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Bird and mammal tracks from the White Narrows Formation (early Pliocene) in the Moapa area, near
Glendale, Nevada, have been known and collected for several decades, furnishing at least four museums with
representative specimens. This easily accessible site contains the best Nevadan examples of early Pliocene ungulate and carnivore tracks. One carn...
Citations
... Additional evidence for the tuff of Napa is closer, approximately 47 km north of the NDRA. Here, west of Moapa, Nevada the White Narrows marl is interpreted as lacustrine and is similar lithologically to white limestone "a" (Gardner 1968;Schmidt et al. 1996;Lockley et al. 2007). The White Narrows marl contains mammalian fauna in a fossiliferous zone near the base of the unit that indicates ages of 4.9-5.0 ...
... The White Narrows marl contains mammalian fauna in a fossiliferous zone near the base of the unit that indicates ages of 4.9-5.0 Ma (Lockley et al. 2007). Schmidt et al. (1996) noted an ash bed above the fossiliferous zone. ...
Lacustrine basins and the sediments within them provide a critical component of regional tectonic development and climate history. Each sub-basin in the landscape records the interplay between active tectonism and shifting climate. Many of the basins in southern Nevada that contain deposits of the Muddy Creek Formation were closed fluvio-lacustrine systems during the latest Miocene and into the Pliocene. They were subsequently integrated into the regional Colorado River system. Lacustrine deposits of the Muddy Creek Formation are exposed along the trace of the Las Vegas Valley shear zone in the Nellis Dunes Recreation Area (NDRA), northeast of Las Vegas, NV. The Muddy Creek Formation in NDRA consists of approximately 130 m of mixed carbonates and clastics, and new tephrochronology suggests an early to late Pliocene age (about 4.7 to about 2.6 Ma) for the lowermost lacustrine deposits and an unknown, but younger age for the uppermost spring-fed lacustrine deposit. These sediments were deposited in an arid to semi-arid lake (Lake Nellis) and alluvial floodplain system based on the interpretation of lithologies and facies associations. The stratigraphic succession coarsens upward and reflects establishment of a lacustrine carbonate system overlain by a clastic succession of peripheral lake sediments of alluvial and floodplain origin. The peripheral lake deposits consist of spring-fed and fluvial wetland mudflats of brown claystone and siltstone. This brown claystone is capped by a succession of yellow and red sandstones deposited by fluvial and minor eolian processes. The second, and youngest freshwater limestone, likely disconformably, overlies the yellow and r ed sandstones, and is thought to be temporally and depositionally distinct from the underlying Muddy Creek deposits. The transition from the lower lacustrine carbonates to clastic fluvial channel and flood plain deposits indicates desiccation of Lake Nellis and possibly occurs soon after about 2.7 to about 2.6 Ma based on the ages of tuffs occurring within the uppermost limestone and marl beds. This is consistent with an interpreted increase in regional aridity after approximately 2.8 Ma (Smith et al. 1993). The second limestone deposit at the top of the section represents an even younger spring/lacustrine deposit of unknown age. Throughout the lower carbonate section, three tuffs were identified, and geochemically correlated, using a discriminant function analysis, to the tuff of Napa (≤ 4.70 ± 0.03 Ma), the Putah Tuff (about 3.3 Ma) and the lower tuffs of the Badlands ( about 2.7 – about 2.6 Ma). The timing of deposition of Lake Nellis sediments post-dates the integration of the three major lake syste ms in this region (Lake Grand Wash, Lake Hualapai, and Lake Las Vegas) into the Colorado River drainage (5.6 – 4.9 Ma), and also postdates the full integration of the river to sea level (4.8 - 4.63 Ma) (Howard et al. 2015; Crow et al. 2021). Lake Nellis represents an isolated lake basin that was one, and maybe the last, lacustrine system to be fully integrated into the Colorado River drainage in this region.
