Article

European glacial relict snails and plants: Environmental context of their modern refugial occurrence in southern Siberia

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Abstract

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.

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... Palynological studies found a high degree of similarity between the Late Pleistocene pollen spectra from central Europe and surface pollen samples from present-day landscapes of the Altai-Sayan Mountains (Kune s et al. 2008;Magyari et al. 2014). Moreover, this and adjacent mountain systems of central Asia are inhabited by extant snails that either represent Beringian allopatric replacements or in some cases the same species known from the European Pleistocene loess record (Meng 2009;Meng & Hoffmann 2009;Hors ak et al. 2010Hors ak et al. , 2015Nekola et al. 2018). Many of these became extremely rare or extinct in Europe after the onset of the Holocene or earlier . ...
... In each plot, we recorded all species of vascular plants and terricolous bryophytes and lichens, along with their cover-abundances estimated using the ninedegree Braun-Blanquet scale (van der Maarel 1979). Snails were searched for by eye by a single observer for 30-60 min per plot in all the appropriate microhabitats within each plot (for details see Hors ak et al. 2010Hors ak et al. , 2015. Small snails (<5 mm in maximum dimension) were recorded by collecting~3 L of leaf litter, mosses and topsoil. ...
... Of the tundra indicators, pollen of Dryas and Betula nana types is produced by Dryas oxyodonta and Betula rotundifolia, the dominant species of the respective habitat types. Plants considered as glacial relicts in Europe according to Hors ak et al. (2015) are most represented in alpine dry grassland, dryas tundra, wet conifer woodland and wet scrub, where they were found in all or nearly all studied plots (Fig. 6A). They are also well represented in steppe habitats, but less represented in mesic and dry conifer woodland. ...
Article
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.
... Because snail shells have a good chance of becoming preserved in calcium-rich sediments (e.g. Ložek, 1964), many land snails are important indicators for quaternary climate reconstructions (Rousseau, 1991;Moine et al., 2002;Horsák et al., 2015) as they appear to have tracked their niche through space and time. This applies in particular to smaller species that disperse readily-albeit passively (Wada et al., 2012;Simonová et al., 2016) -and thus may cover wide ranges such as species of the genus Pupilla occurring in temperate and boreal zones of the Holarctic (Sysoev & Schileyko, 2009;Nekola & Coles, 2010;Nekola et al., 2015). ...
... The suture is deep and the surface has characteristic fine, irregular ribs . It is a typical representative of steppe-tundra communities, currently restricted to cold areas of the central Asian mountain systems (Meng & Hoffmann, 2009;Horsák et al., 2015). From a palaeoecological perspective, it is also a characteristic full-glacial fossil in loess deposits across almost all of temperate Europe east of the Rhine valley (e.g. ...
... Meng & Hoffmann, 2009). Lately, it has been found to occur commonly in the Altai Mountains, in northern Mongolia, near Lake Baikal and occasionally also in the Tien Shan Mountains (Meng & Hoffmann, 2009;Hoffmann et al., 2011;Horsák et al., 2015). Its modern distribution in the Altai is limited to areas with a dry and cold climate . ...
Article
Changing environmental conditions force species either to disperse or to adapt locally either genetically or via phenotypic plasticity. Although limits of plasticity can be experimentally tested, the predictability of genetic adaptation is restricted due to its stochastic nature. Nevertheless, our understanding of evolutionary adaptation has been improving in particular through studies of parallel adaptation. Based on molecular phylogenetic inferences and morphological investigations of both recent and fossil shells we tracked the morphological changes in three land snails, Pupilla alpicola, Pupilla loessica and Pupilla muscorum. These species differ in habitat requirements as well as historical and extant distributions with P. alpicola and P. loessica being more similar to each other than to P. muscorum. Therefore, we hypothesized, that the three species reacted independently and individually to the conditions changing throughout the Pleistocene, but expected that changes within P. alpicola and P. loessica would be more similar compared to P. muscorum. Indeed, intraspecific shell shape differences across time were similar in P. alpicola and P. loessica, suggesting that similar niche shifts have led to similar transformations in parallel. In contrast, extant P. muscorum populations were practically identical in shape to their ancestors. They have probably tracked their ecological niches through time.
... The occurrence of glacial relic Vallonia tenuilabris is characteristic of this assemblage. This is typical loess species usually connected with the severe climatic conditions and open areas of so-called "coldsteppes" (Ložek, 1964), but based on recent data from Asia, it may also occur in alpine grasslands, taiga, hemiboreal forest and wooded fens (Schileyko, 1984;White et al., 2008;Horsák et al., 2010Horsák et al., , 2015. Its highest frequency is noted in sample 21, layer 4. The assemblage is composed mainly of shade-loving gastropods, with the proportion of 53 and 74% of all species and all individuals, respectively ( Fig. 3). ...
... Rather cool climate is also evidenced by the glacial relic V. tenuilabris which usually disappeared in Central Europe in the Pleistocene/Holocene boundary (e.g. Dagnan-Ginter et al., 1992;Wiktor, 2004;White et al., 2008;Horsák et al., 2010Horsák et al., , 2015, but here constitute very characteristic element of the fauna. Accumulation of loess-like deposits with the poor fauna with V. tenuilabris may indicate some transformation towards more open areas around the cave, but due to the occurrence of only few individuals this interpretation is open to discussion. ...
... The most distinctive feature in the Cave above the Słupska Gate is the abundance and long occurrence of the glacial relic Vallonia tenuilabris, unknown both from the Holocene and most of the cave deposits of Central Europe. Nowadays this species occurs in Siberia, Central Asia and northern China (Dagnan-Ginter et al., 1992;Gerber, 1996;Meng, 2008Meng, , 2009White et al., 2008;Horsák et al., 2010Horsák et al., , 2015. It is common in the Late Vistulian loess and loess-loams of southern Poland, but in cave sediments has only been described in the Zawalona Cave and Duża Cave (southern part of the Cracow-Częstochowa Upland) Dagnan-Ginter et al., 1992;Alexandrowicz and Alexandrowicz, 1995a). ...
Article
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The Cave above the Słupska Gate (southern Poland) contains about 2m depth of mollusc-bearing deposits. Radiocarbon and archaeological dating indicate that these deposits accumulated during the Holocene (Preboreal to Subatlantic), although the earliest layers may date from the end of the Pleistocene. Eight layers of silts, sands and loess-like deposits were distinguished at the site. Seven of them contained identifiable snail shells, sometimes in large numbers, and sparse remains of vertebrates and archaeological artefacts. The molluscan assemblages retrieved from the cave contain over 40 taxa and 1,200 specimens. The balance of species distributed among 11 zoogeographical groups enabled us to identify four assemblages which differ in their ecological structure and in the composition of the fauna. The oldest fauna (Late Glacial/Preboreal and/or Preboreal) with many shade-loving species is typical of a cool climate. Episodes of drying are evidenced by the loess-like deposits and the occurrence of open-country snails such as the glacial relic Vallonia tenuilabris. This species disappeared in the younger part of the Early Holocene, which is the most distinctive feature of the Słupsko Hill sequence. The Middle Holocene climatic optimum is characterised by abundantand diverse fauna which is typical of mixed and deciduous forests with distinct oceanic influences. The critical Discus ruderatus and Discus rotundatus succession reflects the general trends in European malacofaunas. The Late Holocene record may bear some hiatuses, but the shift away from a complete forest fauna is evident.
... It has been suggested that the European LGM climate has a large overlap with the present-day climate of Siberia (Fløjgaard et al., 2009). There is also biological evidence indicating ecological similarities between present-day Siberia and European regions during the LGM (Kune s et al., 2008;Meng, 2009;Pel ankov a and Chytrý, 2009;Hors ak et al., 2010Hors ak et al., , 2015Magyari et al., 2014;Pavelkov a Ri c ankov a et al., 2014. Although these studies suggest that, to a certain degree, present-day Siberian vegetation can be used as a model for understanding European vegetation during the LGM, the climatic analogy between the two periods and regions has not been evaluated yet. ...
... Present-day ecosystems of southern Siberia have been considered similar to those occurring in central Europe during the LGM (Kune s et al., 2008;Hors ak et al., 2010Hors ak et al., , 2015Magyari et al., 2014). Our results support that during the LGM the regions adjoining the Scandinavian ice-sheet and non-glaciated parts of the Alps and the Carpathians were suitable for treeless arctic scrub and herbaceous vegetation, including some minerotrophic fens, which may have been similar to the current fens of western Siberia (Lapshina, 2010;Peterka et al., 2017). ...
... Our results indicate that the degree of climatic analogy between current Siberian and LGM European environments is high for eastern and central Europe but much lower for southern and western Europe (Fig. 2). This fact is also supported by biotic evidence of the widespread occurrence of central European glacial relict species in Siberia (Hors ak et al., 2015). The vegetation data compiled from Siberia corresponds well to the most analogue climatic regions between the two study periods (Fig. 3), supporting the appropriateness of the study areas to investigate links with the LGM in Europe. ...
... As a result, the south-eastern Russian Altai represents a unique example of a full-glacial palaeorefugium that extends over landscape scales (macrorefugium; Nekola 2013). It harbours relict populations of many species that were typical of the last full-glacial period including vascular plants (Pelánková and Chytrý 2009;Magyari et al., 2014;Horsák et al., 2015), land snails Hoffmann et al., 2011) and mammals (PavelkováŘičánková et al., 2014(PavelkováŘičánková et al., , 2015. While these characteristic glacial species are common in the south-eastern Russian Altai, they are quite rare or absent from the northern Altai. ...
... On the contrary, they are char-acterized by several species that are rare or absent in the areas with mesic-warm climate. Many of these species are characteristic of European LGM loess assemblages (Table S1 in Appendix A in Supplementary data; Horsák et al., 2010Horsák et al., , 2015Hoffmann et al., 2011). ...
... The values for diagnostic species are phi coefficients of association multiplied by 100. The values for constant species are percentage occurrence frequencies.Asterisks indicate species considered as European glacial relicts byHorsák et al. (2015). -warm areas (low altitudes in the north)Wet-cold areas (high altitudes of the northern mountains) ...
