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Revue de micropaléontologie 52 (2009) 195–218

Original article

Fish otoliths from the Priabonian (Late Eocene) of North Italy andSouth-East France – Their paleobiogeographical significance

Otolithes de poissons priaboniens (Eocène terminal) du nord de l’Italieet du sud-est de la France – Leur intérêt paléobiogéographique

Angela Girone a,∗, Dirk Nolf b

a Dipartimeno di Geologia e Geofisica, University di Bari, via E. Orabona, 70125 Bari, Italyb Institut royal des sciences naturelles de Belgique, 29, rue Vautier, 1000 Bruxelles, Belgium

Abstract

The study of the Late Eocene (Priabonian) otolith associations from Possagno, North-East Italy, and from the Synclinal d’Allons in HauteProvence, South-East France, allows for the reconstruction of a teleost fauna of 55 taxa, which is the most diversified assemblage presentlyknown from the Upper Eocene Paleo–Mediterranean basin. Thirty-six taxa are identified at the species level, and five of those are new: “genusAlepocephalidarum” astrictus, “genus Lophiiformorum” canovae, “genus Agonidarum” sudans, “genus Uranoscopidarum” cochlearis and Aser-aggodes laganum. In the Synclinal d’Allons, the otolith associations reflect a tropical to subtropical neritic environment with a few mesopelagicfishes. At Possagno, the associations indicate an environment that changed from one that was deep and exposed to the pelagic realm and thenevolved to a more shallow sea with a well-diversified benthic life and less mesopelagic fishes. A paleobiogeographical analysis of all known dataon Priabonian otoliths, worldwide, shows clearly a western Atlantic (Louisiana) and an eastern Atlantic–Paleomediterranean association. In theeastern Atlantic–Paleomediterranean association, the Aquitaine association not only differs from the Possagno–Allons association in function ofits more distant geographical position, but also by its stronger oceanic character in the southern part of the basin, and by the occurrence in thenorth, of a very shallow water facies (Saint-Estephe Formation) that contains some taxa which are known nowhere else in the Priabonian. TheUkraine fauna is characterized by a high number of species, which have an Oligocene record in other European sites. The northern geographiclocation of Ukraine, combined with the good connections to both the North Sea Basin and the Turgai street can provide the explanation. ManyOligocene species (or their close relatives) probably already existed at Eocene times in more northern regions, but could penetrate only in moresouthern European seas since the strong cooling at the beginning of the Oligocene.© 2007 Elsevier Masson SAS. All rights reserved.

Résumé

L’étude des otolithes provenant de Possagno (nord-est de l’Italie) et du Synclinal d’Allons (Alpes de Haute-Provence, sud-est de la France) a per-mis la reconstitution d’une faune de téléostéens comptant 55 taxa. C’est l’association la plus diversifiée actuellement recensée de l’Eocène supérieurdu bassin Paléoméditerranéen. Trente-six taxa ont pu être identifiés au niveau de l’espèce et parmi eux, cinq sont nouveaux : « genus Alepocephal-idarum » astrictus, « genus Lophiiformorum » canovae, « genus Agonidarum » sudans, «genus Uranoscopidarum » cochlearis et Aseraggodeslaganum. Dans le Synclinal d’Allons, les associations d’otolithes reflètent un environnement néritique avec peu de poissons mésopélagiques, sousun climat tropical à subtropical. À Possagno, les associations indiquent un environnement néritique profond, bien exposé au domaine pélagique,évoluant progressivement vers une mer moins profonde, avec moins d’éléments mésopélagiques. L’analyse paléogéographique à l’échelle mondialede toutes les données concernant des otolithes priaboniennes distingue clairement une association ouest atlantique (Louisiana) et une associationest atlantique–paléoméditerranéenne. Dans cette dernière, l’association d’Aquitaine ne diffère pas seulement de celles de Possagno-Allons en
fonction de son éloignement géographique plus considérable, mais aussi par son caractère plus franchement océanique dans la partie méridionaledu bassin et par la présence dans le nord, d’un faciès très côtier (Formation de Saint-Estèphe), contenant certains taxa qui ne sont connus nullepart ailleurs au Priabonien. La faune d’Ukraine est caractérisée par un nombre élevé d’espèces qui, en Europe occidentale, ne sont connues qu’àpartir de l’Oligocène. La position plus nordique de l’Ukraine, combinée avec de bonnes connections avec le bassin de la Mer du Nord et le détroit
∗ Corresponding author.E-mail address: [email protected] (A. Girone).

0035-1598/$ – see front matter © 2007 Elsevier Masson SAS. All rights reserved.doi:10.1016/j.revmic.2007.10.006

mailto:[email protected]

dx.doi.org/10.1016/j.revmic.2007.10.006

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96 A. Girone, D. Nolf / Revue de micropaléontologie 52 (2009) 195–218

e Turgai, en fournit sans doute l’explication. Plusieurs « espèces oligocènes » (ou des formes très apparentées) existaient probablement déjà à’Eocène dans des eaux plus nordiques, mais n’ont pu pénétrer dans les mers plus méridionales qu’à partir du refroidissement considérable, auébut de l’Oligocène.

2007 Elsevier Masson SAS. All rights reserved.

eywords: Otoliths; Fishes; Priabonian; Late Eocene; Mediterranean; Paleobiogeography

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ots clés : Otolithes ; Poissons ; Priabonien ; Eocène terminal ; Méditerranée

. Introduction

During the Cenozoic, the climate of Earth underwent a com-lex evolution that was driven by long-term tectonic processesnd by high frequency cyclic changes in orbital force. The mostrastic climatic changes occurred during the Eocene–Oligocenenterval (Salamy and Zachos, 1999; Zachos et al., 1994,996, 2001). These important climatic variations caused deephanges in the biosphere. Otolith-based reconstructions of theocene–Oligocene fish faunas in Europe, mainly concern Lowernd Middle Eocene or Oligocene associations. Knowledge oftolith associations from the Lower and Middle Eocene is mostomplete for the North Sea Basin and its southern extensionsParis Basin and London–Hampshire Basin) where the recog-ized associations are essentially neritic in composition (Stintonnd Nolf, 1970; Stinton, 1975, 1977, 1978, 1980, 1984; Nolfnd Cappetta, 1976; Nolf and Lapierre, 1979; Steurbaut andolf, 1990). The Aquitaine Basin (southwestern France) offersvery good succession from the Early Eocene until the Miocene

Steurbaut, 1984; Nolf, 1988; Nolf and Brzobohaty, 2002; Nolfnd Steurbaut, 2002), including a middle rich, deep neritic faunaith oceanic components in the Upper Eocene (Nolf, 1988).ata for Late Eocene otolith-based fish faunas from the paleo-editerranean Basin are restricted to the Ukraine Basin (Müller

nd Rozenberg, 2003). Nolf and Steurbaut (2004) reported onhe mainly deepwater otolith association of Piemonte, north-rn Italy, and Nolf and Brzobohaty (1994) and Brzobohatynd Krhovsky (1998) on Oligocene faunas of the Paratethys.dditional information regarding Late Eocene otolith-based fish

aunas is available from U.S. Gulf Coast deposits (Nolf andtringer, 2003). Moreover, data on Middle Eocene otoliths from

he marine tropical Indo-west Pacific realm (Nolf and Bajpai,992) are useful for understanding the paleobiogeographic evo-ution of the fish faunas in the Tethys realm.

The present paper deals with the Upper Eocene otolith-ased fish fauna of the “Marne di Priabona” Formation inhe Possagno section (NE Italy) and contemporaneous otolithssemblages from the Synclinal d’Allons in the Haute ProvenceSE France) (Fig. 1). We also attempted to integrate the resultingata with those from the literature and to analyse them withinpaleogeographic framework in order to compare the differ-

nt biogeographic provinces of Ukraine, NE Italy, Alpes deaute-Provence, the Aquitaine Basin, and the U.S. Gulf Coast.

. Studied materials and locality data

Most of the studied material was collected at Possagno,ocated near Bassano de la Grappa (Province of Treviso, NE

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taly), about 60 km NW of Venice (Fig. 1A). An importantdditional collection came from the Synclinal d’Allons in SErance (Département des Alpes de Haute-Provence), locatedbout 80 km north of Saint-Tropez (Fig. 1A).

The Possagno section was studied intensively for variousicropaleontological taxa, and the results of these studies are

resented in a volume edited by Bolli (1975). The most impor-ant papers for the present study are those of Cita (1975),ho provided detailed sections of the whole Possagno section,

rom the Upper Cretaceous until the Oligocene, Proto Decimat al. (1975) on the calcareous nannoplankton stratigraphy,oumarkine and Bolli (1975) on the planktonic foraminiferaltratigraphy, Braga et al. (1975) on the benthic foraminifera, anderb and Hekel (1975) on the nummulites and on the precise

ocations of the Val Organa sites. Most of the examined fos-ils come from the Cunial quarry, located at the east side of theoad leading northwards from the hamlet of Cunial to the Santaiustina Chapel. Some additional material comes from the Colell’Asse (x = 723.469, y = 5080.511), located about 2 km northf the Santa Giustina Chapel.

Garnier (1872) provided the first good description of theocene deposits in the SE–NW oriented Synclinal d’Allons.his description was completed by the observations of Boussac

1912: p.119), who mentioned the following succession:

marly lagunar deposits with Cerithium diaboli, attaining athickness of about 30 m, and easily seen in a small ravine onthe west side of the Verdon, at x = 937.075, y = 3199.625, andin the eastern flank of the river Ivoire, south of a small parkingarea (x = 339.100, y = 3198.350);nummulitic limestone, attaining a thickness of about 20 m,and visible in two small ravines, running into the Ivoire, some500 and 600 m upstream from the previously cited point nearthe parking area;blue marls containing nummulites and some rather poorlypreserved molluscs. These marl deposits, which fill the wholecentral part of the synclinal, attain a thickness of about 50 m,according to Castelain (1956: p. 27). These marls were sam-pled in the banks of the Ivoire torrent and in the badlandcanyons at “la Tuilière”.

.1. Possagno, Cunial quarry

Topographic map 1/25,000, Cavaso del Tomba quadrangle,
= 723.107, y = 5080.728.
The section of the Cunial clay pit was figured by Cita (1975:ig. 9), who provided a metric scale of the pit measured from

he top to the bottom of the section. She located the position of

A. Girone, D. Nolf / Revue de micropaléontologie 52 (2009) 195–218 197

Fig. 1. A. Location of the studied sections. B. Stratigraphic section measured in the Synclinal d’Allons area in La Tuilière locality. C. Measured section at Possagno.Fig. 1. A. Localisation des coupes étudiées. B. Coupe de l’affleurement « La Tuilière », dans le Synclinal d’Allons. C. Coupe de la série stratigraphique de Possagno.

