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Late Middle Miocene) Marine Fishes and Mammals As Proxies for the Onset of the Eastern Paratethys Re-Colonisation by Vertebrate Fauna

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Late Middle Miocene) Marine Fishes and Mammals As Proxies for the Onset of the Eastern Paratethys Re-Colonisation by Vertebrate Fauna Palaeontologia Electronica palaeo-electronica.org The Volhynian (late Middle Miocene) marine fishes and mammals as proxies for the onset of the Eastern Paratethys re-colonisation by vertebrate fauna Pavel Gol’din, Bogdan Stelian Haiduc, Oleksandr Kovalchuk, Marcin Górka, Pavlo Otryazhyi, Mihai Brânzilă, Elena Ionela Păun, Zoltán Barkaszi, Paul Ţibuleac, and Bogdan Gabriel Răţoi ABSTRACT The late Middle Miocene of South-Eastern Europe was the time of major basin- wide physical and biogeographical transformations of marine environments with mass extinction and faunal replacement. We examined new material comprising marine fishes and mammals, re-examined earlier finds, and analysed previously published data on the marine vertebrate assemblage that lived along the western and northern edges of the Eastern Paratethys. The studied materials represent marine fishes that belong to at least four families (Clupeidae, Sparidae, Bothidae, and Scombridae), including Sarmatella doljeana, Sparus brusinai, Bothus parvulus, and indeterminate scombroids; true seals (Phocinae), the earliest seal record in the area; small toothed whales of the family Kentriodontidae, including Kentriodon fuchsii, whose presence in the Volhynian is confirmed here, and two other dwarf forms; a bizarre toothed whale Pachyacanthus, the first record in the area between the Vienna Basin and Central Asia; baleen whales, including an indeterminate member of the family Cetotheriidae, and “Archaeocetus fockii”. The material found so far hints to a low taxonomic diversity and biogeographical connections to both the synchronous Sarmatian fauna of the Cen- tral Paratethys and the later Bessarabian Eastern Paratethyan fauna. The taxonomic composition of the faunal assemblage and its biogeographical connections indicate that it has Central Paratethyan origin and shows an overall west to east direction of gradual dispersal, from the Central Paratethys through the Fore-Carpathian Basin and further east. The taxonomic composition of described Volhynian fish and mammal assemblages supports the idea of faunal connection and transition between the Cen- tral and Eastern Paratethys. Gol’din, Pavel, Haiduc, Bogdan Stelian, Kovalchuk, Oleksandr, Górka, Marcin, Otryazhyi, Pavlo, Brânzilă, Mihai, Păun, Elena Ionela, Barkaszi, Zoltán, Ţibuleac, Paul, and Răţoi, Bogdan Gabriel. 2020. The Volhynian (late Middle Miocene) marine fishes and mammals as proxies for the onset of the Eastern Paratethys re-colonisation by vertebrate fauna. Palaeontologia Electronica, 23(3):a43. https:// doi.org/10.26879/1091 palaeo-electronica.org/content/2020/3152-the-volhynian-late-middle-miocene Copyright: September 2020 Paleontological Society. This is an open access article distributed under the terms of Attribution-NonCommercial-ShareAlike 4.0 International (CC BY-NC-SA 4.0), which permits users to copy and redistribute the material in any medium or format, provided it is not used for commercial purposes and the original author and source are credited, with indications if any changes are made. creativecommons.org/licenses/by-nc-sa/4.0/ GOL’DIN ET AL.: THE VOLHYNIAN (LATE MIDDLE MIOCENE) Pavel Gol’din. Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, 15 Bohdan Khmelnytsky st., 01030, Kyiv, Ukraine. [email protected] Bogdan Stelian Haiduc. Alexandru Ioan Cuza University of Iaşi, Faculty of Geography and Geology, Department of Geology, Iaşi, Romania. [email protected] Oleksandr Kovalchuk. Department of Paleontology, National Museum of Natural History, National Academy of Sciences of Ukraine, 15 Bohdan Khmelnytsky st., 01030, Kyiv, Ukraine and Department of Palaeozoology, Institute of Environmental Biology, University of Wroclaw, Sienkiewicza 21, 50-335 Wroclaw, Poland. [email protected] Marcin Górka. Faculty of Geology, University of Warsaw, Al. Żwirki i Wigury 93, 02-089 Warszawa, Poland. [email protected] Pavlo Otryazhyi. Karazin Kharkiv National University, 4 Svobody sq., 61022, Kharkiv, Ukraine. [email protected] Mihai Brânzilă. Alexandru Ioan Cuza University of Iaşi, Faculty of Geography and Geology, Department of Geology, Iaşi, Romania. [email protected] Elena Ionela Păun. Alexandru Ioan Cuza University of Iaşi, Faculty of Geography and Geology, Department of Geology, Iaşi, Romania. [email protected] Zoltán Barkaszi. Department of Paleontology, National Museum of Natural History, National Academy of Sciences of Ukraine, 15 Bohdan Khmelnytsky st., 01030, Kyiv, Ukraine. [email protected] Paul Ţibuleac. Alexandru Ioan Cuza University of Iaşi, Faculty of Geography and Geology, Department of Geology, Iaşi, Romania. [email