Skeletal microstructure of Stenopterygius quadriscissus (Reptilia,Ichthyosauria) from the Posidonienschiefer (Posidonia Shale,Lower Jurassic) of Germany

Ichthyosaurians (Ichthyosauria) are a major clade of secondarily aquatic marine tetrapods that occupied several major predatory niches during the Mesozoic Era. Multiple lines of evidence including isotopic, body shape and swimming modality analyses suggest they exhibited elevated growth and metabolic rates, and body temperatures. However, applications of osteohistological methods to test hypotheses regarding their physiology are few. Previous studies focused on the humeri, vertebrae and ribs from a small number of taxa. Here, we use osteohistological methods to describe the bone microstructure of over 30 cranial and post‐cranial elements from a nearly complete, articulated individual of Stenopterygius quadriscissus from the Posidonienschiefer Formation (Posidonia Shale, Lower Jurassic) of Germany. The specimen shows highly vascularized primary bone and spongious secondary bone in its limbs, suggesting an overall shift to a lighter spongious‐structured skeleton was achieved through multiple developmental mechanisms. The modified perichondral ossification in elements of the limbs distal to the stylopodium informs our understanding of functional morphology, including hydrodynamic forces on the paddles. The ribs show variation in cortical thickness and trabecular organization along their length. Cyclical growth is inferred from changes in vascularization and osteocyte density as well as the presence of annuli in primary fibrolamellar bone. Cranial elements, due to their relative density and better preservation of growth marks, may prove to be of particular importance in future skeletochronological studies of post‐Triassic ichthyosaurians. We infer and corroborate hypotheses of elevated growth rates and metabolic rates in ichthyosaurians, and the potential for thermoregulation similar to extant homeothermic ectotherms.     

Cranial morphology of an Early Cretaceous Monjurosuchid (Reptilia: Diapsida) from Liaoning Province of China and evolution of the choristoderan palate

A new specimen of Philydrosaurus proseilus from the Early Cretaceous Chiufotang (Jiufotang) Formation preserves the first complete palate of a monjurosuchid choristodere. As in other choristoderes, the palate of Philydrosaurus is akinetic, extended by a broad contact between the vomer and maxilla, and equipped with multiple batteries of palatal teeth. This specimen provides phylogenetically significant information and clarifies the distribution of many apomorphies within Choristodera. Philydrosaurus is primitive relative to the Neochoristodera in that it exhibits only a moderate degree of posterior displacement of the choanae and retains a relatively large interpterygoid vacuity. However, Philydrosaurus also exhibits several derived features previously considered diagnostic of the Neochoristodera, including establishment of a long midline contact of the pterygoids and development of a distinct nasopalatal trough extending from the choana. The choristodere palate exhibits significant modification of the primitive diapsid condition, including elongation of the vomers, establishment of a vomer–maxilla contact, posterior displacement of the choanae, development of the nasopalatal trough, and reduction of the interpterygoid vacuity.     

Preservational modes of some ichthyosaur soft tissues (Reptilia,Ichthyopterygia) from the Jurassic Posidonia Shale of Germany

Konservat-Lagerstätten, such as the Toarcian (Early Jurassic) Posidonia Shale of southwestern Germany, are renowned for their spectacular fossils. Ichthyosaur skeletons recovered from this formation are frequently associated with soft tissues; however, the preserved material ranges from three-dimensional, predominantly phosphatized structures to dark films of mainly organic matter. We examined soft-tissue residues obtained from two ichthyosaur specimens using an integrated ultrastructural and geochemical approach. Our analyses revealed that the superficially-looking ‘films’ in fact comprise sections of densely aggregated melanosome (pigment) organelles sandwiched between phosphatized layers containing fibrous microstructures. We interpret this distinct layering as representing condensed and incompletely degraded integument from both sides of the animal. When compared against previously documented ichthyosaur fossils, it becomes readily apparent that a range of preservational modes exists between presumed ‘phosphatic’ and ‘carbonized’ soft-tissue remains. Some specimens show high structural fidelity (e.g. distinct integumentary layering), while others, including the fossils examined in this study, retain few original anatomical details. This diversity of soft-tissue preservational modes among Posidonia Shale ichthyosaurs offers a unique opportunity to examine different biostratinomic, taphonomic and diagenetic variables that potentially could affect the process of fossilization. It is likely that soft-tissue preservation in the Posidonia Shale was regulated by a multitude of factors, including decay efficiency and speed of phosphatic mineral nucleation; these in turn were governed by a seafloor with sustained microbial mat activity fuelled by high organic matter input and seasonally fluctuating oxygen levels.     

