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Correlations between continental sequences within the Karoo-aged basins of southern and eastern Africa are difficult due to the dearth of shared index fossils and a lack of radioisotopic dates for key formations. Here we describe four sites along the southeastern shoreline of Lake Kariba, Zimbabwe, within the Mid-Zambezi Basin, that yield material of phytosaurs (Archosauromorpha: Phytosauria) from within the informal Tashinga Formation (Upper Karoo Group). These phytosaur remains are the first to be recovered from sub-Saharan mainland Africa, representing a major geographic range extension for this group into high southern latitudes. Furthermore, an LA-ICPMS maximum depositional age of 209.2 ± 4.5 Ma (late Norian/early Rhaetian) derived from detrital zircons provides the first absolute age estimate for any of these sites. The phytosaurs are associated with lungfish and metoposaurid amphibians, forming part of a terrestrial-aquatic dominated biota, a previously undocumented biome from the Late Triassic of southern Africa.

Numerous isolated teeth recovered from the Upper Triassic Tiki Formation of India have high, cylindrical, triangular to conical crowns with serrated mesial and distal carinae, and oval/subcircular and elliptical crown bases. Within this overall morphology, the teeth are subdivided into six morphotypes based on crown proportions, nature of carinae and serration and crown-base ratios. These ziphodont teeth are compared with those of other dominant Late Triassic carnivores of India to show similarity with that of the phytosaurs based on their shape, recurved nature, crown proportions, the presence of carina(e), nature of serrations and serration density. The different morphotypes reflect positional heterodonty, and three upper jaw dental sets are identified. This finding is corroborated by PCA, where the morphotypes form distinctly separate clusters, though partial overlapping of these convex hull polygons suggests that these teeth belonged to a single phytosaur taxon. As tripartite dentition is a derived condition; it may be concluded that these isolated teeth belonged to a more derived phytosaur, yet to be recovered from the Tiki Formation. The study highlights the importance of examining isolated teeth in order to identify the fossil forms which are yet to be discovered.

A monotaxic bonebed containing numerous phytosaur specimens is reported from the Upper Triassic Tiki Formation of India. The comprehensive minimum number of individuals is estimated to be 21, which includes multiple juveniles and subadults, and only a few adults. Such intraspecific aggregation of juvenile-dominated phytosaurs is explained here by parental care and juveniles living in cohorts, as are evident from extant phylogenetic bracketing of either lepidosaurs or crocodiles and birds. The cause of this mass death is probably disease related, after which the carcasses bloated, floated, and disarticulated at the site of death, resulting in an autochthonous assemblage. The carcasses remained submerged under shallow, slow-moving or standing water for a prolonged time and later were subaerially exposed after the recession of water. Subsequently, flooding events resulted in their burial under the overbank fines. Comparison of biostratinomic variables with those of a rhynchosaur-dominated bonebed recovered previously from the same stratigraphic horizon showed distinct differences in their causes of death and fossilization pathways. The latter may be attributed to their habitat differences because the phytosaurs lived in the low-lying areas or ponds and the rhynchosaurs resided on relatively higher and drier areas of the Tiki floodplain.

Herein we describe phytosaurs from thin fluvial overbank sandstones of the Upper Triassic Malmros Klint Formation of the Fleming Fjord Group (central East Greenland). The new sample includes more than 150 disarticulated bones and teeth from small to large specimens belonging to at least four individuals. The fossils mostly consist of teeth and postcranial elements and permit the recognition of a new species of Mystriosuchus, M. alleroq, diagnosed by an L-shaped quadratojugal whose anterior suture trends anterodorsally and a tripartite degree of heterodonty. Humeral diaphyseal histology of one specimen reveals a fairly compact cortex that surrounds a cancellous medullary region followed by a remodeling zone containing scattered secondary osteons. Primary bone tissue is parallel-fibred with a moderate to low vascular density. The cortex is cyclically interrupted by distinct growth marks indicating a seasonal environment. A change in growth rate from moderate to low is documented within the outer cortex, indicating that at least this individual was close to somatic maturity. Mystriosuchus has formerly been known as an exclusively European taxon. The new findings support the European faunal influence in East Greenland during the Late Triassic inferred from other taxa such as temnospondyls and archosaurs. The mid-late Norian age of European Mystriosuchus suggests an additional age constraint for the vertebrate-bearing portion of the Malmros Klint Formation.