... The fragmentary taxon, Gargantuavis is known from the Late Cretaceous (Buffetaut & Le Loeuff 1998;Mayr et al. 2019) of Europe but lacks known foot remains. Despite a useful body fossil record of large flightless birds, including fossil representatives of the Phorusrhacids (Alvarenga & Höfling 2003) and the extant ratites (Yonezawa et al. 2017), the latter the most dominant palaeognathid lineage today (Yonezawa et al. 2017), with the exception of dinornithiform tracks from Tasmania (Rich & Green 1974) and New Zealand's moa represented by a dozen species and several track morphotypes (Lockley et al. 2007b, and references therein), none of these extinct ground dwelling birds left a track record. Incidentally, Andors (1991) compared Diatryma with the New Zealand Takahe (genus Notornis), inferring it to be a slow-moving pedestrian herbivore. ...
... Pachydactylous moa tracks show a polarity between small narrow, large wide morphotypes (Lockley et al. 2007b;Lockley 2009) reminiscent of the allometric spectrum seen in non-avian theropod tracks in the aforementioned Grallator-Eubrontes plexus (Olsen 1980). Could this imply greater diversity among other large flightless birds from earlier epochs such as the Eocene, as the difference between Ornithoformipes and Rivavipes seems to imply? ...
A diverse continental ichnofauna from the Lower Cretaceous (Aptian) Gething Formation occurs at the Ninesting Creek site in northeastern British Columbia. Vertebrate tracks including Wupus and Saurexallopus are inferred to represent medium-sized (footprint length ∼10 cm) tridactyl avian and larger (footprint length ∼15–17 cm), gracile, tetradactyl non-avian theropod trackmakers respectively. Shape, length-to-width and divarication ratios, suggest two morphotypes that are distinct from the poorly known, and infrequently-cited, Gething Formation ichnogenera Columbosauripus and Irenichnites (footprint lengths ∼12.5 and ∼15.0 cm respectively). This suggests a diverse theropod ichnofauna representing size classes which are quite distinct from the better-known large theropod ichnogenus Irenesauripus with footprint lengths in the range of ∼38.0–53.5 cm. Thus, current evidence suggests that the ichnodiversity of smaller avian and non-avian theropod tracks (footprint length ∼<15.0 cm) outnumbers large ‘megatheropod’ morphotypes by about 7:1. The implications of this distribution are consistent with recently proposed palaeoecological models. The assemblage also includes probable pterosaur tracks and distinctive arcuate compound Archaeonassa-Lockeia traces, attributed to freshwater unionid bivalves. The trace fossils comprise a composite Mermia-Scoyenia Ichnofacies assemblage. Palaeoenvironmentally the site represents the low energy margin of an open lake basin or floodplain pond, or a fluvial point bar.
... In this study, based on the morphological characters of this footprint type, the authors suggest that these footprints belong to the Platykopus ichnogenus, because all diagnostic characteristics of this ichnogenus can be recognised on them (see above). All Platykopus footprints are considered to represent those of a large bear-like animal (Sarjeant et al. 2002;Lockley et al. 2007;Abbassi 2010;Diedrich 2011;Aramayo et al. 2015). During the early Miocene, there was no large-sized Ursidae present in Europe (see McLellan and Reiner 1994;Jiangzuo and Flynn 2020). ...
Ipolytarnóc is one of the most important Cenozoic fossil trackway sites in Europe. Most of the discovered footprints were investigated in 1985; however, a considerable period has elapsed since those investigations, and during that time significant advances have been made in the field of 3D imaging. Given this fact, the present study was undertaken to carry out a new analysis of the Ipolytarnóc fossil tracks, with a view to present possible revisions of current knowledge. In line with this, detailed ichnotaxonomical analyses were conducted on two large-sized pentadactyl footprint types using high-quality 3D models. As a result of the investigations presented in this paper, the largest pentadactyl footprint-type (previously defined as Bestiopeda maxima) was reclassified under the Platykopus ichnogenus based on new materials and their 3D models. The P. maxima footprints are believed to represent those of large-sized Amphicyonidae. Thorough ichnotaxonomical analyses were performed on other pentadactyl fossil tracks which had been attributed to Carnivoripeda nogradensis. The aim of the analyses was to suggest an extension of the morphological characters of these ichnospecies. In contrast to the previously suggested Nimravidae origin, we rather suggest that the C. nogradensis footprints belong to a mustelid-like carnivore based on its footprint morphology.
