Catherine A. Forster

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.

New Horned Dinosaurs from Utah Provide Evidence for Intracontinental Dinosaur Endemism

Background During much of the Late Cretaceous, a shallow, epeiric sea divided North America into eastern and western landmasses. The western landmass, known as Laramidia, although diminutive in size, witnessed a major evolutionary radiation of dinosaurs. Other than hadrosaurs (duck-billed dinosaurs), the most common dinosaurs were ceratopsids (large-bodied horned dinosaurs), currently known only from Laramidia and Asia. Remarkably, previous studies have postulated the occurrence of latitudinally arrayed dinosaur “provinces,” or “biomes,” on Laramidia. Yet this hypothesis has been challenged on multiple fronts and has remained poorly tested. Methodology/Principal Findings Here we describe two new, co-occurring ceratopsids from the Upper Cretaceous Kaiparowits Formation of Utah that provide the strongest support to date for the dinosaur provincialism hypothesis. Both pertain to the clade of ceratopsids known as Chasmosaurinae, dramatically increasing representation of this group from the southern portion of the Western Interior Basin of North America. Utahceratops gettyi gen. et sp. nov.—characterized by short, rounded, laterally projecting supraorbital horncores and an elongate frill with a deep median embayment—is recovered as the sister taxon to Pentaceratops sternbergii from the late Campanian of New Mexico. Kosmoceratops richardsoni gen. et sp. nov.—characterized by elongate, laterally projecting supraorbital horncores and a short, broad frill adorned with ten well developed hooks—has the most ornate skull of any known dinosaur and is closely allied to Chasmosaurus irvinensis from the late Campanian of Alberta. Conclusions/Significance Considered in unison, the phylogenetic, stratigraphic, and biogeographic evidence documents distinct, co-occurring chasmosaurine taxa north and south on the diminutive landmass of Laramidia. The famous Triceratops and all other, more nested chasmosaurines are postulated as descendants of forms previously restricted to the southern portion of Laramidia. Results further suggest the presence of latitudinally arrayed evolutionary centers of endemism within chasmosaurine ceratopsids during the late Campanian, the first documented occurrence of intracontinental endemism within dinosaurs.

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 juvenile specimen of a new coelurosaur (Dinosauria: Theropoda) from the Middle–Late Jurassic Shishugou Formation of Xinjiang, People's Republic of China

We describe the anatomy of a new coelurosaurian theropod Aorun zhaoi gen. et sp. nov., from the Middle–Late Jurassic of Xinjiang, China. Histological analysis of the holotype and only known specimen shows that the new taxon is represented by the skeleton of a juvenile individual aged no more than one year. A phylogenetic analysis of theropod relationships places Aorun as a basal member of the Coelurosauria. Although the sole use of a sub-adult ontogenetic exemplar is potentially problematic for phylogenetic reconstruction, we show that the phylogenetic position of Aorun as a member of Coelurosauria is robust to the exclusion of characters known to change during theropod ontogeny. Aorun is the seventh theropod taxon, and temporally oldest coelurosaur, known from the Shishugou Formation, which has one of the most taxonomically diverse Jurassic coelurosaurian theropod faunas in the world. http://zoobank.org/urn:lsid:zoobank.org:pub:5CC73577-9EB3-47AB-9983-1677B278EFFD

Turning semicircular canal function on its head: dinosaurs and a novel vestibular analysis.

Previous investigations have correlated vestibular function to locomotion in vertebrates by scaling semicircular duct radius of curvature to body mass. However, this method fails to discriminate bipedal from quadrupedal non-avian dinosaurs. Because they exhibit a broad range of relative head sizes, we use dinosaurs to test the hypothesis that semicircular ducts scale more closely with head size. Comparing the area enclosed by each semicircular canal to estimated body mass and to two different measures of head size, skull length and estimated head mass, reveals significant patterns that corroborate a connection between physical parameters of the head and semicircular canal morphology. Head mass more strongly correlates with anterior semicircular canal size than does body mass and statistically separates bipedal from quadrupedal taxa, with bipeds exhibiting relatively larger canals. This morphologic dichotomy likely reflects adaptations of the vestibular system to stability demands associated with terrestrial locomotion on two, versus four, feet. This new method has implications for reinterpreting previous studies and informing future studies on the connection between locomotion type and vestibular function.