The first specimen of the Middle Triassic Phalarodon atavus (Ichthyosauria: Mixosauridae) from South China,showing postcranial anatomy and peri‐Tethyan distribution

Phalarodon atavus from the Germanic Muschelkalk Basin was previously represented only by cranial elements. Here we report a nearly complete and articulated specimen of P. atavus from the Middle Triassic Luoping Biota, Yunnan, South China. This is the first specimen of P. atavus from outside the Germanic Basin. This discovery demonstrates a peri‐Tethyan distribution of P. atavus. The new specimen is also the first one preserving the postcranial anatomy of this species, providing the opportunity to evaluate its sustained swimming ability. Inferences made on its functional morphology suggest that this species was probably adapted for active foraging. Tooth crown morphology suggests that P. atavus may have preferred externally soft prey.     

Trophic niche ontogeny and palaeoecology of early Toarcian Stenopterygius (Reptilia: Ichthyosauria)

Reconstructing ecological niche shifts during ontogeny in extinct animals with no living analogues is difficult without exceptional fossil collections. Here we demonstrate how a previously identified ontogenetic shift in the size and shape of the dentition in the early Toarcian ichthyosaur Stenopterygius quadriscissus accurately predicts a particular dietary shift. The smallest S. quadriscissus fed on small, burst‐swimming fishes, with a steady shift towards faster moving fish and cephalopods with increasing body size. Larger adult specimens appear to have been completely reliant on cephalopods, with fish completely absent from gut contents shortly after onset of sexual maturity. This is consistent with a previously proposed ontogenetic niche shift based on tooth shape and body size, corroborating the idea that dental ontogeny may be a useful predictor of dietary shifts in marine reptiles. Applying the theoretical framework used here to other extinct species will improve the resolution of palaeoecological reconstructions, where appropriate sample sizes exist.     

The functional morphology of lingual prey capture in a scincid lizard,Tiliqua scincoides (Reptilia: Squamata)

We investigated the functional morphology of lingual prey capture in the blue‐tongued skink, Tiliqua scincoides, a lingual‐feeding lizard nested deep within the family Scincidae, which is presumed to be dominated by jaw‐feeding. We used kinematic analysis of high‐speed video to characterize jaw and tongue movements during prey capture. Phylogenetically informed principal components analysis of tongue morphology showed that, compared to jaw‐feeding scincids and lacertids, T. scincoides and another tongue‐feeding scincid, Corucia zebrata, are distinct in ways suggesting an enhanced ability for hydrostatic shape change. Lingual feeding kinematics show substantial quantitative and qualitative variation among T. scincoides individuals. High‐speed video analysis showed that T. scincoides uses significant hydrostatic elongation and deformation during protrusion, tongue‐prey contact, and retraction. A key feature of lingual prey capture in T. scincoides is extensive hydrostatic deformation to increase the area of tongue‐prey contact, presumably to maximize wet adhesion of the prey item. Adhesion is mechanically reinforced during tongue retraction through formation of a distinctive “saddle” in the foretongue that supports the prey item, reducing the risk of prey loss during retraction.     

A new choristodere (Reptilia: Diapsida) from the Lower Cretaceous of western Liaoning Province, China, and phylogenetic relationships of Monjurosuchidae

The publication of the scientific name Monjurosuchus splendens in 1940 documented the first tetrapod fossil of the later world-renowned Jehol Biota. For more than half a century since this discovery, however, Monjurosuchus has remained as a monotypic genus of the family Monjurosuchidae, and the relationships of the family with choristoderes have not been correctly recognized until quite recently. In this paper, a new monjurosuchid is named and described based on a nearly complete skull and postcranial skeleton from the Early Cretaceous Chiufotang Formation exposed near Chaoyang, western Liaoning Province, China. This new material documents the first occurrence of monjurosuchid choristoderes outside the type Lingyuan area, and extends the geological range of the family from the Yixian Formation to the younger Chiufotang Formation. Cladistic analyses were conducted with inclusion of monjurosuchids, and the results support the placement of the family Monjurosuchidae as a primitive clade outside the Neochoristodera. A new classification scheme is proposed for choristoderes on the basis of the recovered phylogenetic framework of the group.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 427–444.     

