Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms

Titanosauriforms represent a diverse and globally distributed clade of neosauropod dinosaurs, but their inter‐relationships remain poorly understood. Here we redescribe Lusotitan atalaiensis from the Late Jurassic Lourinhã Formation of Portugal, a taxon previously referred to Brachiosaurus. The lectotype includes cervical, dorsal, and caudal vertebrae, and elements from the forelimb, hindlimb, and pelvic girdle. Lusotitan is a valid taxon and can be diagnosed by six autapomorphies, including the presence of elongate postzygapophyses that project well beyond the posterior margin of the neural arch in anterior‐to‐middle caudal vertebrae. A new phylogenetic analysis, focused on elucidating the evolutionary relationships of basal titanosauriforms, is presented, comprising 63 taxa scored for 279 characters. Many of these characters are heavily revised or novel to our study, and a number of ingroup taxa have never previously been incorporated into a phylogenetic analysis. We treated quantitative characters as discrete and continuous data in two parallel analyses, and explored the effect of implied weighting. Although we recovered monophyletic brachiosaurid and somphospondylan sister clades within Titanosauriformes, their compositions were affected by alternative treatments of quantitative data and, especially, by the weighting of such data. This suggests that the treatment of quantitative data is important and the wrong decisions might lead to incorrect tree topologies. In particular, the diversity of Titanosauria was greatly increased by the use of implied weights. Our results support the generic separation of the contemporaneous taxa Brachiosaurus, Giraffatitan, and Lusotitan, with the latter recovered as either a brachiosaurid or the sister taxon to Titanosauriformes. Although Janenschia was recovered as a basal macronarian, outside Titanosauria, the sympatric Australodocus provides body fossil evidence for the pre‐Cretaceous origin of titanosaurs. We recovered evidence for a sauropod with close affinities to the Chinese taxon Mamenchisaurus in the Late Jurassic Tendaguru beds of Africa, and present new information demonstrating the wider distribution of caudal pneumaticity within Titanosauria. The earliest known titanosauriform body fossils are from the late Oxfordian (Late Jurassic), although trackway evidence indicates a Middle Jurassic origin. Diversity increased throughout the Late Jurassic, and titanosauriforms did not undergo a severe extinction across the Jurassic/Cretaceous boundary, in contrast to diplodocids and non‐neosauropods. Titanosauriform diversity increased in the Barremian and Aptian–Albian as a result of radiations of derived somphospondylans and lithostrotians, respectively, but there was a severe drop (up to 40%) in species numbers at, or near, the Albian/Cenomanian boundary, representing a faunal turnover whereby basal titanosauriforms were replaced by derived titanosaurs, although this transition occurred in a spatiotemporally staggered fashion. © 2013 The Linnean Society of London     

Las cerradicas tracksite (Berriasian,Galve, Spain): Growing evidence for quadrupedal ornithopods

Dinosaur footprints occur in shallow marine sedimentary units of the Berriasian Villar del Arzobispo Formation in the Aliaga basin, NE Spain. Las Cerradicas is a small outcrop (25 m²), near Galve, with four dinosaur trackways. Three tridactyl trackways indicate bipedal animals, and have parallel orientation. A fourth or‐nithopod trackway indicates a quadrupedal animal and is the smallest among any reported in the literature (L = 23 cm, W = 23 cm of pes tracks). It has manus prints that are oval‐shaped impressions. These and other recent discoveries shed much light on the abundance of quadrupedal ornithopods during the early Cretaceous.     

