Boll. Malacol., 45: 61-69(suppl. 2009) Distribution and some biological aspects of cephalopods in the North and Central Adriatic Elisa Ciavaglia (ED) & Chiara Manfredi Laboratoriodi Biologia Abstract Marinae Pesca, Viale This paperreportsdata aboutthedistribution ofcephalopodscaughtduringtrawl surveyscarried outinthe Adriatico 1/N, 61032 North and Central Adriaticfrom 1982 to 2004. Biological notes on some cephalopods are also reported. Fano(PU), Italy, [email protected] Riassunto (El)Corresponding Author Nel corso di 40 campagne di ricerca di pesca a strascico effettuate in Altoe MedioAdriatico, fra il 1982 e il 2004, sono state ritrovate 32 specie di cefalopodi di cui tre molto rare nell'area di studio. In questo lavoro, i dati più recenti vengono utilizzati per aggiornare le informazioni sulla presenza e distribuzione dei cefalo- podi menofrequenti o di minore importanza commerciale. Key words Cephalopods, Adriaticsea, deep-distribution, trawl-survey. Introduction Material and methods The Adriatic Sea extends for about 800 km from NW to The study area covers the Italian territorial waters and SE; its width ranges from 90 to 220 km. It can be subdi- the international waters of the North and Central Adri- vided into threebasins. atic,to thenorthofthe linejoiningtheGarganopromon- The northernbasin has shallow waters and depths that tory (Italy) to Kotor (Montenegro). GRUND surveys gently increase southwards reaching 70 m at maximum were carried out in different seasons up to 1996, after- depth. The central basin reaches the maximum depth in wards they were carried out only in autumn. The gear thePomo (orJabuka)pit(-260 m). Thesouthernbasinis usedwasthetartana,acommercialItaliantrawlnet,with mm almost entirely occupied by a fairly deep pit which the cod-end mesh size of 40 (stretched). Samplings reaches - 1,223 m. The general circulation is dominated in MEDITS surveys were always accomplished in the by a cyclonic gyre: the water masses inflow through the summer, usinganexperimentalgearwiththe mesh cod- mm Strait of Otranto along the eastern side and outflow end size of20 (stretched). m along the western side. The north-western Adriatic is Hauls werecarried out from 10 to 800 ofdepth. Catch characterised by a large freshwater inflow that lowers data inweightwerestandardised to thesurfaceunit(= 1 the salinity and leads to high primary production and km2). h2i00g3h)biological productivity. These features, together Size is reported as dorsal mantle length (ML), ifnototh- with the geological historyofthebasin, trigger the pres- erwisespecified. ence ofsomeendemic species (other than cephalopods), mostlyeuryhalineandofcoldwateraffinity,inthenorth- ernmost part of the Adriatic. However, there is no evi- Results dence of any isolated population of cephalopods in the NorthAdriaticSea (Bello, 1990). Thirty-twospeciesofcephalopodswerefound:3ofthem TheteuthofaunaoftheAdriaticSea includes45ofthe65 belong tothe orderSepiida, 11 totheorder Sepiolida, 10 species known from the Mediterranean basin (Bello, to the orderTeuthida and 8 to theorderOctopoda. In this work some observations are reported IneachsurveytheOctopodawerethepredominantfrac- . about the presence and distribution of the cephalopod tion, in terms of weight: Eledone moschata was the most species caught during several trawl surveys carried out abundant species in autumn and winter surveys (from in the North and Central Adriatic Sea. Such surveys 29% to 60% of the total cephalopod catches), whereas were accomplished in the framework of the demersal Eledone cirrhosa, E. moschata, and Illex coindetii were the GRUND resources evaluation programs: (from 1982 to mostabundant species in spring and summer. 