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Field and laboratory studies on the Coconino Sandstone (Permian) vertebrate footprints and their paleoecological implications
Abstract and Figures
The fossil footprints in the Coconino Sandstone have long been considered to be evidence for eolian deposition. I conducted field study of the fossil footprints in Hermit Basin, the Grand Canyon, and laboratory studies of modern amphibian and reptile footprints on dry sand, damp sand, wet sand, and underwater sand. Five species of salamanders all spent the majority of their locomotion time walking on the bottom, under water, rather than swimming. The experimental animals produced footprints under all test conditions, both up and down the 25° slopes of the laboratory “dunes”. Toe marks and other details were present in over 80% of the fossil tracks, underwater tracks, and wet sand tracks, but less than 12% of the dry sand and damp sand tracks had any toe marks. Dry sand uphill tracks were usually just depressions, with no details. Wet sand tracks were quite different from the fossil tracks in certain features. The fossil tracks were most similar to the underwater tracks. These data suggest that the Coconino Sandstone fossil tracks should not be used as evidence for eolian deposition of dry sand.
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... Interpretations of the Permian Coconino Sandstone began primarily with the work of McKee (1934McKee ( , 1945 and Reiche (1938), followed later by subsequent work (Fisher, 1961;Sorauf, 1962;Lundy, 1973;Elcock, 1993;Sumner, 1999;Maithel et al., 2019Maithel et al., , 2021. Recent studies focused on (1) stratigraphic relationships (Blakey and Middleton, 1983;Blakey, 1990); (2) paleoenvironmental implications of tetrapod trackways (Brand, 1979;Brand and Tang, 1991;Lockley et al., 1992 [see 'Reply,' p. 668-670]; Lockley and Hunt, 1995 [see p. 40-56]; Brand and Kramer, 1996;Millhouse, 2009;Citton et al., 2012); (3) sedimentary properties (Maithel et al., 2019(Maithel et al., , 2021; or (4) soft-sediment deformation (Brand and Maithel, 2021). Most authors consider the Permian Coconino Sandstone to be an eolian dune deposit (McKee, 1934;Reiche, 1938;Baars, 1961;McKee, 1974;McKee and Bigarella, 1979;Loope, 1984;Blakey, 1988;Middleton et al., 2003). ...
Polygonal “cracks” are common in the Coconino Sandstone in Arizona. They have been called desiccation cracks, but several features indicate they are not desiccation cracks. They were never open cracks, but are merely linear depressions, linked to form polygons. They occur only on bounding surfaces, containing almost no clay, and the cracks extend 10 to 15 cm above and below the bounding surfaces. The polygonal patterns continue down from one sandstone lamina to another, for several centimeters. They are persistently continuous across all surfaces within their 20–30 cm vertical range, from the bottomset beds, onto the bounding surface, and continuing into individual cross-beds below the bounding surface. The cracks occur at the Grand Canyon, and are especially numerous and visible in flagstone quarries in the Seligman and Ash Fork area. They occur on some bounding surfaces but not on others, and in some quarries but not in others. The polygonal cracks have been mentioned in passing, but this is the first reported research on these cracks in the Coconino Sandstone. Polygonal cracks have been reported in the Navajo, Page, and Entrada Sandstones, but there are significant differences between these and the Coconino Sandstone cracks, which may indicate differences in their origin.
... McKee (1934) was the first to propose and develop an eolian depositional model for the Coconino SS. Subsequent studies followed this work (e.g., Reiche 1938;McKee 1945;Fisher 1961;Sorauf 1962;Lundy 1973;Elcock 1993;Sumner 1999;Maithel et al., 2021), focused on stratigraphic relationships Blakey 1990), tetrapod trackways (Brand 1979(Brand , 1992Brand and Tang 1991;Lockley and Hunt 1995;Brand and Kramer 1996;Millhouse 2009;Citton et al., 2012), or sedimentary properties (Maithel et al., 2019(Maithel et al., , 2021. Similarly to McKee (1934), most authors interpret the formation as an eolian dune deposit (Reiche 1938;Baars 1961;Mckee 1974;Blakey and Middleton 1983;Loope 1984;Blakey 1988;Middleton et al., 2003). ...
