Jurassic shuotheriids show earliest dental diversification of mammaliaforms.

Shuotheriids are Jurassic mammaliaforms that possess pseudotribosphenic teeth in which a pseudotalonid is anterior to the trigonid in the lower molar, contrasting with the tribosphenic pattern of therian mammals (placentals, marsupials and kin) in which the talonid is posterior to the trigonid1-4. The origin of the pseudotribosphenic teeth remains unclear, obscuring our perception of shuotheriid affinities and the early evolution of mammaliaforms1,5-9. Here we report a new Jurassic shuotheriid represented by two skeletal specimens. Their complete pseudotribosphenic dentitions allow reidentification of dental structures using serial homology and the tooth occlusal relationship. Contrary to the conventional view1,2,6,10,11, our findings show that dental structures of shuotheriids can be homologized to those of docodontans and partly support homologous statements for some dental structures between docodontans and other mammaliaforms6,12. The phylogenetic analysis based on new evidence removes shuotheriids from the tribosphenic ausktribosphenids (including monotremes) and clusters them with docodontans to form a new clade, Docodontiformes, that is characterized by pseudotribosphenic features. In the phylogeny, docodontiforms and 'holotherians' (Kuehneotherium, monotremes and therians)13 evolve independently from a Morganucodon-like ancestor with triconodont molars by labio-lingual widening their posterior teeth for more efficient food processing. The pseudotribosphenic pattern passed a cusp semitriangulation stage9, whereas the tribosphenic pattern and its precursor went through a stage of cusp triangulation. The two different processes resulted in complex tooth structures and occlusal patterns that elucidate the earliest diversification of mammaliaforms.

Fossils document evolutionary changes of jaw joint to mammalian middle ear.

The dual jaw joint of Morganucodon1,2 consists of the dentary-squamosal joint laterally and the articular-quadrate one medially. The articular-quadrate joint and its associated post-dentary bones constitute the precursor of the mammalian middle ear. Fossils documenting the transition from such a precursor to the mammalian middle ear are poor, resulting in inconsistent interpretations of this hallmark apparatus in the earliest stage of mammaliaform evolution1-5. Here we report mandibular middle ears from two Jurassic mammaliaforms: a new morganucodontan-like species and a pseudotribosphenic shuotheriid species6. The morganucodontan-like species shows many previously unknown post-dentary bone morphologies1,2 and exhibits features that suggest a loss of load-bearing function in its articular-quadrate joint. The middle ear of the shuotheriid approaches the mammalian condition in that it has features that are suitable for an exclusively auditory function, although the post-dentary bones are still attached to the dentary. With size reduction of the jaw-joint bones, the quadrate shifts medially at different degrees in relation to the articular in the two mammaliaforms. These changes provide evidence of a gradual loss of load-bearing function in the articular-quadrate jaw joint-a prerequisite for the detachment of the post-dentary bones from the dentary7-12 and the eventual breakdown of the Meckel's cartilage13-15 during the evolution of mammaliaforms.

A putative triradial macrofossil from the Ediacaran Jiangchuan Biota.

The late Ediacaran Jiangchuan biota, from the Dengying Formation in eastern Yunnan, is well-known for its diverse macroalgal fossils, opening a window onto eukaryotic-dominated ecosystems from the late Neoproterozoic of South China. Although multiple lines of evidence suggest that metazoans had already evolved by the late Ediacaran, animal fossils have not yet been formally described from this locality. Here, we report a putative disc-shaped macrofossil from the Jiangchuan biota, Lobodiscus tribrachialis gen. et sp. nov. This specimen shows the triradial symmetry characteristic of trilobozoans, a group of Ediacaran macrofossils previously documented in Australia and Russia. Lobodiscus could record the youngest known occurrence of trilobozoans, strengthening taxonomic and ecological continuities between the Ediacaran "White Sea" and "Nama" assemblages. Our findings may expand the known paleogeographical distribution of trilobozoans and provide data for Ediacaran biostratigraphic correlations across the Yangtze block and globally, helping to track the diversification of early metazoan-grade organisms.

First monotreme from the Late Cretaceous of South America.

Monotremata is a clade of egg-lying mammals, represented by the living platypus and echidnas, which is endemic to Australia, and adjacent islands. Occurrence of basal monotremes in the Early Cretaceous of Australia has led to the consensus that this clade originated on that continent, arriving later to South America. Here we report on the discovery of a Late Cretaceous monotreme from southern Argentina, demonstrating that monotremes were present in circumpolar regions by the end of the Mesozoic, and that their distinctive anatomical features were probably present in these ancient forms as well.

Earliest known Gondwanan bird tracks: Wonthaggi Formation (Early Cretaceous), Victoria, Australia.

