The skull evolution of oviraptorosaurian dinosaurs: the role of niche partitioning in diversification.

Oviraptorosaurs are bird-like theropod dinosaurs that thrived in the final pre-extinction ecosystems during the latest Cretaceous, and the beaked, toothless skulls of derived species are regarded as some of the most peculiar among dinosaurs. Their aberrant morphologies are hypothesized to have been caused by rapid evolution triggered by an ecological/biological driver, but little is known about how their skull shapes and functional abilities diversified. Here, we use quantitative techniques to study oviraptorosaur skull form and mandibular function. We demonstrate that the snout is particularly variable, that mandibular form and upper/lower beak form are significantly correlated with phylogeny, and that there is a strong and significant correlation between mandibular function and mandible/lower beak shape, suggesting a form-function association. The form-function relationship and phylogenetic signals, along with a moderate allometric signal in lower beak form, indicate that similar mechanisms governed beak shape in oviraptorosaurs and extant birds. The two derived oviraptorosaur clades, oviraptorids and caenagnathids, are significantly separated in morphospace and functional space, indicating that they partitioned niches. Oviraptorids coexisting in the same ecosystem are also widely spread in morphological and functional space, suggesting that they finely partitioned feeding niches, whereas caenagnathids exhibit extreme disparity in beak size. The diversity of skull form and function was likely key to the diversification and evolutionary success of oviraptorosaurs in the latest Cretaceous.

A new alvarezsaurid dinosaur from the Nemegt Formation of Mongolia.

Alvarezsaurid diversity has been markedly increased by recent discoveries from China. However, the number of alvarezsaurid specimens in the Nemegt Formation of Mongolia remained low since the initial report on Mononykus olecranus in 1993. Here we report three new alvarezsaurid specimens from this formation, which were associated with each other and also with multiple oviraptorid skeletons in a small multi-species assemblage. Two of the alvarezsaurid specimens represent a new taxon, Nemegtonykus citus gen. et sp. nov., which is mainly distinguished from other alvarezsaurids by the first sacral vertebra with a subtrapezoidal lamina, the second sacral centrum which is directly co-ossified with ilium, the posterodorsally oriented postacetabular process of ilium, and partial co-ossification between metatarsals II and IV. The other specimen is very similar to M. olecranus in morphology and referred to cf. Mononykus sp. Our phylogenetic analysis recovered Nemegtonykus as a parvicursorine forming a polytomy with several other taxa from the Gobi Desert. The presence of three alvarezsaurid individuals in the same locality indicates that the abundance of alvarezsaurids have been greatly underestimated in the Nemegt dinosaur faunas.

A new baby oviraptorid dinosaur (Dinosauria: Theropoda) from the Upper Cretaceous Nemegt Formation of Mongolia.

Recent discoveries of new oviraptorosaurs revealed their high diversity from the Cretaceous Period in Asia and North America. Particularly, at the family level, oviraptorids are among the most diverse theropod dinosaurs in the Late Cretaceous of Mongolia and China. A new oviraptorid dinosaur Gobiraptor minutus gen. et sp. nov. from the Upper Cretaceous Nemegt Formation is described here based on a single holotype specimen that includes incomplete cranial and postcranial elements. The most prominent characters of Gobiraptor are its thickened rostrodorsal end of the mandibular symphysis and a rudimentary lingual shelf on each side of the dentary. Each lingual shelf is lined with small occlusal foramina and demarcated by a weakly developed lingual ridge. This mandibular morphology of Gobiraptor is unique among oviraptorids and likely to be linked to a specialized diet that probably included hard materials, such as seeds or bivalves. The osteohistology of the femur of the holotype specimen indicates that the individual was fairly young at the time of its death. Phylogenetic analysis recovers Gobiraptor as a derived oviraptorid close to three taxa from the Ganzhou region in southern China, but rather distantly related to other Nemegt oviraptorids which, as the results of recent studies, are also not closely related to each other. Gobiraptor increases diversity of oviraptorids in the Nemegt Formation and its presence confirms the successful adaptation of oviraptorids to a mesic environment.

Incubation behaviours of oviraptorosaur dinosaurs in relation to body size.

