An unusual edentulous pterosaur from the Early Cretaceous Romualdo Formation of Brazil

Numerous taxa make up the Early Cretaceous fauna of Brazil, including Ornithocheiroidea, Tapejaridae, Thalassodromidae, Chaoyangopteridae and a purported member of Azhdarchidae. Dsungaripteridae has only been tentatively assumed to be present in the form of ‘Santanadactylus’ spixi. New study of NMSG SAO 251093 (a specimen referred to Thalassodromeus sethi) suggests it is a previously unknown species of dsungaripterid, Banguela oberlii, tax. nov., differing from Thalassodromeus and other pterosaurs from the Early Cretaceous of Brazil by a unique combination of characters, including an upturned jaw tip, a short dorsal mandibular symphyseal shelf (dmss), and an autapomorphic thin crest placed halfway along the fused mandibular symphysis without a keel along the ventral margin of the jaw. B. oberlii, tax. nov., is referred to Dsungaripteridae based on a dmss no longer than the ventral shelf, U-shaped caudal margin of the ventral shelf and lateral margins of the mandibular symphysis concave in dorsal view. B. oberlii, tax. nov., is the youngest known dsungaripterid, and expands known morphological diversity in the clade as well as the Early Cretaceous pterosaur fauna of South America.     

On the cervical vertebrae of the Pterodactyloidea (Reptilia: Archosauria)

Within the Pterodactyloidea, the cervical vertebrae show considerable variation. These elements are also sufficiently common and contain enough anatomical information to make them taxonomically valuable. A survey of these vertebrae concludes that most known pterodactyloids fall into two groups: long-necked forms with attenuated cervical vertebrae that possess low neural spines, and tall-spined forms that possess relatively short neck vertebrae with tall neural spines. These two groups may represent natural taxonomic units. However, this is by no means conclusive and can only be tested by the study of other regions of pterodactyloid skeletons.     

A biomechanical approach on the optimal stance of Anhanguera piscator (Pterodactyloidea) and its implications for pterosaur gait on land

The stance of pterosaurs on land is traditionally a controversial question. Here, we show that pterosaurs like Anhanguera piscator were quadrupeds. An osteological model of A. piscator was three-dimensionally built in digital space. The reconstructed muscles of its pelvic girdle were then placed on their points of origin and insertion to allow the biomechanical calculations to find the most efficient stance on land to be performed. The hindlimb readjustment (i.e. the repositioning of the hindlimb according to the achieved results) led to a pelvic counterclockwise displacement at 10°, which means that the ilium previously placed at 0° regarding an axis parallel to the ground was moved (and so the whole pelvis) 10° up from the preacetabular process. This new position prevents A. piscator from having a fully upright stance. A 10° displacement of the pelvic girdle would compel the forelimbs to be highly sprawled. Therefore, this study affords A. piscator having a quadrupedal gait and demonstrates that a bipedal stance is not viable once the lever arm values decrease abruptly both for extensor and flexor muscles during the femoral extension. This is the first time this approach is used to shed light on this question.     

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.     

A NEW PTEROSAUR FROM THE LIAONING PROVINCE OF CHINA, THE PHYLOGENY OF THE PTERODACTYLOIDEA, AND CONVERGENCE IN THEIR CERVICAL VERTEBRAE

Abstract:  The largest known flying organisms are the azhdarchid pterosaurs, a pterodactyloid clade previously diagnosed by the characters of their extremely elongate middle-series cervical vertebrae. The named species of the Azhdarchidae are from the Late Cretaceous. However, isolated mid-cervical vertebrae with similar dimensions and characters have been referred to this group that date back to the Late Jurassic, implying an almost 60 million year gap in the fossil record of this group and an unrecorded radiation in the Jurassic of all the major clades of the Pterodactyloidea. A new pterosaur from the Early Cretaceous of Liaoning Province of China, Elanodactylus prolatus gen. et sp. nov., is described with mid-cervical vertebrae that bear these azhdarchid characters but has other postcranial material that are distinct from the members of this group. Phylogenetic analysis of the new species and the Pterodactyloidea places it with the Late Jurassic vertebrae in the Late Jurassic–Early Cretaceous Ctenochasmatidae and reveals that the characters of the elongate azhdarchid vertebrae appeared independently in both groups. These results are realized though the large taxon sampling in the analysis demonstrating that the homoplastic character states present in these two taxa were acquired in a different order in their respective lineages. Some of these homoplastic characters were previously thought to appear once in the history of pterosaurs and may be correlated to the extension of the neck regions in both groups. Because the homoplastic character states in the Azhdarchidae and Ctenochasmatidae are limited to the mid-cervical vertebrae, these states are termed convergent based on a definition of the term in a phylogenetic context. A number of novel results from the analysis presented produce a reorganization in the different species and taxa of the Pterodactyloidea.     