High European sauropod dinosaur diversity during Jurassic–Cretaceous transition in Riodeva (Teruel, Spain)

Abstract:  Up to now, more than 40 dinosaur sites have been found in the latest Jurassic – earliest Cretaceous sedimentary outcrops (Villar del Arzobispo Formation) of Riodeva (Iberian Range, Spain). Those already excavated, as well as other findings, provide a large and diverse number of sauropod remains, suggesting a great diversity for this group in the Iberian Peninsula during this time. Vertebrae and ischial remains from Riodevan site RD-13 are assigned to Turiasaurus riodevensis (a species described in RD-10, Barrihonda site), which is part of the Turiasauria clade. This is the first time that a taxon is attributed to Turiasaurus genus out of its type site. A Neosauropod caudal vertebra from the RD-11 site has been classified as Diplodocinae indet., supporting the previous attribution on an ilion also found in Riodeva (CPT-1074) referring to the Diplodocidae clade. New remains from the RD-28, RD-41 and RD-43 sites, of the same age, among which there are caudal vertebrae, are assigned to Macronaria. New sauropod footprints from the Villar del Arzobispo Formation complete the extraordinary sauropod record coming to light in the area. The inclusion of other sauropods from different contemporaneous exposures in Teruel within the Turiasauria clade adds new evidence of a great diversity of sauropods in Iberia during the Jurassic–Cretaceous transition. Turiasauria distribution contributes to the understanding of European and global palaeobiogeography.     

The early evolution of titanosauriform sauropod dinosaurs

Titanosauriformes was a globally distributed, long‐lived clade of dinosaurs that contains both the largest and smallest known sauropods. These common and diverse megaherbivores evolved a suite of cranial and locomotory specializations perhaps related to their near‐ubiquity in Mesozoic ecosystems. In an effort to understand the phylogenetic relationships of their early (Late Jurassic–Early Cretaceous) members, this paper presents a lower‐level cladistic analysis of basal titanosauriforms in which 25 ingroup and three outgroup taxa were scored for 119 characters. Analysis of these characters resulted in the recovery of three main clades: Brachiosauridae, a cosmopolitan mix of Late Jurassic and Early Cretaceous sauropods, Euhelopodidae, a clade of mid‐Cretaceous East Asian sauropods, and Titanosauria, a large Cretaceous clade made up of mostly Gondwanan genera. Several putative brachiosaurids were instead found to represent non‐titanosauriforms or more derived taxa, and no support for a Laurasia‐wide clade of titanosauriforms was found. This analysis establishes robust synapomorphies for many titanosauriform subclades. A re‐evaluation of the phylogenetic affinities of fragmentary taxa based on these synapomorphies found no body fossil evidence for titanosaurs before the middle Cretaceous (Aptian), in contrast to previous reports of Middle and Late Jurassic forms. Purported titanosaur track‐ways from the Middle Jurassic either indicate a substantial ghost lineage for the group or – more likely – represent non‐titanosaurs. Titanosauriform palaeobiogeographical history is the result of several factors including differential extinction and dispersal. This study provides a foundation for future study of basal titanosauriform phylogeny and the origins of Titanosauria. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 624–671.     

A revision of the sauropod dinosaur genus ‘Bothriospondylus’ with a redescription of the type material of the Middle Jurassic form ‘B. madagascariensis’

Abstract: The sauropod dinosaur ‘Bothriospondylus’, originally named on the basis of Late Jurassic remains from England, is demonstrated to be invalid, and the characters used to diagnose it are shown to be obsolescent features which are widespread throughout Sauropoda. Material referred to this genus spans a temporal range from the Middle Jurassic until the early Late Cretaceous and has been described from five different countries, across three continents. These remains represent a wide array of sauropod groups, comprising non‐neosauropod eusauropods, a macronarian, titanosauriforms (including at least one definite brachiosaurid) and a rebbachisaurid. The type material of the Middle Jurassic ‘B. madagascariensis’ represents a derived non‐neosauropod eusauropod and possesses two potential autapomorphies. However, as a result of the fragmentary nature of the material and the uncertainty surrounding its association, a new taxon is not erected. Of the numerous specimens referred to ‘Bothriospondylus’, however, several remains are considered diagnostic: Ornithopsis hulkei (Early Cretaceous, UK), Lapparentosaurus madagascariensis (Middle Jurassic, Madagascar) and Nopcsaspondylus alarconensis (early Late Cretaceous, Argentina). At least three types of sauropod were present in the Bathonian (Middle Jurassic) of north‐west Madagascar, with a basal eusauropod (Archaeodontosaurus), a more derived eusauropod (‘B. madagascariensis’) and a titanosauriform (Lapparentosaurus) all approximately contemporaneous. Palaeocontinental reconstructions suggest that Middle Jurassic Madagascan sauropods would still have been capable of global biotic interchange, and this is perhaps reflected in their diverse assemblage. Re‐evaluation of these Malagasy forms has shed new light on this important time period in sauropod evolution.     

