Permian–Triassic Osteichthyes (bony fishes): diversity dynamics and body size evolution

The Permian and Triassic were key time intervals in the history of life on Earth. Both periods are marked by a series of biotic crises including the most catastrophic of such events, the end‐Permian mass extinction, which eventually led to a major turnover from typical Palaeozoic faunas and floras to those that are emblematic for the Mesozoic and Cenozoic. Here we review patterns in Permian–Triassic bony fishes, a group whose evolutionary dynamics are understudied. Based on data from primary literature, we analyse changes in their taxonomic diversity and body size (as a proxy for trophic position) and explore their response to Permian–Triassic events. Diversity and body size are investigated separately for different groups of Osteichthyes (Dipnoi, Actinistia, ‘Palaeopterygii’, ‘Subholostei’, Holostei, Teleosteomorpha), within the marine and freshwater realms and on a global scale (total diversity) as well as across palaeolatitudinal belts. Diversity is also measured for different palaeogeographical provinces. Our results suggest a general trend from low osteichthyan diversity in the Permian to higher levels in the Triassic. Diversity dynamics in the Permian are marked by a decline in freshwater taxa during the Cisuralian. An extinction event during the end‐Guadalupian crisis is not evident from our data, but ‘palaeopterygians’ experienced a significant body size increase across the Guadalupian–Lopingian boundary and these fishes upheld their position as large, top predators from the Late Permian to the Late Triassic. Elevated turnover rates are documented at the Permian–Triassic boundary, and two distinct diversification events are noted in the wake of this biotic crisis, a first one during the Early Triassic (dipnoans, actinistians, ‘palaeopterygians’, ‘subholosteans’) and a second one during the Middle Triassic (‘subholosteans’, neopterygians). The origination of new, small taxa predominantly among these groups during the Middle Triassic event caused a significant reduction in osteichthyan body size. Neopterygii, the clade that encompasses the vast majority of extant fishes, underwent another diversification phase in the Late Triassic. The Triassic radiation of Osteichthyes, predominantly of Actinopterygii, which only occurred after severe extinctions among Chondrichthyes during the Middle–Late Permian, resulted in a profound change within global fish communities, from chondrichthyan‐rich faunas of the Permo‐Carboniferous to typical Mesozoic and Cenozoic associations dominated by actinopterygians. This turnover was not sudden but followed a stepwise pattern, with leaps during extinction events.     

The Permian–Triassic transition and the onset of Mesozoic sedimentation at the northwestern peri-Tethyan domain scale: Palaeogeographic maps and geodynamic implications

The main aim of this paper is to review Middle Permian through Middle Triassic continental successions in European. Secondly, areas of Middle–Late Permian sedimentation, the Permian–Triassic Boundary (PTB) and the onset of Triassic sedimentation at the scale of the westernmost peri-Tethyan domain are defined in order to construct palaeogeographic maps of the area and to discuss the impact of tectonics, climate and sediment supply on the preservation of continental sediment.At the scale of the western European peri-Tethyan basins, the Upper Permian is characterised by a general progradational pattern from playa-lake or floodplain to fluvial environments. In the northern Variscan Belt domain, areas of sedimentation were either isolated or connected to the large basin, which was occupied by the Zechstein Sea. In the southern Variscan Belt, during the Late Permian, either isolated endoreic basins occurred, with palaeocurrent directions indicating local sources, or basins underwent erosion and/or there was no deposition. These basins were not connected with the Tethys Ocean, which could be explained by a high border formed by Corsica–Sardinia palaeorelief and even parts of the Kabilia microplate. The palaeoflora and sedimentary environments suggest warm and semi-arid climatic conditions.At the scale of the whole study area, an unconformity (more or less angular) is observed almost everywhere between deposits of the Upper Permian and Triassic, except in the central part of the Germanic Basin. The sedimentation gap is more developed in the southern area where, in some basins, Upper Permian sediment does not occur. The large sedimentary supply, erosion and/or lack of deposition during the Late Permian, as well as the variable palaeocurrent direction pattern between the Middle–Late Permian and the Early Triassic indicate a period of relief rejuvenation during the Late Permian. During the Induan, all the intra-belt basins were under erosion and sediment was only preserved in the extra-belt domains (the northern and extreme southern domains). In the northern domain (the central part of the Germanic Basin), sediment was preserved under the same climatic conditions as during the latest Permian, whereas in the extreme southern domain, it was probably preserved in the Tethys Ocean, implying a large amount of detrital components entering the marine waters. Mesozoic sedimentation began in the early Olenekian; the ephemeral fluvial systems indicate arid climatic conditions during this period. Three distinct areas of sedimentation occur: a northern and southern domain, separated by an intra-belt domain. The latter accumulated sediments during the Early–Middle Permian and experienced erosion and/or no-deposition conditions between the Middle–Late Permian and the beginning of Mesozoic sedimentation, dated as Anisian to Hettangian. At the top of the Lower Triassic, another tectonically induced, more or less angular unconformity is observed: the Hardegsen unconformity, which is dated as intra-Spathian and is especially found in the North European basins. This tectonic activity created new source areas and a new fluvial style, with marine influences at the distal part of the systems. During the Anisian and Ladinian, continental sedimentation was characterised by a retrogradational trend. In other words, the fluvial system evolved into fluvio-marine environments, attesting to a direct influence of the Tethys Ocean in the southern and northern domains. Both at the end of the Olenekian (Spathian) and during the Anisian, the presence of palaeosols, micro- and macrofloras indicate less arid conditions throughout this domain.     

