Scopelarchus aff. guentheri Alcock, 1896

Brzobohatý, Rostislav & Nolf, Dirk, 2018, Revision of the Middle Badenian fish otoliths from the Carpathian Foredeep in Moravia (Middle Miocene, Czech Republic), Cybium 42 (2), pp. 143-167 : 148-158

Scopelarchus aff. guentheri Alcock, 1896 ( Pl. 1 ,

Fig. 8a, b). An adult Scopelarchus otolith from Hrušovany nad Jevišovkou is well preserved, but damaged at the anterodorsal part; a second juvenile one from Židlochovice is somewhat eroded. The main features of these otoliths are the clearly divergent dorsal and ventral margins, an oblique anterior margin and a relatively small L/H index (1.3). The ostial colliculum is large; the cauda is elongated and narrow- ing towards the anterodorsal angle. These features are different from the Oligocene and Lower Miocene species S. nolfi Steurbaut, 1982 (e.g. Nolf and Brzobohatý, 1994b: pl. 4, figs 1-6) and from the Recent S. analis (Brauer, 1902) , also known from the Mediterranean Pliocene and Pleistocene ( Nolf and Cappetta, 1989; Girone et al., 2006). The otoliths are most similar to the present-day S. guentheri (see Rivaton and Bourret, 1999: pl. 129, figs 17-20). Additional and bet- ter-preserved material is required to decide about the precise specific affinity of the Badenian otoliths. Scopelarchids seem to be very rare in the Miocene of the Paratethys in contrast to the Oligocene Kiscell Clay and Pouzdřany Formation ( Nolf and Brzobohatý, 1994b; Brzobohatý and Krhovský, 1998). In the Recent fauna, S. guentheri lives mesopelagically (in the upper 150 m at night), mainly in tropical areas of the eastern Atlantic and the Indo-Pacific ( Froese and Pauly, 2016).

Sphagemacrurus ? sp. (Pl. 2, Fig. 9a, b). A single and perfectly preserved macrourine otolith from Drnovice (L = 4.4 mm; H = 3.3 mm; L:H = 1.3), with a strongly developed posterodorsal part. Its features, such as a strong posterodorsal expansion, the regular collicula, the absence of a collicular crest, and a smooth and flat ventral field with a relatively wide ventral furrow, show affinities with otoliths of the Indo-West Pacific Sphagemacrurus richardi (Weber, 1913) ( Nolf and Steurbaut, 1989: pl. 4, fig. G), which seem to be more extended in the dorso-ventral direction. Otoliths of the West Atlantic S. hirundo (Collett, 1896) have a higher and robust posterodorsal part as well as a deeper ventral rim ( Schwarzhans, 2013: pl. 16, fig. 13). More fossil material is required for an adequate taxonomic determination.

Melanonus triangulus ( Robba, 1970) (Pl. 2, Fig. 1a, b).

This species is relatively rare in the Moravian Badenian (see Tab. IC), but some of the otoliths are perfectly preserved and there is no doubt about the presence of M. triangulus in the Middle Badenian Sea. Melanonus triangulus occurs from the Lower Oligocene ( Nolf and Steurbaut, 1988; Brzobohatý and Krhovský, 1998 – both as M. gabbai ) to the Upper Miocene of the Mediterranean ( Robba, 1970: pl. 11, figs 7, 8, pl. 12, figs 1-3; Nolf and Steurbaut, 1983: pl. 3, figs 20, 21). Otoliths of this species show very close relationships with those of the Recent M. zugmayeri Norman, 1930 (e.g. Nolf and Steurbaut, 1983: pl. 4, fig. 1; Nolf, 1985: fig. 49 O; Smale et al., 1995: pl. 28, fig. C1; Campana, 2004: pl. 67). The juvenile and damaged specimen from the Burdigalian of the Aquitaine Basin, illustrated by Nolf and Brzobohatý (2002: pl. 7, fig. 6), may also belong to M. triangulus . A second present-day species of the genus, M. gracilis Günther, 1878 , differs markedly from M. triangulus by the irregular outline of the otoliths ( Nolf and Steurbaut, 1983: pl. 4, fig. 2-7). Two other fossil species, the Paleocene/Eocene M. ellesmerensis Schwarzhans, 1986 and the Pliocene M. paralyconus Schwarzhans, 1986 also present a significantly different outline. In the Recent fauna, Melanonus is widespread (Atlantic, Indian Ocean and Pacific, tropical to temperate or sub Antarctic waters) in deep-water (oceanic, meso- to bathypelagic).

