RUINS OF MEDIEVAL CASTLES AS REFUGES FOR
ENDANGERED SPECIES OF MOLLUSCS
LUCIE JUŘIČKOVÁ 1 AND TOMÁŠ KUČERA 2
1
Department of Zoology, Charles University, Vinicˇná 7, CZ-128 44 Praha 2, Czech Republic; and
Institute of Landscape Ecology, Academy of Sciences of the Czech Republic, Na Sádkach 7, 370 05 Cˇeske´ Budeˇjovice, Czech Republic
2
(Received 20 July 2004; accepted 16 December 2004)
ABSTRACT
Molluscan communities were studied at 114 castles in the Czech Republic. Altogether 70% of the land
fauna species of the Czech Republic were found at these sites. A range of environmental factors were
recorded at each, and analysed for their influence on species diversity. The variability of the molluscan
communities was assessed. It was shown that castles act as islands (positive effect of castle size on species
number) and that a high degree of castle disintegration negatively affected species diversity. Limestone
as a rock substrate was found to be a very important factor, while the effect of calcium in tree litter was
unimportant. Species indicative of some environmental variables were identified. A complete literature
review on castle-related molluscan fauna was performed for the Czech Republic. Castles present a
characteristic set of habitats markedly differing from the surrounding landscape. Thus, they represent
significant refuges for many species, as indicated by the high species diversity and the presence of rare
and geographically isolated species.
INTRODUCTION
The role of man as a geomorphic agent, changing the landscape
since the Neolithic, has become increasingly accentuated in the
last few years (e.g. Larson, Matthes & Kelly, 2000; Hooke,
2000). In early times, human activities entailed only land deforestation that, in turn, triggered geomorphic processes. Beginning
in the Middle Ages, however, man has become a geomorphic
agent himself, actively removing soil and rock and establishing
completely new structures in the landscape. This fragmentation
of the original, largely monotonous, forest landscape of Central
Europe produced open habitats at lower altitudes, which are
not the natural climax kind of habitat (e.g. Ložek, 1964).
During this process, natural ecosystems were turned into islets
or palaeorefugia (Nekola, 1999). This is generally considered
to be one of the causes of landscape degradation and species
diversity reduction (e.g. Haila, 2002). This study attempts to
show that this should not be a general rule.
Starting in the high Middle Ages, castles were built on many
hilltops, elevations or rocky crests in Central Europe. Castles
were the first human constructions made from stone in Middle
Europe, which markedly affected the character of this landscape.
They were built usually in areas with sparse settlements, in highlands and uplands. Now, some of the castles are conserved as
historical monuments. Others are slowly deteriorating, becoming
an integral part of the ambient nature. In the latter case, decaying walls simulate natural eutrophic talus and cliffs. Castles
became new modern anthropogenic neorefugia (according to
the system of biological refugia; Nekola, 1999). The specific
environments generated, full of remarkable plants and animals
often bound to these open habitats, resulted from an ecological
‘experiment’ some seven centuries long. Moreover, these sites
have so-called historical memory, which means that the historical information on these sites is more readily available than that
on nearby hills and elevations. This information usually suggests
that elevations with castle ruins were totally deforested in the
Middle Ages, thus acting as exceptional refuges for heliophilic
organisms bound to natural non-forested areas. Once they had
Correspondence: L. Juřičková; e-mail: [email protected]
been abandoned (they lost their military function in the
15th century), the castles were overgrown by woody plants
and retained their varied vegetational patterns for a long time
(mosaics of shrubs and young forest, rocky outcrops and
boulders resulting from disintegration).
Although castle ruins have always attracted the attention of
scientists, few studies have been published on this unique type
of environment (Brandes, 1996; Ložek & Skalický, 1983).
Papers on molluscs from castle ruins include lists of species
from one or only a few ruins, or deal with different types of
communities at a single castle in detail (Alexandrowicz, 1988;
Berger, 1961; Haldeman, 1990; Jauernig, 1995; Matzke, 1985;
Munzig, 1977; Pokryszko, 1984; Rouse & Evans, 1994; Urbanski,
1973, Wiktor, 1959; Zeissler, 1968, 1975, 1980, 2000; Šteffek,
1986, 1992). The first and, currently, the most extensive study
(Alexandrowicz, 1995) described communities at 20 castles in
the Polish Carpathians. All of these papers clearly show that
castles host communities of high species diversity, which is a
result of the varied mosaics of different types of habitats concentrated on relatively small areas.
The aims of the present study were: (1) to determine the
species diversity for individual zoogeographic groups and the
proportion of species with isolated occurrences at castles
outside of the main area of their distribution, i.e. to document
the real significance of castles as anthropogenic-based neorefuges
(sensu Nekola, 1999) in the cultural landscape; (2) to establish
which environmental variable has the biggest influence on
species diversity and the variability of castle communities;
(3) to test the beneficial effect of limestone substrate on the diversity of castle communities, as generally presumed from nonquantitative evidence (e.g. Ložek, 1956); (4) to test the beneficial
effect of calcium in tree litter on molluscan diversity at the castles
(e.g. Wäreborn, 1969, 1970, 1992; Waldén, 1981); (5) to determine the proportion of variation in a species set explained by
individual environmental factors and look for index species;
(6) to find which species prefer the anthropic environments of
the castles to their natural habitats; and (7) to evaluate the
trends in the evolution of castle communities based on an extensive literature review covering more than 100 years of recording
the occurrence of molluscs at castles in the Czech Republic.
Journal of Molluscan Studies (2005) 71: 233– 246. Advance Access Publication: 28 June 2005
The Author 2005. Published by Oxford University Studies on behalf of The Malacological Society of London, all rights reserved.
#
doi:10.1093/mollus/eyi031
L. JUŘIČKOVÁ & T. KUČERA
terrain, 4. Castles in lowlands – so-called water castles;
Century of destruction (cen-des): 1. Preserved castle,
2. Destroyed in 18th–19th century, 3. Destroyed in 16th–
17th century, 4. Destroyed in 14th– 15th century; Stage of
destruction (st -des): 1. Preserved castle, 2. Ruin of the whole
castle (high walls and towers, preserved interior spaces),
3. Ruin partly disintegrated, only parts of walls without
interior space, 4. Remains of the ruin integrated into nature;
Attendance (attend): 1. Very high, area heavily trampled,
2. High – ruderal habitats, 3. Moderate – a mosaic of
ruderal and natural habitats, 4. Low – natural habitats;
Habitat types (habit): 1. Open habitats, 2. Shrubs, 3. Forest.
