[RT" Italiana cli Paleontologia e Stratigrafia

LAMPADENA IONICA: A NE\T TELEOST FROMTHE 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 statariconosciuta dalla biozona a "la,rge Gephyrocapsa" a quella a Pseudoe

milianìa lacunosa ed apparentemente sembra estinguersi prima dellafine del Pleistocene. Sebbene il genere Lampadena sia rutrora viventcsolo negli oceani Atlantico e Indo-Pacifico, esso è noto nel Mediterra-neo sin dal Miocene inferiore. L. ionica sembra essere I'unica specie diquesto genere esìstente neÌ Pleistocene del Mediterraneo. Sino ad oraè stata rinvenuta associata a fauna bentonica (invertebrati e vertebr,ltì1caratteristica 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 ofsouthern Italy. In particular, L. ionica is known from the "large Gepby-rocapsa" up to the Pseudoemiliania lacunosa biozone. Apparentlr.., thespecies became extinct before the end of the Pleistocene. Although thegenrs 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 thePleistocene deposits of the Mediterranean area.

The new species has been found only associated nith benthicfaunas (invertebrate and vertebrate) indicating an bathyal environmentdeeper than 500m.

Introduction

The genus Lampadena Goode & Bean, 1896 wasknown as a fossil in the Mediterranean realm fromMiocene and Pliocene deposits but is not representedthere in the Recent fauna. The Miocene species are theextincr L. gracile (Schubert 1912) and L. speculigeroidesBrzobohaty 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 Recentfauna, the latter species is restricted to the southern partof the Indian and Pacific Oceans, betsreen the latitudes

of approximately 2O"S and 5O"S (Nafpaktitis & Paxton1968). Recent taxonomic srudies on the MediterraneanPleistocene otoliths revealed the presence of a newspecies, Lampadena ionica, from two sites in southernItaly. The otoliths of the species described here can eas-

i1y be distinguished from the other Lampadena speciesof 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 insouthern Italy.

Montalbano Jonico - A composite succession ofmarine Pleistocene deposits, over 400 m thick, wasreconstructed aiong the internal border of the southernApennines Foredeep (Montalbano Jonico area, Fig. 1)

(Ciaranfi et al. 1997). It consists mainly of terrigenousclayey-silts and silty-clays in the basal and middle part,and of sandy-silts, silty-sands, and sand bodies in theupper part. Nine volcaniclastic layers (V1-V9) are

included at various heights, and a marine conglomeratetops the succession (Fig. 1).

Calcareous nannofossil assemblases indicates a

lower to middle Pleistocene age, and the successron wasproposed 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 "smallGephyrocapsa" nannofossil biozones, and the middle andupper part to the Psewdoemiliania lacunosa nannofossilbiozone (Ciaranfi et al. 2001; Marino 1996).

Palaecological, taphonomical, and sedimentologi-cal analyses suggest a general regressive trend fromupper slope to shoreface environmenrs.

The otoliths studied were collected from the first1.2 m of the section, named Entalina section, whichbelongs to the Lower Pleistocene (Iarge Gephyrocapsanannofossil 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 bybathyal 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-middlePleistocene in age, mainly consists of well stratifiedpelitic sediments with a general slight westward slopingof the beds (Di Geronimo et al. 1997).The depositionoccurred 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 fishotoliths (Girone 2OO0) testify a bathyal palaeoenvrron-ment, 5OO to 1OOO m deep. The occurrence of L. ionicain the section is show in Fie. 1.

Systematic palaeontology

Terminology applied to otolith morphology fol-lows Nolf (1985). The classification adopted is that ofRosen (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 wellmarked 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, leitorolìth': ll-s' p.rr.rrrLrc': r'i1hr orolirh': dl renrrr'ri.*.

495

Fig. l

licular crest is elongated and separated from the cristainferior 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 depressionis deeper just above the crista superior and becomesmore smooth towards the dorsal rim. The ventral area is

slightiy conyex and bears a shallou.. groove near to theventral rim. In large specimens, the dorsal rim is trun-cated in its posterior part (behind rhe posterodorsalangle) where it reaches its maximum height. In sn-iall

specimens the dorsal rim shows a regular curr.e. Theventral rim is curved but not semicircular and bears

irregularly spaced obtuse spines. The anterior spines i3or 4 in number) are more pointed and are closer to each

other. They are separated b). deep furrou's on the innerface. The posterior rim is globally rounded, but the loba-tion shows a marked variability among the availablespecimens. In several llrger ones, it bears two or threelobes near the posterodorsal angle. The outer face is

irregularly convex with the maximal convexity located inthe posterior part. The surface of this face shows a low.

