A transitional snake from the Late Cretaceous period of North America

Snakes are the most diverse group of lizards, but their origins and early evolution remain poorly understood owing to a lack of transitional forms. Several major issues remain outstanding, such as whether snakes originated in a marine or terrestrial environment and how their unique feeding mechanism evolved. The Cretaceous Coniophis precedens was among the first Mesozoic snakes discovered, but until now only an isolated vertebra has been described and it has therefore been overlooked in discussions of snake evolution. Here we report on previously undescribed material from this ancient snake, including the maxilla, dentary and additional vertebrae. Coniophis is not an anilioid as previously thought a revised phylogenetic analysis of Ophidia shows that it instead represents the most primitive known snake. Accordingly, its morphology and ecology are critical to understanding snake evolution. Coniophis occurs in a continental floodplain environment, consistent with a terrestrial rather than a marine origin; furthermore, its small size and reduced neural spines indicate fossorial habits, suggesting that snakes evolved from burrowing lizards. The skull is intermediate between that of lizards and snakes. Hooked teeth and an intramandibular joint indicate that Coniophis fed on relatively large, soft-bodied prey. However, the maxilla is firmly united with the skull, indicating an akinetic rostrum. Coniophis therefore represents a transitional snake, combining a snake-like body and a lizard-like head. Subsequent to the evolution of a serpentine body and carnivory, snakes evolved a highly specialized, kinetic skull, which was followed by a major adaptive radiation in the Early Cretaceous period. This pattern suggests that the kinetic skull was a key innovation that permitted the diversification of snakes.     

The Pleistocene serpent Wonambi and the early evolution of snakes

The Madtsoiidae were medium sized to gigantic snakes with a fossil record extending from the mid-Cretaceous to the Pleistocene, and spanning Europe, Africa, Madagascar, South America and Australia. This widely distributed group survived for about 90 million years (70% of known ophidian history), and potentially provides important insights into the origin and early evolution of snakes. However, madtsoiids are known mostly from their vertebrae, and their skull morphology and phylogenetic affinities have been enigmatic. Here we report new Australian material of Wonambi, one of the last-surviving madtsoiids, that allows the first detailed assessment of madtsoiid cranial anatomy and relationships. Despite its recent age, which could have overlapped with human history in Australia, Wonambi is one of the most primitive snakes known--as basal as the Cretaceous forms Pachyrhachis and Dinilysia. None of these three primitive snake lineages shows features associated with burrowing, nor do any of the nearest lizard relatives of snakes (varanoids). These phylogenetic conclusions contradict the widely held 'subterranean' theory of snake origins, and instead imply that burrowing snakes (scolecophidians and anilioids) acquired their fossorial adaptations after the evolution of the snake body form and jaw apparatus in a large aquatic or (surface-active) terrestrial ancestor.     

Skull of the large non-macrostomatan snake Yurlunggur from the Australian Oligo-Miocene

Understanding the origin and early evolution of snakes from lizards depends on accurate morphological knowledge of the skull in basal lineages, but fossil specimens of archaic snakes have been rare, and either fragmentary or difficult to study as a result of compression by enclosing sediments. A number of Cenozoic fossil snakes from Australia have vertebral morphology diagnostic of an extinct group, Madtsoiidae, that was widespread in Gondwana from mid-Cretaceous (Cenomanian) to Eocene times, and also reached Europe in the late Cretaceous period. Despite this long history, only about half the skull is known from the best-known species Wonambi naracoortensis, and the few known cranial elements of other species have added little further evidence for phylogenetic relationships. Conflicting hypotheses have been proposed for their relationships and evolutionary significance, either as basal ophidians with many ancestral (varanoid- or mosasaur-like) features, or advanced (macrostomatan) alethinophidians of little relevance to snake origins. Here I report two partial skeletons referred to Yurlunggur, from the late Oligocene and early Miocene of northern Australia, which together represent almost the complete skull and mandible. The exceptionally preserved skulls provide new evidence linking Yurlunggur with Wonambi and other madtsoiids, falsifying predictions of the macrostomatan hypothesis, and supporting the exclusion of Madtsoiidae from the clade including all extant snakes.     

