A new traversodontid cynodont from the Santa Maria Formation (Ladinian-Carnian) of southern Brazil, with a phylogenetic analysis of Gondwanan traversodontids

The Traversodontidae is a Triassic family of nonmammalian cynodonts, recorded worldwide, characterized by its bucco-lingually expanded postcanines. A recently discovered fauna in the locality of Santa Cruz do Sul, corresponding to the Santa Maria Formation of the Brazilian Middle/Upper Triassic (Ladinian/Carnian), is made up exclusively of nonmammalian cynodonts with an abundance of traversodontids. A new taxon, Santacruzodon hopsoni gen. et sp. nov . , the most commonly recorded traversodontid cynodont in the fauna, is described here. It is diagnosed by an autapomorphy, a very large posterolabial cusp in the upper postcanine, representing more than half of the labial crest, and a suite of traits present in other traversodontids. Among these features are the descendent flange of the jugal developed as a ball-shaped projection, also known in the Madagascan traversodontid Dadadon isaloi , and the presence of symmetrical incisors with numerous mesial and distal marginal cuspules, feature shared with the North American taxon Arctotraversodon plemmyridon . A phylogenetic analysis of traversodontids from Gondwana, based on a data matrix of 28 characters (mostly dental) and 15 terminals (13 traversodontids, including the new species, plus Diademodon and Trirachodon ) was performed. The new traversodontid appears as the sister taxon of Dadadon . A monophyletic Carnian group composed of Exaeretodon , Menadon and Scalenodontoides , with the Carnian Gomphodontosuchus , and the Ladinian Massetognathus as successive outgroups is also supported. In line with a recent phylogenetic hypothesis, a nonmonophyletic relationship for Scalenodon 'species' from the Anisian Manda beds also results from our analysis. Based on the cladogram obtained, morphological trends of change in the dentition of traversodontids throughout the Triassic are suggested.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139 , 529−545.     

New information on the postcranial skeleton of Massetognathus ochagaviae Barberena, 1981 (Eucynodontia,Traversodontidae), from the Middle Triassic of Southern Brazil

New postcranial remains of Massetognathus ochagaviae are described based on a new specimen collected at the Dinodontosaurus Assemblage Zone from the Middle Triassic of Southern Brazil. Several isolated teeth collected together with the postcranial skeleton allowed a taxonomic assignation of the specimen to M. ochagaviae. Its postcranial morphology is quite similar to Massetognathus pascuali, especially regarding the morphology of the transverse process of presacral vertebrae (i.e. laminar in the anterior most and rounded in the posterior most presacral vertebrae); humerus (i.e. the head is dorsally deflected with a ‘fan-shaped’ distal end a deltopectoral crest abruptly ending at the middle of shaft); and femur (with a bulbous and dorsomedially inclined femoral head, lesser trochanter which abruptly begins near the intertrochanteric fossa and extending up to the middle of the femoral shaft and the medial condyle more pronounced and ventrally projected than the lateral one). On the other hand, the clavicle of M. ochagaviae is less lateromedially elongated than in M. pascuali.     

An unusual new archosauriform from the Middle–Late Triassic of southern Brazil and the monophyly of Doswelliidae

Until now the Doswelliidae was considered a monospecific family including Doswellia kaltenbachi from the Late Triassic of North America. The phylogenetic position of this taxon remained enigmatic until recently, when a sister‐group relationship with the Proterochampsidae was suggested. In the present contribution we describe the new doswelliid species Archeopelta arborensis gen. et sp. nov. from the Middle–Late Triassic of Brazil. A cladistic analysis recovered Archeopelta, Doswellia, and Tarjadia within a monophyletic group of basal archosauriforms, the Doswelliidae. The monophyly of this family is supported by the presence of osteoderm ornamentation that is coarse, incised, and composed of regular pits and the presence of an unornamented anterior articular lamina. Archeopelta is more closely related to Doswellia than to other archosauriforms by the presence of basipterygoid processes anterolaterally orientated, dorsal centra with a convex surface, width of the neural arch plus ribs of the first primordial sacral that are three times the length of the neural arch, and iliac blade laterally deflected, with strongly convex dorsal margin, and a length less than three times its height. The phylogenetic analysis indicates that Doswellidae is the closest large monophyletic entity to Archosauria, which achieved a wide palaeolatitudinal distribution during the late Middle and Late Triassic time span. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 839–871.     