... The fragmentary taxon, Gargantuavis is known from the Late Cretaceous (Buffetaut & Le Loeuff 1998;Mayr et al. 2019) of Europe but lacks known foot remains. Despite a useful body fossil record of large flightless birds, including fossil representatives of the Phorusrhacids (Alvarenga & Höfling 2003) and the extant ratites (Yonezawa et al. 2017), the latter the most dominant palaeognathid lineage today (Yonezawa et al. 2017), with the exception of dinornithiform tracks from Tasmania (Rich & Green 1974) and New Zealand's moa represented by a dozen species and several track morphotypes (Lockley et al. 2007b, and references therein), none of these extinct ground dwelling birds left a track record. Incidentally, Andors (1991) compared Diatryma with the New Zealand Takahe (genus Notornis), inferring it to be a slow-moving pedestrian herbivore. ...
... Pachydactylous moa tracks show a polarity between small narrow, large wide morphotypes (Lockley et al. 2007b;Lockley 2009) reminiscent of the allometric spectrum seen in non-avian theropod tracks in the aforementioned Grallator-Eubrontes plexus (Olsen 1980). Could this imply greater diversity among other large flightless birds from earlier epochs such as the Eocene, as the difference between Ornithoformipes and Rivavipes seems to imply? ...
The Mesozoic and Cenozoic track record of large birds (avian theropods) with footprint lengths (FL) > 10.0 cm is quite limited, whereas small tracks (FL < 10.0 cm) are abundant from the Early Cretaceous onwards. This large versus small threshold value is consistent with the track record of extant birds among which only ˜10% are large, and so is scaled appropriately to Class Aves. The proportion of large pachydactylous (thick-toed) or robust non-avian theropod ichnotaxa reported from the Mesozoic considerably exceeds the few large leptodactylous (thin-toed) or gracile forms such as Archaeornithipus and Magnoavipes, named, albeit controversially, to suggest avian affinity. The Cenozoic record of large avian tracks is quite different, with large pachydactylous tracks limited to a few Eocene-Oligocene (Palaeogene) ichnogenera (Ornithoformipes and Rivavipes), and rare unnamed Neogene and Holocene dinornithiform tracks. Thus, large flightless species are less well represented by tracks than body fossils. This suggests probable preservational bias in favour of waterbird or shorebird-like tracks in shoreline facies. However, the Cenozoic record of large leptodactylous tracks attributable to cranes, herons and related trackmakers is quite extensive, mostly confined to the Neogene and formally named to suggest representatives of extant avian clades capable of flight. These distribution patterns apparently reflect the dominance of large pachydactylous non-avian and avian theropods in the Mesozoic and early Palaeogene, and contrast with the rise of large, as well ongoing diversification of smaller leptodactylous Aequornithes in the Neogene. These distribution patterns in space and time likely reflect changing selection pressures as today's modern avifauna evolved.
... Diagnosis of the ichnogenus Ardeipeda was emended as avian footprints that show four digits, with three large digits (II to IV) that are directed forward, while the fourth (digit I) is directed backward and is somewhat smaller with webbing absent. Lockley et al. (2007) reported Ardeipeda as partly-webbed bird footprints, but ichnospecies of Ardeipeda (A. egretta, A. gigantea, and A. incerta), have no webbing between the digits (Panin and Avram 1962). A. filiportatis, however differs from A. egretta and A. gigantea by short webbing between digits II and III. ...
