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.     

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.     

Jaw biomechanics in the South American aetosaur <Emphasis Type="Italic">Neoaetosauroides engaeus</Emphasis>

The function of the jaw apparatus and the possible dietary habits of the aetosaur Neoaetosauroides engaeus from the Triassic of South America were analyzed in comparison with Northern Hemisphere aetosaurs Desmatosuchus haplocerus and Stagonolepis robertsoni and the living short-snouted crocodile Alligator mississippiensis. The adductor and depressor jaw musculature of these was reconstructed on the basis of dental and skeletal comparisons with living closest relatives’ extant phylogenetic bracket (EPB), followed by the analysis of the moment arms of these muscles to infer feeding habits. The aetosaurian skull design indicates that the total leverage of the inferred jaw musculature provides force rather than speed. However, within aetosaurs, the high ratios of muscle moment arms to bite moments indicate stronger bites in the northern Hemisphere forms, and faster ones in Neoaetosauroides. These differences indicate more developed crushing, chopping, and slicing capacities, especially at the back of the tooth series for D. haplocerus and S. robertsoni; whereas it opens a window to consider different abilities in which speed is involved for N. engaeus. There are differences among aetosaurs in dental characteristics, position of the supratemporal fenestra, location of the jaw joint relative to the tooth row, and shape of the lower jaw. Neoaetosauroides does not show evidence of dental serrations and wear facets, probably consistent with a relatively soft and non-abrasive diet, for example soft leaves and/or larvae and insects without hard structures. It might be possible that Neoaetosauroides represents a tendency towards insectivorous feeding habits, exploiting a food source that was widespread in continental environments throughout the Triassic.     

THE SYSTEMATIC POSITION OF EOGINKGOITES

Eoginkgoites is a fan-shaped, imparipinnate leaf with a short rachis and long petiole. It was first described from the Upper Triassic Newark Group of Pennsylvania by Bock who assigned the fossil to the Ginkgoales. The fossil has also been found in the Upper Triassic Chinle Formation in Utah and Arizona and in the Newark Group in North Carolina. Investigation of the well-preserved specimens found in the Chinle Formation shows that the leaf has anastomosing venation, a marginal vein and paracytic (syndetocheilic) stomata. These characters indicate that the leaf is bennettitalean and Eoginkgoites is reassigned to the Bennettitales although its shape is perplexing. Eoginkgoites may be an important index fossil to the lower Upper Triassic (middle Carnian) rocks of North America.     

A NEW GENUS AND SPECIES OF SPHENODONTIAN FROM THE GHOST RANCH COELOPHYSIS QUARRY (UPPER TRIASSIC: APACHEAN), ROCK POINT FORMATION, NEW MEXICO, USA

Abstract: We document here a new taxon of sphenodontian, Whitakersaurus bermani gen. et sp. nov., that is also the most complete sphenodontian fossil from the Upper Triassic Chinle Group in the south‐western USA and the first Chinle sphenodontian represented by more than a single fragmentary dentulous element. The holotype was recovered during preparation of block C‐8‐82 from the famous Coelophysis (Whitaker) quarry at Ghost Ranch, New Mexico, and is the most complete small vertebrate recovered from the quarry. Detailed lithostratigraphy and geologic mapping demonstrate that the Whitaker quarry is in the Rock Point Formation of the Chinle Group, so Whitakersaurus is the first sphenodontian reported from this unit. Records of the phytosaur Redondasaurus at the quarry and elsewhere in the Chinle Group demonstrate that the quarry, and thus Whitakersaurus, is of Apachean (late Norian–Rhaetian) age. The sphenodontian specimen consists of incomplete left and right dentaries, a partial left? maxilla?, and impressions of a probable palatal element, all of which preserve multiple teeth. Whitakersaurus is distinct from other sphenodontians in possessing a unique combination of the following features: marginal dentition pleurodont anteriorly and posteriorly acrodont; pronounced heterodonty in dentary, with as many as 15 smaller, peg‐like teeth anteriorly and several larger, posterior teeth that are conical and striated; faint radial ornamentation of posterior tooth crowns; presence of c. 19 dentary teeth; and absence of a distinct flange on posterior teeth. Numerous other details distinguish it from both more primitive and more derived taxa. Whitakersaurus, therefore, helps to document further mosaic evolution and an extensive diversification event of sphenodontians during Triassic time. Although sphenodontian taxa are relatively easily recognized, widely distributed, and common small‐ or microvertebrate fossils, the long stratigraphic ranges of taxa known from multiple specimens hinders their utility as index fossils with which to correlate strata across Pangaea.     

