A Wuchiapingian (Late Permian) brachiopod fauna from an exotic block in the Indus–Tsangpo suture zone, southern Tibet, and its palaeobiogeographical and tectonic implications

A brachiopod fauna including 19 species of 17 genera from an exotic block in the Indus–Tsangpo suture zone in southern Tibet is described and illustrated. The brachiopod fauna is dominated by Martinia elegans and two new taxa: Jinomarginifera lhazeensis gen. et sp. nov. and Zhejiangospirifer giganteus sp. nov. The fauna is closely comparable with those from the middle and upper parts of the Wargal Formation and the Chhidru Formation in the Salt Range of Pakistan, the Chitichun Limestone in southern Tibet, and the Basleo area of West Timor, and these correlations suggest a Wuchiapingian age. The fauna exhibits substantial links with both peri–Gondwanan and Cathaysian faunas, which may imply that it is a seamount biota originally located in the southern margin of the Neotethys during the Late Permian, and was later (in the early Cenozoic) displaced and became sandwiched into younger marine deposits in the collision process between India and Eurasia.

Post-extinction brachiopod faunas from the late Permian Wuchiapingian coal series of South China

This paper describes fourteen brachiopod species in eleven genera from the Late Permian Wuchiapingian Coal Series (Lungtan Formation) of South China. Of these, the shell bed fauna from the basal Lungtan Formation is interpreted to represent the onset of the recovery of shelly faunas in the aftermath of the Guadalupian/Lopingian (G/L) mass extinction in South China. The post-extinction brachiopod faunas in the Wuchiapingian are characterized by the presence of numerous Lazarus taxa, survivors, and newly originating taxa. These elements capable of adapting their life habits were relatively more resistant to the G/L crisis. The post-extinction faunas, including survivors and the elements originating in the recovery period, have no life habit preference, but they were all adapted to a variety of newly vacated niches in the Late Permian oceans. Two new species, Meekella beipeiensis and Niutoushania chongqingensis, are described, and two Chinese genera, Niutoushania and Chengxianoproductus, are emended based on re-examination of the type specimens and new topotype materials from the Lungtan Formation.

Earliest Wuchiapingian (Lopingian, Late Permian) brachiopods in southern Hunan, South China : implications for the Pre-Lopingian crisis and onset of Lopingian recovery / radiation

The uppermost 5-15 m of the Douling Formation in the southern Hunan area. South China, yields a diverse fauna comprised of ammonoids, bivalves, and brachiopods. The brachiopods reported in this paper consist of 51 species in 34 genera and are dominated by the Lopingian (Late Permian) species associated with a few species persisting from the underlying Maokouan (Late Guadalupian). This fauna is of earliest Wuchiapingian in age as precisely constrained by the associated conodont Clarkina postbitteri postbitteri and the Guadalupian-type ammonoid fauna of the Roadoceras-Doulingoceras Zone in the brachiopod horizon. The discovery of the Lopingian species-dominated brachiopod fauna in the earliest Wuchiapingian in southern Hunan suggests a much less pronounced effect of the pre-Lopingian crisis (end-Guadalupian mass extinction) than the end-Changhsingian mass extinction in terms of brachiopods, a contemporaneous onset of the Lopingian recovery/radiation during the pre-Lopingian crisis period, and taxonomic selectivity of the pre-Lopingian crisis in terms of different fossil groups. New taxa are Echinauris doulingensis n. sp., Pararigbyella quadrilobata n. gen. and n. sp. and P. doulingensis n. gen. and n. sp.

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.

A Lopingian (Late Permian) brachiopod fauna from the Qubuerga formation at Shengmi in the Mount Qomolangma region of Southern Xizang (Tibet), China

A small fauna of 11 species belonging to 10 genera of Permian Brachiopoda from the lower part of the Qubuerga Formation outcropping near Shengmi village in the Qomolangma region of southern Xizang (Tibet) is figured and new taxa are described. New taxa are Quinquenella semiglobosa and Costatumulus shengmiensis. The fauna is most likely of Wuchiapingian (Djhulfian) age as indicated by the majority of the brachiopod species.

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).

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.

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.

Paleobiogeographical extinction patterns of Permian brachiopods in the Asian-western Pacific region

Spatial and temporal variations in biological diversity are critical in understanding the role of biogeographical regulation (if any) on mass extinctions. An analysis based on a latest database of the stratigraphic ranges of 89 Permian brachiopod families, 422 genera, and 2059 species within the Boreal, Paleoequatorial, and Gondwanan Realms in the Asian–western Pacific region suggests two discrete mass extinctions, each possibly with different causes. Using species/family rarefaction analysis, we constructed diversity curves for late Artinskian–Kungurian, Roadian–Wordian, Capitanian, and Wuchiapingian intervals for filtering out uneven sampling intensities. The end-Changhsingian (latest Permian) extinction eliminated 87–90% of genera and 94–96% of species of Brachiopoda. The timing of the end-Changhsingian extinction of brachiopods in the carbonate settings of South China and southern Tibet indicates that brachiopods suffered a rapid extinction within a short interval just below the Permian/Triassic boundary.

