.
LAND MOLLUSC FAUNAS OF BIA L OWIEZ A FOREST (POLAND),
AND THE CHARACTER AND SURVIVAL OF FOREST FAUNAS
IN THE NORTH EUROPEAN PLAIN
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R. A. D. CAMERON 1 AND B. M. POKRYSZKO 2
1
Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, UK and Department of Zoology, The Natural History Museum,
London SW7 5BD, UK; 2Museum of Natural History, Wrocl aw University, Sienkiewicza 21, 50-335 Wrocl aw, Poland
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(Received 10 March 2003; accepted 26 August 2003)
ABSTRACT
The land mollusc faunas of three plant associations in the Bial owiez.a Forest, Poland, were sampled in
2001/2. Overall 51 species were recorded, of which 45 were found in Circaeo-Alnetum (CA; floodplain
forest), 38 in Tilio-Carpinetum (TC; mixed forests on eutrophic soils), and 35 species in MelittoCarpinetum (MC; mixed forests on meso- to oligotrophic soils). CA forests hold a number of wetland
species but, at site level, their faunas are slightly poorer and more variable than those in TC, which are
rich and uniform. MC faunas are poorer, and many species occur there infrequently and in low numbers. Taken with earlier surveys in the forest, these results give a picture of the undisturbed fauna of
forests off limestone on the North European Plain. They are compared with those from other lowland
forests in Poland, and more widely across northern Europe. While this forest fauna is now confined to
small areas, it appears to have survived intact in sites not subject to gross pollution or disturbance.
There are changes in species composition across the Plain, discussed in terms of historical biogeography. A comparison is also made with tropical forest; where there is a similar pattern of Holocene
habitat change, the faunas are remarkably similar in many aspects of diversity.
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INTRODUCTION
The Bial owiez.a Forest is famous as the largest surviving relic of
the original forests of the North European plain (Faliński,
1994). Although certainly exploited by humans and thus not
completely free of disturbance, the evidence from detailed
botanical studies is that parts of this forest preserve substantially
natural structure and composition. Furthermore, there is a wide
range of forest types present at Bial owiez.a, representing much
of the range of forests originally found over lowland northern
Europe. The flora and fauna of this forest therefore can give us
an indication of the nature of the original communities of the
North European Plain prior to human disturbance, and a yardstick to assess their survival elsewhere. Such studies have been
successful in the case of birds (Tomial ojć & Wesol owski, in
press), beetles (Gutowski, 1986; Wanat, 1993; Buchholz &
Ossowska, 1998), flies (Bańkowska, 1995,1997), Heteroptera
(Gorczyca, 1999) and many other taxa (an extensive bibliography is given by Gutowski & Jaroszewicz, 2001).
The native land mollusc fauna of the lowland areas of north
and central Europe consists mainly of forest and wetland species
(Boycott, 1934; Kerney, Cameron & Jungbluth, 1983). While
many species are tolerant of moderate levels of disturbance, the
richest surviving faunas at local level tend to occur in relatively
unexploited rocky montane forests, and especially in areas
of exposed limestone. Pre-Neolithic, mid-Holocene subfossil
assemblages from such areas show that pre-disturbance faunas
can survive more-or-less intact (Evans, 1972; Wiktor, 1974; Alexandrowicz, 1997). In lowland areas away from limestone, where
such subfossil evidence is lacking, surviving forest fragments
tend to occur on the least fertile soils, and many have very poor
mollusc faunas. Anthropogenic disturbance has a greater effect
in such cases (Waldén, Gärdenfors & Wäreborn, 1991). The
fauna of Bial owiez.a can thus be used to determine the degree of
degradation elsewhere in northern Europe, and to give an
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Correspondence: R. A. D. Cameron; e-mail: [email protected]
J. Moll. Stud. (2004) 70: 149–164
example of a natural lowland forest fauna to compare with those
from other biogeographical regions.
Dyduch (1980) has already given an inventory of land molluscs for the forest, based mainly on studies within the strict
nature reserve within the forest (see below). Dzie˛czkowski
(1988) also provides details for a single site within the reserve. In
this paper we report on a study of the molluscan faunas of three
of the principal plant associations within an adjacent area of
the forest. We use these results, and those of Dyduch and
Dzie˛czkowski, to make comparisons with lowland forest faunas
in northern Europe and elsewhere, and to examine biogeographical and anthropogenic explanations for the differences
and similarities observed.
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THE BIA L OWIEZ A FOREST
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The Bial owiez.a Forest lies in the middle of the North European
Plain, astride the border of Poland and Belarus (Fig. 1), just
south of 53°N. The forested area is a more or less continuous
block of c. 1,100 km2, of which 580 km2 lie within Poland. The
range of elevation is very slight (134–202 m a.s.l.). This topography is a product of Pleistocene glacial and periglacial processes, which have given rise to a variety of soil types ranging
from extreme podsols through a variety of brown earths to gleys
and other wetland soils. Palaeobotanical evidence suggests that
forests similar to those seen today started to form about 7,500 BP
and were initially more completely deciduous than they are now
(Faliński, 1994). The subsequent increase in conifers may be
due both to progressive leaching, and to human intervention.
Forest plant associations recognized within the area range
from communities associated with swamps, through eutrophic
and mesotrophic forests of various kinds to nearly pure stands of
pine on the poorest and most well-drained soils (Faliński, 1994;
Kwiatkowski, 1994).
Although the evidence suggests that there has been forest
cover continuously for many thousands of years, most if not all
of the forest has been subject to human disturbance. At the
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© The Malacological Society of London 2004
R. A. D. CAMERON & B. M. POKRYSZKO
(1) Tilio-Carpinetum (TC). This is the largest and most variable forest association on eutrophic soils in the area. A
number of subdivisions are recognized (Kwiatkowski,
1994). Our sites fall mainly into the typical variety of this
forest type; the community consists of mixed-deciduous
forest containing hornbeam Carpinus betulus, linden Tilia
cordata, oak Quercus robur and other deciduous species
interspersed with spruce Picea abies. These forests occur in
areas where the surface is slightly above the water-table.
Soils are typically eutrophic, brown earth, often underlain
by marly till.
(2) Circaeo-Alnetum (CA). This association is typical of forested
flood plain areas, and its structure and composition is
heavily dependent on seasonal flooding. Soils are typically
gleys and are usually rich in nutrients. The dominant trees
extreme this has resulted either in total clearance or in the
establishment of regularly harvested monocultures. However,
both in the strict nature reserve (47 km2) and in the area immediately around it (Fig. 1) there are patches in which substantially
unmodified natural communities can be recognized. As Dyduch
(1980) had already surveyed the strict nature reserve, we concentrated our sampling effort on areas close to the reserve in
which it was possible to identify the plant communities concerned.
HABITATS SAMPLED
Figure 1 shows the sample locations. The sites fell into three of
the commonest and most easily recognized plant associations
(Kwiatkowski, 1994) of meso- to-eutrophic soils:
Figure 1. A map showing the locations of study sites within Bial owiez.a Forest. Nature Reserves shown stippled, built-up areas with horizontal hatching. Inverted
triangles, sites in Circaeo-Alnetum; circles, sites in Tilio-Carpinetum; squares, sites in Melitto-Carpinetum. Dashed lines denote roads and tracks, solid lines,
rivers and streams. The inset locates the region within Poland in which the study area is situated (B).