New record of a sauropod in the Jurassic–Cretaceous transition of the Iberian Peninsular (Spain): palaeobiogeographical implications

In recent decades a unique association of basal neosauropod and turiasaur sauropods has been described from the Jurassic–Cretaceous transition of Spain. In this context, a sauropod femur from the Tithonian–Berriasian is studied for the first time. The femur in question is an isolated specimen, recovered from the Tera Group in Tera (Soria). It displays a unique mosaic of derived characters as yet undescribed in femora of the Upper Jurassic and Lower Cretaceous of Spain. A prominent lateral bulge, high eccentricity, and a lateromedially flattened proximal end link the femur from Tera with Titanosauriformes. Moreover, it presents a significant distal projection of the tibial condyle, a character observed in Asiatic Titanosauriformes of the Lower Cretaceous. The femur from Tera adds a fifth sauropod taxon to the Tithonian–Berriasian of Spain, and, for the first time, a representative of Titanosauriformes.     

Sauropod tracks from the Middle Jurassic Chuanjie Formation of Yunnan Province and the pre-Cretaceous sauropodomorph trackways from China

The Xixipo dinosaur tracksite in the Chuanjie Formation of Yunnan Province is one of the 14 Chinese sites yielding sauropod tracks from between the Triassic–Jurassic and Jurassic–Cretaceous boundaries, but is only one of the two that represent the Middle Jurassic. Although it is a small site, it adds incrementally to the overall track record of the region and allows comparison with the body fossil record and classification of the Chuanjie Formation as a Type 3b or Type 4b deposit in which both the body fossil and trace fossil record, in this case representing sauropodomorphs, are similar in composition and frequency of occurrence. We argue that the sauropod trace and body fossil records, while based on different categories of evidence, are very important. Integrating and correlating all available data from both records increases our understanding of sauropod communities, and both are equally valuable for this. In addition, we also discuss narrow to wide gauge, coeval sauropod trackways from China, and the relationship between the potential trackmaker of China's Jurassic Brontopodus-type trackways and mamenchisaurids and, beginning from the Late Jurassic, representatives of this type and titanosauriform sauropods.     

Body size evolution in Titanosauriformes (Sauropoda,Macronaria)

Titanosauriformes is a conspicuous and diverse group of sauropod dinosaurs that inhabited almost all land masses during Cretaceous times. Besides the diversity of forms, the clade comprises one of the largest land animals found so far, Argentinosaurus, as well as some of the smallest sauropods known to date, Europasaurus and Magyarosaurus. They are therefore good candidates for studies on body size trends such as the Cope's rule, the tendency towards an increase in body size in an evolutionary lineage. We used statistical methods to assess body size changes under both phylogenetic and nonphylogenetic approaches to identify body size trends in Titanosauriformes. Femoral lengths were collected (or estimated from humeral length) from 46 titanosauriform species and used as a proxy for body size. Our findings show that there is no increase or decrease in titanosauriform body size with age along the Cretaceous and that negative changes in body size are more common than positive ones (although not statistically significant) for most of the titanosauriform subclades (e.g. Saltasaridae, Lithostrotia, Titanosauria and Somphospondyli). Therefore, Cope's rule is not supported in titanosauriform evolution. Finally, we also found a trend towards a decrease of titanosauriform mean body size coupled with an increase in body size standard deviation, both supporting an increase in body size variation towards the end of Cretaceous.     