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.     

Palaeoecological and phylogenetic implications of a new scleractiniamorph genus from Permian sponge reefs,south China

Scleractinian corals are the most important constituents of modern coralgal reefs. For many years, it was thought that they first appeared in the Middle Triassic and subsequently underwent explosive radiation. However, abundant scleractinian-like corals with ancestral morphological traits have recently been recovered from Middle Permian sponge reefs in China, which not only confirms a role in Permian reef ecology but also suggests a possible Palaeozoic origin for the group. Two species of a new Permian scleractiniamorph genus from China are described herein as Houchangocyathus wangi gen. et sp. nov. and Houchangocyathus yaoi gen. et sp. nov. Putative Palaeozoic Scleractinia may have evolved over a substantial time interval and diverged into stem lineages by the end of the Permian. These forms evolved within both the rigid framework of their basic body plan and the morphological constraints characteristic of each lineage. The Middle Permian development of calcisponge reefs was closely related to habitat expansion, which would have provided an ideal dwelling for scleractinian-like corals and enhanced their chances of fossilization. Such scleractiniamorphs disappeared at the end-Permian extinction, but may have survived as progenitors of Triassic Scleractinia.     

Rooting down the aphid's tree – the oldest record of the Aphidomorpha lineage from Palaeozoic (Insecta: Hemiptera)

The new genus and species Lutevanaphis permiana gen.n. et sp.n. , represents the oldest Aphidomorpha, new superfamily Lutevanaphidoidea superfam.n. and new family Lutevanaphididae. This taxon is described from the Middle Permian of the Lodève Basin, southern France. The presence of very small Aphidomorpha in the Middle Permian contradicts the hypothesis of the ‘Lilliput effect’ that is presumed to have affected the insect fauna during the Early to Middle Triassic, after the end‐Permian crisis. Very small insects were present well before the end of the Permian; their relative rarity is attributable probably to taphonomic biases.     

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.     

Why was there a delayed radiation after the end‐Palaeozoic extinctions?

Data from widespread dysaerobic facies, carbon/sulphur ratios and cerium anomalies suggest that the early Triassic was a time when anoxic conditions spread widely over epicontinental seas. These conditions, associated with marine transgression following the latest Permian regression, are likely to be a prime cause of the mass extinction of Palaeozoic marine faunas. The occurrence of many Lazarus taxa in the Middle and Upper Triassic indicates, however, that the extinctions at the end of the Permian were less severe than has been widely assumed, and that the turnover from Palaeozoic to Mesozoic faunas was considerably extended in time, being finally accomplished only after the end‐Triassic mass extinction event.     

Latest Permian Spumellaria and Entactinaria (Radiolaria) from South China

A diverse and well-preserved latest Permian radiolarian fauna was obtained from bedded siliceous rocks and siliceous mudstones of the Dongpan Section, southwest Guangxi, China. All encountered specimens of the order Spumellaria and a few specimens of the order Entactinaria were selected for the taxonomic study of this paper. Thirty-three species, 15 of which are new species, were identified. They belong to 15 genera, three of which (Paracopicyntra, Copiconulus, and Yujingella) are new. The most interesting result of our study is that eight typical Middle Triassic spumellarian genera (Paroertlispongus Kozur and Mostler, Tetrapaurinella Kozur and Mostler, Paurinella Kozur and Mostler, Lithelius Haeckel, Archaeospongoprunum Pessagno, Tamonella Dumitrica, Kozur and Mostler, Bistarkum Yeh, and Tetraspongodiscus Kozur and Mostler) were discovered in the upper Changhsingian strata studied herein. Therefore, the most important and characteristic spumellarian families of the Middle Triassic were already present in the Late Permian of South China.     

