A pterosaurian vertebra from the Upper Cretaceous of the Saratov Region

A dorsal vertebra referred to as Azhdarchidae indet. from the Rybushka Formation (Upper Cretaceous, Lower Campanian) of the Beloe Ozero locality in the Saratov Region is described. Its vertebral centrum has a hypapophysis and, at the base of the neural arch, there is a large pneumatic foramen. The vertebra possibly belongs to Volgadraco bogolubovi Averianov, Arkhangelskii et Pervushov, 2008, described from the Rybushka Formation of the Shirokii Karamysh 2 locality in the Saratov Region.     

The first diplodocid from Asia and its implications for the evolutionary history of sauropod dinosaurs

Abstract: An isolated anterior caudal vertebra from the Qingshan (= Ch'ing shan) Formation (Early Cretaceous) of Shandong Province, China, is redescribed and shown to be an advanced diplodocid sauropod. This specimen possesses several derived character states that are typically observed in advanced diplodocoids or diplodocids, including the following: a mildly procoelous centrum; a deep pit‐like pneumatic fossa immediately below the caudal rib; wing‐ or fan‐shaped caudal ribs; and complex lamination of the neural spine. The neural spine is apomorphically short and the centrum is short relative to its height compared to those of other diplodocids, which, when coupled with the specimen’s unique geographical location and stratigraphical age, suggests that it probably represents a new taxon. This caudal vertebra provides the first convincing evidence that diplodocids were present in Asia, perhaps as a result of the dispersal of neosauropod lineages from Europe and/or North America during the Early Cretaceous. The discovery of a member of the Diplodocidae in the Early Cretaceous also indicates that this clade did not become extinct at the Jurassic/Cretaceous boundary as previously supposed.     

First evidence of an ankylosaur (Dinosauria,Ornithischia) from the Jurassic Qigu Formation (Junggar Basin,NW China) and the early fossil record of Ankylosauria

The first evidence of an ankylosaur from the Late Jurassic Qigu Formation of the southern Junggar Basin (Xinjiang, northwestern China) is described, based on an isolated caudal vertebra that was discovered together with fragmentary remains of other dinosaurs, including stegosaurs, sauropods, and theropods. The caudal vertebra is characterized by the following features: (i) elliptical morphology of the centrum, being wider than high; (ii) short antero-posterior length of the centrum; (iii) pronounced transversely extending ventral groove; (iv) massive transverse process, that is longer than the centrum diameter; (v) transverse process meeting the centrum high at the dorsal half and at a relatively flat angle; (vi) transverse process making a broad contact with the neural arch without forming a proximo-dorsal projection; and (vii) notochordal prominence present in the centre of the anterior articular surface. The study specimen represents only the second record of an ankylosaur from the Jurassic of Asia — aside from the slightly older Tianchisaurus from the early Upper Jurassic Toutunhe Formation, equally from the Junggar Basin. It helps to fill a gap in our knowledge of the early evolution of these armoured dinosaurs. Additionally, this discovery highlights the potential of the southern Junggar Basin to yield a rich vertebrate fauna and thus to provide an important insight into Late Jurassic ecosystems of Central Asia.     

Vertebral pneumatic structures in the Early Cretaceous sauropod dinosaur Pilmatueia faundezi from northwestern Patagonia,Argentina

One of the diagnostic characters of dicraeosaurid sauropods is a reduction of pneumatization of dorsal and caudal vertebrae relative to their Flagellicaudata sister taxon, Diplodocidae. Here, we analyse pneumatic structures in the dicraeosaurid sauropod Pilmatueia faundezi, compare them to those of diplodocoids and report the first record of camerate chambers in a dicraeosaurid. The pneumatic structures are in a posterior cervical centrum (MLL-Pv-002) and consist of lateral pneumatic fossae on the centrum that communicate internally with large camerae. By contrast, Pilmatueia's dorsal and caudal vertebrae (MLL-Pv-005-016) lack pneumatic fossae on the centra, which is consistent with the previously reported reduced pneumaticity in dicraeosaurids. Nevertheless, the base of the neural arch and possibly the base of the bifid neural spines of a posterior dorsal vertebra (MLL-Pv-005) show pneumatic internal chambers. The pneumatic features of the Pilmatueia cervical centrum and dorsal neural arch we describe indicate that the degree of pneumatization is variable within dicraeosaurids.     

