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.

Brachiopods in Early Carboniferous

Changes in Mississippian global paleogeography derived from the reconfiguration of the continents, a reversal in ocean currents and global cooling. Although the tectonic and climatic changes are well-documented, their effects on the distribution of brachiopod fauna are poorly documented. Here we present systematic quantitative analyses on global paleobiogeography based on a global brachiopod database from the Mississippian (i.e., Tournaisian, Visean, and Serpukhovian). The dataset consists of 2123 species of 344 brachiopod genera from 1156 localities. Our results reveal that global provincialism was not evident during the Tournaisian and Visean Stages. Two realms, i.e., the Gondwanan and Paleoequatorial Realms, are recognized during the Tournaisian. The Paleoequatorial Realm dominates during the Visean Stage, whereas the Gondwanan Realm is not documented due to the absence of data points. In contrast to the early and middle Mississippian stages, faunal provincialism is greatly enhanced in the Serpukhovian Stage with Paleotethyan and North American realms easily distinguished. This indicates that the Rheic Ocean was closed before the Serpukhovian due to the collision between Gondwana and Laurussia, that disrupted faunal interchange between the Paleotethys and North America. In addition, the paleolatitude-related thermal gradient was enhanced and the Boreal Realm was distinguished from the Paleotethyan Realm during the onset of the Late Palaeozoic Ice Age (LPIA) in the Serpukhovian. The paleolatitude diversity gradient pattern further shows a distinct shift of diversity center from the southern tropic zone in the Tournaisian and Visean to the northern tropic zone in the Serpukhovian.

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.

The Little Ice Age: glacier variations and climate since AD 1250 [abstract]

EXTRACT (SEE PDF FOR FULL ABSTRACT): During the past hundred years, mountain glaciers throughout the world have retreated significantly from moraines built during the previous several centuries. In the 1930s, Francois Matthes of the U.S. Geological Survey concluded that the moraines represent the greatest advances of glaciers since the end of the last glacial age, some 10,000 years earlier, and informally referred to this late Holocene interval of expanded ice cover as the Little Ice Age.

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.

The Most Extensive Holocene Advance in the Stauning Alper, East Greenland, Occurred in the Little Ice Age

We present glacial geologic and chronologic data concerning the Holocene ice extent in the Stauning Alper of East Greenland. The retreat of ice from the late-glacial position back into the mountains was accomplished by at least 11 000 cal years B.P. The only recorded advance after this time occurred during the past few centuries (the Little Ice Age). Therefore, we postulate that the Little Ice Age event represents the maximum Holocene ice extent in this part of East Greenland.

Possible Effects of Anthropogenically-Increased CO2 on the Dynamics of Climate: Implications for Ice-Age Cycles

A dynamical model, developed to account for the observed major variations of global ice mass and atmospheric CO2 during the late Cenozoic, is used to provide a quantitative demonstration of the possibility that the anthropogenically-forced increase of atmospheric CO2, if maintained over a long period of time (perhaps by tectonic forcing), could displace the climatic system from an unstable regime of oscillating ice ages into a more stable regime representative of the pre-Pleistocene. This stable regime is characterized by orbitally-forced oscillations that are of much weaker amplitude than prevailed during the Pleistocene.

Population genetics of Ice Age brown bears

The Pleistocene was a dynamic period for Holarctic mammal species, complicated by episodes of glaciation, local extinctions, and intercontinental migration. The genetic consequences of these events are difficult to resolve from the study of present-day populations. To provide a direct view of population genetics in the late Pleistocene, we measured mitochondrial DNA sequence variation in seven permafrost-preserved brown bear (Ursus arctos) specimens, dated from 14,000 to 42,000 years ago. Approximately 36,000 years ago, the Beringian brown bear population had a higher genetic diversity than any extant North American population, but by 15,000 years ago genetic diversity appears similar to the modern day. The older, genetically diverse, Beringian population contained sequences from three clades now restricted to local regions within North America, indicating that current phylogeographic patterns may provide misleading data for evolutionary studies and conservation management. The late Pleistocene phylogeographic data also indicate possible colonization routes to areas south of the Cordilleran ice sheet.

