Jørn H. Hurum

Complete Primate Skeleton from the Middle Eocene of Messel in Germany: Morphology and Paleobiology

Background The best European locality for complete Eocene mammal skeletons is Grube Messel, near Darmstadt, Germany. Although the site was surrounded by a para-tropical rain forest in the Eocene, primates are remarkably rare there, and only eight fragmentary specimens were known until now. Messel has now yielded a full primate skeleton. The specimen has an unusual history: it was privately collected and sold in two parts, with only the lesser part previously known. The second part, which has just come to light, shows the skeleton to be the most complete primate known in the fossil record. Methodology/Principal Findings We describe the morphology and investigate the paleobiology of the skeleton. The specimen is described as Darwinius masillae n.gen. n.sp. belonging to the Cercamoniinae. Because the skeleton is lightly crushed and bones cannot be handled individually, imaging studies are of particular importance. Skull radiography shows a host of teeth developing within the juvenile face. Investigation of growth and proportion suggest that the individual was a weaned and independent-feeding female that died in her first year of life, and might have attained a body weight of 650–900 g had she lived to adulthood. She was an agile, nail-bearing, generalized arboreal quadruped living above the floor of the Messel rain forest. Conclusions/Significance Darwinius masillae represents the most complete fossil primate ever found, including both skeleton, soft body outline and contents of the digestive tract. Study of all these features allows a fairly complete reconstruction of life history, locomotion, and diet. Any future study of Eocene-Oligocene primates should benefit from information preserved in the Darwinius holotype. Of particular importance to phylogenetic studies, the absence of a toilet claw and a toothcomb demonstrates that Darwinius masillae is not simply a fossil lemur, but part of a larger group of primates, Adapoidea, representative of the early haplorhine diversification.

Phylogeny and Systematics of Multituberculate Mammals

We present a synopsis of high-rank multituberculate systematicsand a manually generated cladogram illustrating multituberculate interrelationships. We divide the Multituberculata into the paraphyletic suborder ‘Plagiaulacida’, an apparently monophyletic suborder Cimolodonta, and one family incertae sedis. Within ‘Plagiaulacida’ we recognise three informal lines: paulchoffatiid (three families), plagiaulacid (three families) and allodontid (two families and the genus Glirodon). The Cimolodonta are divided into an informal Paracimexomys group; three superfamilies: Ptilodontoidea, Djadochtatherioidea (new), and Taeniolabidoidea (restricted to Taeniolabididae); and five families (superfamily incertae sedis): Eucosmodontidae, Microcosmodontidae, Cimolodontidae, Boffiidae, and Kogaionidae; and some genera incertae sedis. New characters used in our analysis are (1) a tendency of molar cusps to coalesce; and (2) ornamentation of grooves, pits, and ridges on the molars. We argue that the Ptilodontoidea, and less certainly also the Cimolodontidae and Boffiidae, might have originated from amongthe plagiaulacid line, a possible intermediate link being the Paracimexomys group. The remaining Cimolodonta might have originated from unknown members of the Paracimexomys group with separated molar cusps and smooth enamel. The origin of two types of prismatic enamel and a relationship between them are stumbling blocks in understanding the origin of the Cimolodonta; we conclude that microprismatic enamel made its appearance only once. Revised diagnoses of high-rank multituberculate taxa, including lists of all known genera, are given.

Skull structure and evolution in tyrannosaurid dinosaurs

Tyrannosauridae can be subdivided into two distinct subfamilies—the Albertosaurinae and the Tyrannosaurinae. Previ− ously recognized subdivisions Aublysodontinae and Shanshanosaurinae are rejected because they are based on insuffi− cient material and juvenile specimens. Our results are based upon a phylogenetic analysis using PAUP program (Swofford 1999) of 77 skull characters and seven genera (Albertosaurus, Alioramus, Daspletosaurus, Gorgosaurus, Nanotyrannus, Tarbosaurus, and Tyrannosaurus); with Allosaurus as outgroup. Of the 77 characters used, more than half were parsimony informative. A single most parsimonious tree was obtained with the Tree Length being 88. The analysis of cranial characters and comparison of postcranial features reveal that Tarbosaurus bataar is not the sister taxon of Tyrannosaurus rex (contra Holtz 2001). Their similarities are partially due to the fact that both are extremely large ani− mals. Thus, Tarbosaurus should be considered a genus distinct from Tyrannosaurus.

