Discovery of late Changhsingian (latest Permian) brachiopod Attenuatella species from South China

Attenuatella mengi sp. nov. and ?Attenuatella sp. from the Talung Formation, southern Guangxi Zhuang Autonomous Region, South China, are described herein. This discovery represents the first report of Attenuatella from the late Changhsingian (latest Permian) in South China and provides evidence that Attenuatella expanded its range from high-latitude cold-water regions to palaeoequatorial warm water areas in the Late Permian. Attenuatella
species appear to have been pseudoplanktonic, judging from their hair-like spinose ornamentation, which could have contributed to the global palaeogeographical distribution of Attenuatella.

An ecologically mixed brachiopod fauna from Changhsingian deep-water basin of South China: consequence of end-Permian global warming

The Late Permian Shaiwa Group of the Ziyun area of Guizhou, South China is a deep-water facies succession characterized by deep-water assemblages of pelagic radiolarians, foraminifers, bivalves, ammonoids and brachiopods. Here we report 20 brachiopod species in 18 genera from the uppermost Shaiwa Group. This brachiopod fauna is latest Changhsingian in age and dominated by productides. The palaeoecologic and taphonomic analysis reveals that the brachiopod fauna is preserved in situ. The attachment modes and substratum preference demonstrate that the Shaiwa brachiopod fauna comprises admixed elements of deep-water and shallow-water assemblages. The presence of the shallow-water brachiopods in the Shaiwa faunas indicates the involuntary settlement of shallow-water brachiopods. The stressed ecologic pressure, triggered by warming surface waters, restricted ecospace and short food sources, may have forced some shallow-water elements to move to hospitable deep-water settings and others to modify their habiting behaviours and exploit new ecospace in deep-water environments. We infer that the end-Permian global warming and subsequent transgression event may have accounted for the stressed environmental pressure in the shallow-water communities prior to the end-Permian mass extinction.

A late Changhsingian (latest Permian) deep-water brachiopod fauna from Guizhou, South China

A deep-water brachiopod fauna (20 species in 19 genera) is described from the Late Permian Shaiwa Group of Ziyun, Guizhou, South China. New species include Pygmochonetes? shaiwaensis and Martinia ziyunensis. This fauna is associated with deep-water assemblages of pelagic radiolarians, foraminifers, bivalves and ammonoids. The brachiopod faunal correlations and age constraints of the associated fossil groups suggest that the Shaiwa fauna is late Changhsingian (latest Permian) in age. The Shaiwa fauna superficially resembles the coeval deep-water assemblage from Guangxi, South China; both are characterized by a mixture of deep-water brachiopods and shallow-water elements.

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.

A new Changhsingian (Late Permian) brachiopod fauna from the Zhongzhai section (South China) Part 3: Productida

Zhang, Y., He, W.H., Shi, G.R., Zhang, K.X. & Wu, H.T., 26.2.2015. A new Changhsingian (Late Permian) brachiopod fauna from the Zhongzhai section (South China) Part 3: Productida. Alcheringa 39, xxx–xxx. ISSN 0311-5518. As the third and last part of a systematic palaeontological study of the brachiopod fauna from the Permian–Triassic boundary section at Zhongzhai in Guizhou Province (South China), this paper reports 15 species (including three new species: Tethyochonetes minor sp. nov., Neochonetes (Zhongyingia) transversa sp. nov., Paryphella acutula sp. nov.) in Order Productida. In addition, the morphological features and definitions of several key Changhsingian brachiopod taxa (e.g., Paryphella and Oldhamina interrupta) are clarified and revised.

Nearshore-offshore-basin species diversity and body size variation patterns in Late Permian (Changhsingian) brachiopods

