The Luoping biota: exceptional preservation, and new evidence on the Triassic recovery from end-Permian mass extinction

The timing and nature of biotic recovery from the devastating end-Permian mass extinction (252 Ma) are much debated. New studies in South China suggest that complex marine ecosystems did not become re-established until the middle–late Anisian (Middle Triassic), much later than had been proposed by some. The recently discovered exceptionally preserved Luoping biota from the Anisian Stage of the Middle Triassic, Yunnan Province and southwest China shows this final stage of community assembly on the continental shelf. The fossil assemblage is a mixture of marine animals, including abundant lightly sclerotized arthropods, associated with fishes, marine reptiles, bivalves, gastropods, belemnoids, ammonoids, echinoderms, brachiopods, conodonts and foraminifers, as well as plants and rare arthropods from nearby land. In some ways, the Luoping biota rebuilt the framework of the pre-extinction latest Permian marine ecosystem, but it differed too in profound ways. New trophic levels were introduced, most notably among top predators in the form of the diverse marine reptiles that had no evident analogues in the Late Permian. The Luoping biota is one of the most diverse Triassic marine fossil Lagerstätten in the world, providing a new and early window on recovery and radiation of Triassic marine ecosystems some 10 Myr after the end-Permian mass extinction.     

New ophiuroids from the Permian/Triassic boundary beds of South China

A new genus of ophiuroid, Huangzhishania , is created based on new material from the Permian/Triassic boundary beds at the Huangzhishan section, South China. The age of the new genus is constrained as earliest Griesbachian by means of faunal correlation of the associated bivalves and stratigraphical correlation with the Mixed Fauna Beds of the neighbouring Meishan section. Taphonomic and palaeoecological evidence suggest that the collapse of the ophiuroid association was related to a catastrophic event, and Huangzhishania was rapidly buried in life position.     

A new galesaurid (Therapsida: Cynodontia) from the Lower Triassic of South Africa

A new galesaurid cynodont, Progalesaurus lootsbergensis gen. et sp. nov., is described on the basis of a well-preserved skull, lower jaw, right scapula and left atlantal neural arch. Autapomorphies of Progalesaurus include postcanine teeth bearing numerous mesial and distal accessory cusps that flank a recurved main cusp, a post-temporal fenestra bordered by the squamosal ventrally and a large external naris. Progalesaurus is similar to Galesaurus in possessing a poorly defined masseteric fossa on the dentary, a strongly recurved main cusp of the postcanine dentition, an incomplete secondary palate and a similar basisphenoid-parasphenoid morphology. A cladistic analysis of ten early cynodont genera resolves a monophyletic Galesauridae encompassing Cynosaurus , Progalesaurus and Galesaurus , although support for this clade is weak. Procynosuchus and Dvinia are placed at the base of Cynodontia whereas Thrinaxodon and Platycraniellus are positioned higher, but outside of Eucynodontia. The holotype and only known specimen of Progalesaurus was collected during systematic prospecting of Permo/Triassic boundary strata at New Lootsberg Pass, Graaff-Reinet District, South Africa. The discovery of Progalesaurus increases the number of valid Early Triassic cynodonts to four and sheds light on the tempo of early cynodont diversification after the end-Permian mass extinction.     

End-Permian mass extinction of calcareous algae and microproblematica from Liangfengya,South China

A new assemblage of calcareous algae and microproblematica is reported from the Changxing Formation at the Liangfengya section in Chongqing, South China. This assemblage comprises eighteen species of seven genera, including three genera of gymnocodiaceaens (Gymnocodium, Permocalculus, and Tauridium), three genera of dasycladaleans (Epimastopora, Macroporella, and Mizzia), and one genus of microproblematica (Pseudovermiporella). A new algal species, Tauridium elongatum nov. sp., is described. Quantitative analysis indicates that the last occurrences of 22% of the species (4 out of 18) fall into a 52 cm thick interval of the uppermost Changhsingian. No calcareous algae are found in the Permian-Triassic boundary (P-T boundary) beds and the overlying Feixianguan Formation. Four out of 10 species have a stratigraphic abundance greater than 10% and all vanished in the uppermost Changhsingian. An abrupt extinction for calcareous algae occurred in the Clarkina yini conodont Zone.     

