The ‘Giant of Ksour’, a Middle Jurassic sauropod dinosaur from Algeria

Continental strata of Early and Middle Jurassic age are seldom-exposed, and little is known of the history of sauropod dinosaurs prior to the neosauropod radiation of the end of the Middle Jurassic. Here, we report, in the Middle Jurassic of the Occidental Saharan Atlas (Algerian High Atlas), the discovery of a skeleton, including cranial material, of a new cetiosaurid sauropod. Chebsaurus algeriensis n. g., n. sp. represents the most complete Algerian sauropod available to date, only few remains were found before. To cite this article: F. Mahammed et al., C. R. Palevol 4 (2005).     

Cranial anatomy of Shunosaurus, a basal sauropod dinosaur from the Middle Jurassic of China

Shunosaurus, from the Middle Jurassic of China, is probably the best‐known basal sauropod and is represented by several complete skeletons. It is unique among sauropods in having a small, bony club at the end of its tail. New skull material provides critical information about its anatomy, brain morphology, tooth replacement pattern, feeding habits and phylogenetic relationships. The skull is akinetic and monimostylic. The brain is relatively small, narrow and primitively designed. The tooth replacement pattern exhibits back to front replacement waves in alternating tooth position. The teeth are spatulate, stout and show well‐developed wear facets indicative of coarser plant food. Upper and lower tooth rows interdigitate and shear past each other. Tooth morphology, skull architecture, and neck posture indicate that Shunosaurus was adapted to ground feeding or low browsing. Shunosaurus exhibits the following cranial autapomorphies: emargination of the ventral margin of the jugal/quadratojugal bar behind the tooth row; postorbital contains a lateral pit; vomers do not participate in the formation of the choanae; pterygoid is extremely slender and small with a dorsal fossa; quadrate ramus of the pterygoid is forked; quadratojugal participates in the jaw articulation; tooth morphology is a combination of cylindrical and spatulate form; basipterygoid process is not wrapped by the caudal process of the pterygoid; trochlear nerve has two exits; occlusal level of the maxillary tooth row is convex downward, whereas that of the dentary is concave upward, acting like a pair of garden shears; dentary tooth count is 25 or more; and the replacing teeth invade the labial side of the functional teeth. Cranial characters among the basal sauropods are reviewed. As Shunosaurus is the earliest sauropod for which cranial remains are known, it occupies an important position phylogenetically, showing the modification of skull morphology from the prosauropod condition. Although the skull synapomorphies of Sauropoda are unknown at present, 27 cranial synapomorphies are known for the clade Eusauropoda. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society, 2002, 136 , 145?169.     

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.     

The phylogenetic relationships of sauropod dinosaurs

A data-matrix of 205 osteological characters for 26 sauropod taxa is subjected to cladistic analysis. Two most parsimonious trees are produced, differing only in the relationships between Euhelopus, Omeisaurus and Mamenchisaurus. The monophyly of the Euhelopodidae (including Shunosaurus) is supported by seven synapomorphies. The Cetiosauridae (Patagosaurus, Cetiosaurus and Haplocanthosaurus) is paraphyletic with respect to the Neosauropoda. The latter clade divides into two major radiations–the ‘Brachiosauria’ (Camarasaurus, brachiosaurids and titanosauroids), and the Diplodocoidea (nemegtosaurids, dicraeosaurids, diplodocids and Rebbachisaurus). Further evidence for the inclusion of Opisthocoelwaudia in the Titanosauroidea is presented. Phuwiangosaurus, a problematic sauropod from Thailand, may represent one of the most plesiomorphic titanosauroids. ‘Peg’-like teeth have evolved at least twice within the Sauropoda. The postspinal lamina, on the neural spines of middle and caudal dorsal vertebrae, represents a neomorph rather than a fusion of pre-existing structures. Forked chevrons may have evolved convergently in the Euhelopodidae and the diplodocid-dicraeosaurid clade, or they may have been acquired early in sauropod evolution and subsequently lost in the ‘Brachiosauria’. The strengths and weaknesses of the data-matrix and tree topologies are explored using bootstrapping, decay analysis and randomization tests. Several nodes are only poorly supported, but this seems to reflect the large proportion of missing data in the matrix (～46%), rather than an abnormally high level of homoplasy. The results of the randomization tests indicate that the ‘data-matrix’ probably contains a strong phylogenetic ‘signal’. The relationships of some forms, such as Haplocanthosaurus, are influenced by the inclusion or exclusion of certain taxa with unusual combinations of character states. Such a result suggests that there are dangers inherent in the view that ‘higher’ level sauropod phylogeny can be accurately reconstructed using only a small number of well-known taxa.     

Oldest camarasauromorph sauropod (Dinosauria) discovered in the Middle Jurassic (Bajocian) of the Khadir Island, Kachchh, western India

Fragmentary isolated remains of large (up to 20 m or more) sauropods from the Middle Jurassic (Bajocian) Khadir Formation of Khadir Island (Kachchh, W India) are described and compared in detail. Three of the bone fragments (a metacarpal, a first pedal claw and a fibula) can be assigned with confidence to the Camarasauromorpha and represent the oldest known record of that derived dinosaur group. The new finds from western India further close a temporal and geographical gap in our knowledge of sauropods and contribute to understanding their early phylogeny.      