The Mukdadiya Formation (Late Miocene-Pliocene) consists of alternations of red to brown sandstone and claystone layers in the Zagros Mountains Belt, northeastern Iraq. Two track-sites preserving bird and mammal tracks were recorded from the base of the formation in the Chamchamal area of the Kurdistan region in northeast Iraq. Avian tracks are large footprints with an average length of 25 cm that belong to Avipeda filiportatis. Mammal footprints imprinted by terrestrial cetartiodactyls belong to Pecoripeda amalphaea and Bifidipes velox, with cervids the most likely track makers. An unfamiliar large trace fossil on the studied slab, has symmetrical to asymmetrical marks, kidney or number-8-shaped and strong wrinkles on the surface. We interpret trace as having been produced by cetartiodac-tyls wallowing on the soft sediment surface. To accommodate this class of behavioral trace fossils; thus, we introduce a new fossilized behavior class, named "Laspichnia", which includes a vertebrate mud-bathing imprint on a soft sediment surface.
... Nevertheless, avian ichnotaxonomy is still in flux (Lockley et al., 2007), therefore open nomenclature is used. ...
Barrier-islands are common landforms and biodiverse habitats, yet they received scarce neoichnological attention. This gap is tackled by studying the Mula di Muggia barrier-island system (Grado lagoon, Italy), focusing on morphology, ecology and ethology of individual traces. The following incipient ichnotaxa are identified: Archaeonassa, Arenicolites, Bergaueria, ‘diverging shafts’, Helminthoidichnites, Lockeia, Macanopsis, Monocraterion, Nereites, Parmaichnus, Polykladichnus, Skolithos, Thalassinoides and ‘squat burrows’. Vertebrate (Avipeda-/Ardeipeda-like, Canipeda) and invertebrate tracks (‘parallel furrows’) are also described.For each ichnotaxon, tracemaker and behavior are discussed, together with their position with respect to sediment barriers. Results suggest that sediment barriers impose a sharp contrast in terms of ichnological composition. Back-barrier is dominated by branched burrows (i.e. Thalassinoides, Parmaichnus), while the fore-barrier presents vertical and U-shaped burrows (Arenicolites, Skolithos). The environmental conditions of the back-barrier show that low-oxygen substrates favor intense bioturbation, provided that the water column is sufficiently oxygenated.
... Frenchman Mountains are tilted 50-60° east; the stratigraphy in the mountains is very similar to that in the Grand Canyon, ranging in age from Precambrian to Permian (Longwell et al. 1965 (Lockley et al. 2007). This formation fills graben structures within the Muddy Creek Formation that resulted from faulting along the southern edge of the Arrow Canyon Range, which you can see to the north (Lockley et al. 2007). ...
... Frenchman Mountains are tilted 50-60° east; the stratigraphy in the mountains is very similar to that in the Grand Canyon, ranging in age from Precambrian to Permian (Longwell et al. 1965 (Lockley et al. 2007). This formation fills graben structures within the Muddy Creek Formation that resulted from faulting along the southern edge of the Arrow Canyon Range, which you can see to the north (Lockley et al. 2007). Based on fossil rodents, the White Narrows Formation is earliest Blancan (early Pliocene, ~4.7 Ma) (Schmidt et al. 1995;Woodburne and Swisher 1995;Reynolds and Lindsay 1999). ...
... Based on fossil rodents, the White Narrows Formation is earliest Blancan (early Pliocene, ~4.7 Ma) (Schmidt et al. 1995;Woodburne and Swisher 1995;Reynolds and Lindsay 1999). Important vertebrate tracksites, containing tracks of ungulates, proboscideans, carnivorans, and birds have been reported from the White Narrows Formation in the area (Lockley et al. 2007 (Schmidt et al. 1995). The Coconino-Toroweap contact can be seen at milepost 25.5 (Reynolds et al. 2006 (Reynolds et al. 2006). ...
