Hyobranchial and postcranial ontogeny of the temnospondylOnchiodon labyrinthicus (Geinitz, 1861) from Niederhäslich (Dohlen Basin, Autunian, Saxony)

Hyobranchium, postcranial skeleton, and developmental events during ontogeny of the temnospondylOnchiodon labyrinthicus from the Dohlen Basin (Autunian, Saxony) are described. At a skull length of approximately 20 mm, the scapulocoracoid appeared, but ossified very slowly. The ischium ossified distinctly later than the stout ilium and later than the scapulocoracoid. The stapes can be determined in specimens beyond 26 mm skull length, and the exoccipitals started to ossify probably at 40 mm, followed by the quadrate. Vertebral centra ossified in early juveniles at approximately 50 mm skull length. The larval hyobranchium with ossified basibranchial and ceratobranchials indicates strong suction feeding. The bony ceratobranchials were resorbed after the larval period. The poorly ossified postcranium suggests that juveniles lived semiterrestrially.     

Osteology and relationships of the temnospondyl genus Sclerocephalus

The temnospondyl Sclerocephalus from the Permo‐Carboniferous of Germany is one of the most completely preserved and most abundant Palaeozoic tetrapods. Here, we review the complete osteology of the genus, based on a range of fully grown specimens housed in public collections. Among the four valid species, Sclerocephalus haeuseri and Sclerocephalus nobilis reached an adult size of well beyond 1 m in length, and had robust postcranial skeletons. In the skull, the exoccipital and sphenethmoid bones were ossified, completing the well‐known ossification sequence in S. haeuseri. Large adults had an elongate trunk and a laterally compressed tail, and some individuals also retained lateral line sulci: features that taken together suggest an aquatic life. The coracoid, pubis, carpals, tarsals, and the bony tail are fully ossified in the largest specimens. The genus Sclerocephalus forms a weakly supported clade nesting firmly at the base of the Stereospondylomorpha, and the close resemblance between Sclerocephalus, Onchiodon, and Eryops is found to be partially based on shared derived states, but is mostly based on symplesiomorphies. Cladistic analysis of 54 characters and 18 taxa finds more support for the Eryopoidea hypothesis (Eryopidae + Stereospondylomorpha) than for the Euskelia hypothesis (Eryopidae + Zatracheidae + Dissorophoidea). This indicates that the large temnospondyls of the Permian and Mesozoic probably formed a natural group, and that the terrestrial adaptations of Eryops and the dissorophoids probably evolved by convergence.     

Intercentrum versus pleurocentrum growth in early tetrapods: A paleohistological approach

A variety of vertebral centrum morphologies have evolved within early tetrapods which range from multipartite centra consisting of intercentra and pleurocentra in stem‐tetrapods, temnospondyls, seymouriamorphs, and anthracosaurs up to monospondylous centra in lepospondyls. With the present study, we aim to determine the formation of both intercentrum and pleurocentrum and asked whether these can be homologized based on their bone histology. Both intercentra and pleurocentra ossified endochondrally and periosteal bone was subsequently deposited on the outer surface of the centra. Our observations indicate low histological variation between intercentrum and pleurocentrum in microstructural organization and growth which inhibits the determination of homologies. However, intercentrum and pleurocentrum development differs during ontogeny. As previously assumed, the intercentrum arises from ventrally located and initially paired ossification centers that fuse ventromedially to form the typical, crescentic, rhachitomous intercentrum. In contrast, presacral pleurocentra may be ancestrally represented by four ossification centers: a ventral and a dorsal pair. Subsequently, two divergent developmental patterns are observed: In stem‐tetrapods and temnospondyls, the pleurocentrum evolves from the two dorsally located ossification centers which may occasionally fuse to form a dorsal crescent. In some dvinosaurian temnospondyls, the pleurocentrum may even ossify to full rings. In comparison, the pleurocentrum of stem‐amniotes (anthracosaurs, chroniosuchids, seymouriamorphs, and lepospondyls) arises from the two ventrally located ossification centers whereby the ossification pattern is almost identical to that of temnospondyls but mirror‐inverted. Thus, the ring‐shaped pleurocentrum of Discosauriscus ossifies from ventral to dorsal. We also propose that the ossified portions of the intercentrum and pleurocentrum continued as cartilaginous rings or discs that surrounded the notochord in the living animals.     