Article
Site-scale species richness (alpha diversity) patterns are well described for many present-day ecosystems, but they are difficult to reconstruct from the fossil record. Very little is thus known about these patterns in Pleistocene full-glacial landscapes and their changes following Holocene climatic amelioration. However, present-day central Asian ecosystems with climatic features and biota similar to those of the full-glacial periods may serve as proxies of alpha diversity variation through both space and time during these periods. We measured alpha diversity of vascular plants, bryophytes, macrolichens and land snails, as well as environmental variables, in 100-m² plots located in forests and open habitats in the Russian Altai Mountains and their northern foothills. This region contains adjacent areas that possess climatic and biotic features similar to mid-latitude Europe for both the Last Glacial Maximum and contemporaneous Holocene ecosystems. We related alpha diversity to environmental variables using generalized linear models and mapped it from the best-fit models. Climate was identified as the strongest predictor of alpha diversity across all taxa, with temperature being positively correlated to number of species of vascular plants and land snails and negatively correlated to that of bryophytes and macrolichens. Factors important for only some taxa included precipitation, soil pH, percentage cover of tree layer and proportion of grassland areas in the landscape around plots. These results, combined with the high degree of similarity between the current Altai biota and dry-cold Pleistocene ecosystems of Europe and northern Asia, suggest that vascular plant and land snail alpha diversity was low during cold phases of the Pleistocene with a general increase following the Holocene climatic amelioration. The opposite trend probably existed for terricolous bryophytes and macrolichens.
... The snail Vertigo pseudosubstriata Ložek, 1954 was first detected as a fossil in the Weichselian loess at the archaeological site of Dolní Věstonice in Moravia, Czechia (Ložek, 1954). Following its first description, shells of this species were also detected at other findspots in Central and Eastern Europe (Ložek, 1954(Ložek, , 1964, before it was discovered alive in Central Asia (Schileyko, 1984;Meng, 2009;Horsák et al., 2010Horsák et al., , 2015. ...
... Well-known examples are, for instance, Vallonia tenuilabris (A. Braun, 1843), Vertigo parcedentata Braun, 1847 or Pupilla loessica Ložek, 1954, which have discovery histories similar to V. pseudosubstriata and today are likewise found in habitats in Central Asia (Gerber, 1996;Meng 2008Meng , 2009Meng and Hoffmann, 2009;Horsák et al., 2010Horsák et al., , 2015Haase et al., 2021). V. parcedentata was also discovered alive in Norway (Pokryszko, 1993). ...
... Uvalieva (1967) discovered this species later in the southern Altai (northern Kazakhstan) and described it initially as V. laevis (synonym for V. pseudosubstriata) (compare also Schileyko, 1984;Uvalieva 1990). According to currently available data, the distribution of V. pseudosubstriata extends to Tien Shan (Kyrgyzstan, southern Kazakhstan, western China) and to the central and southern Altai (southern Russia, northern Kazakhstan) (Schileyko, 1984;Schileyko and Rymzhanov, 2013;Uvalieva, 1990;Egorov, 2008;Sysoev and Schileyko, 2009;Meng, 2008;Horsák et al., 2010Horsák et al., , 2015Nekola et al., 2018). ...
Article
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The small terrestrial gastropod Vertigo pseudosubstriata Ložek, 1954 is one of the rarest glacial indicator species in the Pleistocene of Central and Eastern Europe. In all, this species has been found at only about 15 sites in Europe. V. pseudosubstriata was initially described as a fossil in Central Europe and was discovered only later alive in Central Asia. With regard to its modern distribution, 25 habitats with V. pseudosubstriata have been examined in Tien Shan and in the central and southern Altai. These findings seem to capture the contemporary distribution of the species and provide information on the boundaries of its ecological requirements. These data are of great significance for the interpretation of the fossil assemblages. Since the few fossil specimens in Europe date from very different glacial periods in the Elsterian, Saalian Complex and Weichselian, it can be concluded that V. pseudosubstriata apparently immigrated in at least three distinct waves. Most of the Pleistocene specimens in eastern Central Europe and Eastern Europe are reported from archaeological sites of the Upper Middle Weichselian (Gravettian), roughly between 33 and 29 ka cal bp. In this paper, we review all reported modern and fossil occurrences and discuss the species' ecological range.
... Furthermore, recent DNA sequence analysis by Nekola et al. (2015) indicates that Pupilla pratensis and Pupilla alpicola are not phylogenetically separated and represent a monophyletic clade for both nDNA and mtDNA . The synonymy between the two taxa has been accepted also by Horsák et al. (2015). The form Pupilla muscorum densegyrata was introduced by Ložek (1954), who suggested its possible affinity with Pupilla alpicola and smaller specimens of Pupilla pratensis. ...
... The form Pupilla muscorum densegyrata was introduced by Ložek (1954), who suggested its possible affinity with Pupilla alpicola and smaller specimens of Pupilla pratensis. Horsák et al. (2015) stated that the subspecies densegyrata is a form of Pupilla alpicola characteristic of the younger loess sediments in Central Europe (e.g. ...
... In the view of the latest studies (e.g. Horsák et al. 2011Horsák et al. , 2015Nekola et al. 2015), we consider our material as representative of Pupilla alpicola and its ecophenotype P. alpicola densegyrata. ...
Article
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We present a taxonomic and palaeoecological analysis of a continental mollusc fauna from a mammoth bearing succession near Bullendorf in Lower Austria. The taxonomic analysis comprises morphological descriptions and SEM documentation of 15 Pleistocene gastropod species. A Principal Component Analysis of the quantitative and qualitative composition of the investigated samples reveals a stratigraphic succession of four mollusc assemblages defined herein as Galba truncatula assemblage, Succinella oblonga assemblage, Pupilla muscorum/loessica and Pu-pilla alpicola/sterrii assemblages. The autecological requirements of the species of each assemblage allow a reconstruction of the palaeoenvironmental history of the section with alternating dry and humid conditions within a general cooling trend. Despite generally low mollusc density and species richness, the Bullendorf site allows important insight into latest Quaternary regional environmental conditions and climate. Based on the comparison with similar mollusc sites across Europe, a biostratigraphic correlation with the Late Pleistocene (~110-12 ka) is proposed.
... Today, most of these species have typical arctic-alpine distributions, being widely dispersed across Fennoscandia with a few relict populations surviving at high elevations in Central Europe (e.g. Horsák et al., 2015). Some isolated populations also persist in environmentally stable habitats, such as mires, in lowland regions (Schenková & Horsák, 2013). ...
... This is not really surprising, since any snail species that specifically avoids calcium-rich environments is very unlikely to be preserved in calcareous deposits such as loess (Ložek, 2001). Its presumed relict status is based largely on its rare and scattered modern distribution in temperate mainland Europe, in contrast to its continuous distribution in northern Europe, where it occurs together with other glacial species characteristic of loess communities (Horsák et al., 2015). The only fossil records from mainland Europe come from Germany and Poland (Jaeckel, 1962), but these records are old, unverified and poorly dated. ...
Article
Vertigo lilljeborgi (Westerlund, 1871) is one of the rarest terrestrial snail species in temperate mainland Europe, where it is traditionally considered a glacial relict. This contrasts with its occurrence in northern Europe where it is a widespread species. This species prefers constantly wet habitats that are neutral to slightly acidic and avoids highly alkaline conditions, which is an extremely rare ecology for a Eurasian mollusc. Until 2012, only five historical records of this species were known in mainland Europe to the south of its main distribution in northern Europe. Since then, 20 new sites have been discovered, mostly located in the Hercynian Mountains (Bohemian Massif in the Czech Republic and Massif Central in France). In comparison with the boreal European and Alpine populations, those from the Hercynian Mountains inhabit acidic, rather soligenous and productive fens, strongly dominated by Sphagnum. Vertigo lilljeborgi does not occur in some sites with apparently suitable habitats as indicated by species composition of the vegetation. We observed a surprising correspondence between the occurrence of V. lilljeborgi and mean July air temperature and we report its first fossil record from the last glacial period from Central Europe. Although the number of its sites has increased recently, these sites represent highly unusual and unique habitats, vulnerable to drainage and destruction from human activities. This highlights the need for conservation efforts in most of the newly discovered isolated sites.
... The malacofauna was divided into different paleoecological groups following the paleoecological classifications of Ložek (1964), Krolopp and Sümegi (1995), Sümegi-Krolopp (2002), Alexandrowicz (2004), Sümegi (2005). The malacological record was also classified according to the recent geographical distribution of the species (Soós, 1943;Evans, 1972;Kerney et al., 1983;Krolopp, 1983;Welter-Schultes, 2012;Horsák et al., 2015), and on the basis of paleoclimatological indicator roles (Sümegi and Krolopp, 2002;Sümegi, 2005;Sümegi et al., 2013). ...
... 33,000-31,500 cal BP years, with a typical south-GHP mollusc fauna Krolopp, 1995, 2002) Succinella oblonga, Pupilla muscorum, Perpolita hammonis Vitrea crystallina, Clausilia dubia association. Besides them the appearance of xeromontane, recently Siberia living (Sümegi, 1996(Sümegi, , 2005White et al., 2008White et al., , 2013Moine et al., 2008;Horsák et al., 2010Horsák et al., , 2015 Vallonia tenuilabris indicates a weak cooling period in this zone, although the dominance rate of thermomesophilous species were still high (Fig. 5). ...
Article
According to the findings of a complex radiocarbon dated sedimentological, geochemical, anthracological, phytolite, malacological study implemented on an open 1036 cm wind-blown sand, loess, sandy loess and fossil soils profile of western part of brickyard at Katymár village in Hungary. Increased boreal mixed-leaved forest steppe and temperate grassland cover characterized the interstadials in the SE Great Hungarian Plain during the terminal phase of MIS3 stage and MIS2 stage. These interstadials correlate well with the Dansgaard-Oeschger (DO) interstadials 8, 7, 6, 5, 4, 3, 2 and the post LGM warm interval (DO1) seen in the Greenland ice core oxygen isotope records. Intervening cold phases, on the other hand, their start correlating with Heinrich event 1, 2, 3, 4 and the LGM. These data overall confirm that millennial scale climate variability during MIS3 and MIS2 stages had profound effect on the terrestrial ecosystems in the continental interior of SE Europe leading to periodic mixed-leaved boreal forest steppe within mosaic structure expansions in the interstadias and cold steppe, taiga and tundra like spots mosaic and mixed vegetation expansion in the stadial phases. 1400–1500 year long drier interstadial and humid, extended vegetation covered stadial periods interchanged at Katymár sequence, therefore the vegetation and fauna development was different from the Atlantic and Central European schemes and inferred an independent local and regional South Carpathian Basin trend.
... In this paper, we describe a new species of Punctum, which was found during surveys of land snails in Siberia, Central Asia (Meng, 2008;Horsák et al., 2013Horsák et al., , 2015Hoffman et al., 2011) and Alaska (J. Nekola and B. Coles, unpublished data). ...
... All recorded shells and live snails were collected and kept dry, and identified under a dissecting microscope using the available literature (Pilsbry, 1948;Meng, 2009;Horsák et al., 2010Horsák et al., , 2013Horsák et al., , 2015 and the authors' comparative collections of Siberian snail specimens. All samples are stored in the collection of Michal Horsák (Brno), Stefan Meng (Greifswald), Lucie Juřičková (Prague), Jeffrey C. Nekola (Albuquerque) and Brian F. Coles (National Museum of Wales) if not otherwise stated for the holotype and paratypes. ...