In comparison, the end-Guadalupian/late Guadalupian extinction is less profound and varies temporally in different realms. Brachiopods in the western Pacific sector of the Boreal Realm nearly disappeared by the end-Guadalupian but experienced a relatively long-term press extinction spanning the entire Guadalupian in the Gondwanan Realm. The end-Guadalupian brachiopod diversity fall is not well reflected at the timescale used here in the Paleoequatorial Realm because the life-depleted early Wuchiapingian was overlapped by a rapid radiation phase in the late Wuchiapingian. The Guadalupian fall appears to be related to the dramatic reduction of habitat area for the brachiopods, which itself is associated with the withdrawal of seawater from continental Pangea and the closure of the Sino-Mongolian seaway by the and-Guadalupian.

Permian brachiopods from the Baoshan and Simao Blocks in Western Yunnan, China

This paper describes and illustrates four Permian brachiopod assemblages (Assemblages A–D) from western Yunnan, southwest China. Assemblage A occurs in the basal part of the Yongde Formation in the Xiaoxinzhai section in the southern Baoshan Block and is assignable to the latest Artinskian or earliest Kungurian. Assemblage B occurs about 50 m stratigraphically above Assemblage A in the same section, and is of Chihsian (Kungurian–Roadian) age. Assemblage C was collected from the Yongde Formation in the Anpaitian section in the southern Baoshan Block and is most likely Roadian to Wordian in age. Assemblage D came from the Longtan Formation in the Simao Block and is of late Wuchiapingian in age. The brachiopod faunas from the Yongde Formation of the Baoshan Block, as a whole, is dominated by species characteristic of the Cathaysian Province, although some links with Peri-Gondwanan faunas are also evident. In contrast, Assemblage D from the Simao Block is characterised exclusively by taxa of the Cathaysian Province. New taxa described and illustrated are Neochonetes (Huangichonetes) inflatus n. sp. Costatumulus minor n. sp. and Celebetus yunnanensis n. sp.

Isogramma Meek and Worthen, 1870 (Dictyonellida, Brachiopoda) from the upper Palaeozoic of East Asia: implications for biogeography and evolutionary trends

Isogramma manchoukuoensis from the Upper Carboniferous of northeast China is redefined based on re-examination of the type specimens. Isogramma specimens from the Middle Permian of northeastern Japan are reassigned to I. aff. paotechowensis. A new family, Schizopleuroniidae, is proposed to include Schizopleuronia, but excludes Megapleuronia, which belongs to the Megapleuroniidae Liao, 1983. The family Isogrammidae is considered to be a transitional group in the eichwaldid-isogrammid-schizopleuronid evolutionary lineage within the Dictyonellida. A review of the global distribution of Isogramma species reveals that the genus has a total of 56 species ranging from the Mississippian (Early Carboniferous) to the Lopingian (Late Permian). Isogramma diversified rapidly after its origination in the middle Viséan and its species diversity remained high throughout the Mississippian. The genus possibly suffered a severe mid-Carboniferous boundary mass extinction, with no Early Carboniferous species surviving this event. Bashkirian Isogramma species show low diversity, followed by a global recovery in the Moscovian. During the latest Carboniferous Isogramma became highly diversified again. At the Carboniferous–Permian (C/P) transition Isogramma underwent another dramatic diversity drop, followed by several stepwise declines in diversity during the Early–Middle Permian. The Wuchiapingian I. sinosa is the last Isogramma species.

Ukraine was the possible centre of origin for Isogramma. From Ukraine Isogramma spread over the Moscow Basin of Russia, Central Europe (Germany, Austria), South Europe (Spain) and West Europe (England, Ireland and Scotland), and migrated to the North American midcontinent and South China during the late Viséan (Early Carboniferous). In Europe, Isogramma migrated to Spain and eastern Europe (Serbia) in the Moscovian, from there it then dispersed into Central Asia (Uzbekistan and Kazakhstan) in the Kasimovian-Gzhelian. In the Palaeo-Tethys Isogramma migrated from South China to northeast and northwest China in the Moscovian, spread over the North China Block during the C/P transition, moved to Russian Siberia, Japan and the Qiangtang terrane of the Palaeo-Tethys during the Early–Late Permian. In North America Isogramma spread over the midcontinent during the Late Carboniferous and Early–Middle Permian and migrated to South America (Bolivia) in latest Carboniferous. Biogeographically, Isogramma was confined principally to the palaeo-tropical and warm to temperate zones throughout the Late Palaeozoic, with the possible exception of the Artinskian, as a questionable species of the genus also occurs in the Transbaikal region of southeast Russia.