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he micropaleontological samples in relation to this scale. Theelevant interval of the scale is used in our Fig. 1B, because ouramples (A–E) are located with reference to the same lithologicaluide horizons:

sample A (about 50 kg of sediment) is from a Turritella hori-zon located just below 170 m and between samples 666 and667 of Cita;sample B (about 50 kg of sediment) is from a shell-rich hori-zon above a somewhat indurated level just below 130 m andbetween samples 670 and 671 of Cita. This point is locatedexactly 24 m below the middle horizon of indurated limoniticsand under our sample D, which constitutes an excellent guidehorizon;sample C (about 500 kg of sediment) is from just above a cal-careous limonitic limestone ledge, which is the same positionas Cita’s sample 675. A loose block of very fossiliferous sed-iment, apparently from the same horizon, also provided somefossils;sample D (about 30 kg of sediment) is from just above themiddle horizon of indurated limonitic sand at about 107 mand is just above Cita’s sample 677;sample E (about 30 kg of sediment) is from 4 m below theupper indurated level and between Cita’s samples 678 and679.

.2. Possagno, Monte Collalto, site 2

Topographic map 1/25,000, Cavaso del Tomba quadrangle,= 724.936, y = 5080.610.

A sample of about 30 kg from the upper part of the Possagnoarls (see southern part of section 3, p. 15 of Cita, 1975) pro-

ided some scarce otoliths, among which was the holotype ofgenus Lophiiformorum” canovae.

.3. Possagno, Val Organa, site 1

Topographic map 1/25,000, Cavaso del Tomba quadrangle,= 724.043, y = 5080.182.

A sample of about 30 kg of sediment and corresponding tobout sample 708 of Herb and Heckel (1975: Fig. 2, p. 115)rovided no otoliths.

.4. Synclinal d’Allons, site B: banks of the torrent Ivoire

Topographic map 1/25,000, 3541 W, Saint-André-des-Alpes,= 939.950, y = 3196.750.

Gray silty marls with nummulites (sample of about 300 kg),xposed in the banks of the river l’Ivoire, about 125 m SE of “Laastide neuve”. The upper course of the Ivoire is called “Torrentes Garets”, which is the name that Boussac (1912) used insteadf l’Ivoire, that appears on the latest 1/25,000 topographic map.

.5. Synclinal d’Allons, La Tuilière (Fig. 1B)

Topographic map 1/25,000, 3541 W, Saint-André-des-Alpes,= 939.775, y = 3197.550.

léontologie 52 (2009) 195–218

Badlands in the east flank of the Ivoire valley, exposing grayilty marls.

Sample (about 200 kg) from + 2 until + 4 m of the section;Sample (about 30 kg) from + 10 until + 12 m of the section.

. Systematic paleontology

A list of all otolith-based taxa represented in our sam-les is given in Table 1. The classification adopted is thene proposed by Nelson (2006). All cited species are shownn Figs. 2–8. Additional comments are given only for newpecies or for those requiring discussion. In those cases wherepecific identity could not be decided unequivocally, the abbre-iation aff. (affinis) is inserted between the name of theenus and the name of the species group. The abbrevia-ion cf. (conformis) is used in cases where the condition ofreservation of the otoliths did not allow conclusive specificdentification. The notations A1a, B1a... and A1b, B1b... etc.n the figures are used to indicate respectively ventral andnner (= mesial) views. Otoliths with only numbers and nomall letters show inner views. For general information abouttoliths (morphological nomenclature, composition, diagnos-ic value, ontogenetic changes, etc.), the reader is referred toolf (1985). Concerning collective (or open) generic nomen-

lature, the current procedure used in numerous papers ontolith taxonomy is applied. For species of uncertain genericosition (i.e., whose systematic position can be identified onlyt familial, subordinal, or ordinal level) the word “genus,”ollowed by the name of the family or higher categoryn plural genitive, followed by the species name is used;.g., “genus Heterenchelyidarum” circularis. See also Nolf1985) for further explanation.

.1. Remarks on taxa requiring comments

(1) “genus Heterenchelyidarum” aff. colei (Müller, 1999)(Fig. 2A) – In the Eocene of Europe, two specieshave been described: “genus Heterenchelyidarum” cir-cularis (Shepherd, 1916) from the Lower and MiddleEocene of the Belgian Basin (Steurbaut and Nolf, 1990)and from the Middle Eocene of the London–HampshireBasin (Shepherd, 1916), and “genus Heterenchelyidarum”richardsi Nolf, 1988, from the Upper Eocene of Aquitaine.“Genus H.” circularis has otoliths with a typical subcir-cular posterodorsal rim and a sulcus that is larger in theanterior region. “Genus H.” richardsi is characterized bya wide sulcus, a well-marked posterodorsal angle and acharacteristic hollowing in the middle portion of the dor-sal rim. The otoliths from Possagno are weakly elongated.The sulcus is completely filled with colliculum and isnearly equally wide across the entire length. The dorsalrim is regularly curved. The highest part of the otoliths is
located in the posterior portion. The posterodorsal angle isprominent. These features allow for the attribution of thePossagno materials to “genus Heterenchelyidarum” colei(Müller, 1999) from the Middle and Upper Eocene deposits


Table 1List of otolith-based fish taxa from Synclinal d’Allons and Possagno. In bold: taxa today absent from the MediterraneanTableau 1Liste des poissons (otolithes) recensés du synclinal d’Allons et de Possagno

Icono-graphy Possagno Allons

A B C D E La Tuilière Allons B

Heterenchelyidae “genus Heterenchelyidarum” aff. colei (Muller, 1999) (1) Fig. 2A 2 24 2Congridae Ariosoma nonsector Nolf and Stringer, 2003 (2) Fig. 2E 1 4 1

Conger glaber Brzobohaty, 1967 (3) Fig. 2F 1Conger sp. Fig. 2F 1 1Pseudophichthys sp. (4) Fig. 2D 1Rhynchoconger transversus (Sulc, 1932) Fig. 2I 2 4 77 4 3 22“genus Congridarum” eocaenicus (Sulc, 1932) Fig. 2G 8 16 29

Nettastomatidae Hoplunnis ariejansseni Nolf, 1988 (5) Fig. 2B 5Clupeidae Clupeidae ind. 1 1 11 1 4Opisthoproctidae Opisthoproctus weitzmani Nolf, 1988 Fig. 2C 1Alepocephalidae “genus Alepocephalidarum” astrictus n. sp. Fig. 6B 4 3Sternoptychidae Valenciennellus brzobohatyi Steurbaut, 1982 Fig. 3J 1 2Phosichthyidae Vinciguerria biarritzensis (Sulc, 1932) Fig. 3F 3 2 7 1 ? 1Synodontidae “genus Synodontidarum” cf. boulangeri Nolf, 1988 (6) Fig. 2K 1

Saurida sp. Fig. 2L 1Scopelarchidae Scopelarchidae ind. Fig. 2J 1Myctophidae Diaphus excavatus (Sulc, 1932) Fig. 3A 12 1 8

Diaphus ind. 1Bregmacerotidae Bregmaceros brihandensis Nolf, 1988 Fig. 3B 220 189 465 6 12 20Carapidae Onuxodon aff. coheni Nolf, 1980 (7) Fig. 3G 1Ophidiidae Hoplobrotula biscaica (Sulc, 1932) Fig. 3D 7 14 204 2 1 5 19

“genus Neobythitinarum” hilgendorfi (Koken, 1891) Fig. 4F 5 11 1“g. aff. Glyptophidium” biarritzensis (Sulc, 1932) Fig. 4E 2 8 1 1 4 24“g. Sirembinorum” aff. arcuatus (Stinton, 1966) (8) Fig. 3H 3

Bythitidae Grammonus pseudoacuminatus (Sulc, 1932) Fig. 3I 1 14Lophiidae “genus Lophiiformorum” canovae n. sp. Fig. 7D 1Antennariidae “g. Antennariidarum” aff. furcatus (Frost, 1933) (9) Fig. 3C 2 27

Antennariidae ind. 1 1 3 1 2Cyprinodontoidei Cyprinodontidae ind. 1Berycidae Centroberyx ingens (Koken, 1884) Fig. 4B 2 4 4 17Holocentridae “genus Myripristinarum” sp. Fig. 5J 1Zenionidae “genus aff. Zenion” sp. Fig. 4D 1Triglidae “genus Triglidarum” vectensis Nolf, 1973 (10) Fig. 3E 1Platycephalidae Platycephalus janeti (Priem, 1911) Fig. 5I 4Agonidae “genus Agonidarum” sudans n. sp. Fig. 6D 1Percoidei incertae sedis “genus Percoideorum” aff. aquitanicus Nolf, 1988 Fig. 4G 3 1 2Acropomatidae Acropomatidae ind. 1Serranidae Chelidoperca elongata (Sulc, 1932) (11) Fig. 4A 4 8 8 43

Serranidae ind. 11Priacanthidae Pristigenys aff. obliquus Nolf and Stringer, 2003 (12) Fig. 4C 1 2Apogonidae Apogon decoratus Stinton, 1980 Fig. 5F 1

Apogon macrolepis Storms, 1898 Fig. 5K 2 2 12 ? 1“g. Apogonidarum” ventrolobatus Schwarzhans, 1973 Fig. 5B 3 4 11 2 1 15Apogon sp. (13) (locality Monte Collalto, Possagno) Fig. 4H

HaemulidaeHaemulidae ind. Fig. 5H 1

Sparidae Sparidae ind. Fig. 4I 3Cepolidae Cepola yrieuensis Steurbaut, 1984 (14) Fig. 5E 12 19 2 5Labridae Labridae ind. 1Champsodontidae Champsodon spinosus Schwarzhans, 1977 (15) Fig. 5D 1 2 1Uranoscopidae “genus Uranoscopidarum” cochlearis n. sp. Fig. 7A 1 3Scombridae Scombridae ind. 1 1Citharidae Citharus cauneillensis Nolf, 1988 Fig. 5C 3 3 34 2

Citharus sp. (16) Fig. 5A 1Soleidae Aseraggodes laganum n. sp. Fig. 8B 2 6

Microchirus sp. Fig. 5G 1total number 260 271 1018 25 7 35 215


Fig. 2. Otoliths from Possagno, and Allons (first series), see appendix A for locality data and collection numbers.Fig. 2. Otolithes de Possagno et d’Allons (première série), voir annexe A pour les données de gisements et numéros de collection.


