Diplodus karrerae Nolf and Steurbaut, 1979

Diplodus karrerae Nolf and Steurbaut, 1979

Figure 7 View FIGURE 7 AG-AL

1979 Diplodus karrerae ; Nolf and Steurbaut, pl. 3, figs. 19–21.

2010 Diplodus karrerae Nolf and Steurbaut, 1979 ; Schwarzhans, pl. 90, figs. 3–4.

2013 Diplodus karrerae Nolf and Steurbaut, 1979 ; Schultz, pl. 86, fig. 9a-b.

2014 Diplodus karrerae Nolf and Steurbaut, 1979 ; Schwarzhans, pl. 6, fig. 4.

2015 Diplodus karrerae Nolf and Steurbaut, 1979 ; Lin et al., fig. 5.21.

2022 Diplodus karrerae Nolf and Steurbaut, 1979 ; Brzobohatý et al., pl. 2, fig. V-W.

Material. 20 otoliths: 1 specimen, NMNHU-P PI

2752, Shydlivshchyna; 11 specimens, NMNHU-P

PI 2753 , Novyi Pliazh; 5 specimens , NMNHU-P PI

2754, Staryi Zavod; 3 specimens, NMNHU-P PI

2755, Skala.

Discussion. The otoliths of Diplodus karrerae are widely distributed in the Middle Miocene of Europe but rarely common. The occurrence of this species in the late Badenian of the Central Paratethys probably indicates a re-immigration from the Mediterranean of this stenohaline marine species. It represents the latest record in the Paratethys.

Part 2. Fish Bones (by Dubikovska and Kovalchuk)

Non-otolith fish remains in the material from the localities studied are considerably less numerous compared to otoliths obtained from the same layers. Almost all of them are represented by isolated teeth, accompanied by a single jaw fragment and one dermal denticle from Lisohirka as well as one vertebra from Skala. Non-otolith remains are also less diverse taxonomically.

Dasyatidae indet. (order Myliobatiformes , class Chondrichthyes) – 1 tooth, NMNHU-P 2771, Shydlivshchyna. The tooth crown bears transversal crest separating the labial and lingual surfaces. The outline of the crown is triangular. Its lingual surface is almost vertical in profile view and concave just below the transversal crest. The labial surface is smooth and curved, with a rounded labial edge. There is a small cusp at the top of the crown, which is crossed in the middle by a small furrow. The hoof-shaped root is shifted posteriorly to the lingual surface, its lobes are well separated, slightly protruding, and triangular in vertical projection. Considering the morphology of the specimen (cf. Cappetta, 2012), it belongs to a female individual of the family Dasyatidae .

Labrodon sp. (family Labridae , order Labriformes , class Actinopterygii) – 1 tooth, NMNHU-P PI 2761, Skala. A small molariform tooth has a triangular outline. Its occlusal surface bears a low central papilla, which is clearly visible in a profile view. This specimen has been identified as Labrodon sp. since it is morphologically like those described and figured in Schultz (1979), Szabó and Kocsis (2020), Szabó et al. (2021) from the Central Paratethys, although we do not exclude the possibility of its belonging to the genus Coris (Carnevale, 2015) . The latter is represented in the same layer by an otolith of C. medoboryensis (Schwarzhans et al., 2022) .

Trigonodon jugleri (Münster, 1846) (family Labridae , order Labriformes , class Actinopterygii) – 2 teeth: NMNHU-P PI 2762, Skala; NMNHU-P PI 2763, Staryi Zavod. Both anterior teeth are rounded triangle (chisel-like) in shape and share incisiform morphology. The specimens are flattened labiolingually and have an even cutting edge. Their labial surface is vertical and slightly concave, lacking ornamentation (as is the smooth lingual surface). The teeth morphologically resemble those in representatives of the genus Trigonodon , and most probably belong to Trigonodon jugleri (Münster, 1846) . This species first appeared in the Early Miocene of Italy and Austria (Schultz and Bellwood, 2004; Marsili et al., 2007), and it was widely distributed within the Central Paratethys during the Middle Miocene (Schultz, 1998; Schultz et al., 2010; Schultz, 2013; Bellwood et al., 2019; Szabó and Kocsis, 2020; Szabó et al., 2021).

