Early Triassic (early Olenekian) life in the interior of East Gondwana: Mixed marine-terrestrial biota from the Kockatea Shale, Western Australia

A new terrestrial-marine assemblage from the lower beds of a thin outcrop section of the Kockatea Shale in the northern Perth Basin, Western Australia, contains a range of fossil groups, most of which are rare or poorly known from the Lower Triassic of the region. To date, the collection includes spinose acritarchs, organic-cemented agglutinated foraminifera, lingulids, minute bivalves and gastropods, ammonoids, spinicaudatans, insects, austriocaridid crustaceans, actinopterygians, a temnospondyl-like mandible, plant remains, and spores and pollen. Of these groups, the insects, crustaceans and macroplant remains are recorded for the first time from this unit. Palynomorphs permit correlation to nearby sections where conodonts indicate an early Olenekian (Smithian) age. The locality likely represents the margin of an Early Triassic shallow interior sea with variable estuarine-like water conditions, at the southwestern end of an elongate embayment within the East Gondwana interior rift-sag system preserved along the Western Australian margin. Monospecific spinose acritarch assemblages intertwined with amorphous organic matter may represent phytoplankton blooms that accumulated as mats, and suggest potentially eutrophic surface waters. The assemblage represents a mixure of marine and terrestrial taxa, suggesting variations in water conditions or that fresh/brackish-water and terrestrial organisms were transported from adjacent biotopes. Some of the lower dark shaly beds are dominated by spinicaudatans, likely indicating periods when the depositional water body was ephemeral, isolated, or subjected to other difficult environmental conditions. The biota of the Kockatea Shale is insufficiently known to estimate biotic diversity and relationships of individual taxa to their Permian progenitors and Triassic successors, but provides a glimpse into a coastal-zone from the interior of eastern Gondwana. Specialist collecting is needed to clarify the taxonomy of many groups, and comparisons to other Lower Triassic sites are required to provide insights into the pattern of biotic decline and recovery at the end-Permian crisis.

A preliminary phylogenetic study of late Palaeozoic spiriferoid brachiopods using cladistic and Bayesian approaches

Abstract The brachiopod Superfamily Spiriferoidea diversified greatly and was widely distributed in the late Palaeozoic (Carboniferous–Permian), and yet its phylogeny has been seldom investigated with analytical methods. This is reflected in the current flux of very different classification schemes for this superfamily. This paper provides the first attempt to investigate the phylogenetic relationships of spiriferoid brachiopods through both cladistic and Bayesian analyses involving 24 discrete and continuous characters. The continuous characters, from morphometric data, have been separately discretized using the gap weighting method, and the ‘as such’ option in TNT. Our results highlight the potential significance of continuous characters in reconstructing and elucidating phylogenies, as much as qualitative characters. Building on the outcomes of the analyses, we also briefly evaluate existing classification schemes of Spiriferoidea. We found that none of the existing classifications fully reflect the phylogeny properly; major families within the superfamily, such as Spiriferidae, Choristitidae, and Trigonotretidae, turned out to be polyphyletic. Although this study is considered preliminary, due to the selection of and restriction to certain taxa, combined with the use of a relatively small number of characters, it nevertheless demonstrates that potentially the true phylogenetic relationships of spiriferoid taxa sharply contrast with any of the existing classification schemes. This highlights the need to develop an alternative scheme that takes into account a more comprehensive range of phylogenetic variables.

Fluctuations of redox conditions across the Permian–Triassic boundary : new evidence from the GSSP section in Meishan of South China

