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Concept: Camarasaurus

341

Diplodocidae are among the best known sauropod dinosaurs. Several species were described in the late 1800s or early 1900s from the Morrison Formation of North America. Since then, numerous additional specimens were recovered in the USA, Tanzania, Portugal, and Argentina, as well as possibly Spain, England, Georgia, Zimbabwe, and Asia. To date, the clade includes about 12 to 15 nominal species, some of them with questionable taxonomic status (e.g., ‘Diplodocus’ hayi or Dyslocosaurus polyonychius), and ranging in age from Late Jurassic to Early Cretaceous. However, intrageneric relationships of the iconic, multi-species genera Apatosaurus and Diplodocus are still poorly known. The way to resolve this issue is a specimen-based phylogenetic analysis, which has been previously implemented for Apatosaurus, but is here performed for the first time for the entire clade of Diplodocidae. The analysis includes 81 operational taxonomic units, 49 of which belong to Diplodocidae. The set of OTUs includes all name-bearing type specimens previously proposed to belong to Diplodocidae, alongside a set of relatively complete referred specimens, which increase the amount of anatomically overlapping material. Non-diplodocid outgroups were selected to test the affinities of potential diplodocid specimens that have subsequently been suggested to belong outside the clade. The specimens were scored for 477 morphological characters, representing one of the most extensive phylogenetic analyses of sauropod dinosaurs. Character states were figured and tables given in the case of numerical characters. The resulting cladogram recovers the classical arrangement of diplodocid relationships. Two numerical approaches were used to increase reproducibility in our taxonomic delimitation of species and genera. This resulted in the proposal that some species previously included in well-known genera like Apatosaurus and Diplodocus are generically distinct. Of particular note is that the famous genus Brontosaurus is considered valid by our quantitative approach. Furthermore, “Diplodocus” hayi represents a unique genus, which will herein be called Galeamopus gen. nov. On the other hand, these numerical approaches imply synonymization of “Dinheirosaurus” from the Late Jurassic of Portugal with the Morrison Formation genus Supersaurus. Our use of a specimen-, rather than species-based approach increases knowledge of intraspecific and intrageneric variation in diplodocids, and the study demonstrates how specimen-based phylogenetic analysis is a valuable tool in sauropod taxonomy, and potentially in paleontology and taxonomy as a whole.

Concepts: Biology, Sauropoda, Dinosaur, Diplodocus, Supersaurus, Jurassic, Brachiosaurus, Camarasaurus

60

Diplodocids are among the best known sauropod dinosaurs. Numerous specimens of currently 15 accepted species belonging to ten genera have been reported from the Late Jurassic to Early Cretaceous of North and South America, Europe, and Africa. The highest diversity is known from the Upper Jurassic Morrison Formation of the western United States: a recent review recognized 12 valid, named species, and possibly three additional, yet unnamed ones. One of these is herein described in detail and referred to the genus Galeamopus. The holotype specimen of Galeamopus pabsti sp. nov., SMA 0011, is represented by material from all body parts but the tail, and was found at the Howe-Scott Quarry in the northern Bighorn Basin in Wyoming, USA. Autapomorphic features of the new species include a horizontal canal on the maxilla that connects the posterior margin of the preantorbital and the ventral margin of the antorbital fenestrae, a vertical midline groove marking the sagittal nuchal crest, the presence of a large foramen connecting the postzygapophyseal centrodiapophyseal fossa and the spinopostzygapophyseal fossa of mid- and posterior cervical vertebrae, a very robust humerus, a laterally placed, rugose tubercle on the concave proximal portion of the anterior surface of the humerus, a relatively stout radius, the absence of a distinct ambiens process on the pubis, and a distinctly concave posteroventral margin of the ascending process of the astragalus. In addition to the holotype specimen SMA 0011, the skull USNM 2673 can also be referred to Galeamopus pabsti. Histology shows that the type specimen SMA 0011 is sexually mature, although neurocentral closure was not completed at the time of death. Because SMA 0011 has highly pneumatized cervical vertebrae, the development of the lamination appears a more important indicator for individual age than neurocentral fusion patterns. SMA 0011 is one of very few sauropod specimens that preserves the cervico-dorsal transition in both vertebrae and ribs. The association of ribs with their respective vertebrae shows that the transition between cervical and dorsal vertebrae is significantly different in Galeamopus pabsti than in Diplodocus carnegii or Apatosaurus louisae, being represented by a considerable shortening of the centra from the last cervical to the first dorsal vertebra. Diplodocids show a surprisingly high diversity in the Morrison Formation. This can possibly be explained by a combination of geographical and temporal segregation, and niche partitioning.

