A New Basal Sauropodomorph Dinosaur from the Lower Jurassic Navajo Sandstone of Southern Utah

Background

Basal sauropodomorphs, or ‘prosauropods,’ are a globally widespread paraphyletic assemblage of terrestrial herbivorous dinosaurs from the Late Triassic and Early Jurassic. In contrast to several other landmasses, the North American record of sauropodomorphs during this time interval remains sparse, limited to Early Jurassic occurrences of a single well-known taxon from eastern North America and several fragmentary specimens from western North America.

Methodology/Principal Findings

On the basis of a partial skeleton, we describe here a new basal sauropodomorph dinosaur from the Lower Jurassic Navajo Sandstone of southern Utah, Seitaad ruessi gen. et sp. nov. The partially articulated skeleton of Seitaad was likely buried post-mortem in the base of a collapsed dune foreset. The new taxon is characterized by a plate-like medial process of the scapula, a prominent proximal expansion of the deltopectoral crest of the humerus, a strongly inclined distal articular surface of the radius, and a proximally and laterally hypertrophied proximal metacarpal I.

Conclusions/Significance

Phylogenetic analysis recovers Seitaad as a derived basal sauropodomorph closely related to plateosaurid or massospondylid ‘prosauropods’ and its presence in western North America is not unexpected for a member of this highly cosmopolitan clade. This occurrence represents one of the most complete vertebrate body fossil specimens yet recovered from the Navajo Sandstone and one of the few basal sauropodomorph taxa currently known from North America.     

A Basal Sauropodomorph (Dinosauria: Saurischia) from the Ischigualasto Formation (Triassic,Carnian) and the Early Evolution of Sauropodomorpha

Background

The earliest dinosaurs are from the early Late Triassic (Carnian) of South America. By the Carnian the main clades Saurischia and Ornithischia were already established, and the presence of the most primitive known sauropodomorph Saturnalia suggests also that Saurischia had already diverged into Theropoda and Sauropodomorpha. Knowledge of Carnian sauropodomorphs has been restricted to this single species.

Methodology/Principal Findings

We describe a new small sauropodomorph dinosaur from the Ischigualsto Formation (Carnian) in northwest Argentina, Panphagia protos gen. et sp. nov., on the basis of a partial skeleton. The genus and species are characterized by an anteroposteriorly elongated fossa on the base of the anteroventral process of the nasal; wide lateral flange on the quadrate with a large foramen; deep groove on the lateral surface of the lower jaw surrounded by prominent dorsal and ventral ridges; bifurcated posteroventral process of the dentary; long retroarticular process transversally wider than the articular area for the quadrate; oval scars on the lateral surface of the posterior border of the centra of cervical vertebrae; distinct prominences on the neural arc of the anterior cervical vertebra; distal end of the scapular blade nearly three times wider than the neck; scapular blade with an expanded posterodistal corner; and medial lamina of brevis fossa twice as wide as the iliac spine.

Conclusions/Significance

We regard Panphagia as the most basal sauropodomorph, which shares the following apomorphies with Saturnalia and more derived sauropodomorphs: basally constricted crowns; lanceolate crowns; teeth of the anterior quarter of the dentary higher than the others; and short posterolateral flange of distal tibia. The presence of Panphagia at the base of the early Carnian Ischigualasto Formation suggests an earlier origin of Sauropodomorpha during the Middle Triassic.     

A new sauropodomorph dinosaur from the Early Jurassic of Patagonia and the origin and evolution of the sauropod-type sacrum

Background

The origin of sauropod dinosaurs is one of the major landmarks of dinosaur evolution but is still poorly understood. This drastic transformation involved major skeletal modifications, including a shift from the small and gracile condition of primitive sauropodomorphs to the gigantic and quadrupedal condition of sauropods. Recent findings in the Late Triassic–Early Jurassic of Gondwana provide critical evidence to understand the origin and early evolution of sauropods.