Palaeoecology of microconchids from microbialites near the Permian–Triassic boundary in South China

Abundant isolated specimens of microconchid tubes have been extracted from a microbialite deposit near the Permian–Triassic boundary (PTB) in the Dajiang section, southern Guizhou Province, South China. They are assignable to Microconchus aff. utahensis, M. aff. aberrans and Helicoconchus aff. elongatus, all of which possess micro‐lamellar tube walls. Quantitative analysis of bulk samples indicates that most microconchids occur in the upper part of the PTB microbialite deposit and show substrate selectivity for bioclastic grainstone–packstones. In contrast, very few microconchids were found in the rocks bearing well‐developed microbialite structures. Both stratigraphical and substrate preferences indicate proliferation of microconchids coincided with an ebb of microbialite development. Microconchids therefore only proliferated in local niches in which microbial activities were not very active within the PTB microbialite ecosystem. The presence of abundant microconchids further strengthens the impression that PTB microbialite metazoans are much more diverse than previously thought. The end‐Permian mass extinction is calibrated to the base of microbialite deposit in South China. Thus, abundant microbialite metazoans, such as ostracods, lingulid brachiopods, microgastropods and microconchids, together with the considerable, temporarily surviving faunas reported from non‐microbialite PTB sections in South China, indicate that metazoans diversified immediately after the first episode of the end‐Permian mass extinction, supporting the scenario that marine ecosystems underwent episodic collapses during the devastating biocrisis over the Permian–Triassic transition.     

Recovery dynamics of benthic marine communities from the Lower Triassic Werfen Formation,northern Italy

The Lower Triassic Werfen Formation of northern Italy represents an important archive for Early Triassic ecosystems. Based on quantitative community analysis using species level identifications, we reconstruct the recovery of benthic ecosystems after the end‐Permian mass extinction throughout this unit. The analysis of benthic macrofossil communities shows that incipient recovery has taken place during the Griesbachian and the Griesbachian–Dienerian transition. A probable deterioration in environmental conditions is observed towards the end of the Dienerian. The Smithian part of the Werfen Formation is characterized by high siliciclastic input, which ceases around the Smithian‐Spathian boundary. The Spathian marks the definitive phase of recovery in the Werfen Formation. The comparison of this pattern with other palaoegeographical regions suggests that both the Griesbachian recovery and the Dienerian decline were of inter‐regional if not global extent, whereas the Smithian diversity low in the Werfen Formation is a local signal. In contrast to the recovery dynamics of ammonoids and conodonts, the Smithian–Spathian boundary interval was no caesura for benthic ecosystems. The Spathian recovery pulse is possibly also an inter‐regional event, at least in the palaeotropics. These results are in contrast with the previously proposed scenario of persistent hostile conditions during the Griesbachian time interval and highlight an initial recovery phase restricted to Griesbachian times. Instead, the apparently sluggish recovery of benthic ecosystems was at least partly shaped by set‐backs due to short‐term environmental perturbations during the Dienerian.     

Additional thylacocephalans (Arthropoda) from the Lower Triassic (upper Olenekian) Osawa Formation of the South Kitakami Belt,Northeast Japan

The Lower Triassic Osawa Formation in the South Kitakami Belt, Northeast Japan, consisting mostly of mudstone of shallow-marine environment, was deposited during the late Olenekian (ca. 250 Ma), and is an important unit through which to examine the biotic recovery process after the end-Permian mass extinction. The Osawa Formation is the only unit in Japan that yields thylacocephalans (Arthropoda). Three species belonging to three genera have been reported before: Ankitokazocaris bandoi, Kitakamicaris utatsuensis and Ostenocaris sp. In addition to the known species, some thylacocephalans, including one new genus and three new species, are described in the present paper: Ankitokazocaris tatensis n. sp., Concavicaris parva n. sp., Miyagicaris costata n. gen. n. sp. and Ostenocaris? sp. Although Thylacocephala have a rather long stratigraphic range (from Silurian to Cretaceous) and are known from a wide geographical region, there are only about thirty genera in this group. The Osawa thylacocephalan fauna comprises at least five genera, making it one of the most diverse in the world at the generic level. During the Triassic Period, the Thylacocephala diversified and spread widely throughout low-latitude regions.     

