<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.     

The oldest known tracks of web-footed birds from the Lower Cretaceous of South Korea

 We describe the oldest tracks of web-footed birds from the Early Cretaceous in South Korea. The tracks are characterized by a wide divarication angle and a long reversed hallux. The web is semipalmate and restricted to the proximal portion of the three forward digits. The tracks from the Early Cretaceous in South Korea are smaller than those of the Late Cretaceous, therefore confirming the trend of size increasing in the early evolution of birds as shown by skeletal fossils. The discovery of web-footed tracks with abundant non-web-footed tracks indicates that there was a considerable diversification of shore birds as early as the Early Cretaceous. Received: 4 November 1999 / Accepted in revised form: 28 February 2000     

Avian-like attributes of a virtual brain model of the oviraptorid theropod <Emphasis Type="Italic">Conchoraptor gracilis</Emphasis>

An almost complete adult endoneurocranium of Conchoraptor gracilis Barsbold 1986 (Oviraptoridae; ZPAL MgD-I/95), discovered at the Hermiin Tsav locality (the Upper Cretaceous) in Mongolia, is analyzed. A virtual model of the endoneurocranial cavity was derived from CT scans and represents the most complete maniraptoran endocast to date. It displays reduced olfactory bulbs, large cerebral hemispheres in contact with the expanded cerebellum, an epiphysial projection, optic lobes displaced latero-ventrally, presumptive cerebellar folia, enlarged cerebellar auricles, and a deep medulla oblongata with a prominent ventral flexure. Contrary to Archaeopteryx, the shortened olfactory tract and cerebellum overtopping cerebral hemispheres of Conchoraptor resemble conditions in modern birds. Calculating brain mass relative to body mass indicates that Conchoraptor falls within the range of extant birds, whereas Archaeopteryx occupies a marginal position. Most of the endoneurocranial attributes, however, have a less birdlike appearance in Conchoraptor than do corresponding structures in Archaeopteryx and modern birds in which 1) postero-laterally expanded hemispheral domains broadly overlap the optic lobes, 2) the epiphysis projects to the posterior cerebrum, 3) lateral extension of the optic lobes substantially decreases a brain length-to-width ratio, 4) optic lobe and anterior hindbrain are superposed in lateral view, and 5) cerebellar and midbrain compartments are in distinct superposition. The endoneurocranial characteristics of Conchoraptor, taken together, suggest that the animal had a keen sense of vision, balance, and coordination. The data presented in this study do not allow an unambiguous assessment whether the avian-like endoneurocranial characteristics of the flightless Conchoraptor evolved convergently to those of avian theropods, or indicate a derivation of oviraptorosaurs from volant ancestors.     

An exquisitely preserved troodontid theropod with new information on the palatal structure from the Upper Cretaceous of Mongolia

Troodontidae is a clade of small-bodied theropod dinosaurs. A new troodontid, Gobivenator mongoliensis gen. et sp. nov., is described based on the most complete skeleton of a Late Cretaceous member of this clade presently known, from the Campanian Djadokhta Formation in the central Gobi Desert. G. mongoliensis is different from other troodontids in possessing a pointed anterior end of the fused parietal and a fossa on the surangular in front of the posterior surangular foramen. The skull was superbly preserved in the specimen and provides detailed information of the entire configuration of the palate in Troodontidae. Overall morphology of the palate in Gobivenator resembles those of dromaeosaurids and Archaeopteryx, showing an apparent trend of elongation of the pterygoid process of the palatine and reduction of the pterygopalatine suture toward the basal Avialae. The palatal configuration suggests that the skull of Gobivenator would have been akinetic but had already acquired prerequisites for later evolution of cranial kinesis in birds, such as the loss of the epipterygoid and reduction in contact areas among bones.     