Methodology/Principal Findings

A new sauropodomorph dinosaur, Leonerasaurus taquetrensis gen. et sp. nov., is described from the Las Leoneras Formation of Central Patagonia (Argentina). The new taxon is diagnosed by the presence of anterior unserrated teeth with a low spoon-shaped crown, amphicoelous and acamerate vertebral centra, four sacral vertebrae, and humeral deltopectoral crest low and medially deflected along its distal half. The phylogenetic analysis depicts Leonerasaurus as one of the closest outgroups of Sauropoda, being the sister taxon of a clade of large bodied taxa composed of Melanorosaurus and Sauropoda.

Conclusions/Significance

The dental and postcranial anatomy of Leonerasaurus supports its close affinities with basal sauropods. Despite the small size and plesiomorphic skeletal anatomy of Leonerasaurus, the four vertebrae that compose its sacrum resemble that of the large-bodied primitive sauropods. This shows that the appearance of the sauropod-type of sacrum predated the marked increase in body size that characterizes the origins of sauropods, rejecting a causal explanation and evolutionary linkage between this sacral configuration and body size. Alternative phylogenetic placements of Leonerasaurus as a basal anchisaurian imply a convergent acquisition of the sauropod-type sacrum in the new small-bodied taxon, also rejecting an evolutionary dependence of sacral configuration and body size in sauropodomorphs. This and other recent discoveries are showing that the characteristic sauropod body plan evolved gradually, with a step-wise pattern of character appearance.     

The braincase of the basal sauropod dinosaur Spinophorosaurus and 3D reconstructions of the cranial endocast and inner ear

Background

Sauropod dinosaurs were the largest animals ever to walk on land, and, as a result, the evolution of their remarkable adaptations has been of great interest. The braincase is of particular interest because it houses the brain and inner ear. However, only a few studies of these structures in sauropods are available to date. Because of the phylogenetic position of Spinophorosaurus nigerensis as a basal eusauropod, the braincase has the potential to provide key evidence on the evolutionary transition relative to other dinosaurs.

Methodology/Principal Findings

The only known braincase of Spinophorosaurus (‘Argiles de l''Irhazer’, Irhazer Group; Agadez region, Niger) differs significantly from those of the Jurassic sauropods examined, except potentially for Atlasaurus imelakei (Tilougguit Formation, Morocco). The basisphenoids of Spinophorosaurus and Atlasaurus bear basipterygoid processes that are comparable in being directed strongly caudally. The Spinophorosaurus specimen was CT scanned, and 3D renderings of the cranial endocast and inner-ear system were generated. The endocast resembles that of most other sauropods in having well-marked pontine and cerebral flexures, a large and oblong pituitary fossa, and in having the brain structure obscured by the former existence of relatively thick meninges and dural venous sinuses. The labyrinth is characterized by long and proportionally slender semicircular canals. This condition recalls, in particular, that of the basal non-sauropod sauropodomorph Massospondylus and the basal titanosauriform Giraffatitan.

Conclusions/Significance

Spinophorosaurus has a moderately derived paleoneuroanatomical pattern. In contrast to what might be expected early within a lineage leading to plant-eating graviportal quadrupeds, Spinophorosaurus and other (but not all) sauropodomorphs show no reduction of the vestibular apparatus of the inner ear. This character-state is possibly a primitive retention in Spinophorosaurus, but due the scarcity of data it remains unclear whether it is also the case in the various later sauropods in which it is present or whether it has developed homoplastically in these taxa. Any interpretations remain tentative pending the more comprehensive quantitative analysis underway, but the size and morphology of the labyrinth of sauropodomorphs may be related to neck length and mobility, among other factors.     

New basal iguanodonts from the Cedar Mountain formation of Utah and the evolution of thumb-spiked dinosaurs

Background

Basal iguanodontian dinosaurs were extremely successful animals, found in great abundance and diversity almost worldwide during the Early Cretaceous. In contrast to Europe and Asia, the North American record of Early Cretaceous basal iguanodonts has until recently been limited largely to skulls and skeletons of Tenontosaurus tilletti.

