The smallest of the largest: a new look at possible dwarfing in sauropod dinosaurs

Franz Baron Nopcsa recognised that the Late Cretaceous titanosaurid sauropod from Transylvania, Magyarosaurus dacus, was much smaller than the better known sauropods from the Upper Jurassic of the United States and even from Late Cretaceous deposits elsewhere in the world. In keeping with his biogeographic interpretations, Nopcsa viewed this difference as a consequence of body-size effects via island habitation. We present a preliminary re-evaluation of Nopcsa's claims and their heterochronic consequences using two approaches, viz. (1) regression analysis of humeral data as a means of establishing patterns in body size among titanosaurids, and (2) optimisation of humeral data onto titanosaurid cladograms to evaluate evolutionary trends within the clade. Our regression analysis is based on twenty species distributed among fourteen genera of neosauropods and uses length and mid-shaft mediolateral width of the humerus of presumed fully adult forms, and of growth samples that consist of postnatal to adult individuals. Linear regression analysis suggests that, among adult neosauropods, M. dacus appears to be represented by the smallest individuals; that M. dacus humeri appear to be more similar to those of subadults than to adults of other taxa; and that this juvenile morphology may constitute dwarfing in M. dacus by paedomorphosis. In order for these regressions to reflect their evolutionary context more fully, we also present optimisation analyses of humeral form within Titanosauroidea. Although many aspects of the phylogeny of this clade are relatively poorly resolved, preliminary results of our analyses are consistent with the conclusion that M. dacus was a heterochronic dwarf.     

记河南汝阳白垩纪一新的巨型晰脚类恐龙

记述了产自河南汝阳地区晚白垩世早期莽川组一新的巨型蜥脚类恐龙--巨型汝阳龙(新属新种):Ruyangosaurus giganteus gen.et sp.nov..它具有以下特征:神经棘低,缺少椎前关节突隔板,神经弓的侧面具有大的、不规则的三角形凹,前关节横突隔板前后伸展,胫骨粗壮,达127 cm等.汝阳龙的发现显示在晚白垩世早期蜥脚类恐龙发生的分异程度要比以前想象的高得多.     

记河南汝阳白垩纪一新的巨型蜥脚类恐龙

记述了产自河南汝阳地区晚白垩世早期莽川组一新的巨型蜥脚类恐龙——巨型汝阳龙（新属新种）：Ruyangosaurus giganteus gen. et sp. nov.。它具有以下特征：神经棘低,缺少椎前关节突隔板,神经弓的侧面具有大的、不规则的三角形凹,前关节横突隔板前后伸展,胫骨粗壮,达127cm等。汝阳龙的发现显示在晚白垩世早期蜥脚类恐龙发生的分异程度要比以前想象的高得多。     

浙江白垩纪恐龙化石

浙江白垩系朝川组、中戴组、金华组、赖家组和赤城山组等地层中,已发现了数以千计保存完好的蛋化石,而恐龙骨骼化石却很少而且大多很残破。其中,目前能鉴别的恐龙有暴龙类（tyrannosaurs）、伤齿龙类（troodontids）、盗蛋龙类（oviraptorids）、鸭嘴龙类（hadrosaurs）、慢龙类（segnosaurs）、泰坦巨龙类（titanosaurs）、甲龙类（ankylosaurs）和结节龙类（nodosaurs）等,分布在天台、永康、东阳、丽水、缙云、金华和江山等广大区域,说明当时浙江存在过种类繁多的恐龙动物群。根据化石埋藏和沉积特征,在白垩纪时期浙江有诸多气候炎热、四面环山的大小盆地,恐龙多沿湖泊、河流筑巢生息,季节性的暴雨导致山洪泛滥,淹没了恐龙巢穴,因泥沙的快速掩埋封闭,巢穴里的蛋形成化石后保存较好。而恐龙或逃离,或尸骨被洪水搬运散落,所以、迄今除了已发现的4具不完整的骨架化石外,绝大多数是零散的骨骼。     

Therizinosauroidea (Dinosauria: Theropoda) from the Upper Cretaceous of Uzbekistan