(3) Categorical: Surrounding vegetation: 1. Spruce (Picea abies )
2. Pine (Pinus sylvestris ), 3. Larch (Larix decidua ), 4. Beech
(Fagus sylvatica ), 5. Oak (Quercus petraea, Q. robur ), 6. Hornbeam
(Carpinus betulus ), 7. Sycamore maple (Acer pseudoplatanus ),
8. Norway maple (Acer platanoides ), 9. Ash (Fraxinus excelsior ),
10. Lime (Tilia cordata ); Phytogeographic zones (zone):
oreophyticum – mountain vegetation (oreo), mesophyticum–
Mid-European zonal vegetation (meso), thermophyticum –
thermophilous vegetation (termo); Geological background:
10 categories were chosen and arranged according to
suitability for molluscs (from extremely suitable to extremely unsuitable) 1. Limestone (limeston), 2. Calcareous
rocks (marlstones, marls, calcareous sandstones etc.),
3. Basalts, 4. Basic crystalline rocks (gabbro, amphibolite,
calc-silicate rock), 5. Trachytic rocks, 6. Shales, 7. Sandstones, 8. Cambrian conglomerates, quartzites, silicites,
9. Granites, 10. Acidic crystalline rocks (aci-cry).
METHODS
The data used in this paper come from a representative set of
castles and castle ruins in the Czech Republic. The castles
studied are characterized by different altitudes, centuries of
destruction and stages of destruction, isolation, rock substrates,
vegetation and numbers of visitors.
Data were gathered on molluscan fauna from studies done
since the latter half of the 19th century to the present. Data
from only 16 castles, where molluscan communities were
studied using standard quantitative methods, were selected
and included in subsequent statistical analyses (Hlaváč, 1998a,
b, 2001, unpublished collections; Horsák, unpublished collections; Ložek, 1994a; Pfleger, 1997).
The first author of this paper then collected and processed
material from a further 98 castles (Fig. 1) using hand sampling
and acquisition of standard litter samples (e.g. Ložek, 1956).
The combined samples of litter and topsoil were always collected
from the whole area of each castle for a total volume of ca 7 l. The
samples were dried, washed and the organic material sorted,
after repeated drying, to separate size categories. Molluscs
were then collected by sorting under a binocular microscope.
Statistical processing involved only snails obtained from sieved
litter samples; slugs obtained by hand sampling only were
excluded from the analysis, but not from the faunal list. The
classification is according to Turner et al. (1998).
Three types of environmental factors were determined:
(1) Numerical: number of species (data were square-root transformed; Jongman, ter Braak & van Tongeren, 1995), and
altitude (alt) (m). Data were divided by 100 to obtain identical
units of variables and their variance for further statistical tests.
(2) Ordinal: Size of castle (size): 1. Fortress, 2. Small castle,
3. Medium castle, 4. Big castle; Isolation of castle (isolatio):
1. Island-castles on isolated lone hills, 2. Spit-castles on rocky
spits above the river, higher than surrounding terrain,
3. Rise-castles at the same or lower altitude as the surrounding
The area and diversity of the environment are herein expressed
by the size of the castle and heterogeneity of the environment –
beside others, and by the stage of castle destruction, which also
depends on the century in which the destruction started.
The shares of the individual environmental variables on
species variability of molluscan communities were expressed by
Figure 1. Position of sampling castles in the area of the Czech Republic. The grid represents rectangles of 0.18 latitude and 0.188 longitude (12 11.2 km).
234
CASTLE RUINS AS REFUGES FOR ENDANGERED MOLLUSCS
In the analysis of environmental variables, the partial
influence of geological substrate and the occurrence of woodland
species making use of citrate lime from the litter were tested in
separate steps. A partial analysis for nominal variables (rock
substrates and woods) as explanatory variables was used, with
the other environmental variables as covariates. In this way,
the additional influence of these variables was determined.
standardized partial regression coefficients (beta weights)
sometimes also used for path analysis
(Legendre & Legendre,
p
1998). The numbers of species ( N) were tested against the
castle size, its isolation, habitat variability and the stage of
destruction as the main explanatory variables.
Statistical analysis
Direct gradient analysis was used to explore multidimensional
data sets of species and environmental variables (Jongman
et al., 1995; Lepš & Šmilauer, 2003) using the program
CANOCO for Windows, v. 4.5 (ter Braak & Šmilauer, 2002).
The species data were log-transformed as Y ¼ log (n þ 1),
where n is the number of individuals, which increases the relative
rate of rare species that are mostly good indicators (Lepš &
Šmilauer, 2003). Centring by species and biplot scaling were
selected, and standardization by species was selected due to
the differences in species quantity. Eigenvalues (l) measure
the explanatory power of axes and express their relative importance. The categorial data on geological substrate and prevailing
woods were nominal ‘dummy’ variables (0, 1).
As a first step, a unimodal model (weighted averaging) of
species response to underlying environmental gradient was
used, because our samples were widely distributed across the territory of the Czech Republic. Simultaneously the gradient
lengths were tested in DCA (see Lepš & Šmilauer, 2003: 50).
From the explanatory variables, a subset of significant predictors
was selected using the Monte Carlo permutation test (MCPT)
with forward selection procedure (for details see Lepš & Šmilauer,
2003: 41, 64), because some of the explanatory variables were
highly correlated. MCPT procedure tests the significance of a
regression (F-statistics and probability of Type I error) under
the null hypothesis of independence of species data from the
environmental variables. The number of permutations was arbitrarily assigned to 999. This test evaluated the power of particular environmental variables. The significant ones were
illustrated by ordination biplots (ter Braak & Šmilauer, 2002).
Limestone, oreophyticum, century of destruction, stage of
destruction, beech, shrubs, pine, acidic crystalline rocks and isolation were treated as factors, while the other variables were
treated as covariates.
RESULTS
Fauna
Samples from 114 castles yielded 110 species of molluscs, which is
70% of the terrestrial species living in the Czech Republic. Of
these, 21% were ranked within the first three categories of the Red
List of IUCN for the Czech Republic (critically endangered – one
species, endangered – seven species, vulnerable – 16 species).
In their zoogeographic composition, castle malacocoenoses
generally corresponded to the composition of fauna in the
Czech Republic as a whole. The list of species found at the
studied castles is given in the Appendix.
