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Pupilla alluvionica Meng & Hoffmann, 2008: A Land Snail Extant in the Altai Refugium Recognised for the First Time in Central European Early-Middle Pleistocene Glacials
Abstract
Pupilla alluvionica Meng & Hoffmann, 2008 is an extant land snail species known until now only from a few sites in the Russian Altai Mountains. We have now identified it in fossil loess assemblages of Early-Middle Pleistocene age at three Central European sites. Fossil materials match those of Altaian populations, being only slightly smaller in mean shell width. In addition to the details of the locations and biometrics of these fossil finds, we list the faunas associated with P. alluvionica in the deposits. These are compared with the ecological conditions and associated faunas of living populations from the Altai. Pupilla alluvionica is a typical inhabitant of xeric habitats: steppes with rocky limestone outcrops. This matches the known habitat preferences of associated species in the fossil deposits that are typical of open loess steppe.
... CRFDs for actual and potential major range axes and areas all exhibit concave shapes characteristic of ecological systems to 4800 km (Vertigo cristata agg., Vertigo modesta) with a median value of 1000 km ( Figure 4). While these minimum values are certainly impacted by underreporting (the actual major and minor range extents are likely at least 1000 km), regional endemics within these genera do rarely exist (Horsák et al., 2016). Median potential major axis extent was 7000 km and that for minor axes was 2800 km, ranging from 2200 to 19,800 and 200 to 5600 km, respectively. ...
... This approach appeared justified as the resultant suitability models accurately portrayed not only the known ranges of these taxa but their better-known European relatives as well. Pupilla alluvionica, although having 14 known occurrences, was also not modelled due to extreme spatial clustering and climatic homogeneity within these records (Horsák et al., 2016), which limits global interpolation due to large potential spatial autocorrelation effects. ...
Aim
We document realized and potential global species ranges based on empirically vetted species concepts in conjunction with global climate databases and climate suitability modelling. From this we investigate the nature of dispersal barriers and illustrate how they generate ecological uniqueness.
Location
Holarctic.
Methods
Fifty‐two small body‐size (i.e. < 5 mm) land snail taxa within the genera Euconulus , Pupilla and Vertigo were considered. These represent ~10% of all small body‐size Holarctic land snails and are among the most proficient known passive dispersers. Their potential climatic ranges were determined using Maxent modelling based on 9205 occurrence records. From these we inferred the location, width and nature of isolating barriers and tested for their effects on regional species pool richness and turnover.
Results
Use of unvetted traditional taxonomic concepts and unverified occurrence records would have created up to threefold higher or lower estimates of species‐specific climatic tolerances than the actual values. Modelling must thus only use high quality occurrence data. All but one taxon were shown at a global scale to possess multiple isolated areas of appropriate climate. While oceans represented the most common barrier (37%), intra‐continental barriers were in total almost twice as frequent (inappropriate climate – 29%, habitat/history – 27% and the Greenland ice sheet – 7%). These barriers restricted taxa to only a subset of their potential range, with European taxa possessing approximately twice the global occupancy rates as North American ones (median scores of 62 vs. 34%). As a result, regional taxa pools were three times smaller than their potential sizes, with 50% change in composition occurring over ~2600‐km distances.
Main conclusions
Even for these readily dispersing taxa, isolation barriers prevented species from saturating their potential global range, reduced the size of regional species pools by 2/3, and generated ecological uniqueness between them.
... limata, V. binneyana, V. chytryi) were not modeled. Pupilla alluvionica, although having 14 known occurrences, was also not modeled as its extreme spatial limitation and climatic homogeneity among known sites 47 limits reliability of global-scale interpolation. We have chosen to not generate multitemporal models for Vertigo oughtoni, which has been reported as LGM fossil, 41,43 because its traditional diagnosis was based on faulty shell identification features leading to confusion with at least two other species. ...
Modeled modern and Last Glacial Maximum (LGM) climate ranges for 47 genetically confirmed small Holarctic land snails documented profound landscape dynamism over the last 21,000 years. Following deglaciation, range areas tended to increase by 50% while isolating barrier widths were cut in half. At the same time, the nature of isolating barriers underwent profound change, with the North American continental ice sheet becoming as important in the LGM as the Atlantic Ocean is today in separating Nearctic and Palearctic faunas. Because appropriate modern climate occurs for these species throughout the Holarctic, with no clear barriers being present—especially for such efficient passive dispersers—the current >90% turnover observed between Eurasian and North American species pools appears at least in part related to the LGM landscape. Understanding current and predicting potential future biodiversity patterns thus requires consideration of the landscape template across at least 15,000 years time scales.