Isotopic records of terrestrial ice age environments in mammoth bioapatite

Palaeoenvironments of the latter half of the Weichselian ice age and the transition to the Holocene, from ca. 52 to 4 ka, were investigated using isotopic analysis of oxygen, carbon and strontium in mammal skeletal apatite. The study material consisted predominantly of subfossil bones and teeth of the woolly mammoth (Mammuthus primigenius Blumenbach), collected from Europe and Wrangel Island, northeastern Siberia. All samples have been radiocarbon dated, and their ages range from >52 ka to 4 ka. Altogether, 100 specimens were sampled for the isotopic work. In Europe, the studies focused on the glacial palaeoclimate and habitat palaeoecology. To minimise the influence of possible diagenetic effects, the palaeoclimatological and ecological reconstructions were based on the enamel samples only. The results of the oxygen isotope analysis of mammoth enamel phosphate from Finland and adjacent nortwestern Russia, Estonia, Latvia, Lithuania, Poland, Denmark and Sweden provide the first estimate of oxygen isotope values in glacial precipitation in northern Europe. The glacial precipitation oxygen isotope values range from ca. -9.2±1.5 in western Denmark to -15.3 in Kirillov, northwestern Russia. These values are 0.6-4.1 lower than those in present-day precipitation, with the largest changes recorded in the currently marine influenced southern Sweden and the Baltic region. The new enamel-derived oxygen isotope data from this study, combined with oxygen isotope records from earlier investigations on mammoth tooth enamel and palaeogroundwaters, facilitate a reconstruction of the spatial patterns of the oxygen isotope values of precipitation and palaeotemperatures over much of Europe. The reconstructed geographic pattern of oxygen isotope levels in precipitation during 52-24 ka reflects the progressive isotopic depletion of air masses moving northeast, consistent with a westerly source of moisture for the entire region, and a circulation pattern similar to that of the present-day. The application of regionally varied δ/T-slopes, estimated from palaeogroundwater data and modern spatial correlations, yield reasonable estimates of glacial surface temperatures in Europe and imply 2-9°C lower long-term mean annual surface temperatures during the glacial period. The isotopic composition of carbon in the enamel samples indicates a pure C3 diet for the European mammoths, in agreement with previous investigations of mammoth ecology. A faint geographical gradient in the carbon isotope values of enamel is discernible, with more negative values in the northeast. The spatial trend is consistent with the climatic implications of the enamel oxygen isotope data, but may also suggest regional differences in habitat openness. The palaeogeographical changes caused by the eustatic rise of global sea level at the end of the Weichselian ice age was investigated on Wrangel Island, using the strontium isotope (Sr-87/Sr-86) ratios in the skeletal apatite of the local mammoth fauna. The diagenetic evaluations suggest good preservation of the original Sr isotope ratios, even in the bone specimens included in the study material. To estimate present-day environmental Sr isotope values on Wrangel Island, bioapatite samples from modern reindeer and muskoxen, as well as surface waters from rivers and ice wedges were analysed. A significant shift towards more radiogenic bioapatite Sr isotope ratios, from 0.71218 ± 0.00103 to 0.71491 ± 0.00138, marks the beginning of the Holocene. This implies a change in the migration patterns of the mammals, ultimately reflecting the inundation of the mainland connection and isolation of the population. The bioapatite Sr isotope data supports published coastline reconstructions placing the time of separation from the mainland to ca. 10-10.5 ka ago. The shift towards more radiogenic Sr isotope values in mid-Holocene subfossil remains after 8 ka ago reflects the rapid rise of the sea level from 10 to 8 ka, resulting in a considerable reduction of the accessible range area on the early Wrangel Island.

