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Home , Gulf of Genoa, Ligurian Sea, Rondeletiola minor, Sepietta, Sepiola, Sepiola aurantiaca

SEPIOLINAE (, CEPHALOPODA) FROM THE LIGURIAN

L. ORSI RELINI and M. BERTULEITI Labaratori di Biologia Marina ed Ecologia Animale, lstituto di Zoologia dell'Universita, Via Ba/bi 5, Genova , /talia

SEPIOLINAE ABSTRACT - A collection of 130 specimens of (ligulata, robusta, rondeletii, DISTRIBUTION intermedia), 115 , 90 (obscura , neglecta) and more abundant MATURITY samples of S. oweniana obtained by trawl fishing in the Ligurian Sea are briefly illustrated in terms of depth distribution, sex ratio and maturity stages.

SEPIOLINAE RESUME - Une collection de 130 specimens de Sepiola (ligulata, robusta, rondeletii, DISTRIBUTION intermedia), 115 Rondeletiola minor, 90 Sepietta (obscura, neglecta) et des nombres plus MATURITE MER LIGU RE importants de S. oweniana recoltes par chalutage en Mer Ligure sont etudies au sujet de Ia distribution selon Ia profondeur, Ia sex-ratio, et les stades de maturite sexuelle.

Mediterranean which has recently INTRODUCTION been brought up to date by Bello ( 1986). The sub-family Sepiolinae in particular, which includes Among the Cephalopods, especially those of II of the 16 of the family, has, as far as we commercial value, the Sepiolinae (family , know, never been studied in the Gulf of . In order Sepioidea or Sepiolioidea Fioroni, 1981) are fact, one has to go back to Verany (1851) to find a one of the lesser known groups with regard to text which deals with this subject, and even that is distribution and life history in the sea. However, in limited to two varieties of , a taxon some of its species it has been possible to observe which at that time included not only different the entire life cycle in captivity (Boletzky et al., species of Sepiola, but also sepiolids betonging to 1971); to study the use and control of bacterial light other genera. Verany states that « ... Les Sepioles ( Herring et al., 1981); to make detailed observations vivent sur toutes nos plages ... ; leur chair est delicate of embryology and comparative anatomy (Fioroni, et estimee : elle porte le nom de Sponcia-courrenti 1981 ). In contrast to these specialized studies, our et malnascui a Genes ... ». approach has more scape for field aspects ; we Naef gave the present status to the subfamily identify the numerous species in the Ligurian Sea describing the genera Sepietta and Rondeletiola with the aim of recording their rote, however small, (1912a; 1916) and seven species (1912b, c; 1916). In in the fishing grounds. the very abundant material which he examined he Taking Naef ( 1923) as her source, Mangold-Wirz mentioned some specimens of , S. ( 1963) lists 15 species of Sepiolidae in the Western ronde/etii and S. steenstrupiana coming from Ville­ Mediterranean; Torchio ( 1968) lists 16, also taking franche, i.e. from the western sector of the Ligurian into account atlantis G . Voss, 1955, Sea (Naef, 1923). For the description of the latter which Voss hirnself ( 1955) reported at Messina. This species Ligurian material was used both by Levy last Iist has remained unchanged in the register of (1912) and by Naef (1912b) almost at the same time ; 184 L. ORSI RELINI and M. BERTULETII

Table I. - Records of Sepiolinae in the Mediterranean subprovince.

GULF OF ADRIATIC CATALAN COAST N.TYRRHENIAN AND CADIZ LIGURIAN SEA

Naef 1923 Rudolph 1932 Mangold Wirz 1963 Lurnare 1970 Ruby & Knudsen Guerra Levy 1912 Lumare 1974 Boletzky et aZ. 1972 1982 Naef 1923 Bello & Mo- 1971 Knudsen 1981 Orsi & Bertu- t olese 1983 Sanchez & Mora- letti 1986 les 1986

SepioLa steenstrupiana Levy + + +

SepioLa aurantiaca Jatta +

SepioLa LiguLatc Naef + + + + Sepia La ronde Leti i Leach + + + Sepiola intermedia Naef + + + + Sepi oLa affinis Naef + + + + SepioLa robusta Naef + + + + RondeLetioLa minor Naef + + +

