ALBERT

Description: We here describe a monospecific assemblage of giant aquatic snakes from the middle Eocene of Kpogamé, Togo. The material, consisting of large isolated vertebrae, is referred to Palaeophis africanus, an enigmatic palaeophiid species, which was so far otherwise known only from a limited number of vertebrae from the middle Eocene of Nigeria and Angola. Material from the late Eocene of the eastern USA that had been referred to the same species, is here instead considered too fragmentary for species-level determination and Palaeophis africanus is thus so far restricted to Africa. With the aid of micro-CT scanning, we present 3D models of 17 vertebrae, pertaining to different portions of the vertebral column. We provide detailed comparisons of the new material with all named African species of the genus Palaeophis. A tentative diagnosis of Palaeophis africanus is provided. With more than 50 vertebrae, the new Togolese specimens represent the most abundant known material attributed to Palaeophis africanus and significantly enhance our knowledge of the vertebral anatomy and intracolumnar variation for this taxon. Furthermore, this adds to the, as yet, extremely scarce fossil record of squamates from central western Africa, a region where Paleogene herpetofaunas are only rather poorly known.

Description: Trace fossils occur in several strata of the Devonian and Carboniferous of the eastern Anti-Atlas, but they are still poorly documented. Here, we describe a fossil swimming trace from strata overlying the Hangenberg Black Shale (correlation largely based on lithostratigraphy; Postclymenia ammonoid genozone, ca. 370 Ma old). We discuss the systematic position of the tracemaker and its body size. This ichnofossil is important for three main reasons: (1) it is considered here to be the first record of Undichna from the Devonian of Gondwana, as far as we know; (2) it is the oldest record of vertebrate trace fossils from Africa; (3) it provides a unique window into the behaviour of Late Devonian fishes for which body-fossils cannot provide direct evidence. Further, we put this discovery into the macroecological context of the palaeoenvironment following the Late Devonian Hangenberg biodiversity crisis.

Description: Large constrictor snakes, referred to the genera Palaeopython and Paleryx, are an ecologically prominent part of the fauna of Europe during the Paleogene. Most species were named over a century ago and their taxonomy is largely based on isolated vertebrae. Furthermore, the majority of named taxa originate from imprecisely known localities within the Phosphorites du Quercy, in southern France, and thus their exact age is not known. We critically review and re-diagnose these genera based on personal examination of all existing type material, an array of new specimens, and a detailed literature review. We consider Palaeopython and Paleryx to be valid and propose vertebral characters to distinguish them. We recognize three valid species of Palaeopython, i.e. Palaeopythoncadurcensis (type species) from the Phosphorites du Quercy, Palaeopythonceciliensis from Geiseltal, and Palaeopythonhelveticus from Dielsdorf (Switzerland), and one valid species of Paleryx, i.e. Paleryxrhombifer (type species) from Hordle Cliff (England). Four other species, which were previously treated as members of Palaeopython and Paleryx, i.e. “Palaeopython” filholii and “Palaeopython” neglectus from the Phosphorites du Quercy, “Palaeopython” fischeri from Messel, and “Paleryx” spinifer from Geiseltal, are also considered as valid but pertain to other genera. Among these four taxa, “Palaeopython” fischeri has been recently assigned to its own genus, Eoconstrictor. A new genus, Phosphoroboa gen. nov. is established to accommodate “Palaeopython” filholii. We designate a lectotype for Palaeopythoncadurcensis and establish that the paralectotype maxilla and dentary are reasonably referred to this species. New material attributed to Palaeopythoncadurcensis is described from the old collections of the Phosphorites du Quercy. Paleryxcayluxi, another species established from the old collections of the Phosphorites du Quercy, is synonymized here with Palaeopythoncadurcensis. We further clarify important errors in the original description and figures of Paleryxcayluxi, identify the exact specimens that comprise the type series, and designate a lectotype. Much new material is described for Palaeopythonceciliensis from its type area in Geiseltal and intracolumnar variation is considered. We describe additional vertebral and cranial material of Paleryxrhombifer from its type area in Hordle Cliff. Based on this cranial material, we suggest non-booid affinities for Paleryxrhombifer. We designate a lectotype for Paleryxdepressus and agree with its previous suggested synonymy with Paleryxrhombifer. We re-describe the lectotype and paralectotypes of “Palaeopython” neglectus and refer and describe new material of this species from the Phosphorites du Quercy, paying special attention to intracolumnar variation; we also defer a decision on its generic relations until more abundant and complete material can be studied. We describe new vertebral material of the booid Eoconstrictor cf. fischeri from Geiseltal; similar material was previously known only from Messel and Dielsdorf. We determine that Eoconstrictorfischeri contains two distinct and unrelated species and describe intracolumnar variation in the nominotype. We clarify certain issues regarding the type series of Paleryxspinifer, designate a lectotype, and report previously unrecognized cranial material associated with the latter specimen; we transfer this species to Eoconstrictor based on cranial features and recombine it as Eoconstrictorspinifer comb. nov. We finally describe much new vertebral and cranial material of Phosphoroboafilholii comb. nov. from the Phosphorites du Quercy (both from the old collections but also from the late Eocene localities of Escamps A and C), paying special attention to intracolumnar variation. Based on this cranial material from Escamps, we identify Phosphoroboa gen. nov. as a booid. An analytical approach is undertaken in many isolated remains in order to quantify vertebral structures and assess intracolumnar variation, as well as associating isolated cranial elements to vertebral-based taxa. 3D models of the type material of the Geiseltal and Messel taxa are presented. The importance of vertebrae in the taxonomy of fossil Constrictores is addressed, although it is acknowledged that it is cranial material that can afford the most reliable phylogenetic conclusions. The diversity, distribution, biogeographic origins, and final demise and extinction of large Constrictores in the Paleogene of Europe are discussed.