2004) and MEDITS(from 1994to2004).Somedataabout Heteroteuthis dispar, Histioteuthis bonnellii, Octopus macro- the first years of these surveys have been already pub- pus, and Pteroctopus tetracirrluis were no longercollected lished (Casali et ah, 1998; Soro & Piccinetti Manfrin, after 1995 inanysurveytakenintoaccountinthispaper. 1989) and are updated here with the results of recent These species, as well as 1. coindetii, E. moschata, E. cir- campaigns. rhosa, Sepia officinalis, Eoligo vulgaris, Octopus vulgaris, kg/km Sepia orbignyana : 2.0 - WIEDITS G autumn 1.5 --X-- G spring - G summer -A- ---®- G winter 1.0 Elisa Ciavaglia 0.5 & Chiara 0.0 ~-X Manfredi 0-50 50-100 100-200 200-300 300-400 400-500 Depth (m) Fig. 1. Biomassindexagainstdepth gradientforSepiaorbignyana in MEDITSSurveysand in differentseasonsofGRUND Surveys. Fig. 1. Indicedi biomassasu gradientedi profondità diSepiaorbignyananei survey MEDITSe in diversestagioni deisurveyGRUND. have been fully discussed previously by Casali et al. size forboth sexesis 12mm. Thesizefor S. elegansinthe (1998) and Soro & Paolini (1994). studied area ranges from 20 to 80 mm in females and mm from 21 to 75 inmales. Sepüdae Sepiolidae Sepia orbignyana and Sepia elegans show a bathymetric range wider than Sepia officinalis, which is not found Sepietta oweniana is the species more frequently caught, deeperthan 100 m. followedby Rondeletiola minor, Sepiola robusta, and Rossia m Sepia orbignyana has been caught from 15 to 450 of macrosoma. m depth, although it is more abundant from 50 to 100 in The latest data corroborate the distribution reported by all seasons (Fig. 1). An increase ofcatches up to 50 m of Guescin and Manfrin (1986) for the species of the genus depth,alongthecoastal areasand inthearea tothenorth Sepietta (Fig. 3). of the Po delta, is recorded in the autumn. Sepiettaobscura, whichprefersshallowwatersandsandy m Ineveryseason, S. elegans ismoreabundantup to 100 bottoms, wasmainlycaughtin thenorthernmostarea of m of depth and its abundance decreases quickly beyond theAdriatic from 30 to 50 whereas only 10 specimens m this depth; occasional catches are recorded till 440 werecaught deeper than 100 m. (Fig. 2). Unlike prior assumptions (Casali et al., 1998), Sepiettaoweniana and Sepietta neglectaweremainlyfound new data allow us to suppose that, in the surveyed area, in the middle-southern part of the study area. Sepietta S. elegans doesnotmigrateseasonally,althoughthisphe- neglecta was occasionally caught (110 specimens) be- nomenon has been observed in other areas (Relini et al., tween 37 and 340 m, that is the same depth range in 1999). which itwas fished in the Strait ofSicily (Jereb & Di Ste- Inbothspecies,maturemalesandfemalesoccurthrough- fano, 1995). out the year, suggesting a long, almost continuous, Sepietta oweniana is common in the whole area across a spawning period, with males reaching maturity at a wide bathymetric range from 15 to 450 m, with catches smallersizethanfemales (Relinietal., 1999). InS. orbign- increasing with depth up to 200 m. Females showed yana, mature males were observed from the size of 38 larger size than males, as reported from the Ionian Sea mm on and females from 40 mm on. In S. elegans the and Aegean Sea (D'Onghia et al., 1995). MLs ranged mm mm mm smallest mature sizes were 29 in males and 31 from 20 to 32 in males and from 12 to45 in females. in females. In the Italian seas, the maximum reported Mature individuals were found throughout the year, mm mm size for S. orbignyana is 96 in females and 93 in with larger percentages in the summer, thus suggesting males (Jereb & Ragonese, 1991;Beicari &Sartor, 1993). In a peak in the reproductive activity in this season (Relini thisstudythe largestcollected femalewas 106mm, larg- et al., 1999). 