 B.V. Body size is a fundamental and defining character of an organism, and its variation in space and time is generally considered to be a function of its biology and interactions with its living environment. A great deal of body size related ecological and evolutionary research has been undertaken, mostly in relation to extant animals. Among the many body size-related hypotheses proposed and tested, the size-bathymetry relationship is probably the least studied. In this study, we compiled a global body size dataset of Changhsingian (Late Permian, ca. 254. Ma-252. Ma) brachiopod species from low-latitude areas (30°S-30°N) and analyzed their species diversity and body size distribution patterns in relation to the nearshore-offshore-basin bathymetric gradient. The dataset contained 1768 brachiopod specimens in 435 species referred to 159 genera and 9 orders, from 135 occurrences (localities) of 18 different palaeogeographic regions. Treating the whole of the Changhsingian Stage as a single time slice, we divided the nearshore-offshore-basin bathymetric gradient into three broad depth-related environments: nearshore, offshore and basinal environments, and compared how the species diversity and body size varied along this large-scale bathymetric gradient.Here, we report an array of complex patterns. First, we found a clear overall inverse correlation between species diversity and water depth along the nearshore-offshore-basin gradient, with most species concentrating in the nearshore environment. Second, when the median sizes of all low-latitude brachiopod species from the three environments were compared, we found that there was no significant size difference between the nearshore and offshore environments, suggesting that neither the wave base nor the hydrostatic pressure exerts a critical influence on the body size of brachiopods. On the other hand, the median sizes of brachiopods from the nearshore environment and, to a lesser extent, the offshore environment were found to be significantly larger than that of basinal brachiopods. This trend of significant size reduction in basinal brachiopods mirrors the relative low species diversity in the basinal environment, and neither can be easily explained by the tendency of decreasing food availability towards deeper sea environments. Rather, both trends are consistent with the hypothesis of an expanding Oxygen Minimum Zone (OMZ) in the bathyal (slope to deepsea) environments, where hypoxic to anoxic conditions are considered to have severely restricted the diversification of benthos and favored the relative proliferation of small-sized brachiopods. Finally, a significant difference was also found between eurybathic and stenobathic species in their body size response to the nearshore-offshore-basin gradient, in that eurybathic species (species found in all three environments) did not tend to change their body size significantly according to depth, whereas stenobathic forms (species restricted to a single environment) exhibit a decline in body size towards the basinal environment. This pattern is interpreted to suggest that bathymetrically more tolerant species are less sensitive to depth control with respect to their body size change dynamics, in contrast to stenobathic species which tend to grow larger in shallower water depths.

Global brachiopod palaeobiogeographical evolution from Changhsingian (Late Permian) to Rhaetian (Late Triassic)

Previous studies suggest that the end-Permian mass extinction caused a dramatic drop of marine biodiversity near the Permian-Triassic boundary. However, it is unclear how profoundly this severe extinction might have changed the global provincialism, and how global provincialism responded to the protracted process of this extinction and subsequent recovery through the Triassic. In this paper, we carried out quantitative time-series analyses of global brachiopod palaeobiogeography over a timespan of nine consecutive stages/substages from the latest Permian Changhsingian to the latest Triassic Rhaetian based on a global brachiopod database of 483 genera and 2459 species from 1425 localities. Our results suggest that the extinction resulted in a global 'biogeographical eclipse' in the ensuing Early Triassic Griesbachian and Dienerian times in that neither biogeographic realm nor province could be recognized. It was characterized by an extreme low-diversity, mostly dwarfed and nearly globally distributed brachiopod fauna, coupled with persistently high sea surface temperature and a flattened global latitudinal thermal gradient. Global provincialization emerged again during the Olenekian at province level and reached its peak stage during the Carnian when three realms and six provinces were clearly recognized. Global provincialism became weakened again in the latest Triassic Rhaetian, marked by three general realms, but no province distinguished. Our analyses suggest that both palaeolatitude-related thermal gradient and the presence of Pangea (a profound geographic barrier) were most effective in explaining the spatial patterns. In addition, oceanic currents along the northwestern coast of Pangea also played an important (albeit regional) role in linking southern North and Central America brachiopod faunas with those of the Boreal Realm. This study also revealed that the brachiopod biodiversity center moved northwards over the studied interval, accompanied and hence accountable for by the northward drift of a large number of tectonic blocks in the Palaeotethys and Neotethys during the Triassic.

Contrasting geochemical signatures on land from the Middle and Late Permian extinction events

The end of the Palaeozoic is marked by two mass-extinction events during the Middle Permian (Capitanian) and the Late Permian (Changhsingian). Given similarities between the two events in geochemical signatures, such as large magnitude negative C-13 anomalies, sedimentological signatures such as claystone breccias, and the approximate contemporaneous emplacement of large igneous provinces, many authors have sought a common causal mechanism. Here, a new high-resolution continental record of the Capitanian event from Portal Mountain, Antarctica, is compared with previously published Changhsingian records of geochemical signatures of weathering intensity and palaeoclimatic change. Geochemical means of discriminating sedimentary provenance (Ti/Al, U/Th and La/Ce ratios) all indicate a common provenance for the Portal Mountain sediments and associated palaeosols, so changes spanning the Capitanian extinction represent changes in weathering intensity rather than sediment source. Proxies for weathering intensity chemical index of alteration, W and rare earth element accumulation all decline across the Capitanian extinction event at Portal Mountain, which is in contrast to the increased weathering recorded globally at the Late Permian extinction. Furthermore, palaeoclimatic proxies are consistent with unchanging or cooler climatic conditions throughout the Capitanian event, which contrasts with Changhsingian records that all indicate a significant syn-extinction and post-extinction series of greenhouse warming events. Although both the Capitanian and Changhsingian event records indicate significant redox shifts, palaeosol geochemistry of the Changhsingian event indicates more reducing conditions, whereas the new Capitanian record of reduced trace metal abundances (Cr, Cu, Ni and Ce) indicates more oxidizing conditions. Taken together, the differences in weathering intensity, redox and the lack of evidence for significant climatic change in the new record suggest that the Capitanian mass extinction was not triggered by dyke injection of coal-beds, as in the Changhsingian extinction, and may instead have been triggered directly by the Emeishan large igneous province or by the interaction of Emeishan basalts with platform carbonates.

