735592

research-article2017
HOL0010.1177/0959683617735592The HoloceneJuřičková et al.

Research paper

The Holocene

Early postglacial recolonisation, refugial

1­–12
© The Author(s) 2017
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DOI: 10.1177/0959683617735592
https://doi.org/10.1177/0959683617735592
journals.sagepub.com/home/hol
biodiversity hotspot. A case study from
the Malá Fatra mountains, Western
Carpathians, Slovakia

Lucie Juřičková,1,2 Petr Pokorný,2 Jan Hošek,2,3 Jitka Horáčková,1

Jiří Květoň,4 Petra Zahajská,5,6 Anna Jansová1 and Vojen Ložek1,2

Abstract
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.

Keywords
climatic changes, forest history, Gastropoda, Holocene, late Glacial, palaeofaunistics, plant macrofossils, pollen analysis, stable isotopes

Received 6 June 2017; revised manuscript accepted 19 August 2017

Introduction
The late Glacial period and the following first few millennia of
the Holocene witnessed major climatic changes on a global scale,
which have been connected to general trends in biotic develop-
ment in periglacial Central Europe. These regional trends are
relatively well known (e.g. Finsinger et al., 2017; Firbas, 1949,
1952; Giesecke et al., 2011; Ložek, 1964; and many others), at
least in comparison with the rest of the world. In particular, mol-
lusc successions have probably been best documented in the
Czech and Slovak Republics (e.g. Horáčková et al., 2015;
Juřičková et al., 2014b; Ložek, 1964, 1982), though some other
Central European countries have also been relatively well stud-
ied (e.g. Alexandrowicz, 1987; Frank, 2006; Fuhrmann, 1973;
Füköh, 1993; Füköh et al., 1995; Mania, 1972, 1973; Meyrick,
2001; Sümegi, 2005).
However, biotic reactions to climatic changes can differ
locally and may show marked deviations from general trends.
For instance, Ložek (1982) described this phenomenon based on
mollusc successions of 15 Central and Eastern European land-
scape types. More recently, Abraham et al. (2016) and Jam-
richová et al. (2017) confirmed striking local differences in
vegetation development trajectories based on analyses of a large
number of pollen successions from the same area (i.e. currently
Czechia and Slovakia). Based on these studies, it seems plausible
that the general trend of postglacial development consists of
interacting patterns of many specific and partly independent
local histories shaped by different geographical settings con-
nected to glacial refugia and postglacial migration routes.
1Department of Zoology, Faculty of Science, Charles University, Czech
Republic
2Center for Theoretical Study, Charles University, Czech Republic
3Czech Geological Survey, Czech Republic
4Faculty of Science, University of South Bohemia, Czech Republic
5Department of Geology, Faculty of Science, Charles University, Czech
Republic
6Department of Geology, Lund University, Sweden
Corresponding author:
Lucie Juřičková, Department of Zoology, Faculty of Science, Charles
University, Viničná 7, CZ-12844 Prague 2, Czech Republic.
Email: lucie.jurickova@seznam.cz
2 The Holocene 00(0)
Molluscs as a tool for uncovering local biogeographic
histories
We believe that the time is now ripe for puzzling out some of
these local histories, with the aim to elucidate the processes
behind the emergence of general biogeographic patterns. Mollusc
(sub)fossil assemblages currently seem to be the best proxy for
this task. Mollusc shells can be identified to species level, whereas
the most commonly used palaeoecological proxy, pollen, is only
partly identifiable into the comparably exact taxonomic level.
Importantly, shells are usually deposited where the animal actu-
ally lived, unlike pollen or vertebrate bones (e.g. Davies, 2008;
Evans, 1972; Ložek, 1964, 2000; Sümegi, 2005).
Just as biogeography or ecology cannot exist without precise
faunistic and floristic knowledge, palaeofaunistic and palaeoflo-
ristic analyses are fundamental not only for palaeontology but
also for understanding the shaping of recent ecosystems. While
palaeofloristic approaches have dealt with data covering a
majority of Europe (e.g. Brewer et al., 2016), mollusc palaeo-
faunistics have just emerged during the past two decades with
studies of more than one succession per particular landscape
domain (Gedda, 2001; Girod, 2011; Juřičková et al., 2013a,
2013b, 2014c; Limondin-Lozouet et al., 2013; Limondin-
Lozouet and Preece, 2004; Meyrick, 2001). Studies in Central
Europe have repeatedly confirmed Ložek’s (1982) division of
the Holocene landscape development into (1) a zonal part with
fully developed woodland faunas (Juřičková et al., 2014b), (2)
an alpine area (Juřičková et al., 2014c) with reduced forest fau-
nas and late-Holocene human impact and (3) a chernozem low-
land zone where woodlands and their faunas were never fully
established during the Holocene due to middle-Holocene (Neo-
lithic) anthropogenic influences and their late-Holocene accel-
eration (Juřičková et al., 2013a, 2013b; Pokorný et al., 2015).
However, there are still substantial differences between indi-
vidual chernozem lowlands heavily impacted by prehistoric
agriculture (Juřičková et al., 2013a, 2013b; Pokorný et al., 2015)
and little-impacted lowlands (Břízová and Juřičková, 2011),
where by contrast fully developed woodlands occurred. In West-
ern Europe, that is, the edge of the continent with oceanic cli-
matic influences, other conditions were established (Gedda,
2001; Limondin-Lozouet and Preece, 2004; Meyrick, 2001),
and only few canopy forest species occurred during the Holo-
cene climatic optimum (Holocene thermal maximum sensu;
Renssen et al., 2009). In the Mediterranean, progressive desic-
cation was a driving factor of Holocene successions (Girod,
2011; Limondin-Lozouet et al., 2013).
In addition to describing local biotic and landscape develop-
ments, palaeofaunistics and palaeofloristics may contribute to
testing hypotheses of general significance. Even in times of
increasingly sophisticated phylogeographic reconstructions, the
location of glacial refugia, postglacial recolonisation processes
and evaluations of landscape changes caused by various forms
of human impact cannot be studied without precise (sub)fossil
data. For example, while the presence of Central European gla-
cial refugia for temperate species has been frequently discussed
(and both accepted and refused) during last two decades (for the
Western Carpathians, e.g. Jamrichová et al., 2014, 2017;
Jankovská and Pokorný, 2008; Juřičková et al., 2014a; Ložek,
2006; Tzedakis et al., 2013), their location remains unclear,
because they likely formed small patches in the landscape
matrix (and have thus been termed cryptic refugia). Even if we