Species diversity
Species diversity, expressed by the number of species, is traditionally related to the area and diversity of the environment
(Rosenzweig, 1995). Pairwise correlation coefficients indicate
mutual correlations among variables. The century in which
the castle was abandoned especially affected the stage of its
destruction (r ¼ 0.7, P , 0.001); the latter variable was therefore used for further analyses (stage of destruction shows a positive correlation also with the number of visitors and castle
isolation, but is negatively correlated with castle size and
biotope variability). The stage of castle destruction had a negative effect on the number of species (r ¼ – 0.32, P ¼ 0.001).
Preserved and especially partly disintegrated castles, which
have the greatest habitat heterogeneity, had the greatest species
diversity. A ruin being integrated into nature does not harbor
sufficient habitat heterogeneity and therefore loses the island
effect. Thus species diversity in these ruins was lower (Fig. 2).
Figure 2. Box-and-whisker plot of dependence of the number of species (square root transformed) on the stage of destruction (STAD, four categories):
1, preserved castle; 2, ruin of the whole castle high walls and towers, preserved interior spaces; 3, ruin partly disintegrated, only parts of the walls
without of interior space; 4, remains of the ruin integrated into the natural environment.
235
L. JUŘIČKOVÁ & T. KUČERA
Figure 3. Box-and-whisker plot of dependence of number of species (square root transformed) to the size of the ruins (four size categories): 1, fortress;
2, small castle; 3, medium castle; 4, big castle.
On the other hand, castle size was positively related to the
number of species (r ¼ 0.44, P , 0.001), which corresponds to
the traditionally perceived dependence of the number of species
on the size of the area (Fig. 3). Larger castles had greater biotope
variability (r ¼ 0.25, P ¼ 0.008). However, larger castles were
usually less ruined (r ¼ – 0.51, P , 0.001) and also less isolated
(r ¼ – 0.43, P , 0.001). Castle size explained 15.8% of the
variability in species numbers [r ¼ 0.319, F(3,110) ¼ 6.89,
P , 0.001] but also countered the negative effect of castle
isolation (r ¼ – 0.332, P , 0.001) and the amount of their
destruction (r ¼ – 0.393, P , 0.001).
Table 1. Environmental variables that explained a significant
proportion of species variability (for abbreviations, see text).
Variable
The effect of environmental variables on species
composition variability
The CCA direct ordination method was used to determine the
effect of the selected environmental variables on the variation
in species composition of castle communities. Of the original
32 environmental variables, nine were chosen using the forward
selection procedure (see Methods) (Table 1). These nine variables
explained 52% of the total variation in species composition
(Table 2). The first CCA ordination axis was dominated by
the effect of limestone on species composition (Fig. 4). Variability
of the species data along the second axis was distributed
according to the position of the castles in the oreophyticum,
isolation of the castle, the century of destruction and stage of
destruction. Thus, the presence of shrubs and pine, which were
absent from castles in the oreophyticum, represent substitute
variables here, probably for warmer lower locations of the mesophyticum, which were not adequately covered by the subdivision into three phytogeographic areas (Fig. 4). The presence of
oreophyticum indicator species increased along the second axis
(Discus ruderatus, Semilimax kotulae and Clausilia cruciata together
with Causa holosericea are typical species of higher-altitude mesophyticum and oreophyticum). These species were complemented by some hydrophilous species (Macrogastra ventricosa, Vertigo
N
Lambda A
P
F
limestone
14
0.13
0.015
5.08
oreo
3
0.07
0.012
2.60
cen-des
8
0.06
0.001
2.65
st-ruin
9
0.06
0.007
2.40
beech
27
0.05
0.001
2.06
shrubs
12
0.04
0.001
1.75
pine
25
0.05
0.009
1.72
aci-cry
23
0.03
0.026
1.40
isolatio
7
0.03
0.040
1.39
Table 2. Summary of the CCA method.
Axes
1
2
3
4
Total
inertia
Eigenvalues
0.142
0.116
0.078
0.050
Species –environment correlations
0.725
0.768
0.766
0.790
of species data
4.7
8.6
11.2
12.8
of species – environment relation
27.2
49.4
64.3
73.9
3.008
Cumulative percentage variance:
Sum of all unconstrained
3.008
eigenvalues
Sum of all canonical eigenvalues
0.522
substriata, Succinea putris, Aegopinella nitens, Zonitoides nitidus ) and
sensitive woodland elements (Bulgarica cana, Petasina unidentata
and Platyla polita ). In addition, there were elements of the
thermophyticum (Oxychilus draparnaudi, Cecilioides acicula and
236
CASTLE RUINS AS REFUGES FOR ENDANGERED MOLLUSCS
Figure 4. Ordination diagram of constrained unimodal CCA method; species and environmental variables in the first factorial plane. The inset shows
the environmental factors on the same co-ordinates. Abbreviations: aci-cry, acidic crystalline; isolatio, isolation of the castle; st-ruin, stage of destruction;
pine, beech, surrounding vegetation; shrubs, habitat type; cen-des, century of castle destruction; oreo, oreophyticum; limeston, geological background.
For species abbreviations see Appendix.
composition was mostly affected by some woody plants, which
do not represent a substitute source of calcium (beech, spruce,
hornbeam). This cannot be obviously explained by the preference of molluscs for such trees. Nevertheless, these trees may
act as substitute variables for other important environmental
variables in the given model (i.e. beech and spruce probably represent the oreophyticum). Interestingly, there was no obvious
effect of deciduous woody plants on mollusc variability. These
tree species are known to provide molluscs with calcium (e.g.
lime, ash, maple). This suggests that the presence of such trees
in castle habitats is due to the specific habitat conditions (soil
enrichment in calcium, origin of secondary talus) that are
equally favourable for molluscs.
Xerolenta obvia ), together with some species of open habitats
(Vertigo pygmaea ) and cliffs (Itala ornata, Clausilia parvula, Faustina
faustina, Pupilla sterrii ). Species positively correlated with limestone as the rock substrate were possible outliers. The presence
of limestone probably imposed a bias on the distribution
defined by the first ordinance axis.
The effect of the individual categories of rock substrate (see
Methods) was tested separately, with other variables entered
as covariates. The variability in species composition proved to
be most affected by limestone as the most favourable rock.
There was also a significant effect of acidic crystalline rocks,
which were the least favourable rock substrate, on the presence
of molluscs (Table 3).
