David M. Unwin

On the phylogeny and evolutionary history of pterosaurs

Abstract Previous cladistic studies of pterosaur relationships suffer from restricted numbers of taxa and characters, incomplete data sets and absence of information on characters, tree structure and the robustness of tress. Parsimony analysis of a new character data set (60 characters, 20 terminal taxa, 97.75% complete) yielded six trees. In the strict consensus tree Preondactylus is the most basal taxon followed, stepwise, by the Dimorphodontidae and the Anurognathidae. Beyond this basal group, more derived pterosaurs (Campylognathoididae (Rhamphorhynchidae + Pterodactyloidea)) share a suite of characters principally associated with elongation of the rostrum. The Pterodactyloidea consists of four major clades. The Ornithocheiroidea is the most basal taxon consisting, stepwise, of Istiodactylus, the Ornithocheiridae, Nyctosaurus and the Pteranodontidae. The remaining taxa, Ctenochasmatoidea, Dsungaripteroidea and Azhdarchoidea, are weakly united in a clade of non-ornithocheiroid pterodactyloids, but their inter-relationships are difficult to resolve. Cycnorhamphus is the basal-most ctenochasmatoid, while the remaining taxa (Pterodactylus, Lonchodectidae, Ctenochasmatidae) form an unresolved trichotomy. The Dsungaripteroidea (Germanodactylus+Dsungaripteridae) is strongly supported by features of the skull and dentition. The Azhdarchoidea (Tapejara [Tupuxuara + Azhdarchidae]) is united by cranial characters such as elevation of the antorbital region, and relative shortening of the wing finger. The pattern of pterosaur evolution suggested by the results of this analysis is broadly similar to traditional ideas, but has greater resolution, more complexity and reveals several previously unrecognized ‘events’.

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.

New pterosaur tracks (Pteraichnidae) from the Late Cretaceous Uhangri Formation, southwestern Korea

Numerous footprints of dinosaurs, pterosaurs and birds, together with arthropod tracks, have been discovered in the upper Cretaceous Uhangri Formation which crops out along the south- western coastline of South Korea. This ichnofauna contains the first pterosaur tracks reported from Asia. The digitigrade tridactyl manus impressions exhibit features of a typical pterosaur hand print. The pes impressions, however, show features that are different from pterosaur footprints reported pre- viously: there is no visible trace of impressions of individual digits, and the toes are triangular or rounded in shape distally without distinct claw impressions. As these features clearly distinguish the Uhangri tracks from Pteraichnus and Purbeckopus, we assign them to a new genus, Haenamichnus which accommodates the new ichnospecies, Haenamichnus uhangriensis. The prints are five to six times larger than those of Pteraichnus, and are currently the largest pterosaur ichnites known. They show virtually no trace of the 5th phalange of the pes, indicating that they were made by pterodactyloids; moreover, features of the tracks suggest that they can be attributed to azhdarchids, the commonest pterosaur of the Late Cretaceous. The longest pterosaur trackway yet known from any track site (length 7.3 m) and consisting of 14 pairs of foot impressions, was also found in the Uhangri Formation and suggests that azhdarchids, at least, were competent terrestrial locomotors. The fossil track site at Uhangri represents the first occurrence of the tracks of pterosaurs, dinosaurs and web- footed birds all on the same level. This demonstrates that pterosaurs and birds visited the same habitat, but the large size disparity suggests that they occupied different ecological niches.

An Egg-Adult Association, Gender, and Reproduction in Pterosaurs

A pterosaur fossil that includes a small, soft-shelled egg indicates a reproductive strategy like that of crocodilians. A sexually mature individual of Darwinopterus preserved together with an egg from the Jurassic of China provides direct evidence of gender in pterosaurs and insights into the reproductive biology of these extinct fliers. This new find and several other examples of Darwinopterus demonstrate that males of this pterosaur had a relatively small pelvis and a large cranial crest, whereas females had a relatively large pelvis and no crest. The ratio of egg mass to adult mass is relatively low, as in extant reptiles, and is comparable to values for squamates. A parchment-like eggshell points to burial and significant uptake of water after oviposition. This evidence for low parental investment contradicts the widespread assumption that reproduction in pterosaurs was like that of birds and shows that it was essentially like that of reptiles.

Respiratory evolution facilitated the origin of pterosaur flight and aerial gigantism.