An investigation into the cladistic relationships and monophyly of therocephalian therapsids (Amniota: Synapsida)

A comprehensive phylogenetic investigation was performed to elucidate the cladistic relationships and possible monophyly of therocephalian therapsids (Amniota: Synapsida). The phylogenetic positions of 30 therapsid taxa were examined under maximum parsimony, including 23 therocephalian genera. The analysis incorporated 110 cranial and postcranial characters in order to assess the interrelationships of basal therocephalians and eutherocephalians and their relationships to Cynodontia, representing the most complete review of therocephalian phylogeny to date. The analysis supports the hypothesis that Therocephalia represents the monophyletic sister taxon to Cynodontia, with as many as 15 morphological synapomorphies, in contrast with other recent analyses of lesser taxon sampling. The results also support the hypothesis that Scylacosauridae is more closely related to Eutherocephalia than to the basal therocephalian family Lycosuchidae, supporting a ‘Scylacosauria’ clade. The taxa suggested here to be neotenic forms (e.g. Ictidosuchoides and Ictidosuchops) are positioned near the base of a monophyletic Baurioidea. Neotenic development of the therocephalian feeding apparatus and evolutionary parallelism with cynodonts are suggested to have been important trends in the early evolution of baurioid therocephalians into the Late Permian and Early Triassic.     

Proharpoceras Chao: a new ammonoid lineage surviving the end-Permian mass extinction

Based on new, bed-rock controlled material from northwestern Guangxi and Oman, the Early Triassic genus Proharpoceras Chao is shown to be a representative of Otocerataceae. Character analysis excludes a direct link with the Griesbachian Otoceratidae and favours a derivation of Proharpoceras from the late Permian Anderssonoceratidae. The biostratigraphic range of Proharpoceras is restricted to the Smithian and its biogeographic distribution comprises Oman, South China, and Primorye, thus indicating an essentially low palaeolatitudinal distribution. Proharpoceras has no apparent relatives among other Early and Middle Triassic Ceratitida and is thus considered to be the last representative of Otocerataceae. This offshoot of the late Permian Anderssonoceratidae implies that an additional ammonoid lineage survived the end Permian extinction and that it dwindled away for some 2 Myr before going extinct.     

A Late Permian flora with Dicroidium from the Dead Sea region, Jordan

Three new species of Dicroidium, D. irnensis, D. jordanensis and D. robustum, are described from the Um Irna Formation (Upper Permian) of the Dead Sea region, Jordan. The plant remains are preserved as compressions with excellent cuticles. These are the earliest unequivocal records of Dicroidium, a genus that is typical for the Triassic of Gondwana. It is also the northernmost occurrence of this genus that apparently originated in the Permian in the palaeotropics. Middle and Late Permian floras from the Arabian Peninsula and adjacent regions show a remarkable mixture of elements from different floral provinces, i.e. Euramerica, Cathaysia and Gondwana. The climatic amelioration in the Early Triassic apparently enabled Dicroidium to migrate southward and eventually colonise the entire Gondwana region. Dicroidium is one of the very few megaplant genera not affected by the end-Permian biotic crisis, the largest Phanerozoic extinction event.     

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.     

Phanerozoic changes in the high-rank suprageneric diversity structure of brachiopods: Linear and non-linear effects

Phanerozoic evolution of brachiopods produced many linear (established by a comparison of successive geologic time units) and non-linear (established by a comparison of non-successive geologic time units) effects, which can be examined quantitatively by using the similarity coefficients (Czekanowski's Quantified Coefficient and Gower Index) and correlation tools. The high-rank suprageneric diversity structure accounts for a number of superfamilies in each of 26 orders for every epoch of geological time. The intensity of turnovers in this structure was generally low during the entire Phanerozoic. It was slightly stronger during the Early Paleozoic, but close to zero during the Cenozoic, when the high-rank suprageneric diversity structure of brachiopods stabilized finally. Significant turnovers took place at the Middle Cambrian–Early Ordovician, the Late Ordovician–Early Silurian, the Late Silurian–Early Devonian, the Middle Devonian–Mississippian, and the Permian–Triassic transitions. Influences of mass extinctions, both major like those End Ordovician or Permian/Triassic and minor like Early Jurassic or Jurassic/Cretaceous, on the high-rank suprageneric diversity structure of brachiopods is registered. The strongest was the consequences of the Permian/Triassic catastrophe, which perhaps even reset the brachiopod evolution. No evident direct relationships are established between intensity of turnovers and eustatic fluctuations. However, the changes in the diversity structure recorded with the Gower Index provide evidence that eustatic lowstands were more favorable for intensification in these changes.     

