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.

Abstract The partial postcrania of Poposaurus gracilis , a bipedal poposauroid convergent with theropod dinosaurs, has been known for nearly a century, but the skull of P. gracilis has proven elusive. P. gracilis is part of a clade of morphologically divergent pseudosuchians (poposauroids) whose members are sometimes bipedal, lack dentition (i.e. beaks) and some have elongated neural spines (i.e. sails). However, the timing and acquisition of these character states is unknown given the uncertainty of the skull morphology of the ‘mid-grade’ poposauroid P. gracilis. Here, we present the first confirmed skull remains of P. gracilis directly associated with diagnostic pelvic elements that overlap with the holotype. The incomplete skeleton (PEFO 34865) from the Chinle Formation of Petrified Forest National Park (Arizona, USA) includes a left maxilla with a large, mediolaterally compressed tooth, left dentary, right prearticular and a partial postcranium. The character states of P. gracilis (bipedal, ‘sail-less’ and toothed) demonstrate that the evolution of bipedalism, the origin/loss of a dorsal ‘sail’ and the shift to an edentulous beak are complex in poposauroids. P. gracilis is widespread in the Upper Triassic formations in the western USA and is restricted temporally prior to the Adamanian–Revueltian faunal turnover during the Norian.