A new genus of Rhynchonellid Brachiopod from the Lower Triassic of South China and Implications for timing the recovery of Brachiopoda after the end-Permian mass extinction

A new genus, Meishanorhynchia, is proposed based on new material from the Lower Triassic of the Meishan section, South China. It is of a late Griesbachian age based on both associated biozones (ammonoids and
bivalves) and radiometric dates of the intercalated volcanic ash beds. Comparison with both Palaeozoic and Mesozoic-Cenozoic-related genera suggests that it may represent the first radiation of progenitor brachiopods in the aftermath of the end-Permian extinction. The lowest brachiopod horizon that contains the genus is estimated to be about 250.1±0.3 Ma. This implies that the initial stage of recovery of Brachiopoda in the Early Triassic was probably about 1.3±0.3 myr after the major pulse of the end-Permian mass extinction (dated as 251.4±0.3 Ma). This is in agreement with Hallam's expectancy that biotic recovery typically begins within one million years or so of major mass extinctions, in contrast to current views on the end-Permian extinction event which propose that the recovery of most if not all biotic groups in the Early Triassic was severely delayed and only began about five million years after the end-Permian extinction.

End-Permian mass extinction pattern in the northern peri-Gondwanan region

The Permian-Triassic extinction pattern in the peri-Gondwanan region is documented biostratigraphically, geochemically and sedimentologically based on three marine sequences deposited in southern Tibet and comparisons with the sections in the Salt Range, Pakistan and Kashmir. Results of biostratigraphical ranges for the marine faunas reveal an end-Permian event comparable in timing with that known at the Meishan section in low palaeolatitude as well as Spitsbergen and East Greenland in northern Boreal settings although biotic patterns earlier in the Permian vary. The previously interpreted delayed extinction (Late Griesbachian) at the Selong Xishan section is not supported by our analysis. The end-Permian event exhibits an abrupt marine faunal shift slightly beneath the Permian-Triassic boundary (PTB) from benthic taxa- to nektic taxa-dominated communities. The climate along the continental margin of Neo-Tethys was cold before the extinction event. However, a rapid climatic warming event as indicated by the southward invasion of abundant warm-water conodonts, warm-water brachiopods, calcareous sponges, and gastropods was associated with the extinction event. Stable isotopic values of δ13Ccarb, δ13Corg and δ18O show a sharp negative drop slightly before and during the extinction interval. Sedimentological and microstratigraphical analysis reveals a Late Permian regression, as marked by a Caliche Bed at the Selong Xishan section and the micaceous siltstone in the topmost part of the Qubuerga Formation at the Qubu and Tulong sections. The regression was immediately followed by a rapid transgression beneath the PTB. The basal Triassic rocks fine upward, and are dominated by dolomitic packstone/wackestone containing pyritic cubes, bioturbation and numerous tiny foraminifers, suggesting that the studied sections were deposited during the initial stage of the transgression and hence may not have been deeply affected by the anoxic event that is widely believed to characterise the zenith of the transgression.

Controls on body size during the Late Permian mass extinction event

This study examines the morphological responses of Late Permian brachiopods to environmental changes. Quantitative analysis of body size data from Permian–Triassic brachiopods has demonstrated significant, directional changes in body size before, during and after the Late Permian mass extinction event. Brachiopod size significantly reduced before and during the extinction interval, increased for a short time in more extinction-resistant taxa in the latter stages of extinction and then dramatically reduced again across the Permian ⁄ Triassic boundary. Relative abundances of trace elements and acritarchs demonstrate that the body size reductions which happened before, during and after extinction were driven by primary productivity collapse, whereas declining oxygen levels had less effect. An episode of size increase in two of the more extinction-resistant brachiopod species is unrelated to environmental change and possibly was the result of reduced interspecific competition for resources following the extinction of competitors. Based on the results of this study, predictions can be made for the possible responses of modern benthos to present-day environmental changes.

Systematics and palaeoecology of Changhsingian (Late Permian) Ambocoeliidae brachiopods from South China and implications for the end-Permian mass extinction

