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.

Permophricodothyris pavlova, 1965 (Brachiopoda, Spiriferida) from the Permian of South China: its morphology, biostratigraphy and distribution

A study of both silicified and nonsilicified specimens of Permian reticularioid brachiopods from South China suggests that Permophricodothyris, a genus previously rarely reported from China, is actually very common and abundant in the Middle and especially Upper Permian of South China. This study also clarifies, for the first time, that many of the reticularioid brachiopod species previously described as Squamularia in fact belong to Permophricodothyris. The new data presented in this paper also allows a critical evaluation of Permophricodothyris in relation to its closest allies: Phricodothyris, Squamularia, Bullarina and Neophricodothyris. The revision reveals that a total of 18 Permophricodothyris species are present in the Middle and Upper Permian of South China, with only one species, P. squamularioides, having survived the Permian-Triassic mass extinction. Two species, P. grandis (Chao) and P. guangxiensis Han, Zhou & Wang, are redescribed here, providing critical new information on the morphology and taxonomy of these species.

Middle permian (Guadalupian) brachiopods from the Xiujimqinqi area, Inner Mongolia, northeast China, and their palaeobiogeographical and palaeogeographical significance

A small brachiopod fauna is described from the lower part of the Xiujimqinqi Formation of the Xiujimqinqi area in central-east Inner Mongolia, northeast China. The age of this fauna is regarded as Wordian (Middle Guadalupian, Middle Permian) by comparison with a similar brachiopod fauna from the Zhesi area of central Inner Mongolia, and by constraints from fusulinaceans associated with the Zhesi fauna. The Xiujimqinqi fauna is typical of mixed Boreal/Palaeoequatorial Middle Permian brachiopod faunas of East Asia. The mixed nature of these faunas is interpreted to have resulted from the combined effects of a middle palaeolatitudinal position, intensified plate convergence between Sino-Korea and Mongolia, and sea surface current connections with both the Arctic Sea in the north and eastern Palaeo-Tethys to the south. Possible Kaninospirifer is reported for the first time from China.

Linshuichonetes gen. nov., a new rugosochonetid (Brachiopoda) genus from the Liangshan Formation (Early Permian) in Sichuan, China, and its ecology

A new genus and species, Linshuichonetes elfinis, belonging to the Rugosochonetidae, is described from the Early Permian (Late Artinskian or early Kungurian) Liangshan Formation of the Yangtze block. The new genus is defined externally by the presence of fine, but delayed, capillation and a weak or absent median sulcus and fold and by the presence of a distinct posteromedian sinus on the ventral umbo; and internally by a lack of median, lateral and accessory septa in the dorsal interior; absence of vascular trunks in the ventral interior and the presence of distinct radiating rows of papillae in the interior of both valves, particularly an unusual clustered arrangement of papillae on the posteromedian portion of the dorsal interior. The local environment during the deposition of the Liangshan Formation appears to have been a restricted tidal flat or lagoon which experienced frequent sealevel fluctuations associated with the onset of the Yanghsingian transgression. The new species, L. elfinis, appears to have several morphological adaptations enabling successful exploitation of this environment. It was typically a very small and thin-valved species with a high surface area to volume ratio, an advantage in an oxygen restricted environment. The small size and numerous body spinules would have aided individuals to remain suspended at the top of the fine, soft substrate. It also dominated the brachiopod assemblage in the Liangshan Formation, comprising up to 94%of specimens within a bed. These factors indicate that the new species appears to be an opportunistic species.

Global distribution of permian conulariids and palaeobiogeographical implications

Previously, minimal work has been carried out on conulariids due to their rare occurrences and resultant biostratigraphical limitations. The palaeobiogeographical distribution of Permian conulariids suggests that they have a marked preference for cold to cool-water regions, that they are significant indicators for migration patterns, and that they can potentially provide information on the palaeogeographical configuration and movement of terranes. Permian conulariids are found in Australia, India, New Zealand, Pakistan, Iran, Afghanistan, Kashmir, China, Japan, Russia, Germany, Canada, United States of America, and Bolivia. The diversity of Permian conulariids is markedly higher in the polar regions than in the palaeoequatorial region.

