A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic

Non-avian theropod dinosaurs attained large body sizes, monopolising terrestrial apex predator niches in the Jurassic–Cretaceous. From the Middle Jurassic onwards, Allosauroidea and Megalosauroidea comprised almost all large-bodied predators for 85 million years. Despite their enormous success, however, they are usually considered absent from terminal Cretaceous ecosystems, replaced by tyrannosaurids and abelisaurids. We demonstrate that the problematic allosauroids Aerosteon, Australovenator, Fukuiraptor and Neovenator form a previously unrecognised but ecologically diverse and globally distributed clade (Neovenatoridae, new clade) with the hitherto enigmatic theropods Chilantaisaurus, Megaraptor and the Maastrichtian Orkoraptor. This refutes the notion that allosauroid extinction pre-dated the end of the Mesozoic. Neovenatoridae includes a derived group (Megaraptora, new clade) that developed long, raptorial forelimbs, cursorial hind limbs, appendicular pneumaticity and small size, features acquired convergently in bird-line theropods. Neovenatorids thus occupied a 14-fold adult size range from 175 kg (Fukuiraptor) to approximately 2,500 kg (Chilantaisaurus). Recognition of this major allosauroid radiation has implications for Gondwanan paleobiogeography: The distribution of early Cretaceous allosauroids does not strongly support the vicariant hypothesis of southern dinosaur evolution or any particular continental breakup sequence or dispersal scenario. Instead, clades were nearly cosmopolitan in their early history, and later distributions are explained by sampling failure or local extinction.     

An abelisaurid from the Late Cretaceous of Egypt: implications for theropod biogeography

Recent paleogeographic scenarios postulate the isolation of continental Africa during the Late Cretaceous. The absence of abelisaurid theropods from Upper Cretaceous African strata was offered as support of hypothesized African isolation with the acknowledgement that the paucity of African abelisaurids may be mostly an issue of sampling. Here we report on a shed theropod tooth from the Upper Cretaceous (Maastrichtian, ∼70 Ma) Duwi Formation of Egypt. The tooth was referred to the Malagasy abelisaurid “Megalosaurus” crenatissimus (=Majungasaurus crenatissimus) in 1921. A discriminant function analysis was run to test for morphological congruence between the Egyptian tooth and the dentitions of 24 theropod taxa. The analysis correctly classified 96.6% of the teeth in the sample and assigned the tooth to Majungasaurus. As current paleogeographic reconstructions posit Madagascar had attained its current position relative to Africa before the Late Cretaceous, it is unlikely that the Egyptian tooth actually pertains to Majungasaurus. Nevertheless, its classification as an abelisaurid supports its referral to the clade. This tooth thus constitutes defensible evidence of an abelisaurid from the post-Cenomanian Cretaceous of mainland Africa. Combined with recent discoveries of abelisaurids in Niger and Morocco, the result indicates that Abelisauridae was a diverse group in Africa during the Cretaceous, existing in multiple places for at least ∼25 Ma and weakens support for hypotheses of an isolated Africa during the Late Cretaceous.Electronic Supplementary Material  Supplementary material is available for this article at

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The first definitive carcharodontosaurid (Dinosauria: Theropoda) from Asia and the delayed ascent of tyrannosaurids

Little is known about the evolution of large-bodied theropod dinosaurs during the Early to mid Cretaceous in Asia. Prior to this time, Asia was home to an endemic fauna of basal tetanurans, whereas terminal Cretaceous ecosystems were dominated by tyrannosaurids, but the intervening 60 million years left a sparse fossil record. Here, we redescribe the enigmatic large-bodied Chilantaisaurus maortuensis from the Turonian of Inner Mongolia, China. We refer this species to a new genus, Shaochilong, and analyze its systematic affinities. Although Shaochilong has previously been allied with several disparate theropod groups (Megalosauridae, Allosauridae, Tyrannosauroidea, Maniraptora), we find strong support for a derived carcharodontosaurid placement. As such, Shaochilong is the first unequivocal Asian member of Carcharodontosauridae, which was once thought to be restricted to Gondwana. The discovery of an Asian carcharodontosaurid indicates that this clade was cosmopolitan in the Early to mid Cretaceous and that Asian large-bodied theropod faunas were no longer endemic at this time. It may also suggest that the ascent of tyrannosaurids into the large-bodied dinosaurian predator niche was a late event that occurred towards the end of the Cretaceous, between the Turonian and the Campanian. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A new scincomorph lizard from the Palaeocene of Belgium and the origin of Scincoidea in Europe