Skeletal remains of indeterminate therizinosauroid dinosaurs are present in the Cenomanian Khodzhakul Formation and common in the Turonian Bissekty Formation of Uzbekistan. At least two taxa are present in the Bissekty Formation based on different frontal and humerus morphotypes. Phylogenetic analysis based on a dataset with 348 morphological characters and including all known therizinosaurian taxa places the Bissekty taxa as derived non-therizinosaurid therizinosauroids. The Bissekty therizinosauroids are more derived than Alxasaurus elesitaiensis in the extensive pneumatization of the postcranial axial skeleton, the absence of teeth in the anterior portion of the dentary, the weak development or even absence of pits for the collateral ligaments on the manual phalanges, and the subtriangular medial aspect of the distal end of the humerus, with the entepicondyle positioned well medial to the ulnar condyle. They are less derived than Therizinosauridae in the presence of a basisphenoid recess, the absence of pneumatic openings on the anterior caudal vertebrae, the distal end of metacarpal III not being ginglymoid in dorsal view, and the separation of the femoral head from the neck of that bone by a raised ventral rim.     

Evidence of iguanodontian dinosaurs from the Lower Cretaceous of Tunisia

The fossil record of ornithischian dinosaurs from Africa is particularly scarce and limited to a few historic localities. In this study we describe new ornithischian remains from the Albian deposits of southern Tunisia (Tataouine Governorate), represented by isolated teeth of large-bodied iguanodontians. Teeth display a wide, diamond-shaped crown with a primary ridge dividing the occlusal surface in two unequal parts and two or more secondary ridges. Hook-like denticles are present on both mesial and distal crown margins and do not display mammillae. In overall morphology, specimens are comparable to those of many Early Cretaceous basal hadrosauriforms, including isolated ornithopodan teeth from comparably-aged levels of Niger. Transversal sections of the crowns permitted identification of dental tissues, which include a thick enamel, and well developed longitudinal and transverse giant tubules. Their relative extents appear to be related to the size, thus developmental age, of the tooth. Teeth are representative of the Oum ed Diab Member, a unit characterized by coastal deposits accumulated under arid to xeric climatic conditions and dominated by fish, crocodilians, and hydraulically transported rebbachisaurid and spinosaurid remains. Sedimentological data and preservation bias strongly support selective taphonomic causes for the fossil distribution of ornithischians in southern Tunisia questioning the purported geographic and paleoecologic distribution of isolated Saharan dinosaurs.     

山东晚白垩世一新的巨龙类恐龙

记述了产自山东诸城上白垩统王氏群中一个较完整的蜥脚类恐龙肱骨(ZJZ-57)。它具有一些进步特征,如肱骨三角胸嵴向远端扩展,肱骨近端外侧1/3处发育隆凸,表明其属于较进步的巨龙类成员。另外,它具有一些独特的鉴定特征组合区别于其他任何巨龙类:近端横向强烈扩展,近端最大宽度与骨干长度之比为0.55;骨干粗壮,粗壮指数为0.39。综合研究表明,ZJZ-57所属个体代表了一种新的巨龙类恐龙:臧家庄诸城巨龙(Zhuchengtitan zangjiazhuangensis gen.et sp.nov.),它可能与Opisthocoelicaudia的亲缘关系较接近。这是山东晚白垩世地层中首次发现的巨龙类恐龙。     

First sauropod bones from Italy offer new insights on the radiation of Titanosauria between Africa and Europe

Here we describe the first sauropod skeletal remains from the Italian peninsula that also represent the earliest record of titanosaurs in Southern Europe. Scattered bones, including an almost complete anterior caudal vertebra, were found in Cretaceous (Aptian–Albian) marine deposits, some 50 km East of Rome. The vertebra shows a bizarre and perhaps unique orientation of the zygapophyseal articular facets that renders their interpretation problematic. Phylogenetic retrofitting tests support the placement of the Italian titanosaur among basal lithostrotians. Palaeobiogeographic analysis based on the resulting phyletic relationships suggests an Afro-Eurasian route for the ancestors of the Italian titanosaur, a scenario compatible with the palaeogeographic evolution of the Italian microplates during the Cretaceous. Together with previously recorded titanosaurian-like ichnites from a Cenomanian locality in Latium, this new find suggests a quite long emersion for the Apenninic carbonate platform. We suggest that the Italian titanosaur was member of a population that crossed the western Tethys Sea through a “filtering bridge” composed of a chain of ephemeral islands and peninsulae, known as Periadriatic (Adria) carbonate platforms, that connected sporadically Africa and Europe since the Early Cretaceous.     

