HABITAT CHOICE IN THE CARPATHIAN LAND SNAILS

MACROGASTRA TUMIDA (ROSSMÄSSLER, 1836) AND VESTIA

TURGIDA (ROSSMÄSSLER, 1836) (GASTROPODA: CLAUSILIIDAE)
ANNA SULIKOWSKA-DROZD
Department of Invertebrate Zoology and Hydrobiology, University of Lodz, Banacha Str. 12/16, 90-237 Lodz, Poland
(Received 21 January 2004; accepted 14 July 2004)

ABSTRACT
The aim of the study was to elucidate on the basis of physical, floristic and edaphic features of habitat, the
distribution in natural montane forests of the two endemic Carpathian clausiliid species, Macrogastra
tumida and Vestia turgida. Sites sampled were in the Bieszczady National Park, East Carpathians, Poland.
Snails were sampled quantitatively at 11 sites located along an elevation gradient in various floristic
communities. The estimated population densities varied from 0 to 49 /m2 of forest floor. In a multivariate
analysis, the first four axes accounted for 82.7% of variation in recorded environmental variation. The
first axis, representing habitat fertility, correlated most strongly with the density of M. tumida. Density of
V. turgida showed a weaker correlation with this axis, as it was relatively less abundant in the wettest sites,
which were also the most fertile. Multiple regression models were used to determine the effects of single
variables: plant species number, herb cover, slope aspect and habitat humidity were the best predictors of
the snails’ abundance at the sites. The interactions and intercorrelations of these factors are discussed in
relation to land snail abundance.

INTRODUCTION are endemic Carpathian snails, common locally but endangered

Two approaches have been used in determining the habitat
preferences of land snails: as a frequency of occurrence in the
particular habitat type (e.g. Cameron, 1973; Coney et al., 1982;
Proschwitz, 1993), or as the abundance of specimens in
standardized samples (e.g. Valovirta, 1968; Wäreborn, 1970;
Outeiro, Agüera & Parejo, 1993; Barker & Mayhill, 1999).
Although habitat distribution can be a product of many
interacting environmental factors (Peake, 1978), malacologists
usually discuss one or a few factors chosen a priori. Most often
these are calcium content of the soil and/or soil acidity, but
moisture, organic matter content, C/N ratio, soil structure,
closure of the vegetation and microclimate have also been taken
into consideration (Bishop, 1977; Burch, 1955; Cameron, 1973;
Dillon, 1980; Gosteli, 1996; Wäreborn, 1970). In mountain
regions, attention has also focused on climatic factors
dependent on elevation and aspect of slope (Cameron, 1973;
Dyduch-Falniowska, 1991; Waldén, 1981; Wäreborn, 1970). The
interrelationships between ecological factors have made it
difficult to detect the true determinants. Local climate, topo-
graphy, humidity and chemical contents of soil influence the
vegetation, which in turn modifies soil properties and micro-
climate. It is still an open question as to which factors can best
indicate distribution of terrestrial gastropods; moreover, the
importance of the same factor varies in different regions
(Wäreborn, 1969; Gosteli, 1996).
The aim of this study was to determine to what extent
environmental factors (topographic, edaphic and floristic)
control abundance of two snail species in a set of forest habitats.
Thus, emphasis was placed on gathering good quantitative
information on habitats for statistical analysis. The collecting
procedure enabled the estimation of the abundance of animals in
samples of soil and ground litter in terms of density/m2 of forest
floor.
The study concentrated on two clausiliids: Macrogastra tumida
(Rossmässler, 1836) and Vestia turgida (Rossmässler, 1836). These
Correspondence: e-mail: sulik@biol.uni.lodz.pl
in Poland as a whole (Wiktor & Riedel, 1992). Macrogastra tumida
inhabits montane mixed forests. Less often it is found in pure
deciduous or coniferous woods and in scrub (Riedel, 1988). It
lives in leaf litter, under stones and between shaded rocks
(Kerney, Cameron & Jungbluth, 1983). According to Kerney
et al. (1983), V. turgida occurs in very moist woodland, under logs
and ground litter, but Urbański (1939) showed that the species
avoided very damp places. Both species ascend to 2100 m in the
Carpathians (the Tatra Mountains) (Dyduch-Falniowska,
1991).
STUDY AREA
The study was conducted in a mountain region of the Bieszczady
Mountains, a Polish part of the East Carpathians (Fig. 1). Since
1973 the area has been protected in the Bieszczady National
Park, one of the largest national parks in the country and famous
for diversity of natural habitats and rich flora and fauna. The
region is one of the least polluted parts of Poland.
Ridges of the Bieszczady Mountains are made of flysch
(sandstones, siltstones and shales) with a locally significant
content of calcium carbonate (up to 35%). The highest summits
slightly exceed 1300 m above sea level. Soils are predominantly
cambisols. The most common are dystric cambisols, occurring on
non-calcareous flysch formation and on deep and decalcified
regolith covering flysch rooks rich in CaCO3. In dystric cambisols
soil reaction is low in a whole profile (pH 4.0 – 5.0), while in eutric
cambisols the reaction is from 5.5 to 6.5. Semi-hydrogenic and
hydrogenic soils occupy small and scattered areas on slope bends
and in valleys. The high soil pH (5.5 – 7.0) is characteristic of
them, due to the influence of nutrient-rich water coming from
underground. Semi-hydrogenic soils provide the most fertile
forest habitats in the Bieszczady Mountains.
Climatic conditions in the Bieszczady Mountains depend
mostly on elevation. Three climatic vertical zones are distin-
guished: moderately warm, moderately cool and cool. The limits
of particular zones are generally higher on southern slopes and on
convex ridges than on northern slopes and in concave valleys.