find some such refugia, the question remains how large they
were, how fast temperate organisms spread from them (if at all)
and to what extent they contributed to colonisation on the (sub)
continental scale. Without a doubt, these questions are essential
for predicting the impact of biotic changes under currently
ongoing climatic changes. Mollusc palaeofaunistics can, there-
fore, provide new independent data for this debate.
The study area
We used a Quaternary mollusc database of Central Europe
(Horáčková et al., 2015) to select a landscape of important bio-
geographical situation and simultaneously covered by a sufficient
network of sampled Holocene mollusc successions. Our previous
palaeofaunistic studies focused on an azonal chernozem area
(Břízová and Juřičková, 2011; Juřičková et al., 2013a; Pokorný
et al., 2015) as well as an azonal mountain area of the Bohemian
Massive (Juřičková et al., 2014c). This study aims to extend this
approach to the key area of the Western Carpathian mountains
where the presence of Last Glacial Maximum (LGM) refugia of
temperate species has repeatedly been hypothesised (Jamrichová
et al., 2014, 2017; Juřičková et al., 2014a; Willis and Van Andel,
2004).
Such a view is strongly supported by current biogeographic
research: At present, the Western Carpathians are an important
interface between biogeographically contrasting regions of
Europe, and the area harbours highly diverse biota that reflects
both climatic and edaphic variability. These characteristics have
been demonstrated to have played a major role in the Quaternary
survival of contrasting biotic elements (Jamrichová et al., 2017;
Mráz and Ronikier, 2016). As the result, the area is currently a
biodiversity hotspot of continental significance and displays a
high degree of endemism. Among vascular plants, the Western
Carpathians are the richest in endemic taxa with the narrowest
distribution ranges (Kliment et al., 2016). High endemism is also
known for many animal groups (e.g. Bálint et al., 2011; Barkasi,
2016). The highest Central European snail endemism has been
found in the Alps as well as Carpathians with the Western Car-
pathians being the northernmost situated area of endemism (Wel-
ter-Schultes, 2012). The survival of some endemic snail species
during the LGM in the Western Carpathians has already been
documented by Ložek (2006) and Juřičková et al. (2014a). How-
ever, endemic species are only rarely encountered in fossil
records, so we have hardly any information about fluctuation of
their ranges.
While studying of species range dynamics in mountain areas,
it is always important to study timberline fluctuations, whose
dynamics markedly differ between Central European Hercynian
mountains (Treml et al., 2006) and the Southern Carpathians
(Feurdean et al., 2016). These fluctuations could have especially
affected the survival of alpine zone species during the Holocene
climatic optimum.
For this study, we selected one of the mountain ranges of the
Western Carpathians – the Malá Fatra mountains – a relatively
temperate area with an especially high degree of endemism, indi-
cating a potential for the presence of a temperate species glacial
refugium. From 24 Western Carpathian endemic snail species, 19
currently live in the Malá Fatra (Welter-Schultes, 2012). Over the
past several decades, we have been able to assemble a complex
fossil record of this area, comprising multi-sited mollusc data,
plus one complex reference record that provides, in addition to
mollusc data, high-resolution vegetation (pollen and plant macro-
fossils) and geochemical (stable isotopes) proxies.
Material and methods
Site characteristics
The six sampled sites are situated in the southern part of the Malá
Fatra mountains (Lúčanská Fatra) in north-western Slovakia (Fig-
ure 1). This mountain chain is built of a mosaic of granite and
Mesozoic carbonaceous rocks (limestones and marlstones,
respectively). At present, the mountain slopes and ridges are cov-
ered by beech forests with an herb-rich undergrowth (Eu–Fage-
nion) and some patches of calciphilous beech woods
(Cephalantero–Fagenion). The vegetation cover is influenced by
the proximity of the warm Upper Nitra and Turiec Basins,
Juřičková et al. 3
Figure 1.  Location of the study area in Central Europe and situation of studied successions in the Malá Fatra mountains (Slovakia). The large
red circle shows the reference site Valča, while smaller black circles show the supporting sites.
as documented, for example, by the common occurrence of the
thermophilous shrub Clematis vitalba in altitudes well above 700
m a.s.l.
The most complex, multi-proxy fossil record used in this study
is from the Valča calcareous tufa deposit, and it was thus used as
a reference site. This site is situated at the bottom of a deep moun-
tain valley, Slovianska dolina, approximately 3.5 km west of the
village Valča (N 49°00′3.8″; E 18°47′44.9″; 550 m a.s.l.). The val-
ley sides at the profile consist of Lower Cretaceous and Jurassic
marlstones and marly limestones forming the bedrock. Higher
upstream, a resurgence with H2S-rich water is located directly in
the stream channel. The spring area of the Sloviansky brook is
situated near the central mountain ridge, which rises here up to
1218 m. The depositional sequence developed in the tufa swamp
at the valley bottom, which was repeatedly dammed by travertine
terraces so that in certain phases, the sedimentation proceeded in
shallow pools where very fine detritus of muddy character, rich in
organic particles, was accumulated. Such sediments alternated
with tufaceous horizons precipitated in running water or marsh-
land vegetation, including a clastic admixture coming from the
slopes built of marlstone bedrock. In light of the position of the
site, the fossil assemblages may also include specimens trans-
ported from up to 3 km away, the length of the upper part of the
Slovianska dolina valley. The malacofauna of tufas in Valča has
been preliminary studied and published (Lisický, 1969; Šilar and
Ložek, 1988). For the lithology of this reference profile, see Sup-
plementary 1 (available online). The lower part of the profile
(below layer 33B) has been excavated and analysed only recently.
This tufa complex is rich not just in mollusc shells but also in
plant macrofossils and pollen – a complex preservation that is
rarely seen elsewhere. Pollen and plant macrofossil data were
obtained just recently.
The Laskár tufaceous footslope accumulation is situated near
the village Valentová on the eastern bank of the middle part of the
Turiec river (N 48°57′43″; E 18°52′37″; 430 m a.s.l.). Alluvium
of this small river is situated in the open agricultural landscape
