The skull of the giant predatory pliosaur Rhomaleosaurus cramptoni: implications for plesiosaur phylogenetics

The predatory pliosaurs were among the largest creatures ever to inhabit the oceans, some reaching gigantic proportions greater than 15 m in length. Fossils of this subclade of plesiosaurs are known from sediments all over the world, ranging in age from the Hettangian (approximately 198 Myr) to the Turonian (approximately 92 Myr). However, due to a lack of detailed studies and because only incomplete specimens are usually reported, pliosaur evolution remains poorly understood. In this paper, we describe the three dimensionally preserved skull of the giant Jurassic pliosaur Rhomaleosaurus cramptoni. The first phylogenetic analysis dedicated to in-group relationships of pliosaurs allows us to hypothesise a number of well-supported lineages that correlate with marine biogeography and the palaeoecology of these reptiles. Rhomaleosaurids comprised a short-lived and early diverging lineage within pliosaurs, whose open-water top-predator niche was filled by other pliosaur taxa by the mid-late Jurassic. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Early loss and multiple return of the lower temporal arcade in diapsid reptiles

The temporal arches of diapsid reptiles have received attention for several decades. In particular, it has been observed that the lower temporal bar at the ventral margin of the cheek is frequently reduced due to the absence of a contact between jugal and quadratojugal. The loss of the arcade was formerly considered to be of high systematic value, but is now often interpreted as being autapomorphic for the respective taxon, and the presence of both arcades is generally regarded as a plesiomorphic feature. Here I show, based on a cladistic analysis as well as on further anatomical evidence, that the lower temporal arcade was lost only once in diapsid evolution, and that the presence of the arch in "higher" diapsids is secondary, which is indicated by the different ratio between jugal and quadratojugal as well as by ontogeny. This result also sheds new light on the understanding of the cheek configuration of enigmatic taxa such as ichthyosaurs and turtles.Electronic Supplementary Material  Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.     

An Upper Cretaceous lizard with a lower temporal arcade

The reduced lower temporal arcade of the skull and the movable quadrate are the most distinctive features of squamates. Up to now, no exception has been documented for any fossil or extant squamates. We report here a new fossil lizard that possesses a complete lower temporal arcade and an unmovable quadrate. The anatomical relationships indicate that those two modifications were secondarily obtained in the new lizard. The complete lower temporal bar and the firm contact between the pterygoid and quadrate may have served as a brace to support the quadrate jaw articulation and thus prevent it from twisting anteriorly rather than posteriorly during the bite cycles. This represents an entirely new pattern of jaw muscle functions within the Squamata.     

No evidence for an afrotherian-like delayed dental eruption in South American notoungulates

The fossil South American ungulates are of great interest relative to the new phylogenetic framework elaborated for living placental mammals. In particular, studies on these endemic taxa can allow for testing congruence between southern placental phylogeny and plate tectonics, beyond what has already been suggested in the Atlantogenata hypothesis based on extant afrotherians and xenarthrans. The presence of delayed dental eruption relative to skull growth is one feature characterizing the extant afrotherians and possibly the xenarthrans. Late dental eruption has been mentioned previously in South American notoungulates, thus suggesting possible resemblance with afrotherians and perhaps xenarthrans. We provide here a detailed study of the dental eruption pattern relative to the skull growth in the notoungulates. In contrast to previous statements, our results demonstrate that there is currently no evidence for an afrotherian-like delayed dental eruption in this group. For now, the inferred absence of a delayed dental eruption in notoungulates does not support atlantogenatan/afrotherian affinities for the Notoungulata, but other atlantogenatan/afrotherian characteristics remain to be explored in more detail in this group and other South American ungulates.     

Juvenile morphology: A clue to the origins of the most mysterious of mysticetes?