The effect of different tree species on the variability of molluscs
was also subjected to separate test because tree litter, i.e. the presence of calcium in citrate bond in leaves of some woody plants,
may be an important source (e.g. Wäreborn, 1969, 1970;
Waldén, 1981). Other environmental variables were taken
as covariates again (Table 4). The variability in species
Species variability of molluscs eliminating the effect
of limestone
Castles built on limestone were outliers in the previous analysis,
based on the shape of the first axis (Fig. 4). Therefore, limestone
Table 3. Effect of the geological background on species variability.
Variable
Lambda A
P
Table 4. The influence of tree litter on species variability.
F
Variable
Lambda A
P
F
limestone
0.13
0.001
5.08
beech
0.06
0.001
2.33
acidic crystalline rocks
0.04
0.002
1.74
spruce
0.05
0.001
1.89
basalts
0.04
0.172
1.22
hornbeam
0.05
0.008
1.73
calcareous rocks
0.02
0.335
1.05
sycamore maple
0.03
0.124
1.31
quartzites, silicites
0.03
0.36
1.05
Norway maple
0.03
0.163
1.26
1.07
granites
0.02
0.425
0.99
pine
0.03
0.329
basic crystalline rocks
0.03
0.521
0.91
larch
0.03
0.332
1.04
sandstones
0.02
0.509
0.81
lime
0.02
0.349
1.07
trachytic rocks
0.02
0.624
0.77
ash
0.03
0.388
1.02
shales
0.02
0.694
0.7
oak
0.02
0.787
0.74
237
L. JUŘIČKOVÁ & T. KUČERA
was excluded as an environmental variable from the analysis, as
well as the three castles built on limestone and all species occurring only at these castles (Truncatellina costulata, Truncatellina
claustralis, Granaria frumentum, Chondrina clienta ). At the same
time, two other species (outliers) found at a single locality only
were also excluded (Bulgarica cana, Semilimax kotulae ).
After the effect of limestone (covariate) was excluded, the
overall proportion of the explained variability of the species
data covered by the first axis increased by ca 3% (Table 5).
This revised model (Fig. 5) showed not only the above mentioned subdivision of species by phytogeographical regions (distributed along the first ordination axis in this case), but also
better revealed the effect of acidic crystalline rocks, which is
tolerated by species of mountain forests and some hydrophilous
elements (Succinea putris, Eucobresia diaphana, Vertigo substriata ).
Species with a high nutrient demand were negatively correlated
with these crystalline rocks (Chondrula tridens, Pupilla sterrii,
Cochlodina orthostoma ). This model also more clearly showed the
effects of castle isolation, and the stage and century of its destruction. While in the model with limestone as the most important
factor (Fig. 4), the stage of destruction seemed to be closely
correlated with castle isolation, the model with the limestone
effect removed clearly showed that this was not the case. Isolated
castles host only small-sized species dwelling on particular
Table 5. Summary of the CCA model, excluding the influence of
limestone.
Axes
1
2
3
4
Total
inertia
Eigenvalues
0.114
0.070
0.050
0.035
Species-environment correlations
0.772
0.768
0.776
0.682
of species data
4.3
7.0
8.9
10.2
of species – environment relation
31.4
50.7
64.5
74.3
2.620
Cumulative percentage variance:
Sum of all unconstrained
2.620
eigenvalues
Sum of all canonical eigenvalues
0.361
surfaces, e.g. the upper edges of half-destroyed walls and other
small habitats (Pupilla muscorum, P. sterrii ). The model also
showed that isolation is better tolerated by species of open habitats than by woodland and hydrophilous elements.
Species characteristic of woodland talus, such as Helicodonta
obvoluta, Sphyradium doliolum and Vitrea diaphana, preferred castle
ruins whose destruction started earlier and whose secondary
Figure 5. Ordination diagram of constrained unimodal CCA method; species and environmental variables excluding the influence of limestone in the
first factorial plane. Abbreviations: aci-cry, acidic crystalline; isolatio, isolation of the castle; st-ruin, stage of destruction; pine, beech, surrounding
vegetation; shrubs, habitat type; cen-des, century of castle destruction; oreo, oreophyticum. For species abbreviations see Appendix.
238
CASTLE RUINS AS REFUGES FOR ENDANGERED MOLLUSCS
a direct RDA method was used for this analysis to provide
pairwise correlation coefficients for species and environmental
variables (values in parentheses).
Phytogeographical regions: The most important elements of
the oreophyticum at the castles were Discus ruderatus and Causa
holosericea, a species of higher-altitude woodland talus. The
only species found at the castles whose occurrence correlated
with the mesophyticum was Oxychilus cellarius, a modern immigrant of later periods of the Holocene. This species was incorporated in woodland as well as strongly anthropogenically affected
communities. The occurrence of the synanthropic species Oxychilus
draparnaudi was correlated with the thermophyticum.
Isolation of the castle: An interesting correlation was that
between the occurrences of the minute species Pupilla muscorum
and Truncatellina cylindrica and isolated castles. These species can
colonize small patches on the upper edges of castle walls, to which
they can be brought by long-distance transport by birds. For
example, an interesting and very isolated population of
P. muscorum was found inside the castle tower of the Hartenštejn
Castle ruin, which stands on a solitary hill in the middle of
fields. White Barn Owls nest in the tower. These birds could
have brought the mollusc species here. Helicigona lapicida was
another species tightly correlated with isolated castles, colonizing
castle walls.
Century and stage of castle destruction: Castles destroyed a
longer time ago correlated best with the woodland talus
species Helicodonta obvoluta, which uses the secondary talus from
deteriorated walls as a suitable habitat. In contrast, the environment of intact castles best correlated with the species Cochlicopa
lubrica, an indicator of early successional stages. More intact
castles provided a wider range of different habitats. These
were correlated with the occurrence of open-habitat species
Vallonia costata and V. pulchella, P. muscorum and the terricolous
Cecilioides acicula.
Attendance (number of visitors): Increasing attendance correlated with the presence of the above mentioned species P. muscorum, which favours trampled grass, and the species Trichia
hispida, one of the key elements of secondary urban habitats. Surprisingly, no species with tight links to infrequently visited castle
ruins were found.
Rock substrate: By far the strongest links were found for calcicolous elements and other species whose occurrence were at their
greatest on limestone substrates. It is generally understood that
species like Pyramidula pusilla, Granaria frumentum, Chondrina clienta,
Pupilla triplicata, Truncatellina claustralis, T. costulata, Oxychilus
inopinatus, Cepaea vindobonensis and Xerolenta obvia favour limestone
habitats, with some of them are restricted solely to this substrate.
