Body mass reconstructions of extinct vertebrates are most robust when complete to near-complete skeletons allow the reconstruction of either physical or digital models. Digital models are most efficient in terms of time and cost, and provide the facility to infinitely modify model properties non-destructively, such that sensitivity analyses can be conducted to quantify the effect of the many unknown parameters involved in reconstructions of extinct animals. In this study we use laser scanning (LiDAR) and computer modelling methods to create a range of 3D mass models of five specimens of non-avian dinosaur; two near-complete specimens of Tyrannosaurus rex, the most complete specimens of Acrocanthosaurus atokensis and Strutiomimum sedens, and a near-complete skeleton of a sub-adult Edmontosaurus annectens. LiDAR scanning allows a full mounted skeleton to be imaged resulting in a detailed 3D model in which each bone retains its spatial position and articulation. This provides a high resolution skeletal framework around which the body cavity and internal organs such as lungs and air sacs can be reconstructed. This has allowed calculation of body segment masses, centres of mass and moments or inertia for each animal. However, any soft tissue reconstruction of an extinct taxon inevitably represents a best estimate model with an unknown level of accuracy. We have therefore conducted an extensive sensitivity analysis in which the volumes of body segments and respiratory organs were varied in an attempt to constrain the likely maximum plausible range of mass parameters for each animal. Our results provide wide ranges in actual mass and inertial values, emphasizing the high level of uncertainty inevitable in such reconstructions. However, our sensitivity analysis consistently places the centre of mass well below and in front of hip joint in each animal, regardless of the chosen combination of body and respiratory structure volumes. These results emphasize that future biomechanical assessments of extinct taxa should be preceded by a detailed investigation of the plausible range of mass properties, in which sensitivity analyses are used to identify a suite of possible values to be tested as inputs in analytical models.
Well-preserved fossils of pivotal early bird and nonavian theropod species have provided unequivocal evidence for feathers and/or downlike integuments. Recent studies have reconstructed color on the basis of melanosome structure; however, the chemistry of these proposed melanosomes has remained unknown. We applied synchrotron x-ray techniques to several fossil and extant organisms, including Confuciusornis sanctus, in order to map and characterize possible chemical residues of melanin pigments. Results show that trace metals, such as copper, are present in fossils as organometallic compounds most likely derived from original eumelanin. The distribution of these compounds provides a long-lived biomarker of melanin presence and density within a range of fossilized organisms. Metal zoning patterns may be preserved long after melanosome structures have been destroyed.
Maximum running speed is an important locomotor parameter for many animals-predators as well as prey-and is thus of interest to palaeobiologists wishing to reconstruct the behavioural ecology of extinct species. A variety of approaches have been tried in the past including anatomical comparisons, bone scaling and strength, safety factors and ground reaction force analyses. However, these approaches are all indirect and an alternative approach is to create a musculoskeletal model of the animal and see how fast it can run. The major advantage of this approach is that all assumptions about the animal's morphology and physiology are directly addressed, whereas the exact same assumptions are hidden in the indirect approaches. In this paper, we present simple musculoskeletal models of three extant and five extinct bipedal species. The models predict top speed in the extant species with reasonably good agreement with accepted values, so we conclude that the values presented for the five extinct species are reasonable predictions given the modelling assumptions made. Improved musculoskeletal models and better estimates of soft tissue parameters will produce more accurate values. Limited sensitivity analysis is performed on key muscle parameters but there is considerable scope for extending this in the future.
Body mass is a critical parameter used to constrain biomechanical and physiological traits of organisms. Volumetric methods are becoming more common as techniques for estimating the body masses of fossil vertebrates. However, they are often accused of excessive subjective input when estimating the thickness of missing soft tissue. Here, we demonstrate an alternative approach where a minimum convex hull is derived mathematically from the point cloud generated by laser-scanning mounted skeletons. This has the advantage of requiring minimal user intervention and is thus more objective and far quicker. We test this method on 14 relatively large-bodied mammalian skeletons and demonstrate that it consistently underestimates body mass by 21 per cent with minimal scatter around the regression line. We therefore suggest that it is a robust method of estimating body mass where a mounted skeletal reconstruction is available and demonstrate its usage to predict the body mass of one of the largest, relatively complete sauropod dinosaurs: Giraffatitan brancai (previously Brachiosaurus ) as 23200 kg.