Journal of Molluscan Studies (2005) 71: 105–112 doi: 10.1093/mollus/eyi013

q The Author 2005. Published by Oxford University Studies on behalf of The Malacological Society of London, all rights reserved.
 A. SULIKOWSKA-DROZDA. SULIKOWSKA-DROZD

Figure 1. The localities described in this paper. A. The Carpathians, showing the location of study area in the Bieszczady National Park.
B. The Bieszczady Mountains, arrows showing the sampling stations (numbered 1 –11); contours in map B are given in metres a.s.l.

In lower parts of the area the average annual temperature
exceeds þ 6 8C and the vegetation season lasts 200– 214 days. On
the highest ridges the average annual temperature is lower than
þ3 8C and the vegetation season lasts 181– 192 days. Minimal
values of temperature occur in January (mean temperature from
2 3.4 8C in valleys to 26.7 8C on the highest ridges), while
maximal in July (þ16.6 8C in valleys and þ 11.5 8C in the highest
ridges). Mean annual precipitation is ca 900 mm in low parts of
the region and probably exceeds 1200 –1300 mm on the highest
ridges. The highest monthly rainfall is in July, the lowest in
January, February and October. Snow cover is present on ridges
from the second half of October and from the second part of
November in valleys. The mean number of days with snow cover
varies from 94 to 180. The upper timberline goes along the
isotherm þ 2 8C (ca 1000– 1200 m a.s.l.).
The Bieszczady Mountains are covered mostly by deciduous
forests; the fertile Carpathian beech wood (Dentario glandulosae-
Fagetum) predominates (80% of forested area). More locally,
acidophilous mountain beech wood, sycamore forests (with Acer
pseudoplatanus) and alder woods (with Alnus incana) occur. There
are subalpine shrubs of Alnus viridis and alpine meadows above
the timberline. Alder woods (plant association Alnetum incanae)
grow along rivers and streams up to 900 m a.s.l. Alder swamps
(plant association Caltho-Alnetum) occur in isolated patches on
low-permeability soil. About 14% of the forested area is occupied
by secondary forest communities. Some of them (with grey alder,
ash, goat willow and hazel) have spontaneously overgrown
unused farmland and some spruce and mixed stands have been
created by tree planting.
METHODS
Sampling procedure
Snails were collected at the 11 stations (each an area of
10 £ 10 m) located along a transect crossing two main mountain
crests, in different forest associations, at elevations from 650 to
1250 m a.s.l. (Fig. 1). At each station snails were collected twice:
in spring (May– June) and in late summer (end of August,
September). Quantitative samples of soil and leaf litter were

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 HABITAT CHOICE IN CARPATHIAN CLAUSILIIDAE

taken with an Ökland frame, 20 £ 20 cm. A total of 550

subsamples were collected: at each sampling occasion 25
subsamples added up to 1 m2 at a station. Since there were
some environmental differences within the study stations (e.g.
dense herb cover/bare ground/deep leaf litter), the subsamples
were collected not at random, but from microhabitats
proportionally to their distribution within each 10 £ 10 m
station. The same procedure was earlier applied by Dziecz-
kowski (1972), who examined the malacofauna of the
Carpathian beech woods in Poland. The forest litter, superficial
soil layer plus herbs were placed in plastic bags. Each
subsample was hand-sorted twice: immediately after bringing
it to the laboratory and then after drying and sieving the
material through sieves (mesh diameters: 5.0, 2.0, 1.0 and
0.5 mm). Only living individuals were counted.

