BULLETIN DE [.'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE SCIENCES DE LA TERRE, 70: 185-206, 2000 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN AARDWETENSCHAPPEN, 70: 185-206, 2000

Diaphus Otoliths from the European Neogene (Myctophidae, Teleostei)

by Rostislav BRZOBOHATY & Dirk NOLF

Abstract and in most species, only adult and large otoliths (larger than 2 mm) show clear diagnostic features, while juve­ The revision of Neogene otoliths of the myctophid genusDiaphus, niles cannot be distinguished. The other main problem is based on extensive otolith collections in the Aquitaine Basin, in South­ eastern France, northern Italy. Mediterranean Spain, the Paratethys and that in several Recent D iaphus species, even at the adult the North Sea Basin, has proved the validity of fourteen nominal stage, the otoliths show only a very generalized morphol­ species, among which two are new: Diaphus hefralai an d D. carallo- ogy (see N o lf & C a p p e t t a , 1989, pi. 9, figs. 12-22). In nis. None of these species has been recorded previously from Oligo­ fossil associations, such otoliths remain predominantly cène deposits, and only four are still living today. unidentified; in many cases, it is impossible to state if Key-words: D iaphus, Myctophidae, otoliths, Neogene. such forms represent species with a more generalized morphology, or if they are juveniles of species that aquire diagnostic features only at larger sizes (see B r z o b o h a t y Résumé & N o l f . 1995, p. 257 for a more extensive discussion). Practically, this means that in most fossil associations, La révision des espèces oligocènes du genre D iaphus, basée sur un only about 5 to 10 % of the D iaphus otoliths can be abondant matériel d'otolithes récoltées dans le Bassin d'Aquitaine, le confidently identified at the species level. In the present Sud Est de la France, l'Italie septentrionale, la bordure méditerrané­ enne de l'Espagne, la Paratethys et le Bassin de la Mer du Nord, a study, we recognise 14 nominal species for the European révélé la validité de quatorze espèces nominales dont deux sont nou­ Neogene, but we are conscious that the true number may v elles: Diaphus hefralai et D. farallonis. Aucune de celles-ci n'est be considerably higher. In many of the studied associa­ connue de dépôts oligocènes et quatre seulement existent encore dans la faune actuelle. tions, otoliths of some species exhibiting generalized morphologies at their adult stage seem to make up an M ots-clefs: D iaphus, Myctophidae, otolithes, Neogene. amalgam with juvenile otoliths of others, but attempts to split up such amalgams can only lead to speculation, and neither biostratigraphy or paleobiogeography are served Introduction with such drudgery. The literature on fossil otoliths is full of citations and illustrations of these non diagnosticD ia­ In a previous paper (B r z o b o h a t y & N o l f , 1996), we p h u sotoliths, and citations of all such cases as negative revised all otolith-based myctophid genera from the synonymies or as doubtful taxa would produce exces­ European Tertiary, except the genusD iaphus. The O li­ sively long lists. Therefore, our synonymies are restricted gocène D iaphus (six species) from various European to only those citations that can be evaluated at the species basins were revised by B r z o b o h a t y & N o l f , 1995. From level (eventual question marks with peculiar numbers of a the European Eocene,D iaphus otoliths are only known given series means that the concerned specimens are not from the Aquitaine basin (six species, discussed in N o l f , diagnostic), and the list of doubtful taxa given in the 1988) The present study constitutes the final part of our appendix is restricted to the primary type material to revision of myctophid otoliths from the European Ter­ which the names apply. tiary and is restricted to the Neogene D iaphus species. In We also were reluctant to give formal names to the fossil associations containing mesopelagic otoliths, espe­ otoliths of two well recognizable species because there cially in those from tropical waters.D iaphus otoliths are are five Messinian D iaphus species based on skeletons usually the most abundant and taxonomically diverse for which the otoliths remain unknown. The eventual elements. Unfortunately, they are also the most proble­ double use of names based either on otoliths or on ske­ matic of all myctophids when it comes to otolith identi­ letons can only be solved by the discovery of skeletons fication. with otoliths in situ. In such cases, however, the otoliths The morphology ofD iaphus otoliths changes markedly are usually so badly preserved that they are unsuitable for during ontogeny (seeN o lf & C a p p e t t a , 1989, pi. 8-10), taxonomic use. Therefore, we preferred to introduce for- 186 Rostislav BRZOBOHATY & Dirk NOLF mai names for taxa that are easily recognizable in several posed north of the quarry, and located about 10 to 15m below associations, because we judge that the use by different the rhodolite limestone. Planktonic foraminifera from our sam­ publications of inaccurate names like “ D iaphus sp. 1” pling point were studied by E. B icchi . The presence of Para­ globorotalia pseudokugleri and P. kugleri (even if rare and generates many more problems. small) and of abundant Globoquadrina dehiscens indicate the lower part of the G. dehiscens Subzone (N4b), corresponding the calcareous nannoplankton Zones NN1/NN2. The nanno­ Geographic and stratigraphie position of the most plankton was studied by L. S vabenicka . Besides reworked important localities material (Campanian, Eocene, Oligocene), the youngest species represented is Helicosphaera carteri, which after Y oung The present study is based on material from nearly 100 different (1998) indicates the basis of the Neogene. Other important localities in the European Neogene which were sampled by the species are H. recta. H. obliqua, H. euphratis. Discoaster authors during the last 30 years and also takes in account all deflandrei, Sphenolithus dissimilis, Pontosphaera discopora. relevant literature data. The major collecting areas are listed After Y oung (1998), H. recta occurs till the NN1/2 boundary, below. and L. S vabenicka concluded to a placement in the upper part of the NN1 Zone. AQUITAINE BASIN (South-West France) For the Langhian, we only know a small fauna of the Tanaro All the sites with abundant myctophid otolith material are Formation in Piemonte(S teurbaut , 1983) and a small fauna located in the Paleocanyon of Saubrigues (neighbourhood of from the basement of the Baldissero Complex, in the hills E of Saubrigues, Saint-Jean de Marsacq, Saint-Martin-de-Hinx, Torino. A productive Serravallian site was sampled near “ Ma­ Saint-Etienne-d’Orthe, Peyrehorade). From the Chattian till donna della Neve” , N of Mondovi, in Piemonte. Because this the Langhian, the Paleocanyon has been progessively fdled site is the only one that provided abundant otolith material of with marls which are often very fossiliferous. The following Serravalian age, it requires some further comments. nannoplankton zones have been recognised (Cahuzac et al., V iolanti & G iraud (1992) mentioned otoliths in clayey 1995; see also this paper for the precise location of the sites): deposits at point 20A which they attributed to the Lower NP25 (Chattian): Numerous points around Saint-Étienne Serravallian, Subzone with Orbulina universa of Iaccarino d’Orthe and Peyrehorade. (1985). Their coordinates (sheet Carru, 1/25.000, x = 06.780, NN1 : Haubernet site at Saint-Martin-de-Hinx. y = 19.980) refer to a point N of the hamlet “ Madonna della NN2-NN3 (Lower to Middle Burdigalian): “ Les Platanes” site Neve” , and the exposure is in a deeply incised southern tribu­ at Saint-André-de-Seignanx. tary of the Rio Branzola. About 1000 kg of clay was sampled NN3-NN4 (Upper Burdigalian): numerous localities east and there in 1998 by Cavallo , Hoffman and N olf, and provided south of Saubrigues. several thousands of otoliths. Because the outcrop was origin­ NN5 (Langhian): numerous localities in the direct surroundings ally mapped as Tortonian, and because these clayey sediments of the village of Saubrigues; among those are the sites “ Jean Tic” were so different from the typical arenaceous Serravallian (presently filled in) and “Tauziets” , which provided the rich facies and looked so much like the regional Tortonian facies, otolith associations described by Steurbaut in 1979 and 1984. a sample was submitted to B. H amrsmid for nannoplankton dating. He identified Cyclicargolithus floridanus, Helico­ SOUTHEAST FRANCE sphaera scissura, H. kamptnerii, H. cf. walbersdorfensis. Coc­ Very rich otolith associations of Pliocene blue marls of this colithus pelagicus, Pontosphaera anisotrema, P. multipora, region have been described by N olf & Cappetta (1989), to Braarudosphaera bigelowii, Calcidiscus carlae, Geminithella which paper one should refer for the precise location of the rotula, Reticulofenestra pseudoumbilica, R. cf. gelida, Spheno­ studied localities. The sampled beds range from the basement lithus abies, little Prinsiaceae, Syracosphaera sp. , Micrantho­ of the planktonic foraminiferal Zone MPI 1 ofC ita (1975) lithus vesper, and reworked species from the Cretaceous and (= Zone 1 or Zone A ofS paak , 1983) up to the top o f Zone Oligocene. Discoasterids are nearly completely absent. Because MPI 5a. One should notice, however, that the locality o f Pichegu, of the presence ofCyclicargolithus floridanus and the absence cited as MPI 4b by N olf & Cappetta (after M agne , 1978, p. 376) ofSphenolithus heteromorphus, he attributed the sample to the is now considered as the lower part of Zone MPI 3 (Zanclean; see NN6 Zone, which confirms the Serravallian age of the site. Clauzon et al., 1990, p. 136). The richest associations come For the Tortonian, we have much material from the stratotype from the Zanclean MPI 2 Zone at Le-Puget-sur-Argens, a locality (hills SE of Sant Agata Fossili and Rio Castellania-Mazzapiedi; that was sampled very intensely by the late H. von Hacht . see N olf & Steurbaut , 1983) and from exposures in the Tanaro at Alba. A problem remains for the locality Borelli, in NORTHERN ITALY the Torino-Monferrato Hills, where turbiditic sediments (Mon- In Northern Italy, Aquitanian and Burdigalian strata were taldo Member) provided an interesting otolith association. The sampled in the hills E of Torino and in the Montferrato hills Montaldo Member was first reported as Messinian (Globoro­ (see B onsignore et al., 1969 and N ovaretti et al., 1995 for talia conomiozea Planktonic Foraminiferal Zone,Turborotalia stratigraphical data and B rzobohaty & N olf, 1996 for locality multiloba Subzone) by Pavia & R obba (1979). Subsequent data). Some additional comments concerning our sample from datation of a nannoplankton sample by E. Martini, reported in Moleto are provided here, because this is the only Mediterra­ Janssen (1999) resulted in a “ very probable Tortonian age” , nean locality of Aquitanian age that provided abundant Diaphus and data from the holoplanctonic molluscs also provide indica­ otoliths. In the paper ofN ovaretti et al. (1995, fig. 8), the tions for a Tortonian age, but without really convincing evi­ section of Moleto was studied only down to the basement of the dence. Therefore, date from Borelli are here reported as ? rhodolite limestone, exposed in the lower part of the quarry. It Messinian. For the Pliocene, our Italian material comes mainly was attributed to planktonic foraminiferal Zone N 5/6, which from the Zanclean of Monticello, near Alba(N olf & Cavallo , suggests that the basement of the section does not go down into 1995) from the lower part of the Vemasca - Castel Arquato the Aquitanian. Our sample, however comes from marls ex­ section (BARBIERI, 1967) and from Orciano, in Toscane. European Neogene Diaphus 187

