Peter Lewis Falkingham

Liverpool John Moores University | LJMU · School of Biological and Environmental Sciences

It is inevitable that some important specimens will become lost or damaged over time, conservation is therefore of vital importance. The Paluxy River dinosaur tracksite is among the most famous in the world. In 1940, Roland T. Bird described and excavated a portion of the site containing associated theropod and sauropod trackways. This excavated tr...

Significance We reconstructed the 3D foot movements of guineafowl traversing a granular substrate from biplanar X-rays, and then incorporated those kinematics into a discrete element simulation. Digital track models permitted visualization of in vivo track formation at the surface and at virtual bedding planes for the first time. Application of the...

As purely sedimentary structures, fossil footprints are all about shape. Correctly interpreting the significance of their surface topography requires understanding the sources of morphological variation. Differences among specimens are most frequently attributed to either taxonomy (trackmaker) or to preservation quality. ‘Well-preserved’ tracks are...

The longitudinal arch of the human foot is viewed as a pivotal adaptation for bipedal walking and running. Fossil footprints from Laetoli, Tanzania, and Ileret, Kenya, are believed to provide direct evidence of longitudinally arched feet in hominins from the Pliocene and Pleistocene, respectively. We studied the dynamics of track formation using bi...

Fossil tracks are important palaeobiological data sources. The quantitative analysis of their shape, however, has been hampered by their high variability and lack of discrete margins and landmarks. We here present the first approach using deep convolutional neural networks (DCNNs) to study fossil tracks, overcoming the limitations of previous stati...

Walking on compliant substrates requires more energy than walking on hard substrates but the biomechanical factors that contribute to this increase are debated. Previous studies suggest various causative mechanical factors, including disruption to pendular energy recovery, increased muscle work, decreased muscle efficiency and increased gait variab...

The identification of presumed tetrapod tracks is not always unequivocal. Other sedimentary structures have been repeatedly mistaken for tracks, including other trace fossils such as arthropod tracks, burrows and fish feeding traces; erosional features; and human-made traces. We here review instances of difficult, ambiguous, or controversial cases...

The stance phase of walking is when forces are applied to the environment to support, propel, and maneuver the body. Unlike solid surfaces, deformable substrates yield under load, allowing the foot to sink to varying degrees. For bipedal birds and their dinosaurian ancestors, a shared response to walking on these substrates has been identified in t...

Limb phase, the timing of the footfalls in quadrupedal locomotion that describes common gaits such as the trot and the pace gait,1,2 is widely believed to be difficult or even impossible to estimate for extinct tetrapods.3-5 We here present a fundamentally new approach that allows for estimating limb phase based on variation patterns in long trackw...

Evidence of Late Triassic large tetrapods from the UK is rare. Here, we describe a track-bearing surface located on the shoreline near Penarth, south Wales, United Kingdom. The total exposed surface is c. 50 m long and c. 2 m wide, and is split into northern and southern sections by a small fault. We interpret these impressions as tracks, rather th...

The dinosaur track record features numerous examples of trackways with elongated metatarsal marks. Such ‘elongate tracks’ are often highly variable and characterized by indistinct outlines and abbreviated or missing digit impressions. Elongate dinosaur tracks are well-known from the Paluxy River bed of Texas, where some have been interpreted as ‘ma...

Differing from the extant physeteroids, macroraptorial sperm whales are currently regarded as apex predators of the Miocene seas based on several morphofunctional observations. Here, we estimate the bite force of Zygophyseter varolai, a macroraptorial physeteroid from lower upper Miocene strata of the Pietra leccese formation (Apulia, Italy) using...

Virtual and augmented reality (VR/AR) are new technologies with the power to revolutionize the study of morphology. Modern imaging approaches such as computed tomography, laser scanning, and photogrammetry have opened up a new digital world, enabling researchers to share and analyze morphological data electronically and in great detail. Because thi...

The emergence of bipedalism had profound effects on human evolutionary history, but the evolution of locomotor patterns within the hominin clade remains poorly understood. Fossil tracks record in vivo behaviours of extinct hominins, and they offer great potential to reveal locomotor patterns at various times and places across the human fossil recor...