The origin of the pygmy right whale (Caperea marginata) has long been one of the most vexing conundrums of marine mammal evolution. The extremely disparate skeletal structure of Caperea and a patchy fossil record have left morphology and molecules at odds: whereas most morphological analyses ally Caperea with right whales (Balaenidae), most molecular studies instead suggest a close relationship with rorquals (Balaenopteridae) and grey whales (Eschrichtiidae). The morphological evidence supporting a Caperea-balaenid clade consists of several shared features of the skull and mandible, as traditionally observed in adult individuals. Here, we show that at least two of these features, the ascending process of the maxilla and the coronoid process, arise from substantially different precursors early during ontogeny and therefore likely do not represent genuine synapomorphies. Both of these juvenile morphologies have adult counterparts in the fossil record, thus indicating that the ontogenetic variation in the living species may be a genuine reflection of differing ancestral states. This new evidence contradicts previous morphological hypotheses on the origins of Caperea and may help to reconcile morphological and molecular evidence.     

Frequency of decompression illness among recent and extinct mammals and “reptiles”: a review

The frequency of decompression illness was high among the extinct marine “reptiles” and very low among the marine mammals. Signs of decompression illness are still found among turtles but whales and seals are unaffected. In humans, the risk of decompression illness is five times increased in individuals with Patent Foramen Ovale; this condition allows blood shunting from the venous circuit to the systemic circuit. This right-left shunt is characteristic of the “reptile” heart, and it is suggested that this could contribute to the high frequency of decompression illness in the extinct reptiles.     

Unexpected skeletal histology of an ichthyosaur from the Middle Jurassic of Patagonia: implications for evolution of bone microstructure among secondary aquatic tetrapods

During the Mesozoic, one of the most significant evolutionary processes was the secondary adaptation of tetrapods to life in water. Several non-related lineages invaded from the terrestrial realms and from the oceans of the entire world. Among these lineages, ichthyosaurs were particularly successful. Advance parvipelvian ichthyosaurs were the first tetrapods to evolve a fish-shaped body profile. The deep skeletal modifications of their bodies, as well as their biology, depict advance ichthyosaurs as the paradigm of secondary adaptation of reptiles to marine life. Functional inferences point to them as off-shore cruising forms, similar to a living tuna, and some of them were capable of deep diving. Bone histology of some genera such as Temnodontosaurus, Stenopterygius, Ichthyosaurus, and Caypullisaurus, characterized by overall cancellous bone, is consistent with the idea of a fish-shaped ichthyosaurs as fast and far cruisers. Here, we provide histological examination of the ribs of the Middle Jurassic parvipelvian Mollesaurus. Contrasting with the bone histology of other parvipelvian, Mollesaurus ribs are characterized by a compact and thick cortex. Our data indicate that the rib cage was heavy and suggest that not all advanced ichthyosaurs were fast cruisers. The compact and dense ribs in these parvipelvian show that advance ichthyosaurs were ecologically more diverse than previously thought and that the lightening of the skeleton reversed, as also occurred in the evolution of cetacean, at least once along the evolutionary history of ichthyosaurs.     

A new bolosaurid parareptile, Belebey chengi sp. nov., from the Middle Permian of China and its paleogeographic significance

A new bolosaurid parareptile, Belebey chengi sp. nov., is described from Dashankou, Gansu Province, China, a Middle Permian locality which is known mostly for its therapsid fauna. The material consists of well-preserved mandibular and anterior skull remains and currently represents the largest and latest surviving member of Bolosauridae. Phylogenetic analysis of bolosaurid interrelationships, the first analysis of any clade of Early Permian parareptiles, indicates that the new taxon groups consistently with the other (Russian) members of the genus Belebey, and forms the sister clade to the genus Bolosaurus from North America. The Early Permian Eudibamus cursoris from Germany falls into the basal most position within Bolosauridae. Our analysis also shows that the split between the main bolosaurid lineages must have occurred near or before the Permo-Carboniferous boundary and that the paleo-equatorial region of Laurasia probably served as the center of origination for these parareptiles. A similar pattern can be found in other clades of Paleozoic amniotes, suggesting that this may be the general trend in early amniote evolution.     