Chaunax aff. pugetensis ( Schwarzhans, 1986) (Pl. 2,

Fig. 3a, b ). In the Badenian of the Carpathian Foredeep, chaunacid otoliths are very rare and are represented mainly by juvenile specimens, not diagnostic at species level. Two otoliths from Drnovice can be compared with the Pliocene species Ch. pugetensis ( Schwarzhans, 1986: pl. 5, figs 55, 56). The Moravian specimens are slightly different from the Pliocene otoliths in their somewhat higher form, the more backwards shifted dorsal tip and a wider and flatter ventral furrow. Other known species of Chaunax from the Paratethys are Ch. niederleisensis ( Schubert, 1906) from the Badenian of the Vienna Basin, which is known from the holotype only (see Nolf, 1981, 2013: pl. 146), and Chaunax sp. from the Badenian Korytnica Clay in Poland ( Radwańska, 1992: pl. 7, fig. 6). In the Recent fauna, the genus includes mainly bathydemersal deep-water fishes in the Atlantic and the Indo-Pacific, but it is absent from the Mediterranean.

Zenion hololepis (Goode & Bean, 1896) (Pl. 3, Fig. 9a,

b). Only a single and relatively well preserved specimen from Borač (L = 1.8 mm; H = 1.45 mm; L:H = 1.24) shows typical, but subadult features of otoliths of the Zenion group (sensu Nolf and Tyler, 2006), especially Z. hololepis (e.g. Smale et al., 1995: pl. 45, figs C1, C2; Nolf and Tyler, 2006: pl. 4, fig. 6). The oldest record of this species is from the Mediterranean Pliocene (SE France, Le-Puget-Sur-Argens; Nolf and Cappetta, 1989: pl. 14, figs 12, 13; Schwarzhans, 1986: pl. 2, fig. 18 – in the last case as Zenion sp. ). The genus Zenion has already been recognized from the Lower Oligocene of northern Italy (Z. sp., Nolf and Steurbaut, 2004: pl. 9, fig. 2). Otoliths of two other Recent species of the genus, Z. japonicus Kamohara, 1934 ( Nolf, 1985: pl. 55B, 2013: pl. 173; Nolf and Tyler, 2006: pl. 4, fig. 1) and Z. longipinnis Kotthaus, 1970 ( Rivaton and Bourret, 1999: pl. 144, figs 1-2; Nolf and Tyler, 2006: pl. 4, fig. 7), differ at first glance from those of Z. hololepis by their shape. Otoliths of the Recent Indo-Pacific species Z. leptolepis (Gilchrist and von Bonde, 1924) are not known. The Moravian record is the first indication of the presence of Zenion in the Miocene of the Central Paratethys and adds new data to the poorly known paleogeography of these fishes.

A list of the otolith-based fossil record of zeiforms was given by Nolf and Tyler (2006). Today, Z. hololepis belongs to the bathydemersal fauna of the Atlantic and the Indo-Pacific, where it lives between 180 and 650 m, but it does not occur in the Mediterranean.

Trachyscorpia aff. cristulata (Goode and Bean, 1896)

(Pl. 5, Fig. 6a, b) A single, large (L = 20.0 mm, H = 10.5 mm, T = 4.0 mm; OL:OH = 1.9), well-preserved and strongly compressed otolith from Brno-Královo Pole is larger than the known otoliths of the extant species of the genus (IRSNB collection; Smale et al., 1995: pl. 51, fig. A1-3). Its dorsal rim is convex and roughly undulated without prominent angles; the ventral rim is regularly concave. The posterior end is largely blunt and short; the rostrum is long and acu- minate. There is no antirostrum or excisura. The inner face is markedly convex, ventrally much higher than dorsally, with a prominent morphology. The dorsal area is flat; the ventral area is high, with a deep and wide ventral furrow. The sulcus is deep and wide, unclearly divided, with unregularly spread colliculum. The ostium is shallower and shorter than the cauda, slightly curved dorsally, unclearly opened ante- riorly. The outer face is flat with five distinct radial furrows and irregular bumps.