Skeletal Morphogenesis of Microbrachis and Hyloplesion (Tetrapoda: Lepospondyli), and Implications for the Developmental Patterns of Extinct,Early Tetrapods

The ontogeny of extant amphibians often is used as a model for that of extinct early tetrapods, despite evidence for a spectrum of developmental modes in temnospondyls and a paucity of ontogenetic data for lepospondyls. I describe the skeletal morphogenesis of the extinct lepospondyls Microbrachis pelikani and Hyloplesion longicostatum using the largest samples examined for either taxon. Nearly all known specimens were re-examined, allowing for substantial anatomical revisions that affect the scoring of characters commonly used in phylogenetic analyses of early tetrapods. The palate of H. longicostatum is re-interpreted and suggested to be more similar to that of M. pelikani, especially in the nature of the contact between the pterygoids. Both taxa possess lateral lines, and M. pelikani additionally exhibits branchial plates. However, early and rapid ossification of the postcranial skeleton, including a well-developed pubis and ossified epipodials, suggests that neither taxon metamorphosed nor were they neotenic in the sense of branchiosaurids and salamanders. Morphogenetic patterns in the foot suggest that digit 5 was developmentally delayed and the final digit to ossify in M. pelikani and H. longicostatum. Overall patterns of postcranial ossification may indicate postaxial dominance in limb and digit formation, but also more developmental variation in early tetrapods than has been appreciated. The phylogenetic position and developmental patterns of M. pelikani and H. longicostatum are congruent with the hypothesis that early tetrapods lacked metamorphosis ancestrally and that stem-amniotes exhibited derived features of development, such as rapid and complete ossification of the skeleton, potentially prior to the evolution of the amniotic egg.     

A NEW TEMNOSPONDYL FROM THE PERMO-TRIASSIC BUENA VISTA FORMATION OF URUGUAY

Abstract:  A partial skull recovered from conglomerates of the Permo-Triassic Buena Vista Formation in Uruguay belongs to a new species, Uruyiella liminea gen. et sp. nov. This species is characterized by a broadly triangular skull with laterally projecting posterior corners, rhytidosteid-like dermal sculpturing, and orbits positioned close to the skull margin. Uruyiella liminea is distinguished from other temnospondyls by a combination of primitive and derived character states, such as the anterior extent of the palatine ramus of the pterygoid, which excludes the ectopterygoid and most of the palatine from the lateral border of the interpterygoid vacuity, and the absence of both tabular horns and otic notches. A phylogenetic analysis places Uruyiella and the enigmatic Early Triassic Laidleria in a clade to which we attach the family name Laidleriidae. The Plagiosauridae and the Laidleriidae form a clade at the base of Dvinosauria, which is the sister group of a clade that includes Stereospondyli and Archegosauroidea. This result is unexpected because Laidleria and Plagiosauridae are nested deeply within Stereospondyli in most phylogenies. The sister-group relationship of Uruyiella and Laidleria suggests that a ghost lineage for the latter genus extends down into the earliest Triassic and perhaps even into the Late Permian, which in turn would suggest survivorship of the Laidleriidae through the Permo-Triassic extinction event.     