Methodology/Principal Findings

Herein we describe two new basal iguanodonts from the Yellow Cat Member of the Cedar Mountain Formation of eastern Utah, each known from a partial skull and skeleton. Iguanacolossus fortis gen. et sp. nov. and Hippodraco scutodens gen. et sp. nov. are each diagnosed by a single autapomorphy and a unique combination of characters.

Conclusions/Significance

Iguanacolossus and Hippodraco add greatly to our knowledge of North American basal iguanodonts and prompt a new comprehensive phylogenetic analysis of basal iguanodont relationships. This analysis indicates that North American Early Cretaceous basal iguanodonts are more basal than their contemporaries in Europe and Asia.     

A New Basal Sauropod Dinosaur from the Middle Jurassic of Niger and the Early Evolution of Sauropoda

Background

The early evolution of sauropod dinosaurs is poorly understood because of a highly incomplete fossil record. New discoveries of Early and Middle Jurassic sauropods have a great potential to lead to a better understanding of early sauropod evolution and to reevaluate the patterns of sauropod diversification.

Principal Findings

A new sauropod from the Middle Jurassic of Niger, Spinophorosaurus nigerensis n. gen. et sp., is the most complete basal sauropod currently known. The taxon shares many anatomical characters with Middle Jurassic East Asian sauropods, while it is strongly dissimilar to Lower and Middle Jurassic South American and Indian forms. A possible explanation for this pattern is a separation of Laurasian and South Gondwanan Middle Jurassic sauropod faunas by geographic barriers. Integration of phylogenetic analyses and paleogeographic data reveals congruence between early sauropod evolution and hypotheses about Jurassic paleoclimate and phytogeography.

Conclusions

Spinophorosaurus demonstrates that many putatively derived characters of Middle Jurassic East Asian sauropods are plesiomorphic for eusauropods, while South Gondwanan eusauropods may represent a specialized line. The anatomy of Spinophorosaurus indicates that key innovations in Jurassic sauropod evolution might have taken place in North Africa, an area close to the equator with summer-wet climate at that time. Jurassic climatic zones and phytogeography possibly controlled early sauropod diversification.     

New Mid-Cretaceous (Latest Albian) Dinosaurs from Winton,Queensland, Australia

Background

Australia''s dinosaurian fossil record is exceptionally poor compared to that of other similar-sized continents. Most taxa are known from fragmentary isolated remains with uncertain taxonomic and phylogenetic placement. A better understanding of the Australian dinosaurian record is crucial to understanding the global palaeobiogeography of dinosaurian groups, including groups previously considered to have had Gondwanan origins, such as the titanosaurs and carcharodontosaurids.

Methodology/Principal Findings

We describe three new dinosaurs from the late Early Cretaceous (latest Albian) Winton Formation of eastern Australia, including; Wintonotitan wattsi gen. et sp. nov., a basal titanosauriform; Diamantinasaurus matildae gen. et sp. nov., a derived lithostrotian titanosaur; and Australovenator wintonensis gen. et sp. nov., an allosauroid. We compare an isolated astragalus from the Early Cretaceous of southern Australia; formerly identified as Allosaurus sp., and conclude that it most-likely represents Australovenator sp.

Conclusion/Significance

The occurrence of Australovenator from the Aptian to latest Albian confirms the presence in Australia of allosauroids basal to the Carcharodontosauridae. These new taxa, along with the fragmentary remains of other taxa, indicate a diverse Early Cretaceous sauropod and theropod fauna in Australia, including plesiomorphic forms (e.g. Wintonotitan and Australovenator) and more derived forms (e.g. Diamantinasaurus).     

