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Dinosaur Tracks

Dinosaur Tracks: The Next Steps

Peter L. Falkingham
Daniel Marty
Annette Richter
Copyright Date: 2016
Published by: Indiana University Press
Pages: 428
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    Dinosaur Tracks
    Book Description:

    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, and a numerical scale for quantifying the quality of track preservation. The high diversity of these up-to-date studies underlines that dinosaur ichnological research is a vibrant field, that important discoveries are continuously made, and that new methods are being developed, applied, and refined. This indispensable volume unequivocally demonstrates that ichnology has an important contribution to make toward a better understanding of dinosaur paleobiology. Tracks and trackways are one of the best sources of evidence to understand and reconstruct the daily life of dinosaurs. They are windows on past lives, dynamic structures produced by living, breathing, moving animals now long extinct, and they are every bit as exciting and captivating as the skeletons of their makers.

    eISBN: 978-0-253-02114-4
    Subjects: Paleontology
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Table of Contents

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  1. Front Matter (pp. i-vi)
  2. Table of Contents (pp. vii-viii)
  3. Acknowledgments (pp. ix-2)
  4. Introduction (pp. 3-11)
    Peter L. Falkingham, Daniel Marty and Annette Richter

    The dinosauria are one of the most morphologically diverse groups of terrestrial vertebrates (Alexander, 1989), spanning several orders of magnitude in size from the smallest hummingbird to the largest sauropods. Ancestrally bipedal, groups within the Dinosauria evolved into a range of habitually and facultatively bipedal and quadrupedal animals. Their skeletons have been found on every continent (Weishampel, Dodson, and Osmólska, 2004), and their fossilized footprints are known from all except Antarctica.

    The public perception of dinosaurs comes almost exclusively via their skeletons, and much of our knowledge about how these enigmatic animals looked and lived comes from osteological information. But...

  5. Part 1. Approaches and Techniques for Studying Dinosaur Tracks
    • 1 Experimental and Comparative Ichnology (pp. 15-27)
      Jesper Milàn and Peter L. Falkingham

      One of the main problems faced in paleoichnology is the delicate relationship between the organism and the sediments it leaves its tracks and traces in. Since the first scientific report of comparisons between fossil and modern tracks, researchers have turned to making experiments and comparing tracks and trackways of modern animals in order to interpret fossil tracks and traces. The easiest experimental approach is simply to make living analogues to the fossil animals walk through soft sediment and directly study the tracks they produce. Modern, more sophisticated experimental procedures include laboratory-controlled settings with sediments of different properties and model feet...

    • 2 Close-Range Photogrammetry for 3-D Ichnology: The Basics of Photogrammetric Ichnology (pp. 29-55)
      Neffra Matthews, Tommy Noble and Brent Breithaupt

      Vertebrate trace fossils reflect the complex interrelationship between an animal’s activities and the substrate (Manning, 2004; Falkingham, 2014), which is well represented in the ichnofaunal record of Mesozoic dinosaurs (Thulborn, 1990; Lockley, 1991; Lockley and Meyer, 2000; Wright and Breithaupt, 2002). As such, these unique three-dimensional (3-D) fossils warrant detailed recordation that captures their multidimensional features to fully understand formation and preservation of the ichnofossils, as well as dinosaur community dynamics (Lockley, 1986; Falkingham, 2014). Currently, the most cost-efficient and high-resolution mechanism to collect 3-D digital data of trace fossils is through the proper use of photogrammetry. Digital ichnological and...