Steppe‐tundra is considered to have been a dominant ecosystem across northern Eurasia during the Last Glacial Maximum. As the fossil record is insufficient for understanding the ecology of this vanished ecosystem, modern analogues have been sought, especially in Beringia. However, Beringian ecosystems are probably not the best analogues for more southern variants of the full‐glacial steppe‐tundra because they lack many plant and animal species of temperate steppes found in the full‐glacial fossil record from various areas of Europe and Siberia. We present new data on flora, land snails and mammals and characterize the ecology of a close modern analogue of the full‐glacial steppe‐tundra ecosystem in the southeastern Russian Altai Mountains, southern Siberia. The Altaian steppe‐tundra is a landscape mosaic of different habitat types including steppe, mesic and wet grasslands, shrubby tundra, riparian scrub, and patches of open woodland at moister sites. Habitat distribution, species diversity, primary productivity and nutrient content in plant biomass reflect precipitation patterns across a broader area and the topography‐dependent distribution of soil moisture across smaller landscape sections. Plant and snail species considered as glacial relicts occur in most habitats of the Altaian steppe‐tundra, but snails avoid the driest types of steppe. A diverse community of mammals, including many species typical of the full‐glacial ecosystems, also occurs there. Insights from the Altaian steppe‐tundra suggest that the full‐glacial steppe‐tundra was a heterogeneous mosaic of different habitats depending on landscape‐scale moisture gradients. Primary productivity of this habitat mosaic combined with shallow snow cover that facilitated winter grazing was sufficient to sustain rich communities of large herbivores.
LOŽEK (1986) is of the opinion that Pupilla loessica has been found recently in Central Asia. The aim of this study is to provide some initial, concrete contributions to the discussion of this subject. It should be understood as a basis for further investigations. During recent visits on site between 1995 and 2006 in Central Asia in the Russian Altay, in Northern Mongolia, in the Baikal region and in the Tien Shan, numerous recent malacocenoses were examined. In many places evidence was found of a form of Pupilla which had not previously been described from this region; its shell morphology cannot be distinguished from Pupilla loessica. Its distribution is concentrated in the strongly continental Khrebet Saylyugem in the South Eastern Altay. The probability of the occurrence of Pupilla loessica in Central Asia is supported, apart from the shell morphology criteria, by the preference of this species for more continental types of habitat with average annual temperatures markedly below 0° C and the corresponding accompanying fauna adapted to the cold, including e.g. Columella columella, Vertigo genesii, Vallonia tenuilabris, etc., which make these fauna easily comparable to the Pleistocene glacial associations of the Central European region.
Using the Pupilla faunas of Europe, North America, the Altai region of central Asia and eastern Asia, we consider whether the existing taxonomy based primarily on shell apertural characteristics correlates with relationships established on the basis of mitochondrial and nuclear DNA-sequence data. We obtained DNA sequence from nuclear ITS1 and ITS2 and mitochondrial COI and CytB from 80 specimens across 22 putative Pupilla taxa. The sequence data were analysed using maximum likelihood, maximum parsimony, Bayesian and neighbour-joining phylogenetic tree reconstruction, as well as base-pair substitution and insertion-deletion analysis. Revised species-level concepts were generated by identifying reciprocally monophyletic clades that exhibited unique conchological features. These analyses document that, although many previously described taxa have biological merit, the highly plastic nature of shell apertural features makes them unreliable indicators of species identity in several independent lineages. However, shell surface sculpture and architecture appear to provide more reliable diagnoses. Because of the traditional reliance of species-level taxonomy in Pupilla on plastic apertural features, too many species-level entities have been described in Europe and the Altai. Also, because taxonomically useful shell sculpture features have tended to be ignored, too few species have been described in eastern Asia and North America. As a result, confusion exists about species ranges, ecological tolerances and interpretation of Quaternary fossils within the genus. Based on these analyses three new species are described: P. alaskensis, P. hudsonianum and P. hokkaidoensis.
