Minute theropod eggs and embryo from the Lower Cretaceous of Thailand and the dinosaur-bird transition

We report on very small fossil eggs from the Lower Cretaceous of Thailand, one of them containing a theropod embryo, which display a remarkable mosaic of characters. While the surficial ornamentation is typical of non-avian saurischian dinosaurs, the three-layered prismatic structure of the eggshell is currently known only in extant and fossil eggs associated with birds. These eggs, about the size of a goldfinch's, mirror at the reproductive level the retention of small body size that was paramount in the transition from non-avian theropods to birds. The egg-layer may have been a small feathered theropod similar to those recently found in China.     

A large Cretaceous theropod from Patagonia, Argentina, and the evolution of carcharodontosaurids

The Cretaceous Carcharodontosauridae is the latest clade of carnosaurs, including the largest predatory dinosaurs yet recorded. Albeit spectacular for their size, the skeletal anatomy of these theropods remains poorly-known, and their diversity was until recently restricted to two Cenomanian species: the highly derived Giganotosaurus carolinii, from southern South America, and the incompletely known Carcharodontosaurus saharicus, from northern Africa. Here we describe an older and basal member of the group, Tyrannotitan chubutensis gen. et sp. nov., from Aptian strata of Patagonia, Argentina. The new taxon gives new insights into the systematics and evolution of carcharodontosaurids and offers a better understanding of the evolution of Southern theropod faunas. We suggest that carcharodontosaurids radiated in Gondwana sharing with spinosaurids the role of top-predators until their extinction in Cenomanian–Turonian times. During this interval, the diplodocoid sauropods and giant titanosaurians went extinct (probably as part of a global-scale crisis), and the smaller abelisaurid theropods took dominance, reigning until the end of the Cretaceous. Electronic Supplementary Material is available.Electronic Supplementary Information  Supplementary material is available for this article at     

First ceratosaurian dinosaur from Australia

The basal theropod dinosaur clade Ceratosauria, and its subclade Abelisauroidea, is characteristic of late Mesozoic terrestrial vertebrate faunas in western Gondwana (South America, Africa, Madagascar, and India) and Europe. Yet unambiguous records of ceratosaurs have hitherto been absent from Australia, where the theropod assemblage appears to include several typically Laurasian clades. Here, we report the first evidence of ceratosaurs (and potentially abelisauroids) from eastern Gondwana––a diagnostic astragalocalcaneum from the Aptian (121–125?Ma) of Victoria, Australia. Ceratosauria thus occurred in both western and eastern Gondwana during the Early Cretaceous. This fossil adds to the poorly known dinosaur fauna of Australia, a major clade of basal theropods, emphasising that its mid-Cretaceous theropod diversity was surprisingly cosmopolitan despite relative geographic isolation, including clades that have been thought to be typical of both Gondwana and Laurasia––Ceratosauria, Spinosauridae, Carcharodontosauria, Tyrannosauroidea, and Deinonychosauria. Such a contemporaneous association of theropod clades is unknown from other Gondwanan continents and questions the views that the late Mesozoic dinosaur fauna of Australia was dominated by Gondwanan or Laurasian elements, extreme isolation, relictualism, and/or novelty as a ‘centre of origin’. The cosmopolitan theropod fauna of Australia probably reflects the global distribution of these clades early in their history, prior to significant continental breakup.     

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.