recently. All layers contain tufa or tufa admixtures together with
malacofauna in all layers, along with vegetation remains in cer-
tain layers. The lowest part of the profile contains limnic sedi-
ments, a fen complex of strata with tufaceous admixture is above
(see Supplementary 2, available online). Preliminary information
about this site has been published before but lacking detailed pro-
cessing and radiocarbon dating (Ložek, 1997). The accumulation
basin of this deposit is relatively high, and fossil assemblages
may include species of flood debris transported from the approxi-
mately 3-km2 area of the Turiec basin.
The Kozol site is a pit on the south footslope of the Kozol hill
(1119 m a.s.l.) situated in the Medzihorská dolina valley (N
49°06′37″; 18°45′10″; 690 m a.s.l.) near the village Rajec. The
vast gravel pit expose footslope sediments from which eastern
part of the pit was sampled (see Supplementary 2, available
online). The area is covered by deciduous forest recently. The
mollusc succession was previously published in a regional collec-
tion (Ložek, 1986). Shells could have been transported from only
a small area of approximately 150 m2.
The Repeš site lies near the village Kl’ačno (N 48°56′15″;
E18°41′50″; 600 m a.s.l.). This tufa accumulation (see Supple-
mentary 2, available online) is situated in the south-oriented allu-
vium of a small brook. The area is covered by deciduous forest
recently. Shells may have been transported from an approximately
0.5-km-long section of the Tmavá dolina valley. The Vyšehradné
site is situated in the Hadvická dolina valley near the village Nitri-
anské Pravno (N 48°53′44″; E18°41′18″; 470 m a.s.l.). This thick
tufa accumulation (see Supplementary 2, available online) is situ-
ated in the south-oriented alluvium of a small brook, and shells
may have been transported from an approximately 0.5-km-long
section of the Hadvická dolina valley. The area is covered by cul-
tural forests and agricultural landscape recently. Both deposits
were sampled in 1961, mentioned in Ložek (1982), but the result-
ing material has been studied only recently.
The last site lies on the rocky top of the Fačkovský Kľak peak
(N 48°58′55″; E18°38’25”; 1300 m a.s.l.). The top of the hill is
covered by oak shrubs and meadows recently, the profile is situ-
ated under the rocky calcareous cliff. Profile was sampled,
4 The Holocene 00(0)
Figure 2.  Comparison of MSS of all six mollusc successions of the Malá Fatra mountains. Ecological groups sensu Juřičková et al. (2014b). On
the left side of spectra are radiocarbon data of particular layers (cal. yr BP; see Table 1). Chronozones sensu (Jäger, 1969) are marked in grey.
processed and published in a local journal (Ložek, 1962; for
lithology, see Supplementary 2, available online). Unfortunately,
this old material was not preserved for later radiocarbon dating.
Thus, the malacospectra from this exposed profile are just sum-
marised here in order to complete the information from the area.
For the lithology of all supporting profiles, see Supplementary 2
(available online).
Sampling and analyses of fossil molluscs
All mollusc successions were sampled by standard methods
(Ložek, 1964) – 8 dm3 of space–discrete samples of the sedi-
ment were taken from the central part of each macroscopically
distinguishable layer, marked in Supplementary 1 (available
online), within 80-cm-wide excavation pits (Repeš and
Fačkovský Kľak) or from naturally eroded exposures, which
were only cleaned. Mollusc shells were extracted from the sedi-
ments by a combination of floating and sieving. After careful
drying, each sample was disaggregated in water and then, if nec-
essary, in hydrogen peroxide. Floating snails were repeatedly
decanted into a 0.5-mm sieve and dried under laboratory condi-
tions. Afterwards, the sediment was dried and sorted by sieving.
Shells were systematically removed from the sediment and
examined under a dissecting binocular microscope. Ecological
groups were used sensu Juřičková et al. (2014b); the nomencla-
ture follows Horsák et al. (2013). Mollusc diagrams (Figure 2)
show the relative proportions of the total number of species
belonging to particular ecological groups (MSS malacospectra)
in separated layers. We use the relative proportion of species
because of the evident local overrepresentation of the spring
prosobranch Bythinella austriaca.
Pollen and plant macrofossil analyses
Besides the palaeomalacological analyses, the reference site
Valča was sampled and analysed for (sub)fossil plant remains.
Plant macrofossils were extracted directly from already processed
mollusc samples (see the processing method above). For pollen
analysis, the profile was newly sampled in the field in intervals
ranging from 10 to 40 cm. A standard acetolysis method (Erdt-
man, 1960) in combination with 10% HCl treatment was used for
preparation of pollen slides. Pollen grains were stained using
0.1% safranin for better identification. During analysis, the mini-
mum terrestrial taxa pollen sum of 500 grains (usually 700, maxi-
mum 800) was reached. Pollen nomenclature follows the
standards of the Czech Quaternary Palynological Database
(PALYCZ; Kuneš et al., 2009; available online at http://botany.
natur.cuni.cz/palycz/).
Radiocarbon dating
Radiocarbon analyses were performed in the Poznań Radiocar-
bon Laboratory, Poland; Izoptech Zrt., Debrecen, Hungary;
and the Center for Applied Isotope Studies of the University of
Georgia, USA. Mollusc shells were measured by the accelera-
tor mass spectrometry (AMS) method and calibrated for vari-
able initial 14C concentration using the OxCal v4.3 calibration
programme (Bronk Ramsey, 2009; see Table 1). Because shells
can potentially contain dead carbon, which can lead to an over-
estimation of their age (Goodfriend and Stipp, 1983), we used
an amalgam of small species shells for radiocarbon dating,
because 78% of these do not contain any dead carbon (Pigati
et al., 2010). The lithology was used as another proxy to con-
trol for the undisturbed development of particular sites. In the
reference site Valča, terrestrial plant macrofossils were also
used for radiocarbon dating in the Poznań Radiocarbon Labo-
ratory, Poland. Results from these confirmed the validity and
precision of radiocarbon measurements from mollusc shell
samples.
Geochemical and stable isotope analyses from the
Valča site
Samples for x-ray fluorescence (XRF) and stable isotope analy-
ses were taken in 5-cm intervals from the cleaned wall. XRF
measurements were performed on approximately 30 g of bulk
sediment using a NITON XL3t 950 GOLDD + (Thermo Scien-
tific) spectrometer with a 50-kV Ag tube and large-area SD
Juřičková et al. 5

