Osteoglossomorphs of the marine Lower Eocene of Denmark – with remarks on other Eocene taxa and their importance for palaeobiogeography

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Abstract The geological, faunal and palaeoecological conditions of the marine deposits from lowermost Eocene in North Jutland are briefly reviewed as background for the descriptions of six species of osteoglossiform fishes from the Stolle Klint Clay and the overlying Mo-clay (Ølst and Fur Formations respectively). Four of these primitive teleosteans are referred to new genera and species (one based on an almost complete skeleton, three others on skull material, one very incomplete), and the two most fragmentary specimens are referred to Brychaetus sp. and an indeterminate osteoglossiform. The phylogenetic relationships of these fossils are evaluated in the framework of two different models of osteoglossomorph phylogeny provided earlier by Taverne and Hilton. Despite differences in both data bases, methodologies and results given by the two models (the former based on c . 300 characters in an intuitive, qualitative phylogenetic parsimony analysis, the latter on 72 characters in a critical, rigorous, quantitative cladistic analysis) the phylogenetic positions of four of the fossil species are very similar in the two models concerning the relations to the recent forms. The other two species, rather fragmentary, but similar in many ways to the Eocene phareodonts (paraphyletic group), end up very differently in relation to extant forms in the two models. The phylogenetic systematics of all the marine, fossil osteoglossiforms (including Brychaetus, Opsithrissops and Monte Bolca forms) is evaluated as background for interpretation of their (palaeo-)biogeographic significance as marine members of a group, Osteoglossomorpha (of which the recent forms are prime examples of ‘primary division freshwater fishes', and of which the extant osteoglossiforms have a classical ‘Gondwana distribution’. There are 9 marine, Eocene taxa (plus an otolith from the Maastrichtian of USA) and none of the 9 appear more closely-related to any other marine form in either model: they might constitute 9 separate migrations from freshwater into the sea. The phylogenetic results strongly suggest instead, that the extant osteoglossiforms have independently entered freshwater from the sea on two, perhaps even three occasions. This may have happened as late as the Eocene, and phareodonts could be yet another independent invasion of freshwater in the Late Cretaceous. The mormyriforms most likely had an independent invasion into freshwater (in one model even with notopterids as a separate migration from the sea by Mid or Early Cretaceous). Because all the closest outgroups of the Osteoglossomorpha are marine, the group obviously originated in the sea, probably by the Late Jurassic, and it is not impossible that Hiodontiforms in NE Asia and North America underwent another independent freshwater invasion very early in the Cretaceous. What then is wanting? The expected Cretaceous, marine osteoglossomorphs are not found (but note the above otolith).