Sepiettc obscum Naef + + + + +

Sepiettc negLecta Naef + + + + +

Sepietta oweniana d'Orbigny + + + +

by only a few days Levy's specific name gained during the 1985 programme on the « Evaluation of priority (*). Demersal Resources » finaanced by the « Ministero Sepiolinae have been the objects of recent obser­ della Marina Mercantile » (Relini, 1985). vations off the Catalan coast (Mangold-Wirz, 1963; We have already provided a table which sum­ Boletzky et al., 1971 ; Sanchez and Morales, 1986), marises our finds of Sepiolinae (Orsi Relini and in the ( Lumare, 1970), off the coasts Bertuletti, 1986) ; here we shall give the essential of Israel and Cyprus (Ruby and Knudsen, 1972; details of the material examined. Knudsen 1981 ), in the Gulf of Cadiz (Guerra, 1982) and in the (Bello and Motolese, 1983; Bello 1984); the species recorded in these areas are shown in Table I. METHODS AND MATERIALS Recently we have had the opportunity of gathe­ ring information about this group thanks to a fishing programme (1982-1983) promoted by the « Minis­ In 1982-83 samples were taken at monthly inter­ tero della Marina Mercantile » on the problems of vals from the trawlable bottoms situated on the the Iimits of trawl fishing and on the protection of Eastern Riviera roughly off the coast at Chiavari. the coastal strip. Catches were carried out at Professional equipment hired for the purpose from monthly intervals, at Ievels between 20 and 90 the Santa Margherita fishing fleet was used : a meters. The use of a tightly-meshed net, which was trawler of 18 tons gross tonnage, equipped with a specifically designed as a commercial net for Cepha­ 150 H p engine, echo-sounding gear and radar and lopod juveniles (Eledone cirrhosa, in particular), a 3-man crew. The net was an otter-trawl, belly 400 made it possible to collect good material, to which meshes of 25 mm; lower body 120 meshes of 30 mm ; were added previous small collections also taken wings 200 meshes of 45 mm; cod end 420 meshes from trawl fishing carried out in the Ligurian Sea. of 6 mm side; lower and upper edge each 70 meshes Finally, more recently, further material was gathered of 35 mm; sweep lines of 200 m; warps of 250-1000 m; otter board of I 00 x 170 cm. Each monthly sampling consisted of one-hour effective trawls carried out during day-light hours at • Levy (1912) also mentioned S. deswigiana from Villefran­ four different depths : 20, 30, 50 and 90 m. Other che-sur-Mer and S. atlantica from Nice. lt is very difficult to establish the identity of the former species (named S. desvigniana occasional samples were taken at the same depth in the original description by Gervais and van Beneden, 1838). du ring the night and at depths of I 0, 85 and 300 m. The latter species is weil identified by the number of rows of Minor samples were also taken on board trawlers suckers on the ventral arms. On the basis of this work (Levy, 1912) S. atlantica seems to belang to the Mediterranean fauna ; which were engaged in commercial trawl fishing in however, since then no other record has appeared in the litera­ various part of the Ligurian Sea; these included : a) ture. samples obtained during the four seasons in 1975-76 SEPIOLINAE FROM LIGURIAN SEA 185 on the neritic bottoms between Genoa and Portofino RESULTS at depths between 30 and 180 m ; b) samples obtained from red shrimp fishing between 500 and 700 m on the Eastern and Western Riviera. Sepiola During the Programme « Evaluation of Demersal Resources ( 1985) », the trawlable bottoms situated Sepiola ligulata between and the mouth of the Magra and The identification of this species on the basis of depths between 0 and 700 m were explored using 56 the hectocotylized arm and of the bursa, respectively, hourly trawls. These were divided into 5 layers by is straightforward. In our material the portion of the depths 50, I 00, 200, 450 m in numbers propor­ bursa that extends on the right side beyond the tional to the extension of each layer. The net had 500 central mantle septum is sometimes almost as large meshes at the mouth and 500 at the cod end of 7.5 as the left portion. mm side. S. /igulata is one of the smallest species of The materials listed were fixed on board in I 0 % Mediterranean sepiolids and its records are gene­ formalin in sea water, with the exception of the last rally scarce. Only in the Adriatic has a group of 45 samples ( 1985), which were frozen. specimens been obtained in a single haul (Lumare, Systematic identification was made according_ to 1974). In the laboratory the embryonie development, the suggestions of Naef (1912b, c, 1916, 1923), 1.e. the hatching and the growth to a final length of mainly by observing the hectocotylized arm of the about 9 mm ML were observed by Boletzky et al. male and the visceral mass of the female in which ( 1971 ). Our records indicate a sexual maturity at the extention and the profile of the empty bursa is only II mm ML in the male and 14 mm ML in the critical. At the Zoological Station of Naples, some female. of Naefs collections of S. atlantica, S. intermedia, The depth range was 65-125 m in the Adriatic S. affinis, S. robusta, S. ligulata, Sepietta owenian~, (Lumare, 197 4) and 50-90 m in the Ligurian Sea. S. obscura and Rondeletiola minor were also exami­ ned. Sepiola robusta Naef gained such wide experience both in fresh and fixed Sepiolids that he was able to recognize the On the characteristic features of the hectocotyli­ species simply on the basis of general external zed arm is the great enlargement of one of the three features and colour. His collections include nume­ basal suckers. The small suckers in the forth and rous small undissected specimens. Unfortunately, by fifth