Ontogeny of the eodiscoid trilobite Tsunyidiscus acutus from the Lower Cambrian of South China

Abstract: The morphology and ontogeny of the eodiscid trilobite Tsunyidiscus acutus Sun is described on the basis of numerous calcified specimens collected from the Lower Cambrian Shuijingtuo Formation in Yichang and Changyang, Hubei Province, South China. An ontogenetic series is established based on the articulated material including the previously unknown protaspides and meraspides (degrees 0 and 1). The material, revealing some prominent morphological changes such as the number of pygidial axial rings, genal spines retained throughout ontogeny and pygidial pleurae from furrowed to unfurrowed, enables a discussion on the trunk segmentation schedule, indicating that somitogenesis and tagmosis occurred independently during the ontogenetic development of T. acutus.     

An unusually diverse mollusc fauna from the earliest Triassic of South China and its implications for benthic recovery after the end-Permian biotic crisis

A single carbonate coquinoid lens from the Griesbachian (Early Triassic) of Shanggan, South China, yielded 11 bivalve species described in this study in addition to four gastropod and one ammonoid species reported elsewhere. This makes the Shanggan fauna one of the richest mollusc faunas from the early post-extinction interval after the end-Permian mass extinction event. Four of the present genera are long-term survivors, five are holdovers that went extinct at the end of the Griesbachian or later in the Early Triassic, and seven first appear in the Griesbachian. Three new bivalve species are described: Myalinella newelli nov. sp., Scythentolium scutigerulus nov. sp., and Eumorphotis shajingengi nov. sp. The genus Astartella, previously assumed to have vanished at the end of the Permian, is reported for the first time from the Early Triassic, which also removes Astartidae from Early Triassic Lazarus taxa. The small growth size of the Astartella specimens supports an earlier hypothesis that many of the Early Triassic Lazarus taxa did not survive in unknown refuges but were simply overlooked due to the scarcity of easily observable large-sized specimens. Ecologically, a comparatively high proportion of infaunal bivalve species (4/11) is remarkable for the early post-extinction interval, supporting the impression of a relatively advanced recovery state. Moreover, abundance-data of the bivalve-gastropod community reveal a remarkably low dominance index (D = 0.17) that is suggestive for advanced recovery and stable environmental conditions. It is proposed that the Shanggan fauna represents a late Griesbachian benthic recovery event that coincided with the appearance of similarly diverse benthic faunas in Oman and Primorye. A high proportion of genera that have previously not been reported from the Early Triassic indicate that the prevalence of poor preservation conditions is a major obstacle in identifying early phases of recovery from the greatest crisis in the history of metazoan life. The early recovery of benthic faunas reported in this study questions previous claims of a prolonged lag phase as a consequence of the extraordinary extinction magnitude or the persistence of adverse environmental conditions.     

Early Triassic conodont clusters from South China: revision of the architecture of the 15 element apparatuses of the superfamily Gondolelloidea

Abstract: Several fused clusters of conodont elements of the genera Neospathodus and Novispathodus were recovered from limestone beds at the Dienerian–Smithian and Smithian–Spathian boundaries, respectively, from several localities in Guangxi province, South China. Conodont clusters are otherwise extremely rare in the Triassic, and these are first described for the Early Triassic. The exceptional specimens partially preserve the relative three‐dimensional position and orientation of ramiform elements and are therefore extremely important for testing hypotheses on the architecture of apparatuses. These specimens partly confirm the previous reconstruction of the Novispathodus apparatus by Orchard. Within apparatuses of members of superfamily Gondolelloidea, elements previously identified as occupying the S₁ and S₂ positions instead occupy the S₂ and S₁ positions. Similarly, within apparatuses of members of the subfamily Novispathodinae, elements previously referred to S₃ and S₄ positions are reinterpreted to have occupied S₄ and S₃ positions, respectively.     