Fig. 3. Otoliths from Possagno and Allons (second series), see appendix A for locality data and collection numbers.Fig. 3. Otolithes de Possagno et d’Allons (deuxième série), voir annexe A pour les données de gisements et numéros de collection.


Fig. 4. Otoliths from Possagno and Allons (third series), see appendix A for locality data and collection numbers.Fig. 4. Otolithes de Possagno et d’Allons (troisième série), voir annexe A pour les données de gisements et numéros de collection.


Fig. 5. Otoliths from Possagno and Allons (fourth series), see appendix A for locality data and collection numbers.Fig. 5. Otolithes de Possagno et d’Allons (quatrième série), voir annexe A pour les données de gisements et numéros de collection.


Fig. 6. Otoliths from Possagno and Allons and present day species (fifth series), see appendix A for locality data and collection numbers.Fig. 6. Otolithes de Possagno et d’Allons et espèces actuelles (cinquième série), voir annexe A pour les données de gisements et numéros de collection.


Fig. 7. Otoliths from Possagno and Allons and present day species (sixth series), see appendix A for locality data and collection numbers.Fig. 7. Otolithes de Possagno et d’Allons et espèces actuelles (sixième série), voir annexe A pour les données de gisements et numéros de collection.

2 cropa
of the U.S. Gulf Coast (Müller, 1999; Nolf and Stringer,2003) rather than to the two above-cited European Eocenespecies;

(2) Ariosoma nonsector Nolf and Stringer, 2003 (Fig. 2E)– Eocene records of Ariosoma have been reported fromU.S. Gulf Coast deposits (Nolf, 2003; Nolf and Stringer,2003). In Europe, the genus Ariosoma was reportedfrom the Lower Oligocene of Atlantic and Mediterranean(Steurbaut, 1984; Nolf and Steurbaut, 2004). The spec-imens from Possagno are most similar to the Americanspecies Ariosoma nonsector;

(3) Conger glaber Brzobohaty, 1967 (Fig. 2F) – One otolithbroken at the anteroventral rim presents a congrid sulcustype. The otolith is elongate in the anteriorposterior direc-tion and presents a pointed posterior rim. The dorsal andventral rims, regularly curved, join each other in a some-what pointed posterior rim. The specimen from Possagnois closely comparable with Conger glaber from the LowerOligocene of Moravia, but our specimen seems to be higherin the central portion and less slender than the holotype ofC. glaber. These features, however, are within the limitsof variability of the C. glaber material from the Rupelianof the Aquitaine Basin figured by Steurbaut (1984);

(4) Pseudophichthys sp. (Fig. 2D) – One small specimen canbe referred to the genus Pseudophichthys. In the UpperEocene, Pseudophichthys elongatus (Sulc, 1932) hasbeen reported from the Aquitaine Basin, Pseudophichthysglaber (Koken, 1888) from the Jackson Group (Missis-sippi, U.S.A.) and Pseudophichthys acer Schwarzhans,1985 from southern Australia. The specimens from Pos-sagno cannot be referred to any of these species due to theirprominent posterodorsal angle and their characteristic pos-terior rim which is nearly subvertical in the inferior portionand concave in the upper portion. Although our speci-men clearly shows these features, more material wouldbe required for a formal description to be made;

(5) Hoplunnis ariejansseni Nolf, 1988 (Fig. 2B) – Thespecimens from Possagno can be attributed to H. arie-jansseni, reported from the Lower Eocene (Ypresian) ofthe Aquitaine Basin (Nolf, 1988). Other fossil records ofthis genus, today represented by eight tropical Atlantic andone eastern Pacific species, are unknown;

(6) “genus Synodontidarum” cf. boulangeri Nolf, 1988(Fig. 2K) – The general shape of this specimen agreeswith that of “genus Synodontidarum” boulangeri figuredby Nolf (1988: Pl. 4, Figs. 15 and 16) from the Priabo-nian of the Aquitaine Basin; however, the crista superiorof our specimen is shorter and less marked than in thosespecimens. This difference may be due to the rather poorpreservation of our specimen; more and better material isrequired to ascertain the presence of “genus Synodonti-darum” boulangeri in the Possagno section;

(7) Onuxodon aff. coheni Nolf, 1980 (Fig. 3G) – A single
small specimen characterized by a perfectly oval sulcus,undivided in ostium and cauda, and by a flat inner facecan be referred to the genus Onuxodon. Two species,O. kiriakoffi Nolf, 1980 and O. coheni Nolf, 1980 are
léontologie 52 (2009) 195–218

known, respectively, from the Lower Eocene (Ypresian)and the Lower Oligocene of the Aquitaine Basin. ThePossagno specimen is most closely related to the LowerOligocene species, based upon its high shape and hol-low posterior portion of the dorsal rim, but it has morerounded anterior and posterior margins and a smootherposterodorsal angle;

(8) “genus Sirembinorum” aff. arcuatus (Stinton, 1966)(Fig. 3H) – This species is characterized by rather robustand elongate otoliths with a spine-like posterior end. Theotoliths are thickest in the upper part of the anterior por-tion with a smooth dorsal rim and a cutting ventral margin.The thickness along the anterodorsal area is accentuatedby a weak expansion. The inner face is strongly convex,whereas the outer face is slightly convex in the anteriorportion but strongly concave in the posterior part, corre-sponding to a thinner region of the otolith. The sulcus,superficially incised, is divided; the cauda is shorter thanthe ostium. The sulcus is filled by colliculum separatedin ostial and caudal portions by a shallow incision that isstrongly inclined in a posteroventral to anterodorsal direc-tion, but the widths of the ostial and caudal collicula arenearly equal. The Possagno specimens closely resemblethe otoliths of “genus S.” arcuatus (Stinton, 1966) knownfrom the Ypresian of Southern England (Stinton, 1966,1977) and from the Middle Eocene of Belgium (figuredby Nolf, 1980 as “genus S.” cappettaensis). The latter arecharacterized by having a sulcus with a subvertical divi-sion of the collicula in which the caudal portion is narrowerthan the ostial portion. In all those figured specimens, theposterior portion of the cauda is curved towards the ven-tral margin. In the specimens from Possagno, the caudais nearly straight and the ostium and cauda are equal inwidth. Moreover, the specimens from Possagno have amore prominent angular posterior end. The sulcus patternof these specimens is very similar to that of the specimenfigured by Nolf and Steurbaut (2002) from the Rupelian ofthe Aquitaine Basin as “genus S.” aff. arcuatus. A defini-tive identification of the material from Possagno and theAquitaine Oligocene is not possible because of the limitedamount of available material;

(9) “genus Antennariidarum” aff. furcatus (Frost, 1933)(Fig. 3C) – In the Eocene, three valid otolith-based speciesof antennariids are known: Antennarius euglyphus Stinton,1966 from the Lower Eocene (Ypresian, southern Eng-land and Aquitaine Basin), Antennarius excavatus Stinton,1978 from the Middle Eocene (Lutetian, southern Eng-land) and “genus Antennariidarum” furcatus Frost, 1933from the Middle Eocene (Bartonian, southern England).Antennariid otoliths have rather significant variability andscarcely any fossil species are represented by enough spec-imens to evaluate intraspecific variability. The otolithsfrom Possagno are characterized by a deeply incised sul-
cus, with the crista inferior divided into ostial and caudalportions by a slight curvature. The crista superior is accen-tuated by a depression just above it. The otoliths are thickand oval in shape. The ventral rim shows a blunt anteroven-

cropaléontologie 52 (2009) 195–218 207

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tral angle and an oblique posterior portion. The dorsal rimis rather variable, as shown in Fig. 3C (1–3). The Possagnomaterial is most similar to the Bartonian material fromsouthern England, but a definitive determination cannot bemade because variability within the Bartonian material ispoorly documented;

10) “genus Triglidarum” aff. vectensis (Nolf, 1973) (Fig. 3E) –A single otolith shows reasonable affinities with the Lute-tian “genus Triglidarum” vectensis (Nolf, 1973) from theParis Basin; however, our specimen is characterized byhaving a wider cauda that is separated from the ostium bya shallower collum. Moreover, the posterior rim is morepointed in the Priabonian specimen than in the Lutetianmaterial;

11) Chelidoperca elongata (Sulc, 1932) (Fig. 4A) – Thisspecies was originally described from the Priabonian ofAquitaine by Sulc (1932) as Centropristis elongatus andby Nolf (1988) as “genus Serranidarum” elongatus. Thecorrect generic allocation for this taxon seems to be theIndo-west Pacific genus Chelidoperca; see Rivaton andBourret (1999: Pl. 11) for pictures of the otoliths of variousRecent species;

12) Pristigenys aff. obliquus Nolf and Stringer, 2003 (Fig. 4C)– The specimens from Possagno are characterized bya manifest anteroposterior convexity, which is morepronounced in the inner face than in the outer face.Furthermore, the otoliths show some distortion along ananteroventral to posterodorsal axis. These features revealsome similarity of the specimens from Possagno withotoliths of Pristigenys obliquus described by Nolf andStringer (2003) from the Priabonian of Louisiana ratherthan with the European Eocene species (see Nolf, 1985:p. 82);

13) Apogon sp. (Fig. 4H) – A small and elongate single otolithcan be referred to the genus Apogon, but it differs fromthe other apogonid species treated herein mainly by theconsiderable height of its anterior part. This otolith seemsto be most similar those of the Plio–Pleistocene A. lozanoiBauza, 1957, but the available material does not permit adefinitive identification;

14) Cepola yrieuensis Steurbaut, 1984 (Fig. 5E) – Thisspecies is characterized by elongate otoliths with aconvex inner face and a nearly flat outer face. Thedorsal rim is regularly curved and shows a slightlysalient posterodorsal angle. The posterior end is acumi-nate. The cauda is shorter than the ostium; there is amarked angle at the junction of the ostial and caudalcrista inferior. These features allow the specimensfrom Possagno to be referred to the Lower OligoceneC. yrieuensis from the Aquitaine Basin. In the Eocene,Cepola otoliths are represented by the Lower EoceneCepola densa (Frost, 1934) and C. robusta Nolf, 1988and by the Middle Eocene C. bartonensis Schubert,
1916 and C. excavata (Stinton, 1978). The otoliths ofC. yrieuensis are quite slender with respect to both LowerEocene species. Cepola yrieuensis has some similaritieswith Cepola bartonensis (Fig. 6F), but in this species, the
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ig. 8. Otoliths from Possagno and Allons and present day species (seventheries), see appendix A for locality data and collection numbers.ig. 8. Otolithes de Possagno et d’Allons et espèces actuelles (septième série),oir annexe A pour les données de gisements et numéros de collection.

otoliths have a longer cauda, no salient posterodorsal angleand a more regularly curved dorsal and upper posterior rim;

15) Champsodon spinosus (Schwarzhans, 1977) (Fig. 5D)– This species, described from the Lower Oligocene ofGermany, is reported for the first time in the Eocene. ThePossagno occurrence represents the earliest fossil recordof Champsodon in Europe, where it is known up to theLate Oligocene. Today, the Champsodontidae are onlyfound in the Indo-Pacific realm;

16) Citharus sp. (Fig. 5A) – A single specimen, broken inthe anterior portion, is even more elongated than C.cauneillensis and probably represents a different species.