Four Ambocoeliidae brachiopod species including one new species (Crurithyris tazawai sp. nov., Crurithyris sp., Paracrurithyris pygmaea and Attenuatella mengi) are described from the Changhsingian (Late Permian) deep-water facies of South China. Analysis of the morphology, palaeoecology and palaeogeographical and temporal distributions of these species revealed that the presence of a delthyrium and/or the micro-ornaments among three of the four species (Crurithyris tazawai sp. nov., Paracrurithyris pygmaea and Attenuatella mengi) favoured an epifaunal (epiphytic) lifestyle. Morphological differences suggest that Paracrurithyris pygmaea may have been more effective metabolically in forming the shell compared with Attenuatella mengi and Crurithyris tazawai. The temporal and palaeogeographical distribution of Attenuatella suggests that A. mengi inhabited cool or cold deep waters. Both Crurithyris tazawai and Attenuatella mengi disappeared earlier in the stratigraphic record than Paracrurithyis pygmaea during the Permian–Triassic mass extinction. These differences in timing of extinction, morphology and palaeogeographical distributions suggest that oxygen deficiency and trophic resource limitation (a consequence of the changing composition of marine phytoplankton in the seas) may have contributed to the end-Permian mass extinction.

Occurrence and Range-through database of functional diversity of marine ecosystems after the Late Permian mass extinction event

The Late Permian mass extinction event about 252 million years ago was the most severe biotic crisis of the past 500 million years and occurred during an episode of global warming. The loss of around two-thirds of marine genera is thought to have had substantial ecological effects, but the overall impacts on the functioning of marine ecosystems and the pattern of marine recovery are uncertain. Here we analyse the fossil occurrences of all known benthic marine invertebrate genera from the Permian and Triassic periods, and assign each to a functional group based on their inferred lifestyle. We show that despite the selective extinction of 62-74% of these genera, all but one functional group persisted through the crisis, indicating that there was no significant loss of functional diversity at the global scale. In addition, only one new mode of life originated in the extinction aftermath. We suggest that Early Triassic marine ecosystems were not as ecologically depauperate as widely assumed. Functional diversity was, however, reduced in particular regions and habitats, such as tropical reefs; at these smaller scales, recovery varied spatially and temporally, probably driven by migration of surviving groups. We find that marine ecosystems did not return to their pre-extinction state, and by the Middle Triassic greater functional evenness is recorded, resulting from the radiation of previously subordinate groups such as motile, epifaunal grazers.

Evolution of the last koninckinids (Athyridida, Koninckinidae), a precursor signal of the early Toarcian mass extinction event in the Western Tethys

Koninckinids are a suitable group to shed light on the biotic crisis suffered by brachiopod fauna in the Early Jurassic. Koninckinid fauna recorded in the late Pliensbachian–early Toarcian from the easternmost Subbetic basin is analyzed and identified as a precursor signal for one of the most conspicuous mass extinction events of the Phylum Brachiopoda, a multi-phased interval with episodes of changing environmental conditions, whose onset can be detected from the Elisa–Mirabile subzones up to the early Toarcian extinction boundary in the lowermost Serpentinum Zone (T-OAE). The koninckinid fauna had a previously well-established migration pattern from the intra-Tethyan to the NW-European basins but a first phase with a progressive warming episode in the Pliensbachian–Toarcian transition triggered a koninckinid fauna exodus from the eastern/central Tethys toward the westernmost Mediterranean margins. A second stage shows an adaptive response to more adverse conditions in the westernmost Tethyan margins and finally, an escape and extinction phase is detected in the Atlantic areas from the mid-Polymorphum Zone onwards up to their global extinction in the lowermost Serpentinum Zone. This migration pattern is independent of the paleogeographic bioprovinciality and is unrelated to a facies-controlled pattern. The anoxic/suboxic environmental conditions should only be considered as a minor factor of partial control since well-oxygenated habitats are noted in the intra-Tethyan basins and this factor is noticeable only in the second westward migratory stage (with dwarf taxa and oligotypical assemblages). The analysis of cold-seep proxies in the Subbetic deposits suggests a radiation that is independent of methane releases in the Subbetic basin.

Significant pre-mass extinction animal body-size changes : evidences from the Permian–Triassic boundary brachiopod faunas of South China

This paper has undertaken a quantitative and statistical analysis of brachiopod body-size changes through the marine Permian–Triassic boundary section at Zhongzhai, Guizhou Province, South China, and found that (1) pre-mass extinction dwarfing is evident for at least the rugosochonetid species chosen for this study; (2) Tethyochonetes species reduced their size earlier than that in the Neochonetes species; and (3) no significant size reduction occurred in the newly evolved species of these two genera. Inter-species competition for resources between Neochonetes species and Tethyochonetes species and the reduction of food supply in the upper part of the uppermost Permian is here proposed to explain these observed stratigraphic patterns of brachiopod body-size changes throughout the Zhongzhai section. In the case of the newly evolved species showing no significant body-size change, morphological innovations (adaptations) in the process of speciation are considered to have significantly enhanced these newly evolved species' flexibility and survival in coping with degrading environmental conditions.