1998; LEFKADITOU, 2006). Conservation Status: Studies on the S. oweniana feeds primarily on crus- state of its exploitation and population taceans, while cephalopod and fish pro- assessment are lacking. portions in their stomach contents vary considerably by geographic area and depth Scaeurgus unicirrhus (Orbigny, 1840) of capture (LEFKADITOU, 2006). S. oweniana has been found in the stomachs General: Scaeurgus unicirrhus is a of Stenella coeruleoalba (SANTOS et al., medium-sized octopus, with mantle length 1995), Galeus melastomus (BERGSTROM reaching 12 cm for females (MANGOLD & SUMMERS, 1983; BELLO, 1997), Chi- –WIRTZ, 1963) and 8 cm for males maera monstrosa (BELLO, 1997), Raya (LEFKADITOU, 2006). Mature males clavata (KARABASAKAL, 2002) and are met throughout the year but mature

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | females have been caught in the Mediter- Conclusion ranean Sea from early spring to early autumn (MANGOLD –WIRTZ, 1963; People who exploit marine resources LEFKADITOU, unpublished data). choose among them according to techno- Potential fecundity of females is about logical, economic, and social considera- 1000 eggs, while counts of spermatophores tions. Prehistoric selectivity in their in the males’ spermatophoric sac do not exploitation has often been postulated. exceed 10. Prey spectrum of the species Many molluscs, although not commercial- indicates its association with the bottom, ly important, have been exploited for made up mainly of carideans crustaceans human consumption or for other uses and in smaller proportions of teleosts, (fish bait, shell collections, the jewellery cephalopods and polychaetes (RELINI et industry). Although for some species al., 1999). there are indications of substantial popu- Distribution: Scaeurgus unicirrhus is a lation decline, there is a lack of scientific cosmopolitan species, scattered in tropical effort towards a better understanding of and warm temperate waters, associated their ecology and assessing the status of with sandy and coralline bottoms their populations. The situation is even (ROPER et al., 1984). It has been report- worse for species of no importance for ed at depths between 30 and 850 m, being humans (not evaluated in this review). Sci- relatively more abundant on lower shelf entific effort and financing targeted a few and upper slope (SANCHEZ et al., 1998; commercial species (KOUTSOUBAS et QUETGLAS et al., 2000; GONZALEZ & al., 2007; LEFKADITOU, 2007), while SANCHEZ, 2002; LEFKADITOU et al., molluscs of minor or no commercial 2003a). It is widespread in Hellenic seas, importance are neglected. For most of presenting a wider depth range in the Ion- these species there is no population ian Sea (43-850 m) in comparison to that assessment in Hellenic seas with only a (30-550 m) in the Aegean Sea, where it few exceptions on a local scale (e.g., Pinna seems to be relatively more abundant nobilis, Lithophaga lithophaga, Donacilla (Table 3). cornea). However, many of these species Exploitation: Scaeurgus unicirrhus is a are important constituents of the Mediter- by-catch of bottom trawlers that is mostly ranean ecosystems and as they are under a discarded by Hellenic trawlers variety of pressures nowadays (marine (MACHIAS et al., 2001) and only occa- pollution, alien species’ invasions, climate sionally marketed with musky change, destructive fishing methods, over- (LEFKADITOU, unpublished data). In exploitation) there is a pressing need to the western Mediterranean, its discard assess the status of their populations and percentage by Italian and Spanish trawlers act towards their conservation. Existing ranges from 0 to 50 % (SARTOR et al., legislation is not enforced in practice and 1998), but its economic value is lower than is extensively violated, thus the conserva- that of other commercial octopod species tion of the molluscan species is not (RELINI et al., 1999). ensured. In addition, a good knowledge of Conservation Status: No population the ecology and status of the aforemen- assessment has been conducted in Hel- tioned species and appropriate conserva- lenic seas or other geographic areas. tion plans are needed.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | An important goal in the next decade structure and growth of the squid is to network targeted collection data Todarodes sagittatus (Cephalopoda: nationally and internationally so this Ommastrephidae) in north-west invaluable source of biodiversity informa- African waters. Journal of the Marine tion can be accessed for conservation and Biological Association of the United planning purposes. Kingdom, 79: 467-477. BELCARI, P., SARTOR, P. & DE Acknowledgements RANIERI, S., 1998. I cefalopodi nello sbarcato commerciale con reti a We would like to thank Dimitris Pour- strasciso del Mar Tirreno settentri- sanidis for the preparation of the maps onale. Biologia Marina Mediterranea, 5: presented in this paper. Evi Vardala- 318-325. Theodorou has provided useful informa- BELLO G., 1985. Su una raccolta di tion for some