[RT" Italiana cli Paleontologia e Stratigrafia LAMPADENA IONICA: A NE\T TELEOST FROM THE MEDITERRANEAN PLEISTOCENE. ANGELA GIRONE'! & DIRK NOLF'I'I Recei.oed December 27,2Aa1; accepted May 3,2Aa2 Key uords: Lampadena, Pleistocene, sourhern ftaly. Riassunto. Viene descritta la nuova specie Lampadena ionica (Myctophidae, teleostei) riscontrata in depositi del Pleistocene infe- riore-medio dell'Italia meridionale. In particolare L. ionica è stata riconosciuta dalla biozona a "la,rge Gephyrocapsa" a quella a Pseudoe milianìa lacunosa ed apparentemente sembra estinguersi prima della fine del Pleistocene. Sebbene il genere Lampadena sia rutrora viventc solo negli oceani Atlantico e Indo-Pacifico, esso è noto nel Mediterra- neo sin dal Miocene inferiore. L. ionica sembra essere I'unica specie di questo genere esìstente neÌ Pleistocene del Mediterraneo. Sino ad ora è stata rinvenuta associata a fauna bentonica (invertebrati e vertebr,ltì1 caratteristica di ambienti batiali indìcante una profondità maggìore di )uu lnetrt. Abstract. The new spectes Lampadena ionica (Mvctophidae, Teleostei) is described from lou.er and middle Pleistocene deposits of southern Italy. In particular, L. ionica is known from the "large Gepby- rocapsa" up to the Pseudoemiliania lacunosa biozone. Apparentlr.., the species became extinct before the end of the Pleistocene. Although the genrs Lampadena Iives only outside the Mediterranean today, ìr is known from the Mediterranean reaim since the early Miocene. Z. lon - lca seems to be the only species of the genus Lampadena existing in the Pleistocene deposits of the Mediterranean area. The new species has been found only associated nith benthic faunas (invertebrate and vertebrate) indicating an bathyal environment deeper than 500m. Introduction The genus Lampadena Goode & Bean, 1896 was known as a fossil in the Mediterranean realm from Miocene and Pliocene deposits but is not represented there in the Recent fauna. The Miocene species are the extincr L. gracile (Schubert 1912) and L. speculigeroides Brzobohaty Ee Nolf, 1996.In the Pliocene, the Recent Z. dea Fraser-Brunner, 1949 is recorded from Zanclean (Lower Pliocene) deposits of the western Mediterranean (Nolf er Cappetta 1988; Nolf et al. 1998). In the Recent fauna, the latter species is restricted to the southern part of the Indian and Pacific Oceans, betsreen the latitudes of approximately 2O"S and 5O"S (Nafpaktitis & Paxton 1968). Recent taxonomic srudies on the Mediterranean Pleistocene otoliths revealed the presence of a new species, Lampadena ionica, from two sites in southern Italy. The otoliths of the species described here can eas- i1y be distinguished from the other Lampadena species of the Mediterranean Miocene and Pliocene as s/ell as from the living species. Otoliths of Lampadena ionica n. sp. were collect, ed from silty Pleistocene deposits ar rwo locations in southern Italy. Montalbano Jonico - A composite succession of marine Pleistocene deposits, over 400 m thick, was reconstructed aiong the internal border of the southern Apennines Foredeep (Montalbano Jonico area, Fig. 1) (Ciaranfi et al. 1997). It consists mainly of terrigenous clayey-silts and silty-clays in the basal and middle part, and of sandy-silts, silty-sands, and sand bodies in the upper part. Nine volcaniclastic layers (V1-V9) are included at various heights, and a marine conglomerate tops the succession (Fig. 1). Calcareous nannofossil assemblases indicates a lower to middle Pleistocene age, and the successron was proposed to locate the GSSP of the Lower-Middle Pleis- tocene (Ciaranfi et ^1. 