rows above the copulatory apparatus give the now these materials have become colourless and appearance of a central narrowing of the arm, which very deformed by the fixatives ; however, they '"?ay in Naefs figure is not shown. still be useful in recognizing the main anatom1cal characteristics mentioned above. The female bursa may be ]arger than that repre­ sented by Naef, and sometimes Jeans on the central With regard to our collections, as we had speci­ septum (it does not however, extend to the right side, mens which were subjected to different kinds of as in S. rondeletit). fixation and for varying length of time, we were not able to use distinctive features based upon colour This « large » species is weil known in many patterns. aspects of its biology having been the object of long and exhaustive observations by Boletzky which he In listing the examined material we give the summarizes in his review on this subject (Boletzky, dorsal mantle length measured as indicated by 1983). In captivity female spawning was observed at Mangold-Wirz ( 1963). The zone of fishing refers to about 170 days after birth and at a M L of 17 mm; the locality on the coast off which the trawl was males are mature at a slightly smaller size. The effected (generally parallel to the ). The notes present field data confirm the same minimum spaw­ about the maturity stage are based upon the storage ning size. The depth range here observed (60-90 m) of spermatophores in the males (few sp. = matu­ appears restricted compared to that of the Catalan ring; abundant sp. = ready to spawn) and the coast (from a few meters to about I 00). diameters of eggs in the females, as suggested by Mangold-Wirz ( 1963) (for the sake of comparison we recall that this author gives the following size for Sepiola rondeletii eggs in advanced maturation : Sepiola rondeletii 2.5-3 Our material is composed only of small females. mm; Sepietta oweniana 2.1-2.6 mm). We have considered critical for the identification the All measurements were effected on specimens presence of a protrusion of the bursa on the right preserved in formalin. side, passing across the ventral septum, but not in the form of a hernia as in S. ligulata. As important samples of S. rondeletii, including adult males and females, have been obtained in 186 L. ORSI-RELINI and M. BERTULETII shallow water and also in lagoons, this species is Sepietta considered strictly littoral (Mangold-Wirz, 1963). However, Guerra ( 1982) recently obtained two im­ Within this , a separation of the group mature females at 190 m. Since also the present oweniana-neglecta from S. obscura is generally made specimens, fished at the least between 20 and 50 m, by the number of suckers at the base of the copu­ are all immature, a greater dispersal of young may latory apparatus, that is four and three, respectively. be supposed. However, we have noted frequent variations in this characteristic. In Sepietta oweniana, the most abun­ Sepiola intermedia dant of the three species, the number of basal suckers may be one, two, three, four, five, six with A very characteristic feature of the hectocotylized a total incidence of anomalaus forms of 7 % compa­ arm is the presence of two large suckers above the red to those bearing four suckers. The anomalaus copulatory apparatus. The enlarged suckers are the forms have the following incidence ( counted in 212 median first and second or, in some cases, the males captured in one hau I of 60 minutes duration) : median second and the third. this is the commonest I basal sucker in 1.4 % of the mal es; 2 b.s. in 2.3 %; species of the genus Sepiola in our material, proba­ 3 b.s. in 1.4%; 5 b.s. in 0.94 %; 6 b.s. in 0.94 %. bly because of its eurybathic distribution. In the However, the distinction of S. obscura is also based present collection we have registered its presence in on other morphological characteristics such as the the range 20-1 35 m, but other data taken from the cl ub structure. « Evalu ation of demersal resource » programme Recent descriptions of the distinctive features of now in progress (1986) extend the range to 220 m. female and male S. neglecta have been given by We obtained a maximum of 15 specimens per hour Bello and Motolese ( 1983) and by Sanchez and of trawling during the night. Moral es ( 1986). This species seems to have a patchy distribution in the Mediterranean; Naef quoted only specimens Sepietta obscura from Naples. Lumare, in extensive surveys on the Tyrrhenian coast, found it only south of Rome. E. S. obscura was studied in captivity by Boletzky et Mauris (pers. comm.) found it at Banyuls-sur-Mer. al. ( 197 1) who observed its development, hatching Sanchez and Moral es ( 1986) recently obtained a and growth to a final length of 24 mm ML in two single specimen from the Catalan coast. The record males. of Guerra ( 1982) at Cadiz seems questionable, since that specimen has a different hectocotylus. The biology of this species is poorly known. We have Table II. - Distribution of Sepiolinae in the Ligurian Sea registered an overall sex-ratio in favour of mal es (I for each month of the year : I Sepiola intermedia, 2 S. to 0.7). Adults in spawning condition are present robusta, 3 S. ligulata, 4 S. rondeletii, 5 Rondeletiota minor, throughout the year. Minimal spawning sizes were 6 Sepietta oweniana, 7 S. neglecta, 8 S. obscura. Blanks II mm ML in the male and 17 mm ML in the female. represent absence of species in the trawl samples, broken lines are for periods and depths at which no sampling was done. Bare numbers represent the respective species taken Rondeletiota in day samples, underlined numbers those taken in night samples. Rondeletiota minor F M A H J J A 0 N D