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150
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B IA L OWIEZ A FOREST FAUNAS
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1
2
3
4
5
6
7
8
9
10
1
2
3
4
5
6
7
8
9
20
1
2
3
4
5
6
7
8
9
30
1
2
3
4
5
6
7
8
9
40
1
2
3
4
5
6
7
8
9
50
1
2
3
4
5
6
7
8
9
60
1
2
3
4
5
6
7
8
These recorded levels of species richness are subject to
sampling error; some species will be missed. This will increase
the apparent differences between sites and habitats. Other
things being equal, this error will depend on sample size, and on
the frequency distribution of individuals amongst species
(Gotelli & Colwell, 2001). Our sampling method does not give
true relative abundances, and it is thus not valid to fit the data to
frequency distribution models. Nevertheless, it is notable that,
in the aggregated data for CA and TC sites, the rank- log abundance relationships tend to be sigmoid, with a downturn
amongst the rarest species (Fig. 2). In relation to the log normal
distribution, this suggests that the sampling ‘veil line’ is excluding only a very small number of very rare species (Magurran,
1988; Southwood & Henderson, 2000). In contrast, the fauna of
MC forest appears to be dominated by a few abundant species,
with a long ‘tail’ of rarities.
Amongst the non-parametric tests for the completeness of
inventories, the Chao estimators are regarded as the most reliable (Colwell & Coddington, 1995; Southwood & Henderson,
2000). Table 2 shows estimates of undiscovered species for each
habitat from Chao 1 (based on numbers) and Chao 2 (based on
frequency). These estimators have very large standard errors,
and any genuine heterogeneity between sites in each category
will distort the estimators upwards. Hence for Chao 1, at least,
they suggest that habitat inventories are close to being complete.
are alder Alnus glutinosa and ash Fraxinus excelsior, with
smaller numbers of many other deciduous trees as well as
spruce. All our sites with significant amounts of alder
appeared to fit inside the CA but some might be classified
in the category Ficario-Ulmetum (Kwiatkowski, 1994), the
composition and properties of which are very similar. The
range of moisture regimes in these forests was large; sites 12
and 13 appeared to be wetter than the remainder, and site
16 to be drier.
(3) Melitto-Carpinetum (MC). This very widespread association is typical of the drier parts of the morainic plateau.
Originally the dominant species was oak Quercus robur, with
significant amounts of pine Pinus sylvestris, spruce Picea abies
and other trees, including birches Betula spp. Soils are
typically leached or slightly podsolized brown soils and, in
the least nutrient-rich areas, there are plants typical of
markedly acid soils, such as Vaccinium myrtillus, found abundantly at site 1. Our sites had mostly been subject to some
forestry activities, with the majority of felled trees being
spruce and with considerable evidence of birch and oak
regeneration. Because of the disturbance it is possible that
some of our sites might be classified within the categories
Querco-Pinetum or Pino-Quercetum (Kwiatkowski, 1994),
which are typically associated with similar soil types and
herbaceous plants.
Species composition: presence and absence data
METHODS
Table 3 summarizes the frequency of occurrence of species by
habitat and overall. Seven species are universal, recorded from
all sites, while a further eight were only recorded in single sites.
A far larger number of species are constant within TC forests
than in MC; CA is intermediate.
Sampling was carried out in September 2001 and August 2002.
Each site was an area of c. 400 m2, chosen to represent a typical
part of the surrounding vegetation, avoiding edges or transitions. Within each site snails and slugs were collected by eye.
Two people searched in this way, in all appropriate microhabitats, for 1 h at each site. In addition, c. 10 l of litter was
collected from patches within each site. The litter was sieved
(10 10 mm mesh); the coarsest fraction was searched by eye in
the field and discarded. The remainder was bagged and taken to
the laboratory, where it was air-dried, sieved and sorted down
to 0.5 mm mesh. Large helicoids were sometimes counted but
not collected; slugs were identified in the field or specimens
preserved for identification in the laboratory. No attempt was
made to count the number of slugs, as daytime sampling in dry
weather is notoriously unreliable (Wäreborn, 1969).
All shells extracted were identified and counted, excluding
only very old or unidentifiable remains. Nomenclature throughout follows Kerney et al. (1983). Authorities are given in
Appendix 1. The samples are kept in the collection of RADC.
Table 1. Data on numbers of samples, species and individual specimens
obtained in the study, together with the estimates of Whittaker’s Index
(S/), and Imax, S/richness of richest site in the category concerned.
Habitat
MC
No. of sites
RESULTS
Species richness
A total of 51 species (45 snails and six slugs) were recorded in
the study. Appendix 2 shows the numbers of each snail species
recorded at each site, and the occurrence of slugs. Table 1 summarizes data on species richness and sample size by habitat type
and overall. No single site or habitat contained all the species
recorded. CA forests collectively have the largest number of
species, and MC forests the least, but TC forest sites are, on
average, marginally richer than those from CA. Table 1 shows
Whittaker’s index, S/, and its variant S/max (Southwood &
Henderson, 2000) for each habitat and overall. On both
measures, TC forests are the most uniform in composition, and
MC the most heterogeneous. Overall heterogeneity is greater
than that within habitats, indicating a differential distribution of
species (see below), but a majority of species are found in all
three habitats.
TC
CA
Total
5
5
7
17
Total species, S
35
38
45
51
Mean species/site, 21.4
31.0
28.1
27.0
SE
1.3
1.4
0.7
1.1
Range
17–24
28–35
26–31
17–35
Whittaker’s I
1.64
1.23
1.60
1.89
Imax
1.46
1.09
1.45
1.46
Slug species
4
3
4
6
Snail species
31
35
41
45
No. of snail specimens
846
1484
2363
4693
Mean specimens/site
169
297
338
276
Range
126–236
165–499
180–725
126–725
Table 2. Chao estimates of the number of undetected species in each habitat
(see text).
Chao 1
Habitat
Singles
Chao 2
Doubles
Estimate
Singles
Doubles
Estimate
4.9
MC
2
4
0.5
7
10
CA
3
2
2.2
9
6
6.7
TC
1
1
0.5
1
8
0.06
For Chao 1, singles number of species represented by one individual, doubles,
number represented by two individuals. For Chao 2, singles number of species
recorded from only one site, doubles, number recorded in only two sites.
151
R. A. D. CAMERON & B. M. POKRYSZKO
Figure 2. Logarithmic rank-abundance relationships for snails in each of the three habitat types. The vertical scale is expressed as log10 percentage of the total.
Inverted triangles, sites in Circaeo-Alnetum; circles, sites in Tilio-Carpinetum; squares, sites in Melitto-Carpinetum. See text for further details.
All of the 13 species found only in one habitat are rare;
eight were found only in one site each, and the remaining five in
two sites each. In CA forests, five of the species concerned,
Succinea oblonga, Cochlicopa nitens, Vallonia pulchella, Deroceras
laeve and Perforatella rubiginosa, are known to have wetland
associations, as do three of the four species universal only in
CA, Carychium minimum, Succinea putris and Zonitoides nitidus.
While the absence of Malacolimax tenellus in CA seems to be a
genuine habitat effect, the distribution of the remaining species
missing from at least one habitat may be largely due to sampling
error. Two of the three species unique to MC, Vertigo alpestris
and Cepaea hortensis, come from the same site (15). It was
more disturbed than others, but had no obviously distinctive
features.
Similarity in composition between sites has been examined
using the Jaccard Index (Southwood & Henderson, 2000).
Figure 3 shows the dendrogram of affinities between sites, and
Table 4 the mean values for within and between habitat comparisons. Sites in TC, and in CA, cluster together with only one ‘mismatch’ (site 16). The MC sites do not do this; apart from one
pair (10 and 14), they attach individually to larger groupings of
other sites, and at low levels of affinity. Overall (Table 4), they
have marginally greater affinity to the TC sites than to those
from CA or amongst themselves.
