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).     

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.     

Extremely distal fluvial sandstone within the playa system of Arnstadt Formation (Norian,Late Triassic), Central Germany

The Wachsenburg Sandstone of Thuringia (Central Germany) occurs within playa deposits of the Arnstadt Formation (Late Triassic, Norian) and furnishes an example of ephemeral river metamorphosis under dryland conditions. Characterized by high flow regime features, the sand-dominated lithofacies constitution exhibits sedimentation by channel processes under the influence of recurring flash floods. Bearing signatures of subaerial exposures, the fining-upward lithofacies cycles are bound by low-angle lateral accretion elements suggesting deposition in a meandering stream. Channel migration in response to point bar expansion and active bank erosion, led to the development of four laterally shifting point bar events. Unimodal palaeocurrent patterns with low variance and azimuthal dispersion support the point bar origin of the Wachsenburg Sandstone. With reduced water budget under largely semi-arid climate, the river progressively became smaller, highly sinuous and ultimately abandoned. The resulting point bar succession was finally covered with sheet flow deposits of over bank origin. The sandstone was deposited during a period of low base-level when the playa system temporarily fell dry.     

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.     

A new crocodylomorph specimen from the Araripe Basin (Crato Member,Santana Formation), northeastern Brazil

The fossil fauna of the Santana Formation (Early Cretaceous) comprises many distinct taxa, but crocodylomorphs are poorly understood. Here we describe a new specimen (MPSC-R1137) that consists of a complete hind limb found in the Crato Member, the basal section of the Santana Formation. Based on the characteristics of the fibula (e.g., pronounced variation of the shaft width) and length proportions of the femur and tibia, this specimen can be distinguished from Caririsuchus camposi and Araripesuchus gomesii, which are known from the Romualdo Member (the upper lithostratigraphic unit of the Santana Formation). The only crocodylomorph formally described from the Crato Member is Susisuchus anatoceps, whose holotype lacks elements of the hind limb. On the basis of a comparative anatomical study of the hind limb, which shows no similarities between MPSC-R1137 and other crocodylomorphs from the Araripe Basin, we tentatively classify this new specimen as cf. Susisuchus sp., and provide new anatomical information for this rather derived crocodylomorph.      

Upper Triassic (Norian) hypercalcified sponges from the Musandam Peninsula (United Arab Emirates and Oman)

Supercalcified sponges, including sphinctozoans, inozoans, chaetetids, spongiomorphids, occurring in Upper Triassic (Norian-Rhaetian) shallow-marine limestones of Musandam Mountains in United Arab Emirates (UAE), are described. The following taxa were determined: sphinctozoans: Hajarispongia osmani Senowbari-Daryan and Yancey, Nevadathalamia arabica n. sp., Nevadathalamia conica n. sp., Fanthalamia milahaensis n. sp., Iranothalamia incrustans (Boiko), Cinnabaria regularis n. sp.; inozoans: Cavsonella triassica n. sp., Molengraaffia regularis Vinassa de Regny, Peronidella? sp., Circopora cf. caucasica Moiseev, Circopora? sp.; spongiomorphids: Spongiomorpha sp.; chaetetids: Lovcenipora chaetetiformis Vinassa de Regny, Lovcenipora musandamensis n. sp., Lovcenipora sp., chaetetid sponge gen. et sp. indet. The most abundant sponge in the studied material is Nevadathalamia arabica n. sp. The described sponge association of the Arabian shelf (Musandam Mountains) shows close affinity to the sponge association known from age-equivalent terranes in the Panthalassa Ocean (Sonora Mountains in Mexico; Pilot Mountains in Nevada, USA), but is remarkably different from sponge associations in carbonates bordering the Tethys. This difference goes along with the biogeography of wallowaconchid bivalves and is most likely attributed to climatic, palaeogeographic or oceanographic factors.     

