Evolution and homology of the astragalus in early amniotes: new fossils, new perspectives

The reorganization of the ankle in basal amniotes has long been considered a key innovation allowing the evolution of more terrestrial and cursorial behavior. Understanding how this key innovation arose is a complex problem that largely concerns the homologizing of the amniote astragalus with the various ossifications in the anamniote tarsus. Over the last century, several hypotheses have been advanced homologizing the amniote astragalus with the many ossifications in the ankle of amphibian-grade tetrapods. There is an emerging consensus that the amniote astragalus is a complex structure emerging via the co-ossification of several originally separate elements, but the identities of these elements remain unclear. Here we present new fossil evidence bearing on this contentious question. A poorly ossified, juvenile astragalus of the large captorhinid Moradisaurus grandis shows clear evidence of four ossification centers, rather than of three centers or one center as posited in previous models of astragalus homology. Comparative material of the captorhinid Captorhinikos chozaensis is also interpretable as demonstrating four ossification centers. A new, four-center model for the homology of the amniote astragalus is advanced, and is discussed in the context of the phylogeny of the Captorhinidae in an attempt to identify the developmental transitions responsible for the observed pattern of ossification within this clade. Lastly, the broader implications for amniote phylogeny are discussed, concluding that the neomorphic pattern of astragalus ossification seen in all extant reptiles (including turtles) arose within the clade Diapsida.     

Limb ossification patterns of the ichthyosaur Stenopterygius, and a discussion of the proximal tarsal row of ichthyosaurs and other neodiapsid reptiles

Limb ossification patterns for the Lower Jurassic (Toarcian) ichthyosaur, Stenopterygius , are described. It is found that limb ossification follows a continuous proximal to distal sequence from the propodial elements through to the terminal elements of 1st to 4th digit in the manus and the 1st to 3rd digit in the pes. The 5th manal and 4th pedal digit begin ossification later than more preaxial digits and also show evidence of proximal addition of elements near the distal mesopodial row in a manner consistent with delayed ossification of the 5th distal mesopodial in other diapsids. Ossification of manal elements in the Supernumerary 3–4 (S3-4) digit and the 5th digit appear interdependent; if one or the other is highly ossified, ossification of the other is retarded. The 1st pedal digit is considered to be lost in Stenopterygius and the 4th pedal digit is identified as the 5th digit. Delayed ossification of the mesopodium is not observed. The most preaxial proximal tarsal is identified as the centralc; the remaining proximal tarsals are the astragalus and calcaneum, and it is inferred that the astragalus and calcaneum ossified from within a single proximal cartilage.     

The tarsus of erythrosuchid archosaurs, and implications for early diapsid phylogeny

The morphology of the erythrosuchid ankle joint is reassessed. Two specimens, recently thought to have been incorrectly referred to Erythrosuchus africanus , are shown without doubt to belong to this taxon. Furthermore, the morphology is essentially similar to that of other early archosaurs. The tarsus of Erythrosuchus is poorly ossified and consists of a calcaneum, astragalus, and two distal tarsals. The calcanea of Erythrosuchus, Vjushkovia triplicostata , and Shansisuchus shansisuchus are all similar in being dorsoventrally compressed, possessing a lateral tuber, and lacking a perforating foramen. The astragalus of V. triplicostata is currently unknown. The astragalus of Shansisuchus is apparently unique in form. The erythrosuchid pes is therefore more derived than has been recently proposed. The tarsal morphology of several other archosauromorph taxa is reviewed and many details are found to be at variance with the literature. The plesiomorphic condition for the Archosauromorpha consists of four distal tarsals and a proximal row of three elements; two of which articulate with the tibia. These proximal elements are interpreted as the astragalus, calcaneum, and a centrale, and the same pattern is retained in the earliest archosaurs. This reassessed tarsal morphology has implications for the homology of the centrale and reconstruction of early diapsid phylogeny.     