Most birds sit on their eggs during incubation, a behaviour that likely evolved among non-avian dinosaurs. Several 'brooding' specimens of smaller species of oviraptorosaurs and troodontids reveal these non-avian theropods sat on their eggs, although little is known of incubation behaviour in larger theropod species. Here we examine egg clutches over a large body size range of oviraptorosaurs in order to understand the potential effect of body size on incubation behaviour. Eggshell porosity indicates that the eggs of all oviraptorosaurs were exposed in the nest, similar to brooding birds. Although all oviraptorosaur clutches consist of radially arranged eggs in a ring configuration, clutch morphology varies in that the central opening is small or absent in the smallest species, becomes significantly larger in larger species, and occupies most of the nest area in giant species. Our results suggest that the smallest oviraptorosaurs probably sat directly on the eggs, whereas with increasing body size more weight was likely carried by the central opening, reducing or eliminating the load on the eggs and still potentially allowing for some contact during incubation in giant species. This adaptation, not seen in birds, appears to remove the body size constraints of incubation behaviour in giant oviraptorosaurs.

Lizards ran bipedally 110 million years ago.

Four heteropod lizard trackways discovered in the Hasandong Formation (Aptian-early Albian), South Korea assigned to Sauripes hadongensis, n. ichnogen., n. ichnosp., which represents the oldest lizard tracks in the world. Most tracks are pes tracks (N = 25) that are very small, average 22.29 mm long and 12.46 mm wide. The pes tracks show "typical" lizard morphology as having curved digit imprints that progressively increase in length from digits I to IV, a smaller digit V that is separated from the other digits by a large interdigital angle. The manus track is 19.18 mm long and 19.23 mm wide, and shows a different morphology from the pes. The predominant pes tracks, the long stride length of pes, narrow trackway width, digitigrade manus and pes prints, and anteriorly oriented long axis of the fourth pedal digit indicate that these trackways were made by lizards running bipedally, suggesting that bipedality was possible early in lizard evolution.

High diversity of the Ganzhou Oviraptorid Fauna increased by a new "cassowary-like" crested species.

A new oviraptorid dinosaur from the Late Cretaceous of Ganzhou, bringing oviraptrotid diversity of this region to seven taxa, is described. It is characterized by a distinct cassowary-like crest on the skull, no pleurocoels on the centra from the second through fourth cervical vertebrae, a neck twice as long as the dorsal vertebral column and slightly longer than the forelimb (including the manus). Phylogenetic analysis recovers the new oviraptorid taxon, Corythoraptor jacobsi, as closely related to Huanansaurus from Ganzhou. Osteochronology suggests that the type specimen of Corythoraptor had not reached stationary growth stage but died while decreasing growth rates. The histology implies that it would correspond to an immature individual approximately eight years old. We hypothesize, based on the inner structure compared to that in modern cassowaries, that the prominent casque of Corythoraptor was a multifunction-structure utilized in display, communication and probably expression of the fitness during mating seasons.

Mosaic evolution in an asymmetrically feathered troodontid dinosaur with transitional features.

Asymmetrical feathers have been associated with flight capability but are also found in species that do not fly, and their appearance was a major event in feather evolution. Among non-avialan theropods, they are only known in microraptorine dromaeosaurids. Here we report a new troodontid, Jianianhualong tengi gen. et sp. nov., from the Lower Cretaceous Jehol Group of China, that has anatomical features that are transitional between long-armed basal troodontids and derived short-armed ones, shedding new light on troodontid character evolution. It indicates that troodontid feathering is similar to Archaeopteryx in having large arm and leg feathers as well as frond-like tail feathering, confirming that these feathering characteristics were widely present among basal paravians. Most significantly, the taxon has the earliest known asymmetrical troodontid feathers, suggesting that feather asymmetry was ancestral to Paraves. This taxon also displays a mosaic distribution of characters like Sinusonasus, another troodontid with transitional anatomical features.

Perinate and eggs of a giant caenagnathid dinosaur from the Late Cretaceous of central China.

The abundance of dinosaur eggs in Upper Cretaceous strata of Henan Province, China led to the collection and export of countless such fossils. One of these specimens, recently repatriated to China, is a partial clutch of large dinosaur eggs (Macroelongatoolithus) with a closely associated small theropod skeleton. Here we identify the specimen as an embryo and eggs of a new, large caenagnathid oviraptorosaur, Beibeilong sinensis. This specimen is the first known association between skeletal remains and eggs of caenagnathids. Caenagnathids and oviraptorids share similarities in their eggs and clutches, although the eggs of Beibeilong are significantly larger than those of oviraptorids and indicate an adult body size comparable to a gigantic caenagnathid. An abundance of Macroelongatoolithus eggs reported from Asia and North America contrasts with the dearth of giant caenagnathid skeletal remains. Regardless, the large caenagnathid-Macroelongatoolithus association revealed here suggests these dinosaurs were relatively common during the early Late Cretaceous.

A Late Cretaceous diversification of Asian oviraptorid dinosaurs: evidence from a new species preserved in an unusual posture.