Article
Punctum lozeki, new species, is described from damp forests and meadows of southern Siberia, Central Siberian plateau, southern Far East (Russia) and Alaska. The species is characterized by a very narrow deep funnel-shaped umbilicus and tumid whorls that expand rapidly in diameter. It was recorded at 21 sites, and it seems to be a rare species, particularly in Asia, being found at only approximately 5% of all explored and potentially suitable sites. It was limited to wet and mesic taiga, mostly inhabiting brook alluviums, wet calcium-rich woodlands, and treeless sedge marshes. It is only the third Punctum species, along with P. pygmaeum and P. ussuriense, currently recognized in extratropical Eurasia. It also seems to represent another example of a species with a Beringian distribution stretching from Alaska in North America to the Altai Mountains in Central Asia.
... Even if we cannot compare absolute commonness in glacial and modern times using the macrofossil database, we can (i) verify the occurrence of a target species in the glacial or early postglacial period, which is a basic prerequisite for assigning species as a glacial or early postglacial relict (this kind of evidence has a long tradition in malacology; Ložek 2001, Horsák et al. 2015a); (ii) reveal past occurrences in the regions where the species does not currently occur, such as the glacial occurrence of mountain and boreal species in the central-European lowlands (e.g. Catoscopium nigritum and Drepanocladus trifarius in the Borská lowland or Sarmentypnum sarmentosum in the Labe river lowland; Hájková et al. 2012b), (iii) trace the disappearance of putative glacial relict species in individual profiles throughout the middle or late Holocene . ...
... Such a tendency is also well documented by the distribution of S. scorpioides in Belarus, where this species occurs only in the western part of the country close to the Baltic, where it is common (Pakalne & Kalnin , a 2000, Maslowsky 2017). This pattern, however, calls into question their frequent occurrence in glacial times, which were characterized by a distinctly continental climate (Horsák et al. 2015a). Although they can grow in distinctly continental landscapes such as those at high altitudes in the Altai Mts (Ignatov 1994), they were probably restricted in their distributions during the full glacial times and spread there as long ago as in late glacial times (Rybníček 1966. ...
Article
Modern databases containing large amounts of botanical data are a promising source of new results based on large data analyses. We used a new database of plant macrofossils of the Czech and Slovak Republics to compare the recent distributions of putative relict species of fen bryophytes with their past distributions since the late glacial. All the species studied occur in lateglacial sediments, but mostly in regions where they are recently recorded (19–21st centuries). There are specific regions rich in putative relict species of fen bryophytes both in late glacial / early Holocene times and recently. In some cases the target species were, however, found outside the recent distribution range where environmental conditions are no longer suitable for their occurrence. We further found that the total number of the glacial and early-Holocene records greatly exceeds the total number of records for the middle Holocene, when succession to woodlands or bogs resulted in a reduction in species of bryophytes that are specific to open rich fens. The observed patterns may imply a relict status of the target species. We especially documented a substantial decline in the abundance of species requiring a high and stable water level (Drepanocladus trifarius, Meesia triquetra and Scorpidium scorpioides), both throughout the Holocene and during the most recent transformations of the landscape. In contrast, those species that tolerate transient decreases in water level persisted into recent times at more localities (Calliergon giganteum, Hamatocaulis vernicosus, Paludella squarrosa). Macrofossil data cannot, however, provide a quantitative analysis of the distribution of a species, because the number of recent data usually greatly exceeds the number of fossil records. The reason is that the area sampled in palaeoecological research is very small as it is time-consuming and expensive; cores or excavations usually are of only a few square centimetres. Despite this shortcoming, macrofossil data are an important, but not the only, source of evidence for the identification of the relict status of a species.
... 11'600 yr BP) the Central European landscape was still cold and arid and sparsely wooded especially by open birch-pine forests (Abraham et al. 2016(Abraham et al. , Šolcová et al. 2018. The best present-day analogue of the Pleistocene environment is believed to occur in cold and dry areas of continental southern Siberia (Horsák et al. 2015, Chytrý, Horsák, et al. 2019. It is likely that many of the temperate woodland taxa went extinct in the Pannonian Basin during the Last Glacial Maximum (ca. ...
... The recent interpretations of paleoecological data suggest that there has always been some proportion of the non-arboreal communities in the landscape, even before the onset of Neolithic agriculture (Magyari et al. 2010, Kuneš et al. 2015, Pokorný et al. 2015, Feurdean et al. 2015, Abraham et al. 2016, Šolcová et al. 2018). These pollen-based assumptions are supported by the fossil evidence of several mollusc species that are not able to survive in a closed forest, especially Chondrula tridens and Helicopsis striata (Ložek 2007, Pokorný et al. 2015 and by the present-day abundance of species considered as glacial relicts (Horsák et al. 2015). Another more direct proxy is the high genetic diversity in Western or Central Europe of populations of some forest-steppe species, e.g. ...
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Forest-steppe landscapes are formed of a complex mosaic of physiognomically contrasting habitats (forests and grasslands) and thus are generally species-richer than the landscapes formed of habitats of similar physiognomy. In the northern Pannonian Basin, the majority of forest-steppes are exposure-related, i.e. steppe is occupying different proportions of the south-facing slopes of the otherwise forested hills. Historically most of these sites were influenced by extensive farming but nowadays most of them are abandoned or managed for conservation purposes. They harbour a high number of rare or endangered species and supply a wide spec-trum of ecosystem services. This study is aiming at: (i) identifying sites in the northern Panno-nian Basin in which the exposure-related forest-steppe vegetation mosaics are well developed; (ii) describing individual habitats of the exposure-related forest-steppe landscapes; (iii) char-acterizing vegetation dynamics at the forest-steppe boundary in the exposure-related forest-steppe by analysing the pattern of woody species regeneration; (iv) comparing plant species diversity of exposure-related forest-steppe sites on different bedrocks. The study was con-ducted in different regions of the northern Pannonian Basin: the loess hilly landscape of south-ern Moravia (CZ), limestone hills of the westernmost Carpathians (AT, CZ, SK), Transdanubian Mts (H) and the Transcarpathian Lowland (UA). Sixteen sites with well-developed exposure-related forest-steppe ecosystems were chosen, and on each of them, all physiognomy-defined habitats of the south-facing slope (i.e. forest, steppe and the ecotones between them) were sampled along a transect. All individuals of woody species including juveniles were sampled in 1 m intervals. The floristic differentiation of forest-steppe habitats was assessed using prin-cipal coordinate analysis (PCoA) and calculating the fidelity of species to certain habitats. The density of woody seedlings was compared among the habitats. It was found that the floristic differentiation between habitats in the forest-steppe mosaic is relatively low, and a high number of species is shared between neighbouring habitats. Species composition of ecotones was intermediate between steppe and forest. The species richness of the ecotones was higher on the sites with hard rocks, while on deep soils, it was similar to those of both steppe and forest. Juveniles of woody species were most abundant in forest, while in steppe they were sparse or missing. Several woody species appeared to have an affinity to some forest-steppe habitat. This study contributes to the understanding of the patterns and processes that maintain plant diversity in the exposure-related forest-steppe landscapes by comparing individual compo-nents of this landscape mosaic in various regions at the western border of the Eurasian forest-steppe biome.
... Today, most of these species have typical arctic-alpine distribu- tions, being widely dispersed across Fennoscandia with a few relict populations surviving at high elevations in Central Europe (e.g. Horsák et al., 2015). Some isolated populations also persist in environmentally stable habitats, such as mires, in lowland regions (Schenková & Horsák, 2013). ...
... This is not really surprising, since any snail species that specifically avoids calcium-rich environments is very unlikely to be preserved in calcareous deposits such as loess (Ložek, 2001). Its presumed relict status is based largely on its rare and scattered modern distribution in temperate mainland Europe, in contrast to its continuous distribution in northern Europe, where it occurs together with other glacial species characteristic of loess communi- ties (Horsák et al., 2015). The only fossil records from mainland Europe come from Germany and Poland (Jaeckel, 1962), but these records are old, unverified and poorly dated. ...
Article
The aims of this study were to analyse whether land snail assemblage patterns reflect the gradient of calcium content on a very small scale within a site. We chose two sites differing in their calcium richness and source of the calcium. The "Tufa site" had abundant tufa in the soil, while the "Boulder site" was on chemically inert chert bedrock where calcium originated from vegetation. On each site a set of 20 quadrats from which snails were extracted was laid down in a line from the calcium rich patch to the calcium poorer surround matrix. At the both sites, available calcium contents of topsoil decreased significantly with samples' position from the patch centre. For the Tufa site, calcium content was a strong controller not only of species composition but also species richness, total abundance, and abundances of almost all species. At the Boulder site only species composition was significantly driven by calcium. Species composition was highly nested along calcium gradient at the Tufa site contrary to the Boulder site where the species had an almost random distribution. We conclude that the best predictor of species composition was in both cases content of carbonate calcium in topsoil. Topsoil pH was positively correlated with calcium content only at the Tufa site.
... Some general areas were targeted for sampling due to prior reports of disjunct European elements 25 . The snail fauna of each site was assessed via direct hand picking, dry or wet sieving 17 . All shells were sorted and identified to the species level based on Nekola et al. ...
... It seems likely a similar mechanism is responsible for the LGM shift to xeric habitats in Columella columella 16,36 , another characteristic full-glacial land snail species 18,31 . Some modern C. columella populations in the high Alps and Carpathians are known to inhabit mesic tundra as well as fens 17,25 . However, because this species was never able to colonize Iceland, there is no way to determine if xeric sites in that landscape could support its occurrence, or to empirically show that its realized niche is able to expand in this way. ...
Article
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The presence of Last Glacial Maximum (LGM) biotic communities without modern counterparts is well known. It is particularly evident in central European fossil LGM land snails whose assemblages represent an odd mix of species that are currently limited to either xeric or wetland habitats. Here we document a genetically verified discovery of the modern calcareous wetland species Pupilla alpicola on Iceland, where it is limited to dry grasslands. This species also represents a common European LGM fossil, and its new records from Iceland help explain puzzling shifts of some glacial land snails of xeric grassland habitats to open wetlands today. Similarities between the climates of modern Iceland and LGM Eurasia suggest that this species did not become limited to wetlands in continental Europe until after the Late Pleistocene–Holocene climate transition. These results are a strong reminder that assumptions of ecological uniformity must be questioned and that the quality and robustness of palaeoecological reconstructions is dependent upon adequate knowledge of the full autecological range of species over time.