First 'Rauisuchian' archosaur (Pseudosuchia,Loricata) for the Middle Triassic Santacruzodon Assemblage Zone (Santa Maria Supersequence), Rio Grande do Sul State,Brazil

The ‘Rauisuchia’ are a group of Triassic pseudosuchian archosaurs that displayed a near worldwide distribution. In Brazil, their fossils are found only in the Santa Maria Formation (Paraná Basin) of the Rio Grande do Sul State, specifically in the Middle Triassic Dinodontosaurus assemblage zone (AZ) and the Late Triassic Hyperodapedon AZ (Rauisuchus tiradentes). Between these two cenozones is the Santacruzodon AZ (Middle Triassic), whose record was, until now, restricted to non-mammalian cynodonts and the proterochampsian Chanaresuchus bonapartei. Here we present the first occurrence of a rauisuchian archosaur for this cenozone, from the Schoenstatt outcrop, located near the city of Santa Cruz do Sul and propose a new species, based on biostratigraphical evidence and a comparative osteological analysis.     

A Primitive Late Triassic 'ictidosaur' from Rio Grande Do Sul, Brazil

A primitive 'ictidosaur' from lower Norian beds of southern Brazil, Riograndia guaibensis gen. et sp. nov., represented by a fragmentary skull and a lower jaw bearing a complete dentition, shows a more generalized morphology than Chaliminia from the Upper Triassic of Argentina and Pachygenelus Diarthrognathus from the Lower Jurassic of South Africa, Canada and Greenland. The frontal bone borders the orbit, and ventrally contacts the dorsal process of the palatine. The secondary bony palate extends back to the last postcanine. I1 and i2 are reduced, whereas I2-3 and i1 are hypertrophied. Both PC 1–7 and pc 1–7 have blade-like crowns without cingula and with 5–9 small sharp cuspules. The upper postcanine crowns are semicircular in labial view with the cuspules around their margins. The lower postcanine crowns are asymmetrical with most of the cuspules dorsodistally distributed. The possible origin of this peculiar dentition is interpreted as the retention of the juvenile dentition of ancestors. The hypothesis that Riograndia guaibensis and the so-called 'ictidosaurs' might have been derived from gomphodont cynodonts is presented.     

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.     

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 specimen of the Triassic pistosauroid Yunguisaurus,with implications for the origin of Plesiosauria (Reptilia,Sauropterygia)

An adult skeleton of the pistosauroid sauropterygian Yunguisaurus liae reveals a number of morphological features not observed in the holotype, such as the complete morphology of the skull roof, stapes, atlas and axis, ventral view of the postcranium, and nearly complete limbs and tail. Size and morphological differences between the two specimens are mostly regarded as ontogenetic variation, and newly added data did not affect the phylogenetic relationships with other pistosauroids significantly. The number of mesopodia (11 carpals and 8 tarsals) exceeds that known in any other Triassic marine reptiles and does not serve as a precursor of the plesiosaurian pattern with fewer mesopodia of different topology; it demonstrates variability of the limb morphology among the Triassic pistosauroids. The pectoral girdles of Corosaurus, Augustasaurus and Yunguisaurus may indicate early stages of the adaptation towards the plesiosaurian style of paraxial limb movements with ventroposterior power stroke.     

