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.

Late Palaeozoic sequence stratigraphy and brachiopod faunas of the Tarim Basin, Northwest China

This thesis deals with the stratigraphy and brachiopod systematic palaeontology of the latest Devonian (Famennian) to Early Permian (Kungurian) sedimentary sequences of the Tarim Basin, NW China. Brachiopod faunas of latest Devonian and Carboniferous age have been published or currently in press in the course of the Ph.D candidature and are herein appendixed, while the Early Permian brachiopod faunas are systematically described in this thesis. The described Early Permian brachiopod faunas include 127 species, of which 29 are new and 12 indeterminate, and six new genera (subgenera) are proposed; Tarimella, Bmntonella, Marginifera (Arenaria), Marginifera (Nesiotia), Baliqliqia and Ustritskia. A new integrated brachiopod biostratigraphical zonation scheme is proposed, for the first time, for the latest Devonian-Early Permian sequences of the entire Tarim Basin on the basis of this study as well as previously published information (including the Candidate's own published papers). The scheme consists of twenty three brachiopod acm biozones, most of which replace previously proposed assemblage or assemblage zones. The age and distribution of these brachiopod zones within the Tarim Basin and their relationships with other important fossil groups are discussed. In terms of regional correlations and biostratigraphical affinities, the Late Devonian to Early Carboniferous brachiopod faunas of the Tarim Basin are closest to those from South China, while the Late Carboniferous faunas demonstrate strong similarities to coeval faunas from the Urals, central Asia, North China and South China. During the Asselian-Sakmarian, strong faunal links between the Tarim Basin and those of the Urals persisted, while at the same time links with central Asia, North China and South China weakened. On the other hand, during the Artinskian-Kungurian times, affinities of the Tarim faunas with the Urals/Russian Platform rapidly reduced, when those with peri-Gondwana (South Thailand, northern Tibet) and South China increased. Thirty lithofacies (or microfacies) types of four facies associations are recognised for the Late Devonian to early Permian sediments. Based on detailed lithostratigraphy, biostratigraphy and facies analysis, 23 third-order sequences belonging to four supcrsequences are identified for the Late Devonian to Early Permian successions, from which sea-level fluctuation curves are reconstructed. The sequence stratigraphical analysis reveals that four major regional regressions, each marking a distinct supersequence boundary, can be recognised; they correspond to the end-Serpukhovian, end-Moscovian, late Artinskian and end-Kungurian times, respectively. The development of these sequences is considered to have been formed and regulated by the interplay of both eustasy and tectonism. Using the system tract of a sequence as the mapping time unit, a succession of 47 palaeogeographical maps have been reconstructed through the Late Devonian to Early Permian. These maps reveal that the Tarim Basin was first immersed by southwest-directed (Recent geographical orientation) transgression in the late Famennian after the Caledonian Orogeny. Since then, the basin had maintained its geometry as a large, southwest-mouthed embayment until the late Moscovian when most areas were the uplifted above sea-level. The basin was flooded again in late Asselian-Artinskian times when a new transgression came from a large epicontinental sea lying to its northwest. Thereafter, marine deposition was restricted to local areas (southwestern and northwestern margins until the late Kungurian, while deposition of continental deposits prevailed and continued through the Middle and late Permian into the Triassic.

Late palaeozoic global changes affecting high-latitude environments and biotas : an introduction

The Late Palaeozoic Ice Age (LPIA), spanning approximately from ~320 Ma (Serpukhovian, late Mississippian) to 290 Ma (mid-Sakmarian, Early Permian), represents the vegetated Earth’s largest and most long-lasting regime of severe and multiple glaciations, involving processes and patterns probably comparable to those of the Last Ice Age. Accompanying the LPIA occurred a number of broadly synchronous global environmental and biotic changes. These global changes, as briefly reviewed and summarized in this introductory paper, comprised (but are not limited to) the following: massive continental reorganization in the lead up to the final assembly of Pangea resulting in profound changes in global palaeogeography, palaeoceanography and palaeobiogeogarphy; substantially lowered global atmospheric carbon dioxide concentrations (pCO₂), coupled with an unprecedented increase in atmospheric oxygen concentrations reaching Earth's all-time high in its last 600 million year history; sharp global temperature and sea-level drops (albeit with considerable spatial and temporal variability throughout the ice age); and apparently a prolonged period of global sluggish macro-evolution with both low extinction and origination rates compared to other times. In the aftermath of the LPIA, the world's climate entered into a transitional climate phase through the late Early to Middle Permian before its transformation into a greenhouse state towards the end-Permian. In recent years, considerable amount of data and interpretations have been published concerning the physical evidence in support of the LPIA, its broad timeframe and eustatic and ecosystem responses from the lower latitudes, but relatively less attention has been drawn to the impact of the ice age on late Palaeozoic high-latitude environments and biotas. It is with this mission in mind that we have organized this special issue, with the central focus on late Palaeozoic high latitude regions of both hemispheres, that is, Gondwana and northern Eurasia. Our aim is to gather a set of papers that not only document the physical environmental changes that had occurred in the polar regions of Gondwana and northern Eurasia during the LPIA, but also review on the biotic responses at different taxonomic, ecological and spatial scales to these physical changes in a refined chronological timeframe.

