Second occurrence of the dinosauriform ichnogenus Atreipus in the western United States,Upper Triassic Chinle Group of Eastern Utah

Abstract

A newly discovered track in the Chinle Group north of Moab, Utah, is attributable to the ichnogenus Atreipus, an ichnotaxon that is relatively common in eastern North America (Newark Supergroup) but very rare in the Late Triassic of the western part of the continent. This is only the second report of the genus from the Chinle Group. Atreipus has been attributed to a silesaurid dinosauriform, and dinosauriform taxa are relatively abundant by skeletal material in the Late Triassic of western North America, but track evidence in the same units is dominated by ichnotaxa attributed to dinosaurs. The rarity of Atreipus is currently an anomaly in the region.     

Grallator-Dominated Fossil Footprint Assemblages and Associated Enigmatic Footprints from the Chinle Group (Upper Triassic), Gateway Area,Colorado

The upper part of the Chinle Group (Late Triassic) of the Gateway area in western Colorado is extraordinarily rich in fossil footprint assemblages. Dominant track types include small Grallator tracks, generally attributed to Coelophysis-like theropods, which often occur in high densities of 50 to 100 per m². Other abundant ichnotaxa that are attributable to dinosaurs or dinosaur-like archosaurs include Pseudotetrasauropus and Tetrasauropus, attributed to prosauropods and sauropods, respectively. Several Pseudotetrasauropus-like tracks appear to be functionally didactyl and may indicate a new ichnotaxon that represents an animal that shows certain unusual features that are convergent with dromeosaurs and certain birds. Such convergence may reflect inherent growth programs as much as functional adaptations. Non-dinosaurian ichnotaxa include Brachychirotherium (probably of aetosaur affinity) and Rhynchosauroides, attributed to a sphenodontid/lizard-like form. Other ichnotaxa include probable therapsid (dicynodont) tracks labeled Pentasauropus sp., mammaloid (non-therian mammal and/or mammal-like reptile) tracks, and the trails of arthropods. Excellent preservation and high track densities mark the Gateway assemblages in a thin stratigraphic interval in the upper part of the Chinle Group (Rock Point Formation). The track assemblages are similar to those reported from the Chinle Group in other parts of the Colorado Plateau and Rocky Mountain region, extending over most of Colorado, Utah, northern Arizona and northern and eastern New Mexico. Some of the Chinle ichnotaxa (Grallator and Brachychirotherium) are found in the overlying Wingate Formation, indicating that it is also Late Triassic in age, at least in the lower part. However, overall the Chinle and Wingate assemblages are quite different, most notably in the rarity of mammaloid/mammal-like tracks in the Chinle Group.     

The “Plastotype Problem” in Ichnological Taxonomy

Abstract

When naming ichnotaxa based on uncollectable trace fossils, the holotype is the actual ichnofossil in the outcrop, though some ichnologists identify the holotype as a replica (cast) held in a museum collection, and refer to it as a “plastotype,” although not all such replicas are made from plaster. Nevertheless, through its Code, the International Commission on Zoological Nomenclature (ICZN) makes it clear that an artificial, human-made replica (plaster cast or otherwise) is not eligible to be the holotype of an ichnotaxon. This directive is potentially destabilizing to much ichnological taxonomy, which is based on holotypes left in the field that have disappeared or will disappear. One possible solution for ichnologists will be to petition the ICZN to recognize that artificial, human-made replicas of ichnofossils can serve as name-bearing types. These may best be called “axiotypes” (from the Greek axios, meaning “of equal value”).     

Stellavelum arborensis igen., sp. nov., Stellavelum uncinum igen., isp. nov. and Termitichnus namibiensis isp. nov.; New Ichnofossils from Cenozoic deposits of Namaqualand,South Africa

Three new ichnotaxa are described from Cenozoic paralic to continental sediments on the west coast of South Africa. Stellavelum arborensis and Stellavelum uncinum are assigned to a mixed Skolithos and Cruziana ichnofacies and Termitichnus namibiensis is assigned to the Termitichnus ichnofacies. An interpretation of the sedimentary record shows that the Stellavelum trace makers were probably sediment ingesting annelids that lived in dense colonies in sheltered lagoonal environments during Neogene times. The Termitichnus namibiensis trace fossils are interpreted as actively backfilled termite nests which form an early to mid‐Pleistocene palimpsest, terrestrial ichnofabric which is superimposed on the older marine sediments.     

Linkichnus Terebrans New Ichnogenus et Ichnospecies,an Insect Boring from the Late Triassic of the Germanic Basin,Southern Germany

The proposed new ichnotaxon Linckichnus terebrans n. ig. et isp., is a small test-tube shaped boring produced by insects in a sphenophyte stem and occurs in Upper Triassic (Keuper) terrestrial deposits.     