Table 1.  Results of AMS radiocarbon age determination. Calibration was performed in the OxCal programme version 4.3 (Bronk Ramsey,
2009).

Sample name and layer Material dated Measured radiocarbon age Age (cal. BP; 95.4% int.) Laboratory code

Laskár 3 Plicuteria lubomirskii shell 3019 ± 31BP 3211 DeA-10021

Laskár 6 Plicuteria lubomirskii shell 4282 ± 33BP 4850 DeA-10022
Laskár 8 shell amalgam 6800 ± 30BP 7636 UGAMS 7671
Laskár 10 Oxyloma elegans shell 8349 ± 40BP 9374 DeA-10023
Laskár 12 Oxyloma elegans shell 8362 ± 36BP 9386 DeA-10024
Laskár 15 Monachoides incarnatus shell 9481 ± 46BP 10828 DeA-10025
Laskár 18 Monachoides incarnatus shell 12400 ± 30BP 14468 UGAMS 7672
Repeš 3 Shell amalgam 3480 ± 30BP 3743 UGAMS 9658
Repeš 9 Shell amalgam 4850 ± 25BP 5567 UGAMS 9659
Valča 3 (190–200 cm) Shell amalgam 4730 ± 25 BP 5456 UGAMS 9650
Valča 10 (475–485 cm) Shell amalgam 5800 ± 25 BP 6589 UGAMS 9651
Valča 18 (685-695 cm) Plant remain (Picea abies needles) 5985 ± 35 BP 6834 Poz-56441
Valča 25 (875–885 cm) Plant remain (Picea abies needles) 7215 ± 35 BP 8060 Poz-56440
Valča 30 (1085–1095 cm) Plant remain (Picea abies needles, Tilia fruit, 7950 ± 40 BP 8816 Poz-56439
Corylus avellana nutshell)
Valča 33B (1295–1305 cm) Plant remain (Picea abies needles, Sambucus 8170 ± 40 BP 9120 Poz-56438
nigra seeds, Alnus glutinosa fruits)
Valča 37 (1425–1435 cm) Plant remain (Corylus avellana nutshells) 8300 ± 60 BP 9298 Poz-63251
Vyšehradné 5 Shell amalgam 2145 ± 30BP 2156 Poz-77698
Vyšehradné 10B Shell amalgam 3580 ± 30BP 3878 UGAMS29906
Vyšehradné 13 Shell amalgam 3780 ± 40BP 4142 UGAMS29907

