An Unusual Pterosaur Specimen (Pterodactyloidea, ?Azhdarchoidea) from the Early Cretaceous Romualdo Formation of Brazil,and the Evolution of the Pterodactyloid Palate

A new and unusual specimen of a probable azhdarchoid pterosaur is described for the Early Cretaceous (Albian) Romualdo Formation of Brazil. The specimen consists of a palate that, although fragmentary, has a unique morphology differing from all other known pterosaurs with preservation of palatal elements. The new specimen probably indicates the presence of a yet undescribed pterodactyloid taxon for Romualdo Formation and brings new information on pterosaur diversity of this sedimentary unity. Mainly due to the rarity of pterodactyloid specimens with palate preservation, this structure has been overlooked in this clade. Here, we reassess the palatal anatomy of Pterodactyloidea, revealing an intriguing variety of morphotypes and evolutionary trends, some of them described here for the first time. The morphological disparity displayed by different pterodactyloid taxa may be further evidence of the presence of diverse feeding strategies within the clade.     

New postcranial elements of the Thalassodrominae (Pterodactyloidea,Tapejaridae) from the Romualdo Formation (Aptian–Albian), Santana Group,Araripe Basin,Brazil

Tapejarids are edentate pterosaurs recovered mainly from Early Cretaceous deposits. They are diagnosed by five synapomorphies, among which only one is postcranial: a broad and well‐developed tubercle at the ventroposterior margin of the coracoid. Regarding the clade Thalassodrominae, most phylogenetic studies are based on cranial elements, as postcranial skeletons of these pterosaurs are rare. Here, new postcranial material from the Romualdo Formation (Aptian–Albian) from the Araripe Basin is described. The material comprises the three posteriormost cervical vertebrae, the first seven dorsal vertebrae (fused into a notarium), both scapulocoracoids, a fragment of a sternum, a partial right humerus, a small fragment of a 4th phalanx of the wing finger, a distal extremity of the right femur and the proximal portions of both tibia and fibula. Comparisons with other specimens and morphological features examined in a phylogenetic context, such as the presence of three foramina lateral and dorsal to the neural canal of the cervical vertebrae, the presence of a notarium and a pneumatic foramen on the ventral side of the proximal portion of the humerus, allow the assignment of this specimen as Thalassodrominae indet. Regarding palaeobiogeographical aspects, to date, this clade is exclusively found in the Romualdo Formation. It is the most complete postcranial material assigned to the Thalassodrominae described so far.     

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.     

A new specimen of the azhdarchoid pterosaur Tapejara wellnhoferi

A new specimen of the Early Cretaceous azhdarchoid Tapejara wellnhoferi is described from the Romualdo Member of the Santana Formation, NE Brazil, providing the first detailed account of the postcranial skeleton. Although limited in its preservation, the osteology is typical of other azhdarchoid pterosaurs from these deposits and represents a juvenile animal with a relatively small wing span of < 1.5 m. The ratios of the pedal elements are identical to those noted for larger, indeterminate azhdarchoids of the Nova Olinda Member of the Crato Formation, where the unguals of the pes are greatly enlarged relative to those of the ornithocheiroids that co-inhabited the Santana lagoon. The ratios of these elements suggest that, as part of a larger suite of characters, these animals were likely better adapted for life on the ground than their ornithocheiroid relatives.     

Partial mandible of a giant pterosaur from the uppermost Cretaceous (Maastrichtian) of the Hațeg Basin,Romania

We describe and interpret a posterior mandibular symphysis of a very large azhdarchid pterosaur. The specimen LPB (FGGUB ) R.2347 exhibits a series of morphological characters present in both azhdarchid and tapejarid pterosaurs, suggesting a more basal position within the clade Azhdarchidae. This fossil was collected from Maastrichtian continental deposits near V?lioara in the Ha?eg Basin, Romania, but cannot be confidently referred to the contemporaneous giant Hatzegopteryx thambema, also from V?lioara, due to the absence of overlapping skeletal elements. Remarkably, this mandibular symphysis shares a number of features the smaller azhdarchoid Bakonydraco galaczi from the Santonian of Hungary. Additional comparisons with previously described large‐sized azhdarchid mandibles indicate a certain degree of morphological and probably ecological disparity within the group. This specimen represents the largest pterosaur mandible ever found and provides insights into the anatomy of the enigmatic giant pterosaurs.     