Description: Especially in Lagerstätten with exceptionally preserved fossils, we can sometimes recognize fossilized remains of meals of animals. We suggest the term leftover fall for the event and the term pabulite for the fossilized meal when it never entered the digestive tract (difference to regurgitalites). Usually, pabulites are incomplete organismal remains and show traces of the predation. Pabulites have a great potential to inform about predation as well as anatomical detail, which is invisible otherwise. Here, we document a pabulite comprising the belemnite Passaloteuthis laevigata from the Toarcian of the Holzmaden region. Most of its soft parts are missing while the arm crown is one of the best preserved that is known. Its arms embrace an exuvia of a crustacean. We suggest that the belemnite represents the remnant of the food of a predatory fish such as the shark Hybodus.

Description: The standard reconstruction of species of Orbitoides d’Orbigny into a single lineage during the late Santonian to the end of the Maastrichtian is based upon morphometric data from Western Europe. An irreversible increase in the size of the embryonic apparatus, and the formation of a greater number of epi-embryonic chamberlets (EPC) with time, is regarded as the main evolutionary trends used in species discrimination. However, data from Maastrichtian Orbitoides assemblages from Central Turkey and the Arabian Platform margin (Southeastern Turkey and Oman) are not consistent with this record. The Maastrichtian Besni Formation of the Arabian Platform margin in Southeastern Turkey yields invariably biconvex specimens, with small, tri- to quadrilocular embryons and a small number of EPC, comparable to late Campanian Orbitoides medius (d’Archiac). The upper Maastrichtian Taraklı Formation from the Sakarya Basin of Central Turkey contains two distinct, yet closely associated forms of Orbitoides, easily differentiated by both external and internal features. Flat to biconcave specimens possess a small, tri- to quadrilocular embryonic apparatus of Orbitoides medius-type and a small number of EPC, whereas biconvex specimens possess a large, predominantly bilocular embryonic apparatus, and were assigned to Orbitoides ex. interc. gruenbachensis Papp–apiculatus Schlumberger based on morphometry. The flat to biconcave specimens belong to a long overlooked species Orbitoides pamiri Meriç, originally described from the late Maastrichtian of the Tauride Mountains in SW Turkey. This species is herein interpreted to be an offshoot from the main Orbitoides lineage during the Maastrichtian, as are forms that we term Orbitoides ‘medius’, since they recall this species, yet are younger than normal occurrence with the accepted morphometrically defined lineage. The consistent correlation between the external and internal test features in O. pamiri implies that the shape of the test is not an ecophenotypic variation, but appears to be biologically controlled. We, therefore, postulate that more than one lineage of Orbitoides exists during the Maastrichtian, with a lineage that includes O. ‘medius’ and O. pamiri displaying retrograde evolutionary features.

Description: Exceptional fossil preservation is required to conserve soft-bodied fossils and even more so to conserve their behaviour. Here, we describe a fossil of a co-occurrence of representatives of two different octobrachian coleoid species. The fossils are from the Toarcian Posidonienschiefer of Ohmden near Holzmaden, Germany. The two animals died in the act of predation, i.e. one had caught the other and had begun to nibble on it, when they possibly sank into hypoxic waters and suffocated (distraction sinking). This supports the idea that primitive vampyromorphs pursued diverse feeding strategies and were not yet adapted to being opportunistic feeders in oxygen minimum zones like their modern relative Vampyroteuthis.

Description: Over the last 25 years, researchers, mostly paleontologists, have developed a system of rank-free, phylogenetically defined names for the primary clades of turtles. As these names are not considered established by the PhyloCode, the newly created nomenclatural system that governs the naming of clades, we take the opportunity to convert the vast majority of previously defined clade names for extinct and extant turtles into this new nomenclatural framework. Some previously defined names are converted with minor adjustments. We also define a number of new clade names to close apparent nomenclatural gaps. In total, we establish 113 clade names, of which 79 had already received phylogenetic definitions and 34 are new.