The sacrum – consisting of those vertebrae that articulate with the ilia – is the exclusive skeletal connection between the hindlimbs and axial skeleton in tetrapods. Therefore, the morphology of this portion of the vertebral column plays a major role in the evolution of terrestrial locomotion. Whereas most extant reptiles only possess the two plesiomorphic sacral vertebrae, additional vertebrae have been incorporated into the sacrum multiple times independently among early‐diverging archosaurian (crocodylians + birds) clades. Phytosauria was a diverse, abundant, and cosmopolitan clade of archosauriforms throughout the Late Triassic, but postcrania of this clade are rarely described and few species‐level taxonomic placements of phytosaurian postcranial material are available, potentially hampering knowledge of morphological disparity in the postcranial skeleton among phytosaurs. Here, we describe the sacrum of Smilosuchus adamanensis, a phytosaur recovered from the Upper Triassic Chinle Formation of Arizona. This sacrum consists of the two primordial sacral vertebrae, but has a vertebra incorporated from the trunk into the sacrum (= a dorsosacral) and is therefore the first Late Triassic phytosaur and one of the first non‐archosaurian archosauromorphs to be described with more than two sacral vertebrae. Our interpretation of this element as a dorsosacral is justified by the lateral extent of the dorsosacral ribs, clear surfaces of articulation between the distal ends of the dorsosacral ribs and the first primordial sacral ribs, and the scar on the medial surface of each ilium for articulation with each dorsosacral rib. Additionally, we provide the first detailed description of the vertebral junction formed by two anteriorly projecting flanges on the first primordial sacral ribs and their corresponding facets on the centrum of the dorsosacral. Computed tomographic (CT) imaging reveals that the two primordial sacrals are not co‐ossified and that the dorsosacral morphology of this specimen is not the result of obvious pathology. We place this incorporation of a trunk vertebra into the phytosaurian sacrum in a broader evolutionary context, with this shift in vertebral identity occurring at least seven times independently among Triassic archosauriforms, including at least three times in early crocodylian‐line archosaurs and at least four times among bird‐line archosaurs. Additionally, anteriorly projecting flanges of sacral ribs which articulate with the anterior‐adjacent centrum have evolved several times in archosauriforms, and we interpret ‘shared’ sacral ribs (= a sacral rib that articulates with two adjacent sacral centra more or less equally) present in some archosaurian clades as a more extreme example of this morphology. In extant taxa the highly conserved Hox gene family plays a central role in the patterning of the axial skeleton, especially vertebral identity; therefore, the independent incorporation of a trunk vertebra into the sacrum across multiple archosauriform lineages may suggest a homologous underlying developmental mechanism for this evolutionary trend.

Following the end-Permian extinction, terrestrial vertebrate diversity recovered by the Middle Triassic, and that diversity was now dominated by reptiles. However, those reptilian clades, including archosaurs and their closest relatives, are not commonly found until ~30 million years post-extinction in Late Triassic deposits despite time-calibrated phylogenetic analyses predicting an Early Triassic divergence for those clades. One of these groups from the Late Triassic, Phytosauria, is well known from a near-Pangean distribution, and this easily recognized clade bears an elongated rostrum with posteriorly retracted nares and numerous postcranial synapomorphies that are unique compared with all other contemporary reptiles. Here, we recognize the exquisitely preserved, nearly complete skeleton of Diandongosuchus fuyuanensis from the Middle Triassic of China as the oldest and basalmost phytosaur. The Middle Triassic age and lack of the characteristically-elongated rostrum fill a critical morphological and temporal gap in phytosaur evolution, indicating that the characteristic elongated rostrum of phytosaurs appeared subsequent to cranial and postcranial modifications associated with enhanced prey capture, predating that general trend of morphological evolution observed within Crocodyliformes. Additionally, Diandongosuchus supports that the clade was present across Pangea, suggesting early ecosystem exploration for Archosauriformes through nearshore environments and leading to ease of dispersal across the Tethys.