CALABRIA (Southern Italy) H olo type : Figured by H o l e c , 1975, pi. 1, fig. 5; PFUK Myctophid otoliths collected by H. Cappetta at several levels University, Bratislava, coli. Geology and Palaeontology, in the reknowned Plio-Pleistocene Vrica section constitutes Nr I. 42 aa; Badenian of Salka, near Sturovo, Slovakia. the only studied material for the Gelasian and the early Pleis­ tocene. Description: Otoliths of this species can grow fairly large SPAIN and show an elongate shape, with a narrow anterior portion Associations with many myctophid otoliths were sampled and a broad posterior part. Their outline is characterised by a mainly in the Messinian Marls of Torremondo near Venta de very prominent rostrum, a clear antirostrum and a strongly la Virgen, east o f Murcia(M ontenat , 1977, p. 46) and in the marked posterodorsal angle. Their ventral margin is strongly Zanclean of Papiol, near Barcelona(N olf et al., 1998). denticulate with usually 8-12 irregular spine-like denticles. The outer face is clearly convex in its posterior part. The MORAVIA (Czech Republic): Central Paratethys inner face is very slightly convex and nearly flat. Its ventral Our material comes from many localities in the vicinity of area shows a well marked, wide ventral furrow. Brno: Karpatian (= Upper Burdigalian, Nannoplankton Zone NN4): see B rzobohaty (1965, 1967a); Lower Badenian R em arks: Ontogenetic change is considerable in otoliths (= Langhian, Nannoplankton Zone NN5): see B rzobohaty o fD. acutirostrum and in addition to his original descrip­ (1982, 1986, 1997), B rzobohaty & Schultz (1978) tion, H o lec (1975) also cited otoliths of the same taxon NORTH SEA BASIN under two other names. Juveniles were attributed to what In the North Sea Basin, identifiable Diaphus otoliths are avail­ is now called Notoscopelus mediterraneus (K o k e n , 1 8 9 1 ) able from Miocene deposits only. In Belgium, material was (see iconography in B r z o b o h a t y & N o l f , 1996, pi. 8, collected from the Edegem Sands (? Early Hemmoorian), An­ fig. 1-8) and medium sized specimens were attributed to a twerpen Sands and the Zonderschot Sands (Late Hemmoorian), taxon originally described by P r o c h a z k a (1893) as O to­ see N olf, 1977; H uyghebaert & N olf, 1979; N olf & S mith, lithus (Berycidarum) splendidus. The holotype of the 1983. Additional Late Hemmorian material was mainly avail­ latter taxon could not be found, and as far as can be able from Miste, near Winterswijk, The Netherlands (see Jans ­ judged from the iconography in P r o c h a z k a (1893, pi. 3, sen , 1984, for locality data). Our Reinbekian material is from fig. 5), is a strongly eroded otolith. One can only guess the Aalten Member near Winterswijk (see V an den B osch et al., 1975) and from the Dingdener Feinsand at Dingden, NW that this holotype could belong to the same species, which Germany (see Ja nssen , 1967). Because of the neritic palaeoen- in that case should be named Diaphus splendidus (P r o­ vironments of all these deposits, Diaphus otoliths are rather c h a z k a , 1893). Such a guess however, should have the scarce and never dominant in the associations. Langenfeldian very nasty result of relegating the well established Recent deposits from Gross Pampau, east of Hamburg (seeH oede- fish species Diaphus splendidus (B r a u e r , 1904) as a makers , 1997) provided an association with abundant otoliths junior homonym ofP r o c h a z k a ’s species. Prochazka’s of Diaphus debilis. Post-Langenfeldian and Pliocene deposits taxon is considered here as a rejected species, because the of the North Sea Basin only provided some scarceDiaphus drawing of the holotype is not diagnostic at the species or otoliths, not identifiable at species level. genus level.

Distribution: D. acutirostrum is a Paratethys species Systematics (Karpatian till Middle Badenian). It is common in Lower Badenian of Moravia. In the Mediterranean sensu stricto, Diaphus acutirostrum (H o l e c , 1975) some scarce juvenile otoliths from the Burdigalian of PI. 3, Figs. 1-6 Sciolze may belong to D. acutirostrum.

? 1905 Otolithus (Berycidarum) mediterraneus Kok. - Schu ­ bert, p. 632, pi. 17, fig. 20 (non 19, non Koken , 1891) Diaphus hefralai n. sp. 1965 Myctophum? splendidum (P rochazka , 1893) - B rzo­ bohaty , p. 112, pi. 1, figs. ? 7, 8a,b (non Prochazka , PI. 6, Figs. 6-11 1893); 1967a Myctophum? splendidum (P rochazka , 1893) - B rzo­ 1986 Aethoprora sp. - B edini , Francalacci & L andini , p. 24, bohaty , p. 234, pi. ID, fig. 6a,b (non Prochazka , pi. 3, figs. 5-6, pi. 4, fig. 6. 1893); 1975 Myctophum mediterraneum Koken , 1891) - H olec , Type material: Holotype: a right otolith (PI. 6, Fig. 11) p. 255, pi. 1, fig. 2; (ÍRSNB P 7407); two paratypes (PI. 6, Figs 9-10) (IRSNB 1975 Myctophum splendidum (P rochazka , 1893) - H olec , P 7405 - P 7406). p. 256, pi. 1, fig. 4a, b (non Prochazka , 1893); 1975 Otol. (Myctophidarum) acutirostrum n. sp. - Holec , Additional material: Madonna della Neve (Serravallian): p. 258, pi. 1, fig. 5a,b; 16 specimens; Alba (Tortonian): 3 specimens; Borelli 1981 Diaphus splendidus (PROCH.) - R ado , pi. l,fig.9a,b, (? Messinian): 4 specimens. pi. 8, fig. 10a,b (non Prochazka , 1893); 1983 Diaphus splendidus (P rochazka ) - B rzobohaty , pi. 4, fig. 1 (non Prochazka , 1893); Dimensions of the holotype: Length: 2.8 mm, height: ? 1998 Diaphus sp. - R eichenbacher , p. 326, pi. 3, fig. 9. 2.4 mm; thickness: 0.7 mm. 188 Rostislav BRZOBOHATY & Dirk NOLF

Stratum typicum'. Tortonian marls, lower N 17 Planktonic 1979 Diaphus cahuzaci n. sp. - Steurbaut , p. 61, pi- 4, figs. I Foraminiferal Zone, E of St. Agata Fossili - 6, pi. 12, fig. 11; 1979 Myctophum pulchrum (Prochazka ) 1893 - Jonet, p. 107, pi. 3, fig. 2 (n o n Procha/ka 1893); Derivado nominis : Acronym of the authors H e d i n i , 1982 Diaphus cahuzaci S teurbaut , 1979 - S teurbaut & F r a n c a l a c c i , L a n d i n i (1986), who first figured an oto­ Jonet , p. 197, pi. 1, fig. 12; lith of this taxon. 1983 Diaphus cahuzaci S teurbaut , 1979 - S teurbaut , p. 257, pi. 1, figs. 4-9; Diagnosis'. This species is characterised by otoliths with a 1983 Diaphus cahuzaci (S teurbaut , 1979) - N olf & S mith, nearly semicircular ventral rim that bears 6-8 (exception­ p. 90. pi. I. figs. 15-18; ally 10) coarse spines. The dorsal rim is high, but narrow; 1984 Diaphus cahuzaci Steurbaut, 1979 - Steurbaut, p. 52, its anterior part is curved, but does not bears a true pi. 8. figs. 15-17: predorsal angle. There is a posterodorsal angle that lies 1992 Diaphus cahuzaci S teurbaut , 1979 - R adw ansk a , close to the center of the dorsal rim. Behind the poster­ p. 178, pi. 4, figs. 4-6, textfig. 26; 1997 Diaphus cahuzaci Steurbaut , 1979 - N olf & B rzobo­ odorsal angle, the dorsal rim is truncate, very slightly haty , pi. 1, fig. 19. concave. The otoliths are also characterised by a strongly incised excissura. The outer face is smooth and somewhat H olo type: Figured by S t e u r b a u t , 1979, pi. 4, fig. 1 and more convex as the inner face. The sulcus is narrow, and pi. 12, fig. 11; here refigured at PI. 5, Fig. 6 (IRSNB its crista superior is marked by a shallow depression in the P 3035); Langhian interval of the Saubrigues Marls at nearly flat dorsal area. The ventral area shows some more Saubrigues, Jean Tic (Aquitaine). convexity and bears a well marked furrow in the middle part. Remarks and description: This small species was origin­ ally described from juvenile otoliths, but seems to be Remarks'. These otoliths apparently belong to the same morphologically well characterised and is abundant in species as the one preserved in a skeleton from the Lower the Lower and Middle Miocene of Europe. On PI. 5, Messinian of Mondaino and figured by B e d i n i et al. Fig. 1-6, we figure an ontogenetic series, including oto­ (1986) as A ethoprora sp. Aethroprora G o o d e & B e a n , liths about twice as long as the holotype. This species is 1896 is a junior synonym ofD iaphus, and its type species characterised by nearly round otoliths with a strong and is D. metopoclampus Cocco, 1829, a common North- acuminate posterodorsal angle, and a more rounded ante- Eastern Atlantic species that also occurs in the Western rodorsal angle which in some specimens reaches higher Mediterranean. Surprisingly, this common species has no than the posterodorsal one. The rostrum is slightly salient. fossil record. Otoliths of D.metopoclampus and D. m e- The ventral rim is nearly semicircular, and bears 7-12 topoclampoides are higher than those ofD. befralai, their little spines. The outer face is convex, with a nearly flat dorsal rim is clearly concave behind the posterodorsal central portion in large specimens. The inner face is angle and their ventral spines are less developed. Some­ slightly convex in both antero-posterior and dorso-ventral what similar looking otoliths are also found inD. holti, sense. but these are more elongate, have a more salient rostrum A somewhat similar looking small specimen from the and a more posteriorly situated posterodorsal angle. Zanclean of Le-Puget-sur-Argens, figured by S c h w a r z -

