William I. Sellers

Trace Metals as Biomarkers for Eumelanin Pigment in the Fossil Record

X-ray maps of fossil feather pigments reveal color patterning in extinct bird species. 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.

New insights into the plantar pressure correlates of walking speed using pedobarographic statistical parametric mapping (pSPM)

This study investigates the relation between walking speed and the distribution of peak plantar pressure and compares a traditional ten-region subsampling (10RS) technique with a new technique: pedobarographic statistical parametric mapping (pSPM). Adapted from cerebral fMRI methodology, pSPM is a digital image processing technique that registers foot pressure images such that homologous structures optimally overlap, thereby enabling statistical tests to be conducted at the pixel level. Following previous experimental protocols, we collected pedobarographic records from 10 subjects walking at three different speeds: slow, normal, and fast. Walking speed was recorded and correlated with the peak pressures extracted from the 10 regions, and subsequently with the peak pixel data extracted after pSPM preprocessing. Both methods revealed significant positive correlation between peak plantar pressure and walking speed over the rearfoot and distal forefoot after Bonferroni correction for multiple comparisons. The 10RS analysis found positive correlation in the midfoot and medial proximal forefoot, but the pixel data exhibited significant negative correlation throughout these regions (p<5x10(-5)). Comparing the statistical maps from the two approaches shows that subsampling may conflate pressure differences evident in pixel-level data, obscuring or even reversing statistical trends. The negative correlation observed in the midfoot implies reduced longitudinal arch collapse with higher walking speeds. We infer that this results from pre- or early-stance phase muscle activity and speculate that preferred walking speed reflects, in part, a balance between the energy required to tighten the longitudinal arch and the apparent propulsive benefits of the stiffened arch.

Human-like external function of the foot, and fully upright gait, confirmed in the 3.66 million year old Laetoli hominin footprints by topographic statistics, experimental footprint-formation and computer simulation

It is commonly held that the major functional features of the human foot (e.g. a functional longitudinal medial arch, lateral to medial force transfer and hallucal (big-toe) push-off) appear only in the last 2 Myr, but functional interpretations of footbones and footprints of early human ancestors (hominins) prior to 2 million years ago (Mya) remain contradictory. Pixel-wise topographical statistical analysis of Laetoli footprint morphology, compared with results from experimental studies of footprint formation; foot-pressure measurements in bipedalism of humans and non-human great apes; and computer simulation techniques, indicate that most of these functional features were already present, albeit less strongly expressed than in ourselves, in the maker of the Laetoli G-1 footprint trail, 3.66 Mya. This finding provides strong support to those previous studies which have interpreted the G-1 prints as generally modern in aspect.

Stride lengths, speed and energy costs in walking of Australopithecus afarensis: using evolutionary robotics to predict locomotion of early human ancestors

This paper uses techniques from evolutionary robotics to predict the most energy-efficient upright walking gait for the early human relative Australopithecus afarensis, based on the proportions of the 3.2 million year old AL 288-1 ‘Lucy’ skeleton, and matches predictions against the nearly contemporaneous (3.5–3.6 million year old) Laetoli fossil footprint trails. The technique creates gaits de novo and uses genetic algorithm optimization to search for the most efficient patterns of simulated muscular contraction at a variety of speeds. The model was first verified by predicting gaits for living human subjects, and comparing costs, stride lengths and speeds to experimentally determined values for the same subjects. Subsequent simulations for A. afarensis yield estimates of the range of walking speeds from 0.6 to 1.3 m s−1 at a cost of 7.0 J kg−1 m−1 for the lowest speeds, falling to 5.8 J kg−1 m−1 at 1.0 m s−1, and rising to 6.2 J kg−1 m−1 at the maximum speed achieved. Speeds previously estimated for the makers of the Laetoli footprint trails (0.56 or 0.64 m s−1 for Trail 1, 0.72 or 0.75 m s−1 for Trail 2/3) may have been underestimated, substantially so for Trail 2/3, with true values in excess of 0.7 and 1.0 m s−1, respectively. The predictions conflict with suggestions that A. afarensis used a ‘shuffling’ gait, indicating rather that the species was a fully competent biped.

Estimating mass properties of dinosaurs using laser imaging and 3D computer modelling.

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.

Estimating dinosaur maximum running speeds using evolutionary robotics

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 animals 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.

Arboreality, terrestriality and bipedalism

The full publication of Ardipithecus ramidus has particular importance for the origins of hominin bipedality, and strengthens the growing case for an arboreal origin. Palaeontological techniques however inevitably concentrate on details of fragmentary postcranial bones and can benefit from a whole-animal perspective. This can be provided by field studies of locomotor behaviour, which provide a real-world perspective of adaptive context, against which conclusions drawn from palaeontology and comparative osteology may be assessed and honed. Increasingly sophisticated dynamic modelling techniques, validated against experimental data for living animals, offer a different perspective where evolutionary and virtual ablation experiments, impossible for living mammals, may be run in silico, and these can analyse not only the interactions and behaviour of rigid segments but increasingly the effects of compliance, which are of crucial importance in guiding the evolution of an arboreally derived lineage.

Archaeopteryx feathers and bone chemistry fully revealed via synchrotron imaging

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.

Predicting the metabolic energy costs of bipedalism using evolutionary robotics

SUMMARY To understand the evolution of bipedalism among the hominoids in an ecological context we need to be able to estimate the energetic cost of locomotion in fossil forms. Ideally such an estimate would be based entirely on morphology since, except for the rare instances where footprints are preserved, this is the only primary source of evidence available. In this paper we use evolutionary robotics techniques (genetic algorithms, pattern generators and mechanical modeling) to produce a biomimetic simulation of bipedalism based on human body dimensions. The mechanical simulation is a seven-segment, two-dimensional model with motive force provided by tension generators representing the major muscle groups acting around the lower-limb joints. Metabolic energy costs are calculated from the muscle model, and bipedal gait is generated using a finite-state pattern generator whose parameters are produced using a genetic algorithm with locomotor economy (maximum distance for a fixed energy cost) as the fitness criterion. The model is validated by comparing the values it generates with those for modern humans. The result (maximum efficiency of 200 J m–1) is within 15% of the experimentally derived value, which is very encouraging and suggests that this is a useful analytic technique for investigating the locomotor behaviour of fossil forms. Initial work suggests that in the future this technique could be used to estimate other locomotor parameters such as top speed. In addition, the animations produced by this technique are qualitatively very convincing, which suggests that this may also be a useful technique for visualizing bipedal locomotion.

Mineralized soft-tissue structure and chemistry in a mummified hadrosaur from the Hell Creek Formation, North Dakota (USA)

An extremely well-preserved dinosaur (Cf. Edmontosaurus sp.) found in the Hell Creek Formation (Upper Cretaceous, North Dakota) retains soft-tissue replacement structures and associated organic compounds. Mineral cements precipitated in the skin apparently follow original cell boundaries, partially preserving epidermis microstructure. Infrared and electron microprobe images of ossified tendon clearly show preserved mineral zonation, with silica and trapped carbon dioxide forming thin linings on Haversian canals within apatite. Furthermore, Fourier transform infrared spectroscopy (FTIR) of materials recovered from the skin and terminal ungual phalanx suggests the presence of compounds containing amide groups. Amino acid composition analyses of the mineralized skin envelope clearly differ from the surrounding matrix; however, intact proteins could not be obtained using protein mass spectrometry. The presence of endogenously derived organics from the skin was further demonstrated by pyrolysis gas chromatography mass spectrometry (Py-GCMS), indicating survival and presence of macromolecules that were in part aliphatic (see the electronic supplementary material).