This species is easily recognized by its peculiar 20 luminous organs. In the trawl catches taken in the l l Ligurian Sea it is usually associated with Sepietta 30 oweniana; in number it represents a maximum of 4 8 .s. l 10 % andin weight a maximum of 5 %. In the eastern l l 50 sector it was registered at deep circalittoral and 6 8 epibathyal Ievels, in the western sector also at 8 l 7 l 6 l 2 l 5 l 6 90 500-700 m, perhaps due to sleeker bottoms. The 2 8 2 7 6 2 8 3 7 overall sex-ratio was I male : I ,4 female. Maturity 5 8 8 3 5 5 may be reached at only 12 mm ML in the male and 100-200 ------6 6 --- at 12 mm M L in the female. The present data suggest a Iong reproductive season (at least from April to 5 200-300 ------6 6 6 --- December). m

300-450 m

450-700 SEPIOLINAE FROM LIGURIAN SEA 187

Table 111. - List of specimens taken of each species in ------,------n dorsal mantle dare depth m reprod. sta~e chronological order, with indication of individual size, lcnAth mm fishing zone, depth of trawling, and stage of sexual 15; 18 01.04 . A5 Camog1 i 50 ab . sp . 7 ; 16;16; 19 01.04 . 85 Tigul1io 26 ab. sp. maturation in both males and females. ab. sp. = abundant 12 29.08.85 5 Terre 40 ;:~h. ~ ·'!. spermatophores; f. sp. = few spermatophores ; imm. 20.03 . 86 P . Manara 135 16 21.03.86 S Te rre 119 ab. sp. = = immature; l.e.d. Jargest egg diameter in mm; mat. 16 02 . 05.86 Camogl i 50 ab. sp. mature. females 19 04.08. 75 Camogli 50- 100 l.e.d. 3 21 25.10.75 Camogl i S0-100 l.e.d. 3 ------11; 16 ; 17;20 1€.05.82 Ch iavari 30 imm.1.e.d. 2.5 night n° dorsal mantle date zone depth m reprod. sta~e length lllm 14 29.07 . 82 Chiavari 50 imm. 14; 14 12.08.82 Chiavari 30 imm.; 1 .e . d . 2 . 5 Sepiola ligulata Naef, 1912 ni~ht mal es 7; 11 12 . 08 .82 Chiavari 50 imm. 1 3 23.09 76 Camogl i 50 ab. sp. night 12 26 .01.83 Chiavari 90 ab . sp . 16 03 . 12 . 82 Chiavari 50 l.e . d. 3 . S 11 08.12 . 83 Chiavari 75 ab . sp. 14 ; 17 26.01.83 Chiavari 30 1 e d 2 , 5 ; 1 ed 3 18 24.02.83 Chiavari 20 l. e .d. 3 females 12 24 . 02 . 83 Chiavari SO imm. 14 26.01.83 Chiavari 90 1 ed3 2-3.5 17 . 5 02.06.83 Chiavar i 20 l.e .d . 3 10 14.09 . 83 Chiavari 90 irrm. 9;14;17 19.07.83 Chiavari 50 2 imm.l.e . d) 10;13 ; 13 ; 14 . 5;15 19.07.83 Chiavari 90 imm. Sepiola robusta Naef, 1912 15 10.08 . 83 Chiavari 90 imm. ma l es 9;10;17 14.09.83 Chi.wari SO 2 imm.l . e . r' 3 13 29 .01.82 Chiavari 90 imm. 13;14;15;16 14.09 . 83 Chiavari 90 3 imm.; l . e.d 3 18 12.08.82 Chiavari 90 f. sp. 17 08.12.83 Chiavari 75 mat. night 16 ; 17;20;21 01.04.85 Tigul1io 25 mat . 12 03.11.82 Chiavari 90 imm . 19 01.04.85 Camog1 i 50 mat 17;22 26.01.83 Chiavari 90 f. sp . 20;21 01.04 . 85 Tigul1io 8S mat . 20 19.07.83 Chiavari 90 ab. sp . 19 29 .08.85 S Terre 62 1. e . d . 1 II 14.09 .83 Ch iavar i 90 inun.; f. sp . 11 . 5 31.08.85 Tigul1 io 30 imm. 15;17 14.09 . 83 Chiavari 90 ab. sp . 20.03.86 P. Manara I 35 fernales 16 ; 19; 20 02.05.86 Camogl i SO 1.1:' . d . 3 14 04.08. 75 Camogl i 50-100 imm . 24 25 . 10. 75 Camogl i 50 l.e.d. 3.5 ~ poor1y preserved eggs 11 23.09. 76 Camogl i 50 irmn. Ro'lf.deletiola minor Naef, 1$12 17 22.02. 78 Camogl i mat.- 23 12.08.82 Chiavari 90 l.e.d. 4.5 mal es n ight 17 11.08 . 75 Camogli 100- 150 ab. sp . 19 . 07.83 Chiavari 90 imm. 13 19.07 . 83 Chiavari 90 ab. sp . 20 14.09.83 Chiavari 90 l.e . d. 5 14 19.07 . 