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.

Early Permian Jakutoproductus (Productida: Brachiopoda) from Patagonia, Argentina

The productid brachiopod genus Jakutoproductus, dominant in the Early Permian marine faunas of northeastern Siberia, is described for the first time from the southern hemisphere. Jakutoproductus australis sp. nov. is described from the Rio Genoa Formation, Chubut Province (Patagonia), Argentina. The age of the Patagonian species is considered to be Sakmarian (Early Permian), possibly Sterlitamakian.

Permian brachiopods from the Tupe formation, San Juan Province, Precordillera, Argentina

The brachiopod fauna from the Tupe Formation at La Herradura Creek, located on the west flank of Perico Hill, San Juan Province, Argentina, palaeogeographically belongs to the western sector of the Paganzo basin ('Guandacol embayment'). The stratigraphical section of the Tupe Formation at La Herradura Creek is the stratotype of the Tivertonia jachalensis-Streptorhynchus inaequiornatus biozone, was previously regarded as being of Late Carboniferous age but here is assigned to the earliest Permian (Asselian). We describe and review the biozone assemblage, which consists of Streptorhynchus inaequiornatus, Tivertonia jachalensis, Kochiproductus sp., Costatumulus sp., Coronalosia argentinensis, Tupelosia paganzoensis, Trigonotreta pericoensis, Septosyringothyris sp. aff. Septosyringothyris jaguelensis and Crurithyris? sp. This brachiopod assemblage is related to Indian and Australian Early Permian faunas and its presence in the La Herradura Creek section provides new evidence in support of an Asselian (Early Permian) age for the Tivertonia jachalensis-Streptorhynchus inaequiornatus biozone. This assemblage is also important for intra- and inter-basinal correlation because several of its characteristic species have been identified from other sections of the Paganzo basin and the Riacuteo Blanco basin. The proposed age for this biozone is consistent with the age of palynological data from slightly above the marine faunas from the stratotype locality.

Lopingian (Late Permian) stratigraphy, sedimentation and palaeobiogeography in southern Tibet

Investigations of the Permian-Triassic sections and limestone blocks scattered in the Indus-Tsangbo Suture Zone in southern Tibet show widespread distribution of the Lopingian strata. The Lopingian deposits mostly contain rich brachiopod fossils and characteristic conodonts of the Mesogondolella shenz Zone of latest Changhsingian age in the topmost part. Brachiopod assemblages are largely comparable with those known from the upper Wargal and Chhidru Formations of the Salt Range, Pakistan, the Zewan Formation of Kashmir, the upper part of the Kuling Group in Spiti of India and the Hardman Formation of Western Australia. A revised Lopingian (Late Permian) age is proposed for the Selong Group and its equivalents in southern Tibet. The Lopingian deposits in southern Tibet can be grouped into three different sedimentary types, each of which reflects different sedimentary environments from coastal to continental shelfal settings on the northern peri-Gondwanan margin. The Qubu-type sequence represents marine coastal and proximal barrier-lagoon sediments during a gradual sea-level rise. Micaceous sandstone and shale of regressive origin, with abundant palynomorphs and acritarches, developed during the Late Lopingian sea-level lowstand, which is followed by a major rapid transgression at the very end of Permian. The Selong-type sequence in the Selong area consists of bioclastic limestone and calcareous shale in the lower part, and crinoid grainstone in the upper part. The latter part is believed to have been formed in a high-energy inner shelf shoal setting. The Chitichun-type sequence, sporadically distributed along the Indus-Tsangbo suture zone as small limestone blocks, consists of pure bioclastic sparite with the ammonoid Cyclolobus fauna. It is interpreted as the break-up products of sea-mounts and/or small isolated carbonate build-ups developed on the outer shelfal settings.

Pennsylvanian (Carboniferous) brachiopods from the Itaituba Formation of the Amazon Basin, Brazil

A new brachiopod fauna is described from the lower Itaituba Formation at the Caima Quarry 2 section in the Itaituba area, Amazon Basin, Brazil. The Amazonoproductus amazonensis-Anthracospirifer oliveirai Assemblage is proposed for this fauna, which is considered early Pennsylvanian (Morrowan) as constrained by associated conodont and fusulinacean faunas. Nine brachiopod taxa are described herein, including Amazonoproductus amazonensis gen. et sp. nov., and Buxtonioides itaitubensis sp. nov. and Linoproductus caima sp. nov. The new tribe Marginovatini of the Linoproductoidea (the Productida) is also proposed.