62 est males did not exceed 88 mm. The smallest collected Species belonging to the genus Sepiola were more often Distribution and some biological aspects of Fig. 2. BiomassindexagainstdepthgradientforSepiaelegansin MEDITS Surveysand in differentseasonsofGRUND Surveys. Fig.2. Indicedi biomassa su gradientedi profondità diSepiaelegansneisurveyMEDITSe in diversestagioni deisurveyGRUND cephalopods in m caught in the area north of Pescara, deeper than 200 mens); in the autumn two individuals were caught at the and along thecoasts (Fig. 4). 120mand twomales werecaught in thePomo Pitat220 Sepioln affinis was occasionally caught in the area be- m, an exceptional depth for this benthic littoral species North m m tween Ancona and Pescara from 20 to 100 (166 speci- thatusually livesat25-30 ofdepth. and Central Adriatic Fig. 3. Distribuzionedel genereSepietta. 63 North MiddleAdriatic Depth Adriatic Species 10- 51- 51- 100- 201- 301- 501- 10-50 min. max. 50 100 100 200 300 500 800 OrderSEPIIDA FamilySEPIIDAE SepiaofficinalisLinnaeus, 1758 + + + + + 10 143 SepiaorbignyanaFérussac, 1826 + + + + + + + - 15 439 Sepiaelegansde Blainville, 1827 + + + + + + + - 11 444 OrderSEPIOLIDA Elisa FamilySEPIOLIDAE SepiolarondeletiiNaef, 1912 + + 21 75 Ciavaglia Sepiola intermediaNaef, 1912 + + + + + - - - 10 114 Sepiola ligulataNaef, 1912 + + + + + + - - 23 201 & SepiolarobustaNaef, 1912 + + - + + + - - 15 229 Chiara SepiolaaffinisNaef, 1912 + + + + + + - - 19 220 Sepiettaoweniana (d'Orbigny, 1841) + + + + + + + - 14 444 SepiettaneglectaNaef, 1916 + + + + + + + - 37 334 Manfredi SepiettaobscuraNaef, 1916 + + + + + - - - 25 113 Rondeletiolaminor(Naef, 1912) + + + + + + + + 23 733 Heteroteuthisdispar(Rüppell, 1844) - - - - - - + - 327 450 Rossia macrosoma (DelleChiaje, 1830) - - - + + + + + 82 742 OrderTEUTHOIDEA Family LOLIGINIDAE LoligovulgarisLamarck, 1798 + + + + + + + 10 412 LoligoforbesiiStreenstrup, 1856 + - - + + + - - 19 260 Alloteuthismedia (Linnaeus, 1758) + + + + + + + - 10 334 FamilyENOPLOTEUTHIADE Abradaverany(Rüppell, 1844) _ _ _ - + + + + 107 750 FamilyONYCHOTEUTHIDAE Onychoteuthisbanskii(Leach, 1817) — _ _ — — - + + 335 830 FamilyHISTIOTEUTHIDAE Histioteuthisbonnellii(Férrusac, 1835) + + 167 581 Histioteuthisreversa (Verril, 1880) - - - - - - - + 564 830 FamilyOMMASTREPHIDAE Illexcoindetii(Verany, 1839) + + + + + + + + 12 733 Todaropsiseblanae(Ball, 1841) - - + + + + + + 19 733 Todarodessagittatus (Lamarck, 1798) - + - + + + + + 77 623 OrderOCTOPODA FamilyOCTOPODIDAE OctopusvidgarisCuvier,1797 + + + + + + 17 396 Octopus macropusRisso, 1826 - - - - - + - - - 107 336 OctopussalutiiVerany, 1851 - - + + + + + - 29 463 OctopusdefilippiVerany, 1851 - - - - + + - - 93 260 Scaeurgusunicirrhus (DelleChiaje, 1830) + + + + + + + - 33 344 Pteroctopustetracirrhus (DelleChiaje, 1830) - - - - + + - - 110 239 Eledonemoschata (Lamarck, 1798) + + + + + - - - 10 163 Eledonecirrhosa (Lamarck, 1798) - + + + + + + - 17 462 Tab. 1. ListofcephalopodscaughtintheNorth and CentralAdriaticduring demersaltrawlsurveyscarriedoutfrom 1982to2004. Tab. 1. Lista deicefalopodi rinvenuti inAltoe MedioAdriaticonell'ambitodi trawlsurveysdal 1982 al2004. Sepiola intermedia was caught only in the spring and in viduals); in the spring and in the summer 17 specimens the summerfrom 10 to 120 m ofdepth (112individuals), were found up to 230 mofdepth. mainly alongcoastal areas. Only four specimens of Sepiola rondeletii were caught: Sepiolaligulatawascaughtbetween20and200mofdepth twoindividualsin thesummerof1999offPesaro (depth: (50 individuals),with catches increasing with depth. 21 m) and two in the summerof2000 offAncona (75 m). Sepiola robusta was regularly found in the northernmost Rondeletiola minor is mainly distributed in the middle 64 part of the studied area, up to 100 m ofdepth (292 indi- partoftheAdriaticbasin (Fig. 5). Previously, itwas doc- Distribution and some biological aspects of cephalopods in Fig.4. DistribuzionedelgenereSepiola. the North and Central Adriatic Fig. 5. Distribution areaofRondeletiolaminorand Rossiamacrosoma. Fig. 5. DistribuzionediRondeletiolaminoreRossiamacrosoma. 