GUODUNITES, A LOW-PALAEOLATITUDE AND TRANS-PANTHALASSIC SMITHIAN (EARLY TRIASSIC) AMMONOID GENUS

Abstract:  Based on new, bed-rock controlled material from Oman and Utah, USA, the Early Triassic genus Guodunites , which was recently erected on the basis of scarce specimens from northwestern Guangxi, South China, is now shown to be a representative of Proptychitidae. This solves the question of the previously unknown phylogenetic affinity of this genus. The genus is restricted to the late middle Smithian, and to date, its biogeographical distribution comprises Oman, South China and Utah, thus indicating an essentially low palaeolatitudinal distribution during the Early Triassic. Its palaeobiogeographical distribution further strengthens the existence of significant equatorial faunal exchanges between both sides of the Panthalassa at that time. It also suggests that, in addition to the potential stepping stones represented by Panthalassic terranes, vigorous equatorial oceanic currents must have contributed largely to the dispersal of ammonoids during such time intervals.     

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.     

Pangea Megasequences of Tethyan Gondwana-margin reflect global changes of climate and tectonism in Late Palaeozoic and Early Triassic times—A review

The maximum concentration of continental crust at the Pangea stage is characterized by a specific depositional sequence generally referred to as the Pangea Megasequence. Extending in time from the Late Carboniferous to the middle of the Triassic, the succession exhibits similar trends across the whole of Gondwana. Invariably, the sequence was initiated by Late Carboniferous to Early Permian glacial and periglacial deposits. Deglaciation occurred in Early Sakmarian time, evidenced by a typical, commonly transgressive facies. The succeeding formations comprise, in ascending order, coal measures, redbeds, some more coal measures and again redbeds with an intercalation of fluviatile sands in the Early Triassic.After deglaciation the basic depositional theme was modified, depending on postglacial adjustments of climate and on the type of regional tectonic regimes. Extension of the tropical climatic belt after deglaciation was one factor that governed the resulting sediment facies. Coal deposition that prevailed in central Gondwana in the Early Permian gave way to dominance of redbeds in the Middle and Late Permian and, in more distal positions, evaporitic deposits were laid down, following deglaciation. Within marine realms, coralline limestones were formed.Within Gondwana the depositional period of the Pangea Megasequence was governed by three distinctive tectonic regimes: collision dominated the Panthalassa margin, transpressional sag controlled the interior basins, and extension and rifting was experienced along the entire Tethyan margin. In the Early Permian, large and complex graben structures commenced to develop between Africa and India (Malagasy Trough) and between India and Australia (West Australian Trough), giving access to Tethyan waters during deglaciation, commencing in the late Early Sakmarian.Rifting along the Tethyan margin commenced in the Early Permian and was associated with active volcanism between Cashmere and Yunnan and in north-western Australia. Spreading of Neo-Tethys and the formation of oceanic crust, leading to the separation of the Cimmerian Blocks from Gondwana, commenced in the late Early Permian and continued into the Triassic. Thus two facies realms developed, an intracratonic rift facies comprising the Cashmere, Lhasa and Baoshan blocks and a facies controlled by detachment, comprising more distal blocks, such as Tengchong, Malay and Sumatra. The present distribution of individual blocks was governed by fold movements of the Himalayan Orogeny, complicated by transpression along the eastern Himalayan Syntaxis.     

Refinements in the Upper Permian to Lower Jurassic stratigraphy of Karakorum, Pakistan

New sampling on critical intervals of the uppermost Permian and Triassic successions of the Northern Karakorum Terrain in the Karakorum Range (Pakistan) has refined the stratigraphy. Two types of successions may be distinguished in the Karakorum Range: a carbonate platform succession, spanning the whole interval from Upper Permian to Upper Triassic, possibly with several gaps; and a basinal succession, deposited from the Middle Permian to Early Carnian (Late Triassic), when the carbonate platform prograded into the basin. With the approaching and later docking of the Karakorum Block against the Asian margin closing the Paleo-Tethys, a portion of Karakorum emerged while another part subsided as a fore-deep, receiving clastics from the emerging Cimmerian Range. Molassic sediments filled the basin, whilst shallow-water carbonates transgressed over the emerged carbonate platform sometime between the latest Triassic and the Pliensbachian (Early Jurassic), with Cimmerian deformation occurring to the north. The age control is provided by conodonts, with assemblages of late Wuchiapingian, Changhsingian, Induan (Griesbachian and Dienerian), late Olenekian, early Anisian, late Ladinian, and early Carnian ages, respectively. Some information on the section around the P/T boundary is provided by palynology and isotopic C¹³ values. The dating of the Norian/Rhaetian platform is provided by foraminifers.     