Molecular phylogeny of cycads inferred from rbcL sequences

The chloroplast gene rbcL was sequenced to elucidate the evolution of the gymnosperm plant order Cycadales. In accordance with traditional systematics, the order Cycadales and the corresponding genera cluster as monophyletic clades. Among them, the genus Cycas forms a basal group. The genetic distances within the genus Encephalartos and between the sister groups Encephalartos, Lepidozamia and Macrozamia, are unexpectedly small, suggesting that the extant species are the result of Miocene and Pliocene speciation. Their distribution in Africa or Australia, respectively, may therefore rather be due to long-distance dispersal than to Cretaceous continental drift, as had previously been assumed. The rbcL sequences also indicate that the colonisation of Madagascar by Cycas thouarsii occurred only recently as the sequences of C. thouarsii and Cycas rumphii from Indonesia are identical. In contrast, the divergence of the Cycadaceae and Zamiaceae apparently occurred in the Mesozoic.     

The origin and early evolution of birds: discoveries,disputes, and perspectives from fossil evidence

The study of the origin and early evolution of birds has never produced as much excitement and public attention as in the past decade. Well preserved and abundant new fossils of birds and dinosaurs have provided unprecedented new evidence on the dinosaurian origin of birds, the arboreal origin of avian flight, and the origin of feathers prior to flapping flight. The Mesozoic avian assemblage mainly comprises two major lineages: the prevalent extinct group Enantiornithes, and the Ornithurae, which gave rise to all modern birds, as well as several more basal taxa. Cretaceous birds radiated into various paleoecological niches that included fish- and seed-eating. Significant size and morphological differences and variation in flight capabilities, ranging from gliding to powerful flight among early birds, highlight the diversification of birds in the Early Cretaceous. There is little evidence, however, to support a Mesozoic origin of modern avian groups. Controversy and debate, nevertheless, surround many of these findings, and more details are needed to give a better appreciation of the significance of these new discoveries.     

New Early Cretaceous fossil from China documents a novel trophic specialization for Mesozoic birds

We report on a new Mesozoic bird, Longirostravis hani, from the Early Cretaceous Jehol Biota of northeastern China. The new taxon has a long, slender rostrum and mandible, and a small number of rostralmost teeth. Postcranial characters such as a furcular ramus wider ventrally than dorsally, a centrally concave proximal margin of the humeral head, and a minor metacarpal that projects distally more than the major metacarpal, support the placement of Longirostravis within euenantiornithine Enantiornithes, the most diverse clade of Mesozoic birds. The morphology of the skull, however, suggests that Longirostravis had a probing feeding behavior, a specialization previously unknown for Enantiornithes. Indeed, this discovery provides the first evidence in support of the existence of such a foraging behavior among basal lineages of Mesozoic birds.     

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.

Bristle-like integumentary structures at the tail of the horned dinosaur Psittacosaurus

A specimen of the horned dinosaur Psittacosaurus from the early Cretaceous of China is described in which the integument is extraordinarily well-preserved. Most unusual is the presence of long bristle-like structures on the proximal part of tail. We interpret these structures as cylindrical and possibly tubular epidermal structures that were anchored deeply in the skin. They might have been used in display behavior and especially if one assumes that they were colored, they may have had a signal function. At present, there is no convincing evidence which shows these structures to be homologous to the structurally different integumentary filaments of theropod dinosaurs. Independent of their homology, however, the discovery of bristle-like structures in Psittacosaurus is of great evolutionary significance since it shows that the integumentary covering of at least some dinosaurs was much more complex than has ever been previously imagined.     

The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the early cretaceous of Laos

Spinosaurids are among the largest and most specialized carnivorous dinosaurs. The morphology of their crocodile-like skull, stomach contents, and oxygen isotopic composition of the bones suggest they had a predominantly piscivorous diet. Even if close relationships between spinosaurids and Middle Jurassic megalosaurs seem well established, very little is known about the transition from a generalized large basal tetanuran to the specialized morphology of spinosaurids. Spinosaurid remains were previously known from the Early to Late Cretaceous of North Africa, Europe, and South America. Here, we report the discovery of a new spinosaurid theropod from the late Early Cretaceous Savannakhet Basin in Laos, which is distinguished by an autapomorphic sinusoidal dorsosacral sail. This new taxon, Ichthyovenator laosensis gen. et sp. nov., includes well-preserved and partially articulated postcranial remains. Although possible spinosaurid teeth have been reported from various Early Cretaceous localities in Asia, the new taxon I. laosensis is the first definite record of Spinosauridae from Asia. Cladistic analysis identifies Ichthyovenator as a member of the sub-clade Baryonychinae and suggests a widespread distribution of this clade at the end of the Early Cretaceous. Chilantaisaurus tashouikensis from the Cretaceous of Inner Mongolia, and an ungual phalanx from the Upper Jurassic of Colorado are also referred to spinosaurids, extending both the stratigraphical and geographical range of this clade.     