Cope's Rule in the Pterosauria, and differing perceptions of Cope's Rule at different taxonomic levels

The remarkable extinct flying reptiles, the pterosaurs, show increasing body size over 100 million years of the Late Jurassic and Cretaceous, and this seems to be a rare example of a driven trend to large size (Cope's Rule). The size increases continue throughout the long time span, and small forms disappear as larger pterosaurs evolve. Mean wingspan increases through time. Examining for Cope's Rule at a variety of taxonomic levels reveals varying trends within the Pterosauria as a whole, as pterodactyloid pterosaurs increase in size at all levels of examination, but rhamphorhynchoid pterosaurs show both size increase and size decrease in different analyses. These results suggest that analyses testing for Cope's Rule at a single taxonomic level may give misleading results.     

Preservational bias controls the fossil record of pterosaurs

Pterosaurs, a Mesozoic group of flying archosaurs, have become a focal point for debates pertaining to the impact of sampling biases on our reading of the fossil record, as well as the utility of sampling proxies in palaeo‐diversity reconstructions. The completeness of the pterosaur fossil specimens themselves potentially provides additional information that is not captured in existing sampling proxies, and might shed new light on the group's evolutionary history. Here we assess the quality of the pterosaur fossil record via a character completeness metric based on the number of phylogenetic characters that can be scored for all known skeletons of 172 valid species, with averaged completeness values calculated for each geological stage. The fossil record of pterosaurs is observed to be strongly influenced by the occurrence and distribution of Lagerstätten. Peaks in completeness correlate with Lagerstätten deposits, and a recovered correlation between completeness and observed diversity is rendered non‐significant when Lagerstätten species are excluded. Intervals previously regarded as potential extinction events are shown to lack Lagerstätten and exhibit low completeness values: as such, the apparent low diversity in these intervals might be at least partly the result of poor fossil record quality. A positive correlation between temporal patterns in completeness of Cretaceous pterosaurs and birds further demonstrates the prominent role that Lagerstätten deposits have on the preservation of smaller bodied organisms, contrasting with a lack of correlation with the completeness of large‐bodied sauropodomorphs. However, we unexpectedly find a strong correlation between sauropodomorph and pterosaur completeness within the Triassic–Jurassic, but not the Cretaceous, potentially relating to a shared shift in environmental preference and thus preservation style through time. This study highlights the importance of understanding the relationship between various taphonomic controls when correcting for sampling bias, and provides additional evidence for the prominent role of sampling on observed patterns in pterosaur macroevolution.     

辽西早白垩世九佛堂组两种新的翼手龙类化石(英文)

简要报道了辽西热河群上部九佛堂组两件新的翼手龙类化石 ,即夜翼龙科(Nyctosauridae)的张氏朝阳翼龙 (新属、新种 )Chaoyangopteruszhangigen .etsp .nov.和古魔翼龙科 (Anhangueridae)的顾氏辽宁翼龙 (新属、新种 )Liaoningopterusguigen .et.sp .nov.。前者为保存较完整的化石骨架 ,后者为一大型翼龙的头骨和部分头后骨骼化石。朝阳翼龙是夜翼龙科在亚洲大陆的首次确切的化石记录 ,也是层位最低和保存最完整的化石骨架。朝阳翼龙具有4节翼指骨 ,手指爪粗大弯曲 ,这些发现补充和修正了前人认为的夜翼龙科只有 3节翼指骨 ,手指爪退化缺失等一些重要的形态学特征。朝阳翼龙与该科的Nyctosaurusgracilis头后骨骼相比 ,具有许多不同的特征 ,如胫骨特长 ,远长于股骨 ,翼掌骨和第 1翼指骨相对较短 ,肩胛骨短于乌喙骨等。辽宁翼龙是我国已发现的个体最大的翼龙化石 ,发育前上颌骨和齿骨弧形脊突这一古魔翼龙科的重要鉴别特征。与该科的其他成员相比 ,辽宁翼龙上、下颌的牙齿较少 ,仅分布在其前部 ,齿列约占上、下颌长度的 1 / 2。上颌第 1、3齿小 ,第 2、4齿巨大 ,其中第 4齿最大 ,为已知翼龙中最大的牙齿。牙齿具有明显的替换现象。夜翼龙科的成员仅分布于美洲大陆的晚白垩世地层中 ,而古魔翼龙科的成员则是     