Carboniferous and Permian Rugosochonetidae (Brachiopoda) from West Spitsbergen

The rugosochonetid brachiopod species Lissochonetes geinitzianus from the Kazimovian of the Nordenskioldbreen Formation, and Dyoros (Dyoros) mucronata sp. nov., Dyoros (Dyoros) spitzbergianus and Lissochonetes superba from the Artinskian to latest Permian Kapp Starostin Formation in West Spitsbergen are described and figured. Dyoros is generally restricted to the Boreal Realm, whereas Lissochonetes is mostly distributed in the Boreal Realm, but occasionally present in the Palaeoequatorial and Gondwanan Realms

Uppermost Mississippian Brachiopods from the Basal Itaituba formation of the Amazon Basin, Brazil

This paper describes 19 brachiopod species (including six indeterminate species) in 15 genera and one indeterminate genus from the basal Itaituba Formation at the Caima Quarry 1 section of Itaituba, Amazon Basin, Brazil. The faunal correlations of the brachiopods and the associated fusulinids and conodonts indicate a late Chesterian (late Serpukhovian) age for the described fauna, therefore confirming for the first time the presence of uppermost Mississippian rocks in the Amazon Basin. A new species, Composita caimaensis, is created, and two species, Inflatia cf. gracilis and Marginovatia cf. catinulus, are described for the first time from the Amazon Basin. The Amazon brachiopods appear to be of strong affinity with coeval faunas of the North American midcontinent.

Carboniferous brachiopods from the "Levipustula Fauna" in central-western Argentina : biostratigraphic, palaeoclimatic, and palaeobiogeographic implications

The “Levipustula fauna” is a relatively diverse fossil assemblage composed of brachiopods, bivalves, bryozoans, gastropods and crinoids that appears in glaciomarine sequences related to the Carboniferous glacial event that affected the central-western Argentinean basins. Brachiopods that characterize this fauna have been studied in some of the classical localities of the Argentine Precordillera in San Juan province.

In the Hoyada Verde Formation, the “Levipustula fauna” is usually located immediately above the glacial diamictite horizons and appears to be associated with the mudstones facies of the postglacial transgression. From a palaeoecological study, three brachiopod subfaunas are distinguished in this formation: the lower “Neospirifer” (“Trigonotretidae” gen. nov.) subfauna, above which the more diversified Kitakamithyris subfauna occurs, and the upper Levipustula subfauna, with the dominant species being attributed to Levipustula levis Maxwell. The Hoyada Verde fauna, as well those identified in the La Capilla Formation (Cerro El Morado locality), have been proposed as the typical “Levipustula fauna”. However, in the Leoncito Formation, the “Levipustula fauna” occurs in sandstones horizons located below the glacial diamictite beds. This fauna is poorly diversified and the brachiopods are characterized by “Neospirifer” (“Trigonotretidae” gen. nov.)-Septosyringothyris assemblage and Levipustula is not abundant. This fauna has been considered a colder “pre-interglacial fauna”.

The significant taxonomic, palaeoecologic and taphonomic variations of the “Levipustula fauna”, as well as its position in the glacial sequences, suggest an important palaeoenvironmental control related to Carboniferous glacial dynamics. From the palaeobiogeographic viewpoint, this fauna shows the highest affinity with the Eastern Australian basins from where it was previously described.

Roadian – Wordian (Guadalupian, Middle Permian) global palaeobiogeography of brachiopods

A database of 4471 Roadian–Wordian (Guadalupian, Middle Permian) occurrences of 381 brachiopod genera in 44 different operational geographical units (stations) was analyzed by both Q-mode and R-mode quantitative methods. Four distinct brachiopod biogeographical realms and nine provinces, and 11 brachiopod associations are recognized. The Boreal Realm in the Northern Hemisphere includes the Verkolyman Province in the northern and northeastern Siberian Platform and the eastern European Province in the Ural seaway between the European and Siberian platforms. Both provinces are characterized by containing typical Boreal cold-water brachiopod associations. The Gondwanan Realm in the south also includes two provinces. The Austrazean Province in eastern Australia and New Zealand is probably the most stable province throughout the Permian and characterized by typical Gondwanan brachiopod associations. The Westralian Province centered in Western Australia is also characterized by typical Gondwanan brachiopods, but also demonstrates biogeographical links with the Tethyan stations. The Palaeoequatorial Realm located mainly in the palaeotropical zone contains highly diverse and abundant brachiopod faunas. Two regions/subrealms and four provinces are recognized within this realm. The North America Subrealm contains a distinct Grandian Province characterized by many endemic brachiopod genera and a few coldwater genera. East-central Alaska and Yukon Territory may constitute another brachiopod province. All the stations in the Tethyan Ocean (both Palaeotethys and Neotethys) constitute a distinct Asian–Tethyan Region/ Subrealm and incorporate three different provinces. The Cathaysian Province is comprised of the stations in South China and its surrounding terranes/blocks and a few stations in the northern and western margin of the Palaeotethys. Two transitional provinces (Sino–Mongolian–Japanese Province and Cimmerian Province) in the northern and southern temperate zones are also recognizable. The brachiopod fauna from the Mino Belt in Japan is well distinguished from those from other regions, and is hence assigned to the palaeoceanic Panthalassan Realm. Principal coordinates analysis and minimum spanning tree analysis suggest that a latitude-related thermal gradient was the major control for the palaeobiogeography of Roadian–Wordian global brachiopod faunas and for the latitudinal of pattern of decreasing brachiopod generic diversities from the equator to the poles. In addition, geographic separation and oceanic currents may also have played some role in the spatial distribution of brachiopods during Roadian–Wordian times.

