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.

Discovery of early paleozoic strata in south of Qiangtang, Northern Tibet and its significance

Due to the lack of fossil evidence, the age of Amugang Group, previously referring to the large area of metamorphic formations developed along the south of Longmucuo-Shuanghu suture in southern Qiangtang area, northern Tibet, has been under heated debate. Some geologists were convinced that the age of metamorphic formations was of Pre-Devonian according to the emergence of nonmetamorphic fossil-beating Devonian in Chasang area, north of the suture. Most geologists believed that its age was of Pre-Sinian. From 2003 to 2004, we collected a large amount of Nautiloid and Graptolite fossils in upper metamorphic formations of Mayigangri area and determined the age to be middle and Late Ordovician-Silurian. It is the first time to discover the Early Paleozoic strata based on reliable fossils, which not only provides evidence to its age ascription, classification of the strata and its correlation to adjacent areas but also offers data to lithofacies, palaeogeography and tectonic pattern of studied area.

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.

Pliocene coastal palaeomorphology and ostracod faunas of the Bass Strait hinterland, southeast Australia

Early to late Pliocene sedimentary strata present across the northern Bass Strait hinterland, southeastern Australia yield extensive fossil proxy data relevant to the interpretation of high sea level coastal palaeomorphology. Within the Pliocene Whalers Bluff Formation exposed in coastal cliffs near the township of Portland, Victoria, marine microfossil faunas delineate two broad cycles of deposition. Both these sedimentary cycles are bound below by unconformity surfaces. Within the lower sedimentary cycle, a basal stress-tolerant (low diversity) marginal marine microfossil fauna devoid of ostracods and suggestive of bottom-water hypoxia, is succeeded by a diverse shallow marine ostracod fauna dominated by stenohaline species indicative of a sheltered (but open) oceanic embayment. This lower sedimentary cycle has an early Pliocene (Zanclean) age. Equivalent shallow marine (e.g. coastal embayment) deposits occur broadly across the coastal hinterland of southeastern Australia-reflecting the generally higher global sea levels of this time. The upper cycle in the cliff exposures at Portland is late Pliocene (Piacenzian) in age. Equivalent deposits across the Bass Strait hinterland are restricted to former incised river valley settings. Euryhaline estuarine/coastal lagoon Ostracoda are present throughout the upper cycle in the Portland cliffs. These are associated with a low diversity microfauna at the base of the upper cycle and a high diversity microfauna towards the top of the cycle. Early Pliocene coastal marine deposits can be distinguished from late Pliocene coastal marine deposits across the northern Bass Strait hinterland on the basis of the presence or absence of certain open marine ('stenohaline') ostracod species.

Early Triassic (early Olenekian) life in the interior of East Gondwana: Mixed marine-terrestrial biota from the Kockatea Shale, Western Australia

A new terrestrial-marine assemblage from the lower beds of a thin outcrop section of the Kockatea Shale in the northern Perth Basin, Western Australia, contains a range of fossil groups, most of which are rare or poorly known from the Lower Triassic of the region. To date, the collection includes spinose acritarchs, organic-cemented agglutinated foraminifera, lingulids, minute bivalves and gastropods, ammonoids, spinicaudatans, insects, austriocaridid crustaceans, actinopterygians, a temnospondyl-like mandible, plant remains, and spores and pollen. Of these groups, the insects, crustaceans and macroplant remains are recorded for the first time from this unit. Palynomorphs permit correlation to nearby sections where conodonts indicate an early Olenekian (Smithian) age. The locality likely represents the margin of an Early Triassic shallow interior sea with variable estuarine-like water conditions, at the southwestern end of an elongate embayment within the East Gondwana interior rift-sag system preserved along the Western Australian margin. Monospecific spinose acritarch assemblages intertwined with amorphous organic matter may represent phytoplankton blooms that accumulated as mats, and suggest potentially eutrophic surface waters. The assemblage represents a mixure of marine and terrestrial taxa, suggesting variations in water conditions or that fresh/brackish-water and terrestrial organisms were transported from adjacent biotopes. Some of the lower dark shaly beds are dominated by spinicaudatans, likely indicating periods when the depositional water body was ephemeral, isolated, or subjected to other difficult environmental conditions. The biota of the Kockatea Shale is insufficiently known to estimate biotic diversity and relationships of individual taxa to their Permian progenitors and Triassic successors, but provides a glimpse into a coastal-zone from the interior of eastern Gondwana. Specialist collecting is needed to clarify the taxonomy of many groups, and comparisons to other Lower Triassic sites are required to provide insights into the pattern of biotic decline and recovery at the end-Permian crisis.

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.