Life crises on land across the Permian–Triassic boundary in South China

The western Guizhou and eastern Yunnan area of southwest China commands a unique and significant position globally in the study of Permian–Triassic boundary (PTB) events as it contains well and continuously exposed PTB sections of marine, non-marine and marginal-marine origin in the same area. By using a range of high-resolution stratigraphic methods including biostratigraphy, eventostratigraphy, chronostratigraphy and chemostratigraphy, not only are the non-marine PTB sections correlated with their marine counterparts in the study area with high-resolution, the non-marine PTB sections of the study area can also be aligned with the PTB Global Stratotype Section and Point (GSSP) at Meishan in eastern China. Plant megafossils (“megaplants”) in the study area indicate a major loss in abundance and diversity across the PTB, and no coal beds and/or seams have been found in the non-marine Lower Triassic although they are very common in the non-marine Upper Permian. The megaplants, however, did not disappear consistently across the whole area, with some elements of the Late Permian Cathaysian Gigantopteris flora surviving the PTB mass extinction and locally even extending up to the Lower Triassic. Palynomorphs exhibit a similar temporal pattern characterized by a protracted stepwise decrease from fern-dominated spores in the Late Permian to pteridosperm and gymnosperm-dominated pollen in the Early Triassic, which was however punctuated by an accelerated loss in both abundance and diversity across the PTB. Contemporaneous with the PTB crisis in the study area was the peculiar prevalence and dominance of some fungi and/or algae species.

The temporal patterns of megaplants and palynomorphs across the PTB in the study area are consistent with the regional trends of plant changes in South China, which also show a long-term decrease in species diversity from the Late Permian Wuchiapingian through the Changhsingian to the earliest Triassic, with about 48% and 77% losses of species occurring respectively in the end-Wuchiapingian and end-Changhsingian. Such consistent patterns, at both local and regional scales, contradict the hypothesis of a regional isochronous extinction of vegetation across the PTB, and hence call into question the notion that the end-Permian mass extinction was a one-hit disaster. Instead, the data from the study area and South China appears more consistent with a scenario that invokes climate change as the main driver for the observed land vegetation changes across the PTB in South China.

Latest Guadalupian brachiopods from the Guadalupian/Lopingian boundary GSSP section at Penglaitan in Laibin, Guangxi, South China and implications for the timing of the pre-Lopingian crisis

A brachiopod fauna comprising nine species in eight genera from three closely spaced stratigraphic horizons of the same stratigraphic section is described for the first time from the Laibin Limestone in the uppermost part of the Maokou Formation in the Guadalupian/Lopingian (G/L) GSSP section at Penglaitan, Guangxi Autonomous Region, South China. The brachiopod assemblages are bracketed between two conodont zones: Jinogondolella xuanhanensis Zone below and Jinogondolella granti Zone above and, therefore, they can be safely assigned to the latest Capitanian in age. However, all but one of the nine brachiopod species from the Laibin Limestone carry strong early Lopingian (Wuchiapingian) aspect. Thus, the discovery of this brachiopod fauna not only suggests that some Lopingian brachiopod species had already appeared in the late Guadalupian (Capitanian); more importantly, it has also highlighted the fact that both the previously noted pre-Lopingian life crisis (or end-Guadalupian or Middle Permian mass extinction) and Lopingian recovery/radiation actually occurred in late Capitanian times, sometime before the G/L chronostratigraphic boundary. So far, the Penglaitan GSSP section provides the highest-resolution disappearance patterns of different fossil groups around the G/L boundary.

Evolution in a cold climate

A brief appraisal of marine fossils from high latitudes and episodically cold climate especially in east Australia and New Zealand during Late Palaeozoic and Early Mesozoic times shows patterns of evolution and survival that differ from those adduced for the palaeotropics and Northern Hemisphere. Examples taken from amongst phyla Scyphozoa, Bryozoa, Brachiopoda and Classes Bivalvia and Class Cephalopoda suggest these attributes:
1. Evolution and demise of species and genera proceeded at a rate close to that known for palaeotropical and Northern Hemisphere macro-invertebrates, but involved fewer families and orders.
2. Possibly, intraspecific variation was greater amongst southern palaeohemisphere Permian species than in those of the Permian palaeotropics.
3. There was no proven diminution of life at the end of the Guadalupian (Middle Permian) at southern high latitudes, where however the fossil record is meagre for this interval. Younger Wuchiapingian and Changhsingian faunas were moderately diverse.
4. There is no evidence for a high latitude Southern Hemisphere anoxic event in the Early Triassic despite claims of a world-wide anoxic interval. Nor has any substantial volcanic eruption or bolide impact left any marked traces in the sedimentary record.
5. As a consequence, some major groups such as Bryozoa and Conulariida (Staurozoa) survived the end- Permian extinction shock in the Southern Hemisphere.
6. Other major groups appear to have survived better in the south than in the north, notably, mollusc Bivalvia and Cephalopoda. It therefore appears likely that Triassic seas were restocked substantially from the Southern Hemisphere and that the Permian extinction shock was asymmetric with respect to latitudes in its distribution and affect.