Description: Due to the lower fossilization potential of chitin, non-mineralized cephalopod jaws and arm hooks are much more rarely preserved as fossils than the calcitic lower jaws of ammonites or the calcitized jaw apparatuses of nautilids. Here, we report such non-mineralized fossil jaws and arm hooks from pelagic marly limestones of continental Greece. Two of the specimens lie on the same slab and are assigned to the Ammonitina; they represent upper jaws of the aptychus type, which is corroborated by finds of aptychi. Additionally, one intermediate type and one anaptychus type are documented here. The morphology of all ammonite jaws suggest a desmoceratoid affinity. The other jaws are identified as coleoid jaws. They share the overall U-shape and proportions of the outer and inner lamellae with Jurassic lower jaws of Trachyteuthis (Teudopseina). We also document the first belemnoid arm hooks from the Tethyan Maastrichtian. The fossils described here document the presence of a typical Mesozoic cephalopod assemblage until the end of the Cretaceous in the eastern Tethys.

Description: In the central Jura Mountains (Western Switzerland), the Urgonien Jaune (UJ) facies with the Marnes de la Russille beds (MRu) have provided very rich nannofloras associated with very rare Tethyan ammonites. A late Early Barremian nannoflora of the Mid-Barremian Event (MBE, following a regional tectonic event of an earliest Barremian synsedimentary tectonic crisis) was found in MRu of the lower UJ and includes 42 genera with 90 species. Among them, Biscutum jurensis De Kaenel, n. sp., Flabellites eclepensensis De Kaenel, n. sp., Palaeopontosphaera giraudii De Kaenel, n. sp., Rhagodiscus buisensis De Kaenel, n. sp., and Vagalapilla rutledgei De Kaenel, n. sp., are recognized as five new species. This nannoflora is a mixture of Boreal and Tethyan taxa with 20 nannofossil markers (Assipetra terebrodentarius, Broinsonia galloisii, Calcicalathina oblongata, Cyclagelosphaera rotaclypeata, Diloma placinum, Ethmorhabdus hauterivianus, Flabellites eclepensensis, Gorkaea pseudoanthophorus, Nannoconus abundans, Nannoconus pseudoseptentrionalis, Palaeopontosphaera giraudii, Palaeopontosphaera pinnata, Placozygus howei, Placozygus reticulatus, Reinhardtites scutula, Rhagodiscus buisensis, Rhagodiscus eboracensis, Tegulalithus septentrionalis, Tubodiscus jurapelagicus, Zeugrhabdotus moulladei) indicating very precisely the nannofossil Zones LK19 (Boreal)–NC5D (Tethyan) as well as the Elegans (Boreal) and Moutonianum (Tethyan) ammonite Zones of the latest Early Barremian. The ammonites in the basal UJ facies of Early Barremian age are reworked Lyticoceras claveli (Nodosoplicatum Zone, Early Hauterivian) and reworked Cruasiceras cf. cruasense (Sayni Zone, early Late Hauterivian), and Pseudometahoplites sp. juv. (Compressissima to Vandenheckii Zones, Early to Late Barremian transition) from the basal MRu. The new palaeontological and sequential results of this study allow a revision of previous data from Godet et al. (2010) precisely assigning a Barremian age to the MRu of the central Jura Mountains (Tethyan Compressissima to lower Sartousiana and Boreal uppermost Fissicostatum to middle Denckmanii ammonite Zones, Boreal LK20B-LK19-LK18 and Tethyan NC5D nannofossil Zones), within the Early to Late Barremian UJ (Tethyan Hugii to lower Sartousiana and Boreal Rarocinctum to middle Denckmanii ammonite Zones, Boreal LK20C to LK18 and Tethyan NC5C-NC5D nannofossil Zones) and below the Late Barremian Urgonien Blanc facies (Tethyan Sartousiana ammonite Zone).

Description: We describe a well-preserved South American Lamini partial skeleton (PIMUZ A/V 4165) from the Ensenadan (~ 1.95–1.77 to 0.4 Mya) of Argentina. The specimen is comprised of a nearly complete skull and mandible with full tooth rows, multiple elements of anterior and posterior limbs, and a scapula. We tested this specimen’s phylogenetic position and hypothesized it to be more closely related to Lama guanicoe and Vicugna vicugna than to Hemiauchenia paradoxa. We formulate a hypothesis for the placement of PIMUZ A/V 4165 within Camelinae in a cladistic analysis based on craniomandibular and dental characters and propose that future systematic studies consider this specimen as representing a new species. For the first time in a morphological phylogeny, we code terminal taxa at the species level for the following genera: Camelops, Aepycamelus, Pleiolama, Procamelus, and Alforjas. Our results indicate a divergence between Lamini and Camelini predating the Barstovian (16 Mya). Camelops appears as monophyletic within the Camelini. Alforjas taylori falls out as a basal member of Camelinae—neither as a Lamini nor Camelini. Pleiolama is polyphyletic, with Pleiolama vera as a basal Lamini and Pleiolama mckennai in a more nested position within the Lamini. Aepycamelus and Procamelus are respectively polyphyletic and paraphyletic. Together, they are part of a group of North American Lamini from the Miocene epoch.