.2. Description of new species

“genus Alepocephalidarum” astrictus nov. sp.Fig. 6B
Type material: Holotype: a left otolith (Fig. 6B (1a, b)), four
aratypes of which two are figured (Fig. 6B (2, 3)).Type locality: Marna di Possagno (Priabonian) at Possagno,

unial quarry.

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Etymology: astrictus, a, um = buttoned: refers to the sulcusf the otoliths that resembles the form of a button hole.

Description: This species is characterized by subcirculartoliths with a slightly flat inner surface that shows a charac-eristic depression in the middle portion of the dorsal area. Thepper portion of the posterior rim is straight to weakly concaveut, in the inferior portion, it shows a blunt angle that may be fol-owed by a small concavity. The ventral rim is regularly curvednd, in large specimens, bears some small lobes. The sulcus hascauda that is longer, narrower, and more deeply incised than thestium. The rostrum and antirostrum are smooth and of similarize. The crista superior is more salient than the crista inferior,hich is smooth in the caudal portion, but sharp in the ostialortion. The outer face is generally smooth and weakly convex.

Affinities: These otoliths are morphological comparableith the extant alephocepalid Photostylus pycnopterus Beebe,933 (Fig. 6 (A1, 2)), but the habitat of this Recent fish isompletely incompatible with the paleoenvironment of theossil assemblage. This fish is bathypelagic at 1000–2000 mnd is associated with steeply inclined slopes (Whitehead etl., 1984). The hypothesis that our fossil represents an extinctpecies of Recent genus Photostylus is not excluded, but werefer to keep it in open generic nomenclature.

“genus Lophiiformorum” canovae nov. sp.

Fig. 7DType material: Holotype: a right otolith (Fig. 7D (2a, b)), one

aratype (Fig. 7D (1a, b)).Type locality: Marna di Possagno (Priabonian) at Possagno,

unial quarry.Etymology: For the sculptor Antonio Canova (1757–1822),

orn in Possagno, alluding to the morphology of the sulcus,hich seems to be sculptured.Description: This species is characterized by robust and

rapezoidal otoliths with a straight ventral rim that is much longerhan the rounded dorsal one. The posterior rim is characterizedy a rather strong concavity, just after the rounded posterodorsalngle. This concavity nearly forms an incision in the paratypehere the inferior portion of the posterior rim is more or less

runcated, and shows a second incision. In the holotype, theosterior rim is smoothly rounded. The outer face is smooth andtrongly convex; the inner face is moderately convex. The sul-us is short and mainly located in the centre of the inner face.owards the anterior rim, the ostium becomes obsolete. Thentirostrum is only marked by some smooth rounding, and theostrum is blunt but salient. The sulcus is rather deeply incised.he division into ostial and caudal portions is only marked by anndulation in the crista inferior. The dorsal area exhibits a well-eveloped depression just below the superior crista. The ventralrea bears a strongly curved ventral furrow in its central portionelow the sulcus. This furrow divides the ventral area into anpper, somewhat swollen part, and a lower, less salient zone.

Affinities: The two specimens from Possagno exhibit eas-
ly recognizable features for purposes of identification, but anyttribution to a Recent family is uncertain. Tentatively, they arettributed to the Lophiiformes because of the irregular shape ofhe otolith and its poorly structured sulcus.
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léontologie 52 (2009) 195–218

“genus Agonidarum” sudans nov. sp.Fig. 6DType material: Holotype: a left otolith (Fig. 6D (1a, b)).Type locality: Marna di Possagno (Priabonian) at Possagno

ection.Etymology: sudare = to sweat, alludes to the modern geo-

raphic distribution of the family Agonidae, which is restrictedo the polar sea and northern North Atlantic and Pacific Oceans,hich leads us to a spontaneous anthropocentric claim: “This

ellow must have sweated very much, in the warm Priabonianaters.”Description: This species is characterized by a robust and

val otolith, with a nearly flat inner face and a strongly convexnd smooth outer face. The sulcus, poorly structured, occupiesssentially the anterior portion of the inner face and is rathereeply incised, especially in its ventral part. The crista superior isearly completely obsolete. In the crista inferior, the division intostial and caudal portions is marked by a flexure. The ventral areaears a narrower furrow just above the rim. Between the furrownd the crista, the ventral area is convex, and the entire dorsalrea is flat and weakly depressed. The ventral rim is regularlyurved; the dorsal area is most expanded in its central part. Theosterior rim is salient but blunt, and the rostrum is well marked.here is no clear antirostrum.

Affinities: The otolith of this species shows a sulcus patternhat can be compared with what is observed in otoliths ofgonids, such as the Recent Aspidophoroides olriki (Lutken,876) (Fig. 6C (1, 4)). This species is also characterized bytoliths with a flat inner face and a strongly convex outer face.lso, their sulcus show a reasonable similarity, but, in theecent species, it is better divided into an ostial and caudalortion. Fossil agonids are known from the Middle Eocenef southern England (Podothecus costulatus Stinton, 1978),rom the Middle Oligocene of Belgium [“genus Agonidarum”inusculus (Nolf, 1977)], from the Middle Miocene of Rouma-ia (Agonus elongatus Weiler, 1950) and from the Pliocene ofelgium (Agonus cataphractus Linnaeus, 1758) (Nolf, 1985),ut no relevant similarity is found between these already knownossil species and the one from Possagno.

“genus Uranoscopidarum” cochlearis nov. sp.Fig. 7AType material: Holotype: a left otolith (Fig. 7A (1a, b)) and

hree paratypes of which two are left otoliths (Fig. 7A (2, 4)),nd one is a right otolith (Fig. 7A (3)) from the Possagno section.

Type locality: Marna di Possagno (Priabonian) at Possagnoection.

Etymology: cochlearis = spoon-like, alludes to the shape ofhe sulcus, which is deeply concave and like a spoon.

Description: This species is characterized by elongatend robust otoliths with an outer face more convex than thenner face. The sulcus is rather deeply incised but is notlearly divided into an ostium and a cauda. The deepest part
f the sulcus courses down behind the strongly marked cristanferior and collicular crest. The crista superior is obsolete,nd the ostium is wider than the cauda. The entire dorsal areas depressed and the ventral area bears a narrow furrow that

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ogether with the crista inferior delimits a very convex area. Theorsal rim bears some lobe furrows that more-or-less incise theorsal area. These furrows continue deeper in the outer surface.he rostrum is present in all specimens, and it ranges from verymooth to pointed. The antirostrum is not always developed,ut when present, it is very short and blunt. The posterior endaries from acuminate to more rounded. In the holotype, theosterior rim bears two pronounced lobes.

Affinities: These otoliths have a reasonable resemblanceo those of the otoliths of uranoscopids. In particular, theyre comparable with the Recent Genyagnus monopterygiusSchneider, 1801) (Fig. 7B) and Gnathagnus innotabilis Waite,904 (Fig. 7C). These extant uranoscopids have a similarulcus pattern and morphology: the sulcus is deeply incisednd is characterized by a well-marked crista inferior and by aoorly marked to absent crista superior; however, in the Recentpecimens the sulcus seems to be more deeply incised. Theorsal area is depressed in both of the Recent species, and theuter face is strongly lobated.

Aseraggodes laganum nov. sp.

Fig. 8BType material: Holotype: a left otolith (Fig. 8B (1a, b));

even paratypes from Possagno, of which one is figured (Fig. 8B2a, b)) and six unfigured specimens from Allons.

Type locality: Marna di Possagno (Priabonian) at Possagno,unial quarry.

Etymology: laganum = pancake, alludes to the fact that theeneral shape and surface of the otoliths is like that of a pancake.

Diagnosis: This species is characterized by globally shapedtoliths, but with irregular outlines. The anterior portion of theentral rim is strongly oblique and delimits a rather narrownteroventral area, much narrower than its dorsal counterpart.he inner face is flat and bears an incised sulcus that is divided

nto an elongated ostium and a slightly shorter, round-shapedauda. The cristae are well marked. In both the dorsal and ven-ral areas of the inner face, there is a wide longitudinal depressionbove and below the sulcus; these areas join each other in theosterior area behind the sulcus. The sulcus is not completelypen on the anterior rim, which bears a smoothed rostrum. Theuter face is entirely smooth and nearly flat. The otoliths arehickest in their ventral portion.

Affinities: These Priabonian otoliths are rather similar tohose of the Recent soleid genus Aseraggodes, which untilow has been unknown in the fossil record. They are hereinompared with those of the Recent A. kaianus (Gunther, 1880)Fig. 8A (1, 2)), which, however, have a more quadrangularutline. Otoliths of other Recent Asseraggodes species also haveeen figured by Schwarzhans (1999); they are all characterizedy a flat inner face with a rather wide circumsulcal depression.toliths of other soleid genera are characterized by a stronger

onvexity of the inner face.

. Composition of the otolith-based fish fauna

The otolith assemblages from Possagno and the Synclinal’Allons are composed mainly of benthic fishes and some

(cas

léontologie 52 (2009) 195–218 209

are additional pelagic taxa. In the Possagno section, the fivexamined samples comprise almost homogeneous fish otolithssemblages, but assemblage C has greater diversity (Table 1).enthic forms dominate in both abundance and diversity, espe-ially eel-shaped fishes that lived on or in the bottom mud.mong these, the congrids Rhynchoconger transversus, “genusongridarum” eocaenicus, and the ophidiid Hoplobrotula bis-aica are the most common taxa. In the Recent fauna, congridsnclude species living on the continental shelf and slope. Theyre distributed worldwide in tropical and temperate seas. Inhe Paleogene, congrid otoliths are very abundant in nearly alloft muddy bottom neritic environments (Nolf, 1985). Many ofhese shallow-water taxa seem to be more similar to present-dayeep-water forms than they do to neritic ones, such as Con-er. Similarly, the Recent ophidiid fauna is composed mainly ofathybenthic fishes with a few neritic and pelagic forms. Duringhe Paleogene, they were represented by a very rich neritic faunahat lived mainly on soft and muddy substrates (Nolf, 1985).

