A new stem-neopterygian fish from the Middle Triassic of China shows the earliest over-water gliding strategy of the vertebrates

Flying fishes are extraordinary aquatic vertebrates capable of gliding great distances over water by exploiting their enlarged pectoral fins and asymmetrical caudal fin. Some 50 species of extant flying fishes are classified in the Exocoetidae (Neopterygii: Teleostei), which have a fossil record no older than the Eocene. The Thoracopteridae is the only pre-Cenozoic group of non-teleosts that shows an array of features associated with the capability of over-water gliding. Until recently, however, the fossil record of the Thoracopteridae has been limited to the Upper Triassic of Austria and Italy. Here, we report the discovery of exceptionally well-preserved fossils of a new thoracopterid flying fish from the Middle Triassic of China, which represents the earliest evidence of an over-water gliding strategy in vertebrates. The results of a phylogenetic analysis resolve the Thoracopteridae as a stem-group of the Neopterygii that is more crown-ward than the Peltopleuriformes, yet more basal than the Luganoiiformes. As the first record of the Thoracopteride in Asia, this new discovery extends the geographical distribution of this group from the western to eastern rim of the Palaeotethys Ocean, providing new evidence to support the Triassic biological exchanges between Europe and southern China. Additionally, the Middle Triassic date of the new thoracopterid supports the hypothesis that the re-establishment of marine ecosystems after end-Permian mass extinction is more rapid than previously thought.     

Reproduction in Middle Triassic actinopterygians; complex fin structures and evidence of viviparity in fossil fishes

New well-preserved specimens of Peltopleurus lissocephalus Brough, 1939, a small actinopterygian fish of the Alpine Middle Triassic, are described. Some of them show a highly modified anal fin, with a brush-shaped holdfast device and a gonopodium-like structure, possibly used during mating behaviour. Additional examples of modified anal fins are reported for the genera Cephaloxenus and Habroichthys. Evidence for viviparity within the genus Saurichthys , found at the same localities, is discussed and compared with earlier reports of the phenomenon. A comparison is made of the different reproductive strategies used in some Middle Triassic actinopterygian fishes.     

A new scanilepiform from the Lower Triassic of northern Gansu Province,China, and phylogenetic relationships of non‐teleostean Actinopterygii

A new scanilepiform, Beishanichthys brevicaudalis gen. et sp. nov. , is named and described based on fossils from the Lower Triassic lake deposits exposed in Beishan area, Gansu Province, China. The discovery documents a new record of this group, which is significantly older than other known scanilepiforms from China, and is slightly younger than Evenkia from the Lowest Triassic of Central Siberia. Although the Beishan beds were previously interpreted as Late Permian in age, based on megaplant fossils, this new discovery supports the reinterpretation of the deposits as Early Triassic in age, based on vertebrate fossils from the same locality and horizon. Phylogenetic analysis was conducted to resolve the relationships of Scanilepiformes with other actinopterygian clades, and the inter‐relationships within Scanilepiformes. Contrary to previous thought that scanilepiforms are closely related to the Amiidae, the phylogenetic results of this study recognize the Scanilepiformes as stem‐group neopterygians. Relationships of the Scanilepiformes and Australosomus with other neopterygians remain unresolved. With a characteristic long‐based dorsal fin, scanilepiforms represent a small group that emerged in Early Triassic freshwater environments, inhabited Eurasia and North America during the Middle–Late Triassic, briefly invaded the marine environment by the Late Triassic in Europe, and became extinct at the end of Triassic. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 595–612.     