As in other archosauriforms, phytosaurs and aetosaurs are characterized by the presence of well-developed osteoderms. Here we provide a comparative study on the microstructure of phytosaur (five taxa) and aetosaur (thirteen taxa) osteoderms. For outgroup comparison, we sampled osteoderms of the sister taxon to Aetosauria, Revueltosaurus callenderi, and the doswelliid Jaxtasuchus salomoni. Phytosaur, aetosaur, and Jaxtasuchus osteoderms are composed of a diploe structure, whereas the Revueltosaurus osteoderm microanatomy is more compact. The external cortex of phytosaurs, Revueltosaurus and Jaxtasuchus osteoderms is mainly composed of parallel-fibered bone. In aetosaurs, the external cortex mainly consists of lamellar bone, with lines of resorption within the primary bone indicating successive cycles of bone erosion and deposition. The basal cortex in all the specimens is composed of parallel-fibered bone, with the cancellous internal core being more strongly developed in aetosaurs than in phytosaurs. Woven or fibro-lamellar bone was recorded in both phytosaurian and aetosaurian taxa, as well as in Jaxtasuchus. Structural fibers, which at least partly suggest metaplastic origin, were only recorded in the internal core of two phytosaurs and in the basal cortex of one aetosaur. Osteoderm thickness and cancellous to compact bone ratios appear to be subject to ontogenetic change. Minimum growth mark counts in osteoderms sampled indicate that some aetosaurs and phytosaurs lived for at least two decades. Bone microstructures are more uniform in phytosaur osteoderms and show a higher level of disparity among aetosaur osteoderms, and at least in the latter, histological features are potentially apomorphic for species/genus level. Anat Rec, 297:240–260, 2014. © 2013 Wiley Periodicals, Inc.

Many recent studies have detailed the morphology of archosaurian endocrania. However, the outgroup to Archosauria, Phytosauria, has yet to be studied with modern techniques that would allow reconstruction of their internal anatomy. Pseudopalatus mccauleyi is a derived phytosaur from the Upper Triassic Chinle Formation, Arizona. A skull of P. mccauleyi, USNM 15839, was imaged using computed tomography in order to create the first high-quality, digitally reconstructed phytosaur endocast. Pseudopalatus mccauleyi exhibits overall endocast morphology that is similar to that of an extant crocodylian. These clades, phytosaurs and extant crocodylians, exhibit convergent Bauplane and similar inferred ecologies. A notable difference between the endocasts of the two clades is a considerable dural expansion in P. mccauleyi that denotes a large pineal body. This expansion, and the overall morphology of the endocast, is consistent with the historic endocranial reconstructions of the phytosaurs Pseudopalatus bu...

The Triassic was first defined based on the characteristic three-fold sequence of rocks that crops out across much of Europe, and many of the first records of Triassic dinosaurs, crocodile-line arc...

A new phytosaur taxon from Petrified Forest National Park, Arizona, is here described based on cranial material from a single individual. This specimen previously was included in an extensive phylogenetic analysis, and it was found to possess a combination of character states that differs from all known phytosaur taxa in addition to two autapomorphies within the braincase and an autapomorphy of the mandible. The new taxon adds to the taxonomic diversity recognized from the Sonsela Member of the Chinle Formation. The continued increase in phytosaur diversity emphasizes the need to more accurately characterize and identify taxa within a phylogenetic systematic context in order to produce a more refined signal for biostratigraphic correlations, biochronologic inferences, and faunal dynamics during the Late Triassic.

Leptosuchus Case, 1922 (Reptilia: Phytosauria) from the Late Triassic of the American West is represented by many specimens. Here, I present complete morphological descriptions of the skull material of a new taxon from the Sonsela Member (Chinle Formation) of Petrified Forest National Park, Arizona, with the first rigorous phylogenetic analysis focused on the interrelationships of Leptosuchus. The new taxon is recovered as the sister taxon to Pseudopalatinae. It possesses one unambiguous synapomorphy (the ‘septomaxillae’ form part of the lateral borders of the nares) and shares the presence of a subsidiary opisthotic process with Pseudopalatinae. The new taxon does not fall within the restricted clade Leptosuchus. In my analysis, the previously proposed, but undemonstrated, sister taxon relationship between Angistorhinus and Rutiodon is not supported, Paleorhinus is recovered as paraphyletic, and a subset of taxa traditionally included within Leptosuchus are found to be more closely related to Pseudopalatinae, rendering Leptosuchus paraphyletic. ‘Leptosuchus’adamanensis emerges as sister taxon to Smilosuchus gregorii and is here referred to as Smilosuchus adamanensis nov. comb., and ‘Machaeroprosopus’lithodendrorum is also transferred to Smilosuchus lithodendrorum nov. comb. Documentation of the variation present within Phytosauria, and specifically within Leptosuchus sensu lato, demonstrates higher diversity within Phytosauria than previously appreciated and places the character states previously proposed for Pseudopalatinae into a broader context of shared characters.