The apogonids are also well diversified and common through-ut the entire section, but they represent only 2% of the fauna inhe samples A–C and 8% in sample D. In the uppermost samplehey only make up 1% of the fauna. In the Eocene, apogo-ids are common in tropical and subtropical neritic deposits.oday they are found in all tropical and many warm temperateeas, mostly in shallow waters but sometime in deeper waters,p to 200 m. Other relatively frequent shallow neritic taxa arentennariids, Pristigenys, Citharus, and Platycephalus, occur-ing only in sample C. In sample C, these taxa are associatedith other shallow neritic groups, such as Onuxodon, agonids,icrochirus and Aseraggodes. Some taxa, such as Chelidop-

rca (samples A and C) and Centroberyx (samples B and C),hat inhabit generally deep shelf habitats comprise less than 1%f these assemblages. Pelagic taxa are rare and are mainly repre-ented by epimesopelagic fishes. Among these Bregmaceros, aenus of small epipelagic fishes (between 0 and 200 m), occursn great numbers. In the lower samples (A–C), it representsrom about 84% (sample A) to about 46% (sample C). Otheresopelagic taxa are represented by percentages less than 1%.The composition of the Possagno assemblages reflects a ner-

tic environment exposed to the pelagic realm. However, alonghe section, the diversity and abundance of shallow neritic ele-

ents increase from the lowermost samples A and B towardsample C. This increase could indicate environmental evolutionrom deeper and more exposed to the pelagic realm setting, inhe lower-middle interval of the section, to a shallow neritic seaith abundant benthic life and less influences from the pelagic

ealm upward. In the uppermost interval of the section (samplesand E) the diversity and abundance of both, neritic and pelagic

axa, decrease, but the very restricted number of the specimenser sample does not permit any relevant palaeoenvironmentalypothesis.

The otolith assemblages from the Synclinal d’Allons and, inarticular from Allons B, are very similar to those of Possagno
Table 1). In Allons B, congrids are represented only by Rhyn-hoconger transversus, and apogonids also have low diversitynd abundance. By contrast, the bythitid Grammonus is repre-ented by just one specimen in Possagno (sample C), but it repre-

2 cropaléontologie 52 (2009) 195–218

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ents 6% of the entire assemblage in Allons B. However, severalhallower taxa like Platycephalus, Pristigenys and antennariidsre lacking or negligible in Allons; the latter is represented onlyy two unidentifiable specimens in Allons B. Deeper neritic taxa,uch as Chelidoperca (20%) and Centroberyx (8%) are repre-ented by higher percentages than in the Possagno assemblages.he pelagic group displays a low diversity. The epipelagic Breg-aceros and the epimesopelagic Valenciennellus are represented

n both samples. These data indicate a neritic environment, withome exposure to the pelagic realm.

Several taxa represented at Possagno and Allons, suchs Hoplobrotula, Centroberyx, Platycephalus, Chelidoperca,hampsodon and Aserragodes, have a present-day distribution

hat is restricted to the Indo-Pacific realm. Today, these fishesre distributed in temperate and tropical–subtropical waters.nother group of taxa with a broader geographic distributionattern, but also absent in the modern Mediterranean, is recog-ized. This group include Hoplunnis (distributed on the shelfnd upper continental slope of the tropical and subtropicalreas of the Atlantic and Pacific Oceans), antennariids (ben-hic fishes distributed in shallow waters of all tropical and warmemperate seas, except the Mediterranean), Pristigenys (occur-ing in tropical–subtropical waters of the Indo-West Pacificrea and western Atlantic) and acropomatids (distributed ineep neritic and continental slope of all tropical-subtropicalceans). Among the taxa, which today are also distributed inhe Mediterranean Sea, heterenchelyids, congrids, apogonidsnd haemulids are indicative of tropical to subtropical climate.ne should also note the presence, at Possagno, of a species

hat we attribute to the agonids, a family which, today, inhabitsold or temperate waters of the northern hemisphere. Pelagicaxa, at Possagno, include some typical bathypelagic forms likepisthoproctus and an alepocephalid. Extant Opisthoproctus

pecies occur below 300 m and have a circumtropical dis-ribution. According to Quéro (1990), the distribution limitsf some Opisthoproctus species are marked by the 8–10 ◦Csotherm.

.1. Paleobiogeographic and stratigraphic implications

Priabonian otolith associations are known from variousasins, all located between 30◦ and 50◦N and extend fromouisiana to Ukraine (Fig. 9). From west to east, they are as

ollows: Louisiana (Nolf and Stringer, 2003), Aquitaine Basin,W France (Nolf, 1988), Alpes de Haute-Provence, SE Franceherein), Possagno (Province of Treviso, NE Italy) (herein) andkraine (Müller and Rosenberg, 2003).In Louisiana, Priabonian otoliths were collected from the

azoo Clay. In total, 43 taxa were recorded of which 27 coulde identified at the species level. The associations mainly reflectshallow neritic environment with muddy bottom, relatively

istant from the oceanic realm, in tropical to subtropical waters.hey are devoid of any typical mesopelagic or deep-water fishes.

In Aquitaine (SW France), otoliths were collected mainly inhe south of the basin, from the Brihande Marls (NP 19/20 Zone)nd their lateral equivalents, the Côte des Basques Marls andouches de Cauneille (64 taxa). These associations essentially

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ig. 9. Localisation géographique des aires ayant fournies les données pour larésente analyse.

eflect deep neritic environments that are well exposed to theceanic realm. Otoliths of mesopelagic fishes account for about9% of all collected specimens. More to the north, in the Borde-ais, the Saint-Estèphe Formation provided otoliths of 14 taxa,ll belonging to very near shore tropical to subtropical fishes.t the time of Nolf’s (1988) report, no precise information on

he age of the Saint-Estèphe Formation was available. On theasis of considerations of sequence stratigraphy, Sztrakos et al.1998) referred these deposits to the Priabonian.

In the Alpes de Haute-Provence (SE France), Priaboniantoliths are from marly deposits in the Synclinal d’Allons (seeStudied material and locality data”). A total of 24 taxa areecorded, reflecting a tropical to subtropical neritic environment,ut some scarce mesopelagic fishes are also represented (seeComposition of the otolith-based fish fauna”).

At Possagno, the Possagno Marls provided otoliths of 52 taxa.he association is mainly made up by a diversified assortmentf rather shallow neritic elements, but some scarce mesopelagicshes occur, and otoliths of Bregmaceros, a group of smallpipelagic fishes (between 0 and 200 m), occurs in great num-ers.

In Ukraine, Müller and Rosenberg (2003) reported a verynteresting otolith association from near Dnepropetrovsk. Withhe exception of a single Bregmaceros specimen, all otoliths arerom neritic fishes. Although the assemblage can be labelled asubtropical to warm moderate, the presence of the gadoids Rani-eps and Phycis suggests a slightly colder environment than thateported for the previous geographic areas. Data from the above-ited paper required some interpretation and revision; therefore,ll the original material in the paper of Müller and Rosenbergas examined by Nolf, whose results are presented in Table 2.

n total, we retain a list of 29 taxa from the paper of Müller andosenberg (2003). Because of the many nomenclatural changesnd improved interpretations since the publication of the listf all nominal otolith-based fossil fish species in Nolf (1985),
nd because of a number of differences between our interpre-ation and the one published by Müller and Rosenberg, somedditional remarks are required, which refer to the numbers inrackets after various taxa in Table 2:


Table 2List of the fish taxa reported from the Priabonian of Dnepropetrovsk (Ukraine) by Müller and Rosenberg (2003), with revised interpretationTableau 2Liste des taxons de poissons décrits par Müller et Rosenberg (2003) des dépôts priaboniens de Dnepropetrovsk (Ukraine), avec une interprétation révisée

Valid name Name in Müller and rosenberg (2003)

Pterothrissus umbonatus Koken, 1884 p. 364 idemAnguilla aff. rouxi (Nolf, 1977) p. 364 Anguilla sp.“genus Congridarum” thevenini (Priem, 1906) (1) p. 365 “genus Congridarum” diagonalis (1)“genus Congridarum” websteri Frost, 1933 p. 365 Muraenesox websteri“genus Ariidarum” germanicus (Koken, 1891) p. 365 idem“genus ? Harpadontidarum” hampshirensis (Schubert, 1916) p. 365 Osmerus hampshirensisAulopidae ind. p. 366 Saurida sp.Bregmaceros cf. minimus (Frost, 1934) p. 366 idemPhycis sp. p. 366 idemRaniceps latisulcatus (Koken, 1884) p. 368 Raniceps ex gr. tuberculosus (Fig. 5/20)Raniceps tuberculosus (Koken, 1884) p. 368 Raniceps ex gr. tuberculosus (Fig. 8/20-21)Hoplobrotula biscaica (Sulc, 1932) p. 369 idem“genus Neobythitinarum” aff. hilgendorfi (Koken, 1891) (2) p. 369 Aequalobythites regularis (2)Bythitidae ind. p. 370 Oligopus pseudoacuminatusBelone aff. hinsberghi (Gaemers, 1984) (3) p. 370 Hemiramphus sosimovichi (3)Gephyroberyx sp. p. 372 Gephyroberyx aff. ostiolatusCentroberyx ingens Koken, 1884 p. 372 idemMyripristidae ind. p. 372 Myripristis sp.Platycephalus sp. p. 372 idem“genus Percoideorum” andreevae Müller and Rosenberg, 2003 p. 378 idem“genus Acropomidarum” lespontinus (Nolf, 1988) (4) p. 373 Parascombrops martini (4)Chelidoperca aff. elongata Sulc, 1932 p. 372 idemEpinephelus cf. plicatus Stinton, 1978 (5) p. 378 Scombrops crenulatus (5)Apogon decoratus Stinton, 1980 p. 377 Apogon cf. decoratus

p. 374 Apogon bargi (holotype only) (6)p. 377 “genus aff. Apogon” ukrainicus (Fig. 7/17, ? 18, not 19) (7)