Reanalysis of Wupus agilis (Early Cretaceous) of Chongqing,China as a Large Avian Trace: Differentiating between Large Bird and Small Non-Avian Theropod Tracks

Trace fossils provide the only records of Early Cretaceous birds from many parts of the world. The identification of traces from large avian track-makers is made difficult given their overall similarity in size and tridactyly in comparison with traces of small non-avian theropods. Reanalysis of Wupus agilis from the Early Cretaceous (Aptian-Albian) Jiaguan Formation, one of a small but growing number of known avian-pterosaur track assemblages, of southeast China determines that these are the traces of a large avian track-maker, analogous to extant herons. Wupus, originally identified as the trace of a small non-avian theropod track-maker, is therefore similar in both footprint and trackway characteristics to the Early Cretaceous (Albian) large avian trace Limiavipes curriei from western Canada, and Wupus is reassigned to the ichnofamily Limiavipedidae. The reanalysis of Wupus reveals that it and Limiavipes are distinct from similar traces of small to medium-sized non-avian theropods (Irenichnites, Columbosauripus, Magnoavipes) based on their relatively large footprint length to pace length ratio and higher mean footprint splay, and that Wupus shares enough characters with Limiavipes to be reassigned to the ichnofamily Limiavipedidae. The ability to discern traces of large avians from those of small non-avian theropods provides more data on the diversity of Early Cretaceous birds. This analysis reveals that, despite the current lack of body fossils, large wading birds were globally distributed in both Laurasia and Gondwana during the Early Cretaceous.     

Vertebral Pneumaticity in the Ornithomimosaur Archaeornithomimus (Dinosauria: Theropoda) Revealed by Computed Tomography Imaging and Reappraisal of Axial Pneumaticity in Ornithomimosauria

Among extant vertebrates, pneumatization of postcranial bones is unique to birds, with few known exceptions in other groups. Through reduction in bone mass, this feature is thought to benefit flight capacity in modern birds, but its prevalence in non-avian dinosaurs of variable sizes has generated competing hypotheses on the initial adaptive significance of postcranial pneumaticity. To better understand the evolutionary history of postcranial pneumaticity, studies have surveyed its distribution among non-avian dinosaurs. Nevertheless, the degree of pneumaticity in the basal coelurosaurian group Ornithomimosauria remains poorly known, despite their potential to greatly enhance our understanding of the early evolution of pneumatic bones along the lineage leading to birds. Historically, the identification of postcranial pneumaticity in non-avian dinosaurs has been based on examination of external morphology, and few studies thus far have focused on the internal architecture of pneumatic structures inside the bones. Here, we describe the vertebral pneumaticity of the ornithomimosaur Archaeornithomimus with the aid of X-ray computed tomography (CT) imaging. Complementary examination of external and internal osteology reveals (1) highly pneumatized cervical vertebrae with an elaborate configuration of interconnected chambers within the neural arch and the centrum; (2) anterior dorsal vertebrae with pneumatic chambers inside the neural arch; (3) apneumatic sacral vertebrae; and (4) a subset of proximal caudal vertebrae with limited pneumatic invasion into the neural arch. Comparisons with other theropod dinosaurs suggest that ornithomimosaurs primitively exhibited a plesiomorphic theropod condition for axial pneumaticity that was extended among later taxa, such as Archaeornithomimus and large bodied Deinocheirus. This finding corroborates the notion that evolutionary increases in vertebral pneumaticity occurred in parallel among independent lineages of bird-line archosaurs. Beyond providing a comprehensive view of vertebral pneumaticity in a non-avian coelurosaur, this study demonstrates the utility and need of CT imaging for further clarifying the early evolutionary history of postcranial pneumaticity.     

