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Chapter 6. Paleozoic Vertebrate Paleontology of Grand Canyon National Park: Research History, Resources, and Potential

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Chapter 6. Paleozoic Vertebrate Paleontology of Grand Canyon National Park: Research History, Resources, and Potential Chapter 6. Paleozoic Vertebrate Paleontology of Grand Canyon National Park: Research History, Resources, And Potential By John-Paul Michael Hodnett1 and David Kenneth Elliott2 1Maryland-National Capital Park and Planning Commission Department of Parks and Recreation, Archaeology Program 8204 McClure Road Upper Marlboro, Maryland 20772 2Northern Arizona University School of Earth and Sustainability, Geology Division Flagstaff, Arizona 86011 Introduction The Grand Canyon is an awe-inspiring place and is a physical testament of deep time. More than a 150 years’ worth of geologic research has yielded a rich paleontological record, documenting approximately a half billion years of life on this planet. Much of the Grand Canyon is composed of Paleozoic rocks, with the oldest going back to the Cambrian. The fossil record of these Paleozoic rocks is dominated by a rich invertebrate heritage. The vertebrate record, however, is typically portrayed as unusually sparse. The Paleozoic vertebrate record at the Grand Canyon is best known for the varied trace fossils of tetrapods from the Pennsylvanian–early Permian Supai Group and the Permian Coconino Sandstone, where multiple different ichnotaxa have been described (e. g., McKee 1982; Spamer 1984; Elliott and Blakey 2005; Hunt et al. 2005). However, no tetrapod body fossils have been identified from these beds. Much of the research and literature on the body fossil record of vertebrates from the Grand Canyon has been focused on Pleistocene vertebrates from cave sites along the length of the canyon (e. g., Lucas and Morgan 2005; Mead 2005). Nevertheless, Paleozoic vertebrate body fossils have been reported from the Grand Canyon. Walcott (1880) made the first report of vertebrate fossils from the Devonian Temple Butte Limestone during the initial surveys made by the United States Geological Survey (USGS). McKee (1938) reported and described the first Permian vertebrate body fossil, a chondrichthyan tooth plate, from the Kaibab Formation. In 1944, the chondrichthyan Megactenopetalus kaibabanus was described from a large tooth collected from the North Rim at Point Sublime, and this is the first designated holotype of a Paleozoic vertebrate fossil from the Grand Canyon (David 1944). Other reports of vertebrate body fossils are from the Mississippian Redwall Limestone (McKee and Gutschick 1969) and the Pennsylvanian–Permian Supai Group (McKee 1982). Reviews of these vertebrate body fossils from the Paleozoic of the Grand Canyon have been made in the past (Spamer 1984; Elliott and Blakey 2005; Hunt et al. 2005). Recently, a rich assemblage of chondrichthyans was described from the Upper Mississippian Surprise Canyon Formation from the western Grand Canyon (Hodnett and Elliott 2018). 237 The purpose of the report presented here is to both collate previous documentation of Paleozoic vertebrate body fossils from the Grand Canyon and present the current physical record of specimens that have been collected from Grand Canyon National Park (GRCA). We have identified the presence of Paleozoic body fossils from seven geologic horizons within the Grand Canyon, which includes records outside of GRCA boundaries. We have included these records as part of our review as a “total record” approach to understanding the taxonomic richness of these seven geologic horizons. This additional information should assist in fossil resource planning, management, and priority assessment for further Paleozoic vertebrate research at GRCA. Institutional Abbreviations—GRCA, Grand Canyon National Park Museum, Grand Canyon, Arizona; MNA, Museum of Northern Arizona, Flagstaff, Arizona; USNM, National Museum of Natural History Paleobiology Collections, Washington, D.C. Devonian Temple Butte Limestone The Temple Butte Limestone is the only Devonian formation exposed at GRCA, and it also extends into southeast Nevada. The Temple Butte Limestone consists of thin, discontinuous lenses no more than 30 m (100 ft) thick deposited in paleo-valleys cut into the underlying Muav Limestone within the eastern Grand Canyon (Beus 2003a). The central and western sections of the Temple Butte Limestone are more continuous (Beus 2003a). The Temple Butte Limestone consists of a westward thickening wedge of interbedded dolomite, sandy dolomite, sandstones, mudstones, and limestones, ranging from purple, reddish-purple, and dark to light gray (Beus 2003a). The eastern paleo-valleys of the Temple Butte strata consist of interbedded mudstones, sandstones, dolomite, and conglomerates (Beus 2003a). The upper and lower contacts of the Temple Butte Limestone represent major unconformities with the