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Otoliths in Situ from Sarmatian (Middle Miocene) Fishes of the Paratethys

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Otoliths in Situ from Sarmatian (Middle Miocene) Fishes of the Paratethys Swiss J Palaeontol (2017) 136:19–43 DOI 10.1007/s13358-016-0114-5 Otoliths in situ from Sarmatian (Middle Miocene) fishes of the Paratethys. Part II: Gadidae and Lotidae 1 2 3,4 Werner Schwarzhans • Giorgio Carnevale • Andriy Bratishko • 5 6 Sanja Japundzˇic´ • Katarina Bradic´ Received: 9 January 2016 / Accepted: 1 March 2016 / Published online: 12 April 2016 Ó Akademie der Naturwissenschaften Schweiz (SCNAT) 2016 Abstract Gadid otoliths are among the most common macropterygius,whileP. anceps does not seem to be present otoliths in the Neogene of Europe. To date, these have been in the Paratethys after the late Badenian/Konkian. The oto- recorded in situ and therefore correlated with the skeletal lith-based species Palimphemus minusculoides (Schubert record only in two cases, Paratrisopterus avus and 1912) is considered as a junior synonym of P. macroptery- Palimphemus anceps. Here, we describe otoliths in situ from gius. Paratrisopterus caspius is regarded as a senior syn- three gadiform taxa from the Sarmatian of the Central onym of P. avus Fedotov 1971, whose otoliths in situ were Paratethys—Palimphemus macropterygius, Paratrisopterus previously described by Fedotov (1976). The new data allow caspius and Enchelyopus susedanus. A number of speci- further synonymization of otolith-based species, such as P. mens previously described by Kramberger (Pala¨ontol. insectus (Weiler 1943) and possibly also P. irregularis O¨ sterr. -Ungar. und des Orients 3:65–85, 1883) and And- (Gaemers 1973). Moreover, Properca sabbai Pauca 1929 is jelkovic´ (Glas. Prir. Muz. A 24:127–154, 1969) are revised. removed from the faunal list of Sarmatian fishes in the Kramberger’s Morrhua macropterygia is reassigned to the Paratethys. Kner’s Brosmius susedanus is reassigned to the extinct gadid genus Palimphemus; moreover, Morrhua extant lotid genus Enchelyopus,withBrosmius elongatus lanceolata is considered a junior synonym of P. anceps Kner Kramberger 1883 representing a junior synonym. There is 1862. All the Palimphemus specimens studied from the no record of isolated otoliths correlating with E. susedanus Sarmatian of the Central Paratethys belong to P. in the literature; however, a number of previously unde- scribed specimens of E. susedanus have been identified from the middle Sarmatian s.l. of Jurkino, Crimea. Editorial handling: L. Cavin & D. Marty. & Werner Schwarzhans Keywords Gadidae Á Lotidae Á Palimphemus Á [email protected] Paratrisopterus Á Enchelyopus Á Otoliths in situ 1 Natural History Museum of Denmark, Zoological Museum, Universitetsparken 15, 2100 Copenhagen, Denmark Introduction 2 Dipartimento di Scienze della Terra, Universita` degli Studi di Torino, via Valperga Caluso 35, 10125 Turin, Italy 3 This is the third study dealing with the otoliths in situ from Faculty of Natural Sciences, Luhansk Taras Shevchenko Sarmatian fishes of the Paratethys. The first of these was National University, Gogolya Sqr. 1, Starobelsk, Luhansk Region 92703, Ukraine devoted to the Atherinidae (Schwarzhans et al. 2016), and the second one to clupeid fishes (Baykina and Schwarzhans 4 BugWare Inc., 1615 Village Square Blvd., Ste 208, Tallahassee, FL 32309, USA 2016). Skeletal remains and otoliths of the families Gadidae 5 Department of Geology and Paleontology, Croatian Natural History Museum, Demetrova 1, 10000 Zagreb, Croatia and, to a lesser extent, of the Lotidae are among the most common teleosts in the Middle Miocene of the Central and 6 Department of Paleontology, Faculty of Mining and Geology, University of Belgrade, Kamenicˇka 6, 11000 Belgrade, Eastern Paratethys. During the early Badenian, most of the Serbia gadid otoliths of the Central Paratethys also occurred in the 20 W. Schwarzhans et al. North Sea Basin (Palimphemus anceps, Gadiculus argen- Morrhua macropterygia Kramberger 1883: Middle Mio- teus, Paratrisopterus labiatus) (Schwarzhans 2010; Nolf cene, Sarmatian s.s. of Dolje, Croatia. Holotype and syn- 2013), where they are the dominant teleosts. In the Central types reviewed in this study and considered as a valid and Eastern Paratethys, gadid and lotid fishes become species of the genus Palimphemus. really abundant only during the Sarmatian, as clearly tes- Morrhua extensa Kramberger 1885: Sarmatian s.s. of tified by articulated skeletons and otoliths. Paratrisopterus Zagorje ob Savi (Szakada´t, Sagor), Slovenia. In need of avus and P. anceps were the first cases of Neogene fossil revision. gadids for which articulated skeletons with otoliths in situ became known (Fedotov 1976; Schwarzhans 2014). Morrhua minima Kramberger 1885: Middle Miocene (late Several nominal skeleton- and otolith-based gadid and Badenian or Sarmatian s.s.) of