A complex hyobranchial apparatus in a Cretaceous dinosaur and the antiquity of avian paraglossalia

The highly specialized feeding apparatus of modern birds is characterized in part by paraglossalia, triangular bones or cartilages in the tongue that constitute part of the rarely fossilized hyobranchial apparatus. Here, we report on a new, juvenile specimen of the ankylosaurid dinosaur Pinacosaurus grangeri Gilmore, 1933 that provides the first evidence of paraglossalia outside of crown group Aves. The specimen is remarkable in preserving a well‐ossified hyobranchial apparatus, including paired paraglossalia, first and second ceratobranchials, epibranchials, and evidence of a median cartilaginous basihyal. Reassessment of Edmontonia, another ankylosaur, also reveals evidence of bony paraglossalia. Ankylosaur paraglossalia closely resemble those of birds, but are relatively larger and bear prominent muscle scars, supporting the hypothesis that ankylosaurs had fleshy, muscular tongues. The other hyobranchial elements, surprisingly, resemble those of terrestrially feeding salamanders. Ankylosaurs had reduced, slowly replacing teeth, as evidenced from dental histology, suggesting that they relied greatly on their tongues and hyobranchia for feeding. Some curved, rod‐like elements of other dinosaur hyobranchia are reinterpreted as second ceratobranchials, rather than first ceratobranchials as commonly construed. Ankylosaurs provide rare fossil evidence of deep homology in vertebrate branchial arches and expose severe biases against the preservation and collection of the hyobranchial apparatus. In light of these biases, we hypothesize that paraglossalia were present in the common ancestor of Dinosauria, indicating that some structures of the highly derived avian feeding apparatus were in place by the Triassic Period. © 2015 The Linnean Society of London     

发酵黄芪的乳酸菌的驯化及其与黄芪相互作用研究

目的驯化能发酵黄芪的优良乳酸菌并研究其与黄芪的相互作用,为开发新型饲料添加剂做准备。方法选取能厌氧发酵黄芪药液的优良乳酸菌,通过在添加黄芪药液的发酵培养基中反复传代对其进行驯化,并用蒽酮-硫酸法测定发酵前后粗多糖的含量,用比浊法测定菌体浓度,并用单因素法研究了最佳黄芪添加量和最佳接种量,初步研究驯化菌株与黄芪药液的相互作用。结果经过6次传代驯化后,驯化菌株发酵黄芪药液后可使其粗多糖得率提高111．9％,而黄芪药液能促进驯化菌株的增殖,可使菌体浓度提高251％。结论驯化菌株与黄芪药液发生了良性的相互作用,有可能开发出新型饲料添加剂。     

Assessing dinosaur growth patterns: a microscopic revolution

Some of the longest standing questions in dinosaur paleontology pertain to their development. Did dinosaurs grow at slow rates similar to extant reptiles or rapidly similar to living birds and mammals? How did some forms attain gigantic proportions? Conversely, how did birds (avian dinosaurs) become miniaturized? New data on dinosaur longevity garnered from bone microstructure (i.e. osteohistology) are making it possible to assess basic life-history parameters of the dinosaurs such as growth rates and timing of developmental events. Analyses of these data in an evolutionary context are enabling the identification of developmental patterns that lead to size changes within the Dinosauria. Furthermore, this rich new database is providing inroads for studying individual and population biology. All in all, paleohistological research is proving to be the most promising avenue towards gaining a comprehensive understanding of dinosaur biology.     

Skeletal adaptations and phylogeny of the oldest mole Eotalpa (Talpidae,Lipotyphla, Mammalia) from the UK Eocene: the beginning of fossoriality in moles

The oldest talpid, Eotalpa, was previously known only from isolated cheek teeth from the European late Middle Eocene to earliest Oligocene. Screenwashing of Late Eocene sediments of the Hampshire Basin, UK, has yielded cranial and postcranial elements: maxilla, dentary, ulna, metacarpals, distal tibia, astragalus, calcaneum, metatarsals and phalanges. In addition to M_1–2 myotodonty, typical talpid features are as follows: ulna with long medially curved olecranon and deep abductor fossa and astragalar body with lateral process. However, Eotalpa retains certain soricid‐like primitive states (M¹ preparacrista, P⁴ with prominent mesiolingual protocone lobe, strongly angled astragalar neck and calcaneum with no space for a cuboid medial process) not found in modern talpids. Eotalpa is more derived than the most primitive living talpid Uropsilus in having lost the M^1–2 talon shelf, developed a convex radial facet on the ulna, an incipient proximal olecranon crest, relatively shorter metapodials and depressed manual unguals. Its astragalus with medial trochlear ridge taller than the lateral one and massive medial plantar process is typical of the Lipotyphla. Eotalpa lacks synostosis of tibia and fibula, found in other Talpidae, Soricidae and Erinaceidae, suggesting that synostosis in these groups has been independently acquired. Cladistic analysis places Eotalpa as stem member of the Talpidae and shows that much homoplasy arose during the early evolution of the family. Ground dwelling in Eotalpa is indicated by the following: astragalus with a medially dipping head, curved in a single plane; calcaneum with distal peroneal process and strongly overlapping ectal and sustentacular facets; and matching sized ectal and sustentacular facets on calcaneum and astragalus. These features would have restricted ankle mobility. Ungual and metatarsal shape and ulnar structure suggest a primitive stage in fossorial evolution and argue against a semiaquatic precursor stage in talpid fossoriality. Shrew‐moles may represent a reversal to surface foraging rather than an intermediate stage in fossoriality.     