Cochlodina commutata, a species in the Czech Republic most frequently reported from sites on opukas (i.e. spiculitic sandy marlstones) from the Orlické hory Mountains piedmont area, proved
to be an indicator of the presence of calcareous rocks at the
castles. Monachoides incarnatus and Causa holosericea were species
capable of withstanding the unfavourable environment of
acidic crystalline rocks.
Trees: In general, trees at castles were a less important
environmental variable, as implied from the above given
overall CCA analysis. The strongest effects perceptible in the
overall analysis were those of beech and pine. However, these
trees were not favourable habitats for molluscs. The only proven
correlations were between sycamore maple and Aegopinella pura
and between Norway maple and Cochlodina laminata.
woodland communities have already passed through the early
stage of succession. The species Daudebardia rufa, Oxychilus depressus
and Platyla polita were correlated with disintegrated castle ruins,
which were reverting to a natural state. In contrast, species of
open secondary habitats (Xerolenta obvia ), synanthropic species
(Oxychilus draparnaudi ) or weed species (Trichia hispida and
Cochlicopa lubrica ) were present in castles that are maintained
and managed. In this model, beech seemed to function as a substitute environmental variable, being positively correlated with
woodland demanding elements and negatively correlated with
open-habitat and weed species.
DISCUSSION
Ecological groups
Ložek (1964) suggested a subdivision of molluscs into 12 ecological
groups (see caption of Fig. 6). Although this subdivision was originally conceived for use in palaeontology, it is currently the most
widely used categorization for living molluscs. A comparison of
the ecological groups found at the castles with the fauna of the
Czech Republic as a whole is given in Figure 6. The castles were
dominated by woodland species of ecological groups 1, 2 and 3
(48% in total). This fact can be explained at least by the partial
overgrowing of most ruined castles by a secondary talus forest,
which provides favourable habitats for woodland species in general
(group 1) as well as for rather hydrophilous woodland species of
group 3. This proportion is, however, slightly lower compared with
that of woodland species (50% in total) in the overall composition
of the terrestrial molluscan fauna of the Czech Republic
(Juřičková, Horsák & Beran, 2001). Open-habitat species of
groups 4, 5 and 6 are also common (22% in total); this proportion
is somewhat higher than that in the overall composition of fauna in
the Czech Republic (19% in total). Group 4 species (xerothermic
steppe species) have a rather low proportion in the castle
communities, while group 5 species (non-forested areas in general)
were, however, quantitatively over – represented in the castle
communities. The catholic species of group 7, generally tolerating
both open areas and forests of different humidities, also occur in a
somewhat higher proportion (23%) than in the Czech fauna as a
whole (19%). Hydrophilous species of groups 8 and 9 are obviously
less frequently represented at the castles (7% relative to 12%).
Do characteristic species exist for particular
environmental variables?
A further aim was to discover which molluscan species were well
correlated with the separate environmental variables. Therefore,
Number of species and island effect
Figure 6. The comparison of the number of species in ecological groups
(Ložek, 1964) in the faunas of the Czech Republic and in the sampled
castles. 1, closed forest; 2, predominantly forest; 3, humid forest;
4, steppe – xerothermic habitats; 5, open grounds in general; 6, woodland
and grassland predominantly dry; 7, wetter woodland and grassland;
8, mainly wetlands; 9, strictly wetland.
An increase in species number with increasing area is a common
rule of island biogeography (MacArthur & Wilson, 1967). It is
therefore not surprising that higher species diversity was encountered at larger castles (Fig. 2). Nevertheless, an increase of area
239
L. JUŘIČKOVÁ & T. KUČERA
(105 sites) and Discus rotundatus (111 sites), had the highest
number of occurrences at castle sites. Punctum pygmaeum is a
pioneer species, which colonizes habitats in the early stages of
succession. It is surprising that this is the only species of this
group notably represented in the castle communities, quantitatively as well as qualitatively. The other two species belong
to the so-called molluscan weeds, i.e. species occurring practically everywhere at favourable sites.
Terricolous and semi-terricolous species – indicators of humic
soils: The presence of these gastropods indicated soils with a deep
humic horizon, in the case of castles a fully developed secondary
Rendzina imitating calcareous soils (Ložek, 1964). Cecilioides
acicula was encountered at 28 ruins and Vitrea contracta at
17 ruins. In only eight cases were the two species found together.
Their occurrences were clearly concentrated at castles lying in
the hill country and upper hill country belts, and were absent
from castles in western Bohemia.
Pioneer species – indicators of early stages of succession: The
term pioneer species refers to a group of catholic species that survived through the glacial period in the Czech Republic. These
can often be found within communities in the initial stages of
succession or as quantitatively indistinct components of communities living in different, mostly shaded habitats. The pioneer
species themselves, together with several slugs (living mostly
on fungi), constitute characteristic communities of central
European oligotrophic forests. These forests are an unfavourable
environment for almost all woodland species and, therefore,
provide no competition to the pioneer species. This group
includes Perpolita hammonis, Punctum pygmaeum, Euconulus fulvus
and Cochlicopa lubrica (in the weed species group these communities
were completed by Vitrina pellucida and Monachoides incarnatus ). The
marginal occurrence of such species (except of Punctum pygmaeum ) in the castle environments can be explained by the
high number of their competitors living in the nutrient-rich set
of castle habitats.
Adventive species – indicators of anthropic stress: Synanthropic
species were generally infrequent at the castles, both quantitatively and qualitatively. More frequent were some hemisynanthropic species (species that live in some natural habitats, but
their optimum condition is in man-made habitats). These
include Deroceras reticulatum (found at 24 castles) and Arion
fasciatus (found at 12 castles). True synanthropic species were
found very rarely.
The invasive slug Boettgerilla pallens, originally from the
Caucasus Mountains, and which has colonized practically the
whole of Europe since the 1960s, lives in both human-influenced
and natural habitats in Bohemia and elsewhere. It did not reach
high abundances in the castle communities and was found at
11 castle ruins. The most recently arrived invasive species in
the Czech Republic, Arion lusitanicus, was found at a single castle.
This locality was, however, managed as a park and A. lusitanicus
was probably introduced there together with plants.
