BackgroundThe classification of gobioid fishes is still under discussion. Several lineages, including the Eleotridae and Butidae, remain difficult to characterize because synapomorphies are rare (Eleotridae) or have not yet been determined (Butidae). Moreover, the fossil record of these groups is scarce.ResultsExceptionally well-preserved fish fossils with otoliths in situ from uppermost Oligocene sediments (≈23–24 Mio. y. ago) in Southern France provide the most in-depth description of a fossil gobioid to date. The species was initially described as Cottus aries Agassiz, then transferred to †Lepidocottus Sauvage, and subsequently assigned to Gobius. Based on a comparative analysis of meristic, osteological and otolith data, this species most likely is a member of the family Butidae. This discovery is important because it represents the first record of a fossil butid fish based on articulated skeletons from Europe.SignificanceThe Butidae and Eleotridae are currently distributed in W-Africa, Madagascar, Asia and Australia, but they do not appear in Europe and also not in the Mediterranean Sea. The new results indicate that several species of the Butidae thrived in Europe during the Oligocene and Early Miocene. Similar to the recent Butidae and Eleotridae, these fishes were adapted to a wide range of salinities and thrived in freshwater, brackish and marginal marine habitats. The fossil Butidae disappeared from Europe and the Mediterranean and Paratethys areas during the Early Miocene, due probably to their lack of competitiveness compared to other Gobioidei that radiated during this period of time. In addition, this study documents the great value of otoliths for gobioid systematics.
The modern Gobioidei (Teleostei) comprise eight families, but the extinct †Pirskeniidae from the lower Oligocene of the Czech Republic indicate that further families may have existed in the past. However, the validity of the †Pirskeniidae has been questioned and its single genus †Pirskenius has been assigned to the extant family Eleotridae in previous works. The objective of this study is to clarify the status of the †Pirskeniidae. Whether or not the †Pirskeniidae should be synonymised with the Eleotridae is also interesting from a biogeographical point of view as Eleotridae is not present in Europe or the Mediterranean Sea today. We present new specimens and reexamine the material on which the two known species of †Pirskenius are based (†P. diatomaceus Obrhelová , 1961; †P. radoni Přikryl, 2014). To provide a context for phylogenetically informative characters related to the palatine and the branchiostegal rays, three early-branching gobioids (Rhyacichthys, Protogobius, Perccottus), an eleotrid (Eleotris) and a gobiid (Gobius) were subjected to micro-CT analysis. The new data justify revalidation of the family †Pirskeniidae, and a revised diagnosis is presented for both †Pirskenius and †Pirskeniidae. Moreover, we provide for the first time an attempt to relate a fossil gobioid to extant taxa based on phylogenetic analysis. The results indicate a sister-group relationship of †Pirskeniidae to the Thalasseleotrididae + Gobiidae + Oxudercidae clade. Considering the fossil record, the arrival of gobioids in freshwater habitats in the early Oligocene apparently had generated new lineages that finally were not successful and became extinct shortly after they had diverged. There is currently no evidence that the Eleotridae was present in the European ichthyofauna in the past.
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