Giant lizards occupied herbivorous mammalian ecospace during the Paleogene greenhouse in Southeast Asia

Mammals dominate modern terrestrial herbivore ecosystems, whereas extant herbivorous reptiles are limited in diversity and body size. The evolution of reptile herbivory and its relationship to mammalian diversification is poorly understood with respect to climate and the roles of predation pressure and competition for food resources. Here, we describe a giant fossil acrodontan lizard recovered with a diverse mammal assemblage from the late middle Eocene Pondaung Formation of Myanmar, which provides a historical test of factors controlling body size in herbivorous squamates. We infer a predominately herbivorous feeding ecology for the new acrodontan based on dental anatomy, phylogenetic relationships and body size. Ranking body masses for Pondaung Formation vertebrates indicates that the lizard occupied a size niche among the larger herbivores and was larger than most carnivorous mammals. Paleotemperature estimates of Pondaung Formation environments based on the body size of the new lizard are approximately 2–5°C higher than modern. These results indicate that competitive exclusion and predation by mammals did not restrict body size evolution in these herbivorous squamates, and elevated temperatures relative to modern climates during the Paleogene greenhouse may have resulted in the evolution of gigantism through elevated poikilothermic metabolic rates and in response to increases in floral productivity.     

The first known chamopsiid lizard (Squamata) from the Upper Cretaceous of Europe (Csehbánya Formation; Hungary,Bakony Mts)

The Upper Cretaceous (Santonian) Iharkút terrestrial vertebrate locality, in Hungary, has yielded several lacertilian taxa since its discovery in 2000. One of these is represented by a left mandible fragment and two dentary fragments. The characters observed on these remains support assignment to the family Chamopsiidae (Scincomorpha). Moreover, the remains belong to a new genus and species, Pelsochamops infrequens n. g. n. sp. Pelsochamops is the first known occurrence of chamopsiids outside North America and adds another scincomorphan lizard to the Iharkút fauna. It represents an additional evidence for dispersal routes between North America and Europe.     

Diversity patterns amongst herbivorous dinosaurs and plants during the Cretaceous: implications for hypotheses of dinosaur/angiosperm co‐evolution

Abstract Palaeobiologists frequently attempt to identify examples of co‐evolutionary interactions over extended geological timescales. These hypotheses are often intuitively appealing, as co‐evolution is so prevalent in extant ecosystems, and are easy to formulate; however, they are much more difficult to test than their modern analogues. Among the more intriguing deep time co‐evolutionary scenarios are those that relate changes in Cretaceous dinosaur faunas to the primary radiation of flowering plants. Demonstration of temporal congruence between the diversifications of co‐evolving groups is necessary to establish whether co‐evolution could have occurred in such cases, but is insufficient to prove whether it actually did take place. Diversity patterns do, however, provide a means for falsifying such hypotheses. We have compiled a new database of Cretaceous dinosaur and plant distributions from information in the primary literature. This is used as the basis for plotting taxonomic diversity and occurrence curves for herbivorous dinosaurs (Sauropodomorpha, Stegosauria, Ankylosauria, Ornithopoda, Ceratopsia, Pachycephalosauria and herbivorous theropods) and major groups of plants (angiosperms, Bennettitales, cycads, cycadophytes, conifers, Filicales and Ginkgoales) that co‐occur in dinosaur‐bearing formations. Pairwise statistical comparisons were made between various floral and faunal groups to test for any significant similarities in the shapes of their diversity curves through time. We show that, with one possible exception, diversity patterns for major groups of herbivorous dinosaurs are not positively correlated with angiosperm diversity. In other words, at the level of major clades, there is no support for any diffuse co‐evolutionary relationship between herbivorous dinosaurs and flowering plants. The diversification of Late Cretaceous pachycephalosaurs (excluding the problematic taxon Stenopelix) shows a positive correlation, but this might be spuriously related to poor sampling in the Turonian–Santonian interval. Stegosauria shows a significant negative correlation with flowering plants and a significant positive correlation with the nonflowering cycadophytes (cycads, Bennettitales). This interesting pattern is worthy of further investigation, and it reflects the decline of both stegosaurs and cycadophytes during the Early Cretaceous.     

