Partial hindlimb of Blikanasaurus cromptoni n. gen. and n. sp.,representing a new family of prosauropod dinosaurs from the upper triassic of South Africa

The holotype of the prosauropod dinosaur Blikanasaurus cromptoni n. gen. and n. sp., a partial hindlimb, is described from the lower Elliot Formation (= Red Beds) (upper Carnian or lower Norian; Upper Triassic) of Herschel district, Transkei, South Africa. The new prosauropod family Blikanasauridae is characterized by a hindlimb that is extremely stocky, especially the metatarsus, with the distal tarsals medially situated. Blikanasaurus was an early experiment in the direction of heavily-built, quadrupedal saurischians, but it was not on the evolutionary line that gave rise to the Sauropoda.     

On the anatomy and relationships ofEfraasia diagnostica (Huene) n. gen., a prosauropod dinosaur (Reptilia: Saurischia) from the Upper Triassic of Germany

Palaeosaurus (?) diagnosticus Huene from the Stubensandstone (Upper Triassic, Norian) of Nordwürttemberg, Germany is made the type species of a new genus of prosauropod dinosaurEfraasia, which is referred to the family Anchisauridae. The anatomy ofEfraasia diagnostica is conservative and it was an ideal ancestor for the later and more specializedAnchisaurus polyzelus of North America. The slenderness of the manus and pes ofEfraasia andAnchisaurus is matched amongst prosauropods only by the feet ofGyposaurus capensis andThecodontosaurus antiquus. Staurikosaurus pricei Colbert from the Upper Triassic of Brazil is probably a very primitive theropod dinosaur.     

Are Spondylosoma and Staurikosaurus (Santa Maria Formation, Middle-Upper Triassic, Brazil) the oldest saurischian dinosaurs?

Spondylosoma absconditum (Ladinian, Middle Triassic) lacks certain Dinosauriformes-Dinosauria characters (sigmoid curve and epipophyses in neck, distally placed deltopectoral crest on humerus) and those present (accessary hyposphene-hypantra vertebral articulations, three sacral vertebrae, elongate pubis) also occur in the Rauisuchia (Crurotarsi, Suchia, Rauisuchiformes).Spondylosoma is tentatively referred to the Rauisuchidae because, as indicated by the ventrolaterally directed sacral ribs, the ilium was held obliquely (and not vertically as in the Poposauridae).Staurikosaurus pricei (Carnian, Upper Triassic), one of the oldest dinosaurs, possesses autapomorphies of Dinosauriformes, Dinosauria, Saurischia, and Herrerasauridae, but it retains certain plesiomorphic characters (subcircular distal end to tibia, ?proximally placed deltopectoral crest, axially unshortened neural spine to sacral 1, ?scapular spine unreduced distally) not present inHerrerasaurus (Carnian, Ischigualasto Formation, Argentina). However, the femur is more derived in the loss of the trochanteric shelf and reduction of the lesser trochanter (as also occurs in some prosauropods). The two vertebrae sacrum ofStaurikosaurus is a plesiomorphic character for Dinosauria (also in other herrerasaurids, German Upper Triassic prosauropodSellosaurus gracilis — ?males), to which was added either a caudosacral (e.g. Santa Maria prosauropodSaturnalia, Sellosaurus gracilis — ?females) or a dorsosacral (e.g. Ischigualasto theropodEoraptor and prosauropodRiojasaurus). Agrosaurus (Upper Triassic, Durdham Down, England — not Australia) is not a herrerasaurid; it is referred to the prosauropod dinosaurThecodontosaurus antiquus (Upper Triassic, Durdham Down).     

The skull ofvelociraptor mongoliensis,a small cretaceous theropod dinosaur from Mongolia

The skull of the small theropod dinosaurVelociraptor mongoliensis is described in detail, using the holotype from the Djadokhta Formation (Campanian) of Mongolia. A reconstruction of the skull is attempted.Velociraptor is closely related toDeinonychus andDromaeosaurus; however, it is distinguished from all other dromaeosaurids described by its relatively long, low skull and a depressed muzzle. The affinities ofVelociraptor are discussed and the taxonomic status of all other previously described East Asian dromaeosaurids is examined.     

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.     