Mystriosuchus westphali is based on a large, well-preserved cranium and a snout fragment from the Stubensandstein (Norian) of south-west Germany. The hypodigm is redescribed and new or poorly known cranial structures in phytosaurs are discussed. For the first time, the presence of a premaxillary crest is substantiated in a phytosaur. The type specimen shows a supernumerary occipital element (=‘tabular’) that is probably fused to the parietal in other phytosaurs, and an orbitosphenoid. A computerised parsimony analysis confirms the hypothesis that Mystriosuchus is nested within Pseudopalatinae, the most derived clade of phytosaurs, and thus does not fall within basal phytosaurs. Mystriosuchus is characterised by five unique features (slit-like interpremaxillary fossa, triangular cross-section of the postorbito-squamosal bar, strongly reduced posttemporal fenestra, and two features of the cranial sculpture), plus eight synapomorphies that also occur in some more distantly related taxa. Mystriosuchus westphali is diagnosed by, among other apomorphies, a distinct premaxillary crest, a squamosal-proo¨tic contact, absence of a posterior process of the squamosal, and a slit-like posttemporal fenestra. The type species Mystriosuchus planirostris shows, most significantly, the naris facing forward anteriorly and upward posteriorly, and the longest rostrum and the highest degree of depression of the supratemporal opening in any phytosaur. Mystriosuchus exemplifies a common pattern in phytosaurids in being a genus that includes a gracile, elongated, slender-snouted and a more robust species with a broader, often crested snout. This study demonstrates that a detailed analysis of the cranial anatomy and the rigorous application of cladistic principles to identified character states help to clarify current inconsistencies in the taxonomy and nomenclature of phytosaurs.

“Zanclodon”arenaceus has been suggested to represent the oldest reliably dated phytosaur. The type and only specimen, a mandibular fragment of an archosaur from the Schilfsandstein (Carnian; Late Triassic) of Southwest Germany, is described in detail for the first time. “Z.”arenaceus is characterized by a great elongation of the mandible and symphysis, and labiolingually flattened, serrated teeth, a combination that is so far unique among Triassic archosaurs. The previous assignment to the enigmatic reptilian genusZanclodon is unsubstantiated. “Z.”arenaceus shares with Phytosauria both these mandibular characters, but differs in numerous other characters, including most of the autapomorphic mandibular and dental features of Phytosauria. Thus, “Z.”arenaceus is not a phytosaur as currently defined. A cladistic analysis identified “Z.”arenaceus as the sister-taxon of Phytosauria, but because of the limited data available and numerous homoplasies that occur among mandibular characters of archosaurs a closer relationship with other archosaur taxa is a reasonable alternative. It seems unjustified to redefine Phytosauria, or to propose a more inclusive taxon to include “Z.”arenaceus, and therefore the species is assessed as Archosauriaincertae sedis.

The holotype and only specimen of Paleorhinus (= Arganarhinus) magnoculus is most likely a juvenile, as demonstrated by absolute size and skull proportions such as large orbits, a short snout and a narrow rostrum in comparison with other primitive (non-phytosaurid) phytosaurs. The specimen is the most complete juvenile phytosaur known, and may shed light on the poorly known ontogenetic development of phytosaurs. On the basis of the data available at present, P. magnoculus, and therefore Arganarhinus, cannot be regarded as valid taxa.

The dentition of Nicrosaurus kapffi, a phytosaur from the Norian of central Europe, consists of three distinct dental sets in the upper jaw, and two in the mandible. Dental sets are defined by: (1) gradual, linear transformations of dental characters within an array that results in characteristic end members; and (2) the position of each array within the jaws. The tip-of-snout set in the upper jaw comprises mainly the large fang-like anteriormost teeth. In the premaxilla set, the teeth grade from a conical, unspecialized form anteriorly to high, D-shaped and bicarinate teeth posteriorly. In the maxilla set the teeth grade from stout, conical, unspecialized forms anteriorly to triangular forms with expanded flanges posteriorly. The tip-of-mandible and the dentary set correspond to the tip-of-snout, and to an elongated maxilla set, respectively, but were studied in less detail. The presence of tripartite and bipartite dentitions (three versus two dental sets in the upper jaw) is suggested as a crit...