First procolophonid (Reptilia,Parareptilia) from the Lower Triassic of Madagascar

Lasasaurus beltanae nov. gen. nov. sp., a new procolophonid (Reptilia, Parareptilia) from the Lower Triassic of Madagascar, is represented by a single partial skeleton preserved in a ferro-calcareous nodule from the Middle Sakamena Formation, in the North of the island. This new taxon is unique in possessing peculiar, fine and dendritic crests running along the posterolateral side of the squamosal, widely spaced maxillary teeth, subparallel mesiodistal ridges connecting maxillary teeth to the tooth row, and a strongly acute anterior margin of the copula (hyoid bone). This well-preserved specimen belongs to a juvenile individual. The inclusion of L. beltanae nov. gen. nov. sp. in a phylogenetic analysis suggests that it is close to Theledectinae, Procolophoninae, and Leptopleurinae, though their respective relationships are uncertain. This specimen is the first procolophonid described from Madagascar and represents a minor terrestrial component of a coastal vertebrate assemblage dominated by amphibious to fully-aquatic taxa.     

An unusual lizard (Reptilia: Squamata) from the Early Cretaceous of Las Hoyas, Spain

The Early Cretaceous (Barremian) lepidosaurian assemblage of Las Hoyas (Cuenca Province, Castilla-La Mancha), Spain, resembles that of roughly contemporaneous localities at Montsec (Catalonia), Una (Castilla-La Mancha) and Galve (Aragon) in being dominated by the scincomorph lizard Meyasaums ( Ilerdaesaurus ). It differs in the presence of two previously unknown taxa, one of which, described here, shows a strikingly unusual morphology in its ribs and distal limb skeleton suggestive of climbing specialization. This form is sufficiently distinct from other known taxa to warrant the erection of a new genus and species, Scandensia ciervensis gen. et sp. nov. Cladistic analysis of Scandensia suggests that it forms a sister taxon to living squamates (Iguania + Scleroglossa and all descendants of their most recent common ancestor).     

A new aquatic pythonomorph (Reptilia,Squamata) from the Turonian (Late Cretaceous) of France

Disarticulated vertebrae from the Turonian of France display a distinctive suite of characters and probably represent a new pythonomorph. This taxon displays some degree of vertebral pachyostosis s.s., often observed in varanoid squamates from the Cenomanian-Turonian interval of the ‘Mediterranean’ portion of the Tethys. The discovery of this new material highlights the importance of also describing possibly new taxa based on isolated remains.     

Cranial functional morphology of fossil dogs and adaptation for durophagy in Borophagus and Epicyon (Carnivora,Mammalia)

Morphological specialization is a complex interplay of adaptation and constraint, as similarly specialized features often evolve convergently in unrelated species, indicating that there are universally adaptive aspects to these morphologies. The evolutionary history of carnivores offers outstanding examples of convergent specialization. Among larger predators, borophagine canids were highly abundant during the tertiary of North America and are regarded as the ecological vicars of Afro‐Eurasian hyenas. Borophaginae is an extinct group of 60+ species, the largest forms evolving robust skulls with prominently domed foreheads, short snouts, and hypertrophied fourth premolars. These specializations have been speculated to enhance bone cracking. To test the extent that the skulls of derived borophagines were adapted for producing large bite forces and withstanding the mechanical stresses associated with bone cracking relative to their nonrobust sister clades, we manipulated muscle forces in models of six canid skulls and analyzed their mechanical response using 3D finite element analysis. Performance measures of bite force production efficiency and deformation minimization showed that skulls of derived borophagines Borophagus secundus and Epicyon haydeni are particularly strong in the frontal region; maximum stresses are lower and more evenly distributed over the skull than in other canids. Frontal strength is potentially coupled with a temporalis‐driven bite to minimize cranial stress during biting in the two derived genera, as tensile stress incurred by contracting temporalis muscles is dissipated rostro‐ventrally across the forehead and face. Comparison of estimated masticatory muscle cross section areas suggests that the temporalis‐masseter ratio is not strongly associated with morphological adaptations for bone cracking in Borophagus and Epicyon; larger body size may explain relatively larger temporalis muscles in the latter. When compared with previous studies, the overall cranial mechanics of the derived borophagines is more similar to bone‐cracking hyaenids and percrocutids than to their canid relatives, indicating convergence in both morphological form and functional capability. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Endocranial morphology of psittacosaurs (Dinosauria: Ceratopsia) based on CT scans of new fossils from the Lower Cretaceous, China