    • 3 The Early Cretaceous Dinosaur Trackways in Münchehagen (Lower Saxony, Germany): 3-D Photogrammetry as Basis for Geometric Morphometric Analysis of Shape Variation and Evaluation of Material Loss during Excavation (pp. 57-71)
      Oliver Wings, Jens N. Lallensack and Heinrich Mallison

      Lower cretaceous sandstones in lower saxony, northern Germany, are well known for their abundant fossil dinosaur tracks. One of the most productive sites is Münchehagen, which is well known for the only German Cretaceous sauropod trackways and hundreds of tracks of ornithopods and theropods, often forming long individual trackways with dozens of consecutive footprints. The largest theropod trackway T3 from the layer that has produced the best preserved true tracks (Lower Level) shows variations in the footprint morphology that allow use of this data as an example for studying the variability of tridactyl dinosaur track measurements.

      Photogrammetric high-resolution three-dimensional (3-D)...

    • 4 Applying Objective Methods to Subjective Track Outlines (pp. 73-81)
      Peter L. Falkingham

      Formally communicating the morphology of a track generally occurs via a two-dimensional (2-D) medium (i.e., paper). For this reason, track outlines are often used to convey the geometry and morphology of a track. However, these track outlines are routinely subjective, based on the interpreter’s opinion of where the track ends and the surrounding undeformed substrate begins. Although such outlines are not a problem themselves, any subsequent application of numerical objective methods such as multivariate analyses or equations using track parameters can be strongly influenced by the subjective nature of the outline. This effect is compounded in deeper tracks with sloping...

    • 5 Beyond Surfaces: A Particle-Based Perspective on Track Formation (pp. 83-91)
      Stephen M. Gatesy and Richard G. Ellis

      Fossil footprints record unrivaled evidence of behavior in long extinct species. For students of dinosaur locomotion, tracks offer clues about gait, speed, limb posture, foot motion (kinematics), foot loading (kinetics), and social behavior (e.g., Ostrom, 1972; Alexander, 1976; Thulborn and Wade, 1984, 1989; Padian and Olsen, 1989; Gatesy et al., 1999; Milàn, 2006; Graversen, Milàn, and Loope, 2007; Pérez-Lorente and Herrero Gascón, 2007; Ishigaki and Lockley, 2010; Avanzini, Piñuela, and Garcia-Ramos, 2011; Falkingham, 2014). Yet tracks must be studied differently from body fossils. Although complementary to skeletal remains, footprints are purely sedimentary structures that preserve traces of anatomy only indirectly....

    • 6 A Numerical Scale for Quantifying the Quality of Preservation of Vertebrate Tracks (pp. 93-99)
      Matteo Belvedere and James O. Farlow

      From its beginning, vertebrate ichnology has described fossilized footprints in a qualitative, descriptive way. At the same time, considerable effort has gone into illustrating footprint morphology. In recent years, new technologies (e.g., laser-scanning and close-range photogrammetry) and methods (e.g., geometric morphometrics) have allowed more objective, quantitative approaches to vertebrate ichnology. However, quantitative shape analyses need to be based on data of high quality, and comparisons are best made between tracks comparable in quality of preservation. Thus, determining which footprints constitute the most reliable sample for quantitative analyses is fundamental for the progress of ichnology.

      We introduce here a numerical scale...

    • 7 Evaluating the Dinosaur Track Record: An Integrative Approach to Understanding the Regional and Global Distribution, Scientific Importance, Preservation, and Management of Tracksites (pp. 101-117)
      Luis Alcalá, Martin G. Lockley, Alberto Cobos, Luis Mampel and Rafael Royo-Torres

      Many papers on fossil tracks, from many regions of the world have been published in the last two decades, and this rapid increase in documentation has itself generated the idea of a dinosaur “footprint renaissance” marked by a landslide of new discoveries and documentation. Many of these papers mention the significance of these sites in terms of selected variables such as size of site, number of tracks, new or unknown ichnotaxa, new stratigraphic or geographic occurrence, trackmaker behavioral implications, and so forth. However, the significance of fossil tracksites is often not comprehensively discussed or evaluated in such a way as...