Allometric shape vector projection: a new method for the identification of allometric shape characters and trajectories applied to the human astragalus (talus)

The surface morphology of the human astragalus (talus) is difficult to represent accurately using landmarks as it is essentially globular in shape. Advances in laser scanning technology allow fast and accurate capture of bone surface morphology. However, methodologies to utilise these new accurate 3D data have not been fully developed. The present study uses canonical sampling of whole surface morphology attained through laser scanning and for the first time applies the technique to analysis of bone morphology. We introduce a new technique for identifying allometric shape characters in whole bone surface morphology. In a sample of adult human astragalus the new technique is successful in identifying and isolating intra-specific allometric shape characters in a bone which typically lacks landmarks and has, consequently, proved difficult to analyse using traditional 3D morphometric methods.     

Astragalar morphology of Afradapis, a large adapiform primate from the earliest late Eocene of Egypt

The ～37 million-year-old Birket Qarun Locality 2 (BQ-2), in the Birket Qarun Formation of Egypt's Fayum Depression, yields evidence for a diverse primate fauna, including the earliest known lorisiforms, parapithecoid anthropoids, and Afradapis longicristatus, a large folivorous adapiform. Phylogenetic analysis has placed Afradapis as a stem strepsirrhine within a clade of caenopithecine adapiforms, contradicting the recently popularized alternative hypothesis aligning adapiforms with haplorhines or anthropoids. We describe an astragalus from BQ-2 (DPC 21445C), attributable to Afradapis on the basis of size and relative abundance. The astragalus is remarkably similar to those of extant lorises, having a low body, no posterior shelf, a broad head and neck. It is like extant strepsirrhines more generally, in having a fibular facet that slopes gently away from the lateral tibial facet, and in having a groove for the tendon of flexor fibularis that is lateral to the tibial facet. Comparisons to a sample of euarchontan astragali show the new fossil to be most similar to those of adapines and lorisids. The astragali of other adapiforms are most similar to those of lemurs, but distinctly different from those of all anthropoids. Our measurements show that in extant strepsirrhines and adapiforms the fibular facet slopes away from the lateral tibial facet at a gradual angle (112-126°), in contrast to the anthropoid fibular facet, which forms a sharper angle (87-101°). Phylogenetic analyses incorporating new information from the astragalus continue to support strepsirrhine affinities for adapiforms under varying models of character evolution.     

The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania

The rise of dinosaurs was a major event in vertebrate history, but the timing of the origin and early diversification of the group remain poorly constrained. Here, we describe Nyasasaurus parringtoni gen. et sp. nov., which is identified as either the earliest known member of, or the sister–taxon to, Dinosauria. Nyasasaurus possesses a unique combination of dinosaur character states and an elevated growth rate similar to that of definitive early dinosaurs. It demonstrates that the initial dinosaur radiation occurred over a longer timescale than previously thought (possibly 15 Myr earlier), and that dinosaurs and their immediate relatives are better understood as part of a larger Middle Triassic archosauriform radiation. The African provenance of Nyasasaurus supports a southern Pangaean origin for Dinosauria.     

The first record of a dinosaur from Bulgaria

Mateus, O., Dyke, G.J., Motchurova-Dekova, N., Kamenov, G.D. & Ivanov, P. 2010: The first record of a dinosaur from Bulgaria. Lethaia, Vol. 43, pp. 88–94.
A portion of a left humerus from the Upper Maastrichtian of Vratsa district (NW Bulgaria) is shown to be from a non-avian theropod dinosaur: this is the first record of a dinosaur from Bulgaria. We describe this bone, suggest that it most likely pertains to an ornithomimosaur, and discuss the fossil record of other similar taxa of Late Cretaceous age that have been reported from Europe. To investigate the taphonomy of this fossil, rare earth element (REE) analysis is combined with strontium (Sr) isotope data to confirm that this Bulgarian dinosaur bone was initially fossilized in a terrestrial environment, then later re-worked into late Maastrichtian marine sediments. □ Bulgaria , Dinosauria , Late Cretaceous , Ornithomimosauria , rare earth elements , Sr isotopes , taphonomy , Theropoda .     

