Mediterranean Marine Science Vol. 4/1, 2003, 79-86

Cephalopods distribution in the southern Aegean Sea

E. LEFKADITOU1, P. PERISTERAKI2, P. BEKAS1, G. TSERPES2, C-Y. POLITOU1 and G. PETRAKIS1

1 Hellenic Centre for Marine Research, Agios Kosmas, Helliniko, 16604 Athens, Greece e-mail: [email protected]

2 Institute of Marine Biology of Crete, PO Box 2214, 71003 Iraklio, Greece e-mail: [email protected]

Abstract

The present study concerns faunal composition and distribution of cephalopods in the southern Aegean Sea (35Æ 13’ 19’’- 37Æ 55’ 25’’ N, 23Æ 00’ 15’’ - 28Æ 15’ 37’’ E). Samples were collected from 708 hauls, obtained by an experimental bottom trawl net during eight surveys carried out in the summers of the years 1994-2001, as well as by commercial trawl net during four surveys carried out in September 1995, December 1995, May and September 1996. The hauls were performed at depths ranging from 16 to 778 m. A total of 34 species of cephalopod in 12 families were identified, including 11 oegopsid squid, 3 myopsid squid, 7 octopod, 3 cuttlefish and 10 sepiolid. Trawling with the experimental net resulted in the capturing of some uncommon pelagic species, such as Ctenopteryx sicula and Octopoteuthis sicula, which were recorded for the first time in the Aegean Sea. Most of the species showed a wide depth and geographical range. The species: Sepia officinalis, Sepietta neglecta, Sepietta obscura and Sepiola rondeleti were caught only on the continental shelf, whereas the Ancistroteuthis lichtensteini, Bathypolypous sponsalis, Brachioteuthis riisei, Chiroteuthis veranyi, Ctenopteryx sicula, Heteroteuthis dispar, Histioteuthis reversa, Neorossia caroli and Pyroteuthis margaritifera were found only on the slope. The rest of the species extended in both continental shelf and slope. The spatial distribution of different species groups is discussed in relation to the hydrology and topography of the study area and the species ecology.

Keywords: Cephalopoda, Eastern Mediterranean.

Introduction not only for fisheries, but also for the understanding of their significance in energy Cephalopods constitute increasingly and material flow in marine ecosystems important resources for human consumption (PIATKOWSKI et al., 2001). and a principal food for many top predators The earliest available scientific literature (CLARKE, 1983).The knowledge of their on the cephalopod fauna of the Aegean Sea abundance and distribution is fundamental, comes from the Danish Oceanographical

Medit. Mar. Sci., 4/1, 2003, 79-86 79 Expeditions to the Mediterranean and adjacent in September 1995, December 1995, May and seas in 1910 (DEGNER, 1926). Nevertheless September 1996 in the context of a national prior to the 1990s references to cephalopods project for the "Stock Assessment of are scanty (STERGIOU et al., 1997) and only Commercially Important Demersal Species in within the last decade their distribution has the southern Aegean Sea". The area been regularly monitored, based mainly on investigated and the sampling stations for each trawl surveys carried out in the northern and one of the projects are shown in Figure 1. eastern part of the Aegean Sea (D’ ONGHIA During the eight MEDITS surveys 582 et al., 1991, 1996; SALMAN et al., 1997, 2002). hauls in total were performed, in depths This is a preliminary study on the species ranging between 34 and 778 m. The composition and distribution of cephalopods experimental trawl net used had a mesh size in the southern Aegean Sea. of 10 mm at the cod end and its horizontal and vertical net opening, measured by means of a Material and Methods SCANMAR sonar system, ranged between 13- 18.8 m and 1.9-2.8 m respectively. The data analyzed come from bottom trawl During the 4 surveys of the national project surveys carried out in summer during 1994- 161 hauls were carried out by a commercial 2001 in the framework of the "International trawl net having a cod end of 16 mm mesh size, Bottom Trawl Survey in the Mediterranean horizontal net opening 11.2-18.2 m and vertical (MEDITS)" and from four surveys carried out net opening 1.1-1.3 m. The minimum and

Fig. 1: Map of the southern Aegean Sea showing the position of the hauls performed during the trawl surveys of "MEDITS" (s) and the national project (l).