The first occurrence of the aetosaurParatypothorax andressi (Reptilia,Aetosauria) in the western United States and its biochronological significance

A partial skeleton of the aetosaurParatypothorax andressi from Petrified Forest National Park, Arizona, is the first documentation of this species in North America.Paratypothorax is one of the few tetrapod taxa that occur in both the Keuper of Germany and the Chinle Group of the western United States, but it is of limited biochronological utility, because it occurs in strata that range in age from late Carnian to middle Norian.     

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.     

A new non-mammalian eucynodont from the Chinle Formation (Triassic: Norian), and implications for the early Mesozoic equatorial cynodont record

The Upper Triassic tetrapod fossil record of North America features a pronounced discrepancy between the assemblages of present-day Virginia and North Carolina relative to those of the American Southwest. While both are typified by large-bodied archosaurian reptiles like phytosaurs and aetosaurs, the latter notably lacks substantial representation of mammal relatives, including cynodonts. Recently collected non-mammalian eucynodontian jaws from the middle Norian Blue Mesa Member of the Chinle Formation in northeastern Arizona shed light on the Triassic cynodont record from western equatorial Pangaea. Importantly, they reveal new biogeographic connections to eastern equatorial Pangaea as well as southern portions of the supercontinent. This discovery indicates that the faunal dissimilarity previously recognized between the western and eastern portions of equatorial Pangaea is overstated and possibly reflects longstanding sampling biases, rather than a true biogeographic pattern.     

A new species of the enigmatic archosauromorph Doswellia from the Upper Triassic Bluewater Creek Formation,New Mexico,USA

Abstract: Doswellia sixmilensis is a new species of the doswelliid archosauromorph genus Doswellia named for an incomplete skeleton from the Upper Triassic Bluewater Creek Formation of the Chinle Group in west‐central New Mexico, USA. D. sixmilensis differs from D. kaltenbachi Weems, the type and only other known species of Doswellia, in its larger size, higher tooth count and greater heterodonty, possession of keels on the cervical centra and the presence of discrete knobs or spikes on some osteoderms. The holotype of D. sixmilensis is the fourth occurrence of Doswellia and only the second occurrence of a Doswellia skull, which includes the previously unknown premaxilla and maxilla (and therefore the best dentition) and has the best‐preserved cervical vertebrae. Although it adds to our knowledge of the anatomy of Doswellia, this new information does not alter previous concepts of the phylogenetic relationships of the doswelliid genera, largely because they are so poorly known anatomically. The genus Doswellia is known from the Newark Supergroup in Virginia, and the Chinle Group in Texas, New Mexico and Utah, in strata of Otischalkian–Adamanian age. The type locality of D. sixmilensis is c. 43 m stratigraphically below a bed from which U‐Pb dating of detrital zircons yields a maximum depositional age of c. 220 Ma, so this is a reasonable approximate numerical age for D. sixmilensis.     

Intravital Sex Identification of Adult Sterlets <Emphasis Type="Italic">Acipenser ruthenus</Emphasis> (Acipenseridae) Based on the Morphological Structure of Dorsal Scutes

We studied the morphological structure of dorsal scutes in adult individuals of Acipenser ruthenus. We determined the morphometric indices of scutes, the values of which are different in males and females. We created a system for sex identification in sterlets based on indices and criteria indicative of the shape of the first five scutes. The results can be used for developing a method for intravital sex identification in sturgeons at earlier stages of development.     