Synopsis Substrate supportiveness is linked to the metabolic cost of locomotion, as it influences the depth to which the foot of a moving animal will sink. As track depth increases, animals typically reduce their speed to minimize any potential energetic imbalance. Here, we examine how self-selected speed in the Svalbard rock ptarmigan is affected...

Foot skeletons of small (young) hadrosaurid dinosaurs were compared with those of large (adult) hadrosaurids to assess the extent of pedal shape change during ontogeny. Foot skeletons of juvenile hadrosaurids were also compared with those of similar-sized adult, bipedal, non-hadrosaurian ornithischians to which the juvenile hadrosaurids were closer...

Bird necks display unparalleled levels of morphological diversity compared to other vertebrates, yet it is unclear what factors have structured this variation. Using three-dimensional geometric morphometrics and multi-variate statistics, we show that the avian cervical column is a hierarchical morpho-functional appendage, with varying magnitudes of...

Many parasites and hosts are embroiled in an on-going arms race that affects the evolution of each participant. One such battle is between parasitic nematodes and terrestrial gastropods which have co-evolved for 90–130 MY. Recently, snails have been shown to encase and kill invading nematodes using their shell as a defence mechanism. However, there...

The Vértesszőlős quarry, the Palaeolithic site where the “Samu” hominin fossil remains (Homo heidelbergensis) were found, is located in North West Hungary. The site is dated between the Early and Middle Pleistocene (ca. 310 ka). A short distance from where the Samu remains were found is an exposed surface of calcareous mudstone, preserving numerous...

Two new small tridactyl dinosaur tracks are found in the Middle Jurassic Bagå Formation of Bornholm and are interpreted as ornithopodian in origin. A skeletal fragment is identified as a crocodilian skull fragment. Previous finds of dinosaur tracks from the locality consist of two sizes of sauropods, a medium sized theropod and thyreophorans. The a...

Starting with his first report on fossil footprints from the Connecticut Valley over 180 years ago, Edward Hitchcock described what he interpreted as a burgeoning ancient fauna founded on ever-increasing nominal track diversity. For three decades, Hitchcock made countless contributions to ichnology, but his inference of thin-toed animals (Leptodact...

Whilst bones present a static view of extinct animals , fossil footprints are a direct record of the activity and motion of the track maker. Deep footprints are a particularly good record of foot motion. Such footprints rarely look like the feet that made them; the sediment being heavily disturbed by the foot motion. Because of this, such tracks ar...

Terrestrial vertebrate trace fossils are relatively abundant in mid-to-late Triassic and early Jurassic deposits in the British Isles but to date none at all have been recorded from the Rhaetian, the final stage of the Triassic. This represents a persistent gap in the terrestrial ichnological record. We present the first Rhaetian track to be recogn...

The feet of ground-dwelling birds retain many features of their dinosaurian ancestry. Experiments with living species offer insights into the complex interplay among anatomy, kinematics and substrate during the formation of Mesozoic footprints. However, a key aspect of the track-making process, sub-surface foot movement, is hindered by substrate op...

Three-dimensional (3D) surface scanning includes techniques of image acquisition and image processing. Among the former, hard- ware devices (e.g., portable and non-portable scanners, camera) capture images from the target, whereas image processing is conducted via specialized software, in which acquired images are processed to merge them into a sin...

Three parallel, manus-only sauropod trackways from the Coffee Hollow A-Male tracksite (Glen Rose Formation, Kendall County, Texas) were studied separately by researchers from the Heritage Museum of the Texas Hill Country and the Houston Museum of Natural Sciences. Footprint and trackway measurements generally show good agreement between the two gro...

Constraints of time, personnel, and financial resources made it necessary to devise a system for rapid, salvage screen-washing of a heterogeneous mixture of unconsolidated fossiliferous sediments and intermixed, unrelated sediments from a Neogene continental sinkhole site in northern Indiana, USA. We constructed three sets of an apparatus that used...