83 Chiavari 300 ab. sp. 13 10.08.83 C:hiavari 300 ab. sp. Sepiola rondeletii Leach, 1817 12;13;14;14 14.09 . 83 Chiavari 90 ab. sp. 10;11;11;15;17 . 5 08 . 12 . 83 Chiavari 90 females ab . sp . 9 ; 9 08 . 12 . 83 Chiavari 90 inun . 14 04.08 . 75 Camogli 50-100 imm . --- 15 ; 17.5 01.04.85 P. Manara 180 ab. sp . 13 23 . 09. 76 Camogl i 20-50 imm . 12 ; 16 ; 16;17;19 26.04 .85 220 ab . sp. 15 18.05.82 Chiavari 30 imm. 16 ; 18 27.07.85 550-700 spent? night 14 27 . 07 . 85 Ventimiglia 550-700 f. sp. 15 18.05.82 Chiavari 20 imm. 12 29.08.85 Oeiva 190 imm. night 14;16;16 ; 17 29.08.85 Oeiva 190 ab . sp.; spent? "' poorly preserved eggs 14;16;17;18 29.08.85 Tigu11io 322 ab . sp. 15 29 .08.85 Portofine 380 ab . sp. Sepiola intermedia Naef, 1912 15;20;20 09.09.85 Magra estu~ 117 ab. sp. 16 ; 17; 18 09.09 . 85 Magra estu. 117 spent? ; ab. sp. ma l es 18;18;18 ; 18 12 . 09 . 85 Cinque Ter. 225 ab . sp. 15;17; 17 .5 04.08.75 Camogl i 50-100 ab . sp. 16;17 ; 17 12 . 09 . 86 Cinque Ter, 225 ab . sp. 14 09.10 . 75 Camogl i 50- 100 ab . sp. 10; 14 23.09 . 76 Camogl i 20-50 ab. sp. fema1es 13 24 .II. 76 Camogl i 100-150 ab. sp . 16;19 11.08. 75 Camogli 100-150 l.e.d . 2 ; 2.5 16.5 10 .03. 78 Camog1i ab. sp. 13 19.07 . 83 Chiavari 90 inun. 13;14 ; 15;16; 17 29.01.82 Chiavari 90 ab . sp. 19 19.07.83 Chiavari 300 l.e . d. 2 20 24.03.82 Chiavari 90 ab. sp . 12; 13; 15 19.07 . 83 Ch iavar i 300 inun. 12 28.04.82 Ch iavari 30 ab . sp. 18 10.08 . 83 Chiay-ari 300 l.e.d . 11 18.05.82 Chiavari 30 ab . sp. 12; 12; 16 14.09.83 Chiavari 90 imm. night 12;13;13;15 14.09.83 Chiavari 90 l. e . d. 2 15;15;18 29 . 06.82 Chiavari 50 ab . sp. 15 ; 17 ; 17 08. 12.83 Ch i avari 90 l.e . d . 10 29.07.82 Chiavari 50 imm. 14; 15; 19 08.12.83 Chiavari 90 l.e .d. 17. ~ 12 ."~ . f1/ 17 .5 ; 18 ; 18 ; 18 01.04. 85 P . Manar a 180 1 . e . d. nigllt 14; 17; 20; 22 01.04. 85 P. Manara 180 l. e .d. 14;14;15;16;16 12.08.82 Chiavari 30 ab. sp. 19 26.04 . 85 P . Mesco 220 l.e . d. 2 night 15; 17 26.04.85 La Spez ia 220 inun. 12 ; 15 ; 17 ; 17 12.08. 82 Chiavari 30 ab. sp. 17;18;21 ; 21 ; 21 ; 2127.04.85 Levante 240 l.e . d. 2.5 ni~ht 1J;l6 27 . 04 . 85 Levante 240 inun. 14 ; 15 12.08.82 Chiavari 50 f . sp. 16 04 .05.85 Portofine 250 l.e . d. 2.5 night 14 27.07.85 Ventimiglia 5.50- 700 itt'l11. 13 ; 16; 16; 17 04.10.82 Chiavari 50 f. sp . 19;20;21 29 . 08 . 85 Deiva 190 l.e.d. 2.5 8;8;9; 13 04.10.82 Chiavari 50 imm. 17 29 .08.85 Oeiva 190 imm . 13.5 04.10.82 Chiavari 90 f . sp. 12 29 . 08.85 Chiavari 322 l.e.d. 2 . 5 15; 16 ; 16 03.11.82 Chiavari 50 ab . sp.;imm . 18 09 . 09 . 85 La Spezia 240 l.e.d. 2 13 03.11.82 Chiavari 90 a b. sp. 1 2 09.09.85 Magra estu. 117 inun. 10 24.02.83 Chiavari 30 imm. 17; 17; 17 ; 20 09.09.85 Magra estu. 117 l.e.d. 11 24.02.83 Chiavari 50 ab . sp. 21 09.09.85 Hagra estu. 117 l.e . d. 2.5 9; 14 02.06.83 Chiavari 20 imm . ; f. sp . 16; 16 22 ;22 12.09.85 Cinque Ter, 22.5 l.e.d. 2 15 19.07 . 83 Chiavari 50 imm. 17; 19 20; 20 12 . 9.85 Cinque Ter. 22.5 l.e.d. 2.5 11; 17 19 .07.83 Chiavari 90 imm . or spent? 15; 17 18 12 . 09.85 Cinque Te r. 22.5 l.e .d. 2 9 10.08.83 Chiavar i 90 imm. 15 20.03.86 Levante 82 l. e . d . 12; 13 14.09.83 Chiavari 50 imm.; ab . sp. 20.09.86 P , Manara 13.5 10; 13 14.09 .83 Chiavar i 90 irrna. 16 ; 16; 18; 18 (.•8 l2 .C..:3 Chiavuri 90 ab. sp. 17 oe.t~.8J Chiavar i 75 ab. sp . 188 L. ORSI RELINI and M. BERTULETII