... Steppes on southern and south-western hill slopes in warm regions of the Czech Republic (thermophyticum sensu Skalický 1988) are unique biotopes across the whole of Europe and represent the most important remnants of the communities that have been widespread across central European lowlands since the last glacial period (Ellenberg 1988;Čížek et al. 2012;Pokorný et al. 2015;Divíšek et al. 2020). These habitats host several steppe species of plants and animals characteristic for glacial loess steppes during the cold intervals of the Pleistocene (Horsák et al. 2015). Human activities have maintained their occurrence since the Neolithic, keeping the landscape open (Ložek 1964;Horsák et al. 2010;Pokorný et al. 2015). ...
Article
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Bees and wasps inhabiting steppe formations are, according to recent red lists, among the most endangered species, quickly disappearing from local faunas of central European countries. Several species, which are specialised nesters in empty gastropod shells, show the opposite pattern. Based on their distribution maps, we found that these species are recently more common and widespread than in the past. In contrast, the bee species nesting in the ground showed clearly the opposite result. They are much rarer now than in the past. Climatic data suggest that the bees nesting in gastropod shells have expanded towards higher altitudes and/or areas of lower mean July temperature. The main reasons of these distributional shifts are (i) unfavourable nature conservation management practices during the twentieth century, resulting in the successional shifts of steppe formations towards shrubs and mesic grasslands, (ii) human-driven change of many sites towards wood plantations or crop fields, and (iii) the increase of the number of available gastropod shells (i.e. nesting resources for shell-nesting bees) in steppe habitats as the response to the mentioned successional changes.
... Consequently, the total species richness per se may not be a sufficient criterion in biodiversity conservation, as well as in general ecology and biogeography when attempting to disentangle relationships between the species richness, environment and history. Despite that, specialists and matrix-derived species have been treated separately in only a few regional studies (e.g., Bergamini et al., 2009;Hájek et al., 2007;Horsák et al., 2015). ...
Article
We present the first continental-scale study of factors controlling the species richness of groundwater-fed fens, comparing land snails, vascular plants and bryophytes. We separately analyse two ecologically distinct groups differing in conservation value and colonization/extinction dynamics, that is habitat specialists, and matrix-derived species. Considering the island-like nature of fen habitats, we hypothesize larger differences in the species richness–environment relationships between habitat specialists and matrix-derived species than among the taxonomic entities.
... Palynological studies have indicated that the pollen spectra from eastern Central Europe from the last glacial period are very similar to the modern surface pollen spectra from the Altai-Sayan Mountains, suggesting a high similarity between European palaeovegetation and modern vegetation of these mountains (Kuneš et al., 2008;Magyari et al., 2014). Indeed, whole extant assemblages of snails that were typical of European Late Pleistocene loess sediments but which strongly retreated or went extinct in the Holocene were recently discovered in the Altai and the adjacent mountain ranges (Meng, 2009;Horsák et al., 2010Horsák et al., , 2015Hoffmann et al., 2011;Nekola et al., 2015). Furthermore, a comparison of fossil and recent mammal faunas across Eurasian regions revealed that the southeastern Altai-Sayan region experienced the lowest rate of extinction of the Pleistocene fauna and that its modern mammal fauna is most similar to the Pleistocene fauna (Pavelková Řičánková et al., 2014). ...
... The phenomenon of isolated occurrences of boreal species in the fens of south-eastern Europe [6,36] is fascinating and inspiring, because it may give a hint to understanding European Quaternary history [10]. Isolated occurrences of mire species are further extraordinarily important in biodiversity conservation [7]. ...
Article
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The boreal-arctic species Carex vaginata Tausch (Sheathed Sedge) has been found in Romania in 2014 during research on fen ecology. The species grows in the Rarau Mts., in an alkaline sloping fen of the Caricion davallianae alliance, transitional to Cratoneurion commutati springs (SE margin of Mt. Popii Rarăului, close to Plaiul Todirescu grasslands) and in a subalpine grassland at Mt. Popii Rarăului. Rarau Mts is the fourth locality for this species in the entire Carpathians, underlining its relictual distribution pattern in temperate Europe that may represent a remnant of formerly more continuous distribution during the glacial period. We further present a vegetation-plot record with Carex vaginata and basic chemical parameters of the water.
... *Corresponding author: horsak@sci.muni.cz estimated for the full-glacial of Central Europe had very similar assemblages to those of the latter in terms of species composition (Meng, 2008;Horsák et al., 2010aHorsák et al., , 2015. ...
Article
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.
... S. kotulae is still noted in southern Poland (Wiktor, 2004;Welter-Schultes, 2012), whereas it appears to be the last appearance of V. tenuilabris in the region. Now it occurs only in Siberia, Central Asia and northern China (Gerber, 1996;Meng, 2008Meng, , 2009White et al., 2008;Horsák et al., 2010Horsák et al., , 2015. Progressing warming and some oceanic influences favoured the appearance of species typical of warm and humid conditions Sphyradium doliolum, Discus rotundatus, Aegopinella pura, Helicigona faustina, Cochlodina orthostoma and others Krajcarz et al., 2020). ...
Article
The Holocene has always been considered a crucial epoch where the major cultural steps of humankind took place. Understanding past Holocene climatic variability, shifts in vegetation, and faunal communities are among crucial challenges in predicting the upcoming changes of the natural environment. In Central Europe, the Kraków-Częstochowa Upland is one of the best study areas for detecting faunal, environmental, and climate shifts. For that reason, we choose to study the small vertebrate and mollusc communities of three archaeological sites with Holocene horizons: Sąspowska Zachodnia Cave, Ciasna Cave, and Małe Rockshelter. We analysed the faunal community shifts, and we compared our results with the major Holocene sites in the area with the correspondence and cluster analyses. The environment and climate were reconstructed through Habitat Weighting, Quantified Ecology, malacological spectra and Bioclimatic methods. Environment and climate values showed a temporary replacement of the Late Glacial tundra and grasslands with boreal forest mixed with open wetlands during the Preboreal and then stable landscapes dominated by temperate forests. Models based on rodent data show that temperature and precipitation generally increased, starting from the Boreal, to reach values near, or sometimes overwhelm, the current climate values of the area during the Atlantic and Subboreal. The climate values obtained from herpetofaunal analyses show very slight changes between the Preboreal and the Atlantic (Ciasna Cave layers 3 and 4), except for the winter temperatures, whereas results obtained for Sąspowska Zachodnia Cave are probably biased by a very low number of recovered remains. Among rodents, a major negative oscillation detected in layers related to the edge of Early Holocene was related to the Preboreal Oscillation or long-lasting effect of the Younger Dryas.
... This region represents one of the most important biogeographical crossroads in Eurasia, playing a major role in the Quaternary survival of contrasting biogeographical elements (B alint et al., 2011;Mr az & Ronikier, 2016). Indeed, in a pan-European synthesis of fossil pollen data (Fyfe, Woodbridge, & Roberts, 2015) this region represented, during the entire Holocene, the contact zone between temperate mountain landscapes with needle-leaved forests and Eastern European lowland landscapes with broadleaved forests, and during the Middle Holocene also between grassland-rich south-eastern Europe and more (Kliment, Sib ıkov a, & Sib ık, 2011), steppe-tundra (Hors ak et al., 2015) or forest-steppe (Feurdean et al., 2015;Kune s et al., 2015). Furthermore, the rarity of some species associated with boreal coniferous forests may reflect a refugial distribution of this habitat in the Late Holocene prior to the industrial era (Ku cera, 2012). ...
Article
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Aim: The regional co-occurrence of contrasting bioclimatic elements (warm-temperate , continental, boreal, arctic-alpine) may be shaped by the distribution of their glacial or post-glacial refugia. We tested this hypothesis using pollen proxies in a region where such refugia are expected, but not unequivocally demonstrated. Location: East-Central Europe (Western Carpathians and adjacent regions). Methods: We compiled pollen spectra from 112 sites distributed across various landscapes for six time-periods from the Late Glacial to the present. Compositional patterns were assessed by principal coordinates analyses (PCoA) with a sensitivity analysis based on a bootstrap technique. Site PCoA scores were interpolated geographically and correlated with palaeoclimatic models. Results: Consistently over the last 15,000 years, the first ordination axis sorted samples according to the proportion of deciduous temperate trees, while the second axis consistently followed an altitudinal gradient that coincided with temperature. The principal gradient was more important than the altitudinal gradient except for the Late Glacial and Bronze & Iron Ages, when both gradients were of similar importance. The fine-grained pattern in the present mountain landscape was formed as late as during early modern colonization. Main conclusions: Since the Late Glacial, the landscape has been differentiated into temperate, continental and cold regions. This finding supports the hypothesis that refugia are a key factor for understanding current biogeography in Central Europe. The Late Glacial occurrence of temperate trees is unlikely to be explained only by gradual migrations from southern Europe. Humid but relatively warm mountains hence might have acted as glacial refugia of temperate forest species, while low-lands and leeward basins might have acted as post-glacial refugia of steppe grasslands. The strong contrast between forested (temperate) and more open continental landscapes during the Early Holocene seems to correspond with recent diversity patterns. Our results highlight the relevance of integrating past landscape trajecto-ries into modern biogeographical models.
... For Siberia (the Altai, the Western Sayan) we used samples collected for another project on recent similarities with European glacial landscapes (e.g. Horsák et al. 2015). We included Siberia because of the possible effect of Pleistocene history on the genetic structure of S. warnstorfii, that is acknowledged by Mikulášková et al. (2015). ...
Article
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Peat mosses are a key functional group in peatlands, driving biogeochemical cycles, habitat development and changes in species composition. They are generally intolerant of calcium and magnesium bicarbonate, but some species are adapted to mineral-rich fens. A previous study found a coincidence between genetic variation and the ability to tolerate high pH/calcium levels in Sphagnum warnstorfii. Here we compare its microsatellite variation with that of two rarer calcium- tolerant species (Sphagnum subnitens, S. contortum), using a novel data set from Eurasia. Because physiological experiments indicate that S. warnstorfii can tolerate high magnesium levels, we included also samples from dolomite and serpentinite. Genetic diversity of S. warnstorfii was higher than that of other species. The Bayesian analysis in program Structure resulted in two population groups of S. warnstorfii. One group coincided with dolomite (Italy, Austria, Estonia) and moderately magnesium-rich (but calcium-poor) rocks (serpentinite, metadolerite, cordieritebearing migmatite on the Bohemian Massif), while the second one coincided with magnesiumpoor bedrock across Eurasia. The principal coordinate analysis revealed a cline between populations from magnesium-rich and magnesium-poor bedrocks, with populations from dolomite and serpentinite forming one extreme. Populations from magnesium-poor bedrock located far from any dolomite or serpentinite formed the opposite extreme of the cline. We demonstrate for the first time that magnesium toxicity may drive bryophyte microevolution, as has repeatedly been shown for vascular plants, including ferns.