The postcranial anatomy of Coloradisaurus brevis (Dinosauria: Sauropodomorpha) from the Late Triassic of Argentina and its phylogenetic implications

Abstract: Basal sauropodomorphs from the Upper Triassic Los Colorados Formation of north‐western Argentina have been known for several decades but most of them are only briefly described. New postrcanial remains of Coloradisaurus brevis, the most gracile sauropodomorph from this unit, are described here and evaluated within a phylogenetic context. These materials belong to a single individual and include elements of the vertebral column, pectoral girdle, incomplete forelimb, pelvis and hindlimb. These elements share an autapomorphic feature with the type specimen of Coloradisaurus brevis and provide novel and unique features that diagnose this taxon, such as pubic apron with semicircular cross‐section and with a depression on its anterior surface, distal surface of the tibia deflected and facing posterodistally and well‐developed pyramidal dorsal process of the posteromedial corner of the astragalus. Several postcranial characters of Coloradisaurus are exclusively shared with Lufengosaurus, from the Lower Jurassic of China. The inclusion of this information in two recent phylogenetic data sets depicts Coloradisaurus as closely related to Lufengosaurus and well nested within Plateosauria. Both data sets used indicate strong character support for the inclusion of Coloradisaurus and Lufengosaurus within Massospondylidae.     

First Jurassic dinosaur tracksite from Guizhou Province,China: morphology,trackmaker and paleoecology

Abstract

Sauropod tracks from the Early Jurassic Maanshan Member of the Ziliujing Formation in the Dazhuanwan area of Guizhou represent the first Jurassic dinosaur track record for this province. The best preserved and longest trackway is narrow gauge (Brontopodus type), and indicates a relatively small trackmaker (footprint length ~35.0 cm). In conjunction with sauropodomorph skeletal remains these tracks suggest that basal sauropodomorphs and primitive sauropods coexisted in this region during the Early Jurassic. This pattern is same in the Lufeng and Sichuan basins. Thus, sauropod tracks from southwest China are diverse in the Early Jurassic, and include narrow gauge Parabrontopodus, wide gauge Brontopodus-type, and basal sauropodomorph tracks.     

A new crocodylian from the late Maastrichtian of Spain: implications for the initial radiation of crocodyloids

Background

The earliest crocodylians are known primarily from the Late Cretaceous of North America and Europe. The representatives of Gavialoidea and Alligatoroidea are known in the Late Cretaceous of both continents, yet the biogeographic origins of Crocodyloidea are poorly understood. Up to now, only one representative of this clade has been known from the Late Cretaceous, the basal crocodyloid Prodiplocynodon from the Maastrichtian of North America.

Methodology/Principal Findings

The fossil studied is a skull collected from sandstones in the lower part of the Tremp Formation, in Chron C30n, dated at −67.6 to 65.5 Ma (late Maastrichtian), in Arén (Huesca, Spain). It is located in a continuous section that contains the K/P boundary, in which the dinosaur faunas closest to the K/P boundary in Europe have been described, including Arenysaurus ardevoli and Blasisaurus canudoi. Phylogenetic analysis places the new taxon, Arenysuchus gascabadiolorum, at the base of Crocodyloidea.

Conclusions/Significance

The new taxon is the oldest crocodyloid representative in Eurasia. Crocodyloidea had previously only been known from the Palaeogene onwards in this part of Laurasia. Phylogenetically, Arenysuchus gascabadiolorum is situated at the base of the first radiation of crocodyloids that occurred in the late Maastrichtian, shedding light on this part of the cladogram. The presence of basal crocodyloids at the end of the Cretaceous both in North America and Europe provides new evidence of the faunal exchange via the Thulean Land Bridge during the Maastrichtian.     

