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’.
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.
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.
This paper outlines simple techniques for determining rates of heat gain and loss in relation to the weight and reflectance of insects caught in their natural habitats. In particular the construction of a new 'reflectometer' is described. The results thus obtained permit estimates of the relative importance of size and colour in determining rates of heat exchange and temperature excesses, so allowing better predictions of heat budgets for a given species.
The geological and paleoenvironmental setting and the vertebrate taxonomy of the fossiliferous, Cenomanian-age deltaic sediments in eastern Morocco, generally referred to as the “Kem Kem beds”, are reviewed. These strata are recognized here as the Kem Kem Group, which is composed of the lower Gara Sbaa and upper Douira formations. Both formations have yielded a similar fossil vertebrate assemblage of predominantly isolated elements pertaining to cartilaginous and bony fishes, turtles, crocodyliforms, pterosaurs, and dinosaurs, as well as invertebrate, plant, and trace fossils. These fossils, now in collections around the world, are reviewed and tabulated. The Kem Kem vertebrate fauna is biased toward large-bodied carnivores including at least four large-bodied non-avian theropods (an abelisaurid, Spinosaurus, Carcharodontosaurus, and Deltadromeus), several large-bodied pterosaurs, and several large crocodyliforms. No comparable modern terrestrial ecosystem exists with similar bias toward large-bodied carnivores. The Kem Kem vertebrate assemblage, currently the best documented association just prior to the onset of the Cenomanian-Turonian marine transgression, captures the taxonomic diversity of a widespread northern African fauna better than any other contemporary assemblage from elsewhere in Africa.
Abstract. 1. This paper describes field observations on diel changes in the nectars of Crataegus, Tilia and Echium in relation to microclimate and insect visits.2. Nectar concentration is highly correlated with ambient relative humidity, but the concentrative properties of the nectars differ from those of pure sugar solutions in ways that could be accounted for by microclimate or chemical effects.3. The microclimate inside flowers may influence the rate of equilibration of nectar with the relative humidity of the air, or the equilibrium concentration itself.4. The vapour pressure relationships of nectars may be influenced by the presence of components with low r/P values (that is, solutes in which solution show a relatively large lowering of vapour pressure, P, for a small change in refraction, r). Such components might be released by micro‐organisms in the nectar.5. The quantity of sugar per flower depends on the relative rates of secretion and reabsorption, among other things. The flowers we studied showed evidence for morning and evening peaks of secretion, and in Crataegus substantial quantities of sugar were ‘sequestered’ around midday.6. The pattern of visits by bumblebees to the flowers of Tilia and Echium can be related to the changing concentration of sugar in the nectar; caloric reward was probably not limiting for bumblebees visiting Tilia.
A partial rostrum of a new species of scaphognathine pterosaur, distinguished by a thin median crest along its dorsal margin and a deep embayment of the dental margin, is the first identifiable cranial fragment of a pterosaur from the Upper Jurassic Morrison Formation of western North America. By contrast with pterodactyloids, cranial crests are rare in “rhamphorhynchoids” and this is the first record of such a structure. The new material provides fresh insights into the taxonomic diversity of Late Jurassic North American pterosaurs. Based on the ratio of the skull and skeleton of Scpahognathus, the fragment represents an individual with an estimated wing span of 2.5 m. Consequently, this is one of the largest “rhamphorhynchoids” found so far. A mandible fragment from the same quarry has closely spaced alveoli, therefore cannot be referred to the rostrum. Its large size indicates another large “rhamphorhynchoid” in the Morrison Formation.
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 previously: 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.
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