80 Medit. Mar. Sci., 4/1, 2003, 79-86 maximum investigated depths were 30 and 635 Onychoteuthis banksii. Such pelagic species are m respectively. very rarely caught by conventional fishing gear The species were identified following the although they form a considerable part in the keys in MAGNOLD & BOLETZKY (1987) and diet of marine mammals, sharks and large Bello (1995). Preliminary analysis of the pelagic fish (CLARKE, 1983; ORSI-RELINI et composition and distribution of the al., 1994; BELLO, 1996). cephalopod catches in the southern Aegean Trawling with the experimental net resulted Sea was carried out. in the capture of 32 species, whereas 25 species were present in the commercial net catches Results and Discussion (Table 1). Bathypolypous sponsalis and Sepietta obscura were the only caught species exclusively by the commercial trawl net. The higher Cephalopods were collected in 645 of the number of species found in the catches of the 708 valid hauls. A total of 34 species of experimental trawl net results from the capture cephalopod were identified, including 11 of the pelagic species Ancistroteuthis oegopsid squid, 3 myopsid squid, 7 octopod, 3 lichtensteini, Brachioteuthis riisei, Chiroteuthis cuttlefishe and 10 sepiolid. Remains of female veranyi, Ctenopteryx sicula, Heteroteuthis dispar, shells of Argonauta argo were also recorded. The Histioteuthis reversa, Pyroteuthis margaritifera, frequencies of the species appearance in the Octopoteuthis sicula and Todarodes sagittatus. catches of the different trawl nets are listed in These mesopelagic species carry out diel Table 1, where indications of the depth range vertical migrations, remaining close to the and the geographical rectangle of capture for bottom during the daytime (ROPER & each species are included. Previous records of YOUNG, 1975). The wider vertical opening of cephalopod species in the southern and the the experimental net seems to be more effective northern Aegean Sea are also reported in in capturing mesopelagic cephalopods, as Table 1. suggested by MAIORANO et al. (1999). A total of 25 species have been known from Twenty-four and 29 species were recorded the southern Aegean Sea (Table 1). Among respectively at the continental shelf and slope, the 34 species recorded during our surveys, 16 of which 20 were distributed in both depth are new findings for the studied area (Table 1) zones (Table 1). Four species, Sepia officinalis, and two of them, Ctenopteryx sicula and Sepietta neglecta, Sepietta obscura and Sepiola Octopoteuthis sicula, have been caught for the rondeleti, were caught only at depths shallower first time in the Aegean Sea. Thus with this than 200 m, however other species typically study the cephalopod fauna has increased to distributed on the continental shelf, like 49 species for the Aegean Sea and 41 for the Eledone moschata, Octopus vulgaris, Sepiola southern part, reaching 96% and 80% intermedia and Loligo vulgaris were fished down respectively of the eastern Mediterranean to 223, 270, 293 and 335 meters respectively. fauna which is represented by 51 species The species fished exclusively on the slope were (SALMAN et al., 2002). the pelagic sepiolid Heteroteuthis dispar, the The nine species previously recorded in the mesopelagic squids Ancistroteuthis lichtensteini, southern Aegean Sea but not found during Brachioteuthis riisei, Chiroteuthis veranyi, these surveys (Table I), include a species of the Ctenopteryx sicula, Histioteuthis reversa, littoral zone, Octopus macropus, the epipelagic Pyroteuthis margaritifera and the bathy-benthic Thysanoteuthis rhombus, Ocythoe tuberculata, Neorossia caroli and Bathypolypous sponsalis; Tremoctopus violaceus and the mesopelagic these latter fished very close to the shelf-break. species Abraliopsis pfefferi, Ancistrocheirus Pteroctopus tetracirrhus, Sepia orbignyana, Sepia lesueuri, Ommastrephes bartrami and elegans and Rondeletiola minor were found

