Edina Prondvai

Rethinking the nature of fibrolamellar bone: an integrative biological revision of sauropod plexiform bone formation

We present novel findings on sauropod bone histology that cast doubt on general palaeohistological concepts concerning the true nature of woven bone in primary cortical bone and its role in the rapid growth and giant body sizes of sauropod dinosaurs. By preparing and investigating longitudinal thin sections of sauropod long bones, of which transverse thin sections were published previously, we found that the amount of woven bone in the primary complex has been largely overestimated. Using comparative cellular and light‐extinction characteristics in the two section planes, we revealed that the majority of the bony lamina consists of longitudinally organized primary bone, whereas woven bone is usually represented only by a layer a few cells thin in the laminae. Previous arguments on sauropod biology, which have been based on the overestimated amount, misinterpreted formation process and misjudged role of woven bone in the plexiform bone formation of sauropod dinosaurs, are thereby rejected.

Phylogeny, Histology and Inferred Body Size Evolution in a New Rhabdodontid Dinosaur from the Late Cretaceous of Hungary

Background Rhabdodontid ornithopod dinosaurs are characteristic elements of Late Cretaceous European vertebrate faunas and were previously collected from lower Campanian to Maastrichtian continental deposits. Phylogenetic analyses have placed rhabdodontids among basal ornithopods as the sister taxon to the clade consisting of Tenontosaurus, Dryosaurus, Camptosaurus, and Iguanodon. Recent studies considered Zalmoxes, the best known representative of the clade, to be significantly smaller than closely related ornithopods such as Tenontosaurus, Camptosaurus, or Rhabdodon, and concluded that it was probably an island dwarf that inhabited the Maastrichtian Haţeg Island. Methodology/Principal Findings Rhabdodontid remains from the Santonian of western Hungary provide evidence for a new, small-bodied form, which we assign to Mochlodon vorosi n. sp. The new species is most similar to the early Campanian M. suessi from Austria, and the close affinities of the two species is further supported by the results of a global phylogenetic analysis of ornithischian dinosaurs. Bone histological studies of representatives of all rhabdodontids indicate a similar adult body length of 1.6–1.8 m in the Hungarian and Austrian species, 2.4–2.5 m in the subadults of both Zalmoxes robustus and Z. shqiperorum and a much larger, 5–6 m adult body length in Rhabdodon. Phylogenetic mapping of femoral lengths onto the results of the phylogenetic analysis suggests a femoral length of around 340 mm as the ancestral state for Rhabdodontidae, close to the adult femoral lengths known for Zalmoxes (320–333 mm). Conclusions/Significance Our analysis of body size evolution does not support the hypothesis of autapomorhic nanism for Zalmoxes. However, Rhabdodon is reconstructed as having undergone autapomorphic giantism and the reconstructed small femoral length (245 mm) of Mochlodon is consistent with a reduction in size relative to the ancestral rhabdodontid condition. Our results imply a pre-Santonian divergence between western and eastern rhabdodontid lineages within the western Tethyan archipelago.

Life History of Rhamphorhynchus Inferred from Bone Histology and the Diversity of Pterosaurian Growth Strategies

Background Rhamphorhynchus from the Solnhofen Limestones is the most prevalent long tailed pterosaur with a debated life history. Whereas morphological studies suggested a slow crocodile-like growth strategy and superprecocial volant hatchlings, the only histological study hitherto conducted on Rhamphorhynchus concluded a relatively high growth rate for the genus. These controversial conclusions can be tested by a bone histological survey of an ontogenetic series of Rhamphorhynchus. Methodology/Principal Findings Our results suggest that Bennetts second size category does not reflect real ontogenetic stage. Significant body size differences of histologically as well as morphologically adult specimens suggest developmental plasticity. Contrasting the ‘superprecocial hatchling’ hypothesis, the dominance of fibrolamellar bone in early juveniles implies that hatchlings sustained high growth rate, however only up to the attainment of 30–50% and 7–20% of adult wingspan and body mass, respectively. The early fast growth phase was followed by a prolonged, slow-growth phase indicated by parallel-fibred bone deposition and lines of arrested growth in the cortex, a transition which has also been observed in Pterodaustro. An external fundamental system is absent in all investigated specimens, but due to the restricted sample size, neither determinate nor indeterminate growth could be confirmed in Rhamphorhynchus. Conclusions/Significance The initial rapid growth phase early in Rhamphorhynchus ontogeny supports the non-volant nature of its hatchlings, and refutes the widely accepted ‘superprecocial hatchling’ hypothesis. We suggest the onset of powered flight, and not of reproduction as the cause of the transition from the fast growth phase to a prolonged slower growth phase. Rapidly growing early juveniles may have been attended by their parents, or could have been independent precocial, but non-volant arboreal creatures until attaining a certain somatic maturity to get airborne. This study adds to the understanding on the diversity of pterosaurian growth strategies.