Basal titanosauriform (Dinosauria, Sauropoda) teeth from the Lower Cretaceous Yixian Formation of Liaoning Province, China

The Yixian Formation of Liaoning Province, People's Republic of China has yielded a diverse fauna of non-avian dinosaurs, but is dominated by small-bodied taxa. Here, we describe a series of isolated teeth from the Lujiatun Beds of the formation that are referable to a basal titanosauriform sauropod. Some of the teeth possess a distinctive circular boss on the lingual surface, which suggests that they are referable to cf. Euhelopus sp. This identification provides some additional support for biostratigraphical correlations between the Jehol Group and the Mengyin Formation of Shandong Province that suggest an Early Cretaceous age for the latter unit. Moreover, the titanosauriform affinities of the teeth provide further evidence for the dominance of this sauropod clade in eastern Asia during the Cretaceous.     

Osteology and phylogenetic relationships of Tehuelchesaurus benitezii (Dinosauria,Sauropoda) from the Upper Jurassic of Patagonia

The diversification and early evolution of neosauropod dinosaurs is mainly recorded from the Upper Jurassic of North America, Europe, and Africa. Our understanding of this evolutionary stage is far from complete, especially in the Southern Hemisphere. A partial skeleton of a large sauropod from the Upper Jurassic Cañadón Calcáreo Formation of Patagonia was originally described as a ‘cetiosaurid’ under the name Tehuelchesaurus benitezii. The specimen is here redescribed in detail and the evidence presented indicates that this taxon is indeed a neosauropod, thus representing one of the oldest records of this clade in South America. A complete preparation of the type specimen and detailed analysis of its osteology revealed a great number of features of phylogenetic significance, such as fully opisthocoelous dorsal vertebrae, the persistence of true pleurocoels up to the first sacral vertebra, associated with large camerae in the centrum and supraneural camerae, and an elaborate neural arch lamination, including two apomorphic laminae in the infradiapophyseal fossa. The phylogenetic relationships of this taxon are tested through an extensive cladistic analysis that recovers Tehuelchesaurus as a non‐titanosauriform camarasauromorph, deeply nested within Neosauropoda. Camarasauromorph sauropods were widely distributed in the Late Jurassic, indicating a rapid evolution and diversification of the group. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 605–662.     

A theropod–sauropod track assemblage from the ?Middle–Upper Jurassic Shedian Formation at Shuangbai,Yunnan Province,China, reflecting different sizes of trackmakers: Review and new observations

A dinosaur tracksite at Hemenkou (Shuangbai County, Yunnan Province) in the ?Middle–Upper Jurassic Shedian Formation that consists mainly of gray-purple feldspathic quartz sandstones was previously reported incorrectly as being in the Lower Cretaceous Puchanghe Formation. The previous assignment is also inconsistent with two regional geological maps. Although mostly yielding poorly preserved tracks, the site nevertheless indicates a diversity of theropod and sauropod trackmakers partly consistent with the Late Jurassic body fossils from the region. Purported ornithopod are re-evaluated here as those of theropods. The theropod tracks and trackways show distinct similarities to those of the Grallator–Eubrontes plexus and can be subdivided into three morphotypes that may reflect different pes anatomy and/or substrate conditions. Two sizes of tracks (small, large) indicate the presence of different size classes or species in this area in the Late Jurassic. Similarly, the sauropod trackways document three differently sized trackmakers (small–medium–large) showing a typical wide-gauge (Brontopodus) pattern. The track record is the first evidence of theropods in the ?Middle–Late Jurassic of central Yunnan, whereas the sauropod tracks suggest a relation to the coeval basal eusauropods known from this region by skeletal remains.     

