Louis Taverne ¹, Dirk Nolf ² & Annelise Folie ²

¹ Université Libre de Bruxelles (U. L. B.), Faculté des Sciences, Département de Biologie des Organismes, Unité de Morphologie fonctionnelle (C. P. 160/11), avenue F. D. Roosevelt 50, B-1050 Brussels, Belgium

² Royal Institute of Natural Sciences of Belgium, Department of Paleontology, Section of Vertebrate Paleontology, Vautier street 29, B-1000 Brussels, Belgium

Corresponding author : Louis Taverne, e-mail : louis.taverne@gmail.com

ABSTRACT. Some bony remains, otoliths and squamules belonging to the osteoglossid genus Scleropages are described from the continental Paleocene of Hainin (Mons Basin, Belgium). The hypotheses to explain the occurrence of such a freshwater fish in Europe at that time are discussed.

KEY WORDS : Teleostei, Osteoglossidae, Scleropages, fossil remains, continental Paleocene, Belgium.

RÉSUMÉ. Quelques restes osseux, des otolithes et des squamules appartenant à l’ostéoglossidé Scleropages sont décrits dans le Paléocène continental de Hainin (Bassin de Mons, Belgique). Les hypothèses expliquant la présence de ce poisson dulçaquicole en Europe à cette époque sont discutées.

MOTS-CLÉS : Teleostei, Osteoglossidae, Scleropages, restes fossiles, Paléocène continental, Belgique.

Continental deposits of Paleocene age, overlying the marine Calcaire de Mons (Danian), were described by RUTOT (1886) from the railway section at Hainin, about 13km West of Mons. In 1970, a test hole (HA 1) was drilled (at x = 107.250, y = 124.890) by the Geological Laboratory of the “Faculté Polytechnique de Mons” at the municipal football field of Hainin, about 600m North of the railway section, with the aim to realize a complete section through the lacustrine continental deposits (observed between 8.5m and 38.4m depth). Another test hole, HA 2 (at x = 107.045, y = 124.990) was drilled, some 250m North West of HA 1, with the intention to explore the underlying marine Danian deposits (GODFRIAUX & ROBASZYNSKI, 1974).

According to STEURBAUT (1998 : 144-145), the concerned “continental Montian” deposits, now called Hainin Formation, represents the lowstand systems tract of sequence TP 1.4, which corresponds to the oldest Seelandian of Europe (about 60 million years old).

Sediments from the HA 1 hole provided the oldest mammal teeth from the European Cenozoic (GODFRIAUX & THALER, 1972; VIANEY-LIAUD, 1979; SUDRE & RUSSEL, 1982; CROCHET & SIGÉ, 1983; SIGÉ & MARANDAT, 1997). This fauna has been chosen as reference-level MP1-5 of the mammalian biochronological scale for the European Paleogene (SCHMIDT-KITTLER, 1987; AGUILAR et al., 1997). The “Montian” fauna of Hainin also contains fossil amphibians (GROESSENS-VAN DYCK, 1981), turtles (GROESSENS-VAN DYCK, 1984; SCHLEICH et al., 1988), crocodiles (GROESSENS-VAN DYCK, 1986), lizards (FOLIE et al., 2005) and many fish remains. Additional fossils were obtained in 1974, from a hole with a diameter of 1m, known as “Puits Hainin”, located at 25m North of the mentioned exploration drilling. This hole was burrowed till a depth of 25m and provided the bulk of the now available Hainin fossils.

In 1994, MAGIONCALDA published additional data on three new boreholes made at Hainin in 1986 and located in the area immediately East of the HA 1 hole. He mentioned otoliths of osmerids (?), Thaumaturus Reuss, 1844, percoids, umbrids and esocids, based on otolith identification by Nolf. The list of Magioncalda does not mention osteoglossids, because his work did not considered the “Puits Hainin”, from where a small sample examined by Nolf about 20 years ago yielded osteoglossid otoliths, identified in the collection as Scleropages Günther, 1864 but never published until now.