elongate umbo in the central part and numerous radiallobes 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 ttNolf, 1996 from the Mediterranean Miocene bv theirnotched 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 stomachcontenr of r pigmv sperm whale (.Kogia simus) caughtoff Taiji, Japan, are different from all other presentlyknori,'n Lamapadena species and apparentlv represent e

ninth species (P1. 1, tig.22), but the entire fishes havenot been causht. Otoliths of all known taxa have beenfigured bv various authors and those pictures are scar-tered through the literature. References to rhese sourcesare listed in Table 1, which also provides a sur\-ey of rhegeographic distribution of each taxon. Otoliths of al1

those Recent taxa are also figured on Pl. 1, which forseveral of species, provides gro\\rth series that never havebeen 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, inadult specimens, a less notched superior part of thepostrior rim. In the medium-sized otoliths of the pres-ent Pleistocene species and the Recent L. luminosa, thetwo lrtter features appear to be very similar, but themedium-sized as q.ell as the large specimens of L, ionìcaare less oblongate and have a wider-shaped anrerior parrthan 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.9-10Cor-strl,a.N & Nappaxrtr:s, 1912, fig. 4C, p, 7NAFpAKrns & P,qxroN, 1968, fig. 10.8, p. 24

Pl. l. Fig. 5-8Nappa,rrrrrs & P,txroN, 1968, fig. 10.7 , p. 24Suele et al., 1995, pI.21, fig. E1-3

PÌ. 1, Fig. 18

Napparrrrrs & PaxroN, 1968, fig. 10.6,p.24Reow, nsra,1992, fig. 36, p. 186

Pì. 1, Fig. 1-4Napperrrrrs & P,q.xroN, 1968, fig. 10.1, p. 24RrveroN & BounRer, 1999, pl. I 19, figs. 1-13Sua.r-E et al., i995, pl. 21, fig. F1-2

Pl. 1, Fig. 1 i-12NnppRrrrrrs & PaxroN, 1968, fig. 10.5,p.24Suers et al., 1995, pl. 2I, fig. G2,not Gl(?: Bolinichthys)

Pl. 1, Fig. 20-2iKRrppr, 1970, fig. 3, p.280

Pl. I, Fig. 13-15Nerpe.rtrrrs & Pe,xroN, 1968, fig. 4, p. 12, frg.10.1 , p. 24Svers er al,, 1995, pi. 21, fig. A1-3

Pl. 1, Fig. 16-17KotLyeR & P,qnlN, 1990, fig. 2, p. 101

Nepperrlrrs & PAXroN, 1968, fig. \0.2,p.24RlvaroN & Bounnrr, 1999, pl. 1 19, figs. 14-19

Pl. 1, Fig. 22Nerparcrrrrs & Paxrou. 1968, fig

19FioP1

10.3

Atlantic, Indian Ocean,east and central Pacific

N and S Atiantic,antitropical; southemIndian and Pacificoceans

Southem parts ofallthree oceans, between20 and 50"S

Atlantic, eastern Pacifi c

Southern Ocean

Atlantic, mainlyMauritanian upwelling

N Atlantic, S Atlantic,S Indian ocean, SEPacifrc

Eastern Pacific

N Atlantic

Off Japan, otoliths fromstomach content ofsperm whale Kogiasinasl 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 toother 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 NorthAtlantic L. notialis Nafpaktitis & Paxton, 1968 (P1. 1,

fig. 11-12), the Pleistocene specimens differ from thelatter by having a larger sulcus, a more salient pos-terodorsal angle and a more expanded anteroventralarea. In L. notialis the antirostrum is well developedwhile in L. ionica it is almost absent. These differencescan also be observed in the medium sized-otolith of Z.

notialis from the south Atlantic figured by NafpaktitisEr Paxton (1968, fig. 10) in which the anterior part of thedorsal 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 fossilfrom the Lower Pliocene deposits of the Mediterraneanrealm, by their marked rostrum and their more obtuseposterodorsal angle which, in the recent species, is near-1y right. Moreoveq L. dea generally shows a well distinctrostrum and antirostrum (Nafpaktitis Er Paxton 1968).

The more obtuse posterodorsal angle is also thefeature that distinguishes otoliths of L, ionica fromthose 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) LampadenaanomalaParr,1928,left otoliths. Recent: Atlantic (Researchvessel Knorq station 65). Fig. 11-12) Lampadenanotìalis Nafpaktitis & Paxton, 1968, leftotoliths. 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 speculigeraGoooE & BEAN, 1896

lmm-\ -J tt

,/f- -\""/rl€l+;gl

,,r'.\iMtoLampadena nolialis

NAFPAKTITIS &PAXTON, 1968

Lampadena pontiÉr KREFFT, 1970

Lampadena urophaos PAxroN, 1963

Lampadena dea

FRASER-BRUNNER, 1949

Lampadena urophaos atlanticaMAUL, 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 posteriormargin 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 fromthe Archi Section (Calabria, Southern Itaiy), referable tothe "sma11 Gephyrocapsa" and the Pseudoemilianialacwnosa 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 Lampadenaappear to be among the deepest-dwelling myctophids;they are merely captured below 600-700 m. L pontifex

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L. ionica was collected from beds which, accord-ing to their content of benthic invertebrates and benth-ic fish assemblages, indicate a bathyal environment ofmore than 500 m deep (Girone 2OOOt. In the Montal-bano sections, L. ionica is absent in the more shallow

associations referable to the "small Gephyrocapsa" bio-zone.

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