Oviraptorosaurs are a bizarre group of bird-like theropod dinosaurs, the derived forms of which have shortened, toothless skulls, and which diverged from close relatives by developing peculiar feeding adaptations. Although once among the most mysterious of dinosaurs, oviraptorosaurs are becoming better understood with the discovery of many new fossils in Asia and North America. The Ganzhou area of southern China is emerging as a hotspot of oviraptorosaur discoveries, as over the past half decade five new monotypic genera have been found in the latest Cretaceous (Maastrichtian) deposits of this region. We here report a sixth diagnostic oviraptorosaur from Ganzhou, Tongtianlong limosus gen. et sp. nov., represented by a remarkably well-preserved specimen in an unusual splayed-limb and raised-head posture. Tongtianlong is a derived oviraptorid oviraptorosaur, differentiated from other species by its unique dome-like skull roof, highly convex premaxilla, and other features of the skull. The large number of oviraptorosaurs from Ganzhou, which often differ in cranial morphologies related to feeding, document an evolutionary radiation of these dinosaurs during the very latest Cretaceous of Asia, which helped establish one of the last diverse dinosaur faunas before the end-Cretaceous extinction.

New Material of the Pterosaur Gladocephaloideus Lü et al., 2012 from the Early Cretaceous of Liaoning Province, China, with Comments on Its Systematic Position.

Although there are nine genera of ctenochasmatoids reported from the Jehol Biota, at present each is known from a specimen that has either a skull or a relatively complete postcranial skeleton. A nearly complete juvenile specimen of Gladocephaloideus from the Lower Cretaceous Jiufotang Formation of Sihedang, Lingyuan of Liaoning Province is the most complete ctenochasmatoid preserved to date with a skull and postcranial skeleton. Based on the holotype (IG-CAGS 08-07) and the nearly complete new specimen (JPM 2014-004), the diagnosis of Gladocephaloideus is amended: approximately 50 teeth in total with sharp tips; small nasoantorbital opening, occupying approximately 13% of total skull length; ratio of prenarial rostrum length to skull length approximately 0.63; deep groove along the mid-line of the mandibular symphysis; length to width ratio of the longest cervical vertebra = 4.1; ratio of femur length to tibia length = 0.61; tibia as long as the wing-phalange 1. Phylogenetic analysis recovers Gladocephaloideus within the clade Ctenochasmatidae. Gladocephaloideus has a closer relationship to the Chinese Pterofiltrus rather than to other ctenochasmatid pterosaurs. Microstructure of limb bones implies that JPM 2014-004 represents an early juvenile of Gladocephaloideus jingangshanensis, and that the type specimen is not a fully grown specimen either. We assume that the holotype may equate to the late juvenile or sub-adult developmental stage of Gladocephaloideus.

Largest known Mesozoic multituberculate from Eurasia and implications for multituberculate evolution and biology.

A new multituberculate, Yubaartar zhongyuanensis gen. and sp. nov., is reported from the Upper Cretaceous of Luanchuan County, Henan Province, China. The holotype of the new taxon is a partial skeleton with nearly complete cranium and associated lower jaws with in situ dentitions. The new species is the southern-most record of a Late Cretaceous multituberculate from outside of the Mongolian Plateau in Asia and represents the largest known Mesozoic multituberculate from Eurasia. The new specimen displays some intriguing features previously unknown in multituberculates, such as the first evidence of replacement of the ultimate upper premolar and a unique paleopathological case in Mesozoic mammals in which the animal with a severely broken right tibia could heal and survive in natural condition. The phylogenetic analysis based on craniodental characters places Yubaartar as the immediate outgroup of Taeniolabidoidea, a group consisting of a North American clade and an Asian clade. This relationship indicates at least a faunal interchange of multituberculates before the K-Pg transition. The new evidence further supports the hypothesis that disparity in dental complexity, which relates to animal diets, increased with generic richness and disparity in body size, and that an adaptive shift towards increased herbivory across the K-Pg transitional interval.

A New Oviraptorid Dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of Southern China and Its Paleobiogeographical Implications.

The Ganzhou area of Jiangxi Province, southern China is becoming one of the most productive oviraptorosaurian localities in the world. A new oviraptorid dinosaur was unearthed from the uppermost Upper Cretaceous Nanxiong Formation of Ganzhou area. It is characterized by an anterodorsally sloping occiput and quadrate (a feature shared with Citipati), a circular supratemporal fenestra that is much smaller than the lower temporal fenestra, and a dentary in which the dorsal margin above the external mandibular fenestra is strongly concave ventrally. The position of the anteroventral corner of the external naris in relation to the posterodorsal corner of the antorbital fenestra provides new insight into the craniofacial evolution of oviraptorosaurid dinosaurs. A phylogenetic analysis recovers the new taxon as closely related to the Mongolian Citipati. Six oviraptorid dinosaurs from the Nanxiong Formation (Ganzhou and Nanxiong) are distributed within three clades of the family. Each of the three clades from the Nanxiong Formation has close relatives in Inner Mongolia and Mongolia, and in both places each clade may have had a specific diet or occupied a different ecological niche. Oviraptorid dinosaurs were geographically widespread across Asia in the latest Cretaceous and were an important component of terrestrial ecosystems during this time.