A new Hyperodapedon (Archosauromorpha,Rhynchosauria) from the Upper Triassic of India: implications for rhynchosaur phylogeny

A new species of the rhynchosaur genus Hyperodapedon, namely H. tikiensis, is described from well‐preserved skeletal elements that were collected from the Upper Triassic Tiki Formation of India. Hyperodapedon tikiensis is diagnosed on the basis of several cranial and postcranial features including longer than wide basipterygoid process, crest‐shaped maxillary cross section lateral to the main longitudinal groove, deeply excavated neural arches of mid‐dorsal vertebrae, long scapular blade, a pronounced deltopectoral crest, proximal humeral end much broader than distal end, iliac length greater than iliac height, equal pre‐ and postacetabular iliac lengths and circular femoral cross section. Two distinct morphotypes of the maxillary tooth plates can be discerned, which are attributed to ontogenetic variations. A maximum‐parsimony analysis was carried out to show that the order Rhynchosauria is characterized by nine cranial and one postcranial character states. The analysis reveals that Otischalkia elderae is invalid and the basal forms, Howesia and Mesosuchus, are closely related. The Mid‐Triassic genus Ammorhynchus is more derived and forms a sister group to the Late Triassic subfamily Hyperodapedontinae. Isalorhynchus and Teyumbaita are basal to the pandemic genus Hyperodapedon. Twenty‐four characters that are not homoplasious document major patterns of skeletal evolution in rhynchosaurs. From laterally oriented scapula and slender propodials, the postcranial skeleton evolved into a more robust form as is evident from nearly vertical scapula and increase in the robustness of the propodials. Shortening of the femur is noted in the derived Late Triassic forms as exemplified in Hyperodapedon gordoni, Hyperodapedon huxleyi and H. tikiensis.     

Evidence for Prosauropod Dinosaur Gastroliths in the Bull Run Formation (Upper Triassic,Norian) of Virginia

Definitive criteria for distinguishing gastroliths from sedimentary clasts are lacking for many depositional settings, and many reported occurrences of gastroliths either cannot be verified or have been refuted. We discuss four occurrences of gastrolith-like stones (category 6 exoliths) not found within skeletal remains from the Upper Triassic Bull Run Formation of northern Virginia, USA. Despite their lack of obvious skeletal association, the most parsimonious explanation for several characteristics of these stones is their prolonged residence in the gastric mills of large animals. These characteristics include 1) typical gastrolith microscopic surface texture, 2) evidence of pervasive surface wear on many of these stones that has secondarily removed variable amounts of thick weathering rinds typically found on these stones, and 3) a width/length-ratio modal peak for these stones that is more strongly developed than in any population of fluvial or fanglomerate stones of any age found in this region. When taken together, these properties of the stones can be explained most parsimoniously by animal ingestion and gastric-mill abrasion. The size of these stones indicates the animals that swallowed them were large, and the best candidate is a prosauropod dinosaur, possibly an ancestor of the Early Jurassic gastrolith-producing prosauropod Massospondylus or Ammosaurus.

Skeletal evidence for Upper Triassic prosauropods is lacking in the Newark Supergroup basins; footprints (Agrestipus hottoni and Eubrontes isp.) from the Bull Run Formation in the Culpeper basin previously ascribed to prosauropods are now known to be underprints (Brachychirotherium parvum) of an aetosaur and underprints (Kayentapus minor) of a ceratosaur. The absence of prosauropod skeletal remains or footprints in all but the uppermost (upper Rhaetian) Triassic rocks of the Newark Supergroup is puzzling because prosauropod remains are abundant elsewhere in the world in Upper Triassic (Carnian, Norian, and lower Rhaetian) continental strata. The apparent scarcity of prosauropods in Upper Triassic strata of the Newark Supergroup is interpreted as an artifact of ecological partitioning, created by the habitat range and dietary preferences of phytosaurs and by the preservational biases at that time within the lithofacies of the Newark Supergroup basins.     

