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.

中文翻译：

---

来自Paratethys的Sarmatian（中新世中期）鱼类的原位耳石。第三部分：蓬图里海虾虎虾的摇篮故事

带有原位耳石的铰接化石鱼骨骼为链接这两个否则独立的骨骼和耳石数据集提供了独特的机会。它们为耳石提供了校准点，还为鱼类的进化解释添加了重要信息。在这里，我们回顾了克罗地亚多尔杰和塞尔维亚贝尔格莱德的早期萨尔马提人的九个具关节状的虾虎鱼骨骼，这些骨骼以前被视为单个戈壁鱼和一个and虫物种的成员。我们发现它们代表了属于五个不同属的五个不同的戈壁类物种，其中四个与现存的特有的蓬图里海戈壁类谱系有关。该物种是：大蚜虫n.sp.，Proneogobius n.gen。Pullus（以前唯一公认的物种），Protobenthophilus n.gen。鳞片 Economidichthys triangleis（首先根据耳石描述的物种）和Hesperichthys n.gen。减少量 五个标本原位包含耳石，第六个标本显示了耳石的印记，不幸的是，这些印石在过去很可能是在化石制备过程中丢失的。它们共同代表了由骨架识别的所有五个物种，其中三个与基于耳石的物种相关。已对来自奥地利，保加利亚，捷克，罗马尼亚和斯洛伐克的萨尔马地层的各种收集物中的分离出的耳石进行了审查，得出了五个基于耳石的新物种的描述：嗜睡菌？ovisulcus n.sp.，stythcuscus styriacus n.sp.，Protobenthophilus strashimirovi n.sp.，Economidichthys altidorsalis n.sp. 和保加利亚烟草（Knipowitschia bulgarica）我们的评论表明，所有主要的地方性蓬图里海沿岸戈壁血统都已经存在于萨尔马特时期，因此与以前公布的二分法日期相比，它们的起源被推迟了约5-10 myr。在我们的评估中，这些血统的起源与Paratethys从世界海洋中分离出来的早期阶段以及当时发生的生态变化有关。这些地质事件伴随着戈壁辐射和物种形成的急剧增加，产生了许多世系，但并不是所有世系一直持续到今天。这些血统的起源与Paratethys从世界海洋中分离的早期阶段以及当时发生的生态变化有关。这些地质事件伴随着戈壁辐射和物种形成的急剧增加，产生了许多世系，但并不是所有世系一直持续到今天。这些血统的起源与Paratethys从世界海洋中分离的早期阶段以及当时发生的生态变化有关。这些地质事件伴随着戈壁辐射和物种形成的急剧增加，产生了许多世系，但并不是所有世系一直持续到今天。