Specialization of bone structure in Pachyvaranus crassispondylus Arambourg, 1952, an aquatic squamate from the Late Cretaceous of the southern Tethyan margin

The histological organization of the vertebrae of the Maastrichtian squamate Pachyvaranus crassispondylus Arambourg, 1952 , was compared to that of various extant squamates, in order to further document the causes and functional consequences of the so-called 'pachyostosis', frequently observed in Late Cretaceous squamates. The vertebrae of Pachyvaranus are composed of the same basic bone tissue types as those of extant lizards and snakes. In particular, periosteal cortices are made of a pseudolamellar (or 'parallel-fibred') tissue, with radial vascular canals, Sharpey's fibres and conspicuous cyclic growth marks that are strictly identical to that found in extant varanids. Conversely, the vertebrae of Pachyvaranus are extremely compact, whereas those of extant squamates are very cancellous and lightly built. This difference is due to the absence in Pachyvaranus of a broad internal resorption field that, in extant lizards and snakes, transforms compact cortices into loose spongy formations. This absence of inner bone resorption typically corresponds to an osteosclerotic process. In Pachyvaranus , cortical hyperplasy, or pachyostosis stricto sensu , was restricted to the walls of the neural spine. Extreme vertebral porosity is likely to be a primitive condition in squamates, because all lizards and snakes examined in this study display this feature. Therefore, the high vertebral compactness observed in Pachyvaranus would be a derived condition arising from the loss (or de-differentiation) of a morphogenetic process: the broad internal resorption of the vertebrae. Possible palaeoecological bearings of these results are discussed.     

Cenomanian-Turonian carbon isotope events recorded in terrestrial organic matter from northern Japan

Carbon isotopic fluctuations recorded in terrestrial organic matter from the Upper Cretaceous Cenomanian-Turonian succession of Hokkaido, Japan show a remarkably similar pattern to isotopic fluctuation in carbonate carbon reported from southern England and the Italian Apennines. Chronological correlation for the Cenomanian-Turonian section of Japan is based on planktonic foraminiferal biostratigraphy with some additional data from macrofossils, allowing a detailed comparison of isotope stratigraphies for Japan and Europe. Three distinct features of the isotope profile, namely a positive spike near the Cenomanian/Turonian boundary, a shoulder in the lower-middle Turonian and a level segment in the middle-upper Turonian are observed in Japan. These features are biostratigraphically comparable and correlative with isotope profiles reported from Europe. The single factor influencing isotopic composition of both terrestrial organic carbon and marine carbonate carbon is isotopic change in the global CO₂ reservoir of the ocean-atmosphere system. If these three carbon isotope features are synchronous, they may be inferred as having been caused by global phenomena and as providing global high-resolution chemo-stratigraphic markers. Carbon isotope stratigraphy could be an important bridge between terrestrial inland sequences and marine sequences.     

Sauropod dinosaurs from the Lower Cretaceous of eastern Asia: taxonomic and biogeographical implications

Sauropod dinosaurs are poorly represented in the Lower Cretaceous of eastern Asia. Here, we describe a number of isolated sauropod teeth from the Kuwajima Formation (?Berriasian–?Hauterivian) of Shiramine, Japan. The mosaic of shared derived characters and symplesiomorphies displayed by the teeth indicate that they are referable to a basal member of the titanosauriform radiation. A taxonomic review of previously described sauropod specimens from eastern and south–eastern Asia reveals that a diversity of sauropods (including a titanosaurian, a basal titanosauriform and a ?euhelopodid, as well as several forms of indeterminate systematic position) was present in this region in the Early Cretaceous. This diversity conflicts with previous suggestions that eastern Asia was biogeographically isolated from the rest of Laurasia until the late Early Cretaceous and that the sauropod fauna was limited to the endemic East Asian clade Euhelopodidae. The presence of titanosauriform sauropods in the basal Cretaceous of Japan and Thailand indicate that the proposed faunal isolation of eastern Asia ended approximately 20 myr earlier than usually suggested.     

