Jonah N. Choiniere

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.

Tyrannosaur Paleobiology: New Research on Ancient Exemplar Organisms

Tyrannosaurs Revisited Tyrannosaurs represent some of the most successful and largest carnivores in Earths history. An expanding fossil record has allowed studies of their evolution and behavior that now allow broader comparisons with other groups, not just dinosaurs. Brusatte et al. (p. 1481) review the biology and evolutionary history of tyrannosaurs and update their phylogenetic relations to include several new fossils. The analysis suggests that tyrannosaurs remained relatively small (less than about 5 meters long) until the Late Cretaceous (about 80 million years ago). Tyrannosaurs, the group of dinosaurian carnivores that includes Tyrannosaurus rex and its closest relatives, are icons of prehistory. They are also the most intensively studied extinct dinosaurs, and thanks to large sample sizes and an influx of new discoveries, have become ancient exemplar organisms used to study many themes in vertebrate paleontology. A phylogeny that includes recently described species shows that tyrannosaurs originated by the Middle Jurassic but remained mostly small and ecologically marginal until the latest Cretaceous. Anatomical, biomechanical, and histological studies of T. rex and other derived tyrannosaurs show that large tyrannosaurs could not run rapidly, were capable of crushing bite forces, had accelerated growth rates and keen senses, and underwent pronounced changes during ontogeny. The biology and evolutionary history of tyrannosaurs provide a foundation for comparison with other dinosaurs and living organisms.

A basal alvarezsauroid theropod from the early Late Jurassic of Xinjiang, China.

Big Finger Alvarezsauroidea are an enigmatic group of theropods presumed to be closely related to birds, though most specimens are younger than Archaeopteryx. Choiniere et al. (p. 571; see news story by Stone) now describe a more complete early specimen, dating to about 160 million years ago, which supports the conclusion that Alvarezsauroidea are a basal group of the clade containing both birds and their close theropod relatives. The fossil also helps to reveal the evolution of this groups peculiar forelimb, which includes one enlarged functional finger. The Alvarezsauroidea group extends the clade containing birds and their theropod relatives back to 160 million years ago. The fossil record of Jurassic theropod dinosaurs closely related to birds remains poor. A new theropod from the earliest Late Jurassic of western China represents the earliest diverging member of the enigmatic theropod group Alvarezsauroidea and confirms that this group is a basal member of Maniraptora, the clade containing birds and their closest theropod relatives. It extends the fossil record of Alvarezsauroidea by 63 million years and provides evidence for maniraptorans earlier in the fossil record than Archaeopteryx. The new taxon confirms extreme morphological convergence between birds and derived alvarezsauroids and illuminates incipient stages of the highly modified alvarezsaurid forelimb.

A basal coelurosaur (Dinosauria: Theropoda) from the Late Jurassic (Oxfordian) of the Shishugou Formation in Wucaiwan, People's Republic of China

ABSTRACT We describe a new coelurosaurian theropod, Zuolong salleei, gen. et sp. nov., from exposures of the upper part of the Shishugou Formation at the Wucaiwan locality, Xinjiang Autonomous Region, Peoples Republic of China. Zuolong has a large, inclined quadrate foramen that extends onto the medial surface of the quadratojugal, an unusually large fovea capitis on the femoral head, and an apomorphically large distal condyle of metatarsal III with a medially projecting flange on the extensor surface. Radiometric dating of the Shishugou Formation constrains the age of the specimen to the beginning of the Late Jurassic (Oxfordian). A cladistic analysis of Zuolong salleei in a broadly sampled theropod data matrix recovers it as a basal coelurosaur. These data make Zuolong one of the oldest coelurosaur fossils yet known that preserves both cranial and postcranial bones.

A monodactyl nonavian dinosaur and the complex evolution of the alvarezsauroid hand

Digital reduction is a striking evolutionary phenomenon that is clearly exemplified in theropod dinosaurs by the functionally didactyl manus of tyrannosaurids, the flight-adapted manus of birds (Aves), and the tridactyl but digit II-dominated manus of alvarezsauroids. The enlargement of manual digit II in alvarezsauroids and the concurrent reduction of the lateral digits have been interpreted as adaptations for digging, although no detailed biomechanical analysis of hand function has so far been carried out for this group. In the derived alvarezsauroid clade Parvicursorinae, the lateral digits are so small as to be presumably vestigial. Here we report a new alvarezsauroid, Linhenykus monodactylus gen. et sp. nov., based on a specimen from the Upper Cretaceous Wulansuhai Formation of Inner Mongolia, China. Cladistic analysis identifies Linhenykus as the most basal parvicursorine, and digit II of the manus retains a slender morphology and other primitive features. However, Linhenykus is also highly apomorphic in exhibiting the most extreme reduction of the lateral manual digits seen in any alvarezsauroid. Phalanges are retained only on the most medial digit (digit II), making Linhenykus the only known functionally monodactyl nonavian dinosaur. Other parvicursorines are more primitive in retaining a tridactyl manus but more derived in that digit II is highly robust and shows other apomorphic features in both of its phalanges. The unexpected combination of features seen in the hand of Linhenykus points to a complex mosaic pattern of manual evolution in alvarezsauroids, with loss of the presumably vestigial outer digits being decoupled from change in the form of digit II.