Osteohistological insight into the early stages of growth in Mussaurus patagonicus (Dinosauria,Sauropodomorpha)

Here, we describe the bone histology of juvenile specimens of the basal sauropodomorph Mussaurus patagonicus and interpret its significance in terms of the early growth dynamics of this taxon. Thin sections from three juvenile specimens (femur length, 111–120 mm) of Mussaurus were analysed. The sampled bones consist of multiple postcranial elements collected from the Late Triassic Laguna Colorada Formation (El Tranquilo Group, Patagonia). The cortical bone is composed of fibrolamellar bone tissue. Vascularisation is commonly laminar or plexiform in the long bones. Growth marks are absent in all the examined samples. The ‘epiphyses’ of long bones are all formed by well-developed hypertrophied calcified cartilage. The predominance of woven-fibred bone matrix in cortical bones indicates a fast growth rate in the individuals examined. Moreover, given the existence of growth marks in adult specimens of Mussaurus, as in other sauropodomorphs, and assuming that the first lines of arrested growth was formed during the first year of life, the absence of growth marks in all the bones suggest that the specimens died before reaching their first year of life. Compared with the African taxon Massospondylus carinatus (another basal sauropodomorph for which the bone histology has been previously studied), it appears that Mussaurus had a higher early growth rate than Massospondylus.     

A subadult specimen of Rubeosaurus ovatus (Dinosauria: Ceratopsidae), with observations on other ceratopsids from the two medicine formation

Background

Centrosaurine ceratopsids are well known from the middle Campanian Upper Two Medicine Formation of Montana. Four taxa have been named: Brachyceratops montanensis, Rubeosaurus ovatus, Einiosaurus procurvicornis, and Achelousaurus horneri. Rubeosaurus has been historically the most enigmatic of these taxa; only two specimens, the holotype caudal parietal bar and a referred incomplete skull, have been assigned to Rubeosaurus.

Methodology/Principal Findings

A revised interpretation of the parietal processes of USNM 14765, the partial skeleton of a subadult centrosaurine formerly referred to Brachyceratops, indicates that it shares a P5 spike with the holotype of Rubeosaurus ovatus and should therefore be referred to that taxon. Brachyceratops is considered a nomen dubium.

Conclusions/Significance

USNM 14765 provides additional anatomical information for Rubeosaurus ovatus. These new data are incorporated into a recent phylogenetic analysis of centrosaurine relationships; Rubeosaurus appears as the sister taxon of a clade composed of Einiosaurus, Achelousaurus, and Pachyrhinosaurus.     

New information on the cranial anatomy of Acrocanthosaurus atokensis and its implications for the phylogeny of Allosauroidea (Dinosauria: Theropoda)

Background

Allosauroidea has a contentious taxonomic and systematic history. Within this group of theropod dinosaurs, considerable debate has surrounded the phylogenetic position of the large-bodied allosauroid Acrocanthosaurus atokensis from the Lower Cretaceous Antlers Formation of North America. Several prior analyses recover Acrocanthosaurus atokensis as sister taxon to the smaller-bodied Allosaurus fragilis known from North America and Europe, and others nest Acrocanthosaurus atokensis within Carcharodontosauridae, a large-bodied group of allosauroids that attained a cosmopolitan distribution during the Early Cretaceous.

Methodology/Principal Findings

Re-evaluation of a well-preserved skull of Acrocanthosaurus atokensis (NCSM 14345) provides new information regarding the palatal complex and inner surfaces of the skull and mandible. Previously inaccessible internal views and articular surfaces of nearly every element of the skull are described. Twenty-four new morphological characters are identified as variable in Allosauroidea, combined with 153 previously published characters, and evaluated for eighteen terminal taxa. Systematic analysis of this dataset recovers a single most parsimonious topology placing Acrocanthosaurus atokensis as a member of Allosauroidea, in agreement with several recent analyses that nest the taxon well within Carcharodontosauridae.

Conclusions/Significance

A revised diagnosis of Acrocanthosaurus atokensis finds that the species is distinguished by four primary characters, including: presence of a knob on the lateral surangular shelf; enlarged posterior surangular foramen; supraoccipital protruding as a double-boss posterior to the nuchal crest; and pneumatic recess within the medial surface of the quadrate. Furthermore, the recovered phylogeny more closely agrees with the stratigraphic record than hypotheses that place Acrocanthosaurus atokensis as more closely related to Allosaurus fragilis. Fitch optimization of body size is also more consistent with the placement of Acrocanthosaurus atokensis within a clade of larger carcharodontosaurid taxa than with smaller-bodied taxa near the base of Allosauroidea. This placement of Acrocanthosaurus atokensis supports previous hypotheses of a global carcharodontosaurid radiation during the Early Cretaceous.     

