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Chapter 9. Paleozoic Vertebrate Ichnology of Grand Canyon National Park

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Chapter 9. Paleozoic Vertebrate Ichnology of Grand Canyon National Park Chapter 9. Paleozoic Vertebrate Ichnology of Grand Canyon National Park By Lorenzo Marchetti1, Heitor Francischini2, Spencer G. Lucas3, Sebastian Voigt1, Adrian P. Hunt4, and Vincent L. Santucci5 1Urweltmuseum GEOSKOP / Burg Lichtenberg (Pfalz) Burgstraße 19 D-66871 Thallichtenberg, Germany 2Universidade Federal do Rio Grande do Sul (UFRGS) Laboratório de Paleontologia de Vertebrados and Programa de Pós-Graduação em Geociências, Instituto de Geociências Porto Alegre, Rio Grande do Sul, Brazil 3New Mexico Museum of Natural History and Science 1801 Mountain Road N.W. Albuquerque, New Mexico, 87104 4Flying Heritage and Combat Armor Museum 3407 109th St SW Everett, Washington 98204 5National Park Service Geologic Resources Division 1849 “C” Street, NW Washington, D.C. 20240 Introduction Vertebrate tracks are the only fossils of terrestrial vertebrates known from Paleozoic strata of Grand Canyon National Park (GRCA), therefore they are of great importance for the reconstruction of the extinct faunas of this area. For more than 100 years, the upper Paleozoic strata of the Grand Canyon yielded a noteworthy vertebrate track collection, in terms of abundance, completeness and quality of preservation. These are key requirements for a classification of tracks through ichnotaxonomy. This chapter proposes a complete ichnotaxonomic revision of the track collections from GRCA and is also based on a large amount of new material. These Paleozoic tracks were produced by different tetrapod groups, such as eureptiles, parareptiles, synapsids and anamniotes, and their size ranges from 0.5 to 20 cm (0.2 to 7.9 in) footprint length. As the result of the irreversibility of the evolutionary process, they provide useful information about faunal composition, faunal events, paleobiogeographic distribution and biostratigraphy. Of note, these vertebrate trace fossils are always in situ (i.e., not transported before burial) and preserved in lithofacies representing different continental paleoenvironments (marginal marine, alluvial plain and desert), therefore they can provide useful paleoecological information. Also, the occurrence of surfaces with long trackways provides 333 fundamental insights into the locomotion of the trackmakers on inclined planes, such as the paleosurfaces of dune foresets. Vertebrate Ichnology Vertebrate ichnology is the discipline that principally studies the traces left on the substrate by vertebrates during their lives. These traces can be produced by different behaviors, such as: walking, running, hopping, resting, swimming, etc. The main subject of vertebrate ichnology is thus the study of tracks left on the sediment during locomotion. The tracks can be preserved as concave epirelief (natural mold) and convex hyporelief (natural cast) and can be preserved on multiple layers, forming undertracks, underprints and overtracks. On the actual trampled surface, these ichnofossils sometimes preserve expulsion rims (i.e., a marginal rim of displaced sediment produced by the footfall; Allen 1997). Their occurrence generally indicates subaerial exposure, the presence of a substrate adequate to record the footprint impression, and a non-erosive and rapid burial of the actual track surface. These conditions are common in intermittently wet environments such as lake margins, alluvial plains, tidal flats, fluvial channels, and/or in desert environments characterized by rapid sedimentary cover (dune foreset surfaces). In intermittently wet environments, they are often associated with mud cracks, rain drops, wave ripples, tool marks, and microbial structures, among other sedimentary structures, and with invertebrate tracks and burrows. In desert environments, they are often associated with sand avalanches, wind ripple crests and invertebrate tracks and burrows. With regard to the Paleozoic, only footprints of quadrupedal vertebrates are known, which include front (manus) and hind (pes) foot imprints. Tail, body, and digit drag marks may be present, but are not relevant for footprint classification. These footprints are usually arranged in sequences of pes- manus couples, arranged in two parallel rows that form a trackway. From the footprint and trackway parameters, it is possible to infer the size, weight, gait and speed of the trackmakers (e.g., Leonardi 1987). Generally, the Paleozoic terrestrial vertebrates were not fast moving, and most of them adopted a sprawling gait as seen in modern lizards and salamanders. Their size was also not very large, especially before the Guadalupian (maximum footprint length of about 20 cm or 7.9 in). Tetrapod footprints are classified by their morphology, and especially by the anatomy-consistent