Osteology of the Late Cretaceous alvarezsauroid Linhenykus monodactylus from China and comments on alvarezsauroid biogeography

The alvarezsauroid theropod Linhenykus monodactylus from the Upper Cretaceous of Inner Mongolia, China is the first known monodactyl non-avian dinosaur, providing important information on the complex patterns of manual evolution seen in alvarezsauroids. Here we provide a detailed description of the osteology of this taxon. Linhenykus shows a number of features that are transitional between parvicursorine and non-parvicursorine alvarezsauroids, but detailed comparisons also reveal that some characters had a more complex distribution. We also use event-based tree-fitting to perform a quantitative analysis of alvarezsauroid biogeography incorporating several recently discovered taxa. The results suggest that there is no statistical support for previous biogeographic hypotheses that favour pure vicariance or pure dispersal scenarios as explanations for the distributions of alvarezsauroids across South America, North America and Asia. Instead, statistically significant biogeographic reconstructions suggest a dominant role for sympatric (or “within area”) events, combined with a mix of vicariance, dispersal and regional extinction. At present the alvarezsauroid data set is too small to completely resolve the biogeographic history of this group: future studies will need to create larger data sets that encompass additional clades.

Theropod Teeth from the Middle-Upper Jurassic Shishugou Formation of Northwest Xinjiang, China

ABSTRACT Sixteen isolated theropod teeth were discovered in two areas in the upper Middle-lower Upper Jurassic Shishugou Formation of the Xinjiang Uyghur Autonomous Region, northwest China. This sample can be divided on the basis of qualitative features and simple quantitative metrics into seven tooth morphotypes, each of which probably represents a distinct taxon. Three of the morphotypes may be attributable to theropods already known from the Shishugou Formation, namely the alvarezsauroid Haplocheirus and the basal tetanurans Monolophosaurus and Sinraptor. The other four morphotypes, however, appear to represent new taxa, taking the known theropod diversity of the Shishugou Formation from six species to at least ten. One of the new taxa is probably a dromaeosaurid. Another appears to represent a troodontid or a relative of the potentially troodontid-like Paronychodon, itself so far known only from isolated teeth. Of the remaining two taxa, one appears to be a basal tetanuran or tyrannosauroid, whereas the other either belongs to one of these same groups or represents a ceratosaur. The probable deinonychosaurian teeth in our sample are among the oldest fossils known for this clade, and highlight the diversity of coelurosaurs in the Shishugou Formation.

Cranial Osteology of Haplocheirus sollers Choiniere et al., 2010 (Theropoda: Alvarezsauroidea)

ABSTRACT The basalmost alvarezsauroid Haplocheirus sollers is known from a single specimen collected in Upper Jurassic (Oxfordian) beds of the Shishugou Formation in northwestern China. Haplocheirus provides important data about the plesiomorphic morphology of the theropod group Alvarezsauroidea, whose derived members possess numerous skeletal autapomorphies. We present here a detailed description of the cranial anatomy of Haplocheirus. These data are important for understanding cranial evolution in Alvarezsauroidea because other basal members of the clade lack cranial material entirely and because derived parvicursorine alvarezsauroids have cranial features shared exclusively with members of Avialae that have been interpreted as synapomorphies in some analyses. We discuss the implications of this anatomy for cranial evolution within Alvarezsauroidea and at the base of Maniraptora.

An articulated pes from a small parvicursorine alvarezsauroid dinosaur from Inner Mongolia, China

A near complete and articulated parvicursorine pes from the Campanian Wulansuhai Formation is described. This pes is referred to the genus Linhenykus and is one of the first foot skeletons to be described for a derived alvarezsaur, providing new information on the first digit of the pes. The evolution of a laterally directed flange of the anterior face of the distal third metatarsal in arctometatarsalian taxa is described and discussed. This flange may have increased stability of the foot during cursorial locomotion and may also provide useful taxonomic and systematic data.

Reply to Dyke and Naish: European alvarezsauroids do not change the picture

Dyke and Naish (1) draw attention to three points that they consider to be “serious shortcomings” of our recent paper on a monodactyl nonavian dinosaur (2). Here, we respond to each point in turn.