A large, short-armed, winged dromaeosaurid (Dinosauria: Theropoda) from the Early Cretaceous of China and its implications for feather evolution.

The famous 'feathered dinosaurs' from the Early Cretaceous of Liaoning Province, northeastern China, include several dromaeosaurids, which are among the closest relatives of birds. Most of these are small-bodied taxa with long arms and broad wings comprised of vaned feathers, but a single specimen (the holotype of Tianyuraptor) belongs to a much larger individual with reduced forelimbs, which unfortunately lacks any preserved integument. We describe a new specimen of large-bodied, short-armed Liaoning dromaeosaurid, which we designate as a new genus and species, Zhenyuanlong suni. The integument is well preserved and provides the first evidence of feather morphologies and distribution in a short-armed (and probably non-volant) dromaeosaurid, indicating that these rare and aberrant taxa had large wings consisting of pennaceous feathers on the arms and long pennaceous feathers on the tail very similar to their smaller and longer-armed relatives, but potentially lacked vaned feathers on the legs. Zhenyuanlong adds yet more diversity to the Liaoning dromaeosaurid fauna, helps further reveal a distinct short-armed bauplan among dromaeosaurids, and illuminates previously-unrecognized homoplasy that complicates dromaeosaurid phylogeny and suggests that the Liaoning taxa may not have formed their own clade.

A new rhamphorhynchid pterosaur (Pterosauria) from Jurassic deposits of Liaoning Province, China.

Compared to pterosaurs from the Early Cretaceous from China, Late Jurassic pterosaurs are relatively rare. A new rhamphorhynchid pterosaur, Orientognathus chaoyngensis gen. et sp. nov., is erected based on an incomplete skeleton from the Upper Jurassic Tuchengzi Formation of Chaoyang, Liaoning Province, China. It is identified by the following characters: the toothless tip of the lower jaw is slightly pointed; the length ratio of wing metacarpal to humerus is 0.38, the ulna is shorter than each wing phalanx and the tibia is nearly equal to femur in length. A phylogenetic analysis recovers Orientognathus chaoyngensis as a rhamphorhynchid pterosaur. Orientognathus chaoyngensis is perhaps the youngest Jurassic pterosaur from western Liaoning Province of China. 

A new clade of Asian late Cretaceous long-snouted tyrannosaurids.

The iconic tyrannosaurids were top predators in Asia and North America during the latest Cretaceous, and most species had deep skulls that allowed them to generate extreme bite forces. Two unusual specimens of Alioramus from Mongolia seem to indicate a divergent long-snouted body plan among some derived tyrannosaurids, but the rarity and juvenile nature of these fossils leaves many questions unanswered. Here, we describe a remarkable new species of long-snouted tyrannosaurid from the Maastrichtian of southeastern China, Qianzhousaurus sinensis. Phylogenetic analysis places Qianzhousaurus with both species of Alioramus in a novel longirostrine clade, which was geographically widespread across latest Cretaceous Asia and formed an important component of terrestrial ecosystems during this time. The new specimen is approximately twice the size as both Alioramus individuals, showing that the long-snouted morphology was not a transient juvenile condition of deep-snouted species, but a characteristic of a major tyrannosaurid subgroup.

A new oviraptorosaur (Dinosauria: Oviraptorosauria) from the late cretaceous of southern China and its paleoecological implications.

A new oviraptorosaur Nankangia jiangxiensis gen. et sp. nov. is described on the basis of a partial postcranial skeleton with a partial lower jaw collected from the Upper Cretaceous Nanxiong Formation of Ganzhou, in Jiangxi Province of southern China. The new taxon is diagnosed by: (1) a mandibular symphysis that is not turned down; (2) neural spines of the cranial caudal vertebrae that are wider transversely than anteroposteriorly, forming a large posterior fossa with rugose central areas; (3) a femoral neck extending at an angle of about 90 to the shaft; and (4) a ratio of femur to tibia length of 0.95. Phylogenetic analysis recovers Nankangia as basal to the oviraptorid Yulong, but more derived than Caenagnathus, which also has a mandibular symphysis that is not turned down. The coexistence of Nankangia jiangxiensis, Ganzhousaurus nankangensis, Jiangxisaurus ganzhouensis, an unnamed oviraptorid from Nanxiong Basin and Banji long suggests that they occupied distinct ecological niches. Nankangia may have been more herbivorous than carnivorous.