A fresh look at ancient dungs: the Brazilian Triassic coprolites revisited

Terrestrial tetrapod coprolites are abundant in the Middle to Late Triassic of southern Brazil, but only few specimens have been described in just a couple of papers. Here we revisit the already‐known specimens and describe new materials, including their size, shape, external modifications, matrix composition and rare inclusions. Regarding size and shape, the measurements of 152 specimens show that the Triassic coprolites of Brazil follow a normal distribution, in which length and width are positively correlated variables, so that they are not useful for taxonomic purposes. Notwithstanding, two ichnotaxa, Santamariacopros elongatus and Rhynchocopros soutoi, were previously described from the Middle/Upper Triassic Santa Maria Formation, based mainly on morphological (length/width ratio) and preservational features, but we discuss here the validity of such taxa. Once the Santa Maria Formation encompasses at least three tetrapod Assemblage Zones (AZ), Dinodontosaurus AZ, Santacruzodon AZ and Hyperodapedon AZ (from the oldest to the youngest, respectively), it is difficult to attribute these coprolites to any specific animal producers. In addition, the rarity of inclusions and the carbonated composition of the coprolites, related to early diagenetic processes, complicate this attribution too. Nevertheless, some rare inclusions as a hair‐like structure and a parasite egg were found in this study. Although the coprotaxonomy fails to represent the Santa Maria Formation coprolites and the recognition of their producers is a problematic task, these fossil dungs are important elements in the knowledge of the Middle to Late Triassic environments of southern Brazil.     

The Triassic fish faunas of the Cuyana Basin,Western Argentina

Abstract: The continental deposits of the Cuyana Basin, western Argentina, have yielded the most diverse, but so far almost unstudied, Triassic ichthyofaunas of South America. Here, we review these faunas and show that only eight of the 29 named taxa can be considered valid, including the chondrostean Neochallaia, the acrolepid Challaia, Guaymayenia, a taxon of uncertain affinities, and five species of the perleidiform family Pseudobeaconiidae. The first three taxa most probably come from Middle Triassic sediments, while the pseudobeaconiids are of Late Triassic age. Other material, although not diagnostic, probably represents other species, and thus, the diversity of actinopterygians in the Cuyana basin is certainly higher than currently recognized. For the Late Triassic fish fauna, the absence of crown‐group neopterygians and a single record of a sarcopterygian is noteworthy and probably indicates some degree of endemism in this fauna, also supported by the high abundance of pseudobeaconiids, which are unknown from other areas. Furthermore, on the basis of the age indicated by the fishes and the available geological information, we discuss the age of the local fauna of the Cerro Bayo, close to the city of Mendoza, and the Agua de la Zorra Formation, Uspallata.     

Bite traces on dicynodont bones and the early evolution of large terrestrial predators

Nied?wiedzki, G., Gorzelak, P. & Sulej, T. 2010: Bite traces on dicynodont bones and the early evolution of large terrestrial predators. Lethaia, Vol. 44, pp. 87–92. Dicynodont (Synapsida: Anomodontia) bones from the Late Triassic (late Norian/early Rhaetian) of Poland yield characteristic tooth marks that can be attributed to three ichnotaxa (Linichnus serratus, Knethichnus parallelum and Nihilichnus nihilicus). The general shape and dimension of these traces perfectly match the dental morphology of a co‐occurring carnivorous dinosaur. It is therefore concluded that early carnivorous dinosaurs were feeding on dicynodonts. This discovery constitutes one of the oldest evidence of dinosaur predator–prey interaction. It is suggested that an evolutionary increase in the size of dicynodonts across the Late Triassic may have been driven by selection pressure to reach a size refuge from early dinosaur predators. □Bite traces, dicynodonts, dinosaurs, predation, Triassic.     

Halorina cryptica nov. ichnogen., nov. ichnosp., mass-occurrence of Upper Triassic crustacean microcoprolites from neptunian dikes and sills cutting the Dachstein-type carbonate platform and their paleoenvironmental significance (Northern Apuseni Mountains,Romania)

The new crustacean microcoprolite Halorina cryptica nov. ichnogen., nov. ichnosp., is reported from cryptic cavities cutting the Upper Triassic (Carnian to lower Rhaetian) Dachstein-type limestones from the Northern Apuseni Mountains, Romania. The new ichnotaxon is extremely abundant in cavities, neptunian dikes and sills filled with red ferruginous carbonate sediment. The associated microfauna consists of ostracods and rare foraminifera. The microfacies is represented by bioclastic coprolite-bearing wackestone-packstone to grainstone. The red ferruginous carbonate fillings are strongly bioturbated. The neptunian sill located at the top of the studied section contains a rich ichnofauna associated with brachiopod accumulations. It is dominated by the dimerelloid rhynchonellid Halorella, indicating a late Norian-early Rhaetian age. Although neptunian dikes and sills are rather common in the Dachstein-type carbonate platform that extended on the northwestern Tethyan margin, we report here the first record of mass-occurrence of crustacean microcoprolites in neptunian dikes and sills filled with red carbonate sediments of marine origin. They are interpreted here as cryptic cavities with specific palaeoenvironmental features (e.g., lack of light, abundant nutrients, intensive microbial activity) where crustacean arthropods thrived.     