This introductory paper is designed to provide a global context for the special issue, with a brief review of key late Palaeozoic global environmental changes (including: changes in global land-sea configurations, atmospheric chemistry, global climate regimes, global ocean circulation patterns and sea levels) and large -scale biotic (biogeographic and evolutionary) responses, followed by a summary of what we see as unresolved scientific issues and various working hypotheses concerning late Palaeozoic global changes and, in particular, the LPIA, as a possible reference to future research.

A preliminary phylogenetic study of late Palaeozoic spiriferoid brachiopods using cladistic and Bayesian approaches

Abstract The brachiopod Superfamily Spiriferoidea diversified greatly and was widely distributed in the late Palaeozoic (Carboniferous–Permian), and yet its phylogeny has been seldom investigated with analytical methods. This is reflected in the current flux of very different classification schemes for this superfamily. This paper provides the first attempt to investigate the phylogenetic relationships of spiriferoid brachiopods through both cladistic and Bayesian analyses involving 24 discrete and continuous characters. The continuous characters, from morphometric data, have been separately discretized using the gap weighting method, and the ‘as such’ option in TNT. Our results highlight the potential significance of continuous characters in reconstructing and elucidating phylogenies, as much as qualitative characters. Building on the outcomes of the analyses, we also briefly evaluate existing classification schemes of Spiriferoidea. We found that none of the existing classifications fully reflect the phylogeny properly; major families within the superfamily, such as Spiriferidae, Choristitidae, and Trigonotretidae, turned out to be polyphyletic. Although this study is considered preliminary, due to the selection of and restriction to certain taxa, combined with the use of a relatively small number of characters, it nevertheless demonstrates that potentially the true phylogenetic relationships of spiriferoid taxa sharply contrast with any of the existing classification schemes. This highlights the need to develop an alternative scheme that takes into account a more comprehensive range of phylogenetic variables.

Palaeobiogeographic distribution patterns and processes of neochonetes and fusichonetes (brachiopoda) in the late palaeozoic and earliest mesozoic

The global palaeobiogeographic distributions of two resembling genera, Neochonetes and Fusichonetes (Brachiopoda), from the Carboniferous to Griesbachian are analysed. This analysis provides insight into the biotic response of two related genera to changing palaeoclimate, regional tectonics, and environmental crises. Neochonetes originated in the equatorial area in the Mississippian, and it mostly retained this position during the peak of the glaciation in the Carboniferous–Permian ice age (namely in the Pennsylvanian). Neochonetes then dispersed globally during the Cisuralian when the climate became warmer and the ice sheet started to retreat. In the Guadalupian and Lopingian, following the closure of the Ural seaway at the end of the Cisuralian and the regression at the end-Guadalupian, Neochonetes almost disappeared in the western part of Gondwana. Subsequently during the Lopingian the genus retracted to the middle- and low-latitude Palaeo-Tethys and Tethys. In comparison, Fusichonetes originated in the equatorial area in the late Guadalupian and was still present in that area in the Lopingian. Both genera occurred only in South China in the Griesbachian. It is inferred that this could be related, not only to the deteriorated palaeoenvironmental conditions (e.g., anoxia, global warming) leading up to the extinction of most of the Neochonetes and Fusichonetes species in other areas, but also to the better physiological adaptation of the smaller shells of Neochonetes and Fusichonetes species in South China.

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.

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.