Medit. Mar. Sci., 4/1, 2003, 79-86 81 4, 10, 11 10, 4, us 11 13 5 1 1515 13, 6 1 11 9, 2, S. AegeanS. Aegean N. ”05”15’ 23Æ 35”59’ 26Æ 1113 11, 10, 4, 9”20”11’ 23Æ 35”59’ 26Æ 4,10,11 21”27”19’ 23Æ 49”14’ 27Æ 33”31”00’ 11 23Æ 37”45’ 27Æ 9 3, 11 10, 4, 11 10, 4, 3, 22”21”19’ 23Æ 58”14’ 27Æ 119, 2, 13 11, 10, 4, 55” 58”40’ 23Æ 37”45’ 27Æ 30” 23”15’ 23Æ 41”04’ 27Æ 11 13 11 10, 4, 25” 25” 23”10’ 23Æ 11”25’ 27Æ 23”10’ 23Æ 58”14’ 27Æ 11 9, 119, 2, 11 10, 4, 11 10, 4, 54’ 12”54’ 07”15’ 23Æ 43”16’ 27Æ 1110 4, Æ 36’ 16”36’ Æ 01”59’ 23Æ 53”59’ 23Æ 11 Æ 11’ 22”11’ Æ 15”00’ 23Æ 35”00’ 23Æ 13 10, rimental and commercial trawl nets. trawl commercial and rimental ’ 52”’ 17”39’ 37Æ 44”’ 54”18’ 36Æ 48”13’ 23Æ 39” 5’ 23Æ 39”47’ 24Æ 54”17’ 25Æ 10,11 4, 07”’ 25”55’ 37Æ 23”10’ 23Æ 43”16’ 27Æ 11 9, 11 ’ 02”’ 11”52’ 37Æ 48”15’ 23Æ 22”25’ 27Æ 11 9, 10 4, 4’ 07”4’ 25”55’ 37Æ 23”10’ 23Æ 58”14’ 27Æ 19”2’ 05”08’ 37Æ 1511, 9, 3, 31”00’ 23Æ 11 10, 4, 3, 38” 26’ 23Æ 16”2’ 50”32’ 37Æ 00”17’ 24Æ 32”04’ 26Æ 11 10, 4, 08’ 48”08’ 17”39’ 37Æ 15”00’ 23Æ 39”15’ 23Æ 13 37Æ 08’ 42”08’ 37Æ 15”39’ 37Æ 15”00’ 23Æ 51”14’ 23Æ 14 Table 1 Table 257 77037”57’ 37Æ 04”09’ 37Æ 31”00’ 23Æ 37”42’ 23Æ 14 15 trawl trawl Frequency of occurrence inoccurrence of Frequency (m)depth Haul Latitude Longitude Previo Cephalopod species collected in the southern Aegean Sea by expe by Sea Aegean southern the in collected species Cephalopod Abraliopsis pfefferi (Verany, 1837) (Verany, pfefferi Abraliopsis 1839) (Orbigny, lesueuri Ancistrocheirus 1848)(Ruppell, margaritifera Pyroteuthis (Verrill,1880)reversa Histioteuthis 1798) (Lamarck, subulata Alloteuthis 1856)(Steenstrup, forbesi Loligo 0,21798)(Lamarck, vulgaris Loligo 7561798)(Lamarck, sagittatus Todarodes 14,9 1,2 7581841)(Ball, eblanae Todaropsis 29,81821) (Leueur, bartramii Ommastrephes 56148”08’ 37Æ 7,4 8,8 37 18,91817) (Leach, banksi Onychoteuthis 694 35 7,5 40 1604”30’ 36Æ 58’ 36Æ 549 7701798)(Lamarck, 335cirrhosa Eledone 6,229”19’ 35Æ 1798)(Lamarck, moschata Eledone 27’ 37Æ 26”14’ 35Æ 1826) (Risso, macropus Octopus 11051’ 37Æ 1839)(Verany, salutii Octopus 20,51798)(Cuvier, vulgaris Octopus 766 58,407”14’ 35Æ 1830) Chiaje, (Delle tetracirrhus Pteroctopus 51’ 37Æ 1840)(Orbigny, unicirrhus 33,8Scaeurgus 05”31’ 36Æ 5,6 25,04 53’ 37Æ 3,1 44 36,0 4,0 16 28,0 548 1,9 19,1 223 88 15,016”14’ 35Æ 13558”18’ 35Æ 70655’ 16 37Æ 9755’ 37Æ 52017 35Æ 270 54902”17’ 35Æ 55”14’ 35Æ 24 53’ 37Æ 29”14’ 35Æ 53’ 37Æ 37Æ Family Species Commercial MEDITS min max min max min max references Brachioteuthidaeriisei(Steenstrup,1882) Brachioteuthis ChiroteuthidaeCtenopterygidae1835)(Ferussac, veranyi Chiroteuthis 1851)(Verany, sicula Ctenopteryx Enoploteuthidae1844)(Ruppell, veranyi Abralia Histioteuthidae 0,31835) (Ferussac, bonnellii Histioteuthis 1,9 0,3Loliginidae 756 0,2 404 3,6 758 1758)(Linnaeus, media Alloteuthis 404 758Octopoteuthidae 881844)sicula(Ruppel, Octopoteuthis Ommastrephidae 4091839)(Verany, coindetii Illex 37Æ 68,9 72036 37Æ 37,423 35Æ Onychoteuthidae1839)(Orbigny, lichtensteini Ancistroteuthis 30 60,9Thysanoteuthidae1857) (Troschel, rhombus Thysanoteuthis 0,5Argonautidae 585Octopodidae 47,61758)(Linnaeus, argo Argonauta 7071 35Æ 1,2 1892)sponsalis(Fischer, Bathypolypous 37 778 766 0,60 37Æ 14 35Æ * 211 131 494 1394 36Æ 01”36’ 37Æ 37