65 Elisa Ciavaglia & Chiara Manfredi Fig. 6. DistribuzionediLoligoforbesii, Abraliaverany, HistioteuthisreversaeOnychoteuthisbanksii. m umented up to 150-200 ofdepth (Bello, 1990;Guescini Allotenthis media is widely found in the Middle and & Manfrin, 1986). Recent data pointout that this species NorthAdriatic from 50 to200m ofdepth with occasion- is common from 150 to 300 m of depth, but four speci- alcatchesdownto500m. Inall seasonsA. media is more m m mens were caught also at depth from 20 to 80 to the concentrated to the depthof50 and itsabundancede- m north of Ancona and four specimens beyond 450 in creases with depth. Unlike the North Tyrrhenian Sea the area in front of the Gargano promontory. It was al- (Mannini & Volpi, 1989), seasonal migrations towards ways caught together with S. oweniam, as observed in thecoastal waters werenot observed during the spawn- otherMediterraneanareas (Mannini &Volpi, 1989;Jereb ingperiod. Thelargestabundanceindiceswerereported & Di Stefano, 1995). in theautumnsurvey (Soro & Piccinetti-Manfrin, 1989). Rossiamacrosomaismainlydistributedintheareainfront m of the Gargano, from 100 to 450 of depth (192 speci- Ommastrephidae m mens); three specimens were netted at 82 offGiulian- ova (Fig. 5). As itwasobservedin theAegean and in the Todarodes sagittatus and Todaropsis eblanae share a simi- Ionian Seas (D'Onghia et al., 1995), in the northern and lar distribution (Fig. 7). Todarodes sagittatus was rarely m central Adriatic as well mature individuals were found caught,mainlyinthecentralAdriaticfrom90to610 of throughout all seasons. The smallest mature female depth,althoughnineindividualswerecaughtat70m;in measured 47 mm and the smallest male 31 mm. Mantle all, 54 individuals were caught. Catches increased with length ranged from 26 to 58 mm in males, from 30 mm depth,from0.08kg/km2 inthe50-100mstratum,to 1.57 , on in females. The latterwere found toreachlargersizes kg/km2 in the500-800 mstratum. , thantheformer,asobservedintheAegeanSea(D'Onghia Todaropsis eblanae was fished only in the central Adriatic m et al, 1994). (1,014 specimens), from 20 to 750 of depth, often to- getherwith lllexcoindetii, as also found byGamulin-Bri- Loligimdae da & Ilijanic (1972). It showed the largest abundance m from 200 to 300 ofdepth. Biological data of Todaropsis Loligoforbesii is an uncommon species in the surveyed eblanaepointoutawidereproductiveperiod:intheAdri- m area. It prefers depths over 50 (Relini et al., 1999) but atic Sea individualsare found atdifferentmaturitystag- 13 specimens were caught at shallower depths in the es all year though, as in the Tyrrhenian Sea (Favilla, northern Adriatic, especially in the spring and in the 1996). Mature specimens were found throughout the summer (Fig. 6); the same depth ofcapture was already year although they were more abundant in the spring 66 documented in the southern AdriaticSea (Bello, 1990). and autumn. This is in agreement with Mangold-Wirz Distribution and some biological aspects of cephalopods in Fig. 7. Distribuzionedi Todarodessagittatuse Todaropsiseblanae. the North and Central Adriatic Fig. 8. Distribuzionedi Octopusdefilìppi, OctopussalutaeScaeurgusunicirrhus. 