The evolutionary continuum of limb function from early theropods to birds

The bipedal stance and gait of theropod dinosaurs evolved gradually along the lineage leading to birds and at some point(s), flight evolved. How and when did these changes occur? We review the evidence from neontology and palaeontology, including pectoral and pelvic limb functional morphology, fossil footprints/trackways and biomechanical models and simulations. We emphasise that many false dichotomies or categories have been applied to theropod form and function, and sometimes, these impede research progress. For example, dichotomisation of locomotor function into ‘non-avian’ and ‘avian’ modes is only a conceptual crutch; the evidence supports a continuous transition. Simplification of pelvic limb function into cursorial/non-cursorial morphologies or flexed/columnar poses has outlived its utility. For the pectoral limbs, even the classic predatory strike vs. flight wing-stroke distinction and separation of theropods into non-flying and flying—or terrestrial and arboreal—categories may be missing important subtleties. Distinguishing locomotor function between taxa, even with quantitative approaches, will always be fraught with ambiguity, making it difficult to find real differences if that ambiguity is properly acknowledged. There must be an ‘interpretive asymptote’ for reconstructing dinosaur limb function that available methods and evidence cannot overcome. We may be close to that limit, but how far can it be stretched with improved methods and evidence, if at all? The way forward is a combination of techniques that emphasises integration of neontological and palaeontological evidence and quantitative assessment of limb function cautiously applied with validated techniques and sensitivity analysis of unknown variables.     

Dietary characterization of the hominoid Khoratpithecus (Miocene of Thailand): evidence from dental topographic and microwear texture analyses

The genus Khoratpithecus, a hominoid thought to be related to the orangutan lineage, is represented by two known fossil species K. chiangmuanensis and K. piriyai. Both were discovered in Southeast Asia (Thailand) and are dated to the Middle and Late Miocene, respectively. In this study, dental topographic and microwear texture analyses were used to examine molars from both of these species, with the goal of understanding their dietary preferences. Although sample sizes are small for Khoratpithecus, available data are compared to that collected for extant apes. Environmental evidence, such as botanical remains and sedimentological data, is also considered for comparisons with dietary reconstruction. Results from dental topographic analysis suggest that the two fossil species were better adapted to a diet of fruits than to one of leaves, much like the living orangutan or chimpanzee. Results from microwear texture analysis further support this, suggesting that Khoratpithecus preferred soft fruits to hard fruits or seeds. And finally, the botanical and sedimentological evidence point to environments for Khoratpithecus that would have been compatible with a fruit-eating species. Given the small sample sizes available for analysis, however, definitive judgments are not yet possible at this time.     

The last polar dinosaurs: high diversity of latest Cretaceous arctic dinosaurs in Russia

A latest Cretaceous (68 to 65 million years ago) vertebrate microfossil assemblage discovered at Kakanaut in northeastern Russia reveals that dinosaurs were still highly diversified in Arctic regions just before the Cretaceous–Tertiary mass extinction event. Dinosaur eggshell fragments, belonging to hadrosaurids and non-avian theropods, indicate that at least several latest Cretaceous dinosaur taxa could reproduce in polar region and were probably year-round residents of high latitudes. Palaeobotanical data suggest that these polar dinosaurs lived in a temperate climate (mean annual temperature about 10°C), but the climate was apparently too cold for amphibians and ectothermic reptiles. The high diversity of Late Maastrichtian dinosaurs in high latitudes, where ectotherms are absent, strongly questions hypotheses according to which dinosaur extinction was a result of temperature decline, caused or not by the Chicxulub impact.     

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.     

Direct evidence of hybodont shark predation on Late Jurassic ammonites

Sharks are known to have been ammonoid predators, as indicated by analysis of bite marks or coprolite contents. However, body fossil associations attesting to this predator–prey relationship have never been described so far. Here, I report a unique finding from the Late Jurassic of western France: a complete specimen of the Kimmeridgian ammonite Orthaspidoceras bearing one tooth of the hybodont shark Planohybodus. Some possible tooth puncture marks are also observed. This is the first direct evidence of such a trophic link between these two major Mesozoic groups, allowing an accurate identification of both organisms. Although Planohybodus displays a tearing-type dentition generally assumed to have been especially adapted for large unshelled prey, our discovery clearly shows that this shark was also able to attack robust ammonites such as aspidoceratids. The direct evidence presented here provides new insights into the Mesozoic marine ecosystem food webs.     