h a n s (1986, pi. 3, fig. 32) as D. cahuzaci must be con­ Distribution: This is an uncommon species in the Medi­ sidered as a non-identifiable juvenile otolith of another terranean Serravallian, Tortonian and Messinian. species; we do not have any record ofD. cahuzaci am ong the many thousands of otoliths available from that local­ Diaphus cahuzaci S t e u r b a u t , 1979 ity. PI. 5, Figs. 1-6 Distribution: Northern Italy: Aquitanian (Antognola 1965 Scopelus pulcher Prochaska - R ado , p. 60, pi. 1, Marls at Moleto), Burdigalian (Termo Fora Complex fig. 5a,b (non fig. 3a,b; non Prochazka , 1893); and Pietra da Cantone), Langhian (Tanaro Formation), 1965 Myctophum excavatum (S ulc , 1932) - B rzobohaty , Serravallian (Madonna della Neve locality); Paratethys: p. 111, pi- 1, fig- 10 (non S ulc 1932); Ottnangian, Karpatian and Lower Badenian; Portugal: 1966 Myctophum excavatum (S ulc ) - S migielska , p. 234, Middle Miocene; Aquitaine: Aquitanian till Langhian pi. 13, figs. 4, 5,? 6 (non S ulc 1932); interval of the Saubrigues Marls; North Sea Basin: Hem­ 1967a Myctophum excavatum (S ulc , 1932) - B rzobohaty , p. 234, pi. ID, fig. 6a,b. (non S ulc , 1932); moorian (Edegem Sands). 1973 Myctophum debile (K oken ), 1891 - Jonet , p. 137, pi. 2, figs. 9, 10 (non 8, non Koken 1891); 1973 Myctophum pulchrum (P rochazka ), 1893 - Jonet , Diaphus cavallonis n. sp. p. 139, pi. 2, figs. 11-13 (non Prochazka , 1893); 1973 Diaphus cf. excavatus (S ulc ) - B rzobohaty & PI. 5, Fig. 7-14 S chultz , pi. 5, fig. 6 (non S ulc , 1932); 1979 Diaphus? aff. metopoclampus (Cocco, 1829)- S t e u r ­ 1976 Myctophum tuberculatum (Bassoli) - Anfossi & Mos- baut , p. 62, pi. 5, fig. 8; n a , p . 21, pi. 2, fig. 3a,b (non B a s s o l i , 1906); European Neogene Diaphus 189

1979 Diaphus splendidus (P rochazka 1893) - Schwarz - Messinian skeleton-based D iaphus species with unknown hans , p. 8, pi. 3, figs. 17, 18 (non 16, non Prochazka , or only poorly preserved otoliths in situ (see introduc­ 1893); tion). However, because the above described type of 1986 Diaphus sulcatus (B assoli 1906) s. 1. - Schw arzhans , otolith is very common in Mediterranean Pliocene depos­ p. 223, pi. 4, figs. 47-49 (non B assoli , 1906); its, we judged that a formal otolith-based name is by far 1986 Diaphus aff. rossiae (R o b b a , 1970) - S c h w a r z h a n s , more preferable to a forced guess for one of the skeleton- p. 223, pi. 4, flg. 46 (non R o b b a , 1970); non pi. 4. flg. 43 and 45 (= D. aff. splendidus); based taxa, or to further designation as “ sp. 1” . A some­ 1989 Diaphus sp. 1 - N o l f & C a p p e t t a , p. 218, pi. 10, what similar looking otolith was figured byB e d i n i et al. figs. 11 - 16; (1980, pi. 1, fig. 1) from a skeleton that they attributed to 1994 Diaphus sp. 1 - N olf & C avallo, pi. 3, figs 1, 2; Ceratoscopelus dorsalis (S a u v a g e , 1870). This otolith 1998 Diaphus sp. 1 - N olf, M a n e & Lopez, pi. 2, fig. 11. apparently belongs to a D iaphus, but does not show the anterior narrowing which is typical forD. cavallonis. Type material: Holotype: a left otolith (PI. 5, Fig. 7) (IRSNB P 7389); more than 500 paratypes, of which three Distribution: Otoliths of D. cavallonis are rare in the are figured (PI. 5, Figs. 8-10) (IRSNB P 7390 - P 7392). Serravallian and Tortonian. They become more common in the ? Messinian of Borelli and especially in the Plio­ Additional material'. Madonna della Neve (Serravallian, cene of the Mediterranean realm. not common), St. Agatha Fossili (Tortonian, not com­ mon), Borelli (? Messinian, common), Mediterranean Pliocene: common at many localities (e.g. Papiol, Mon- Diaphus debilis (K o k e n , 1891) ticello, Rio Torsero, Vrica, Bussano La Cava, Bussano PI. 2, Figs. 13-18 Cusinaire, Vezza d’Alba). 1891 Otolithus (Berycidarum) debilis Koken - Ko­ ken , p. 122, pi. 6, fig. 3, 3a; Dimensions o f the holotype: Length: 5.0 mm, height: 1905 Otolithus (Berycidarum) austriacus Koken - 3.9 mm; thickness: 1.1 mm. S chubert , p. 630, pi. 17, fig. l(non 2-7, non Koken , 1891); Stratum typicum: Zanclean marls, Foraminiferal Zone M 1905 Otolithus (Berycidarum) Kokeni Proch . - S chu ­ PI. 2, at Le-Puget-sur-Argens. bert, p. 631, pi. 17, fig. 8, (non 9, 10, 11, non Prochazka 1893); Derivado nominis: This species is named after Oreste ? 1942 Scopelus debilis Kok. - W eiler, p. 21, pi. 1, Cavallo (Alba, Italy) in honour of his many contributions figs. 33, 37 (non 30-32, 34-36); to the knowledge of the palaeontology of Piemonte. 1966 Myctophum debile (K oken ) - S migielska , p. 229, pi. 12, figs. 6, 7 (non 8 = juv. Ex., non D iagnosis: This species is characterised by relatively pi. 13, fig. 1 = D. aff. taaningi); 1967a Ot. (Myctophidarum) kokeni (P rochazka , high, irregular elliptical otoliths with a somewhat narrow­ 1893) - B rzobohaty , p. 235, pi. ID, fig. 4a,b ing anterior portion. The anterior part of the ventral rim is (non Prochazka , 1893); straighter than the central and posterior zone, and 1969 Myctophum debile (K oken , 1891) - H einrich , stretches obliquely towards the rostrum. Between the p. 16, pi. 2, fig. 3 (? non pi. 3, fig. 1, pi. 17, rostrum and the posterodorsal angle, the dorsal rim shows figs. 1-12); a regular curving in small and medium sized specimens, 1971 Myctophum debile (K oken , 1891) - R ado , but has an obtuse angular expansion in large otoliths. The p. 184, pi. 2, fig. 7a,b, pi. 9, fig. 68 (? non posterodorsal angle is blunt but well marked, and accen­ pi. 1, fig. 5, pi. 9, fig. 67); tuated by a little hollowing just behind, towards the 1979 Diaphus debilis (K oken , 1891) - Huygebaert & N olf, p. 70; posterior rim. In small specimens, the posterior rim is 19 81 Diaphus austriacus (S chub .) - R a d o , pi. 6, fig. 4 regularly rounded, but in larger ones, it is more oblique (non Koken , 1891); and runs straight towards a posteroventral angle, which 1984 Otolith - PFEIL, textfig. 2; marks the transition with the ventral rim. The ventral rim 1986 Diaphus debilis (K oken 1891 ) - S chw arzhans , usually bears more than ten obtuse spines. pi. 3, fig. 28-29; The outer face has a smooth surface, and is convex in 1990 Diaphus debilis (K oken 1891 ) - M üller, p. 440, the dorso-ventral sense. In a ventral profile, the otoliths pi. 4, figs. 2, 6,?3-4,? 5; are thickest near to the posterior rim. The inner face is 1992 Diaphus debilis (K oken , 1891) - R adw ansk a , much less convex. The ostial coniculum often narrows p. 178, pi. 5. fig. 1-2 (non figs. 3-6 = D. kokeni), anteriorly. textfig. 27 a-b (non 27 c-f = D. kokeni); 1992 Diaphus sp. 2 - R adw ansk a , p. 181, pi. 5, fig. 10, 11, (non 7-9 = D. aff. cahuzaci, Rem arks: Although otoliths of this species do not imme­ textfig. 29 b, c, (non a = D. aff. taaningi); diately attract one’s attention, they are morphologically 1992 Diaphus sp. 3 - R adw ansk a , p. 181, pi. 6, figs. 7, well defined and distinct from those of other D iaphus 8,? 9, textfig. 30 a, b,?c; species. The taxon was previously cited as D iaphus sp. 1 partim 1994 Diaphus debilis (K oken 1891) - B rzobohaty , in several papers (see synonymy) because there are five pi. 2, figs. 6, 7, (non 5, 8 = D. taaningi); 190 Rostislav BRZOBOHATY & Dirk NOLF