... Both A. phoenix and A. sirin are small tracks (lengths of 1.6 and 2.5 cm, respectively); the latter is most comparable to Koreanaornis in size. A. filiportalis is huge by comparison (footprint length up to 19 cm) and is morphologically distinct because it possesses a caudally-oriented hallux trace -in this and other respects, it is similar to the earlier named, heron-like footprint Ardeipeda egretta (the type ichnospecies) and Ardeipeda gigantea (Panin and Avram, 1962;Lockley et al., 2007c). Thus, A. filiportalis is justifiably considered a synonym of Ardeipeda. ...
Fossil footprints provide important evidence regarding the morphology, behavior, distribution, and ecology of ancient animals. In recent years, the ichnological record (pertaining to fossils other than skeletal or body parts, most familiarly and commonly tracks) of major tetrapod clades has been studied intensively. The body fossil record amply demonstrates that the origin of birds lies within the theropod dinosaur lineage (making birds extant dinosaurs, in an evolutionary sense), but the ichnological record contributes much valuable information concerning behavioral shifts during both this evolutionary transition and the early diversification of birds. Here, for the first time, we review the entire avian track record, including its specialized ichnotaxonomy, from the Mesozoic (the "Age of Reptiles," 250-65 million years ago) and Cenozoic (the "Age of Mammals and Birds," 65 million years ago through the present, including the Holocene) and consider how the evidence impacts the understanding of avian evolution and ecology. Growing evidence from both the skeletal and track records indicates that the initial avian taxonomic, morphological, and ecological radiations took place around the Jurassic-Cretaceous boundary (about 145 million years ago). Tracks similar to, and in some cases indistinguishable from, those made by modern shorebirds (Charadriiformes), small ducks (Anseriformes), small herons (Ciconiidae), and even roadrunners (Cuculiformes) appeared, and were even regionally abundant only 15-20 million years thereafter. In contrast, the oldest body fossil records of anseriforms and possibly charadriiforms occur very close to the end of the Cretaceous (roughly 70 million years ago), and later still for ciconiiforms and cuculiforms. This strongly implies that the early track makers were members of extinct, early avian lineages with which later taxa converged in foot morphology. Feeding traces associated with some of these tracks demonstrate that behaviors reminiscent of extant herons and spoonbills had also evolved by this time. However, despite significant skeletal and footprint finds, there is little correspondence between the records - few footprints match the feet of birds represented by skeletal remains. In short, the familiar morphologies and behaviors of many modern birds actually evolved convergently with many of their extinct, Mesozoic relatives. Footprints thus have the dual benefits of providing an important, and unexpected, complementary record of early avian morphological and ecological diversity while highlighting the importance of morphological and behavioral convergence. Although the skeletal record suggests an avian taxonomic shift at the "dinosaur-killing" Cretaceous-Paleogene (K-Pg) boundary extinction event, the track record provides insufficient evidence to support or refute such a shift because the dominance of shorebird-like tracks continues uninterrupted from Mesozoic to Cenozoic. Early Paleogene tracks provide evidence of large, Diatryma- or Gastornis-like, ground dwelling birds in addition to typical shorebirds and waterbirds like the Eocene anseriform Presbyornis. Neogene tracks include those of a few large ratites and a turkey-like species; Holocene tracks include those of several species of moa. Unlike its Mesozoic counterpart, the Cenozoic avian body fossil and ichnological records correspond much more closely. Tracks of perching birds, raptors, and other groups that do not habitually frequent wet shorelines - the most suitable environment for track preservation - are rare. Indeed, the avian track record is dominated by the footprints of shorebirds, with a minor component attributable to large flightless and cursorial forms. Nevertheless, the body of literature on fossil bird tracks is still relatively small (~200 reports), describing about 6 ichnofamilies, comprising about 38 named ichnogenera and 65 ichnospecies.
... As summarized by Lucas (2007) Oleg Stepanovich Vyalov published a landmark, but problematic paper (Vyalov, 1965) in which he ''proposed that all bird tracks be assigned to the single ichnogenus Avipeda'' (Lucas, 2007, p. 113). This is clearly both impractical and unjustified on morphological grounds: i.e., bird tracks show considerable morphological diversity which, in the case of modern species, correlate with diagnostic morphological variation at the genus and higher taxonomic levels (Lockley et al., 2007c). Although Lucas (2007, p. 114) credited Vyalov with being conceptually ahead of his time and also affirmed that Vyalov's ichnospecies were ''correctly introduced in accordance with the International Code of Zoological Nomenclature'' he noted that many of his illustrations are of poor or indifferent quality and lack scales. ...