Diversity trends in Neogene European ungulates and rodents: large-scale comparisons and perspectives

The ungulate and rodent fossil records are often used independently to understand mammalian evolutionary history. Few studies have directly compared both records over long time periods and large geographic areas. Here, we compiled two datasets of European fossil localities containing rodents and/or ungulates over 20 My (Early Miocene–Early Pliocene) and processed the data with the same methodology. We counted the raw diversity and calculated a measure of evenness (Pielou’s index). After controlling for potential biases on diversity estimators, we identify the evenness index as a more reliable estimator bringing interesting insights into the way both mammal groups are structured by biotic or abiotic factors. In this study, we consider that an uneven distribution of the species richness among families, when only some families successfully diversify within the “continental-scale community”, represents a lower adaptability of this community to the environmental context. Pielou’s index is used to estimate this adaptability through time. The responses of ungulates and rodents to environmental changes are very divergent, especially facing the climatic changes known since the Middle Miocene. The general patterns suggest that rodent broad-scale assemblages are affected by all kinds of perturbations, even short biotic and abiotic events, but show a better adaptability when facing long-term abiotic changes. Unlike rodents, the ungulate assemblages show more stability in periods of relative environmental stability but show less adaptability to long-term climatic changes. Life-history traits of mammals can help explain patterns of diversity and biogeography at different spatial scales and may probably partly explain the opposite patterns evidenced here.     

The mammal assemblage of the hominid site TM266 (Late Miocene, Chad Basin): ecological structure and paleoenvironmental implications

Characterizing the paleoenvironmental context of the first hominids is a key issue for understanding their behavioral and morphological evolution. The present study aims at reconstructing the paleoenvironment of the TM266 vertebrate assemblage (Toros-Menalla, Northern Chad) that yielded the earliest known hominid Sahelanthropus tchadensis (7 Ma). For the first time, a quantitative analysis is carried out on the fossil mammal assemblage associated with that hominid. Two complementary approaches were applied: (1) the analysis of the relative abundances of taxa and their habitat preferences; and (2) the study of the distribution of taxa within three meaningful ecovariables: locomotion, feeding preferences, and body mass. The resulting taxonomic and paleoecological structures are used to reconstruct the diversity and the relative extent of the habitats in that part of northern Chad seven million years ago. The paleoenvironment was composed of open areas with dry and humid grasslands, prevailing over wooded habitats. Water was also widely available as freshwater bodies and certainly swamps. It appears that the high habitat diversity of the landscape is a common feature among paleoenvironments associated with early hominids.     

Can the envisaged reductions of fossil fuel CO<Subscript>2</Subscript> emissions be detected by atmospheric observations?

The lower troposphere is an excellent receptacle, which integrates anthropogenic greenhouse gases emissions over large areas. Therefore, atmospheric concentration observations over populated regions would provide the ultimate proof if sustained emissions changes have occurred. The most important anthropogenic greenhouse gas, carbon dioxide (CO(2)), also shows large natural concentration variations, which need to be disentangled from anthropogenic signals to assess changes in associated emissions. This is in principle possible for the fossil fuel CO(2) component (FFCO(2)) by high-precision radiocarbon ((14)C) analyses because FFCO(2) is free of radiocarbon. Long-term observations of (14)CO(2) conducted at two sites in south-western Germany do not yet reveal any significant trends in the regional fossil fuel CO(2) component. We rather observe strong inter-annual variations, which are largely imprinted by changes of atmospheric transport as supported by dedicated transport model simulations of fossil fuel CO(2). In this paper, we show that, depending on the remoteness of the site, changes of about 7-26% in fossil fuel emissions in respective catchment areas could be detected with confidence by high-precision atmospheric (14)CO(2) measurements when comparing 5-year averages if these inter-annual variations were taken into account. This perspective constitutes the urgently needed tool for validation of fossil fuel CO(2) emissions changes in the framework of the Kyoto protocol and successive climate initiatives.     