Osteoglossiformes are one order of the primitive teleostean super-order Osteoglossomorpha, commonly known as the “bony tongue” fishes by reason of their peculiar bite between parasphenoid, pterygoid bones, dermobasihyal and dermobasibranchials. Osteoglossiformes are heavily ossified teleosts, with a medio-parietal skull, very large posterior infraorbitals, a more or less prognathous lower jaw, and generally large acute teeth on both jaws and a reduced or lost subopercle (TAVERNE, 1977; 1978). Their scales frequently offer a reticular pattern formed by small bony plates, the squamules, separated from each other by very narrow grooves and resting on the basal calcified collagenous layer of the scale (GAYET & MEUNIER, 1983; MEUNIER, 1984).

Osteoglossiformes appear in the fossil record as soon as the Early Cretaceous with species such Laelichthys ancestralis Silva-Santos, 1985 from Brazil, Chanopsis lombardi Casier, 1961 from the Democratic Republic of Congo and Nieerkunia liae Su, 1992 and Xixiaichthys tongxinensis Zhang, 2004 both from the South East of China. Most fossil and all Recent Osteoglossiformes are freshwater fishes but there are also a few fossil marine species in the Paleocene and Eocene of Europe, North America, Central Asia and Africa. “Genus Osteoglossidarum” tavernei Nolf & Stringer, 1996 from the Maastrichtian of Mississipi is the first known marine occurrence for the order. Recent freshwater Osteoglossiformes generally tolerate brackish waters and sometimes enter in marine waters near the estuaries.

The fossil record of the suborder Osteoglossoidei shows a worldwide distribution, essentially because of the wide expansion of marine species during the Lower Paleogene. Recent species offer a remarkable example of Gondwanan distribution with Osteoglossum Cuvier, 1829 and Arapaima Müller, 1843 in tropical South America, Heterotis Rüppel, 1829 and Pantodon Peters, 1877 in Africa and Scleropages in Australia, New Guinea and Southeastern Asia. Such a Gondwanan distribution already occurs for the oldest osteoglossoid species in the continental Lower Cretaceous deposits.

Recent Osteoglossoidei are moderately large fishes with the exception of the small Pantodon reaching only a few centimetres. Arapaima, the giant of the group, grows to about 3 or 4 metres in length. Those Recent genera are grouped in three families, Osteoglossidae with Osteoglossum and Scleropages, Arapaimidae (= Heterotidae) with Heterotis and Arapaima and Pantodontidae for the sole Pantodon (Taverne, 1979). Some consider Arapaimidae as a simple subfamily of Osteoglossidae and also include in this family the fossil subfamily Phareodontinae (NELSON, 2006 : 104-106).

In the present paper, we describe fossil fish remains belonging to Scleropages from the continental Paleocene of Hainin in the Mons Basin (Belgium) and we discuss the occurrence of such an osteoglossid genus in those geological layers of that Belgian locality.

The present day distribution of Scleropages is restricted to Southeastern Asia, Australia and New Guinea, with a total of six species : Scleropages formosus (Müller & Schlegel, 1844) or Asian arowana (Fig. 1) from Indonesia, Malaysia, Thailand and Cambodia, Scleropages leichardti Günther, 1864 or spotted barramundi from Eastern Australia, Scleropages jardinii (Saville-Kent, 1892) or northern barramundi from Southern New Guinea and Northern Australia, Scleropages aureus Pouyaud et al., 2003 from the Siak river in Sumatra, Scleropages legendrei Pouyaud et al., 2003 from West Borneo and Scleropages macrocephalus Pouyaud et al., 2003 from Borneo too (POUYAUD et al., 2003). Scleropages is known as a fossil in the Neogene of Queensland, Australia (HILLS, 1943) and in the Eocene of Sumatra where it is contemporaneous with Musperia radiata (Heer, 1874), another fossil osteoglossid from Sumatra (SANDERS, 1934; TAVERNE, 1979). “Genus Osteoglossidarum” deccanensis Rana, 1988 and “genus Osteoglossidarum” intertraptus Rana, 1988, both from the Maastrichtian “Deccan Intertrappean Beds” of India (RANA, 1988), can also confidently be attributed to Scleropages or to a closely related genus and provides the oldest fossil record of the taxon. KUMAR et al. (2005) further mention the presence of an osteoglossid skull, Taverneichthys bikanericus Kumar et al., 2005, that seems close to Osteoglossum and Scleropages, in the continental Paleocene of Rajasthan, India.

