Skull of Grippia Longirostris: no contradiction with a diapsid affinity for the Ichthyopterygia

The skull of Grippia longirostris , one of the basal ichthyopterygians, is redescribed, based on a newly prepared referred specimen and the examination of others. Grippia had been considered to be the most basal ichthyopterygian until recently, and it has been suggested that the quadratojugal enters the upper temporal fenestra in this species. This and other unique features, which were considered plesiomorphic for ichthyopterygians, obscured the relationship of the group to other amniotes. It is shown here that the quadratojugal is much smaller than previously described, and does not enter the upper temporal fenestra. The skull configuration of the species is very similar to the condition in the recently-described skull of Utatsusaurus hataii , another ichthyopterygian belonging to the basal grade of the group. The close similarity between the two help clarify ichthyopterygian plesiomorphies. There is no feature in the skull of Grippia that particularly contradicts the emerging consensus that ichthyopterygians are diapsids.     

A new lizard-like reptile (Diapsida: Lepidosauromorpha) from the Middle Jurassic of England

The Middle Jurassic (Late Bathonian) Mammal Bed at Kirtlington, Oxfordshire, has yielded a rich assemblage of small vertebrates including mammals, frogs, salamanders and small reptiles. This paper describes the skull of a new diapsid reptile, Marmorelta oxoniensis, which was common within the fauna. The skull is gracile with large orbits and relatively long external nares. The frontals and parietals are single in the adult and almost separated on the roof of the skull by the postfrontals. The temporal region shows a mosaic of primitive and derived character states. The quadrate was firmly held to the skull by a large squamosal and a small, free, quadratojugal was retained. However, reduction of the anterior process of the quadratojugal and of the posterior process of the jugal left the lower temporal fenestra broadly open. Comparison with other diapsids leads to the conclusion that Marmoretta was a lepidosauromorph, most probably the sister taxon of Lepidosauria.     

The evolution of the reptilian hindfoot and the homology of the hooked fifth metatarsal

The homology of the fifth metatarsal in turtles and in diapsid reptiles is reassessed in the light of new phylogenetic studies. The two nearest outgroups to turtles — pareiasaurs and procolophonoids — both lack a fifth distal tarsal but retain a normal fifth metatarsal. The fifth distal tarsal was therefore lost at the base of the clade that contains turtles. Thus, the hooked fifth metatarsal in turtles must consist entirely of a modified fifth metatarsal: it does not include contributions from the fifth distal tarsal, as commonly supposed. In diapsids, loss of the fifth distal tarsal appears to have occurred at the base of crown-clade diapsids, hooking of the fifth metatarsal subsequently occurring within lepidosauromorphs, and in archosauromorphs. If so, the hooked fifth metatarsal in archosauromorphs, and some lepidosauromorphs, consists entirely of a modified fifth metatarsal. In both turtles and diapsids, integration of the elements distal to the mesotarsal joint precedes evolution of the hooked fifth metatarsal, supporting the view that the latter element evolved to perform a lever function (analogous to the “heel bone” of mammals).     

Morphometry of transport tissues in a freshwater crustacean

Gills and fenestra dorsalis of the syncarid crustacean Allanaspides helonomus were examined with the electron microscope. Their epithelia were characterized using morphometric measurement, with a view to assessing potential for ion transport and respiratory activity. The results obtained show distinctions between the paired gills and allow the possibility that one gill of each pair is respiratory and the other osmoregulatory. The fenestra dorsalis has been tentatively assigned a transport function, although the nature of its transport load remains uncertain. The detailed morphometric analysis provided useful insights into the nature of the tissues studied.     

MicroRNAs support a turtle + lizard clade

Despite much interest in amniote systematics, the origin of turtles remains elusive. Traditional morphological phylogenetic analyses place turtles outside Diapsida-amniotes whose ancestor had two fenestrae in the temporal region of the skull (among the living forms the tuatara, lizards, birds and crocodilians)-and allied with some unfenestrate-skulled (anapsid) taxa. Nonetheless, some morphological analyses place turtles within Diapsida, allied with Lepidosauria (tuatara and lizards). Most molecular studies agree that turtles are diapsids, but rather than allying them with lepidosaurs, instead place turtles near or within Archosauria (crocodilians and birds). Thus, three basic phylogenetic positions for turtles with respect to extant Diapsida are currently debated: (i) sister to Diapsida, (ii) sister to Lepidosauria, or (iii) sister to, or within, Archosauria. Interestingly, although these three alternatives are consistent with a single unrooted four-taxon tree for extant reptiles, they differ with respect to the position of the root. Here, we apply a novel molecular dataset, the presence versus absence of specific microRNAs, to the problem of the phylogenetic position of turtles and the root of the reptilian tree, and find that this dataset unambiguously supports a turtle + lepidosaur group. We find that turtles and lizards share four unique miRNA gene families that are not found in any other organisms' genome or small RNA library, and no miRNAs are found in all diapsids but not turtles, or in turtles and archosaurs but not in lizards. The concordance between our result and some morphological analyses suggests that there have been numerous morphological convergences and reversals in reptile phylogeny, including the loss of temporal fenestrae.     