A longirostrine tyrannosauroid from the Early Cretaceous of China

The fossil record of tyrannosauroid theropods is marked by a substantial temporal and morphological gap between small-bodied, Barremian taxa, and extremely large-bodied taxa from the latest Cretaceous. Here we describe a new tyrannosauroid, Xiongguanlong baimoensis n. gen. et sp., from the Aptian–Albian Xinminpu Group of western China that represents a phylogenetic, morphological, and temporal link between these disjunct portions of tyrannosauroid evolutionary history. Xiongguanlong is recovered in our phylogenetic analysis as the sister taxon to Tyrannosauridae plus Appalachiosaurus, and marks the appearance of several tyrannosaurid hallmark features, including a sharp parietal sagittal crest, a boxy basicranium, a quadratojugal with a flaring dorsal process and a flexed caudal edge, premaxillary teeth bearing a median lingual ridge, and an expanded axial neural spine surmounted by distinct processes at its corners. Xiongguanlong is characterized by a narrow and elongate muzzle resembling that of Alioramus. The slender, unornamented nasals of Xiongguanlong are inconsistent with recent hypotheses of correlated progression in tyrannosauroid feeding mechanics, and suggest more complex patterns of character evolution in the integration of feeding adaptations in tyrannosaurids. Body mass estimates for the full-grown holotype specimen of Xiongguanlong fall between those of Late Cretaceous tyrannosaurids and Barremian tyrannosauroids, suggesting that the trend of increasing body size observed in North American Late Cretaceous Tyrannosauridae may extend through the Cretaceous history of Tyrannosauroidea though further phylogenetic work is required to corroborate this.     

The composition of the caudal skeleton of teleosts (Actinopterygil: Osteichthyes)

The diural caudal skeleton of teleostean actinopterygians develops phylogeneticaily and ontogenetically from a polyural skeleton. The reduction of the polyural anlage to four, three, two or fewer centra in the adult caudal skeleton takes different pathways in different genera (e.g. compare Elops and Albula) and groups of teleosts. As a result, ural centra are not homologous throughout the teleosts. By numbering the ural centra in a homocercal tail in polyural fashion, one can demonstrate these and the following differences. The ventral elements (hypurals) always occur in sequential series, whereas the dorsal elements (epurals and uroneurals) may alter like the ural centra. The number of epurals, five or four in fossil primitive teleosts, is reduced in other primitive and advanced teleosts, but the same epurals are not always lost. The number of uroneurals, seven in fossil teleosts, is reduced in living teleosts, but it has not been demonstrated that the first uroneural is always derived from the neural arch of the same ural centrum. The landmark in the homocercal tail is the preural centrum I which can be identified by (1) bifurcation of the caudal artery and vein in its ventral element, the parhypural, (2) its position directly caudal to the preural centrum (PU2) which supports the lowermost principal caudal ray with its haemal spine, (3) carrying the third hypaxial element ventral to the course of arteria and vena pinnalis, and (4) by carrying the first haemal spine (parhypural) below the dorsal end of the ventral cartilage plate. The study of the development of the vertebral column reveals that teleosts have different patterns of centrum formation. A vertebral centrum is a complete or partial ring of mineralized, cartilaginous or bony material surrounding at least the lateral sides of the notochord. A vertebral centrum may be formed by arcocentrum alone, or arcocentral arcualia and chordacentrum, or arco-, chorda- and autocentrum, or arcocentral arcualia and autocentrum. This preliminary research demonstrates that a detailed ontogenetic interpretation of the vertebral centra and of the caudal skeleton of different teleosts may be useful tools for further interpretations of teleostean interrelationships.     

A Diplodocid Sauropod Survivor from the Early Cretaceous of South America

Diplodocids are by far the most emblematic sauropod dinosaurs. They are part of Diplodocoidea, a vast clade whose other members are well-known from Jurassic and Cretaceous strata in Africa, Europe, North and South America. However, Diplodocids were never certainly recognized from the Cretaceous or in any other southern land mass besides Africa. Here we report a new sauropod, Leikupal laticauda gen. et sp. nov., from the early Lower Cretaceous (Bajada Colorada Formation) of Neuquén Province, Patagonia, Argentina. This taxon differs from any other sauropod by the presence of anterior caudal transverse process extremely developed with lateroventral expansions reinforced by robust dorsal and ventral bars, very robust centroprezygapophyseal lamina in anterior caudal vertebra and paired pneumatic fossae on the postzygapophyses in anterior-most caudal vertebra. The phylogenetic analyses support its position not only within Diplodocidae but also as a member of Diplodocinae, clustering together with the African form Tornieria, pushing the origin of Diplodocoidea to the Middle Jurassic or even earlier. The new discovery represents the first record of a diplodocid for South America and the stratigraphically youngest record of this clade anywhere.     

