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.     

Dinosaurs of France

The French dinosaur record is one of the most extensive in Europe; it ranges stratigraphically from the Late Triassic to the Latest Cretaceous. All major clades of dinosaurs but marginocephalians are known. About 20 species are based on significant material; the theropods are the best represented. Most of these taxa have been described or revised in recent years. Important specimens have been discovered in the Late Triassic of eastern France, the Middle Jurassic of Normandy, and the Late Cretaceous of Provence and Languedoc. The ichnological record is good for the Late Triassic-Early Jurassic, and the Late Cretaceous egg sites are among the richest in the world. To cite this article: R. Allain, X.P. Suberbiola, Palevol 2 (2003) 27–44.     

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     

A large pholidosaurid in the Phu Kradung Formation of north‐eastern Thailand

In the early 1980s, the remains of a large crocodilian, consisting of a nearly complete lower jaw, were referred to a distinct species of Sunosuchus, S. thailandicus. The specimen was recovered from a road‐cut near Nong Bua Lamphu, north‐eastern Thailand, in the upper part of the continental Phu Kradung Formation, and then considered Early to Middle Jurassic in age. Since then, this age has been revised and most of the formation is now considered Early Cretaceous, although a Late Jurassic age is possible for its lowermost part. Here, we report for the first time cranial elements associated with mandibular remains assignable to ‘S’. thailandicus. An attribution to Pholidosauridae is proposed on the basis of premaxillary morphology, and the original referral of this taxon to the goniopholidid Sunosuchus is discarded. A new genus name Chalawan now designates the originally described material of S. thailandicus. Nevertheless, the newly described specimen shares a characteristic with both ‘traditional’ Goniopholididae and Pholidosauridae: the presence of a depression located on the lateral wall of the maxilla and jugal. A phylogenetic analysis confirms the inclusion of both Goniopholididae and Pholidosauridae into a common clade, Coelognathosuchia tax. nov. Although the new Thai skull is much fragmented, its original shape is reconstructed and is compared with other pholidosaurid genera, namely Elosuchus, Meridiosaurus, Oceanosuchus, Pholidosaurus, Sarcosuchus and Terminonaris. The presence of the genus Sunosuchus being highly questionable in Thailand, it cannot be used as evidence to link the Chinese and Indochinese blocks. Instead, the recognition of a freshwater pholidosaurid in a continental formation of the Indochinese block suggests that early in their evolutionary history, these crocodilians, already known from Europe, Africa and South America, were more widely distributed along the northern margin of the Tethys than previously recognized.     

MUSOPSIS N. GEN.: A BANANA-LIKE LEAF GENUS FROM THE EARLY TERTIARY OF EASTERN NORTH GREENLAND

A fossil flora from the Late Paleocene-Early Eocene Thyra Ø Formation of eastern North Greenland (paleolatitude 77° N) has yielded monocotyledon leaf impressions with characters seen only in the closely related modem species in the families of Heliconiaceae, Musaceae, and Strelitziaceae. The combination of large costae widths and parallel, nonanastomosing, lateral veins that depart at right angles from the costae in the fossil material are features present only in leaves of extant species from these families. Three basic venation patterns also are recognized in the modem species of these families, but except for the genera Strelitzia and Phenakospermum, none of these patterns are present exclusively in any one family. Musopsis n. gen. is created for the fossil material from Greenland, but it is considered a form genus due to the lack of gross morphological features that can be used for separating leaves of the modem genera in Heliconiaceae, Musaceae, and Strelitiziaceae. It is the first known Arctic occurrence of fossil leaf material resembling this modem group of taxa.     

New species of the genus <Emphasis Type="Italic">Macclintockia</Emphasis> (Angiosperms) from the Campanian of the Ugol’naya Bay (Northeastern Russia)

New species of angiosperms are described as members of the genus Macclintockia from the Ugol’naya Bay area, in northeastern Russia: M. barykovensis Moiseeva, sp. nov., M. gigantea Moiseeva, sp. nov., and M. terranea Moiseeva et Herman, sp. nov. Leaves of this genus are considerably variable, which hampers differentiation between particular species. The study of new material has allowed the author to trace the morphological variability of these species and revised on this ground the composition of the species M. ochotica Vachr. et Herman and M. beringiana Herman. The composition of Macclintockia is discussed, as well as its occurrence during the Late Cretaceous.     

