Lucie Juřičková

Mid-Holocene bottleneck for central European dry grasslands: Did steppe survive the forest optimum in northern Bohemia, Czech Republic?

Revisiting the classical Gradmann’s ‘steppe theory’ for central Europe, we examine whether the early Holocene steppe habitats survived the critical period of maximum Holocene afforestation: the mid-Holocene bottleneck. Despite the undisputable fact that afforestation was a dominant ecological factor in this period, our parallel analyses of pollen and molluscs from sedimentary sequences discovered in the dry lowland area of northern Bohemia, Czech Republic (Zahájí and Suchý potok sites, lower Ohře area) provide strong evidence for uninterrupted local occurrence of steppe grasslands throughout the Holocene. At the onset of the Neolithic agriculture, this area was covered by forest-steppe. Analogously to the present forest-steppe landscapes of eastern Europe and south-western Siberia, dry areas of northern Bohemia were dominated by open-canopy pine–birch forests that enabled continuous survival of many light-demanding plant species from the late Glacial and early Holocene to the Neolithic. Later on, anthropogenic deforestation and livestock grazing created a semi-natural steppe. Our data suggest that this secondary steppe can be viewed as a direct continuation of the late Pleistocene and early Holocene natural steppe rather than a purely cultural steppe developed only after deforestation of a continuously forested mid-Holocene landscape by humans. At the same time, we provide evidence supporting Gradmann’s ‘steppe theory’, assuming that in central Europe, Neolithic farming started in those areas that were not completely forested but contained remnants of natural steppes. This finding has important implications for the interpretation of present biodiversity patterns in central Europe.

Diversity and Biotic Homogenization of Urban Land-Snail Faunas in Relation to Habitat Types and Macroclimate in 32 Central European Cities

The effects of non-native species invasions on community diversity and biotic homogenization have been described for various taxa in urban environments, but not for land snails. Here we relate the diversity of native and non-native land-snail urban faunas to urban habitat types and macroclimate, and analyse homogenization effects of non-native species across cities and within the main urban habitat types. Land-snail species were recorded in seven 1-ha plots in 32 cities of ten countries of Central Europe and Benelux (224 plots in total). Each plot represented one urban habitat type characterized by different management and a specific disturbance regime. For each plot, we obtained January, July and mean annual temperature and annual precipitation. Snail species were classified into either native or non-native. The effects of habitat type and macroclimate on the number of native and non-native species were analysed using generalized estimating equations; the homogenization effect of non-native species based on the Jaccard similarity index and homogenization index. We recorded 67 native and 20 non-native species. Besides being more numerous, native species also had much higher beta diversity than non-natives. There were significant differences between the studied habitat types in the numbers of native and non-native species, both of which decreased from less to heavily urbanized habitats. Macroclimate was more important for the number of non-native than native species; however in both cases the effect of climate on diversity was overridden by the effect of urban habitat type. This is the first study on urban land snails documenting that non-native land-snail species significantly contribute to homogenization among whole cities, but both the homogenization and diversification effects occur when individual habitat types are compared among cities. This indicates that the spread of non-native snail species may cause biotic homogenization, but it depends on scale and habitat type.

Mollusc succession of a prehistoric settlement area during the Holocene: A case study of the České středohoří Mountains (Czech Republic)

Some Central European areas were attractive for the first agricultural settlements due to their suitable natural conditions. The Holocene development of such areas was thus under long-term human pressure, whose impact on the whole landscape is still poorly understood. One of such areas is the České středohoří Mountains. While pollen analyses can provide the general pattern of the landscape development, the analyses of mollusc succession provide landscape details, which are important primarily in landscapes with high habitat diversity. Based on the study of 11 mollusc successions situated at the České středohoří Mountains, we describe the postglacial development of the area and show the moderate fluctuation of woodland, wetland and open country habitats without any distinct succession peaks of particular habitat types during the whole Holocene. However, a detailed look of species exchange has provided additional information of succession pattern. The impoverishment of woodland communities is probably caused by human pressure, not natural processes, because fully developed woodland assemblages had occurred there during the past interglacials. It seems for now that humans had affected the whole landscape of the prehistoric settlement area, including hard-to-access sites, not only the nearby surroundings of their settlement.