New and little known blattogryllidae (Insecta: Grylloblattida) from the Permian and Triassic of Russia,Kazakhstan, and Kyrgyzstan

New members of the family Blattogryllidae are described, including Permoblattogryllus praecox gen. et sp. nov. from the Middle Permian Soyana locality in Russia, Protoblattogryllus nedubrovensis from the Lower Triassic Nedubrovo locality in Russia, Madygenocephalus micropteron gen. et sp. nov. (the second known case of brachyptery in grylloblattids), and Costatoviblatta similis sp. nov. from the Middle Triassic Madygen locality in Kyrgyzstan. Protoblattogryllus zajsanicus Storozhenko, 1990 from the Upper Permian Karaungir II locality in Kazakhstan is redescribed. The genus Microblattogryllus Storozhenko, 1990 from Madygen is considered to be a synonym of Protoblattogryllus Storozhenko, 1990. Protoblattogryllus abruptus Storozhenko, 1990 from the Madygen locality is transferred to the genus Mesoblattogryllus Storozhenko, 1990.     

云南墨江上三叠统歪古村组燧石质砾石中的晚古生代放射虫化石及其地质意义

云南哀牢山缝合带由于长期未找到晚石炭世至二叠纪深海环境的化石及沉积地层记录,对哀牢山古特提斯盆地演化历史存在着不同认识。文中报道了采自云南墨江坝留地区上三叠统歪古村组底砾岩中的早石炭世和中二叠世放射虫化石,所有放射虫化石发现于4件燧石质砾石中,共计11属9种和7未定种及1属种未定放射虫。其中,3件砾石含有Albaillella deflandrei Gourmelon,Albaillella sinuosa Won and Seo等早石炭世放射虫化石组合,另1件砾石含有Pseudoalbaillella spp.,Quadricaulis scalae Caridroit and De Wever,Cauletella sp.和Ishigaum sp.等中二叠世放射虫化石组合。由此表明,哀牢山缝合带存在着早石炭世和中二叠世深海盆地沉积地层记录,哀牢山深海盆地应该在中二叠世之后封闭。该成果为探讨哀牢山古特提斯盆地演化提供了放射虫古生物学证据,进而说明哀牢山缝合带与金沙江缝合带一样,也存在石炭纪和二叠纪深水洋盆沉积地层记录,指示其演化历史是相同的。     

New beetles (Insecta,Coleoptera) from the Lower Triassic of European Russia

Fossil remains of beetles are described from two Lower Triassic localities: Entala (Induan) and Tikhvinskoe (Olenekian). Only one beetle fossil was previously known from the Lower Triassic of Tikhvinskoe. The fossils are rather few and poorly preserved, but they are worth describing as finds rare for the Lower Triassic. Five fossils from Entala most probably belong to beetles of the same species of the formal genus Pseudochrysomelites. Beetles of this genus are especially abundant in deposits close to the Permian–Triassic boundary and can be considered “disaster taxa.” There are no known cases, either in the Permian or in the Middle–Upper Triassic, of a random sample of five specimens belonging to a single species. This suggests that in the Entala oryctocenosis the species diversity of beetles is extremely low. All three beetle fossils found in Tikhvinskoe belong to beetles of different species, showing that diversity had already started to increase. However, it remained low, and all fossils belong to the formal family Schizocoleidae, and two of the three belong to the same genus, Pseudochrysomelites. The Khei-Yaga locality, which immediately follows Tikhvinskoe in time (topmost Olenekian or early Anisian), already contains beetles of the families Asiocoleidae and Permosynidae. In the Lower Anisian of the Buntsandstein, such typical Mesozoic beetles as Cupedidae and Coptoclavidae have been recorded. The appearance of such advanced beetles as early as the Lower Anisian suggests that the famous Permian–Triassic crisis was not as deep as it is usually believed, and many beetles survived it, disappearing, however, from the fossil record in the Early Triassic.     

巴基斯坦盐岭地区二叠纪生物地层研究的新进展

近30年来,巴基斯坦盐岭地区的Wargal组和Chhidru组以及在喜马拉雅地区相当地层的地质时代被许多学者视为中二叠世(瓜达鲁普世)。根据华南与盐岭地区瓜达鲁普统和乐平统的牙形类化石带对比,可以确定Amb组和Wargal组底部的时代为瓜达鲁普世晚期,瓜达鲁普统与乐平统的界线位于Wargal组下部,吴家坪阶与长兴阶的界线位于Chhidru组的下部,而二叠与三叠系的界线位于Mianwali组下部的Kathwai段之内。我国西藏南部色龙群及相当地层含有与巴基斯坦盐岭地区Kalabagh段和Chhidru组可对比的乐平世腕足动物群以及二叠系—三叠系界线附近连续的牙形类化石带,由此可以推定色龙群的时代应为乐平世。