Ecological site description

The following features were noted at each sampling station.
Physiographic: ALT, altitude a.s.l. (m); SLO, slope angle; ASP,
slope aspect (northern, southern, etc.).
Floristic composition and cover of the different vegetation strata:
PLANT, number of plant species; CANOP, canopy cover (%);
SHRU, shrub cover (%); HERB, herb cover (%); HUM, soil
moisture index (measured by occurrence of hydrophilous plant
species).
Edaphic factors (the average of the values measured in two soil
samples from each station were used in analysis): MORG,
organic matter content in superficial soil layer (after logarithmic
transformation); C/N, carbon/nitrogen ratio in the leaf litter; Ca,
calcium content in the leaf litter; pH, soil acidity; POR, soil
porosity.
The values of these parameters are presented in the Appendix.
The values were estimated in spring, thus the statistical analysis
was also based on spring densities of clausiliids. However, if both Figure 2. Distribution of Clausiliidae over sampling stations. A. Mean
spring and summer densities were averaged and used for analysis, density of Macrogastra tumida. B. Mean density of Vestia turgida.
the statistical results would be very similar and make no
difference to the conclusions. (15.5 ind./m2) in alder wood at the stream bank (Stn 8). The
correlation between spring and summer densities was significant
Statistical methods (r 2 ¼ 0.50; P , 0.05).
A PCA of the recorded habitat characteristics showed that the
The following methods were adopted: principal components first four principal components (eigenvalues . 1) explained
analysis (PCA) for environmental factors in the 11 sampling 82.7% of the total variation in environmental factors (Table 1).
stations; Pearson correlation coefficient between the first four PC I was strongly influenced by plant species richness, herb
components and the density of clausiliids; multivariate regression cover, Ca content and soil pH (Table 2). It can be called ‘habitat
of each set of clausiliid densities on the environmental factors. fertility’. The highest value of habitat fertility was recorded at Stn
The latter method can only be applied when the dependent
8 (alder wood at the stream bank) and the lowest in beech woods
variable has a normal distribution. Thus, densities of Macrogastra
(Stns 1, 3, 4) (Fig. 3). PC II was connected with soil porosity and
tumida were logarithmically transformed ½xt ¼ lnðx þ 1Þ
: Multi-
canopy cover; PC III with shrub cover; and PC IV with soil
variate regression was conducted by the stepwise method (Sokal
& Rohlf, 1995). moisture (positively) and angle of slope (negatively). PC IV can
be called ‘habitat humidity’. The lowest value of PC IV was
recorded at Stn 7, which was situated on a very steep slope. For
other stations the differences of habitat humidity were rather
RESULTS minor.
Between six and 32 species of land snails were recorded at a single
station on one sampling occasion (see Appendix). The abun- Table 1. Principal component analysis for the environmental factors.
dance of snails in the samples varied from 18 to 1069. The highest
species diversity and abundance occurred in alder woods (Stns 9, Principal components Eigenvalues Percentage of
8). Macrogastra tumida was found at eight stations (Fig. 2). The assignable variation
spring and summer densities of M. tumida were significantly
correlated (r 2 ¼ 0.88; P , 0.05). The highest density was I 5.33 41.03
assessed in alder wood at the stream bank (Stn 8) (average II 2.44 18.81
42 ind./m2) and in alder swamp (Stn 9) (38 ind./m2). Vestia III 1.87 14.41
turgida was found at nine sampling stations: in sycamore forests, IV 1.10 8.45
alder woods, secondary alder forest, subalpine shrubs and in some I – IV 82.70
beech woods. The highest density (30 ind./m2) was estimated in
sycamore forest with Allium ursinum (Stn 6) and the second highest Eigenvalues of the first four principal components.