Among the vertebrate remains from the Triassic of the western states no other group is so abundantly represented as the Phytosauria. Relatively abundant as fossils of this group are, however, there is much to be learned of each of the several distinct types that have been described. For the most part it is the skull that is available for study, but even this is imperfectly known. Some time ago the writer described a well-preserved phytosaur skull from Arizona, now in the geological museum of the University of Wisconsin. This skull was considered a new form and was made the type of the genus Machaeroprosopus.' While several of the doubtful details of the phytosaurian skull were made known by the study of the specimen, especially the relations of the bones of the posterior side, the palate, as is usually the case, was left in doubt. Through the kindness of the University of Chicago the writer was permitted some time ago to study a phytosaur skull in the collections of Walker Museum; a skull very similar to the University of Wisconsin specimen in many points. The study of this material has made evident several pointed suggestions especially concerning the structure of the palate. The specimen herein described is No. 396 of the Walker Museum Vertebrate Paleontology Collections. It is the gift of Professor J. E. Anderson, formerly of the School of Mines, at Socorro New, Mexico. The name of the collector is unknown and the exact locality has not been recorded. However, the skull is known to

The phytosaur of the Triassic exhibits many interesting adaptive modifications of the diapsid skull. The modifications of the muscle attachment areas for the jaw muscles have been examined in comparison with the musculature of the recent forms Sphenodon and Alligator. It has been determined that the development of the fleshy anterior pterygoid and of the intramandibular muscles is an adaptation concomitant with the elongation of the snout. The modification of the temporal region in the phytosaur with the posterior extension of the squamosal is also seen to be an adaptation to gain speed in closing the great mouth. The forward position of the head-neck joint would appear to gain advantage in the support of the heavy head. Such similarities as are seen between the phytosaur and the crocodilian may be accounted for as parallel adaptation.

The red arkosic sandstones of the Cenger Formation, in the allochthonous units of the Lycian Taurus (western Turkey), which had already yielded a small ichthyofauna, have yielded fragmentary remains (teeth, jaw fragments, vertebrae) of archosaurian reptiles. Among these, phytosaur remains can be recognised; they are too incomplete to warrant an accurate identification, but they do provide biostratigraphical data, since they indicate a Late Triassic age. The reptile remains from the Cenger Formation thus confirm the stratigraphical conclusions previously drawn from the study of lungfishes, and increase our knowledge of the geographical distribution of Gondwanan phytosaurs.

A skull of the phytosaur Machaeroprosopus, discovered in the Sloan Canyon formation, confirms the Triassic age of the rocks lying beneath the Exeter (Jurassic) in the valley of the dry Cimarron River of Union County, New Mexico, and Cimarron County, Oklahoma. The underlying red beds, the Sloan Canyon formation, and the Sheep Pen Canyon formation form a continuous and conformable succession of strata in this region, and the conclusion is drawn that these three units should be included in the Dockum group.

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The phylogenetic relationships of the Phytosauria are discussed, and a new species of Angistorhinus is described from Big Spring, Texas.

We describe here new Late Triassic haramiyidan mammaliaform and reptile fossils from near the classic ‘Microlestes’ Quarry’ at Holwell, Somerset, U.K., where Charles Moore discovered a huge collection of microvertebrates in the 1850s. Moore’s discoveries included the haramiyid Thomasia (formerly ‘Microlestes’ and Microcleptes) for which he achieved worldwide fame. Subsequently, despite much fossicking by researchers at Holwell, few new identifiable specimens of mammaliamorphs and lepidosaurs have been recorded and these were by Kuhne in 1939. The new finds described here from a bedded sequence, not from a fissure, add significantly to our knowledge of the Holwell tetrapods and to the Rhaetian terrestrial faunas of the SW U.K. Our discovery of haramiyidan teeth includes a previously unknown type of Theroteinus, a genus not previously recorded from outside of the St-Nicholas-de-Port locality in France. An archosaur tooth, possibly from a phytosaur, is also recorded. The new lepidosaur specimens add significant detail to the recently described ‘basal’ rhynchocephalian Penegephyrosaurus curtiscoppi as well as demonstrating that the global diversity of Lepidosauria in the Late Triassic remains incompletely known.