The ontogenic development of the vertebrae in some gekkonoid lizards

The ontogeny of amphicoelous vertebrae was studied in Ptyodactylus hasselquistii and Hemidactylus turcicus, and that of procoelous vertebrae, in Sphaerodactylus argus. The embryos were assigned arbitrary stages, drawn to scale, and mostly studied in serial sections. Resegmentation occurs as in all amniotes. A sclerocoel divides each sclerotome into an anterior “presclerotomite” and a denser posterior “postsclerotomite.” Tissue surrounding the intersegmental boundary forms the centrum, which is intersegmental. Tissue around the sclerocoel builds the intervertebral structures, which are midsegmental. In the trunk and neck, postsclerotomites form neural arches, and presclerotomites build zygapophyses. The adult centrum consists of the perichordal primary centrum, plus neural arch bases (= secondary centrum). Between the latter and the arch proper, a neurocentral suture persists until obliterated in maturity. A dorso-ventral central canal persists on either side of the primary centrum, between the latter and the secondary centrum. The notochord becomes true cartilage midvertebrally in all vertebrae, and elastic cartilage intervertebrally in the posterior caudal region. Elsewhere its characteristic tissue persists. Intervertebrally, cervical hypapophyses, caudal chevrons and chevron-bases in the trunk are preformed early in cartilage. Directly ossifying median intercentra are added later in all regions. The first cervical presclerotomite is absent: the hypapophysis (= corpus) of the atlas consists exclusively of postsclerotomitic tissue, there is no proatlas, and the odontoid lacks the apical half-centrum present in other lepidosaurians. In the autotomous caudal region presclerotomites are as prominent as postsclerotomites. Both build neural arches, the two arches of each vertebra remaining distinct and ossifying separately, so that the intersegmental autotomy split persists between them. The last sclerotome is complete, its postsclerotomite forming a half centrum which ossifies. In Sphaerodactylus, while the vertebrae ossify, each intervertebral ring becomes concave anteriorly, convex posteriorly; it remains as a cushion between the condyle and a facet formed by differential growth of the centra. Thus these procoelous centra resemble the amphicoelous centra of Ptyodactylus and Hemidactylus, rather than the procoelus centra of other squamates. The vertebral column of Gekkonoidea closely resembles in its development and microscopical structure that of Sphenodon.     

Reevaluation of the caudal skeleton of some actinopterygian fishes: II. Hiodon,Elops, and Albula

The vertebral centra of Hiodon, Elops, and Albula are direct perichordal ossifications (autocentra) which enclose the arcocentra as in Amia. An inner ring of ovoid cells forms in late ontogeny from the intervertebral space inside the autocentrum. The chordacentrum is reduced or completely absent in centra of adult Elops, whereas it forms an important portion of the centra in adult Hiodon. The posterior portion of the compound ural centrum 3+4+5 is partially (Hiodon) or fully formed by the chordacentrum (Elops, Albula). The haemal arches and hypurals are fused medially by cartilage or bone trabecles of the arcocentrum with the centra, even though they appear autogenous in lateral view in Elops and Albula. The composition of the caudal skeleton of fossil teleosts and the ontogeny of that of Hiodon, Elops, and Albula corroborate a one-to-one relationship of ural centra with these dorsal and ventral elements. The first epural (epural 1) of Elops relates to ural centrum 1, whereas the first epural (epural 2) of Hiodon and Albula relates to ural centrum 2. In Albula, the first ural centrum is formed by ural centrum 2 only. With 4 uroneurals Hiodon has the highest number within recent teleosts. Juvenile specimens of Hiodon have eight, the highest number of hypurals within recent teleosts; this is the primitive condition by comparison with other teleosts and pholidophorids. Reduction of elements in the caudal skeleton is an advanced feature as seen within elopomorphs from Elops to Albula. Such reductions and fusions occur in osteoglossomorphs also, but the lack of epurals and uroneurals separates most osteoglossomorphs (except Hiodon) from all other teleosts.     