All of these data show that castles host their own unique
mollusc communities with a structure (dominance relation,
composition, frequency) completely different from other known
mollusc communities.
also implies an increase in habitat diversity, often functioning as
an intermediary between species number and area (Kohn &
Walsh, 1994). This, however, could not be proved from the
present dataset.
Castles as refuges for species in the landscape
Castle species: ‘Castle species’ are the species that display a
more frequent occurrence at castles or are more abundant at
the castles, relative to their natural habitats, over most of the
territory of the Czech Republic. A known castle species for
over 100 years is Balea perversa, which is continuously distributed only in the Pálava area (S Moravia) as an element of
rock-steppe communities. Elsewhere, it lives on islets of noncalcareous coarse scree without matrix or very loosely scattered
on cliffs (Ložek, 1956). This species was found at 23 studied
castles, inhabiting shaded castle walls and talus. Another,
Laciniaria plicata, is relatively abundant only on marlstone substrates in the NE quadrant of Bohemia, and less abundant but
distributed over a wider range of sites in Moravia. It shows a
tendency towards overpopulation at the castles, is often vicarious with the otherwise common species Alinda biplicata (in
the very NE quadrant), or is much more abundant compared
with the latter species. Laciniaria plicata was found at 21 studied
castles. Clausilia parvula has a similar natural distribution to that
of L. plicata. It was, however, much more abundant in Bohemia
in the period of the climatic optimum, i.e. at higher humidity
(Ložek, 1964). Its occurrences at Andělská hora Hill near
Karlovy Vary or Bezděz Hill are isolated introductions. It
was found at 12 castle ruins. Clausilia dubia favours the walls
of ruined castles. In nature, this species prefers limestone cliffs
of moderate humidity but can be rather rarely found also on
tree trunks in talus forests. Calcium-enriched shaded walls of
ruined castles provide an ideal environment for this species.
It was found at 43 castle ruins. The common species Alinda
biplicata is the last species of the family Clausiliidae with a tendency to proliferate at castles. It occurs in a variety of habitats
ranging from forests to gardens and parks in towns and villages.
It was found at 82 castles, ranking quantitatively as the second
most common gastropod of castle ruins. Open habitats at the
castles were inhabited by small gastropods. Pupilla muscorum is
an open-country species, which survived on cold steppes in
the glacial periods and still occurred in the earliest Holocene.
After its disappearance in the period of climatic optimum, it
re-appeared in the Middle Ages (Ložek, 1964). Subsequently,
it is found mainly in secondary open habitats, the existence
of which is conditioned by human activity, and is practically
absent from habitats of a purely natural character. In the
last few years, however, this species has declined in most of
its usual habitats in Bohemia. Large and viable populations
can still be found on castle ruins. It was found at 70 castle
ruins, dwelling in open habitats in castle interiors and on the
upper edges of castle walls, where its populations often reach
striking abundances. Most of the castle ruins hosted both of
the common species of the family Valloniidae: Vallonia pulchella
and V. costata. The latter species was a typical castle species
because it was markedly dominant at the castles, as opposed
to most other favourable habitats where the numbers of the
two species were more or less equal (V. pulchella is heliophilic
but tolerates higher humidity levels while V. costata tolerates
slight shade but is more xerophilous). Vallonia costata was
found in 91 castle ruins and was the most abundant species
of these habitats. Helicigona lapicida was a typical inhabitant
of castle walls. It cannot be considered as a typical castle
species, because it probably colonized the castle sites before
their construction. Still, it found optimum conditions in
shaded wall habitats and was found at 86 castle ruins. Three
other species, Punctum pygmaeum (101 sites), Vitrina pellucida
Molluscs and phytogeographic regions
In his recent study, Ložek (2000) classified molluscan species
using the phytogeographic zonation of the Czech Republic
(Skalický in Hejný & Slavı́k, 1988). The influence of this phytogeographical zonation on castle faunas was tested in this study.
Most castles lie in the mesophyticum at a mean altitude of
around 490 m a.s.l. Thirteen castles lying in the thermophyticum were studied quantitatively. The only species found in
higher abundances here, and which is typical for the thermophyticum and lower parts of the mesophyticum, was the terricolous
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CASTLE RUINS AS REFUGES FOR ENDANGERED MOLLUSCS
of castle walls are very often covered by xerothermic vegetation,
creating a thin layer of calcium-rich soil. This is where characteristic communities of small gastropods can be found (especially
Pupilla muscorum, Truncatellina cylindrica and Vallonia costata ).
Shrubs within the walls are inhabited by species of forest-edge
communities, such as Helix pomatia or Euomphalia strigella. Sometimes these are colonized by species making use of cold, shaded
corners of the former residential spaces, such as the relatively
recent immigrant Oxychilus cellarius. Open habitats on secondary
Rendzinas can be also found in the castle area. Rendzinas can
develop (owing to weathered mortar) even on silicate substrates
and can support the terricolous blind gastropod Cecilioides acicula.
On the outer side of the walls, the ruins are turned into artificial
talus accumulations of different exposure, either bare or overgrown by a secondary talus forest that includes a number of
sensitive woodland species, such as Vertigo alpestris. Talus
accumulations more deeply incorporated into the woodland
environment host the species Helicodonta obvoluta and Daudebardia
rufa, index species of destroyed castle ruins. All of these types
of habitats gradiate into one another with different transitions, actually small ecotones, that generally increase species
diversity.
Cecilioides acicula. At the other end of this scale, eight castles from
the oreophyticum were studied quantitatively. The most frequently encountered species of those, characteristic for the
oreophyticum according to Ložek (2000), were Discus ruderatus
and Causa holosericea, an element of the higher-altitude mesophyticum and the oreophyticum. Significant elements of the
mesophyticum include most woodland species of moderate altitudes, i.e. species whose proportions in the castle communities
were the highest. The most important mesophyticum elements,
Clausilia parvula and Daudebardia rufa, were relatively rare in
Bohemia, although their abundances at the studied castles
were relatively high. In summary, castles in the two extrazonal
phytogeographic regions (thermophyticum and oreophyticum)
generally hosted only weak or no populations of elements indicative of these regions, and the respective index species contributed
only weakly to the castle communities in this particular region.
In spite of this fact, phytogeographic regions, and the oreophyticum in particular, proved to be one of the most important
environmental variables to explain the species variability of
molluscan communities. These differences are defined, with
regard to castles, by relative abundances in the malacocoenoses
rather than by the presence of indicator species. Nevertheless,
the evidence for the influence of phytogeographic regions on
malacocoenoses supports the validity of the proposed zonation
(Ložek, 2000).
