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On the presence of Ichniotherium in the Coconino Sandstone (Cisuralian) of the Grand Canyon and remarks on the occupation of deserts by non-amniote tetrapods

  • Heitor FrancischiniEmail author
  • Spencer G. Lucas
  • Sebastian Voigt
  • Lorenzo Marchetti
  • Vincent L. Santucci
  • Cassandra L. Knight
  • John R. Wood
  • Paula Dentzien-Dias
  • Cesar L. Schultz
Research Paper


The colonization of deserts by tetrapods occurred for the first time in the late Paleozoic. In spite of amniotes being the most abundant and diverse taxon in such environments, fossil tracks indicate that anamniotes also inhabited late Paleozoic deserts. In this paper, the presence of the tetrapod-footprint ichnotaxa Ichniotherium sphaerodactylum and cf. Ichniotherium is documented in the eolian Coconino Sandstone (early Permian), based on 16 trackways found at several localities in Arizona, USA. Because there is a strong association between Ichniotherium and different species of diadectomorphs, we also discuss some aspects of the ichnotaxonomy of this ichnogenus. Diadectomorpha is considered the sister taxon of Amniota and, as a consequence, the tracks described in this paper represent the oldest evidence of occupation of deserts by non-amniote tetrapods. The presence of Ichniotherium in this environmental context also sheds light on the paleobiology of Diadectomorpha and, as a result, the emergence of features typically related to Amniota. The ichnofauna of the Coconino Sandstone has been used as a model for the Chelichnus ichnofacies, which supposedly indicates a low-diversity desert fauna. On the other hand, the tracks described here demonstrate that diadectomorphs were also important faunal components of such deserts and suggests that the significance of this ichnofacies should be reconsidered.


Leonardian Permian Arizona Ichnology Diadectomorpha Ichniotherium sphaerodactylum 



We thank Ms. Anne Miller (NPS, GRCA) and Mr. Bill Ludlow (Mesa, Arizona) for their field assistance. We are also indebted to Mrs. Coleen Hyde (NPS, GRCA), Mrs. Janet Gillette, Dr. David Gillette (both MNA), Dr. Andrew Farke, Mr. Gabriel Santos (both RAM), Dr. Patricia Holroyd (UCMP), Dr. Daniel Brinkman (YPM), Dr. Robert Emry and Amanda Millhouse (both USNM) for allowing the analysis of specimens under their care. We also thank Dr. Martin G. Lockley (UCD), Dr. Jahn J. Hornung and Dr. Mike Reich (both formerly at Georg-August-Universität Göttingen) for their comments which greatly improved the early version of this manuscript. Voltaire Paes Neto (UFRGS) is acknowledged by the artwork presented in Fig. 8. This work was supported by the Conselho Nacional de Desenvolvimento Científico e Tecnológico and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior with grants to Heitor Francischini (Process numbers: 150623/2018-6 and 88881.133764/2016-01, respectively).

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Copyright information

© Paläontologische Gesellschaft 2019

Authors and Affiliations

  • Heitor Francischini
    • 1
    Email author
  • Spencer G. Lucas
    • 2
  • Sebastian Voigt
    • 3
  • Lorenzo Marchetti
    • 3
  • Vincent L. Santucci
    • 4
  • Cassandra L. Knight
    • 5
  • John R. Wood
    • 6
  • Paula Dentzien-Dias
    • 7
  • Cesar L. Schultz
    • 1
  1. 1.Programa de Pós-Graduação em GeociênciasInstituto de Geociências, Universidade Federal do Rio Grande do SulPorto AlegreBrazil
  2. 2.New Mexico Museum of Natural History and ScienceAlbuquerqueUSA
  3. 3.Urweltmuseum GEOSKOP. Burg Lichtenberg (Pfalz)ThallichtenbergGermany
  4. 4.Geologic Resources DivisionNational Park ServiceWashingtonUSA
  5. 5.PaleoWorks ConsultingPortlandUSA
  6. 6.Geologic Resources DivisionNational Park ServiceLakehoodUSA
  7. 7.Laboratório de Geologia e PaleontologiaInstituto de Oceanografia, Universidade Federal do Rio GrandeRio GrandeBrazil

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