1997 Diaphus debilis (K o k e n , 1891)- H o e d e m a k e r s , species, and is compared to various other specimens from p. 53, pi. 3, Figs. 1-6. the Lower Miocene of Northern Italy and Aquitaine on PI. 1. H olotype : Figured by K o k e n , 1891, pi. 6, fig. 3; from the “ Miocene of Langenfelde” . The original material of Description'. The presently figured material was chosen K o k e n seems to be lost, but H o e d e m a k e r s (1997, p. 52) in a series of otoliths with a length ranging from 1.5 - 5.6 recorded 1600 specimens from the Langenfeldian of mm, and illustrates clearly their variability and onto­ Gross Pampau, near Hamburg, where no otherD iaphus genetic changes, while the original description was only species occur. Six specimens from Gross Pampau are based on adult specimens. Otoliths of this species grow figured here. fairly large, and are characterised by their elongate shape, with a narrowing anterior portion. The post­ Description: The otoliths are slightly elongate, nearly dorsal angle is well developed and looks somewhat round, (length/height proportion varies between 1.1 and spiny. The ostial coniculum becomes narrower in its 1.2). They are characterised by an obtuse but well marked anterior portion, especially in larger specimens. The posterodorsal angle and a strong antirostrum. Most speci­ inner face is slightly convex, both in the antero-posterior mens show only a very obtuse anterodorsal angle, so that and in the dorso-ventral sense. The outer side is clearly the dorsal area narrows from the posterodorsal angle convex, with maximal thickness situated in the posterior towards the antirostrum. The rostrum is more prominent part. The ventral rim generally bears 7 to 9 irregular than the antirostrum, and both are separated by a sharply spines, exceptionally 10, and sometimes only 6 in juve­ incised excissura. The ventral rim is regularly rounded niles. but not semicircular, and bears 6 to 8 irregularly disposed obtuse spines. The dorsal area shows a well marked Remarks'. Otoliths ofD. haereticus show most similarity depression just above the sulcus, and most specimens to those of D.m arw icki (F r o s t , 1933) from the Middle show a rather wide ventral furrow. The outer face is and Upper Miocene of New Zealand (see S c h w a r z h a n s , smooth and convex; the inner face is nearly flat. 1980, figs. 167-169), but in this species, the junction of the dorsal and posterior rim is more angular, and the Remarks'. Otoliths of this species are nearest to those of posterior rim extends higher in dorsal direction. Diaphus kokeni and D. taaningi, and otolith series of the three species are figured together on PI. 2, for direct Distribution: Northern Italy: Aquitanian (Antognola comparison. Otoliths ofD. debilis differs from those of Marls at Moleto) Burdigalian (Termo Fora Complex, D. taaningi by their narrow antero-dorsal area and more Pietra da Cantone); Paratethys: Lower Badenian of Mor­ straight posterodorsal rim, while otoliths ofD. kokeni avia; Aquitaine: NN 2 - N N 3 interval o f the Saubrigues show a very strong posterodorsal angle, and a nearly M arls. vertical posterior rim.

Distribution: Paratethys: From the Eggenburgian till the Diaphus holti T a a n i n g , 1918 Badenian; In Northern Italy, some juvenile specimens PI. 3, Figs. 7-10 from the Serravallian can only tentatively be referred to this species; Aquitaine: Langhian interval of the Saubri­ 1971 Diaphus praerafinesquii n. sp. - W eiler, p. 11, pi- 1, gues Marls; North Sea Basin: from the Hemmoorian till fig. 15; the Langenfeldian; very common in the Langenfeldian. 1976 Diaphus theta E igenmann & E igenmann - Anfossi & M o sn a , p. 18, pi. 1, fig. 4ab (non Eigenmann & Eigen ­ m ann , 1890); 1986 Diaphus holti T aaning 1918 - Schw arzhans , p. 222, Diaphus haereticus (Brzobohaty & Schijltz, 1978) pi. 4, fig. 40-41; PI. 1. F ig s. 1-14 1989 Diaphus holti T aaning , 1918 - N olf & Cappetta , pi. 8, figs. 22-24; aaning olf ane opez 1978 Symbolophorus haereticus n. sp. - B r z o b o h a t y & 1997 Diaphus holti T , 1918 - N , M & L , S c h u l t z , p. 450, pi. 4, fig. IO, pi. 5, fig. 1; pi. 3, figs. 2-4. 1983 Symbolophorus haereticus B rzobohaty & Schultz - B rzobohaty , pi. 4, fig. 2. L ecto typ e: A Recent fish from the Mediterranean, 36°53’N, 3°9’E; ZMUC P2339204. Otoliths from Recent Holotype'. Figured by B r z o b o h a t y & S c h u l t z , 19 7 8 , specimens of D. holti are illustrated by N o l f & M a r t i - pi. 5, fig. 1; Coli. Katedra Geologie a Paleontologie, n e l l , 1980, pi. 2, figs. 15-20 and N o l f & C a p p e t t a , Masarykovy University Brno, inv. nr. 0251, re-figured 1989, pi. 8, figs. 22-24. here on PI. 1, Fig. 8; Badenian of Zabcice, Moravia. This specimen is a somewhat eroded and very short otolith that Remarks'. The otoliths of this species can only be con­ unfortunately, is marginal in the variability of the species; fused with those of another Mediterranean D iaphus spe­ the paratype MUB 0250, refigured here (PI. 1, fig. 9 ) cies, D. rafinesquei, but in the latter species, otoliths are represents much better the usual aspect of otoliths in this more elongate. Growth series of Recent otoliths, for European Neogene Diaphus 191

direct comparison, are figured by N o l f & M a r t i n e l l Diaphus metopoclampoides S t e u r b a u t , 1983 (1980, pi. 2). PI. 4, Figs. 7-14

Distribution'. Fossil otoliths of this present day species are 1979 “genus Myctophidarum” sp. S- teurbaut , p. 63, pi. 5, only known from the Mediterranean (Serravallian, Tor­ fig. l í ; tonian, ? Messinian of Borelli and Pliocene), but they are 1981 Notoscopelus sp. - R ado , pi. 8, fig. 2; never common in any of the associations. 1983 Diaphus metopoclampoides n.sp. - S teurbaut , p. 257, pi. 1, figs. 22-28; 1984 Diaphus metopoclampoides Steurbaut 1983 - S teur­ baut , p. 53, pi. 7, fig. 23.

Diaphus kokeni (P r o c h a z k a , 1893) PI. 2, Figs. 1-6 H olotype : Figured by S t e u r b a u t , 1979, pi. 5, fig. 22, and refigured here PI. 4, Fig. 7; IRSNB P 3704; from the

1893 Otolithus (Berycidarum) Kokeni nov. spec. - Prochaz ­ Tanaro Formation, Langhian of Marrone, Piemonte. ka , p. 81, pi. 3, fig. 3a, b; 1905 Otolithus (Berycidarum) Kokeni Proch . - S chubert , Rem arks: These high otoliths are relatively common in p. 631, pi. 17, figs. 9-11 (non 8); the Mediterranean Miocene and can be distinguished 1958 Scopelus debilis (K oken ) - W eiler, p. 327, pi. 1, fig. 10 from otherD iaphus species by their relatively long cauda, (non fig. 9 = D. taaningi, non Koken , 1891); which somewhat evocates the genusBenthosema. H ow ­ 1966 Myctophum kokeni (P rochazka ) - S migielska , p. 232, ever, the ventral rim with its regular ornamentation of pi. 13, fig. 2a, b; small obtuse spines, and the general aspect of the large 1967b Ot. (Myctophidarum ) kokeni kokeni (P roch .) - B rzo­ specimens (till 4 mm), makes this doubtless a D iaphus bohaty , p. 130. pi. 2, fig. 11; 1977 Diaphus debilis (K oken , E., 1891) - N olf, p. 18, pi. 2, species. Among Recent species, otoliths ofD. brachyce­ figs. 11, 12, 15, 16,? 13,? 14,? 18, (non 17, non Koken , p h a lu s T a a n i n g , 1928 (see N o l f & S t r i n g e r , 1992, 1891); pi. 10, figs 4-9) and D. metopoclampus (Cocco, 1829) 1983 Diaphus kokeni (P rochazka ) - B rzobohaty , pi. 6, (see N o l f & M a r t i n e l l , 1980, pi. 2, figs. 9-14) are fig. 3; closest to D. metopoclampoides, but have a markedly 1992 Diaphus debilis (K oken , 1891) - R a dw ansk a , p. 178, more prominent rostrum. pi. 5, figs. 3-6 (non 1-2 = D. debilis), textfig. 27 c, d, e, f (non a-b = D debilis), (non Koken , 1891). Distribution : Northern Italy: Upper Burdigalian (Termo Fora Complex), Langhian, Serravallian, Tortonian, ? H olotype: Figured by P r o c h a z k a , 1993, pi. 3, fig. 3; Messinian of Borelli; Paratethys: Lower Badenian; Aqui­ from the Badenian of Zidlochovice, Moravia. The origi­ taine: Upper Burdigalian (Zone NN 3 - NN 4) and Lan­ nal specimen seems to be lost, but otoliths of this species ghian interval of the Saubrigues Marls. are so characteristic that they can easily be judged from Prochazka’s figure, which fits perfectly in our series from the Badenian of Moravia (PI. 2 , Figs 1-6). Diaphus pedemontanus (R o b b a , 1970) Description'. These otoliths are characterised by the fol­ PI. 4, Figs 1-6 lowing features: almost quadrangular outline, with a 1970 Porichthys pedemontanus sp. n. - R o b b a , p. 151, nearly vertically oriented posterior rim; very well devel- pi. 16, fig. 8. opped, often spine-like posterodorsal angle, which is 1979 Diaphus pulcher (P r o c h a z k a 1893) - S c h w a r z h a n s , located at the highest point of the dorsal rim; moderately p. 9, pi. 2, fig. 25 (non figs. 22-24, non P r o c h a z k a curved ventral rim, bearing 5 to 7 (exceptionally 8) strong 1893). spines; crista superior salient in the anterior part, but 1979 Diaphus sp. - S c h w a r z h a n s , p. 10, pi. 2, fig. 26, sometimes becoming obsolete posteriorly; ventral rim pi. 11, fig. 128: clear and well delimited. 1980 “genus Neobythitinorum” pedemontanus( R o b b a , The length of the rostrum and antirostrum varies con­ 1970) - N olf. p. 137; siderably during ontogenesis: those structures become 1983 Diaphus pedemontanus (R o b b a , 1970) - N o l f & much more pronounced in large specimens, while otoliths S t e u r b a u t , p. 154, pi. 1, figs. 26-37; 1983 Diaphus sp. 1 - N o l f & S t e u r b a u t , p. 156, pi. 1, of juveniles are more rounded, and often difficult to fig. 25; separate from D. debilis. ? 1983 Diaphus pedemontanus (R o b b a , 1970) - S t e u r b a u t , p. 257, pi. 1, figs. 13-19; R em arks: Otoliths of this species are nearest to those of 1994 Diaphus sp. 2 - N o l f & C a v a l l o , pi. 3, figs. 6,? 5; Diaphus debilis and D. taaningi ; see under D. debilis for 1998 Diaphus sp. 2 - N o l f , M a n e & L o p e z , pi. 2, fig. 1 distinctive features. (non fig. 2 = abnormal specimen?).