There are six extant species of this genus of marine deep-water scorpionfishes, living below 200 m. The Badenian specimen is rather similar to the otoliths from the East.

7 Moravia 1

Alpine-Carpathian Foredeep (Molasse) Poland P? P LA Lower Austria LA 24 Židlochov. 2 2 2 48 1 23 Železné 22 Žabčice 1 1 21 Vodȅrady 20 Šlapanice 19 Sudice 18 Rájec-Jestř. 17 Podbřežice 16 Přemyslov. 1 1 14 15 Nedvȅdice 14 Myslejovic.) paper 13 Lysice this 12 Lomnička, specimens 11 Lomnice 10 Kurim of number 9 Kralice n. O. 8 Hrušovany (Localities 7 Drnovice 6 Drahanovice? 1? 1 4 6 6 1 1 1 2 4 6 14 6 3 1 4 1 5 Cerná Hora 4 Čebín 3 Brno 2 Boskovice 1 Borač 7 5 1 3 15 3 6 2 1 1 Iconography Pl. 3, Fig. 11 Pl. 4, Fig. 4 Pl. 4, Fig. 5 Pl. 4, Fig. 8 Pl. 4, Fig. 7 Pl. 4, Fig. 3 Pl. 4, Fig. 2 Pl. 4, Fig. 1 Pl. 4, Fig. 6 Pl. 4, Fig. 12 Pl. 4, Fig. 13 Pl. 4, Fig. 10 Pl. 4, Fig. 11 Taxa * taxa mentioned in section 4.1 LABRIDAE Thalassoma sp. GOBIIDAE Deltentosteus telleri ( Schubert, 1906) Gobius sp. * Gobiidae sp. 1 * Gobiidae sp. 2 * Pomatoschistus ? sp. * Istigobius ? sp. Lesueurigobius vicinalis ( Koken, 1891) * Priolepis ? sp. SPHYRAENIDAE Sphyraena sp. TRICHIURIDAE Aphanopus ? sp. CAPROIDAE * Antigonia aff. capros Lowe, 1843 Capros ? sp. SOLEIDAE * Aseraggodes sp.

Continued

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I

Table Atlantic and West Mediterranean species T. cristulata (see Nolf, 2013: pl. 177), but differs from them by the more rounded posterior margin and the perspicuous thickness of the posterior part of the otolith.

Our specimen represents the first otolith-based fossil record of the genus and the whole tribus Sebastolobini ( Nolf, 2013) . The fish must have measured about 50 cm of TL (compare data by Smale et al., 1995; Quéro et al., 2003).

Antigonia aff. capros Lowe, 1843

(Pl. 4, Fig. 13a, b). Only subadult otoliths of an Antigonia type were found in the Badenian of the Carpathian Foredeep. The largest, from Židlochovice (Pl. 4, Fig. 13), could be fully compared with the holotype of “ Ot. (Monocentridarum) altus ” described by Weiler (1950: pl. 2, fig. 7) from the same stratigraphic level of the Transylvanian Basin and synonymized by Steurbaut (1984) with A. aff. capros .

Recent A. capros lives demersal (with juveniles in the midwater) in the tropical and subtropical waters of the Western and Eastern Atlantic and the Pacific between 50-900 m.

Gobiidae (notice): Gobiid otoliths are very rare in the studied calcareous clays, in comparison with myctophid or macrourid ones, and are represented mainly by juvenile or eroded specimens. There are only two nominally valid species support- ed by a collection of well-preserved material: Lesueurigobius vicinalis ( Koken, 1891) and Deltentosteus telleri ( Schubert, 1906) – see pl. 4, figs 1, 4. Considering the restricted knowledge of Recent gobiid otoliths ( Girone et al., 2010) the following taxa are mentioned only at generic level.

Gobius sp. (Pl. 4, Fig. 5a, b). Material: Borač (7 ex.), Černá Hora (1 ex.), Drnovice (6 ex.), Kralice n. O. (1 ex.), Lomnička u T. (7 ex.), Židlochovice (2 ex.). Moder- atly elongated, predominant juvenile or corroded gobiid otoliths with outstanding postdorsal and praeventral projection and relatively clear postcaudal iugum. In the Central Paratethys this type of otoliths was usually assigned to the taxon “ Gobius ex gr. multipinnatus (H. v. M.)” (e.g. Brzobohatý et al., 2007) whose taxonomical status is based only on the skeletons after the revision published by Gierl and Reichenbacher (2015). Their affinity with the species G. reichenbacherae from the Serravallian of the Karaman Basin ( Turkey) has also been stated by Schwarzhans (2014). The unsufficient Moravian material does not allow for any more precise systematic conclusions.