1,997). The basal part of the sec- tion belongs to the "Iarge Gephyrocapsatt and "small Gephyrocapsa" nannofossil biozones, and the middle and upper part to the Psewdoemiliania lacunosa nannofossil biozone (Ciaranfi et al. 2001; Marino 1996). Palaecological, taphonomical, and sedimentologi- cal analyses suggest a general regressive trend from upper slope to shoreface environmenrs. The otoliths studied were collected from the first 1.2 m of the section, named Entalina section, which belongs to the Lower Pleistocene (Iarge Gephyrocapsa nannofossil biozone). The benthic invertebrate associa- 'lDipartimento di Geologia e Geofisica, Unìversità di Bari, r.ia E. Orabona 4,7A125 Bari : email girone@geo.uniba.ir 'r'rInstìtut royal des Sciences naturelles de Belgique, rue Vautier 29, B-1000 Bruxelles: email Dirk.Nolf@natuurwetenschappen.be 494 A. Girone & D. Nolf tions and the otolith associations are dominated by bathyal species suggesting a palaeodepth of 500-600 m (Ciaranfi et al. 2001; Girone 2000). Archi - This section, 9 m thick, is located near Reg- gio Calabria, southern Italy, (Fig. 1) at about 90 m above the sea level and is particularly well-exposed along a N- S abandoned quarry front. The section, lower-middle Pleistocene in age, mainly consists of well stratified pelitic sediments with a general slight westward sloping of the beds (Di Geronimo et al. 1997).The deposition occurred during a cold phase, as inferred from the plank- tonic foraminiferal assemblages. The invertebrate benth- ic faunas (foraminifers, moiluscs, bryozoans, and ser- pulids) (Di Geronimo et al. op. cit.) as well as the fish otoliths (Girone 2OO0) testify a bathyal palaeoenvrron- ment, 5OO to 1OOO m deep. The occurrence of L. ionica in the section is show in Fie. 1. Systematic palaeontology Terminology applied to otolith morphology fol- lows Nolf (1985). The classification adopted is that of Rosen (1973) Subsection CTENOSQUAMATA Rosen, 1973 Sept SCOPELOMORPHA Rosen, 1923 Order Myctophiformes Regan, 1911 Family Myctophifdae E Gill, 1893 Genus Lampadena Goode & Bean, 1896 Fio I Loc.rrion of Monrelb:no Jon- ico and Archi sections. Type species. - Lampadena speculigera Goode tr Bean, 1896. The otolìths of the species of the genus Lampadena generally have large and oval-shaped otoliths, considerabiy longer thrn high. They can show a slight or absent posterodorsal notch. Only one Recent species L. anomala has a small and more round-shaped otolith. Lampadena ionica new species Fig. 2 a-e Etymology, - From the Montalbano Jonico Section (Basilicata, southern Italv), the type locality. Type materiaì, - Holorrpe: a left oroìith 'Fig. 2ar /DGCUB AG 1) from the Entalina section (Montalbano Jonico Composite Sec- tion) Lon er PÌeistocene in age; 1O parat,vpes! of which four are figured (Fig. 2b-e) (DGGUB AG 2-s). Type locality. - Montalbano Jonico section, southern ltalv. Repository. - Dipartimento di Geologia e Geofisica, Università degli Studi di Bari, Italy (DGGUB AG1-s). Dìagnosis. A species characterised by moderately thin, oval shaped otoliths becoming more elongated with the growth. The ros- trum is large and salient with no antirostrum. The posterodorsal angle is r-ide and moderately notched. The sulcus is t-ide. Description This species is characterized