Distribution-. Paratethys: from Karpatian till the Middle H olotype: Figured by R o b b a , 1970, pi. 16, fig. 8, IPUM Badenian. North Sea Basin: Hemmoorian (Zonderschot PE 385, from his locality 7 in the Tortonian of the Rio Sands); Reinbekian (Dingdener Feinsand). Mazzapiedi-Castellania. 192 Rostislav BRZOBOHATY & Dirk NOLF

Remarks'. This species grows to a very large size. The H olo t\pe\ A Recent fish from off New Caledonia, figured holotype otolith attains a length of 6.9 mm; the specimen by T a a n i n g , 1932, fig. 12 (ZMUC P 2329213). A series figured here at PI. 4, Fig. 1 attains 7.3 mm, and must come of Recent otoliths is figured byN o l f & A g u i l e r a , 1998, from a fish approximately 20 cm in length (a D. adeno­ pi. 5, figs. 1-6. m us otolith of 10.5 mm, the largest D iaphus otolith among our Recent comparative material comes from a Remarks'. The present day distribution of D. regani is fish of 26.5 cm total length). The growth series shown at restricted to the Indo-Pacific realm (W i s n e r , 1976, N a f - PI. 4, Figs. 1-6 shows strong ontogenetic changes, and in p a k t i t i s , 1978), but in the Lower Miocene, it frequently very large otoliths, the sulcus becomes deeply incised, the occurs in Northern Italy, and has also been recorded from antirostrum is strongly developed and prominent, and Aquitaine, and from Venezuela (N o l f & A g u i l e r a , there is a backwards expansion in the lower part of the 1998). These Lower Miocene records must be interpreted posterior rim. These otoliths are probably from fishes that as a case of an ancient circumtropical Tethys distribution. changed their mesopelagic habitat into a benthic way of In the Paleocanyon of Saubrigues, Aquitaine, the first life, when they attained a very large size (which is also appearance of D. regani is a good marker for the begin­ the case in D. adenomus). Variability in these specimens ning of the Miocene, while the last occurrence ofD. is also considerable. Many Zanclean otoliths, like those perspicillatoides (see B r z o b o h a t y & N o l f , 1996, pi. 4, figured here at PI. 4, Figs. 2-6 have a more undulant figs. 17-23) marks the end of the Oligocene. At first dorsal rim than the average Tortonian specimens, figured glance, both species appear to be similar, but otoliths of by N o l f & S t e u r b a u t (1983, pi. 1, figs. 26-37). D. regani always have a strongly expanded anterodorsal area and a narrow posterodorsal area, while inD. p ersp i­ Distribution'. Northern Italy: Upper Burdigalian (Termo cillatoides the dorsal area is much more symmetrical. Fora Complex), Langhian (basis of the Complex of Bal- dissero), Serravallian, Tortonian, ? Messinian of Borelli; Distribution'. Otoliths of this Recent Indo-Pacific species Zanclean of SE France and Catalania. Some juvenile are relatively common in the Lower Miocene of Northern otoliths from the Lower Badenian of the Paratethys Italy: Aquitanian (Marls of Antognola); Burdigalian (Ter­ may also belong toD. pedemontanus, but unquestionable mo Fora Complex, Pietra da Cantone). There is no con­ records of this species are known from the Mediterranean vincing evidence for the presence of this species in realm only. younger deposits, although some juvenile otoliths from the Mediterranean Tortonian and the Pliocene looks much like those ofD. regani. Aquitaine: Aquitanian Diaphus rafinesquii (Cocco, 1838) (NN 1 - 2) and Lower Burdigalian (NN 2 - NN 3) interval PI. 3, Figs. 11-14 of the Saubrigues Marls.

1972 Myctophum splendidum (P r o c h a z k a , 1893) - A n f o s s i & M o s n a , p. 101, pi- 17, fig. 1 (non P r o c h a z k a , 1893); Diaphus aff. splendidus (B r a u e r , 1904) species complex 1985 Diaphus rafinesquei (Cocco, 1838) - Bossio,L a n d in ii , PI. 6, Figs. 1-5 M a z z e i , S a l v a t o r in i & V a r ó l a , p. 57, pi. 2, fig. 5; 1989 Diaphus rafinesquei (Cocco, 1838) -N o l f & C a p p e t ­ 1906 Otolithus (Berycidarum) pulcher Prochazka - B assoli , t a , pi. 10, fig. 21; p. 49, pi. 2, figs. 19-20 (non Prochazka ); 1994 Diaphus rafinesquei (Cocco, 1838) -N o l f & C a v a l l o , 1906 Otolithus (Berycidarum) sulcatus B a ss . - B assoli , p. 50, pi. 2, fig. 5. pi. 2, figs. 23-24; 1906 Otolithus (Berycidarum) tuberculatus B a ss . - B assoli , Holotype'. A Recent fish from off Messina, figured by p. 50, pi. 2, figs. 25-26; Cocco, 1838, pi. 7, fig. 7, asNyctophus rafinesquii', no 1970 Myctophum debile ( Koken , 1891 ) - R o bba , p. 104, pi. 8, type material preserved according to E s c h m e y e r (1998). fig. 2 (non fig. 1 = D. aff. holti, non Koken , 1891 ); A series of Recent otoliths is figured byN o l f & M a r t i ­ 1970 Myctophum tuberculatum (B assoli , 1906) - R o bba , n e l l , 1980, pi. 2, figs. 21-26. p .'109, pi. 8, figs. 8-9; 1970 Myctophum rossiae sp. n. - R o bba , p. 112, pi. 9, figs. 2- Remarks', see under Diaphus holti. 4; 1976 Otolithus (Myctophidarum) sp. 2 - Anfossi & Mo sna , p. 23, pi. 3, fig. 2a,b; Distribution'. L ike D. holti, this Recent species is also 1983 Diaphus sulcatus (B assoli , 1906) - N olf & S teur­ present in the Mediterranean realm since the Serravallian, baut , p. 156, pi. 2, figs. 8-15; but it is never common in any of the associations. 1986 Diaphus rossiae (R o bba , 1970) - S chw arzhans , p. 222, pi. 4, figs. 43, 45; 1986 Diaphus sulcatus (B assoli , 1906) -S chw arzhans , pi. 4, Diaphus repalii T a a n i n g , 1932 fig. 44 (non pi. 4, figs. 47-49 = D. cavallonis)', PI. 3, Figs. 15-20 1989 Diaphus sulcatus (B assoli , 1906) - N olf & Cappetta , p. 218, pi. 10, figs. 1-10; 1998 Diaphus aff. regani T a a n in g , 1932 - N o l f & A g u il e r a , 1994 Diaphus aff. splendidus (B rauer , 1904) - N olf & p. 239, pi. 5, figs. 7-12. Cavallo , pi. 2, figs. 8-11; European Neogene Diaphus 193

1998 Diaphus aff. splendidus (Brauer, 1904) - N olf, Mane figs. 5, 8 (non 6-7, non Koken , 1891); & Lopez, pi. 2, fig. 10. 1998 Diaphus sp. 2 - N olf, Mane & Lopez, pi. 2, figs. 3-8.