Chemosensory responses to foods by an herbivorous acrodont lizard, <Emphasis Type="Italic">Uromastyx aegyptius</Emphasis>

Actively foraging lizards use the lingual-vomeronasal system to identify prey by chemical cues, but insectivorous ambush foragers do not. The major clade Iguania includes numerous herbivores and omnivores; among them, two iguanid and one agamine species identify plant and animal foods by tongue flicking, and data suggest that the leiolepidine Uromastyx acanthinurus may as well. We conducted experiments on chemosensory response to food by the herbivorous U. aegyptius. When chemical stimuli were presented on cotton balls in experiment 1, the lizards exhibited greater responsiveness (tongue-flick attack scores) to chemical stimuli from crickets and a preferred plant food (dandelion flowers) than from deionized water. When chemical stimuli were on ceramic tiles in experiment 2, the lizards exhibited greater total tongue flicks to cricket stimuli than to any other stimuli, and to dandelion than to deionized water. Lizards bit more frequently in response to cricket and dandelion cues than to stimuli from a nonpreferred plant (carrot) and deionized water. Tongue-flick attack scores were greater in response to cricket and dandelion stimuli than to carrot or water stimuli. These findings are consistent with the hypothesis that herbivores, even those having ambush-foraging ancestors, use chemical cues to identify potential foods. The data support the hypothesis that chemosensory responses correspond to diet. Because most lizards are generalist predators, studies of herbivorous species can provide important information on possible evolutionary adjustment of chemosensory response to dietary shifts. Electronic Publication     

Responses by a generalist predator,the Balearic lizard <Emphasis Type="Italic">Podarcis lilfordi</Emphasis>, to chemical cues from taxonomically diverse prey

Specialist predators may respond strongly to sensory cues from preferred prey, but responses by generalist predators, although predicted to be less specific, are poorly known. Among squamate reptiles, diet and strength of response to chemical prey cues covary geographically in snakes that are specialist predators. There have been no previous studies of correspondence between diet and chemosensory response in lizards that are prey generalists. Actively foraging lizards discriminate between prey chemicals and control substances. It has been speculated that differential responses among prey species are unlikely in typical species that are dietary generalists. We examined this relationship in Podarcis lilfordi, an omnivorous lacertid that consumes a wide variety of animal prey. In experiments in which chemical stimuli were presented on cotton swabs, lizards responded more strongly to chemicals from a broad spectrum of prey types than to deionized water, an odorless control. These findings plus previous data showing that P. lilfordi is capable of prey chemical discrimination suggest that P. lilfordi can identify a wide range of potential prey using chemical cues. However, there was no evidence of differential response to stimuli among prey species, even in comparisons of prey included in the natural diet and potential prey not in the diet. The results, although limited to a single species, are consistent with the hypothesis that lizard species that are prey generalists do not exhibit the differential response strengths to chemical prey cues observed in snakes that have more specialized diets. Received in revised form: 17 July 2001 Electronic Publication     

Necrosaurus or Palaeovaranus? Appropriate nomenclature and taxonomic content of an enigmatic fossil lizard clade (Squamata)

Necrosaurus cayluxi is an enigmatic lizard from the Paleogene of the Phosphorites of Quercy, France that was first mentioned in the 19th century. Although it is generally believed that Filhol was the author who established this taxon, I am herein demonstrating that authorship should in fact be attributed to Zittel, a fact that also influences not only its generic nomenclature, but also its appropriate type material. As such, the valid name for this taxon should be Palaeovaranus cayluxi and its holotype is a left maxilla. Additionally, Ophisauriscus eucarinatus from the middle Eocene of Geiseltal, Germany, another taxon that was previously assigned to Necrosaurus, is herein shown to be a nomen dubium, whereas Melanosauroides giganteus from the same locality, is considered a valid species and is recombined as Palaeovaranus giganteus comb. nov. The suggested changes in nomenclature also affect “Necrosauridae”, a poorly defined clade of lizards from the Cretaceous–Paleogene of Europe, North America, and Asia. In order to maintain nomenclatural stability and define a monophyletic lineage, I am here establishing the new family Palaeovaranidae fam. nov., which includes solely the genus Palaeovaranus. The known occurrences of Palaeovaranus across the Paleogene of Western Europe are discussed.     