Many land snail species of the European Pleistocene have recently been reported from Central Asia. The ecological conditions in which they live, however, are little known. We studied vascular plants and land snails along a climatic gradient in the Russian Altai Mountains. We recorded 566 plant and 40 mollusc species. In the most continental part of the Altai we discovered living Pupilla loessica, a land snail now extinct in Europe and which is an important indicator of palaeoenvironments of the European Pleistocene. We inferred thermal ranges of the snail species in the Altai and observed considerable differentiation among the species. Moisture requirements of the species were indirectly estimated by using moisture indicator values of plant species co-occurring with the snails. They reveal local moisture requirement patterns that cannot be inferred from climate data. Snails are sorted along our transect; only a few species occur throughout the Altai. Many snails are able to occupy various plant communities. A few species may be indicators of vegetation types. For palaeorecon-structions, mollusc species are apparently good indicators for climate but less so for vegetation types.
New distribution data for Vertiginidae (Gastropoda: Pulmonata) in Central Asia. – In the context of expeditions and collections during private visits in Central Asia (1995–2006), the author investigated numerous malacocenoses in the Altai, Tien Shan, Mongolia and the Baikal region. This article presents new insights concerning the dispersal of the Vertiginidae (Gastropoda: Pulmonata) comprising Vertigo substriata, Vertigo lilljeborgi, Vertigo genesii, Vertigo geyeri, Vertigo alpestris and Vertigo parcedentata in the northern palaearctic region with special reference to Central Asia. > Kurzfassung Im Rahmen von Expeditionen und privaten Sammelreisen (1995–2006) untersuchte der Verfasser in Zentralasien (Altai, Tienschan, Mongolei und Baikal-Gebiet) zahlreiche Malakozönosen. Zur Verbreitung der Vertiginidae (Gastropoda: Pulmo-nata) im nördlichen paläarktischen Raum, unter besonderer Berücksichtigung Zentralasiens, erbrachten die Untersuchungen wesentliche neue Erkenntnisse. Dargestellt werden in dieser Arbeit Ergebnisse zur Verbreitung von Vertigo substriata, Ver-tigo lilljeborgi, Vertigo genesii, Vertigo geyeri, Vertigo alpestris sowie Vertigo parcedentata.
Pleistocene mammalian communities display unique features which differ from present-day faunas. The paleocommunities were characterized by the extraordinarily large body size of herbivores and predators and by their unique structure consisting of species now inhabiting geographically and ecologically distinct natural zones. These features were probably the result of the unique environmental conditions of ice age ecosystems. To analyze the ecological structure of Last Glacial and Recent mammal communities we classified the species into biome and trophic-size categories, using Principal Component analysis. We found a marked similarity in ecological structure between Recent eastern Altai-Sayan mammalian assemblages and comparable Pleistocene faunas. The composition of Last Glacial and Recent eastern Altai-Sayan assemblages were characterized by the occurrence of large herbivore and predator species associated with steppe, desert and alpine biomes. These three modern biomes harbor most of the surviving Pleistocene mammals. None of the analyzed Palearctic Last Glacial faunas showed affinity to the temperate forest, taiga, or tundra biome. The Eastern part of the Altai-Sayan region could be considered a refugium of the Last Glacial-like mammalian assemblages. Glacial fauna seems to persist up to present in those areas where the forest belt does not separate alpine vegetation from the steppes and deserts.
Aim Shells of fossil molluscs are important for palaeoecological reconstructions. However, the habitat requirements of snail species typical of central European full‐glacial loess sediments are poorly known because most of them became very rare or extinct in Europe. The recent discovery of an almost complete extant assemblage of such species in mountainous regions of central Asia enables more precise characterization of their habitats, which may significantly improve reconstructions of Pleistocene environments.
Location Altai Mountains, Russia.
Methods Terrestrial snail assemblages, vegetation composition and selected environmental variables were recorded at 118 sites along a gradient of climatic continentality in the Russian Altai. Habitat characteristics of sites where species typical of the full‐glacial period occurred were described using a classification tree.
Results Seven of the eight species that are typical of central European full‐glacial loess sediments were found in the study area. They were confined to cool areas with January mean temperatures below −17 °C, but occurred mainly in sheltered habitats with a warmer microclimate, such as scrub or open woodland. Pupilla loessica and Vallonia tenuilabris had the broadest habitat range, occurring from woodland to dry steppe. Unexpectedly, Columella columella , Pupilla alpicola , Vertigo genesii , V. parcedentata and V. pseudosubstriata were found mainly in wooded fens and shrubby tundra rather than in open steppe. Most of these seven species were recorded in base‐rich wooded fens. Very dry open steppe habitats usually supported no snails.
