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.

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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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Mudskippers are amphibious gobies (Teleostei: Gobioidei, Oxudercinae) that have served as models for the specialised physiology and behaviour of fishes out of water. In this study, a comparative analysis of the posterior vertebral column and the caudal skeleton of ten mudskipper species was conducted on the basis of X‐ray imaging. The species considered were Apocryptes bato, Apocryptodon madurensis, Boleophthalmus dussumieri, Oxuderces dentatus, Periophthalmodon freycineti, Pn. schlosseri, Periophthalmus novemradiatus, Ps. waltoni, Pseudapocryptes borneensis, and Scartelaos tenuis. For the osteological description the new term 'modified caudal vertebra' is used for all those vertebrae that display visibly modified neural and/or haemal spines compared to the spines of a 'usual' caudal vertebra, but are not involved in the support of caudal rays. The results reveal that the most terrestrial forms (Pn. freycineti, Pn. schlosseri, Ps. novemradiatus, Ps. waltoni) possess distinct traits that are seldom found in the other species. Among these features are (a) the existence of at least two modified caudal vertebrae (also present in S. tenuis), (b) a particularly close, dovetailing association between the neural spines of the preural vertebrae two and three (restricted to Ps. novemradiatus and Ps. waltoni), and (c) thickening and shortening of the ventralmost principal caudal rays (also present in B. dussumieri and S. tenuis). These findings support the idea that the posterior caudal vertebrae and caudal skeleton of the mentioned species are modified to enhance locomotion on land. Moreover, a relationship between character development and degree of terrestrial adaptation is probable, as all three traits are most pronounced in Ps. waltoni, which correlates with its strikingly high level of adaptation to amphibious life. A further aspect of this study is that the newly recognized skeletal structures have good fossilization potential and could therefore facilitate recognition of fossil species of mudskippers, which are currently unknown.

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The suborder Gobioidei is among the most diverse groups of vertebrates, comprising about 2310 species. In the fossil record gobioids date back to the early Eocene (c. 50 m.y. ago), and a considerable increase in numbers of described species is evident since the middle Miocene (c. 16 m.y. ago). About 40 skeleton-based gobioid species and > 100 otolith-based species have been described until to date. However, assignment of a fossil gobioid species to specific families has often remained tentative, even if well preserved complete specimens are available. The reasons are that synapomorphies that can be recognized in a fossil skeleton are rare (or absent) and that no phylogenetic framework applicable to gobioid fossils exists. Here we aim to overcome this problem by developing a phylogenetic total evidence framework that is suitable to place a fossil skeleton-based gobioid at family level. Using both literature and newly collected data we assembled a morphological character matrix (48 characters) for 29 extant species, representing all extant gobioid families, and ten fossil gobioid species, and we compiled a multi-gene concatenated alignment (supermatrix; 6271 bp) of published molecular sequence data for the extant species. Bayesian and Maximum Parsimony analyses revealed that our selection of extant species was sufficient to achieve a molecular ‘backbone’ that fully conforms to previous molecular work. Our data revealed that inclusion of all fossil species simultaneously produced very poorly resolved trees, even for some extant taxa. In contrast, addition of a single fossil species to the total evidence data set of the extant species provided new insight in its possible placement at family level, especially in a Bayesian framework. Five out of the ten fossil species were recovered in the same family as had been suggested in previous works based on comparative morphology. The remaining five fossil species had hitherto been left as family incertae sedis. Now, based on our phylogenetic framework, new and mostly well supported hypotheses to which clades they could belong can be presented. We conclude that the total evidence framework presented here will be beneficial for all future work dealing with the phylogenetic placement of a fossil skeleton-based gobioid and thus will help to improve our understanding of the evolutionary history of these fascinating fishes. Moreover, our data highlight that increased sampling of fossil taxa in a total-evidence context is not universally beneficial, as might be expected, but strongly depends on the study group and peculiarities of the morphological data.