Sauropod dinosaurs from the Lower Cretaceous of eastern Asia: taxonomic and biogeographical implications

Sauropod dinosaurs are poorly represented in the Lower Cretaceous of eastern Asia. Here, we describe a number of isolated sauropod teeth from the Kuwajima Formation (?Berriasian–?Hauterivian) of Shiramine, Japan. The mosaic of shared derived characters and symplesiomorphies displayed by the teeth indicate that they are referable to a basal member of the titanosauriform radiation. A taxonomic review of previously described sauropod specimens from eastern and south–eastern Asia reveals that a diversity of sauropods (including a titanosaurian, a basal titanosauriform and a ?euhelopodid, as well as several forms of indeterminate systematic position) was present in this region in the Early Cretaceous. This diversity conflicts with previous suggestions that eastern Asia was biogeographically isolated from the rest of Laurasia until the late Early Cretaceous and that the sauropod fauna was limited to the endemic East Asian clade Euhelopodidae. The presence of titanosauriform sauropods in the basal Cretaceous of Japan and Thailand indicate that the proposed faunal isolation of eastern Asia ended approximately 20 myr earlier than usually suggested.     

A new sauropodomorph ichnogenus from the Lower Jurassic of Sichuan,China fills a gap in the track record

A dinosaur tracksite in the Lower Jurassic Ziliujing Formation of Sichuan Province, China consists of a spectacular sub-vertical exposure, with multiple track-bearing levels and trackways showing parallel and bimodal orientations. Based on well-preserved material, the new ichnogenus and ichnospecies, Liujianpus shunan ichnogen. nov. ichnosp. nov. is erected to accommodate distinctive sauropodomorph trackways occurring in this assemblage. Liujianpus has a unique combination of features, some relating to the early Jurassic basal sauropodomorph (prosauropod in traditional usage) ichnogenus Otozoum, others to the sauropod ichnogenus Brontopodus. Despite such a mix of basal sauropodomorph- and sauropod-like features, the trackmaker of Liujianpus is likely a basal sauropodomorph. This identification is consistent with the occurrence of basal sauropodomorph skeletons from geographically and chronologically close localities. The other distinct morphotype from the tracksite is linked to a sauropod trackmaker. As such, the ichnofauna consisting of two distinct foot morphotypes reflects the diversity of sauropodomorph dinosaurs in the Early Jurassic of Asia.     

First Jurassic dinosaur tracksite from Guizhou Province,China: morphology,trackmaker and paleoecology

Abstract

Sauropod tracks from the Early Jurassic Maanshan Member of the Ziliujing Formation in the Dazhuanwan area of Guizhou represent the first Jurassic dinosaur track record for this province. The best preserved and longest trackway is narrow gauge (Brontopodus type), and indicates a relatively small trackmaker (footprint length ~35.0 cm). In conjunction with sauropodomorph skeletal remains these tracks suggest that basal sauropodomorphs and primitive sauropods coexisted in this region during the Early Jurassic. This pattern is same in the Lufeng and Sichuan basins. Thus, sauropod tracks from southwest China are diverse in the Early Jurassic, and include narrow gauge Parabrontopodus, wide gauge Brontopodus-type, and basal sauropodomorph tracks.     

A diverse Upper Jurassic dinosaur ichnofauna from central‐west Portugal

Mateus, O. & Milàn, J. 2009: A diverse Upper Jurassic dinosaur ichnofauna from central‐west Portugal. Lethaia, Vol. 43, pp. 245–257. A newly discovered dinosaur track‐assemblage from the Upper Jurassic Lourinhã Formation (Lusitanian Basin, central‐west Portugal), comprises medium‐ to large‐sized sauropod tracks with well‐preserved impressions of soft tissue anatomy, stegosaur tracks and tracks from medium‐ to large‐sized theropods. The 400‐m‐thick Lourinhã Formation consists of mostly aluvial sediments, deposited during the early rifting of the Atlantic Ocean in the Kimmeridgian and Tithonian. The stratigraphic succession shows several shifts between flood‐plain mud and fluvial sands that favour preservation and fossilization of tracks. The studied track‐assemblage is found preserved as natural casts on the underside of a thin bivalve‐rich carbonate bed near the Tithonian–Kimmeridgian boundary. The diversity of the tracks from the new track assemblage is compared with similar faunas from the Upper Jurassic of Asturias, Spain and the Middle Jurassic Yorkshire Coast of England. The Portuguese record of Upper Jurassic dinosaur body fossils show close similarity to the track fauna from the Lourinhã Formation. □Dinosaur tracks, Lusitanian Basin, Portugal, skin impressions, Upper Jurassic.     

