ABSTRACT. 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 ®rst 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 con®rms 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 ®ve 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 Ètic contact, absence of a posterior process of the squamosal, and a slit-like posttemporal fenestra. The type species Mystriosuchus planirostris shows, most signi®cantly, 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 exempli®es 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 identi®ed character states help to clarify current inconsistencies in the taxonomy and nomenclature of phytosaurs.KEY WORDS: Triassic, Germany, Phytosauria, Pseudopalatinae, cranial anatomy, phylogeny. PH Y T O S A U R IA is a basal clade of crurotarsans, the line of archosaurs that culminates in modern crocodiles, which mimic crocodiles in both the overall morphology and the supposed semi-aquatic lifestyle. Phytosaurs are entirely restricted to the Late Triassic, but represent the most abundant terrestrial vertebrates in many¯uvial deposits of that age in the Northern Hemisphere. The Stubensandstein of Wu Èrttemberg (Middle Keuper, Norian) is the most proli®c source of phytosaurs in Europe, and specimens from these deposits played a pivotal role in the early history of scienti®c exploration of that group. The ®rst phytosaur taxa that were described are Phytosaurus cylindricodon Jaeger, 1828 and Belodon plieningeri von Meyer, in von Meyer and Plieningeri 1844 from the Stubensandstein, but both are based on inadequate mandibular fragments and isolated teeth and are now unanimously considered nomina dubia (Ballew 1989;Hunt and Lucas 1989;Long and Murry 1995; Hungerbu Èhler 1998a In 1909 and 1911 von Huene described a large phytosaur skull, GPIT 261/001 (for abbreviations,...

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The skull anatomy of a new species of the phytosaur Machaeroprosopus is described for the first time on the basis of two specimens from the Upper Triassic Cooper Canyon Formation of Texas. Additional information is provided by a third specimen referred to Machaeroprosopus sp. A paranasal bone, an additional paired element of the narial region, is identified. Important new data are presented for the braincase, including the morphology of the epipterygoid and presphenoid, an anterior process of the prootic, an anteroventral process of the laterosphenoid, and a parasphenoid process. Machaeroprosopus lottorum n. sp. is characterised by four apomorphies: a supratemporal fenestra closed on the skull roof with bevelled anterior rim, a comparatively short squamosal, a flat and rugose narial rim, and medially extended palatines that come close to form an ossified secondary palate. With respect to the supratemporal fenestra, the supraoccipital–parietal complex and several features of the squamosal, Machaeroprosopus lottorum n. sp. bridges the morphological gap between species previously referred to the genera Pseudopalatus and Redondasaurus. A parsimony analysis of known species of Machaeroprosopus supports the hypothesis that the development of the rostral crest in Machaeroprosopus is a sexually dimorphic feature, and questions the validity of the genus Redondasaurus. Consequently, Redondasaurus is here considered a junior synonym of Machaeroprosopus.

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 underly...