New Interpretation of the Palate of Pterosaurs

On the basis of a new, three‐dimensionally preserved specimen of the Early Jurassic pterosaur Dorygnathus banthensis we present a reinterpretation of the pterosaur palate. The hard palate is formed by the extensive palatal plate of the maxilla and not by the palatine as has been generally reconstructed. This palatal plate of the maxilla emarginates the choana rostrally and rostrolaterally as in other archosaurs and lepidosaurs. The longitudinally elongate and dorsoventrally flat palatine in Dorygnathus is an isolated bone caudal to the palatal plate of the maxilla and morphologically and topographically it resembles that of crocodilians and birds, respectively. The palatine separates the choana laterally from the suborbital fenestra demonstrating the homologous nature of the (primary) choana in all archosaurs and lepidosaurs. Our study indicates that in basal pterosaurs the pterygo–ectopterygoid fenestra existed caudal to the suborbital fenestra, which became confluent with the adductor chamber in pterodactyloids thereby increasing the relative size of the adductor chamber and hence the mass of the jaw adductors. The choana in basal pterosaurs was relatively small compared with the interpterygoid vacuity. With increasing rostroventral inclination of the quadrates in more derived pterosaurs, the interpterygoid vacuity was reduced considerably, whereas the choana increased in size. This exceptional Dorygnathus specimen also shows a hitherto unknown pair of fenestrae situated at the palatal contact of the premaxilla–maxilla and might represent the aperture for the vomeronasal organ. Anat Rec, 2010.

New models for the wing extension in pterosaurs

All powered flying animals have to face the same energetic problems: operating the wings during steady flight with muscles that require constant energy input and neural control to work. Accordingly the extant flying vertebrates have apparently found very similar solutions to parts of these issues – the biomechanical automatism built in their skeletal, muscular and connective tissue system. Based on these extant analogues (birds and bats) two new models are presented here for the mechanism of the distal wing extension in pterosaurs, an extinct group of flying vertebrates. The elongate fourth finger which solely supported their extensive flight membrane was a long lever arm that experienced significant loads and for which a reduction in muscle mass through automatisation would have been strongly beneficial. In the first model we hypothesize the presence of a propatagial ligament or ligamentous system which, as a result of the elbow extension, automatically performs and maintains the extension of the wing finger during flight and prohibits the hyperextension of the elbow. The second model has a co-operating bird-like propatagial ligamentous system and bat-like tendinous extensor muscle system on the forearm of the hypothetical pterosaur. Both models provide strong benefits to an animal with powered flight: (1) reduction of muscles and weight in the distal wing; (2) prevention of hyper extension of the elbow against drag; (3) automating wing extension and thereby reducing metabolic costs required to operate the pterosaurian locomotor apparatus. These models, although hypothetical, fit with the existing fossil evidence and lay down a basis for further biomechanical and/or aerodynamical investigations.

Does morphology reflect osteohistology-based ontogeny? A case study of Late Cretaceous pterosaur jaw symphyses from Hungary reveals hidden taxonomic diversity

Abstract With a single complete mandible and 56 mandibular symphyseal fragments of various sizes, the Late Cretaceous Hungarian azhdarchid material has been considered one of the most extensive monospecific pterosaur assemblages in the world. Representing a broad size range, these elements have been thought to demonstrate a developmental series of Bakonydraco galaczi. As such, they were ideal to test whether absolute size and/or morphology reliably indicate relative ontogenetic stages in this pterosaur. Forty-five specimens were selected for multivariate morphometrics and classified into four size classes. After acquiring the morphometric data set, we thin-sectioned eight symphyses representing all size groups and classified them into relative ontogenetic stages based on qualitative microstructural inspection prior to quantitative histological analyses. Microstructural characters suggestive of developmental state were then quantified for intra- and interindividual uni- and multivariate analyses to test the correspondence among the results of qualitative and quantitative analyses. In contrast to our expectations, histological features identified the smallest specimen as an adult and not an early juvenile. The substantial size difference between this specimen and other adults, along with its distinct microanatomical and histological features, implies the presence of at least two pterosaur taxa in this symphysis assemblage. This hypothesis is further supported by multivariate morphometrics, which separate the smallest symphyses from all other specimens that form one continuous group. Although the latter group also shows considerable size variability in corresponding ontogenetic stages, this suggests developmental plasticity rather than the presence of even more taxa, and indicates that symphysis size and morphology are poor indicators of skeletal maturity in these animals. Hence, bone histology is an important independent test of the assessment of ontogenetic stage using size and morphology.