THE CRUSHING BITE OF TYRANNOSAURIDS

Tyrannosaurids are a group of large meat eating theropod dinosaurs. Newly prepared material of Gorgosaurus libratus, Tarbosaurus bataar, and Tyrannosaurus rex provides additional knowledge of the structure and movement of the intramandibular jaw joint in the family. Paradoxically the theories of jaw movement and the fossil evidence in tyrannosaurids have not been comparable. It has been suggested that the joint had a wide range of movement to allow the gullet to expand, and to allow adjustment of the pitch and position of the mandibular teeth relative to the upper teeth (Maleev, 1974; Barsbold, 1983; Bakker et al., 1988; Molnar, 1991). A moveable joint suggested a feeding behaviour like lizards with a weak bite. On the other hand, the shortening and elevation of the skull in tyrannosaurids for insertion of powerful jaw muscles (Molnar and Farlow 1990), increased tooth robustness (Farlow et al., 1991), bite marks suggesting bone fragmentation (Erickson and Olson 1996), estimation of bite force from tooth-marked bone (Erickson et al., 1996), and bone fragments in coprolites (Chin et al. 1998) all suggests high bite force. Here we describe the intramandibular jaw joint and report on the discovery of the first completely preserved and fused supradentary/coronoid bones in three different tyrannosaurids. The fused supradentary/coronoid bone crosses the intramandibular jaw joint restricting its movement. This leads to a rigid lower jaw in tyrannosaurids that is a secondary specialization for a powerful crushing bite. Institutional Abbreviations-BHI, Black Hills Institute, South Dakota, USA; TMP, Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta, Canada; ZPAL, Institute of Palaeobiology, Academy of Sciences, Warszawa, Poland.

An ankylosaurid dinosaur from Mongolia with in situ armour and keratinous scale impressions

A Mongolian ankylosaurid specimen identified as Tarchia gigantea is an articulated skeleton including dorsal ribs, the sacrum, a nearly complete caudal series, and in situ osteoderms. The tail is the longest complete tail of any known ankylosaurid. Remarkably, the specimen is also the first Mongolian ankylosaurid that preserves impressions of the keratinous scales overlying the bony osteoderms. This specimen provides new information on the shape, texture, and arrangement of osteoderms. Large flat, keeled osteoderms are found over the pelvis, and osteoderms along the tail include large keeled osteoderms, elongate osteoderms lacking distinct apices, and medium-sized, oval osteoderms. The specimen differs in some respects from other Tarchia gigantea specimens, including the morphology of the neural spines of the tail club handle and several of the largest osteoderms.

Large Onychites (Cephalopod Hooks) from the Upper Jurassic of the Boreal Realm

We report on the discovery of large cephalopod arm hooks (mega−onychites) from the Kimmeridgian and Volgian of Spitsbergen (Agardhfjellet Formation). This includes a largely uncompressed hook in a seep carbonate,with preservation of surface sculpture. We suggest the use of logarithmic spirals as morphological descriptors for the outer part of cephalopod arm hooks, with implications for systematics and functional morphology. Comparison with Upper Jurassic material from Greenland, northern Norway and the North Sea demonstrates a remarkably consistent morphology, which we assign to the same form species, Onychites quenstedti. Considering the relatively small stratigraphic (Kimmeridgian— Volgian) and biogeographic (Boreal) range of this large form, it is likely that it represents a single biological species or genus.

Postcranial Skeleton of a Cretaceous Multituberculate Mammal Catopsbaatar

We describe an incomplete postcranial skeleton of Catopsbaatar catopsaloides from the?late Campanian red beds of Hermiin Tsav I, in the Gobi Desert, Mongolia. The skeleton is fragmentary and the preservation of bone surface does not permit reconstruction of the musculature. The studied skeleton contains some parts not preserved or incompletely known in other multituberculate genera, such as a long spinous process in a single lumbar vertebra, which together with long transverse processes preserved in Nemegtbaatar, might indicate that at least some multituberculates had jumping ability. The calcaneus of Catopsbaatar is unusual, differing from most other multituberculates (where known) and other mammals by having a short tuber calcanei, with a large proximal anvil-shaped process strongly bent laterally and ventrally, arranged obliquely with respect to the distal margin of the calcaneus, rather than arranged at 90° to it, as in other mammals. This suggests the presence of strong muscles that attached to the tuber calcanei, perhaps further attesting to jumping abilities in Catopsbaatar. We also describe an unfused pelvic girdle and the first extratarsal spur bone (os cornu calcaris) known in multituberculates.