Arctic charr from Ennerdale water - relicts of the ice age

The work discussed in this report deals with aspects of the ecology of Arctic charr (Salvelinus alpinus, L.). The main aims of the study were: (1) To assess the relative abundance of migrants entering the River Liza and Smithy Beck in the English Lake District). (2) To assess the degree of stream specificity. (3) To determine the period of residency in the streams. (4) To obtain a better understanding of migratory behaviour. (5) To determine the growth rate of mature fish. (6) To determine the morphometric and meristic attributes. In conjunction with this work a study was carried out to investigate the feasibility of using a video recording system to monitor the migration of charr in Smithy Beck.

Evidence from northwest European bogs shows 'Little Ice Age' climatic changes driven by variations in solar activity

Fluctuations in Holocene atmospheric radiocarbon concentrations have been shown to be due to variations in solar activity. Analyses of both Be-10 and C-14 nuclides confirm that production-rate changes during the Holocene were largely modulated by solar activity. Analyses of peat samples from two intact European ombrotrophic bogs show that climatic deteriorations during the 'Little Ice Age' are associated with transitions to increasing atmospheric C-14 content due to greater C-14 production. Both ombrotrophic mires, which are positioned c. 800 km apart, register reactions to globally recorded C-14 fluctuations between AD 1449 and 1464 and an almost identical reaction between AD 1601 and 1604.

Palaeobiology of an extinct Ice Age mammal: Stable isotope and cementum analysis of giant deer teeth

The extinct giant deer, Megaloceros giganteus, is among the largest and most famous of the cervids. Megaloceros remains have been uncovered across Europe and western Asia. but the highest concentrations come from Irish bogs and caves Although Megaloceros has enjoyed a great deal of attention over the centuries, paleobiological study has focused oil morphometric and distributional work until now. This paper presents quantitative data that have implications for understanding its sudden extirpation in western Europe during a period of global climate change approximately 10.600 C-14 years ago (ca 12,500 calendar years BP). We report here the first stable isotope analysis of giant deer teeth. which we combine with dental cementum accretion in order to document age, diet and life-history seasonality from birth until death Enamel delta C-13 and delta O-18 measured in the second and third molars from seven individual giant deer Suggest a grass and forbbased diet supplemented with browse in a deteriorating. possibly water-stressed, environment, and a season of birth around spring/early summer Cementurm data indicate that the ages of the specimens ranged from 6.5 to 14 years and that they possessed mature antlers by autumn, similar to extant cervids. In addition. the possibility for combining these two techniques in future mammalian paleoccological studies is considered. The data presented in this study imply that Megoloceros would have indeed been vulnerable to extirpation during the terminal Pleistocene in Ireland. and this information is relevant to understanding the broader pattern of its extinction.

Sediment magnetic properties of glacial till deposited since the Little Ice Age maximum for selected glaciers at Svartisen and Okstindan, northern Norway

Samples of glacial till deposited since the Little Ice Age (LIA) maximum by two glaciers, North Bogbre at Svartisen and Corneliussen-breen at Okstindan, northern Norway, were obtained from transects running from the current glacier snout to the LIA (c. AD 1750) limit. The samples were analysed to determine their sediment magnetic properties, which display considerable variability. Significant trends in some magnetic parameters are evident with distance from the glacier margin and hence length of subaerial exposure. Magnetic susceptibility (X) decreases away from the contemporary snout, perhaps due to the weathering of ferrimagnetic minerals into antiferromagnetic forms, although this trend is generally not statistically significant. Trends in the ratios of soft IRM/hard IRM which are statistically significant support this hypothesis, suggesting that antiferromagnetic minerals are increasing relative to ferrimagnetic minerals towards the LIA maximum. Backfield ratios (IRM -100 mT/SIRM) also display a significant and strong trend towards magnetically harder behaviour with proximity to the LIA maximum. Thus, by employing a chronosequence approach, it may be possible to use sediment magnetics data as a tool for reconstructing glacier retreat in areas where more traditional techniques, such as lichenometry, are not applicable.