Trace 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.

Landmark-based Principal Component Analysis was performed on multiple phytosaur and crocodilian skulls. Multiple landmark points were selected defining suture points of bone tissues and regions of ...

Regurgitalites (fossilized regurgitates) can provide insight into the behavioral ecology and physiology of extinct species but they are rarely reported because they are difficult to identify and distinguish from coprolites. A compact mass of skeletal material from the Owl Rock Member of the Upper Triassic Chinle Formation of Arizona reveals features that identify it as a regurgitalite. Characteristics of the teeth and osteoderms in the specimen indicate that these remains belong to the pseudosuchian archosaur Revueltosaurus. Chemical and microstructural analysis revealed a dearth of gastric etching, the preservation of muscle fibers, and the absence of a phosphatic matrix, indicating that this bone mass is a regurgitalite and not a coprolite. It was probably produced by a phytosaur, rauisuchid, or temnospondyl, all of which occur in the Owl Rock Member. We offer an identification key to assist in distinguishing between different types of digestive remains produced by vertebrate carnivores.

Heteropelta boboi is a new archosauriform reptile from the upper Anisian of northeastern Italy represented by a fragment of dorsal armor with a row of neural arches of the dorsal vertebrae. The dorsal armor of the new taxon is composed of two columns of paramedian osteoderms and at least six columns of lateral osteoderms. Unlike other armored archosaurs, the osteoderms are imbricated with the posterior osteoderm overlapping the anterior one. The low neural arches bear small neural spines and long postzygapophyses. The osteoderms of the lateral columns increase in size and change in shape from the most medial to the most lateral columns. Among the Archosauriformes, only the non-archosaur proterochampsians Vancleavea campi, Litorosuchus somnii, and the doswelliids have dorsal armor comprised of more than two columns of osteoderms per side, but the morphology and arrangement of their osteoderms is unlike those of the new Italian taxon. A cladistic analysis of Archosauromorpha positions Heteropelta boboi as either a basal phytosaur or a basal suchian. However, a second cladistic analysis focused on armored archosaurs alternatively positions the new taxon as a basal archosauriform, basal suchian, basal loricatan or crocodylomorph. Better resolution of the phylogenetic relationships of Heteropelta boboi will likely be obtained only with the discovery of cranial and postcranial remains associated with its diagnostic armor elements.

The Late Triassic vertebrate assemblages from Hallau, Switzerland, have been collected since 1915 and include sharks, bony fish (actinopterygians and lungfish), amphibians and archosau- riformes (a phytosaur and the dinosaur Plateosaurus). Many of the microtetrapods, including the haramyid and morganucodontid mammaliamorphs, are well known but the lepidosaurian reptiles have not been figured or described in any detail. We address that gap here reporting four distinct taxa in the 20th century collections made by Bernhard Peyer and emil Schutz. We describe a new basal rhynchocephalian, Deltadectes elvetica gen. et. sp. nov., which is of Gephyrosaurus-type grade rather than a sphenodontian. All the lepidosaurs in the fauna have teeth showing little or no replacement including a suppressed pleurodont and/or pleuracrodont dentition or acrodonty. A Diphydontosaurus- like sphenodontian is present and we record Paleollanosaurus,previously only known from the type locality in Texas U. S. A., together with an acrodont rhynchocephalian of uncertain affinities. The Hal- lau basal rhynchocephalians add further insight into the Norian-Rhaetian lepidosaur faunas found in Europe and North America by providing important new information, particularly to our knowledge of non-sphenodontian rhynchocephalians.