Pterosaurs, enigmatic extinct Mesozoic reptiles, were the first vertebrates to achieve true flapping flight. Various lines of evidence provide strong support for highly efficient wing design, control, and flight capabilities. However, little is known of the pulmonary system that powered flight in pterosaurs. We investigated the structure and function of the pterosaurian breathing apparatus through a broad scale comparative study of respiratory structure and function in living and extinct archosaurs, using computer-assisted tomographic (CT) scanning of pterosaur and bird skeletal remains, cineradiographic (X-ray film) studies of the skeletal breathing pump in extant birds and alligators, and study of skeletal structure in historic fossil specimens. In this report we present various lines of skeletal evidence that indicate that pterosaurs had a highly effective flow-through respiratory system, capable of sustaining powered flight, predating the appearance of an analogous breathing system in birds by approximately seventy million years. Convergent evolution of gigantism in several Cretaceous pterosaur lineages was made possible through body density reduction by expansion of the pulmonary air sac system throughout the trunk and the distal limb girdle skeleton, highlighting the importance of respiratory adaptations in pterosaur evolution, and the dramatic effect of the release of physical constraints on morphological diversification and evolutionary radiation.

A survey of pterosaurs from the Jurassic and Cretaceous of the former Soviet Union and Mongolia

Remains of pterosaurs, the dominant aerial vertebrate throughout much of the Mesozoic were, until relatively recently, almost exclusively known from marine and marginal marine sediments of western Europe and North America. Prior to the 1960s Mesozoic deposits in the former Soviet Union and Mongolia had produced very few pterosaurs, but, in the last thirty years, many remains, including some from continental environments, have been found. Most important among these are Sordes and Batrachognathus from the Late Jurassic of Karatau in Kazakhstan, Azhdarcho from the Late Cretaceous of the Kyzylkum desert in Uzbekistan, and a number of new Mongolian pterosaurs, including a possible anurognathid from the Mid Jurassic of Bakhar, a dsungaripterid from the early Early Cretaceous of Tatal and an ornithocheirid from the late Early Cretaceous of Khuren‐Dukh. Although already documented to some extent, the significance of these discoveries remains underappreciated. The discovery and collection of these pterosaurs is de...

On Zhejiangopterus and the relationships of pterodactyloid pterosaurs

A summary of recent studies on the interrelationships of pterodactyloid pterosaurs is used as a framework for reassessing the taxonomic status of Zhejiangopterus, a new, long‐necked, Late Cretaceous pterosaur from China that has been assigned to the Nyctosauridae. Characters cited in support of this decision include: a notarium, edentulous jaws, and lack of a cranial crest. However, none of these is diagnostic of the Nyctosauridae. Zhejiangopterus exhibits a number of derived characters (orbit relatively small and located in a low position, posteroventrally facing occiput, features of the humerus and ‘T‐shaped’ cross‐section of wing phalanges two and three) only otherwise found in azhdarchids, thus we propose that Zhejiangopterus be reassigned to the Azhdarchidae.

A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution

Toothless pterosaurs played a key role in broadening the taxonomic, morphological and ecological diversity of Cretaceous pterosaurs. Here we report a complete, articulated skeleton of a 1.4-m-wingspan pterosaur from the Lower Cretaceous Jiufotang Formation of Liaoning Province, China which is identified as a new genus and species, Shenzhoupterus chaoyangensis gen. et sp. nov. The new taxon is edentulous, with a relatively large skull and a remarkably large, tall nasoantorbital fenestra that extends well above the main part of the braincase. Phylogenetic analysis shows that Shenzhoupterus gen. nov. belongs in a distinct clade of azhdarchoid pterosaurs, formally recognised here as a new family, Chaoyangopteridae, that also includes Chaoyangopterus, Jidapterus and Eoazhdarcho from the Jiufotang Formation and Eopteranodon from the Yixian Formation. These new data clarify recent confusion surrounding the systematics of these Lower Cretaceous taxa and provide new insights into the evolutionary history of pterosaurs.

Pterosaurs: back to the traditional model?

For most of the past 200 years, pterosaurs have been reconstructed with wing membranes attached to both the forelimbs and hindlimbs and with a quadrupedal stance and gait when grounded. In the early 1980s, this traditional model was replaced by a new design with narrow, stiff wings confined to the forelimbs and an upright, bipedal, digitigrade stance and gait. However, new studies of complete, uncrushed skeletal remains, well preserved wing membranes and extensive new pterosaur tracks, combined with reanalysis of the relationships of pterosaurs to other reptiles, suggest that a return to the traditional model is now overdue.

On the Nature of the Pteroid in Pterosaurs

The nature of the pteroid, a rod-like bone projecting from the carpus in pterosaurs, has long been disputed. Three lines of evidence, morphological, developmental and histological, indicate that the pteroid is a true bone, rather than ossified cartilage. The origin of the pteroid is unclear: it may be a modified carpal, the first metacarpal, or a neomorph.