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.

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.

Global review of permian Tyloplecta muir-wood and cooper, 1960 (Brachiopoda): morphology, palaeobiogeographical and palaeogeographical implications

A global review of the stratigraphical and geographical distribution of Tyloplecta reveals that the genus ranges in age from Kungurian to Changhsingian (Middle to Late Permian). Tyloplecta first evolved in South China in the Kungurian (late Early Permian). The genus went through its first diversification in the Guadalupian, suffered a major extinction at the end of the Guadalupian, and re-diversified in the Wuchiapingian. T. yangtzeensis persisted into the Changhsingian as the only survivor of the genus involved in the end-Permian mass extinction. Palaeogeographically, South China is not only the centre of origin for the genus but also an area of diversification and evolution. In addition to South China, Tyloplecta has also been recorded from the Far East Russia, Japan, central Thailand, Laos, Cambodia, Qiangtang Terrane of Tibet, Salt Range, Iran, Armenia, Hungary, Yugoslavia, and Slovenia. This geographic spread suggests that Tyloplecta was primarily restricted to the Palaeotethys and is indicative of warm-water palaeoequatorial conditions. Its presence in some of the northeast Asian terranes (e.g., parts of Japan and Far East Russia) and in the Salt Range (Pakistan) and central and north Iran (part of the Cimmerian microcontinents) demonstrate that the genus invaded the middle palaeolatitudinal regions in both hemispheres during the late Middle Permian in response to increased shallow marine biotic communications between Cathaysia in the eastern Palaeotethys and southern Angaraland, and between Cathaysia and Peri-Gondwanaland. The invasion of Tyloplecta (and some other taxa) into the southern shore waters of Angaraland may be explained by assuming ocean surface current connections and close palaeogeographical proximities between the South China, Sino-Korea and Bureya blocks. In comparison, the invasion of Tyloplecta into the Peri-Gondwanaland region is more likely a result of reduced palaeogeographical distance between South China and Peri-Gondwanaland and the appearance of the Cimmerian microcontinents as migratory stepping stones.

Capitanian (late Guadalupian, permian) global brachiopod palaeobigeography and latitudinal diversity pattern

A global database containing 3365 occurrences, 821 species and 251 genera of the Capitanian (Late Guadalupian, Permian) brachiopod faunas from 24 stations has been analyzed by cluster analysis using the Jaccard and Otsuka coefficients and the probabilistic index of similarity, nonmetric multidimensional scaling and minimum spanning tree. Two supergroups, three groups and six subgroups are revealed and interpreted as representing, respectively, two biotic realms (the Palaeoequatorial and Gondwanan Realms), two regions and six provinces. An additional realm (the Boreal Realm), based on the fauna from Spitsbergen, also appears recognizable although it also shows considerable similarities with southwestern North America and the northern margin of Gondwana as revealed by the statistical analysis. The Palaeoequatorial Realm can be further subdivided into the North America Region and the Asian Tethyan Region. The six biotic provinces are the Cathaysian Province in the Palaeotethys and Mesotethys, the Greenland-Svalbard Province in the Arctic region, the Austrazean Province in eastern Australia and New Zealand, the Grandian Province in western North America and the two transitional zones (the Himalayan Province in the southern temperate zone and the Sino–Mongolian–Japanese Province in the northern temperate zone). Polynomial regression analysis and rarefaction analysis indicate that the generic diversities of brachiopod faunas during the Capitanian peaked in the Palaeoequatorial Cathaysian Province and the two transitional zones (Himalayan Province and Sino–Mongolian–Japanese Province), but fell dramatically in the polar regions. The generic diversity of the Palaeoequatorial Grandian Province is apparently lower than in the two transitional zones of temperate palaeolatitudes, suggesting that the generic diversity of Capitanian brachiopod faunas does not exhibit a strict negative correlation with palaeolatitudes. This in turn would suggest that biogeographical determinants (such as geographical barriers, inhabitable area and ocean currents) other than latitude-related temperature control may also have played an important role in the dispersal of some brachiopods and the characterization of some local provinces and high diversities. The Capitanian global brachiopod palaeobiogeography is generally comparable with those in the Wuchiapingian and Changhsingian, but with some notable differences. These include: (1) that the Grandian Province of the Capitanian in western North America vanished after the end-Guadalupian regression, (2) that the western Tethyan Province of the Lopingian could not be distinguished in the Capitanian, and (3) that the Austrazean Province was larger in area than either in the Wuchiapingian or in the Changhsingian.