The neotype skull of the Indian phytosaur Parasuchus hislopi Lydekker, 1885 (ISI R42) is re-evaluated and compared with the type material of other basal phytosaurs. Parasuchus hislopi is extremely similar to species previously placed in Paleorhinus (P. bransoni and P. angustifrons), sharing with them such characters as a series of nodes on the lateral surface of the jugal, paired ridges on the squamosal and a frontal depression. Parasuchus hislopi represents a valid species: it can be distinguished from P. bransoni by a relatively low narial eminence and P. angustifrons by the absence of paired nasal depressions. Inclusion of Parasuchus hislopi in a phylogenetic analysis of phytosaurs recovers it in a well-supported clade with P. bransoni and P. angustifrons. Parasuchus is considered the senior synonym of Paleorhinus and Arganarhinus. Parasuchus (here considered to include P. hislopi, P. angustifrons, P. bransoni and P. magnoculus) has a broad circum-Pangaean distribution, with species occurring in the south-western United States, Morocco, central Europe and India. Phytosaur higher-level taxonomy is also revised: Parasuchidae is redefined to include ‘Paleorhinus-grade’ phytosaurs and the later-diverging Mystriosuchinae (the group formerly known as Phytosauridae), and Pseudopalatinae is renamed Mystriosuchini for reason of priority.

A rostrum fragment from the marginal marine upper Lunz Formation of Austria (early Late Triassic: late Carnian) was previously identified as a new species of phytosaurian archosauriform, ‘Francosuchus’ trauthi. ‘Francosuchus’ trauthi was subsequently synonymized with the non-phytosaurid phytosaur Paleorhinus, and this synonymy was used as evidence to correlate the ‘Paleorhinus biochron’ and the Otischalkian land-vertebrate faunachron to the marine timescale. Here, I provide a redescription of ‘Francosuchus’ trauthi and document anatomical features that differ substantially from all known species of non-phytosaurid phytosaur. There is no evidence to support synonymy of ‘Francosuchus’ trauthi with Paleorhinus, and no unambiguous features to support a phytosaurian identification. However, ‘Francosuchus’ trauthi possesses a unique combination of characters that distinguish it from all other Triassic tetrapods, and necessitates referral of the species to a new genus, Dolerosaurus, gen. nov. Rejection ...

The genus Machaeroprosopus has long been considered invalid because the type specimen of the Late Triassic phytosaur species, M. validus, has been lost. Re-examination of the primary literature regarding the establishment of the Late Triassic phytosaur genus Machaeroprosopus demonstrates that M. buceros is the correct type species, not M. validus. Thus, the genus level name Machaeroprosopus has priority over the genera Pseudopalatus and Arribasuchus and all nominal species should be reassigned. Reassignment of these species to Machaeroprosopus satisfies the requirements of the International Code of Zoological Nomenclature (ICZN) and preserves historical context. The name Pseudopalatinae is retained as the valid clade name for these phytosaurs because its usage falls outside of the ICZN.

The paraphyletic genus ‘Paleorhinus’ is understood currently as a cosmopolitan phytosaur taxon from the Late Triassic. There is no consensus regarding the number of species of ‘Paleorhinus,’ with multiple species and genera synonymised into a single genus or even a single species at various points in its published history. The taxonomy is confounded by historical descriptions without the benefit of comparisons to more recently collected specimens, emphasis on plesiomorphic cranial morphology as diagnostic features of the genus, and lack of cladistic analyses. When included in a recent explicitly cladistic phylogenetic analysis, the holotype of ‘Paleorhinus’ scurriensis (TTU P-00539) was found to be the earliest-branching phytosaur with respect to other North American specimens previously referred to ‘Paleorhinus,’ and is generically distinct from Paleorhinus. ‘Paleorhinus’ scurriensis differs from all known phytosaurs in five unambiguous characters: basitubera widely separated mediolaterally; ridge present on lateral surface of jugal; thickened shelf present along posteroventral edge of expanded pterygoid-quadrate wing; ‘septomaxillae’ separated and excluded from internarial septum; and nasal swelling present posterior to posterior borders of nares. This detailed morphological description of an early-branching phytosaur taxon is a first step towards resolving long-standing issues surrounding specific anatomical features and relationships among early members of the clade.