Gobiidae sp. 1 (Pl. 4, Fig. 8a, b). Material: Borač (5 ex.), Drnovice (14 ex.), Lomnice u Tišnova (4 ex.). Elongated gobiid otoliths with a perspicuous notch in the dorsal margin seem to be very close to the ones depicted as “genus Gobiidarum ” sp. 3 from the Catalunyan Miocene ( Hoedemakers and Batllori, 2005: pl. 11, figs 5-8). But the praedorsal part of the Badenian specimens is more developed and their sulcus is clearly located horizontally.

Gobiidae sp. 2. (Pl. 4, Fig. 7a, b). Material: Borač (1 ex.), Drnovice (6 ex.). Well preserved rectangular otoliths (length ~ 2.5 mm) with rounded postero- and anterodorsal angles, notched middle part of the dorsal margin and a nearly horizontal sulcus. Otoliths with a very similar morphology are figured under different names, e.g., from the Italian Messinian ( Girone et al., 2010: pl. 10, figs j1a,b, j4, j5). By no means are our otoliths of Gobius sp. 2 conspecific with the ones described as Thorogobius intimus (Prochazka, 1893) by Schwarzhans (2010: p. 265, pl. 106, figs 11-13) from the Badenian of Slovakia and Austria, which have an oblique anterodorsal margin.

Pomatoschistus ? sp. (Pl. 4, Fig. 3a, b ). Material: Židlochovice – two well preserved and (?) juvenile specimens. These otoliths seem to be conspecific with those described by Radwanska (1992: p. 292, pl. 35, figs 3-4, texfig. 150a-d) as “ Ot. Gobiidarum sp. 5 ” from the Badenian of Korytnica.

Istigobius ? sp. (Pl. 4, Fig. 2a, b ). Material: Borač (three specimens), Žabčice (1 ex.). One figured otolith is well preserved and comparable with the otoliths of the present day species I. decoratus (Herre, 1927) and I. ornatus (Rüppel, 1830) – see Smale et al., 1995: pl. 129, figs C and D, respectively, and Rivaton and Bourret, 1999: pl. 79, figs 9-10. Both species are widespread in the Indo-Pacific waters including the Red Sea and the presence of this genus in the Middle Badenian Sea can be considered as probable. Otoliths described as “ genus Gobiidarum ” sp. 2 from the Burdigalian/Langhian of the Catalunya ( Hoedemakers and Batllori, 2005: pl. 12, figs 3-7) also show a very similar morphology.

Priolepis ? sp. (Pl. 4, Fig. 6a, b, c). Three gobiid otoliths from Borač show the following characters: a prominent anterior margin; a nearly vertical posterior margin with a small notch in its center; a horizontal sulcus; a flat ventral field with a ventral ridge and a deep and wide depression in the dorsal area. These characters, together with the ventral and posterior views, are quite similar to what is seen in otoliths of the West Atlantic P. hippoliti (Metzelaar, 1922) (see Nolf and Brzobohatý, 2009: pl. 8, fig. 4) and of P. cincta (Regan, 1908) (see Smale et al., 1995: pl. 129 H) from the Indo-Pacific realm, including the Red Sea. The otoliths figured by Brzobohatý et al. (2007: pl. 8, figs 9-11) under the name Priolepis sp. from the Badenian of the Vienna Basin belong to another genus (see Nolf and Brzobohatý, 2009: p. 329).

Aseraggodes sp. (Pl. 4, Fig. 11a, b). A slightly eroded flatfish otolith from Borač has a sulcus, with more or less cir- cular colliculi and shows some similarity with otoliths of the present day A. cyaneus (Alckock, 1890) (see Schwarzhans, 1999: figs 859-861) and may belong to the same genus. There is only one other fossil species of Aseraggodes : A. laganus Girone & Nolf, 2009 from the North Italian Upper Eocene, which is characterised by a clearly longer ostium. The genus Aseraggodes does not live in the present day Mediterranean, but many Recent species are widely distributed throughout the Indo West Pacific.