... Shells and their fragments were hand sorted under a dissecting stereomicroscope and identified using the methodology devised by Ložek (1964) and Horsák et al. (2013). The latter was also used for mollusc nomenclature with some updates according to Horsák et al. (2015). ...
Article
The Middle Pleniglacial on the northern hemisphere is characterized by millennial scale, frequent, and high amplitude environmental climatic shifts. In loess-paleosol sequences (LPSs) the transition from the Lower (MIS 4) to the Middle Pleniglacial (MIS 3) was accompanied by significant erosion events, as recorded in various terrestrial archives across Central Europe. As a result, potentially existing paleosol horizons of the particular period have been widely erased from the LPSs and only little is known about pedogenesis in this vast area. This study tries to fill this gap in our knowledge through studying a pedo-sedimentary record from Bíňa, a site situated in a relatively poorly investigated area on the northwestern edge of the Carpathian Basin. We investigated a 12-m-high and more than 500-m-wide outcrop along the Hron River, where a well-structured sequence of fluvio-lacustrine sediments, paleosols, and loess is preserved. The paleoenvironmental development within the studied area is presented and discussed on the basis of soil micromorphology, rock-magnetic and geochemical measurements, grain size analyses, and a malacological record accomplished by luminescence dating. Based on the OSL dating of the central profile, the sequence provides a high resolution record of the time period 60-20. ka (MIS 3-MIS 2). The most developed paleosol horizons (Phaeozem and two Bw horizons of Gleyic Cambisols) were dated to the early stage of the MIS 3 (60-50. ka) and correlated with the Greenland interstadials GI-17/16, GI-14/13 and/or GI-12 (the northwestern European interstadials Oerel, Glinde and Moershoofd). The development of the Bw horizon of a Cambisol recorded within the younger loess body was dated to ca. 35. ka and it probably corresponds to the late MIS 3 intestadials GI-8-5 (Denekamp). The preservation as well as the development of the well-stratified record were closely related to the specific hydro- and geomorphological settings of the paleo-channel structure in which the sequence developed. Although soil development benefited from the specific local settings, we suggest that the recorded paleosols can also be related to the interregional climate differences of the Carpathian Basin: within the northerly located pericarpathian zones, a moister climate predominated during the last Pleniglacial, in contrast to the drier continental areas to the South. Thus, a sharp climatic transition existed separating a semi-arid steppe region from a climatic zone under the persistent influence of Atlantic air masses.
... This poor malacocoenosis contains from 5 to 60 shells per layer (Fig 7), mostly of open-country gastropods, which constitute 35-73% of all individuals. Vallonia tenuilabris, a glacial relic that occupies the open areas of so-called "cold-steppes", the alpine grasslands, taiga, hemiboreal forest and wooded fens [77,[131][132][133][134], is the most numerous in the central zone, being accompanied by the shade- loving species Discus ruderatus and Semilimax kotulae, both characteristic of a cool and continental climate. In the entrance zone, D. ruderatus slightly outnumber V. tenuilabris (Fig 7). ...
Article
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A cave site Shelter in Smoleń III (southern Poland) contains an approximately 2-m-thick stratified sequence of Upper Pleistocene and Holocene clastic sediments, unique for Central Europe. The sequence contents abundant fossil fauna, including mollusk, rodent and bat remains. The cave sites with long profiles of subfossil fauna present a great value for reconstructions of regional terrestrial paleoenvironment. We explore the stratigraphy of this site through analyses of the lithology and geochemistry of sediments, radiocarbon dating of faunal and human remains and charcoals, and archaeological study, as well as the paleoecology derived from the taxonomic composition of fossil faunal assemblages. Our data show that the entire period of the Holocene is recorded in the rockshelter, which makes that site an exceptional and highly valuable case. We present paleoenvironmental reconstructions of regional importance, and we propose to regard Shelter in Smoleń III as a regional stratigraphic stratotype of Holocene clastic cave sediments.
... During the coldest and driest phases of the last glacial cycles, the common land snail fauna of Central Europe differed substantially from that of interglacials and included a number of cold-adapted species (Ložek, 1964(Ložek, , 1973Mania, 1973;Alexandrowicz, 1987;Fűköh et al., 1995). Given the present habitats of typical species of the coldest phases (Horsák et al., 2010(Horsák et al., , 2015, survival of the Roman snail seems unlikely even in sheltered locations. ...
Article
The Holocene postglacial expansions offer a possibility to assess how quickly snails, with proverbially slow active dispersal and unclear passive dispersal capabilities, can naturally spread. We explore the possibilities and limitations of such an approach on an iconic European snail. We locate probable sources of postglacial expansion of Helix pomatia and date its earliest postglacial subfossil occurrences in Central Europe, close to presumed natural northern distribution limits. With dense sampling of the species native range, we found most of its mitochondrial diversity at c. 45 °N or more southerly. A hotspot in Bosnia and Serbia contributed to colonization of the Pannonian Basin, Carpathians and Bohemian Massif; however, the most widely distributed lineage has a more westerly centre of diversity, with a possible refugium in Italy. We revised the oldest occurrences post-dating the Late Pleniglacial from Czechia and Slovakia suggested by literature and radiocarbon-dated the preserved shell fragments. No reliably identified fragment yielded a date earlier than 10121–9695 cal BP. Stratigraphy-based records presumed older turned out to be unreliable, making the argument for a direct dating approach. However, our results confirm that in the absence of northern refugia, H. pomatia must have been able to colonize the newly emerging postglacial habitats rapidly.
... Especially in recent decades, the study of the loess snail in the world has made great progresses, including snail's ecological distribution, high resolution fossil record and stable isotopes analysis of terrestrial snail shell etc. (e.g. Metref et al., 2003;Balakrishnan et al., 2005;Zanchetta et al., 2005;Antoine et al., 2009;Yanes et al., 2009;Kehrwald et al., 2010;Zaarur et al., 2011;Huang et al., 2012;Colonese et al., 2013;Eagle et al., 2013;Yanes and Romanek, 2013;Horsák et al., 2015). ...
Article
The terrestrial mollusk fossils found in Chinese loess strata have been studied for over one hundred years. However, the greatest progress in these studies has been made only in the last two decades. In this paper, we review the advancements, advantages and limitations of terrestrial mollusk studies in Chinese loess deposits. Improvements in research methods and approaches have allowed the extraction of more detailed paleoenvironmental and paleoclimatic information from mollusk assemblages. The broadened research scope and content have yielded many new findings and results. The mollusk record has thus become one of the most important proxies in the paleoenvironmental and paleoclimatic reconstruction of loess-paleosol sequences in China. The greatest progress in the studies of terrestrial mollusks in Chinese loess sequences can be summarized as follows: (1) modern mollusk assemblages can be classified into four ecotypes, based on their temperature and humidity requirements, including eurytopic, semi-aridiphilous and sub-humidiphilous, cold-aridiphilous, and thermohumidiphilous types; (2) Quaternary mollusk assemblages can be modified into the following three ecological types: glacial loess, interglacial paleosol, and interstadial weakly-developed paleosol assemblages; (3) mollusk records successfully reveal long-term climatic and environmental changes reflective of the history of East Asian monsoonal variations since the Late Cenozoic, and the succession of mollusk species also indicate short-term environmental changes such as millennial climate variability during Last Glacial Maximum and unstable climatic fluctuations during glacial and interglacial periods; and (4) more recently, new analytical approaches have offered increased research potential in areas such as paleotemperature reconstruction using the isotopic compositions of modern and fossil mollusk shells, combined with higher accuracy 14C dating of Quaternary loess deposits, which will greatly improve future loess paleoenvironmental research.
... In this context, the main shortcomings of fossil mollusc assemblages are related to the fact that most of these samples are poorly defined in space and time, which can bias interpretation for two principal reasons. First, there are differences among species in their potential to enter the fossil record and also in the capacity of sediments to preserve calcareous fossils ( Hors ak et al., 2010Hors ak et al., , 2015). Second, the analysed layers of the sediment certainly differ in their accumulation time and thus also in the shell accumulation of species living at the site over a variable period. ...
Article
This study investigates the effects of sample volume, resolution and accumulation time on the interpretation of a mollusc record from terrestrial deposits. We tested (i) if, and to what extent, small sample sizes impoverish the reconstructed mollusc species richness, and (ii) whether fine-resolution sampling is worth the effort. We analysed three mollusc sequences, covering continuously the entire Holocene, which were collected in a tufa-forming spring fen and differed in sample size and resolution. More than 36 000 specimens of 76 species were processed. Using a rarefaction method, we observed that different sample volumes had a significant effect on the recorded species richness in this type of environment, and 100-cm3 samples seemed to be sufficient for a reliable reconstruction in tufa deposits. Although the thickness of the sampled layers had no influence on the palaeoecological interpretation, we observed a shift in the mollusc diversity peak once different resolutions were applied. Furthermore, the layers of finer resolution allowed detailed radiocarbon dating and better understanding of trends in species dynamics. We also observed significant confounding relationships among the number of species, number of specimens and accumulation time that can be disentangled if a precise depth–age model is available.
... From the previous work we know that the response can fundamentally differ between habitat specialists, with high affinity to a given ecosystem, and ubiquitous matrix-derived species, i.e. those inhabiting also adjacent surroundings with ability to colonize various types of habitats (e.g. Bergamini et al., 2009;Horsák et al., 2015;Horsáková et al., 2018). To account for such potential differences, we opt for testing these two groups separately, however, followed by some analyses with undivided dataset to uncover drivers of the overall species composition. ...
Article
Human activities have enormous impact on current biodiversity distribution across all spatial scales. Despite the numerous studies showing the difference between preserved and impaired sites, only little is known about the regional scale. Therefore, we selected four European regions differing in habitat conservation status (HCS) to explore if the variation in land snail communities reflects regional differences. We collected quantitative land snail samples at 169 isolated spring fen sites and measured environmental parameters. The species richness of habitat specialists expressed low variation and weak associations with local conditions in the regions of adequate-HCS, presumably because of their common occurrence throughout most sites. In contrast, the richness of matrix-derived species, i.e. predominantly habitat generalists, was highly variable in the two regions of adequate-HCS and also tightly associated with local conditions, especially moisture. In both the intermediate and the inadequate-HCS region, these associations were much weaker as the fens are less extreme and allow for infiltration of matrix-derived species. Population densities of Vertigo geyeri , an umbrella species being internationally protected by the E.U. Habitat Directive, were highest in the two high-HCS regions. Species composition was primarily controlled by moisture in the regions of adequate-HCS, while in the remaining regions, those predictors that are less easily jeopardized by human impact, such as climate, water chemistry and terrain topography, gained importance. In the inadequate-HCS region, none of the analysed predictors was associated with the main compositional gradient, suggesting a complete disruption of community-environment relationships. Our results suggest that the species richness and community responses to natural gradients might be substantially modified by human impact, although the effect of some other regionspecific factors cannot be easily disentangled because of inevitably low number of study regions.