A new species of Williamsonia from the Upper Triassic Chinle Formation of New Mexico

Williamsonia nizhonia sp. nov., the first undoubted bennettitalean flower known from the Chinle Formation of Upper Triassic age in the south - western United States, is described in detail. The species is based on a single vertically compressed specimen collected from the Lower Red Member of the Chinle Formation near Fort Wingate in west-central New Mexico. It is rather small, the gynoecium is about 5 mm in diameter and is surrounded by a whorl often persistent sterile bracts about 1–5 cm long. The cuticles of the bracts and interseminal scales are 1 to 2 u thick. Stomata are only slightly sunken on the cuticles of the bracts and interseminal scales. Only a few hairs occur on the bracts. Williamsonia nizhonia cannot be attributed with any degree of confidence to either of the two bennettitalean leaves presently known in the Chinle flora.     

Variation in the Late Triassic Canjilon quarry (Upper Chinle Group, New Mexico) phytosaur skulls: a case for sexual dimorphism

The Canjilon quarry, located in north-central New Mexico near Ghost Ranch, contains a death assemblage of phytosaurs located stratigraphically high within the Petrified Forest Formation of the Chinle Group (Revueltian = early-mid Norian). The site has yielded numerous fossils ofPseudopalatus-grade phytosaurs, including at least 10 skulls collected from the locality byCharles Camp in 1928 and 1933 and another collected more recently byAlex Downs, curator of paleontology at Ghost Ranch. A re-examination of these skulls reveals two morphotypes that differ only in the relative lengths and relative robustness of their premaxillae. In these two morphotypes, the premaxillae define the shape and length of the rostral crest, the dimensions of which are independent of skull size. In one morphotype, the premaxillae are long, thin bones that lead to an abrupt, volcano-like narial crest. In the second morphotype, the premaxillae are of approximately the same length, but expand dorsoventrally halfway along their lengths, creating a longer and more robust crest. The most probable explanation of these two variants in rostral crest morphology in a Single, catastrophic death assemblage is thatPseudopalatus-grade phytosaurs are sexually dimorphic. Thus, the larger, more robust crest of the first morphotype is probably a display feature, most likely of the male animal. The more gracile snout änderest characterize the female morph. In the sample of phytosaurs examined, there are three individuals of the more robust (male) morphotype, five individuals of the gracile (female) morphotype, and three individuals (2 adult, 1 juvenile) that cannot be assigned to either morphotype because the skulls are too damaged to make an aecurate assessment. This is the first clear evidence of sexual dimorphism in phytosaurs, and has important implications for phytosaur species-level taxonomy, as well as for understanding aspects of their paleobiology (e.g., population dynamics).     

Novel data on aetosaur (Archosauria,Pseudosuchia) osteoderm microanatomy and histology: palaeobiological implications

One of the most striking features of aetosaurs is the possession of an extensive bony armour composed of dorsal, ventral and appendicular osteoderms. With the purpose of establishing the main histological changes during ontogeny and the degree of histological variation within the armour, we analysed the bone histology of dorsal (paramedian and lateral), ventral and appendicular osteoderms from different taxa from the Late Triassic of South America, including Aetosauroides scagliai, Aetobarbakinoides brasiliensis and Neoaetosauroides engaeus. Histological data support an intramembranous origin for osteoderms. Nevertheless, evidence for metaplastic ossification (i.e. structural fibres) at advanced ontogenetic stages, in at least some elements, is also present. A variant type of parallel fibred bone, which we have named ‘crossed parallel fibred bone’, is characterized for aetosaurs. In this pseudosuchian group, osteoderms exhibit very important microstructural changes during ontogeny, which can be useful for determining ontogenetic stages from isolated elements. Histological data suggest a relatively early onset of sexual maturity among aetosaurs. Microanatomical analysis from different taxa reveal that having high values of compactness is the plesiomorphic condition for Aetosauria. The notably increased compactness of the osteoderms does not appear to be related to size, ontogeny, sex or reproductive status of the individuals. Although a high degree of compactness of osteoderms and other bones has been considered as evidence for an aquatic lifestyle in vertebrates, such an inference contradicts the current concept of a fully terrestrial lifestyle in aetosaurs.     

Application of morphologic burrow interpretations to discern continental burrow architects: Lungfish or crayfish?