New insights into the base of the Emsian (Lower Devonian) in South China

Traditionally, the lower boundary of the Emsian (Lower Devonian) has been defined as the base of the dehiscens Zone marked by the first entry of the conodont Polygnathus dehiscens. However, following taxonomic revision of P. dehiscens, the current practice is to use the first appearance of P. kitabicus to define the base of the Emsian. As a result, specimens previously identified as P. dehiscens from South China need to be thoroughly reviewed. Careful examination of the oral and aboral features of the platform reveal that most specimens previously described as P. dehiscens from nine sections in Guangxi and Yunnan actually refer to P. excavatus excavatus or P. excavatus ssp. 114. In the South China Block, widespread and massive deposition of Devonian carbonates began during the time of the excavatus Zone, whereas the kitabicus Zone is probably located in the underlying strata, which mainly comprise marine siliciclastic rocks. Thus, the precise level of the lower boundary of the Emsian in South China is still unknown.     

New evidence for laurasian corystosperms: Umkomasia from the Upper Triassic of Northern China

Recent finds of remarkable fossil plants from the Upper Triassic Yangcaogou Formation in Liaoning Province, PR China include branched, cupule-bearing structures referable to the corystosperm ovulate organ Umkomasia. This material is described and assigned to the proposed new species Umkomasia asiatica. The collection includes numerous isolated cupules and fragments of ultimate cupule-bearing axes. Two specimens consisting of portions of the main axis with attached, cupulate lateral axes have also been found. The main axis was at least 6.5 cm long, with each lateral axis bearing one to at least three pairs of stalked, ovoid cupules. The new Umkomasia is similar to U. franconica from the Jurassic of Germany, which is the only other known laurasian species, but the cupules are smaller and more elongated. It is also similar to many gondwanan forms, including the type species U. macleanii. Leaves associated with the Chinese Umkomasia species are tentatively referred to Thinnfeldia, and may have been produced by the same plant. Associated ovoid seeds with elongated, curved micropyles are similar to those of gondwanan species of Umkomasia. The fossils described here are, therefore, significant in representing the first report of corystosperm reproductive structures from Asia, and only the second report of Umkomasia from the entire northern hemisphere. The new Chinese fossils also support leaf-based evidence that the Corystospermales were present in Laurasia as early as the Late Triassic.     

New Genus and Species of Chirotheroid Tracks in the Detfurth-Formation (Middle Bunter,Lower Triassic) of Central Germany

Archosaur tracks are described that have been found in 1999 in a single track bed in the Dethfurth-Formation of the Middle Bunter (Lower Triassic) near Wolfhagen (Central Germany). The taxonomic evaluation of these tracks requires the erection of the new genus and species Protochirotherium wolfhagense ichnogen. n. ichosp. n. Apart from this a rearrangement of groups of chirotheroid tracks is proposed due to the digital proportions.     

The skull of a large Lower Triassic ichthyosaur from Spitzbergen and its implications for the origin of the Ichthyosauria

An incomplete ichthyosaur skull from the Lower Triassic (Spathian) Sticky Keep Formation of Spitzbergen is one of the earliest ichthyosaurs known and by far the largest of the most basal ichthyosaurs. The specimen shows an extremely plesiomorphic cheek region of the skull and allows for the first time a confident reconstruction of the grundplan condition of the lower cheek region of ichthyosaurs. It unequivocally demonstrates the presence of a deep emargination of the lower cheek, which is bordered dorsally by the quadratojugal, squamosal and jugal, and therefore a plesiomorphic lack of contact between jugal and quadratojugal. The specimen also allows an evaluation of recent skull reconstructions of several Lower Triassic taxa which probably contain severe misinterpretations. The structure of the temporal and cheek region of the specimen can be most parsimoniously explained by the initial presence of a lower temporal fenestra. This provides important new evidence that the ancestry of ichthyosaurs might have lain within the Diapsida or their stem-group.     