no dorsal man tle date depth m reprod stage n° dorsal mant t e' date depth m repr od stage _____ len th ______c l.ength__

Sepietta obscura Naef , 1916 73 1 7-)1 14 . 09.83 Chi.war i 300 25 08.12.83 Chiav.:1ri 90 ma l es 14 18-25 0 1.04.85 Nervi 180 23.09 . 76 Camogli 25- 50 ab . sp . 11.5 8 14-26 27.04.85 Geno.1 240 29 .01. 82 Chiavari 13;13 90 ab. sp. 22:23; 24 ; 26 : 28 04 .O S. 85 Portofine 250 12; 13; 16 ; l7 18.05.82 Chiavari 30 ab . sp . '!.I I 3 .05 . 85 S;mremo night 17 ; 22 ; 22 . 5 30 .0).85 J.a Sp ez i.a 455 10 18 . 05.82 Chiavar i 30 i rmn . 34 16- 23 31 .05 .85 La Spezi;t 430 night 4 17;18;22;22 0 ).08 .85 Cinque Terr 13; 15 12 .08.82 Chiavari 30 ab . sp . 4 22;22;22;29 17.08.85 Portofino 430-700 nigh t 1]1 14-28 22.08.85 Cinque Tcrr425 14-16 imm . 16 ; 17 ; 18 ; 18; 19 12 . 08 . 82 Ch iavari 50 ab . s p . 15- 20 f . sp. nigh t 17-28 .1b . sp . 16 03 .12 . 82 Ch iavari 50 i rmn. 123 14-]0 22 .08.85 Cinqu(' TC'rr468 14-16 irran . 15 26 . 0 1.83 Ch iavar i 20 ab . sp . 15-20 f. sp. 17 24.02 . 83 Ch iavar i 20 a b . sp. 17-::!R ;Jb. SI>, 11 24.02.83 Ch iavari 50 a b. s p. 10 22 .08.85 Cinqu..: Ter 520 9 02 . 06 . 83 Chiav a r i 20 imm. 15 11.08.85 Sestri L. 500 18 19 .07.83 Ch i avari 50 ab . s p . 36 18- 27 22.08 . 85 Cinquc Tcr 520 9;9 10 . 08 . 83 Ch iava r i 50 iffi!11 . 112 16- 29 22 .0!: . 85 L.:1 Spezi.1 4 70 12 14 .09. 8 3 Ch i avari 90 a b. sp . 3 20- 23- 26 23 .08.85 Chiavari )JO Chi .1var i <>Jn 11 14 .09.83 Ch i avari 90 i mm . 22 23.08.85 22;23 Chiav,ni ]Q(l 11 08. 12 . 83 Chiavari 50 imm . 29.08.85 13 16- 25 29.08.85 P••r t n t 1!\t• J.'.' :~l> . s p . 15 0 1 .04 . 85 T igu ll io 26 ab . s p . 19 31 .08. A5 im:n . 14; 14 20. 03 . 86 Lev ant e 82 ab. sp. 12 15- 18 09. 09.85 : .. 1 Spt•zi,l 117 fema les 80 14 - 29 09.09 . 85 l....t Sf'cl.id 240 12 04.08. 75 Camogl i 50-100 i mm . 36 18-29 oq .C'o .R 'i I 'I <:.r•·~ i I !..F, ) 15-17 29 . 01.82 Chiavari 90 imm. 18 I 7-21 12.09 . 85 Ci nq ue Ter l25 15 . 5 18 . 05 . 82 Ch iavari 50 imm. 2 25 ; 27 12 . 09 . 8S p . ~1(.' sco 42 5 .1b . sp. n i gh t 19- 20 08 .05 . 86 La SpezL1 220 .1b. sp. 22 18.05 . 82 C:hiava r i 30 l.e .d. fema 1es night 31 ot. . 0 8 75 Camogl i 50- 100 i.t'.d . 3 12 15; 15; 18 18.05.82 Chiavad 30 imm.l.e . d. 2 18;23 LI .08 75 Camogl i 100- 180 invn. 23 29 .11fl . 82 Chiavari 50 ma t. 23;24 ; 28 15 .01 76 Camo~1 i 100- 180 I .e.d . 3 19 12 .r.$ , 32 Chiavar i 20 l.e . d . 2.5 18 15 .01. 76 Camo~l i 100-180 ni~h t 21 18 .02 76 CamoAI i 50-100 I .P .d. 3.5 }f,· J9 1? ll!l A? 10 l. e .rl . ] 26;37 18 .02 . 7f. CamoJal i 100-180 1 .e.d . 3 nigh r: 20;22;24 30.06. 76 Camo~ l i 300 2 irran. ;l.l·.d .2 15 ; 18 l 2.0ö.132 Chiavari 30 l. e . d. 2 26 20.08. 76 Camo g l i l00-L80 I .e.d. 3 . 5 night 18;26 '22 .02 . 78 Camog1 i 100-180 .e.d . 3 15 ; 15: 17 ;1 7 ; )2.08 . 82 Chiavari 90 imm .l . e . d..2 . 5 29 1o .o :-. . 7F> C;ttni'IP\ i ],)0-180 .e . d. 1 19 ; 19 ni~ht 32 29.0i' . 62 Ch iavari ! IJ 1 . .... 1. j 15 26.01.83 Chiavari 90 i mm. 12 ; 18 ; 18 ; 19; 2102 . 06 . 83 Chiavari 300 13 ; 15; 18 19 . 07.83 Chiavari 50 imm . 2 l.e.d. 2 53 10- 34 19.07.83 Ch iavari 300 15 10.08.83 Chiavari 50 imm . 16 15- 28 10.08 . 83 Chiavari 300 16-34 r:h i r i 12 10.08 . 83 Chiavari 90 imm . 50 14.09.83 ava 300 6 18- 31 0 I . ·)I • ~:~, ;~Hvi 180 l. e . d . 3.5 12; \8 14 . 09 . 83 Chiavari 50 imm. in 27 - 11 17 . 5 14 . 09 . 83 Chiavari 90 24 ;26 ; 28 26.04 . 85 Ci nque Ter 23 ; 24; 28 \0; I L; 14; 17 14 . 09 . 83 Chiavari 90 i mm. 26.04.85 La Sp ez ia 220 l.e.d. 1; 4 23 \0 08.11.83 Chi avari 20 imm . 26.04.85 La Spezia 220 22-30 27.04.85 Gt:noa 240 12 08.11 . 83 Ch iavari 50 imm. 24 ; 27 Po rtof ine 250 17 23 01 .04 . 85 Tigullio 26 l.e .d . 3 04.05 . 85 26 17 01 .04 85 Camogl i 50 imm . 13.05 . 85 19 0 1. 04 85 T i gul lio 8 5 l.e . d. 3 16 17.05 . 85 Cinque Ter 423 17 24.08.85 70 imm. 30 17 .OS .85 Cinque Ter 4 39 20 02.05.86 Camogl i 50 l.e .d . 3 18 ; 25 ; 28 30.05.85 La Spez ia 455 16 02.05.86 Camog1 i 50 imm . 33 15- 25 31.05.85 La Spezi<~ 430 1.e . d. 3- 3.7 20 03 .08.85 Cinque Ter in 22-25 21 ;29 ; 31 17.08.85 Portof ine 430- Jn0 86 10-32 mal es 22 .08 . 85 CinCJUC Te r 425 \0-22 : irran. l. 7 ; 20 30.04. 76 Camog 1 i 50-100 ab . s p. 22-27 : \.•· .d. '} 11 26.01 .83 Chiavari 90 ab. s p . 25 -30: l.c .d. '3 . 5 94 14-30 - 14 24.02 . 83 Chiavari 30 f . sp . 22.08.85 Cinqul: Ter 468 14 - 25 : i1Tl1'1. 11 19.07.83 Chiavari 90 ab. sp . 18 - 25: I ~·d 2 5-'J 11 08. 12.33 Chiavari 75 ab . sp . 2:.- 'J<~ : I . , . . d . } . j 12- 32 14 26 . 04 . 85 La Spezia 220 i mm. 18 3 22 . 08 . 85 L.1 Spezi a 4 70 12-2f. : imm . 15 09.09 . 85 Magr a estu 117 i mm. 20-32: I.C'.d, 3 21 - '32 : I . l' .d. 4 females 46::!6 22.08.85 Sestri L. 'iOO 14 04.10.82 Chiavari 90 imm . 37 14-29 22.08.85 Cinqut· Ter 520 12 28.12 . 82 Chiavari 50 i mm. 3 23;21 ; 27 23.08.85 Chiav:1r i 530 14 ; 14;18;17; 17 26.01.83 . 2 l.e .d . 2 . 5 Chiavari 90 l mrrL 2 l. e .d . 2 24 23 . 08.85 Chiavari 5JO imm. 17 02 . 06 . 83 Ch iavari 300 l. e . d . 2 . 5 31 24.08 . 85 CinquC' Tn L 10 I .t' .d. 4 11 08 11.83 Chiavari 90 l.e . d . 2 16 ; 21 29.08.85 Chiav:~ri 190 17 08. 12.83 Ch iavari 90 l.e.d . 3 20;27;29;29 29.08.85 Portufino ]22 20- 23 29.08.85 Po rtof ino 322