A methodology for trace fossil identification using burrowing signatures is tested by evaluating ancient and modern lungfish and crayfish burrows and comparing them to previously undescribed burrows in a stratigraphic interval thought to contain both lungfish and crayfish burrows. Permian burrows that bear skeletal remains of the lungfish Gnathorhiza, from museum collections, were evaluated to identify unique burrow morphologies that could be used to distinguish lungfish from crayfish burrows when fossil remains are absent. The lungfish burrows were evaluated for details of the burrowing mechanism preserved in the burrow morphologies together forming burrowing signatures and were compared to new burrows in the Chinle Formation of western Colorado to test the methodology of using burrow signatures to identify unknown burrows.

Permian lungfish aestivation burrows show simple, nearly vertical, unbranched architectures and relatively smooth surficial morphologies with characteristic quasi‐horizontal striae on the burrow walls and vertical striae on the bulbous terminus. Burrow lengths do not exceed 0.5 m. In contrast, modern and ancient crayfish burrows exhibit simple to highly complex architectures with highly textured surficial morphologies. Burrow lengths may reach 4 to 5 m.

Burrow morphologies unlike those identified in Gnathorhiza aestivation burrows were found in four burrow groups from museum collections. Two of these groups exhibit simple architectures and horizontal striae that were greater in sinuosity and magnitude, respectively. One of these burrows contains the remains of Lysoro‐phus, but the burrow surface reveals no reliable surficial characteristics. It is not clear whether Lysorophus truly burrowed or merely occupied a pre‐existing structure. The other two groups exhibit surficial morphologies similar to those found on modern and ancient crayfish burrows and may provide evidence of freshwater crayfish in the Permian.

Burrows from the Upper Triassic Chinle Formation in western Colorado exhibit simple to moderately complex architectural morphologies, ranging from predominantly vertical, unbranched, with little or no chamber development to predominantly vertical, few branches, and with minor chamber development. Surficial burrow morphologies are moderate to highly textured. The burrows have scrape marks, scratch marks, mud and lag‐liners, knobby surfaces, pleopod striae, and body impressions.

Although no fossil remains of the burrowing organism were found within or associated with the Chinle burrows from western Colorado, the similarity of architectural and surficial burrow morphologies to those in the Chinle of Canyonlands, Utah and to modern crayfish burrows, clearly indicates that the Colorado burrows are the product of burrowing crayfish rather than lungfish. Evaluation of burrowing signatures preserved in the architectural and surficial burrow morphologies is a very useful tool to compare and contrast Chinle burrows from different regions on the Colorado Plateau. Documentation of crayfish burrows in the Chinle of Utah and Colorado strongly suggests that other large‐diameter Chinle burrows elsewhere on the Colorado Plateau and in stratigraphically equivalent units may also be the product of crayfish activity.     

Dermal armour histology of aetosaurs (Archosauria: Pseudosuchia), from the Upper Triassic of Argentina and Brazil

Cerda, I.A. & Desojo, J.B. 2010: Dermal armour histology of aetosaurs (Archosauria: Pseudosuchia), from the Upper Triassic of Argentina and Brazil. Lethaia, Vol. 44, pp. 417–428. One of the most striking features documented in aetosaurs is the presence of an extensive bony armour composed of several osteoderms. Here, we analyse the bone microstructure of these elements in some South American Aetosaurinae aetosaurs, including Aetosauroides scagliai. In general terms, Aetosaurinae osteoderms are compact structures characterized by the presence of three tissue types: a basal cortex of poorly vascularized parallel‐fibred bone tissue, a core of highly vascularized fibro‐lamellar bone, and an external cortex of rather avascular lamellar bone tissue. Sharpey’s fibres are more visible at the internal core, toward the lateral margins and aligned parallel to the major axis of the dermal plate. No evidence of metaplastic origin is reported in the osteoderms, and we hypothesize an intramembranous ossification for these elements. The bone tissue distribution reveals that the development of the osteoderm in Aetosaurinae starts in a position located medial to the plate midpoint, and the main sites of active osteogenesis occur towards the lateral and medial edges of the plate. The osteoderm ornamentation is originated and maintained by a process of resorption and redeposition of the external cortex, which also includes preferential bone deposition in some particular sites. Given that no secondary reconstruction occurs in the osteoderms, growth marks are well preserved and they provide very important information regarding the relative age and growth pattern of Aetosaurinae aetosaurs. □Aetosauria, Aetosauroides, Archosauria, bone microstructure, integumentary skeleton, osteoderm.     