A new sphenodontian (Lepidosauria: Rhynchocephalia) from the Late Triassic of Argentina and the early origin of the herbivore opisthodontians

Sphenodontians were a successful group of rhynchocephalian reptiles that dominated the fossil record of Lepidosauria during the Triassic and Jurassic. Although evidence of extinction is seen at the end of the Laurasian Early Cretaceous, they appeared to remain numerically abundant in South America until the end of the period. Most of the known Late Cretaceous record in South America is composed of opisthodontians, the herbivorous branch of Sphenodontia, whose oldest members were until recently reported to be from the Kimmeridgian–Tithonian (Late Jurassic). Here, we report a new sphenodontian, Sphenotitan leyesi gen. et sp. nov., collected from the Upper Triassic Quebrada del Barro Formation of northwestern Argentina. Phylogenetic analysis identifies Sphenotitan as a basal member of Opisthodontia, extending the known record of opisthodontians and the origin of herbivory in this group by 50 Myr.     

High diversity of rabies viruses associated with insectivorous bats in Argentina: presence of several independent enzootics

Background

Rabies is a fatal infection of the central nervous system primarily transmitted by rabid animal bites. Rabies virus (RABV) circulates through two different epidemiological cycles: terrestrial and aerial, where dogs, foxes or skunks and bats, respectively, act as the most relevant reservoirs and/or vectors. It is widely accepted that insectivorous bats are not important vectors of RABV in Argentina despite the great diversity of bat species and the extensive Argentinean territory.

Methods

We studied the positivity rate of RABV detection in different areas of the country, and the antigenic and genetic diversity of 99 rabies virus (RABV) strains obtained from 14 species of insectivorous bats collected in Argentina between 1991 and 2008.

Results

Based on the analysis of bats received for RABV analysis by the National Rabies system of surveillance, the positivity rate of RABV in insectivorous bats ranged from 3.1 to 5.4%, depending on the geographic location. The findings were distributed among an extensive area of the Argentinean territory. The 99 strains of insectivorous bat-related sequences were divided into six distinct lineages associated with Tadarida brasiliensis, Myotis spp, Eptesicus spp, Histiotus montanus, Lasiurus blosseviilli and Lasiurus cinereus. Comparison with RABV sequences obtained from insectivorous bats of the Americas revealed co-circulation of similar genetic variants in several countries. Finally, inter-species transmission, mostly related with Lasiurus species, was demonstrated in 11.8% of the samples.

Conclusions

This study demonstrates the presence of several independent enzootics of rabies in insectivorous bats of Argentina. This information is relevant to identify potential areas at risk for human and animal infection.     

Floral features,pollination biology and breeding system of Chloraea membranacea Lindl. (Orchidaceae: Chloraeinae)

Background and Aims

The pollination biology of very few Chloraeinae orchids has been studied to date, and most of these studies have focused on breeding systems and fruiting success. Chloraea membranacea Lindl. is one of the few non-Andean species in this group, and the aim of the present contribution is to elucidate the pollination biology, functional floral morphology and breeding system in native populations of this species from Argentina (Buenos Aires) and Brazil (Rio Grande do Sul State).

Methods

Floral features were examined using light microscopy, and scanning and transmission electron microscopy. The breeding system was studied by means of controlled pollinations applied to plants, either bagged in the field or cultivated in a glasshouse. Pollination observations were made on natural populations, and pollinator behaviour was recorded by means of photography and video.