R em arks: In this species complex, we include all the H o lotyp e: A Recent fish from off French Congo, figured otoliths which show a reasonable similarity to Recent by Norman, 1930, fig. 30; BMNH 1930.1.12.835. A D. splendidus otoliths (see N o l f & C a p p e t t a , 19 8 9 , series of Recent otoliths is figured by S t e u r b a u t pi. 9 , figs. 9 -1 1 ). Such otoliths occur from the Serraval­ (1979, pi. 3, figs. 11-16). lian till the end of the Pliocene. Only some large otoliths from the Tortonian (e.g. PI. 6, Fig. 1) and some Pliocene R em arks: Relationships (or identity) of this Recent spe­ specimens from Monticello are almost identical to Recent cies w ith D. debilis and D. kokeni were suggested by material of similar size, but many other fossils have a less N o l f (1977) and S t e u r b a u t (1979), but the availability expanded posterodorsal angle (e.g. the Tortonian series of otoliths from a wide range of stratigraphically and figured by N o l f & S t e u r b a u t , 19 8 3 , pi. 2 , figs. 8 -1 5 and geographically distinct associations now reveals that the Zanclean population from Le-Puget-sur-Argens fig­ three distinct species are involved; see PI. 2 and com­ ured by N o l f & C a p p e t t a , 1 9 8 9 , pi. 10, figs. 1 -1 0 ), and ments under D. debilis for distinctive features. Moreover, many intermediate conditions exist. Up to a length of 3 the present revision also suggest thatD. taaningi is fairly mm, otoliths of this species complex readily fall in the common in the Mediterranean Pliocene, where it was category of elliptical otoliths with a very generalized previously cited as D iaphus sp. ind. by N o l f & C a p p e t t a morphology, non-diagnostic at species level. Another (1989) and D iaphus sp. 2 by N o l f et al. (1998), but the problem is that Recent otolith material from various average Pliocene specimens tend to be slightly more populations of this worldwide ranging species is not elongated than the recent material available to us. Such available, which makes it very difficult to judge the elongate morphology was also observed in specimens meaning of small differences between fossil populations. from the Lower Pliocene of Dar Bel Hamri, Atlantic M orocco. Distribution: Very common in the Mediterranean realm, from the Serravallian till the Upper Pliocene. Distribution: Mediterranean: Tortonian, no Messinian records; known from the Zanclean till the Gelasian; Paratethys: from the Karpatian till the Upper Badenian; Diaphus taaningi N o r m a n , 1930 Aquitaine:Langhian interval of the Saubrigues Marls. PI. 2, Figs. 7-12 North Sea Basin: Hemmoorian (Miste Bed, Winterswijk, The Netherlands; Reinbekian (Dingdener Feinsand, NW 1905 Otolithus (Berycidarum) austriacus K oken - S c h u b e r t , Germany; Stemerdinck Bed, Winterswijk, The Nether­ p. 630 (partim), pi. 17, fig. 6 (non 1-5, 7, non Koken lands). 1891); 1943 Scopelus debilis K o k e n 1891 - W e il e r , p. 90 (partim), pi. 1, fig. 8 (non 4-7, 9-13, non K o k e n , 1891); Otoliths erroneously referred to Diaphus 1949 Scopelus debilis K o k e n - W e il e r (partim), pi. 1, fig. 8 (non 4-7, pi. 2, figs. 9-13, non K o k e n , 1891); Bolinichthys italicus (A n f o s s i & M o s n a , 1971) 1950 Scopelus debilis (K o k e n 1891) - W e il e r , p. 211, pi. 1, T extfig. 1 fig. 2 (non pi. 9, figs. 66, 67, nonK o k e n , 1891); 1958 Scopelus debilis (K oken ) - W eiler, p. 327, pi. I, fig. 9 (non 10 = D. kokeni, non Koken , 1891); 1971 Diaphus italicus n. sp. - A n f o s s i & M o s n a , p. 141, 1965 Myctophum debile (K o k e n , 1891) - B r z o b o h a t y , pi. 2, figs. 1-3; p. 109, pi. 1, figs. 1, 6 (non 2-3, non K o k e n , 1891); 1976 Diaphus italicus Anfossi & M osna - Iaccarino & 1966 Myctophum debile (K o k e n ) - S m ig ie l s k a , p. 229 (par­ M osna , pi. 1, fig. 5; tim), pi. 13, fig. 1 (non pi. 12, figs. 6-8, non K o k e n , 1983 Diaphus italicus A n f o s s i & M o s n a , 1971 - N o l f & 1891); S t e u r b a u t , p. 154; 1967 Myctophum debile (K o k e n , 1891) - B r z o b o h a t y , ? 1986 Diaphus sp. 2 - B e d in i , F r a n c a l a c c i & L a n d in i , p. 232, pi. ID, fig. 3a,b (non K o k e n , 1891); p. 38, pi. 2, fig. 9. 1972 Myctophum debile (K o k e n ) - B a u z a , p. 63, p i., fig s . 18- 19,? 20-21 (non 22, 23, 25-31, non K o k e n , 1891); R em arks: Those little otoliths (length of the holotype 1978 Diaphus austriacus (K o k .) - B r z o b o h a t y , pi. 1, fig. 10 = 1.8 mm), which were considered originally to represent (non K o k e n , 1891); a sm all D iaphus species, seems to be much more closely 1979 Diaphus debilis( K o k e n , 1 8 9 1 )-S t e u r b a u t , p. 62, pi. 3, related to the Recent Bolinichthys supralateralis (P a r r , figs. 9-10 (non K o k e n , 1891); 1928), a tropical to subtropical Atlantic species, which is 1983 Diaphus debilis (K o k e n ) - B r z o b o h a t y , pi. 6, fig. 2 figured here for direct comparison (Fig. 1). The fossil (non K o k e n , 1891); otoliths differ from the Recent ones by their more quad­ 1984 Diaphus debilis (K o k e n 1891 ) - S t e u r b a u t , p. 53, pi. 7, figs. 24, 25 (non K o k e n , 1891); rangular outline, their less curved ventral rim, which 1985 Diaphus debilis (K o k e n 18 9 1 ) - N o l f , fig. 47G; bears stronger spines, even at small size, by their more 1989 Diaphus sp. ind. - N o l f & C a p p e t t a , pi. 10, figs. 17-20; salient and acuminate rostrum, and by their ostium that is 1994 Diaphus debilis (K o k e n , 1891) - B r z o b o h a t y , pi. 2, more opened anteriorly. 194 Rostislav BRZOBOHATY & Dirk NOLF

Distribution : Known by the type material (four otoliths) I mm from the Tortonian of Cascine Valle Sorda near Alessan­ dria, Italy and by specimens from Messinian deposits near Reggio.

Conclusions

The present revision documents the presence of at least

Bolinichthys supralateralis (PARR. 1928) fourteen otolith-based D iaphus species in the European Neogene. As already stated in the introduction, this num­ ber is minimal, and it is even probable that several species with generalized elliptical otolith morphology are hidden in the large amount of non-diagnostic material (many thousands of otoliths) that was left aside. All fourteen recognizable taxa occur in the Mediterranean and Para­ I mm tethys Basin (Fig. 2). On the Atlantic side (Figure 3), where identifiable D iaphus otoliths are known from the

Bolinichthys italicus (ANFOSSI & MOSNA. 1971) Lower and Middle Miocene only, five species are re­ corded from the Paleocanyon of Saubrigues, Aquitaine, Fig. 1 — Otoliths o f (a-b) Bolinichthys supralateralis (Recent, and only four from the North Sea Basin. offPortugal) and (c-d) Bolinichthys italicus (Tortonian It is evident that the main interest of the present study of Cascine Valle Sorda, c = holotype, d = paratype). concerns the Mediterranean Basin and its Paratethys de-

& 3 _) CL CENTRAL AGE O Ma II ü 2 PARATETHYS S ’S j z < < STAGES RECENT A C u z NN 21 QUATERNARY I NN 19 GELASIAN NN 18 PIACENZIAN NN 14 ZANCLEAN NN 13 C 5 - >. NN 12 2 MESSINIAN 1 NN II -5

TORTONIAN NN IO NN 9 10- NN 8 ..M NN 7 .5S? ^ S Q NN 6 SERRAVALLIAN 3 2

NN 5 BADENIAN 1 5 - 3 LANGHIAN S, C i

NN 4 KARPATIAN

OTTNANGIAN BURDIGALIAN M = M editerranean NN 3 EGGENBURGIAN 2 0 - p = Paratethys

= real occurence NN 2 AQUITANIAN = virtual occurence EGER1AN J = recent outside Mediterranean NN I A * recent in M editerranean CHATTIAN

Fig. 2 — Stratigraphie range of NeogeneDiaphus species in the Mediterranean realm (including the Paratethys) (extinct species which are only known by osteological remains excluded). Correlation of the international and Paratethys stratigraphie scales after R ögl, 1998. European Neogene Diaphus 195

D iaphus. Four Late Oligocene species,D. alcoholicus NORTH SEA B r z o b o h a t y & N o l f , 1995, D. longirostris (B r z o b o ­ STAGES h a t y , 1967), D. perspicillatoides B r z o b o h a t y & N o l f ,

1995, and D. pristismetallis N o l f & B r z o b o h a t y , 1994 SERRAVALLIAN LANGENFELDIAN do not cross the boundary, and both in the Mediterranean Basin and in Aquitaine the appearance of D. haereticus, D. cahuzaci and D. regani marks the beginning of the | A , I M iocene. BURDIGALIAN HEMMOORIAN Fig. 2 also suggests the simultaneous appearence of five taxa in the Upper Burdigalian, but this faunal reno­ A * Aquitaine N = North Sea vation may well be an artifact. There are almost no lower — * real occurence VIERLANDIAN — • = virtual occurence NN I Burdigalian deposits investigated for otoliths in the Med­ f recent outside North Atlantic iterranean Basin, and in the Paratethys, the interval from ’ recent in N onh Atlantic NP 25 neochattian the basal Miocene to basal Karpatian does not provide Fig. 3 — Stratigraphie range of NeogeneDiaphus species in very appropriate facies for collecting mesopelagic fish the Aquitaine and the North Sea Basin. Tentative otoliths. Moreover, the Upper Burdigalian first appear­ correlation of the international and North Sea Basin ances constitute a mixture of Mediterranean and Para­ stratigraphie scales after L a u r s e n et aí. (1998), tethys records, and some of the Paratethys species may L o u w y e & L a g a , 1998 and V a n d e n b e r g e et al., well be more northern, or pseudoceanic (= living near to 1998. continental margins, see N a f p a k t i t i s et al., 1977) spe­ cies. A pseudoceanic habitat is known for the Recent D. taaningi, which is one of the species recorded in the Paratethys. We also do not exclude that D. debilis and pendence. Five of the recorded species are still living D. kokeni , which otoliths are somewhat similar to those of today, but three of them, D. regani. D. taaningi and D. D. taaningi, had a pseudoceanic way of life. The presence splendidus, only occur outside the Mediterranean. The of these three species in the more shallow and isolated Recent Mediterranean D. holti and D. rafinesquii are Miocene North Sea Basin may even validate such a point represented since the Serravallian, but they are uncom­ o f view . mon in all the investigated associations. Finally, D. m e­ The first appearance of five species in the Mediterra- topoclampus, a Recent western Mediterranean inhabitant, nian Serravallian is an important renovation, especially has no fossil record. Pliocene associations are dominated because four of them continue into the Pliocene, while by the extinct D. cavallonis, and by D iaphus aff. sp len­ only two of the nine Lower Miocene species do so. didus, whose Recent relatives live outside the Mediterra­ Dating of the event, however, is less precise and it could nean. All this stresses that till the end of the Pliocene already have happened in the Langhian, for which we (only few data are available for the Pleistocene), the only have good otolith localities in the Paratethys, but not Mediterranean mesopelagic fauna was very different, in the Mediterranean Basin sensu stricto. and essentially more oceanic, than the present day one. In the North Sea Basin, the only relevant “ myctophid The same conclusion could already be drawn for the non- event” seems to be an important acme ofD. debilis, in myctophids (B r z o b o h a t y & N o l f , 1996) and a D iaphus Langenfeldian deposits. more oceanic character was also demonstrated for Oligo­