Using creation science to demonstrate evolution 2: morphological continuity within Dinosauria

Creationist literature claims that sufficient gaps in morphological continuity exist to classify dinosaurs into several distinct baramins (‘created kinds’). Here, I apply the baraminological method called taxon correlation to test for morphological continuity within and between dinosaurian taxa. The results show enough morphological continuity within Dinosauria to consider most dinosaurs genetically related, even by this creationist standard. A continuous morphological spectrum unites the basal members of Saurischia, Theropoda, Sauropodomorpha, Ornithischia, Thyreophora, Marginocephalia, and Ornithopoda with Nodosauridae and Pachycephalosauria and with the basal ornithodirans Silesaurus and Marasuchus. Morphological gaps in the known fossil record separate only seven groups from the rest of Dinosauria. Those groups are Therizinosauroidea + Oviraptorosauria + Paraves, Tazoudasaurus + Eusauropoda, Ankylosauridae, Stegosauria, Neoceratopsia, basal Hadrosauriformes and Hadrosauridae. Each of these seven groups exhibits within‐group morphological continuity, indicating common descent for all the group’s members, even according to this creationist standard.     

A palaeoequatorial ornithischian and new constraints on early dinosaur diversification

Current characterizations of early dinosaur evolution are incomplete: existing palaeobiological and phylogenetic scenarios are based on a fossil record dominated by saurischians and the implications of the early ornithischian record are often overlooked. Moreover, the timings of deep phylogenetic divergences within Dinosauria are poorly constrained owing to the absence of a rigorous chronostratigraphical framework for key Late Triassic–Early Jurassic localities. A new dinosaur from the earliest Jurassic of the Venezuelan Andes is the first basal ornithischian recovered from terrestrial deposits directly associated with a precise radioisotopic date and the first-named dinosaur from northern South America. It expands the early palaeogeographical range of Ornithischia to palaeoequatorial regions, an area sometimes thought to be devoid of early dinosaur taxa, and offers insights into early dinosaur growth rates, the evolution of sociality and the rapid tempo of the global dinosaur radiation following the end-Triassic mass extinction, helping to underscore the importance of the ornithischian record in broad-scale discussions of early dinosaur history.     

中药及其水煎液中微量元素含量研究

采用火焰原子吸收光谱法(flame atomic absorption spectrometry,FAAS)测定了黄芪、白术、防风及玉屏风散各次水煎液中铁、铜、锰、铅4种微量元素的含量。结果表明,中药各次水煎液中微量元素的浸出率各不相同,应合理用药,以更好地发挥中药的疗效。     

On the existence of non‐microbiotherian Australidelphian marsupials (Diprotodontia) in the Eocene of Patagonia

A diverse assemblage of extinct mammals of early–middle Eocene age (Ypresian–Lutetian boundary) come from the Patagonian localities of La Barda and Laguna Fría around Paso del Sapo in northwestern Chubut Province (Argentina). Metatherians are well represented, mostly by dental remains of ‘Didelphimorphia’, Paucituberculata, Sparassodonta, Microbiotheria, and Polydolopimorphia. Here we analyse three calcanea and one astragalus referable to the same, indeterminate taxon, from La Barda, showing the fusion of their ectal and sustentacular facets. This facet arrangement characterizes the Australidelphia, a marsupial clade represented by Microbiotheria from South America and Antarctica, plus all Australasian lineages. Other australidelphian features shown by these tarsals include: in the calcanea, a reduced peroneal process and a tripartite cuboid facet; in the astragalus, a round, very small astragalar head in relation to the body; a convex trochlea with a half for the tibia and half, in slightly lower position, for the fibula. Their size and other anatomical features suggest that the new tarsals cannot be referred to the Microbiotheria. Phylogenetic analysis suggests that the La Barda taxon lies within the Australidelphia, and that it is either closely related or belongs to, the Diprotodontia. The existence of advanced australidelphians in Patagonia adds further evidence of the close relationship between Patagonia, Antarctica, and Australia during the Late Cretaceous – early Palaeogene.     