A New Ophthalmosaurid (Ichthyosauria) from Svalbard, Norway, and Evolution of the Ichthyopterygian Pelvic Girdle

In spite of a fossil record spanning over 150 million years, pelvic girdle evolution in Ichthyopterygia is poorly known. Here, we examine pelvic girdle size relationships using quantitative methods and new ophthalmosaurid material from the Slottsmøya Member Lagerstätte of Svalbard, Norway. One of these new specimens, which preserves the most complete ichthyosaur pelvic girdle from the Cretaceous, is described herein as a new taxon, Keilhauia nui gen. et sp. nov. It represents the most complete Berriasian ichthyosaur known and the youngest yet described from the Slottsmøya Member. It is diagnosed on the basis of two autapomorphies from the pelvic girdle, including an ilium that is anteroposteriorly expanded at its dorsal end and an ischiopubis that is shorter or subequal in length to the femur, as well as a unique character combination. The Slottsmøya Member Lagerstätte ichthyosaurs are significant in that they represent a diverse assemblage of ophthalmosaurids that existed immediately preceding and across the Jurassic–Cretaceous boundary. They also exhibit considerable variation in pelvic girdle morphology, and expand the known range in size variation of pelvic girdle elements in the clade.

The Slottsmøya marine reptile Lagerstätte: depositional environments, taphonomy and diagenesis

Abstract The Late Jurassic Slottsmøya Member Lagerstätte on Spitsbergen offers a unique opportunity to study the relationships between vertebrate fossil preservation, invertebrate occurrences and depositional environment. In this study, 21 plesiosaurian and 17 ichthyosaur specimens are described with respect to articulation, landing mode, preservation, and possible predation and scavenging. The stratigraphic distribution of marine reptiles in the Slottsmøya Member is analysed, and a correlation between high total organic content, low oxygen levels, few benthic invertebrates and optimal reptile preservation is observed. A new model for 3D preservation of vertebrates in highly compacted organic shales is explained. Supplementary material: A taphonomic description of each marine reptile specimen is available at https://doi.org/10.6084/m9.figshare.c.2133549

FIRST DRYOLESTOID (MAMMALIA, DRYOLESTOIDEA, MERIDIOLESTIDA) FROM THE CONIACIAN OF PATAGONIA AND NEW EVIDENCE ON THEIR EARLY RADIATION IN SOUTH AMERICA

Abstract. We report on a new dryolestoid (Mammalia, Dryolestoidea, Meridiolestida) from the Los Bastos Formation (Coniacian), Neuquén Province, Patagonia, Argentina, consisting of an edentulous left dentary (MCF-PVPH 412). The alveoli preserved suggest the presence of three incisors, one double-rooted canine, and six double-rooted postcanines (probably three premolars and three molars). Based on comparisons with previously known dentaries and isolated teeth, MCF-PVPH 412 would have been about the same size as Reigitheriurn Bonaparte. Among Dryolestoidea, MCF-PVPH 412 is assigned to Meridiolestida because there were probably three molars, the roots of the posterior molars are anteroposteriorly compressed, and there is no Meckelian groove. In addition, the penultimate lower premolar would be the largest in the tooth series, which is also true in other meridiolestidans. The position of the mandibular foramen, the probable presence of three premolars, and the outline of the posteroventral part of the jaw suggest affinities with the Mesungulatoidea (e.g ., Coloniatherium Rougier, Forasiepi, Hill and Novaceck; Peligrotherium Bonaparte, Van Valen and Kramarz; and Reigitheriurn). The Coniacian specimen represents the oldest Mesungulatoidea and fills the gap in the record between the oldest South American dryolestoid (i.e., Cenomanian) and the better known Campanian—Maastrichtian taxa. The discovery of MCF-PVPH 412 in the Coniacian of Patagonia is consistent with the dryolestoid diversification during the Late Cretaceous that makes them the most abundant mammals during that period in South America.