The Permian Coconino Sandstone of northern Arizona contains numerous small-scale, soft-sediment deformation structures (SSDSs). These novel structures may be indicators of paleoenvironment or sedimentary processes. These SSD are generally shallow and occur on the surfaces of cross-beds, in contrast to convoluted bedding up to tens of meters thick commonly observed in some other eolian sandstones. These differences in structures imply differences in the processes that formed the Coconino Sandstone, or differences in the underlying depositional conditions. These SSDSs occur in outcrops at the Grand Canyon, and farther south in quarries near the towns of Seligman and Ash Fork. Size, orientation, structure, sedimentary context, clay content, and porosity of the structures are described. The SSDSs occur as small folds and ridges on the paleo lee side of otherwise undisturbed cross-beds. Some are associated with small rotated sandstone blocks within the cross-beds. The structures are exposed on bedding plane surfaces and in cross-section on vertical quarry walls. A few SSDSs up to a meter thick also occur in the Coconino Sandstone, but the others are only up to a few cm thick, 2–10 cm wide, and 20 cm to 10 m long. Evidence is presented that liquidization (as fluidization or liquefaction) may have been involved in producing these features, implying a high water content in scattered locations at time of deformation, but this process also requires some stressor to trigger the deformation. Seismic events may provide a triggering mechanism. The Coconino Sandstone SSDSs represent unusual or previously overlooked small-scale features related to individual foreset surfaces.
... In typical eolian facies of Botucatu Formation, tracks were interpreted as "cut undertracks", which are formed with the autopodia crossing through the superficial layer of dry sand and reaching damper layers below. Several deposits of ancient dunes were previously interpreted as subaqueous or at least moist paleoenvironments based on preservation of fossil tracks in it, as the Jurassic Navajo Sandstone and the Permian Coconino Sandstone (e.g., Brand, 1979;Brand and Tang, 1991;Mckeever, 1991;Rainforth, 2001), both in the USA. Abruptly starting and stopping trackways were interpreted by Brand and Tang (1991) as the result of the animal alternately walking on and swimming over the substrate, but it also could be explained if the animal moved between distinct grain flow lobes (Loope, 2006). ...
This work aimed to investigate the ichnofossiliferous sites in Nioaque and surrounding region, Mato Grosso do Sul state, Brazil, analyzing their paleontological, stratigraphic and paleoenvironmental context, because dinosaur tracks were found in an area previously mapped as Permo–Carboniferous age. Some sedimentary facies previously assigned to the glacial Aquidauana Formation were herein reinterpreted as part of the basal section of the Botucatu Formation (Late Jurassic/Early Cretaceous) and include floodplain and residual channel deposits, possibly representing the lower half of this formation's sequence in the studied area. The ichnofossils that have been found include both isolated tracks of Theropoda and Ornithopoda, a trackway of Eubrontes isp., a vertebrate burrow and invertebrate traces (Palaeophycus) possibly related to the Entradichnus ichnofacies. Sedimentological interpretation suggests the existence of a river system just before the deposition of the typical Botucatu eolian facies in this region. This is the first documented record of fluvial deposits transitionally to the traditional eolian Botucatu Formation. Noteworthy, the dinosaur tracks have played a key role for the interpretation of these rocks as Botucatu Formation and for better understanding the whole system due to the impossible Paleozoic age (dinosaur tracks).
... The formation and preservation of tetrapod tracks and trackways are closely related to the trackmaker (identity, size and gait), sedimentary substrate (grain size, moisture content and dip), sedimentary environments and fluid dynamics (water and wind) (Brand, 1979;McKeever, 1991;Gatesy et al., 1999;Manning, 2004;Jackson et al., 2009Jackson et al., , 2010Mancuso et al., 2016;Marchetti et al., 2019). In general, track preservation requires: (1) impression into a slightly moist substrate; (2) formation of surface cohesion and scour resistance by the means of subaerial exposure or cementation; (3) rapid burial ensured by high sedimentary rate, without enough hydrodynamic force to rework the track-bearing sediment. ...
... The formation and preservation of tetrapod tracks and trackways are closely related to the trackmaker (identity, size and gait), sedimentary substrate (grain size, moisture content and dip), sedimentary environments and fluid dynamics (water and wind) (Brand, 1979;McKeever, 1991;Gatesy et al., 1999;Manning, 2004;Jackson et al., 2009Jackson et al., , 2010Mancuso et al., 2016;Marchetti et al., 2019). In general, track preservation requires: (1) impression into a slightly moist substrate; (2) formation of surface cohesion and scour resistance by the means of subaerial exposure or cementation; (3) rapid burial ensured by high sedimentary rate, without enough hydrodynamic force to rework the track-bearing sediment. ...