Psittacosaurs, small basal ceratopsians with a parrot-like beak, are among the most abundant dinosaurs, but occur only in the Early Cretaceous of East Asia. Although the general morphology of psittacosaurs is fairly well understood, the endocranial anatomy of the group has never been described. New discoveries of well-preserved skulls from the celebrated Liaoning beds in northeastern China provide the material for conducting research on psittacosaur endocranial morphology. Using computed tomography scans of three-dimensionally preserved skulls, this study reveals basic endocranial anatomy of psittacosaurs and provides the first palaeoneurological evidence of psittacosaurs in relation to their behaviour. Although commonly believed to have had a small brain and small eyes, psittacosaurs had relatively high brain/body size ratios that are comparable to those in the large theropod Tyrannosaurus, and probably had a keen sense of smell and acute vision, as evidenced by their enlarged olfactory lobes and bulbous optic lobes. The configuration of the semicircular canals agrees with limb proportions to suggest that psittacosaurs were agile animals, perhaps better able to escape predation by carnivorous dinosaurs on that account. The behavioural adaptations implied by this study may have been crucial for the successful radiation of psittacosaurs during the Early Cretaceous of East Asia.     

A new genus of pliosaur (Reptilia: Sauropterygia) from the Lower Jurassic of Holzmaden,Germany

Abstract: The exquisitely preserved holotype of the pliosaur ‘Rhomaleosaurus’victor (SMNS 12478) is described from the Toarcian Posidonien‐Schiefer (Upper Lias, Lower Jurassic) of Holzmaden (Baden‐Württemberg), Germany. The specimen presents a novel combination of synapomorphies and unique morphometric proportions separating it from Rhomaleosaurus sensu stricto and warranting the erection of a new genus, Meyerasaurus gen. nov. Historically, the name ‘Thaumatosaurus’ has been interchangeable with Rhomaleosaurus and is frequently associated with SMNS 12478 in the literature. However, this is an invalid taxon and cannot be reinstated. The anatomy of Meyerasaurus victor is compared in detail with other pliosaurs, and its taxonomic affinity is reviewed. M. victor belongs to the family Rhomaleosauridae and shares several anatomical characters with Rhomaleosaurus including a short and robust premaxillary rostrum (length‐to‐width ratio c. 1.0), parallel premaxilla–maxilla sutures anterior to the nares, vomers contacting the maxillae posterior to the internal nares, and c. 28 cervical vertebrae minus the atlas–axis. The known geographical distribution of Rhomaleosaurus, which previously extended across the German and English palaeobiogeographical zones, is reduced to the English zone as a consequence of the referral of SMNS 12478 to a new genus. This is significant because it contributes to an ongoing trend of increasing generic separation between the German and English zones, while increasing the generic diversity within the German zone itself.     

Cretaceous origins of the vibrotactile bill-tip organ in birds

Some probe-foraging birds locate their buried prey by detecting mechanical vibrations in the substrate using a specialized tactile bill-tip organ comprising mechanoreceptors embedded in densely clustered pits in the bone at the tip of their beak. This remarkable sensory modality is known as ‘remote touch’, and the associated bill-tip organ is found in probe-foraging taxa belonging to both the palaeognathous (in kiwi) and neognathous (in ibises and shorebirds) clades of modern birds. Intriguingly, a structurally similar bill-tip organ is also present in the beaks of extant, non-probing palaeognathous birds (e.g. emu and ostriches) that do not use remote touch. By comparison with our comprehensive sample representing all orders of extant modern birds (Neornithes), we provide evidence that the lithornithids (the most basal known palaeognathous birds which evolved in the Cretaceous period) had the ability to use remote touch. This finding suggests that the occurrence of the vestigial bony bill-tip organ in all modern non-probing palaeognathous birds represents a plesiomorphic condition. Furthermore, our results show that remote-touch probe foraging evolved very early among the Neornithes and it may even have predated the palaeognathous–neognathous divergence. We postulate that the tactile bony bill-tip organ in Neornithes may have originated from other snout tactile specializations of their non-avian theropod ancestors.     