New data from Oman indicate benthic high biomass productivity coupled with low taxonomic diversity in the aftermath of the Permian–Triassic Boundary mass extinction

A new Early Triassic marine fauna is described from an exotic block (olistolith) from the Ad Daffah conglomerate in eastern Oman (Batain), which provides new insights into the ecology and diversity during the early aftermath of the Permian–Triassic Boundary mass extinction. Based on conodont quantitative biochronology, we assign a middle Griesbachian age to the upper part of this boulder. It was derived from an offshore seamount and yielded both nektonic and benthic faunas, including conodonts, ammonoids, gastropods and crinoid ossicles in mass abundance. This demonstrates that despite the stratigraphically near extinction at the Permian–Triassic Boundary, Crinoidea produced enough biomass to form crinoidal limestone as early as middle Griesbachian time. Baudicrinus, previously placed in Dadocrinidae, is now placed in Holocrinidae; therefore, Dadocrinidae are absent in the Early Triassic, and Holocrinidae remains the most basal crown‐group articulates, originating during the middle Griesbachian in the Tethyan Realm. Abundant gastropods assigned to Naticopsis reached a shell size larger than 20 mm and provide another example against any generalized Lilliput effect during the Griesbachian. Whereas the benthic biomass was as high as to allow the resumption of small carbonate factories, the taxonomic diversity of the benthos remained low compared to post‐Early Triassic times. This slow benthic taxonomic recovery is here attributed to low competition within impoverished post‐extinction faunas.     

The First Articulated Trematosaur 'amphibian' from the Lower Triassic of Madagascar: Implications for the Phylogeny of the Group

The temnospondyl Wantzosaurus elongatus Lehman, 1961, from the Lower Triassic of Madagascar, is redescribed on the basis of a new specimen, which is the most complete trematosaur ever found. Detailed osteological observations and comparisons provide new data on the anatomy, ontogeny, palaeobiology and palaeoecology of this peculiar marine 'amphibian'. The morphology of this aquatically readapted taxon is compared to that of marine 'reptiles': Wantzosaurus might have been able to swim by undulation. A phylogenetic analysis of the trematosaurs is performed for the first time and suggests that Wantzosaurus is a derived taxon within the clade Trematosauridae. The family is defined on the basis of derived character states and is shown to be monophyletic.     

四川江油渔洞子下三叠统飞仙关组巨鲕灰岩中疑似蓝细菌及早三叠世疑似蓝细菌的双幕式爆发

二叠纪末海洋生态环境的恶化导致海洋底栖生物的大灭绝及早三叠世蓝细菌的爆发,有关这一时期我国华南蓝细菌化石的报道主要来自早三叠世早期的微生物岩。四川江油渔洞子剖面下三叠统飞仙关组下部巨鲕灰岩中首次发现有丰富的疑似蓝细菌,在种类和结构上与以往所报道的蓝细菌有着明显区别。疑似蓝细菌在巨鲕内和围岩中的富集,表明巨鲕灰岩的成因与疑似蓝细菌的活动有关,与飞仙关组底部微生物岩在成因上有着密切联系,显示早三叠世早期在这一地区发生了疑似蓝细菌的双幕式爆发,同时,暗示着这一地区二叠纪-三叠纪之交至少发生了两次海洋环境恶化及动物灭绝事件。疑似蓝细菌的爆发性生长,对于海洋生态环境的修复和海洋含氧条件的改善,进而为早三叠世末、中三叠世的生物全面复苏、辐射有着重要意义。     

A New Lower Devonian Genus from Yunnan,with Notes on the Origgin of Leaf

A new genus and species, Eophyllophyton bellum, is described from the Posongchong Formation (Siegenian) of the Lower Devonian in Wenshan cfistrict, Yunnan, China This plant bears lateral branching laminate structures interpreted as leaves, which are believed to derive from lateral axial branching systems In addition similarly organised sporophylls are considered homologous to these leaves. In the possession of these vegetative leaves and sporophylls Eophyllophyton, is significantly more advanced than any other Siegenian taxon.