cene Mediterranean fish faunas (N o l f & S t e u r b a u t , 1988; N o l f & B r z o b o h a t y , 1994). The present day distribution of D. regani is restricted to the Indo-Pacific Acknowledgements realm (W i s n e r , 1976, N a f p a k t i t i s , 1978), but in the Lower Miocene, its presence in Northern Italy, Aquitaine, It is a pleasure for us to express our sincere gratitude to HP.o f fm a n , and Venezuela (N o l f & A g u i l e r a , 1998) must be inter­ who helped us in preparing the iconography of the present paper, and to preted as a case of ancient circum-tropical Tethys dis­ M . H o w e ll (University of South Carolina, Columbia) for the time he tribution. spent in discussing a first draft of our manuscript. We also are greatly indebted to Erica B icchi (University of Torino), B. H a m r sm id (M o r­ From the biostratigraphic point of view, one can state avian Oil and Gas Company, Hodonin) and to L. S v a b e n ic k a (C zech that the Oligo-Miocene boundary is well marked by Geological Survey, Prague) for their micropaleontological analyses. 196 Rostislav BRZOBOHATY & Dirk NOLF

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P o s t h u m u s , O ., 1923. Bijdrage tot de kennis der tertiaire V andenberghe , N ., L a g a , P., S t e u r b a u t , E., H a r d e n b o l , J. & vischfauna van Nederland. Verhandelingen van het Geolo- V a il , P.R., 1998. Tertiary sequence stratigraphy at the southern gisch-Mijnbouwkundig Genootschap voor Nederland en Kolo­ border of the North Sea Basin in Belgium. Society o f Economic nien, Geologische Serie, 7: 105-142. Paleontologists and Mineralogists, special publication, 60: 119-154.

Prochazka , V.J., 1893. Das Miozän von Seelowitz in Mähren V a n d e n B o s c h , M., C a d e e , M.C. & J a n s s e n , A.W., 1975. und dessen Fauna. Rozpravy Ceské Akademie Cisare Frantiska Lithostratigraphical and biostratigraphical subdivision of Ter­ Josefa pro Vedy Slovesnost a Umeni v Praze, 2 (2): 6588. tiary deposits (Oligocene - Pliocene) in the Winterswijk - R a do , G., 1965. Otolite din depozitele tortoniene de la costeiul Almelo region (eastern part of the Netherlands).Scripta Geo­ de Sus. Analele Universitatii Bucuresti, Serie Stiintele naturii, lógica, 29: 1-167.

Géologie Geografie , 14: 55-71. V i o l a n t i , D. & G ir a u d , V., 1992. Contributi allo studio del R ado , G., 1971. Otolite in depozitele miocene din forajele de la Neogene delle Langhe sud-occidentali (Mondovi). Rivista Itali­ Islaz. Buletinul Societatii de Stiinte Geologice din R.S. Roma­ ana di Paleontología e stratigrafia, 97 (3-4): 639-659. nia,, 13: 179-194. W e il e r , W., 1942., Die Otolithen des rheinischen und nord­ R ado , G., 1981. Otolite din depozitele langhiene (Badenian westdeutschen Tertiärs. Abhandlungen des Reichsamts fur Bod­ inferior) de la Lapugiul de Sus.Analele Universitatii Bucuresti, enforschung, 206: 1-140. Série Stiintele naturii, Géologie Geografie, 30: 29-49. W e il e r , W., 1943. Die Otolithen aus dem Jungtertiär Süd- R adw ansk a , U., 1992. Fish otoliths in the Middle Miocene Rumäniens. I. Buglow und Sarmat.Senckenbergiana Lethaea, (Badenian) deposits of southern Poland.Acta Geológica Polo­ 26 (1-3): 87-115. nica , 42 (34): 141-328. W e il e r , W., 1949. Die Otolithen aus dem Jungtertiär Süd- R eichenbacher , B., 1998. Fish-Otolithen aus dem Karpat des Rumäniens. Ergänzende Tafeln zu den Otolithen des Buglow Korneuburger Beckens. Beiträge zur Paläontologie von Öster­ und Sarmat. Senckenbergiana Lethaea, 30: 291-293. reich, 23: 352-345. W e il e r , W., 1950. Die Otolithen aus dem Jungtertiär Süd- R obba , E., 1970. Otoliti del Tortoniano-tipo (Piemonte).Riv- Rumäniens. 2. Mittel-Miozän, Torton, Buglow und Sarmat. ista Italiana di Paleontología, 76 (1): 89-172. Senckenbergiana Lethaea, 31: 209-258. Rögl, F., 1998. Palaeogeographic considerations for Mediter­ W e il e r , W., 1958. Fisch-Otolithen aus dem Ober-Oligozän und ranean and Paratethys seaways (Oligocene to Miocene).Annal­ dem Mittelmiozän der Niederrheinischen Bucht. Fortschritte in en des Naturhistorischen Museums in Wien. 99A: 279-310. der Geologie von Rheinland und Westfalen, 1: 323-361. Schubert , R.J., 1905. Die Fischotolithen des österr-ungar. Ter­ W e il e r , W., 1971. Fisch-Otolithen aus dem Jungtertiär Süd- tiärs. II. Macruriden und Bi#yciden. Jahrbuch der Kaiserlich- Siziliens. Senckenbergiana Lethaea, 52 (1): 5-37. Königlichen Geologischen Reichsanstalt, Wien, 55: 613-638. W is n e r , R.L., 1976. The taxonomy and distribution of Lantern- S c h w a r z h a n s , W., 1979. Otolithen aus dem Unter-Pliozän von fishes (Family Myctophidae) of the Eastern Pacific Ocean. Navy Süd-Sizilien und aus der Toscana. Berliner Geowissenschaf- Ocean Research and Development Activity Report, 3: 1-229. tliche Abhandlungen, A, 8: 1-52. Y o u n g , 1998., Neogene. In Brown, P.R. Calcareous Nannofos- Schw arzhans , W., 1980. Die tertiäre Teleosteer-Fauna Neu­ sil Biostratigraphy. Cambrige University Press, Cambridge, seelands, rekonstruiert anhand von Otolithen. Berliner geowis- pp. 225-265. senschafiliche Abhandlungen, A, 26: 1-211. Z il c h , A., 1965. Die Typen und Typoide des Natur-Museums Schwarzhans , W., 1986. Die Otolithen des Unter-Pliozän von Le Puget, S-Frankreich. Senckenbergiana lethaea, 67 (1-3): 219-273. Senckenberg, 31: Fossile Fisch-Otolithen. Senckenbergiana Lethaea, 46a: 453-490. S migielska , T., 1966: Otoliths of fishes from the Tortonian of Southern Poland. Annales de la Société géologique de Pologne, R. B r z o b o h a t y 36: 205-275. Institute of Geology and Paleontology S paak , P., 1983. Accuracy in correlation and ecological aspects Masaryk University of the planktonic foraminiferal zonation of the Mediterranean Kotlarska 2 Pliocene. Utrecht Micropaleontological Bulletins, 28: 1-159. 611 37 Brno S teurbaut , E., 1979. Les otolithes de Téléostéens des Marnes Czech Republic de Saubrigues (Miocène d’Aquitaine méridionale, France). Pa- (e-mail: [email protected]) laeontographica, A, 166: 50-91. D. N o l f S teurbaut , E., 1983: Les Otolithes de Téléostéens de la For­ Departement Paleontologie mation de Tanaro (Langhien inférieur du Piémont, Italie sep­ Koninklijk Belgisch Instituut tentrionale). Geológica et Paleontológica, 17: 255-263. voor Natuurwetenschappen Steurbaut , E., 1984: Les otolithes de téléostéens de l’Oligo- Vautierstraat 29 Miocène d’Aquitaine (Sud-Ouest de la France ). Palaeontogra- B-1000 Brussel phica A, 186 (1-6): 1-162. Belgium S teurbaut , E. & Jonet , S., 1982. Révision des otolithes de (e-mail: [email protected] ) téléostéens du Miocène portugais.Bulletin de la Société Belge de Géologie, 90 (1981), fasc. 3: 191-229. Typescript submitted 30 June 1999

S u l c , J., 1932. Preliminary report on the otoliths from the Corrected typescript received I December 1999 European Neogene Diaphus 199

Appendix 1: doubtful species and synonyms in the European Tertiary

D iaphus an n. sp. in S ulc (1932, p. 2, fig. 1) = not c h a z k a (1893, p, 57, pi. 3, fig. la,b) = doubtful species. identifiable juvenile specimen. Based on a not identifiable eroded juvenile specimen.