A gravid lizard from the Cretaceous of China and the early history of squamate viviparity

Although viviparity is most often associated with mammals, roughly one fifth of extant squamate reptiles give birth to live young. Phylogenetic analyses indicate that the trait evolved more than 100 times within Squamata, a frequency greater than that of all other vertebrate clades combined. However, there is debate as to the antiquity of the trait and, until now, the only direct fossil evidence of squamate viviparity was in Late Cretaceous mosasauroids, specialised marine lizards without modern equivalents. Here, we document viviparity in a specimen of a more generalised lizard, Yabeinosaurus, from the Early Cretaceous of China. The gravid female contains more than 15 young at a level of skeletal development corresponding to that of late embryos of living viviparous lizards. This specimen documents the first occurrence of viviparity in a fossil reptile that was largely terrestrial in life, and extends the temporal distribution of the trait in squamates by at least 30 Ma. As Yabeinosaurus occupies a relatively basal position within crown-group squamates, it suggests that the anatomical and physiological preconditions for viviparity arose early within Squamata.     

Gastroliths in<Emphasis Type="Italic"> Yanornis</Emphasis>: an indication of the earliest radical diet-switching and gizzard plasticity in the lineage leading to living birds?

Yanornis martini is an Early Cretaceous basal ornithurine bird. Its fish-eating diet was previously recognized from a discrete mass of disarticulated fish remains discovered in its abdominal region. A new complete and articulated specimen of Yanornis martini preserves abundant in-situ gastroliths such as have been associated with a herbivorous diet. We suggest that the occurrence of gastroliths in this specimen, fish remains in a second, and the lack of gastroliths in three others, is consistent with diet-switching in Yanornis martini. Incompatibility of the preserved data with explanations of the grit as an artifact of preservation or result of accidental ingestion is discussed. This discovery indicates the earliest presence of intermittent diet change (and associated gizzard plasticity) observed in extant birds seasonally and in response to changes in available food sources.     

The hand of birds revealed by early ostrich embryos

The problem of resolving the homology of the digits of the avian hand has been framed as a conflict between paleontological and embryological evidence, the former thought to support a hand composed of digits I, II, III, because of similarity of the phalangeal formulae of the earliest known bird Archaeopteryx to that of Mesozoic pentadactyl archosaurs, while embryological evidence has traditionally favored a II, III, IV avian hand. We have identified the critical developmental period for the major features of the avian skeleton in a primitive bird, the ostrich. Analysis of digit anlagen in the avian hand has revealed those for digits/metacarpals I and V, thus confirming previous embryological studies that indirectly suggested that the avian hand comprises digits II, III, IV, and was primitively pentadactyl.     

The first multituberculate mammal from India

Mesozoic deposits of the former Gondwanaland are depauperate in early mammals, in general, and multituberculate mammals, in particular. Until now, the oldest multituberculate mammals known from the Gondwanan continents come from the Early Cretaceous of Morocco, NW Africa. Here, we report the presence of a new multituberculate mammal, Indobaatar zofiae gen. et sp. nov., from the Lower/Middle Jurassic Kota Formation, Pranhita-Godavari valley in peninsular India. This is the first record of a multituberculate from the Mesozoic rocks of India and possibly predates the oldest known multituberculates from Gondwanan continents. The new specimen, representing an upper premolar (P⁴), compares well with the upper premolar morphology of Eobaatariinae multituberculates known from the Early Cretaceous of Mongolia, China, England, and Spain. Together with the recent findings of cimolodontan multituberculates from the Early Cretaceous of Australia and Late Cretaceous of South America, the new discovery indicates a wide temporal and spatial distribution for multituberculate mammals in the former Gondwanaland.     