Additional non-avian theropod and bird remains from the early Late Cretaceous (Santonian) of Hungary and a review of the European abelisauroid record

Hitherto unpublished remains of non-avian and avian theropods from the Late Cretaceous (Formation Csehbánya, Santonian) Iharkút locality (western Hungary) are described. Non-avian theropod remains include an abelisaurid femur, which confirms the presence of this theropod family at Iharkút, and a metacarpal and a tibiotarsus from a paravian which may belong to Pneumatoraptor fodori, previously described from Iharkút. Birds are represented by two femora which clearly belong to enantiornithines, possibly to Bauxitornis, previously described from Iharkút. The abelisauroid record from the Cretaceous of Europe is reviewed.     

Zapalasaurus bonapartei, un nuevo dinosaurio saurópodo de La Formación La Amarga (Cretácico Inferior), noroeste de Patagonia, Provincia de Neuquén, Argentina

An incomplete skeleton from Puesto Morales (Neuquén Province, Argentina) is described as a new species of sauropod, Zapalasaurus bonapartei. The unit that yielded the holotype of this dinosaur is the Piedra Parada Member of the La Amarga Formation, whose age is regarded as Barremian-lower Aptian. Several characters are interpreted as autapomorphies of Zapalasaurus bonapartei: cervical vertebrae with a lamina uniting the prezygapophysis and the zygapophyseal portion of the postzygodiapophyseal lamina, cervical vertebrae with the diapophyseal portion of the postzygodiapophyseal lamina reduced, cervical vertebrae with poorly developed spinoprezygapophyseal laminae, mid and posterior caudal vertebrae with anteroposteriorly elongated neural spines, whose anterodorsal corners are higher than their posterodorsal ones, and caudal centrum length doubles over first 20 vertebrae. Zapalasaurus bonapartei is considered as the sister group of the other diplodocoids (excluding Haplocanthosaurus). Diplodocoids were abundant in the Early Cretaceous, becoming extinct by the early Late Cretaceous. The record of Zapalasaurus bonapartei shows that, at least in the Neuquina Basin, basal diplodocoids were more diverse than previously thought.     

Shape variation in presacral vertebrae of saltasaurine titanosaurs (Dinosauria,Sauropoda)

Titanosaurs were small- to giant-sized sauropods, highly derived and highly pneumatic. Using morphometric analyses, we studied differences in shape of the presacral vertebral centra in some of these sauropods, especially in saltasaurines, and compared asymmetry patterns in lateral pneumatic foramina (LPF) between these titanosaurs and avian and non-avian theropods. Geometric morphometric analyses showed that the cervical centra tend to be elongated and dorsoventrally short, with an elliptical LPF located in the middle of the centrum; dorsal centra tend to be short and higher than the cervical centra, with the LPF slightly displaced to the anterior region. Shape variation can be described as a result of the ordering of the vertebrae within both the cervical and dorsal sequences, and therefore these methods can be applied to predict the position of isolated vertebrae. A persistent pattern of asymmetry among LPF was observed when length–height indexes were plotted. The right LPF are usually larger than those on the left side in the cervical vertebrae (except in Saltasaurus loricatus) but variable in the dorsal vertebrae. We propose an explanation of this asymmetry based on the asymmetric arrangement of viscera and late development of the respiratory (and air sacs) system.     

Ecology,Systematics and Biogeographical Relationships of Dinosaurs,Including a New Theropod,from the Santana Formation (?Albian,Early Cretaceous) of Brazil

Although rare, dinosaurs are well preserved in calcareous nodules of the Santana Formation (Early Cretaceous, ?Albian) of the Araripe Basin, in northeastern Brazil. So far, including only a spinosauroid and three coelurosaurs, the dinosaur fauna appears depauperate. High theropod diversity in assemblages where other dinosaurs are rare or absent is not unique to the Santana Formation. It is seen also in several other assemblages, including Solnhofen and the Maevarano Formation of Madagascar. We consider several factors, including the occurrence of intraguild predation, the possibility that small theropods could subsist in marginal environments, and reliance on coastal resources, that may have been responsible for this apparent ecological imbalance. A new coelurosaur from the Santana Formation, here formally named Mirischia asymmetrica, is shown to be distinct from Santanaraptor placidus [Kellner, A.W.A. () “Short note on a new dinosaur (Theropoda, Coelurosauria) from the Santana Formation (Romualdo Member, Albian) northeastern Brazil”, Boletim do Museu Nacional, Nova Serie, Rio de Janeiro, Brasil 49, 1–8]. Other theropods from the Santana Formation are briefly reviewed. Mirischia is a compsognathid, more similar to the European Compsognathus than to the Asian Sinosauropteryx.     