Phylogenetic perspectives in the evolution of parental care in ray-finned fishes

Among major vertebrate groups, ray-finned fishes (Actinopterygii) collectively display a nearly unrivaled diversity of parental care activities. This fact, coupled with a growing body of phylogenetic data for Actinopterygii, makes these fishes a logical model system for analyzing the evolutionary histories of alternative parental care modes and associated reproductive behaviors. From an extensive literature review, we constructed a supertree for ray-finned fishes and used its phylogenetic topology to investigate the evolution of several key reproductive states including type of parental care (maternal, paternal, or biparental), internal versus external fertilization, internal versus external gestation, nest construction behavior, and presence versus absence of sexual dichromatism (as an indicator of sexual selection). Using a comparative phylogenetic approach, we critically evaluate several hypotheses regarding evolutionary pathways toward parental care. Results from maximum parsimony reconstructions indicate that all forms of parental care, including paternal, biparental, and maternal (both external and internal to the female reproductive tract) have arisen repeatedly and independently during ray-finned fish evolution. The most common evolutionary transitions were from external fertilization directly to paternal care and from external fertilization to maternal care via the intermediate step of internal fertilization. We also used maximum likelihood phylogenetic methods to test for statistical correlations and contingencies in the evolution of pairs of reproductive traits. Sexual dichromatism and nest construction proved to be positively correlated with the evolution of male parental care in species with external fertilization. Sexual dichromatism was also positively correlated with female-internal fertilization and gestation. No clear indication emerged that female-only care or biparental care were evolutionary outgrowths of male-only care, or that biparental care has been a common evolutionary stepping stone between paternal and maternal care. Results are discussed in the context of prior thought about the evolution of alternative parental care modes in vertebrates.     

A new rauisuchian archosaur from the Middle Triassic of India

Abstract:  Yarasuchus deccanensis gen. and sp. nov. is a new addition to the poorly known Middle Triassic terrestrial vertebrates. It is described on the basis of the fossil material of at least two incompletely preserved individuals and several isolated bones. It was a long-necked, gracile animal and might have had a facultatively bipedal gait . Apart from its characteristic elongate cervical vertebrae, other osteological features of Y. deccanensis resemble those of Prestosuchus, Saurosuchus, Ticinosuchus and ' Mandasuchus '. It is therefore referred to the Rauisuchian family Prestosuchidae despite the debated validity of this taxon.     

Tracing the genetic footprints of vertebrate landing in non-teleost ray-finned fishes

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A new sphenodontian (Lepidosauria: Rhynchocephalia) from the Late Triassic of Argentina and the early origin of the herbivore opisthodontians

Sphenodontians were a successful group of rhynchocephalian reptiles that dominated the fossil record of Lepidosauria during the Triassic and Jurassic. Although evidence of extinction is seen at the end of the Laurasian Early Cretaceous, they appeared to remain numerically abundant in South America until the end of the period. Most of the known Late Cretaceous record in South America is composed of opisthodontians, the herbivorous branch of Sphenodontia, whose oldest members were until recently reported to be from the Kimmeridgian–Tithonian (Late Jurassic). Here, we report a new sphenodontian, Sphenotitan leyesi gen. et sp. nov., collected from the Upper Triassic Quebrada del Barro Formation of northwestern Argentina. Phylogenetic analysis identifies Sphenotitan as a basal member of Opisthodontia, extending the known record of opisthodontians and the origin of herbivory in this group by 50 Myr.     

陕西鄂尔多斯盆地中三叠统一离片椎类牙齿化石

陕西省鄂尔多斯盆地渭北油田中三叠统铜川组发现一枚牙齿化石。该枚牙齿具有如下特征:牙齿细长,外形呈圆锥形,齿冠基部横截面为圆形,齿冠整体较直,沿长轴稍微向后缘弯曲,齿冠表面无齿脊和锯齿分布,发育纵向延伸的沟槽状纹饰。通过与鳄型类、蛇颈龙类和离片椎类的牙齿比较,该标本可以归入离片椎类。这枚牙齿化石为这一类群在中三叠统铜川组的首次发现,代表了离片椎类在华北板块最晚的化石记录。     

The oldest ionoscopiform from China sheds new light on the early evolution of halecomorph fishes