Key Results

Both Argentinean and Brazilian plants were very consistent regarding all studied features. Flowers are nectarless but scented and anatomical analysis indicates that the dark, clavate projections on the adaxial labellar surface are osmophores (scent-producing glands). The plants are self-compatible but pollinator-dependent. The fruit-set obtained through cross-pollination and manual self-pollination was almost identical. The main pollinators are male and female Halictidae bees that withdraw the pollinarium when leaving the flower. Remarkably, the bees tend to visit more than one flower per inflorescence, thus promoting self-pollination (geitonogamy). Fruiting success in Brazilian plants reached 60·78 % in 2010 and 46 % in 2011. Some pollinarium-laden female bees were observed transferring pollen from the carried pollinarium to their hind legs. The use of pollen by pollinators is a rare record for Orchidaceae in general.

Conclusions

Chloraea membrancea is pollinated by deceit. Together, self-compatibility, pollinarium texture, pollinator abundance and behaviour may account for the observed high fruiting success. It is suggested that a reappraisal and re-analysis of important flower features in Chloraeinae orchids is necessary.     

Evolution of High Tooth Replacement Rates in Sauropod Dinosaurs

Background

Tooth replacement rate can be calculated in extinct animals by counting incremental lines of deposition in tooth dentin. Calculating this rate in several taxa allows for the study of the evolution of tooth replacement rate. Sauropod dinosaurs, the largest terrestrial animals that ever evolved, exhibited a diversity of tooth sizes and shapes, but little is known about their tooth replacement rates.

Methodology/Principal Findings

We present tooth replacement rate, formation time, crown volume, total dentition volume, and enamel thickness for two coexisting but distantly related and morphologically disparate sauropod dinosaurs Camarasaurus and Diplodocus. Individual tooth formation time was determined by counting daily incremental lines in dentin. Tooth replacement rate is calculated as the difference between the number of days recorded in successive replacement teeth. Each tooth family in Camarasaurus has a maximum of three replacement teeth, whereas each Diplodocus tooth family has up to five. Tooth formation times are about 1.7 times longer in Camarasaurus than in Diplodocus (315 vs. 185 days). Average tooth replacement rate in Camarasaurus is about one tooth every 62 days versus about one tooth every 35 days in Diplodocus. Despite slower tooth replacement rates in Camarasaurus, the volumetric rate of Camarasaurus tooth replacement is 10 times faster than in Diplodocus because of its substantially greater tooth volumes. A novel method to estimate replacement rate was developed and applied to several other sauropodomorphs that we were not able to thin section.

Conclusions/Significance

Differences in tooth replacement rate among sauropodomorphs likely reflect disparate feeding strategies and/or food choices, which would have facilitated the coexistence of these gigantic herbivores in one ecosystem. Early neosauropods are characterized by high tooth replacement rates (despite their large tooth size), and derived titanosaurs and diplodocoids independently evolved the highest known tooth replacement rates among archosaurs.     

Heterodichogamy in Kingdonia (Circaeasteraceae, Ranunculales)

Background and Aims

Preliminary field observations in 2001 and 2002 suggested that Kingdonia uniflora (Circaeasteraceae, Ranunculales) exhibits heterodichogamy, an unusual kind of reproductive heteromorphy, hitherto unreported in Ranunculales and known from only one other genus in basal eudicots.

Methods

During several subsequent years flowers were observed in the field. Flowers were fixed in FAA and studied with microtome sections series and with the scanning electron microscope.

Key Results

The flowers proved to be heterodichogamous, with protandrous and protogynous morphs, which have a 1 : 1 ratio. Both morphs equally set fruit. Each year a single flower is formed at the tip of a rhizome or more rarely two flowers. The flowers are already open when they appear at the soil surface, before they are receptive and before pollen is dispersed. In both floral morphs the styles elongate early and the stigmas are positioned above the anthers before anthesis begins. In protogynous flowers the stigmas become receptive in this position; later the styles become reflexed and then the anthers dehisce. In contrast, in protandrous flowers the stamen filaments elongate during early anthesis such that the dehiscing anthers come to lie above the (still unreceptive) stigmas; after dehiscence of all anthers in a flower the styles begin to elongate and become receptive.

