David W. E. Hone

Dinosaurs and the Cretaceous Terrestrial Revolution

The observed diversity of dinosaurs reached its highest peak during the mid- and Late Cretaceous, the 50 Myr that preceded their extinction, and yet this explosion of dinosaur diversity may be explained largely by sampling bias. It has long been debated whether dinosaurs were part of the Cretaceous Terrestrial Revolution (KTR), from 125–80 Myr ago, when flowering plants, herbivorous and social insects, squamates, birds and mammals all underwent a rapid expansion. Although an apparent explosion of dinosaur diversity occurred in the mid-Cretaceous, coinciding with the emergence of new groups (e.g. neoceratopsians, ankylosaurid ankylosaurs, hadrosaurids and pachycephalosaurs), results from the first quantitative study of diversification applied to a new supertree of dinosaurs show that this apparent burst in dinosaurian diversity in the last 18 Myr of the Cretaceous is a sampling artefact. Indeed, major diversification shifts occurred largely in the first one-third of the groups history. Despite the appearance of new clades of medium to large herbivores and carnivores later in dinosaur history, these new originations do not correspond to significant diversification shifts. Instead, the overall geometry of the Cretaceous part of the dinosaur tree does not depart from the null hypothesis of an equal rates model of lineage branching. Furthermore, we conclude that dinosaurs did not experience a progressive decline at the end of the Cretaceous, nor was their evolution driven directly by the KTR.

A Jurassic ceratosaur from China helps clarify avian digital homologies

Theropods have traditionally been assumed to have lost manual digits from the lateral side inward, which differs from the bilateral reduction pattern seen in other tetrapod groups. This unusual reduction pattern is clearly present in basal theropods, and has also been inferred in non-avian tetanurans based on identification of their three digits as the medial ones of the hand (I-II-III). This contradicts the many developmental studies indicating II-III-IV identities for the three manual digits of the only extant tetanurans, the birds. Here we report a new basal ceratosaur from the Oxfordian stage of the Jurassic period of China (156–161 million years ago), representing the first known Asian ceratosaur and the only known beaked, herbivorous Jurassic theropod. Most significantly, this taxon possesses a strongly reduced manual digit I, documenting a complex pattern of digital reduction within the Theropoda. Comparisons among theropod hands show that the three manual digits of basal tetanurans are similar in many metacarpal features to digits II-III-IV, but in phalangeal features to digits I-II-III, of more basal theropods. Given II-III-IV identities in avians, the simplest interpretation is that these identities were shared by all tetanurans. The transition to tetanurans involved complex changes in the hand including a shift in digit identities, with ceratosaurs displaying an intermediate condition.

Macroevolutionary trends in the Dinosauria: Cope's rule

Copes rule is the tendency for body size to increase over time along a lineage. A set of 65 phylogenetically independent comparisons, between earlier and later genera, show that Copes rule applied in dinosaurs: later genera were on average about 25% longer than the related earlier genera to which they were compared. The tendency for size to increase was not restricted to a particular clade within the group, nor to a particular time within its history. Small lineages were more likely to increase in size, and large lineages more likely to decrease: this pattern may indicate an intermediate optimum body size, but can also be explained as an artefact of data error. The rate of size increase estimated from the phylogenetic comparisons is significantly higher than the rate seen across the fauna as a whole. This difference could indicate that within‐lineage selection for larger size was opposed by clade selection favouring smaller size, but data limitations mean that alternative explanations (which we discuss) cannot be excluded. We discuss ways of unlocking the full potential usefulness of phylogenies for studying the dynamics of evolutionary trends.

The vertebrates of the Jurassic Daohugou Biota of northeastern China

ABSTRACT The Early Cretaceous Jehol Biota of northeastern China has become famous over the last two decades as a source of feathered avialan and non-avialan theropods, preserved alongside an array of other fossil vertebrates, invertebrates, and plants. Still more recently, a rich assemblage referred to in this paper as the Daohugou Biota has begun to emerge from Jurassic strata in the same region. Like their counterparts from the Jehol Biota, Daohugou Biota vertebrate specimens are typically preserved in fine-grained lacustrine beds and often retain feathers and other soft-tissue features. At present, 30 vertebrate taxa (five salamanders, one anuran, two lizards, 13 pterosaurs, five dinosaurs, and four mammals) are known from the Daohugou Biota, which was first recognized at the Daohugou locality in Inner Mongolia. The presence of the salamander Chunerpeton tianyiensis, proposed in this paper as an index fossil for the Daohugou Biota, links the Daohugou locality to five other fossil-producing areas in the provinces of Hebei and Liaoning. The strata containing the Daohugou Biota are close to the Middle—pper Jurassic boundary and belong at least partly to the regionally widespread Tiaojishan Formation. In general, the vertebrate fauna of the Daohugou Biota is strikingly different from that of the Jehol Biota, although paravian dinosaurs, anurognathid pterosaurs, and salamanders with cryptobranchid and hynobiid affinities occur in both. Nevertheless, the Daohugou Biota and the Jehol Biota are two successive Lagerstätte assemblages that collectively offer a taphonomically consistent window into the Mesozoic life of northeast Asia over a significant span of geologic time.