安徽巢湖早三叠世原始鱼龙类的结构和行为特征及其对鱼龙类起源研究的启示

鱼龙类(Ichthyosaurs)起源于早三叠世晚期, 以往发现的鱼龙类已高度适应水生生活, 其身体结构无任何能将其与可能的陆地祖先相联系的陆生适应特征。针对近年在安徽巢湖下三叠统奥伦尼克阶采集的柔腕短吻龙(Cartorhynchus)和含胚胎巢湖龙两件鱼龙类标本, 通过对柔腕短吻龙的荐前椎数目(31节)、吻长与头骨长之比(0.35)、前肢长与荐前椎长之比(0.45)的研究以及与后期鱼龙类相关数据和骨骼形态学的对比, 认为柔腕短吻龙结构更为原始, 可两栖生活。结合对早三叠世含胚胎巢湖龙、侏罗纪含胚胎狭翼鱼龙幼体保存情况的对比分析, 认为巢湖龙虽比柔腕短吻龙身体结构更进化, 且完全适应水生生活, 但其生殖行为仍然保持陆生爬行动物头位胎生的特征。鱼龙类在其陆地祖先逐渐演化至适应水生生活的过程中, 结构的适应性特征(如肢体的鳍状化、吻部的增长等)和行为特征(如生产方式的变化)的变化不同步, 前者的改变更早。     

Bone histology indicates insular dwarfism in a new Late Jurassic sauropod dinosaur

Sauropod dinosaurs were the largest animals ever to inhabit the land, with truly gigantic forms in at least three lineages. Small species with an adult body mass less than five tonnes are very rare, and small sauropod bones generally represent juveniles. Here we describe a new diminutive species of basal macronarian sauropod, Europasaurus holgeri gen. et sp. nov., and on the basis of bone histology we show it to have been a dwarf species. The fossils, including excellent skull material, come from Kimmeridgian marine beds of northern Germany, and record more than 11 individuals of sauropods 1.7 to 6.2 m in total body length. Morphological overlap between partial skeletons and isolated bones links all material to the same new taxon. Cortical histology of femora and tibiae indicates that size differences within the specimens are due to different ontogenetic stages, from juveniles to fully grown individuals. The little dinosaurs must have lived on one of the large islands around the Lower Saxony basin. Comparison with the long-bone histology of large-bodied sauropods suggests that the island dwarf species evolved through a decrease in growth rate from its larger ancestor.     

A Middle Jurassic 'sphenosuchian' from China and the origin of the crocodylian skull

The skull of living crocodylians is highly solidified and the jaw closing muscles are enlarged, allowing for prey capture by prolonged crushing between the jaws. Living species are all semi-aquatic, with sprawling limbs and a broad body that moves mainly from side-to-side; however, fossils indicate that they evolved from terrestrial forms. The most cursorial of these fossils are small, gracile forms often grouped together as the Sphenosuchia, with fully erect, slender limbs; their relationships, however, are poorly understood. A new crocodylomorph from deposits in northwestern China of the poorly known Middle Jurassic epoch possesses a skull with several adaptations typical of living crocodylians. Postcranially it is similar to sphenosuchians but with even greater adaptations for cursoriality in the forelimb. Here we show, through phylogenetic analysis, that it is the closest relative of the large group Crocodyliformes, including living crocodylians. Thus, important features of the modern crocodylian skull evolved during a phase when the postcranial skeleton was evolving towards greater cursoriality, rather than towards their current semi-aquatic habitus.     

Hydatellaceae identified as a new branch near the base of the angiosperm phylogenetic tree

Although the relationship of angiosperms to other seed plants remains controversial, great progress has been made in identifying the earliest extant splits in flowering-plant phylogeny, with the discovery that the New Caledonian shrub Amborella trichopoda, the water lilies (Nymphaeales), and the woody Austrobaileyales constitute a basal grade of lines that diverged before the main radiation in the clade. By focusing attention on these ancient lines, this finding has re-written our understanding of angiosperm structural and reproductive biology, physiology, ecology and taxonomy. The discovery of a new basal lineage would lead to further re-evaluation of the initial angiosperm radiation, but would also be unexpected, as nearly all of the approximately 460 flowering-plant families have been surveyed in molecular studies. Here we show that Hydatellaceae, a small family of dwarf aquatics that were formerly interpreted as monocots, are instead a highly modified and previously unrecognized ancient lineage of angiosperms. Molecular phylogenetic analyses of multiple plastid genes and associated noncoding regions from the two genera of Hydatellaceae identify this overlooked family as the sister group of Nymphaeales. This surprising result is further corroborated by evidence from the nuclear gene phytochrome C (PHYC), and by numerous morphological characters. This indicates that water lilies are part of a larger lineage that evolved more extreme and diverse modifications for life in an aquatic habitat than previously recognized.     