Main conclusions Habitat ranges of the studied snails in the Altai indicate that the full‐glacial landscapes of central European lowlands that harboured these species were not completely dominated by open and dry loess steppe. Most probably they contained a significant component of shrubby vegetation, patches of wet habitats, and probably also areas of woodland at sites with a favourable mesoclimate.
Pupilla pratensis (Clessin, 1871) was recently confirmed as a distinct species based on morphological, ecological and molecular evidence. The main purpose of this study is to publish the first reliable data on the occurrence of P. pratensis in the Czech Republic and Slovakia. The second goal is to analyse conchometry of P. pratensis, P. muscorum (L., 1758), and P. alpicola (Charpentier, 1837) to find out whether it is possible to reliably distinguish these species solely based on shell measurements. For multidimensional analysis of shell measurement variation we used principal components analysis (PCA). We documented six populations of P. pratensis in the Czech Republic and one in SW Slovakia. The revision of voucher material showed that all previously reported records of P. alpicola from the Czech Republic belonged in fact to P. pratensis. This requires the exclusion of P. alpicola from the list of Czech molluscs. Based on multidimensional analysis of shell measurements it was possible to distinguish P. pratensis from P. muscorum with no overlapping specimens. Pupilla alpicola was almost completely different from P. muscorum with only few overlapping specimens, contrary to P. pratensis which was mostly impossible to distinguish from P. alpicola based on analysed shell measurements. Shell width was the best single shell measurement for distinguishing P. pratensis and P. muscorum. Shell measurements of two Swedish populations of P. pratensis did not differ from shell variation of Czech and Slovak populations. However, Scandinavian populations displayed some differences from central European populations in apertural barriers which are discussed in detail. Czech and Slovak populations of P. pratensis occurred in calcium-rich fen meadows which perfectly matches with site characteristics reported from Scandinavia. We assume that the observed morphometric differences between P. pratensis and P. muscorum can be useful for distinguishing the species also outside the Czech territory and for palaeoecological studies where only empty shells are available. Since these species occupy ecologically different habitats their reliable identification in fossil material can improve the reconstructions of past environments.
We developed interpolated climate surfaces for global land areas (excluding Antarctica) at a spatial resolution of 30 arc s (often referred to as 1-km spatial resolution). The climate elements considered were monthly precipitation and mean, minimum, and maximum temperature. Input data were gathered from a variety of sources and, where possible, were restricted to records from the 1950-2000 period. We used the thin-plate smoothing spline algorithm implemented in the ANUSPLIN package for interpolation, using latitude, longitude, and elevation as independent variables. We quantified uncertainty arising from the input data and the interpolation by mapping weather station density, elevation bias in the weather stations, and elevation variation within grid cells and through data partitioning and cross validation. Elevation bias tended to be negative (stations lower than expected) at high latitudes but positive in the tropics. Uncertainty is highest in mountainous and in poorly sampled areas. Data partitioning showed high uncertainty of the surfaces on isolated islands, e.g. in the Pacific. Aggregating the elevation and climate data to 10 arc min resolution showed an enormous variation within grid cells, illustrating the value of high-resolution surfaces. A comparison with an existing data set at 10 arc min resolution showed overall agreement, but with significant variation in some regions. A comparison with two high-resolution data sets for the United States also identified areas with large local differences, particularly in mountainous areas. Compared to previous global climatologies, ours has the following advantages: the data are at a higher spatial resolution (400 times greater or more); more weather station records were used; improved elevation data were used; and more information about spatial patterns of uncertainty in the data is available. Owing to the overall low density of available climate stations, our surfaces do not capture of all variation that may occur at a resolution of 1 km, particularly of precipitation in mountainous areas. In future work, such variation might be captured through knowledge-based methods and inclusion of additional co-variates, particularly layers obtained through remote sensing.