An unusual basal Therizinosaur dinosaur with an ornithischian dental arrangement from northeastern China.

Therizinosauria are an unusual group of theropod dinosaurs, found mostly in the Cretaceous deposits in Mongolia, China and western USA. The basal forms of this group are represented by incomplete or disarticulated material. Here, we report a nearly complete, articulated skeleton of a new basal therizinosaur from the Early Cretaceous Yixian Formation of Jianchang County, western part of Liaoning Province, which sheds light on our understanding of anatomy of basal therizinosaurs. This new dinosaur shows some typical therizinosaur features, such as neural spines of the anterior caudal vertebrae that possess anterior and posterior alae, a rectangular buttress on the ventrolateral side of the proximal end of metacarpal I, and appressed metatarsal shafts. Our phylogenetic analysis suggests that it is a basal therizinosaur (sister taxon to Therizinosauroidea) because it bears many basal therizinosaur characters in the dentition, pelvis and hind limbs. The new therizinosaur described here has unique tooth and jaw characters such as the offsetting of the tooth row by a shelf and dentary teeth with labially concave and lingually convex dentary teeth, similar to ornithopods and ceratopsians.

Chicken-sized oviraptorid dinosaurs from central China and their ontogenetic implications.

Oviraptorids are a group of specialized non-avian theropod dinosaurs that were generally one to 8 m in body length. New specimens of baby oviraptorids from the Late Cretaceous of Henan Province are some of the smallest individuals known. They include diagnostic characters such as the relative position of the antorbital fenestra and the external naris, distinct opening in the premaxilla anteroventral to the external naris, antorbital fossa partly bordered by premaxilla posterodorsally, lacrimal process of premaxilla does not contact the anterodorsal process of the lacrimal, parietal almost as long as frontal; in dorsal view, posterior margin forms a straight line between the postzygapophyses in each of the fourth and fifth cervicals; femur longer than ilium. They also elucidate the ontogenetic processes of oviraptorids, including fusion of cranial elements and changes in relative body proportions. Hind limb proportions are constant in oviraptorids, regardless of absolute body size or ontogenetic stage. This suggests a sedentary lifestyle that did not involve the pursuit of similar-sized prey. The functional implications for bite force and therefore dietary preferences are better understood through the study of such small animals. The comparison of the measurements of 115 skeletons indicates that oviraptorids maintain their hind limb proportions regardless of ontogenetic stage or absolute size, which is a pattern seen more commonly in herbivores than in carnivores. This may weakly support the hypothesis that oviraptorids are herbivores rather than active carnivores.

An egg-adult association, gender, and reproduction in pterosaurs.

A sexually mature individual of Darwinopterus preserved together with an egg from the Jurassic of China provides direct evidence of gender in pterosaurs and insights into the reproductive biology of these extinct fliers. This new find and several other examples of Darwinopterus demonstrate that males of this pterosaur had a relatively small pelvis and a large cranial crest, whereas females had a relatively large pelvis and no crest. The ratio of egg mass to adult mass is relatively low, as in extant reptiles, and is comparable to values for squamates. A parchment-like eggshell points to burial and significant uptake of water after oviposition. This evidence for low parental investment contradicts the widespread assumption that reproduction in pterosaurs was like that of birds and shows that it was essentially like that of reptiles.

Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull.

The fossil record is a unique source of evidence for important evolutionary phenomena such as transitions between major clades. Frustratingly, relevant fossils are still comparatively rare, most transitions have yet to be documented in detail and the mechanisms that underpin such events, typified by rapid large scale changes and for which microevolutionary processes seem insufficient, are still unclear. A new pterosaur (Mesozoic flying reptile) from the Middle Jurassic of China, Darwinopterus modularis gen. et sp. nov., provides the first insights into a prominent, but poorly understood transition between basal, predominantly long-tailed pterosaurs and the more derived, exclusively short-tailed pterodactyloids. Darwinopterus exhibits a remarkable 'modular' combination of characters: the skull and neck are typically pterodactyloid, exhibiting numerous derived character states, while the remainder of the skeleton is almost completely plesiomorphic and identical to that of basal pterosaurs. This pattern supports the idea that modules, tightly integrated complexes of characters with discrete, semi-independent and temporally persistent histories, were the principal focus of natural selection and played a leading role in evolutionary transitions.