DIVERSITY, MORPHOLOGY, AND PHYLOGENY OF COLEOID CEPHALOPODS FROM THE UPPER CRETACEOUS PLATTENKALKS OF LEBANON–PART II: TEUDOPSEINA DIRK FUCHS 1 AND NEAL LARSON 2 1 Freie Universita ¨ t Berlin, Institute of Geological Sciences, Branch Palaeontology, Malteserstr. 74-100, D-12249 Berlin, Germany, ,drig@zedat.fu-berlin.de.; and 2 Black Hills Institute of Geological Research and Black Hills Museum of Natural History, PO Box 643, 117 Main Street, Hill City, SD 57745, USA, ,ammoniteguy@bhigr.com. ABSTRACT—Morphologic analyses of a large quantity of teudopseid coleoids from the Upper Cretaceous (Cenomanian) of Lebanon has yielded a much higher diversity than previously assumed and revealed numerous extraordinarily well-preserved soft-part characters. The Teudopseina is represented by three families (only the Muensterellidae are still unknown in the Lebanon Plattenkalks). The Teudopsidae is represented by one species (Teudopsinia haasi), the Trachyteuthididae by five species (Trachyteuthis bacchiai n. sp., Glyphiteuthis libanotica, Gl. abisaadiorum, Gl. freijii n. sp., and Glyphidopsis waagei n. gen. n. sp.), and the Palaeololiginidae by one species (Rachiteuthis donovani). Gl. freijii n. sp. represents the first record of a coleoid cephalopod from Na ˆ mmoura. A detailed comparison with Jurassic gladii shows that the Teudopseina is a very homogenous group that can be easily differentiated from other Mesozoic, gladius-bearing groups (Prototeuthidina, Loligosepiina). Despite comparatively large gaps in the fossil record, each of the Late Cretaceous gladii can be readily associated with Jurassic precursors. Particularly, the first record of two pairs of fins in Glyphiteuthis exemplifies that in addition to gladius similarities, similarities in soft-parts exist as well. A previously undescribed specimen of Rachiteuthis exhibits unusually well- preserved gills. In general, the soft-part morphology of the Teudopseina clearly indicates octobrachiate affinities (i.e., absence of tentacles, circular suckers, cirri, two pairs of fins, octopod-like eye capsules); no existing evidence supports a decabrachiate relationship. INTRODUCTION T EUDOPSEINA STAROBOGATOV, 1983 is an extinct group of coleoid cephalopods, that appeared in the Early Jurassic and became extinct during the latest Late Cretaceous (Fuchs, 2006b, 2009, 2010; Fuchs et al., 2007, 2010; Fuchs and Schultze, 2008; Fuchs and Weis, 2009, 2010). This highly diverse and widely distributed group regularly attracts interest because teudopseid sub-groups have been regarded to represent the stem-groups of phylogenetically non-related present-day groups. Bizikov (2008, fig. 208) hypothesized ancestor-descen- dent relationships between trachyteuthidid teudopseids and sepiids; between muensterellid teudopseids and sepiolids, sepiadariids, as well as idiosepiids; and between palaeololi- ginid teudopseids and loliginids as well as oegopsids. Ideas of close relationships between Trachyteuthididae and Sepiidae are not new, particularly with the Sepia-like granulation on the dorsal surface of the trachyteuthidid gladius, such as in Trachyteuthis and Actinosepia that led many nineteenth century workers to assume similar phylogenetic relationships (Mu ¨nster, 1837; Ferrusac and Orbigny, 1835; Whiteaves, 1897). Instead, the soft-parts of Jurassic teudopseids provide evidence of octobrachiate affinities (Bandel and Leich, 1986; Donovan et al., 2003; Klug et al., 2005; Fuchs, 2006b; Fuchs et al., 2003, 2007). Additional material from the Cenomanian Plattenkalks of Lebanon, which are well-known for their extraordinary soft- part preservation (Woodward, 1883; Roger, 1946; Fuchs, 2006b; Fuchs et al., 2009; Fuchs and Weis, 2009; Larson et al., 2010; Fuchs and Larson, 2011), to date, provide data on the soft-part morphology of Late Cretaceous teudopseids that has been neglected in this important debate. This present study, which is based on a large set of specimens, yields three new species as well as numerous, previously unknown or poorly known, soft-part characters with phylo- genetic significance. MATERIAL AND METHODS One hundred twenty one (121) teudopseid specimens from various collections were involved in this study. The material includes the type specimens, previously described teudopseids and 91 unexamined specimens from the BHI collection. All but one of the specimens comes from the Cenomanian of Ha ˆ kel and Ha ˆdjoula; a single, additional specimen is from Na ˆmmoura. A brief outline of the geological settings and the methods used in this study is given in Fuchs and Larson (2011). The gladius terminology used in the text follows that of Fuchs (2006b, 2006c), Fuchs et al. (2007), and Fuchs and Weis (2010). The general morphology of the teudopseid gladius and its terminology is given in Figure 1. Institutional abbreviations.—BHI, Black Hills Institute of Geological Research, Hill City (the repository for types and figured specimens is the Black Hills Museum of Natural History, Hill City); BMNH, British Museum of Natural History London; MfN, Museum fu ¨ r Naturkunde, Berlin; MNHNL, Muse ´e National D’Histoire Naturelle Luxembourg; MSNMi, Museo Civico di Storia naturale di Milano; SHK, Sammlung Helmut Keupp; SMNS, Staatliches Museum fu ¨r Naturkunde Stuttgart. SYSTEMATIC PALEONTOLOGY Subclass COLEOIDEA Bather, 1888 Superorder VAMPYROPODA Boletzky, 1992 Order OCTOBRACHIA Fioroni, 1981 Suborder TEUDOPSEINA Starobogatov, 1983 Diagnosis after Fuchs and Weis (2009).—Gladius with clearly reduced, opened, and spoon-shaped conus. Lateral fields and hyperbolar zones 55% and less of the total gladius length. Hyperbolar zones located between lateral and median field as well-developed broad furrows. Anterior median field more or less pointed. Journal of Paleontology, 85(5), 2011, p. 815–834 Copyright ’ 2011, The Paleontological Society 0022-3360/11/0085-0815$03.00 815