Spine anatomy reveals the diversity of catfish through time: a case study of Synodontis (Siluriformes)

Synodontis (Mochokidae, Siluriformes) is a freshwater catfish endemic to Africa. The 118 extant species are present in almost all hydrographic basins. Some species are restricted to a single stream, whereas others have a vast distribution. Synodontis is known in the fossil record since the Miocene, and its history depends on the connections among African basins through time. The identification of species in the fossil record is essential to reconstruct this historical pattern. Catfish pectoral and dorsal spines are robust, they preserve well and they form most of the fossil remains for the genus Synodontis. Unfortunately, the criteria for the identification of extant Synodontis species are not applicable to fossil specimens. Here, we define 11 original morphological characters that permit to discriminate four extant species from the Chad-Chari hydrographic system. Six of these characters are defined on pectoral spines and five on dorsal spines. We then show that these characters can be used successfully for identifying fossil specimens. In particular, we present a case study in which we identify Synodontis cf. schall and Brachysynodontis cf. batensoda in the hominid-bearing sector Toros-Menalla (Late Miocene, northern Chad). We show that spine anatomy can be a powerful tool to recognise catfish species through time and thus to identify historical diversity pattern.     

Soil biodiversity for agricultural sustainability

We critically highlight some evidence for the importance of soil biodiversity to sustaining (agro-)ecosystem functioning and explore directions for future research. We first deal with resistance and resilience against abiotic disturbance and stress. There is evidence that soil biodiversity does confer stability to stress and disturbance, but the mechanism is not yet fully understood. It appears to depend on the kind of stress and disturbance and on the combination of stress and disturbance effects. Alternatively, community structure may play a role. Both possible explanations will guide further research. We then discuss biotic stress. There is evidence that soil microbial diversity confers protection against soil-borne disease, but crop and soil type and management also play a role. Their relative importance as well as the role of biodiversity in multitrophic interactions warrant further study. Henceforth, we focus on the effects of plant and soil biodiversity on nutrient and water use efficiencies as important ecological functions in agroecosystems. The available evidence suggests that mycorrhizal diversity positively contributes to nutrient and, possibly, water use efficiency. Soil fauna effects on nutrient and water use efficiencies are also apparent, but diversity effects may be indirect, through effects on soil structure. We present a conceptual diagram relating plant and soil biodiversity with soil structure and water and nutrient use efficiencies as a framework for future studies. We then consider how cropping systems design and management are interrelated and how management options might be interfaced with farmers’ knowledge in taking management decisions. Finally, we attempt to express some economic benefits of soil biodiversity to society as part of a wider strategy of conserving and using agrobiodiversity.     

A juvenile coelurosaurian theropod from China indicates arboreal habits

Here we report an unequivocal arboreal coelurosaur, Epidendrosaurus ningchengensis gen. et sp. nov. This juvenile coelurosaur's third manual digit is extremely elongated, distinctively different from that of other known dinosaurs and birds. It represents certainly a type of adaptation previously unreported from the Mesozoic although the exact function of the third manual digit is unclear. The relatively long forelimb, penultimate phalanx of manual digit II, and pedal penultimate phalanges, are interpreted as evidence for the arboreal habit of Epidendrosaurus. Because Epidendrosaurus is more similar to advanced birds in some arboreal features than to Archaeopteryx, we suggest that the initial appearance of tree-adaptation in theropods was probably not directly related to flight but to other functions, such as seeking food or escaping from predators. Electronic Supplementary Material is available if you access this article at http://dx.doi.org/10.1007/s00114-002-0353-8. On that page (frame on the left side), a link takes you directly to the supplementary material.     

Skeletal remains of a diminutive primate from the Paleocene of Germany

Most living mammal orders, including our own, started their career during the first 10 million years of the Cenozoic, the Age of Mammals. The fossil record documents that early Paleogene adaptive radiations of various clades included tiny species of the size of living shrews. Remains of particularly diminutive limb bones are described from the late Paleocene site of Walbeck, Sachsen-Anhalt. Discovered in 1939, it has remained the only known Paleocene mammal-bearing locality from Germany. The remains are referred to the family Adapisoriculidae, which is considered on the basis of the present postcranial evidence to represent plesiadapiform primates rather than alleged lipotyphlan insectivores as previously proposed. The Walbeck fossils compete with the Early Eocene species Toliapina vinealis from Europe and Picromomys petersonorum from North America for the status of the smallest known primate, fossil and living. Their estimated body weights are as small as 10 g. The limb bones show features related to enhanced flexion at the elbow and hip joint, suggesting arboreal habits and environments such as terminal branches. The diminutive size and tooth morphology suggest feeding on small insects and other invertebrates. Postcranials are important to assess early radiations, such tiny specimens as the present ones are extremely scarce in the fossil record, however.     