67 : (1963) who documented the reproductive period from Only few species show anomalous occurrences. Some March to November. Males reach maturity at a smaller specimens of Octopus salutii were fished at depths shal- ML than females (Relini et al., 1999). In the study area lower than 50 m, Todarodes sagittatus at 70 m, whereas mm the minimum size for a mature male was 100 and two specimens of Sepiola ajfinis, which is considered a for a mature female 110 mm. Females attained larger shallow waterspecies, werefound deeper than200 m. sizes than males, in agreement with Relini et al. (1999). Inthestudyarea,littlebiologicalinformationisavailable Themaximumsizeobservedwas 185mminfemalesand about rarely caught species as well as low commercial mm 140 in males. value ones, compared to fishery targeted cephalopods. This study makes us suppose a broad reproductive pe- Octopodidae riod for some species such as Todaropsis eblanae, Sepia el- egans, Sepiaorbignyana, Sepiettaoweniana,and Rossia mac- Herein only data about Octopus salutii, Octopus defilippi, rosoma. We expect to widen the knowledge of many as- Elisa and Scaeurgus unicirrhus are reported; for the other spe- pectsoftheecologyandreproductivebiologyofallthese cies of the order Octopoda, refer to Casali et al. (1998) species with the data tobe collected in thenextsurveys. Ciavaglia and Soro & Paolini (1994). & These species inhabit sandy and muddy bottoms (Man- gold-Wirz, 1963; Boletzky, 1977). The most abundant Acknowledgements Chiara catcheswererecordedintheareain frontoftheGargano promontory (Fig.8),probablybecauseofsuitabledepths We would like to thank Dr. Gabriella Manfrin for her Manfredi and type ofbottom. help and support during the fieldwork and Dr. Giam- Amongthesespecies,O.salutiishowedthewidestbathy- battista Bello for constructive suggestions and language metric range: 279 specimens were caught from 50 to 450 improvement of the text. Thanks are also due to two m of depth; in the summer of 2004 an individual was anonymous referees for their useful comments on the caughtat30 m. Catches increased with depth. manuscript. In other areas of the Mediterranean S. unicirrhus is occa- sionally found, usually deeper than 100 m (Mannini & Volpi., 1989; Bello, 1990; Pastorelli et al., 1998). In the References studyareaitwascollectedinthebathymetricrangefrom m 50 to400 (102specimens);some individualswere also Belcari P. & Sartor R, 1993. Bottomtrawling teuthofauna of caught in thenorthern partoftheAdriatic. theNorthernTyrrheniansea. SdenticiMarina, 57 145-152. : Octopus defilippi showed the lowest abundance; it was Bello G., 1990. The cephalopod fauna of the Adriatic. Acta rarely caught (33 specimens) and only in the southern Adriatica,31: 275-291. area ofthe MiddleAdriatic atdepths from 110 to260m. Bello G., 2003. The biogeography of Mediterranean cephalo- pods. Biogeographia,24 209-226. : Boletzky S.v., 1999. Biologie et biogéographie des céphalop- Rare species odes actuéis. Bulletin de la Société Géologique de France, 170: 205-215. Abraliaveranyisapelagicspecieswhichwascaughtonly Boletzky S.v., 1977. Le développement embryonnare de Scae- in summer off the Gargano promontory. The largest urgus unicirrhus contribution a l'étude du "Macrotritopus m catcheswere recorded from 300 to400 ofdepth; how- problem" (Mollusca, Cephalopoda). Rapports et procès-ver- everitwasfound inawiderbathymetricrange,from 100 baux des reunions de la Commission internationale pour to 750 m (Fig. 6). VexplorationscientifiquedelaMerMediterranée,24(5): 53-63. Only 12individualsofOnychoteuthis banskiiwerecaught CasaliR,ManfrinPiccinettiG. &SoroS., 1998.Distribuzi- in three summer surveys at depths from 330 to 830 m one di cefalopodi inAltoeMedioAdriatico. BiologiaMarina H(iFisgt.io6u)t.euthisreversawascaughtinthesummer.Thefour D'MOendigtheirraanGe.a,,M5at(2a):r3r0e7s-e317A.., Tursi A., Maiorano P. & Pan- m etta R, 1995. Osservazioni sulla teutofauna epi e mesoba- nettedindividualswerefoundbetween550and830 of tialenelMediterraneoorientale(MarIonioeMarEgeo). Bio- depth (Fig. 6). logiaMarinaMediterranea,2(2): 199-204. 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