protected] Bogdan Gabriel Răţoi. Alexandru Ioan Cuza University of Iaşi, Faculty of Geography and Geology, Department of Geology, Iaşi, Romania. Corresponding author. [email protected] Keywords: Paratethys; Miocene; fishes; seals; cetaceans; dispersal Submission: 21 April 2020. Acceptance: 25 August 2020. INTRODUCTION Sarmatian s.l. fauna developed into the assem- blages of Bessarabian and, later, Khersonian The late Middle Miocene of South-Eastern regional stages. Europe was the time of major physical and biogeo- There are numerous records of marine verte- graphic transformations of marine environments brates from the Middle/Late Miocene boundary, (Rögl, 1998; Popov et al., 2004; Palcu et al., 2015, particularly from the Sarmatian s.s. of the Central 2017). The Paratethys, a great epicontinental sea Paratethys (Kazár, 2006), and the Bessarabian of of Eurasia, gradually lost its connection with the the Eastern Paratethys (Brandt, 1873; Koretsky, World Ocean due to the uplift of the Alpine- 2001). However, few localities in the Eastern Para- Dinarides, Carpathians and Pontides, and broke tethys yielded marine vertebrates from pre-Volhyn- into a number of inner seas (Rögl, 1998; Popov et ian times. The fossil record of fishes from the al., 2004; Palcu et al., 2015, 2017; Sant et al., Volhynian beds of the Eastern Paratethys is well- 2017). At that time, there were two major basins, documented (Bogachev, 1929, 1938; Paucă, 1938; the Central and the Eastern Paratethys, with Ionko, 1954; Bogachev, 1955; Rückert-Ülkümen, changing contours and environmental conditions. 1965; Bannikov, 1986, 1989, 1993; Carnevale et These basins were divided by the uplifting land- al., 2006; Bannikov, 2009, 2010; Baykina, 2012, mass of the Carpathians, and by the smaller Fore- 2013a; Bannikov and Kotlyar, 2015; Baykina and Carpathian Basin connected to both basins (Kováč Schwarzhans, 2017; Schwarzhans et al., 2017a-d). et al., 2017). Their progressing isolation led to the Bogachev (1955) provided a list of fishes including formation of specific local faunas and, respectively, clupeids and a few other marine fish taxa based on regional stratigraphic units. During the Volhynian materials from the Volhynian deposits near Amvro- age (or Sarmatian s.s.), ca. 12.82 (or 12.65) – 11.8 siivka (eastern Ukraine). However, his material is Ma, which corresponds to the late Serravallian considered problematic (Bannikov, 2010) since it (Palcu et al., 2017), they formed an internally con- was not described properly and also was probably nected basin. Further isolation of the Central Para- lost thus unavailable for revision. Macarovici tethys led to the development of the Pannonian (1970) reported a small fish assemblage from the fauna, while in the Eastern Paratethys the endemic 2 PALAEO-ELECTRONICA.ORG Moldavian Platform; those specimens are re- sented by the regional stratigraphic units of the described here. Bannikov (2009) described the Konkian (or late Badenian in the Central Para- early Volhynian fish assemblage from Naslavcea, tethys) and Volhynian (or Sarmatian s.s. in the Republic of Moldova, and noticed that it was differ- Central Paratethys) (Figure 2). During the trans- ent from the contemporary fauna of the North Cau- gressive phases both basins had fully marine con- casus. ditions; however, they were separated during Contrary to fish records, data on Volhynian regressive phases that led to a gradual decrease in marine mammals are scarce. A fish-bearing locality salinity with certain salinity crises (Popov et al., and a few toothed whales (“dolphins”) were 2004; Ter Borgh et al., 2014; Vernyhorova, 2015). reported from the Caucasus and the Caspian The newly discovered and the revised material region (Dombrovsky, 1927; Kirpichnikov, 1954; mostly comes from the Fore-Carpathian Basin Mchedlidze, 1964; Bannikov, 2009), whereas the (Podolian Platform): from the Ukrainian localities of western part of the basin was less studied (Kudrin Kolubaivtsi, Hrushivtsi, and Vilkhovets, and from and Tatarinov, 1966). A few cetaceans were the Romanian localities of the Eastern Carpathian reported from the Moldavian Platform in Romania Foreland (Moldavian Platform): Stăuceni, (Macarovici, 1947; Macarovici and Zaharia, 1968; Leucuşeşti, Stâncești, Basarabi, and Drăguşeni Ionesi and Galan, 1988; Figure 1). (Figures 1, 2). Here we report newly collected marine verte- A number of other findings and localities were brate specimens from Romania and Ukraine with reported from these areas (Khotin: Macarovici, their tentative classification, a review of previous 1947; Khonkivtsi: Hofstein, 1965; Brykiv: Kudrin finds, a revision of the localities, and respective and Tatarinov, 1966; Ghireni, Cordăreni, Râng- biogeographical implications. hileşti, Mitoc-Prut, Hăneşti, Săveni-Ghireni, Drăguşeni: Macarovici and Zaharia, 1967, 1968; GEOLOGICAL FRAMEWORK Fălticeni, Voineşti, Erbiceni, Aroneanu,
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