New crested specimens of the Late Cretaceous pterosaurNyctosaurus

Two recently collected specimens of the Late Cretaceous pterosaurNyctosaurus differ from all previously known specimens in the possession of a large branching cranial crest. The crest extends upward and backward from the posterior skull roof and is nearly three times the length of the skull proper. Despite the large crest, the specimens do not differ significantly in morphology from previously known specimens ofNyctosaurus, and do not seem to represent a new species ofNyctosaurus. The specimens suggest that the cranial crest was developed late in ontogeny, which is consistent with the interpretation of pterosaur cranial crests as intraspecific display structures.      

Taxonomic review of the Ornithocheirus complex (Pterosauria) from the Cretaceous of England

Over a decade after the last major review of the Cambridge Greensand pterosaurs, their systematics remains one of the most disputed points in pterosaur taxonomy. Ornithocheiridae is still a wastebasket for fragmentary taxa, and some nomenclatural issues are still a problem. Here, the species from the Cretaceous of England that, at some point, were referred in Ornithocheirus, are reviewed. Investigation of the primary literature confirmed that Criorhynchus should be considered an objective junior synonym of Ornithocheirus. Taxonomic review of more than 30 species known from fragmentary remains showed that 16 of them are undiagnosable (nomina dubia): Palaeornis cliftii, Cimoliornis diomedeus, Pterodactylus compressirostris, Pterodactylus fittoni, Pterodactylus woodwardi, Ornithocheirus brachyrhinus, Ornithocheirus carteri, Ornithocheirus crassidens, Ornithocheirus dentatus, Ornithocheirus enchorhynchus, Ornithocheirus eurygnathus, Ornithocheirus oxyrhinus, Ornithocheirus scaphorhynchus, Ornithocheirus tenuirostris, Ornithocheirus xyphorhynchus, and Pterodactylus sagittirostris. Fourteen species are considered valid, and diagnoses are provided to all of them: Ornithocheirus simus, Lonchodraco giganteus comb. n., Lonchodraco machaerorhynchus comb. n., Lonchodraco(?) microdon comb. n., Coloborhynchus clavirostris, ‘Ornithocheirus’ capito, Camposipterus nasutus comb. n., Camposipterus(?) sedgwickii comb. n., Camposipterus(?) colorhinus comb. n., Cimoliopterus cuvieri comb. n., ‘Ornithocheirus’ polyodon, ‘Ornithocheirus’ platystomus, ‘Pterodactylus’ daviesii, and ‘Ornithocheirus’ denticulatus. These species are referred in the genera Ornithocheirus, Lonchodraco gen. n., Coloborhynchus, Cimoliopterus gen. n., and Camposipterus gen. n., but additional genera are probably present, as indicated by the use of single quotation marks throughout the text. A cladistic analysis demonstrates that Anhangueridae lies within a newly recognized clade, here named Anhangueria, which also includes the genera Cearadactylus, Brasileodactylus, Ludodactylus, and Camposipterus. The anhanguerian ‘Cearadactylus’ ligabuei belongs to a different genus than Cearadactylus atrox. Lonchodraconidae fam. n. (more or less equivalent to Lonchodectidae sensu Unwin 2001) is a monophyletic entity, but its exact phylogenetic position remains uncertain, as is the case of Ornithocheirus simus. Therefore, it is proposed that Ornithocheiridae should be constricted to its type species and thus is redundant. Other taxa previously referred as “ornithocheirids” are discussed in light of the revised taxonomy.