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.     

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.     

Necks for sex: sexual selection as an explanation for sauropod dinosaur neck elongation

The immensely long neck of a sauropod is one of the most familiar and striking of anatomical specializations among dinosaurs. Here, I use recently collected neontological and paleontological information to test the predictions of two competing hypotheses proposed to explain the significance of the long neck. According to the traditional hypothesis, neck elongation in sauropods increased feeding height, thereby reducing competition with contemporaries for food. According to the other hypothesis, which is advanced for the first time here, neck elongation in sauropods was driven by sexual selection. Available data match the predictions of the sexual selection hypothesis and contradict the predictions of the feeding competition hypothesis. It is therefore more plausible that increases in sauropod neck lengths were driven by sexual selection than by competition for foliage.     

Osteology and relationships of a new theropod dinosaur from the Middle Jurassic of Patagonia

Abstract:  A new taxon of theropod dinosaur is described as Condorraptor currumili gen. et sp. nov., from the Middle Jurassic (Callovian) Cañadón Asfalto Formation of Chubut Province, Argentinean Patagonia. The taxon is represented by a single fragmentary postcranial skeleton. Although incompletely known, Condorraptor is the second most complete theropod from the Middle Jurassic of Gondwana. The new taxon is characterized by the absence of a posterior incision between the fibular condyle and the medial side of the proximal articular end of the tibia, the pleurocoels in the anterior cervicals being situated posteroventral to the parapophyses, and the presence of a pronounced 'step' between the distal articular facet and shaft of Mt IV. Pneumatic features of the vertebral column show strong variation between the left and right side. Condorraptor gen. nov. can be referred to the Tetanurae and is a representative of a global radiation of basal tetanurans in the Early to Mid Jurassic.     

A sauropod dinosaur megatracksite from the Late Jurassic of northern Switzerland

The largest known sauropod trackway site from the Upper Jurassic in Europe has been found in the northern Jura Mountains of Switzerland. Since the initial discovery of the site in 1988, detailed ichno‐facies mapping of the sites has been undertaken and completed. Six separate sites (from 5 to 45km apart) have been located to date; the largest one, displaying 345 single imprints, extends over a surface of 7000 m². As all of the reported sites occur within the same bed, these vertebrate prints form a megatracksite covering an area of more than 360 km². All localities are stratigraphically within the Reuchenette Formation. Two biostratigraphically diagnostic ammonites have been found within the sequence (Aulacostephanus; Gravesia), indicating an Upper Kimmeridgian age (sensu gallico). In the easternmost sites, imprints occur on mud‐cracked tidal pond deposits, whereas the western tracksites are found in supratidal algal marsh deposits. The track‐bearing horizon is immediately below a trans‐gressive event at all the sites, suggesting that the ichnofaunas represent dinosaur activity during a relative sea‐level low‐stand. Footprint size and morphology, as well as trackway dimensions, suggest that the track producers belong to the largest sauropods yet recorded in the European Jurassic, closely resembling the Breviparopus tracks from the Middle Jurassic of Morocco.     

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.     

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     

The earliest known sauropod dinosaur and the first steps towards sauropod locomotion

A partial dinosaur skeleton from the Upper Triassic (Norian) sediments of South Africa is described and named Antetonitrus ingenipes. It provides the first informative look at a basal sauropod that was beginning to show adaptations towards graviportal quadrupedalism such as an elongated forelimb, a modified femoral architecture, a shortened metatarsus and a changed distribution of weight across the foot. These adaptations allowed the clade to produce the largest-ever terrestrial animals. However, A. ingenipes lacked specializations of the hand found in more derived sauropods that indicate it retained the ability to grasp. Antetonitrus is older than the recently described Isanosaurus from Thailand and is the oldest known definitive sauropod.     

The northernmost sauropod record in the Northern Hemisphere

Isolated sauropod teeth from the Early Cretaceous Teete locality in Yakutia (Eastern Siberia, Russia) are the only evidence that sauropods lived in high latitudes (palaeolatitude estimate of N 62°) in the Northern Hemisphere. The spatulate broad tooth crowns of adult individuals lack marginal denticles while these are present in a juvenile tooth. The teeth have overlapping facets and likely belong to a basal macronarian. The juvenile tooth indicates that sauropods reproduced in high latitudes and possibly stayed there around the year. The Teete vertebrate assemblage comprises both endothermic, or presumably endothermic tetrapods (theropod dinosaurs, tritylodontids and mammals), and ectothermic tetrapods (salamanders, turtles, choristoderes and lizards), but no crocodyliforms. This suggests a temperate climate, with an annual mean temperature well above freezing level but below 14°C.     