The Tetrapod Ichnogenus Protochirotherium Fichter and Kunz 2004, a Characteristic Early Triassic Morphotype of Central Pangea

Early Triassic chirotherian footprint assemblages from Poland, Germany, and Morocco are important for understanding archosaur evolution in the aftermath of the Permian-Triassic crisis. However, their ichnotaxonomy is confusing because various authors have interpreted their diversity differently. After an analysis and ichnotaxonomic re-assessment, the presence of the ichnogenera Brachychirotherium, Isochirotherium, and Chirotherium in these assemblages is not supported. Distant similarities with these ichnotaxa are functions of extra morphological variation and substrate-related factors. Instead, Early Triassic chirotherian footprints described under these names are assigned here to the ichnogenus Protochirotherium and to a more slender morphotype identified as Synaptichnium. In particular, Protochirotherium appears to be more widely distributed in central Pangea as a characteristic morphotype reflecting a distinct stage in archosaur evolution. Trackmakers were nonarchosaurian archosauriforms or, alternatively, stem-group crocodylians. Morphologically and temporally these footprints match the hypothetical ancestor of the Chirotherium barthii trackmaker. Chirotherium barthii appears by the beginning of the Middle Triassic. Because of its restricted stratigraphic range, and its wider distribution in central Pangea, Protochirotherium also has biostratigraphic significance for this region and can be considered as an indicator of Early Triassic-aged strata.     

Plant Traces Resembling Skolithos

The ichnogenus Skolithos Haldeman 1840 is a simple tubular trace fossil that was initially described as a “fucoid” or seaweed impression, i.e., a fossilized marine plant. Today the tracemaker is commonly considered to have been a vermiform marine invertebrate, although this interpretation is not acceptable to all. It is the name giver to the archetypal, Seilacherian Skolithos ichnofacies, which is a widely acknowledged indicator of relatively high energy, shallow water, nearshore to marginal marine environments. Here we record and describe analogous cylindrical structures, unquestionably reflecting plant tracemakers. These have been recognized in Quaternary consolidated “coffee” sandrock and Recent coastal dune, marsh or swamp, and salt meadow settings from northern New Zealand and Sapelo Island, USA.

We conclude that Skolithos is not an unequivocal indicator of shallow marine settings. Our observations confirm the opinions of a number of previous workers who have expressed the need for caution when using this ichnotaxon as a shallow-water, higher energy palaeoenvironmental indicator, especially when supporting evidence is otherwise lacking. In particular we note that in high intertidal and non-aquatic settings, Skolithos may be a plant trace fossil rather than serve as evidence for invertebrate activities. At the present time there is pressing need for a thorough systematic revision of the Skolithos ichnotaxon.     

A Late Triassic Invertebrate Ichnofauna from Ghost Ranch,New Mexico

Invertebrate trace fossils are reported from the Late Triassic Chinle Formation at Ghost Ranch, New Mexico, for the first time. They occur in beds higher in the section than the Coelophysis Quarry. Six ichnotaxa are recognized: Steinichnus milfordensis, Planolites montanus, Palaeophycus tubularis, Taenidium serpentium, ?Arenicolites sp. and Skolithos sp. This ichnofauna is consistent with the Scoyenia ichnofacies, considered typical of lake margins in semiarid to arid regions.     

Parataenidium,a new taenidium‐like ichnogenus from the carboniferous of Ireland

The new ichnogenus Parataenidium is erected for backfilled tubular trace fossils that can appear superficially similar to Taenidium, but are divided horizontally into two distinct levels. Two ichnospecies are recognised: Parataenidium mullaghmorensis isp. nov. and Parataenidium moniliformis (Tate 1859). The latter ichnospecies is transferred from Eione Tate 1859, which is a junior homonym of Eione Rafinesque 1814, and therefore unavailable for Tate's ichnotaxon. The ichnogenus is an important component of late Paleozoic shallow‐water siliciclastic sediments, and can be considered as a “guide”; indicator for the Carboniferous.     

A Revised Systematic Ichnotaxonomy and Review of the Vertebrate Footprint Ichnofamily Chirotheriidae from the British Triassic

Since 1838, many discoveries of the tetrapod footprint taxon Chirotherium and closely similar forms have been reported from the British Middle Triassic. These have been assigned to dozens of ichnotaxa, so that the identification and interpretation of members of the ichnofamily Chirotheriidae have become confused.

