A new long-tailed pterosaur, Wukongopterus lii gen. et sp. nov, is described based on an almost complete skeleton (IVPP V15113) representing an individual with an estimated wing span of 730 mm. The specimen was discovered in strata that possibly represent the Daohugou Bed (or Daohugou Formation) at Linglongta, Jianchang, Liaoning Province, China. Wukongopterus lii is a non-pterodactyloid pterosaur diagnosed by the first two pairs of premaxillary teeth protruding beyond the dentary, elongated cervical vertebrae (convergent with Pterodactyloidea), and a strongly curved second pedal phalanx of the fifth toe. The specimen further has a broken tibia that indicates an injury occurred while the individual was still alive. Taphonomic aspects provide indirect evidence of an uropatagium, supporting the general hypothesis that at least all non-pterodactyloid pterosaurs show a membrane between the hind limbs. A phylogenetic analysis including most non-pterodactyloid pterosaurs shows that Wukongopterus lii gen. et sp. nov. lies outside the Novialoidea, being cladistically more primitive than the Rhamphorhynchidae and Campylognathoides. This analysis differs from previous studies and indicates that more work is needed before a stable picture of non-pterodactyloid pterosaur relationships is achieved.

Two almost complete long-tailed pterosaurs from the Linglongta, Jianchang County, western Liaoning, China, are described and represent new taxa referred to the non-pterodactyloid clade Wukongopteridae. Kunpengopterus sinensis gen. et sp. nov. differs from other members of this clade mainly by the rounded posterior region of the skull, thick lacrimal process of the jugal and lack of a bony premaxillary crest. This species further shows a soft tissue crest above the frontal, a comparatively larger wing finger, and the proximal segment of the second pedal phalanx of the fifth toe shorter than in other wukongopterids. The second new species is referred to the genus Darwinopterus, D. linglongtaensis sp. nov. based on the posterior region of the skull. It further differs from other wukongopterid pterosaurs by the thin lacrimal process of the jugal, foramen on nasal process rounded, and by having the second pedal phalanx of the fifth toe less curved (115 • ). Several differences among the Wukongopteridae can be found in the dentition and the feet, suggesting that they might have occupied slightly different ecological niches. The long-tailed Changchengopterus pani is tentatively referred to this clade and new diagnosis for the wukongopterids Wukongopterus lii and Darwinopterus modularis is provided.

tapejaridae is a clade of toothless pterosaurs typically characterized by a large nasoantorbital fenestra and a premaxillary crest that extends from the anterior to the posterior part of the skull. Sinopterus dongi was the first reported Chinese tapejarid. Another genus "Huaxiapterus" was described, but it was later on determined that "Huaxiapterus" is in fact an invalid name. here, we report on a new juvenile specimen of Sinopterus atavismus from the Jiufotang Formation of western Liaoning, China, and revise the diagnosis of this species. Sinopterus atavismus does not present a square-like crest. Moreover the feature that groove in the ventral part of the second or third phalanx of manual digit IV is not diagnostic of the species. In the new material, the skull preserves a pointed process in the middle part of the dorsal margin of the premaxillary crest, which is different from other Chinese tapejarids. Considering the new specimen is known from a large skeleton that differed from the holotype, this difference may be related to ontogeny, as the premaxillary crest of the holotype is short and does not extend as long as that of the new specimen.

Despite the great increase in pterosaur diversity in the last decades, particularly due to discoveries made in western Liaoning (China), very little is known regarding pterosaur biogeography. Here, we present the description of a new pterosaur from the Jiufotang Formation that adds significantly to our knowledge of pterosaur distribution and enhances the diversity of cranial anatomy found in those volant creatures. Guidraco venator gen. et sp. nov. has an unusual upward-directed frontal crest and large rostral teeth, some of which surpass the margins of the skull and lower jaw when occluded. The new species is closely related to a rare taxon from the Brazilian Crato Formation, posing an interesting paleobiogeographic problem and supporting the hypothesis that at least some early Cretaceous pterosaur clades, such as the Tapejaridae and the Anhangueridae, might have originated in Asia. The association of the new specimen with coprolites and the cranial morphology suggest that G. venator preyed on fish.

These fossils shed new light on the reproductive strategy, ontogeny, and behavior of pterosaurs. The cranial crests show sexually dimorphic morphologies, with presumed males and females differing in crest size, shape, and robustness. Ontogenetic variation is reflected mainly in the expansion of the rostrum. The eggs have some external rigidity of the general pliable eggshell, and the microstructure of the eggshell is similar to that of some modern "soft" snake eggs. We suggest that this new pterosaur nested in colonies and thus exhibited gregarious behavior, a possible general trend for at least derived pterodactyloid pterosaurs.

Fossil eggs and embryos that provide unique information about the reproduction and early growth of vertebrates are exceedingly rare, particularly for pterosaurs. Here we report on hundreds of three-dimensional (3D) eggs of the species from a Lower Cretaceous site in China, 16 of which contain embryonic remains. Computed tomography scanning, osteohistology, and micropreparation reveal that some bones lack extensive ossification in potentially late-term embryos, suggesting that hatchlings might have been flightless and less precocious than previously assumed. The geological context, including at least four levels with embryos and eggs, indicates that this deposit was formed by a rare combination of events, with storms acting on a nesting ground. This discovery supports colonial nesting behavior and potential nesting site fidelity in the Pterosauria.

The Aptian Jiufotang Formation of northeast China is a Konservat Lagerstätte particularly rich in pterosaurs, notably azhdarchoids. Here we describe a new genus and species of toothed pteranodontoid pterosaur, Ikrandraco avatar gen. et sp. nov., based on two laterally flattened specimens. Ikrandraco avatar is diagnosed by a suite of features, including a very low and elongate skull, strongly inclined quadrate, and a deep, blade-like bony mandibular crest with a hook-like process on its posterior edge, an unusual structure so far unique to this taxon. The particular skull shape hints at a distinct feeding habit for pterosaurs that potentially includes temporary skimming and an extensible skin acting as a throat pouch that was more developed than in any other pterosaur known so far. The presence of two other taxa of purported piscivorous pterosaurs in the Jiufotang Formation suggests distinct resource exploitation in this part of China during the Early Cretaceous.

The Wukongopteridae compose a non-pterodactyloid clade of pterosaurs that are the most abundant flying reptiles in the deposits of the Middle-Late Jurassic Yanliao Biota. Until now, five species of three genera and two additional unnamed specimens have been described. Here we report on a new material, IVPP V 23674, that can be referred to the wukongopterid Kunpengopterus sinensis due to several features such as a comparably short nasoantorbital fenestra, the dorsally rising posterodorsal margin of the ischium, and the very short first pedal phalanx of digit V relative to metatarsal IV. IVPP V 23674 provides the first view of a wukongopterid palate, which differs from all other pterosaurs by having a very large postpalatine fenestra and laterally compressed choanae, indicating that the evolution of the pterosaur palate was more complex than previously thought. Sesamoid bones at the dorsal side of manual unguals are present and are reported for the first time in a wukongopterid suggesting an arboreal life-style for these pterosaurs.