Podsused, Croatia. Kram- lotid species have been described from the Sarmatian s.s. berger mentioned about 16 specimens, none of which could of the Central Paratethys and the Sarmatian s.l. to Pontian be located at CNHM. Based on the meristic data provided of the Eastern Paratethys (see below). A thorough revision in the original description, this is probably a species of the of both skeletal and otolith data, when possible, would extinct genus Paratrisopterus. likely reduce the large number of established taxa in this Gadus caspius Bogatshov 1929: Late Miocene, middle to group, even if this is beyond the scope of this study. Our late Sarmatian s.l. of Azerbaijan. The two type specimens aim is to provide link between skeletal and otolith finds as are probably lost. Based on the original description by much as possible, and for this reason we restrict our Bogatshov, the species is considered herein as Para- efforts to review the taxa with otoliths in situ. Never- trisopterus caspius. theless, below we list the key data for all relevant taxa from the Sarmatian and younger strata of the Central and Gadus kiplingi Bogatshov 1929: Middle to late Miocene, Eastern Paratethys in order to provide a general overview, middle to late Sarmatian s.l. of Azerbaijan. The type and add short comments on their current status. The list specimens are probably lost. Based on the original follows the sequence of description of the individual description by Bogatshov, the species can be considered as species, first skeletons, then otoliths, with their original a species of the extinct genus Paratrisopterus. designation. Gadus kwitkae Bogatshov 1933: Middle to late Miocene, Pontian of Azerbaijan. The type specimen is probably lost. Based on the original description by Bogatshov, it probably Annotated list of skeleton-based Gadidae represents a species of the genus Gadiculus. Palimphemus anceps Kner 1862: Middle Miocene, late Paratrisopterus avus Fedotov 1971: Middle Miocene, Sar- Badenian of St. Margarethen, Austria. Recently reviewed matian s.l. of Moldavia. Reviewed by Prokofiev (2004)and and re-described by Carnevale et al. (2012); otoliths in situ considered valid as Gadiculus avus. Fedotov (1976)provided described in a specimen from the middle Badenian of a figure of an otolith found in situ in the holotype, but Pro- Poland by Schwarzhans (2014), including synonymization kofiev (2004) could not locate the otolith. In this study, the of several otolith-based species, notably Colliolus sculptus genus Paratrisopterus is considered valid and the species is (Koken 1891). considered as a junior synonym of the otolith-based species Paratrisopterus insectus (Weiler 1943) and also of the Morrhua aeglefinoides Kner & Steindachner 1863: Middle skeletal-based species P. caspius (Bogatshov 1929). Miocene (late Badenian or Sarmatian s.s.) of Podsused, Croatia. In need of revision; based on the meristic data provided in the original description, it appears to be a Annotated list of otolith-based Gadidae species of the genus Trisopterus. Morrhua szagadatensis Steindachner 1863: Middle Mio- Otolithus (Gadidarum) minusculus Schubert 1906: Middle cene, Sarmatian s.s. of Zagorje ob Savi (Szakada´t, Sagor), Miocene, late Burdigalian and Sarmatian s.s. of localities Slovenia. In need of revision; based on the short descrip- in Slovakia and Hungary. Reviewed by Nolf (1981) who tion by Steindachner, it appears to be a problematic taxon selected a lectotype and rejected the species because of based on an incomplete specimen. inadequate preservation and non-diagnostic juvenile status of the specimens. Bratishko et al. (2015) considered the Morrhua lanceolata Kramberger 1883: Middle Miocene lectotype as a juvenile of Palimphemus anceps. (late Badenian or Sarmatian s.s.) of Podsused, Croatia. Holotype reviewed in this study and considered as a junior Otolithus (Gadidarum) minusculoides Schubert 1912:Middle synonym of Palimphemus anceps. Miocene, Sarmatian s.s. of Hungary. Reviewed by Nolf Otoliths in situ from Sarmatian fishes of the Paratethys. Part II: Gadidae and Lotidae 21 (1981) and rejected because of inadequate preservation of the Macrurus simplex Smigielska 1966: Middle Miocene, late type specimen. Re-established and re-described by Bratishko Badenian of Gliwice, Poland. Considered as a doubtful et al. (2015)asPalimphemus minusculoides;considered species based on not interpretable juvenile specimens by herein as a junior synonym of Palimphemus macropterygius. Nolf (1985); considered as a synonym of Paratrisopterus rumanus by Schwarzhans (2010). Otolithus (Gadidarum) insectus Weiler 1943: Middle Miocene, late Badenian of Melicesti, Romania. Reviewed Macrurus planus Smigielska 1966: Middle Miocene, late by Nolf (1985), Schwarzhans (2010) and in this study, and Badenian of Gliwice, Poland. Considered as a doubtful considered as a junior synonym of Paratrisopterus
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