Paleogeographic significance of Early Permian crinoids and blastoids from Oman

Early Permian crinoids and blastoids from Oman show relationship with Late Paleozoic Tethyan faunas of Timor and Western Australia and support an Early Permian age for part of the Basleo fauna of Timor. The camerate crinoidPlatycrinites omanensis n. sp. and the blastoidsTimoroblastus andDeltoblastus are reported for the first time from Sakmarian strata of northeastern Oman, doubling the known Permian echinoderms from Oman. The blastoids suggest an offshore lower energy shelf environment of deposition.      

The Origin and Early Radiation of Archosauriforms: Integrating the Skeletal and Footprint Record

We present a holistic approach to the study of early archosauriform evolution by integrating body and track records. The ichnological record supports a Late Permian–Early Triassic radiation of archosauriforms not well documented by skeletal material, and new footprints from the Upper Permian of the southern Alps (Italy) provide evidence for a diversity not yet sampled by body fossils. The integrative study of body fossil and footprint data supports the hypothesis that archosauriforms had already undergone substantial taxonomic diversification by the Late Permian and that by the Early Triassic archosauromorphs attained a broad geographical distribution over most parts of Pangea. Analysis of body size, as deduced from track size, suggests that archosauriform average body size did not change significantly from the Late Permian to the Early Triassic. A survey of facies yielding both skeletal and track record indicate an ecological preference for inland fluvial (lacustrine) environments for early archosauromorphs. Finally, although more data is needed, Late Permian chirotheriid imprints suggest a shift from sprawling to erect posture in archosauriforms before the end-Permian mass extinction event. We highlight the importance of approaching palaeobiological questions by using all available sources of data, specifically through integrating the body and track fossil record.     

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.     

Reinvestigation of conchostracans (Crustacea: Branchiopoda) from the Permian–Triassic transition in Southwest China

Sedimentary deposits of the Permian–Triassic transition are well-exposed in numerous outcrops of South China. Depending on the palaeogeographic positions of the sections, their lithofacies vary from fully marine, shallow marine, lagoonal, lacustrine, and fluvial to alluvial. In the present study, conchostracans (Crustacea: Branchiopoda) were newly collected from the continental deposits of the Kayitou and Jialingjiang formations around the Kangdian Highland elevated by the Emeishan Basalt in the southern Sichuan, western Guizhou, and northeastern Yunnan provinces. The conchostracan fauna of the Kayitou Formation is composed of Pseudestheria chatangensis, Euestheria fuyuanensis, and Euestheria sp. aff. E. gutta. These species occur in varying lithofacies types of different sections. In particular, the late Permian Pseudestheria chatangensis occurs in grey siltstones interbedded with pebbly sandstones, which are interpreted as lacustrine deposits. Euestheria fuyuanensis and Euestheria sp. aff. E. gutta were obtained from yellowish to greenish–grey clay- and siltstones, interpreted as coastal plain deposits. In comparison to other regions, occurrences of Euestheria gutta are indicative of an early Induan to Olenekian (Early Triassic) age. The fossil record of Euestheria fuyuanensis is so far restricted to a few occurrences in the Kayitou Formation of Southwest China and, therefore, using this species for long-distance biostratigraphic correlation requires further investigation. The distribution of late Permian pseudestheriid and Early Triassic euestheriid species in the respective sections possibly depends on the local lithofacies and, therefore, a diachronous age of the Kayitou Formation within the study area can be assumed. Additionally, Magniestheria sp. aff. M. mangaliensis and Magniestheria sp. aff. M. subcircularis were recorded in the Jialingjiang Formation, which represents a lithostratigraphic unit considered as late Early Triassic (Olenekian). Further investigations of both taxonomy and the real stratigraphic ranges of the conchostracan species as well as cross-correlations with other age data are recommended, in order to better constrain the position of the Permian–Triassic boundary and the specific timing of the terrestrial end-Permian mass extinction in continental deposits of Southwest China.     