Description of new species

Order ATELEOPODIFORMES Berg, 1937 Family ATELEOPODIDAE Kaup, 1858 Genus Ijimaia Sauter, 1905 Ijimaia rara n. sp. Pl. 5, Fig. 7a, b

Ijimaja sp. – Nolf, 2013: p. 48, pl. 53

Derivatio nominis: rarus, a, um = scarce, refers to the solitary occurrence of the holotype.

Holotype: a left otolith (Pl. 5, Fig. 7), deposited in the collections of the Masaryk University ( DGS MU Inv. Nr. O 373).

Stratum typicum: calcareous clay of the Badenian (= Langhian, Middle Miocene) at Borač (Carpathian Foredeep, Middle Moravia, Czech Republic).

Dimensions: L = 10.9 mm; H = 8.4 mm; T = 2.9 mm; OL:OH = 1.3

Diagnosis

This species is characterized by thick, sub rhomboidal otoliths showing a nearly rectangular ventral half with a smooth surface. The dorsal half shows a strong posterior expansion, and the whole upper rim is lobated. The lobes are separated by well-incised radial furrows that continue down- wards to near the crista superior. The sulcus is not divided in an ostial and a caudal portion and both the crista superior and the crista inferior are very salient, resulting in a sulcus, that is somewhat extruding from the inner face. The crista inferior has a spine-like anterior expansion.

Figures

Plate 1. – Explanation of the plates:All figured specimens are deposited in the collections of the Department of Geological Sciences, Masa- ryk University (DGS MU), Brno, Kotlářská 2, Czech Republic.The fossil otoliths bear numbers of the collection of types and figured fossil fish specimens in the DGS MU (Inv. Nr. O 315-377). The annotations a, b and c are used to indicate ventral, inner (= mesial) and posterior views, respectively. Figures with only numbers and no letter show inner views. In the captions, L stands for left otolith and R for right oto- lith. 1: Conger aff. conger (Linnaeus, 1758); Drnovice, Badenian, L (DGS MU Inv. Nr. O 315). 2: Serrivomer sp.; Lomnička, Badenian, R (DGS MU Inv. Nr. O 316). 3: Pterothrissus umbonatus (Koken, 1884); Drnovice, Badenian, R (DGS MU Inv. Nr. O 317). 4: Bathycongrus nagymarosyi Nolf & Brzobohaty, 1994; Voděrady, Badenian, R (DGS MU Inv. Nr. O 318). 5: Glossanodon cf. leioglossus (Valenciennes, 1848); Lomnice u Tišnova, Badenian, R (DGS MU Inv. Nr. O 319). 6: Sagamichthys? sp.; Voděrady, Badenian, R (DGS MU Inv. Nr. O 320). 7: Sarmatella aff. pulchra (Smigielska, 1966); Lomnice u Tišnova, Badenian, L (DGS MU Inv. Nr. O 321). 8: Scopelarchus aff. guentheri Alcock, 1896; Hrušovany nad Jevišovkou, Badenian, R (DGS MU Inv. Nr. O 326). 9: Etrumeus? weileri (Smigielska, 1966); Borač, Badenian, R (DGS MU Inv. Nr. O 323). 10: Bregmaceros albyi (Sauvage, 1880); Lomnice u Tišnova, Badenian, R (DGS MU Inv. Nr. O 324). 11: Physiculus aff. huloti Poll, 1953; Lomnice u Tišnova, Badenian, L (DGS MU Inv. Nr. O 325). 12: Nansenia sp.; Borač, Badenian, L (DGS MU Inv. Nr. O 322). 13: Merluccius aff. merluccius (Linnaeus, 1758); Lomnice u Tišnova, Badenian, L (DGS MU Inv. Nr. O 327).

Figure 2. – Stratigraphic charts of the Badenian (after Hohenegger et al., 2014) and position of the localities studied.

Figure 3. – Present-day bathymetric range of benthic and benthopelagic taxa (without pelagic taxa) (Jardas, 1996; Quéro et al., 2003; Froese and Pauly, 2016) represented in the Badenian clay of the Borač Section (compare Brzobo- hatý, 1997).