Early tetrapod relationships revisited

In an attempt to investigate differences between the most widely discussed hypotheses of early tetrapod relationships, we assembled a new data matrix including 90 taxa coded for 319 cranial and postcranial characters. We have incorporated, where possible, original observations of numerous taxa spread throughout the major tetrapod clades. A stem‐based (total‐group) definition of Tetrapoda is preferred over apomorphy‐ and node‐based (crown‐group) definitions. This definition is operational, since it is based on a formal character analysis. A PAUP* search using a recently implemented version of the parsimony ratchet method yields 64 shortest trees. Differences between these trees concern: (1) the internal relationships of aistopods, the three selected species of which form a trichotomy; (2) the internal relationships of embolomeres, with Archeria crassidisca and Pholiderpeton scutigerum collapsed in a trichotomy with a clade formed by Anthracosaurus russelli and Pholiderpeton attheyi; (3) the internal relationships of derived dissorophoids, with four amphibamid species forming an unresolved node with a clade consisting of micromelerpetontids and branchiosaurids and a clade consisting of albanerpetontids plus basal crown‐group lissamphibians; (4) the position of albenerpetontids and Eocaecilia micropoda, which form an unresolved node with a trichotomy subtending Karaurus sharovi, Valdotriton gracilis and Triadobatrachus massinoti;(5) the branching pattern of derived diplocaulid nectrideans, with Batrachiderpeton reticulatum and Diceratosaurus brevirostris collapsed in a trichotomy with a clade formed by Diplocaulus magnicornis and Diploceraspis burkei. The results of the original parsimony run ‐ as well as those retrieved from several other treatments of the data set (e.g. exclusion of postcranial and lower jaw data;character reweighting; reverse weighting) ‐ indicate a deep split of early tetrapods between lissamphibian‐ and amniote‐related taxa. Colosteids, Crassigyrinus, Whatcheeria and baphetids are progressively more crownward stemtetrapods. Caerorhachis, embolomeres, gephyrostegids, Solenodonsaurus and seymouriamorphs are progressively more crownward stem‐amniotes. Eucritta is basal to temnospondyls, with crown‐lissamphibians nested within dissorophoids. Westlothiana is basal to Lepospondyli, but evidence for the monophyletic status of the latter is weak. Westlothiana and Lepospondyli form the sister group to diadectomorphs and crown‐group amniotes. Tuditanomorph and microbrachomorph microsaurs are successively more closely related to a clade including proximodistally: (1) lysorophids; (2) Acherontiscus as sister taxon to adelospondyls; (3) scincosaurids plus diplocaulids; (4) urocordylids plus aïstopods. A data set employing cranial characters only places microsaurs on the amniote stem, but forces remaining lepospondyls to appear as sister group to colosteids on the tetrapod stem in several trees. This arrangement is not significantly worse than the tree topology obtained from the analysis of the complete data set. The pattern of sister group relationships in the crownward part of the temnospondyl‐lissamphibian tree re‐emphasizes the important role of dissorophoids in the lissamphibian origin debate. However, no specific dissorophoid can be identiffed as the immediate sister taxon to crown‐group lissamphibians. The branching sequence of various stem‐group amniotes reveals a coherent set of internested character‐state changes related to the acquisition of progressively more terrestrial habits in several Permo‐Carboniferous forms.     

A new species of Dyrosaurus (Crocodylomorpha, Dyrosauridae) from the early Eocene of Morocco: phylogenetic implications

A new dyrosaurid is described from the Ypresian of the phosphatic deposits of the Oulad Abdoun Basin of Morocco. It is based on numerous cranial and postcranial remains, allowing an almost complete reconstruction. This new Dyrosaurus species, Dyrosaurus maghribensis sp. nov. , is currently only known from Morocco. It differs from D. phosphaticus , present in contemporaneous levels of Algeria and Tunisia, by several autapomorpies, including a smooth dorsal margin of the parietal and widely opened choanae. A phylogenetic analysis, using 47 taxa and 234 morphological characters, shows the dyrosaurids as the sister taxon of pholidosaurids, which include Elosuchus , Sarcosuchus , Terminonaris and Pholidosaurus , and the thalattosuchians. Goniopholididae is a non-monophyletic group; however, if dyrosaurids are not included in the analysis, the result differs and the goniopholidids form a distinct clade. If Thalattosuchia is excluded, both Goniopholididae and Pholidosauridae become paraphyletic assemblages. Thus, phylogenetic problems remain with respect to longirostrine clade, and more attention should be paid to resolving their evolutionary relationships amongst the crocodyliforms.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 603–656.     