... For example, wetland habitats and vegetation types with Trichophorum pumilum, rich in minerals and derived solutes, are widespread in dry and cold mountain regions of Central Asia, whereas in temperate Central Europe such vegetation is restricted to isolated habitats such as small travertine fens in West Carpathian locations where fen patches have persisted throughout the entire Holocene (Dítě et al., 2013;Hájková et al., 2020). Furthermore, steppe areas in West and Central Asia have been used as recent analogues for palaeoreconstructions of full-glacial conditions (Horsák et al., 2010;Horsák et al., 2015;Janská et al., 2017). It is likewise suggested that highland mires in Iran, as a country in West Asia, can be regarded as "interglacial refugia" for boreal, glacial relict vascular plant species (Kürschner et al., 2015). ...
Article
Aims To develop the first comprehensive syntaxonomic classification for patchy montane mire and spring vegetation across the Irano‐Turanian phytogeographical region in Iran, Tajikistan and Kyrgyzstan and to explore the effects of the main environmental and geographic gradients on their distribution. Location Alborz Mountain range (Iran), Pamir‐Alai Mountains (Tajikistan) and Tian Shan Mountains (Kyrgyzstan); total area of about 3,000,000 km². Methods A database of 1,015 vegetation relevés including a total of 675 vascular and bryophyte taxa was established, covering the large mountains ranges of the Irano‐Turanian regions in Iran, Tajikistan and Kyrgyzstan, at altitude ranging from 1,300 to 4,505 m a.s.l. A semi‐supervised k‐means analysis was performed. Additional analyses were carried out to show differences among predefined vegetation groups in terms of phytogeographical, climatic and compositional factors. Results Two large groups of mire communities, referable to mires and springs were identified. Twenty associations, one subassociation and four alliances of these wetlands, belonging to three orders and three classes, were defined. Among them, ten associations, one subassociation and two alliances were formally described as new syntaxa. Moreover, a new order Caricetalia orbicularis, that comprises phytogeographically unique vegetation of the Irano‐Turanian mires, is proposed. Conclusions Based on our results and comparison between the Irano‐Turanian and other phytogeographical regions, we propose a first comprehensive syntaxonomic synopsis for the Irano‐Turanian mires and springs. Despite some identical character species and their vicariant nature, the Irano‐Turanian, Euro‐Siberian and Mediterranean mires and springs show considerable compositional differences. Irano‐Turanian wetlands contain a large and distinct set of endemic plant species, which are mixed with Euro‐Siberian species in the west and north, but predominate in the eastern locations. They have patchy pattern and scattered distribution and serve as inland hotspots of wetland vegetation in the vast dry landscape of SW and Central Asia.
... For example, the terrestrial mollusk Vertigo genesii (Gredler), which is common in European deposits of the late Pleistocene age, is now widespread in Northern Europe and has recently been found in England and the Alpine highlands (Coles, Colville, 1980;Schenková, Horsák, 2013 and references therein). Mollusks and plants characteristic of the glacial-age deposits still inhabit the mountains of southern Siberia (Horsák et al., 2015) and North America (Miller, 1996). Similar examples are especially numerous in earlier publications (review: Birks, 2008). ...
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In this study, we tried to understand why the biota of northern regions is similar to that of southern mountain regions. Phylogeographic studies of several Arctic-alpine plants (Arabis alpina, Bistorta vivipara, Carex atrofusca, Gentiana sect. Cruciata, Koenigia islandica, Oxyria digyna, Ranunculus glacialis, Saxifraga oppositifolia, Sibbaldia procumbens, Trollius europaeus, Veronica alpina, Lagotis spp., and Pedicularis spp.), insects (Oeneis spp. and Arcynopteryx dichroa), and a mammal species (Ovibos moschatus) indicate that the respective groups emerged in the mountains of the temperate climatic zone and then migrated to the Arctic. As paleontological findings indicate, the mountains of the temperate and tropical zones provided habitats for the ancestors of several Salmonidae genera and at least some of the mammalian species common to the Eurasian mammoth steppe (Mammuthus primigenius, Coelodonta antiquitatis, Bos (Poëphagus) baikalensis, Alopex lagopus, and Panthera spelaea). A hypothesis is suggested to explain the crucial role of mountain regions in the evolution of northern forms. Additionally, colonization events by Arctic taxa in the mountains of temperate climatic zones have been demonstrated in a few studies.
... Land snail communities of temperate and boreal zones are well known to be driven by several ecological gradients, mainly related to the calcium level (Wäreborn, 1970;Hylander et al., 2005;Juřičková et al., 2008), soil moisture and architecture (Wäreborn, 1992;Nekola, 2003;Hettenbergerová et al., 2013) and habitat history (Cameron et al., 1980;Kappes, 2006;Horsák et al., 2012). Additionally, the effect of macroclimate (Horsák et al., 2013a;Nicolai & Ansart, 2017;Horsák et al., 2018) and the existence of shelters to overwinter (Millar & Waite, 1999;Ansart et al., 2001;Horsák et al., 2015) have been found to be important. Although we know a good amount about the bottom-up limitation of snail communities, little is known about top-down mechanisms constituted by snail predators, pathogens, and other natural enemies (see Barker, 2004). ...
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Shell formation is the main defensive strategy against predation for the majority of snails. Therefore, various predators have had to develop a variety of techniques how to overcome this barrier. As shells can persist in a calcium-rich environment for a long time, specific external or internal traces on shells left by predators indicate whether and who killed the snail. Based on litter samples collected at 30 sites of five different habitat types, the intensity and type of predation were assessed. The minimal predation rate varied between 0.0 and 21%, with an average of 8%. The highest rate was observed at limestone steppes, on average 15%. Beetles were found to be the most common predators of snails; however, predation by snails was more common in calcareous fens. Predation by some vertebrates and dipteran flies was also recognised. To test the role of mouth barriers as a means to reduce predation by carabid beetles that break the shell from an aperture, we analysed the predation rate separately on adult and juvenile shells using 24 populations of the steppe snail Granaria frumentum (Draparnaud, 1801). As expected, carabid beetles chiefly preferred juveniles compared to adult shells (Wilcoxon test, p < 0.001). On the contrary, the parasitoid fly Pherbellia limbata (Meigen, 1830) and Drilus beetles preferred adults. We found that predation by carabid beetles positively increased with prey abundance (R ² = 42.8%, p = 0.021), while no relation was observed for the parasitoid (p = 0.703), likely due to their feeding specialisation.
... Land-snails are well preserved in carbonate loess sequences over the Northern Hemisphere, especially in Europe (Ložek, 1990;Rousseau et al., 1990;Markovi c et al., 2007;Gerasimenko and Rousseau, 2008;Moine, 2008Moine, , 2014, North America (Rousseau and Kukla, 1994;Rossignol et al., 2004) China, and Central Asia (Rousseau and Wu, 1997;Li et al., 2006;Wu et al., 2018;Zong et al., 2020), and are highly sensitive to environmental changes. Analysis of terrestrial mollusc assemblages reveals several advantages: (1) they are widely distributed geographically and are present in most of the stratigraphic horizons; (2) many species have not become extinct since the Pleistocene, but instead have moved geographically in response to climate and environmental changes; and (3) the abundance of modern analogue assemblages can help to improve the understanding of distribution patterns and species assemblages of past communities, and to provide information on ecological processes across multiple habitats (Horsák et al., 2015). ...
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Loess are geological deposits of silt-size mineral dust that together with paleosols (buried soils) form some of the most important terrestrial records of Quaternary climatic change. In the simplest terms, loess units are formed during cold and arid periods when dust accumulation peaks, and are therefore identified as glacial/stadial units. Conversely, paleosols indicate interglacial/interstadial periods as they develop under milder climates when dust accumulation is significantly reduced, and landscape is stable. Within them a number of sedimentological, biological, and geochemical proxies are preserved that when supported by geochronological approaches, offer an opportunity to reconstruct past climates and environments. Extensive loess-paleosol sequences can be found in Europe, Asia, and North and South America, while smaller patches also cover parts of Africa and Oceania. Therefore loess has the potential to improve our understanding of Quaternary changes on local, regional, and continental scales.
... For this reason, the origin and development of Central European forest-steppe has always been a source of controversy (Sümegi et al., 2012). According to current understanding, some steppe patches in this region are relicts of more extensive late-Pleistocene and early-Holocene steppe and forest-steppe environments (Chytrý, Horsák, et al., 2019;Horsák et al., 2015;Niklfeld, 1993;Novák et al., 2019;Zólyomi, 1964). Although the regional macroclimate may have been unsuitable for steppe vegetation in certain periods of the Holocene, particularly during the moist mid-Holocene (Dabkowski et al., 2019;Mauri et al., 2015), the current interpretations of the palaeoecological record support its continuous persistence throughout the Holocene (Feurdean et al., 2015;Kuneš et al., 2015;Magyari et al., 2010;Novák et al., 2019;Pokorný et al., 2015;Šolcová et al., 2018) BP (Feurdean et al., 2018;Magyari et al., 2010;Novák et al., 2019;Šolcová et al., 2018). ...
Article
The occurrence and origin of dry grasslands and their rich biota in the moderately humid Central‐European climate have fascinated scientists for over a century. Modern palaeoecological and phylogeographical data support earlier hypotheses that these grasslands are late Pleistocene relicts and can therefore be considered part of the Eurasian forest–steppe biome. However, it is still unclear which factors fostered the maintenance of steppe patches in Central Europe throughout the Holocene. Here, we provide an overview of the main hypotheses, which stress, respectively, the effects of climate, edaphic conditions and disturbances. We then develop a general conceptual framework on how these three factors interact to form forest–steppe mosaics. We thereby emphasize the role of topography as a crucial control on forest–steppe patterns at the landscape scale. Topography is related to several mechanistic drivers that influence vegetation processes, such as near‐surface microclimate and soil formation. Consequently, topographic variation allows both forest and steppe patches to occur beyond their macroclimatic niche, favouring the development of forest–steppe mosaics. To illustrate our framework, we demonstrate the interactive effect of macroclimate and topography on the occurrence of steppe patches at 108 selected Central European forest–steppe sites. Although we developed our framework focusing on the current distribution of Central European forest–steppe, we suggest that it contributes to the understanding of similar transitions between temperate forest and steppe biomes in the past as well as elsewhere in the world.