detector. We included data on the elemental concentrations of
strontium (Sr) and calcium (Ca). A total of 5–8 g of dried bulk
sediment was leached in 30% hydroxide peroxide to remove
any organic carbon in the sample. Stable carbon and oxygen
isotope compositions of carbonate samples were determined
using isotope relative mass spectrometry (IRMS). Finely pow-
dered (milled) samples (200–300 µg) were decomposed with
anhydrous phosphoric acid (~100 µL) in tightly sealed 12-mL
vials made from borosilicate glass, using helium for the atmo-
sphere inside the vials. Evolved carbon dioxide was transferred
through a gas sampling system (GasBench II; Thermo Finni-
gan, Bremen, Germany) into a continuous flow stable isotope
ratio mass spectrometer (Deltaplus XL; Thermo Finnigan) for
analysis. Isotope compositions of the samples were initially
compared with the known composition of a working standard
(carbon dioxide). Final results were then expressed with respect
to the International Atomic Energy Agency (IAEA) standard
Vienna Pee Dee Belemnite (VPDB; IAEA, Vienna, Austria).
Relative isotope abundances δ (δ13C for carbon and δ18O for
oxygen) were calculated according to the formula δ =
(Rsample/Rstandard − 1)1000 (‰), where R stands for the ratios of
isotope amounts 13C/12C or 18O/16O. Standard deviations of both
the isotope abundances δ13C and δ18O, determined using a labo-
ratory standard barium carbonate, were mostly smaller than
0.2‰. We also included results on carbon isotope composition
(δ13) of n-alkanes of the spruce Picea abies, which is usually
used as a proxy indicator of local variability in water availabil-
ity. The n-alkanes in pine waxes were analysed following the
method of Eley et al. (2012, 2014).
Results
Mollusc successions of six profiles in the Malá Fatra were stud-
ied (Figure 2, Supplementary materials 3, 4, 5, and 6, available
online). Two of them, Valča and Laskár, contained malacofauna
of time spans from the late Glacial to the Epiatlantic and the Pre-
Boreal to the Sub-Boreal (sensu Jäger, 1969). The remaining
four profiles contained only middle- to late-Holocene snail
assemblages. Other proxies (pollen, plant macrofossils, oxygen
and carbon stable isotopes) were available only for the Valča
reference site (Figures 3 and 4, Supplementary materials 7, 8,
and 9, available online).
Valča – Reference site of the Malá Fatra
Malacofauna. The mollusc succession of the Valča site can be
divided in two main biostratigraphic zones (Figure 2, Supplemen-
tary materials 3, 4, 5, and 6, available online). The underlying com-
plex of strata (layers 38–26; 1485–925 cm depth) was dominated by
indifferent species (ecogroup C: 7–8) and wetland species (eco-
group D: 9–10) with an admixture of some more tolerant woodland
species in very low numbers. Of particular importance is the high
abundance of Discus ruderatus (9100 cal. yr BP) as well as of the
demanding terrestrial prosobranch Platyla polita (ca. 9300 cal. yr
BP) in association with Aegopinella pura and Clausilia pumila,
which may indicate a favourable woodland environment already in
this early-Holocene (Pre-Boreal) period. This interpretation is also
supported by the higher abundances of the hygrophilous species
Carychium tridentatum and Vitrea crystallina, which occur in humid
woodland leaf litter. However, it is difficult to determine whether
these scattered records indicate a minor climatic oscillation or tem-
porary change in the sedimentary environment. In contrast, the
whole upper group of strata (layers 22–1; 925–0 cm depth) was
characterised by a dramatic increase in species richness, predomi-
nantly due to the expansion of woodland species (ecogroups 1–3)
and partially by inhabitants of canopy forest (ecogroup 1). Their
arrival began, however, already 8100 cal. yr BP (layer 25), with dis-
persed occurrences of several woodland elements in low abun-
dances. The early occurrences of Cochlodina orthostoma,
Isognomostoma isognomostomos and Petasina unidentata (layer
26) indicate a canopy forest still during the late-Holocene. There
was a dramatic boom of silvicolous (canopy forest) species in the
middle-Holocene (Atlantic period), where from almost one layer to
the next, the number of these species doubled (11 woodland species
in layer 23 vs 34 ones in layer 21) and their abundances multiplied
(33 woodland specimens in layer 23 vs 1407 ones in layer 21). Of
particular interest is the late (late Atlantic; Epiatlantic) occurrence of
the dead wood dwelling Carpathian endemic species Argna bielzi on
the westernmost border of its range. The appearance of Discus per-
spectivus indicates warming during the second half of the Atlantic
 6

Figure 3.  Pollen diagram of Valča (Slovakia).

The Holocene 00(0)
Juřičková et al. 7

Figure 4.  Graphs of δ18O (proxy for temperature), δ13C, Sr/Ca (proxy for precipitation amount) and variation of arboreal pollen at the Valča
site plotted by age (ka) and compared with the δ18O record from North Greenland Ice Core Project (NGRIP Members, 2004).