Well-preserved jaw fragments and skull bones of a new scincomorphan lizard are described from the continental Hainin Formation (Middle Palaeocene) of the Mons Basin in southwestern Belgium. This taxon, Scincoideus haininensis gen. et sp. nov., is characterised by vermiculate dermal rugosities and represents the earliest formally recognised scincoid in Europe. It differs from cordylids (girdled lizards) by lacking the parietal downgrowth and differs from scincids (skinks) by having robustly built bones and a well-developed lingual cusp on tooth crowns. By its mosaic characters, S. haininensis is considered here as being a basal scincoid lizard close to the origin of cordylids and scincids. Palaeobiogeographic analysis suggests that scincoids were present in Europe from the Palaeocene or maybe even before the Cretaceous/Palaeogene boundary and that they were replaced by modern scincoids such as true scincids and cordylids during the Eocene. The origin of the latter two families in Europe was very likely related with the immigration of the first modern mammals.Electronic supplementary material  Supplementary material is available for this article at and accessible for authorised users.     

An unusual archosaurian from the marine Triassic of China

A new Triassic archosaurian from China shows a number of aquatic specializations, of which the most striking is the extreme lateral compression of the long tail. Others that may also reflect aquatic adaptations include platelike scapula and coracoid, elongate neck with extremely long and slender ribs, and reduction of osteoderms. In contrast, its pelvic girdle and hind limb have no aquatic modifications. Anatomic features, taphonomy, and local geological data suggest that it may have lived in a coastal–island environment. This lifestyle, convergent with some Jurassic marine crocodyliforms that lived at least 40 million years later and the saltwater species of extant Crocodylus, contradicts with the prevailing view that Triassic archosaurians were restricted to nonmarine ecosystems. Its mosaic anatomy represents a previously unknown ecomorph within primitive archosaurians.Electronic Supplementary Material  Supplementary material is available for this article at     

Cannibalistic feeding of larval Trichogramma carverae parasitoids in moth eggs

Wasps of the genus Trichogramma parasitise the eggs of Lepidoptera. They may deposit one or many eggs in each host. Survival is high at low density but reaches a plateau as density increases. To reveal the mechanism by which excess larvae die we chose a lepidopteran host that has flattened, transparent eggs and used video microscopy to record novel feeding behaviours and interactions of larval Trichogramma carverae (Oatman and Pinto) at different densities. Single larvae show a rapid food ingestion phase, followed by a period of extensive saliva release. Ultimately the host egg is completely consumed. The larva then extracts excess moisture from the egg, providing a dry environment for pupation. When multiple larvae are present, the initial scramble for food results in the larvae consuming all of the egg contents early in development. All larvae survive if there is sufficient food for all to reach a threshold developmental stage. If not, physical proximity results in attack and consumption of others, continuing until the surviving larvae reach the threshold stage beyond which attacks seem to be no longer effective. The number of larvae remaining at the end of rapid ingestion dictates how many will survive to emerge as adults. Electronic Supplementary Material  Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Evidence for contact sex recognition pheromone of the Asian longhorned beetle,<Emphasis Type="Italic"> Anoplophora glabripennis</Emphasis> (Coleoptera: Cerambycidae)

Field observations of the Asian longhorned beetle (Anoplophora glabripennis) mating behavior in China suggested that a female-produced contact pheromone was almost certainly involved in sex recognition. Gas chromatography–mass spectrometry (GC-MS) analysis of A. glabripennis adults' whole body cuticular extracts indicates that a series of long-chain hydrocarbons comprise the cuticular waxes of both sexes. Although for the most part the GC profiles are similar for the two sexes, five monounsaturated compounds were consistently more abundant in samples from females than in those from males. These compounds were identified as (Z)-9-tricosene, (Z)-9-pentacosene, (Z)-7-pentacosene, (Z)-9-heptacosene, and (Z)-7-heptacosene in the approximate ratio of 1:2:2:8:1, respectively. Antennal and palpi contact to a polypropylene micro-centrifuge tube coated with a synthetic mixture of the five compounds stimulated copulatory behavior in males.Electronic Supplementary Material  Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.     