... The second PCA axis followed conservatism, rarity and disjunctivity in distribution ranges, independently of habitat preferences, and correlated partially with high final scores of species. Species showing rather wide moisture tolerances that are characteristic of Siberian and Central Asian steppe and forest steppe, disjunctively occurring in Europe as putative full-glacial or later relicts (Holub 1999;Horsák et al. 2015;Roleček et al. 2015), reached extreme scores along both the first and the second axis (Veronica paniculata, Ligularia glauca, L. sibirica, Dendrathema zawadskii, Helictotrichum desertorum, Stipa eriocaulis, Dracocephalum ruyshiana). ...
Article
Glacial relicts have been regionally more common in glacial than in recent times. A rigorous assessment of which species are indeed glacial relicts is extremely difficult because direct evidence is untraceable or equivocal for many species. We aimed to identify species of the Western Carpathian flora (vascular plants, bryophytes and terrestrial lichens) that display apparent biogeographical and ecological symptoms, suggesting a wider regional or supra-regional distribution during glacial times, or at least before the middle-Holocene climate optimum. We worked with the premise that exemplary relict species should tolerate continental and/or arctic climates, should have large distribution ranges with disjunctions, being regionally rare and ecologically conservative nowadays, should be associated with habitats that occurred during glacial times (tundra, steppe, peatland, open coniferous forest) and should display a restriction of ecological niches in the study region. The assessed species were primarily those with boreo-continental or artcic-alpine distribution. We demonstrated a conspicuous gradient of glacial-relict symptoms, with Carex vaginata, Betula nana, Trichophorum pumilum, Nephroma arcticum, Saxifraga hirculus and Cladonia stellaris topping the ranking. Based on the arbitrary ranking, 289 taxa can be considered high-probability relicts. For only a minority of them, there are any phylogeographical and/or palaeoecological data available from the study area. Biogeographical and ecological symptoms of 144 taxa suggest that they retreated rapidly after the Last Glacial Maximum whereas other species probably retreated later. The first principal component of biogeographical symptoms sorted species from circumpolar arctic-alpine species of acidic peatlands and wet tundra to strongly continental species of steppe, steppe-tundra and mineral-rich fens. This differentiation may mirror the altitudinal zonation of glacial vegetation in the Western Carpathians.
... Effects of glacial periods in speciation and geographic expansion in the Iranian plateau is discussed in some taxonomic groups (Djamali et al. 2011, Manafzadeh et al. 2014, Kürschner et al. 2015 and introduced as a reason for disjunct occurrence of some species from main distribution realm (Noroozi et al. 2008, Doostmohammadi & Kilian 2017. Generally, steppe areas in West and Central Asia have been used as recent analogues for paleoreconstructions of full-glacial conditions (Horsák et al. 2010, Horsák et al. 2015, Janská et al. 2017. Like to highland wetlands, scree vegetation of the nival parts of the mountains could be regarded as interglacial refugia for many relict plants (see Kürschner et al. 2015, Kürschner & Djamali 2008). ...
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Trisetum spicatum, a widespread arctic and alpine species, is reported for the first time in the flora of Iran. This species was collected from scree habitats above 4000 m a.s.l. of Mt. Damavand (Central Alborz, Alborz range, N. Iran). This new occurrence of the species in the alpine area of Iran highlights the role of high mountains as migration corridors and refugee during interglacial periods. This new record is different from the other species by its habitat characteristics, compact inflorescence and having dense trichomes on whole parts especially on the peduncle. In addition, ecological and floristic characteristics of the habitat of the species are discussed.
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Biological invasions are common among freshwater molluscs, with the North American planorbid gastropod Gyraulus parvus being reported from Europe (Germany) by the 1970s. It has since spread across Central and Western Europe, mostly living in artificial and highly modified habitats. However, considerable conchological and anatomical similarity exists between it and the native European G. laevis. Using four other European and one North American Gyraulus species as outgroups, separate phylogenetic analyses using mitochondrial and nuclear DNA sequences show that G. parvus and G. laevis are in fact part of the same species-level clade, with the former having nomenclatural priority. However, the structure within the mitochondrial tree suggests a North American origin of the invasive populations. It also makes it possible to track down the distribution of both races. Although native and non-native races in Europe tend to possess some differences in conchology and ecology, the degree of overlap makes it impossible to accurately distinguish between them without the DNA barcode data. Our results change the outlook on the conservation of the rare native race. While interspecific competition among snail species is rare, invasion on an intraspecific level may represent a serious threat for native populations.
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An overview of the diversity of vascular plants in the Czech Republic is presented. This country is situated at the intersection of several assumed important European migration routes. Consequently, the flora is composed of almost all the floristic elements that occur in Central Europe, of which the Central European geoelement is dominant. The occurrence of various sorts of relicts is discussed in the context of the changes in vegetation caused by Pleistocene climatic fluctuations. An account of Czech endemics includes 82 species and subspecies, which is 2.2% of the total vascular plant diversity in this country. Patterns in the distribution and occurrence of endemics in different habitat types are described. Groups of species with similar ecogeographical features within the Czech Republic are distinguished as regional types of distribution. Phytogeographical division of the country is described and the phytogeographical units distinguished are shown on a map.
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Naive use of molecular data may lead to ambiguous conclusions, especially within the context of “cryptic” species. Here, we integrated molecular and morphometric data to evaluate phylogenetic relationships in the widespread terrestrial micro-snail genus, Euconulus. We analyzed mitochondrial (16S + COII) and nuclear (ITS1 + ITS2) sequence across 94 populations from Europe, Asia and North America within the nominate species E. alderi, E. fulvus and E. polygyratus, and used the southeastern USA E. chersinus, E. dentatus, and E. trochulus as comparative outgroups. Phylogeny was reconstructed using four different reconstruction methods to identify robust, well-supported topological features. We then performed discriminant analysis on shell measurements between these genetically-identified species-level clades. These analyses provided evidence for a biologically valid North American “cryptic” species within E. alderi. However, while highly supported polyphyletic structure was also observed within E. fulvus, disagreement in placement of individuals between mtDNA and nDNA clades, lack of morphological differences, and presence of potential hybrids imply that these lineages do not rise to the threshold as biologically valid cryptic species, and rather appear to simply represent a complex of geographically structured populations within a single species. These results caution that entering into a cryptic species hypothesis should not be undertaken lightly, and should be optimally supported along multiple lines of evidence. Generally, post-hoc analyses of macro-scale features should be conducted to attempt identification of previously ignored diagnostic traits. If such traits cannot be found, i.e. in the case of potentially “fully cryptic” species, additional criteria should be met to propound a cryptic species hypothesis, including the agreement in tree topology among both mtDNA and nDNA, and little (or no) evidence of hybridization based on a critical analysis of sequence chromatograms. Even when the above conditions are satisfied, it only implies that the cryptic species hypothesis is plausible, but should optimally be subjected to further careful examination.
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The Czech Republic belongs to the temperate broad-leaved deciduous forest biome for its largest part, but two of its dry lowland areas belong to the forest-steppe biome. Landscapes corresponding to the coniferous forest biome and alpine tundra occur mainly as temperate orobiomes in small areas. Eight altitudinal vegetation belts from lowland to alpine are distinguished. There are high diversities of different vegetation types mainly in deep river valleys in the Bohemian Massif, karst areas, sandstone pseudokarst areas, on solitary volcanic hills, in glacial cirques, lowland riverine landscapes and serpentinite areas. Potential natural vegetation across most of the country is deciduous and mixed forests of beech, oak, hornbeam and noble hardwoods, and coniferous forests of spruce and fir. However, large areas of these forests have been cleared or converted into forest plantations. Open landscape is covered mainly by arable land and perennial grassland. Diversity, ecology, distribution, history and dynamics of the different vegetation types defined in the national vegetation classification are described here in detail.
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Impacts of invasive species on rare species are relevant to conservation. We studied the response of Leedy’s roseroot Rhodiola integrifolia subsp. leedyi, a subspecies listed as threatened under the Endangered Species Act, to removal of the invasive species Japanese knotweed Fallopia japonica. Japanese knotweed has invaded the largest known population of Leedy’s roseroot, affecting about 10% of all Leedy’s roseroot in the world. Japanese knotweed shaded Leedy’s roseroot, but the two did not share belowground resources because of their position on cliffs. To study this interaction and, ultimately, to restore Leedy’s roseroot habitat to an open, high-light condition, we removed Japanese knotweed in a three-treatment block design. We measured Leedy’s roseroot abundance, growth, and reproduction in treatment blocks and in uninvaded areas before and after treatment. Compared with uninvaded areas, Japanese knotweed invasion negatively affected Leedy’s roseroot abundance, growth, and reproduction. Light interception by Japanese knotweed degraded the habitat for Leedy’s roseroot. Herbicide removal of Japanese knotweed resulted in increased light and temperature compared with untreated invaded plots but did not affect Leedy’s roseroot abundance, growth, or reproduction over the 2 y of our study. These results show that invasive species removal is conducive to restoring Leedy’s roseroot habitat, but recovery in the subspecies may lag behind restoration of the habitat, suggesting that additional action or time may be required to restore preinvasion performance of Leedy’s roseroot. Results of this study may inform restoration efforts for other systems and contribute to the literature on interspecific interactions.
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The genus Allium is one of the largest genera in the family of Amaryllidaceae. It comprises over 1000 species, and the number is still increasing. The new intergeneric classification system, based on molecular sequence data, subdivided the genus into 15 subgenera and more than seventy sections. In some sections, the phylogenetic position of species remained unresolved. This is also the case regarding the sections Caespitosoprason and Rhizomatosa. These two sections are confined to Central Asia, where they inhabit steppe habitats. For this study we conducted a phylogenetic analysis of most of the taxa of sections Caespitosoprason and Rhizomatosa with the nuclear marker ITS and three different chloroplast regions (trnQ-rps16, trnL-trnF and rpl32-trnL). Taxonomical remarks, identification key and distribution maps for all species of section Rhizomatosa (including all species of section Caespitosoprason) are presented. A newly discovered Allium bellulum from the Katun Valley (Altai, Russia) was cytologically compared (flow cytometry and karyology) with the other two known localities from Tuva, Russia, and the Buchtarma Valley, Altai, Kazakhstan, and the disjunctive distribution discussed. Divergence time estimates of the species of section Rhizomatosa and their distribution are in agreement with the origin and climate/landscape history of the Central Asian steppe since the Lower Miocene.