Early Permian brachiopod faunas from Peninsular India: Their Gondwanan relationships

The Early Permian (Late Asselian? to Aktastinian?) brachiopod faunas of Peninsular India are revised in terms of current taxonomy. Genera such as Semilingula, Arctitreta, Etherilosia, Strophalosia, Aulosteges, Bandoproductus, Cyrtella, Neospirifer, Crassispirifer, Tomiopsis and Gilledia confirm the Gondwanan aspect of the faunas and a close relationship, at the generic level, to the Early Permian brachiopod faunas of Western Australia. Peninsular Indian Early Permian brachiopod faunas belong to a complex of Gondwanan and peri‐Gondwanan faunas from Oman and the Pamirs in the West to Australasia in the east. This distribution implies relative freedom of migration for the faunas along the northern margin of Gondwana during the Early Permian.     

波环孢属(Sinulatis porites)在华北地区的时空分布及其地层学意义

对鲁西晚古生代含煤岩系以及华北各地石炭、二叠系孢粉组合中已经发现的Sinulatisporites一属的时空分布进行详细研究,结果表明：其时空分布仅限于华夏植物区的北方亚区,在时间上始于早二叠世早期(太原组),灭绝于中二叠世中、晚期,可视为鲁西乃至华北地区早二叠世晚期、中二叠世早期中期华夏植物区北方亚区的典型分子。其母体植物可能为一种喜湿热的真蕨植物或种子蕨植物。该属产出层位稳定,延续时间较短,在月门沟统山西组、石盒子统下石盒子组含量丰富,又常产出于月门沟统山西组的主要含煤岩段及其上部石盒子统下石盒子组的砂、泥岩段,故可作为二叠系划分和对比煤层的重要依据。     

Upper Permian coral reef and colonial rugose corals in northwest Hunan, South China

Summary The roles of Permian colonial corals in forming organic reefs have not been adequately assessed, although they are common fossils in the Permian strata. It is now known that colonial corals were important contributors to reef framework during the middle and late Permian such as those in South China, northeast Japan, Oman and Thailand. A coral reef occurs in Kanjia-ping, Cili County, Hunan, South China. It is formed by erect and unscathed colonies ofWaagenophyllum growing on top of one anotherin situ to form a baffle and framework. Paleontological data of the Cili coral reef indicates a middle to late Changhsing age (Late Permian), corresponding to thePalaeofusulina zone. The coral reef exposure extends along the inner platform margin striking in E-S direction for nearly 4 km laterally and generally 35 to 57 m thick. The Cili coral reef exhibits a lateral differentiation into three main reef facies; reef core facies, fore-reef facies, and marginal slope facies. The major reef-core facies is well exposed in Shenxian-wan and Guanyin-an sections where it rests on the marginal slope facies. Colonial corals are dispersed and preserved in non-living position easward. Sponges become major stabilizing organisms in the eastern part of Changhsing limestone outcrop in Kanjia-ping, but no read sponge reefs were formed. Coral reefs at Cili County in Human are different distinctly from calcisponge reefs in South China in their palaeogeography, lithofacies development, organic constitutuents, palaeoecology and diagenesis. The Cili coral reef also shows differences in age, depositional facies association, reef organisms and diagenesis from coral reefs in South Kitakami of Japan, Khorat Plateau of Thailand, and Saih Hatat of Oman. Although some sponge reefs and mounds can reach up to the unconformable Permian/Triassic boundary, coral reef at Kanjia-ping, Cili County, is the latest Permian reef known. This reef appears to had been formed in a palaeoenvironment that is different from that of the sponge reefs and provides an example of new and unique Permian reef type in South China, and could help us to: 1) understand the significance of colonial corals in Permian carbonate buildups; 2) evaluate the importance of coral community evolution prior to the collapse of reef ecosystems at the Permian/Triassic boundary; 3) better understand the effects of the biotic extinction events in Palaeotethys realm; 4) look for environmental factors that may have controlled reefs through time and space, and 5) provide valuable data for the study of Permian palaeoclimate and global evolutionary changes of Permian reefs and reef community.     