Occurrence ofByronia Matthew andSphenothallus Hall in the Lower Cambrian of China

Rare phosphatic tubular fossils from the Lower-Middle Cambrian Kaili Formation of Guizhou Province, southern China were originally identified as non-calcified algae or ‘worms’ (ScoleciellusLiu). Re-examination of these fossils indicates that specimens identified as non-calcified algae areSphenothallus taijiangensis n. sp., while specimens identified asScoleciellus belong toByronia natus (Liu).Sphenothallus taijiangensis andByronia natus from Lower Cambrian strata in the Kaili Formation are the oldest known representatives of their genera. In addition,B. natus (Liu) is the only known species ofByronia with the exception ofB. annulata Matthew (Middle Cambrian, British Columbia). CambrovitusMao et al., a tubular fossil from Middle Cambrian strata in the Kaili Formation, originally was classified as a hyolithid. However, the discovery of a nearly complete specimen possessing an apical attachment disk shows thatCambrovitus, likeByronia andSphenothallus, probably was a thecate cnidarian polyp.     

OnStaurikosaums pricei,an early saurischian dinosaur from the Triassic of Brazil,with notes on the Herrerasauridae and Poposauridae

The anatomy of the saurischian dinosaurStaurikosaums pricei Colbert from the Santa Maria Formation (Middle or Upper Triassic) of Rio do Sul, Brazil is redescribed.Staurikosaurus has 9 (or possibly 10) short cervicals, 15 dorsals and 2 sacral vertebrae, a modified brachyiliac pelvic girdle and an elongate hindlimb with the tibia longer than the femur. The new family Staurikosauridae is erected, the Herrerasauridae is redefined, and both families are considered to be Saurischia incertae sedis. The distal part of the “femur” ofPoposaurus gracilis Mehl (Upper Triassic, North America) is the ventral part of the pubes with a footed distal end but, because some members of the Poposauridae have an imperforate acetabulum, this family is transferred from the Saurischia (Theropoda, Carnosauria) to the Thecodontia (Pseudosuchia).     

Taxonomic and biochronological significance of specimens of the Triassic dicynodontDinodontosaurus Romer 1943 in the Tübingen collection

Two species of the dicynodontDinodontosaurus from the Middle Triassic interval of the Santa Maria Formation in Rio Grande do Sul, southern Brazil,D. tener (von Huene 1935) andD. turpior (von Huene 1935), are based on undiagnostic lectotypes and thus arenomina dubia. The oldest valid, available name for a species ofDinodontosaurus isD. oliveirai Romer 1943, the type species of the genus. (The unused senior subjective synonymDiodontosaurus pedroanum Caldas, 1936 has been suppressed by the International Commission on Zoological Nomenclature).Chanaria platyceps Cox 1968 andDinodontosaurus brevirostris Cox 1968 are junior subjective synonyms ofD. oliveirai Romer 1943.Dinodontosaurus thus is a monospecific genus known from the Santa Maria Formation and the Ischichuca (= Chañares) Formation of northwestern Argentina. This dicynodont and associated tetrapods characterize the Chanarian land-vertebrate faunachron, which is of Middle Triassic age, probably Ladinian.     

A new specimen of Orthosuchus stormbergi (Nash 1968) and a review of the distribution of Southern African Lower Jurassic crocodylomorphs

Orthosuchus stormbergi (Nash 1968) is a rare member of the Lower Jurassic crocodylomorph fauna of Southern Africa. Here we report on a new specimen, consisting of a nearly complete dentary bone, and a previously collected specimen that can confidently be referred to this taxon. The dentary presents several features of utility for generic-level identification of Southern African crocodylomorphs, and furthermore the anatomy of this specimen allows for a modest revision of the generic diagnosis. Furthermore, it provides an opportunity to reassess differences in dentary morphology between basal South African crocodylomorphs with the intention of refining taxonomic identification. Finally, we review the stratigraphic provenance of all of Southern Africa’s known crocodylomorph taxa, and show that most are confined to the uppermost Elliot Formation. Protosuchus, Litargosuchus, Notochampsa, and Orthosuchus have first appearance data in uppermost Elliot Formation. Only Notochampsa has a confirmed last appearance datum in the Lower Clarens Formation. These findings have implications with regards to the true position of the Triassic-Jurassic Boundary, and paleoenvironmental pressures acting on the Crocodylomorpha over the boundary of the Upper Elliot and Lower Clarens Formations.     