Concepts: Cervical vertebrae, Dinosaur, Diplodocus, Supersaurus, Jurassic, Brachiosaurus, Camarasaurus, Allosaurus

47

High megaherbivore species richness is documented in both fossil and contemporary ecosystems despite their high individual energy requirements. An extreme example of this is the Late Jurassic Morrison Formation, which was dominated by sauropod dinosaurs, the largest known terrestrial vertebrates. High sauropod diversity within the resource-limited Morrison is paradoxical, but might be explicable through sophisticated resource partitioning. This hypothesis was tested through finite-element analysis of the crania of the Morrison taxa Camarasaurus and Diplodocus. Results demonstrate divergent specialization, with Camarasaurus capable of exerting and accommodating greater bite forces than Diplodocus, permitting consumption of harder food items. Analysis of craniodental biomechanical characters taken from 35 sauropod taxa demonstrates a functional dichotomy in terms of bite force, cranial robustness and occlusal relationships yielding two polyphyletic functional ‘grades’. Morrison taxa are widely distributed within and between these two morphotypes, reflecting distinctive foraging specializations that formed a biomechanical basis for niche partitioning between them. This partitioning, coupled with benefits associated with large body size, would have enabled the high sauropod diversities present in the Morrison Formation. Further, this provides insight into the mechanisms responsible for supporting the high diversities of large megaherbivores observed in other Mesozoic and Cenozoic communities, particularly those occurring in resource-limited environments.

Concepts: Sauropoda, Dinosaur, Diplodocus, Jurassic, Brachiosaurus, Camarasaurus, Allosaurus, Morrison Formation

5

Sauropod dinosaurs were the largest vertebrates ever to walk the Earth, and as mega-herbivores they were important parts of terrestrial ecosystems. In the Late Jurassic-aged Morrison depositional basin of western North America, these animals occupied lowland river-floodplain settings characterized by a seasonally dry climate. Massive herbivores with high nutritional and water needs could periodically experience nutritional and water stress under these conditions, and thus the common occurrence of sauropods in this basin has remained a paradox. Energetic arguments and mammalian analogues have been used to suggest that migration allowed sauropods access to food and water resources over a wide region or during times of drought or both, but there has been no direct support for these hypotheses. Here we compare oxygen isotope ratios (δ(18)O) of tooth-enamel carbonate from the sauropod Camarasaurus with those of ancient soil, lake and wetland (that is, ‘authigenic’) carbonates that formed in lowland settings. We demonstrate that certain populations of these animals did in fact undertake seasonal migrations of several hundred kilometres from lowland to upland environments. This ability to describe patterns of sauropod movement will help to elucidate the role that migration played in the ecology and evolution of gigantism of these and associated dinosaurs.

Concepts: Water, Earth, Sauropoda, Dinosaur, Diplodocus, Jurassic, Brachiosaurus, Camarasaurus

2

The Late Cretaceous titanosauriform sauropod Huabeisaurus allocotus Pang and Cheng is known from teeth and much of the postcranial skeleton. Its completeness makes it an important taxon for integrating and interpreting anatomical observations from more fragmentary Cretaceous East Asian sauropods and for understanding titanosauriform evolution in general.

Concepts: Sauropoda, Dinosaur, Sauropodomorpha, Titanosaur, Camarasaurus, Macronaria

1

A partial skeleton from the Little Snowy Mountains of central Montana is the first referable specimen of the Morrison Formation macronarian sauropod Camarasaurus. This specimen also represents the northernmost occurrence of a sauropod in the Morrison. Histological study indicates that, although the specimen is relatively small statured, it is skeletally mature; this further emphasizes that size is not a undeviating proxy to maturity in dinosaurs, and that morphologies associated with an individual’s age and stature may be more nebulous in sauropods.