A peculiar climbing Megalonychidae from the Pleistocene of Peru and its implication for sloth history

The Xenarthra, particularly the Tardigrada, are with the Notoungulata and Marsupialia among the most diversified South American mammals. Lujanian South American Land Mammal Age localities from the coastal Piedra Escrita site and Andean Casa del Diablo Cave, Peru, have yielded three specimens of the Megalonychidae Diabolotherium nordenskioldi gen. nov. This singular fossil sloth exhibits a peculiar mosaic of cranial and postcranial characters. Some are considered convergent with those of other sloths (e.g. 5/4 quadrangular teeth, characteristic of Megatheriidae), whereas others clearly indicate climbing capabilities distinct from the suspensory mode of extant sloths. The arboreal mode of life of D. nordenskioldi is suggested by considerable mobility of the elbow, hip, and ankle joints, a posteriorly convex ulna with an olecranon shorter than in fossorial taxa, a radial notch that faces more anteriorly than in other fossil sloths and forms an obtuse angle with the coronoid process (which increases the range of pronation–supination), a proximodistally compressed scaphoid, and a wide range of digital flexion. D. nordenskioldi underscores the great adaptability of Tardigrada: an arboreally adapted form is now added to the already known terrestrial, subarboreal, and aquatic (marine and freshwater) fossil sloths. A preliminary phylogenetic analysis of the Tardigrada confirmed the monophyly of Megatherioidea, Nothrotheriidae, Megatheriidae, and Megalonychidae, in which Diabolotherium is strongly nested.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 179–235.     

A NEW SEMIONOTIFORM (ACTINOPTERYGII, NEOPTERYGII) FROM UPPER JURASSIC – LOWER CRETACEOUS DEPOSITS OF NORTH-EAST THAILAND, WITH COMMENTS ON THE RELATIONSHIPS OF SEMIONOTIFORMS

Abstract:  A new semionotiform fish, Isanichthys palustris gen. et sp. nov., is described from the Late Jurassic – Early Cretaceous Phu Kradung Formation, north-east Thailand. I. palustris is known from a single, nearly complete specimen found alongside abundant Lepidotes specimens at the Phu Nam Jun locality. I. palustris shows a mixture of semionotid-like characters, such as the pattern of cheek ossifications, and lepisosteid-like characters, such as the body shape and a dorsal fin opposed by an anal fin. I. palustris possesses only some of the characters currently used to define the Semionotidae. Cladistic analyses including various semionotid and gar taxa, together with Amia calva and Leptolepis coryphaenoides , suggest that the Semionotiformes (Lepisosteidae and 'Semionotidae') form a monophyletic clade, but the 'Semionotidae' taxa form an unresolved polytomy. The relationships between Semionotiformes, Halecomorphi and Teleostei are unresolved. When restricted to the best-known taxa, however, the analysis shows the monophyly of the Semionotidae sensu stricto ( Semionotus + Lepidotes ) and a sister-group relationship between halecomorphs and teleosts. These last two results are regarded as the preferred hypothesis for further studies. I. palustris is the only known example of a predaceous, probably piscivorous, 'semionotid'. It illustrates the great diversity and ecological adaptation of the semionotiforms during the Late Jurassic – Early Cretaceous. We question the phylogenetic relationships of 'ancient fishes' founded on molecular-based trees because we suspect that the use of very few Recent taxa as representatives of previously diverse lineages is an inevitable, but important, bias in the construction of such trees.     

The phylogeny of the 'higher' temnospondyls (Vertebrata: Choanata) and its implications for the monophyly and origins of the Stereospondyli

A parsimony analysis of 'higher' temnospondyls (all temnospondyls descended from the common ancestor of Eryops and Parotosuchus ) was performed using 37 terminal taxa and 121 osteological characters. Bremer support values for each internal node were calculated as a measure of clade strength. Additionally, the shortest trees that conformed to some alternative hypotheses were searched for. The following new taxa are established on the basis of the results: Euskelia (the clade containing the Eryopoidea and Dissorophoidea), Limnarchia (the clade containing Trimerorhachidae, Dvinosauroidea, Archegosauroidea and Stereospondyli), Dvinosauria (the clade containing Trimerorhachidae and Dvinosauroidea), Stereo-spondylomorpha (the clade containing Archegosauroidea and Stereospondyli), Capitosauria (the clade containing Lydekkerina and 'capitosauroids'), and Trematosauria (the clade containing Trematosauroidea, Rhytidosteidae, Plagiosauroidea, Metoposauroidea and Brachyopoidea). The monophyly of the assemblage of Mesozoic families called the Stereospondyli by Romer is supported. The dominance of the Stereospondyli in the Mesozoic and its rarity in the Palaeozoic is discussed. It is suggested that the radiation of the diverse stereospondyl clades, the Capitosauria and Trematosauria, began in the Late Permian of Gondwana, in a 'safe haven' that was less severely affected by the Late Permian extinction event. It is further speculated that the 'safe haven' was located in Antarctica, or possibly Australia.     