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 A. SULIKOWSKA-DROZDA. SULIKOWSKA-DROZD

Table 2. Principal component analysis. Table 3. Pearson correlation coefficients (r) between principal
components and DENSITY of clausiliids.
Environmental PC PC PC PC
factor I II III IV Principal components DENSITY of
habitat fertility habitat humidity
Macrogastra tumida Vestia turgida
ALT 2 0.17 0.21 2 0.13 2 0.17
I 0.79* 0.61*
SLO 2 0.21 0.11 0.21 2 0.87
II 0.04 2 0.22
ASP 2 0.19 2 0.08 0.14 0.02
III 2 0.23 2 0.22
PLANT 0.92 2 0.05 2 0.14 0.15
IV 0.38 2 0.38
CANOP 2 0.33 2 0.81 0.20 0.09
SHRU 0.15 0.07 2 0.94 0.18 *P , 0.01.
HERB 0.82 0.02 0.01 0.08
C/N 2 0.54 2 0.31 0.59 2 0.09 range of site humidities studied, V. turgida appeared to be a less
Ca 0.71 0.28 0.08 0.27 hydrophilous species than M. tumida.
pH 0.82 0.11 2 0.05 0.45
POR 0.08 0.84 0.15 0.02
DISCUSSION
HUM 0.37 0.06 0.01 0.76
MORG 0.16 0.21 0.01 0.10 Data on the density of clausiliid populations are sparse. In
general, recorded densities do not exceed 50 ind./m2, but the
Coordinates of environmental factors on the first four principal components. highest densities are about 400 ind./m2 (Agócsy, 1968; Piechocki,
Factor loadings . 0.7 are in bold. 1982). The density of Macrogastra tumida population in beech
woods of the Tatra Mountains was as low as 1 – 6 ind./m2
Habitat fertility (PC I) was positively correlated with the (Dyduch-Falniowska & Tobis, 1989; Dzieczkowski, 1972), and
number of land snail species recorded at the stations (r 2 ¼ 0.56) was higher only near a stream bank where Dyduch-Falniowska &
and with the total snail abundance (r 2 ¼ 0.44). Habitat fertility Tobis (1989) recorded 40 ind./m2 for M. tumida. Density of Vestia
turgida populations in the Carpathian beech forests was usually ca
of the habitats (PC I) also significantly favoured high abundance
10 ind./m2 (Dyduch-Falniowska & Tobis, 1989; Dzieczkowski,
of both studied species but density of M. tumida was more strongly
1972) and was rarely higher: 30 ind./m2 near stream banks
associated with PC I (r 2 ¼ 0.62) than the density of V. turgida
(Dyduch-Falniowska & Tobis, 1989). These data correspond
(r 2 ¼ 0.37) (Table 3). Other PCs showed no significant well with the results of this study.
associations, but PC IV showed quite substantial, but opposite, Because of the habitat complexity it is difficult to single out
trends in the two species. Macrogastra tumida was not found at the which factors play a crucial role in controlling molluscs
least humid station (Stn 7), and was most abundant at stations 8 distribution and abundance. Many parameters involved are in
and 9, where PC IV reached higher values. a functional relation and are correlated at sampling stations: e.g.
In the multiple regression analyses the number of plant species soil pH and calcium content (positively); herb cover, soil pH
and moisture related best to density of M. tumida (Table 4). The (positively); and all the above and C/N ratio (negatively); herb
highest density of M. tumida populations were at the sites with cover and plant species richness (positively); hydrophilous plant
very diverse flora (more than 40 plant species/100 m2 of richness and steepness of slope (negatively).
mountain forests), which also contain a significant number of Relations between abundance of land snails and environmen-
hydrophilous plants (Fig. 4). tal factors have been investigated with the help of multi-
For V. turgida, high density of populations could be expected at dimensional statistical analysis (Barker & Mayhill, 1999; Cowie
sites with dense herb cover, cool microclimate but not very damp et al., 1995; Dillon, 1980; Tattersfield et al., 2001), but they have
(Table 5). It was less abundant at temporary submerged alder not proceeded to multiple regression using real environmental
swamp (Stn 9) than at sites of moderate moisture. Within the factors. This study is the first to do so. Since the number of

Figure 3. Principal component analysis: ordering of sampling stations (1–11) based on environmental factors.

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 HABITAT CHOICE IN CARPATHIAN CLAUSILIIDAE

Table 4. Results of multiple regression analysis.

Environmental factor B SE B beta Partial correlation T P

PLANT 0.095 0.022 0.683 0.840 4.373 0.0024

HUM 3.285 1.414 0.363 0.633 2.323 0.0487
Intercept 213.326 4.699 2 2.836 0.0219
R ¼ 0.927 R 2 ¼ 0.859
Analysis of variance
Source of variation df Sum of squares Mean square F P
Regression 2 18.2207 9.1104 24.31776 0.0004
Residual 8 2.9971 0.3746

The regression of Macrogastra tumida densities on the environmental factors. PLANT, HUM, environmental factors (see explanation in text); B, regression
coefficient; SE B, standard error of B; beta, standardized regression coefficient; T, F, test values; P, significance level; R2, coefficient of multiple determination;
df, degrees of freedom.