The surface area of feet in contact with the ground is a key morphological feature that influences animal locomotion. Underfoot pressures (and consequently stresses experienced by the foot), as well as stability of an animal during locomotion, depend on the size and shape of this area. Here we tested whether the area of a skeletal foot could predic...

Plesiosaurs are an enigmatic, diverse extinct group of Mesozoic marine reptiles well known for their unique body plan with two pairs of flippers and usually an elongated neck. The long neck evolved several times within the clade, yet the evolutionary advantages are not well understood. Previous studies have mainly focused on swimming speeds or flip...

Piscivorous birds have a unique suite of adaptations to forage under the water. One method aerial birds use to catch fish is the plunge dive, wherein birds dive from a height to overcome drag and buoyancy in the water. The kingfishers are a well-known clade that contains both terrestrially foraging and plunge-diving species, allowing us to test for...

We present the results of a two-year design study to developing virtual reality (VR) flow visualization tools for the analysis of dinosaur track creation in a malleable substrate. Using Scientific Sketching methodology, we combined input from illustration artists, visualization experts, and domain scientists to create novel visualization methods. B...

We present a two-year design study of developing virtual reality (VR) flow visualization tools for the analysis of dinosaur track creation using the Scientific Sketching design methodology. We involved 25 art and computer science students from a VR design course in a rapid visualization sketching cycle, guided by paleontologist collaborators throug...

Functional reconstructions of extinct animals represent a crucial step towards understanding palaeocological interactions, selective pressures and macroevolutionary patterns in the fossil record. In recent years, computational approaches have revolutionised the field of 'evolutionary biomechanics' and have, in general, resulted in convergence of qu...

Owing to an error in our muscle physiological cross-sectional area calculations, the range of bite force estimates for four models in our original analysis (maximum and minimum values presented in table 2 of [1]) are approximately 6% too high. This relates specifically to error in our calculation of the effect of a 208 pennation angle on muscle phy...

The collection and dissemination of vertebrate ichnological data is struggling to keep up with techniques that are becoming commonplace in the wider palaeontological field. A standard protocol is required to ensure that data is recorded, presented and archived in a manner that will be useful both to contemporary researchers, and to future generatio...

We documented trackways of free-living Crocodylus acutus on beaches at the mouths of Tamarindo and Ventanas estuaries, Costa Rica. Our crocodiles had estimated total lengths of 1–3 meters or more. Manus prints have five digits, with digits I–III bearing claw marks. Pes prints have four digits, with claw marks on digits I–III. The pes is plantigrade...

Trackways and tracemakers preserved together in the fossil record are rare. However, the co-occurrence of a drag mark, together with the dead animal that produced it, is exceptional. Here, we describe an 8.5 m long ammonite drag mark complete with the preserved ammo-nite shell (Subplanites rueppellianus) at its end. Previously recorded examples pre...

Over the past two decades, the development of methods for visualizing and analysing specimens digitally, in three and even four dimensions, has transformed the study of living and fossil organisms. However, the initial promise that the widespread application of such methods would facilitate access to the underlying digital data has not been fully a...

In 2015, a group of small predominantly tridactyl tracks was discovered in the lower Shawan Member of the Lufeng Formation during an expedition to the Dalishu area, Lufeng County, Yunnan Province, China. The tracks are attributable to Anomoepus, and although mostly tridactyl, they include a few examples with characteristic tetradactyl morphology. A...

Some primarily aquatic vertebrates make brief forays onto land, creating traces as they do. A lack of studies on aquatic trackmakers raises the possibility that such traces may be ignored or misidentified in the fossil record. Several terrestrial Actinopterygian and Sarcopterygian species have previously been proposed as possible models for ancestr...

Autochthonous Triassic sediments of the Vieux Emosson Formation near Lac d’Emosson, southwestern Switzerland, have yielded assemblages with abundant archosaur footprints that are assigned to chirotheriids based on pentadactyl pes and manus imprints with characteristic digit proportions. Tridactyl footprints formerly considered as those of dinosaurs...