Evolution of the Pedolateral Foot in Ground Sloths: Patterns of Change in the Astragalus

The rotation of the pes or pedolateral stance in the extinct ground sloths so the body weight of the animal is primarily supported by the fifth metatarsal and the calcaneum occurred independently at least three times and is present in the Megatheriidae, Nothrotheriidae, and Mylodontidae. In contrast, the pes in the Megalonychidae more closely resembles the primitive eutherian pattern. The pedolateral rotation of the pes thus represents an excellent example of parallel evolution in a closely related group of mammals. While the rotation of the foot occurs as a functional complex resulting in the modification of many bones in the pes, the astragalus is the one bone that shows the highest degree of departure from the primitive mammalian condition and the most distinctive changes in morphology. The morphological transition from a plantigrade foot as occurs in xenarthran anteaters and is essentially retained in the megalonychid sloths to the highly derived condition seen in the megathere, nothrothere, and mylodont sloths follows a similar pattern in all groups but there is still significant variation in the foot structure between the lineages. Despite these variations there are consistent patterns of change in the astragalus in all groups related to the progressive rotation of the pes and a change from dorso-plantar flexion and extension to a medio-lateral rotation of the pes relative to the tibia.     

Tarsal development in birds: evidence for homology with the theropod condition

Theropod dinosaurs and birds share a specialized ankle joint in which the proximal tarsal series. the astragalus and calcaneum is braced against the tibia by an ascending process. This feature has been used since T. H. Huxley's time (1870) to support the proposal that birds evolved from dinosaurs. However, according to Martin, Stewart & Whetstone (1980), the avian tarsus is not homologous with that of theropods. They argue that while the ascending process in theropods is continuous with the astragalus in Archaeopteryx and all later birds, it is an independent ossification associated primarily with the calcaneum. A preliminary study of tarsal ontogeny in birds (McGowan, 1984), undertaken to resolve this problem, revealed two developmental pathways, one exemplified in ratites and the other in carinates. The ratite condition corresponded to that of theropods, the bony ascending process being part of the astragalus, while in carinates the corresponding process was part of the calcaneum. The present study, based on more extensive material, reveals that, although the carinate process becomes associated with the calcaneum during later development, there is evidence that it originates as a cartilaginous process from the astragalus and is therefore homologous with the ratite condition. As the avian tarsus is homologous with that of theropods, and of Archaeopteryx , it may be used to support a close phylogenetic relationship among them.     

New Data on Dinosaurs of the Crimean Peninsula

Reexamination of the holotype of Riabininohadros weberae from the Upper Cretaceous (upper Maastrichtian) of the Crimean Peninsula (Besh-Kosh) allowed determination of previously unknown elements of the femur, astragalus, and calcaneus. This taxon shows a set of primitive characters observed in iguanodontids and basal ornithischians and is referred to as Styracosterna indet. The second dinosaur specimen from Crimea (Aleshino) is a fragmentary skeleton, including cervical and dorsal vertebrae. It possibly belongs to advanced iguanodontids or primitive hadrosauroids. Thus, in the Maastrichtian of the Crimean Peninsula, at least two dinosaur species coexisted.     

A beaked basal ornithurine bird (Aves, Ornithurae) from the Lower Cretaceous of China

We report here one of the earliest known beaked ornithurine birds from the Lower Cretaceous deposits in Liaoning, northeast China. The new basal ornithurine, Archaeorhynchus spathula gen. et sp. nov., has a rhynchokinetic skull with toothless jaws. It also contains over three dozen preserved gizzard stones, suggesting an herbivorous diet. The distal end of the tibiotarsus is unfused, enabling recognition of the astragalus with a broad ascending process, generally similar to that of Archaeopteryx . The new discovery sheds new light on our understanding of the early radiation and diet diversification of early birds in the Lower Cretaceous.     

New Study on Dental and Skeletal Features of the Cretaceous “Symmetrodontan” Mammal Zhangheotherium

A new partial skeleton of the Cretaceous “symmetrodontan” mammal Zhangheotherium quinquecuspedens from the Yixian Formation of Liaoning, China has shed light on the dental and skeletal features of this taxon. The new fossil is a juvenile individual of late growth stage, preserved with interesting features of the premolar replacement. This fossil also provides new information on the vertebral column, the pelvis, the hindlimb and pes. Zhangheotherium has a typical diphyodont replacement of its premolars that is characterized by an alternating pattern (p1 → p3 → p2). This alternating replacement of premolars is a derived condition shared by Dryolestes, Slaughteria, and some basal eutherians, and differs from the plesiomorphic sequential replacement of anterior postcanines in eutricondontans, in most multituberculates and in stem mammaliaforms. The calcaneus and astragalus in the ankle joint of Zhangheotherium lack superposition. This shows that the trechnotherian clade, of which Zhangheotherium is a basal taxon, has retained the primitive condition of mammaliaforms in which the astragalus is in juxtaposition with the calcaneus. Coupled with recent evidence from the earliest metatherians and eutherians, this suggests that the superposition of astragalus and calcaneus evolved in parallel in metatherians and eutherians.