period (6800 cal. yr BP). Wetland species (ecogroup 9) disappeared
during the late Atlantic (Epiatlantic) period (5450 cal. yr BP) and on
the contrary the steppe species Chondrula tridens emerged. Subse-
quently, the rapid sedimentation and formation of tufa ended.
Pollen and plant macrofossil records.  The state of pollen preserva-
tion enabled successful pollen analyses from the lowest
sedimentary layer (1470 cm) to the level of 390 cm. The pollen
diagram (Figure 3) shows a fully developed temperate broadleaf
forest already for the oldest recorded period, between ca. 9300
and 9000 cal. yr BP (from the bottom-up to the 1200 cm depth),
dominated by hazel (Corylus avellana). Despite the elevated posi-
tion of the area (the sampling site is located at 550 m a.s.l., while
surrounding slopes reach much higher altitudes up to 1000 m
8 The Holocene 00(0)
a.s.l.), this forest already had a thermophilous character in this
early period, as indicated by the regular occurrence of Hedera
helix and Viscum pollen grains. Towards the end of this period,
hazel progressively declined and Norwegian spruce (Picea abies)
expanded.
In the oldest phases of the record, pollen grains of beech
(Fagus) already occurred in low quantities. Such an early occur-
rence could either be the result of early immigration from LGM
refugia located in the south, or the result of LGM refugia in the
proximity, as proposed for the area of the Western Carpathians by
Magri et al. (2006).
At the level of 1090 cm (radiocarbon-dated to 8800 cal. yr
BP), maple (Acer) expanded and became another important ele-
ment of regional forests (note that maple is a relatively very low
pollen producer; Broström et al., 2008). Shortly after, other
broadleaf thermophilous tree taxa (mainly Ulmus and Quercus)
reached their absolute maxima. The very high pollen percentages
of spruce (Picea) and alder (Alnus glutinosa-type) can be attrib-
uted to local overrepresentation caused by their presence in the
wetland or around its edges. This interpretation was verified by
the results of the plant macrofossil analysis (Supplementary mate-
rials 7 and 8, available online) that shows abundant findings of
these two taxa. Shortly after 8000 cal. yr BP, the abundance of
silver fir (Abies alba) started to increase.
The pollen record ended at 390 cm, a period roughly dated to
6500 cal. yr BP. Above this level, heavy corrosion of the pollen
grains occurred, and poor preservation makes the pollen record of
younger periods unreliable.
The upland plant taxa macrofossil record (Supplementary 7,
available online) shows the common occurrence of temperate for-
est taxa – either trees or herbs of forest undergrowth – for the entire
profile, further supporting the above interpretation of the pollen
analyses, indicating the prevalence and continuity of the forested
environment since the very start of the record (9300 cal. yr BP) to
its end in the late Atlantic (Epiatlantic) period.
Palaeoclimatic interpretations of the stable isotope record. Since
the variability of δ18O in calcareous tufa is driven mainly by
changes in water body temperature during tufa precipitation, the
δ18O value measured in this sediment can be used as a proxy for
palaeotemperatures (Andrews, 2006). Tufa calcite δ13C reflects
the relative sources of carbon that contribute to the dissolved inor-
ganic carbon of groundwater – low δ13C derived from soil organic
matter and higher δ13C carbon derived from the dissolution of the
aquifer limestone (Andrews et al., 1993).
For a palaeoclimatic interpretation of the δ13C record, we used
the following syllogism: if degassing in aquifer air pockets is
enhanced during decreased recharge when groundwater levels are
low, variation in δ13C may represent an index of recharge and,
therefore, precipitation intensity (Andrews and Brasier, 2005).
Decreased recharge (dry conditions) increases aquifer water resi-
dence time, allowing longer contact and the dissolution of aquifer
limestone increasing δ13C and vice versa. As demonstrated by a
number of previous studies, variation in the Sr/Ca ratio in tufa
deposits is mostly controlled by the same mechanism – a higher
Sr/Ca ratio has been found in calcite precipitated in an aquifer
(during stagnation of groundwater circulation, i.e. dry conditions)
in comparison with calcite precipitated out of the aquifer (Garnett
et al., 2004; Ihlenfeld et al., 2003). A strong positive correlation
detected between δ13C and the Sr/Ca ratio (r2 = 0.7) supports the
idea that δ13C variation in the studied profile represents a proxy
for past precipitation amounts.
The Sr/Ca and δ13C records (Figure 4) showed a general trend
towards decreasing values from the base of the deposit, suggest-
ing decreased aquifer water residence time because of wetter con-
ditions. Several episodes of apparently dry stable conditions
(increased δ13C and Sr/Ca values due to increased aquifer water
residence time) were found within the time intervals ca. 8900–
8700, 8350–8500, 8000–7650, 7000–6650 and 6400–5800 cal. yr
BP. The most significant increase in humidity occurred after ca.
7650 cal. yr BP. These changes are consistent with an increasing
canopy of woodland vegetation, as was indicated by the pollen
and plant macrofossil data (Figure 3, Supplementary materials 7
and 8, available online) and from the distinct increase of number
in the number of forest mollusc species (Figure 2, Supplementary
materials 3, 4, 5, and 6, available online). However, woodlands
provided leaf litter and allowed the development of humus,
increasing root respiration and microbial organic decay, which in
turn increased the generation of isotopically light soil carbon.
Results on the compound-specific isotope analysis of δ13C
from Picea abies sub-fossil needles were used as a proxy indica-
tor of the local variability in water availability. Contrary to the
above-described regional variations in the precipitation amount,
the local variability in wetness showed quite stable conditions
during the whole study period (see Supplementary 5, available
online). This suggests that continuous wetland conditions domi-
nated at the study sites with no significantly drier episodes and
that consequently spruce benefited from the local hydrological
conditions, independent of regional climatic changes.
The δ18O record showed a continuous increasing trend from
the base of the profile, suggesting progressive warming from the
start of the Holocene. This trend was interrupted by several short-
term cool oscillations, of which the most significant occurred
between ca. 8200 and 8000 cal. yr BP. This oscillation can be
correlated with the 8.2 kyr event, although in the Northern Hemi-
sphere, this period is broadly associated with significantly dry
conditions (Alley and Ágústsdóttir, 2005), whereas at our site, a
rather wetter environment prevailed (indicated by increased δ13C