Turanoceratops tardabilis—sister taxon, but not a ceratopsid

The ceratopsian dinosaur Turanoceratops tardabilis, from the Upper Cretaceous of Uzbekistan, has been assigned variously as a member of Ceratopsidae (the clade including Triceratops and relatives; Nessov et al. 1989), sister taxon to Ceratopsidae (e.g., Sereno 1997), or a nomen dubium (e.g., Dodson et al. 2004). Cladistic affiliation is of considerable importance for this problematic taxon because all other members of the diverse Ceratopsidae are restricted to western North America. In a recent paper published in Naturwissenschaften, (Sues and Averianov 2009) described newly recovered specimens of Turanoceratops and conducted a cladistic analysis of Ceratopsia. Based on their study, the authors concluded that Turanoceratops is indeed a member of the less inclusive Ceratopsidae and, thus, represents “the first definite ceratopsid dinosaur recorded from Asia.” This result has important implications for the biogeography, origin, and evolution of horned dinosaurs. The ingroup taxa used for their phylogenetic analysis of Turanoceratops (Sues and Averianov 2009) included a thorough representation of non-ceratopsid ceratopsians, but ceratopsids were restricted to two genera (Centrosaurus and Triceratops). Additionally, the source matrix (Makovicky and Norell 2006) was not focused on Ceratopsidae, so many informative phylogenetic characters pertaining to the clade were not included. In order to better evaluate the phylogenetic relationships of Turanoceratops relative to Ceratopsidae, we scored the taxon for the matrix previously published by Dodson et al. (2004). This matrix includes Protoceratops as an outgroup, Zuniceratops, and 13 ceratopsid genera. Eight out of 73 characters were scored for Turanoceratops as follows (character number followed by state in parentheses): 21(1), 22(0), 25(0), 29(0), 57(0), 60(1), 61(1), and 62(0). The matrix was analyzed in PAUP* (Swofford 2003), and three equally parsimonious trees (length=85, consistency index=0.9059, retention index=0.9437) were recovered (Fig. 1). In all three trees, Turanoceratops fell outside of Ceratopsidae (defined as all taxa descended from the most recent common ancestor of Triceratops and Centrosaurus; Dodson et al. 2004), and an additional two steps were required to move Turanoceratops within Ceratopsidae. Turanoceratops lacks at least two features shared by all ceratopsids in this data set: a true supracranial sinus complex and reduced subsidiary ridges on the teeth. The postorbital is remarkably similar to that of the derived neoceratopsian Zuniceratops, particularly in the presence of Naturwissenschaften (2009) 96:869–870 DOI 10.1007/s00114-009-0543-8

A Late Cretaceous (Maastrichtian) Avifauna from the Maevarano Formation, Madagascar