Table 5. Results of multiple regression analysis.

Environmental factor B SE B beta Partial correlation T P

HERB 0.180 0.057 0.697 0.766 3.155 0.0160

ASP 2 4.122 1.150 2 0.679 2 0.804 2 3.584 0.0089
HUM 214.257 5.888 2 0.546 2 0.675 2 2.421 0.0460
Intercept 40.399 18.834 2.145 0.0691
R ¼ 0.873 R 2 ¼ 0.762
Analysis of variance
Source of variation df Sum of squares Mean square F P
Regression 3 134.516 44.839 7.4678 0.0138
Residual 7 42.0296 6.004

The regression of Vestia turgida densities on the environmental factors. HERB, ASP, HUM, environmental factors (see explanation in text); B, regression
coefficient; SE B, standard error of B; beta, standardized regression coefficient; T, F, test values; P, significance level; R 2, coefficient of multiple determination;
df, degrees of freedom.

sampling sites was not very high, the results of the analysis should
be interpreted with much care. Nevertheless, they are concordant
with many data on ecology of land snails collected throughout the
world.
Some differences were found in moisture preferences of both
species, with M. tumida being more hydrophilous. Contrary to
its ecological description published by Kerney et al. (1983),
V. turgida had a quite wide moisture tolerance. This result is
in concordance with the remarks of Urbański (1939). He
claimed that V. turgida did not inhabit very damp places,
where it was usually replaced by Vestia gulo (E.A. Bielz, 1859).
Densities of both species studied were best correlated with
number of plant species (PLANT) at the sampling stations. At
least two causes for this relationship can be considered. First,
natural forests are a mosaic of microhabitats, which differ in
terms of moisture, shade and soil properties, providing suitable
conditions for many species of plants. The diversity of flora may
control fauna diversity or both are similarly influenced by
climatic and edaphic factors. Plant diversity (which is easier to
assess than animal diversity) can be applied as a user-friendly
indicator of overall habitat diversity. Secondly, the importance of
dead plants (especially palatable herbs) as the food source for
most snails can be stressed. The correlation between plant
diversity and mollusc richness is seldom discussed in malaco-
logical papers (Barker & Mayhill, 1999). More often, the view is
that the physical nature of the habitat, rather than floristic

Figure 4. Relationship between DENSITY of Macrogastra tumida on sampling stations (1–11) and environmental factors.

109
 A. SULIKOWSKA-DROZDA. SULIKOWSKA-DROZD
composition, is most important (e.g. Bishop, 1977). Peake (1978)
claimed that there was no conclusive evidence that the
proportion of palatable vegetation is a limiting factor for snails.
Later Proschwitz (1993) criticized this attitude, suggesting that
the composition of vegetation, which took part in litter formation,
was highly significant. He noticed that the number of coexisting
vertiginid species was the highest where several species of
deciduous trees contributed to the litter content. Since leaves of
different tree species are processed at different rates, higher tree
species diversity provided more stable food supply over time for
snails (Getz & Uetz, 1994). According to Barker & Mayhill
(1999), mollusc abundance may also be regulated by niche
availability, and this with increasing floral richness via site
successional development and higher microhabitat
differentiation.
However, at the sampling sites, ecological factors such as
calcium content in leaf litter and soil pH were positively
correlated with number of plant species. The highest population
densities were recorded at the sites with the highest pH and
calcium content. Thus, it is possible that these factors (and not
directly plant diversity) favour high abundance of clausiliids. In
the Bieszczady Mountains soils are mostly decalcified with the
exception of places of groundwater discharge, which have a pH
higher than 5.5 (Skiba et al., 1998). Consequently, advantageous
factors (humidity and low acidity) operate in the same places. It
is clear that a separation of these various factors requires detailed
studies in areas in which one or more of the variables remains
relatively constant, while others vary.
It is supposed that the litter and bacteria decomposing dead
wood are the main component of the diet of clausiliids (Fog, 1979).
Thus, decomposition rate may affect the abundance of snails.
Differences in consumption of different kinds of litter are related to
the C/N ratio in fallen leaves (Ghilarov, 1971). Leaves of Alnus,
which contain a lot of nitrogen, are mineralized very rapidly and
are a rich food source for saprophagous soil animals (Zimka &
Stachurski, 1976). High abundance of clausiliids, and of snails
generally, in alder woods is probably a joint effect of favourable
moisture, low acidity and high decomposition rate. Alder woods,
therefore, stand out in an otherwise oligotrophic area, but in
calcareous areas, e.g. the Pieniny Mountains further west in the
Carpathian chain, mixed woods on well-drained soils have very
rich clausiliid faunas (Urbański, 1939; Dzieczkowski, 1972). On
the contrary, beech leaves have a high C/N ratio and decompose
very slowly, especially in places with a short vegetation season (e.g.
at higher altitude). The paucity of snail populations in these sites
can be attributed to the low value of beech litter as a food source
(see Mason, 1970). Conversely, Cameron (1973) found highest
diversities in beech woods with least field layer vegetation, but this
related to highly calcareous sites, in which the relative unpalat-
ability of beech litter was mitigated. The properties of sycamore
litter as a food lay between these two extremes. The chemical form
of calcium in leaf litter may also be of importance (Wäreborn,
1970).
ACKNOWLEDGEMENTS
I thank Professor A. Piechocki, the supervisor of my PhD project,
for helpful comments and support and Dr Z. Wojciechowski
(University of Lodz) for advice on statistical analyses. I am also
indebted to referees for comments on the manuscript. The project
has been supported by grant from Polish State Committee for
Scientific Research.
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 APPENDIX