and Sr/Ca during this time period).
Supporting records
The succession at Laskár (Figure 2, Supplementary 3, available
online) extended to the late Glacial, when aquatic and wetland
malacofauna predominated (Valvata piscinalis, Gyraulus crista,
Bathyomphalus contortus, Perforatella bidentata, Vertigo anti-
vertigo, Vertigo geyeri, etc.), but a few forest (Faustina faustina,
Fruticicola fruticum and Clausilia pumila), while open-country
species (Vallonia costata and Vallonia pulchella) indicated a tran-
sitional mosaic landscape character. During the Pre-Boreal period,
the data infer the presence of parkland with wetland, ephemeral
water bodies (Anisus leucostoma and Galba truncatula) and open
country with scattered forest (Discus ruderatus and Monachoides
incarnatus). A boom of silvicolous species was dated to the Atlan-
tic period (7600 cal. yr BP), but there was also forest parkland
with ephemeral water bodies and calcareous seepages. During the
Epiatlantic (4850 cal. yr BP), steppe species (Chondrula tridens)
and even the thermophilous East-European Caucasotachea vin-
dobonensis appeared. From the Sub-Boreal period (3200 cal. yr
BP onwards), the number of forest and wetland species rapidly
declined and open steppe patches prevailed. The West Carpathian
endemic species Plicuteria lubomirskii occurred very early in the
lowlands (from layer 13 Laskár, see Supplementary 2, available
online), but appeared only sub-recently in the mountains
(Fačkovský Kľak hill).
Together with the mollusc succession of the Kozol profile,
the lithology documents the development of slope sedimenta-
tion during the Holocene in this small mountain valley. While
the oldest parts (layers 10–6) showed evidence of gravel sedi-
mentation on the partly open hill slope with typical rock dwell-
ing species (Pyramidula pusilla and Faustina cingulella), from
layer 5 sedimentation ceased and then altered into a flow of
coarse scree developing into a humid talus slope forest with a
fully developed canopy forest fauna (Acicula parcelineata,
Juřičková et al. 9
Cochlodina orthostoma, Daudebardia rufa, Discus perspecti-
vus, Isognomostoma isognomostomos, Ruthenica filograna,
etc.). Some parts of the footslope may have even been water-
logged, as indicated by few Carychium tridentatum and espe-
cially Bythinella austriaca shells. Small open patches, indicated
by few shells of Cochlicopa lubricella and Truncatellina cylin-
drica, may have been the consequence of natural desiccation or
pastures.
Profiles of tufaceous sediments at Repeš and Vyšehradné are
younger. In both profiles, the dominance of woodland species
(ecogroup 1–3) indicated a humid forest during the middle- and
late-Holocene at Repeš and only from the late-Holocene at
Vyšehradné. Ephemeral wetlands and small springs, with a high
abundance of the spring prosobranch Bythinella austriaca,
occurred at both sites. A slowing of sedimentation and the end of
tufa formation in both profiles, together with the appearance of
steppe species (Pupilla muscorum in Repeš and Granaria fru-
mentum in Vyšehradné) during the late-Holocene, may indicate
the influence of pastoral farming penetrating into small mountain
valleys, thanks to a drying out of the landscape.
Discussion
As recently demonstrated by Feurdean et al. (2014) and Jam-
richová et al. (2014, 2017), the Inner Carpathian Basin, including
its northern edge in the territory of the present Slovakia, was
already occupied in the first millennia of the Holocene by well-
developed temperate broadleaf forests. This is probably the
result of the proximity of LGM refugia for temperate forest taxa
(Jankovská and Pokorný, 2008; Juřičková et al., 2014a; Ložek,
2006; Willis and Van Andel, 2004), and this view is further sup-
ported by our study that shows the dominance of a forested envi-
ronment at the Valča site since the very start of the record (9300
cal. yr BP). This finding also supports the hypothesis (Jam-
richová et al., 2017; Willis and Van Andel, 2004) that the early
establishment of temperate forests is likely a key factor for
understanding the current biogeography of Eastern–Central
Europe. This early occurrence of temperate trees is unlikely to be
explained only by gradual postglacial migrations from Southern
Europe. Humid but relatively warm mountains might hence have
acted as glacial refugia of temperate forest species, while adja-
cent lowlands and leeward basins might have rather acted as
postglacial refugia for steppe grasslands (as shown in recent
study by Jamrichová et al., 2017). Unfortunately, the existence of
LGM refugia for temperate forest taxa cannot be directly demon-
strated by this study, due to the lack of a sufficiently old fossil
record (i.e. going back to the LGM).
Postglacial mollusc succession and climatic
development of the Malá Fatra
In general, the mollusc succession of the Malá Fatra corresponds
to the standard pattern of development in mid-European upland
areas (e.g. Juřičková et al., 2014b; Ložek, 1964) but with some
biogeographically related peculiarities. The late Glacial (starting
at 14,500 cal. yr BP) occurrence of some forest species (Clausilia
pumila, Faustina faustina and Fruticicola fruticum; for detail, see
Supplementary 3, available online) provides early evidence of
scattered woodlands in the Malá Fatra and again the likelihood of
nearby or local glacial refugia for these species. In the Pre-Boreal,
other woodland species completed the mollusc assemblages. Spe-
cies such as Acanthinula aculeata, Aegopinella pura, Columella
edentula, Discus ruderatus, Platyla polita, Vertigo pusilla and
Monachoides incarnatus indicate the occurrence of a canopy for-
est in lowlands and mountain valleys in this transitional period.
Steppe and open-country patches were characteristic of lowlands
in the Turiec Basin throughout the whole Holocene, likely owing
to the impact of human settlements (Veliačik, 1989; Žaár, 2015). In
mountain valleys, the rare records of open-country species may
likely results shell transport from nearby higher situated exposed
rocky habitats. Open patches in the Slovianska dolina valley
(Valča site), which are directly associated with human impacts in
the Turiec Basin, appeared only later (5450 cal. yr BP), as indi-
cated by the steppe snail species Chondrula tridens (and supported
by the occasional macrofossil find of cereal remains in the same
period – charred chaff of Triticum monococcum; Supplementary 7,
available online). The boom of silvicolous species was dated to
7600 cal. yr BP in the lowland (Laskár) and 7400 cal. yr BP in the
mountain valley (Valča) and can be associated with the mid-Holo-
cene humidity pulse, visible in many palaeoenvironmental records
throughout Central Europe (see Roberts, 2014). At the Valča site,
this supraregional trend is reflected in the distinct increase of δ18O
values (proxy for temperature rise) accompanied by the decrease
