Typothorax Research Articles

Incongruent ontogenetic maturity indicators in a Late Triassic archosaur (Aetosauria: Typothorax coccinarum).

Maximum individual body size in pseudosuchian archosaurs is not well constrained in the fossil record, but it may be influenced by a variety of factors including basal metabolic rate, evolutionary relationships, and environmental conditions. Body size varies among the Aetosauria in which estimated total length ranges between 1 m (e.g., Coahomasuchus kahleorum) and 5 m (e.g., Desmatosuchus spurensis). A new, very large specimen of the aetosaurian Typothorax coccinarum from Petrified Forest National Park in northeastern Arizona is nearly twice the size of all other known specimens of Typothorax and is the largest aetosaur specimen currently known worldwide. The specimen lacks co-ossified neurocentral sutures in the trunk vertebrae which may suggest that the individual had not yet reached skeletal maturity, yet smaller specimens of T. coccinarum exhibit partially or fully co-ossified neurocentral sutures in the same region. If body size correlates with skeletal maturity in aetosaurs, this discrepancy warns that timing of neurocentral suture co-ossification in aetosaurs may not be a reliable indicator of ontogenetic stage. Osteohistological observations of a trunk rib demonstrate that although PEFO 42506 showsa large body size, the specimen did not deposit an external fundamental system despite depositing as many as 19 growth lines, further indicating that it had not yet reached skeletal maturity. Thus, at least within Aetosauria, neurocentral suture co-ossification and skeletal maturity may correlate, whereas body size can be incongruent in comparison. Furthermore, this specimen indicates that non-desmatosuchin aetosaurs could exhibit large body sizes and suggests that some aetosaurs may have experienced indeterminate growth.

BITE MARKS ON AN AETOSAUR (ARCHOSAURIA, SUCHIA) OSTEODERM: ASSESSING LATE TRIASSIC PREDATOR-PREY ECOLOGY THROUGH ICHNOLOGY AND TOOTH MORPHOLOGY

ABSTRACTTrace fossils such as bite marks provide rare, direct evidence of animal behavior, including predator-prey interactions. We present an osteoderm of the aetosaur Typothorax coccinarum from the Late Triassic Chinle Formation of Arizona with several punctures and scores, interpreted here as bite marks, preserved as evidence of predation/scavenging by a large carnivore. The marks include a single bite producing four subparallel fusiform pits on the ventral surface and several additional marks, including striated scores, on the dorsal surface. These traces are described and compared with known contemporaneous carnivorous taxa to determine the source of the bite marks. Some Triassic carnivores, including theropod dinosaurs can be ruled out because of tooth shape and serration densities. Phytosaurs and large paracrocodylomorphs remain as likely candidates based on tooth morphology. Although some phytosaur teeth are too rounded to produce the marks seen in this specimen, we demonstrate that the more lingually flattened teeth typically found in the posterior section of the snout are sufficiently mediolaterally compressed to produce a fusiform pit. A protective function for aetosaur osteoderms cannot be confirmed presently, but the extensive carapace these bones formed would have been a major barrier to both scavengers and active predators and may preserve more feeding/predation traces than previously thought. The bite marks described herein support the hypothesis that aetosaurs were prey items of large archosauromorphs, expanding our understanding of the complex, and seemingly carnivore dominated Late Triassic terrestrial ecosystems of North America.

Cranial anatomy and dentition of the aetosaur Typothorax coccinarum (Archosauria: Pseudosuchia) from the Upper Triassic (Revueltian–mid Norian) Chinle Formation of Arizona

ABSTRACT Our understanding of Typothorax coccinarum is primarily based on postcranial material, along with a few isolated cranial elements. Here we describe the first complete articulated skull of Typothorax coccinarum from the Owl Rock Member of the Late Triassic Chinle Formation in Petrified Forest National Park. We assessed the relationships and defining synapomorphies for some of the major clades within Aetosauria by using a combination of morphological characters in the cranium of Typothorax coccinarum. Based on the descriptions of skulls from a variety of aetosaur species including Aetosaurus ferratus, Aetosauroides scagliai, Coahomasuchus chathamensis, Neoaetosauroides engaeus, Paratypothorax andressorum, Stagonolepis olenkae, Stenomyti huangae, Scutarx deltatylus, and now Typothorax coccinarum, it is apparent that the two major aetosaurian clades vary in their overall skull morphologies. The stagonolepidoids share a more robust, elongate skull, with varying degrees in the lateral expansion of the premaxillary tips. On the other hand, aetosaurines share an overall shorter, more gracile skull with tapered premaxillae. These new characters are mapped onto a revised phylogenetic tree for the Aetosauria. Historically, aetosaurs were interpreted as being predominantly herbivorous. In recent years, the discovery of new aetosaur cranial material, like that of Neoaetosauroides engaeus and Aetosauroides scagliai, suggested an omnivorous diet for at least some aetosaur taxa. The dentition of Typothorax coccinarum possesses the most strongly developed heterodonty for an aetosaur, therefore challenging the historical interpretation by expanding the range of dentition types within Aetosauria. This supports the more recent hypothesis that aetosaurs exhibited diverse feeding strategies beyond just herbivory.