Climate change in the past has led to significant changes in species' distributions. However, how individual species respond to climate change depends largely on their adaptations and environmental tolerances. In the Quaternary, temperate-adapted taxa are in general confined to refugia during glacials while cold-adapted taxa are in refugia during interglacials. In the Northern Hemisphere, evidence appears to be mounting that in addition to traditional southern refugia for temperate species, cryptic refugia existed in the North during glacials. Equivalent cryptic southern refugia, to the south of the more conventional high-latitude polar refugia, exist in montane areas during periods of warm climate, such as the current interglacial. There is also a continental/oceanic longitudinal gradient, which should be included in a more complete consideration of the interaction between species ranges and climates. Overall, it seems clear that there is large variation in both the size of refugia and the duration during which species are confined to them. This has implications for the role of refugia in the evolution of species and their genetic diversity.
Eight glaciations of Brunhes age are recorded in deep-sea sediments but only four are recognized on land. Sequences of loess beds and soils resting on river terraces in Czechoslovakia and Austria, close to the former fronts of the Scandinavian and Alpine glaciers, shed more light on the dilemma as they enable a correlation to be made between oceanic record and the classical glacial stages of Europe. They show seventeen major glacial-interglacial shifts within the last 1.6 million years (cf. Figure 6). Eight of them are of Brunhes age. However, only four times did the continental glacier advance far enough to leave behind end moraines undestroyed by subsequent glaciation. According to the best estimates available at present, this occurred 20,000 years ago during the Weichsel advance; about 140,000 years ago during the Warthe advance; about 350,000 years ago during the Saale, sensu stricto (Drenthe), and about 550,000 years ago during the main Elster sensu stricto advance. Weichsel moraines broadly correlate with the oceanic 0 ¹⁸ stage 2, Warthe with stage 6, Saale with stage 10, and Elster with stage 16 (cf. Figure 29). Unlike the short-lived ice advances representing the north European classical glacial stages, the Würm terraces in the Alps apparently formed during a long interval covering the last glacial, a substantial part of the next-to-last glacial, the last interglacial, and the Holocene. It probably correlates with O ¹⁸ stages 1 through 6. Correspondingly the Riss terrace probably correlates with 0 ¹⁸ stages 12 through 7; Mindel with 18 through 13, and Giinz terrace with 22 through 19. The RW, MR, GM, and DG erosional episodes of the alpine rivers are of glacial rather than interglacial origin. They seem to represent morphostratigraphic correlation horizons of wide regional significance with approximate ages of 180, 450, 650, and 850 millenia (within 0 ¹⁸ stages 6, 12, 18 and 22). They are supposed to be primarily of tectonic rather than climatic origin. Evidently the classical glacial stages of Penck and Briickner based on the alpine terraces cover the last 850,000 years fully, but their climatic concept is in error. Interglacials are defined as intervals of continuous presence of mixed broadleaf forest in northwestern and central Europe. There were at least three such interglacials locally labeled as Eem (120, 230, and about 330 thousand years ago), two which were described as Holstein, and three labeled, or mislabeled, as Cromer. Our conclusions are based on three highly probable assumptions that still cannot be directly proved: (1) that the eight glacial cycles of Brunhes age, recognized in the semicontinuous loess-soil sequences around Brno and Prague and labeled as L-B to L-I in an order of increasing age, are the products of roughly synchronous, gross global climate changes which left a record of eight similar cycles in deepsea sediments; (2) that the development of river terraces around Brno was roughly synchronous with that of the terraces around Ulm and Munich; and (3) that the development of terraces around Prague was roughly synchronous with those around Leipzig. Needless to say, the misinterpreted climatic and time stratigraphic concept of classical European glacial stages and the broad use of corresponding labels in world-wide correlations far away from the original type localities contributed greatly to the remarkable confusion which reigns in the jungles of Pleistocene stratigraphy today. The use of classical Pleistocene stage names should be restricted to their type areas and to the intervals of time truly represented by their type units. The meaningful world-wide climatostratigraphic subdivision of the Pleistocene must be based on continuous depositional sequences.
