Andrea Cau

Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds

Turning large dinosaurs into small birds Most paleontologists agree that birds are descended from dinosaurs. How did such large terrestrial or aquatic animals evolve into small feathered fliers? Lee et al. used two large databases of theropod morphology to explore possible evolutionary patterns that may have driven this dramatic transformation (see the Perspective by Benton). They found no clear pattern of miniaturization across the entire clade of Theropoda. However, several lines of evidence suggested that the lineage leading to birds underwent sustained miniaturization. Within that lineage, body sizes decreased and species evolved faster. They also developed ecological and morphological innovations linked to smaller body sizes. Science, this issue p. 562; see also p. 508 Birds evolved from their dinosaurian ancestors through a period characterized by sustained and repeated decreases in size. [Also see Perspective by Benton] Recent discoveries have highlighted the dramatic evolutionary transformation of massive, ground-dwelling theropod dinosaurs into light, volant birds. Here, we apply Bayesian approaches (originally developed for inferring geographic spread and rates of molecular evolution in viruses) in a different context: to infer size changes and rates of anatomical innovation (across up to 1549 skeletal characters) in fossils. These approaches identify two drivers underlying the dinosaur-bird transition. The theropod lineage directly ancestral to birds undergoes sustained miniaturization across 50 million years and at least 12 consecutive branches (internodes) and evolves skeletal adaptations four times faster than other dinosaurs. The distinct, prolonged phase of miniaturization along the bird stem would have facilitated the evolution of many novelties associated with small body size, such as reorientation of body mass, increased aerial ability, and paedomorphic skulls with reduced snouts but enlarged eyes and brains.

A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds.

The recent discovery of small paravian theropod dinosaurs with well-preserved feathers in the Middle–Late Jurassic Tiaojishan Formation of Liaoning Province (northeastern China) has challenged the pivotal position of Archaeopteryx, regarded from its discovery to be the most basal bird. Removing Archaeopteryx from the base of Avialae to nest within Deinonychosauria implies that typical bird flight, powered by the forelimbs only, either evolved at least twice, or was subsequently lost or modified in some deinonychosaurians. Here we describe the complete skeleton of a new paravian from the Tiaojishan Formation of Liaoning Province, China. Including this new taxon in a comprehensive phylogenetic analysis for basal Paraves does the following: (1) it recovers it as the basal-most avialan; (2) it confirms the avialan status of Archaeopteryx; (3) it places Troodontidae as the sister-group to Avialae; (4) it supports a single origin of powered flight within Paraves; and (5) it implies that the early diversification of Paraves and Avialae took place in the Middle–Late Jurassic period.

Morphological Clocks in Paleontology, and a Mid-Cretaceous Origin of Crown Aves

Birds are among the most diverse and intensivelystudied vertebrate groups, but many aspects of theirhigher-level phylogeny and evolution still remaincontroversial. One contentious issue concerns theantiquity of modern birds (=crown Aves): the ageof the most recent common ancestor of all livingbirds (Gauthier 1986). Very few Mesozoic fossilsare attributable to modern birds (e.g., Clarke et al.2005; Dyke and Kaiser 2011; Brocklehurst et al. 2012;Ksepka and Boyd 2012) suggesting that they diversifiedlargely or entirely in the early Paleogene, perhaps in theecologicalvacuumcreatedbytheextinctionofnon-aviandinosaurs, pterosaurs, and many archaic (stem) birds(e.g.,Longrichetal.2011).Incontrast,molecularstudiesindicate that modern birds commenced radiating deepwithin the Mesozoic, for example ∼130Ma(Cooperand Penny 1997; Haddrath and Baker 2012)or∼113 Ma(Jetz et al. 2012), with ratites, galliforms, anseriforms,shorebirds, and even passerines surviving acrossthe KPg boundary (∼66 Ma). The oldest moleculardates further imply an extraordinarily rapid earlybird evolution, with the modern birds appearingonly 20 myr after

A new sauropod dinosaur from the Early Cretaceous of Tunisia with extreme avian-like pneumatization