Morphology and phylogenetic relationships of a new eschrichtiid genus (Cetacea: Mysticeti) from the Early Pliocene of northern Italy

A new eschrichtiid, Eschrichtioides gastaldii gen. nov., comb. nov. , is established based on a specimen previously assigned to Balaenoptera gastaldii Portis, 1885. The holotype is from the Early Pliocene of north-east Italy. It represents a fossil mysticete closely related to the living grey whale, Eschrichtius robustus . Comparative morphology and phylogenetic analysis support the monophyly of Eschrichtiidae and Cetotherium -like mysticetes and a sister group relationship between this clade and Balaenopteridae. Eschrichtiid fossils previously described are all from the Pleistocene and Late Pliocene while Eschrichtioides gastaldii is from the Early Pliocene. The recognition of this new eschrichtiid genus suggests that the Mediterranean trophic web of the Early Pliocene was more complex than at present and that the Neogene mysticete family-level biodiversity of the Mediterranean was higher than that currently observed in this basin.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 161–186.     

New archaeocynipids (Hymenoptera,Cynipoidea, Archaeocynipidae) from the Lower Cretaceous of Transbaikalia

A new genus and three new species of parasitic hymenopterans of the family Archaeocynipidae are described from the Lower Cretaceous of Transbaikalia: Khasips alisectus gen. et sp. nov., K. sculptus sp. nov., and K. kovalevi sp. nov. The placement of archaeocynipids in the system of Cynipoidea is discussed.     

Cranial anatomy,taxonomic implications and palaeopathology of an Upper Jurassic Pliosaur (Reptilia: Sauropterygia) from Westbury,Wiltshire, UK

Abstract: Complete skulls of giant marine reptiles of the Late Jurassic are rare, and so the discovery of the 1.8‐m‐long skull of a pliosaur from the Kimmeridge Clay Formation (Kimmeridgian) of Westbury, Wiltshire, UK, is an important find. The specimen shows most of the cranial and mandibular anatomy, as well as a series of pathological conditions. It was previously referred to Pliosaurus brachyspondylus, but it can be referred reliably only to the genus Pliosaurus, because species within the genus are currently in need of review. The new specimen, together with another from the same locality, also referred to P. brachyspondylus, will be crucial in that systematic revision, and it is likely that the genus Pliosaurus will be found to include several genera. The two Westbury Pliosaurus specimens share many features, including the form of the teeth, but marked differences in the snout and parietal crest suggest sexual dimorphism; the present specimen is probably female. The large size of the animal, the extent of sutural fusion and the pathologies suggest this is an ageing individual. An erosive arthrotic condition of the articular glenoids led to prolonged jaw misalignment, generating a suite of associated bone and dental pathologies. Further damage to the jaw joint may have been the cause of death.     

Sensory structures involved in prey detection on the labial palp of the ant‐hunting beetle Siagona europaea Dejean 1826 (Coleoptera,Carabidae)

Giglio, A., Ferrero E.A., Perrotta, E., Talarico, F.F. and Zetto Brandmayr, T. 2010. Sensory structures involved in prey detection on the labial palp of the ant‐hunting beetle Siagona europaea Dejean 1826 (Coleoptera, Carabidae). —Acta Zoologica (Stockholm) 91 : 328–334 The ultrastructure and distribution of sensilla on the labial palps of a myrmecophagous carabid beetle, Siagona europaea, were investigated using scanning and transmission electron microscopy techniques. Five types of sensilla were identified: three types of sensilla basiconica on the apical sensory area and two types, one sensillum trichodeum and one coeloconicum, on the external palp surface. On morphological grounds, the s. basiconica type 1 were considered as olfactory, the type 2 as gustatory, the type 3 and the s. trichodeum as mechanoreceptive, and the s. coeloconicum as a thermo/hygroreceptor. Their function is discussed in relation to prey detection and habitat adaptations.