The caudal skeleton of ostariophysan fishes (Teleostei): Intraspecific variation in trichomycteridae (Siluriformes)

The different elements of the caudal skeleton of the South American catfish genera Nematogenys (Nematogenyinae) and Trichomycterus, Hatcheria, and Bullockia (Pygidiinae) (Siluriformes, Trichomycteridae) show Ontogenetic transformation of the second ural centrum in Trichomycteridae separates the subfamilies Nematogenyinae and Pygidiinae. In the former, the second ural centrum is aligned with the first ural centrum in early stages of ontogeny; it is not fused with the bases of hypurals 3 and 4 in any stage of development. In the Pygidiinae, in contrast, the second ural centrum is connected with the base of hypural 3 from an early stage of development on. One of the most noteworthy features of the Pygidiinae is the epural, a polymorphic element with three or four morphotypes that are species specific. The primitive catfish Nematogenys shows intraspecific variation in the ural centra, segmentation of procurrent caudal rays, and principal caudal ray formulae. Species of Trichomycterus, Hatcheria, and Bullockia are characterized by great intraspecific variability that involves ural centra, the epural, hypurapophyses, and the neural arches of the compound centrum. There is intraspecific variation in the fusion of the hypurals in some species of Trichomycterus. Intraspecific variation of the caudal skeleton of fishes of the family Trichomycteridae involves the presence and frequency of different morphotypes of the epural, neural arch of the compound centrum, fusion of hypurals, and principal caudal ray formulae. Ontogenetic changes of the first and second ural centra, hypurapophyses (with the exception of Nematogenys), and segmentation of procurrent caudal rays (in Nematogenys) are involved also.     

Small theropod teeth from the Upper Jurassic coal mine of Guimarota (Portugal)

Isolated teeth of small theropod dinosaurs from the Upper Jurassic lignite coal mine of Guimarota (near Leiria, Portugal) are described and illustrated. The well known Upper Jurassic theropods from Europe,Archaeopteryx andCompsognathus, are the most common taxa in the Guimarota assemblage. One morphotype is closely related to an allosaurid theropod. Six further morphotypes of theropod teeth are also described, which are closely related to Cretaceous theropods such as dromaeosaurids, troodontids, tyrannosaurids,Richardoestesia andParonychodon. A Late Jurassic origin of these groups of theropods, which is very often postulated, is discussed.     

Vestiges,rudiments and fusion events: the zebrafish caudal fin endoskeleton in an evo‐devo perspective

The vertebral column results from a controlled segmentation process associated with two main structures, the notochord and the somites. Pathological fusion of vertebral bodies can result from impaired segmentation during embryonic development or occur postnatally. Here, we explore the process of formation and subsequent fusion of the caudalmost vertebral bodies in zebrafish, where fusion is a normal process, mechanically required to support the caudal fin. To reveal whether the product of fusion is on an evolutionary or a developmental scale, we analyze the mode of formation of vertebral bodies, identify transitory rudiments, and characterize vestiges that indicate previous fusion events. Based on a series of closely spaced ontogenetic stages of cleared and stained zebrafish, parasagittal sections, and detection methods for elastin and mineral, we conclude that the formation of the urostyle involves four fusion events. Although fusion of preural 1 (PU1⁺) with ural 1 (U1) and fusion within ural 2 (U2⁺) are no longer traceable during centrum formation (phylogenetic fusion), fusion between the compound centrum [PU1⁺+U1] and U2⁺ (ontogenetic fusion) occurs after individualization of the centra. This slow process is the last fusion and perhaps the latest fusion during the evolution of the zebrafish caudal fin endoskeleton. Newly described characters, such as a mineralized subdivision within U2⁺, together with the reinterpretation of known features in an evolutionary–developmental context, strongly suggest that the zebrafish caudal fin endoskeleton is made from more fused vertebral bodies than previously assumed. In addition, these fusion events occur at different developmental levels depending on their evolutionary status, allowing the dissection of fusion processes that have taken place over different evolutionary times.