Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy) vol. 124(3): 535-571. November 2018 NEOGENE PARATETHYAN CROAKERS (TELEOSTEI, SCIAENIDAE) ALEXANDRE F. BANNIKOV1, WERNER SCHWARZHANS2* & GIORGIO CARNEVALE3 1Borisyak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, Moscow 117647, Russia. E-mail: [email protected] 2*Corresponding author. Natural History Museum of Denmark, Zoological Museum, Universitetparken 15, 2100 Copenhagen, Denmark. E-mail: [email protected]; http://orcid.org/0000-0003-4842-7989 3Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via Valperga Caluso 35 10125 Torino, Italy. E-mail: [email protected]; http://orcid.org/0000-0002-3433-4127 To cite this article: Bannikov A.F., Schwarzhans W. & Carnevale G. (2018) - Neogene Paratethyan croakers (Teleostei, Sciaenidae). Riv. It. Paleontol. Strat., 124(3): 535-571. Keywords: Sciaenidae; new genera and species; Neogene; Tarkhanian; North Caucasus; Sarmatian; Croatia; Paratethys. Abstract. The fossil record of sciaenid fishes (based on both otoliths and osteological finds) from the Neoge- ne of the Paratethys is reviewed. The species Labrax (=Morone) multipinnatus Gorjanović-Kramberger, 1882 from the Sarmatian of Croatia (Sv. Nedelja) that was originally described as a sea bass of the family Moronidae is re-assigned to the extant croaker genus Argyrosomus. Two new genera and species of sciaenid fishes are described based on a single skeleton each, namely Landinisciaena popovi gen. et sp. nov. from the Tarkhanian (Lower/Middle Miocene) North Shir- vanskaya Formation exposed along the Pshekha River, North Caucasus, Russia, and Croatosciaena krambergeri gen. et sp. nov. from the Sarmatian s.s. (Middle Miocene) deposits of Dolje in the nearby of the city of Zagreb, Croatia. The holotype of the former taxon possesses a well-preserved saccular otolith in situ (first record of in situ croaker otolith in Europe) similar to the otoliths of the extant genus Atractoscion. The otolith-based Ottnangian species Atractoscion elongatissimus Schwarzhans, 1993 is transferred to the genus Landinisciaena gen. nov. A comprehensive revision of the Neogene otholith-based sciaenid record from Paratethys resulted in the recognition of three new genera (Chaoia gen. nov., Pontosciaena gen. nov. and Leptosciaena gen. nov.) as well as in the description of a new species L. caputoi sp. nov. from the uppermost Messinian (Lago-mare phase) of Italy. A number of known otholith-based species were either synonymized or assigned to a new generic affiliation. The complex biogeographic history of the Neogene Paratethyan sciaenids (based on both otoliths and skeletal records) is discussed, resulting in the recognition of a vanished “sciaenid bioprovince”. INTRODUCTION in the suborder Sciaenoidei together with the Em- melichthyidae (e.g., Betancur-R. et al. 2013; Near Members of the family Sciaenidae, also et al. 2013; Nelson et al. 2016), or as incertae se- known as croakers are shallow-dwelling supraben- dis within the Eupercaria (Betancur-R. et al. 2017). thic fishes widespread in tropical and subtropical The fossil record of croakers is mostly based on coastal and estuarine waters worldwide. A number their large and thick otoliths (Schwarzhans 1993), of species of this family are restricted to fresh- whereas articulated fossil skeletons of the Sciaeni- waters, especially in the Atlantic drainages of the dae are quite uncommon (see Bannikov et al. 2009). Americas (Nelson et al. 2016). The Sciaenidae is a The earliest record of the Sciaenidae is a single oto- highly diverse percomorph family, including about lith from the Ypresian Bashi Marls of Mississippi 280 recent species belonging to approximately 70 (Nolf 1995), although croaker otoliths become rel- genera (Nelson et al. 2016). While sciaenids have atively abundant in the late Eocene, Oligocene and been traditionally regarded as percoid perciforms Neogene shallow water sediments of Europe and (Nelson 2006), recent broad-scale molecular stud- the Americas (e.g., Schwarzhans 1993; Nolf 1985, ies placed them within the order Acanthuriformes 2013; Aguilera et al. 2016). Skeletal remains of ex- tinct croakers are relatively more common in the Received: July 19, 2018; accepted: September 25, 2018 Eastern Paratethyan record compared to other re- 536 Bannikov A.F., Schwarzhans W. & Carnevale G. gions, at least since the beginning of the Miocene. pterygiophore located posterior to the neural spine The extinct genus Caucasisciaena (Bannikov et al. of the third abdominal vertebra), and was assigned 2009; Bannikov 2010), with the species C. ignota is to the subfamily Carnevalellinae (Bannikov 2013). known from the Sakaraulian (Lower Miocene, Up- One of the two species of this genus, C. impropria, per