Recent interpretations of the postcranial anatomy of sauropod dinosaurs differ about pneumatic features supporting an avian-like ventilatory system; the most conservative workers reject most postcranial pneumatizations as being unambiguous evidence of abdominal air sacs. Here we describe the first articulated dinosaur skeleton from Tunisia and refer it to a new rebbachisaurid sauropod, Tataouinea hannibalis gen. et sp. nov. The Tunisian specimen shows a complex pattern of caudosacral and pelvic pneumatization--including the first report of an ischial pneumatic foramen among Dinosauria--strongly supporting the presence of abdominal air sacs. Character optimization among Rebbachisauridae indicates that in the caudal vertebrae, pneumatization of the neural arches preceded that of the centra; in the pelvis, pneumatization of the bones adjacent to the sacrum preceded that of more distal elements. Tataouinea was more closely related to European nigersaurines than to otherwise Gondwanan rebbachisaurids; this supports an Afro-European route for rebbachisaurid dispersal.

New Information on Tataouinea hannibalis from the Early Cretaceous of Tunisia and Implications for the Tempo and Mode of Rebbachisaurid Sauropod Evolution

The rebbachisaurid sauropod Tataouinea hannibalis represents the first articulated dinosaur skeleton from Tunisia and one of the best preserved in northern Africa. The type specimen was collected from the lower Albian, fluvio-estuarine deposits of the Ain el Guettar Formation (southern Tunisia). We present detailed analyses on the sedimentology and facies distribution at the main quarry and a revision of the vertebrate fauna associated with the skeleton. Data provide information on a complex ecosystem dominated by crocodilian and other brackish water taxa. Taphonomic interpretations indicate a multi-event, pre-burial history with a combination of rapid segregation in high sediment supply conditions and partial subaerial exposure of the carcass. After the collection in 2011 of the articulated sacrum and proximalmost caudal vertebrae, all showing a complex pattern of pneumatization, newly discovered material of the type specimen allows a detailed osteological description of Tataouinea. The sacrum, the complete and articulated caudal vertebrae 1–17, both ilia and ischia display asymmetrical pneumatization, with the left side of vertebrae and the left ischium showing a more extensive invasion by pneumatic features than their right counterparts. A pneumatic hiatus is present in caudal centra 7 to 13, whereas caudal centra 14–16 are pneumatised by shallow fossae. Bayesian inference analyses integrating morphological, stratigraphic and paleogeographic data support a flagellicaudatan-rebbachisaurid divergence at about 163 Ma and a South American ancestral range for rebbachisaurids. Results presented here suggest an exclusively South American Limaysaurinae and a more widely distributed Rebbachisaurinae lineage, the latter including the South American taxon Katepensaurus and a clade including African and European taxa, with Tataouinea as sister taxon of Rebbachisaurus. This scenario would indicate that South America was not affected by the end-Jurassic extinction of diplodocoids, and was most likely the centre of the rapid radiation of rebbachisaurids to Africa and Europe between 135 and 130 Ma.

A gigantic bird from the Upper Cretaceous of Central Asia.

We describe an enormous Late Cretaceous fossil bird from Kazakhstan, known from a pair of edentulous mandibular rami (greater than 275 mm long), which adds significantly to our knowledge of Mesozoic avian morphological and ecological diversity. A suite of autapomorphies lead us to recognize the specimen as a new taxon. Phylogenetic analysis resolves this giant bird deep within Aves as a basal member of Ornithuromorpha. This Kazakh fossil demonstrates that large body size evolved at least once outside modern birds (Neornithes) and reveals hitherto unexpected trophic diversity within Cretaceous Aves.

The phylogenetic affinities of the bizarre Late Cretaceous Romanian theropod Balaur bondoc (Dinosauria, Maniraptora): dromaeosaurid or flightless bird?