“genus Apogonidarum” ventrolobatus Schwarzhans, 1973 p. 377 “genus aff. Apogon” ukrainicus (Fig. 19 only) (7)Apogon aff. macrolepis Storms, 1898 p. 374 Apogon bargi (all specimens, except holotype) (6)Orthopristis bartonensis (Priem, 1884) (8) p. 373 Orthopristis goodyi (8)Cepola bartonensis Schubert, 1916 p. 374 idemTrachinus cf. biscissus Koken, 1884 p. 380 idem

Doubtful taxa and obsolate citations in Müller and Rosenberg, 2003:

• “genus Merlucciidarum” sp. (p. 368): Very eroded otolith belonging to a Merlucciid or to Gadiculus;• Epinephelus sp. (p. 373): strongly eroded juvenile percoid otoliths;• Dentex ex gr. gregarius (p. 373): not identifiable eroded juvenile sparid otoliths;•• h;• diagn

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“genus Sparidarum” sp. (p. 373): not identifiable eroded sparid otolith;“genus Sparidarum” noetlingi (p. 373): not identifiable eroded percoid otolit“genus Percoideorum” sp. (p. 380): incomplete juvenile percoid otolith, non

1) “genus Congridarum” diagonalis (Stinton and Nolf, 1970),cited in many papers on Eocene otoliths, is a junior syn-onym of “genus Congridarum” thevenini (Priem, 1906).The latter taxon was considered as doubtful by Nolf (1975:p. 208; Nolf, 1985: p. 132), but the numerous topotypesfrom Hérouval that are now available leave no doubt aboutthe identity of the holotype;

2) “genus Neobythitinarum” regularis (Priem, 1911) and“genus Neobythitinarum” hilgendorfi (Koken, 1891) aretaxa belonging to a single lineage, starting with the firsttaxon in the Early Lutetian of Belgium and ending withthe second in the Early Oligocene of the North Sea basin,the Oligocene of Castellane (Alpes de Haute-Provence) and
Aquitaine. There is a gradual change from more slenderand elongate otoliths in the Lutetian towards higher andmore robust ones in the Early Oligocene. There is alsoa gradual increase of otolith size through time, but any
(

ostic.

manner of defining a clear break between the initial andthe final morphologies is arbitrary or corresponds to theimportance of stratigraphic gaps between successive pop-ulations. Therefore, the taxon is herein considered to be asingle species, and the older name hilgendorfi is applied,but we agree that this may be interpreted as a speciescomplex by some. “Genus Neobythitinarum” boulangeriNolf, 1980 from the Early Eocene of Aquitaine is also asynonym;

3) Hemiramphus sosimovichi Müller and Rosenberg, 2003, isknown only by a single specimen that looks very much likethe otoliths of Belone hinsberghi (Gaemers, 1984), a com-mon species in the Rupelian of the Paris Basin and Belgium.
More material from Ukraine is required to decide if it isreally distinct from that species;
4) the specimens described as Parascombrops martini byMüller and Rosenberg are different from the type mate-

2 cropaléontologie 52 (2009) 195–218

(

(

(

(

dfi

osaeoctflaTsepAoaoAobd

Fig. 10. Dendrograms of Priabonian faunal sites derived from UPGMA, respec-tively based on (A) Simpson index and (B) Jaccard coefficient processing thematrices including all Priabonian nominal taxa. Similarity values are reportedon the scale bars.Fig. 10. Dendrogramme de sites priaboniens, résultant d’analyses UPGMAbasées respectivement sur (A) les indices de Simpson et (B) les coefficients Jac-card appliqués à des matrices incluant la totalité des taxa nominaux de téléostéens(otolithes) priaboniens représentés. Les valeurs de similarité obtenues sontindiquées sur les échelles.

Fig. 11. Two-dimensional ordination plot, based on Jaccard coefficient, of theselected faunal sites on the principal axes of non-metric multidimensional scal-ing. The resulting graph displays an alignment, according to the positive valuesof the second component of Ukraine, Allons, Possagno and Aquitaine in agree-ment with the cluster analysis outcome.


rial of “genus Acropomidarum” martini Gaemers and VanHinsbergh, 1978 from the Rupelian of the Netherlands.They seem to us to belong to the taxon from the Priabo-nian of Aquitaine, described by Nolf (1988) as “genusApogonidarum” lespontinus, that today we consider to bean acropomatid;

5) Scombrops crenulatus Stinton, 1980 is a junior synonymof Epinephelus plicatus Stinton, 1978, from the Lutetianof southern England. The otolith from Ukraine figured byMüller and Rosenberg may belong to the same species, butthis is difficult to judge based upon this single incompletespecimen;

6) the holotype of Apogon bargi Müller and Rosenberg, 2003seems to be referable to Apogon decoratus Stinton, 1980,but all the paratypes are much higher bodied, and apparentlybelong to a different species, probably Apogon macrolepisStorms, 1898;

7) the paratype of “genus aff. Apogon” ukrainicus, fig. 7/17 ofMüller and Rosenberg (2003), can be referred to Apogondecoratus, and probably also the holotype (fig. 7/18), arather worn eroded specimen. The paratype fig. 7/19 belongsto a different species, “genus Apogonidarum” ventrolobatusSchwarzhans, 1973;

8) the holotype of Otolithus (Percidarum) bartonensis Priem,1912 could not be traced, and the taxon could not be evalu-ated on the basis of the original iconography by Nolf (1985).A good series of topotypes from the Barton Clay, however,have allowed us to validate the taxon and to recognize itas an Orthopristis species. Orthopristis goodyi Nolf, 1975from the Bartonian of Le Guépelle in the Paris Basin is ajunior synonym of O. bartonensis.

The studied associations, together with the already publishedata, allow for a list of 139 valid marine Priabonian otolith-basedsh taxa for the above-cited basins (Appendix B - Table 3).

A palaeobiogeographical analysis was performed by meansf multivariate methods using PAST (Hammer et al., 2001)oftware. The analysis was made applying the Simpson indexnd the Jaccard coefficient to a matrix consisting of the pres-nce/absence (1/0) of 104 nominal taxa (all taxa recognizednly at family or genus level were removed) from the alreadyited geographic areas. The matrix was processed by multipleechniques consisting of both classification (cluster analyses,exible UPGMA = unweightened pair-group arithmetic aver-ging) and ordination (non-metric multidimensional scaling).he cluster analysis based on the application of both the Simp-on index and the Jaccard coefficient provides the same resultxpressed in the dendrograms (Fig. 10). In the two-dimensionallot of non-metric scaling (Jaccard analysis) (Fig. 11), Ukraine,llons, Possagno and Aquitaine are located in the positive valuesf the second coordinate, and Allons, Possagno and Aquitainere clustered again in more similar positive values of the sec-nd coordinate and the positive values of the first coordinate.
ll this, however, expresses no more than relationships basednly on presence or absence of taxa in the various cited areas,ut disregarding geographical distance, local ecological con-itions, bathymetry, etc. Another disregarded fact is that, for
Fig. 11. Représentation à deux dimensions, basée sur les coefficients Jaccard dessites sélectionnés sur les axes principaux de graduation multi-dimensionnellenon métrique. Le graphique montre un alignement des valeurs positives pourl’Ukraine, Allons, Possagno et l’Aquitaine, en accord avec l’analyse des groupe-ments.

cropaléontologie 52 (2009) 195–218 213

aaiteheeate

tnoawsCishoAeEPAal

ccPnAtff

ooohagattiDP

tEao

Fig. 12. Paleobiogeographical map of the Paleomediterranean basin during theUF(

aimtHRofgwwvboa

tssEibe

A

tSiJot


given stratum and area, a taxon which is known from oldernd younger strata but unrecorded in the analyzed associations just recorded as absent, while this absence is very rela-ive and probably due to local ecological circumstances, orven to sample size. A glance at the dendrograms of Fig. 10,owever, shows clearly a western Atlantic (Louisiana) and anastern Atlantic–Paleomediterranean association. In the east-rn Atlantic–Paleomediterranean association, the groupings,lthough purely based on absence/presence, seem to be consis-ent with other factors, such as geographical position and localnvironmental conditions.

The western Atlantic association, identified as the most dis-inct one in the dendrograms, comprises 43 taxa of which 27 areominal species. Among those, 22 are exclusively American andnly five occurs in the eastern Atlantic–Paleomediterranean Pri-bonian. At generic level, 22 Recent genera are represented, ofhich 17 are also known from the European Priabonian (or from

lightly older or younger deposits). Among the remaining five,entropristis, Anisotremus, and Citharichthys are typical Amer-

can genera, although in the Recent fauna, one Citharichthyspecies dispersed into the West African shores. The genus Nibeaas a present day Indo-west Pacific distribution, and Urocongerccurs on both sides of the Atlantic and in the Indian Ocean.bove generic level, all taxa have an Eocene European record,

xcept the sciaenids, which only seems to appear there after theocene and the paralichthyids, a typical American and Indo-acific group of flatfishes. The difference between the westerntlantic association and the European ones is the more accentu-

ted because the first one is entirely devoid of oceanic elementsike mesopelagic and deep-water fishes.

In the eastern Atlantic–Paleomediterranean associations,lustering seems in the first instance a function of the geographi-al distance between the sites. The geographically closest places,ossagno (51 taxa, 35 nominal species) and Allons (25 taxa, 19ominal species), share 21 taxa, of which 18 at species level.llons, the association with the lowest diversity, has 84% of its

axa and 95% of its species in common with Possagno; the dif-erence between both associations is essentially due to a higheraunal diversity at Possagno.

The Aquitaine association (69 taxa, 47 nominal species) notnly differs from the Possagno–Allons association in functionf its more distant geographical position but also by its strongerceanic character in the southern part of the basin (e.g., the ratherigh diversity of congrid eels, sternoptychids and myctophids,nd the presence of deep-water genera like Polymixia, Bathy-adus, Beryx Epigonus and Bembrops) and by the occurrence ofvery peculiar shallow water facies, the Saint Estèphe Forma-

ion, characterized by abundant small echinoids in the north ofhe Basin. This deposit has a peculiar otolith association, whichs dominated by the apogonid Fowleria, and contains taxa likeipulus and Glaucosoma which are known nowhere else in theriabonian.