The fossil record for Cretaceous birds in Australia has been limited to rare skeletal material, feathers, and two tracks, a paucity shared with other Gondwanan landmasses. Hence the recent discovery of 27 avian footprints and other traces in the Early Cretaceous (Barremian-Aptian, 128-120 Ma) Wonthaggi Formation of Victoria, Australia amends their previous rarity there, while also confirming the earliest known presence of birds in Australia and the rest of Gondwana. The avian identity of these tracks is verified by their tridactyl forms, thin digits relative to track lengths, wide divarication angles, and sharp claws; three tracks also have hallux imprints. Track forms and sizes indicate a variety of birds as tracemakers, with some among the largest reported from the Early Cretaceous. Although continuous trackways are absent, close spacing and similar alignments of tracks on some bedding planes suggest gregariousness. The occurrence of this avian trace-fossil assemblage in circumpolar fluvial-floodplain facies further implies seasonal behavior, with trackmakers likely leaving their traces on floodplain surfaces during post-thaw summers.

First ceratosaurian dinosaur from Australia.

The basal theropod dinosaur clade Ceratosauria, and its subclade Abelisauroidea, is characteristic of late Mesozoic terrestrial vertebrate faunas in western Gondwana (South America, Africa, Madagascar, and India) and Europe. Yet unambiguous records of ceratosaurs have hitherto been absent from Australia, where the theropod assemblage appears to include several typically Laurasian clades. Here, we report the first evidence of ceratosaurs (and potentially abelisauroids) from eastern Gondwana--a diagnostic astragalocalcaneum from the Aptian (121-125 Ma) of Victoria, Australia. Ceratosauria thus occurred in both western and eastern Gondwana during the Early Cretaceous. This fossil adds to the poorly known dinosaur fauna of Australia, a major clade of basal theropods, emphasising that its mid-Cretaceous theropod diversity was surprisingly cosmopolitan despite relative geographic isolation, including clades that have been thought to be typical of both Gondwana and Laurasia--Ceratosauria, Spinosauridae, Carcharodontosauria, Tyrannosauroidea, and Deinonychosauria. Such a contemporaneous association of theropod clades is unknown from other Gondwanan continents and questions the views that the late Mesozoic dinosaur fauna of Australia was dominated by Gondwanan or Laurasian elements, extreme isolation, relictualism, and/or novelty as a 'centre of origin'. The cosmopolitan theropod fauna of Australia probably reflects the global distribution of these clades early in their history, prior to significant continental breakup.

First spinosaurid dinosaur from Australia and the cosmopolitanism of Cretaceous dinosaur faunas.

A cervical vertebra from the Early Cretaceous of Victoria represents the first Australian spinosaurid theropod dinosaur. This discovery significantly extends the geographical range of spinosaurids, suggesting that the clade obtained a near-global distribution before the onset of Pangaean fragmentation. The combined presence of spinosaurid, neovenatorid, tyrannosauroid and dromaeosaurid theropods in the Australian Cretaceous undermines previous suggestions that the dinosaur fauna of this region was either largely endemic or predominantly 'Gondwanan' in composition. Many lineages are well-represented in both Laurasia and Gondwana, and these observations suggest that Early-'middle' Cretaceous theropod clades possessed more cosmopolitan distributions than assumed previously, and that caution is necessary when attempting to establish palaeobiogeographic patterns on the basis of a patchily distributed fossil record.

The oldest platypus and its bearing on divergence timing of the platypus and echidna clades.

Monotremes have left a poor fossil record, and paleontology has been virtually mute during two decades of discussion about molecular clock estimates of the timing of divergence between the platypus and echidna clades. We describe evidence from high-resolution x-ray computed tomography indicating that Teinolophos, an Early Cretaceous fossil from Australia's Flat Rocks locality (121-112.5 Ma), lies within the crown clade Monotremata, as a basal platypus. Strict molecular clock estimates of the divergence between platypus and echidnas range from 17 to 80 Ma, but Teinolophos suggests that the two monotreme clades were already distinct in the Early Cretaceous, and that their divergence may predate even the oldest strict molecular estimates by at least 50%. We generated relaxed molecular clock models using three different data sets, but only one yielded a date overlapping with the age of Teinolophos. Morphology suggests that Teinolophos is a platypus in both phylogenetic and ecological aspects, and tends to contradict the popular view of rapid Cenozoic monotreme diversification. Whereas the monotreme fossil record is still sparse and open to interpretation, the new data are consistent with much slower ecological, morphological, and taxonomic diversification rates for monotremes than in their sister taxon, the therian mammals. This alternative view of a deep geological history for monotremes suggests that rate heterogeneities may have affected mammalian evolution in such a way as to defeat strict molecular clock models and to challenge even relaxed molecular clock models when applied to mammalian history at a deep temporal scale.

The bone microstructure of polar "hypsilophodontid" dinosaurs from Victoria, Australia.

High-latitude (i.e., "polar") Mesozoic fauna endured months of twilight and relatively low mean annual temperatures. Yet non-avian dinosaurs flourished in this taxing environment. Fossils of basal ornithopod dinosaurs ("hypsilophodontids") are common in the Early Cretaceous high-latitude sediments of Victoria, Australia, and four taxa have been described; although their ontogenetic histories are largely unexplored. In the present study, eighteen tibiae and femora were utilized in the first multi-specimen ontogenetic histological analysis of Australian polar hypsilophodontids. The sample consists of eleven individuals from the Flat Rocks locality (Late Valanginian or Barremian), and five from the Dinosaur Cove locality (Albian). In both groups, growth was most rapid during the first three years, and skeletal maturity occurred between five and seven years. There is a weak asymptotic trend in a plot of growth mark count versus femur length, with considerable individual variation. Histology suggests two genera are present within the Dinosaur Cove sample, but bone microstructure alone could not distinguish genera within the Flat Rocks sample, or across the two geologically separate (~ 26 Ma) localities. Additional histologic sampling, combined with morphological analyses, may facilitate further differentiation between ontogenetic, individual, and species variation.