Natural assemblages of Hindeodus conodonts from a Permian–Triassic boundary sequence,Japan

Hindeodus parvus and Hindeodus typicalis occur in a deep‐water chert and claystone section in the Mino Terrane, Japan, which has been identified as a Jurassic accretionary complex. Conodont fossils are preserved as natural assemblages of impression fossils on bedding planes in claystone. In this study, 13 assemblages of Hindeodus species were recognized, comprising at most 13 elements which generally maintain the original composition and structure of an apparatus. We discriminated pairs of carminiscaphate P₁, angulate P₂ and makellate M elements, as well as a single alate S₀ element and two digyrate and four bipennate elements constituting the S array. Although the digyrate and bipennate elements are preserved in the S₂ and S_3–4 positions, respectively, a pair of S₁ elements was not found due to incompleteness in the natural assemblages. The conodont biostratigraphy indicates that the lithological boundary between chert and claystone units in the study section corresponds exactly to the Permian–Triassic boundary.     

Taphonomy and palaeoecology of Late Triassic (Carnian) ammonoid concentrations from the Taurus Mountains,Turkey

The deposits of the Carnian Kas?mlar Formation within the Taurus Platform Units of south‐western Turkey represent an important archive of a Late Triassic ecosystem. New palaeontological information was obtained by analysing the Kasimlarceltites mass occurrence, located within the Kas?mlar Formation and named after the Lower Carnian (Julian) ammonoid genus Kasimlarceltites. This is the dominant taxon (> 94%) within the mass occurrence: nearly 775 million ammonoids and 50 million gastropods were extrapolated for the whole extension (at least 5 km²) of the Kasimlarceltites beds. This calculation is one of the main findings within this study, as it is the first time that such a fossil mass occurrence was quantified. Additionally, orientation measurements of the planispiral ammonoids and the helical gastropods enabled reconstructing the history of the mass occurrence and interpreting the underlying transport mechanisms. Further taphonomic aspects (e.g. biofabric, preservation, bioerosion or genetic classification) as well as comparisons with samples of the same acme zone from different localities near A?a?iyaylabel (AS IV, KA I‐II) point to a two‐phased genetic history. Accordingly, local mass mortality within the Kasimlarceltites fauna due to oxygen fluctuations or methane degassing may have initially led to a primary accumulation. These deposits were then reworked and redeposited basinward by gravity flows to create the present‐day secondary allochthonous concentrations.     

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.     

Re‐evaluation of the ontogeny and reproductive biology of the Triassic fish Saurichthys (Actinopterygii,Saurichthyidae)

Viviparity has evolved independently at least 12 times in ray‐finned fishes. However, the fossil record of actinopterygian viviparity is poor, with only two documented occurrences. Both of these are from the non‐teleost actinopterygian Saurichthys, and include S. curionii and S. macrocephalus from the Middle Triassic Meride Limestone (Monte San Giorgio, Switzerland). Here, we present new data on the reproductive biology of these species, giving unprecedented insights into their life‐history. Based on positional and preservational criteria, six specimens were identified as unambiguously gravid. Embryos were positioned dorsal to the gastrointestinal tract, parallel to the axial skeleton and to each other, in the posterior two‐thirds of the abdominal region. A minimum of 16 embryos are preserved in the most fecund females and, based on the largest preserved embryos and smallest preserved neonates, birth must have occurred at 7–12% of maternal fork length. Embryonic crania and teeth are relatively well‐ossified, however ossification of the parietal region is delayed. In the postcranium, the median scale rows and lepidotrichia are ossified, but not the lateral scale rows. Ossified squamation and gradual allometric growth suggests that neonates did not undergo metamorphosis and were relatively precocial. When considered in a phylogenetic context, neither live birth nor internal fertilization appears to represent the primitive state for saurichthyid fishes.     