A new fossil thryonomyid from the Late Miocene of the United Arab Emirates and the origin of African cane rats

Cane rats (Thryonomyidae) are represented today by two species inhabiting sub-Saharan Africa. Their fossil record is predominately African, but includes several Miocene species from Arabia and continental Asia that represent dispersal events from Africa. For example, Paraulacodus indicus, known from the Miocene of Pakistan, is closely related to living Thryonomys. Here we describe a new thryonomyid, Protohummus dango, gen. et sp. nov., from the late Miocene Baynunah Formation of the United Arab Emirates. The new thryonomyid is less derived than “Thryonomys” asakomae from the latest Miocene of Ethiopia and clarifies the origin of crown Thryonomys and the evolutionary transition from Paraulacodus. A phylogenetic analysis shows Protohummus dango to be morphologically intermediate between Paraulacodus spp. and extinct and living Thryonomys spp. The morphological grade and phylogenetic position of Protohummus dango further supports previous biochronological estimates of the age of the Baynunah Formation (ca. 6–8 Ma).     

Artiodactyls from the Pondaung Formation (Myanmar): new data and reevaluation of the South Asian Faunal Province during the Middle Eocene

Although Asia is thought to have played a critical role in the radiation of artiodactyls, the fossil record of stem selenodonts (“dichobunoids”) remains dramatically poor in tropical Asian regions. In this study, we report a new dichobunid genus and species Cadutherium kyaukmagyii and a new basal ruminant genus and species Irrawadymeryx pondaungi, from the late Middle Eocene Pondaung Formation, Central Myanmar. Although the scarcity of the present material prevents any attempts to investigate the phylogenetic relationships of Cadutherium with contemporaneous forms from other Holarctic landmasses, this new form shed new light on the diversity of these small rabbit-like ungulates during a key period of their evolutionary history. Reexamination of the small-bodied artiodactyls from Pondaung leads us to propose new identifications of certain published specimens and, in turn, to investigate the temporal and geographic distribution of taxa recognized in the Pondaung Formation. Although fragmentary, these potential new taxa reveal an unsuspected diversity of small forms among artiodactyls of Pondaung. This addition to the Eocene record of dichobunoids and early ruminants provides further insight in the diversity of dental patterns among small artiodactyls from the Pondaung Formation and attests to the antiquity of these groups in Southeast Asia.     

Salt glands in the Jurassic metriorhynchid <Emphasis Type="Italic">Geosaurus</Emphasis>: implications for the evolution of osmoregulation in Mesozoic marine crocodyliforms

The presence of salt-excreting glands in extinct marine sauropsids has been long suspected based on skull morphology. Previously, we described for the first time the natural casts of salt-excreting glands in the head of the Jurassic metriorhynchid crocodyliform Geosaurus araucanensis from the Tithonian of the Vaca Muerta Formation in the Neuquén Basin (Argentina). In the present study, salt-excreting glands are identified in three new individuals (adult, a sub-adult and a juvenile) referable to the same species. New material provides significant information on the salt glands form and function and permit integration of evolutionary scenarios proposed on a physiological basis in extant taxa with evidence from the fossil record. G. araucanensis represents an advanced stage of the basic physiological model to marine adaptations in reptiles. G. araucanensis salt glands were hypertrophied. On this basis, it can be hypothesized that these glands had a high excretory capability. This stage implies that G. araucanensis (like extant pelagic reptiles, e.g. cheloniids) could have maintained constant plasma osmolality even when seawater or osmoconforming prey were ingested. A gradual model of marine adaptation in crocodyliforms based on physiology (freshwater to coastal/estuarine to estuarine /marine to pelagic life) is congruent with the phylogeny of crocodyliforms based on skeletal morphology. The fossil record suggests that the stage of marine pelagic adaptation was achieved by the Early Middle Jurassic. Salt gland size in the juvenile suggests that juveniles were, like adults, pelagic.     