Phytosaurs are a diverse and morphologically distinctive clade of superficially crocodile-like archosauriforms that had a near global distribution during the Late Triassic. Because their remains are among the most abundant vertebrate remains recovered in many Upper Triassic terrestrial formations, phytosaurs are used extensively in long-range biochronological and biostratigraphic correlations. The biochronologically oldest and earliest branching known phytosaurs include an array of nominal species from the early Late Triassic of the United States, Germany, Poland, Morocco, and India that have been synonymized within the genus Paleorhinus, and subsequently used to define a global ‘Paleorhinus biochron’. However, recent phylogenetic work suggested that the North American species previously referred to Paleorhinus are paraphyletic. Here, we reassess the systematics and anatomy of putative specimens of Paleorhinus from southern Germany. Two well-preserved basal phytosaur skulls from the Blasensandstein (Carnian) of Bavaria form the holotypes of Francosuchus angustifrons and Ebrachosuchus neukami, both of which were synonymized with Paleorhinus by previous workers. We demonstrate that Francosuchus angustifrons shares unique synapomorphies with specimens referred to Paleorhinus bransoni from the Late Triassic of Texas, and thus refer the species to Paleorhinus. By contrast, the longirostrine Ebrachosuchus is highly distinctive in morphology, and our new cladistic analysis of Phytosauria demonstrates that it represents a valid taxon that is more closely related to Phytosauridae than to Paleorhinus. We provide the first autapomorphy-based support for a monophyletic but restricted Paleorhinus (supported by a nodal row on the jugal, and low paired ridges on the squamosal) and confirm that previous broader conceptions of Paleorhinus are likely to be paraphyletic. © 2013 The Linnean Society of London

The vomeronasal (VN) system is a pheromone-processing sensory system of tetrapods. Tetrapods use pheromones to communicate territorial boundaries, reproductive status, sex, and species identity. Presumed impressions of VN bulbs on phytosaur frontals led to a claim that phytosaurs possessed the VN system. However, in extant crocodilians, which lack the VN system, the corresponding impressions are associated not with cerebral tissue but with the ophthalmic nerves. Phytosaur head morphology was not conducive to pheromone collection. The extant phylogenetic bracket suggests that all extinct archosaurs, including phytosaurs, lacked the VN system. Without the pheromonal sense, they would not have used chemical means to convey territorial boundaries, reproductive status, sex, and species identity. Instead, they would have used visual, acoustic, and tactile cues, as in extant archosaurs and other tetrapods in which the VN sense is reduced or absent.

The Upper Triassic Tiki Formation of India is a rich storehouse of varied vertebrate fossil assemblages. So far, there is no information on taphonomic signatures of the Tiki vertebrate assemblages in comparison to that of other fossiliferous horizons of India. We report a new, low diversity, mono-dominant, multitaxic vertebrate accumulation where rhynchosaurs constitute the dominant component. The formation of the rhynchosaur bonebed is attributed to biological aggregation with a hydraulic overprint resulting in a mixed-origin concentration. Other taxa include rauisuchid, phytosaur, small indeterminate archosaur and cynodonts. Taphonomic study based on 617 skeletal specimens of rhynchosaurs collected from nine sites within an area of about 250 × 217 m 2 shows that most of the specimens are disarticulated and disassociated but in close spatial proximity to one another; some are associated specimens and few are articulated. About 13 to 20 individuals of rhynchosaur at different ontogenetic stages are estimated from the specimens collected, suggesting gregarious behavior, possibly herding. These specimens show varying degree of weathering, breakage, encrustation, abrasion and deformation. The bonebed is preserved within the Tiki red mudstone unit and is found in association with paleosol profiles, suggesting prolonged subaerial exposure. Spatial distribution and relative bone frequencies show differential susceptibility of the skeletal specimens to fluvial transport. 55.4% of the collected skeletal specimens belonged to Voorhies Group I, whereas 12.4% and 24% constituted Voorhies Groups II and III respectively, and 8.2% of the collected specimens belonged to the intermediate Groups I and II. It appears that the animals concentrated in the vicinity of the water sources during prolonged period of aridity and died possibly during high seasonal rainfall that resulted in a major flood event. Subsequently, the soft tissues decomposed, and the skeletons suffered prolonged subaerial exposure when the water receded leading to disarticulation and fragmentation followed by minor dispersion by low velocity water currents. This resulted in segregation of skeletal specimens, which were gradually covered by mud deposited during later flooding events. Based on the known flora and fauna, the Tiki Upper Triassic ecosystem is reconstructed for the first time. In the aquatic ecosystem, the metoposaurid labyrinthodonts occupied the top of the food pyramid together with the semi-aquatic parasuchids, which occupied an ecological niche similar to that of the present day crocodilians. The abundance of herbivorous rhynchosaurs at the base with few large and carnivorous rauisuchids and parasuchids at the top suggest a trophic structure similar to that of a modern day terrestrial ecosystem.