Mammalian biochronology of the Paleogene-Neogene boundary at Aktau Mountain, eastern Kazakhstan

At Aktau Mountain in the Ili depression of eastern Kazakstan, fossil mammals that encompass the Paleogene-Neogene boundary occur at three stratigraphic levels. The lowest level is in the lower Kyzylbulak Formation and produces Brontotheriidae and the hyracodontidArdynia and is tentatively assigned a late Eocene (Ergilian) age. The lower part of the overlying Aktau Formation produces fossils of the giant rhinocerosParaceratherium and is tentatively assigned a late Oligocene (Tabenbulukian) age. The upper part of the Aktau Formation yields a fossil mammal assemblage that includesGomphotherium,Stephanocemas, Brachypotherium andLagomeryx. It is clearly of Miocene age, probably late early Miocene (late Burdigalian), a correlative of European Reference Level MN 5 and the late Shanwangian of China. The Paleogene-Neogene boundary at Aktau Mountain thus is in the Aktau Formation.     

Foraminifera from the Silberberg Formation («Lower Oligocene») at Silberberg,near Helmstedt (Germany)

From the Silberberg Quarry, type locality of the Silberberg Formation (Martini &Ritzkowski 1968), some planktonic foraminifera are described:Globigerina officinalis Subbotina, Globigerina ouachitaensisHowe &Wallace,G. praebulloides leroyi Blow &Banner, G.praebulloides occlusa Blow &Banner which taxa are here suggested to form intergrading morphotypes within the range of variation of a population dominated byG. officinalis. In addition,G. cf.danvillensis Howe &Wallace is found. The planktonic fauna, together with some elements in the benthonic fauna, suggests a possible correlation with the Danville Landing Member of the Jackson Formation, Louisiana, traditionally ascribed to the Upper Eocene. There is some similarity with European Uppermost Eocene — Lower Oligocene faunas described by various authors. The Silberberg Formation was suggested to be a Neostratotype for the Latdorfian Stage of the Lower Oligocene byMartini &Ritzkowski (1968). It is suggested that the Silberberg Formation may well be coeval with the Upper Priabonian as proposed byCita 1969.     

First documentation of Llandovery (Silurian) conodont genus Astropentagnathus in China (Langao,Shaanxi Province) and the age of Baiyaya Formation

Age of the Baiyaya Formation in the Ziyang-Langao region of Shaanxi Province remains debatable. Ten samples were taken for conodont biostratigraphical studies from the Tianwancun section. The first discovery of Astropentagnathus in China indicates that the genus has a wider geographical distribution than previously known. Conodont fauna from the Baiyaya Formation suggests that the lower part of the unit corresponds to the Pterospathodus eopennatus Superbiozone, and its upper part to the lower Wenlock (probably mainly to the Kockelella ranuliformis Superbiozone). Diachroneity of the Baiyaya Formation in the region is confirmed by conodont studies from the Tianwancun and Qiaoxi sections. Further studies are required to find out age-variation of the formation more precisely.     

Brachiopods from the Cisuralian–Guadalupian of Darvaz,Tajikistan and implications for Permian stratigraphic correlations

In this paper, we describe the upper Cisuralian Safetdara and Gundara formations of the Darvaz mountains, North Pamir, which were part of the Kunlun Arc, developed along the active Eurasian margin. The Safetdara Formation comprises massive limestones (mainly cyanobacterial, Tubiphytes and Archaeolithoporella boundstones) alternating with well-bedded bioclastic and oncoidal limestones and an interval of recessive shales. The formation crops out above the Chelamchi Formation consisting of turbiditic siltstones and sandstones with bioclastic silty limestones yielding massive limestone olistoliths. The Gundara Formation consists of fine sandstones at the base, followed by well-bedded marly bioclastic, oncoidal and microbial limestones, bearing a rich silicified brachiopod fauna in life-position. Two new taxa have been identified in this association: the cemented coralliform Gundaria insolita n. gen. n. sp. and the pedicle attached Hemileurus politus n. sp. The inferred environmental setting is that of shoal deposits of warm, shallow, high energy, clear marine waters for the Safetdara Formation. The agglutinated microbial reefs to cluster reefs of the Gundara Formation were probably growing in a muddier, quieter and probably slightly deeper setting.The foraminifers of the Brevaxina Zone suggest a Bolorian age for the top of the Chelamchi Formation, the Safetdara Formation and the base of the Gundara Formation. Kungurian conodonts have been found in the lower part of the Safetdara Formation. The biostratigraphic data from the sedimentary succession of North Pamir, integrated with those already obtained from Southeast Pamir, allow to refine the correlations between the Tethyan regional scale and the International Time Scale. In particular, it seems now clear that the Bolorian and the lower part of the Kubergandian correlate to the Kungurian.     