Of the two outlying sites, 1 has much the poorest fauna in the
whole array and one unique species, while 15 has two unique
species. Unique species can contribute only to difference, and
variation in species richness reduces the maximum value the
index can attain. Table 5 shows aggregate between habitat
Jaccard Indices with and without unique species, and as modified by considering the species in common as a proportion of
the species in the poorer habitat only. This evidence indicates
that the fauna of MC forests is simply a reduced version of the
others, but it does not distinguish between real impoverishment
Figure 3. Dendrogram of Jaccard affinities between sites (UPGMA). Habitat
symbols as in Figure 1.
and a sampling artefact. Allowing for variations in recorded
species richness, all the sites have very similar faunas.
Species composition: quantitative data
On presence and absence data alone, MC forest faunas seem to
be merely impoverished versions of the others. Inspection of the
quantitative data in Appendix 2 shows that this is not the whole
152
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B IA L OWIEZ A FOREST FAUNAS
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1
2
3
4
5
6
7
8
9
10
1
2
3
4
5
6
7
8
9
20
1
2
3
4
5
6
7
8
9
30
1
2
3
4
5
6
7
8
9
40
1
2
3
4
5
6
7
8
9
50
1
2
3
4
5
6
7
8
9
60
1
2
3
4
5
6
7
8
Table 3. A. Species occurring in all sites, overall and in each habitat. B. Species missing in one or more habitats, and those
not universal in any habitat.
A
Occurrence
Melitto-Carpinetum
Universal
Tilio-Carpinetum
Circaeo-Alnetum
Punctum pygmaeum, Nesovitrea hammonis, N. petronella, Euconulus fulvus,
Cochlodina laminata, Macrogastra plicatula, Arion subfuscus
Universal
Discus ruderatus, D. rotundatus, Limax cinereoniger
in habitats indicated
Cochlicopa lubrica, Vitrea crystallina, Aegopinella pura,
Bradybaena fruticum, Perforatella bidentata
Carychium tridentatum,
Carichium minimum,
Vertigo pusilla,
Succinea putris,
Acanthinula aculeata,
Columella edentula,
Vitrina pellucida,
Zonitoides nitidus
Cochlodina orthostoma,
Macrogastra ventricosa,
Laciniaria plicata,
Bulgarica cana,
Perforatella vicina,
Malacolimax tenellus
B
No. of universal species
10
25
16
Occurrence
Melitto-Carpinetum
Only in habitats
Vertigo alpestris, 1
Macrogastra tumida, 2
Succinea oblonga, 2
indicated
Cepaea hortensis, 1
Clausilia dubia, 2
Cochlicopa nitens, 1
Lehmannia nyctelia, 1
.
Vallonia pulchella, 1
.
.
Clausilia bidentata, 1
.
.
Ruthenica filograna,
.
.
Perforatella rubiginosa, 1
.
.
Arion fasciatus, 2
.
Deroceras laeve, 2
Tilio-Carpinetum
.
Circaeo-Alnetum
Malacolimax tenellus, 7
Acicula polita, 4; Cochlicopa lubricella, 4;
Macrogastra ventricosa, 7; Clausilia pumila, 4;
Trichia hispida, 4; Isognomostoma isognomostoma, 2
Helix pomatia, 2
Present in all,
Absent
Helix pomatia, 2
Vertigo substriata, 11; Aegopinella minor, 7.
universal in none
The number after each species indicates the number of sites overall in which it was found.
Table 5. Values of the Jaccard Index, and for the proportion of species in
common relative to the least rich habitat, for comparisons between the total
faunas of each habitat.
Table 4. Mean values ( SE) of the Jaccard Index within and between habitats, on a site-by-site basis.
Habitat
MC
TC
CA
MC
54.5 2.04
58.2 1.62
52.7 1.50
TC
.
78.9 1.96
63.4 0.96
CA
.
.
67.8 1.30
Jaccard (%)
Joint/least rich (%)
MC
TC
MC
TC
CA
64.5
74.5
88.6
92.1
MC
.
69.8
.
85.7
CA
75.6
81.4
88.6
92.1
MC
.
75.9
.
93.8
All
Without uniques
story. Despite much variation in numbers between sites both
within and amongst habitats, some species are at a maximum
in MC in both relative and absolute terms. Figure 4 shows the
relative abundances (based on mean numbers per site) of the
23 most abundant species of snail (more than 50 individuals
collected). Nesovitrea petronella scarcely discriminates between
habitats at all; Nesovitrea hammonis and Euconulus fulvus are more
abundant in MC than elsewhere; and, most notably, the two
species of Discus and Laciniaria plicata are more abundant in MC
than in CA. Conversely, there is a suite of species which are
much more abundant in CA than elsewhere, including extreme
Values are given with and without the inclusion of species found only at one site
(uniques).
cases such as Succinea putris and Carychium minimum. Such
extremes are missing from the TC pole, but only the two cases
mentioned above give values of more than 10% in TC: contrast
this with four cases for CA, and 11 for MC.
153
R. A. D. CAMERON & B. M. POKRYSZKO
gradient from poor to rich sites in which species are added as a
greater variety of microhabitats become available. No species
specializes in the poorer sites, although some (e.g. Discus
species) avoid wetland. The positive correlations between abundances of congeneric species (Discus, Nesovitrea) show conditions between sites varying in general suitability for similar
molluscs, rather than offering a distinctive range of niches.
There is, however, a residue of variation between sites and
habitats that cannot be ascribed to known environmental differences. Sampling is bound to be incomplete, and sampling error
will be confounded with genuine differences between sites. TC
and CA sites have rank-abundance distributions that resemble
‘Broken Stick’ or log-normal models (Southwood & Henderson, 2000). In such cases, around 10 times as many specimens as
species need to be seen to record more than 95% of the fauna
(R. A. D. Cameron & B. M. Pokryszko, in preparation). MC
samples have a longer tail of rare species, and are probably
under-recorded; this ‘times ten’ rule is often not met in this
habitat, and the season and method of sampling will also affect
reliability (R. A. D. Cameron & B. M. Pokryszko, in preparation). Because numbers are larger, estimates of richness for
each habitat in total should be more reliable, since the number
of specimens is comfortably more than 10 times the species
recorded. Nevertheless, the effects of sampling error need to be
remembered when wider comparisons are made. On evidence
from elsewhere, for example, there is no reason to think that
Vertigo alpestris or Cepaea hortensis have an affinity for poorer soils.
Their restriction to a single MC site is a chance encounter
with very patchily distributed species. Both have been recorded
earlier from richer forests in the region.
Figure 4. The proportion of the total numbers of each of the 23 commonest
species (represented by more than 50 specimens) found in each habitat,
adjusted for the number of samples in each. Abbreviations: AA, Acanthinula
aculeata; AP, Aegopinella pura; BF, Bradybaena fruticum; CE, Columella edentula;
CL, Cochlicopa lubrica; CLA, Cochlodina laminata; CM, Carychium minimum;
CT, Carychium tridentatum; DRO, Discus rotundatus; DRU, Discus ruderatus;
EF, Euconulus fulvus; LP, Laciniaria plicata; MP, Macrogastra plicatula;
NH, Nesovitrea hammonis; NP, Nesovitrea petronella; PB, Perforatella bidentata;
PP, Punctum pygmaeum; PV, Perforatella vicina; SP, Succinea putris; VC, Vitrea
crystallina; VP, Vertigo pusilla; VS, Vertigo substriata; ZN, Zonitoides nitidus.