Permian conulariid genera include Notoconularia Thomas 1969, Gondaconularia Waterhouse 1986, Cheliconularia Waterhouse 1986, Neoconularia Sugiyama 1942, Calloconularia Sinclair 1952, Diconularia Sinclair 1952, Paraconularia Sinclair 1940, Mesoconularia Boucek 1939 and Conularia Sowerby 1821. This paper describes two new species of conulariids: Diconularia meadepeakensis sp. nov. from the Phosphoria Formation (Guadalupian), Idaho, USA and Paraconularia kazanensis sp. nov. from the Sokian Horizon (?Roadian), Volga Region, Russia.

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.

A Guadalupian-Lopingian (Middle to Late Permian) brachiopod fauna from the Juripu formation in the Yarlung-Zangbo suture zone, Southern Tibet, China

A brachiopod fauna of 16 species belonging to 11 genera and three genera and species indeterminate from the middle and upper parts of the Juripu Formation in the Yarlung-Zangbo (Indus-Tsangbo) Suture zone (=Yarlung-Zangbo River zone), southern Tibet, is described and figured for the first time. A new species, Taeniothaerus zhongbaensis, is described. The fauna is comparable with that in the Kalabagh Member of the Wargal Formation of the Salt Range, Pakistan, and is considered to be most likely Capitanian (late Guadalupian/Middle Permian) to Wuchiapingian (early Lopingian/early Late Permian) in age, as indicated by the majority of the brachiopod species and by being constrained by an underlying fusulinacean fauna (Parafusulina Zone) and an overlying ammonite fauna (Cyclolobus fauna).

The Gondwanan Carboniferous-Permian Boundary Revisited: New Data from Australia and Argentina

The identification and correlation of the Carboniferous-Permian (Gzhelian-Asselian) boundary within the sedimentary sequences of Gondwana has always been a topic of debate. Type latest Carboniferous and earliest Permian marine sequences are characterised by warm tropical faunas and come from the Uralian Region of Russia and Kazakhstan. Faunas include conodonts and fusulinid foraminiferids which are prime tools for correlation. Such faunal groups are absent from most Gondwanan sequences where reliance for correlations must be placed primarily on brachiopods, bivalve molluscs and palynology. The Western Australian marine sequences, with their contained ammonoids, provide a pivotal link for the dating and correlating of Early Permian Gondwanan sequences with those of the type regions and their palynostratigraphical record is essential for trans-Australian correlations and correlations elsewhere throughout Gondwanaland.

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.

New lopingian (late permian) rugosochonetid species from Sichuan, South China

Two rugosochonetid species, Neochonetes (Huangichonetes) geniculatus sp. nov. and Neochonetes (Zhongyingia) linshuiensis sp. nov., are described from the Lopingian (Late Permian) of the Chuanmu section, Sichuan, South China. Ecological changes from the diverse upper Changhsingian brachiopod palaeocommunity to the depauperate post-extinction brachiopod community are briefly discussed.

High-resolution terrestrial permian-triassic eventostratigraphic boundary in western Guizhou and eastern Yunnan, southwestern China

The adjoining area of western Guizhou and eastern Yunnan Provinces in southwest China is an ideal place to investigate the feasibility of correlating marine and nonmarine Permian–Triassic boundary (PTB) sequences, as it contains outcrop sections of shallow marine, marginal marine (or paralic), and terrestrial PTB sections, all in close geographic proximity. This paper documents for the first time multiple stratigraphic data from several well-preserved terrestrial PTB sections in the area and attempts to use these data to define, locate, and correlate the PTB in the area. A study of the spores and pollen and vegetation types across the terrestrial PTB sections in the study area suggests three distinct evolutionary stages across the boundary: Stage 1 (Xuanwei Formation) is characterised by Late Permian or Paleozoic-type ferns and pteridosperms (85.0%), with a few gymnosperms (15.0%); stage 2 is marked by an abrupt drop of sporopollen elements of Late Permian aspects, coupled with the appearance of fungal spores and limited Early Triassic palynomorphs; stage 3 (top Xuanwei Formation and Kayitou Formation) is dominated by gymnosperm pollen (58.8%) of clearly Early Triassic aspect, although still retaining limited ferns and pteridosperms. The three biotic stages seem to well correspond with the changing trend of the δ13Corg curves from the same sections, which is characterized by a sharp drop just before the PTB, followed by a short term partial recovery across the boundary, and then succeeded by a gradual decline after the PTB in the Early Triassic. Combining evidence from eventostratigraphic (i.e., the succession of boundary clay beds), biostratigraphic (using both macroplants and palynomorphs), and chemostratigraphic (i.e., organic carbon isotope excursion signals), we propose that a high-resolution PTB succession, closely correlatable to its marine counterpart at the Meishan section in eastern China, is recognisable at the terrestrial PTB sections in the western Guizhou–eastern Yunnan area in southwest China.