中国二叠纪—三叠纪陆生植物多样性变化的研究

本文基于华南、华北地区二叠纪—三叠纪陆生植物大化石和孢粉的数据库， 对中国二叠纪—三叠纪陆生植物的多样性变化进行了统计分析研究，并重点探讨了在二叠纪—三叠纪界线（Permian-Triassic Boundary，PTB ）陆生植物是否与同期的海洋动物一样发生了同步的集群灭绝事件。 统计分析表明，华南、华北陆生植物大化石的分异度穿过PTB 均显示了较长时续（约37.8Ma）的下降和残存阶段，而孢粉化石在早三叠世的分异度则是上升的。总体上，陆生植物分异度穿过PTB 的变化较同期的海洋动物平稳缓慢。华南地区陆生植物大化石在晚二叠世末长兴期（Changhsingian）虽然伴随着最高的属灭绝率85.94% 和最低的属新生率28.12%，发生了最大的灭绝事件，但在晚二叠世早期和早三叠世的属的灭绝率也较高，分别为61.02% 和66.67% 。种的灭绝率在晚二叠世早期从早二叠世晚期的39%大幅度上升到80.36%，晚二叠世晚期达峰值97%，早三叠世稍降为93%，显然高于其它时段灭绝率范围（30—70%）。种和属的灭绝率呈现了同样的高峰阶段，从晚二叠世早期至早三叠世，时续为20.8 百万年（Ma）。基于更替率分析，华南地区陆生植物的高更替率事件分别发生在早二叠世晚期（93.75%）、早三叠世（90.92%）和晚三叠世（91.38%），但陆生植物在穿越早二叠世晚期—晚三叠世的整个过程中，更替率波动不大、比较平稳。华北地区陆生植物大化石穿越PTB 的灭绝率比华南地区低，属级高灭绝率事件集中在晚二叠世早期（67.31%）和晚二叠世晚期（63.89%）， 时续为14.8Ma，种级高灭绝率事件与华南地区类似，集中在晚二叠世早期（85.67%）、晚二叠世晚期（90.86%）和早三叠世（80.28% ）三个阶段，时续为20.8Ma 。显而易见，这比同期海洋动物集群灭绝的时续（3—11Ma ）要长。 本文基于这些分析结果，仔细考虑了集群灭绝的4 个特点（即量值、广度、幅度和时续），认为华南、华北陆生植物在PTB 并未发生集群灭绝事件，而是发生了演化替代，即陆生植物穿过PTB 经历了大的植物群重组和新种的演化。总体上，中国二叠纪—三叠纪陆生植物中选择性灭绝非常明显，古生代占优势的种子蕨、真蕨类、木本石松类和楔叶类逐渐被早中生代比较进化的裸子植物和真蕨类植物所替代，陆生植物穿过PTB 显示了危机（灭绝）—残存—复苏—辐射的宏演化式样。

Peri-gondwanan permian correlations : the meso-tethyan margins

Correlations of the Permian sequences for sixteen regions of north eastern Gondwana during the Permian are presented in this review. These correlations are compared with Permian sequences of the Australian continent. Broad conclusions on palaeoclimatic change and tectonic events are summarised for six time intervals of the Permian Period.

The Asselian-Sakmarian-early Artinskian time interval indicates a change from cold to temperate depositional environments. Glacial deposits and low diversity Gondwanan marine faunas are succeeded by younger, warmer water, clastic and bioclastic sequences with moderately diverse marine faunas. Deposition of these sequences is occasionally associated with basaltic volcanism and initial rifting of the peripheral northern Gondwanan margin.

During the Late Artinskian-Kungurian (including Early Ufimian) time interval, climate amelioration occurred with the onset of carbonate deposition in several Cimmerian terranes. Basaltic volcanism in several terranes is indicative of significant rifting and the opening of the Meso-Tethys.

The Roadian (Late Ufimian) and Wordian-Capitanian (including Kazanian-Midian) time intervals were characterised by widespread, subtropical, marine carbonate depositional sequences. These occurred throughout the Cimmerian blocks as they drifted northward and on the more northerly parts of the Meso-Tethyan southern margin. These transgressive sequences may rest on significant unconformity surfaces. Equivalent carbonate units are known in the offshore and subsurface sequences of western Australia. Andesitic, convergent plate margin volcanism and volcaniclastic sequences are present in eastern Australia.

The Wuchiapingian time slice is characterised by widespread marine transgressions which extended into the north western basins of Australia.

The Changhsingian time slice is represented by relatively minor marine transgressive events in the Trans-Himalaya with the Selong section of Tibet being probably the most complete Permo-Triassic sequence for the southern margin of the Meso-Tethys.