我国蜥蜴能量生态学的研究概况

蜥蜴是爬行动物进化的一个关键类群.有关蜥蜴能量生态学的研究是倍受关注的领域.本文概括了蜥蜴能量生态学主要研究内容,重点介绍了我国蜥蜴能量生态学的研究进展.     

Discovery of novel intermediate forms redefines the fungal tree of life

Fungi are the principal degraders of biomass in terrestrial ecosystems and establish important interactions with plants and animals. However, our current understanding of fungal evolutionary diversity is incomplete and is based upon species amenable to growth in culture. These culturable fungi are typically yeast or filamentous forms, bound by a rigid cell wall rich in chitin. Evolution of this body plan was thought critical for the success of the Fungi, enabling them to adapt to heterogeneous habitats and live by osmotrophy: extracellular digestion followed by nutrient uptake. Here we investigate the ecology and cell biology of a previously undescribed and highly diverse form of eukaryotic life that branches with the Fungi, using environmental DNA analyses combined with fluorescent detection via DNA probes. This clade is present in numerous ecosystems including soil, freshwater and aquatic sediments. Phylogenetic analyses using multiple ribosomal RNA genes place this clade with Rozella, the putative primary branch of the fungal kingdom. Tyramide signal amplification coupled with group-specific fluorescence in situ hybridization reveals that the target cells are small eukaryotes of 3-5?μm in length, capable of forming a microtubule-based flagellum. Co-staining with cell wall markers demonstrates that representatives from the clade do not produce a chitin-rich cell wall during any of the life cycle stages observed and therefore do not conform to the standard fungal body plan. We name this highly diverse clade the cryptomycota in anticipation of formal classification.     

Tail-assisted pitch control in lizards, robots and dinosaurs

In 1969, a palaeontologist proposed that theropod dinosaurs used their tails as dynamic stabilizers during rapid or irregular movements, contributing to their depiction as active and agile predators. Since then the inertia of swinging appendages has been implicated in stabilizing human walking, aiding acrobatic manoeuvres by primates and rodents, and enabling cats to balance on branches. Recent studies on geckos suggest that active tail stabilization occurs during climbing, righting and gliding. By contrast, studies on the effect of lizard tail loss show evidence of a decrease, an increase or no change in performance. Application of a control-theoretic framework could advance our general understanding of inertial appendage use in locomotion. Here we report that lizards control the swing of their tails in a measured manner to redirect angular momentum from their bodies to their tails, stabilizing body attitude in the sagittal plane. We video-recorded Red-Headed Agama lizards (Agama agama) leaping towards a vertical surface by first vaulting onto an obstacle with variable traction to induce a range of perturbations in body angular momentum. To examine a known controlled tail response, we built a lizard-sized robot with an active tail that used sensory feedback to stabilize pitch as it drove off a ramp. Our dynamics model revealed that a body swinging its tail experienced less rotation than a body with a rigid tail, a passively compliant tail or no tail. To compare a range of tails, we calculated tail effectiveness as the amount of tailless body rotation a tail could stabilize. A model Velociraptor mongoliensis supported the initial tail stabilization hypothesis, showing as it did a greater tail effectiveness than the Agama lizards. Leaping lizards show that inertial control of body attitude can advance our understanding of appendage evolution and provide biological inspiration for the next generation of manoeuvrable search-and-rescue robots.     