  6. Part 2. Paleobiology and Evolution from Tracks
    • 8 Iberian Sauropod Tracks through Time: Variations in Sauropod Manus and Pes Track Morphologies (pp. 121-137)
      Diego Castanera, Vanda F. Santos, Laura Piñuela, Carlos Pascual, Bernat Vila, José I. Canudo and José Joaquin Moratalla

      The iberian sauropod track record has yielded more than 100 sauropod tracksites ranging in age from the Middle Jurassic (Bathonian) to the Late Cretaceous (Maastrichtian). During this wide range of time, four different types of manus prints can be differentiated, changing in morphology from (1) speech-bubble–shaped with a prominent claw mark in digit I (Middle Jurassic), (2) kidney-shaped with a claw mark in digit I or (3) without a claw mark in digit I (Late Jurassic and Early Cretaceous), to (4) horseshoe-shaped (Cretaceous). Pes prints are slightly more conservative in morphology through the Mesozoic and are generally subtriangular. They...

    • 9 The Flexion of Sauropod Pedal Unguals and Testing the Substrate Grip Hypothesis Using the Trackway Fossil Record (pp. 139-151)
      Lee E. Hall, Ashley E. Fragomeni and Denver W. Fowler

      Sauropod pedes exhibit a unique, highly derived pedal ungual morphology and articulation. During plantar flexion of the pes, the spade-like, laterally compressed unguals are rotated ventrally and deflected laterally across the front of the pes so the claws overlap in an en-echelon fashion; this positions the dorsal margins ventrally and the medial sides face posteriorly, creating a hoe-like structure oriented perpendicular to the plane of limb movement. Several functional hypotheses have been erected in an attempt to explain this feature. One, the miring-avoidance via substrate grip hypothesis, suggests orientation of the unguals during plantar flexion was utilized to generate traction...

    • 10 Dinosaur Swim Track Assemblages: Characteristics, Contexts, and Ichnofacies Implications (pp. 153-181)
      Andrew R. C. Milner and Martin G. Lockley

      Traces made by swimming tetrapods are simply known as “swim tracks.” These trace fossils are of interest to paleontologists because they provide insight into the behavior of past vertebrates in aquatic environments. However, swim tracks have always been a controversial subject for several reasons. Often swim tracks show irregular morphologies and are incomplete, so interpretation of them can be problematic. Unlike tracks made by animals walking on firm ground, which supports most or all of their weight, swimming tetrapods are fully or partially buoyant, and if their feet or hands come into contact with the subaqueous substrate, they will register...

    • 11 Two-Toed Tracks through Time: On the Trail of “Raptors” and Their Allies (pp. 183-201)
      Martin G. Lockley, Jerald D. Harris, Rihui Li, Lida Xing and Torsten van der Lubbe

      The two-toed, or didactyl, tracks of deinonychosaurian dinosaurs, popularly known as “raptors,” are among the most distinctive theropod tracks known. Including the first confirmed report from China in 1994, a total of 16 tracksites have been recognized, all from Cretaceous strata. These include nine Chinese, two Korean, three North American, and two European occurrences. Many of these tracks have been assigned to four ichnogenera:Velociraptorichnus(two ichnospecies),Dromaeopodus, Menglongipus, andDromaeosauripus(three ichnospecies). Most of the tracks have been attributed to dromaeosaurid theropods, but in the case of the largest sample, from Germany, a troodontid trackmaker is inferred.

      Here we...

    • 12 Diversity, Ontogeny, or Both?: A Morphometric Approach to Iguanodontian Ornithopod (Dinosauria: Ornithischia) Track Assemblages from the Berriasian (Lower Cretaceous) of Northwestern Germany (pp. 203-225)
      Jahn J. Hornung, Annina Böhme, Nils Schlüter and Mike Reich

      Identifying the causes of morphological variation (including taxonomic diversity, ontogeny, sexual dimorphism, and individual variation) observed in a set of vertebrate tracks – especially from different closely related trackmaker species – is difficult and often not straightforward due to imperfect knowledge of biological variation in the autopodia of the trackmakers, and a number of ethological, preservational, and taphonomical influences. Here we use multivariate data sets obtained from 14 homologous twodimensional (2-D) landmarks to evaluate the range and potential causes of variation in iguanodontian ornithopod pes tracks from the Berriasian of northwestern Germany.