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.

Pennsylvanian brachiopods from the Geumcheon-Jangseong formation, Pyeongan supergroup, Taebaeksan Basin, Korea

We provide the first detailed systematic taxonomy and paleoecological investigation of late Paleozoic brachiopod faunas from Korea. Specifically, we focus on the brachiopods from the Geumcheon-Jangseong Formation, the lower part of the Pyeongan Supergroup in the Taebaeksan Basin. The formation yields a variety of marine invertebrate fossils, including brachiopods, molluscs, echinoderms, corals, fusulinids, and conodonts. Diverse brachiopods are described from six siliciclastic horizons of the formation at three localities, including 23 species belonging to 20 genera with two new species: Rhipidomella parva n. sp. and Stenoscisma wooi n. sp. Three brachiopod assemblages of the late Moscovian (Pennsylvanian) age are recognized based on their species compositions and stratigraphic distributions, namely the Choristites, Rhipidomella, and Hustedia assemblages. The brachiopod faunal composition varies within each assemblage as well as between the Assemblages, most likely reflecting local paleoenvironmental and hence paleoecological differences. The Choristites Assemblage includes relatively large brachiopods represented by Derbyia, Choristites, and Stenoscisma and may have inhabited open marine to partly restricted marine environments, whereas the Rhipidomella and Hustedia Assemblages consist of a small number of small-sized brachiopods living in lagoonal environments. The Choristites Assemblage shows a close affinity with Moscovian brachiopod assemblages in the eastern Paleo-Tethys regions, especially the Brachythyrina lata–Choristites yanghukouensis–Echinoconchus elegans Assemblage of North China, whereas the Rhipidomella and Hustedia assemblages both exhibit strong endemism.

Controls on body size during the Late Permian mass extinction event

This study examines the morphological responses of Late Permian brachiopods to environmental changes. Quantitative analysis of body size data from Permian–Triassic brachiopods has demonstrated significant, directional changes in body size before, during and after the Late Permian mass extinction event. Brachiopod size significantly reduced before and during the extinction interval, increased for a short time in more extinction-resistant taxa in the latter stages of extinction and then dramatically reduced again across the Permian ⁄ Triassic boundary. Relative abundances of trace elements and acritarchs demonstrate that the body size reductions which happened before, during and after extinction were driven by primary productivity collapse, whereas declining oxygen levels had less effect. An episode of size increase in two of the more extinction-resistant brachiopod species is unrelated to environmental change and possibly was the result of reduced interspecific competition for resources following the extinction of competitors. Based on the results of this study, predictions can be made for the possible responses of modern benthos to present-day environmental changes.

Discovery of Early Devonian strata in northern Qiangtang, Tibet, and the establishment of the Pingshagou Formation

Although it is important to determine whether there exist sedimentary strata older than the Upper Permian in the northern Qiantang area in Tibet, there has been no report of such old strata in this area. During the geological mapping of the Mayigangri area, we discovered strata that contain the bivalves Eoschizodus roemeri (Beushausen), E. minor (Beushausen) , E. infiatus (Roemer), Actinodesma (Actinodesma) cf . maneiforme Sandberger, A . (Actindesma) cf . vespertilio Maurer, and the brachiopod Huananochonetes subquadratus Sun & Chen. These fossils indicate a late Early Devonian age (Emsian) , thus the strata represent the first discovered Lower Devonian rocks in northen Qiangtang, confirming the existence of strata older than the Upper Permian. A new stratigraphic term, the Pingshagou Formation , is introduced. The new data provide constraints on the tectonic, palaeogeographic, and palaeobiogeographic history of the north Qiangtang area in the Early Devonian.

The Lopingian series in the Lhasa Block, Tibet and its palaeogeographical implications

Latest investigation indicates that the Lopingian Series including both terrestrial and marine deposit s are developed in the Lhasa Block. The marine Lopinigian Series in the Lhasa Block contains the compound coral Waagenophyllum, fusulinid Reichelina, and foraminifer Colaniella faunas , and the terrestrial Lopingian Series is characterized by both Cathaysian floras and mixed floras consisting of Gondwanan element s such as Glossopteris , Noeggerathiopsis, Phyllotheca and Cathaysian elements such as Pecopteris ,Sphenopteris. An anlaysis of the Lopingian sequences in the Lhasa Block reveals that it experienced a regression stage f rom Guadalupian to Lopingian. By contrast , the Himalayan Tethys Zone south to the Lhasa Block is characterized by typical Gondwanan Glossopteris flora , coldwater brachiopod and solitary coral faunas. In addition, the Lopingian sequence in the Himalayan Tethys Zone reflect s a transgressive process from the terrestrial Qubu Formation to the shallow marine Qubuerga Formation. Therefore, the Lhasa Block shows significant differences in both biota and depositional features from the Himalayan Tethys Zone during the Lopingian, which implies that the Lhasa Block had rifted from the northern periGondwanan margin before the Lopingian.