Abstract The Wukongopteridae is an important pterosaur clade from the Yanliao Biota, combining features of basal and derived pterosaurs. So far, the Wukongopteridae consists of five species divided into three genera: Wukongopterus lii, Darwinopterus modularis, Darwinopterus linglongtaensis, Darwinopterus robustodens, and Kunpengopterus sinensis. Here we report a new species, Darwinopterus camposi sp. nov., based on an almost complete skeleton (IVPP V 17957). The new species is referred to Darwinopterus due to the presence of an elongated posterior region of the skull and the bony premaxillary crest that starts about the anterior margin of the nasoantorbital fenestra. It differs from all other wukongopterids by having the dorsal margin of the premaxillary crest straight, without an extensive dorsal projection and presenting a smooth lateral surface. Furthermore, D. camposi sp. nov. has eighteen and fourteen teeth on each side of the upper and lower jaws, respectively, and the fourth phalanx of the wing finger shorter than the first. IVPP V 17957 shows some fused postcranial bones, like the extensor tendon process to the first wing finger phalanx, but also has unfused premaxilla and frontal, which provides further information about wukongopterid ontogeny.

Abstract Pterosaur remains are rare from the lowermost Cretaceous, hampering our understanding of the taxonomic and morphological diversities of pterosaurs during this period. The Lower Cretaceous Tugulu Group in Wuerho, China is renowned for hosting the Wuerho Pterosaurian Fauna (WPF), which has so far yielded numerous fossil remains of two dsungaripterid pterosaurs, Dsungaripterus weii and Noripterus complicidens. Here we report a partial ornithocheiromorph humerus from the WPF, representing a deeply divergent clade from Dsungaripteridae. The scarcity of ornithocheiromorphs from the WPF might be interpreted by niche partitioning with dsungaripterids. Meanwhile, we also report a U-Pb zircon age of 134.27 ± 0.36 Ma dated by LA-ICP-MS for the tuffaceous layer at the uppermost part of the Shengjinkou Formation, confirming the Valanginian age of the WPF presented by a previous study. The Wuerho region is one of the few localities producing abundant pterosaur fossils and the only one with an earliest Cretaceous age. The new finding here also suggests that Ornithocheiromorpha had rapidly diversified and achieved a global distribution during the earliest Cretaceous, presumably through a series of modifications on the locomotor apparatus including the warped deltopectoral crest of the humerus, which might substantially improve their flight efficiency.

Abstract The Wukongopteridae is a transitional clade between the long- and short-tailed pterosaur groups, and at least ten specimens have been studied without a determined juvenile specimen. Here, we described a small-sized Kunpengopterus sinensis, less than half the size of the holotype, which is the smallest specimen in wukongopterids. Based on unossified small elements, unfused cranial and postcranial elements, and grooves on the bone surface, this specimen is thought to be at least an early juvenile or even a late hatchling. By comparing the juvenile and subadult specimens of K. sinensis, we have found that the mid region of the upper and lower jaws had a higher growth rate than the anterior part, and that the growth rates were similar in most postcranial elements except for a higher rate in the caudal vertebrae. We revised the previous diagnosis of K. sinensis and specified that two characteristics, nasoantorbital fenestra approximately 40% of the skull length and a thin and relatively short maxillary process of the jugal, should be diagnostic in subadult or adult specimens. We have also found that pedal features are stable during ontogeny and can be diagnostic in juvenile, subadult or adult specimens in K. sinensis.

Abstract: 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.