Osteologic development of the viscerocranial skeleton in sea bream: alternative ossification strategies in teleost fish

The ontogeny of the viscerocranial skeleton of sea bream Sparus aurata larvae was studied from 1 to 90 days post-hatching. In the smallest specimens analysed at 2·7 mm L _N no cephalic elements were present and at 3·1 mm L _N the following cartilaginous structures were visible: trabecula cranii, auditory capsule, Meckel's cartilage, quadrate, hyosymplectic cartilage, sclerotic, hypohyal, ceratohyal epihyal cartilage, interhyal, hypobranchial 1 and ceratobranchial 1. The only structure ossified at this size is the maxillary and the next ossified structures to appear are the preopercle and opercle at about 3·7 mm L _N. The last bones to appear are infraorbital 2 and 6 at 15·1 mm L _S. The first cartilaginous elements and structures to ossify in S. aurata appear to be related with functional requirements, so that structures involved directly in feeding and breathing generally appear and ossify before those that are not. The ontogeny of different regional structures revealed that generally the dermal bones ossify before the cartilage replacement bones. Comparison of S. aurata viscerocranial skeleton ontogeny with that of phylogenetically distant fish demonstrates that different ossification strategies exist in higher and lower teleost fish.     

白枕鹤的呼吸系统及其生态适应

白枕鹤的呼吸系统由喉头、气管、鸣管、肺及气囊组成。喉头有淋巴小结分布。气管在龙骨突起内盘旋,并随年龄而增长,软骨环逐渐骨化。鸣管由左右两个支气管特化而成,呈膜状扁管入肺。肺的长度约占躯干的1/2。气囊几遍布全身,高度发达。整个呼吸系统的结构,与其高空飞翔生活相适应。     

A NEW PTEROSAUR FROM THE LIAONING PROVINCE OF CHINA, THE PHYLOGENY OF THE PTERODACTYLOIDEA, AND CONVERGENCE IN THEIR CERVICAL VERTEBRAE

Abstract:  The largest known flying organisms are the azhdarchid pterosaurs, a pterodactyloid clade previously diagnosed by the characters of their extremely elongate middle-series cervical vertebrae. The named species of the Azhdarchidae are from the Late Cretaceous. However, isolated mid-cervical vertebrae with similar dimensions and characters have been referred to this group that date back to the Late Jurassic, implying an almost 60 million year gap in the fossil record of this group and an unrecorded radiation in the Jurassic of all the major clades of the Pterodactyloidea. A new pterosaur from the Early Cretaceous of Liaoning Province of China, Elanodactylus prolatus gen. et sp. nov., is described with mid-cervical vertebrae that bear these azhdarchid characters but has other postcranial material that are distinct from the members of this group. Phylogenetic analysis of the new species and the Pterodactyloidea places it with the Late Jurassic vertebrae in the Late Jurassic–Early Cretaceous Ctenochasmatidae and reveals that the characters of the elongate azhdarchid vertebrae appeared independently in both groups. These results are realized though the large taxon sampling in the analysis demonstrating that the homoplastic character states present in these two taxa were acquired in a different order in their respective lineages. Some of these homoplastic characters were previously thought to appear once in the history of pterosaurs and may be correlated to the extension of the neck regions in both groups. Because the homoplastic character states in the Azhdarchidae and Ctenochasmatidae are limited to the mid-cervical vertebrae, these states are termed convergent based on a definition of the term in a phylogenetic context. A number of novel results from the analysis presented produce a reorganization in the different species and taxa of the Pterodactyloidea.     