Changes in lifestyle and habitat of Zoophycos‐producing animals related to evolution of phytoplankton during the Late Mesozoic: geological evidence for the ‘benthic‐pelagic coupling model’

The trace fossil Zoophycos characterized by complex, three‐dimensional morphology with systematic internal structures occurs throughout the Phanerozoic marine sediments. The specimens of Zoophycos examined herein consist of a downward and helical spreite and are a product of the excretory behaviour of endobenthic detritus feeders. They are divided into two basic types: pre‐Jurassic and post‐Cretaceous types on the basis of whorls of spreiten in a single specimen. The pre‐Jurassic type has fewer than four whorls. In contrast, most of the post‐Cretaceous specimens exhibit spreite with multiple coils more than ten whorls. The abrupt increase in whorl number during the Cretaceous suggests that the sedentary lifestyle of the producer should change from a short‐term stay to long‐term or permanent occupation of the same burrow. Timing of the lifestyle change the Zoophycos producers seems to be closely related to the deep‐seaward migration of their habitats. The change in lifestyle and migration of Zoophycos‐producing animals during the Cretaceous might be attributable to the establishment of eutrophic bottom conditions in the deep sea. These changes seem to be associated with the flux of large amounts of phytodetrital food produced by phytoplankton, which experienced an explosive increase in species diversity during the Late Jurassic to the Late Cretaceous. The series of changes in lifestyle and habitat of the Zoophycos animals during the Late Mesozoic can serve as one piece of geological evidence for the ‘benthic‐pelagic coupling model’.     

Normapolles plants: a prominent component of the Cretaceous rosid diversification

The Normapolles complex, characterised by its oblate and triaperturate pollen, constitutes an important and diverse element of many Late Cretaceous and Early Cainozoic floras of the Northern Hemisphere. Based on the dispersed pollen record alone it has been difficult to assess systematic affinities, but relationships with Fagales have been proposed. Over the past twenty years several exquisitely preserved Late Cretaceous reproductive structures with Normapolles type pollen in situ have been described. In this study we provide a summary and new information of these floral structures. Further, a new genus, Dahlgrenianthus, is described from the Late Cretaceous of southern Sweden. The genus includes the type species Dahlgrenianthus suecicus, a number of reproductive structures referred to Dahlgrenianthus sp., and Dahlgrenianthus trigonus (Knobloch et Mai) comb. nov. from the Maastrichtian flora of Walbeck, Germany. Dahlgrenianthus comprises small flowers with pentamerous perianth and androecium and a tricarpellate gynoecium. It is distinguished from all other Normapolles floral structures in its hypogynous floral organisation. All Normapolles floral structures described so far are thought to be related to various members of the core Fagales, but the group is obviously not monophyletic. The stratigraphic range of the Normapolles taxa and other fagalean fossils strongly suggests that all major fagalean lineages were present by the Cenomanian or earlier.     

The first definitive Middle Jurassic atoposaurid (Crocodylomorpha,Neosuchia), and a discussion on the genus Theriosuchus

Atoposaurids were a clade of semiaquatic crocodyliforms known from the Late Jurassic to the latest Cretaceous. Tentative remains from Europe, Morocco, and Madagascar may extend their range into the Middle Jurassic. Here we report the first unambiguous Middle Jurassic (late Bajocian–Bathonian) atoposaurid: an anterior dentary from the Isle of Skye, Scotland, UK. A comprehensive review of atoposaurid specimens demonstrates that this dentary can be referred to Theriosuchus based on several derived characters, and differs from the five previously recognized species within this genus. Despite several diagnostic features, we conservatively refer it to Theriosuchus sp., pending the discovery of more complete material. As the oldest known definitively diagnostic atoposaurid, this discovery indicates that the oldest members of this group were small‐bodied, had heterodont dentition, and were most likely widespread components of European faunas. Our review of mandibular and dental features in atoposaurids not only allows us to present a revised diagnosis of Theriosuchus, but also reveals a great amount of variability within this genus, and indicates that there are currently five valid species that can be differentiated by unique combinations of dental characteristics. This variability can be included in future broad‐scale cladistics analyses of atoposaurids and closely related crocodyliforms, which promise to help untangle the complicated taxonomy and evolutionary history of Atoposauridae. © 2015 The Authors. Zoological Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of The Linnean Society of London     

Evolutionary relationships and systematics of Atoposauridae (Crocodylomorpha: Neosuchia): implications for the rise of Eusuchia