ABSTRACT A new saurolophine hadrosaurid, Laiyangosaurus youngi gen. et sp. nov. is described and phylogenetically analyzed based on several cranial elements from the Jingangkou Formation, Wangshi Group, Upper Cretaceous of Laiyang, Shandong, China. Laiyangosaurus youngi differs from other members of the saurolophine clade on the basis of a number of autapomorphies, including a prominent and narrow ridge on the lateral side of the nasal which forms the posterodorsal and posterior margin of the circumnasal depression, a primary ridge that runs along most of the maxillary tooth row that is slightly deflected posteriorly, a retroarticular process of the surangular that is dorsolateroposteriorlly recurved, and orbital margins that are wider than the infratemporal margins of the jugal. This new taxon can be further distinguished by a number of unique combination of characters, including dorsal margin of nasal is flat, absence supracranial crest, a relatively shallow and rostrodorsally directed caudal margin of the lacrimal process of the jugal, and one or more foramina present on the rostral surface of the premaxilla. A phylogenetic analysis indicates that L. youngi comprises a monophyletic clade, which is known as Edmontosaurini.

ABSTRACT Cranial crests show considerable variation within the Pterosauria, a group of flying reptiles that developed powered flight. This includes the Wukongopteridae, a clade of non-pterodactyloids, where the presence or absence of such head structures, allied with variation in the pelvic canal, have been regarded as evidence for sexual dimorphism. Here we discuss the cranial crest variation within wukongopterids and briefly report on a new specimen (IVPP V 17957). We also show that there is no significant variation in the anatomy of the pelvis of crested and crestless specimens. We further revisit the discussion regarding the function of cranial structures in pterosaurs and argue that they cannot be dismissed a priori as a valuable tool for species recognition.

A contraparte de um espécime anteriormente descrito, contendo uma fêmea de pterossauro não-pterodactiloide com um ovo, revela a existência de um segundo ovo, este situado dentro da cavidade corporal desta fêmea prenhe. O exemplar demonstra claramente que os pterossauros possuíam dois ovidutos funcionais, e evidencia que a redução de um oviduto não foi um pré-requisito para o desenvolvimento do voo ativo pelo menos neste grupo. A análise da composição de um dos ovos sugere a ausência de uma camada externa formada por carbonato de cálcio. Seções histológicas de um fêmur não registram osso medular e demonstram que este pterossauro atingiu maturidade reprodutiva antes da maturidade óssea. O presente estudo evidencia que os pterossauros punham ovos de tamanho ainda menor do que se supunha e que desenvolveram uma estratégia reprodutiva mais similar a dos répteis basais do que a das aves. Se pterossauros eram altamente precociais ou necessitavam de cuidado parental ainda está aberto à discussão.

The counterpart of a previously described non-pterodactyloid pterosaur with an egg revealed the presence of a second egg inside the body cavity of this gravid female. It clearly shows that pterosaurs had two functional oviducts and demonstrates that the reduction of one oviduct was not a prerequisite for developing powered flight, at least in this group. Compositional analysis of one egg suggests the lack of a hard external layer of calcium carbonate. Histological sections of one femur lack medullary bone and further demonstrate that this pterosaur reached reproductive maturity before skeletal maturity. This study shows that pterosaurs laid eggs even smaller than previously thought and had a reproductive strategy more similar to basal reptiles than to birds. Whether pterosaurs were highly precocial or needed parental care is still open to debate.

Uma nova espécie de ovo de tartaruga fóssil foi estabelecida: Emydoolithus laiyangensis oogen. et oosp. nov. Esta possui forma elíptica, uma casca rígida e outras características da casca que diferem dos demais ovos fósseis de quelônios conhecidos, todos de forma arredondada. Trata-se da segunda espécie de ovo fóssil de quelônio encontrado no Cretáceo na China. Esta descoberta mostra que o ecossistema Laiyang no Cretáceo Superior é mais diversificado do que se pensava.

A new type of turtle egg fossil was established: Emydoolithus laiyangensis oogen. et oosp. nov.. Based on its elliptical morphological shape, rigid eggshells, and eggshell characteristics, it is different from other types of round chelonian egg fossils. It is the second chelonian egg fossil found in Cretaceous in China. This discovery shows the Laiyang ecosystem in Late Cretaceous is more diversified than previously thought.