The discovery of a largely complete and well preserved specimen of Poposaurus gracilis has provided the opportunity to generate the first phylogenetically based reconstruction of pelvic and hindlimb musculature of an extinct nondinosaurian archosaur. As in dinosaurs, multiple lineages of basal archosaurs convergently evolved parasagittally erect limbs. However, in contrast to the laterally projecting acetabulum, or "buttress erect" hip morphology of ornithodirans, basal archosaurs evolved a very different, ventrally projecting acetabulum, or "pillar erect" hip. Reconstruction of the pelvic and hindlimb musculotendinous system in a bipedal suchian archosaur clarifies how the anatomical transformations associated with the evolution of bipedalism in basal archosaurs differed from that of bipedal dinosaurs and birds. This reconstruction is based on the direct examination of the osteology and myology of phylogenetically relevant extant taxa in conjunction with osteological correlates from the skeleton of P. gracilis. This data set includes a series of inferences (presence/absence of a structure, number of components, and origin/insertion sites) regarding 26 individual muscles or muscle groups, three pelvic ligaments, and two connective tissue structures in the pelvis, hindlimb, and pes of P. gracilis. These data provide a foundation for subsequent examination of variation in myological orientation and function based on pelvic and hindlimb morphology, across the basal archosaur lineage leading to extant crocodilians.
Tracks can potentially offer unique sources of information, providing insight into the environments, gait and posture, locomotion and behaviour. Track preservation can yield important information on substrate consistency and enable the recognition of transmitted subsurface tracks. The ability to recognize transmitted tracks has broad implications for the understanding of palaeoenvironments and interpretation of ichnological assemblages. In order to gain an understanding of how tracks are formed in three dimensions, and of their variability of expression in different substrates, controlled laboratory simulations were undertaken. Experiments were designed to recover subsurface track layers, yielding for the first time detailed information on subsurface morphology that could be related to ‘true’ surface track features. It was found that subsurface track relief can be correlated with the magnitude and distribution (across a foot) of load acting on the surface sediment. This pressure is transmitted through the sediment, and deforms successive layers at depth, producing an undertrack. The most significant factor controlling track morphology, whether surface or subsurface, was found to be the moisture/density relationship within the substrate at the time of track formation. Variability in the dimensions of simulated tracks, relative to the ‘true’ surface track, indicates that caution should be exercised when using fossil tracks to calculate hip height, speed, age, and population dynamics. In addition, comparison of experimental tracks with dinosaur tracks from the Yorkshire coast suggests that many morphological differences between vertebrate ichnotaxa reflect sediment rheology and taphonomy rather than taxonomy of the track-maker.
Evolution of flight in maniraptoran dinosaurs is marked by the acquisition of distinct avian characters, such as feathers, as seen in Archaeopteryx from the Solnhofen limestone. These rare fossils were pivotal in confirming the dinosauria-avian lineage. One of the key derived avian characters is the possession of feathers, details of which were remarkably preserved in the Lagerstätte environment. These structures were previously simply assumed to be impressions; however, a detailed chemical analysis has, until now, never been completed on any Archaeopteryx specimen. Here we present chemical imaging via synchrotron rapid scanning X-ray fluorescence (SRS-XRF) of the Thermopolis Archaeopteryx, which shows that portions of the feathers are not impressions but are in fact remnant body fossil structures, maintaining elemental compositions that are completely different from the embedding geological matrix. Our results indicate phosphorous and sulfur retention in soft tissue as well as trace metal (Zn and Cu) retention in bone. Other previously unknown chemical details of Archaeopteryx are also revealed in this study including: bone chemistry, taphonomy (fossilization process), and curation artifacts. SRS-XRF represents a major advancement in the study of the life chemistry and fossilization processes of Archaeopteryx and other extinct organisms because it is now practical to image the chemistry of large specimens rapidly at concentration levels of parts per million. This technique has wider application to the archaeological, forensic, and biological sciences, enabling the mapping of "unseen" compounds critical to understanding biological structures, modes of preservation, and environmental context. trace elements | X-ray absorption spectroscopy A rchaeopteryx (1) are rare but occupy a pivotal place in the development of Darwinian evolution because of their possession of both reptilian (jaws with teeth and a long bony tail) and avian (feathered wings) characters (2). The specimen used in this study is considered to be the most complete and best preserved archaeopterygid (3) belonging to the species A. siemensii (4). Previous analyses of this fossil have relied upon visual inspection, X-ray computer tomography, scanning electron, and ultraviolet/ visible light microscopy. Structural studies have been extensive and strongly indicate that this organism is transitional between dinosaurs and birds; however, detailed chemical analysis has never been performed. Here we apply state-of-the-art synchrotron rapid scanning X-ray fluorescence (SRS-XRF) imaging to this remarkably well-preserved specimen revealing striking and previously unknown details about the chemical preservation of soft tissue, elemental distribution patterns most likely related to the organism's life processes, insights into the chemistry of the fossilization process, and details of curation history. In addition, quantitative chemical analyses and X-ray absorption spectroscopy are presented that not only corroborate the imaging results but also give further det...
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