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Most studies of mammal extinctions during the Pleistocene–Holocene transition explore the relative effects of climate change vs. human impacts on these extinctions, but the relative importance of the different environmental factors involved remains poorly understood. Moreover, these studies are strongly biased towards megafauna, which may have been more influenced by human hunting than species of small body size. We examined the potential environmental causes of Pleistocene–Holocene mammal extinctions by linking regional environmental characteristics with the regional extinction rates of large and small mammals in 14 Palaearctic regions. We found that regional extinction rates were larger for megafauna, but extinction patterns across regions were similar for both size groups, emphasizing the importance of environmental change as an extinction factor as opposed to hunting. Still, the bias towards megafauna extinctions was larger in Southern Europe and smaller in central Eurasia. The loss of suitable habitats, low macroclimatic heterogeneity within regions and an increase in precipitation were identified as the strongest predictors of regional extinction rates. Suitable habitats for many species of the Last Glacial fauna were grassland and desert, but not tundra or forest. The low-extinction regions identified in central Eurasia are characterized by the continuous presence of grasslands and deserts until the present. In contrast, forest expansion associated with an increase in precipitation and temperature was likely the main factor causing habitat loss in the high-extinction regions. The shift of grassland into tundra also contributed to the loss of suitable habitats in northern Eurasia. Habitat loss was more strongly related to the extinctions of megafauna than of small mammals. Ungulate species with low tolerance to deep snow were more likely to go regionally extinct. Thus, the increase in precipitation at the Pleistocene–Holocene transition may have also directly contributed to the extinctions by creating deep snow cover which decreases forage availability in winter. This article is protected by copyright. All rights reserved.
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In the article we inform about occurrence of the species Triglochin maritima in the Slovakia, and review historical and current locations, which we have confirmed during growing periods in years 2001 – 2003. The species has been confirmed at 18 localities restricted to the north part of Slovakia. We made phytosociological relevés at these sites, which are summarised in the table. Ecological requirements of the species in Slovakia and current status of the localities is also presented.
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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.
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This study presents results of the comprehensive paleontological study of the reference Paleolithic sites of the Altai Mountains. Unique materials of the major problems of the earliest history of the Paleolithic man and natural environment in a key region of northern Asia. The combined study of geoarcheological objects and the natural complex of the Pleistocene allowed the reconstruction of conditions of the development of relief, fauna, and flora, the formation of paleolandscapes and paleoclimates. Physical and radiological methods allowed a reconstruction of the chronological sequence of cultural and paleogeographic events in the Paleolithic of Altai.
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Introduction To include small terrestrial gastropod species (under 5 mm) in faunistic and ecological studies on the malacofauna it is necessary to take samples of the litter layer. The established practise is to take 3–5 litres of litter or foerna. The method how to separate shells from other mineral matter has been well known for many years (e.g. GEYER 1927, LOŽEK 1956). In particular this method is based on the fact that empty shells are filled with air and therefore float on the water surface, whereas other mineral matter sinks down. However, in mires this method is not optimal. Moreover in fens with a higher portion of fine clay particles it is not applicable at all. Of course, the alternative method of direct picking is insufficient in mires, due to the small size of the majority of species (maximum adult size of ca. 10 snail species ranges from 1.6 to 2 mm). Besides that, most of the rare, endangered and indicator species are of smaller size (e.g. Vertigo spp.). In mires, empty shells are filled with water. Therefore, both empty shells and living animals sink down in the course of washing. The present method is based on this fact. Procedure of mire sample reduction After sampling the upper soil layer including litter and herbaceous vegetation in the mires, a first washing is conducted. The sample is washed through a bowl-shaped sieve (mesh size 0.5 mm) to wash out the fine clay particles (otherwise they would cause all the material to stick together after drying). The coarse plant matter is picked out during this step as well. The washing of the sample has to be done separately for small portions of the sample. The removal of coarse plant matter has to be done with extreme care to avoid the loss of molluscs potentially clinging to them. Due time has to be given for the shells or live molluscs to sink down to the bottom of the sieve. Finally, the fine plant matter is washed out in clear standing water. This is done by submerging the sieve with the cleaned material into the water several times. As the fine plant matter sinks down more slowly than the molluscs, we can separate these two fractions by moving the sieve sidewards in the right moment. This final step needs some practise to avoid loosing some of the molluscs in the process. For instance one can produce a weak whirl-pool with the other hand by which the separation of the lighter plant fraction is facilitated. The cleaned sample (Figs 1 and 4) is dried well so that the mollusc bodies are desiccated. In case of samples with a low content of other inorganic particles we can subsequently pick the mollusc shells (Fig. 4). Especially in samples from fens with strong tufa precipitation, a previous separation of tufa and shells comes in useful (cf. Fig. 1 and 2). This is best done by the usual method for normal litter samples: the dry sample is put into a vessel and then water is sharply poured on it; the floating shells are carefully skimmed of the water surface by a sieve. After drying, we may collect the shells by hand-sorting under a binocular microscope (cf. Figs 2 and 3). 11 Conclusions The described method presents a substantial facilitation of the processing of samples from mires (and other types of wetlands with substrates of high water content) compared to the very labour-intensive picking of shells by hand. Thanks to this increased efficiency we can process large samples allowing to study qualitative as well as quantitative aspects of mollusc communities in such habitats. In case of petrifying springs the author was able to reduce 12 litres of sampled substrate to a volume of 1–0.2 litres (depending on the tufa content). More than 3000 mollusc individuals were collected per sample from the petrifying springs of highest species richness and abundance of molluscs. In mires without tufa 12 litre samples were reduced to 2–0.1 decilitre (depending on the calcium content in water, which determines mollusc abundance and species diversity). In malacozoologically poor Sphagnum-fens we often get a pure sample of mollusc shells already after the first washing (see Fig. 4). Another significant advantage of the present method is the possibility to substantially reduce the sample volume already in the field.
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The analyses of low-resolution models simulations of the last glacial maximum (LGM, 21 kyr BP) climate have revealed a large discrepancy between all the models and pollen-based palaeoclimatic reconstructions. In general, the models are too warm relative to the observations, especially in winter, where the difference is of the order of 10°C over western Europe. One of the causes of this discrepancy may be related to the low spatial resolution of these models. To assess the impact of using high-resolution models on simulated climate sensitivity, we use three approaches to obtain high-resolution climate simulations over Europe: first an atmospheric general circulation model (AGCM) with a stretched grid over Europe, second a homogeneous T106 AGCM (high resolution everywhere on the globe) and last a limited area model (LAM) nested in a low-resolution AGCM. With all three methods, we have performed simulations of the European climate for present and LGM conditions, according to the experimental design recommended by the Palaeoclimate Modeling Intercomparison Project (PMIP). Model results have been compared with updated pollen-based palaeoclimatic indicators for temperature and precipitation that were initially developed in PMIP. For each model, a low-resolution global run was also performed. As expected, the low-resolution simulations underestimate the large cooling indicated by pollen data, especially in winter, despite revised slightly warmer reconstructions of the temperatures of the coldest month, and show results in the range of those obtained in PMIP with similar models. The two high-resolution AGCMs do not improve the temperature field and cannot account for the discrepancy between model results and data, especially in winter. However, they are able to reproduce trends in precipitation more closely than their low-resolution counterparts do, but the simulated climates are still not as arid as depicted by the data. Conversely, the LAM temperature results compare well with climate reconstructions in winter but the simulated hydrological cycle is not consistent with the data. Finally, these results are discussed in regard of other possible causes for discrepancies between models and palaeoclimatic reconstructions for the LGM European climate.
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Chapter
Large and small tracts of steppe and tundra-steppe vegetation are scattered widely in the more continental parts of northeastern Siberia. Soils in these steppe and tundra-steppe areas share common features, including base saturation, lime-accumulation horizons, and a humic horizon involving saturation with plant roots rather than accumulation of surficial peat. A study of the character and distribution of these arctic steppe tracts provides insight into the former character of Beringian vegetation during late Pleistocene cold intervals when glaciers were expanded and sea level was at its lowest. Steppe vegetation covers large areas in the Yana and Indigirka drainage basins. In the most arid and continental part of this region, larch forests are restricted to north slopes and especially moist sites with other exposures. Steppe vegetation commonly replaces burned-over larch woodlands, suggesting that climatic conditions may have been more optimal for woodlands at some past time in the Holocene. Steppe patches diminish in size and abundance eastward, but nevertheless, steppe and tundra-steppe vegetation are found on many sunny, well-drained sites in the Kolyma and Anadyr River basins and in an arctic belt extending eastward through the Anyui Mountains and northern Chukotka. The surprising presence of islets of steppe vegetation on favorable sites amidst the hypoarctic tundra of northern Chukotka may reflect the absence of larch and stone pine, which otherwise might be expected to colonize these drier, warmer sites. Several endemic species of very local distribution and several spectacular range disjunctions suggest that the small steppe patches of Chukotka may have expanded recently from even more restricted areas during an earlier, more mesic part of the Holocene. Especially significant are large tracts of tundra-steppe on Wrangel Island, a large island on the continental shelf well north of the northeast Siberian mainland. The interior and the southwestern parts of the island are covered largely by a mosaic of tundra-steppe and cryophytic meadow steppe with low willow thickets in the valleys. A recent pollen spectrum from one of the steppe-tundra tracts resembles the herb zone spectra encountered in pollen sequences that extend back to late Pleistocene levels. The vegetation of Wrangel Island may be the closest living analog to the sort of vegetation that clothed northern Beringia during Pleistocene time (abstract by D. M. Hopkins).
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The malacofauna of 37 topogenous, soligenous and mixed fens in Dovrefjell consists of 18 snail species and three slugs, with 1-10 species per locality and 5-358 individuals per 10 l litter; at most localities single species dominate strongly. Number of species and individuals and the dominance structure of the malacocenoses depend on fen type, litter pH and Ca content, and altitude. Two of the four records of Vertigo parcedentata are new. -Author
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At the time being, the glacial stratigraphy prevails in this region. It is based on the glacial series and on events geologically connected with them. It does n't consider, or only very little, the sequences in the periglacial and their biology. But, these sequences can fill the gaps of the glacial stratigraphy existing since the Elster glaciation. Important sequences are the terrace-travertin sequence of Bilzingsleben, the interglacial-glacial basin sequence of Schöningen, the complex Quaternary from the Geisel valley and the cyclically divisioned basin sequence from the Aschersleben depression. The period of the Elster complex is present with two glacial cycles. The Holstein complex comprises four glacial-interglacial sequences. The first interglacial is apperently the Holstein s.str. The Saale complex follows with three glacial cycles additionnally subdivisioned. Two intra Saalian interglacials could be determined lying between them. The Eem interglacial exhibits two late interglacial oscillations leading into the main cycle of the Weichselian. With Eem and Holocene it comprises twently climatical small cycles with interstadials/intervalls and stadials. Fauna and flora of the interglacials and glacials and of the interstadials/intervalls are described. Thus, the history of fauna and vegetation of the middle and younger Quaternary and the corresponding palaeoecology can be reconstructed. The most important Palaeolithic and palaeoanthropological sites are mentioned.