Um crânio quase completo de um novo pterossauro ctenochasmatídeo, Pterofiltus qiui gen. et sp. nov., de depósitos do Cretáceo Inferior de Liaoning, China é descrito. O exemplar (IVPP V12339) foi coletado de um argilito da parte inferior da Formação Yixian (125 Ma) na localidade Zhangjiagou. A nova espécie possui a seguinte combinação de caracteres: aproximadamente 112 dentes (incluindo ambas as arcadas); dentição ocupando mais de 50% do comprimento do crânio; dentes anteriores de tamanho variado; sínfise mandibular ocupando mais da metade do comprimento da mandíbula; abertura ventral na parte média da sínfise mandibular.

A nearly complete skull of a new ctenochasmatid pterosaur, Pterofiltus qiui gen. et sp. nov., from the Lower Cretaceous deposits of Liaoning, China, is described here. The specimen (IVPP V12339), was collected from the shale of the lower Yixian Formation (125 Ma) at the Zhangjiagou locality. It has the following combination of characters: about 112 teeth in total (including the upper and lower jaws); the dentition occupies more than 50% of the skull length; the anterior teeth vary in size; the mandibular symphysis is longer than half of the whole mandible length; in ventral view, an apparent symphyseal trough in the median part of the symphysis.

Dois novos pterossauros de cauda longa procedentes de Linglongta, Jianchang County da região oeste de Liaoning, China, são descritos e referidos ao clado Wukongopteridae. Kunpengopterus sinensis gen. et sp. nov. difere dos demais integrantes desse clado basicamente pela região mais arredondada da parte posterior do crânio, pelo espesso processo lacrimal do jugal e pela ausência de uma crista óssea premaxilar. Esta espécie apresenta uma crista formada por tecido mole acima da região frontal e possui o dedo alar proporcionalmente maior do que nos demais wukongopterídeos. A segunda espécie é referida ao gênero Darwinopterus, D. linglongtaensis sp. nov., com base na região posterior do crânio. Difere dos deais pterossauros wukongopterídeos por possuir o processo lacrimal do jugal delgado, o forâmen no processo nasal arredondado e pela condição menos curvada da segunda falange do quinto dígito do pé. As principais diferenças entre os Wukongopteridae são encontradas na dentição e nos pés, o que sugere que estes ocupavam nichos ecológicos ligeiramente distintos. O pterossauro de cauda longa Changchen-gopterus pani também é tentativamente referido a esse clado. Novas diagnoses para os wukongopterídeos Wukongopterus lii e Darwinopterus modularis são apresentadas.

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.

Um novo pterossauro de cauda longa, Wukongopterus lii gen. et sp. nov., é descrito baseado em um esqueleto quase completo (IVPP V15113) de um indivíduo com abertura alar estimada em 730 mm. O exemplar foi encontrado nas camadas Daohugou (ou Formação Daohugou) em Linglongta, Jianchang, Província de Liaoning, China. Wukongopterus lii é um pterossauro não-pterodactilóide diagnosticado pela presença de dois pares de dentes pré-maxilares posicionados antes do início do dentário, vertebras cervicais alongadas (convergente com os Pterodactyloidea) e a segunda falange do quinto dígito do pé fortemente curvada. Este espécime também apresenta uma tíbia quebrada indicando que a quebra ocorreu com o animal em vida. Evidências tafonômicas apresentam dados indiretos da presença de um uropatágio, corroborando com a hipótese de que pelo menos os não-pterodactilóides possuíam uma membrana entre os seus membros posteriores. Uma análise filogenética incluindo vários pterossauros não-pterodactilóides resulta no posicionamento de Wukongopterus lii gen. et sp. nov. fora dos Novialoidea, sendo cladisticamente mais primitivo do que os Rhamphorhynchidae e Campylognathoides. Esta nova análise filogenética difere de resultados anteriores, indicando que mais trabalhos são necessários até que uma estabilidade da relação de parentesco entre os pterossauros não-pterodactilóides seja alcançada.

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 Capylognathoides. This analysis differs from previous studies and indicates that more work is needed before a stable picture of non-pterodactyloid pterosaur relationships is achieved.