Comparisons with earlier work in Bial owiez.a
/
Dyduch (1980) gives a detailed account of the Bial owiez.a
mollusc fauna, incorporating earlier records. However, she
does not give lists for individual sites, nor the numbers collected
in each habitat she distinguishes. She does provide a figure
showing the proportional abundance of species in each of seven
habitat types. For purposes of comparison, we have combined
her types I–III (all Tilio-Carpinetum) to compare with our TC,
and her types IV and VI (both Alnetum) to compare with our
CA. Her type V (Querceto-Betulinum) is compared with our
MC. We have no comparison to make with data from her type
VII, which categorizes the highly modified forest of the Palace
Park, containing both occupied and ruined buildings. The
combination of some of her categories means that the spread of
environmental conditions within each is wider than ours, and it
is evident that she included data from forest edge, from open,
and from highly disturbed sites in some of her categories. Her
sampling methods are also likely to reduce the apparent abundance and frequency of some small litter-dwelling species (R. A.
D. Cameron & B. M. Pokryszko, in preparation)
Overall, Dyduch (1980) lists 55 species of which one, Vertigo
genesii, is an error (Pokryszko, 1998). Dzie˛czkowski (1988) adds
two, and Pokryszko (1998) three species to this list. It is probable
that at least three further species, Euconulus alderi (open wetland), Vertigo ronnebyensis and Columella aspera (very oligotrophic
forest) will be found (Pokryszko, 1998). Of these 59 species, 12
were not found by us, but seven of these are not typically forest
species (Table 6). We recorded four species for the first time
(Table 6). There are thus 63 species known from the area, of
which 56 are forest species; 47 of these were found both in our
survey and earlier (Jaccard Index, 84%). Although there are
some minor ambiguities in Dyduch’s account, it is also possible
to make comparisons habitat-by-habitat. In the TC/TilioCarpinetum comparison, there are 35 species in common compared with 45 recorded overall (Jaccard Index, 78%); in the
CA/Alnetum comparison there are 36 species in common
/
Amongst the most abundant species, there are three pairs
of congenerics (Carychium, Discus, Nesovitrea). In Carychium,
there is no sign of any association between their abundances
(r –0.12), but between Discus species (r 0.63, P < 0.01),
and between Nesovitrea species (r 0.58, P < 0.05), there are
significant positive associations. In the case of Discus, this is
largely due to the low abundance of both species in CA sites.
DISCUSSION
Ecological differences and sampling efficiency
There are differences in the abundance and occurrence of
species at our sites that reflect ecological differences between
sites and habitats. Many of the species confined to CA, and those
which are at maximum frequency and abundance there, have
known wetland affinities (Kerney et al., 1983; Riedel, 1988). The
dry and oligotrophic MC sites are species poor, and have low
overall abundances. This reduction in abundance and frequency
is not random or uniform; it affects most the larger, heavy
shelled clausiliids and helicoids. Some of the differences
between sites within habitats also have an ecological basis: site 1
was clearly the most oligotrophic of the MC sites, and has the
poorest fauna; the presence of Succinea putris in site 14 reflects
the presence of a small marsh within the site. Amongst CA sites,
site 16 appeared to be the driest (less Alnus, more Quercus), and
it associated with TC sites in the Jaccard dendrogram. In contrast, the TC sites show a remarkable uniformity in composition.
Overall, the faunas show considerable uniformity, with 29 out of
51 species occurring in all three habitats, albeit in varying frequencies and abundance.
Excepting the wetland element, these faunas fit the pattern
described by Waldén (1981) for Swedish forests: there is a
154
.
B IA L OWIEZ A FOREST FAUNAS
/
1
2
3
4
5
6
7
8
9
10
1
2
3
4
5
6
7
8
9
20
1
2
3
4
5
6
7
8
9
30
1
2
3
4
5
6
7
8
9
40
1
2
3
4
5
6
7
8
9
50
1
2
3
4
5
6
7
8
9
60
1
2
3
4
5
6
7
8
quantitative volume method used by us (Wäreborn, 1969; R. A.
D. Cameron & B. M. Pokryszko, in preparation), at least in terms
of effort. Areas searched by eye in some of these studies clearly
included open and wetland patches, and sample areas sometimes included more than one forest type.
We have selected a set of 31 sites from such studies for comparison, restricting them to those made in lowland forest, with
large numbers of quadrats sampled, not less than 100 specimens
retrieved (usually far more, ranging up to more than 4,000),
and spanning a similar range of forest types (thus excluding
swamp alder forests, for example). Figure 5 shows the general
distribution of these sites, and Appendix 3 gives sources.
Collectively, these sites contain 68 species; 14 of these are not
recorded from Bial owiez.a, which in turn contains nine species
not found in the others. Fifty-four species are held in common
(Jaccard, 70%), and the aggregate total of 77 species represents
77% of the known fauna of northern, lowland Poland (Riedel,
1988). This gross comparison underestimates the similarity of
specifically forest faunas, as both lists contain casually encountered species of open wetland and forest margins. Also some of
the species not held in common have geographical ranges that
exclude them from one or the other series (Table 7). Bial owiez.a
lacks some western species, but adds a number with eastern
or southern/montane distributions. Accounts of early 20th
century surveys suggest that some eastern species (e.g. Discus
ruderatus) have retreated from western sites in recent times
(Drozdowski, 1979, 1988). If the geographically impossible
species (and Vertigo moulinsiana) are excluded, the Jaccard
Index rises to 79%, a degree of similarity not much less than that
recorded between successive surveys within Bial owiez.a alone.
Table 6. A. Species not found by us, and omitted from comparison between
surveys. B. Forest species not found by us. C. New records of forest species
(see text).
Species
Comments
A. Species not recorded in our study, but not usually found in closed forest
Vertigo moulinsiana
Open wetland (Pokryszko, 1998)
Vertigo antivertigo
Open wetland (Pokryszko, 1998)
Vertigo pygmaea
Open habitats (Pokryszko, 1998)
Vertigo angustior
Open wetland margins (Pokryszko, 1998)
Deroceras reticulatum
Found only once in very anthropogenic site
(Dyduch, 1980)
Oxyloma elegans
Only in open wetland (Dyduch, 1980)
Pupilla muscorum
Only on ruins in Palace Park (Dyduch, 1980)
/
B. Forest species not found by us
Ena obscura
One site only (Dyduch, 1980)
Arion silvaticus
In one site only (Dyduch, 1980) (an unidentifiable
juvenile, possibly this species, was found by us)
Macrogastra latestriata
Three sites (Dyduch, 1980; Dzie˛czkowski, 1988)
Perforatella incarnata
Recorded only by Dzie˛czkowski (1988)
Euomphalia strigella
Two sites (Dyduch, 1980; Dzie˛czkowski, 1988)
/
C. Species reported for the first time here
Macrogastra tumida
Recorded from 2 sites
Clausilia bidentata
Only a single specimen from 1 site
Deroceras laeve
Recorded from 1 site
Succinea oblonga
Recorded from 2 sites
/
Table 7. Species unique to other forest sites (Appendix 3) or to Bial owiez.a,
arranged by apparent cause of exclusion.
/
out of a total of 47 (Jaccard Index, 76%), and for the MC/
Quercetum comparison, 25 species in common out of 40
(Jaccard Index, 62%). These values of the Index are lower than
that for the overall comparison, especially for MC. In general,
the species missed, in one or the other survey, are rare and infrequent or, in the case of absences from Dyduch’s list for
Quercetum, are small, litter-dwelling species. It is also possible
to compare Dzie˛czkowski’s (1988) list for an area (mainly TilioCarpinetum) within the reserve with our TC: Jaccard Index
71%.