Concepts: Sauropoda, Dinosaur, Sauropodomorpha, Diplodocus, Titanosaur, Brachiosaurus, Camarasaurus, Macronaria

1

Attempts to reconstruct the neutral neck posture of sauropod dinosaurs, or indeed any tetrapod, are doomed to failure when based only on the geometry of the bony cervical vertebrae. The thickness of the articular cartilage between the centra of adjacent vertebrae affects posture. It extends (raises) the neck by an amount roughly proportional to the thickness of the cartilage. It is possible to quantify the angle of extension at an intervertebral joint: it is roughly equal, in radians, to the cartilage thickness divided by the height of the zygapophyseal facets over the centre of rotation. Applying this formula to published measurements of well-known sauropod specimens suggests that if the thickness of cartilage were equal to 4.5%, 10% or 18% of centrum length, the neutral pose of the Apatosaurus louisae holotype CM 3018 would be extended by an average of 5.5, 11.8 or 21.2 degrees, respectively, at each intervertebral joint. For the Diplodocus carnegii holotype CM 84, the corresponding angles of additional extension are even greater: 8.4, 18.6 or 33.3 degrees. The cartilaginous neutral postures (CNPs) calculated for 10% cartilage-the most reasonable estimate-appear outlandish. But it must be remembered that these would not have been the habitual life postures, because tetrapods habitually extend the base of their neck and flex the anterior part, yielding the distinctive S-curve most easily seen in birds.

Concepts: Knee, Sauropoda, Dinosaur, Diplodocus, Brachiosaurus, Camarasaurus, Othniel Charles Marsh, Apatosaurus

0

Titanosauriformes is a conspicuous and diverse group of sauropod dinosaurs that inhabited almost all land masses during Cretaceous times. Besides the diversity of forms, the clade comprises one of the largest land animals found so far, Argentinosaurus, as well as some of the smallest sauropods known to date, Europasaurus and Magyarosaurus. They are therefore good candidates for studies on body size trends such as the Cope’s rule, the tendency towards an increase in body size in an evolutionary lineage. We used statistical methods to assess body size changes under both phylogenetic and nonphylogenetic approaches to identify body size trends in Titanosauriformes. Femoral lengths were collected (or estimated from humeral length) from 46 titanosauriform species and used as a proxy for body size. Our findings show that there is no increase or decrease in titanosauriform body size with age along the Cretaceous and that negative changes in body size are more common than positive ones (although not statistically significant) for most of the titanosauriform subclades (e.g. Saltasaridae, Lithostrotia, Titanosauria and Somphospondyli). Therefore, Cope’s rule is not supported in titanosauriform evolution. Finally, we also found a trend towards a decrease of titanosauriform mean body size coupled with an increase in body size standard deviation, both supporting an increase in body size variation towards the end of Cretaceous.

Concepts: Statistics, Standard deviation, Sauropoda, Dinosaur, Sauropodomorpha, Titanosaur, Camarasaurus, Macronaria

0

A vertebral element assigned to an Apatosaurus cf. ajax from the Late Jurassic Morrison Formation is described. The specimen exhibits an unusual morphology where two vertebrae are nearly seamlessly fused together, including the haemal arch that spans them. This morphology is thought be the result of a developmental abnormality. CT scans of the specimen reveal a thin zone of dorsoventral thickening between the two neural arches consistent with cortical bone. Contrast in internal morphology differentiates the anterior and posterior vertebral bodies with the anterior expressing greater porosity, which increased accommodation for barite-rich calcite precipitation. No vacuities are observed to suggest the former presence of an intervertebral disk or intervertebral joints: the absence of an intervertebral disc or intervertebral joints is indicative of a condition known as block vertebra. Block vertebrae occur with the loss, or inhibition, of somitocoele mesenchyme early in embyogenesis (i.e., during resegmentation of the somites responsible for the formation of the affected vertebra). The derivatives of somitocoele mesenchyme include the intervertebral disc and joints. Although vertebral paleopathologies are not uncommon in the fossil record, this specimen is the first recognized congenital malformation within Sauropoda. Anat Rec, 2014. © 2014 Wiley Periodicals, Inc.

Concepts: Developmental biology, Vertebral column, Vertebra, Intervertebral disc, Dinosaur, Diplodocus, Jurassic, Camarasaurus