Oceanic anoxia has long been considered as one of the main causes for the end-Permian mass extinction. However, the results obtained by different researchers are rather divergent from different sections, or even on the same section using the same redox proxy. This study aims to examine the causes for some of these divergent results using high-resolution pyrite framboid sampling at the Meishan GSSP section in South China. Detailed microfacies analysis shows that the uppermost Late Permian strata comprises two significantly different sedimentary facies: one characterized by silicious muddy limestone and recognized as representing autochthonous background sediments; the other distinguished by bioclastic grainstone, interpreted to be allochthonous in origin and have been transported from the nearby platform margin. These two different sedimentary facies represent two distinctly different redox conditions. Together with the facies analysis, a statistical analysis of pyrite framboids was carried out to evaluate the redox evolution across the Permian-Triassic boundary. Abundant framboids with average diameters of about 6μm are found in background sediments beneath the extinction boundary, indicating generally anoxic bottom water conditions. But this condition was punctuated by transient intervals of rapid oxygenation interpreted to have been caused by intrusion of intermittent turbidity flows. Our study also showed that anoxic conditions persisted into the immediate aftermath of the mass extinction, thereafter it was quickly followed by a relatively long period of oxic conditions (with rare framboids). However, the redox conditions returned to anoxia (with abundant pyrite framboids averaging about 5μm in diameter), accompanied by a rapid global transgression. The oxygenation manifested near the Permian-Triassic boundary coincides with the negative excursion of carbon isotope. This would imply that, contrary to previous interpretations, this great δ13C negative excursion was probably not caused by the upwelling of anoxic deep ocean waters.

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.

Early Triassic disaster and opportunistic foraminifers in South China

Survival and recovery are important dynamic processes of biotic evolution during major geological transitions. Disaster and opportunistic taxa are two significant groups that dominate the ecosystem in the aftermath of mass extinction events. Disaster taxa appear immediately after such crises whilst opportunists pre-date the crisis but also bloom in the aftermath. This paper documents three disaster foraminiferal species and seven opportunistic foraminiferal species from Lower Triassic successions of South China. They are characterized by extreme high abundance and low diversity and occurred occasionally in Griesbachian, Smithian and Spathian strata. The characteristics (small size, simple morphology) and stratigraphic ranges of these groups suggest that r-selection is a commonly used strategy for survivors to cope with either harsh post-extinction conditions and/or environments lacking incumbents.

Taphonomy and palaeobiology of early Middle Triassic coprolites from the Luoping biota, southwest China: implications for reconstruction of fossil food webs

Abundant, exceptionally preserved coprolites are documented from the Luoping biota (Anisian, Middle Triassic) of Yunnan Province, southwest China. These coprolites can be categorized into fourmorphological types: A) bead to ribbon-shaped, B) short to long cylindrical-shaped, C) flattened, disk-like, and D) segmented faeces. Detailed multi-disciplinary studies reveal that coprolite type A was likely produced by invertebrate animals,while coprolite types B to D could be faeces generated by carnivorous fishes or marine reptiles, perhaps from different taxonomicgroups. When compared with coprolites reported from the Lower Triassic, the Luoping forms indicate more complicated predation-prey food web networks. These evidences, combined with body fossil discoveries fromLuoping, suggest the emergence of complex trophic ecosystems in the Anisian,marking the full biotic recovery following the Permian–Triassic Mass Extinction.

Palaeobiogeographic distribution patterns and processes of neochonetes and fusichonetes (brachiopoda) in the late palaeozoic and earliest mesozoic

The global palaeobiogeographic distributions of two resembling genera, Neochonetes and Fusichonetes (Brachiopoda), from the Carboniferous to Griesbachian are analysed. This analysis provides insight into the biotic response of two related genera to changing palaeoclimate, regional tectonics, and environmental crises. Neochonetes originated in the equatorial area in the Mississippian, and it mostly retained this position during the peak of the glaciation in the Carboniferous–Permian ice age (namely in the Pennsylvanian). Neochonetes then dispersed globally during the Cisuralian when the climate became warmer and the ice sheet started to retreat. In the Guadalupian and Lopingian, following the closure of the Ural seaway at the end of the Cisuralian and the regression at the end-Guadalupian, Neochonetes almost disappeared in the western part of Gondwana. Subsequently during the Lopingian the genus retracted to the middle- and low-latitude Palaeo-Tethys and Tethys. In comparison, Fusichonetes originated in the equatorial area in the late Guadalupian and was still present in that area in the Lopingian. Both genera occurred only in South China in the Griesbachian. It is inferred that this could be related, not only to the deteriorated palaeoenvironmental conditions (e.g., anoxia, global warming) leading up to the extinction of most of the Neochonetes and Fusichonetes species in other areas, but also to the better physiological adaptation of the smaller shells of Neochonetes and Fusichonetes species in South China.