Maikopian, Lower Burdigalian) of the north- was collected from the Pontian (=Messinian) de- ern Caucasus, Crimea, and Azerbaijan; the fossils posits of Abkhazia (Gabelaia 1976) associated to belonging to this genus were initially assigned to a brackish fish fauna. The second species, C. (?) Perca (Smirnov 1936) and, subsequently to Larim- tmutarakanica Bannikov, 2013, was found in the us (Daniltshenko 1960, 1980) and were referred Maeotian (=Upper Tortonian) diatomites exposed to Caucasisciaena based on a unique combination along the southern coast of the Taman Peninsula, of features (Bannikov et al. 2009). Several extinct SW Russia (Bannikov 2013). An additional croaker croakers are known from the Sarmatian s.l. (see specimen from the same locality (Bannikov 2013: Bannikov 2010, 2013), of both the Eastern and fig. 2) was not identified at genus and species level Western Paratethys. Two species are known from due to its poor preservation and incompleteness. the Sarmatian of Croatia, one of which from Dol- Apart from the known fossil croakers listed above, je not far from Zagreb, is described herein for the there is an undescribed largely incomplete articu- first time (see below); the other, Labrax (=Morone) lated skeleton from the Miocene of Coşniţa, Mol- multipinnatus from Sv. Nedelja, Samoborsko gorje, davia, housed in the Natural History Museum of was described as a member of the family Moro- Kishinev. Unfortunately, up to now none of the nidae (Anđelković 1989; Gorjanović-Kramberger fossil Paratethyan croaker skeletons were report- 1882) and is currently referred to the Sciaenidae ed to possess the otolith in situ, thereby prevent- (Bannikov 1992, 2010, 2013) and to the extant ing the recognition of any possible link between croaker genus Argyrosomus (see below). Sciaena pi- the skeletal and the abundant otolith records (see menovae was briefly described (but not illustrated) Schwarzhans & Carnevale 2017). from the Lower Sarmatian of Amvrosievka (south- The initial impetus for this paper was a dis- eastern Ukraine) together with an incomplete spec- covery by one of us (AFB) of a nearly complete imen of Sciaena sp. (?) (Bogatshov 1955). The loca- articulated skeleton of a croaker with the otolith tion of the storage of the specimens described by in situ in the right bank of the Pshekha River up- Bogatshov (1955) is currently unknown, and since stream of the Shirvanskaya settlement, Krasnodar that time no additional specimens were collected Region, NW Caucasus, SW Russia during the field at Amvrosievka. Another Sarmatian species, Sciae- campaign that took place in summer 2017. na knyrkoi, was described based on a single skeletal The fossil sciaenid skeleton exhibits an oto- impression from the vicinity of Grozny, Chechen lith in situ and pertains to a late juvenile or to a sub- Republic (Daniltshenko 1980; Bannikov 2010). adult individual exhibiting a unique combination Menner (1948) described an incomplete croaker of morphological features that allow the creation skeleton from the Kimmerian (=Zanclean) of the of a new previously undescribed taxon. The com- Kerch Peninsula (Crimea) as the extinct genus and parative analysis of the skeletal features of this fos- species Pseudoumbrina pcelini Menner, 1948. The ge- sil with other Paratethyan croakers revealed the ex- nus Pseudoumbrina is impossible to diagnose based istence of another undescribed croaker taxon from on the original description and figures because of the Sarmatian of Dolje, Croatia (mentioned above) incomplete preservation of available material, and and confirmed the sciaenid affinities of Labrax the Kimmerian taxon was tentatively referred by (=Morone) multipinnatus evidenced by Bannikov et Bannikov (2010) to an indeterminate taxon of the al. (2009). At the same time, the comparative anal- family Sciaenidae, together with “Sciaena pimenovae”. ysis of the otolith in situ highlighted that a taxo- The late Neogene sciaenid genus Carnevalella from nomic revision of the Neogene sciaenid otoliths the eastern Black Sea Region strongly differs from from the Paratethys cannot be furtherly delayed. the other known croakers by having a set of apo- This paper thus has three goals: (1) the description morphic characters (possession of 29 or 30 verte- of new Miocene croaker taxa from the Paratethys brae, four supraneurals, fourth hypural fused with based on articulated skeletal remains, in one case the terminal compound centrum, first dorsal-fin with the otolith in situ; (2) a comprehensive analysis Neogene Paratethyan croakers (Teleostei, Sciaenidae) 537 of the Neogene otolith record of the Paratethys, diagnoses of the family are based on a combina- with the description of three new genera and one tion of both derived