      In order to minimize the nonanatomical-related effects associated...

    • 13 Uncertainty and Ambiguity in the Interpretation of Sauropod Trackways (pp. 227-243)
      Kent A. Stevens, Scott Ernst and Daniel Marty

      Trackway interpretation, the drawing of inferences about a trackmaker and its movements from a pattern of trace impressions, is examined from the perspective of the information in the pattern of individual tracks along a trackway, with emphasis here on sauropod trackways. Although trackways are commonly regarded as direct records of locomotion behavior, their interpretation is in fact less straightforward than is often expected. Even the basic estimation of trackmaker size (e.g., glenoacetabular distance, a common proxy for trackmaker size) is not generally valid. Moreover, without knowledge of trackmaker size, any observed pattern of manus and pes tracks has arbitrarily many...

    • 14 Dinosaur Tracks as “Four-Dimensional Phenomena” Reveal How Different Species Moved (pp. 245-255)
      Alberto Cobos, Francisco Gascó, Rafael Royo-Torres, Martin G. Lockley and Luis Alcalá

      Although thousands of dinosaur tracks have been found worldwide, three-dimensional (3-D) natural track casts are still relatively poorly documented. Those few that have been published, however, sometimes show impressions of reticulated skin, toe pads, and scratch marks made by scales and may even record how the sole of the foot bore the trackmaker’s weight. In very exceptional circumstances, such casts can even preserve evidence of distal limb kinematics of the trackmaker by recording the movement of the feet during track-making: in other words, footfall or footfall registration dynamics. Here we present a description of natural track casts that show all...

  7. Part 3. Ichnotaxonomy and Trackmaker Identification
    • 15 Analyzing and Resolving Cretaceous Avian Ichnotaxonomy Using Multivariate Statistical Analyses: Approaches and Results (pp. 259-309)
      Lisa G. Buckley, Richard T. McCrea and Martin G. Lockley

      Several new ichnotaxa of avian tracks have been described in recent years, adding to the known ichnodiversity of Cretaceous avians. The naming of new avian ichnospecies and ichnogenera has resulted in the creation of several avian ichnofamilies, but due to the challenges of documenting bird tracks, there are several ichnogenera that to date remain unassigned to any ichnofamily. Multivariate statistical analyses can be used to quantitatively test avian ichnotaxonomic assignments. Ichnospecies within the ichnofamilies Avipedidae (Aquatilavipes swiboldae, A. izumiensis, andA. curriei), Ignotornidae (Ignotornis mcconnelli, I. yangi, I. gajinensis, Hwangsanipes choughi, Goseongornipes ichnosp., G. markjonesi), Koreanaornipodidae (Koreanaornis hamanensis, K. dodsoni,...

    • 16 Elusive Ornithischian Tracks in the Famous Berriasian (Lower Cretaceous) “Chicken Yard” Tracksite of Northern Germany: Quantitative Differentiation between Small Tridactyl Trackmakers (pp. 311-331)
      Tom Hübner

      The “Chicken Yard” tracksite in Northern Germany, stratigraphically located in the Berriasian (lowermost Cretaceous) Bückeberg Formation, is famous for the extraordinary abundance of typical tridactyl theropod dinosaur tracks and the first didactyl footprints of deinonychosaurian theropods from Europe. Among the vast number of theropod tracks, there are a few small tridactyl tracks, which usually differ from the former by weaker mesaxony, wider divarication angles, stronger symmetry, broader digits, and a rounder and more consistent metatarsal imprint. These features indicate ornithischian track makers, but the strong morphological variation often hampers unambiguous differentiation from less well-preserved theropod tracks. Thus, measurements of the...