The pterosaurian remains from the Grünbach Formation (Campanian, Gosau Group) of Austria: A reappraisal of 'Ornithocheirus buenzeli'

The fragmentary pterosaur material from the Campanian Grunbach Formation (Gosau Group) of Muthmannsdorf (Austria), previously identified as Ornithocheirus buenzeli Bunzel, 1871, is revised. A lower jaw fragment shows a helical type of articulation, which is known in several families of pterosaurs, and cannot be identified with great accuracy. The proximal part of a humerus shows distinctive features that allow it to be referred to as a member of the family Azhdarchidae, which is widespread in the Late Cretaceous Period of Europe. Ornithocheirus buenzeli is considered a nomen dubium . The pterosaur material from the Grunbach Formation cannot be used as evidence for the presence of ornithocheirids in the Late Cretaceous of Europe.

The history of Late Jurassic pterosaurs housed in Hungarian collections and the revision of the holotype of Pterodactylus micronyx Meyer 1856 (a 'Pester Exemplar')

Abstract The history and scientific significance of three Late Jurassic pterosaur specimens housed in different Hungarian palaeontological collections are described. One of these is the holotype of Pterodactylus micronyx Meyer 1856 that was thought to be lost, but with its rediscovery in the 1980s the ‘Pester Exemplar’ becomes the name-bearing type again. The second specimen is an articulated, partially three-dimensional skeleton of a Rhamphorhynchus muensteri; and the third is an articulated right hindlimb of a Pterodactylus sp. – both donated by Andor Semsey to the Hungarian Geological Institute. The anatomical revision of the holotype of P. micronyx indicated the osteological immaturity of the specimen; however, there is insufficient data on this taxon to assess its taxonomic validity.

Potential For Intracranial Movements in Pterosaurs

Based on comparative anatomical, morphological, and phylogenetic considerations the potential of pterosaurs for cranial kinesis is assessed. Our investigation shows that whereas skeletally mature derived pterodactyloids have completely fused, rigid and doubtlessly akinetic skulls, skeletally immature derived pterodactyloids and more basal pterosaurs possess key features in the morphology of their otic and basal joints that are suggestive of cranial kinesis, namely streptostyly. In addition, pterosaurs exhibit an evolutionarily informative trend in the degree of cranial ossification, where it is low in most nonpterodactyloids (here named bifenestratans), intermediate in Rhamphorhynchus and Archaeopterodactyloidea, and high in derived pterodactyloids. Incomplete fusion could also indicate loose connections between skull elements. However, another crucial anatomical requirement of a kinetic skull, the permissive kinematic linkage is absent in all pterosaurian taxa. The fact, that the presence of permissive kinematic linkages in the skull is also a prerequisite of all types of cranial kinesis, provides hard evidence that all members of Pterosauria had akinetic skulls. Thus, the presence of the morphological attributes indicative of intracranial movements in some pterosaurs must be explained on grounds other than real potential for cranial kinesis. It could either be of mechanical or ontogenetic importance, or both. Alternatively, it might be considered as the morphological remnant of a real, kinetic skull possessed by the diapsid ancestors of pterosaurs. Anat Rec, , 2011.

Diversity and convergences in the evolution of feeding adaptations in ankylosaurs (Dinosauria: Ornithischia)*

Abstract Ankylosaurian dinosaurs were low-browsing quadrupeds that were traditionally thought of as simple orthal pulpers exhibiting minimal tooth occlusion during feeding, as in many extant lizards. Recent studies, however, have demonstrated that effective chewing with tooth occlusion and palinal jaw movement was present in some members of this group. Qualitative and quantitative analysis of feeding characters (i.e. craniodental features, tooth wear patterns, origin and insertion of jaw adductors) reveal at least three different jaw mechanisms during the evolution of Ankylosauria. Whereas, in basal members, food processing was restricted to simple orthal pulping, in late Early and Late Cretaceous North American and European forms a precise tooth occlusion evolved convergently in many lineages (including nodosaurids and ankylosaurids) complemented by palinal power stroke. In contrast, Asian forms retained the primitive mode of feeding without any biphasal chewing, a phenomenon that might relate to the different types of vegetation consumed by these low-level feeders in different habitats on different landmasses. Further, a progressive widening of the muzzle is demonstrated both in Late Cretaceous North American and Asian ankylosaurs, and the width and general shape of the muzzle probably correlates with foraging time and food type, as in herbivorous mammals.