Respective Role of the Axialand Appendicular Systems in Relation to the Transition to Limblessness

In lower quadrupedal vertebrates locomotor efficiency seems to result from the associate movements of the axial and appendicular systems, which are totally independent in structure and embryological origin. The curvature of the trunk, produced by a standing wave, magnifies the propulsive action of the limbs. In intermediate forms, the association of an elongate trunk with limbs reduced in size brings about functional consequences which may be noticeably diverse according to the degree of trunk elongation and limb reduction. According to environmental constraints, animals search for better locomotor efficiency, which implies the maintenance or breakage of this association of both locomotor systems. In some cases, limb action on the ground is added to the axial wave action through a perfect mutual adjustment of rhythmic activity, until mechanical inefficiency of the limbs is reached by possible loss of contact with the ground. In other cases, the limbs dragged on the ground during the stance phase act against the axial action or, on the contrary, are inhibited by the axial system. A review of available data tries to contribute to an understanding of the respective roles of both systems in the transition to limblessness.     

Composition and spatial distribution of microencrusters and microbial crusts in upper Jurassic–lowermost Cretaceous reef limestone (Torinosu Limestone,southwest Japan)

Tethyan microencrusters and microbial crusts, most of them previously unknown in Japanese Mesozoic biotas, are present in the uppermost Jurassic–lowermost Cretaceous Torinosu Limestone distributed in southwestern Japan. They construct reefal facies together with reef-forming metazoans. Bacinella irregularis and Lithocodium aggregatum are quantitatively most important, while subordinate constituents include Thaumatoporella parvovesiculifera, Koskinobullina socialis , Iberopora bodeuri , Girvanella sp. and Tubiphytes morronensis. They are especially common in the shallow-water reefal facies, but appear micritic in outcrops. Microencrusters and microbial crusts can only be recognized in thin sections, and they grow around the reef building metazoans and form bindstone. Each microencruster exhibits some specific spatial distribution associated with its paleoecology. Similarities with the taxonomic composition of the upper Jurassic Tethyan microencruster association imply that the community extended geographically at least to the Tethyan gateway where the Japanese Island Arc was located.     

Pinus MUTOI (Pinaceae), a new species of permineralized seed cone from the upper Cretaceous of Hokkaido,Japan

Pinus mutoi is described as a new species on the basis of a permineralized seed cone from the Upper Cretaceous of Hokkaido, Japan. The cone is at least 20 cm long and up to 6 cm in diameter, consisting of a cone axis and numerous cone-scale complexes that are arranged helically around the axis. Two winged seeds are borne on the adaxial surface of each ovuliferous scale. Each complex receives a single trace from the vascular cylinder of the cone axis. In the scale base, all the resin canals occur abaxially to the vascular strand. The spatulate bract of the fossil is unique to the specimen among the cones of both living and fossil Pinus. The central umbo, broad sclerotic cortex of cone axis, and absence of serotinous features of the fossil cone suggest affinity with the subsection Sylvestres of the section Pinus, subgenus Pinus. This is the first record of permineralized preserved Pinus cone from the Cretaceous of Eastern Eurasia.     

福建梅花山17种常绿阔叶植物叶片虫食状况

从福建梅花山自然保护区海拔450m和1 200m的常绿阔叶林中选取17种常绿乔、灌木,对其叶片虫食状和取食强度进行分析,结果如下:(1)17种植物叶片共有15种虫食状类型,每种叶片虫食状类型为8~14;虫食状出现频率0.02%~37.01%,其中缘食状出现频率最高(37.01%),叶中脉食状出现频率最低(0.02%)。(2)木荷(Schima superba)的取食强度最大(9.60%),栲树(Castanopsis fargesii)次之(6.61%),油茶(Camellia Oleifera)最小(0.34%)。(3)15种虫食状类型中有6对虫食状呈显著正相关,2对呈显著负相关。(4)海拔1 200m处的甜槠(Castanopsis eyrei)、木荷、栲树和连蕊茶(Camellia fraterna)比海拔450m处的同种植物叶片的虫食强度高,其中木荷差异最大,从13.81%降至2.93%。