  8. Part 4. Depositional Environments and Their Influence on the Track Record
    • 17 Too Many Tracks: Preliminary Description and Interpretation of the Diverse and Heavily Dinoturbated Early Cretaceous “Chicken Yard” Ichnoassemblage (Obernkirchen Tracksite, Northern Germany) (pp. 335-357)
      Annette Richter and Annina Böhme

      The moderately to heavily dinoturbated Berriasian Chicken Yard level from the Obernkirchen tracksite (Lower Saxony, northern Germany) is preliminarily described and analyzed. Its ichnoassemblage is characterized by an extraordinary high track density composed of several different morphotypes and size classes of theropod and ornithopod true tracks with an overall similar preservation quality. The occurrence of didactyl tracks of a new, so far unnamed ichnotaxon that can be attributed to deinonychosaurian dinosaurs is particularly remarkable. Despite the high track density and associated frequent overprinting of tracks, several trackways were identified. Their orientation analysis tends toward a primarily bimodal orientation pattern despite...

    • 18 Dinosaur Tracks in Eolian Strata: New Insights into Track Formation, Walking Kinetics, and Trackmaker Behavior (pp. 359-365)
      David B. Loope and Jesper Milàn

      Dinosaur tracks are abundant in wind-blown Mesozoic deposits, but the nature of loose eolian sand makes it difficult to determine how they are preserved. This also raises the questions: Why would dinosaurs be walking around in dune fields in the first place? And, if they did go there, why would their tracks not be erased by the next wind storm?

      Most dunes today form only in deserts and along shorelines – the only sandy land surfaces that are nearly devoid of plants. Normally plants slow the wind at the ground surface enough that sand will not move even when the...

    • 19 Analysis of Desiccation Crack Patterns for Quantitative Interpretation of Fossil Tracks (pp. 367-379)
      Tom Schanz, Maria Datcheva, Hanna Haase and Daniel Marty

      This chapter presents a conceptual approach to interpret fossil track environments employing the progress made in soil mechanics regarding understanding and modeling of soil desiccation cracks. It must be emphasized that the thorough analysis of soil desiccation phenomena is crucial for the understanding of track formation and preservation processes and the interpretation of the paleoenvironmental setting associated with fossil track-bearing strata. The basis for the proposed method is the generally accepted fact that crack formation depends on the intrinsic state properties of the soil and on the boundary conditions the soil layer is exposed to during desiccation. Both the observations...

    • 20 A Review of the Dinosaur Track Record from Jurassic and Cretaceous Shallow Marine Carbonate Depositional Environments (pp. 381-391)
      Simone D’Orazi Porchetti, Massimo Bernardi, Andrea Cinquegranelli, Vanda Faria dos Santos, Daniel Marty, Fabio Massimo Petti, Paulo Sá Caetano and Alexander Wagensommer

      An extensive literature on dinosaur ichnology is available today, with hundreds of papers describing dinosaur tracks in different depositional settings. In recent years, it has become common practice in paleontology to gather data in databases to ease organization, managing, and analysis of large amounts of information. A review of the occurrences of dinosaur tracks in shallow marine carbonate depositional environments is presented here, based on 131 papers published between 1917 and 2013 and describing a total of 212 tracksites. The raw data set, which adopts the tracksite as the basic unit, reveals an abundance of saurischian footprints and a relative...

  9. Paleoenvironment Reconstructions of Vertebrate Tracksites in the Obernkirchen Sandstone, Lower Cretaceous of Northwest Germany (pp. 393-398)
    Jahn J. Hornung, Annette Richter and Frederik Spindler
  10. Dinosaur Track Terminology: A Glossary of Terms (pp. 399-402)
    Daniel Marty, Peter L. Falkingham and Annette Richter
  11. Index (pp. 403-412)
  12. List of Contributors (pp. 413-414)
  13. Back Matter (pp. 415-417)