Cycads: their evolution,toxins, herbivores and insect pollinators

Palaeobiological evidence indicates that gymnosperms were wind-pollinated and that insect pollination began in angiosperms in the Lower Cretaceous (ca. 135 mya) leading to close associations between higher plants and their pollinators. Cycads, which were widespread and pervasive throughout the Mesozoic (250-65 mya) are among the most primitive living seed-plants found today. Because pollination by beetles and by thrips has now been detected in several modern cycads, it is attractive to speculate that some insects and cycads had already developed similar mutualistic interactions in the Triassic (250-205 mya), long before the advent of angiosperms. We also draw attention to another key factor in this insect-plant relationship, namely secondary, defensive plant substances which must always have controlled interspecific interactions. Cycads mainly produce toxic azoglucosides and neurotoxic non-protein amino acids (e.g. BMAA), which apparently are crucial elements in the development and maintenance of mutualism (pollination) and parasitism (herbivory) by cycad-linked herbivores. We now add new results on the uptake and storage of the main toxin, cycasin, of the Mexican cycad Zamia furfuracea by its pollinator, the weevil Rhopalotria mollis, and by a specialist herbivore of Zamia integrifolia, the aposematic Atala butterfly Eumaeus atala.     

Split-second escape decisions in blue tits (Parus caeruleus)

Bird mortality is heavily affected by birds of prey. Under attack, take-off is crucial for survival and even minor mistakes in initial escape response can have devastating consequences. Birds may respond differently depending on the character of the predator's attack and these split-second decisions were studied using a model merlin (Falco columbarius) that attacked feeding blue tits (Parus caeruleus) from two different attack angles in two different speeds. When attacked from a low attack angle they took off more steeply than when attacked from a high angle. This is the first study to show that escape behaviour also depends on predator attack speed. The blue tits responded to a high-speed attack by dodging sideways more often than when attacked at a low speed. Escape speed was not significantly affected by the different treatments. Although they have only a split-second before escaping an attack, blue tits do adjust their escape strategy to the prevailing attack conditions.     

Food remains in Confuciusornis sanctus suggest a fish diet

Despite hundreds of excellent fossils of Confuciusornis, the most abundant group of birds in the Early Cretaceous, ‘Jehol Biota’ in China, there is yet no indication of the food choice of these birds. Here, we describe fish remains preserved in the alimentary system of a specimen of Confuciusornis sanctus from the Jiufotang Formation. This find is about five million years younger than all previously published confuciusornithid birds from the Yixian Formation. Although it is unknown how common fish was in the diet of Confuciusornis, the find does not support previous hypotheses that it fed on plants or grain.     

Europe's last Mesozoic bird

Birds known from more than isolated skeletal elements are rare in the fossil record, especially from the European Mesozoic. This paucity has hindered interpretations of avian evolution immediately prior to, and in the aftermath of, the Cretaceous-Tertiary (K-T) extinction event. We report on a specimen of a large ornithurine bird (closely related to Ichthyornis) from the uppermost Cretaceous (Maastricht Formation) of Belgium. This is the first record of a bird from these historic strata and the only phylogenetically informative ornithurine to be recovered from the Mesozoic of Europe. Because this new specimen was collected from 40 m below the K-T boundary (approximate age of 65.8 Ma), it is also the youngest non-neornithine (=non-modern) bird known from anywhere in the world.     

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.     

Phylogeny of cooperatively breeding cuckoos (Cuculidae,Crotophaginae) based on mitochondrial gene sequences

The Crotophaginae is a subfamily of New World cuckoos comprising the monotypic genus Guira and three ani species (Crotophaga). All exhibit a rare form of cooperative breeding known as plural female joint-nesting, whereby two or more females lay eggs in a single nest. I reconstructed the phylogeny of Crotophaginae using the mitochondrial genes cytochrome oxidase I, II, and III, ATPase 6 and 8, and cytochrome b. The subfamily was monophyletic, implying a single origin of cooperative breeding in New World cuckoos. Crotophaga was also monophyletic with Guira as its sister taxon. Within Crotophaga, the smooth-billed (C. ani) and groove-billed (C. sulcirostris) anis formed the internal clade with the greater ani (C. major) basal to this pair. This phylogeny is consistent with differences in reproductive patterns and social organization exhibited by crotophagine cuckoos, and will serve as a framework for future study of the evolution of cooperative breeding in this subfamily.