species. cephalopodi pescati nel mesobatiale del golfo di Taranto. Bolletino Malaco- References logico, 21/10-12: 275-280. BELLO, G., 1997. Cephalopods from the ABBOTT, T.R., 1960. The Strom- stomach contents of demersal chon- bus in the Indo-Pacific. Indo-Pacific drichthyans caught in the Adriatic Sea. Mollusca, 1: 33–146. Vie Milieu, 47: 221-227. ALIDROMITI, C., 2007. Study of age and BELLO, G. & PIPITONE, C., 2002. Preda- growth of Alloteuthis media tion on cephalopods by the giant red (Cephalopoda: Loliginidae) in the shrimp Aristaeomorpha Foliacea. Jour- North Aegean Sea. Investigation of nal of the Marine Biological Association geographical and gender differences of the United Kingdom, 82: 213-218. in the statolith morphology and the BERGSTROM, B. & SUMMERS, W., mantle growth. MSc dissertation, 1983. Sepietta oweniana. In: P.R. Boyle University of Athens, Athens, Hellas, (ed.) Cephalopod life cycles. Volume 68 pp (in Hellenic, English abstract). 1, Academic press: 75-91. AL-MADFA, H., ABDEL-MOATI, BIAGI, F., SARTOR, P., ARDIZZONE, M.A.R. & AL-GIMALY, F.H., 1998. G.D., BELCARI, P., BELLUSCIO, Pinctada radiata (Pearl Oyster): a A. & SERENA, F., 2002. Analysis of bioindicator for metal pollution moni- demersal fish assemblages of the Tus- toring in the Qatari waters (Arabian cany and Latium coasts (north- Gulf). Bulletin of Environmental Cont- Western Mediterranean): community amination and Toxicology, 60: 245-251. structure and biodiversity. Scientia ANTONIADOU C., KOUTSOUBAS D. Marina, 66 (suppl. 2): 233-242. & CHINTIROGLOU, C., 2005. Mol- BORGES, T.C. & WALLACE, J.C., 1993. luscan fauna from infralittoral hard Some aspects of the fishery biology of substrate assemblages in the North the Ommastrephid squid Todarodes Aegean Sea. Belgian Journal of Zoolo- sagittatus (Lamarc, 1798) from the gy, 135 /2: 119-126. Northeast Atlantic. In: Recent ARKHIPKIN, A., LAPTIKHOVSKY, V. advances in Fisheries Biology, edited by & GOLUB, A., 1999. Population T. Okutani, R.K. O’Dor & T.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 27/09/2021 17:44:35 | Kubodera, Tokay University Press, NOAA Technical Report NMFS, 35: 1-9. Tokyo, pp. 25-36. CATSIKI, V.A., KATSILIERI, C. & BOUDOURESQUE, C.F., 2004. Marine GIALAMAS, V., 1994. Chromium biodiversity in the Mediterranean: sta- distribution in benthic species from a tus of species, populations and com- gulf receiving tannery wastes (Gulf of munities. Travaux scientifiques du Parc Geras - Lesbos Island, Greece). national de Port-Cros, 20: 97-146. Science of the Total Environment, 145: BROMLEY, R.G. & ASGAARD, U., 173-185. 1990. Solecurtus strigilatus: a jet-pro- CHEN, C.S., PIERCE, G.J., WANG, J., pelled burrowing bivalve. p. 313-320. ROBIN, J-P., POULARD, J.C., In: The Bivalvia – Proceedings of a PEREIRA, J., ZUUR, A.F., BOYLE, Memorial Symposium in Honor of Sir P.R., BAILEY, N., BEARE, D.J., Charles Maurice Yonge, Edinburg, JEREB, P., RAGONESE, S., 1986, B. MORTON, Hong Kong MANNINI, A., & ORSI RELINI, L., University Press, Hong Kong. 2006. The apparent disappearance of BROWN, K.M. & RICHARDSON, T.D., Loligo forbesi from the south of its 1987. Foraging ecology of the south- range in the 1990s: trends in Loligo spp. ern oyster drill Thais haemastoma abundance in the northeast Atlantic (Gray): constraints on prey choice. and possible environmental influences. Journal of Experimental Marine Biology Fisheries Research, 78: 44-54. and Ecology, 114: 123-141. CHINTIROGLOU, C., ANTONIADOU, BROWN, K.M. & ALEXANDER Jr, J.E., C., VAFIDIS, D. & KOUTSOUBAS, 1994. Group foraging in a marine gas- D., 2005. A review of the biodiversity tropod predator: Benefits and costs to of hard substrate invertebrate commu- individuals. Marine Ecology Progress nities in the Aegean Sea. Mediter- Series, 112: 97-105. ranean Marine Science, 6/2: 51-62. BUSTAMANTE, P & MIRAMAND, P., CONTI, M.E. & CECCHETTI, G., 2003. 2005. Subcellular and body distribu- A biomonitoring study: trace metals in tions of 17 trace elements in the varie- algae and molluscs from Tyrrhenian gated scallop Chlamys varia from the coastal areas. Environmental Research, French coast of the Bay of Biscay. 93: 99-112. Science of the Total Environment, 337: DALY, H. I., PIERCE, G. J., SANTOS, 59-73. M. B., ROYER, J., CHO, S. K., G., S., BUSTAMANTE, P., GERMAIN, P., ROBIN, J.-P. & HENDERSON, S., LECLERC, G. & MIRAMAND, P., 2001. Cephalopod consumption by 2002. Concentration and distribution fish in UK waters. Fisheries Research, of 210Po in the tissues of the scallop 52: 51-64. Chlamys varia and the mussel Mytilus DELAMOTTE, M. & VARDALA- edulis from the coasts of Charente- THEODOROU, E., 2007. Shells from Maritime (France). Marine Pollution Greek seas. Goulandris Natural Histo- Bulletin, 44: 997-1002. ry Museum, Athens. BUTLER, P.A., 1985. Synoptic review of DEL PIERO, D. & DACAPRILE, R., the literature on the southern oyster 1998. The alternating recruitment pat- drill Thais haemastoma floridana. tern in Ensis minor, an exploited

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