Pterosaur remains from the Cretaceous of Morocco

New pterosaur remains consisting of jaw fragments of toothless taxa and isolated teeth are described from the red beds of the Kern Kern region of southern Morocco. The stratigraphic position of those red beds is discussed and it is concluded that they are in all likelihood early Cenomanian in age. At least four taxa of pterodactyloid pterosaurs are present. The toothless jaw fragments are referred to the families ?Pteranodontidae, ?Azhdarchidae and Tapejaridae. Four different morphotypes can be distinguished among the isolated teeth. They are tentatively referred to the Ornithocheiridae. This assemblage reveals a high diversity of pterosaurs in Africa during the early Upper Cretaceous. The possible occurrence of tapejarids and anhanguerids indicates relationships with the somewhat older pterosaur assemblage from the Santana Formation (Aptian/Albian) of Brazil. If confirmed, the presence of azhdarchids and pteranodontids in the early Cenomanian suggests an early differentiation of these essentially late Late Cretaceous groups of large pterosaurs.     

Postcranial skeletal pneumaticity and air-sacs in the earliest pterosaurs

Patterns of postcranial skeletal pneumatization (PSP) indicate that pterosaurs possessed components of a bird-like respiratory system, including a series of ventilatory air-sacs. However, the presence of PSP in the oldest known pterosaurs has not been unambiguously demonstrated by previous studies. Here we provide the first unequivocal documentation of PSP in Late Triassic and earliest Jurassic pterosaurs. This demonstrates that PSP and, by inference, air-sacs were probably present in the common ancestor of almost all known pterosaurs, and has broader implications for the evolution of respiratory systems in bird-line archosaurs, including dinosaurs.     

A new species of Neoblattella Shelford (Blattellidae: Pseudophyllodromiinae) from Amazonas State, Brazil

Neoblattella mista sp. nov. is described and illustrated based on male genitalia and morphological characters of a single specimen collected in the town of Coari, State of Amazonas, Brazil. By studying the literature on the genus, we determined that N. mista sp. nov. differs from the other four known complexes in the morphology of its genital structures, including the supra-anal plate, subgenital plate, right and left phallomeres, median genital sclerite and tergal modification in the abdomen, and is placed in the new "mista complex".     

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     

The phylogenetic position of the Pterosauria within the Archosauromorpha

In recent years the hypothesis that pterosaurs were the major sister-group of dinosaurs and a closely-linked hypothesis that pterosaurs evolved flight from the ground up have gained general acceptance. A cladistic analysis of the Archosauromorpha using characters presented by previous workers results in a single most parsimonious tree with the Pterosauria as the major sister-group of the Dinosauria. However, that sister-group relationship is supported only by a suite of hindlimb characters that are correlated with bipedal digitigrade locomotion in dinosaurs. In pterosaurs the characters have been interpreted as correlates of bipedal cursorial locomotion, arboreal leaping, or involvement of the hindlimb in the wing. The homology of those characters in dinosaurs and pterosaurs cannot be supported. Reanalysis of the data after exclusion of those hindlimb characters results in most parsimonious trees with the Pterosauria as the sister-group of the Erythrosuchidae + Proterochampsidae + Euparkeria + Archosauria, in that order. This sister-group relationship is supported by a diverse assemblage of functionally independent skeletal characters from all regions of the skeleton. The results of the analysis cast doubt on the hypothesis that pterosaurs evolved flight from the ground up.     