Atoposaurids are a group of small‐bodied, extinct crocodyliforms, regarded as an important component of Jurassic and Cretaceous Laurasian semi‐aquatic ecosystems. Despite the group being known for over 150 years, the taxonomic composition of Atoposauridae and its position within Crocodyliformes are unresolved. Uncertainty revolves around their placement within Neosuchia, in which they have been found to occupy a range of positions from the most basal neosuchian clade to more crownward eusuchians. This problem stems from a lack of adequate taxonomic treatment of specimens assigned to Atoposauridae, and key taxa such as Theriosuchus have become taxonomic ‘waste baskets’. Here, we incorporate all putative atoposaurid species into a new phylogenetic data matrix comprising 24 taxa scored for 329 characters. Many of our characters are heavily revised or novel to this study, and several ingroup taxa have never previously been included in a phylogenetic analysis. Parsimony and Bayesian approaches both recover Atoposauridae as a basal clade within Neosuchia, more stemward than coelognathosuchians, bernissartiids, and paralligatorids. Atoposauridae is a much more exclusive clade than previously recognized, comprising just three genera (Alligatorellus, Alligatorium, and Atoposaurus) that were restricted to the Late Jurassic of western Europe, and went extinct at the Jurassic/Cretaceous boundary. A putative Gondwanan atoposaurid (Brillanceausuchus) is recovered as a paralligatorid. Our results exclude both Montsecosuchus and Theriosuchus from Atoposauridae. Theriosuchus is polyphyletic, forming two groupings of advanced neosuchians. Theriosuchus (restricted to Theriosuchus pusillus, Theriosuchus guimarotae, and Theriosuchus grandinaris) spanned the Middle Jurassic to early Late Cretaceous, and is known from Eurasia and North Africa. Two Cretaceous species previously assigned to Theriosuchus (‘Theriosuchus’ ibericus and ‘Theriosuchus’ sympiestodon) are shown to be nested within Paralligatoridae, and we assign them to the new genus Sabresuchus. The revised phylogenetic placement of Theriosuchus has several implications for our understanding of eusuchian evolution. Firstly, the presence of fully pterygoidean choanae, previously regarded as a defining characteristic of Eusuchia, is not found in some basal members of Eusuchia. However, eusuchians can be distinguished from Theriosuchus and other basal neosuchians in that their choanae are posteriorly positioned, with an anterior margin medial to the posterior edge of the suborbital fenestra. This feature distinguishes eusuchians from Theriosuchus and more basal neosuchians. Secondly, our refined understanding of Theriosuchus implies that this taxon possessed only amphicoelous presacral vertebrae, and therefore fully developed vertebral procoely is likely to have evolved only once in Crocodylomorpha, on the lineage leading to Eusuchia. These and other findings presented herein will provide an important framework for understanding the neosuchian–eusuchian transition.     

The morphology,taxonomy, and occurrence of the genus <Emphasis Type="Italic">Pseudoprotophyllum</Emphasis> Hollick (Platanaceae) in Late Cretaceous floras of Northern Asia

An extended generic diagnosis of Pseudoprotophyllum is given on the basis of new data from Late Cretaceous floras of Northern Asia. Both peltate and apeltate morphological types are included in the genus. The morphological diversity is described in the type species P. boreale (Dawson) Hollick from the Cenomanian-Turonian of North America and four North Asiatic species: P. minimum I. Lebed. , P. sibiricum I. Lebed., P. hatangaense Abramova, and P. giganteum Sveshn. et Budants. The geographical and stratigraphic analyses show that Pseudoprotophyllum evolved in Northern Asia since the Cenomanian until Campanian and was restricted to the Siberian-Canadian floristic region with a warm-temperate humid climate.     

Ants of the genus <Emphasis Type="Italic">Plagiolepis</Emphasis> Mayr (Hymenoptera,Formicidae) from Late Eocene ambers of Europe

The genus Plagiolepis Mayr (Hymenoptera, Formicidae) from the Late Eocene Baltic, Bitterfeld, Rovno, and Scandinavian ambers is revised. Two new species, Plagiolepis wheeleri sp. nov. and P. paradoxa sp. nov., are described. A lectotype of Plagiolepis klinsmanni Mayr, 1868 and neotypes of P. kuenowi Mayr, 1868 and P. solitaria Mayr, 1868 are designated. Rhopalomyrmex pygmaeus Mayr, 1868 (= Plagiolepis balticus Dlussky, 1997) is recognized as a new synonym of P. kuenowi Mayr, 1868. A key for identification of the Late Eocene species of Plagiolepis from the European ambers is provided.     