Conclusions

This is the first record of heterodichogamy in a representative of Ranunculales, in an herbaceous eudicot, and in a plant with uniflorous ramets. The occurrence of heterodichogamy in Kingdonia in which clonal reproduction appears to be dominant might be an adaptation to avoid mating between the ramets from a common mother individual (genet).Key words: Kingdonia, Circaeasteraceae, Ranunculales, heterodichogamy, reproductive heteromorphy     

Phylogeny of Thaumastodermatidae (Gastrotricha: Macrodasyida) inferred from nuclear and mitochondrial sequence data

Background

Phylogenetic relationships within Gastrotricha are poorly known. Attempts to shed light on this subject using morphological traits have led to hypotheses lacking satisfactory statistical support; it seemed therefore that a different approach was needed.

Methodology/Principal Findings

In this paper we attempt to elucidate the relationships within the taxonomically vast family Thaumastodermatidae (Macrodasyida) using molecular sequence data. The study includes representatives of all the extant genera of the family and for the first time uses a multi-gene approach to infer evolutionary liaisons within Gastrotricha. The final data set comprises sequences of three genes (18S, 28S rDNA and COI mtDNA) from 41 species, including 29 thaumastodermatids, 11 non-thaumastodermatid macrodasyidans and a single chaetonotidan. Molecular data was analyzed as a combined set of 3 genes and as individual genes, using Bayesian and maximum likelihood approaches. Two different outgroups were used: Xenotrichula intermedia (Chaetonotida) and members of the putative basal Dactylopodola (Macrodasyida). Thaumastodermatidae and all other sampled macrodasyidan families were found monophyletic except for Cephalodasyidae. Within Thaumastodermatidae Diplodasyinae and Thaumastodermatinae are monophyletic and so are most genera. Oregodasys turns out to be the most basal group within Thaumastodermatinae in analyses of the concatenated data set as well as in analyses of the nuclear genes. Thaumastoderma appears as the sister taxon to the remaining species. Surprisingly, Tetranchyroderma is non-monophyletic in our analyses as one group of species clusters with Ptychostomella while another appears as the sister group of Pseudostomella.

Conclusions/Significance

Results in general agree with the current classification; however, a revision of the more derived thaumastodermatid taxa seems necessary. We also found that the ostensible COI sequences from several species do not conform to the general invertebrate or any other published mitochondrial genetic code; they may be mitochondrially derived nuclear genes (numts), or one or more modifications of the mitochondrial genetic code within Gastrotricha.     

A new troodontid theropod dinosaur from the lower Cretaceous of Utah

Background

The theropod dinosaur family Troodontidae is known from the Upper Jurassic, Lower Cretaceous, and Upper Cretaceous of Asia and from the Upper Jurassic and Upper Cretaceous of North America. Before now no undisputed troodontids from North America have been reported from the Early Cretaceous.

Methodology/Principal Findings

Herein we describe a theropod maxilla from the Lower Cretaceous Cedar Mountain Formation of Utah and perform a phylogenetic analysis to determine its phylogenetic position. The specimen is distinctive enough to assign to a new genus and species, Geminiraptor suarezarum. Phylogenetic analysis places G. suarezarum within Troodontidae in an unresolved polytomy with Mei, Byronosaurus, Sinornithoides, Sinusonasus, and Troodon + (Saurornithoides + Zanabazar). Geminiraptor suarezarum uniquely exhibits extreme pneumatic inflation of the maxilla internal to the antorbital fossa such that the anterior maxilla has a triangular cross-section. Unlike troodontids more closely related to Troodon, G. suarezarum exhibits bony septa between the dental alveoli and a promaxillary foramen that is visible in lateral view.

Conclusions/Significance

This is the first report of a North American troodontid from the Lower Cretaceous. It therefore contributes to a fuller understanding of troodontid biogeography through time. It also adds to the known dinosaurian fauna of the Cedar Mountain Formation.