A new species of basal rhynchosaur (Diapsida: Archosauromorpha) from the early Middle Triassic of South Africa,and the early evolution of Rhynchosauria

Rhynchosauria was an important clade of herbivorous archosauromorph reptiles during the Triassic, with a worldwide distribution. We describe a new genus and species of early rhynchosaur, E ohyosaurus wolvaardti gen. et sp. nov. , from the early Middle Triassic (early Anisian) Cynognathus Assemblage Zone (Subzone B) of the Karoo Supergroup, South Africa. Eohyosaurus wolvaardti is known from a single skull, and is recovered as the sister taxon of Rhynchosauridae in a new phylogenetic analysis. Cynognathus Subzone B has previously yielded the stratigraphically oldest well‐understood rhynchosaur species, Mesosuchus browni and Howesia browni. Eohyosaurus wolvaardti increases the rhynchosaur diversity within this stratigraphical horizon to three species. Intriguingly, all currently confirmed rhynchosaur occurrences from the Early Triassic to earliest Middle Triassic are from South Africa. This may suggest a relatively restricted palaeogeographical distribution for early rhynchosaurs, followed by a global dispersal of rhynchosaurids during the Middle Triassic. © 2015 The Linnean Society of London     

Basal dinosauriform and theropod dinosaurs from the mid–late Norian (Late Triassic) of Poland: implications for Triassic dinosaur evolution and distribution

The rise of dinosaurs during the Triassic is a widely studied evolutionary radiation, but there are still many unanswered questions about early dinosaur evolution and biogeography that are hampered by an unevenly sampled Late Triassic fossil record. Although very common in western North America and parts of South America, dinosaur (and more basal dinosauriform) remains are relatively rare in the Upper Triassic deposits of Europe, making any new discoveries critically important. One of the most diverse dinosauriform assemblages from Europe comes from the Por?ba site in Poland, a recently described locality with exposures of the Zb?szynek Beds, which have a palynomorph assemblage characteristic for the mid–late Norian in the biostratigraphic schemes of the Germanic Basin. Using a synapomorphy‐based approach, we evaluate several isolated dinosauriform specimens from Por?ba. This assemblage includes a silesaurid, a herrerasaurid and remains of another type of theropod (potentially a neotheropod). The Por?ba herrerasaurid is the first record of this rare group of primitive dinosaurs from Europe and one of the youngest records worldwide, whereas the silesaurid is the youngest record of a silesaurid from Europe. These findings indicate that silesaurids persisted alongside true dinosaurs into the mid–late Norian of Europe and that silesaurid–herrerasaurid–neotheropod assemblages (which are also known from the Norian of North America, at low latitudes) were more widespread geographically and latitudinally than previously thought. Silesaurid–herrerasaurid–neotheropod assemblages may have been a common ecological structuring of dinosaurs during their early evolution, and their widespread distribution may indicate weak palaeolatitudinal controls on early dinosaur biogeography during the latest Triassic.     

Prozostrodon brasiliensis,a probainognathian cynodont from the Late Triassic of Brazil: second record and improvements on its dental anatomy

Probainognathian cynodonts are conspicuous elements of the Assemblage Zones of the Triassic Santa Maria Supersequence in southern Brazil. Within this group, the derived clade Prozostrodontia, in which the crown group Mammalia is included, is taxonomic diverse in the Hyperodapedon and Riograndia AZs. We describe here the second known specimen (CAPPA/UFSM 0123) of Prozostrodon brasiliensis, until now only represented by its holotype. CAPPA/UFSM 0123 includes a right dentary with dentition. As in the holotype of P. brasiliensis, it has four lower incisors, pc4 with conspicuous cusp a, and small cusps b, c, and d, pc5-pc6 of ‘triconodont’ type with cusps a > c > b > d, with continuous lingual cingulum bearing up to six small discrete cusps, length of the lower tooth row more than half the length of the dentary, and relatively deep horizontal ramus of the dentary. The new specimen is about 25% smaller than the holotype and there is not a diastema between the canine and postcanine teeth, indicating its juvenile condition. Based on both known specimens of P. brasiliensis, a discussion on tooth replacement is presented, showing that the adult individual has more postcanine tooth morphotypes than the juvenile one.     