Molecular phylogeny of the genus Helix (Pulmonata: Helicidae)

The vast diversity of land snail forms is insufficiently understood even in seemingly familiar taxa. This holds for Helix Linnaeus, 1758, a genus with several common edible species which comprises the largest Western Palearctic snails. The taxonomy of this genus, which has a centre of diversity in the eastern Mediterranean, has recently undergone significant changes, in both the delimitation of the genus itself and its species‐level systematics. Here, we compare the lineage diversity of Helix, as revealed by two mitochondrial markers, with the conclusions of the recently published morphology‐based taxonomic revision. For the molecular analysis, we assembled a representative data set covering almost all species of the genus as recognized by the mentioned revision. We obtained sequences not only from fresh and preserved soft tissues but also from dried tissue remains (some of them decades old) from shell collections. Our results show that the genus Helix, in the narrow sense proposed by recent studies, is paraphyletic because the genus Tacheopsis was unambiguously revealed as one of the tip branches of Helix. The monophyly of several species, as presently recognized, was not supported; partly, this may be attributed to a lineage diversity overlooked so far. This holds also for the type species of the genus, H. pomatia, which comprises at least one additional lineage. Greece, the Aegean and western Turkey is the core area for the diversity of Helix and its relatives, and the region is probably a major long‐term refuge for large Helicidae. The highest species diversity is found along the Alpide belt from the western Balkans to southern Turkey. The diversity of Helix in Europe, north of Greece and the Apennines, is a result of a single European radiation. Our data also suggest that past human activities likely influenced the present‐day distributions of some species.

Invasiveness Does Not Predict Impact: Response of Native Land Snail Communities to Plant Invasions in Riparian Habitats

Studies of plant invasions rarely address impacts on molluscs. By comparing pairs of invaded and corresponding uninvaded plots in 96 sites in floodplain forests, we examined effects of four invasive alien plants (Impatiens glandulifera, Fallopia japonica, F. sachalinensis, and F.×bohemica) in the Czech Republic on communities of land snails. The richness and abundance of living land snail species were recorded separately for all species, rare species listed on the national Red List, and small species with shell size below 5 mm. The significant impacts ranged from 16–48% reduction in snail species numbers, and 29–90% reduction in abundance. Small species were especially prone to reduction in species richness by all four invasive plant taxa. Rare snails were also negatively impacted by all plant invaders, both in terms of species richness or abundance. Overall, the impacts on snails were invader-specific, differing among plant taxa. The strong effect of I. glandulifera could be related to the post-invasion decrease in abundance of tall nitrophilous native plant species that are a nutrient-rich food source for snails in riparian habitats. Fallopia sachalinensis had the strongest negative impact of the three knotweeds, which reflects differences in their canopy structure, microhabitat humidity and litter decomposition. The ranking of Fallopia taxa according to the strength of impacts on snail communities differs from ranking by their invasiveness, known from previous studies. This indicates that invasiveness does not simply translate to impacts of invasion and needs to be borne in mind by conservation and management authorities.

Field and laboratory studies on the life-cycle, growth and feeding preference in the hairy snail Trochulus hispidus (L., 1758) (Gastropoda: Pulmonata: Hygromiidae)

For the first time the life cycle of the common land snail Trochulus hispidus was completely described in Central Europe (Poland). This is a semelparous species predominantly with an annual life cycle and the reproductive period lasting from April till October. The first young snails hatch in spring, grow rapidly in summer and reach ca. 4 whorls until winter. In spring of the next year they mature and reproduce. After that they die. There is hardly any growth from late autumn till early spring. The average proportional growth rate is ca. 0.3 whorl/month in the wild. The fastest growth is present in the youngest snails and then gradually decreases over the course of their age. Laboratory and field observations allowed for establishing the following life cycle parameters: eggs calcified, almost spherical, ca. 1.5 mm, laid in spring and summer in batches of between 1 and 47. Time to hatching is 6–24 days, hatching is asynchronous; newly-hatched snails have approximately 1.5 whorls. Analysis of food preferences revealed, that T. hispidus tends to restrict its diet during the life. Generally the youngest snails equally consumed leaves of all four tree species offered (Fraxinus excelsior, Acer pseudoplatanus, Tilia cordata and A. platanoides) whereas adults preferred F. excelsior over A. pseudoplatanus and A. platanoides.