The first evidence of osteomyelitis in a sauropod dinosaur

Osteomyelitis is reported for the first time in a sauropod dinosaur. The material (MCS‐PV 183) comes from the Anacleto Formation (Campanian, Late Cretaceous), at the Cinco Saltos locality, Río Negro Province, Argentina. The specimen consists of 16 mid and mid‐distal caudal vertebrae of a titanosaur sauropod. Evidence of bacterial infection is preserved in all of these vertebrae. The main anomalies are as follows: irregular ‘microbubbly’ texture of bone surfaces produced by periosteal reactive bone, abscesses on the rims of the anterior articular surfaces of two centra, numerous pits on centra anterior articulation surfaces, erosions on the anterior articulation of the vertebral centra, a vertical groove in posterior articular face of all the centra and disruption of the prezygapophysis and postzygapophysis (mainly the articular face) from the vertebra 19 and beyond. The last anomaly is increasingly pronounced in more distal elements of the series. Thin sections reveal that the anomalous cortical tissue is composed of avascular and highly fibrous bone matrix. The fibres of the bone matrix are organized into thick bundles oriented in different directions. Both morphological and histological abnormalities in the MCS‐PV 183 specimen are pathognomonic for osteomyelitis.     

Comparative vegetative anatomy and classification of Diseae (Orchidaceae)

The leaf, stem, root, tuber and dropper anatomy of the orchid tribe Diseae (including the subtribes Satyriinae, Disinae, Brownlecinac, Huttonaeinae and Coryciinae) is reviewed. The study is largely based on investigations of 123 species, and data from several previous publications have also been incorporated. Two characters were identified as being taxonomically valuable: (1) the presence of sclerenchyma caps associated with leaf vascular bundles, and (2) the degree of dissection of the siphonostele of the tuber (‘polystelic’ or ‘monostelic’). The phylogenetic analysis shows that anatomical characters do not change the basic structure of a cladogram that is based on morphological characters. The taxa of Diseae are discussed on the basis of anatomical data. Subtribes Satyriinae (excluding the anatomically unusual genus Pachites), Brownleeinae, Huttonaeinae, and Coryciinae are uniform in. critical anatomical characters. However, subtribe Disinae is rather diverse in vegetative anatomy. Disa sect. Micranthae differs from the rest of the genus in its leaf anatomy. The occurrence of foliar sclerenchyma bundle caps and ‘polystelic’ tubers supports the incorporation of Herschelianthe in Disa sect. Stenocarpa.     

Comparative vegetative anatomy and classification of Diseae (Orchidaceae)

The leaf, stem, root, tuber and dropper anatomy of the orchid tribe Diseae (including the subtribes Satyriinae, Disinae, Brownlecinac, Huttonaeinae and Coryciinae) is reviewed. The study is largely based on investigations of 123 species, and data from several previous publications have also been incorporated. Two characters were identified as being taxonomically valuable: (1) the presence of sclerenchyma caps associated with leaf vascular bundles, and (2) the degree of dissection of the siphonostele of the tuber ('polystelic' or 'monostelic'). The phylogenetic analysis shows that anatomical characters do not change the basic structure of a cladogram that is based on morphological characters. The taxa of Diseae are discussed on the basis of anatomical data. Subtribes Satyriinae (excluding the anatomically unusual genus Pachites), Brownleeinae, Huttonaeinae, and Coryciinae are uniform in. critical anatomical characters. However, subtribe Disinae is rather diverse in vegetative anatomy. Disa sect. Micranthae differs from the rest of the genus in its leaf anatomy. The occurrence of foliar sclerenchyma bundle caps and 'polystelic' tubers supports the incorporation of Herschelianthe in Disa sect. Stenocarpa.     

Phylogenetic analysis and revision of the linyphiid spider genus Solenysa (Araneae: Linyphiidae: Erigoninae)

In this paper we carry out a taxonomic revision and phylogenetic analysis of the linyphiid spider genus Solenysa Simon, 1894. A total of 12 species is treated here, including five new species collected from China and Japan: Solenysa akihisai Tu sp. nov., Solenysa lanyuensis Tu sp. nov., Solenysa retractilis Tu sp. nov., Solenysa tianmushana Tu sp. nov. , and Solenysa yangmingshana Tu sp. nov. Solenysa circularis Gao, Zhu & Sha, 1993 is a junior synonym of Solenysa protrudens Gao, Zhu & Sha, 1993. We have assembled two different character matrices to reconstruct the phylogenetic relationships of Solenysa. In the first matrix (Matrix 1), five representative species of Solenysa were added to the morphological dataset of Miller & Hormiga to test the monophyly of the genus and its placement within Linyphiidae. The genitalic structures and somatic morphology of Solenysa were studied by means of scanning electron microscopy for the first time. To infer the species‐level phylogenetic relationships of Solenysa we produced a second matrix (Matrix 2) that includes all 12 Solenysa species and six outgroup species chosen from the results of the analysis of the first matrix. The two most parsimonious trees from the analysis of Matrix 1 support the monophyly of Solenysa and its placement within the ‘Distal Erigonines’ clade. The single most parsimonious tree resulting from the analysis of the second matrix suggests that the Solenysa clade includes four monophyletic groups, each group represented by a distinct genitalic pattern. The morphology of Solenysa, both somatic and genitalic, is highly autapomorphic. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 484–530.     

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.