A new sauropod dinosaur from the Lower Cretaceous Ilek Formation,Western Siberia,Russia

Sibirotitan astrosacralis nov. gen., nov. sp., is described based on isolated but possibly associated cervical and dorsal vertebrae, sacrum, and previously published pedal elements from the Lower Cretaceous (Barremian?) Ilek Formation at Shestakovo 1 locality (Kemerovo Province, Western Siberia, Russia). Some isolated sauropod teeth from the Shestakovo 1 locality are referred to the same taxon. The phylogenetic parsimony analyses place Sibirotitan astrosacralis nov. gen., nov. sp., as a non-titanosaurian somphospondyl titanosauriform. The new taxon exhibits four titanosauriform and one somphospondylan synapomorphies, and one autapomorphy – a hyposphene ridge that extends between the neural canal and the postzygapophyses. It differs from all other Somphospondyli by having only five sacral vertebrae. The new taxon shares with Euhelopus and Epachtosaurus sacral ribs that converge towards the middle of the sacrum in dorsal view. Sibirotitan astrosacralis nov. gen., nov. sp., is only the second sauropod taxon from Russia and one of the oldest titanosauriform described so far in Asia.     

Large sauropod and theropod tracks from the Middle Jurassic Chuanjie Formation of Lufeng County,Yunnan Province and palaeobiogeography of the Middle Jurassic sauropod tracks from southwestern China

Tracks of large theropods and a single sauropod footprint are reported from red beds at Beikeshan locality in the Middle Jurassic Chuanjie Formation, of Lufeng County, near the large World Dinosaur Valley Park complex. The Chuanjie theropod tracks are assigned to the ichnogenus Eubrontes and the large sauropod track is given the provisional label Brontopodus. All occur as isolated tracks, i.e., trackways are not preserved. Saurischian dominated ichnofaunas are relatively common in the Jurassic of China. The producers of the Chuanjie tracks may have been similar to the basal tetanuran theropod Shidaisaurus and to mamenchisaurid sauropods, which were widely distributed throughout China, during the Jurassic, and are known from skeletal remains found in the same unit. Other potential sauropod trackmakers include titanosauriforms or as-yet-unknown basal eusauropods. The ichno- and skeletal records from the Jurassic of the Lufeng Basin are largely consistent, and both document the presence of middle-large sized theropods and sauropods.     

湖北淅川盆地上白垩统蜥脚类幼年个体新材料

描述湖北郧县淅川盆地晚白垩世地层中产出的蜥脚类恐龙化石。标本不关联保存于马家村组中段灰绿色含钙泥质粉砂岩中,材料包括牙齿、脊椎和肢骨化石等。从愈合特征（如分离的椎体和椎弓）分析,这些化石至少由2个未成年个体组成。这些化石体现出的特征组合,如股骨近端外侧突出,近端外缘向内侧倾斜,荐前椎气腔构造发育,以及牙齿呈细长的棒状,表明湖北郧县马家村组地层发现的蜥脚类可归入巨龙形类中的多孔椎龙类或者更进步的类群。新材料的发现对于进一步了解中国乃至东亚地区白垩纪晚期的蜥脚类恐龙分布和演化具有一定的意义。     

<Emphasis Type="Italic">Arkharavia heterocoelica</Emphasis> gen. et sp. nov., a new sauropod dinosaur from the Upper Cretaceous of the Far East of Russia

A new sauropod dinosaur, Arkharavia heterocoelica gen. et sp. nov., from the Maastrichtian (Udurchukan Formation) of the Amur Region, Russia, is described based on a tooth and several isolated anterior caudal vertebrae. It is distinguished by the saddle-shaped centrum and high neural spine of the anterior caudal vertebrae. Certain structural characters of the new genus are in common with Chubutisaurus insignis (Titanosauriformes) from the Upper Cretaceous of Argentina.