A new galesaurid (Therapsida: Cynodontia) from the Lower Triassic of South Africa

A new galesaurid cynodont, Progalesaurus lootsbergensis gen. et sp. nov., is described on the basis of a well-preserved skull, lower jaw, right scapula and left atlantal neural arch. Autapomorphies of Progalesaurus include postcanine teeth bearing numerous mesial and distal accessory cusps that flank a recurved main cusp, a post-temporal fenestra bordered by the squamosal ventrally and a large external naris. Progalesaurus is similar to Galesaurus in possessing a poorly defined masseteric fossa on the dentary, a strongly recurved main cusp of the postcanine dentition, an incomplete secondary palate and a similar basisphenoid-parasphenoid morphology. A cladistic analysis of ten early cynodont genera resolves a monophyletic Galesauridae encompassing Cynosaurus , Progalesaurus and Galesaurus , although support for this clade is weak. Procynosuchus and Dvinia are placed at the base of Cynodontia whereas Thrinaxodon and Platycraniellus are positioned higher, but outside of Eucynodontia. The holotype and only known specimen of Progalesaurus was collected during systematic prospecting of Permo/Triassic boundary strata at New Lootsberg Pass, Graaff-Reinet District, South Africa. The discovery of Progalesaurus increases the number of valid Early Triassic cynodonts to four and sheds light on the tempo of early cynodont diversification after the end-Permian mass extinction.     

A new trirachodontid cynodont from the lower levels of the Burgersdorp Formation (Lower Triassic) of the Beaufort Group, South Africa and the cladistic relationships of Gondwanan gomphodonts

We describe a new trirachodontid cynodont from the base of the Burgersdorp Formation (Subzone A fauna of the Cynognathus Assemblage Zone), of the South African Karoo Basin. Langbergia modisei gen. et sp. nov. is characterized by circular to ovoid in outline upper postcanines and the absence of a maxillary platform lateral to the postcanine series. Apart from the new taxon, we recognize two other species of this family in the Cynognathus Assemblage Zone: Trirachodon berryi and Cricodon metabolus , the latter also represented in the Tanzanian Manda Formation. A phylogenetic analysis of gomphodont cynodonts was conducted using a data matrix of 43 craniodental characters and 18 terminals. Trirachodontidae appears as a monophyletic assemblage, with Langbergia appearing as the sister taxon of Cricodon . The monophyly of trirachodontids is weakly supported, however, with one extra step breaking it. Traversodontid cynodont relationships were also inspected and compared with a recent phylogeny proposed for this group. Considering the resulting phylogeny and the sudden appearance and diversification of Cynognathia representatives, the origin of basal Cynognathia (i.e. Cynognathus , Diademodon , and trirachodontids) is suggested to predate their first appearance in the fossil record.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 147 , 383–413.     

Cyclotosaurus cf. PosthumusFraas (Capitosauridae,Stereospondyli) from the Huai Hin Lat Formation (Upper Triassic), Northeastern Thailand: With a note on capitosaurid biogeography

Part of a large capitosaurid skull, similar to that of Cyclotosaurus posthumus from the Upper Triassic of Germany, has been discovered in the upper part of the Huai Hin Lat Formation near Chulabhorn (Nam Phrom) Dam. This discovery is consistent with the presumed Norian age of this formation. Although the phylogeny of the Capitosauridae is still unclear, the group of Upper Triassic Cyclotosaurus species to which C. posthumus belongs is monophyletic and seems to be known only from Laurasia or Northwestern Gondwana (Morocco). The occurrence of C. cf. posthumus in Thailand is consistent with the hypothesis previously put forward, that this part of Southeast Asia was bound to Laurasia in Mesozoic times.     