... Vyalov's inclusion of many morphologically distinct and much larger tracks (up to 13 cm in length) in Avipeda, dilutes the concept of Avipeda to the point where it lacks any diagnostic morphological significance. For example, Vyalov included morphotypes in Avipeda which are obvious junior synonyms of ichnogenera such as Ardeipeda (a large heron track) named by Panin and Avram (1962) and discussed by Lockley et al. (2007c). Sarjeant and Langston (1994) considered Avipeda representative of a poorly-defined 'morphofamily' which they named Avipedidae. ...
Although body fossils of shorebirds and shorebird-like species are extremely rare from the Cretaceous, rapid increase in the discovery of bird footprint sites provides valuable alternate evidence to help fill gaps in the story of the early evolution of shorebird-like species. Newly discovered bird tracks from the Albian-Cenomanian Dakota Formation in northeastern Utah represent the first report of the ichnogenus Koreanaornis from North America and only the second report of bird tracks from this formation. These tracks are not attributable to Aquatilavipes as previously claimed. Three well-preserved trackways are described and provisionally referred to Koreanaornis cf. hamanensis (Kim). However, a review of the ichnotaxonomy of shorebird ichnites reveals that this ichnotaxon also closely resembles the Miocene ichnospecies Avipeda sirin (Vyalov). This latter comparison points to the need for a thorough evaluation of the similarity between Mesozoic and Cenozoic avian ichnotaxa, which may be over-split in some cases and under-differentiated in others.The new material helps distinguish ichnogenus Koreanaornis from the larger bird track Aquatilavipes, which is more abundant and widespread in North America. In some cases Aquatilavipes has been incorrectly used as a catch-all ichnogenus both in North America and Asia. The Dakota Formation stratigraphy at the tracksite indicates that the track makers lived in a marginal marine paleoenvironment. However, despite the widespread distribution of such facies, often replete with dinosaur tracks, the bird track record of the Dakota Formation, and the Cretaceous of the western USA remains relatively sparse in comparison with other areas such as east Asia.
... Frenchman Mountains are tilted 50-60° east; the stratigraphy in the mountains is very similar to that in the Grand Canyon, ranging in age from Precambrian to Permian (Longwell et al. 1965 (Lockley et al. 2007). This formation fills graben structures within the Muddy Creek Formation that resulted from faulting along the southern edge of the Arrow Canyon Range, which you can see to the north (Lockley et al. 2007). ...
... Frenchman Mountains are tilted 50-60° east; the stratigraphy in the mountains is very similar to that in the Grand Canyon, ranging in age from Precambrian to Permian (Longwell et al. 1965 (Lockley et al. 2007). This formation fills graben structures within the Muddy Creek Formation that resulted from faulting along the southern edge of the Arrow Canyon Range, which you can see to the north (Lockley et al. 2007). Based on fossil rodents, the White Narrows Formation is earliest Blancan (early Pliocene, ~4.7 Ma) (Schmidt et al. 1995;Woodburne and Swisher 1995;Reynolds and Lindsay 1999). ...
... Based on fossil rodents, the White Narrows Formation is earliest Blancan (early Pliocene, ~4.7 Ma) (Schmidt et al. 1995;Woodburne and Swisher 1995;Reynolds and Lindsay 1999). Important vertebrate tracksites, containing tracks of ungulates, proboscideans, carnivorans, and birds have been reported from the White Narrows Formation in the area (Lockley et al. 2007 (Schmidt et al. 1995). The Coconino-Toroweap contact can be seen at milepost 25.5 (Reynolds et al. 2006 (Reynolds et al. 2006). ...