Troodontidae (Dinosauria: Theropoda) from the Upper Cretaceous of Uzbekistan

To date three taxa of troodontid theropod dinosaurs have been recognized from Upper Cretaceous strata in two regions of the Kyzylkum Desert in Uzbekistan. The Cenomanian Khodzhakul Formation in the southwestern Kyzylkum Desert has yielded isolated serrated teeth and some postcranial bones of an indeterminate troodontid. In the central Kyzylkum Desert troodontids are known from the Cenomanian Dzharakuduk Formation (Urbacodon itemirensis) and the Turonian Bissekty Formation (Urbacodon sp.). Urbacodon itemirensis is known from a single dentary whereas Urbacodon sp. is represented by isolated teeth, maxilla and dentary fragments, a partial braincase, and some postcranial bones. The troodontid affinities of Urbacodon are supported by several synapomorphies: presence of a subotic recess; reduced basal tubera placed directly under the occipital condyle; maxilla participating in the margin of the external naris; nutrient foramina on dentary situated within a deep lateral groove; dentary without distinct interdental plates; large number of small dentary and maxillary teeth; teeth constricted between root and crown; anterior dentary teeth smaller, more numerous, more closely spaced than those in the middle of the tooth row, and implanted in a groove; posterior dorsal vertebrae with tall and posterodorsally tapering neural spines; and presence of a midline sulcus on the neural arches of distal caudals. Among Troodontidae, Urbacodon resembles Byronosaurus, Gobivenator, and Xixiasaurus in the absence of serrations on the tooth crowns and having premaxillary teeth that are D-shaped in cross-section. However, phylogenetic analysis did not recover a clade of Asiatic troodontids with unserrated teeth.     

Multiple neoplasms in a single sauropod dinosaur from the Upper Cretaceous of Brazil

Tumors are thought rare in dinosaurs, previously limited in distribution to a single family – hadrosaurs. Here we recognized two different neoplastic lesions, osteoma and hemangioma, in a single caudal vertebra of a titanosaur dinosaur from the Upper Cretaceous of Brazil, diagnosed on the basis of macroscopic, radiological and histological analysis. This is the first occurrence of neoplasia in a non-hadrosaur dinosaur. Therefore, neoplasia is not restricted to hadrosaur dinosaurs, as previously suggested.     

Sauropod Teeth from the Lower Cretaceous Luohandong Formation of Ordos Basin, Inner Mongolia

The Early Cretaceous Zhidan Group in the northern Ordos Basin, Inner Mongolia, yielded a large number of tetrapods, including turtles, choristoderes, crocodyliforms, psittacosaurs, stegosaurs, theropods and birds. Well-preserved sauropod teeth have been found in the Luohandong Formation, a middle-upper unit of the Zhidan Group. The large V-shaped wear facet, low slenderness index value, labial grooves, lingual ridge and concavity on the tooth crown suggest that these teeth are from titanosauriforms. Moreover, the presence of the prominent bosses on the lingual side of the tooth crown indicates these teeth should be identified as Euhelopus teeth further. The existence of Euhelopus in Ordos Basin (Inner Mongolia), Shandong Province and western Liaoning Province shows some connections about vertebrate faunas during Early Cretaceous in these areas. Other tetrapod groups such as turtles (Sinemys, Ordosemys), choristoderes (Ikechosaurus), psittacosaurs (Psittacosaurus) and birds (Cathayornis) provide more evidences for this viewpoint.     

吉林九台白垩系一新的蜥脚类恐龙

记述了蜥脚类恐龙(Sauropoda)中一新的属种—西地九台龙(Jiutaisaurusxidiensisgen.etsp.nov.),包括18个相关节的尾椎及13个脉弧。化石产于吉林九台白垩纪泉头组的紫色-杂色砂岩中。其主要特征为:中部尾椎椎体两侧出现前后延长的棱脊,前-中部尾椎双凹型,神经弓位于椎体前半部,脉弧不分叉,血管孔长度约占脉弧长度的1/2以上。新属种的发现对进一步研究白垩纪蜥脚类恐龙的系统演化、古地理分布等提供了新的材料。     

Ornithomimidae (Dinosauria: Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan

The stratigraphically oldest remains of ornithomimid theropod dinosaurs are known from the Cenomanian Khodzhakul Formation and the Turonian Bissekty Formation of Uzbekistan. The ornithomimid from the Bissekty Formation is documented by more than 800 isolated bones that represent much of the skeleton. It shows at least three unambiguous synapomorphies of Ornithomimidae: length of anterior cervical centra three to five times greater than transverse width; low and rounded fibular crest of tibia; metatarsal III pinched between metatarsals II and IV; and proximal end of metatarsal III not visible in anterior view. Phylogenetic analysis, based on a dataset with 568 morphological characters and including all known ornithomimosaurian taxa, places the ornithomimid from the Bissekty Formation near the base of the ornithomimid radiation, between Archaeornithomimus asiaticus and Sinornithomimus dongi.     

