Swiss Journal of Palaeontology (2019) 138:99–108 https://doi.org/10.1007/s13358-018-0179-4 (0123456789().,-volV)(0123456789().,-volV) REGULAR RESEARCH ARTICLE Diversity and distribution of Miocene–Pliocene sepiids (Cephalopoda) in the Mediterranean area, with new records from Italy and Turkey 1 2 3 Martin Kosˇt’a´k • John W. M. Jagt • Jan Schlo¨ gl Received: 14 September 2018 / Accepted: 17 November 2018 / Published online: 30 November 2018 Ó Akademie der Naturwissenschaften Schweiz (SCNAT) 2018 Abstract Two new records of Cenozoic sepiids from the Mediterranean area are described and discussed. While the single, fragmentary specimen from Pliocene (Zanclean) strata in northwestern Italy can only be identified generically as Sepia sp., the Turkish Sepia vandervoorti sp. nov., of late Langhian–earliest Serravallian (Middle Miocene) age, constitutes a new species. The diversity and disparity of Cenozoic sepiids from the Mediterranean area are discussed within palaeobio- geographical and stratigraphical contexts. Current cuttlefish records from the Central Paratethys, the Atlantic coast and the North Sea Basin provide indications of migration patterns of the genus Sepia during the Miocene. A synthesis of published data, complemented by the new records presented here, allows us to reconstruct sepiid distribution in time and space. The highest diversity and morphological disparity are seen during the Middle Miocene; this is followed by a rapid decrease during the Late Miocene and a renewed radiation during the Pliocene. Keywords Sepia Á Palaeobiogeography Á Southern Europe Á Near east Á Cenozoic Á New species Introduction the Central Paratethys, Australia and the North Sea Basin (Kosˇt’a´k et al. 2016, 2017; Kosˇt’a´k and Jagt 2018), as well In general, cuttlebones of extinct sepiids related to the as from the Lower Pleistocene of central Italy (Pasini et al. genus Sepia Linnaeus 1758 are considered to be rare in the 2014). Extant cuttlefish evolved from a stem lineage of fossil record. In this respect, each new record of fossil Recent Sepia, which derived from Oligocene representatives constitutes a notable contribution to the archaeosepiids (Kosˇt’a´k et al. 2017). The majority of Oli- evolutionary history and palaeobiogeography of sepiids. gocene records (archaeosepiids) come from Hungary and During recent years, new records have been described from Italy (Lorenthey} 1899; Szo¨re´nyi 1933; Wagner 1938; Fornasiero and Vicariotto 1995), where these forms made their first appearance approximately following the extinc- Editorial Handling: D. Fuchs. tion of another group of extinct sepiids, the belosaepiids. Consequently, the origin of extant cuttlefish probably is & Martin Kosˇt’a´k linked to the Paratethys and Mediterranean areas. The [email protected] highest diversity, as well as the greatest morphological John W. M. Jagt disparity amongst fossil sepiids, is seen during the Middle [email protected] Miocene (Kosˇt’a´k et al. 2016). It is notable that the highest Jan Schlo¨gl sepiid diversity worldwide is seen in the Recent (Reid et al. [email protected] 2005); the same holds true for a very high morphological 1 Institute of Geology and Palaeontology, Faculty of Science, disparity of cuttlebones (Neige 2003a, b, 2006; Neige and Charles University, Prague, Albertov 6, 128 43 Prague 2, von Boletzky 1997). Czech Republic Historically, ancestral sepiids from the Mediterranean 2 Natuurhistorisch Museum Maastricht, De Bosquetplein 6-7, area were studied by several scholars during the nineteenth 6211 KJ Maastricht, The Netherlands century (e.g. Gastaldi 1866; Bellardi 1872; Ponzi 1876; 3 Department of Geology and Paleontology, Faculty of Natural Tiberi 1880; Issel 1889; Foresti 1890; Parona 1892). Later Sciences, Comenius University in Bratislava, Mlynska work includes that by Sacco (1904), Lipparini (1934), dolina, Ilkovicˇova 6, 842 15 Bratislava, Slovakia 100 M. Košt’ák et al. A´ lvarez Ramis and Mele´ndez (1966), Hewitt and Pedley the presence of median and lateral keels, the character of (1978), Abad et al. (1981), Cavallo and Repetto (1992) and the apical part [‘‘rostrum’’], the shape of septal suture lines others. The most recent papers are those by Gaudant et al. and the arrangement of granules on the dorsal shield) fol- (2007), Okan and Hos¸go¨r(2009) and Pasini et al. (2014). lows previous systematic treatments by Bizikov (2008), On the basis of published data and new records from Italy Kosˇt’a´k et al. (2016) and Kosˇt’a´k and Jagt (2018). Sche- and Turkey, we here summarise the stratigraphical distri- matic general shapes of cuttlebones are shown in Fig. 1. bution and palaeobiogeographical aspects of extinct sepiids from the Mediterranean area. Systematic palaeontology Geological, geographical and stratigraphical