On the size and flight diversity of giant pterosaurs, the use of birds as pterosaur analogues and comments on pterosaur flightlessness

The size and flight mechanics of giant pterosaurs have received considerable research interest for the last century but are confused by conflicting interpretations of pterosaur biology and flight capabilities. Avian biomechanical parameters have often been applied to pterosaurs in such research but, due to considerable differences in avian and pterosaur anatomy, have lead to systematic errors interpreting pterosaur flight mechanics. Such assumptions have lead to assertions that giant pterosaurs were extremely lightweight to facilitate flight or, if more realistic masses are assumed, were flightless. Reappraisal of the proportions, scaling and morphology of giant pterosaur fossils suggests that bird and pterosaur wing structure, gross anatomy and launch kinematics are too different to be considered mechanically interchangeable. Conclusions assuming such interchangeability--including those indicating that giant pterosaurs were flightless--are found to be based on inaccurate and poorly supported assumptions of structural scaling and launch kinematics. Pterosaur bone strength and flap-gliding performance demonstrate that giant pterosaur anatomy was capable of generating sufficient lift and thrust for powered flight as well as resisting flight loading stresses. The retention of flight characteristics across giant pterosaur skeletons and their considerable robustness compared to similarly-massed terrestrial animals suggest that giant pterosaurs were not flightless. Moreover, the term 'giant pterosaur' includes at least two radically different forms with very distinct palaeoecological signatures and, accordingly, all but the most basic sweeping conclusions about giant pterosaur flight should be treated with caution. Reappraisal of giant pterosaur material also reveals that the size of the largest pterosaurs, previously suggested to have wingspans up to 13 m and masses up to 544 kg, have been overestimated. Scaling of fragmentary giant pterosaur remains have been misled by distorted fossils or used inappropriate scaling techniques, indicating that 10-11 m wingspans and masses of 200-250 kg are the most reliable upper estimates of known pterosaur size.     

A second specimen of the pterosaur<Emphasis Type="Italic">Anurognathus ammoni</Emphasis>

A new complete and fully articulated specimen of the anurognathid pterosaurAnurognathus ammoni from the Upper Jurassic Solnhofen Limestone of southern Germany provides new information about the species. The skull is broader than long and quite tall. The naris and antorbital fenestra are both small and anteriorly placed, whereas the orbit is very large. The palatal elements are slender struts of bone separating large openings. The tail is short, but is neither pygostyle-like nor like that of pterodactyloids. The wingfinger, unlike that of almost all pterosaurs, is reduced to three phalanges. Pedal digit V bears two long phalanges. Reexamination of the holotype specimen revealed evidence of bony bumps on the premaxilla and dentary that may have been associated with a fringe of bristles around the mouth.Anurognathus and the other anurognathids were probably adapted to aerial insectivory in low light conditions like extant caprimulgids and insectivorous bats, and may have spent little time on the ground. The taxonomy of the Anurognathidae is reviewed and new diagnoses are presented. A cladistic analysis supports the interpretation that the Anurognathidae is the sister-group to all other pterosaurs.      

Terrestrial locomotion in pterosaurs

On the basis of a well‐preserved pelvis of Anhanguera sp. from the Lower Cretaceous (Aptian) of the Chapada do Araripe, Brazil, the problem of terrestrial locomotion in pterosaurs is discussed. A three‐dimensional reconstruction of the pelvis led to a lateral, dorsal and posterior orientation of the acetabula. By use of the preserved proximal ends of the femora of the same individual, the articulation in the hip socket could be tested. The normal articulation of the femur resulted in a horizontal position of the femur shaft, probably during flight. For constructional reasons the femur could not be brought down to a vertical position. Therefore, a parasagittal swing of the femora necessary for a bird‐like stance and gait must have been impossible. It is suggested that in pterosaurs the wing membrane was attached to the upper leg, which helped in stretching, steering and cambering.