Recent Osteoglossiformes no more exist in Europe today but Scleropages is not the unique fossil osteoglossiform fish on that continent. However, most other European fossil Osteoglossiformes are marine teleosts. They include Brychaetus muelleri Woodward, 1901, “genus Osteoglossidarum” rhomboidalis (Stinton, 1977) and “genus Arapaimidarum” acutangulus (Stinton, 1977), all three from the Lower or Middle Eocene of the southern North Sea Basin, Monopteros gigas Volta, 1796, Thrissopterus catullii Heckel, 1856 and Foreyichthys bolcensis, Taverne 1979, all three from the lowermost Lutetian of Monte Bolca, and four still unpublished new osteoglossoid genera in the Eocene of Denmark (BONDE, 1966, 1987, in press; TAVERNE, 1998).

Fig. 1. – Scleropages formosus (Müller & Schlegel, 1844) from Southeast Asia (modified from TAVERNE, 1977 : fig. 70). This species can grow to about 90cm in length.

IRSNB P 8238 : A right premaxilla, Hainin, at 1717.7m depth (level R1).

IRSNB P 8239 : An incomplete right maxilla, Hainin, at 17-17.7m depth (level R1).

IRSNB P 8240 : A fragment of a right entopterygoid, Hainin, at 13.7-14m depth (level N2).

IRSNB P 8241 : A fragment of a left palato-ectopterygoid, Hainin, at 13.3-13.7m depth (level N1).

IRSNB P 8242 : An abdominal vertebra, Hainin, at 1414.6m depth (level O1).

IRSNB P 8243 : A left saccular otolith, Hainin, precise depth unknown, and a fragmentary saccular otolith, Hainin, at 17-17.7m depth (level R1).

IRSNB P 8244-8251 : Squamules, Hainin, at 13.3-13. m (level N1), 13.7-14m (level N2), 16-16.6m (level Q1), 16.6-17m (level Q2), 17-17.7m (level R1) and 17.7-18m depth (level R2).

Levels N1, N2 and O1 are grey clays with calcareous concretions broken by peaty ruptures, while levels Q1, Q2, R1 and R2 are rough tufaceous grey marls (GODFRIAUX & ROBASZYNSKI, 1974).

The material has been studied with a Wild M5 stereomicroscope supplied with a camera lucida used for the drawings.

Order OSTEOGLOSSIFORMES Regan, 1909

Suborder OSTEOGLOSSOIDEI Regan, 1909

The discovery of squamules ornate with granulations in the Hainin continental Paleocene deposits attests the presence there of a fossil osteoglossoid fish. The African osteoglossomorph family Mormyridae also possesses scales with squamules but their ornamentation is formed by normal circuli and not by granulations as in Osteoglossoidei.

Fig. 3. – Scleropages sp. from the Paleocene of Hainin. (A) Fragment of a right entopterygoid (IRSNB P 8240) in external (A1) and internal view (A2). (B) Schematic drawing of the right entopterygoid and palato-ectopterygoid of Scleropages in internal view to show the location of the Hainin pterygoid fragments (IRSNB P 8241). (C) Fragment of a left ectopterygoid in external (C1) and internal view (C2). Legend : cr. : crest on the external face of the ectopterygoid; g. ps. : groove on the external face of the entopterygoid to receive the basipterygoid process of the parasphenoid.