CONCLUSIONS
Castles represent characteristic sets of habitats markedly different from the surrounding landscape. Castles function as important refuges for a number of species, as indicated by the high
species diversity and the presence of rare and uncommon
species. The effect is much less for castles built on a limestone
substrate, where the surrounding area may support a similar
range of microhabitats.
Evolution of castle malacocenoses
Although some of the studied castles were sampled previously,
the sampling methods used then were non-quantitative, mostly
searching and handpicking, which makes quantitative comparision impossible. Nevertheless, the presence or absence of previously found species at a given site allows conclusions to be
drawn about the possible changes taking place. Such a comparison leads to the conclusion that castle communities have probably not been subjected to any major changes and, unless
local conditions change dramatically, they will maintain their
species diversity. Only Cochlodina orthostoma was found at much
fewer numbers than 50 years ago at the castles. This is a vulnerable species that is generally rare in Bohemia.
Specific comparisons of molluscan community compositions
were performed at two castle ruins that were studied by quantitative methods more than 50 years ago (Jandečka, 1935;
Culek, 1944). Neither of the two sites showed any major signs
of degradation of the local communities. This finding is significant, especially as tourist pressure on these castle ruins is
heavy. Similar conclusions were also made by Pfleger (1997),
who studied the malacofauna of Kašperk Castle 80 years after
Z. Frankenberger.
(1) Altogether 110 species of molluscs were identified at
114 castles. In its zoogeographic composition, the fauna of
the castles roughly corresponds to that of the Czech Republic
as a whole. For about 26% of the molluscan species, their
castle occurrences form islets outside the main area of their
distribution.
(2) The strongest effect of rock substrate was shown for limestone as a rock most favoured by molluscs, as opposed to
acidic crystalline rocks, which was an extremely unfavourable substrate. The size and isolation of castles are factors
implying their island character. The century in which a
castle was abandoned and the stage of its destruction are
the factors that most affect the occurrence of sensitive
species of woodland talus (at ruined castles) in the communities. Conversely, intact castles host open-habitat species
and molluscan weeds. Castle ruins, which have been fully
incorporated into the local environment, show low species
diversity. This is caused by the disappearance of habitat
diversity and by the fact that the secondary woodland
talus is still in an early stage of succession. The restoration
of woodland communities is generally a very slow process.
(3) There are distinctive species found on castles built on limestone. The effect of rock substrate type outweighs the effect
of other environmental factors of castle ruins in such cases.
(4) A significant effect of trees, whose litter contains citrate lime,
on the diversity and variability of molluscan communities at
castles could not be shown. Therefore, it can be assumed
that enrichment of the substrate with calcium from ruined
walls is a more important calcium source for molluscs than
tree litter.
(5) The study revealed that many of the molluscan species at
castles were indicators of the studied environmental
factors. These included species indicative of phytogeographic regions, confirming the views of Ložek (2000),
Castles and habitat mosaics
Castle ruins are sets of habitats characterized by a specific relief
formation and peculiar properties, not of the rock substrate but a
secondary structure created by human activity. In their hydrological and atmospheric regimes, castle ruins differ from their
surroundings. Their flora and fauna contrast with those of the
average character of the surrounding landscape. Thus, they
increase the local species diversity.
A castle ruin comprises several lower-order habitats differing
in their orientational direction and structure. Wall remains
simulate either insolated or, on the contrary, shaded artificial
cliffs, either protected or exposed, and enriched in calcium
from mortar. The presence of calcium, even in mortar-free
joints between stones, is indicated by the presence of wall rue.
The castle species Helicigona lapicida and often also Balea perversa
are almost always found living on wall surfaces. The upper edges
241
L. JUŘIČKOVÁ & T. KUČERA
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talus species in castle ruins and, alternatively, xerothermic
species on intact castles. The degree of isolation of a castle
was indicated by the presence of small, open-country
species. Visitor pressure at a castle was correlated with the
presence of the weed species Trichia hispida.
(6) Of the 110 molluscan species identified at the studied castles,
21% are ranked in the first three categories of the IUCN
Red List for the Czech Republic. Synanthropic species
were almost absent from the castles. The species which
were more frequent or especially abundant in the castles
than in their natural habitats are herein called ‘castle
species’: Balea perversa, Laciniaria plicata, Clausilia parvula,
C. dubia, Alinda biplicata, Pupilla muscorum, Vallonia costata,
Helicigona lapicida.
(7) No general trends towards degradation of castle communities of molluscs over the last 100–50 years were observed.
Our study shows that human influence need not be always
directly associated with landscape degradation and reduction
of species diversity. The very opposite was observed in the case
of the castles.
ACKNOWLEDGEMENTS
We especially thank V. Ložek for his critical reading of the
manuscript and also M. Horsák and J. Č. Hlaváč for providing
unpublished materials. We thank also referees J. Nekola
(Wisconsin), B. Pokryzsko (Wroclaw) and R. Cameron for
their valuable comments on the manuscript. We thank
K. Edwards for improving the English. The study originated
with support from the Grant Agency of Charles University,
Project No. 112/1998/Bbio/PřFUK, and Research Plan MSM
113100003. T. Kučera was supported by the Institutional
Research Plan Z6087204.
=
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L. JUŘIČKOVÁ & T. KUČERA
APPENDIX
Molluscan species identified at 114 castles in the Czech Republic.