Middle and Late Triassic radiolarians from northern Tibet: Implications for the Bayan Har Basin evolution

During the Triassic, the Bayan Har Basin is a huge triangular basin surrounded by the North China Platform, South China Platform and Qingtang Terrane. It is filled by a Triassic turbidite sequence, the Bayan Har Group. For a long time, the series of Bayan Har Group in the eastern part of the basin were considered to be a Lower to Upper Triassic sequence, and in the western part, was attributed to the Upper Triassic. A well-preserved diversified radiolarian fauna was recovered from radiolarian chert and tuffite interbeds of the Bayan Har Group turbidites and adjacent stratigraphic units in the Hoh Xil area, northern Tibet. Sixty-seven species are identified and subdivided into two assemblages: late Anisian and early Carnian. Combined with the discovery of the Late Permian and Early Triassic turbidite in the Bayan Har Group in this area by Huang et al., it proves that all the Triassic is also present in the Bayan Har Group sequence in the western part. The evolution of the Bayan Har Basin may be traced back to the Late Permian. The massive sequence of the Bayan Har Group and its provenance indicate that the Kunlun and Qinling orogenic belts rapidly rise during the Middle-Late Triassic. The basin extended to the end of the Triassic, possibly locally to the Jurassic.     

Late Paleozoic–early Mesozoic continental biostratigraphy — Links to the Standard Global Chronostratigraphic Scale

Nonmarine biostratigraphic/biochronologic schemes have been created for all or parts of the late Carboniferous–Middle Triassic using palynomorphs, megafossil plants, conchostracans, blattoid insects, tetrapod footprints and tetrapod body fossils, and these provide varied temporal resolution. Cross correlation of the nonmarine biochronologies to the Standard Global Chronostratigraphic Scale has been achieved in some parts of the late Carboniferous–Middle Triassic in locations where nonmarine and marine strata are intercalated, the nonmarine strata produce biochronologically significant fossils and the marine strata yield fusulinids, conodonts and/or ammonoids. Other cross correlations have been aided by magnetostratigraphy, chemostratigraphy and a growing database of radioisotopic ages. A synthetic nonmarine biochronology for the late Carboniferous–Middle Triassic based on all available nonmarine index fossils, integrated with the Standard Global Chronostratigraphic Scale, is presented here. The focus is on the nonmarine biostratigraphy/biochronology of blattoid insects, conchostracans, branchiosaurid amphibians, tetrapod footprints and tetrapod body fossils within the biochronological framework of land-vertebrate faunachrons. Correlation to the Standard Global Chronostratigraphic Scale presented here is divided into seven time intervals: Pennsylvanian, Carboniferous–Permian boundary, Cisuralian, Guadalupian, Lopingian, Permian–Triassic boundary and Early to Middle Triassic. The insects, conchostracans and branchiosaurs provide robust nonmarine correlations in the Pennsylvanian–Cisuralian, and the footprints and tetrapod body fossils provide robust correlations of varied precision within the entire Pennsylvanian–Middle Triassic. Radioisotopic ages are currently the strongest basis for cross correlation of the nonmarine biostratigraphy/biochronology to the Standard Global Chronostratigraphic Scale, particularly for the Pennsylvanian–Cisuralian. Chemostratigraphy and magnetostratigraphy thus far provide only limited links of nomarine and marine chronologies. Improvements in the nonmarine-marine correlations of late Paleozoic–Triassic Pangea require better alpha taxonomy and stratigraphic precision for the nonmarine fossil record integrated with more reliable radioisotopic ages and more extensive chemostratigraphic and magnetostratigraphic datasets.     

Palaeoecological significance of a new Griesbachian (Early Triassic) gastropod assemblage from Oman

A new Early Triassic (Griesbachian) gastropod fauna from the Al Jil Formation of Oman is described. Early Triassic faunas from elsewhere (e.g. the Italian Dolomites and the western USA) are typically of low diversity and high dominance, usually attributed to environmental stress in the immediate aftermath of the end-Permian mass extinction event. The new Oman fauna has, by contrast, relatively high diversity, low dominance and a more even spread of individuals between taxa. It is the most diverse Griesbachian fauna known to date. This is attributed to the favourable (i.e. well-oxygenated) conditions under which the fauna lived. This uncharacteristic Griesbachian gastropod fauna demonstrates that, in the absence of oceanic anoxia, biotic recovery after the end-Permian extinction event may occur surprisingly quickly (within one conodont zone). The fauna is also partially silicified, which has increased its preservation potential relative to other Griesbachian gastropod assemblages. However, only one reappearing Lazarus genus is present in the Oman fauna. This suggests that there was some other control on the abundance of Lazarus genera at this time, other than the absence of silicified faunas as previously suggested.     