The Permian brachiopod genus Trigonotreta Koenig and its occurrence in Argentina

A review of the Permian spiriferid brachiopod genus Trigonolrela and its occurrence in the Early Permian of Argentina is provided herein. Several species are known from the Late Palaeozoic sequences of the Argentinean Precordillera. These include Trigonolrela sp. and Trigonolrela riojanensis (Lech and Acenolaza), from the Rio del Peii.on Formation (Rio Blanco Basin), Trigonolrela pericoensis (Leanza), from the Tupe Formation at the La Herradura creek locality (Paganzo Basin) and Trigonolrela sanjuanensis (Lech and Acenolaza), from Del Salto Formation (Calingasta-Uspallata Basin). These species are characterised by being small to medium sized, relatively transverse, with cardinal extremities often strongly angular. Costae are weakly bifurcated and superimposed on weakly developed lateral flank plications adjacent to the fastigium and sulcus. The Argentinean species are close to the oldest known Indian species of the genus, Trigonotreta hesdoensis (Salmi and Dutt), particularly with respect to the nature of its weakly fasciculated costae. Further study will refine the details of the relationship of the South American species with those from elsewhere in Gondwana and may permit the recognition of a distinctive lineage. The presence of the genus in Argentina in the earliest Permian is an important palaeobiogeographical observation that raises questions about the probable migration routes of the genus from the western Gondwanan South American margin to eastern Australia and India. The Precordilleran region appears to be the likely site of the first appearance of Trigonolrela. Species with relatively simple costae appeared first. These gave rise to more complex species with a more elaborate costal pattern indicating an evolutionary progression through time.

Palaeontology, the biogeohistory of Victoria

The broad-scale distribution of fossils within Victoria is controlled by general global patterns in the biological evolution of life on Earth, the local development and environmental evolution of habitats, and the occurrence of geological processes conducive to the preservation of fossil floras and faunas. Early Palaeozoic fossils are mostly marine in origin because of the predominance of marine sedimentary rocks in Victoria and because life on land was not significant during most of this time interval. Middle Palaeozoic sequences have both terrestrial and marine fossil records. Within Victoria, marine rocks are only very minor components of strata deposited during the late Palaeozoic, so that few marine fossils are known from this time period. A similar situation existed during most of the Mesozoic except towards the end of this era when marine conditions began to prevail in the Bass Strait region. During long intervals in the Cainozoic, large areas of Victoria were flooded by shallow-marine seas, particularly in the southern basins of Bass Strait, as well as in the northwest of the State (Murray Basin). Cainozoic sediments contain an extraordinary range of animal and plant fossils. During the Quaternary, the landscape of Victoria became, and continues to be, dominated by continental environments including, at times, extensive freshwater lake systems. Fossil floras and faunas from sediments deposited in these lake systems and from other continental sediments, as well as from Quaternary sediments deposited in marginal marine environments, collectively record a history of rapid fluctuations in climate and sea level.

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.

Taxonomic review and evolutionary trends of Levipustulini and Absenticostini (Brachiopoda) from Argentina : palaeobiogeographic and palaeoclimatic implications

The diagnosis and composition of the brachiopod Tribe Levipustulini Lazarev, 1985 is reviewed, leading to a detailed revision of the genera Levipustula Maxwell, 1951 and Lanipustula Klets, 1983, as well as a review of previous records of the species Levipustula levis Maxwell from Australia and Argentina. The presence of Lanipustula patagoniensis Simanauskas in Patagonia is confirmed with additional topotypic material described and illustrated. Based on this review, we reassign Levipustula levis from New South Wales, Australia to Lanipustula. Two new species, Lanipustula kletsi from the middle Pennsylvanian of Patagonia and the Absenticostinin Absenticosta bruntoneileenae from the latest Viséan of western Argentina, are proposed. Abstenticosta bruntoneileenae is suggested as a possible ancestral stock of the Patagonian Levipustulini through the lineage Lanipustula-Verchojania-Jakutoproductus-Piatnitzkya (Serpukhovian-middle Artinskian). The development of similar phylogenetic lineages of Levipustulini in high latitude regions of both northern and southern hemispheres (such as Siberia in Northeast Asia and Patagonia in southwestern Gondwana) is here interpreted as a consequence of parallel evolution. The progressive palaeobiogeographic isolation of Patagonia from mainland South America, coupled with its southward drift under cold palaeoclimatic conditions during middle Carboniferous-earliest Permian times, is proposed to have triggered the Levipustulini vicariance.

An integrated multidisciplinary palaeontological study of selected permian arctic brachiopods

This research discussed various topics related to Late Palaeozoic spiriferid brachiopod fossils. The outcomes include taxonomic and biostratigraphic revision of Permian brachiopods from Spitsbergen, palaeogeographical/palaeoenvironmental reconstruction of northern margin of Pangea during the Permian, interpretation of phylogeny among spiriferoidean brachiopods, and analysis of brachiopod shell morphology through new three-dimensional techniques.