Ancient desert, characterized by low biotic diversity as well as low preservation potential, has long been assumed as devoid of biotic activity. However, recent works from Mesozoic trace fossils preserved in the eolianites in North America, South America and East Asia indicate that the ancient desert can also have diverse inhabitants. This paper presents a diversified preserved dinosaur track assemblage from the Jurassic-Cretaceous transition eolian dune deposits in western Shandong Province, China. Based on the ichnological analysis, tracks are assigned to ichnogenus Anomoepus, Eubrontes-like and Ornithopodichnus-like forms, representing the oldest and second example of the Chelichnus ichnofacies reported from China as well as Asia. Sedimentological analysis of the tracksites indicates that tracks occurred in linear dunes under arid climatic conditions. Well-preserved tracks with detailed anatomical information are more likely to be preserved in moist sands, contrary to the dry cohesionless sands. Anomoepus and Ornithopodichnus-like tracks cooccurring with northwestward winds were made during summer wet season, while Eubrontes-like track was present during winter dry season characterized by strong northeastward wind, implying different habits between ornithopod and theropod dinosaurs. The preserved Jurassic-Cretaceous transition dinosaur tracks in eolian dune deposits not only expand the spatial-temporal distribution of dinosaur fauna in North China, but also greatly improve our understanding of the adaptive capacity of dinosaurs in severe living conditions, such as the desert environment.
... Although aquatic animals that engage in underwater pedestrianism may do so while completely submerged (cf. Zug 1971;Brand 1979;Martinez 1996;Martinez, Full, and Koehl 1998;Coughlin and Fish 2009;Grigg and Kirshner 2015;Farlow, Robinson, Turner, et al. 2018a, b), the likely pneumaticity of their bodies makes it unlikely that sauropods could have done the same (Henderson 2004). This in turn sets an upper limit to how deep the water through which hypothetical punting sauropods moved could have been: probably no more than a few to several metres (Figs. ...
Three parallel, manus-only sauropod trackways from the Coffee Hollow A-Male tracksite (Glen Rose Formation, Kendall County, Texas) were studied separately by researchers from the Heritage Museum of the Texas Hill Country and the Houston Museum of Natural Sciences. Footprint and trackway measurements generally show good agreement between the two groups’ data sets. Footprints appear to be shallowly impressed true tracks rather than undertracks. One of the Coffee Hollow trackways shows marked asymmetry in the lengths of paces that begin with the left as opposed to the right forefoot, and two of the Coffee Hollow trackways are unusually broad. The Coffee Hollow trackways differ enough from the manus portions of other Glen Rose Formation sauropod trackways to suggest that they were made by a different kind of sauropod. Greater differential pressure exerted on the substrate by the forefeet than the hindfeet probably explains the Coffee Hollow trackways, like other manus-only sauropod trackways, but the possibility that they indicate unusual locomotion cannot at present be ruled out.
In their review article in this book, Marchetti et al. (2024) note that
Permian tetrapod footprints are abundant, relatively diverse, and widespread
in Pangea. They consider 22 ichnogenera to be valid and attributed
them to temnospondyls and lepospondyls (Batrachichnus, Limnopus, Matthewichnus),
seymouriamorphs (Amphisauropus), diadectomorphs (Ichniotherium),
nontherapsid synapsids (Dimetropus, Tambachichnium), nonmammalian
therapsids (Brontopus, Capitosauroides, Dicynodontipus, Dolomitipes, Karoopes),
captorhinomorphs (Hyloidichnus, Notalacerta), diapsids (Dromopus, Paradoxichnium,
Protochirotherium, Rhynchosauroides) and parareptiles (Erpetopus,
Pachypes, Procolophonichnium, Varanopus) (Fig. 5).
Marchetti et al. note that Permian footprints are more diverse than
Carboniferous footprints, likely due to the ecological and morphological
diversification of tetrapods. The Permian tracks indicate a generally raised,
sprawling mode of locomotion, although some Permian ichnogenera are
consistent with a semi-erect posture that clearly evolved before the Triassic.
Permian tracks include examples of locomotion on inclined surfaces and
penetrative traces, which markedly altered their morphological preservation.