The Halecomorphi are a major subdivision of the ray-finned fishes. Although living halecomorphs are represented solely by the freshwater bowfin, Amia calva, this clade has a rich fossil history, and the resolution of interrelationships among extinct members is central to the problem of understanding the origin of the Teleostei, the largest clade of extant vertebrates. The Ionoscopiformes are extinct marine halecomorphs that were inferred to have originated in the Late Jurassic of Europe, and subsequently dispersed to the Early Cretaceous of the New World. Here, we report the discovery of a new ionoscopiform, Robustichthys luopingensis gen. et sp. nov., based on eight well-preserved specimens from the Anisian (242–247 Ma), Middle Triassic marine deposits of Luoping, eastern Yunnan Province, China. The new species documents the oldest known ionoscopiform, extending the stratigraphic range of this group by approximately 90 Ma, and the geographical distribution of this group into the Middle Triassic of South China, a part of eastern Palaeotethys Ocean. These new data provide a minimum estimate for the split of Ionoscopiformes from its sister clade Amiiformes and shed new light on the origin of ionoscopiform fishes.     

云南中三叠世(安尼期)预言鱼目(全骨鱼类:近鲱形类)一新属种

预言鱼目是近鲱形类的一个绝灭支系,被认为是弓鳍鱼目的姐妹群.预言鱼目化石过去主要发现于欧洲的中三叠世拉丁期和晚侏罗世地层以及新大陆的早白垩世地层.近年来,在云南和贵州的中三叠世安尼期地层中分别发现了一种预言鱼目鱼类化石(强壮鱼和盘县鱼),代表了该目在中国的首次发现.根据产于云南罗平中三叠世安尼期(～244 Ma)海相地层中的6块保存良好的鱼化石,命名了预言鱼目一个新的属种,三叠复兴鱼(Subortichthys triassicus gen.et sp.nov.).三叠复兴鱼是罗平生物群中发现的第二种预言鱼目鱼类,代表了预言鱼目最古老的化石记录之一,为研究该目的起源和早期分异提供了重要信息.复兴鱼无疑可以归入近鲱形类,因为它具有近鲱形类两个共近裔性状,续骨与下颌关节以及上颌骨后缘具有凹缺.分支分析结果表明,复兴鱼位于预言鱼目的基部,因为它具有该目的重要鉴定特征,上颌骨具有感觉管,但它不具有其他预言鱼目鱼类的进步特征.特别的是,复兴鱼具有一些独特的性状,如额外肩胛骨3或4对、第三眶下骨特别宽大并与前鳃盖骨前缘相接等.复兴鱼的发现表明预言鱼目鱼类在华南地区(三叠纪时期位于古特提斯洋的东部)的早期分异至少发生在中三叠世早期(安尼期).     

云南罗平中三叠世意外裸鱼的再研究

意外裸鱼(Gymnoichthys inopinatus)于2010年由Tintori等首次记述并归入基干新鳍鱼类。依据在云南省曲靖地区罗平县大凹子村中三叠世关岭组二段发现的新材料,对意外裸鱼进行了补充描述,并重新讨论了其系统发育关系。在意外裸鱼的新标本上,可见其续骨前端很可能与关节骨相关节,并有确切无疑的单一辅上颌骨,表明意外裸鱼应为鲱亚部(Halecomorphi)鱼类。此外,意外裸鱼的体表无鳞、椎体未骨化、髓棘和髓弓的结构和排列方式以及牙齿和尾脉棘的形状都与金尾鱼超科鱼类(caturoids)一致。系统发育分析结果表明意外裸鱼属于金尾鱼超科的基干类群。之前金尾鱼类主要见于欧洲和北美的侏罗系,现在公认的金尾鱼超科鱼类主要包括Liodesmidae中的Liodesmus属和金尾鱼科(Caturidae)中的Caturus和Amblysemius两属。意外裸鱼的发现不仅使金尾鱼类的出现提前了40Ma,而且填补了我国相关材料的空缺。     

Echinoid Spines from the Anisian (Middle Triassic) of Qingyan, South- western China