A chicken-sized crane precursor from the early Oligocene of France

A skeleton of a chicken-sized crane precursor is described from the Lower Oligocene of the Lubéron in Southern France. Parvigrus pohli gen. et sp. nov. is the most substantial Paleogene fossil record of the Grues (Aramidae [limpkin] + Gruidae [cranes]), and among its oldest representatives. The fossil species is classified in the new taxon Parvigruidae, which is shown to be the sister group of extant Grues. It is the first fossil record of a stem lineage representative of the Grues and, among others, differs from modern Grues in its smaller size, shorter beak, and rail-like limb proportions. Size increase in the stem lineage of the Gruidae may be related to the spread of grasslands during the Oligocene and Miocene. Occurrence of stem lineage Grues in the Lower Oligocene of Europe is in concordance with the fact that there is no evidence for the presence of crown group members of modern avian “families” in pre-Oligocene fossil deposits. Electronic Supplementary Material  Supplementary material is available for this article at     

Floral CO<Subscript>2</Subscript> emission may indicate food abundance to nectar-feeding moths

As part of a study of the roles of the sensory subsystem devoted to CO₂ in the nectar-feeding moth Manduca sexta, we investigated CO₂ release and nectar secretion by flowers of Datura wrightii, a preferred hostplant of Manduca. Datura flowers open at dusk and wilt by the following noon. During the first hours after dusk, when Manduca feeds, the flowers produce considerable amounts of nectar and emit levels of CO₂ that should be detectable by moths nearby. By midnight, however, both nectar secretion and CO₂ release decrease significantly. Because nectar production requires high metabolic activity, high floral CO₂ emission may indicate food abundance to the moths. We suggest that hovering moths could use the florally emitted CO₂ to help them assess the nectar content before attempting to feed in order to improve their foraging efficiency.Electronic Supplementary Material  Supplementary material is available in the online version of this article at     

<Emphasis Type="Italic">Jeholornis</Emphasis> compared to<Emphasis Type="Italic"> Archaeopteryx</Emphasis>, with a new understanding of the earliest avian evolution

The recently reported Jeholornis represents the only known bird with a complete long skeletal tail except for Archaeopteryx. Two newly discovered specimens referable to Jeholornis provide some important new information about its anatomy. The tail of Jeholornis is much longer than that of Archaeopteryx and comprises a maximum of 27 caudal vertebrae compared with only 23 in Archaeopteryx. More interestingly, the tail feathers are shaped more like those of dromaeosaurs than those of Archaeopteryx. We conclude that the common ancestor of birds must have a more primitive tail than that in Archaeopteryx, confirming the side branch position of Archaeopteryx in the early avian evolution. The synsacrum is composed of six sacrals, representing a transitional stage between Archaeopteryx and more advanced birds. The scapula of Jeholornis has a dorso-laterally exposed glenoid facet, and the coracoid has a supracoracoid foramen. The presence of a pair of fenestrae in the sternum of Jeholornis has further implications for the air-sac system in early birds. Electronic Supplementary Material  The online version of this article (doi: contains supplementary material, which is available to authorized users.     

Palaeoenvironmental controls on the distribution of Cretaceous herbivorous dinosaurs

Previous attempts to determine palaeoenvironmental preferences in dinosaurs have generally been qualitative assessments based upon data from restricted geographical areas. Here, we use a global database of Cretaceous herbivorous dinosaurs to identify significant associations between clades and broad palaeoenvironmental categories (‘terrestrial’, ‘coastal’, ‘marine’). Nodosaurid ankylosaurs and hadrosaurids show significant positive associations with marine sediments, while marginocephalians (Ceratopsia, Pachycephalosauria), saurischians (herbivorous theropods, Sauropoda) and ankylosaurid ankylosaurs are significantly positively associated with terrestrial sediments. These results provide quantitative support for the hypothesis that some clades (Nodosauridae, Hadrosauridae) were more abundant in coastal and/or fluvial environments, while others (e.g. Marginocephalia, Ankylosauridae) preferentially inhabited more distal environments. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. An erratum to this article can be found at     

Isolation and characterization of mesophilic,oxalate-degrading<Emphasis Type="Italic"> Streptomyces</Emphasis> from plant rhizosphere and forest soils

The present work was aimed at the isolation of additional new pure cultures of oxalate-degrading Streptomyces and its preliminary characterization for further work in the field of oxalate metabolism and taxonomic studies. Mesophilic, oxalate-degrading Streptomyces were enriched and isolated from plant rhizosphere and forest soil samples. Strains were examined for cultural, morphological (spore chain morphology, spore mass colour, diffusible and melanin pigment production), physiological (antibiosis, growth in the presence of inhibitory compounds, assimilation of organic acids and enzyme substrates) and chemotaxonomic characters (cellular lipid components and diagnostic cell-wall diamino acid). The taxonomic data obtained were analysed by using the simple matching (S_SM) and Jaccard (S_J) coefficients, clustering was achieved using the UPGMA algorithm. All strains were able to utilize sodium-, potassium-, calcium- and ammonium-oxalate salts. Based on the results of numerical taxonomy, isolates were grouped into five cluster groups with a 70% S_SM similarity level. Streptomyces rochei was the most common of the cluster groups, with a Willcox probability of P>0.8. Streptomyces antibioticus, S. anulatus, S. fulvissimus, S. halstedii and S. violaceusniger are newly reported as oxalate-utilizing Streptomyces.Electronic Supplementary Material  Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.     