O. austriacus in K o k e n (1891, p. 122, fig. 14, p. 123) = not Scopelus obliquus n. sp. in W eiler (1943, p. 91, pi. 1, interpretable drawing; lectotype SMF 2410 indicated and figs. 20, 21 ) = doubtful species. Based on not identifiable figured by Zilch (1965, pi. 37, fig. 1) is a not identifiable juvenile otoliths. juvenile specimen. Diaphus poignantae n. sp. in S t e u r b a u t (1979, p. 62, Diaphus crassus n. sp. in S c h w a r z h a n s (1979, p. 10, pi. 5, figs. 1-6, pi. 12, fig. 4) = doubtful species. Based on pi. 2, figs. 19-21, pi. 11, fig. 126-127) = probably aberrant not identifiable juvenile otoliths. otoliths of D.holti T a a n in g , 1918. Diaphus praerafimesquii n. sp. in W eiler (1971, p. 11, Diaphus dirknolfi n. sp. in S c h w a r z h a n s (1986, p. 231, pi. 1, fig. 15) = D. holti T a a n in g , 1918 (see synonymy). pi. 3, figs. 34-35) = doubtful species. Based on type O tolithus (Berycidarum) p u lc h er nov. spec, in P r o c h a z ­ material without diagnostic features. The holotype SMF k a ( 1893, p. 58, pi. 3, fig. 17a,b) = not identifiable, eroded P 7225 looks very similar to a Recent specimen of specimen, nomen dubium. Diaphus garmani G il b e r t, 1906 in the IRSNB collection, Myctophum regulare n. sp. in S m ig ie l sk a (1966, p. 236, another species with very generalized and poorly diag­ pi. 13, figs. 7, 8, Textfig. 2) = doubtful species. Based on nostic otolith morphology. Type material ofD. dirknolfi an eroded, non-diagnostic specimen. comes from Dar Bel Hamri, Atlantic Morocco, which is O tolithus (Berycidarum) splendidus nov. spec, in P ro­ outside the geographic area treated here, but the name is c h a z k a (1893, p. 59, pi. 3, fig. 5) = rejected species; mentionned because S c h w a r z h a n s (1986, p. 223) also based on an eroded, non-diagnostic specimen. See also attributed 11 non-diagnostic otoliths from Le-Puget-sur remarks under Diaphus acutirostrum (H o l e c , 1975). Argens to this taxon. Diaphus rossiae sp. n. in R o b b a (1970, p. 112, pi. 9, Otolithus (Berycidarum)insoletus nov. spec, in P r o c h a z ­ figs. 2-4) = D. aff. splendidus (B r a u e r , 1904) species k a (1893, p. 60, pi. 3, fig. 8a,b) = doubtful species, based com plex. on an eroded specimen, may be a Notoscopelus. Diaphus vohnhachti n. sp. in S c h w a r z h a n s ( 1986, p. 2 3 0 , Diaphus italicus n. sp . in An f o ssi & M o s n a ( 1 9 7 1 , p. 141, pi. 3, figs. 37-38) = Lobianchia aff. dofleini (Z u g m a y e r , pi. 2, figs. 1-3) = Bolinichthys italicus. 1911). Scopelus latirostratus n. sp. in W eiler (1 9 5 0 , p. 215, pi. 7, Otolithus (Berycidarum) sulcatus B a s s , in B a sso l i fig s. 44-45, pi. 9, fig. 68a-d) = doubtful species; based on (1906, p. 50, pi. 2, figs. 23-24) = D iaphus aff. splendidus not identifiable juvenile otoliths. (B rauf .r, 1904) species complex. Ot. (Berycidarum) latus in B a sso l i (1 9 0 9 , p. 41, pi. 2, O tolithus (Berycidarum) tuberculatus B a s s , in B a sso l i fig. 27) = doubtful species (seeN o lf & S t e u r b a u t 1983). (1906, p. 50, pi. 2, figs. 25-26) = Diaphus aff. splendidus O tolithus (Berycidarum) M oravicus nov. spec, in P ro­ (B r a u e r , 1904) species complex.

Explanation of the plates

All figured specimens are deposited in the collections of the Institut Royal des Sciences Naturelles de Belgique (IRSNB), with the exception of the holotype and paratype ofDiaphus haereticus, which belong to the collection of the Department of Geology and Paleontology of the Masaryk University in Brno (Czechia) (MUB). The abbreviations F and (F) in the upper right corner of each compartment of the plates indicate if the figured specimens in that compartment are fossil [F] or belong to Recent species known as fossils [(F)]. In the captions, L stands for left otolith and R for right otolith. The notations Fig. a, b and c are used to indicate respectively ventral and inner (=mesial) views. Figures with only numbers and no letter show inner views.

P l a t e 1

Figs. 1-14 — Diaphus haereticus (B r z o b o h a t y & S c h u l t z , 1978). 1-7 = R, Antognola Marls. Aquitanian. Moleto-Prera (IRSNB P 7328 - P7334), 8-9 = R, Langhian (Badenian), Zabcice, 8 = holotype (MUB 0251), 9 = paratype (MUB 0250), 10-12 = R, Pietra da Cantoni, Upper Burdigalian, Ponzano (IRSNB P 7335 - P 7337), 13-14 = Lower Burdigalian interval of the Saubrigues Marls, 13 = L. 14 = R (IRSNB P 7338, P 7339).

P late 2

Figs. 1-6 — Diaphus kokeni (P r o c h a z k a , 1893). Langhian (Badenian), Drnovice (IRSNB P 7340 - P 7345).

Figs. 7-12 — Diaphus taaningi N o r m a n , 1930. L, Langhian (Badenian), Brno, Kralovo Pole (IRSNB P 7346 - P 7351). Figs. 13-18 — Diaphus debilis (K o k e n , 1891 ). 13-15 = L, 16-18 = R, Glimmerton, Langenfeldian, Gross Pampau (IRSNB P 7032 - P 7037). 2 0 0 Rostislav BRZOBOHATY & Dirk NOLF

P late 3

Figs. 1-6 — Diaphus acutirostrum (H olec , 1975). 1 = Langhian (Badenian), Cema Hora (IRSNB P 7352), 2-6 = L, Langhian (Badenian), Drnovice (IRSNB P 7353 - P 7257). Figs. 7-10 — Diaphus holti T a a n in g , 1918. 7 = L, 8-10 = R, Tortonian, Sant Agata Fossili (IRSNB P 7358 - P 7361). Figs. 11-14 — Diaphus rafinesquii (Cocco, 1838). 11 = L, Pleistocene, Santernian, Vrica section, between sapropel e and f (IRSNB P 7362), 12 = L, 13-14 = R, Zanclean, Rio Torsero (IRSNB P 7363 - P 7365). Figs. 15-20 — Diaphus regani T a a n in g , 1932. 15-16 = L, 17-20 = R, 15-16 and 18-20 = Termo Fora Complex, Burdigalian, Sciolze (IRSNB P 7366 - P 7370), 17 = basement of the Baldissero Complex, Langhian, Baldissero, point 5 (IRSNB P 7371).

P l a t e 4

Figs. 1-6 — Diaphus pedemontanus (R obba , 1970). 1 = L, Tortonian, Rio Mazapiedi-Castellania (IRSNB P 3782), 2-6 = R, Zanclean, Le-Puget-sur-Argens (IRSNB P 7372 - P 7376). Figs. 7-14 — Diaphus metopoclampoides S t e u r b a u t , 1983. 7 = R, Holotype (IRSNB P 3704), Tanaro Formation, Lower Langhian, Marrone., 8-10 = R, 11-14 = L, basement of the Baldissero Complex, Langhian, Baldissero, point 5 (IRSNB P 7377 - P 7378).

Plate 5

Figs. 1-6 — Diaphus cahuzaci Steurbaut , 1979.1-5 = L, Antognola Marls, Aquitanian, Moleto-Prera (IRSNB P 7384 - P 7388), 6 = R, holotype (IRSNB P 3035), Langhian interval of the Saubrigues Marls. Figs. 7-14 — Diaphus cavallonis n. sp. L, 7 = holotype (IRSNB P 7389), 8-10 = paratypes. Zanclean, Le-Puget-sur-Argens (IRSNB P 7390 - P 7392), 11-14 = Serravallian, Madonna della Neve near Mondovi (IRSNB P 7393 - P 7396).

Plate 6

Figs. 1-5 — Diaphus aff. splendidus (B rauer , 1904) species complex. 1 = L, Tortonian, Sant Agata Fossili, Point c ofN olf & Steurbaut , 1983 (IRSNB P 7387), 2-5 = R, Torremondo Marls, Messinian, Venta de la Virgen (IRSNB P 7398 - P 7401). Figs. 6-11 — Diaphus befralai n. sp. L, 6-8 = Serravallian, Madonna della Neve near Mondovi (IRSNB P 7402 - P 7404), 9-11 = Tortonian, Sant Agata Fossili, Point c ofN olf & Steurbaut (1983), 9-10 = paratypes (IRSNB P 7405, P 7406), 11 = holotype (IRSNB P 7407). European Neogene Diaphus 201

8b mm

mm

Zabice, Langhian Ponzano, Burdigalian

mm

mm

Saint-Martin-de-Seignanx, iMoleto. Aquitanian Les Platanes". Early Burdigalian

Diaphus haereticus (BRZOBOHATY & SCHULTZ, 1978)

P late 1 2 0 2 Rostislav BRZOBOHATY & Dirk NOLF

(F)

7a 13a

13b 7b

1b

Q

O

Diaphus taaningi Diaphus debilis Diaphus kokeni (PROCHAZKA. 1893) NORMAN. 1930 (KOKEN. 1891)

P late 2 European Neogene Diaphus 2 0 3

Diaphus acutirostrum Diaphus holti (HOLEC. 1975) TAANING, 1918

Diaphus regani TAANING, 1932

Diaphus rafinesquii (COCCO, 1838)

P late 3 Rostislav BRZOBOHATY & Dirk NOLF

Diaphus pedemontanus (ROBBA. 1970) Diaphus metopoclampoides STEURBAUT, 1983

P late 4 European Neogene Diaphus 205

11a

7b 11b

1 mm 1 mm

6b •■■•.Y.-

Diaphus cahuzaci Diaphus cavallonis n. sp STEURBAUT, 1979

P late 5 206 Rostislav BRZOBOHATY & Dirk NOLF

Diaphus aff. splendidus (BRAUER,

mm

9b Diaphus befralai n. sp

P l a t e 6