Although these comparisons show a high degree of consistency, the differences between surveys, habitats and sites suggest
that a number of the rarer species are distributed only patchily
within the forest. As indicated above, there is a problem in discriminating between genuine patchiness and the consequences
of sampling error, which deserves further detailed study.
Dzie˛czkowski’s samples, and most of Dyduch’s, were made in
the strict nature reserve, in which we did not sample. The high
level of overall similarity in the results of the two surveys gives
some reassurance about the status of molluscan habitats outside
the reserve, at least in areas of richer soils. We did not attempt to
sample the substantial areas outside the reserve that gave the
impression of being spruce or pine monocultures.
.
Species present in others, but not Bial owieza
/
Aegopinella nitidula
Forest species excluded by geographical
Aegopinella nitens
distribution (Riedel, 1988)
Arion rufus
Arion intermedius
Truncatellina costulata
Non-forest species unlikely for reasons
Chondrula tridens
of habitat or geography (Riedel, 1988)
Vallonia excentrica
Cepaea nemoralis
Vitrea contracta
.
Forest species for which Bial owieza is
Oxychilus alliarius
geographically marginal (Riedel, 1988)
/
Helicigona lapicida
Arianta arbustorum
No evident reason for absence
Deroceras agreste
Deroceras sturanyi
.
Species present in Bial owieza, but not in others
/
Comparison with other forest faunas in the Polish lowlands
Aegopinella minor
Montane species absent from western
Lehmannia nyctelia
and central lowlands (Riedel, 1988)
Macrogastra tumida
There are many published studies detailing the molluscan faunas of lowland forest sites across Poland. These often relate to
nature reserves, or to other sites believed to preserve elements
of natural forest vegetation. Typically, such studies refer to areas
of between a few hundred square metres and 50 ha. They
involve searching by eye, and the sampling of randomly placed
quadrats (20 20 or 25 25 cm), from which soil and litter
are removed for detailed examination (Oekland samples:
Dzie˛czkowski, 1988). While truly quantitative, such quadrat
sampling is less efficient for inventory work than the semi-
Perforatella vicina
Vertigo alpestris
Montane/boreal species generally rare in
Discus ruderatus
lowlands. May be retreating eastwards
Macrogastra latestriata
(Drozdowski, 1979, 1988)
Isognomostoma isognomostoma
Vertigo moulinsiana
Not a forest species, and very rare in Poland
(Pokryszko, 1990)
155
R. A. D. CAMERON & B. M. POKRYSZKO
Figure 5. The distribution of other forest study sites in northern Poland (see text). Site numbers refer to the list in Appendix 3.
The sites from elsewhere can be assigned to four habitat
categories, combining some of those used by Dzie˛czkowski
(1988), and corresponding as much as possible to those used by
us:
Table 8. Diversity data for 31 lowland forest sites in central and western
Poland, arranged by forest type (see text).
Habitat
A. Beech
(A) Typical beech forests, combining two categories both on
well-drained and mostly oligotrophic soils. These are most
comparable to our MC sites.
(B) Mixed deciduous forests, generally Carpineta, but with a
range of dominant deciduous species, including beech on
eutrophic soil. Soils are generally richer than in A, but not
flooded or gleyed. Together with type C, these overlap
our TC sites, but span a wider range of ecological conditions.
(C) Ficario-Ulmeta (Dzie˛czkowski, 1988), damper forests on
eutrophic soils, marking a transition between B and D
(below).
(D) Circaeo-Alneta, directly comparable to CA sites in
Bial owiez.a; eutrophic soils with seasonal flooding.
No. of sites
B. Mixed
C. Fic-Ulm
D. Circ-Aln
6
13
6
6
Total species
32
55
55
49
Mean species/site
19.5
26.4
27.3
28.3
SE
1.1
1.4
2.5
2.4
Range
16–23
19–36
19–36
19–36
Whittaker’s I
1.64
2.08
2.01
1.73
Imax
1.39
1.53
1.53
Snail species
Slug species
Jaccard
25
46
7
9
46% MC
48% TC
49
6
55% TC
1.36
41
8
62% CA
.
Jaccard comparisons are made with Bial owieza equivalents as indicated.
/
/
Table 8 shows diversity data for these forests by habitat category,
and Jaccard indices for appropriate comparisons with
Bial owiez.a. Comparison with Table 1 shows the remarkably
similar mean levels of species richness in comparable forest types
between studies. The greater heterogeneity (Whittaker’s I) of
B and C relative to Bial owiez.a TC reflects the wider range of
habitats and sizes of sample area represented, and the inclusion
of non-forest species. A similar range of I (1.7–2.0) is found in
other forest series from north and western Europe (Cameron,
1995). The low Jaccard index for the poorest habitat comparison (paralleled by the same comparison within Bial owiez.a,
noted above) is attributable both to greater sampling errors and
to more patchy distributions of the rarer species.
Broader geographical comparisons
There is a large literature on forest molluscan faunas across
northern, central and northwestern Europe. Surveys vary considerably in aims and methods, and in some cases the passage of
time and varying taxonomic opinions make direct comparisons
inexact. We have selected for comparison studies that seem to
contain complete accounts of forest faunas in habitats similar to
those encountered at Bial owiez.a (Table 9). Given the potential
sources of error, the figures for species in common should be
regarded as merely indicative.
A clear pattern emerges. Species richness varies rather little
between studies (taking account of the varying scales and ranges
/
/
/
/
156
.
B IA L OWIEZ A FOREST FAUNAS
/
1
2
3
4
5
6
7
8
9
10
1
2
3
4
5
6
7
8
9
20
1
2
3
4
5
6
7
8
9
30
1
2
3
4
5
6
7
8
9
40
1
2
3
4
5
6
7
8
9
50
1
2
3
4
5
6
7
8
9
60
1
2
3
4
5
6
7
8
Table 9. Similarities between the Bial owiez.a faunas and those from selected studies across northern Europe.
/
Distance
Species
Bialowieza
Species
Species in
Jaccard
(km)
listed
comparison
listed
common
(%)
Source
W/C Poland
250–600
68
All
63
54
70
This study
Valdai, W Russia, 1
800
45
All
63
40
59
Shikov, 1981
Valdai, W Russia, 2
800
50
All
63
43
61
Shikov, 1982
Rügen, NE Germany
700
48
TC, CA, MC
51
35
55
Körnig, 1980
Stockholm, Sweden
800
37
TC, MC, Dy, Dz
47
28
50
Waldén, 1954
Meadow, woods, S Sweden
700
41
TC, MC, Dy, Dz
47
28
46
Wäreborn, 1969
Hyperite hills, Finland
1100
31
TC, MC
43
22
42
Valovirta, 1968
Study
Netherlands, summary for forest
1200
49
All
63
33
41
Bruijns et al., 1959
C Scotland, summary for forest
1750
42
TC, MC Dy, Dz
47
24
35
Bishop, 1977a
NE Scotland, Alders
1750
38
CA
45
21
34
Cameron & Greenwood, 1992
C England, non-limestone woods
1700
44
All forest species
56
24
31
Tattersfield, 1990
C England, coal measures
1700
44
TC, MC
43
20
30
R. A. D. Cameron, unpublished
W Ireland, acid woodland
2100
42
TC, MC
43
18
27
Bishop, 1977b
Czech Republic alder
650
32
CA
45
22
40
Flasar, 1995
Hungary, Danube gallery forests
750
62
all
63
37
42
Bába, 1977
In each case, the most appropriate comparison has been made, relative to habitat. Dy, Dz indicates the inclusion of species recorded in TC and MC by Dyduch and
Dzie˛czkowski, but not by us. Note that variations in taxonomy and sampling methods make these figures indicative only.
in all these respects from faunas in older, and much more
oligotrophic, tropical forests, which have richer, but much
more locally variable faunas (de Winter & Gittenberger, 1998;
Schilthuisen & Rutjes, 2001). Evidence for the role of competition, or saturation, in determining richness is notoriously hard
to find (Cameron, Mylonas, Triantis et al., 2003). In this case,
similar topography and history have resulted, independently, in
two remarkably similar faunas.
of habitats involved), but there is distance decay in similarity
(Nekola & White, 1999) in all directions across the plain, or
towards its equivalents in the south (Hungary and the Czech
Republic). The striking similarity with forest faunas from western Russia emphasizes the transitional position of Bial owiez.a
between eastern and western sections of the plain in botanical
terms (Faliński, 1994). Over greater distances to the west,
similarity declines to 30% or less relative to sites in the British
Isles.