Scientific name
Diagram
Conservation
Number of castles with
Total number of
abbreviation
status
occurrence of species
collected individuals in
litter samples
Platyla polita
Plat-pol
VU
17
74
Carychium tridentatum
Car-tri
LC
40
4041
Carychium minimum
Car-min
LC
3
252
Cochlicopa lubrica
Co-rica
LC
60
1248
Cochlicopa lubricella
Co-cella
LC
48
1221
Pyramidula pusilla
Pyr-pus
VU
2
136
Columella edentula
Col-ede
LC
34
968
Truncatellina cylindrica
Tru-cyl
LC
96
10,346
Truncatellina claustralis
Tru-cla
EN
1
42
Truncatellina costulata
Tru-cos
CR
1
21
Vertigo pusilla
Ver-pus
NT
64
1136
Vertigo substriata
Ver-sub
NT
3
61
Vertigo pygmaea
Ver-pyg
NT
12
25
Vertigo alpestris
Ver-alp
VU
42
548
Granaria frumentum
Gra-fru
NT
2
62
Chondrina clienta
Chon-cli
VU
1
3
Sphyradium doliolum
Sphy-dol
NT
14
3431
Pupilla muscorum
Pup-mus
NT
70
7958
Pupilla triplicata
Pup-tri
VU
1
100
Pupilla sterri
Pup-ster
VU
1
786
Acanthinula aculeata
Acan-acu
NT
46
1573
Vallonia costata
Vall-cos
LC
91
15,979
Vallonia pulchella
Vall-pul
LC
70
1920
Vallonia excentrica
Vall-ex
DD
25
158
Ena montana
Ena-mon
NT
43
1222
Merdigera obscura
Mer-obs
LC
12
346
Chondrula tridens
Cho-tri
VU
1
1
Cochlodina orthostoma
Coch-ort
VU
3
46
Cochlodina laminata
Coch-lam
LC
49
1147
Cochlodina commutata
Coch-com
VU
1
28
Cochlodina dubiosa corcontica
Coch-dub
EN
2
7
Itala ornata
It-orn
VU
4
212
Macrogastra ventricosa
Mac-ven
NT
2
21
Macrogastra plicatula
Mac-pli
NT
33
1294
Clausilia parvula
Clau-par
NT
12
1317
Clausilia dubia
Clau-dub
LC
43
1712
Clausilia cruciata
Clau-cru
VU
2
18
Clausilia pumila
Clau-pum
LC
4
61
Ruthenica filograna
Ruth-fil
VU
10
304
Laciniaria plicata
Lac-pli
NT
21
2,249
Alinda biplicata
Ali-bip
LC
82
12,528
Balea perversa
Bal-per
VU
23
634
Bulgarica cana
Bul-can
EN
2
13
Bulgarica nitidosa
Bul-nit
VU
1
36
Succinea putris
Suc-put
LC
1
6
Succinella oblonga
Suc-obl
LC
5
31
Cecilioides acicula
Cec-aci
LC
28
245
Punctum pygmaeum
Punc-pyg
LC
101
7,459
Discus rotundatus
Dis-rot
LC
111
7,350
Discus ruderatus
Dis-rud
NT
4
36
Discus perspectivus
Dis-per
VU
3
24
Zonitoides nitidus
Zon-nit
LC
1
3
244
CASTLE RUINS AS REFUGES FOR ENDANGERED MOLLUSCS
Continued
Scientific name
Diagram
Conservation
Number of castles with
Total number of
abbreviation
status
occurrence of species
collected individuals in
litter samples
Euconulus fulvus
Eu-ful
LC
55
377
Vitrina pellucida
Vit-pel
LC
105
5,001
Semilimax semilimax
Sem-sem
LC
34
229
Semilimax kotulae
Sem-kot
VU
1
5
Eucobresia diaphana
Euc-dia
LC
6
78
Eucobresia nivalis
Euc-niv
EN
1
1
Vitrea diaphana
Vit-dia
NT
18
224
Vitrea subrimata
Vit-sub
VU
10
361
Vitrea crystallina
Vit-cry
LC
9
216
Vitrea contracta
Vit-co
LC
17
167
Aegopis verticillus
Aego-ver
VU
6
48
Aegopinella pura
Aeg-pur
LC
53
3,000
Aegopinella nitens
Aeg-nit
LC
8
483
Aegopinella minor
Aeg-min
LC
95
2,607
Perpolita hammonis
Per-ham
LC
42
361
Oxychilus cellarius
Oxy-cell
LC
68
708
Oxychilus draparnaudi
Oxy-drap
LC
8
54
Oxychilus glaber
Oxy-glab
NT
20
168
Oxychilus depressus
Oxy-dep
NT
7
31
Oxychilus inopinatus
Oxy-ino
NT
1
1
Daudeberdia rufa
Dau-ruf
NT
21
113
Daudebardia brevipes
Dau-brev
EN
1
9
Tandonia rustica
Tan-rus
NT
5
Limax maximus
Lim-max
LC
5
Limax cinereoniger
Lim-cin
LC
37
Malacolimax tenellus
Mal-ten
LC
28
Lehmania marginata
Leh-mar
LC
35
Boettgerilla pallens
Boet-pal
LC
11
Deroceras laeve
Der-lae
LC
1
Deroceras sturanyi
Der-stu
LC
2
Deroceras agreste
Der-ag
LC
7
Deroceras reticulatum
Der-ret
LC
24
Arion rufus
Ari-ruf
LC
8
Arion lusitanicus
Ari-lus
LC
1
Arion subfuscus
Ari-sub
LC
36
Arion distinctus
Ari-dis
LC
9
Arion fasciatus
Ari-fas
LC
12
Arion circumscriptus
Ari-cir
NT
1
Arion silvaticus
Ari-sil
LC
14
Fruticicola fruticum
Fru-fru
LC
14
202
Trichia hispida
Tri-his
LC
30
1,042
Trichia sericea
Tri-ser
LC
6
135
Plicutheria lubomirskii
Pli-lub
NT
6
33
Petasina unidentata
Pet-uni
NT
18
1,010
Xerolenta obvia
Xer-ob
LC
7
106
Urticicola umbrosus
Urti-um
LC
27
1,239
Monachoides incarnatus
Mon-inc
LC
68
1,759
Monachoides vicinus
Mon-vic
NT
2
10
Euomphalia strigella
Euo-stri
LC
22
239
Helicodonta obvoluta
Hel-obvo
NT
24
121
Arianta arbustorum
Aria-arb
LC
47
690
Helicigona lapicida
He-lapi
LC
86
1,190
Faustina faustina
Fau-fau
VU
3
36
Isognomostoma isognomostomos
Iso-iso
LC
40
307
245
L. JUŘIČKOVÁ & T. KUČERA
Continued
Scientific name
Diagram
Conservation
Number of castles with
Total number of
abbreviation
status
occurrence of species
collected individuals in
litter samples
Causa holosericea
Cau-hol
NT
11
72
Cepaea hortensis
Cep-hor
LC
46
406
Cepaea vindobonensis
Cep-vin
NT
1
1
Helix pomatia
H-poma
LC
7
336
Conservation status in the Czech Republic (Juřičková et al., 2001; IUCN, 2001). Abbreviations: CR, critically endangered; EN, endangered;
VU, vulnerable; NT, near threatened; LC, least concern; DD, data deficient.
246