by oval-shaped otoliths with a wide sulcus. The ostium is somewhat longer than the cauda. The caudal colliculum is narrow- er than the ostial one, which becomes slender towards the anterior end. The cristae become obsolete in the posterior part of the otolith; the superior one is well marked and more linear than the inferior one. The col- Lampadena ioníca, a nea teleost Lampadena lozlca, lVlontalbano Jonico, Ìou'er Pleistocene (large Gephyrocapsa biozone): a) holotl-pe; left otolith; b-c) paratypes, leit orolìth': ll-s' p.rr.rrrLrc': r'i1hr orolirh': dl renrrr'ri.*. 495 Fig. l licular crest is elongated and separated from the crista inferior by a deep furrow The dorsal and ventral areas are very similar in size; the former is characterized b1- :r depression covering most of its surface. This depression is deeper just above the crista superior and becomes more smooth towards the dorsal rim. The ventral area is slightiy conyex and bears a shallou.. groove near to the ventral rim. In large specimens, the dorsal rim is trun- cated in its posterior part (behind rhe posterodorsal angle) where it reaches its maximum height. In sn-iall specimens the dorsal rim shows a regular curr.e. The ventral rim is curved but not semicircular and bears irregularly spaced obtuse spines. The anterior spines i3 or 4 in number) are more pointed and are closer to each other. They are separated b). deep furrou's on the inner face. The posterior rim is globally rounded, but the loba- tion shows a marked variability among the available specimens. In several llrger ones, it bears two or three lobes near the posterodorsal angle. The outer face is irregularly convex with the maximal convexity located in the posterior part. The surface of this face shows a low. elongate umbo in the central part and numerous radial lobes in the marginai zone. Affinities. Otoliths of Lampaclena ionica can be dis- tinguished from those of the fossil species L. gracile (Schubert, 1912) and L. speculigeroides Brzobohaty tt Nolf, 1996 from the Mediterranean Miocene bv their notched and wide posterodorsal angle, their less pro- nounced rostrum and a more rounded and convex ocrs- terior mrrgin. In Lhe tn o M iocene species, this nrargin is linear and subvertical (Brzobohaty & Nolf 1996, pI. 4, figs. 1-6 and figs. 12-1o1. ln the Recent fauna, the genus Lampadena is rep- resented by eight nominal species and one subspecies (Paxton 1979). Orcliths collected from the stomach contenr of r pigmv sperm whale (.Kogia simus) caught off Taiji, Japan, are different from all other presently knori,'n Lamapadena species and apparentlv represent e ninth species (P1. 1, tig.22), but the entire fishes have not been causht. Otoliths of all known taxa have been figured bv various authors and those pictures are scar- tered through the literature. References to rhese sources are listed in Table 1, which also provides a sur\-ey of rhe geographic distribution of each taxon. Otoliths of al1 those Recent taxa are also figured on Pl. 1, which for several of species, provides gro\\rth series that never have been published before. Otoliths of L. ionica differ from those of L. lumi- nosa (Garman 1899) (P1. 