In a few cases, differences arise as a result of congeneric
replacement (e.g. in Acicula, Aegopinella, less often Clausilia),
but in general they are caused by geographical variation in the
richness of disparate groups. Thus the Bial owiez.a fauna is rich
in clausiliids (12 species), as are those of western Russia (11
species, 10 in common), poor in slugs, and it lacks Oxychilus
(Zonitidae) altogether. Clausiliids decline to the west, and
become more dependent on limestone substrates (Bruijns,
Altena and Butot, 1959), while slugs (especially Arion species)
and Oxychilus increase. The great majority of species found
throughout the plain are small litter-dwelling species in the genera Carychium, Columella, Vertigo, Acanthinula, Punctum, Vitrina,
Vitrea, Aegopinella, Nesovitrea and Euconulus.
These faunas have all developed by immigration during the
Holocene. They lack restricted endemic species and, although
there is distance decay it is much less rapid than that seen in
Mediterranean faunas, which are rich in endemics and survived
the Pleistocene in situ (Cameron, Mylonas & Vardinoyannis,
2000). Different sources, as much as any environmental differences across the plain, may account for the distance decay. In its
clausiliid fauna, and in the presence of Perforatella vicina,
Bial owiez.a shows strong affinities to the Carpathians, the nearest source of immigrants.
In this context, a comparison can be made with Tattersfield’s
(1996) study of Kakamega Forest in Kenya. It shares with
Bial owiez.a a subdued topography, a division into riverine
forests and others, and a Holocene origin, the area having been
open savannah in the late Pleistocene. While taxonomically very
different, the fauna is identical in recorded richness (51
species) to that of our study. Very similar sampling methods
were used, and the number of shells found is very similar. Rankabundance distributions are much the same, as is the overall
value of Whittaker’s I (2.1 in Kakamega, 1.9 here). Both differ
/
Human disturbance and the status of the lowland forest fauna
in northern Europe
The forest molluscan fauna of Bial owiez.a is rich and, in the
more eutrophic habitats at least, consistent from site to site. On
the evidence of other, arguably more sensitive groups (Gutowski & Jaroszewicz, 2001), it seems that the forest maintains the
full pre-disturbance faunas, despite spasmodic human intervention. While poorer than the faunas of montane forests on limestone further south (e.g. Pieniny: Urbański, 1939; Riedel, 1988),
it matches anything seen elsewhere on the northern plain where
there is no exposed limestone, and indeed preserves around
two-thirds of the fauna of all northern Poland (Riedel, 1988).
Comparisons across northern Europe show that while there
are rather few small sites that can rival the richest in Bial owiez.a,
many forest fragments on eutrophic and wetland soils contain
the great majority of the original fauna; collectively, perhaps all
of it. Mollusc faunas can survive more-or-less intact in very small
fragments of suitable habitat. This is not to minimize the consequences of disturbance, which has fragmented many distributions, and caused local extinctions. Planted forests tend to be
species-poor (Skrzypczak & Umiński, 1979); rarer, more stenotopic species find it hard to disperse across the fragmented landscape. Apart from complete clearance, the primary threats
come from acidification and drainage. While long-term acidification of soils of glacial origin may be a natural process, it is
greatly enhanced by conversion to conifer monocultures and,
more locally, by acid rain (Waldén et al., 1991). Some sites in
western Poland show significant declines in diversity over the
20th century (Drozdowski, 1979, 1988). Clausiliids may be especially sensitive to airborne pollution (Holyoak, 1978) and it is
noticeable that they are absent in a number of the Polish and
other sites discussed earlier. It is fortunate that Bial owiez.a is far
/
/
/
/
/
/
157
R. A. D. CAMERON & B. M. POKRYSZKO
from sources of pollution, and that many of its soils are sufficiently eutrophic to be buffered.
DROZDOWSKI, A. 1988. Untersuchungen über Veränderungen der
Malakofauna des Eibenreservats Wierzchlas (Pomorze, VR Polen).
Malakologische Abhandlungen Staatliches Museum für Tierkunde Dresden,
13: 39–48.
ACKNOWLEDGEMENTS
DYDUCH, A. 1980. Ślimaki la˛dowe (Gastropoda terrestria) wybranych
zbiorowisk roślinnych puszczy Bial owieskiej i puszczy Niepol omickiej. Ochrona Przyrody, 43: 223–272.
/
R.A.D.C.’s work in Poland was supported by a Royal Society
short-term visit grant.
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9
20
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30
1
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8
9
40
1
2
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4
5
6
7
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/
/
/
/
/
159
R. A. D. CAMERON & B. M. POKRYSZKO
APPENDIX 1
Names and authorities for all species mentioned in the text, tables and other appendices (sequence and nomenclature follows Kerney et al., 1983).
Acicula polita (Hartmann, 1840)
Aegopinella nitens (Michaud, 1831)
Carychium minimum Müller, 1774
Aegopinella nitidula (Draparnaud, 1805)
Carychium tridentatum (Risso, 1826)
Nesovitrea hammonis (Ström, 1765)
Succinea oblonga Draparnaud, 1801
Nesovitrea petronella (L. Pfeiffer, 1853)
Succinea putris (Linnaeus, 1758)
Oxychilus alliarius (Miller, 1822)
Oxyloma elegans (Risso, 1826)
Zonitoides nitidus (Müller, 1774)
Cochlicopa lubrica (Müller, 1774)
Limax cinereoniger Wolf, 1803
Cochlicopa lubricella (Porro, 1838)
Malacolimax tenellus Müller, 1774
Cochlicopa nitens (Gallenstein, 1848)
Lehmannia nyctelia (Bourguignat, 1861)
Columella edentula (Draparnaud, 1809)
Deroceras laeve (Müller, 1774)
Columella aspera Waldén, 1966
Deroceras sturanyi (Simroth, 1894)
Truncatellina costulata (Nilsson, 1823)
Deroceras agreste (Linnaeus, 1758)
Vertigo pusilla Müller, 1774
Deroceras reticulatum (Müller, 1774)
Vertigo antivertigo (Draparnaud, 1801)
Euconulus fulvus (Müller, 1774)
Vertigo pygmaea (Draparnaud, 1801)
Euconulus alderi (Gray, 1840)
Vertigo moulinsiana (Dupuy, 1849)
Cochlodina laminata (Montagu, 1803)
Vertigo ronnebyensis (Westerlund, 1871)
Cochlodina orthostoma (Menke, 1830)
Vertigo genesii (Gredler, 1856)
Ruthenica filograna (Rossmässler, 1836)
Vertigo alpestris Alder, 1838
Macrogastra ventricosa (Draparnaud, 1801)
Vertigo substriata (Jeffreys, 1833)
Macrogastra latestriata (A. Schmidt, 1857)
Vertigo angustior Jeffreys, 1830
Macrogastra plicatula (Draparnaud, 1801)
Pupilla muscorum (Linnaeus, 1758)
Macrogastra tumida (Rossmässler, 1836)
Vallonia pulchella (Müller, 1774)
Clausilia bidentata (Ström, 1765)
Vallonia costata (Müller, 1774)
Clausilia dubia Draparnaud, 1805
Vallonia excentrica Sterki, 1892
Clausilia pumila C. Pfeiffer, 1828
Acanthinula aculeata (Müller, 1774)
Laciniaria plicata (Draparnaud, 1801)
Chondrula tridens (Müller, 1774)
Bulgarica cana (Held, 1836)
Ena obscura (Müller, 1774)
Bradybaena fruticum (Müller, 1774)
Punctum pygmaeum (Draparnaud, 1801)
Perforatella bidentata (Gmelin, 1788)
Discus ruderatus (Férussac, 1821)
Perforatella incarnata (Müller, 1774)
Discus rotundatus (Müller, 1774)
Perforatella vicina (Rossmässler, 1842)
Arion rufus (Linnaeus, 1758)
Perforatella rubiginosa (A. Schmidt, 1853)
Arion subfuscus (Draparnaud, 1805)
Trichia hispida (Linnaeus, 1758)
Arion silvaticus Lohmander, 1937
Euomphalia strigella (Draparnaud, 1801)
Arion fasciatus (Nilsson, 1822)
Arianta arbustorum (Linnaeus, 1758)
Arion intermedius Normand, 1852
Helicigona lapicida (Linnaeus, 1758)
Vitrina pellucida (Müller, 1774)
Isognomostoma isognomostoma (Schröter, 1784)
Vitrea crystallina (Müller, 1774)
Cepaea nemoralis (Linnaeus, 1758)
Vitrea contracta (Westerlund, 1871)
Cepaea hortensis (Müller, 1774)
Aegopinella pura (Alder, 1830)
Helix pomatia Linnaeus, 1758
Aegopinella minor (Stabile, 1864)
160
.