River Floodplains as Habitat and Bio-Corridors for Distribution of Land Snails: Their Past and Present

Abstract River floodplains of Czech rivers serve as refugia to woodland or hydrophilous gastropods, in current intensively agriculturally utilised, urbanised and largely fragmented landscape. This habitat often form one of the last refuge and replace the natural habitat of these species. River floodplains also represent linear bio-corridors in landscape and allow gastropods to spread through the landscape in both directions, up and down the stream. We showed based on available fossil mollusc successions that development of the floodplain mollusc fauna took place quite different way in various river floodplains, depending on their specifics and geographical location, because especially the ones situated in the chernozem area of the Czech Republic had very different history in comparison with those in higher altitudes. The species richness and composition of recent floodplain malacofauna arises from historical development of particular area/site and depends also on environmental factors such as an elevation, humidity gradient, vegetation type and its biomass, light conditions of the site and soil reaction. Recently, the invasive plants represent a serious problem for current floodplain ecosystems; species richness and abundances of terrestrial mollusc floodplain assemblages are changing due to their effect. The impact on gastropods is species-specific and was described for the following species: Impatiens glandulifera, Fallopia japonica subsp. japonica, F. sachalinensis, F. ×bohemica.

The spread of non-native Cepaea nemoralis and Monacha cartusiana (Gastropoda: Pulmonata) in the Czech Republic with comments on other land snail immigrants

The aim of our study is to describe and visualise the spread of two non-indigenous land snail species Cepaea nemoralis and Monacha cartusiana in the Czech Republic during more than 100 years period. Several factors play an important role in changes of the distribution of these species: ecological (climate change), ethological (passive dispersal potencial) and economic (increasing traffic as a vector of spreading). The spreading of M. cartusiana has a rapidly increasing trend. More than half sites in the Czech Republic were colonised by this species in 2000–2010. While the spread of C. nemoralis has been continuous during the last century, the rapid range extension was recorded in the last two decades.

Early postglacial recolonisation, refugial dynamics and the origin of a major biodiversity hotspot. A case study from the Malá Fatra mountains, Western Carpathians, Slovakia

While general trends in Central European postglacial recolonisation dynamics are relatively well known, we often lack studies on intermediate (meta-population, landscape) scales. Such studies are needed to increase our understanding of, for example, the location of refugia; emergence of endemism, rates and trajectories of postglacial migrations; and anthropogenic landscape changes. Here, we focused on the outer Western Carpathian mountain chain Malá Fatra, which is currently characterised by high biodiversity and endemism and is thus considered a likely refugium of the Last Glacial period for the temperate biota of Eastern–Central Europe. We used molluscs and vascular plants as reference taxonomic groups and supported palaeoenvironmental interpretations of their (sub)fossil assemblages using high-resolution geochemical data. Generally, postglacial biotic successions from the study region fit the standard developmental pattern well in Middle and Eastern European uplands. Nevertheless, we found important biogeographically based peculiarities. In total, more than 50 species per (sub)fossil community at the reference site Valča, including 30 woodland species and 11 Carpathian endemites, make site of the highest known Holocene mollusc species diversity in Europe. Our palaeoecological analysis of this long-term biodiversity hotspot suggests that the Western Carpathians were likely an important source of the postglacial recolonisation of Central Europe by forest biota and, at the same time, an area of refugium-based endemism.

Pupilla alluvionica Meng & Hoffmann, 2008: A Land Snail Extant in the Altai Refugium Recognised for the First Time in Central European Early-Middle Pleistocene Glacials

ABSTRACT Pupilla alluvionica Meng & Hoffmann, 2008 is an extant land snail species known until now only from a few sites in the Russian Altai Mountains. We have now identified it in fossil loess assemblages of Early-Middle Pleistocene age at three Central European sites. Fossil materials match those of Altaian populations, being only slightly smaller in mean shell width. In addition to the details of the locations and biometrics of these fossil finds, we list the faunas associated with P. alluvionica in the deposits. These are compared with the ecological conditions and associated faunas of living populations from the Altai. Pupilla alluvionica is a typical inhabitant of xeric habitats: steppes with rocky limestone outcrops. This matches the known habitat preferences of associated species in the fossil deposits that are typical of open loess steppe.