1, fig. 1-4) and L. urophaosPax- ton,1963 (Pl. 1, fig. 16-17) by a wider sulcus, a pos- terodorsal angle which is conspicuously obtuse and, in adult specimens, a less notched superior part of the postrior rim. In the medium-sized otoliths of the pres- ent Pleistocene species and the Recent L. luminosa, the two lrtter features appear to be very similar, but the medium-sized as q.ell as the large specimens of L, ionìca are less oblongate and have a wider-shaped anrerior parr than I. lumirtosa. The features that distinguish the Pleis- tocene species from the Pacific L. urophaos are much NOMINAL SPECiES FIGURED OTOLITHS DISTRIBUTION Lampadena anomala Pan. I 928 Lampadena chavesí Collet, 1905 Lampadena dea Fraser-Brunner, 1949 Lampadena luminosa (Garman. 1899) Lampadena notialís Nafpaktitis & Paxton, 1968 Lampadena pontifex Krefft, 1970 Lampadena speculigera Goode & Bean, i896 Lampadena urophaos urophaos Paxton, 1963 Lampadena urophaos atlantica Maul, 1969 Lampadena sp. A Pl. 1, Fig.9-10 Cor-strl,a.N & Nappaxrtr:s, 1912, fig. 4C, p, 7 NAFpAKrns & P,qxroN, 1968, fig. 10.8, p. 24 Pl. l. Fig. 5-8 Nappa,rrrrrs & P,txroN, 1968, fig. 10.7 , p. 24 Suele et al., 1995, pI.21, fig. E1-3 PÌ. 1, Fig. 18 Napparrrrrs & PaxroN, 1968, fig. 10.6,p.24 Reow, nsra,1992, fig. 36, p. 186 Pì. 1, Fig. 1-4 Napperrrrrs & P,q.xroN, 1968, fig. 10.1, p. 24 RrveroN & BounRer, 1999, pl. I 19, figs. 1-13 Sua.r-E et al., i995, pl. 21, fig. F1-2 Pl. 1, Fig. 1 i-12 NnppRrrrrrs & PaxroN, 1968, fig. 10.5,p.24 Suers et al., 1995, pl. 2I, fig. G2, not Gl(?: Bolinichthys) Pl. 1, Fig. 20-2i KRrppr, 1970, fig. 3, p.280 Pl. I, Fig. 13-15 Nerpe.rtrrrs & Pe,xroN, 1968, fig. 4, p. 12, frg. 10.1 , p. 24 Svers er al,, 1995, pi. 21, fig. A1-3 Pl. 1, Fig. 16-17 KotLyeR & P,qnlN, 1990, fig. 2, p. 101 Nepperrlrrs & PAXroN, 1968, fig. \0.2,p.24 RlvaroN & Bounnrr, 1999, pl. 1 19, figs. 14-19 Pl. 1, Fig. 22 Nerparcrrrrs & Paxrou. 1968, fig 19FioP1 10.3 Atlantic, Indian Ocean, east and central Pacific N and S Atiantic, antitropical; southem Indian and Pacific oceans Southem parts ofall three oceans, between 20 and 50"S Atlantic, eastern Pacifi c Southern Ocean Atlantic, mainly Mauritanian upwelling N Atlantic, S Atlantic, S Indian ocean, SE Pacifrc Eastern Pacific N Atlantic Off Japan, otoliths from stomach content of sperm whale Kogia sinasl fish unknown 496 A. Girone & D. Nolf Table 1 - List of Recent species of the genus Lampadena. The references on iconography and the geographic distribution are also reported. more marked in the Atlantic subspecies L. urophaos atlantica Maul, 1969 (Pl. 1, fig. 19) that, in addition to other features, has a more elongated antirostrum and a more salient rostrum than L. ionica. Although the general shape of otoliths of L. ioni- ca shows some similarity to those of the Recent North Atlantic L. notialis Nafpaktitis & Paxton, 1968 (P1. 1, fig. 11-12), the Pleistocene specimens differ from the latter by having a larger sulcus, a more salient pos- terodorsal angle and a more expanded anteroventral area. In L. notialis the antirostrum is well developed while in L. ionica it is almost absent. These differences can also be observed in the medium sized-otolith of Z. notialis from the south Atlantic figured by Nafpaktitis Er Paxton (1968, fig. 10) in which the anterior part of the dorsal area is wider than in L. ionica. Otoliths of the I. ionica differs from those of Z. deaFraser-Brunner, 1949 (P1. 