B IA L OWIEZ A FOREST FAUNAS
/
1
2
3
4
5
6
7
8
9
10
1
2
3
4
5
6
7
8
9
20
1
2
3
4
5
6
7
8
9
30
1
2
3
4
5
6
7
8
9
40
1
2
3
4
5
6
7
8
9
50
1
2
3
4
5
6
7
8
9
60
1
2
3
4
5
6
7
8
APPENDIX 2
Numbers of each species of snail found in each site. A. In Circaeo-Alnetum. B. In Melitto-Carpinetum, Tilio-Carpinetum and in aggregate. C. The occurrence of
slugs by site and overall. F, frequency of occurrence; N, aggregate numbers recorded.
A.
Circaeo-Alnetum
Sites
Species
Acicula polita
2
5
9
11
12
13
16
F
N
.
.
.
.
.
15
2
2
17
269
4
14
22
61
65
18
7
453
Carychium tridentatum
4
8
.
4
11
5
2
6
34
Succinea oblonga
5
.
.
.
.
2
.
2
7
Succinea putris
4
6
6
17
37
29
29
7
128
Carychium minimum
Cochlicopa lubrica
45
30
15
15
47
31
15
7
198
Cochlicopa lubricella
.
.
.
3
.
.
2
2
5
Cochlicopa nitens
.
.
.
.
1
.
.
1
1
16
2
5
1
11
6
11
7
52
Vertigo pusilla
5
.
3
6
1
3
14
6
32
Vertigo alpestris
.
.
.
.
.
.
.
.
.
Vertigo substriata
59
.
7
.
6
3
3
5
78
Vallonia pulchella
.
1
.
.
.
.
.
1
1
Vallonia costata
.
.
.
5
.
14
11
3
30
Columella edentula
Acanthinula aculeata
12
1
8
.
3
19
3
6
46
Punctum pygmaeum
108
13
13
6
25
24
23
7
212
Discus ruderatus
4
7
4
2
.
3
4
6
24
Discus rotundatus
1
6
12
.
.
.
.
3
19
Vitrina pellucida
2
2
1
2
2
.
4
6
13
Vitrea crystallina
55
11
18
12
9
22
10
7
137
Aegopinella pura
18
9
37
5
6
31
4
7
110
.
2
.
.
.
1
.
2
3
Nesovitrea hammonis
40
6
31
3
4
7
5
7
96
Nesovitrea petronella
Aegopinella minor
27
24
14
6
9
15
4
7
99
Zonitoides nitidus
1
1
6
6
46
11
4
7
75
Euconulus fulvus
18
4
4
5
3
5
1
7
40
Cochlodina laminata
6
6
6
13
8
4
26
7
69
Cochlodina orthostoma
.
.
.
.
.
2
.
1
2
Macrogastra ventricosa
.
.
1
2
3
.
3
4
9
Macrogastra plicatula
3
7
6
19
1
9
1
7
46
Macrogastra tumida
.
.
.
.
.
.
.
.
Clausilia dubia
.
.
.
.
.
.
.
.
.
Clausilia pumila
.
.
.
.
3
.
4
2
7
Clausilia bidentata
.
.
1
.
.
.
.
1
1
Laciniaria plicata
.
.
1
.
.
1
2
3
4
Bulgarica cana
.
.
.
.
.
.
4
1
4
Ruthenica filograna
.
.
.
.
.
14
.
1
14
Bradybaena fruticum
6
6
8
11
9
14
25
7
79
Perforatella bidentata
106
14
17
14
11
16
15
19
7
Perforatella vicina
3
.
4
21
6
24
6
6
64
Perforatella rubigiosa
.
.
.
.
18
.
.
1
18
Trichia hispida
.
5
.
20
.
.
.
2
25
Isognomostoma isognomostoma
.
.
3
.
.
.
.
1
3
Cepaea hortensis
.
.
.
.
.
.
.
.
.
Helix pomatia
.
2
.
.
.
.
.
1
2
No. of species
No. of individuals
24
24
26
24
25
28
29
41
.
725
180
242
217
346
394
259
.
2363
161
R. A. D. CAMERON & B. M. POKRYSZKO
Appendix 2. (Continued).
B.
Species
Melitto-Carpinetum
Tilio-Carpinetum
Sites
Sites
1
4
10
14
15
F
N
3
Total
17
6
7
8
F
N
F
N
Acicula polita
.
.
.
.
.
.
.
.
1
.
2
.
2
3
4
20
Carychium minimum
.
.
5
1
.
2
6
13
4
.
5
.
3
22
12
481
Carychium tridentatum
1
.
21
1
20
4
43
24
9
12
130
15
5
190
15
267
Succinea oblonga
.
.
.
.
.
.
.
.
.
.
.
.
.
.
2
7
Succinea putris
.
.
.
4
.
1
4
.
.
.
.
.
.
.
8
132
Cochlicopa lubrica
.
4
6
4
.
3
14
4
8
8
19
6
5
45
15
257
Cochlicopa lubricella
.
.
.
.
.
.
.
.
2
.
8
.
2
10
4
15
Cochlicopa nitens
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
1
Columella edentula
.
1
.
2
4
3
7
.
2
3
3
1
4
9
14
68
Vertigo pusilla
.
2
1
.
2
3
5
2
1
3
7
9
5
22
14
59
Vertigo alpestris
.
.
.
.
2
1
2
.
.
.
.
.
.
.
1
2
Vertigo substriata
1
2
.
1
2
4
6
.
3
.
6
.
2
9
11
93
Vallonia pulchella
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
1
Vallonia costata
.
.
.
.
3
1
3
4
1
.
.
.
2
5
6
38
Acanthinula aculeata
.
1
.
.