First dinosaurs from Saudi Arabia.

Dinosaur remains from the Arabian subcontinent are exceedingly rare, and those that have been documented manifest indeterminate affinities. Consequently the discovery of a small, but diagnostic, accumulation of elements from Campanian-Maastrichtian (~ 75 Ma) deposits in northwestern Saudi Arabia is significant because it constitutes the first taxonomically identifiable dinosaur material described from the Arabian Peninsula. The fossils include a series of possible lithostrotian titanosaur caudal vertebrae, and some isolated theropod marginal teeth that share unique character states and metric parameters (analyzed using multivariate statistical methods) with derived abelisaurids - this is the first justifiable example of a non-avian carnivorous dinosaur clade from Arabia. The recognition of titanosaurians and abelisaurids from Saudi Arabia extends the palaeogeographical range of these groups along the entire northern Gondwanan margin during the latest Cretaceous. Moreover, given the extreme paucity of coeval occurrences elsewhere, the Saudi Arabian fossils provide a tantalizing glimpse into dinosaurian assemblage diversity within the region.

Theropod fauna from southern Australia indicates high polar diversity and climate-driven dinosaur provinciality.

The Early Cretaceous fauna of Victoria, Australia, provides unique data on the composition of high latitude southern hemisphere dinosaurs. We describe and review theropod dinosaur postcranial remains from the Aptian-Albian Otway and Strzelecki groups, based on at least 37 isolated bones, and more than 90 teeth from the Flat Rocks locality. Several specimens of medium- and large-bodied individuals (estimated up to ~8.5 metres long) represent allosauroids. Tyrannosauroids are represented by elements indicating medium body sizes (~3 metres long), likely including the holotype femur of Timimus hermani, and a single cervical vertebra represents a juvenile spinosaurid. Single specimens representing medium- and small-bodied theropods may be referrable to Ceratosauria, Ornithomimosauria, a basal coelurosaur, and at least three taxa within Maniraptora. Thus, nine theropod taxa may have been present. Alternatively, four distinct dorsal vertebrae indicate a minimum of four taxa. However, because most taxa are known from single bones, it is likely that small-bodied theropod diversity remains underestimated. The high abundance of allosauroids and basal coelurosaurs (including tyrannosauroids and possibly ornithomimosaurs), and the relative rarity of ceratosaurs, is strikingly dissimilar to penecontemporaneous dinosaur faunas of Africa and South America, which represent an arid, lower-latitude biome. Similarities between dinosaur faunas of Victoria and the northern continents concern the proportional representatation of higher clades, and may result from the prevailing temperate-polar climate of Australia, especially at high latitudes in Victoria, which is similar to the predominant warm-temperate climate of Laurasia, but distinct from the arid climate zone that covered extensive areas of Gondwana. Most dinosaur groups probably attained a near-cosmopolitan distribution in the Jurassic, prior to fragmentation of the Pangaean supercontinent, and some aspects of the hallmark 'Gondwanan' fauna of South America and Africa may therefore reflect climate-driven provinciality, not vicariant evolution driven by continental fragmentation. However, vicariance may still be detected at lower phylogenetic levels.

Growth dynamics of Australia's polar dinosaurs.

Analysis of bone microstructure in ornithopod and theropod dinosaurs from Victoria, Australia, documents ontogenetic changes, providing insight into the dinosaurs' successful habitation of Cretaceous Antarctic environments. Woven-fibered bone tissue in the smallest specimens indicates rapid growth rates during early ontogeny. Later ontogeny is marked by parallel-fibered tissue, suggesting reduced growth rates approaching skeletal maturity. Bone microstructure similarities between the ornithopods and theropods, including the presence of LAGs in each group, suggest there is no osteohistologic evidence supporting the hypothesis that polar theropods hibernated seasonally. Results instead suggest high-latitude dinosaurs had growth trajectories similar to their lower-latitude relatives and thus, rapid early ontogenetic growth and the cyclical suspensions of growth inherent in the theropod and ornithopod lineages enabled them to successfully exploit polar regions.

Independent origins of middle ear bones in monotremes and therians.

A dentary of the oldest known monotreme, the Early Cretaceous Teinolophos trusleri, has an internal mandibular trough, which in outgroups to mammals houses accessory jaw bones, and probable contact facets for angular, coronoid, and splenial bones. Certain of these accessory bones were detached from the mandible to become middle ear bones in mammals. Evidence that the angular (homologous with the mammalian ectotympanic) and the articular and prearticular (homologous with the mammalian malleus) bones retained attachment to the lower jaw in a basal monotreme indicates that the definitive mammalian middle ear evolved independently in living monotremes and therians (marsupials and placentals).