DO THE LATERAL FIELDS EXIST IN SOME GALEASPIDS (JAWLESS FISHES)?

<正> Pan et Wang (1981) reported that the dorsal shield of the galeaspid, Qingmenaspis, held a large opening on each side, but owing to the ill preservation they reservedly assumed it likely to be a dorsal branchial opening. This structure was also recognized later in Lungmenshanaspis by Wang Nian-zhong (1991). He considered its function to remain obscure. More recently Pan (1992) described the structure in a number of new galeaspids and renamed it fenestra. He distingushed the structure into two kinds: the lateral dorsal fenestra, which is narrow and small, existing in Pentathyraspis and Microhoplonaspis, and the dorsal fenestra, which is broad, found in Macrothyraspis, Sinoszechuanaspis as well as Qingmenaspis. Pan assumed the fenestra to be coverd by the skin when the creatures were alive, and to play a role in hydrodynamic. He regarded this structure as a unique character for the fenestra galeaspids.     

Unidirectional flow in lizard lungs: a paradigm shift in our understanding of lung evolution in Diapsida

Conventional wisdom has held that unidirectional pulmonary airflow is unique to birds and is an adaption enabling high rates of gas exchange, essential for sustaining flight as well as an endothermic metabolism. Recent visualizations and measurements of flow in the lungs of monitor and iguanid lizards show a bird-like pattern of unidirectional flow in these lineages. These findings call for a paradigm shift in our understanding of lung evolution in diapsids. This pattern of flow is not unique to birds. It is much older than previously believed, and it may be advantageous to the low-energy lifestyles typical of ectothermic animals.     

Morphology of the interorbital region of Saimiri sciureus

The skull of the platyrrhine primate Saimiri sciureus is distinguished by a large interorbital fenestra. Juvenile skulls still show a bony interorbital septum with some small gaps. A morphogenetic study was undertaken to better understand the structures of the interorbital region, which represents a linkage between the base of the braincase and the nasal skeleton. Already in early ontogenetic stages a reduction of the posterior portion of the nasal capsule and of the cartilaginous interorbital septum are observed, resulting in the formation of a primary interorbital fenestra. A bony interorbital septum is mainly formed in perinatal age stages by ossification of the presphenoid and by medial fusion of the frontals; the primary interorbital fenestra is retained as a small opening. It only occurs in late juvenile stages when the definitive interorbital fenestra develops by by secondary transformation of bone into a membrane of dense connective tissue; this process is most probably caused by mechanical friction of the very closely approximated eyes of both sides.     

The phylogeny of early eureptiles: comparing parsimony and Bayesian approaches in the investigation of a basal fossil clade

For the first time the phylogenetic relationships of early eureptiles, consisting of captorhinids, diapsids, and protorothyridids, are investigated in a modern phylogenetic context using both parsimony and Bayesian approaches. Ninety parsimony-informative characters and 25 taxa were included in the analyses. The Bayesian analysis was run with and without a gamma-shape parameter allowing for variable rates across characters. In addition, we ran two more Bayesian analyses that included 42 autapomorphies and thus parsimony-uninformative characters in order to test the effect of variable branch lengths. The different analyses largely converged to the same topology, suggesting that the "protorothyridid" Coelostegus is the sister taxon of all other eureptiles and that the remaining "protorothyridids" are paraphyletic. Also, there is a close relationship between diapsids and Anthracodromeus, Cephalerpeton, and Protorothyris, a grouping of Thuringothyris with captorhinids, and a variable position of the "protorothyridids" Brouffia, Hylonomus, and Paleothyris. The lack of resolution in some parts of the tree might be due to "hard polytomies" and short divergence times between the respective taxa. The tree topology is consistent with the hypothesis that the temporal fenestrations of diapsid reptiles appear to be the consequence of a more lightly built skeleton, indicating a significant ecological shift in the early stages of diapsid evolution. Bayesian analysis is a very useful additional approach in studies of fossil taxa in which more traditional statistical support like the bootstrap is often weak. However, the exclusive use of the Mk model appears suitable only if autapomorphic characters are included, whereas the Mk+gamma model performed well with or without autapomorphies.     