.
1
1
3
3
3
18
2
5
29
12
76
Punctum pygmaeum
3
18
5
3
28
5
57
21
16
21
105
23
5
186
17
455
Discus ruderatus
7
22
7
39
29
5
104
27
38
38
7
6
5
116
16
244
Discus rotundatus
17
22
21
26
43
5
129
36
16
22
27
24
5
125
13
273
Vitrina pellucida
1
1
.
.
.
2
2
2
2
1
8
3
5
14
13
29
Vitrea crystallina
2
.
.
5
7
3
14
27
4
10
15
3
5
59
15
210
Aegopinella pura
.
3
2
.
2
3
7
12
8
5
14
5
5
44
15
161
Aegopinella minor
.
1
.
.
1
2
2
1
.
.
8
2
3
11
7
16
Nesovitrea hammonis
36
42
11
9
26
5
124
40
18
5
9
6
5
78
17
298
Nesovitrea petronella
31
25
5
11
10
5
82
11
16
18
11
5
5
61
17
242
Zonitoides nitidus
.
.
2
13
.
2
15
3
8
1
.
.
3
12
12
102
Euconulus fulvus
22
21
2
7
10
5
62
11
8
6
20
5
5
50
17
152
Cochlodina laminata
11
6
8
4
20
5
49
25
20
14
18
7
5
84
17
202
Cochlodina orthostoma
.
1
.
2
.
2
3
5
5
2
3
1
5
16
8
21
Macrogastra ventricosa
.
.
.
.
.
.
.
1
1
1
2
2
5
7
9
16
Macrogastra plicatula
5
2
20
14
10
5
51
37
31
15
15
3
5
101
17
198
Macrogastra tumida
.
.
.
.
.
.
.
1
2
.
.
.
2
3
2
3
Clausilia dubia
.
.
.
.
.
.
.
.
.
2
.
1
2
3
2
3
Clausilia pumila
.
.
.
.
.
.
.
.
1
.
1
.
2
2
4
9
Clausilia bidentata
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
1
Laciniaria plicata
.
3
5
14
2
4
24
7
9
10
13
5
5
44
12
72
Bulgarica cana
.
2
1
1
.
3
4
2
4
7
5
5
5
23
9
31
Ruthenica filograna
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
14
Bradybaena fruticum
1
.
.
2
2
3
5
3
2
3
4
1
5
13
15
97
Perforatella bidentata
.
.
2
.
.
1
2
8
4
9
6
6
5
35
13
143
Perforatella vicina
.
1
2
3
5
4
11
5
7
9
9
19
5
49
15
124
Perforatella rubiginosa
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
18
Trichia hispida
.
.
.
.
.
.
.
2
.
.
1
.
2
3
4
28
Isognomostoma isognomostoma
.
.
.
.
.
.
.
.
1
.
.
.
1
1
2
4
Cepaea hortensis
.
.
.
.
1
1
1
.
.
.
.
.
.
.
1
1
Helix pomatia
.
.
.
.
7
1
7
.
.
.
.
.
.
.
2
9
No. of species
13
20
18
21
22
31
.
28
32
25
30
25
35
.
45
138
180
126
166
236
.
846
337
255
228
499
165
.
1484
.
No. of individuals
162
4693
.
B IA L OWIEZ A FOREST FAUNAS
/
1
2
3
4
5
6
7
8
9
10
1
2
3
4
5
6
7
8
9
20
1
2
3
4
5
6
7
8
9
30
1
2
3
4
5
6
7
8
9
40
1
2
3
4
5
6
7
8
9
50
1
2
3
4
5
6
7
8
9
60
1
2
3
4
5
6
7
8
Appendix 2. (Continued).
C.
Species
Circaeo-Alnetum
Melitto-Carpinetum
Tilio-Carpinetum
Site
Site
Site
2
5
9
Arion subfuscus
Arion fasciatus
.
.
.
Limax cinereoniger
11
12
13
16
.
F
1
4
10
7
2
.
6
.
14
.
Limax tenellus
.
.
.
.
Lehmannia nyctelia
.
.
.
.
.
15
F
3
5
.
Total
17
.
6
7
8
F
F
5
17
16
.
2
5
5
.
2
5
1
.
.
.
.
7
1
Deroceras laeve
.
.
2
.
No. of species
2
2
2
3
4
3
1
4
4
3
2
2
2
4
3
3
3
3
3
4
6
Total molluscs
26
26
28
27
29
31
30
45
17
23
20
23
24
35
31
35
28
33
28
38
51
.
.
163
.
.
.
.
2
R. A. D. CAMERON & B. M. POKRYSZKO
APPENDIX 3
Names, locations and sources for forest faunas from lowland Poland.
Number on map refers to Figure 5 (NR, Nature Reserve; WNP, Wielkopolski
National Park, near Poznań).
No. on
Location
map
Source
A. Beech forests
Zygmunt Czubiński NR, Wolin
1
Dzie˛czkowski, 1988
Bohdan Dyakowski NR, Wolin
2
Dzie˛czkowski, 1988
Mścice, nr Koszalin
3
Dzie˛czkowski, 1988
Mścice, nr Koszalin
4
Dzie˛czkowski, 1988
Buczyna NR, nr Sl omowo
5
Dzie˛czkowski, 1988
Mielenko, nr Koszalin
6
Dzie˛czkowski, 1989
Dzie˛czkowski, 1988
/
B. Mixed forests
Zygmunt Czubiński NR, Wolin
7
Buki nad Jeziorem Lutomskim NR
8
Dzie˛czkowski, 1988
De˛bina NR, nr Wa˛growiec
9
Dzie˛czkowski, 1988
Wia˛zy w Nowym Lesie NR, nr Gniezno
10
Dzie˛czkowski, 1988
Adam Wodziczko Grabina NR, WNP
11
Dzie˛czkowski, 1988
Wyspa Zamkowa NR, WNP
12
Dzie˛czkowski, 1988
Puszczykowskie Góry NR, WNP
13
Dzie˛czkowski, 1963
Jabukowo, nr Poznań
14
Koralewska-Batura, 1989
Opalenica, nr Poznań
15
Koralewska-Batura, 1993
Morasko, nr Poznań
16
Dzie˛czkowski, 1974
Buki nad Jeziorem Lutomskim NR
17
Szybiak, 2002
Wierzchlas, Pomerania
18
Drozdowski, 1988
Szczerkowo, Osie, nr Bydgoszcz
19
Drozdowski, 1979
Wielki Las NR, nr Lwówek
20
Dzie˛czkowski, 1988
Wia˛zy w Nowym Lesie NR, nr Gniezno
21
Dzie˛czkowski, 1988
Kolno Mie˛dzychodzkie, nr Mie˛dzychód
22
Dzie˛czkowski, 1988
Jeziory-Widok, WNP
23
Dzie˛czkowski, 1988
Radunia gorge, nr Kartuzy
24
Drozdowski, 1981
Lake Klasztorne, nr Kwidzyń
25
Drozdowski, 1966
C. Ficario-Ulmeta
D. Circaeo-Alneta
Buki nad Jeziorem Lutomskim NR
.
Uroczysko Bazantarnia NR,
26
Dzie˛czkowski, 1988
27
Dzie˛czkowski, 1988
Zwierzyniec-Korabka, nr Skierniewice
28
Dzie˛czkowski, 1988
Mielenko, nr Koszalin
29
Dzie˛czkowski, 1989
De˛biniec, nr Poznań
30
Jankowiak, Bl oszyk and
nr Skierniewice
/
Jackiewicz, 1991
Buki nad Jeziorem Lutomskim NR
31
Szybiak, 2002
164