The Ukraine association (29 taxa, 23 nominal species), iden-
ified in the dendrogram as the most different one from otheruropean associations, is also the geographically most distantnd most northern one. Only nine of the 23 nominal speciesccur in the Possagno–Allons–Aquitaine area, but this is partly
Rbrc

pper Eocene (Priabonian) (modified from Rögl, 1999).ig. 12. Paléogéographie du basin Paléoméditerranéen à l’Eocène supérieurPriabonien) (d’après Rögl, 1999, modifié).

distorted picture because all 14 of the remaining species occurn slightly older or younger west European strata. Part of this

ay be due to local environmental conditions in Ukraine, ando differences in sampling intensity at all our Priabonian sites.owever, one thing is remarkable: five (Anguilla aff. rouxi,aniceps tuberculosus, Belone aff. hinsberghi, “genus Percoide-rum” andreevae, and Trachinus cf. biscissus) are known onlyrom Oligocene west European sites, and the same is true for theenus Phycis. Moreover, the total number of Ukrainian specieshich have an Oligocene record in other European sites is 13,hich constitute 57% of the nominal species. This situation isery different from the Aquitaine one, where a strong faunalreak characterizes the Eocene–Oligocene transition (only 9%f all Priabonian and Early Oligocene species cross the bound-ry, see Nolf, 1988: fig. 8, p. 106).

The northern geographic location of Ukraine, combined withhe good connections to both the North Sea Basin and the Turgaitreet (Fig. 12) can provide the explanation. Many Oligocenepecies (or their close relatives) probably already existed atocene times in more northern regions, but could penetrate only

n more southern European seas since the strong cooling at theeginning of the Oligocene (Salamy and Zachos, 1999; Zachost al., 1996).

cknowledgements

It’s a pleasure for us to thank Giampetro Braga for guiding uso various less known exposures around Possagno and to Etienneteurbaut, who assisted with a first exploration and sampling

n both the Possagno and Allons area. We also thank Diane andames Tyler for a critical reading and correction of a first draftf our manuscript and Anne Wauters who helped in preparinghe otolith drawings. Careful reviews from G. Stringer and
. Brzobohaty are greatly acknowledged. Angela Gironeenefited from a grant of the project “Synthesis of systematicesources” of the European community for studying the IRSNBollections.

2 cropa

A

M

8

8

8

P

I

C8

C

(

B

B

8

8

pP

(

P

8

P

8

s

BBB

s

s

p

s

s

DDc

P

8

s

s

s

s

(

AA8

(

I

II

8

P

(

8

P

(

(

p

(P


ppendix A. : repository of the figured specimens.

Institutional abbreviations:IRSN = Institut royal des sciences naturelles de Belgique;BMNH = The natural History Museum (formerly British

useum [Natural History]);Apogon decoratus, Fig. 5F: Possagno, sample C (IRSNB P

345);Apogon macrolepis, Fig. 5K: Possagno, sample C (IRSNB P

351–8353);Apogon sp., Fig. 4H: Possagno, Monte Collalto (IRSNB P

328);Ariosoma nonsector, Fig. 2E: Possagno, sample C (IRSNB

8269–8271);Aseraggodes kaianus, Fig. 8A: Recent, NW Australia (coll.

RSNB);Aseraggodes laganum n. sp. Fig. 8B: Possagno, sample

, B1 = holotype (IRSNB P 8366), B2 = paratype (IRSNB P367);

Aspidophoroides olriki, Fig. 6C: Recent, (loc. unknown, coll.haine in IRSNB);

Bregmaceros brihandensis, Fig. 3B: Possagno, sample CIRSNB P 8288–8290);

Centroberyx ingens, Fig. 4B: B1 = Allons B (IRSNB P 8313),2 = Possagno, sample C (IRSNB P 8314);

Cepola bartonensis, Fig. 6F: Bartonian, Barton Clay, loc.arton, (F 1–4 = BMNH P 23570, F 5 = BMNH P 14426);

Cepola yrieuensis, Fig. 5E: Possagno, sample C (IRSNB P342–8344);

Cepola yrieuensis, Fig. 6E: Possagno, sample B (IRSNB P358–8359);

Champsodon spinosus, Fig. 5D: D1 and D3 = Possagno, sam-le B (IRSNB P 8339–8340, D2 = Possagno, sample A (IRSNB8341);Chelidoperca elongata, Fig. 4A: Possagno, sample C

IRSNB P 8310–8312);Citharus cauneillensis, Fig. 5C: Possagno, sample C (IRSNB

8335–8338);Citharus sp., Fig. 5A: Possagno, sample B (IRSNB P 8331);Conger glaber, Fig. 2F: Possagno, sample C (IRSNB P

272);Conger sp., Fig. 2H: H1 = Possagno, sample B (IRSNB

8277), H2 = Possagno, sample C (IRSNB P 8278);Diaphus excavatus, Fig. 3A: Possagno, sample C (IRSNB P

286–8287);“genus Agonidarum” sudans n. sp., Fig. 6D: holotype, Pos-

agno, sample C (IRSNB P 8357);“genus Alepocephalidarum” astrictus n. sp., Fig. 6B:

1 = holotype, Possagno, sample C (IRSNB P 8354),2 = paratype, Possagno, sample B (IRSNB P 8355),3 = paratype, Possagno, sample C (IRSNB P 8356);

“genus Antennariidarum” aff. furcatus, Fig. 3C: Possagno,ample C (IRSNB P 8291–8293);

“genus Apogonidarum” ventrolobatus, Fig. 5B: Possagno,ample C (IRSNB P8332–8334);

“genus Congridarum” eocaenicus, Fig. 2G: Possagno, sam-le C (IRSNB P 8273–8276);

P

(

léontologie 52 (2009) 195–218

“genus aff. Glyptophidium” biarritzensis, Fig. 4E: Possagno,ample C (IRSNB P 8318–8320);

“genus Heterenchelyidarum” aff. colei, Fig. 2A: Possagno,ample C (IRSNB P 8261–8264);

“genus Lophiiformorum” canovae n. sp., Fig. 7D:1 = paratype, Possagno, sample C (IRSNB P 8364),2 = holotype, from loose block from stratigraphic layer very

lose to sample C (IRSNB P 8365);“genus aff. Zenion” sp., Fig. 4D: Possagno, sample B (IRSNB

8317);“genus Myripristinarum” sp., Fig. 5J: Allons B (IRSNB P

350);“genus Neobythitinarum” hilgendorfi, Fig. 4F: Possagno,

ample C (IRSNB P 8321–8326);“genus Percoideorum” aff. aquitanicus, Fig. 4G: Possagno,

ample C (IRSNB P 8327);“genus Sirembinorum” aff. arcuatus, Fig. 3H: Possagno,

ample C (IRSNB P 8303–8304);“genus Synodontidarum” cf. boulangeri, Fig. 2K: Possagno,

ample C. (IRSNB P 8284);“genus Triglidarum” vectensis, Fig. 3E: Possagno, sample D.

IRSNB P 8299);“genus Uranoscopidarum” cochlearis n. sp., Fig. 7A:

1 = holotype, Possagno, sample B (IRSNB P 8360),2-4 = paratypes, Possagno, sample C (IRSNB P 8361–363);

Genyagnus monopterygius, Fig. 7B: Recent, New Zealandcoll. IRSNB);

Gnathagnus innotabilis, Fig. 7C: Recent, New Zealand (coll.RSNB);

Grammonus pseudoacuminatus, Fig. 3I: I1 and3 = Possagno, sample C (IRSNB P 8305-8306), I2 and4 = Allons B (IRSNB P 8307-8308);

Haemulidae ind., Fig. 5H: Possagno, sample C (IRSNB P347);

Hoplobrotula biscaica, Fig. 3D: Possagno, sample C (IRSNB8294–8298);Hoplunnis ariejansseni, Fig. 2B: Possagno, sample C

IRSNB P 8265–8266);Microchirus sp., Fig. 5G: Possagno, sample C (IRSNB P

346);Onuxodon aff. coheni, Fig. 3G: Possagno, sample C (IRSNB

8302);Opisthoproctus weitzmani, Fig. 2C: Possagno, sample C

IRSNB P 8267);Photostylus pycnopterus, Fig. 6A: Recent, New Caledonia

coll. IRSNB);Platycephalus janeti (Priem, 1911), Fig. 5I = Possagno, sam-

le C (IRSNB P 8348–8349);Pristigenys aff. obliquus, Fig. 4C: C1 = Possagno, sample A

IRSNB P 8315), C2 = Possagno, sample C sample A (IRSNB8316);Pseudophichthys sp., Fig. 2D: Possagno, sample C (IRSNB

8268);Rhynchoconger transversus, Fig. 2I: Possagno, sample C

IRSNB P 8279–8282);Saurida sp., Fig. 2L: Allons B (IRSNB P 8285);

cropa

P

8

(

A

List of upper Eocene otolith-based teleost taxa recorded

TLtTLs

A

A

H

M

C

N

C

A

O

A

S

P

S

SM

PB

M

P

G

C


Scopelarchidae ind., Fig. 2J: Possagno, sample B (IRSNB8283);

Sparidae ind., Fig. 4I: Allons B (IRSNB P 8329-8330);
Valenciennellus brzobohatyi, Fig. 3J: Allons B (IRSNB P
309);Vincinguerria biarritzensis, Fig. 3F: Possagno, sample C

IRSNB P 8300–8301).

fa(

able 3ist of Upper Eocene otolith-based fish taxa. Taxa in bold: taxa disappeared from the m

o Indo-Pacific areasableau 3iste des taxons de poissons de l’Eocène supérieur basée sur les otolithes. Taxons enoulignés = taxons limités aujourd’hui aux régions indo-pacifiques