Permian footprints are sometimes associated with resting traces, scaly skin
impressions, and swimming traces, and they have been found in various
depositional environments, such as alluvial plains, lake margins, tidal flats,
and deserts. The composition of the ichnoassemblages changes between
different settings, especially the relative abundance of ichnotaxa, consistent
with the ichnofacies concept. Nevertheless, a thorough review of Permian
tetrapod ichnofacies is needed.
During the early Permian (Cisuralian), footprints are very wellrepresented
at low paleolatitudes in western Europe, northern Africa, and
North America. And, during the middle-late Permian (Guadalupian-
Lopingian) they are well known as at mid-high paleolatitudes latitudes,including South Africa and Russia. They are generally more diverse at low
paleolatitudes. Tetrapod footprints are valuable for Permian chronostratigraphy;
five tetrapod footprint biochrons are currently defined (ascending): Dromopus,
Erpetopus, Brontopus, Dicynodontipus, and Paradoxichnium. Permian
footprints are also key to the understanding of major faunal events such as
the Artinskian Warming Event and the end-Guadalupian and end-Permian
extinctions.
This bibliography is comprehensive for the entire volume. In addition to references that discuss particular systematic taxa , it also includes citations of references that in some way discuss paleontology in the Grsad Canyon region without naming any systematic taxa. Citations provide complete bibliographical data in Mfioort of abbreviated citations in the indexes that follow in Part 2. References that are flagged by a triangle () indicate publications that wholly or in part discuss fossils derived from Tertiary that are important in the interpretation of the development of the Grand Canyon; these occurrences are not neceeaarily at or near the Grand Canyon.
These are lists of genera and species reported from publications th. cover areas greater than that covered by' these indaexcluded from these lists (thus thes- lists do not necessarily cite every). The taxa sre listed alphabeticaity, first listing Animalia,then AcriIIrdaa, dofouill. Taxa that are cited only as genera (e.g., Pinus sp.) are iDduded 0lIl,IaaYe included in them (in the cited publication). The systematic placemelll of tau foIlowidaelCbeme explainedin the "Scopeand Organization" section of this volume. In I few inItaDceI, it to update the identification under which the taxon was originally described; piItiaaArIy in the taIeI of Proterozoic fossils that were originally described under metazoan groups that fint evolved durin, the Paleozoic.
Mainstream scientists, including Woodward, Buckland, Prestwich, Suess, and Ryan and Pitman, have proposed a variety of theories to explain the biblical deluge. The extent of the flood in these theories has decreased as empirical knowledge of global geology has increased. In contrast, contemporary flood geology attempts to explain most of the geologic record in terms of a single, year-long, global catastrophe. Flood geology exists in the context of an alternate scientific universe with its own institutions, organizations, journals, and meetings. The views of Leonard Brand, Steven Austin, and Walt Brown, representative of flood geology, are discussed.
Lower Permian strata of the Boskovice Basin in the Czech Republic have yielded hundreds of largely complete seymouriamorph individuals, some of which are exceptionally preserved and have noticeable soft tissues, such as external gills and eye structures. The vast majority of these finds are referable to Discosauriscus austriacus, and almost exclusively represent larvae and juveniles tied to aquatic environments. Only one of the specimens discovered to date has been interpreted as an early adult, although late juvenile and early sub-adult stages have been proposed for it as well. Regardless, no other seymouriamorph adults have been reported from the Boskovice Basin so far. Here we present the first comprehensive assessment of seymouriamorph tracks and a trackway from the Asselian (lowermost Permian) of the Boskovice Basin. The morphology of the tracks is congruent with that of the ichnotaxon Amphisauropus and indicates an attribution to Discosauriscidae or that they belong to as yet unrecognized large seymouriamorphs. All tracks were made by individuals much larger than the largest discosauriscid known from bodily preserved material and are interpreted as those of adults tied to terrestrial environments, thus apparently documenting a habitat shift that occurred relatively late in the ontogenetic development of these seymouriamorphs. Furthermore, the largest track is preserved in extraordinary detail and includes soft part impressions such as digital, palmar and plantar flexion creases, making the specimen one of the best preserved seymouriamorph tracks described to date. Finally, one of the tracks originates from the lowermost Asselian and is therefore among the oldest known records of seymouriamorphs worldwide.