从浙江天台白垩纪蛋化石复原恐龙类群

浙江天台盆地白垩纪地层中,已发现了甲龙类及慢龙类等恐龙骨骼化石及数以千计的蛋化石,其中许多为恐龙蛋化石。根据国内外相同类型的蛋化石中迄今已发现的恐龙胚胎化石研究成果,可大致确定其中一部分蛋化石分别是由鸭嘴龙类、巨龙类、慢龙类、盗蛋龙类、伤齿龙类和暴龙类等恐龙所产。并发现鸭嘴龙类、巨龙类和慢龙类的圆形蛋、副圆形蛋在数量上占绝大多数,盗蛋龙的长形蛋和伤齿龙的棱柱形蛋较少,暴龙类的巨型长形蛋最少,从而反映了他们各自在生态金字塔中的位置。     

Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia

Patagonia has yielded the most comprehensive fossil record of Cretaceous theropods from Gondwana, consisting of 31 nominal species belonging to singleton taxa and six families: Abelisauridae, Noasauridae, Carcharodontosauridae, Megaraptoridae nov. fam., Alvarezsauridae, and Unenlagiidae. They provide anatomical information that allows improved interpretation of theropods discovered in other regions of Gondwana. Abelisauroids are the best represented theropods in Patagonia. They underwent an evolutionary radiation documented from the Early Cretaceous through to the latest Cretaceous, and are represented by the clades Abelisauridae and Noasauridae. Patagonian carcharodontosaurids are known from three taxa (Tyrannotitan, Giganotosaurus and Mapusaurus), as well as from isolated teeth, collected from Aptian to Cenomanian beds. These allosauroids constituted the top predators during the mid-Cretaceous, during which gigantic titanosaur sauropods were the largest herbivores. Megaraptorans have become better documented in recent years with the discovery of more complete remains. Megaraptor, Aerosteon and Orkoraptor have been described from Cretaceous beds from Argentina, and these taxa exhibit close relationships with the Aptian genera Australovenator, from Australia, and Fukuiraptor, from Japan. The Gondwanan megaraptorans are gathered into the new family Megaraptoridae, and the Asiatic Fukuiraptor is recovered as the immediate sister taxon of this clade. Although megaraptorans have been recently interpreted as members of Allosauroidea, we present evidence that they are deeply nested within Coelurosauria. Moreover, anatomical information supports Megaraptora as more closely related to the Asiamerican Tyrannosauridae than thought. Megaraptorans improve our knowledge about the scarcely documented basal radiation of Gondwanan coelurosaurs and tyrannosauroids as a whole. Information at hand indicates that South America was a cradle for the evolutionary radiation for different coelurosaurian lineages, including some basal forms (e.g., Bicentenaria, Aniksosaurus), megaraptorans, alvarezsaurids less derived than those of Laurasia, and unenlagiids, revealing that Gondwanan coelurosaurs played sharply differing ecological roles, and that they were taxonomically as diverse as in the northern continents. The unenlagiids represent an endemic South American clade that has been recently found to be more closely related to birds than to dromaeosaurid theropods. Analysis of the theropod fossil record from Gondwana shows the highest peak of origination index occurred during the Aptian–Albian and a less intense one in the Campanian time spans. Additionally, peaks of extinction index are recognized for the Cenomanian and Turonian–Coniacian time spans. In comparison, the Laurasian pattern differs from that of Gondwana in the presence of an older extinction event during the Aptian–Albian time-span and a high origination rate during the Cenomanian time-bin. Both Laurasian and Gondwanan theropod records show a peak of origination rates during the Campanian.     

New record of titanosaurian (Dinosauria: Sauropodomorpha) osteoderms from the Upper Cretaceous of North Patagonia

The idea that titanosaurs had osteoderms was proposed in the late nineteenth century by Depéret. However, this idea was given little credence by other researchers until 1980 when unequivocal evidence of armoured titanosaurs was reported. Since then, many discoveries of titanosaurian osteoderms have been made worldwide. In this work, seven osteoderms are described from the Allen Formation (upper Campanian–lower Maastrichtian), Salitral Moreno locality, Río Negro Province, Argentina. Among the described osteoderms it was possible to recognize three morphologies (keeled, ellipsoidal and cylindrical), with the first of these being most prevalent. Although the osteoderms from Salitral Moreno resemble osteoderms found in other parts of the world, no osteoderms of a similar morphotype are known from this locality, or from Lago Pellegrini or Cinco Saltos.     