Discovery of a short-necked sauropod dinosaur from the Late Jurassic period of Patagonia

Sauropod dinosaurs are one of the most conspicuous groups of Mesozoic terrestrial vertebrates. They show general trends towards an overall increase in size and elongation of the neck, by means of considerable elongation of the length of individual vertebrae and a cervical vertebra count that, in some cases, increases to 19 (ref. 1). The long neck is a particular hallmark of sauropod dinosaurs and is usually regarded as a key feeding adaptation. Here we describe a new dicraeosaurid sauropod, from the latest Jurassic period of Patagonia, that has a particularly short neck. With a neck that is about 40% shorter than in other known dicraeosaurs, this taxon demonstrates a trend opposite to that seen in most sauropods and indicates that the ecology of dicraeosaurids might have differed considerably from that of other sauropods. The new taxon indicates that there was a rapid radiation and dispersal of dicraeosaurids in the Late Jurassic of the Southern Hemisphere, after the separation of Gondwana from the northern continents by the late Middle Jurassic.     

黑眉锦蛇和乌梢蛇几种组织中乳酸脱氢酶同工酶的比较研究

本文采用聚丙烯酰胺凝胶电(冫永)方法,研究了黑眉锦蛇和乌梢蛇血清、心肌和肝脏的乳酸脱氢酶同工酶。结果表明:该二种蛇的血清仅具有LDH_1一种同工酶;心肌有三种LDH同工酶,它们以各自的特异性比例而存在,以B亚基占优势为特征。在肝脏中,两种蛇都具有四种LDH同工酶主带,乌梢蛇还有一条附属于LDH_2的亚带而不同于黑眉锦蛇。根据实验结果,本文初步探讨了该二种蛇的分类地位,同时还对乳酸脱氢酶(LDH)同工酶与遗传、生理、生化、生态的关系作了讨论。     

Dual origin of tribosphenic mammals

Marsupials, placentals and their close therian relatives possess complex (tribosphenic) molars that are capable of versatile occlusal functions. This functional complex is widely thought to be a key to the early diversification and evolutionary success of extant therians and their close relatives (tribosphenidans). Long thought to have arisen on northern continents, tribosphenic mammals have recently been reported from southern landmasses. The great age and advanced morphology of these new mammals has led to the alternative suggestion of a Gondwanan origin for the group. Implicit in both biogeographic hypotheses is the assumption that tribosphenic molars evolved only once in mammalian evolutionary history. Phylogenetic and morphometric analyses including these newly discovered taxa suggest a different interpretation: that mammals with tribosphenic molars are not monophyletic. Tribosphenic molars evolved independently in two ancient (holotherian) mammalian groups with different geographic distributions during the Jurassic/Early Cretaceous: an australosphenidan clade endemic to Gondwanan landmasses, survived by extant monotremes; and a boreosphenidan clade of Laurasian continents, including extant marsupials, placentals and their relatives.     

脆蛇转录组序列的分析和系统发育定位(英文)

脆蛇是一种类似于蛇但是又具有蜥蜴的特征的爬行动物,这些矛盾的特性与脆蛇的系统发育分类学有密切的关系.脆蛇在传统中药中也有很多治疗作用.基于这些方面,本研究利用高通量测序方法对脆蛇胃肠道的转录组序列进行检测,组装和注释.最终获得4.6 Gbp高质量的数据,组装获得58 959个单一的基因,其中35 708(60.56%)个单一基因与数据库的基因相匹配.为了确定脆蛇与蛇和蜥蜴的同源进化关系,对同源基因家族和系统进化树进行了分析,结果显示脆蛇的转录组序列更接近于蛇而不是蜥蜴.除此之外还鉴定了10 613 cSSR标签,其中1 644个标签根据严格的标准能够用至少一个引物获得.本研究第一次揭示了脆蛇胃肠道的转录组序列,确定其在系统发育学上的定位.这些序列和标签为脆蛇的研究提供了重要的资源.     

Developmental basis of limblessness and axial patterning in snakes.