Knowledge of present-day communities and ecosystems resembling those reconstructed from the fossil record can help improve our understanding of historical distribution patterns and species composition of past communities. Here, we use a unique data set of 570 plots explored for vascular plant and 315 for land-snail assemblages located along a 650-km-long transect running across a steep climatic gradient in the Russian Altai Mountains and their foothills in southern Siberia. We analysed climatic and habitat requirements of modern populations for eight land-snail and 16 vascular plant species that are considered characteristic of the full-glacial environment of central Europe based on (i) fossil evidence from loess deposits (snails) or (ii) refugial patterns of their modern distributions (plants). The analysis yielded consistent predictions of the full-glacial central European climate derived from both snail and plant populations. We found that the distribution of these 24 species was limited to the areas with mean annual temperature varying from −6.7 to 3.4 °C (median −2.5 °C) and with total annual precipitation varying from 137 to 593 mm (median 283 mm). In both groups there were species limited to areas with colder and drier macroclimates (e.g. snails Columella columella and Pupilla loessica, and plants Kobresia myosuroides and Krascheninnikovia ceratoides), whereas other species preferred areas with relatively warmer and/or moister macroclimates (e.g. snails Pupilla turcmenica and P. alpicola, and plants Artemisia laciniata and Carex capillaris). Analysis of climatic conditions also indicated that distributional shifts of the studied species during the Pleistocene/Holocene transition were closely related to their climatic tolerances. Our results suggest that the habitat requirements of southern Siberian populations can provide realistic insights into the reconstruction of Eurasian, especially central European, glacial environments. Data obtained from modern populations also highlight the importance of wet habitats as refugia in the generally dry full-glacial landscape.
During the research of fen mollusc assemblages in Switzerland carried out in August 2012, we recorded the first occurrence of Vertigo lilljeborgi (Westerlund, 1871) in the Alps and several new occurrences of V. genesii (Gredler, 1856), substantially extending its Alpine distribution island towards the western part of the mountain range. Both these Euro-Asian species are mainly distributed in northern Europe and have a strong ecological affinity to groundwater-fed fens. With respect to the rarity, refugial character and high conservation value of central European populations of these species, we review in detail their ecological preferences and present European distribution, and also comment on the possible conservation implications for their globally threatened habitats.
In the European context, the malacofaunas from the loess series in Bohemia and Moravia provided relatively rich evidence of environmental conditions in distinct phases of the climatic cycle. This is particularly true of the Last Interglacial–Glacial cycle in which a number of minor documented phases have made it possible to reconstruct the sequence of detailed environmental changes. The major conclusions may be summarized as follows: (1) Loess malacocoenoses differ in their composition from all known molluscan assemblages in present-day Europe and show a relatively uniform character in space and time. (2) They reflect specific environments that may be called as loess steppe, characterized by severe cold-continental climate, grassland with discontinuous turf cover and raw soils rich in salts. (3) Malacocoenoses of the loess steppe differ considerably from those of the modern subpolar zone. (4) Differences in species distribution correspond to particular loess facies due to various depositional and climatic conditions. (5) Loess is the product of pleniglacial phases: in early glacial phases its occurrence is very limited, and the malacofauna of these loess horizons is characterized by an absence of most subpolar or alpine species. (6) Interstadials of the Early Glacial phase are characterized by assemblages of the chernozem steppe and indicate strongly continental conditions. (7) In the pedocomplex I (PK I=δ18O stage 3, Stillfried B) no malacological records are available which would indicate a warmer climate: its assemblages are very similar to those from the loess. (8) Interglacials have fully developed woodland malacocoenoses which reflect a warm and humid climate, even in areas where such assemblages did not develop during the Postglacial. (9) Loess formation and specific loess malacocoenoses decline in the earlier phase of the Late Glacial, being gradually replaced by much more diversified communities with a higher proportion of moisture-loving elements. (10) The above-mentioned development corresponds to the situation in dry loess areas that may be considered typical loess landscapes. At higher elevations and in moister areas, different sediments, soils and molluscan assemblages existed at the same time. Their correlation with sequences of the loess series is poorly documented due to the lack of reliable fossil and stratigraphic evidence.
At least seventeen glacials and seventeen interglacials occurred in Europe over the last 1.7 million years. Eight glacials and eight interglacials are of Brunhes age. Only four glacials and three interglacials are recognized by classical Alpine and north European subdivisions of the Pleistocene. The classical units are correlated with continuous oxygen isotope records from the oceans using loess sections and terraces as a link (cf. Fig. 21). It is found that: (1) the terraces representing the four Alpine “glacial” stages fully cover the last 0.8 million years but correspond to both glacial and interglacial climates; (2) the Alpine “interglacial” stages do not represent episodes of interglacial climate but probably intervals of accelerated crustal movements; (3) the physical evidence on which the north European classical subdivision is based accounts for only about 15% of the time represented. This has led to serious miscorrelations.It is urgently recommended to abandon the classical terminology in all interregional correlations and to base the chronostratigraphic subdivision of Pleistocene on the18O record of deep-sea sediments.
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