Amazonsaurus maranhensis gen. et sp. nov. (Sauropoda, Diplodocoidea) from the Lower Cretaceous (Aptian–Albian) of Brazil

A new genus and species of an Aptian–Albian sauropod from the Itapecuru Formation, northern Brazil, Amazonsaurus maranhensis, is described. It is known from an incomplete, but diagnostic postcranial skeleton. The new taxon possesses several autapomorphies, such as the anterior caudal vertebrae with lateral laminae formed by the coalescence of the spinoprezygapophyseal and postzygodiapophyseal laminae and, to a lesser extent, of the postzygodiapophyseal laminae. It exhibits many synapomorphies supporting its inclusion in the Diplodocoidea. These include high caudal neural arches and anterior caudal neural arches with spinoprezygapophyseal laminae on the lateral aspect of the neural spine. This record is consistent with previous hypotheses on the existence of a community of Afro-South American dinosaurs.     

Description of a juvenile titanosaurian dinosaur from the Upper Cretaceous of Brazil

We describe a juvenile specimen of a titanosaurian sauropod dinosaur consisting of two dorsal and three caudal vertebral centra, an ilium fragment, and an ischium unearthed in 1991 from Site Km 153.5 at BR-050 highway in the Serra da Galga region, municipality of Uberaba, state of Minas Gerais, Brazil. The outcrop at the site is assigned to the Serra da Galga Member of the Marília Formation (Bauru Basin: Bauru Group; Upper Cretaceous: Maastrichtian). Although the material is very incomplete, features such as strongly procoelous caudal centra suggest an affinity with the titanosaurian clade Lithostrotia. The extensive vertebral pneumaticity with deep pleurocoels and well-developed camerae supports the hypothesis that, in titanosaurs, the air sac system was already present and fully developed even at early ontogenetic stages.     

First report of lacertiform (lizard) tracks from the Cretaceous of Asia

The well-preserved trackway of a lacertiform, lizard-like trackmaker from the Haman Formation (Cretaceous) of Korea is described as Neosauroides koreaensis ichnogen. et ichnosp. nov. This is the only example of a Cretaceous lacertiform or lizard-like trackway currently known in the global track record. Although lacertiform trackways, mostly assigned to the ichnogenus Rhynchosauroides, are common in the global Triassic, they are almost entirely absent in the Jurassic and Cretaceous. Moreover, ichnological classification criteria allow that Neosauroides is morphologically distinct from Rhynchosauroides at the genus level, and more like the tracks of the extant lizard Sceloporus. The reasons for the conspicuous lack of post-Triassic occurrences are not certain, but not due to a post-Triassic lack of potential lizard trackmakers. Thus, the preservation biases are likely due to paleobiological factors such as trackmaker ecology and paleoenvironmental preference.     

Diversity of hadrosauroid dinosaurs from the Late Cretaceous Ibero-Armorican Island (European Archipelago) assessed from dentary morphology

A substantial part of the European record of hadrosauroid ornithopod dinosaurs comes from the Maastrichtian (Upper Cretaceous) strata of the south-central Pyrenees (northern Catalunya and Aragón, northeastern Spain). Yet, our understanding on the taxonomy and phylogenetic systematics of these animals in this region is hindered by the lack of complete specimens, paucity of diagnostic material, and poor preservation of the fossils. The Basturs Poble site (Lleida Province, northern Catalunya) has yielded over 900 bones attributed to a hadrosauroid monodominant population. The dentary is the most common, and one of the better-preserved elements found in this locality. Here, we evaluate the utility of dentary morphology as an indicator of hadrosauroid taxonomic diversity. We do so via principal component analysis, cluster analysis, and mean comparison tests of various morphometric data from the available sample of dentaries collected so far in the Basturs Poble locality and other Maastrichtian sites in northeastern Spain. Three different dentary morphotypes are recognized: two different hadrosauroids, one more derived than the other, and one lambeosaurine type. One of the hadrosauroid morphotypes corresponds to relatively small individuals that may represent insular dwarf species. Concerning Basturs Poble locality, our results suggest the coexistence of hadrosauroid and lambeosaurine dinosaurs during the late Maastrichtian on the Ibero-Armorican Island.