The exceptionally well-preserved Romanian dinosaur Balaur bondoc is the most complete theropod known to date from the Upper Cretaceous of Europe. Previous studies of this remarkable taxon have included its phylogenetic interpretation as an aberrant dromaeosaurid with velociraptorine affinities. However, Balaur displays a combination of both apparently plesiomorphic and derived bird-like characters. Here, we analyse those features in a phylogenetic revision and show how they challenge its referral to Dromaeosauridae. Our reanalysis of two distinct phylogenetic datasets focusing on basal paravian taxa supports the reinterpretation of Balaur as an avialan more crownward than Archaeopteryx but outside of Pygostylia, and as a flightless taxon within a paraphyletic assemblage of long-tailed birds. Our placement of Balaur within Avialae is not biased by character weighting. The placement among dromaeosaurids resulted in a suboptimal alternative that cannot be rejected based on the data to hand. Interpreted as a dromaeosaurid, Balaur has been assumed to be hypercarnivorous and predatory, exhibiting a peculiar morphology influenced by island endemism. However, a dromaeosaurid-like ecology is contradicted by several details of Balaur’s morphology, including the loss of a third functional manual digit, the non-ginglymoid distal end of metatarsal II, and a non-falciform ungual on the second pedal digit that lacks a prominent flexor tubercle. Conversely, an omnivorous ecology is better supported by Balaur’s morphology and is consistent with its phylogenetic placement within Avialae. Our reinterpretation of Balaur implies that a superficially dromaeosaurid-like taxon represents the enlarged, terrestrialised descendant of smaller and probably volant ancestors.

Evidence of a new carcharodontosaurid from the Upper Cretaceous of Morocco

We report an isolated frontal of a large-bodied theropod from the Cenomanian “Kern Kern beds” of Morocco with an unusual morphology that we refer to a new carcharodontosaurid distinct from the sympatric Carcharodontosaurus. The specimen shows an unique combination of plesiomorphic and potentially autapomorphic features: very thick and broad bone with a complex saddle-shaped dorsal surface, and a narrow vertical lamina between the prefrontal and lacrimal facets. This study supports the hypothesis that a fourth large theropod was present in the Cenomanian of Morocco together with Carcharodontosaurus, Deltadromeus, and Spinosaurus.

Application of numerical cladistic analyses to the Carnian–Norian conodonts: a new approach for phylogenetic interpretations

The high intraspecific variability of conodont platform elements in the upper Carnian–lower Norian interval and the proliferation of numerous species in this relatively short time have generated many problems for the understanding of Late Triassic conodont phylogeny, systematics and taxonomy. Since Late Triassic natural assemblages are still unknown, we have applied cladistic methodologies to investigate the evolution of Carnian–Norian conodont platforms and to infer more precise phylogenetic relationships among taxa. Numerical cladistic analysis was undertaken of species belonging to the five Late Triassic genera Paragondolella, Carnepigondolella, Metapolygnathus, Epigondolella and Norigondolella. A taxon–character data matrix describing the distribution of 64 characters amongst two outgroups and 31 ingroup taxa was compiled and processed using PAUP* 4.1. Our analyses show the evolutionary and systematic value of certain morphological characters, and lead to a reinterpretation of the phylogenetic position of the genera considered: Metapolygnathus, Epigondolella and Norigondolella are monphyletic taxa, Paragondolella represents a polyphyletic assemblage of basal members of the ingroup, and Carnepigondolella a paraphyletic group including primitive forms of the Metapolygnathus and Epigondolella lineages. Our analyses also clarify the systematic position of ‘Metapolygnathus communisti B’ and confirm the existence of four new species: Carnepigondolella angulata sp. nov., Epigondolella heinzi sp. nov., Epigondolella miettoi sp. nov. and Norigondolella trinacriae sp. nov.

The affinities of ‘Steneosaurus barettoni’ (Crocodylomorpha, Thalattosuchia), from the Jurassic of Northern Italy, and implications for cranial evolution among geosaurine metriorhynchids

In 1787, a fragmentary fossil skull of a crocodylian was discovered in the Altopiano dei Sette Comuni (Veneto, Northern Italy). In 1883, the specimen was referred to a new species of the teleosauroid Steneosaurus, S. barettoni. In this study, the specimen is redescribed and its taxonomic status reviewed. S. barettoni fails to conform to Article 12 of the International Code of Zoological Nomenclature and is considered as a nomen nudum. The specimen lacks synapomorphies of both Teleosauroidea and Steneosaurus, whereas it is referred to Metriorhynchidae based on the presence of an elongate median depression between the dorsal surface of nasals, and a large teardrop-shaped prefrontal that is expanded laterally. The combination of cranial and dental features present in this specimen supports its referral to Neptunidraco, an interpretation that adds new information on the morphology of the Italian metriorhynchid. Although total evidence analysis placed Neptunidraco as a basal divergence among Geosaurinae, it independently evolved some craniomandibular features previously considered exclusive of the macrophagous Geosaurini. This specimen is historically relevant because it may represent the first metriorhynchoid found worldwide.