生物卵化石研究进展简述

卵生是动物界最普遍的繁殖方式,卵的孵化对繁殖季节突发的气候变化响应敏感,孵化成败又与生物种群数量变动和群落演替密切相关,因此卵化石具有重要的古生物学、古气候学、古生态学研究价值。但由于卵生动物种类十分丰富,卵形态特征多样,研究领域广泛,有必要总结梳理卵化石的研究进展。本文通过对主要卵生动物类群卵的形态特征,以及卵化石在古生物学、古气候学、古生态学方面研究进展的回顾,发现卵化石的研究主要集中在脊椎动物类群,为脊椎动物演化提供了重要证据,但是由于脊椎动物卵化石通常难以建立连续的长序列记录,其古生态学和古气候学研究极为不足。然而,无脊椎动物卵化石由于数量丰富,具有统计意义,其在古生态学、古气候学研究领域具有重要意义,但是其在古生物学,包括种属鉴定等方面的研究需要加强。在无脊椎动物中,蜗牛及其卵化石在黄土-古土壤地层中保存丰富且地层分布连续,最有可能建立卵化石的长序列。通过现代过程研究,明确蜗牛卵的气候指示意义,建立第四纪蜗牛卵化石长序列,发挥蜗牛卵化石的古气候学和古生态学研究价值,是亟需开展的研究工作。     

Possible postcranial pneumaticity in the last common ancestor of birds and crocodilians: evidence from Erythrosuchus and other Mesozoic archosaurs

Birds and crocodilians (extant archosaurs) have differing, distinctive morphologies. Birds have respiratory airsacs with diverticula that pneumatize the postcranial skeleton, a feature absent in crocodilians. Bony correlates of pneumatic sinuses are known in the vertebrae of some non-avian dinosaurs and in pterosaurs - taxa more closely related to birds than crocodilians. This and the apparent absence of pneumatic postcranial bones in fossil archosaurs more closely related to crocodilians than to birds, has been interpreted as evidence that postcranial pneumaticity is a derived character of birds and their nearest fossil relatives. The presence of apparent osteological correlates of postcranial pneumaticity is here reported in some non-crown-group archosaurs, and some of the fossil taxa more closely related to crocodilians than to birds. This suggests that the last common ancestor of birds and crocodilians might have had a pneumatized postcranium, and that the absence of this feature in crocodilians might be derived.     

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     

Direct evidence of trophic interactions among apex predators in the Late Triassic of western North America

Hypotheses of feeding behaviors and community structure are testable with rare direct evidence of trophic interactions in the fossil record (e.g., bite marks). We present evidence of four predation, scavenging, and/or interspecific fighting events involving two large paracrocodylomorphs (=‘rauisuchians’) from the Upper Triassic Chinle Formation (～220–210 Ma). The larger femur preserves a rare history of interactions with multiple actors prior to and after death of this ～8–9-m individual. A large embedded tooth crown and punctures, all of which display reaction tissue formed through healing, record evidence of a failed attack on this individual. The second paracrocodylomorph femur exhibits four unhealed bite marks, indicating the animal either did not survive the attack or was scavenged soon after death. The combination of character states observed (e.g., morphology of the embedded tooth, ‘D’-shaped punctures, evidence of bicarination of the marking teeth, spacing of potentially serial marks) indicates that large phytosaurs were actors in both cases. Our analysis of these specimens demonstrates phytosaurs targeted large paracrocodylomorphs in these Late Triassic ecosystems. Previous distinctions between ‘aquatic’ and ‘terrestrial’ Late Triassic trophic structures were overly simplistic and built upon mistaken paleoecological assumptions; we show they were intimately connected at the highest trophic levels. Our data also support that size cannot be the sole factor in determining trophic status. Furthermore, these marks provide an opportunity to start exploring the seemingly unbalanced terrestrial ecosystems from the Late Triassic of North America, in which large carnivores far outnumber herbivores in terms of both abundance and diversity.