Guadalupian (Middle Permian) brachiopods from Sungai Toh, a Leptodus Shale locality in the Central Belt of Peninsular Malaysia. Part I: Lower Horizons

This work presents a systematic study of Permian Brachiopoda from the Sungai Toh Leptodus Shale locality, Pahang State, Peninsular Malaysia. This locality lies within the Central Belt of Peninsular Malaysia, a tectonic unit characterised.by tuffaceous sediments and limestones of Late Palaeozoic age. Two brachiopod-bearing horizons were studied in detail at this locality, the lower one (Horizon 2) bearing a mixed plant and invertebrate assemblage, including the brachiopods Urushtenoidea chaoi (CHING), Leptodus richthofeni KAYSER, Anidanthus cf. sinosus HUANG, Acosarina dorashamensis (SOKOLSKAJA), A. minuta (ABleH) and unidentifiable species of Linoproduetus, Neochonetes, and Strophalosiina. Horizon 3 contains a more abundant and diverse brachiopod fauna, comprising a total. of 57 species representing 47 genera, including Vediproductus punetatiformis (CHAO), Permianella typica HE & ZHU, Tranrennatia gratiosa (WAAGEN), Leptodus richthofeni KAYSER, Leptodus cf. tenuis (WAAGEN) and "Semibrachythyrina" [= Alphaneospirifer] cf. pyramidiformis LIANG. It is
suggested in this study that the age of the Sungai Toh locality is Capitanian (late Guadalupian) to possibly Wuchiapingian (early Lopingian),
as it appears to correlate well with the Lengwu fauna from Zhejiang in eastern China. The palaeobiogeographical affinities of the Sungai Toh fauna are interesting, mainly indicating strong Palaeo-equatorial affinities, while there are also some elements more typical of the cooler periGondwana
Region.

Lower permian oncolites from South China: implications for equatorial sea-level responses to late palaeozoic Gondwanan glaciation

The oncoid-bearing Chuanshan Formation is a regionally extensive carbonate deposit of predominantly Asselian to early Sakmarian (Early Permian) age in South China, occupying an area of some 500,000 km2. Throughout South China, the oncoid-bearing horizons are generally stable and broadly comparable in lithology, fossil content and the morphology of the oncoid grains. Four types of microfacies are recognized from the oncolite succession and overall they suggest a moderate- to high-energy, wave-agitated shallow marine carbonate platform environment. An analysis of the stratigraphic distribution of oncoid grain size, density, thickness and the bedding structures of the oncolite beds and the number of coating laminae indicate the presence of metre-scale cyclothems, suggestive of possible high-frequency cycles of sea-level fluctuation. Compared to carbonate successions above and below that lack oncolites, and in conjunction with evidence from sequence stratigraphic and isotopic geochemical analyses of coeval carbonate deposits in South China and elsewhere, the origin of the Chuanshan oncolites is linked to a drastic drop in global sea-level at the Pennsylvanian–Permian boundary, that can be correlated closely in timing with the zenith of the Late Palaeozoic Gondwanan glaciation. It is further suggested that the eustatic changes apparent from the deposition of the Chuanshan oncolites and similar coeval deposits in lower palaeolatitudes were coupled with, and influenced by, the contemporaneous high-latitude Gondwanan glaciation, the largest and longest known such event in Phanerozoic Earth history.