Age of the Silurian Wuxiahe Formation in Langao,Northwest China: New conodont data

A recent study of conodonts from the Wuxiahe Formation (lower to middle part) in the Ziyang-Langao region suggested its age of middle Telychian (Llandovery) to lower Sheinwoodian (Wenlock), contradicted by subsequent graptolite studies indicating an age of late Telychian for the same interval. New samples from the Qiaoxi section for conodonts to re-access the age of the Wuxiahe Formation collected in this study show that the lower to middle part of the formation belongs to the Pterospathodus amorphognathoides amorphognathoides Biozone, suggesting the age of late Telychian; thus, the Llandovery–Wenlock boundary in the section is most probably higher than previously estimated, but its precise position is not determined since the identification of the Wenlock graptolite Cyrtograptus cf. lundgreni in the section is to be further confirmed. Based on the conodont faunas recognized in the Qiaoxi and Tianwancun sections, the base of the Wuxiahe Formation in the Ziyang-Langao region is diachronous, i.e., not higher than the upper Telychian Pterospathodus amorphognathoides amorphognathoides Biozone at Qiaoxi, but not lower than the lower Sheinwoodian Kockelella ranuliformis Biozone at the Tianwancun setion.     

Notes on the Melanorosauridae,a family of large Prosauropod Dinosaurs (Saurischia: Sauropodomorpha)

The femur of the anchisaurid (= plateosaurid) prosauropoddinosaur Euskelosaurus browniiHuxley, 1866 from the lower Elliot Formation (Upper Triassic) of South Africa is sigmoidal in posterior view with the fourth trochanter well removed from the medial edge. The sympatric Melanorosaurus readiHaughton, 1924 is not a junior synonym of Euskelosaurus brownii because the femur is straight in posterior view with the fourth trochanter close to the medial edge. In addition to Melanorosaurus, the Melanorosauridae Huene, 1929 includes Camelotia borealis nov. gen. and nov. sp. from the Upper Triassic of England, Riojasaurus incertusBonaparte, 1969 from the Upper Triassic of Argentina, and possibly some of the posteranial material referred to the? rauisuchian the codontian Sinosaurus triassicusYoung, 1948 from the Lower Jurassic of China.     

Age constraints of the Hungshihyen Formation (Early to Middle Ordovician) on the western margin of the Yangtze Platform,South China: New insights from chitinozoans

The Hungshihyen Formation represents the Lower to Middle Ordovician near-shore siliciclastic deposits on the western margin of the Yangtze Platform, South China. However, its age is still debated, largely because of insufficient studies. Here we documented the Bursachitina maotaiensis–B. qianbeiensis chitinozoan assemblage from the lower part of the Hungshihyen Formation at Songliang, Qiaojia County, eastern Yunnan Province. This assemblage is remarkably similar to an assemblage that roughly correlates to the middle part of the Corymobograptus deflexus graptolite Biozone in northern Guizhou Province, indicating that the Hungshihyen Formation at Songliang is probably of late Floian age and may extend into the early Dapingian. Additionally, a revision on the chitinozoans previously documented from the strata assigned to the same formation in the Wuding area was made according to the latest classification. The revision allows the recognition of the Euconochitina symmetrica–Eremochitina brevis assemblage, and therefore suggests a late Tremadocian age for this interval based on regional and global chitinozoan biostratigraphic correlations. Considering also its lithologic succession, we argue that the chitinozoan-bearing strata in the Wuding area previously assigned to the Hungshihyen Formation correspond most likely to the uppermost part of the underlying Tangchi Formation or the lowermost part of the Hungshihyen Formation. The base of the Hungshihyen Formation appears therefore to be diachronous in eastern Yunnan Province.     