1, fig. 18), known as fossil from the Lower Pliocene deposits of the Mediterranean realm, by their marked rostrum and their more obtuse posterodorsal angle which, in the recent species, is near- 1y right. Moreoveq L. dea generally shows a well distinct rostrum and antirostrum (Nafpaktitis Er Paxton 1968). The more obtuse posterodorsal angle is also the feature that distinguishes otoliths of L, ionica from those of the recent L. pontifex (Pl. 1, [rg. 20-21) and Pl. 1 Lampadena ionica, a nezu teleost 497 PLATE 1 The Recent specimens figured in this plate are from the collection IRSNB (Institut Ro1'al des Sciences Naturelles de Belgique). Fig. 1-a) Lampadena lumi- nosa,leftotoliths.Recent: Gulf ofMexico.Fig.5-B)Lampadenacha'",esiCollet,1905, leftotoliths.Recent:Atlantic,off theAzores.Fig.g-10) Lampade naanomalaParr,1928,left otoliths. Recent: Atlantic (Researchvessel Knorq station 65). Fig. 11-12) Lampadenanotìalis Nafpaktitis & Paxton, 1968, left otoliths. Recent: 11) South \West Atlantic (Research vessel Valter Herwìg statìon 362), coll. SCH\IARZHANS; 12) off New South \Wales, Australia. Fig. 13-1.5)Lampadenaspeculi.geraGoode&Bean, 1896, leftotoliths.Recent:13)Atiantic,offNlllreland;14-15)GulfofBiscaye.Fig. 16-17) Lampadena uropbaosPuton, 1963, leftotoliths:fig. 16) NewCaledonia,fig. 1Z)California,Recent:16) offNewCaledonia;12)offCalifornia.Fig. 18 Lampadena deaFraser-Brunner,I949,left otolìths. Recent: SE Pacific. Fig. 79) Lampadena urophaos atkiltic.lMaul, 1969, right otoliths. Recent: off Portugal, Atlanric. Fig. 20-21) Lampadenapontifex lkeflì, 1970, left otoliths. Recent: Atlantic (Research vessel Atlantis II sration 59). Fig.22) Lampadena sp. A,left otolith. Recent: from the stomach content of a pigmy sperm whale (Kogia simus) caught offTaiji,Japan. Redraq afterNafpaktitis Er Paxton (1968, fig. 10.3) Lampadena luminosa (Genmr, 1899) Lampodena chavesi CoLLErr, 1905 PARR, i928 :.--=--_---1t" Lampadena speculigera GoooE & BEAN, 1896 lmm-\ -J tt ,/f- -\"" /r l€l+;gl ,,r'.\iMto Lampadena nolialis NAFPAKTITIS & PAXTON, 1968 Lampadena pontiÉr KREFFT, 1970 Lampadena urophaos PAxroN, 1963 Lampadena dea FRASER-BRUNNER, 1949 Lampadena urophaos atlantica MAUL, 1969 498 A. Girone & D. Nolf REFERENCES Lampadena sp. A (P1. l, frg. 22). In the Recent L, speculigera and L. anomald, rhe general shape of the otoliths is quite different; the otoliths oÍ L. speculigera have a nearly vertical posterior margin and a narrower sulcus with a very short cauda. According to Nafpaktitis & Paxton (1968) and Coleman & Nafpaktitis (1,972), the otoliths of L, ano- mala can be considered as intermediate between the more elongated otoliths of all other Lampadena species and the almost round, smooth-edged otoliths of the genus Thaningichthys (Myctophidae). Otoliths of L. ion- ica c\ear\y belong ro the Lampadena group with elongate otoliths and are quite different from the high-shaped L. anomala morphology. Distribution Lampadena ionica first appears in the Lower Pleis- tocene beds ("large Gepbyrocaps4" nannofossil biozone) in the basal part of the Montalbano Jonico section. The presence of L. ionica in the bathyal assemblages from the Archi Section (Calabria, Southern Itaiy), referable to the "sma11 Gephyrocapsa" and the Pseudoemiliania lacwnosa nannofossil biozones (Di Geronimo et al. 1997;, suggests that this species survived in the Mediter- ranean basin up to the Middle Pleistocene. Most Recent species of the gents Lampadena appear to be among the deepest-dwelling myctophids; they are merely captured below 600-700 m. L pontifex and L. lwminosa occur also at shallower depths between 275 and 450 m (Nafpaktitis et al. 1977). 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