The soft tissue of Jeholopterus (Pterosauria, Anurognathidae, Batrachognathinae) and the structure of the pterosaur wing membrane

The soft tissue preserved in the holotype (IVPP V12705) of Jeholopterus ningchengensis from the Daohugou Bed (Late Jurassic or Early Cretaceous) of China is described in detail. The plagiopatagium can be divided into the distal, comparatively more rigid actinopagatium and a proximal, more tensile tenopatagium. The actinopatagium extends from the wing finger to the articulation between the humerus and the forearm, and shows the presence of at least three layers containing actinofibrils. In each layer, the actinofibrils are parallel to subparallel, but this direction diverges from layer to layer. When distinct layers of actinofibrils are superimposed (owing to taphonomic compression), a reticular pattern is generated. The presence of layers with differently oriented actinofibrils is widespread in this pterosaur. A well-developed integumental covering formed by fibres (here named pycnofibres) that are thicker than the actinofibrils is present. Ungual sheaths that extend the length of the pedal and manual claws of this taxon are also observed. Although the understanding of the mechanical properties of the wing membrane is hampered by the lack of knowledge regarding the composition of the actinofibrils, the configuration observed in Jeholopterus might have allowed subtle changes in the membrane tension during flight, resulting in more control of flight movements and the organization of the wing membrane when the animal was at rest.

Sexual selection in prehistoric animals: detection and implications

Many fossil animals bear traits such as crests or horns that probably functioned as sexually selected signals or weapons. Interpretations of these structures as functioning in mate choice or intrasexual contests are often controversial, with interpretations based on biomechanics or physiology being favoured by many. Although testing hypotheses based on sexual selection can be difficult, especially given that there is no single, reliable means of recognising sexual selection, we argue that it is not impossible; indeed, there are now several cases where sexual selection is strongly supported. In other cases, a careful study of features such as sexual dimorphism, ontogeny, and allometry, coupled with testing of alternative hypotheses, will be necessary to distinguish between possible explanations for exaggerated features.

The first definitive carcharodontosaurid (Dinosauria: Theropoda) from Asia and the delayed ascent of tyrannosaurids

Little is known about the evolution of large-bodied theropod dinosaurs during the Early to mid Cretaceous in Asia. Prior to this time, Asia was home to an endemic fauna of basal tetanurans, whereas terminal Cretaceous ecosystems were dominated by tyrannosaurids, but the intervening 60 million years left a sparse fossil record. Here, we redescribe the enigmatic large-bodied Chilantaisaurus maortuensis from the Turonian of Inner Mongolia, China. We refer this species to a new genus, Shaochilong, and analyze its systematic affinities. Although Shaochilong has previously been allied with several disparate theropod groups (Megalosauridae, Allosauridae, Tyrannosauroidea, Maniraptora), we find strong support for a derived carcharodontosaurid placement. As such, Shaochilong is the first unequivocal Asian member of Carcharodontosauridae, which was once thought to be restricted to Gondwana. The discovery of an Asian carcharodontosaurid indicates that this clade was cosmopolitan in the Early to mid Cretaceous and that Asian large-bodied theropod faunas were no longer endemic at this time. It may also suggest that the ascent of tyrannosaurids into the large-bodied dinosaurian predator niche was a late event that occurred towards the end of the Cretaceous, between the Turonian and the Campanian.

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 (Copes 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 Copes 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 Copes Rule at a single taxonomic level may give misleading results.

The Extent of the Pterosaur Flight Membrane

The shape and extent of the membranous brachioptagium in pterosaurs remains a controversial topic for those attempting to determine the aerodynamic performance of the first vertebrate fliers. Various arguments in favour of the trailing edge terminating against either the torso or hip, the femur, the ankle, or different locations for various taxa, has resulted in several published reconstructions. Uncertainty over the correct model is detrimental to both aerodynamic and palaeoecological studies that are forced to simultaneously consider multiple and highly variable configurations for individual taxa. A review of relevant pterosaur specimens with preserved soft tissues or impressions of the wing membrane, however, strongly suggests that the trailing edge of the wing extended down to the lower leg or ankle in all specimens where the brachiopatagium is completely preserved. This configuration is seen across a phylogenetically broad range of pterosaurs and is thus likely to have been universally present throughout the Pterosauria. Support for opposing hypotheses where the trailing edge terminates against the body, hip, or knee are based on several specimens where the wing membrane is either incomplete or has undergone post-mortem contraction. An ankle attachment does not rule out a high aspect ratio wing as the curvature of the trailing edge and the ratio of the fore to hind limbs also play a major role in determining the final shape of the membrane.

The asymmetry of the carpal joint and the evolution of wing folding in maniraptoran theropod dinosaurs.

In extant birds, the hand is permanently abducted towards the ulna, and the wrist joint can bend extensively in this direction to fold the wing when not in use. Anatomically, this asymmetric mobility of the wrist results from the wedge-like shape of one carpal bone, the radiale, and from the well-developed convexity of the trochlea at the proximal end of the carpometacarpus. Among the theropod precursors of birds, a strongly convex trochlea is characteristic of Coelurosauria, a clade including the highly derived Maniraptora in addition to tyrannosaurs and compsognathids. The shape of the radiale can be quantified using a ‘radiale angle’ between the proximal and distal articular surfaces. Measurement of the radiale angle and reconstruction of ancestral states using squared-change parsimony shows that the angle was small (15°) in primitive coelurosaurs but considerably larger (25°) in primitive maniraptorans, indicating that the radiale was more wedge-shaped and the carpal joint more asymmetric. The radiale angle progressively increased still further within Maniraptora, with concurrent elongation of the forelimb feathers and the forelimb itself. Carpal asymmetry would have permitted avian-like folding of the forelimb in order to protect the plumage, an early advantage of the flexible, asymmetric wrist inherited by birds.