Echinoid remains from the Triassic of China are described for the first time. The Middle Triassic of Qingyan in Guizhou Province, south-western China, has yielded nine taxa of primary spines; seven of these come from the lower Upper Anisian. Because all of the spines are disarticulated, the material is treated only parataxonomically. 'Cidaris' lineola (Bather) and 'Cidaris' wissmanni (Desor) are species originally described from the alpine Triassic of Europe. 'Cidaris' bangtoupoensis sp. nov., 'Cidaris' wachangpoensis sp. nov., 'Cidaris' venustespinosa sp. nov., 'Cidaris' fangchui sp. nov., 'Cidaris' gu sp. nov., 'Cidaris' mafengpoensis sp. nov. and 'Cidaris' sp. A are so far known only from Qingyan. Palaeoautecological interpretations based on the shapes of the spines fit well with the fact that the echinoids from Qingyan are part of highly diverse fossil associations which represent the remains of communities that lived in a protected, shallow-marine, stable soft-bottom habitat. The rather high diversity of the echinoid spines in terms of the number of both taxa and general types of shape shows that the echinoids, after nearly becoming extinct at the end of the Palaeozoic, began a new phase of adaptive radiation in the Anisian at the latest.     

Comparative morphology and cytology of the male sperm-transmission organs in viviparous species of clinid fishes (Clinidae: Teleostei, Perciformes)

This work comprises the first comparative study of the morphology and cytology of the sperm transmission organs in males of 14 species of viviparous clinid fishes (Clinidae, Blennioidei, Teleostei). The form and dimensions of these organs differ among the various species studied. The organs are composed of intra-abdominal ampullae, into which the sperm ducts and urinary bladder anchor, and an external protruding intromittent papilla used for insemination. The form of the ampullae differs among the various species, from pear-shaped to horseshoe-shaped. It increases in dimensions with increasing length of the male. In all the species this organ is covered by a connective-tissue tunic that encompasses both circular and longitudinal striated muscle bundles. The lumina of the ampullae harbor the epididymis, a strongly convoluted and plicated duct, which becomes filled with spermatozeugmata during reproduction. From here, the epididymis continues into the protruding intromittent papillae, where its folds gradually straighten at the apical part of the intromittent organ. The form and dimensions of this copulatory organ also differ in the various species. Papillae bearing taste buds are found on the apical parts of the intromittent organ, and it is probable that these, together with the difference in forms of the organ, help to prevent interspecific copulation.     

渤海沿岸临清坳陷堂古5井中三叠统轮藻化石与地层划分对比

在临清坳陷堂古5井发现轮藻化石Stenochara和Stellatochara两属7种,证实该区存在三叠世地层。根据地层层序、岩石地层特征和区域地质资料分析,该区亦应存在下三叠统,并建议参照区域方案将该区中、下三叠统红层划分为刘家沟组、和尚沟组和二马营组。     

Pseudoplanktonic lifestyle of the Triassic crinoid Traumatocrinus from Southwest China

Colonies of Traumatocrinus (Echinodermata, Crinoidea) attached to driftwood from the lower Upper Triassic (Carnian) Xiaowa Formation of the Guanling area (Guizhou, Southwest China) document a pseudoplanktonic lifestyle for this specialized offshoot of the otherwise benthic Middle Triassic Encrinidae. After the end-Carnian decline of Traumatocrinus, its ecological niche was subsumed in Norian by genera derived from another benthic family (Holocrinidae) with convergent morphological modifications. After the Toarcian (Lower Jurassic), this niche disappeared, possibly due to the emergence of wood-boring bivalves.     