Diplodus sp. (family Sparidae , order Spariformes , class Actinopterygii) – 1 tooth, NMNHU-P PI 2764, Lisohirka. The anterior tooth has an incisiform morphology. Its crown is quadrangular, flattened to slightly convex labially, concave lingually, and has a smooth surface. The left cutting edge of the tooth is higher than the right one in lingual view. The root was broken just at the base.

Pshekharus yesinorum Bannikov and Kotlyar, 2015 (family Sparidae , order Spariformes , class Actinopterygii) – 1 tooth, NMNHU-P PI 2765, Skala. A single tooth has a central papilla and probably represents an intermediate morphotype. Its crown is conical, as high as wide, what is a characteristic feature of the genus Pshekharus (see Bannikov and Kotlyar, 2015 for details).

Sparidae indet. (order Spariformes , class Actinopterygii) – 1 jaw fragment, NMNHU-P PI 2766, Lisohirka. 47 teeth: 8 specimens, NMNHU-P PI 2767 , Lisohirka ; 1 specimen, NMNHU-P PI 2768 , Shydlivshchyna ; 31 specimens, NMNHU-P PI 2769 , Staryi Zavod ; 7 specimens, NMNHU-P PI 2770 , Skala. There is a rounded opening on the labial side of the jaw. The latter bears one conical (intermediate) tooth and numerous hemispherical (molariform) teeth. Anterior and intermediate teeth are conical, higher than wide, circular in cross-section, and have a rounded pulp cavity. Some of them have a cylindro-conical crown. The apical and basal parts of the latter are separated by a furrow. Molariform teeth with hemispherical or oval crowns are of different sizes .

In addition to the specimens mentioned above, there are some fossils whose precise taxonomic identification is impossible due to their poor preservation or too general morphology (i.e., lacking reliable diagnostic characters). Those are one dermal denticle of a shark (Selachimorpha indet.) NMNHU-P PI 2773, one ray tooth (Batoidea indet.) NMNHU-P PI 2774 from Lisohirka, a single anterior conical tooth of actinopterygian fish NMNHU-P PI 2772 from Shydlivshchyna, as well as one actinopterygian vertebra NMNHU-P PI 2775 from Skala.

Part 3. Other Biota (by Ryabokon, Kovalenko, and Klots)

In addition to the otoliths and other fish skeletal elements described above, rich foraminiferal and ostracod assemblages have also been obtained from the respective rock samples.

Foraminiferan fauna was studied to clarify the stratigraphic position of the samples obtained from the new localities. The list of identified species is presented in Table 2 along with data on the distribution of particular taxa in the Badenian of the Central Paratethys (Cicha et al., 1998; Garecka and Olszewska, 2011; Peryt et al., 2021), Konkian of the Eastern Paratethys (Krasheninnikov et al., 2003; Vernyhorova, 2018; Vernyhorova et al., 2023), and Middle Miocene of the Volyn –Podolia region (Kudrin, 1966; Didkovsky and Satanovskaya, 1970; Goretsky and Didkovsky, 1975; Andreeva-Grigorovich et al., 1996; Gedl and Peryt, 2011; Peryt et al., 2021; Schwarzhans et al., 2022). The studied foraminifers are similar in composition and consist exclusively of benthic forms, including various and numerous miliolids ( Quinqueloculina , Triloculina , Pseudotriloculina, Pyrgo , and others) and representatives of the genera Elphidium , Ammonia, Lobatula, Heterolepa, Cibicidoides, Asterigerinata , Globulina , Guttulina , frequent Cancris, Melonis, Reussella, Porosononion , also present Borelis and Sphaerogipsina. All of them are marine species including those strictly associated with reefal environments while brackish taxa are absent. In addition, several permanently motile foraminifers recognised in the studied samples (in particular, Elphidium crispum , E. macellum , Globulina gibba, Nonion depressulum, Reussella spinulosa , Spiroloculina canaliculata , and Triloculina gibba ) and representatives of the genera Ammonia, Anomalinoides, Cibicides, Discorbis, Pyrgo , Quinqueloculina , and Rosalina are commonly associated with seagrass or seaweeds (Langer, 1993; Moissette et al., 2007).