Moreover, on the basis of comparisons of the fossil preservation of pterosaurs Compsognathus and Archaeopteryx in the Solnhofen limestone, it is concluded that the femora of pterosaurs were splayed out laterally, and that they had a semi‐erect gait. They were not bipedal animals, but had to use their fore limbs as well on the ground. Nevertheless, as vertebrates extremely adapted to flight, they could not have been able quadrupeds, either.     

Palaeogenomics of pterosaurs and the evolution of small genome size in flying vertebrates

The two living groups of flying vertebrates, birds and bats, both have constricted genome sizes compared with their close relatives. But nothing is known about the genomic characteristics of pterosaurs, which took to the air over 70 Myr before birds and were the first group of vertebrates to evolve powered flight. Here, we estimate genome size for four species of pterosaurs and seven species of basal archosauromorphs using a Bayesian comparative approach. Our results suggest that small genomes commonly associated with flight in bats and birds also evolved in pterosaurs, and that the rate of genome-size evolution is proportional to genome size within amniotes, with the fastest rates occurring in lineages with the largest genomes. We examine the role that drift may have played in the evolution of genome size within tetrapods by testing for correlated evolution between genome size and body size, but find no support for this hypothesis. By contrast, we find evidence suggesting that a combination of adaptation and phylogenetic inertia best explains the correlated evolution of flight and genome-size contraction. These results suggest that small genome/cell size evolved prior to or concurrently with flight in pterosaurs. We predict that, similar to the pattern seen in theropod dinosaurs, genome-size contraction preceded flight in pterosaurs and bats.     

The shape of pterosaur evolution: evidence from the fossil record

Abstract Although pterosaurs are a well‐known lineage of Mesozoic flying reptiles, their fossil record and evolutionary dynamics have never been adequately quantified. On the basis of a comprehensive data set of fossil occurrences correlated with taxon‐specific limb measurements, we show that the geological ages of pterosaur specimens closely approximate hypothesized patterns of phylogenetic divergence. Although the fossil record has expanded greatly in recent years, collectorship still approximates a sigmoid curve over time as many more specimens (and thus taxa) still remain undiscovered, yet our data suggest that the pterosaur fossil record is unbiased by sites of exceptional preservation (lagerstätte). This is because as new species are discovered the number of known formations and sites yielding pterosaur fossils has also increased – this would not be expected if the bulk of the record came from just a few exceptional faunas. Pterosaur morphological diversification is, however, strongly age biased: rarefaction analysis shows that peaks of diversity occur in the Late Jurassic and Early Cretaceous correlated with periods of increased limb disparity. In this respect, pterosaurs appear unique amongst flying vertebrates in that their disparity seems to have peaked relatively late in clade history. Comparative analyses also show that there is little evidence that the evolutionary diversification of pterosaurs was in any way constrained by the appearance and radiation of birds.     

On a theropod scapula (Upper Cretaceous) from the Marília Formation, Bauru Group, Brazil

The record of theropod dinosaurs in Brazil is very scarce. One of the most promising lithostratigraphic units for those reptiles is the Bauru Group. The dinosaur remains found in this unit are mainly those of sauropods, while theropods are represented mostly by teeth. Here we describe a right scapula (housed at the Earth Science Museum of the Departamento Nacional de Produção Mineral/Rio de Janeiro) that is the first theropod scapula reported from the Cretaceous of Brazil and only the second osteological evidence of the Tetanurae from the Bauru Group. The specimen was recovered from the outskirts of Peirópolis, in Minas Gerais State. Comparisons with other theropod dinosaurs are limited, but the overall morphology of the new specimen indicates that it is neither a member of the Abelisauridae nor a member of the Avialae. It also differs from more basal members of the Theropoda, but its overall shape is consistent with several derived members of the Tetanurae, likely a non-avialan maniraptoran. Despite the lack of precision in its taxonomic position, the new specimen confirms the presence of non-avialan Maniraptora in the Bauru Group.     