underlying middle Cambrian Muav Limestone and the overlying Lower Mississippian Redwall Limestone (Beus 2003a). The model for deposition of the Temple Butte Limestone is similar to that of the younger Surprise Canyon Formation (Beus 2003a). The Temple Butte Limestone is a contemporaneous equivalent to the Martin Formation found in central and southern Arizona, which is known for a diverse vertebrate assemblage (Elliott and Blakey 2005). Both the Temple Butte Limestone and the Martin Formation are considered to be early Late Devonian (Frasnian) in age (Elliott and Blakey 2005). History The first report of a vertebrate fossil from the Grand Canyon was by Charles Doolittle Walcott (1880), who reported: “The purple sandstones deposited in the hollows of the Silurian limestones are characterized by the presence of Placoganoid fishes of a Devonian type,” which he identified during a field excursion in 1879. Schuchert (1918) communicated with L. Noble on a series of scales and plates of fish collected from the Temple Butte Limestone in 1916. These fish fossils were identified by J. W. Gidley of the Smithsonian Institution as the plates of the antiarch placoderm fish Bothriolepis nitidens and the scales of the sarcopterygian Holoptychius (Schuchert 1918; Noble 1922). Noble (1922) made no report of Holoptychius scales in the summary he published from 238 Gidley’s analysis. Denison (1951) accepted the occurrence of Holoptychius and revised the Temple Butte placoderm to Bothriolepis coloradensis based on similar outer dermal plate ornamentation morphology. GRCA Resources At present there are only two specimens identified in the National Museum of Natural History Paleobiology collections that come from the Temple Butte Limestone of the Grand Canyon (Figure 6-1). USNM PAL 328639 consists of part and counterpart of the distal part of a pectoral appendage (Figures 6-1A and 1B) and fragments and impressions of indeterminate plates of Bothriolepis cf. B. coloradensis that were collected by C. D. Walcott on January 17, 1883 as part of the United States Geological Survey study. This specimen is here considered to be the first discovered and oldest fossil vertebrate to be catalogued from the Grand Canyon. In addition, USNM PAL 328643 (Figures 6-1C– E) is a collection of approximately 30+ elements consisting of pectoral appendage, dorsal and ventral shield, and cephalic plates of Bothriolepis coloradensis collected by L. F. Noble in April 1916 and later mentioned by Denison (1951). At present, no specimens of Holoptychius from the Temple Butte Limestone have been identified in a museum collection. Potential Notes by Noble (1922) suggest that vertebrate remains from the Temple Butte are common but fragmentary approximately 1 m (3 ft) above the base of the Temple Butte Limestone and its contact with the Muav Limestone, in an impure dolomitic limestone. Noble (1922) speculated that other deposits within the Temple Butte Limestone could similarly be rich in vertebrate material. 239 Figure 6-1. Antiarch placoderm, Bothriolepis, fossils from the Temple Butte Limestone, Grand Canyon (JOHN-PAUL HODNETT). A–B. USNM PAL 328639, Bothriolepis cf. B. coloradensis, the proximal part of a pectoral appendage collected by C. D. Walcott. C–E. USNM PAL 328643, Bothriolepis coloradensis, a selection of plates collected by L. F. Noble. Scale equals 1 cm (0.4 in). 240 Mississippian Redwall Limestone The Redwall Limestone is a cliff-forming unit of early to middle Mississippian age sedimentary rocks (Beus 2003b). It consists primarily of light-olive-gray to light gray, thin- to thick bedded cherty limestones, interbedded crystalline dolomites, and limestone layers with chert lenses (Beus 2003b). The Redwall Limestone is divided into four lithologic units starting with the Whitmore Wash Member at the base, and ascending up to the Thunder Springs Member, Mooney Falls Member, and Horseshoe Mesa Member, respectively (McKee and Gutschick 1969). The Redwall Limestone was formed through two marine transgression and regression events from a seaway to the west (McKee and Gutschick 1969). The Whitmore Wash Member represents the first transgression event, and the Thunder Springs Member represents the first regression event, followed by the second transgression event of the Mooney Falls Member, and the final regression of the Horseshoe Mesa Member (McKee and Gutschick 1969). The Redwall Limestone has an upper unconformable contact with the Surprise Canyon Formation, which represents a time of uplift and karstic paleo-valleys cutting into the Redwall Limestone (Beus 2003). The Redwall Limestone has a rich invertebrate record. Regional variation of fossil faunas are noted within it, particularly of the corals and foraminifers (McKee and Gutschick 1969). History McKee and Gutschick (1969) reported a number of durophagous holocephalan chondrichthyan tooth plates and a spine from multiple localities within the Redwall
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