西藏申扎地区中二叠世(竹蜓)类动物群

结合新采集的材料与前人研究,综合分析拉萨地块申扎永珠乡下拉组中类动物群的构成与时代,并将其与同期华南类动物群进行对比。分析结果表明,申扎下拉组的类动物群共16属(含1亚属,2存疑属),70种(含比较种与未定种),时代为茅口期,大致相当于Roadian阶上部至Wordian阶。与同期华夏区华南的类动物群相比,下拉组的类动物群在属种构成,尤其是优势类群方面差异显著,其主要特征为Neoschwagerinidae与Verbeekinidae两科的种级分异度很低,Chusenella与Nankinella占优势地位,且缺失了约60%华南的常见属。     

Evolution of the Permian and Triassic tetrapod communities of Eastern Europe

The Permo-Triassic terrestrial and freshwater tetrapod communities of Eastern Europe are reconstructed as food-webs. The Late Permian theriodont-dinocephalian community (Ocher, Mezen, Isheyevo) changes to a latest Permian theriodont-pareiasaur community (North Dvina, Vyazniki). After a major extinction, the Triassic thecodontian-dicynodont communities appear, a lystrosaurid one in the Early Triassic (Lower and ?Upper Vetluga), and a kannemeyerid one in the later Early Triassic (?Yarenga) and the Mid Triassic (Donguz, Bukobay). Similar stages are represented in the evolution of aquatic communities: the Late Permian temnospondyl community (Ocher, Isheyevo), the latest Permian chroniosuchian one (North Dvina, Vyazniki), the Lower and Middle Triassic new temnospondyl one (from Vetluga to Bukobay). The faunal changes in Eastern Europe are mirrored in other parts of the world, although there are some endemic Russian forms.     

A new holocrinid (Articulata) from the Paris Biota (Bear Lake County,Idaho, USA) highlights the high diversity of Early Triassic crinoids

After the end-Permian crisis and the extinction of their four Paleozoic subclasses, crinoids rapidly recovered. This group is classically believed to have radiated from a small surviving clade and to have diversified during the Middle and Upper Triassic from two lineages. Nevertheless, recent findings suggested that several lineages of crinoids had already diversified during the Early Triassic, and that their diversity has been overlooked. Here we describe a new form of holocrinid, Holocrinus nov. sp., from the earliest Spathian (Early Triassic) of southeastern Idaho (USA). So far, the exceptional completeness of sampled specimens, with skeletal elements of arms and stem in connection, is unique for the Early Triassic. They show that derived morphological features had already evolved ∼1.3 million years after the Permian–Triassic boundary, supporting the scenario of a rapid Early Triassic diversification of crinoids.     

Annalepis, a pioneering lycopsid genus in the recovery of the Triassic land flora in South China

Fossil plants are scarce in the Earliest Triassic marine deposits of western Guizhou and eastern Yunnan. Only Annularia shirakii, Lobatannularia sp., Paracalamites stenocostatus, Gigantopteris sp., Pecopteris sp. were reported from the base of the Kayitou Formation dated as Early Induan by marine fauna. Recently, we discovered numerous representatives of the genus Annalepis in the same Lowermost Triassic beds: A. latiloba, A. brevicystis, A. angusta, Annalepis spp. occur associated with a basal Triassic marine fauna. This discovery fills the biostratigraphic gap between the Late Permian “Gigantonoclea guizhouensis-Ullmannia cf. bronnii-Annularia pingloensis” and the late Lower Triassic “Neuropteridium–Albertia–Voltzia” assemblages reported from South China. It represents an important datum dealing with the very beginning of a new terrestrial flora installation after the Permian flora disappearance following the Permian–Triassic boundary mass extinction. This “starting point” of a new vegetal cover in South China is to be taken into account in reconstructing through space and time the settlement process of the Mesozoic floristic provinces.     

昆虫口器及其进化简史

昆虫口器有着很高的分化率,可细分为34个现生和2个化石类型,大部分出现在自泥盆纪以来的5个历史发展时期:早泥盆、中石炭、早二叠、三叠-侏罗、侏罗-白垩。文章依据时间线索,介绍每种口器的特征以及所属主要类群,对整个昆虫纲的口器做出了归类和描述。