Isolation and characterization of melanopsin and pinopsin expression within photoreceptive sites of reptiles

Non-mammalian vertebrates have multiple extraocular photoreceptors, mainly localised in the pineal complex and the brain, to mediate irradiance detection. In this study, we report the full-length cDNA cloning of ruin lizard melanopsin and pinopsin. The high level of identity with opsins in both the transmembrane regions, where the chromophore binding site is located, and the intracellular loops, where the G-proteins interact, suggests that both melanopsin and pinopsin should be able to generate a stable photopigment, capable of triggering a transduction cascade mediated by G-proteins. Phylogenetic analysis showed that both opsins are located on the expected branches of the corresponding sequences of ortholog proteins. Subsequently, using RT-PCR and RPA analysis, we verified the expression of ruin lizard melanopsin and pinopsin in directly photosensitive organs, such as the lateral eye, brain, pineal gland and parietal eye. Melanopsin expression was detected in the lateral eye and all major regions of the brain. However, different from the situation in Xenopus and chicken, melanopsin is not expressed in the ruin lizard pineal. Pinopsin mRNA expression was only detected in the pineal complex. As a result of their phylogenetic position and ecology, reptiles provide the circadian field with some of the most interesting models for understanding the evolution of the vertebrate circadian timing system and its response to light. This characterization of melanopsin and pinopsin expression in the ruin lizard will be important for future studies aimed at understanding the molecular basis of circadian light detection in reptiles.     

Phylogeny of Collembola based on cuticular compounds:inherent usefulness and limitation of a character type

The phylogeny of Collembola, originally discussed from a morphological point of view, has more recently benefited from novel insights brought by molecular analyses. Both morphological and molecular characters produced a well-resolved phylogenetic hypothesis including all orders, most families, and a large number of genera. However, several conflicting points exist between molecular and morphological data, and new characters are clearly needed to resolve these inconsistencies. In this study the usefulness of a new character type not previously used in the phylogenetic study of Collembola was tested: the epicuticular chemical compounds. Our phylogenetic analysis was based on 380 compounds from 26 Collembola species. The results show good resolution for terminal branches but not for internal nodes. This is probably due to the partial involvement of epicuticular lipids in ecological functions such as water conservation and sexual attraction. Thus, this character type is appropriate for reconstructing phylogenetic relationships among recently diversified groups. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The sea-finding behavior of hatchling olive ridley sea turtles, Lepidochelys olivacea, at the beach of San Miguel (Costa Rica)

Newly hatched olive ridley sea turtles (Lepidochelys olivacea) were tested for their directional preferences in a sand-filled circular arena in total darkness. Hatchlings that had crawled about 5 m on the beach, toward the sea preferred the southwesterly direction that would have brought them to the water line, whereas hatchlings that had been denied this experience headed eastward, a direction of unclear origin. These data suggest that a short crawl across the natural beach can set the direction in which the young turtles subsequently move. The crawling experience was sufficient to acquire the compass course that they later follow, probably with the help of a magnetic compass, not only in the water, but already while still on land.     

Juvenile morphology in baleen whale phylogeny

Phylogenetic reconstructions are sensitive to the influence of ontogeny on morphology. Here, we use foetal/neonatal specimens of known species of living baleen whales (Cetacea: Mysticeti) to show how juvenile morphology of extant species affects phylogenetic placement of the species. In one clade (sei whale, Balaenopteridae), the juvenile is distant from the usual phylogenetic position of adults, but in the other clade (pygmy right whale, Cetotheriidae), the juvenile is close to the adult. Different heterochronic processes at work in the studied species have different influences on juvenile morphology and on phylogenetic placement. This study helps to understand the relationship between evolutionary processes and phylogenetic patterns in baleen whale evolution and, more in general, between phylogeny and ontogeny; likewise, this study provides a proxy how to interpret the phylogeny when fossils that are immature individuals are included. Juvenile individuals in the peramorphic acceleration clades would produce misleading phylogenies, whereas juvenile individuals in the paedomorphic neoteny clades should still provide reliable phylogenetic signals.     

What drove reversions to quadrupedality in ornithischian dinosaurs? Testing hypotheses using centre of mass modelling

The exceptionally rare transition to quadrupedalism from bipedal ancestors occurred on three independent occasions in ornithischian dinosaurs. The possible driving forces behind these transitions remain elusive, but several hypotheses—including the development of dermal armour and the expansion of head size and cranial ornamentation—have been proposed to account for this major shift in stance. We modelled the position of the centre of mass (CoM) in several exemplar ornithischian taxa and demonstrate that the anterior shifts in CoM position associated with the development of an enlarged skull ornamented with horns and frills for display/defence may have been one of the drivers promoting ceratopsian quadrupedality. A posterior shift in CoM position coincident with the development of extensive dermal armour in thyreophorans demonstrates this cannot have been a primary causative mechanism for quadrupedality in this clade. Quadrupedalism developed in response to different selective pressures in each ornithischian lineage, indicating different evolutionary pathways to convergent quadrupedal morphology.     