Tithonian-Berriasian foraminiferal faunas from the Torinosu-type calcareous blocks of the southern Kanto Mountains, Japan: their implications for post-accretionary tectonics of Jurassic to Cretaceous terranes

The Torinosu-type limestones, having many lithologic characters showing their original deposition on shallow shelves, are widely distributed in the Jurassic to Cretaceous terranes of Japan. The foraminiferal faunas from the Jurassic to the lowermost Cretaceous of Japan were first revealed in the calcareous blocks of the southern Kanto Mountains. Distinguished microfaunas consist of 39 species including many marker species of the Upper Jurassic to Lower Cretaceous in Europe, West Asia, and North Africa such as Melathrokerion spirialis, Charentia evoluta, Freixialina planispiralis, Nautiloculina oolithica, Everticyclammina cf. virguliana, Haplophragmium lutzei and Pseudocyclammina lituus. These faunas suggest a Tithonian to Berriasian age of Torinosu-type limestones. They are contained in four tectonostratigraphic units (Kamiyozawa, Hikawa and Gozenyama Formations; Ogouchi Group) continuously accreted from Middle Jurassic to Late Cretaceous. The younger deposition age of Torinosu-type limestones than the accretion age (Bajocian to Bathonian) in the Kamiyozawa Formation and their older age than the accretion age of the Ogouchi Group (late Albian to middle Maastrichtian) are important to date the post-accretionary tectonics of Jurassic to Cretaceous terranes of Japan and to explain the emplacement process of Torinosu-type limestones.     

The oldest species of the genus <Emphasis Type="Italic">Glaphyrus</Emphasis> Latr. (Coleoptera: Scarabaeoidea: Glaphyridae) from the Mesozoic of China

Glaphyrus ancestralis sp. nov. is described from the Yixian Formation (Upper Jurassic or Lower Cretaceous). The species is not only one of the earliest records of the family Glaphyridae but also the oldest representative of an extant genus of the family.     

A molecular timescale for the evolution of the African freshwater fish family Kneriidae (Teleostei: Gonorynchiformes)

We re-examine the phylogenetic relationships of the family Kneriidae using whole mitogenome sequences across all four kneriid genera including the two recently recognized species of Cromeria (C. nilotica and C. occidentalis), and we provide a timescale to discuss the evolution of the family. The resulting phylogeny supports the monophyly of the family Kneriidae and the monophyly of the genus Cromeria. The two Cromeria species exhibit large genetic divergence (18.2%) that is comparable to those between Grasseichthys gabonensis and each two Cromeria species (16.9 and 19.0%). The three paedomorphic kneriid species (C. occidentalis, C. nilotica and G. gabonensis) do not form a monophyletic group, but the alternative hypothesis in which they are monophyletic cannot be statistically rejected. Two alternative relaxed molecular-clock Bayesian analyses, differing on how we time-calibrated the phylogenetic tree using the fossil record, support a Late Jurassic or Late Cretaceous origin of the African freshwater gonorynchiforms. The early diversification of the family Kneriidae is concomitant with the reductions or loss of several morphological characters that took place in a relatively short time interval of about 12–21 million years either during the Eocene or at the end of the Late Cretaceous.     

A new lobster <Emphasis Type="Italic">Paraclytia valashtensis</Emphasis> (Crustacea,Decapoda, Nephropidae) from the Late Cretaceous of the central Alborz Range,Iran

A new species of Late Cretaceous clawed lobster, Paraclytia valashtensis, is described. The discovery is a notable addition to the sparse decapod fossil record of Iran, and this is the first record of the genus outside central Europe. The four previously known species of Paraclytia are from Germany and the Czech Republic, so this discovery represents a significant expansion of the palaeogeographic range of the genus.     

New water scavenger beetles (Coleoptera: Hydrophilidae) from the mesozoic of Mongolia

A new genus and two new species of water scavenger beetles, Hydrophilopsia bontsaganica, sp. nov. and Prospercheus cristatus, gen. et sp. nov., are described from the Upper Jurassic and Lower Cretaceous of Mongolia. The systematic position of the new genus is discussed.     

The belemnites described by Baron Ernst Friedrich von Schlotheim (1764–1832)

E. F. v. Schlotheim described 46 species of Jurassic and Cretaceous belemnites in 1813,1820, and 1832. Their type specimens are in part housed at the Humboldt University of Berlin. The holotype ofBelemnites irregularis is here refigured. Lectotypes forB. acuarius, B. canaliculatus, B. clavatus, B. giganteus, andB. tripartitus are designated and figured for the first time. The lectotype ofBelemnites giganteus is designated to be the neotype ofBelemnites suevicus Klein 1773. 36 species are nomina nuda and nomina dubia, because none of their type specimens could be detected, among them the invalidB. paxillosus v. Schlotheim 1820, which is, according to its lectotype designated herein, likeBelemnites giganteus, a junior synonym ofMegateuthis suevica (Klein 1773). An original type specimenof Belemnites mucronatusv. Schlotheim 1813, for which a neotype had already been ruled by the ICZN in 1985, was re-discovered in thev. Schlotheim collection in 1996.v. Schlotheim’s belemnites were up to now designated to be type species of 12 genera.