Aeronian (Llandovery, Lower Silurian) palynomorphs and graptolites from the Lipeón Formation, Eastern Cordillera, north-west Argentina

The lower levels of the Lipeón Formation, in the Eastern Cordillera, north-west Argentina, yield a marine-dominated palynomorph assemblage, together with graptolites of mid to late mid Llandovery age (Demirastrites convolutus and probably Stimulograptus sedgwickii zones). The palynomorph assemblage is dominated by acritarchs, but also contains algae and terrestrial cryptospores. Crassiangulina variacornuta, considered a potentially good global biostratigraphical marker for the Upper Llandovery is recovered for the first time from the Silurian of Argentina. The occurrence of this species in strata not younger than late Aeronian, and independently dated by graptolites, indicates an early first appearance for Crassiangulina variacornuta, in the Lipeón Formation, below the Aeronian/Telychian boundary. The lower part of the unit corresponds to a quiet marine environment; thus supporting that Crassiangulina variacornuta is a facies-sensitive acritarch.     

Conodonts from the Permian Jisu Honguer (Zhesi) Formation of Inner Mongolia, China

The Jisu Honguer Formation (“Zhesi Formation”) is a North China marine carbonate unit of Permian age containing a mixed fauna of Tethyan, Boreal and endemic elements. The age of the Jisu Honguer Formation has been thought to range from Artinskian to Kazanian based on previous studies using mostly benthic macrofossils. A typical Mesogondolella aserrata conodont fauna is reported from the lower part of the upper member of the Jisu Honguer Formation in the Ulanqub District, Zhesi area, of northern China. The fauna indicates a Middle Permian (Guadalupian) age, most likely late Wordian to early Capitanian, for the strata yielding conodonts. The whole Jisu Honguer Formation could be assigned to the Wordian to early Capitanian age. The overlying Yihewusu Formation is probably of Capitanian age. There are no Wuchiapingian marine deposits in the Zhesi area. As characteristic of the open sea Guadalupian conodont faunas of the Tethys, smooth Mesogondolella dominate the fauna. The view that all Tethyan Guadalupian conodont faunas consist exclusively of serrated Mesogondolella cannot be confirmed. Three new species are described: Mesogondolella neoprolongata C. - y. Wang, Mesogondolella mandulaensis C. - y. Wang and Wardlawella jisuensis C. - y. Wang.     

Discovery of the Jehol Biota from the Celaomiao region and discussion of the Lower Cretaceous of the Bayingebi Basin,northwestern China

Some typical components of the Jehol Biota, including conchostracans Eosestheria sp., the mayfly Ephemeropsis trisetalis Eichwald, 1864, the aquatic beetle Coptoclava longipoda Ping, 1928, and a fragmentary dragonfly, are reported for the first time from the Bayingebi Formation in the Celaomiao region, western Inner Mongolia, China. This discovery indicates that the middle Upper Member of Bayingebi Formation can be correlated with the upper Yixian and the lower Jiufotang formations in western Liaoning Province. Combining it with the radio-isotopic dating result, we further believed that the Upper Member of Bayingebi Formation could be roughly correlated with the Yixian, Jiufotang, and Shahai formations, and the overlying Suhongtu Formation with the Fuxin Formation in western Liaoning Province.In the major Bayingebi Basin, palaeontological and radio-isotopic dating evidence shows that the Bayingebi Formation has a long depositional history of over 30 Ma: its Upper Member bearing the Jehol Biota and the early Fuxin Biota is probably coeval to the Yixian, Jiufotang and Shahai formations and has a Barremian–early Albian age; its Lower Member may be Berriasian–Hauterivian in age and could be correlated with the upper Tuchengzi, Zhangjiakou, and Dabeigou formations in northern Hebei Province. This suggests that the Bayingebi Formation should be promoted to the stratigraphic rank of group and subdivided into several secondary units (formations). Unlike the previous result, the Yingen Formation is considered across the Lower Cretaceous–Upper Cretaceous boundary and being late Albian–early Turonian in age.