The red-fin Decapterus group (Perciformes: Carangidae) with the description of a new species, Decapterus smithvanizi

The carangid genus Decapterus can be defined by having a single finlet behind both the second dorsal and anal fins, and lacking scutes on the anterior curved part of the lateral line. We revised taxonomically the species of Decapterus with red-colored caudal fins (the red-fin Decapterus group) and established that the group consisted of the following four species: Decapterus akaadsi Abe 1958, distributed in the eastern Indian Ocean and West Pacific from the Andaman Sea to Indonesia, north to central Japan; Decapterus kurroides Bleeker 1855, distributed in the Indo-West Pacific from the Red Sea and eastern coast of Africa to eastern Australia, north to the Philippines; Decapterus smithvanizi sp. nov., occurring in the Andaman Sea, the South China Sea, and Indonesia; and Decapterus tabl Berry 1968, distributed circumglobally in tropical and subtropical seas. The diagnostic characters of these species are as follows: D. akaadsi—curved part of lateral line with 43–53 cycloid scales, straight part of lateral line with 26–29 scutes, head length 26.7–30.1 % SL, and body depth 24.0–27.9 % SL; D. kurroides—curved part of lateral line with 45–51 cycloid scales, straight part of lateral line with 30–32 scutes, head length 30.3–33.0 % SL, and body depth 23.4–26.4 % SL; D. smithvanizi—lower gill rakers 25–31, curved part of lateral line with 54–62 cycloid scales, body depth 19.4–22.5 % SL, pectoral-fin tip usually beyond the level of second dorsal-fin origin; D. tabl—tip of upper jaw usually hooked and opercular membrane partly serrated in larger specimens, lower gill rakers 28–33, curved part of lateral line with 61–72 cycloid scales, body depth 16.6–23.0 % SL, pectoral-fin tip not reaching to the level of second dorsal-fin origin.     

Rioarribasaurus,a new name for a Late Triassic dinosaur from New Mexico (USA)

The Late-Triassic-dinosaur generic nameCoelophysis Cope 1889 (type species C.bauri [Cope 1887]) is a nomen dubium because the lectotype of C.bauri, AMNH 2722, is four sacral vertebrae and a pubic process of the ilium that are not diagnostic. Dinosaur specimens from the famous Whitaker (“Coelophysis”) quarry in the Rock Point Member of the Chinle Formation at Ghost Ranch, New Mexico thus lack a valid name. We create a new genus and species name,Rioarribasaurus colberti, for these specimens.     

Genetic diversity of the harmful family Kareniaceae (Gymnodiniales,Dinophyceae) in France,with the description of Karlodinium gentienii sp. nov.: A new potentially toxic dinoflagellate

The family Kareniaceae is mostly known in France for recurrent blooms of Karenia mikimotoi in the Atlantic, English Channel, and Mediterranean Sea and for the unusual green discoloration in the saltwater lagoon of Diana (Corsica) caused by Karlodinium corsicum in April 1994. In terms of diversity, this taxonomic group was long overlooked owing to the difficult identification of these small unarmored dinoflagellates. In this study, thanks to the molecular characterization performed on single cells from field samples and cultures, twelve taxonomic units were assigned to the known genera Karenia, Karlodinium and Takayama, whereas one could not be affiliated to any described genus. The molecular phylogeny inferred from the D1–D2 region of the LSU rDNA showed that five of them formed a sister taxon of a known species, and could not be identified at species-level, on the basis of molecular analysis only. Among these latter taxa, one Karlodinium which was successfully cultured was investigated by studying the external morphological features (using two procedures for cells fixation), ultrastructure, pigment composition, and haemolytic activity. The results of our analyses corroborate the genetic results in favour of the erection of Karlodinium gentienii sp. nov., which possesses an internal complex system of trichocysts connected to external micro-processes particularly abundant in the epicone, and a peculiar pigment composition. In addition, preliminary assays showed a haemolytic activity.