Earliest zygodactyl bird feet: evidence from Early Cretaceous roadrunner-like tracks

Fossil footprints are important in understanding Cretaceous avian diversity because they constitute evidence of paleodiversity and paleoecology that is not always apparent from skeletal remains. Early Cretaceous bird tracks have demonstrated the existence of wading birds in East Asia, but some pedal morphotypes, such as zygodactyly, common in modern and earlier Cenozoic birds (Neornithes) were unknown in the Cretaceous. We, herein, discuss the implications of a recently reported, Early Cretaceous (120–110 million years old) trackway of a large, zygodactyl bird from China that predates skeletal evidence of this foot morphology by at least 50 million years and includes the only known fossil zygodactyl footprints. The tracks demonstrate the existence of a Cretaceous bird not currently represented in the body fossil record that occupied a roadrunner (Geococcyx)-like niche, indicating a previously unknown degree of Cretaceous avian morphological and behavioral diversity that presaged later Cenozoic patterns. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.

Phylogeography of Australia’s king brown snake (Pseudechis australis) reveals Pliocene divergence and Pleistocene dispersal of a top predator

King brown snakes or mulga snakes (Pseudechis australis) are the largest and among the most dangerous and wide-ranging venomous snakes in Australia and New Guinea. They occur in diverse habitats, are important predators, and exhibit considerable morphological variation. We infer the relationships and historical biogeography of P. australis based on phylogenetic analysis of 1,249 base pairs from the mitochondrial cytochrome b, NADH dehydrogenase subunit 4 and three adjacent tRNA genes using Bayesian, maximum-likelihood, and maximum-parsimony methods. All methods reveal deep phylogenetic structure with four strongly supported clades comprising snakes from New Guinea (I), localities all over Australia (II), the Kimberleys of Western Australia (III), and north-central Australia (IV), suggesting a much more ancient radiation than previously believed. This conclusion is robust to different molecular clock estimations indicating divergence in Pliocene or Late Miocene, after landbridge dispersal to New Guinea had occurred. While members of clades I, III and IV are medium-sized, slender snakes, those of clade II attain large sizes and a robust build, rendering them top predators in their ecosystems. Genetic differentiation within clade II is low and haplotype distribution largely incongruent with geography or colour morphs, suggesting Pleistocene dispersal and recent ecomorph evolution. Significant haplotype diversity exists in clades III and IV, implying that clade IV comprises two species. Members of clade II are broadly sympatric with members of both northern Australian clades. Thus, our data support the recognition of at least five species from within P. australis (auct.) under various criteria. We discuss biogeographical, ecological and medical implications of our findings.Electronic Supplementary Material  Supplementary material is available in the online version of this article at     

European Atlantic: the hottest oil spill hotspot worldwide

Oil spills caused by maritime transport of petroleum products are still an important source of ocean pollution, especially in main production areas and along major transport routes. We here provide a historical and geographic analysis of the major oil spills (>700 t) since 1960. Spills were recorded from several key marine ecosystems and marine biodiversity hotspots. The past four decades have been characterized by an overall decrease in the number of accidents and tonnes of oil spilled in the sea, but this trend was less distinct in the European Atlantic area. Recent black tides from the Erika and Prestige vessels provided new evidence for the high risk of accidents with serious ecological impact in this area, which according to our analysis is historically the most important oil spill hotspot worldwide. The English Channel and waters around Galicia in Spain were the areas with most accidents. Maritime transport in European Atlantic waters has been predicted to continue increasing. Together with our own results this suggests that, in addition to measures for increased traffic safety, deployment of emergency capacities in the spill hotspot areas may be crucial for a sustainable conservation of sea resources and ecosystems.Electronic Supplementary Material  Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.     