A poorly prcserved e ~gs 26;2f. 31 .08.85 Re cco 450 15 13-28 L1 Spf.•Zi:J 117 S.pi.·t ta

The Ligurian specimens were fished both on this species extend from 1905 to 1966, but are mainly infralittoral and circalittoral bottoms. Minimum from the oldest time period, when artisanal coastal spawning sizes were 12 mm M L in the male and 16 fisheries were probably more common than now. mm ML in the female (fixed spec.); the sex-ratio was Therefore, this species can be definitely added to the in favour of females ( 1.4 : I) ; the spawning season cephalopods of the Ligurian Sea and considered was long (at least April-September). quite common. S. steenstrupiana, described on the basis of speci­ Sepietta neg/ecta mens from Villefranche, was found also in the · and near Naples (Naef, 1923) in the S. neglecta was kept in the aquarium from hat­ Adriatic (Rudolph, 1932) and in Haifa Bay (Knud­ ching to a final length of 15 mm M L by Boletzky et sen, 1981 ). Considering these few records, it seems al. ( 1971 ). We have occasionally fished this species a rare species which Jives at infralittoral Ievels (5-20 from infralittoral to bathyal grounds, as could be m in the Adriatic; 47 m in Haifa Bay on sandy mud). expected from recent records in the Adriatic (30-50 No Ligurian records are to be found at the m), in the Gulf of Cadiz (70-475 m) and off M.N.H.G. Taragona (315-363 m). S. aurantiaca was described by Jatta ( 1896). Naef Minimum spawning sizes were 17 mm ML in the (1923) saw only preserved specimens coming from male and 17 mm ML in the female. Naples and the Atlantic coasts (Roscoff, Firth of Forth, Bergen, ); he stated that it probably Sepietta oweniana lived in deep waters at about 200-400 m, although it is not clear on which Observations he based this Mangold-Wirz ( 1963) and Bergstrom and Sum­ remark. This species seems the rarest in the Mediter­ mers ( 1983) give exhaustive accounts of the life-cycle ranean, with no records after Naef, and it represents of S. oweniana on the basis of many observations the only species actually lacking till now in the made both in the field and in the laboratory. Ligurian Sea. S. oweniana is very abundant in some fishing From the data collected in our area the ecological grounds of the eastern sector of the Ligurian Sea, distribution of the five commoner species of Sepiola where we have recorded a maximum of 395 indivi­ may be summarized as follows : duals per hour of trawling. The denser samples were obtained at 400-500 m at a distance from the coast a) shallow water species : S. rondeletii. S. affinis of about 20-25 miles. b) circalittoral species : S. robusta. S. ligulata c) eurybathic species : S. intermedia Males range from 14 to 30 mm ML and females from 10 to 35 mm ML (fixed specimens). Minimum The scarce records of S. steenstrupiana have so far spawning sizes are 18 and 20, respectively, in male suggested that it belongs to group a). On the other and female. Mature are present throughout hand, if Naers supposition proves correct, S. au­ the year. In contrast, the bathyal population is not rantiaca will be the only species of Sepiola living stable, because the spring sampling gave a nurober entirely along the slope. of animals corresponding to only I 0 % of the sum­ The distribution of S. robusta in areas different mer sampling. A peak of hatching and/ or a migra­ from the Ligurian Sea seems !arger than that above tion toward deeper waters takes place in the late indicated. Data of Mangold-Wirz ( 1963) and Bo­ spring or at the beginning of the summer. letzky ( 1983) state its presence on infralittoral bottoms along the Catalan coast, i.e. in the same zone where the absence (or a very scarce presence) of S. intermedia was registered. An enlargement of DISCUSSION the niche of S. robusta in this particular case may perhaps be supposed. Considering the Mediterranean records (Table I), In general, the species of the genus Sepietta seem three species, S. affinis, S. steenstrupiana and S. to be even more eurytopic than Sepiola. S. obscura aurantiaca, are found missing in this collection of Jives in neritic waters, from the shore to about I 00 Sepiolinae from the . m, exposed to a range of temperatures and other We suppose that our sampling methods were not environmental factors similar to S. intermedia. S. suitable for the capture of S. affinis, which Jives « at neglecta has a !arger range, reaching bathyal Ievels. depth of a few metres on sandy bottoms » (Naef, S. oweniana is weil known for its presence from 1923). One of us has ascertained its presence in the about 50 to more than 900 m (Mangold-Wirz, 1963). Gulf of Genoa by examining the small sepiolids, The eight species in our collection are all found generally Iabeiied as S. rondeletii, which form the in a belt between the depths of 50 and I 00 m (Table collection of the Museum of Natural History in II). In this particular range the temperature is almost Genoa (M.N.H.G.). Out of a total of 35 specimens constant (13-I4°C) from December to April, and of Sepiola, 14 are S. affinis; the Ligurian records of variable with the thermocline ( 13-l9°C) from May to 190 L. ORSI-RELINI and M. BERTULETII