Growth curve of Aetosauroides scagliai Casamiquela 1960 (Pseudosuchia: Aetosauria) inferred from osteoderm histology

Abstract Recent palaeohistological studies on paramedian osteoderms of aetosaurs revealed the presence of growth lines (lines of arrested growth or LAGs) and a minimal or nonexistent secondary remodelling in the bone matrix of these elements. This feature allows the age of individuals to be estimated through growth line count. In the present contribution we study the growth curve of the South American aetosaur Aetosauroides scagliai . We estimated the age (obtained from LAG counting) and body size (body length and body mass were used as proxies) of different aetosaur specimens in order to reconstruct the growth curve of the South American species. The data obtained for Aetosauroides scagliai were compared with that of other aetosaurs, such as Neoaetosauroides engaeus , Aetosaurus ferratus , Aetobarbakinoides brasiliensis , Typothorax coccinarum and Paratypothorax sp. Our results indicate that, if body length is considered as proxy, all studied aetosaur specimens have a similar or almost identical growth rate. However, important variations arose among aetosaur taxa if body mass is considered as proxy, which would be related to a body morphology ranging from slender (e.g. Aetobarbakinoides brasiliensis ) to very wide ( Typothorax coccinarum ) morphotypes. In comparison with extant pseudosuchians (i.e. crocodylians), Aetosauroides scagliai possesses a relatively lower growth rate.

A taxonomic and biostratigraphic re-evaluation of the Post Quarry vertebrate assemblage from the Cooper Canyon Formation (Dockum Group, Upper Triassic) of southern Garza County, western Texas

ABSTRACTThe Post Quarry, within the lower part of the type section of the Upper Triassic Cooper Canyon Formation in southern Garza County, western Texas, contains a remarkably diverse vertebrate assemblage. The Post Quarry has produced: the small temnospondylRileymillerus cosgriffi; the metoposauridApachesaurus gregorii; possible dicynodonts and eucynodonts; a clevosaurid sphenodontian; non-archosauriform archosauromorphs (Trilophosaurus dornorum, simiosaurians, and possiblyMalerisaurus); the phytosaurLeptosuchus; several aetosaurs (Calyptosuchus wellesi,Typothorax coccinarum,Paratypothorax, andDesmatosuchus smalli); the poposauroidShuvosaurus inexpectatus(“Chatterjeea elegans”); the rauisuchidPostosuchus kirkpatricki; an early crocodylomorph; several dinosauromorphs (the lagerpetidDromomeron gregorii, the silesauridTechnosaurus smalli, a herrerasaurid, and an early neotheropod); and several enigmatic small diapsids. Revised lithostratigraphic correlations of the lower Cooper Canyon Formation with the Tecovas Formation, the occurrence ofLeptosuchus, and the overall composition of the assemblage indicate that the Post Quarry falls within the Adamanian biozone, and not the Revueltian biozone. Stratigraphic subdivision of the Adamanian biozone may be possible, and the Post Quarry may be correlative with the upper part of the Adamanian biozone in Arizona. The age of the Post Quarry assemblage is possibly late Lacian or earliest Alaunian (late early Norian or earliest middle Norian), between 220 and 215 Ma.

Redescription and taxonomic status of specimens of Episcoposaurus and Typothorax, the earliest known aetosaurs (Archosauria: Suchia) from the Upper Triassic of western North America, and the problem of proxy “holotypes”

ABSTRACTHistoric type and referred material of the aetosaurian taxa Typothorax coccinarum, Episcoposaurus horridus and Episcoposaurus haplocerus are redescribed and the non-aetosaurian material identified and removed, a task previously considered “hopeless”. Reexamination of the original material reveals that the holotypes of E. haplocerus and probably T. coccinarum are not diagnosable at the species level and therefore are nomena dubia. The next available names for material referred to these taxa are Desmatosuchus spurensis and E. horridus respectively, although it may be more desirable for reasons of taxonomic stability to attempt to petition for a neotype in the latter case. The redescription of historical specimens is necessary to determine their nomenclatural validity. The use of referred specimens as proxy “type” specimens is problematic, as these referrals were originally made not on the basis of apomorphies, but rather on biostratigraphic and/or geographical assumptions which are inherently circular and cannot be unambiguously supported.

Late Triassic Aetosaurs as the Trackmaker of the Tetrapod Footprint Ichnotaxon Brachychirotherium

Brachychirotherium is the common ichnogenus of Late Triassic chirothere footprints well known from western Europe, North America, Argentina and South Africa. Although it has long been agreed by most workers that the trackmaker of Brachychirotherium was a derived crurotarsan archosaur, the trackmaker has been identified as either a rauisuchian or an aetosaur, and some workers attribute it to a primitive crocodylomorph (sphenosuchian). New knowledge of the osteology of the manus and pes of a large aetosaur, Typothorax coccinarum, indicates a close correspondence between the manus and pes structure of aetosaurs and the morphology of Brachychirotherium. Furthermore, functional analysis of complete skeletons indicates aetosaurs plausibly placed their feet in the narrow gauge, nearly the overstepped walk characteristic of Brachychirotherium. Brachychirotherium and aetosaurs have matched distributions, that is, they were Pangea-wide during the Late Triassic. The manus and pes morphology of rauisuchians and early crocodylomorphs (sphenosuchians) deviate from Brachychirotherium footprint morphology in key features, thus excluding their identification as trackmakers. Aetosaurs made Brachychirotherium footprints.