Osteology of Rauisuchus tiradentes from the Late Triassic (Carnian) Santa Maria Formation of Brazil,and its implications for rauisuchid anatomy and phylogeny

As the eponymous type species for rauisuchians, Rauisuchus tiradentes von Huene, 1938 represents an important but inadequately known specimen. The revision of the type material from the Upper Triassic (Carnian) Santa Maria Formation of Brazil reveals new anatomical data and previously unrecognised characters, including three new possible autapomorphies: a knob‐like thickening on the base of the posterior process of the premaxilla; short and ventrally keeled cervicals lacking postzygodiapophyseal laminae; and mid‐caudal vertebrae with an accessory neural spine and a postspinal lamina. Several elements are re‐identified, including a postorbital (originally identified as postfrontal) and a pterygoid (originally identified as a prefrontal), and additional material from a secondary fossil site, originally assigned to R. tiradentes, including a left ilium, is excluded. Based on the recovered information and new morphological data, the systematic position of R. tiradentes is tested in a comparison of two phylogenetic reanalyses. Both analyses differ in respect to the phylogenetic position of R. tiradentes, and recover ‘rauisuchians’ as a paraphyletic assemblage of non‐crocodylomoprh archosaurs. © 2014 The Linnean Society of London     

A new gomphodont cynodont (Traversodontidae) from the Middle–Late Triassic Dinodontosaurus Assemblage Zone of the Santa Maria Supersequence,Brazil

The tetrapod faunas from the terrestrial Middle–Late Triassic basins of Africa and South America are among the richest in the world, especially in non‐mammalian cynodonts. Despite the great abundance of cynodont specimens found in these basins, there are few known taxa that exhibit interbasinal distributions. Here we describe a new species of traversodontid cynodont of the genus Scalenodon from the Triassic Dinodontosaurus Assemblage Zone of the Santa Maria Supersequence, from the state of Rio Grande do Sul, Brazil. Scalenodon ribeiroae sp. nov. is based on a partial skull that possesses a combination of features not observed in any other South American traversodontid: ellipsoid upper postcanines with the transverse crest formed by three cusps, lacking a mesiobuccal accessory cusp, and with lingual cusp projected lingually creating a concave lingual surface on the upper postcanines; the paracanine fossa is positioned medially to the upper canine, and jugal lacks a suborbital process. A phylogenetic analysis places the new taxon in a basal position within the Family Traversodontidae, with the African Scalenodon angustifrons as sister‐taxon. The new specimen of Scalenodon represents the first record of this genus outside of the Manda Beds of Tanzania, and reinforces the biostratigraphical and biogeographical connection between Gondwanan Middle–Late Triassic tetrapod faunas. Although recent advances have been made, our current knowledge of these faunas is limited by the lack of absolute dates for most units and by uncertainties in the taxonomy and stratigraphical provenance of key fossils.     

Two new parrot species (Psittaciformes) from the early Pliocene of Langebaanweg,South Africa,and their palaeoecological implications

Two new parrot species (Psittaciformes) are described from the early Pliocene Varswater Formation at Langebaanweg, South Africa, an area where no parrots currently are found. A coracoid, humeri, ulnae, carpometacarpi, tibiotarsi and tarsometatarsi of at least four individuals are assigned to a new species of lovebird Agapornis. Additional tarsometatarsi of at least five individuals including a nestling are referred to a new genus and species of Psittacinae, a taxon endemic to Africa comprising the extant genera Poicephalus and Psittacus. Both species form the as yet earliest geological record of parrots in Africa and document the early diversification of the taxa Agapornis and Psittacinae. Evidence for parrots in general, and a putative graminivorous species of lovebird in particular, indicates that woodlands as well as grasslands were present at Langebaanweg during the early Pliocene, which is consistent with current hypotheses on the palaeoenvironment at and around this site.     

A new,well-preserved genus and species of fossil Glaphyridae (Coleoptera,Scarabaeoidea) from the Mesozoic Yixian Formation of Inner Mongolia,China

A new genus and species of fossil Glaphyridae, Cretohypna cristata gen. et sp. n., is described and illustrated from the Mesozoic Yixian Formation. This new genus is characterized by the large body; large and strong mandibles; short labrum; elytra without longitudinal carina; and male meso- and possible metatibia apically modified. A list of described fossil glaphyrids of the world is provided. This significant finding broadens the known diversity of Glaphyridae in the Mesozoic China.