Fig. 4. – (A) Left sagitta (IRSNB P 8243) of the Scleropages sp. from the Paleocene of Hainin, in dorsal (A1), profile (A2) and external view (A3). (B) Left sagitta of Scleropages formosus (Müller & Schlegel, 1844) in external view (modified from TAVERNE, 1977 : fig. 81). (C) Left sagitta of Scleropages leichardti Günther, 1864 in external view (modified from NOLF, 1993 : Fig. 4A).

Fig. 5. – Scleropages sp. from the Paleocene of Hainin. Squamules (IRSNB P 8244-8251) of the posterior field of the scale in external (A1, B1, C1, E1, G1) and internal view (A2, B2, C2, E2, G2). Three united squamules from the lateral field in external (D1) and internal view (D2), with enlargement of (D2) in (D3) and (D4). Two united squamules of the anterior field of the scale in external (F1) and internal view (F2). A squamule of the anterior field of the scale in external (H1) and internal view.

A deep groove is present at the top of the external face process of the parasphenoid (BRIDGE, 1895 : pl. 22, fig. 4) of the entopterygoid in the Hainin fossil fish. Such a which is firmly fixed there by an elongated anterior pro-groove is a unique autapomorphy of the osteoglossoid cess of the hyomandibula, except in Pantodon where that families Osteoglossidae, Arapaimidae and Pantodontidae hyomandibular process is missing. That very peculiar (TAVERNE, 1977 : Figs 53, 54, 82, 83, 107, 134; 1978 : structure allows a supplementary articulation between the Figs 40, 41). No other teleost offers such a specialized neurocranium and the hyopalatine arch that strengthens structure. That groove receives the paired basipterygoid the palato-lingual bite in those fishes.

The pattern of the dentition on the inner face of the entopterygoid, with one dorsal row of large teeth and a ventral area with much smaller and irregularly arranged teeth, agrees with a belonging to Osteoglossidae (TAVERNE, 1977 : Figs 54, 83) but not to Arapaimidae and Pantodontinae. Indeed, there are only some large teeth on the upper part of the entopterygoid in Heterotis and Pantodon (TAVERNE, 1977 : Fig. 107, 1978 : Fig. 41) and very small denticles on practically the whole inner surface of the bone in Arapaima (TAVERNE, 1977 : Fig. 134).

Fossil and Recent Osteoglossidae offer premaxillae with a well developed symphyseal ascending process (TAVERNE, 1977 : Figs 42, 47, 49, 1978 : Figs 2, 7, 20; GAYET, 1991 : Fig. 7a; LI, 1996 : Fig. 3; LI et al., 1997 : Figs 3, 8). In Scleropages however, this symphyseal process is very short (TAVERNE, 1977 : Fig. 71). The premaxilla of the Hainin osteoglossid, with its very short symphyseal process, completely looks like the corresponding bone of Scleropages and more particularly to the one of the Recent Asian Scleropages formosus which also possesses ten or eleven premaxillar teeth. In the Recent Scleropages leichardti from Australia, there are only three to five teeth on the premaxilla.

The identification of the Hainin fossil fish with the genus Scleropages is further confirmed by its saccular otoliths that look very much like those of Scleropages leichardti (NOLF, 1993 : fig. 4A). It differs only from them by a wider sulcus and a thicker posterior portion. Otoliths of Sleropages formosus (Fig. 4B2) are more elongate and have a narrower sulcus.

The Scleropages from Hainin is neither the first mentioned, nor the oldest Osteoglossidae present in the European continental deposits. A fragment of a toothed jaw bone and some squamules of an undetermined osteoglossid have already been described in the continental Campanian (Late Cretaceous) of Champ-Garimond, in southern France (SIGÉ et al., 1997). This French fossil may also belong to the genus Scleropages, but those remains are too fragmentary for decisive conclusions.