of δ13C values and the Sr/Ca ratio (a proxy for precipitation
decrease) that occurred here after 8000 cal. yr BP (Figure 4). The
predominantly forested environment and dominance of thermoph-
ilous broadleaf trees at Valča are also supported by the pollen and
plant macrofossil analyses, as well as by the presence of the high-
est known canopy forest mollusc diversity in the entire European
Holocene record, at least to our knowledge. The desiccation (as
demonstrated locally by the geochemical data, see Figure 4)
started immediately after the silvicolae boom in the lowland site
but culminated there around 4850 cal. yr BP. While almost all pro-
files contain tufa sediments, desiccation resulted in a slowing rate
of sedimentation and an end of tufa formation in the Sub-Boreal
period. This desiccation is also apparent from the declines in large
populations of the spring species Bythinella austriaca. During this
period, the Malá Fatra consisted of a mosaic of woodland, open
(mostly anthropogenic) and wetland habitats.
Holocene treeline shifts
The survival of a forest-free alpine belt in Central European
mountains is the subject of current discussion, and the likelihood
differs substantially for particular mountains (e.g. Treml et al.,
2006). Sometime around the end of the late Atlantic (Epiatlantic)
period, peaks of the Malá Fatra were probably completely cov-
ered by forest, as indicated by the fully developed woodland com-
munities on the Fačkovský Kľak peak (1300 m a.s.l.; Ložek,
1962). Steppe and shrubby beech habitats occurred there recently,
being approximately 100 m under the recent treeline in the West-
ern Carpathians (approximately 1400 m a.s.l.; Kozak et al., 2007).
Unfortunately, material from the Fačkovský Kľak profile was not
preserved since its extraction, so radiocarbon dating of the highest
altitude of the timberline is not possible. Nevertheless, the precise
dating of the Holocene treeline shifts of this Carpathian area
needs to be critically revised. Ložek (1981) attempted to resolve
this problem using three successions in the nearby Kriváňská
Fatra mountains but lacked radiocarbon dating. A succession in
the profile from the Rozsutec peak (1500 m a.s.l.) in this area was
considered to represent the whole Holocene (Ložek, 1978, 1981),
but we have dated the base of this profile as being much younger
(3700 cal. yr BP; unpublished data), so it cannot document
treeline shifts during the critical period of the Holocene, that is,
before grazing and other possible human influences started.
The zoogeographic border between the Bohemian
Massive and the Western Carpathians
While the geological border between the Carpathians and the Bohe-
mian Massive is quite clear, the westernmost borders of Carpathian
species ranges and the easternmost borders of some Central Euro-
pean snail species ranges differ and shifted during the Holocene.
Interestingly, while the Malá Fatra is not the westernmost mountain
10 The Holocene 00(0)
belt of the Carpathians, many West Carpathian endemics have their
recent westernmost border there (Argna bielzi, Chondrina tatrica,
Cochlodina cerata and Faustina cingulella – the last two also in the
nearby Súľovské vrchy mountains). From 24 Western Carpathian
endemic snail species, 19 currently live in the Malá Fatra (Welter-
Schultes, 2012). In addition, the Malá Fatra represents the eastern-
most border of Cepaea hortensis. Together these species ranges
provide evidence of the important zoogeographic position of this
mountain belt for molluscs. However, some Central European spe-
cies such as Alinda biplicata, Semilimax semilimax and Urticicola
umbrosus that occur farther to the east are locally lacking in the
Malá Fatra mollusc successions. This may help us evaluate the
shifting of this border during postglacial successions. Of particular
interest is the appearance of Balea perversa during the late Atlantic
(Epiatlantic) in Vyšehradné, which is the second-known Holocene
record of this species from Slovakia (according to Czech and Slo-
vak Holocene mollusc database – unpublished data). A more recent
occurrence of Balea perversa is not known from the Malá Fatra.
The only known Holocene record of the Central–Western European
species Helicodonta obvoluta in the Malá Fatra also comes from
the Epiatlantic section of this profile, but this species has survived
there to the present. One of the oldest records of the Western Car-
pathian endemic subterraneous species Alzoniella slovenica (6800
cal. yr BP) was obtained from the Valča section. Another Western
Carpathian endemic species, Faustina rossmaessleri, is recently
most abundant just in the Malá Fatra. This species has a docu-
mented LGM occurrence at the Farkašovo site approximately 70
km to the east (Juřičková et al., 2014a; Ložek, 2006), and thus, this
part of the Western Carpathians seems to be its glacial refugium.
Conclusion
Hardly anyone doubts the role of the Western Carpathians as a
glacial macro-refugium (Jamrichová et al., 2014, 2017; Mráz and
Ronikier, 2016; Wielstra et al., 2015), being documented as the
easternmost border of various studied species or genetic lineages
being traced to the area of the Western Carpathians (Magri et al.,
2006; Pinceel et al., 2005; Wielstra et al., 2015). In concordance
with this evidence, and despite the pronounced early- to middle-
Holocene climatic changes (in both temperature and precipita-
tion), our data suggest that the Malá Fatra mountains could have
been a glacial (LGM) refugium for several Western Carpathian
endemic snail species. However, the precise location of these
hypothetical LGM refugia remains a challenge. In any case, in
this study, the Western Carpathians were demonstrated to have
undergone substantial late Glacial and early-Holocene afforesta-
tion. Therefore, the area could have served as a significant step-
ping stone for the postglacial spread of closed-canopy temperate
forest biota to the rest of Central Europe (at the least). The bio-
logical diversity within this type of biome has currently reached
locally maximum values, clearly the result of long-term regional
continuity of the forested environment.
Acknowledgements
We appreciate Nicole Limondin-Lozouet and an anonymous re-
viewer for their valuable comments on the previous version of the
manuscript and David Hardekopf for the English revision. Petr
Vorm is acknowledged for performing the XRF analysis.
Funding
This work was supported by the Czech Science Foundation
(GAČR) grant 13-08169S and 17-05696S, and the HACIER proj-
ect funded from the Norwegian Financial Mechanism 2009–2014,
and the Ministry of Education, Youth and Sports under project
contract no. MSMT-28477/2014. Access to instruments and other
facilities was supported by the Czech research infrastructure for
systems biology C4SYS (project no. LM2015055).
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