morphology (e.g., Marchetti et al. 2019a). This is the morphology that reflects the actual lower side of the trackmaker foot, and thus it is strongly connected with the trackmaker’s anatomical structure, which is subject to irreversible evolution. Tracks showing these anatomy-consistent features are regarded as well-preserved. This kind of preservation, named morphological preservation, can be evaluated through a numerical scale (Marchetti et al. 2019a). In the study of tetrapod tracks, it is necessary to exclude from the analysis those tracks that are not anatomy-consistent due to loss of information or deformation during (e.g., footprint wall collapse) and after track registration in the sediment (e.g., superimposition of other footprints and sedimentary structures and partial erosion). The footprints preserved on the dune foreset surfaces show a peculiar process that causes loss of information of the anatomy: the oblique direction of the gravity force due to the dune surface inclination (usually more than 30°). This causes the sliding of the trackmaker digits during locomotion, and asymmetrical trackway patterns and footprint preservation in trackways oblique to the slope (Figure 9-1A–D). Also, the trackmaker speed was influenced, with very close pes-manus 334 couples and secondary overstep in trackways directed upslope (slow gait) and well-spaced pes-manus couples and primary overstep (fast gait) in trackways directed downslope (Figure 9-1A–D). Overstep is the preservation of the pes in front of the manus, different than what is usually seen (manus in front of the pes). This has been observed in trackways of the same track type (Varanopus), therefore these different gaits and morphologies are likely not due to different trackmakers (Marchetti et al. 2019b, 2019c). Figure 9-1. Track formation and track measurements (LORENZO MARCHETTI & SPENCER LUCAS). A. Track formation on a dune foreset surface with different kind of progression: B. directly upslope, C. directly downslope and D. transversely upslope. E. Track and trackway measurements. FL=footprint length, FW=footprint width, psL=palm/sole length, psW=palm/sole width, DL=digit length, div=digit divergence, V=digit V, SLp=stride length of the pes, SLm=stride length of the manus, PLp=(oblique) pace length of the pes, PAp=pace angulation of the pes, PAm=pace angulation of the manus, PLm=(oblique) pace length of the manus, LPp=length of pace of the pes, LPm=length of pace of the manus, WPp=width of pace of the pes, WPm=width of pace of the manus, Divp=divarication from midline of the pes, Divm=divarication from midline of the manus, Dmp=manus-pes distance, BL=gleno-acetabular length. After the selection of well-preserved material, the footprints and trackways are described and measured. The digit impressions are counted with Roman numbers, starting from the inner (medial) side of the trackway. Tracks with longer external digit impressions (commonly digit IV) are named ectaxonic. Common track measurements (Figure 9-1E) include the foot length and width, and the 335 sole/palm impression length and width; all compared to the digit III axis. Other measurements are the digit length and the interdigital angles. The distance between two consecutive tracks of the same kind (manus or pes) on the same trackway side is named stride length. The distance between two consecutive tracks of the same kind (manus or pes) on opposite trackway sides is termed (oblique) pace length, and the angle between two consecutive (oblique) pace lengths is termed pace angulation. The line equidistant from the manus-pes couples is the trackway midline. Other measurements are the trackway width, the manus-pes distance and the manus and pes divarication (i.e., its rotation), all compared to the trackway midline. Ichnotaxonomy is the discipline that classifies tetrapod tracks and other traces. It is considered a parataxonomy because it is disentangled from the taxonomic classification of the producers. In fact, it is quite difficult to relate a track type directly to a trackmaker genus or species, with some exceptions (e.g., Voigt et al. 2007; Marchetti et al. 2017a). Tetrapod track ichnotaxa usually correspond to higher taxonomic levels than species and genera, because similar species and genera may produce very similar footprints (e.g., all of the “pelycosaur” synapsid genera, except for varanopids, may be attributed to a single ichnogenus, Dimetropus). The tetrapod tracks are classified with a binomial nomenclature (ichnogeneric epithet + ichnospecific epithet), following the procedures of the International Code of Zoological Nomenclature (1999). Ichnofamilies can be used as well, but are generally less extensively studied. The relevant criteria (ichnotaxobases) are the anatomy-consistent foot morphology and the trackway pattern, including measurements. The list of ichnotaxa from a formation
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