Abstract
The diverse ichnofaunas of the Late Triassic have been studied for almost 200 years. During the Late Triassic, facies favorable for the preservation of trace fossils were the result of low sea levels, monsoonal climates and the development of extensive depositional basins as Pangea began to fragment. The most abundant vertebrate trace fossils in the Late Triassic are tetrapod tracks, including Brachychirotherium, Chirotherium, “Parachirotherium,” Synaptichnium, Atreipus, Grallator, Eubrontes, Banisterobates, Trisauropodiscus, Evazoum, Tetrasauropus, Pseudotetrasauropus, Eosauropus, Apatopus, Batrachopus, Rhynchosauroides, Gwyneddichnium, Procolophonichnium, Chelonipus, Brasilichnium and Dicynodontipus. There are five tetrapod footprint biochrons of Triassic age that can be identified across the Pangaean footprint record. Coprolites are the second most abundant vertebrate trace fossils in the Late Triassic and include Heteropolacopros, Alococoprus, Dicynodontocopros, Liassocoprus, Saurocoprus, Strabelocoprus, Malericoprus, Falcatocoprus and Revueltobromus. Coprolites are useful in biochronology in the Late Triassic. Consumulites, dentalites (new term for bite marks), and burrows are moderately common in the Late Triassic. Nests and gastroliths are rare. All groups of vertebrate trace fossils demonstrate different diversity and abundance patterns through the Phanerozoic. Most vertebrate trace fossils have their earliest occurrences in the Devonian. The early Permian is an acme for both tracks and coprolites. The Late Triassic yields abundant tracks and coprolites, and tracks are also common in the Early Jurassic. The Jurassic and Cretaceous represent the times with the greatest diversity of vertebrate traces (tracks, coprolites, consumulites, dentalites, nests and gastroliths). The Quaternary also represents a time of vertebrate ichnological diversity (tracks, coprolites, regurgitalites, nests and burrows).
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Acknowledgments
We thank those in charge of collections in North and South America, Europe and Asia who have allowed us access to collections over the years to study vertebrate trace fossils. Paula Denzien-Dias and Larry Tanner provided insightful and helpful reviews.
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Appendix
Appendix
Systematic Ichnotaxonomy
Gaspeichnus, ichnogen. nov.
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Type ichnospecies: Gaspeichnus complexus.
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Included ichnospecies: Known only from the type ichnospecies.
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Etymology: From the Gaspé Peninsula and the Greek ichnos (trace).
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Distribution: Late Devonian to Early Triassic.
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Diagnosis: Elongate and irregularly sinuous borings of small size (diameter 0.1–0.2 mm) with irregular width in a coprolite substrate.
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Discussion: This is possibly the oldest example of coprophagy and is certainly the oldest example of coprolite modification by another organism.
Brachaniec et al. (2015: fig. 6A–B) reported sinuous surface borings of Early Triassic coprolites from Poland that we refer to Gaspeichnus. They noted that Gaspeichnus occurs in 57% of the coprolites that contain fossils.
The size and sinuosity are similar to the morphology of nematode trails, although Gaspeichnus is more irregular in its undulations (Moussa 1970; Baliński et al. 2013; Brachaniec et al. 2015). We tentatively interpret Gaspeichnus as being produced by a nematode. The coprolite substrate alone differentatiates it from other sinuous trace fossils.
Gaspeichnus complexus, ichnosp. nov. (Fig. 12.17)
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Holotype: RSM 2002. 59.166, coprolite with borings.
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Etymology: From Latin complexus in allusion to the complicated structure of the borings.
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Type locality: Escuminac Bay, Gaspé Peninsula, Quebec, Canada.
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Type horizon: Escuminac Formation (Upper Devonian).
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Distribution: As for ichnogenus.
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Referred specimens: None.
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Diagnosis: As for ichnogenus.
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Description: RSM 2002. 59.166 is a vertebrate coprolite of approximately cylindrical shape with an irregular margin. It is 22 mm long and 6 mm wide. It contains two elongate and sinuous borings that do not overlap. The borings vary in width from 0.1 to 0.2 mm in width and from 8 to 12 mm in length. The sinuosity is irregular.
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Discussion: The Escuminac Formation yields a diverse bromalite ichnofauna that has been described by McAllister (1996).
Holotype of Gaspeichnus complexus Hunt et al. (2017) (RSM 2002.59.166) from the Upper Devonian Escuminac Formation of Quebec, Canada
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Hunt, A.P., Lucas, S.G., Klein, H. (2018). Late Triassic Nonmarine Vertebrate and Invertebrate Trace Fossils and the Pattern of the Phanerozoic Record of Vertebrate Trace Fossils. In: Tanner, L. (eds) The Late Triassic World. Topics in Geobiology, vol 46. Springer, Cham. https://doi.org/10.1007/978-3-319-68009-5_12
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