Disulfide connectivity and reduction in pheromone-binding proteins of the gypsy moth, Lymantria dispar

Males of the gypsy moth, Lymantria dispar, are attracted by a pheromone released by females. Pheromones are detected by olfactory neurons housed in specialized sensory hairs located on the antennae of the male moth. Once pheromone molecules enter the sensilla lymph, a highly abundant pheromone-binding protein (PBP) transports the molecule to the sensory neuron. The PBPs are members of the insect odorant-binding protein family, with six conserved cysteine residues. In this study, the disulfide bond connectivities of the pheromone-binding proteins PBP1 and PBP2 from the gypsy moth were found to be cysteines 19–54, 50–109, and 97–118 for PBP1, and cysteines 19–54, 50–110, and 97–119 for PBP2, as determined by cyanylation reactions and cyanogen bromide chemical cleavage. We have discovered that the second disulfide linkage is the most easily reduced of the three, and this same linkage is missing among four cysteine-containing insect odorant-binding proteins (OBPs). We are the first to identify the unique steric and electronic properties of this second disulfide linkage. Electronic Supplementary Material  Supplementary material is available for this article at

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Sound strategy: acoustic aposematism in the bat–tiger moth arms race

The night sky is the venue for an ancient arms race. Insectivorous bats with their ultrasonic sonar exert an enormous selective pressure on nocturnal insects. In response insects have evolved the ability to hear bat cries, to evade their hunting maneuvers, and some, the tiger moths (Arctiidae), to utter an ultrasonic reply. We here determine what it is that tiger moths say to bats. We chose four species of arctiid moths, Cycnia tenera, Euchaetes egle, Utetheisa ornatrix, and Apantesis nais, that naturally differ in their levels of unpalatability and their ability to produce sound. Moths were tethered and offered to free-flying naïve big brown bats, Eptesicus fuscus. The ability of the bats to capture each species was compared to their ability to capture noctuid, geometrid, and wax moth controls over a learning period of 7 days. We repeated the experiment using the single arctiid species E. egle that through diet manipulation and simple surgery could be rendered palatable or unpalatable and sound producing or mute. We again compared the capture rates of these categories of E. egle to control moths. Using both novel learning approaches we have found that the bats only respond to the sounds of arctiids when they are paired with defensive chemistry. The sounds are in essence a warning to the bats that the moth is unpalatable—an aposematic signal.Electronic Supplementary Material  Supplementary material is available for this article at     

Cranial pneumatization and auditory perceptions of the oviraptorid dinosaur <Emphasis Type="Italic">Conchoraptor gracilis</Emphasis> (Theropoda,Maniraptora) from the Late Cretaceous of Mongolia

The distribution of air-filled structures in the craniofacial and neurocranial bones of the oviraptorid ZPAL MgD-I/95, discovered at the Hermiin Tsav locality, Mongolia, is restored. Based on the complete obliteration of most of the cranial sutures, the specimen is identified as an adult individual of Conchoraptor gracilis Barsbold 1986. Except for the orbitosphenoids and epipterygoids, the preserved bones of the neurocranium are hollow. Three types of tympanic recess are present in Conchoraptor, a characteristic shared with troodontids, dromaeosaurids, and avian theropods. The contralateral middle ear cavities are interconnected by the supraencephalic pathway that passes through the dorsal tympanic recesses, the posterodorsal prootic sinuses and the parietal sinus. The spatial arrangements of the middle ear cavity and a derived neurocranial pneumatic system in Conchoraptor indicate enhancements of acoustic perception in the lower-frequency registers and of auditory directionality. We further speculate that this improvement of binaural hearing could be explained as an adaptation required for accurate detection of prey and/or predators under conditions of low illumination. The other potentially pneumatic structures of the Conchoraptor cranium include (1) recessus-like irregularities on the dorsal surface of the nasal and frontal bones (a putative oviraptorid synapomorphy; pos); (2) a subotic recess; (3) a sub-condylar recess; and (4) a posterior condylar recess (pos). Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Comparative morphology and secretion chemistry of the scoli in caterpillars of<Emphasis Type="Italic"> Hyalophora cecropia</Emphasis>

The morphology of the variously coloured scoli (bristle-bearing structures on the integument, producing an exocrine secretion) on caterpillars of Hyalophora cecropia and the secondary chemistry of the discharged secretions have been investigated for the first time and compared. According to our scanning electron microscopic study, the red/orange, yellow and blue coloured groups of these glands differ morphologically. Gas chromatographic-mass spectrometric analyses showed that the patterns of secondary compounds in the respective glandular secretions are also different. Furthermore, the secretion of the penultimate larval instar is chemically distinct from that of the last instar, as are both secretions from the respective haemolymph. The results favour the idea that the differences in scoli colour, morphology and chemistry could affect various predator species differently.Electronic Supplementary Material  Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.