Euarchontan affinity of Paleocene Afro-European adapisoriculid mammals and their origin in the late Cretaceous Deccan Traps of India

The controversial family Adapisoriculidae, a group of shrew-sized Paleocene mammals, had proposed relationships with insectivores, marsupials and more recently to plesiadapiforms. Adapisoriculid remains are numerous in the early Paleocene locality of Hainin, Belgium, and allow us a test of these different phylogenetic hypotheses. Here, we identify the first tarsal bones of adapisoriculid mammals. The highly specialised bones indicate an arboreal mode of life with euarchontan affinity. Moreover, the tarsal bones are morphologically very close to those of the late Cretaceous Deccanolestes from the Deccan intertrappean beds of India, and also share several characters with the Paleocene plesiadapiforms and the extant cynocephalid dermopterans. The adapisoriculid affinities of Deccanolestes are also confirmed by tooth morphology, indicating that Deccanolestes is a primitive member of this family. These phylogenetic affinities suggest a paleobiogeographic scenario for the family with dispersal either via East Africa or across the Tethys area.     

A gravid lizard from the Cretaceous of China and the early history of squamate viviparity

Although viviparity is most often associated with mammals, roughly one fifth of extant squamate reptiles give birth to live young. Phylogenetic analyses indicate that the trait evolved more than 100 times within Squamata, a frequency greater than that of all other vertebrate clades combined. However, there is debate as to the antiquity of the trait and, until now, the only direct fossil evidence of squamate viviparity was in Late Cretaceous mosasauroids, specialised marine lizards without modern equivalents. Here, we document viviparity in a specimen of a more generalised lizard, Yabeinosaurus, from the Early Cretaceous of China. The gravid female contains more than 15 young at a level of skeletal development corresponding to that of late embryos of living viviparous lizards. This specimen documents the first occurrence of viviparity in a fossil reptile that was largely terrestrial in life, and extends the temporal distribution of the trait in squamates by at least 30 Ma. As Yabeinosaurus occupies a relatively basal position within crown-group squamates, it suggests that the anatomical and physiological preconditions for viviparity arose early within Squamata.     

An enormous Jurassic turtle bone bed from the Turpan Basin of Xinjiang, China

A spectacular new terrestrial Konzentratlagerst?tte is introduced from the Turpan Basin of Xinjiang, China that probably belongs to the late Middle Jurassic Qigu Formation. It contains a mass accumulation of “xinjiangchelyid” turtles preliminarily identified as Annemys sp. In the zone with the highest turtle concentration, complete and articulated turtle skeletons are tightly packed at a density of up to 36 turtles per square meter. The fossiliferous layer is thickened here and shows an erosional base. This high concentration zone outcrops approximately 10?m in length and shows no decrease in turtle density after exposing 2?m of the layer into the hillside. Adjacent is a more expansive zone of at least 10?m by 30?m. In this region, the fossiliferous layer is evenly thick, and approximately five, fully disarticulated turtles are present per square meter. A conservatively estimated 1,800 turtles may, therefore, have been deposited at this site. It is likely that these aquatic turtles gathered in a retreating water hole in a riverine environment during a drought, much as some aquatic turtles will do today, but perished when the habitat dried up completely. A following catastrophic rainfall event caused a debris flow, possibly channelized in a dry river bed, which transported complete turtles, disarticulated turtles, and mudstone clasts and deposited them after a short distance. This taphonomic model is consistent with previous environmental reconstructions of the Turpan Basin during the late Middle Jurassic in predicting the episodic breakdown of regional monsoonal circulation resulting in a seasonally dry climate with severe episodic droughts.