A New Pterosaur from the Jurassic of Cuba

Cacibupteryx caribensis gen. et sp. nov. is a new pterosaur of the family Rhamphorhynchidae found in western Cuba, in rocks of the Jagua Formation (Middle–Upper Oxfordian). The holotype, a skull and part of the left wing, is one of the few Jurassic pterosaurs that is well preserved in three dimensions. The new taxon shares characters with early and late Jurassic pterosaurs, and is one of the few late Jurassic taxa from western Laurasia and Gondwana. Furthermore, Cacibupteryx joins Nesodactylus hesperius Colbert from Cuba, and Sordes pilosus Sharov, from Kazakhstan as the most complete pterosaur recorded from the Middle–Upper Oxfordian. Cacibupteryx caribensis is one of the largest Jurassic pterosaurs known, and its skull possesses several distinct characters, including relatively broad roof elements (mainly frontal and parietal bones), a jugal with a prominent recess, occipital table trapezoidal in shape with the maximum width between the quadrate bones, and a small fenestra located in the posterior part of the pterygoid bones. In the Oxfordian, the Caribbean Corridor separated Laurasia and western Gondwana. The diversity of the marine herpetofauna found in the Jagua Vieja Member (Jagua Formation), and of teleostean fish, confirms that the corridor was an effective seaway over which flew at least Nesodactylus and Cacibupteryx .     

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.     

A new species of Megalonychidae (Mammalia, Xenarthra) from the Quaternary of Poço Azul (Bahia, Brazil)

The new genus and species Ahytherium aureum (Mammalia, Xenarthra, Megalonychidae) from the Quaternary of Poço Azul (Bahia, Brazil) is described. It is the first Brazilian megalonychid known from reasonably complete and well-preserved remains. Purported Brazilian megalonychids described in the past, such as Ocnopus gracilis and Xenocnus cearesis, are noted as belonging to other sloth clades, and the acceptance by past paleontologists of the existence of ‘strange’ megalonychids in Brazil is shown to be erroneous. Ahytherium aureum, in fact, exhibits typical megalonychid morphology. It differs from other known members of Megalonychidae in several characters, including a markedly shortened, but high rostral region, with dorsally inflated frontals, wide zygomatic processes of the frontal, narrow, blade-like and anterolaterally oriented lacrimals, curved, slender and oval caniniforms, gracile humerus with less developed deltopectoral shelf, and relatively distal position of the greater trochanter of the femur. A second specimen from São Paolo state is tentatively assigned to the new genus and species.     

Prozostrodon brasiliensis,a probainognathian cynodont from the Late Triassic of Brazil: second record and improvements on its dental anatomy

Probainognathian cynodonts are conspicuous elements of the Assemblage Zones of the Triassic Santa Maria Supersequence in southern Brazil. Within this group, the derived clade Prozostrodontia, in which the crown group Mammalia is included, is taxonomic diverse in the Hyperodapedon and Riograndia AZs. We describe here the second known specimen (CAPPA/UFSM 0123) of Prozostrodon brasiliensis, until now only represented by its holotype. CAPPA/UFSM 0123 includes a right dentary with dentition. As in the holotype of P. brasiliensis, it has four lower incisors, pc4 with conspicuous cusp a, and small cusps b, c, and d, pc5-pc6 of ‘triconodont’ type with cusps a > c > b > d, with continuous lingual cingulum bearing up to six small discrete cusps, length of the lower tooth row more than half the length of the dentary, and relatively deep horizontal ramus of the dentary. The new specimen is about 25% smaller than the holotype and there is not a diastema between the canine and postcanine teeth, indicating its juvenile condition. Based on both known specimens of P. brasiliensis, a discussion on tooth replacement is presented, showing that the adult individual has more postcanine tooth morphotypes than the juvenile one.