Timing of development of the middle ear of Anura (Amphibia)

Summary The opercularis system and tympanum-stapes complex of the anuran middle ear develop at different times relative to metamorphosis. In early larvae, the fenestra ovalis is represented by a large lateral opening in the otic capsule filled with connective tissue. At later larval stages, but well before metamorphosis, a cartilaginous operculum begins to form at the posterior margin of the fenestra ovalis, and proceeds to expand to fill all except the anterior part of the fenestra. The opercularis muscle forms along with the levator scapulae superior muscle at the anteromedial edge of the developing suprascapular cartilage of the shoulder girdle. The muscle fibers extend anteroventrally towards the operculum and otic capsule, and, just before emergence of the forelimbs, that portion that will form the opercularis muscle inserts on the lateral surface of the operculum. At this stage, when the metamorphosing frogs first show terrestrial habits, the opercularis system is complete and presumably functional. Timing of development of the tympanum-stapes complex is more variable. The stapes begins as a cartilaginous condensation in the anterior part of the fenestra ovalis, and develops laterally to eventually contact the epidermis and dermis that together will form the tympanum. Meanwhile a middle ear cavity and tympanic annulus form to complete the complex. In several species, especially those that metamorphose at a smaller body size, the tympanum-stapes complex is quite incomplete by the end of metamorphosis, and in Hyla crucifer it takes about 60 days to fully develop. The presence of a complete opercularis system by the start of terrestrial activity is consistent with an hypothesized seismic function of the system. The independent timing of development of the opercularis system and tympanum-stapes complex does not support functional hypotheses linking the opercularis system with modulation of responsiveness of the tympanum-stapes complex to aerial sound. Newly metamorphosed frogs with poorly developed tympanum-stapes complexes are presumably either insensitive to aerial sound or employ alternate mechanisms for transmission of sound energy to the inner ear, possibly involving the opercularis system.     

A new Early Permian reptile and its significance in early diapsid evolution

The initial stages of evolution of Diapsida (the large clade that includes not only snakes, lizards, crocodiles and birds, but also dinosaurs and numerous other extinct taxa) is clouded by an exceedingly poor Palaeozoic fossil record. Previous studies had indicated a 38 Myr gap between the first appearance of the oldest diapsid clade (Araeoscelidia), ca 304 million years ago (Ma), and that of its sister group in the Middle Permian (ca 266 Ma). Two new reptile skulls from the Richards Spur locality, Lower Permian of Oklahoma, represent a new diapsid reptile: Orovenator mayorum n. gen. et sp. A phylogenetic analysis identifies O. mayorum as the oldest and most basal member of the araeoscelidian sister group. As Richards Spur has recently been dated to 289 Ma, the new diapsid neatly spans the above gap by appearing 15 Myr after the origin of Diapsida. The presence of O. mayorum at Richards Spur, which records a diverse upland fauna, suggests that initial stages in the evolution of non-araeoscelidian diapsids may have been tied to upland environments. This hypothesis is consonant with the overall scant record for non-araeoscelidian diapsids during the Permian Period, when the well-known terrestrial vertebrate communities are preserved almost exclusively in lowland deltaic, flood plain and lacustrine sedimentary rocks.     

New Evidence for a Discrete Supratemporal Bone in the Jurassic Ichthyosaur Temnodontosaurus

The temporal region in Temnodontosaurus trigonodon and Temnodontosaurus platyodon consists of three discrete ossifications. These bones are identified as the supratemporal, bordering much of the temporal opening, and in mid-cheek, the squamosal, which is in contact with the quadratojugal along its ventral margin. Interpretation of the bone in mid-cheek position as a neomorphic "supernumerary" ossification is rejected. The supratemporal bone of ichthyosaurs is functionally convergent with the squamosal of diapsids. The size and configuration of the supratemporal casts doubt on the supposition that ichthyosaurs were diapsids.     

Quantitative microanatomy of jaw muscle attachment in extant diapsids

Muscular reconstructions in vertebrate paleontology have often relied heavily on the presence of "muscle scars" and similar osteological correlates of muscle attachment, a practice complicated by the fact that approximately half of tendinous muscle attachments to bone in extant vertebrates do not leave readily interpretable scars. Microanatomical and histological correlates of tendinous muscle attachment are much less ambiguous. This study examines the microanatomical correlates of muscle attachment for the mandibular adductors in six species of diapsids. Most prominent tendinous or aponeurotic muscle attachments display a high density of extrinsic fibers (similar to Sharpey's fibers). There is also some indication that the density of extrinsic fibers at an attachment may be directly related to the amount of stress exerted on that attachment. The presence of comparable densities of extrinsic fibers in fossil tissue constitutes strong and readily interpretable positive evidence for the presence of adjacent fibrous connective tissue in life. Microanatomy and histology provide reliable data about muscle attachments that cannot be gleaned from gross observation alone. These additional data, when coupled with existing muscular reconstruction techniques, may be essential to the resolution of ambiguous character states, and will provide more severe tests for long-standing hypotheses of musculature in extinct diapsids. Increasing the accuracy and precision of muscular reconstructions lends greater strength to any phylogenetic, paleobiological, or paleoecological inferences that draw upon these reconstructions as important lines of evidence.     