setting Subclass Coleoidea Bather, 1888 Order Sepiida Gray, 1849 Turkey: marine strata in the Mut Basin have been subdi- Family Sepiidae Keferstein, 1866 vided into five units, namely the Mut, Ko¨selerli, Dag˘pazarı, Tirtar and Ballı formations (Atabey et al. 2000). The Mut Genus Sepia Linnaeus, 1758 and Ko¨selerli formations are the oldest units in this Sepia vandervoorti sp. nov. sequence and frequently demonstrate lateral interfingering; they have been dated as late Burdigalian and Langhian Figures 2, 3a–g (Mut Formation) to earliest Serravallian (Ko¨selerli For- Diagnosis Smaller-sized sepiid with rhomboid cuttlebone mation). The Mut Formation comprises mainly limestones, possessing fine granulation, relatively wider outer cone, while the Ko¨selerli Formation consists of clayey limestone oval-rounded inner cone with fork and well-developed and marl. The new sepiid was collected from grey marls of prong. Median and lateral fields without ornamentation. the latter formation and can be dated as late Langhian– earliest Serravallian. This age assignment of the marls is Types The holotype, and sole specimen known to date, is confirmed by the abundant material of the pteropod Vagi- RGM.1351051.a (part) and RGM.1351051.b (counterpart), nella austriaca Kittl, 1886 and the gastropod Sulcogladius in the collections of Cenozoic Mollusca at Naturalis Bio- collegnoi (Bellardi and Michelotti, 1841) var. pluridentic- diversity Center (Leiden, the Netherlands; formerly ulata Sacco, 1893, which was originally described from the Rijksmuseum van Geologie en Mineralogie). Turin hills in Italy as characteristic of the ‘‘Helvetian’’; this Derivatio nominis In honour of Mr. Jaap van der Voort currently corresponds to the Langhian/Serravallian. (Ostercappeln-Venne, Germany), who found the specimen Italy: Montaldo Roero (Trinita`, province of Cuneo, in May 1989. Italy; co-ordinates: 44°4504400N–7°5601600E) from Lower Pliocene (Zanclean) strata. Locus typicus Outcrop on the eastern side of the road (D- 715) from Karaman to Mersin, between the northern margin of the village of Palantepe with the mosque and the village Materials and methods itself (co-ordinates: 36°36053.5000N–33°26028.6000E). This locality is within the Mut Basin, which comprises strata of The Turkish specimen (part and counterpart, in matrix) was Miocene age (see below). discovered and collected (May 1989) near the village of Stratum typicum Ko¨selerli Formation (late Langhian–ear- Palantepe (near Mut, Mersin Province, southern Turkey) by liest Serravallian). Mr. Jaap van der Voort (Ostercappeln, Germany) from grey marls of the Ko¨selerli Formation, which is dated as late Description: Complete specimen with an oval to rather Langhian–earliest Serravallian. The fragmentary specimen rhomboid cuttlebone (Figs. 2, 3a, b) with partially pre- from northwestern Italy (Montaldo Roero, province of served shell (Fig. 3a); a smaller-sized member of Sepia Cuneo) originates from Lower Pliocene (Zanclean) strata (L = 51 mm), the widest part (* 18 mm) being situated and is completely free of matrix. approximately in the middle part. Dorsal surface with Specimens were measured and studied using the com- slightly developed median keel, widening anteriorly under puter graphic programs CorelDRAW X5 and CorelPho- a low angle (7°). Where preserved, the cuttlebone surface toPaint X5. Photographs were taken with a digital camera (shell remnants preserved in the posterior part only) shows (Canon EOS D600D). The terminology used herein is regularly arranged fine granules (Fig. 3f). No prominent based mainly on the preserved dorsal parts of the cuttle- granules are visible. bone (i.e. more specifically, the shape of the dorsal shield, Miocene–Pliocene sepiids (Cephalopoda) from the Mediterranean area 101 Fig. 1 General cuttlebone shape (dorsal view; not to scale); a prolonged oval; b oval; c oval- rhomboid; d rhomboid; e oval- rhomboid to elliptical with posterior wings expanded (from Kosˇt’a´k et al. 2016) Lateral fields (diverging at an angle less than 38°) are from the flattened lateral sides. Anterior margins slightly well developed, lacking ornamentation or ‘false’ lateral splayed. Septal suture lines parallel and rhomboidal keels. Lateral fields only slightly separate the median field (Fig. 3e) in the anterior part, semi-circular–oval in early 102 M. Košt’ák et al. ontogenetic stages showing significant changes in shape during ontogeny. Originally chitinous rims are absent. Calcareous rims are only slightly indicated. Well-calcified remains of inner and outer cones are quite well preserved (Fig. 3d). Outer cone relatively wide, possessing fine striation. Pillar structure imprints (intracameral walls) are seen only in one place of the cuttlebone (Fig. 3e–g), revealing the typical arrangement