The studied foraminifera are characteristic of the Badenian (Middle Miocene, early Serravallian) of the Central Paratethys (Cicha et al., 1998). A specific feature of these assemblages is the presence of some endemic taxa distributed in the Volyn –Podolia region and the Carpathian Foredeep: Nodobaculariella podolica , Sigmomorphina karpatica , Pseudopatellinoides primus, Cibicides menneri, Nonionella ventragranosa, Ammonia galiciana , A. pseudobeccarrii , Elphidium stellans , and E. ukrainicum (Didkovsky and Satanovskaya, 1970; Goretsky and Didkovsky, 1975).

Nearly half of the foraminifera species identified in the studied samples are also common in the Konkian of the Eastern Paratethys, and two-thirds of them ( Table 2) are known from the Ternopil and Vyshgorod beds (=upper Badenian) of the Volyn- Podolia region (Kudrin, 1966; Didkovsky and Satanovskaya, 1970; Goretsky and Didkovsky, 1975; Schwarzhans et al., 2022). We therefore conclude that the studied samples date the late Badenian.

Ostracods were recognised in rock samples obtained from Lisohirka, Skala, and Staryi Zavod. The general species list includes 20 taxa ( Table 3), of which the most common (i.e., present in the material from all the studied localities) are Aurila cicatricosa and A. novata ; less frequent are Cyclocypris regularis , Aurila convexa , Loxoconcha spongiosa , Cytheridea muelleri , and Cnestocythere truncata , while all other species each come from a single locality ( Table 3).

Most ostracods from the Middle Miocene of the Medobory backreef preferred normal marine conditions. The presence of Cytheridea muelleri and Leptocythere ex gr. canaliculata in the material from Staryi Zavod and Lisohirka indicates higher than normal salinity (ca. 15–17‰) and warm waters. Only two species ( Cyclocypris regularis from Lisohirka and Skala and Cyclocypris laevis from Skala) are indicators of a limited freshwater influx. The presence of five species of the genus Aurila , two species of the genus Loxoconcha , and two species of the genus Xestoleberis in the material from Lisohirka, Skala, and Staryi Zavod ( Table 2), in addition to epiphytic foraminifers mentioned above, suggest a seagrass environment (Pisera, 1985; Aiello and Szczechura, 2004; Moissette et al., 2007; Cornée et al., 2009; Forsey, 2016). The studied ostracod assemblages comprise a number of species distributed in the Central Paratethys during the Early Miocene (Kollmann, 1971; Zorn, 1998, 2003; Tunoğlu and Bilen, 2001), Badenian (Paruch-Kulczycka, 1992; Szuromi-Korecz and Szegő, 2001; Aiello and Szczechura, 2004; Zorn, 2004; Szczechura, 2006), Sarmatian ( Tóth, 2008), and Pannonian (Stancheva, 1962, 1963), although they are more similar taxonomically to those from the Konkian of the Eastern Paratethys (Schneider, 1953, 1959; Didkovsky, 1959, 1964; Ljuljev, 1967, 1969; Bondar, 2006; Kovalenko, 2013).

Gastropods are dominated by turitellid forms throughout the sampled localities, particularly species of the genera Terebralia, Thericium , and Tiaracerithium ( Figure 8 View FIGURE 8 Terebralia and Tiaracerithium are typical for mudflats (Harzhauser et al., 2023); the other observed genera are widely distributed in sublittoral environments. Lucinid bivalves of the genera Lucina , Lucinoma ( Figure 8 View FIGURE 8 ), and Loripes are common in seagrass environments (e.g., van der Heide et al., 2012; Stanley, 2014) and are common in the material from Novyi Pliazh, Skala, and Horodok being represented by both large (more than 5 mm) and small shells. These molluscs were also observed, albeit rarely, in all other studied localities (Kozatskyi Yar, Lisohirka, Mlyntsi, Shydlivshchyna, and Staryi Zavod). Their distributon and abundance are not entirely congruent with those of foraminifera, ostracods, and fishes, which, however, in combination are more indicative in our opinion than the distribution pattern of the Lucinidae in the Medobory backreef environments.