Osseous inner ear structures and hearing in early marsupials and placentals

Based on the internal anatomy of petrosal bones as shown in radiographs and scanning electron microscopy, the inner ear structures of Late Cretaceous marsupials and placentals (about 65 Myr ago) from the Bug Creek Anthills locality of Montana, USA, are described. The inner ears of Late Cretaceous marsupials and placentals are similar to each other in having the following tribosphenic therian synapomorphies: a fully coiled cochlea, primary and secondary osseous spiral laminae, the perilymphatic recess merging with the scala tympani of the cochlea, an aqueductus cochleae, a true fenestra cochleae, a radial pattern of the cochlear nerve and an elongate basilar membrane extending to the region between the fenestra vestibuli and fenestra cochleae. The inner ear structures of living therians differ from those of their Late Cretaceous relatives mainly in having a greater number of spiral turns of the cochlea and a longer basilar membrane. Functionally, a coiled cochlea not only permits the development of an elongate basilar membrane within a restricted space in the skull but also allows a centralized nerve system to innervate the elongate basilar membrane. Qualitative and quantitative analyses show that, with a typical therian inner ear, Late Cretaceous marsupials and placentals were probably capable of high-frequency hearing.     

似哺乳类爬行动物和哺乳类动物脑颅侧壁构造类型的演替

本文对从原始似哺乳爬行动物进化到高级兽类哺乳动物过程中脑颅侧壁所发生的形态演化进行了分析,根据构造上的不同,将这些动物的脑颅侧壁的构造方式划分为四个构造类型,代表四个进化阶段。     

Brr6 drives the Schizosaccharomyces pombe spindle pole body nuclear envelope insertion/extrusion cycle

The fission yeast interphase spindle pole body (SPB) is a bipartite structure in which a bulky cytoplasmic domain is separated from a nuclear component by the nuclear envelope. During mitosis, the SPB is incorporated into a fenestra that forms within the envelope during mitotic commitment. Closure of this fenestra during anaphase B/mitotic exit returns the cytoplasmic component to the cytoplasmic face of an intact interphase nuclear envelope. Here we show that Brr6 is transiently recruited to SPBs at both SPB insertion and extrusion. Brr6 is required for both SPB insertion and nuclear envelope integrity during anaphase B/mitotic exit. Genetic interactions with apq12 and defective sterol assimilation suggest that Brr6 may alter envelope composition at SPBs to promote SPB insertion and extrusion. The restriction of the Brr6 domain to eukaryotes that use a polar fenestra in an otherwise closed mitosis suggests a conserved role in fenestration to enable a single microtubule organizing center to nucleate both cytoplasmic and nuclear microtubules on opposing sides of the nuclear envelope.     

The development of the chondrocranium of the lacertid lizard,Acanthodactylus boskiana

The trabeculae cranii are at first quite separate from each other, after few days their anterior two fifths are connected by a trabecular plate which is obliterated throughout development. The paired origin of the parachordal plate is not observed. The fused posterior orbital cartilages chondrify in the form of a wide short plate, traversed by the oculomotor and trochlear nerves. The basicranial fenestra and fenestra ovalis are formed by the degeneration of pre-existing cartilage. The cochlear portion is completely fused with the parachordal plate from the very beginning. The elements of the pterygoquadrate are fused together. The quadrate and Meckel's cartilage are in close contact from the very beginning. While the lower part of the interorbital septum is derived from the trabecula communis, its upper part is derived from the anterior orbital cartilages. The lateral parts of the fused posterior orbital cartilages give rise to most of the taeniae and pilae of the orbitotemporal region. There is only one commissure between the auditory capsule and parachordal plate. A cartilaginous connection between the distal portion of the columella auris and ceratohyal persists for some time. The parietotectal and paranasal cartilages are fused together from the very beginning. The processus paroticus originates from the columella auris. In the fully formed stage the notochord is completely embedded in the occipital condyle. The union between the condyle and odontoid process persists. The auditory capsules and occipital arches contribute to the formation of the tectum synoticum plus posterius. The prefacial commissure and facial foramen lie in front of the cochlear portion. The columella auris possesses a processus internus (connected with the quadrate), but the processes a dorsalis has completely disappeared. The orbitotemporal region is quite complete. A medial fenestra is formed in the planum supraseptale. A fenestra is observed in each of the interorbital and nasal septa. The lamina transversalis anterior is fused with the parietotectal cartilage. A complete zona annularis is present. The outer wall of the paranasal cartilage is perforated by a large fenestra lateralis. The parietotectal and paranasal cartilages and the posterior process of the lamina transversalis anterior contribute to the formation of the concha nasalis. There is a contact between the planum antorbitale and nasal septum. The pterygoid process has disappeared. The common characters of the lacertid chondrocranuium are deduced.