The Braincase of Eocaecilia micropodia (Lissamphibia,Gymnophiona) and the Origin of Caecilians

The scant fossil record of caecilians has obscured the origin and evolution of this lissamphibian group. Eocaecilia micropodia from the Lower Jurassic of North America remains the only stem-group caecilian with an almost complete skull preserved. However, this taxon has been controversial, engendering re-evaluation of traits considered to be plesiomorphic for extant caecilians. Both the validity of the placement of E. micropodia as a stem caecilian and estimates of the plesiomorphic condition of extant caecilians have been questioned. In order to address these issues, the braincase of E. micropodia was examined via micro-computed tomography. The braincase is considered to be a more reliable phylogenetic indicator than peripheral regions of the skull. These data reveal significant new information, including the possession of an ossified nasal septum, ossified anterior wall of the sphenethmoid, long anterolateral processes on the sphenethmoid, and paired olfactory nerve foramina, which are known only to occur in extant caecilians; the latter are possibly related to the evolution of the tentacle, a caecilian autapomorphy. A phylogenetic analysis that included 64 non-amniote taxa and 308 characters represents the first extensive test of the phylogenetic affinities of E. micropodia. The results place E. micropodia securely on the stem of extant caecilians, representing a clade within Temnospondyli that is the sister taxon to batrachians plus Gerobatrachus. Ancestral character state reconstruction confirms the braincase of E. micropodia to be largely representative of the plesiomorphic condition of extant caecilians. Additionally, the results refine the context within which the evolution of the caecilian form can be evaluated. The robust construction and pattern of the dermal skull of E. micropodia is interpreted as symplesiomorphic with advanced dissorophoid temnospondyls, rather than being autapomorphic in its robust construction. Together these data increase confidence in incorporating E. micropodia into discussions of caecilian evolution.     

New postcranial elements of the Thalassodrominae (Pterodactyloidea,Tapejaridae) from the Romualdo Formation (Aptian–Albian), Santana Group,Araripe Basin,Brazil

Tapejarids are edentate pterosaurs recovered mainly from Early Cretaceous deposits. They are diagnosed by five synapomorphies, among which only one is postcranial: a broad and well‐developed tubercle at the ventroposterior margin of the coracoid. Regarding the clade Thalassodrominae, most phylogenetic studies are based on cranial elements, as postcranial skeletons of these pterosaurs are rare. Here, new postcranial material from the Romualdo Formation (Aptian–Albian) from the Araripe Basin is described. The material comprises the three posteriormost cervical vertebrae, the first seven dorsal vertebrae (fused into a notarium), both scapulocoracoids, a fragment of a sternum, a partial right humerus, a small fragment of a 4th phalanx of the wing finger, a distal extremity of the right femur and the proximal portions of both tibia and fibula. Comparisons with other specimens and morphological features examined in a phylogenetic context, such as the presence of three foramina lateral and dorsal to the neural canal of the cervical vertebrae, the presence of a notarium and a pneumatic foramen on the ventral side of the proximal portion of the humerus, allow the assignment of this specimen as Thalassodrominae indet. Regarding palaeobiogeographical aspects, to date, this clade is exclusively found in the Romualdo Formation. It is the most complete postcranial material assigned to the Thalassodrominae described so far.     

中国广西晚白垩世一新的巨龙类恐龙

记述了采自广西南宁市郊晚白垩世地层中一巨龙类恐龙新属种:右江清秀龙(Qingxiu- saurus youjiangensis gen.et sp.nov.)。新属种正型标本包括以下不关联的头后骨骼:一段较完整的前部尾椎神经棘、左右胸骨板、左右肱骨。它以以下独特特征组合区别于其他巨龙类:前部尾椎神经棘板状结构不发育、相对较高并呈桨状;胸骨与肱骨最大长之比值较低(约0.65)。广西发现的新属种以及近年来报道的巨龙类恐龙材料表明,亚洲巨龙类恐龙的头后骨骼形态变异度高,白垩纪时期这类恐龙曾在亚洲广为分布。