Part 1 of this article deals with ichnotaxonomy. Revision of all the available material shows that there are at least three valid ichnospecies of Chirotherium, one of Synaptichnium, and one of Isochirotherium. The presence of a fourth related ichnogenus, Brachychirotherium, cannot yet be confirmed in Britain. Emended diagnoses are proposed for all four ichnogenera; they may be readily distinguished by the shape of the hand (manus) and foot (pes) impression and especially by the phalangeal counts. The natural cast of a probable tail impression, associated with Chirotheriidae footprints from the British Triassic is figured for the first time, as is a rare detailed impression of skin and the tarsal region of an Isochirotherium lomasi pes cast.

Part 2 provides a critical history of the earliest discoveries of each of these ichnogenera in the British Triassic. It is subdivided into divisions, depending on the quality of data and existence of specimens, the localities in Britain which have yielded these ichnogenera; a detailed history and discussion is provided of the earliest discoveries made at each locality. The stratigraphic distribution and paleological significance of the ichnogenera/species are assessed.     

Commentary

A new ichnotaxon is recognized in the Perkasie Member of the Passaic Formation (Upper Triassic) at the Smith Clark Quarry, Milford, New Jersey. Spongeliomorpha milfordensis is a small trace fossil possessing sharply incised partial wall striations which form a distinctive angle with the axis of the burrow. It is construed as a deposit‐feeding structure possibly formed by aquatic insect larvae along the margin of a lake.     

Revision and re-evaluation of the Early Jurassic dinosaurian ichnogenus Otozoum

Otozoum moodii Hitchcock, 1847 is one of the classic Connecticut Valley ichnotaxa established by Edward Hitchcock between 1836 and 1865. The taxon is redescribed, and AC 4/1a re-established as the holotype specimen. Otozoum minus and O. caudatum are synonymised with O. moodii . The syntypes of Kalosauropus pollex ( nomen nudum ) are described as O. pollex sp. nov. Cladistic, quantitative, and comparative methods of trackmaker identification suggest that Otozoum was made by a prosauropod dinosaur. In quantitative analyses, only those phalanges likely to contribute to footprint morphology were considered; claws were also excluded due to their highly variable styles of impression. Several diagnostic characters distinguish Otozoum from Brachychirotherium , Chirotherium , Batrachopus , Tetrasauropus , and Pseudotetrasauropus ; these ichnotaxa are probably attributable to crurotarsan archosaurs. Otozoum is found primarily in Hettangian strata of the Hartford, Deerfield, and Fundy rift basins of eastern North America; it is also present in the Early Jurassic Navajo Sandstone (Colorado Plateau, USA) and Clarens Formation (Lesotho). The taxon is unknown in older strata; Triassic material previously referred to Otozoum (including O. grandcombensis ) instead belongs in Pseudotetrasauropus . Correlation of sediments and comparison of faunal assemblages, particularly within the Newark Supergroup, suggest that the Otozoum trackmakers were restricted to arid environments.     

Ichnologic note: in defence of Thatchtelithichnus Zonneveld,Bartels, Gunnell and McHugh, 2015

Abstract

The validity of the monotypic ichnogenus Thatchtelithichnus has been called into question due to an inference that it is a bioclaustration. Close analysis of the Thatchtelithichnus holmani holotype reveals that no part of this ichnotaxon involves a bioclaustration, and thus it remains a valid ichnotaxon. Thatchtelithichnus holmani has been observed on the bottom shell (plastron) of cryptodiran and pleurodiran turtles in North America and Egypt where it is interpreted to represent an attachment trace of aquatic ectoparasites. It has also been observed on dinosaur long bones where it may represent the activity of beetle larvae.     

Caedichnus,a New Ichnogenus Representing Predatory Attack on the Gastropod Shell Aperture

Predatory traces, in which the tracemaker has damaged the prey animal's skeleton to kill and consume it, have a deep fossil history and have received much scientific attention. Several types of predatory traces have been assigned to ichnotaxa, but one of the most studied predatory traces, the wedge-shaped excision produced as a result of attacks mainly by crustaceans on the apertures of gastropod shells, has yet to be described as an ichnotaxon. We propose the ichnogenus Caedichnus to describe the shell damage produced by aperture peeling behavior. Caedichnus is produced by predators that are unable to crush their prey's shells outright. Depending on the predator's peeling ability and the prey's withdrawal depth within the shell, the trace can extend through several whorls of the shell. Aperture peel attacks may fail, allowing such damage to be repaired by surviving gastropods. Thus, the types of attacks that produce Caedichnus may exert selective pressure on prey to evolve better-defended shells (in the case of gastropods) or to inhabit better-defended shells (in the case of hermit crabs). The identification of these trace fossils will enhance our understanding of how predation influences the morphological, and even behavioral, evolution of prey organisms.