Taxonomy of the proterosuchid archosauriforms (Diapsida: Archosauromorpha) from the earliest Triassic of South Africa,and implications for the early archosauriform radiation

Proterosuchidae is one of the first clades of Archosauriformes (archosaurs and closely related species) to appear in the fossil record, with the richest sample of the group coming from the Lystrosaurus Assemblage Zone (earliest Triassic) of South Africa. Four nominal proterosuchid species were described from South Africa during the twentieth century (Proterosuchus fergusi, Chasmatosaurus vanhoepeni, Chasmatosaurus alexanderi and Elaphrosuchus rubidgei), but interpretations of their taxonomy have been widely disparate. The most recent taxonomic revision concluded that P. fergusi is the only valid species and that the other nominal species are junior subjective synonyms of this taxon. This proposal was based on the interpretation that anatomical differences between the nominal species could be explained as a result of ontogenetic changes and/or post‐mortem deformation. The recent discoveries of multiple new South African proterosuchid specimens provide an impetus to revisit their taxonomy. Based upon a comprehensive re‐examination of all known specimens, as well as examination of other proterosuchid taxa in collections worldwide, we conclude that the holotype of Proterosuchus fergusi is undiagnostic. As a result, we propose a neotype (RC 846) for the species. ‘Chasmatosaurus vanhoepeni’ and ‘Elaphrosuchus rubidgei’ are considered subjective junior synonyms of P. fergusi. ‘Chasmatosaurus’ alexanderi is considered a valid species, for which we propose the new combination P. alexanderi comb. nov. A third species, P. goweri sp. nov., is erected on the basis of a single specimen (NMQR 880). All three species recognized here are taxonomically distinct from a previously described archosauriform maxilla from the lower Lystrosaurus AZ. As a result, we recognize a minimum of four archosauriform species following the Permo‐Triassic mass extinction in South Africa. Our results suggest a greater species richness of earliest Triassic archosauriforms than previously appreciated, but that archosauriform morphological disparity remained low and did not expand until the late Early Triassic – early Mid‐Triassic.     

Biostratigraphy and geometric morphometrics of conchostracans (Crustacea,Branchiopoda) from the Late Triassic fissure deposits of Cromhall Quarry,UK

The enigmatic fissure deposits of south‐western England and southern Wales are famous for their unique assemblage of Late Triassic vertebrates, although their age is contentious. While recent studies of palynomorphs have dated some as Rhaetian, their conchostracan (Crustacea, Branchiopoda) assemblages have not been described in detail nor used in biostratigraphy. We find that species determination of British Late Triassic conchostracans requires detailed observations of size, shape and ornamentation. We provide evidence that although Euestheria brodieana is invariably smaller than E. minuta, with some slight differences in carapace ornamentation, the traditional view that they are very similar is upheld. The use of conchostracans as a biostratigraphical tool is here tested by application to the British Triassic fissures at Cromhall quarry where the usual stratigraphical evidence provided by superposition is absent. We find no distinction between conchostracans from bedded Rhaetian deposits of the UK and specimens collected from the fissure deposits of Cromhall Quarry, Gloucestershire, supporting a late Rhaetian age for these deposits.     

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.     

Evidence of osteomyelitis in the dentary of the late Triassic rhynchocephalian Clevosaurus brasiliensis (Lepidosauria: Rhynchocephalia) from southern Brazil and behavioural implications

Clevosaurus was a cosmopolitan Rhynchocephalia genus restricted to the Late Triassic and the Early Jurassic. In Brazil, C. brasiliensis is one of the most conspicuous species collected from the Candelária Sequence (Riograndia Assemblage Zone, Norian age) of the Santa Maria Supersequence. Several jaws of C. brasiliensis are housed in the Laboratório de Paleontologia de Vertebrados of the Instituto de Geociências-UFRGS. Some of these jaws bear a relatively small protuberant bony callus on the anterolateral margin of the dentary, evidenced by a different tissue pattern incorporating small pits and discrete grooves. This pattern closely resembles a common bone infection known in the mandible of the extant Sphenodon punctatus. Although this similarity, the infection may be the result of two possible processes: as consequence of orthal jaw shearing movements during feeding at the moments that the dentary impacts with the enlarged premaxillary tooth or due to injuries produced after fights between conspecific individuals (as is the case for S. punctatus). If the second hypothesis is correct, the same pathological processes probably occurred in the Late Triassic C. brasiliensis indicating that similar ethological conditions were already present at the beginning of the Mesozoic, during the initial radiation of the lepidosaurian clade.