Articulated skeletons of the aetosaur Typothorax coccinarum Cope (Archosauria: Stagonolepididae) from the Upper Triassic Bull Canyon Formation (Revueltian: early-mid Norian), eastern New Mexico, USA

ABSTRACT We report two nearly complete, articulated skeletons of the crurotarsan archosaur Typothorax coccinarum from the Upper Triassic Bull Canyon Formation of east-central New Mexico. These are the most complete, articulated aetosaurs from North America and provide a wealth of new anatomical and paleobiological data, including articulated presacral armor that confirms the distinctiveness of T. coccinarum from the closely related T. antiquum and from Redondasuchus. Cervical vertebrae are small, but the corresponding reduction in armor is accomplished by a reduced number of cervical osteoderms. The third row of osteoderms includes a thin, elongate, lateral spike. The ventral armor consists of 10 thoracic columns and four caudal columns of osteoderms. Spiked osteoderms near the cloacal vent are the first spikes reported in aetosaurian ventral osteoderms. The forelimb of T. coccinarum was very short, only ∼0.65 the length of the hind limb, possesses some adaptations found in digging taxa, and was held in a sprawling or ‘semi-erect’ position. In contrast the hind limb is much more robust, ‘pillar erect,’ and functionally mesotarsal. The articulated pes, including unguals, has, minimally, the phalangeal formula 2-3-3?-4?-3? with relative digit lengths III > II > IV > I > V, digits I–IV equally as wide as long and other characteristics of the footprint ichnogenus Brachychirotherium, often attributed to an aetosaurian trackmaker. Both specimens are ∼2.5 m long and the preserved armor and limb bones are as large or larger than known Typothorax fossils, suggesting that this approximates the upper size limit of T. coccinarum, and we calculate body mass estimates of ∼100–104 kg for both specimens.

The Triassic timescale based on nonmarine tetrapod biostratigraphy and biochronology

Abstract The Triassic timescale based on nonmarine tetrapod biostratigraphy and biochronology divides Triassic time into eight land-vertebrate faunachrons (LVFs) with boundaries defined by the first appearance datums (FADs) of tetrapod genera or, in two cases, the FADs of a tetrapod species. Definition and characterization of these LVFs is updated here as follows: the beginning of the Lootsbergian LVF=FAD of Lystrosaurus ; the beginning of the Nonesian=FAD Cynognathus ; the beginning of the Perovkan LVF=FAD Eocyclotosaurus ; the beginning of the Berdyankian LVF=FAD Mastodonsaurus giganteus ; the beginning of the Otischalkian LVF=FAD Parasuchus ; the beginning of the Adamanian LVF=FAD Rutiodon ; the beginning of the Revueltian LVF=FAD Typothorax coccinarum ; and the beginning of the Apachean LVF=FAD Redondasaurus . The end of the Apachean (= beginning of the Wasonian LVF, near the beginning of the Jurassic) is the FAD of the crocodylomorph Protosuchus . The Early Triassic tetrapod LVFs, Lootsbergian and Nonesian, have characteristic tetrapod assemblages in the Karoo basin of South Africa, the Lystrosaurus assemblage zone and the lower two-thirds of the Cynognathus assemblage zone, respectively. The Middle Triassic LVFs, Perovkan and Berdyankian, have characteristic assemblages from the Russian Ural foreland basin, the tetrapod assemblages of the Donguz and the Bukobay svitas, respectively. The Late Triassic LVFs, Otischalkian, Adamanian, Revueltian and Apachean, have characteristic assemblages in the Chinle basin of the western USA, the tetrapod assemblages of the Colorado City Formation of Texas, Blue Mesa Member of the Petrified Forest Formation in Arizona, and Bull Canyon and Redonda formations in New Mexico. Since the Triassic LVFs were introduced, several subdivisions have been proposed: Lootsbergian can be divided into three sub-LVFs, Nonesian into two, Adamanian into two and Revueltian into three. However, successful inter-regional correlation of most of these sub-LVFs remains to be demonstrated. Occasional records of nonmarine Triassic tetrapods in marine strata, palynostratigraphy, conchostracan biostratigraphy, magnetostratigraphy and radioisotopic ages provide some basis for correlation of the LVFs to the standard global chronostratigraphic scale. These data indicate that Lootsbergian=uppermost Changshingian, Induan and possibly earliest Olenekian; Nonesian=much of the Olenekian; Perovkan=most of the Anisian; Berdyankian=latest Anisian? and Ladinian; Otischalkian=early to late Carnian; Adamanian=most of the late Carnian; Revueltian=early–middle Norian; and Apachean=late Norian–Rhaetian. The Triassic timescale based on tetrapod biostratigraphy and biochronology remains a robust tool for the correlation of nonmarine Triassic tetrapod assemblages independent of the marine timescale.