Stratigraphic range

Lower

Eocene

Middle

Eocene

Upper

Eocene

lbulidae Albula sp. UE

Pterothrissus umbonatus (Koken, 1884) UE

Pterothrissus sp. UE

nguillidae Anguilla aff. rouxi (Nolf, 1977) UE

Anguilla sp. UE

eterenchelyidae “genus Heterenchelyidarum” colei (Muller, 1999) ME UE

“genus Heterenchelyidarum” richardsi Nolf, 1988 UE

uraenesocidae Muraenesox sp. UE

ongridae Ariosoma nonsector Nolf and Stringer, 2003 UE

Conger glaber Brzobohaty, 1967 UE

Conger vetustus Frizzel and Lamber, 1962 ME UE

Conger sp. UE

Gnathophis dissimilis (Frizzel and Lamber, 1962) ME UE

Paraconger sauvagei (Priem, 1906) ME UE

Paraconger sector (Koken, 1888) UE

Paraconger yazooensis Nolf and Stringer, 2003 UE

Pseudophichthys elongatus (Sulc, 1932) UE

Pseudophichthys glaber (Koken, 1888) UE

Pseudophichthys sp. UE

Rhynchoconger transversus (Sulc, 1932) ME UE

Rhynchoconger sp. UE

Rhechias sp. UE

“genus” Congridarum” biarritzensis Nolf, 1988 UE

“genus Congridarum” eocaenicus (Sulc, 1932) LE UE

“genus Congridarum” thevenini (Priem, 1906) LE UE

“genus Congridarum” websteri Frost, 1933 LE ME UE

“genus aff. Uroconger” sp. UE

ettastomatidae Hoplunnis ariejansseni Nolf, 1988 LE UE

lupeidae Clupeidae ind. UE

riidae Arius cf. crassus (Koken, 1884) UE

“genus Ariidarum” germanicus (Koken, 1891) UE

Ariidae ind. UE

pisthoproctidae Opisthoproctus weitzmani Nolf, 1988 UE

lepocephalidae “genus Alepocephalidarum” astrictus n. sp. UE

ternoptychidae Danaphos gibbsi Nolf, 1988 UE

Valenciennellus brzobohatyi Steurbaut, 1982 UE

Valenciennellus reniculus Nolf, 1988 UE

“genus Sternoptychidarum” sp. UE

hosichthyidae Vinciguerria biarritzensis (Sulc, 1932) UE

ynodontidae “genus Synodontidarum” boulangeri Nolf, 1988 UE

“g. ? Harpadontid.” hampshirensis (Schubert, 1916) UE

Synodus sp. UE

Saurida sp. UE

copelarchidae Scopelarchidae ind. UE

yctophidae Diaphus excavatus (Sulc, 1932) UE

Diaphus quadrangularis (Sulc, 1932) LE UE

Diaphus sulci Nolf, 1988 LE UE

olymixiidae Polymixia sp. UE

regmacerotidae Bregmaceros brihandensis Nolf, 1988 UE

Bregmaceros cf. minimus (Frost, 1934) UE

Bregmaceros sp. UE

acrouridae Bathygadus mauli Nolf, 1988 UE

hycidae Phycis sp. UE

adidae Raniceps latisulcatus (Koken, 1884) UE

Raniceps tuberculosus (Koken, 1884) UE

arapidae Onuxodon aff. coheni Nolf, 1980 ? Pr

“genus Carapinarum” novus (Sulc, 1932) UE

Carapidae ind. UE

léontologie 52 (2009) 195–218 215

ppendix B

rom Possagno, Allons, Aquitaine, Louisiana and Ukrainend their stratigraphic and geographic distributionTable 3).

odern Mediterranean basin. Taxa in bold and underlined: taxa today restricted

gras = taxons disparus dans le bassin méditerranéen actuel. Taxons en gras et

Priabonian localities

Lower

Oligocene

Upper

Oligocene

Miocene Possagno Allons Aquitaine Louisiana Ukraine

L

LO UO M Aq U

L

LO U

Aq

? P ? A L

Aq

Aq

P L

LO P

L

P

L

LO Aq

L

L

LO UO M Aq

L

P

LO P A Aq

L

Aq

Aq

P A Aq

Aq ?L U

U

L

P

P A Aq

Aq

U

L

UO P Aq

P

LO Aq

LO UO M A Aq

Aq

Aq

P ? A Aq

? P Aq

U

L

A

P

P A Aq

Aq

Aq

Aq

P A Aq

U

L

Aq

U

LO U

LO U

LO P

Aq

L


Table 3 ( Continued )Stratigraphic range Priabonian localities

Lower

Eocene

Middle

Eocene

Upper

Eocene

Lower

Oligocene

Upper

Oligocene


Ophidiidae Hoplobrotula biscaica (Sulc, 1932) UE P A Aq U

Brotula aquitanica Nolf, 1980 LE UE L

Glyptophidium polli (Casier, 1946) LE UE Aq

“genus aff. Glyptophidium” biarritzensis (Sulc,

1932)

UE P A Aq

“genus Neobythitinarum” hilgendorfi (Koken, 1891) UE LO P A ?U

“genus Neobythitinarum” meyeri (Koken, 1888) UE L

“genus Neobythitinarum” aequaloides N. and S.,

2003

UE L

“genus Sirembinorum” ringeadei Nolf, 1980 UE Aq

“genus Sirembinorum” arcuatus (Stinton, 1986) LE ME ? UE ? LO P

“genus Sirembinorum” granus Muller, 1999 ME UE L

Bythitidae Grammonus pseudoacuminatus (Sulc, 1932) ME UE ? UO P A Aq

Dipulus bliecki Nolf, 1988 UE Aq

Bythitidae ind. UE U

Batrachoididae Batrachoididae ind. UE Aq

Lophiidae “genus Lophiiformorum” canovae n. sp. UE P

Antennariidae “genus Antennariidarum” furcatus (Frost, 1933) ME ? UE P

Antennarius sp. UE Aq

Chaunacidae Chaunax sp. UE Aq

Ogcocephalidae Ogcocephalus cirrhosus Stinton, 1978 ME UE Aq

Atherinidae “genus Atherinidarum” debilis (Koken, 1888) UE L

Belonidae Belone aff. hinsberghi (Gaemers, 1984) UE LO U

Cyprinodontidae Cyprinodontidae ind. UE P

Trachichthydae Gephyroberyx sp. UE U

Berycidae Centroberyx ingens (Koken, 1884) UE LO P A U

Centroberyx sp. UE Aq L

Beryx sp. UE Aq

Holocentridae “genus Myripristinarum” sp. LE UE A Aq U

“genus Myripristinarum” cajun (Frizz. and Lam.,

1961)

ME UE L

Zeniontidae “genus aff. Zenion” sp. UE P

Caproidae Antigonia sp. LE UE Aq L

Triglidae “genus Triglidarum” vectensis Nolf, 1973 L UE ? P

Platycephalidae Platycephalus janeti (Priem, 1911) LE ME UE P Aq

Platycephalus sp. UE U

Agonidae “genus Agonidarum” sudans n. sp. UE P

Percoidei inc. sedis “genus Percoideorum” pseudolestidiops Nolf, 1988 UE Aq

“genus Percoideorum” sp. 3 UE Aq

“genus Percoideorum” aquitanicus Nolf, 1988 UE ? P ? A Aq

“g. Percoideorum” andreevae Mull. and Rosen.,

2003

UE LO U

Ambassidae Ambassis meldertensis (Nolf, 1973) ME UE Aq

Acropomatidae Parascombrops brzobohatyi Nolf, 1988 UE Aq

“genus Acropomidarum” lespontinus (Nolf, 1988) UE Aq U

Serranidae Chelidoperca elongata (Sulc, 1932) UE P A Aq ? U

Centropristis priaboniana Nolf and Stringer, 2003 UE L

Epinephelus cf. plicatus Stinton, 1980 ME ? UE U

Priacanthidae Pristigenys obliquus Nolf and Stringer, 2003 UE ? P L

Pristigenys aff. caduca Nolf, 1973 ME UE Aq

Apogonidae Fowleria blayensis (Nolf, 1988) UE Aq

Apogon decoratus Stinton, 1980 UE P U

Apogon macrolepis Storms, 1898 ME UE P ? A Aq ? U

“genus Apogonidarum” ventrolobatus Schw., 1973 UE LO P A U

Epigonidae Epigonus polli Nolf, 1988 UE Aq

Malacanthidae “genus Malacanthidarum” sulcatus (Koken, 1888) UE L

“genus Malacanthidarum” cadenati Steurbaut, 1984 UE LO Aq

Menidae Mene sp. UE L

Haemulidae Anisotremus sp. UE L

Orthopristis americana (Koken, 1888) UE LO L

Orthopristis bartoniensis (Priem, 1884) UE U

Orthopristis kokeni (Leriche, 1905) ME UE Aq

“genus Haemulidarum” obliquus (Muller, 1999) ME UE L

“genus Haemulidarum” aff. pulcher (Frost, 1934) ME UE Aq

Haemulidae ind. UE P

Sparidae “genus Sparidarum” elegantulus (Koken, 1888) UE L

Sparidae ind. UE A

Dentex sp. UE Aq

Sciaenidae “genus aff. Nibea” sp. UE L

Sciaena aff. pseudoradians (Dante and Frizzel,

1965)

UE L

“genus Sciaenidarum” claybornensis Koken, 1888 ME UE L

Glaucosomatidae Glaucosoma sp. UE Aq

Bathyclupeidae Bathyclupea arribaoutensis Nolf, 1988 UE Aq

Cepolidae Cepola yrieuensis Steurbaut, 1984 UE LO P A

Cepola bartoniensis Schubert, 1916 ME UE U

Owstonia brihandensis Nolf, 1988 UE Aq

“genus Cepolidarum” longissimus Nolf, 1988 UE Aq


Table 3 ( Continued )Stratigraphic range Priabonian localities

Lower

Eocene

Middle

Eocene

Upper

Eocene

Lower

Oligocene

Upper

Oligocene


Labridae Labridae ind. UE P

Scaridae Sparisoma sp. UE Aq

Trachinoidei inc. sedis “genus Trachinoideorum” schwarzhansi Nolf, 1988 UE Aq

Champsodontidae Champsodon spinosus Schwarzhans, 1977 UE LO P A

Percophidae Bembrops sp. UE Aq

Ammodytidae “genus Ammodytidarum” sp. UE Aq

Trachinidae “genus Trachinidarum” laevigatus (Koken, 1888) UE L

Trachinus cf. biscissus Koken, 1884 UE LO U

Uranoscopidae “genus Uranoscopidarum” cochlearis n. sp. UE P

Blenniidae “genus Blenniidarum” cor (Koken, 1888) UE L

“genus Blenniidarum” sp. UE Aq

Gobiidae “genus Gobiidarum” vetustus Nolf and Stringer,

2003

UE L

Scombridae Scombridae ind. UE P A L

Psettodidae Psettodes sp. UE L

Citharidae Citharus cauneillensis Nolf, 1988 UE P A Aq

Citharus sp. UE P

“genus Citharidarum” hoffmani Nolf and Stringer,

2003

UE L

Paralichthyidae Citharichthys altissimus Nolf and Stringer, 2003 UE L

Pleuronectidae “genus Pleuronectidarum” sp. UE Aq

B E

S E

E

R

B

B

B

B

C

C

G

H

H

M

M

NN

N

N

N

N

N

N

N

N

N

N

N

N

Geological Pamphlet 13, 1–23.Proto Decima, F., Roth, P.H., Todesco, L., 1975. Nannoplancton Calcareo del

Paleocene e dell’Eocene della Sezione di Possagno. In: Bolli, H. (Ed.),

othidae “genus Bothidarum” sp. U

oleidae Aseraggodes laganum n. sp. U

Microchirus sp. U

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