Diet of prosauropod dinosaurs from the late Triassic and early Jurassic

Prosauropods were not scavenger-predators, rather they were the dominant large terrestrial herbivores during the late Triassic and early Jurassic. The herbivorous adaptations of anchisaurids include spatulate teeth with anteroposteriorly expanded crowns (maximum width apical to base of crown) which are obliquely inclined with respect to the jaws so each slightly overlaps the tooth behind it, and which have coarse marginal serrations at 45° to the cutting edges. Most of the teeth of yunnanosaurids lack serrations and resemble those of sauropod dinosaurs in form and in having self-sharpening surfaces, formed by tooth-to-tooth wear, which increased the efficiency of dealing with more resistant plant material. Anchisaurids and yunnanosaurids had a ventrally set jaw articulation; the teeth and skull of melanorosaurids are unknown. All prosauropods were high browsers that extended the feeding range with a long neck and tripodal feeding (long hindlimbs and stout tail for support). They used herding and the enormous claw on the pollex for defense, and probably had a muscular gastric mill with stones that was used for grinding the food. They account for at least 95% of the biomass in their respective faunas.     

An early isocrinid sea lily from the middle to late Anisian boundary (Middle Triassic) of south‐west China – evidence for a far‐east Tethyan origin of the family Isocrinidae

Abstract: The isocrinid sea lily Tyrolecrinus wugangi sp. nov. from uppermost middle to lowermost upper Anisian (lower Middle Triassic) strata of Leidapo near Qingyan, Guizhou Province, south‐west China, provides new data on the early phylogeny of the order Isocrinida. The new species is the earliest unequivocal representative of the family Isocrinidae. It predates all other known species of the genus Tyrolecrinus that come from upper Ladinian to Rhaetian strata and also all other known taxa of the Isocrinidae. Nevertheless, its stem is of fully developed isocrinid type and is characteristic of the genus, with consistently synostosial, rarely cryptosymplectial, distal nodal articular facets. These findings indicate that the Isocrinidae evolved prior to the late middle Anisian in the far‐eastern part of the Palaeotethys ocean, perhaps in the region forming today’s south‐west China, and spread to various other regions later in the Middle and Late Triassic. The genus Tyrolecrinus is revised and the new genus Bakonycrinus gen. nov. is erected.     

云南罗平中三叠世翼鳕属一新种及早期辐鳍鱼类系统发育关系（英文）

辐鳍鱼亚纲是现存脊椎动物中最大的类群,包括腕鳍鱼次亚纲、辐鳍鱼次亚纲(包括软骨硬鳞类和新鳍鱼类)和亲缘关系密切的化石类群。已灭绝的翼鳕属(Pteronisculus)是隶属于辐鳍鱼亚纲的一个干群,包括产于马达加斯加、欧洲和北美下三叠统的11个种和中国中三叠统的一个种。根据滇东罗平中三叠世(安尼期)海相地层中发现的5块保存完好的化石,命名翼鳕属一新种,张氏翼鳕(Pteronisculus changae sp. nov.)。这是翼鳕属在中三叠世的第二个确切种,最大体长达295 mm,代表了罗平生物群中已知体型最大的辐鳍鱼亚纲干群物种。新种具有翼鳕属的独特衍征,泪骨具有牙齿,但它又有明显区别于本属其他种的自近裔特征,如间颞骨中部有一个内突起,21根上神经骨,83列侧线鳞。分支分析结果为早期辐鳍鱼类系统发育关系提供了新的见解,认为翼鳕属是Cyranorhis的姐妹群。根据体型和口缘牙齿等特征推测张氏翼鳕是一个快速游动的捕食者,以浮游无脊椎动物和体型较小的鱼类或鱼类幼体为食。作为翼鳕属最年轻的成员之一,张氏翼鳕的发现进一步表明翼鳕的多样性比我们过去认识的要高,古特提斯洋东缘可能是该属在中三叠世早期的避难所。     

A theropod tooth from the Late Triassic of southern Africa

An isolated, large recurved and finely serrated tooth found associated with the prosauropodEuskelosaurus from the Late Triassic part of the Elliot Formation is described here. It is compared to the Triassic thecodonts and carnivorous dinosaurs and its possible affinity is discussed. The tooth possibly belongs to a basal theropod and shows some features similar to the allosauroids. This tooth is of significance, as dinosaur remains except for some footprints and trackways, are poorly known in the Late Triassic horizons of southern Africa.