Abstract
Articulated fossil fish skeletons with otoliths in situ provide a unique opportunity to link these two, otherwise independent data sets of skeletons and otoliths. They provide calibration points for otoliths also adding important information for the evolutionary interpretation of fishes. Here, we review nine articulated skeletons of gobies from the early Sarmatian of Dolje, Croatia, and Belgrade, Serbia, which were previously regarded as members of a single gobiid and a callionymid species. We found them to represent five different gobiid species belonging to five different genera, four of which are related to extant endemic Ponto-Caspian gobiid lineages. The species are: Aphia macrophthalma n.sp., Proneogobius n.gen. pullus (the only previously recognized species), Protobenthophilus n.gen. squamatus n.sp., Economidichthys triangularis (a species first described based on otoliths) and Hesperichthys n.gen. reductus n.sp. Five specimens contained otoliths in situ and a sixth shows imprints of otoliths which unfortunately must have been lost in the past, probably during preparation of the fossil. Together, they represent all five species recognized by skeletons, and three are linked to otolith-based species. Isolated otoliths have been reviewed from a variety of collections from Sarmatian strata in Austria, Bulgaria, Czechia, Romania and Slovakia resulting in the description of five new otolith-based species: Benthophilus? ovisulcus n.sp., Benthophilus styriacus n.sp., Protobenthophilus strashimirovi n.sp., Economidichthys altidorsalis n.sp. and Knipowitschia bulgarica n.sp. Our review demonstrates that all major endemic Ponto-Caspian gobiid lineages were already present during Sarmatian times, thereby pushing back their origin by approximately 5–10 myr in comparison to previously published dates for dichotomies. In our assessment, the origination of these lineages is linked to the early stage of separation of the Paratethys from the world oceans and the ecological changes that occurred during that time. These geological events parallel a dramatic increase in gobiid radiation and speciation, giving rise to many lineages, not all of which have persisted until today.
Highlights
Articulated fossil fish skeletons with otoliths in situ provide a unique opportunity to link these two, otherwise independent data sets of skeletons and otoliths
Proneogobius pullus was originally described by Kramberger (1882) as Gobius pullus based on two specimens
Otoliths of Proterorhinus vasilievae are characterized by a compressed shape (OL:OH = 0.9–0.95); sharp and pronounced predorsal and preventral angles; distinct postdorsal angle followed by a short postdorsal projection, which is only slightly bent outwards; and a distinctly sole-shaped sulcus with a long, rather narrow subcaudal iugum
Summary
Articulated fossil fish skeletons with otoliths in situ provide a unique opportunity to link these two, otherwise independent data sets of skeletons and otoliths. It has only been very recently that fossil otoliths of gobies have been described from Middle Miocene strata of the Paratethys from Kazakhstan (Bratishko et al 2015) and Serbia (Schwarzhans et al 2015) and were identified as related to those endemic Ponto-Caspian goby groups. The oldest report of gobies from the Sarmatian of the Paratethys was provided by Steindachner (1860), who described three species from a particular paleoenvironment in Austria (Gobius elatus, G. oblongus and G. viennensis) They are not part of this review and are currently being studied by Reichenbacher and Gierl in Munich. Additional material currently being studied by Bratishko and Schwarzhans from the Sarmatian of the Crimea will further increase the taxonomic diversity This amazing diversity compares to the nine skeleton-based gobiid species recognized from the same area and time interval. Reveals that the material that was assigned to Gobius pullus comprises five different species allocated to five different genera
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