The evolution of snakes involved major changes in vertebrate body plan organization, but the developmental basis of those changes is unknown. The python axial skeleton consists of hundreds of similar vertebrae, forelimbs are absent and hindlimbs are severely reduced. Combined limb loss and trunk elongation is found in many vertebrate taxa, suggesting that these changes may be linked by a common developmental mechanism. Here we show that Hox gene expression domains are expanded along the body axis in python embryos, and that this can account for both the absence of forelimbs and the expansion of thoracic identity in the axial skeleton. Hindlimb buds are initiated, but apical-ridge and polarizing-region signalling pathways that are normally required for limb development are not activated. Leg bud outgrowth and signalling by Sonic hedgehog in pythons can be rescued by application of fibroblast growth factor or by recombination with chick apical ridge. The failure to activate these signalling pathways during normal python development may also stem from changes in Hox gene expression that occurred early in snake evolution.     

草原沙蜥和避役骨骼解剖分析——游舌部蜥蜴(Ascalabotans)骨骼比较

以草原沙蜥和避役的骨骼解剖为素材,证明它们是较原始的蜥蜴,两者共同的原始性为:颞窝宽大,顶骨呈板状,牙齿三尖,下颌板骨前端参加下颌联合,环椎的间椎体同神经弧松散结合,荐椎无分化,尾椎脉弧呈Y型,腕骨超过9枚。各自的原始性状:草原沙蜥为顶骨孔发达,上颞骨存在,腭部骨片未退缩,而且排列位置未变。避役为上颞弧由后眶鳞骨构成。归纳喙头蜥、(?)鳞蜥、草原沙蜥和避役的骨骼结构,为原始蜥蜴特征提供新的证据。同时分析了游舌部蜥蜴的系统发育关系。     

A bizarre tandem-horned elasmothere rhino from the Late Miocene of northwestern China and origin of the true elasmothere

Although the modern Indian and Javan rhinos have a single horn on their noses,the extinct one-horned rhino Elasmotherium was a source for the legendary unicorn,because the latter had a very long horn on its forehead and lived with the prehistoric human beings who drew its images on cave paintings.Elasmothere rhinos first appeared in South Asia in the Early Miocene,but the origin of Elasmotherium has been unclear.All other elasmotheres have a weak or strong nasal horn,whereas Elasmotherium seems to lose the nasal horn of its ancestors and to get a huge frontal horn apparently abruptly.Here we report the first discovered skull of Sinotherium lagrelii from the Late Miocene red clays in the Linxia Basin,northwestern China.This skull has an enormous nasofrontal horn boss shifted posteriorly and a smaller frontal horn boss,which are connected to each other,indicating an intermediate stage for the single frontal horn of Elasmotherium.Morphological and phylogenetic analyses confirm that Sinotherium is a transitional taxon between Elasmotherium and other elasmotheres,positioned near the root of the giant unicorn clade and originated in a subarid steppe.The posteriorly shifted nasal horn has a more substantial support and the arched structure of the nasofrontal area is an adaptation for a huge horn.     

西域沙虎骨骼系统的解剖研究

观察研究了西域沙虎(Teratoscincus przewalskii)的骨骼系统,对其各部分骨骼的构成、形状及位置等均作了详细的描述,并与其它蜥蜴类动物进行了比较。结果表明,西域沙虎既有蜥蜴类共有的结构,也有其自身的特征。     

Large Cretaceous sphenodontian from Patagonia provides insight into lepidosaur evolution in Gondwana

Sphenodontian reptiles successfully radiated during Triassic and Jurassic times, but were driven almost to extinction during the Cretaceous period. The sparse Early Cretaceous record of sphenodontians has been interpreted as reflecting the decline of the group in favour of lizards, their suspected ecological successors. However, recent discoveries in Late Cretaceous beds in Patagonia partially modify this interpretation. Numerous skeletons of a new sphenodontian, Priosphenodon avelasi gen. et sp. nov., were collected from a single locality in the Cenomanian-Turonian Candeleros Formation, where it is more abundant than any other tetrapod group recorded in the quarry (for example, Crocodyliformes, Serpentes, Dinosauria and Mammalia). Adult specimens of Priosphenodon reached one metre in length, larger than any previously known terrestrial sphenodontian. Here we propose, using available evidence, that sphenodontians were not a minor component of the Cretaceous terrestrial ecosystems of South America, and that their ecological replacement by squamates was delayed until the early Tertiary. The new discovery helps to bridge the considerable gap in the fossil record (around 120 million years) that separates the Early Cretaceous sphenodontians from their living relatives (Sphenodon).