The occurrence of Scleropages, a freshwater genus actually living in Southeast Asia and Oceania, in the continental Paleocene of Belgium and perhaps already in the continental Campanian of France, can be explained by two different ways.

In the first hypothesis, the osteoglossoid distribution is considered as worldwide since the origin of the suborder during the Cretaceous thanks to the tolerance of those fishes for brackish and even marine waters. The present day distribution (see map in BERRA, 2001 : 53), which suggests an ancient “Gondwana pattern”, is then viewed as a fallacious picture. In that case, the European fossil Scleropages simply is a relic of that old cosmopolite distribution.

Another interpretation of the osteoglossoid geographic distribution is possible. Indeed, the Lower Cretaceous osteoglossoids only are known in Gondwanan regions. It is during the Upper Cretaceous that a few Osteoglossoidei appear for the first time in continental deposits of the northern continents. At that time, only one species is known in Europe, the undetermined Campanien osteoglossid of Champ-Garimont, and two species are reported in the western part of North America, the osteoglossid Cretophareodus alberticus Li, 1996 from the Campanian of Alberta, and the osteoglossoid incertae sedis Chandlerichthys strickeri Grande, 1986 from the Cenomanian of Alaska. A real worldwide distribution of the Osteoglossoidei only occurs during the Lower Tertiary and is principally due to the numerous marine species then present in the oceans. The disappearance of all those marine Paleogene osteoglossoids, of Scleropages in Europe, and of Phareodus acutus Leidy, 1873 and Phareodus testis (Cope, 1877) in North America after the Eocene has brought back the dispersal of the suborder to Gondwanan frontiers as in the Lower Cretaceous.

That second hypothesis puts a question about the European fossil Scleropages. How this freshwater genus has succeeded to reach Europe?

An Indian origin is questionable. During the Upper Cretaceous and the Lower Tertiary, India still was isolated in the ocean and separated from Western Europe by all the length of the Mesogea, a too long marine distance to pass over for a freshwater fish.

A connection between North America and Europe is possible via Greenland and Britain during practically all the Cretaceous times. The marine expanses between those areas during that geological period are not very wide and we know that Osteoglossidae are able to tolerate brackish and even marine waters. However, the two Upper Cretaceous osteoglossoid species are located westward to the large Cretaceous epicontinental seaway then bisecting North America into two land masses. To go over such a wide marine expanse is practically impossible for freshwater teleosts and moreover no freshwater osteoglossoid has ever been found located eastward to that sea in North America during the Cretaceous. Thus, a North American origin for the European Scleropages also is improbable.

A connection between North Africa and Europe during the Cretaceous is possible via the Iberian and/or the Apulian platforms and the surrounding islands. The marine expanses between those emerged territories at that time were not very wide. We know also that the occurrence of Osteoglossidae in Africa is attested as soon as the Early Cretaceous with Chanopsis lombardi from the Valanginian-Barremian (CAHEN et al., 1959), a geological period during which South-America and Africa were still connected. Other fossil Osteoglossiformes are known in Africa during the Cretaceous but they are younger : the arapaimid Paradercetis kipalaensis Casier, 1965 and the kipalaichtyid Kipalaichthys sekirksyi Casier, 1965, both from the Cenomanian of the Democratic Republic of Congo (TAVERNE, 1976). In addition to that it is obvious that the Recent South-American Osteoglossum and the Southeast Asian and Oceanian Scleropages are very close relatives, sharing an almost similar osteology and differing only by a few details (TAVERNE, 1977). Even if no fossil remains of Sleropages were found until now in Africa, the actual Gondwanan distribution of those two freshwater genera implicates that an Osteoglossum-Scleropageslike osteoglossid was present in Africa in the earliest period of the Cretaceous, before a too important break up of Gondwana. An African origin for the European fossil Scleropages is thus the most probable hypothesis.

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Received: January 27, 2006 Accepted: December 18, 2006