Station number 1 2 3 4 5 6 7 8 9 10 11

Locality Wielka Suche Połonina Wielka Wielka Czoło Połonina Dolina Dolina Bereżki Wielka Rawka
Rawka Rzeki Caryńska Rawka Rawka Caryńska Rzeczycy Rzeczycy
Altitude above sea level (m) 800 725 1060 730 1175 1120 930 665 670 650 1250
Slope aspect SE N S SSE NE N ENE SE E NE NE
Slope angle 58 58 158 108 258 108 408 18 28 108 158
Relief form convex concave convex convex convex convex convex concave concave convex concave
Average annual temp. ( 8C) 5.3 4.7 4.0 5.6 3.2 3.4 4.3 5.1 5.1 5.6 2.8

Plant community Class Querco-Fagetea Betulo-

A. SULIKOWSKA-DROZDA. SULIKOWSKA-DROZD
Adenostyletea

Alliance Fagion silvaticae Alno-Padion Secondary forest

with Alnus incana
Suballiance Eu-Fagion Acerion pseudoplatani

Association Dentario glandulosae-Fagetum Aceri-Fagetum Lunario-Aceretum Alnetum incanae Caltho-Alnetum Pulmonario-

carpathicum Alnetum viridis
typicum lunarietosum allietosum festucetosum typicum allietosum

Canopy cover 80% 90% 80% 95% 75% 80% 65% 70% 70% 85% –
112

Shrub cover 40% 30% 15% 15% 5% 50% 20% 20% 25% 15% 40%
Herb cover 60% 60% 65% 80% 65% 90% 80% 100% 100% 95% 97%
No. of plant species 28 28 20 16 27 42 29 50 41 41 39
Soil moisture index 3.44 3.53 3.48 3.52 3.55 3.70 3.22 3.58 3.86 3.62 3.64
Organic matter content (%) 14.1 8.15 14.77 11.19 15.13 16.68 13.13 9.67 36.6 19.57 17.2
Soil Ph H2O 3.6 5.0 3.6 4.0 4.0 5.0 3.7 6.2 5.4 4.5 5.2
Soil porosity (%) 68.85 69.7 69.24 68.53 74.3 65.2 73.17 64.9 82.83 77.17 86.3
C/N in leaf litter 35.32 33.37 33.65 30.68 21.9 32.43 28.92 23.62 17.54 18.14 24.65
Calcium in leaf litter (mg/dm3) 295 500 235 285 310 400 400 690 935 605 550
No. of species
Terrestrial Gastropoda 22 18 6 15 11 23 24 31 19 28 14
(spring)
Terrestrial Gastropoda 20 21 7 13 11 24 21 32 18 27 18
(late summer)
Density (ind./m2)
Terrestrial Gastropoda 130 119 18 118 28 304 283 546 1069 296 265
(spring)
Terrestrial Gastropoda 184 242 24 108 35 670 233 800 871 340 275
(late summer)
M. tumida (spring) 4 0 0 0 1 16 0 34 40 4 9
M. tumida (late summer) 3 2 0 0 0 27 0 49 36 6 1
V. turgida (spring) 0 6 0 0 1 11 0 7 3 9 4
V. turgida (late summer) 0 1 1 0 3 49 10 24 10 6 9