The colossal size and body plan of sauropod dinosaurs are unparalleled in terrestrial vertebrates. However, to date, there have been only limited attempts to examine temporal and phylogenetic patterns in the sauropod bauplan. Here, we combine three-dimensional computational models with phylogenetic reconstructions to quantify the evolution of whole...

For more than 70 years unusual sauropod trackways have played a pivotal role in debates about the swimming ability of sauropods. Most claims that sauropods could swim have been based on manus-only or manus-dominated trackways. However none of these incomplete trackways has been entirely convincing, and most have proved to be taphonomic artifacts, e...

Sivatherium giganteum is an extinct giraffid from the Plio-Pleistocene boundary of the Himalayan foothills. To date, there has been no rigorous skeletal reconstruction of this unusual mammal. Historical and contemporary accounts anecdotally state that Sivatherium rivalled the African elephant in terms of its body mass, but this statement has never...

Details of skeletal reconstruction, scaling equations used, and convex hull volumes

The latest advances in dinosaur ichnology are showcased in this comprehensive and timely volume, in which leading researchers and research groups cover the most essential topics in the study of dinosaur tracks. Some assess and demonstrate state-of-the-art approaches and techniques, such as experimental ichnology, photogrammetry, biplanar X-rays, an...

Reconstructing the feeding mode of the latest common ancestor of deutero-stomes is key to elucidating the early evolution of feeding in chordates and allied phyla; however, it is debated whether the ancestral deuterostome was a tentaculate feeder or a pharyngeal filter feeder. To address this, we evaluated the hydrodynamics of feeding in a group of...

Estimates of body mass often represent the founding assumption on which bio-mechanical and macroevolutionary hypotheses are based. Recently, a scaling equation was applied to a newly discovered titanosaurian sauropod dinosaur (Dreadnoughtus), yielding a 59 300 kg body mass estimate for this animal. Herein, we use a modelling approach to examine the...

the substrate will not deform. Any given substrate can therefore only preserve a range of tracks from those animals which exert an underfoot pressure at approximately the yield strength of the sediment. However, rarely are substrates vertically homogeneous for any great depth, varying either due to heterogeneity across sediment layers, or from mech...

An ichnological and sedimentological study of the El Frontal dinosaur tracksite (Early Cretaceous, Cameros basin, Soria, Spain) highlights the pronounced intra-trackway variation found in track morphologies of four theropod trackways. Photogrammetric 3D digital models revealed various and distinct intra-trackway morphotypes, which reflect changes i...

An ichnological and sedimentological study of the El Frontal dinosaur tracksite (Early Cretaceous, Cameros basin, Soria, Spain) highlights the pronounced intra-trackway variation found in track morphologies of four theropod trackways. Photogrammetric 3D digital models revealed various and distinct intra-trackway morphotypes, which reflect changes i...

Fossil tracks represent a direct window onto the lives of extinct organisms, being formed and preserved in situ. Because track morphology is determined by limb motion, foot anatomy and substrate consistency, studies of fossil tracks can provide insight into producer, behaviour and palaeoenvironment. However, each determining factor is subject to va...

Ornithischian dinosaurs were primitively bipedal with forelimbs modified for grasping, but quadrupedalism evolved in the clade on at least three occasions independently. Outside of Ornithischia, quadrupedality from bipedal ancestors has only evolved on two other occasions, making this one of the rarest locomotory transitions in tetrapod evolutionar...

This is a meshed version of the Tyrannosaurid track described by Manning et al 2008. It's pretty high res, downsampled to 2,000,000 faces, created from 8 photos taken at the time. If using, please cite the paper, and acknowledge Falkingham.

This is a point cloud of the Tyrannosaurid track described by Manning et al 2008. It's pretty high res, created from 8 photos taken at the time. If using, please cite the paper, and acknowledge Falkingham.

The Las Cerradicas site (Tithonian-Berriasian), Teruel, Spain, preserves at least seventeen dinosaur trackways, some of them formerly attributed to quadrupedal ornithopods, sauropods and theropods. The exposure of new track evidence allows a more detailed interpretation of the controversial tridactyl trackways as well as the modes of locomotion and...