ABSTRACT Recent field efforts in the Mahajanga Basin of northwestern Madagascar have recovered a diverse Late Cretaceous terrestrial and freshwater vertebrate fauna, including a growing diversity of avialans. Previous work on associated bird skeletons resulted in the description of two named avialans (Rahonavis, Vorona). Other materials, including two synsacra and numerous appendicular elements, represent at least five additional taxa of basal (non-neornithine) birds. Among the materials described herein are two humeri tentatively referred to Rahonavis and numerous elements (e.g., humeri, ulnae, tibiotarsi, tarsometatarsi) assigned to Vorona. A near-complete carpometacarpus exhibits a minor metacarpal that exceeds the major metacarpal in length, documenting an enantiornithine in the fauna. Moreover, two additional, small humeri, an ulna, a femur, and a tarsometatarsus also compare favorably with enantiornithines. Finally, two other isolated humeri and a synsacrum are referable to Ornithurae. The latter specimen is notable in the presence of distinct, transversely oriented lumbosacral canals along the inner surface of the bony neural canal. This reveals for the first time a hard-tissue correlate of an anatomical specialization related to increased sensorimotor integration, one likely related to the unique form of avialan bipedal locomotion. Bird fossils recovered from the Maevarano Formation document one of the most size- and phylogenetically diverse Cretaceous-age Gondwanan avifaunas, including representative (1) basal pygostylian, (2) enantiornithine, (3) nonenantiornithine, ornithothoracine, and (4) ornithurine taxa. This Maastrichtian avifauna is notable in that it demonstrates the co-existence of multiple clades of basal (non-neornithine) birds until at least the end of the Mesozoic.

Cranial anatomy of Yinlong downsi (Ornithischia: Ceratopsia) from the Upper Jurassic Shishugou Formation of Xinjiang, China

ABSTRACT Yinlong downsi, from the Upper Jurassic Shishugou Formation, Xinjiang, northwestern China, is the oldest known ceratopsian dinosaur. Here we provide a detailed description of the skull and mandible based on the holotype, three partial skulls, and disarticulated materials from several other specimens. Yinlong can be diagnosed by six autapomorphies: a distinct fossa along the midline of the frontals; a slit-like carotid canal bordered by laminae; premaxillary teeth with a vertical wear facet and a basal shelf; a deep sulcus on the ventral surface of the quadratojugal; large oval nodules concentrated on the lateral surface of the jugal; and a squamosal with an expanded dorsal surface and a long, constricted quadrate process. A detailed comparison with other basal ceratopsians suggests that Yinlong downsi may belong within Chaoyangsauridae. Yinlong also shares many derived characters both with Psittacosaurus and neoceratopsians, suggesting that character evolution in early ceratopsians is complex. Additionally, a caniniform premaxillary tooth is present in one small specimen (IVPP V18636), which may suggest sexual dimorphism or individual variation.

Postcranial anatomy of Yinlong downsi (Dinosauria: Ceratopsia) from the Upper Jurassic Shishugou Formation of China and the phylogeny of basal ornithischians

Ceratopsia includes some of the best-known ornithischian dinosaurs. Many species are erected based on cranial elements alone, and the postcranial skeletons are either missing or undescribed in many taxa. Here we provide the first detailed postcranial description of Yinlong downsi based on the holotype and eight other well-preserved skeletons. Yinlong downsi from the early Late Jurassic Shishugou Formation of the Wucaiwan area, Xinjiang, China, represents one of the most basal ceratopsians. The detailed study of the postcranial skeleton reveals one feature unique to it among ceratopsians: a blade-like prepubic process of the pubis with an elongate notch near its ventral margin. The postcranial material of Yinlong shares some unique features with that of the ornithischian Stenopelix valdensis from the Early Cretaceous of Germany, and provides further evidence that the latter is a basal ceratopsian. A comprehensive phylogenetic analysis of basal ornithischians was built based on 72 taxa and 380 characters. Most of the characters are illustrated for the first time in order to clarify character states. The new ornithischian phylogeny confirms that Yinlong belongs to Chaoyangsauridae. Chaoyangsaurids and Psittacosaurus form a monophyletic group that is sister to all other ceratopsians. The new phylogeny also supports Stenopelix valdensis as a basal ceratopsian, and Mosaiceratops to be close to Coronosauria. Additionally, the new phylogeny agrees with other recent analyses that place heterodontosaurids as the most basal ornithischians rather than with marginocephalians. Furthermore, Isaberrysaura, which has been hypothesized to be a basal ornithopod, is recovered as one of the most basal stegosaurs for the first time. The former ‘hypsilophodontid’ taxa are recovered within Ornithopoda rather than outside Cerapoda, and Jeholosauridae is shown to be valid in this analysis.