82 Medit. Mar. Sci., 4/1, 2003, 79-86 shallower than previously reported in the Aegean Sea (D’ ONGHIA et al., 1991,1996; KALLIANIOTIS et al., 1995; 4, 10 4, 11 SALMAN et al., 1997). The narrow continental shelf and the generally steep waters of the southern Aegean 83 715 Sea probably favour the wide depth range observed for most of the species. Concerning the geographical range, the species caught frequently (f>5 %) (1999); 14 - Lefkaditou et al. (2000); 15 - Salman - 15 (2000); al. et Lefkaditou - 14 (1999);

(1992); 7 - Vardala - Theodorou (1994); 8 - Corsini - 8 (1994); Theodorou - Vardala - 7 (1992); by trawl net presented a wide et al. et

et al. et longitudinal and latitudinal range 2’ 34”2’ 44”52’ 26Æ 15 (Table 1). The capture of some 3Æ 11’ 20”11’ 3Æ 55”01’ 27Æ 10 4, 23Æ 30’ 06”30’ 23Æ 44”52’ 26Æ 12 11, uncommon pelagic squids, like Ancistroteuthis lichtensteini, ”04”29’ 23Æ 07”02’ 27Æ 11 10, 2”53”16’ 23Æ 05”01’ 27Æ 11 10, 4, 34”20”11’ 23Æ 22”25’ 27Æ 11 3, 11 10, 4,

’ 25”’ 16”15’ 23Æ 37”15’ 28Æ 119, 3, 25”’ 11 10, 4, 20”11’ 23Æ 46”04’ 27Æ 11 ,10, 4 3, Brachioteuthis riisei, Chiroteuthis veranyi, 3’ 22”3’ 40”19’ 23Æ 58”14’ 27Æ 11 9, 11 10, 4, 06”5’ 15”29’ 23Æ 16”53’ 26Æ 11 10, 4,

7Æ 53’ 24”53’ 7Æ 07”15’ 23Æ 37”45’ 27Æ 1511 10, 4, Ctenopteryx sicula, Pyroteuthis margaritifera and Octopoteuthis sicula ”25”55’ 37Æ 23”10’ 23Æ 55”27’ 27Æ 119, 3, 11 10, 4,

57”32”09’ 37Æ 31”00’ 23Æ 55”27’ 27Æ 111 3, was limited at the westernmost part of the study area (TABLE 1). The strong anticyclone appearing in the Myrtoan deep water basin during winter (NITTIS & LASCARATOS, 1993) may result in (1991); 5 -Vardala - Theodorou et al., (1991); 6 - Katagan - 6 (1991); al., et Theodorou - -Vardala 5 (1991); (1997); 12 - Lefkaditou and Kaspiris (1998); 13 – Lefkaditou – 13 (1998); Kaspiris and Lefkaditou - 12 (1997); high biological productivity along the et al. et et al. et abrupt coasts of eastern Peloponnisos, which could be the reason for a higher abundance of such species known to be usually associated with

(1996); 11 - Salman - 11 (1996); hydrographic regimes (COELHO,

et al. et 1985). Kocatas (1990), 4 -D’Onghia 4 (1990), Kocatas

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