November. If egg laying takes place here, different KNUDSEN 1., .1981. Three Sepiolidae new to the eastern clusters of eggs laid at slightly different depths in Mediterranean (Mollusca : Cephalopoda). Argamon, spring, may hatch at very different times. This fact Isr. J. Malac., 7 : 45-50. may help to maitain a continuity of spawning LEVY F., 1912. Observations sur les Sepioles des cötes de conditions during the year in shortlived semelparous . Archs. Zoo/. Exp. Gen., 9, Notes et Revue, 3 : species. 54-59. LUMARE F., 1970. Note sulla distribuzione di aleuni Cefalopodi del Mar Tirreno. Bol/. Pesca Pisci. Idrobiol., ACKNOWLEDGEMENT. - We are indebted to Or. 25 : 313-344. G. Bello (Laboratorio di Biologia Marina and LUMARE F., 1974. Occurrence .of Cephalopods Sepio/a Acquario, Bari) for valuable discussions and Iitera­ ligulata (Naef, 1912) and Rondeletiota minor (Naef, ture supply. 1912) in the Adriatic. Biljeske-Notes Inst. oceanogr. rib. Split., 33: 1-7. MANGOLD-WIRZ K., 1963. Biologie des Cephalopodes benthiques et nectoniques de Ia Mer Catalane. Vie Milieu, suppt. 13 : 285 p. NAEF A., 1912a. Teuthologische Notizen. Zoo/. Anz., 39 : LITERATURE CITED 241-248. NAEF A., 1912b. Teuthologische Notizen. Zoo/. Anz., 39: BELLO G., 1984. Sepioidei del litorale di Bari. Nova 262-271. Thalassia, 6: 707-709. NAEF A., 1912c. Teuthologische Notizen. Zoo/. Anz., 40 : BELLO G., 1986. Catalogo dei Molluschi Cefalopodi 78-85. viventi nel Mediterraneo. Bol/. Malacologico, 22 (9112) : NAEF A., 1916. Ueber neue Sepiatiden aus dem Golfvon 197-214. Neapel. Pubbl. Staz. zoo/. Napoli, I : 1-10. BELLO G. and MOTOLESE G., 1983. Sepialids from the NAEF A., 1923. Die Cephalopoden. Fauna Flora Golf Adriatic Sea (Mollusca, Cephalopoda). Rapp. Comm. Neapel; 35 (1. Teil, I. Band : Systematik) 863 p., 19 tavv. int. Mer Medit., , 28 (5) : 281-282. Traduzione in inglese di A. Mercado, 1972. Smithso­ BERGSTROM B. and SUMMERS W.C., 1983 . Sepietta nian Inst., Washington, 917 p. oweniana. In : Life Cycles, I, P.R. Boyle, ORSI RELINI L. and M. BERTULETII, 1986. Notes on ed., Academic Press, N.Y. & London: 75-91. the presence and distribution of the Sepiolinae in the BOLETZKY (von) S., 1983 . Sepio/a robusta. In: Cephalo­ Ligurian Sea. Rapp. Comm. int. Mer Medit., 30 (2) : 247. pod Life Cycles, I, P.R. Boyle, ed., Academic Press, RELINI G., 1985. Programme of the Italian Ministry of N.Y. & London : 53-67. the Merchant Marine for the demersal resources survey BOLETZKY (von) S., BOLETZKY (von) M.V., FROSCH in the area 37/ 3. FAO Fish Rep., 336 : 119-123. D. and GATZI V., 1971 . Laborator rearing of Sepio/inae RUBY G. and KNUDSEN J., 1972. Cephalopoda from (Mollusca : Cephalopoda). Mar. Bio!., 8 : 82-87. the Eastern Mediterranean. Isr. J. Zoo/., 21 : 83-97. FIORONI P., 1981. Die Sonderstellung der Sepioliden, ein RUDOLPH H., 1932. Die Sepiolinen der Adria. Zoo/. Vergleich der Ordnungen der rezenten Cephalopoden. Anz., 101 : 112-120. Zoo/. Jb . Syst., I 08 : 178-228. · SANCHEZ P. and MORALES E., 1986. Nota sobre Ia GERVAIS M.P. and VAN BENEDEN P.J ., 1838. Sur les presencia de cuatro especies de Sepiolidae (Mollusca : malacozoaires du genre Sepiole. Bull. Acad. Sei. Bruxel­ Cephalopoda) en el Mediterraneo nordoccidental es­ /es, 5 : 421-430. panol. Inv. Pesq., 50(1): 137-144. GUERRA A., 1982. Cefalopodos capturados en Ia cam­ TORCHIO M., 1968. Eieneo dei cefalopodi del Mediter­ pana « Golfo de Cadiz-81 ». Res. Exp. Cient., I 0 : raneo con considerazioni biogeografiche ed ecologiche. 17-49. Ann. Civ. Mus. St. Nat. Genova, 77 : 257-269. HERRING P.J., CLARKE M.R., BOLETZKY (von) S. VERANY J.B., 1851. Mollusques mediterraneens obser­ and RYAN K.P., 1981. The light argans of Sepiola ves, decrits, figures et chromolithographies d'apres le at/antica and Spirula spirula (Mollusca : Cephalo· vivant. I. Cephalopodes de Ia Mediterranee. Genes : poda) : bacterial and intrinsic systems in the order 132 p., 41 pls. Sepioidea. J. Mar. bio/. Ass. U.K., 61(4) : 901-916. VOSS G.L., 1955. The Cephalopoda obtained by the JATIA G., 1896. I Cefalopodi viventi nel Golfo di Napoli. Harvard Havana-Expedition off the Coast of Cuba in Fauna jlora Golf. Naepel : 268 p.; 31 tavv. 1938-1939. Bull. Mar. Sei. Gulf , 5 (2) : 81-115.

Re~u l.e 18 fevrier 1988; received February 18, 1988 Accepte le 28 jevrier 1988; accepted February 28, 1988

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