Plant-Dwelling Spider Assemblages in Managed and Protected Primeval Deciduous Stands of Białowieża Forest, Poland
Institute of Biological Sciences, Faculty of Natural Sciences, University of Siedlce, Ul. Prusa 14, 08-110 Siedlce, Poland
*
Author to whom correspondence should be addressed.
Forests 2025, 16(7), 1093; https://doi.org/10.3390/f16071093
Submission received: 13 May 2025
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Revised: 27 June 2025
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Accepted: 30 June 2025
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Published: 1 July 2025
(This article belongs to the Special Issue Species Diversity and Habitat Conservation in Forest)
Abstract
The Białowieża Forest is home to both primary forests under strict protection and commercial forests, which provides an opportunity to compare them in terms of the number of individuals, number of species, and composition of different animal assemblages. The main goal of our study was to compare spider assemblages inhabiting herbaceous vegetation in these two types of forest stands. Spiders were sampled using a sweep net in an oak–lime–hornbeam forest, an ash–alder forest, and an alder carr. More spiders were found in unmanaged stands compared to managed stands, but a significant difference was found only in the alder carr. Total species richness did not significantly vary between managed and unmanaged stands in all forest types. In the oak–lime–hornbeam forest, more species per sample were found in commercial stands compared to primeval stands, while the result was the opposite for the alder carr. Our research did not show a clear negative impact of forest management on plant-dwelling spiders. The impact of forest management was evident in the case of the riparian forest, where the managed stand was characterized by low canopy cover as a result of logging carried out years ago, which is likely to have resulted in differences in family composition.
1. Introduction
Protecting the entire Białowieża Forest, a valuable forest complex in Europe, where our research was conducted, is one of the priorities for many scientists and nature conservation organizations [1,2]. Despite years of public debate in Poland on expanding the Białowieża National Park to cover the entire area of the Białowieża Forest, there is still conflict between proponents of this solution and foresters, and intensive logging and elimination of dead trees continues [3,4,5]. These practices cause serious threats to many organisms connected with mature, dead wood-rich forests, such as woodpeckers [6,7,8,9] and saproxilic beetles [10].
Unlike the commercial stands of the Białowieża Forest, human activity in stands protected within the Białowieża National Park is limited to scientific studies and guided tourist excursions on selected routes. The stands found in the national park are among the most natural forests in Europe. They are characterized by a diverse tree species composition, a vertically stratified and uneven-aged tree structure, significant tree height, and an abundance of dead wood [11,12].
The presence of both commercial and primary stands in the Białowieża Forest provides an opportunity to compare them in terms of number of species, abundance, and species diversity of the various groups of organisms inhabiting them. Studies on this topic involved birds, especially woodpeckers [6,8,9], Collembola [13], carabid beetles [14], and spiders. In the case of the latter, however, they were limited to species that inhabit tree branches, as well as rare and endangered species [15,16]. Most of these studies showed a negative impact of forest management on both individual species and entire animal assemblages [9,13,16,17].
Spiders are widely recognized as one of the model groups of organisms used to study the impact of human activity on the environment [18,19]. The reason for this is their sensitivity to disturbances in habitat structure and changes in microclimatic conditions [20,21,22]. Forest-dwelling spider assemblages are influenced by, among others, the openness of the tree canopy [23,24], the type of stand, and the diversity of tree species [25,26,27,28], as well as the presence of woody debris [29]. Moreover, in the case of plant-dwelling spiders, the structure and complexity of herbaceous vegetation may be the main factor affecting their assemblages [30,31].
Research on spider fauna in the Białowieża Forest has been carried out before, both in the Polish and Belarusian parts [16,27,32]. None of the published studies compared plant-dwelling spider assemblages in managed and primary stands of the Białowieża Forest, which is the focus of this study.
The main goals of our study were as follows: (1) to identify the species and family composition of plant-dwelling spider assemblages in three types of deciduous stands: oak–lime–hornbeam forest, ash–alder riparian forest, and alder carr located in both commercial and primeval stands of the Białowieża Forest; (2) to assess differences between spider assemblages in commercial and unmanaged stands with respect to their abundance and species richness, as well as the proportion of individual families in the assemblages. Our hypothesis was that the number of spider individuals, number of species, and number of spider families would be higher in primeval forests compared to managed forests. This hypothesis was supported by earlier studies demonstrating that forest management in the Białowieża Forest had detrimental effects on specific species as well as on entire animal groups [9,13,16,17].
2. Materials and Methods
2.1. Study Site
The study was conducted in the Białowieża Forest (1500 km2) located on the Polish–Belarusian border, representing one of the last remaining fragments of the primeval forest that once covered extensive areas of Europe. Part of the Białowieża Forest in Poland had been protected since 1921 as a national park, which, after its enlargement in 1996, now covers 105 km2. Forest stands in the Białowieża National Park (BNP) exhibit key structural and compositional characteristics of primeval forests, such as vertically stratified and heterogenous age structure, multispecies tree communities, considerable tree heights, and a substantial accumulation of dead wood [11,12]. The rest of the Polish part of the Białowieża Forest (525 km2) is managed by the “State Forests”, which carries out typical forest management involving timber extraction, dead wood clearance, reforestation, and pest management, mainly by removing infected trees. As a result of these practices, forests in the managed zone are younger, less diverse in terms of structure and much less abundant in dead wood [6,7,9]. Our study was conducted in three deciduous forest stands: an oak–lime–hornbeam forest (Tilio-Carpinetum), an ash–alder riparian forest, (Circaeo-Alnetum) and an alder carr (Carici elongatae-Alnetum). Two plots were located in each type of forest: one in the managed part of the Białowieża Forest and one in the primeval forest of the BNP. The location of the study plots is presented in Figure 1 and their characteristics in Table 1.
2.2. Vegetation Measurements and Spider Sampling
To characterize and compare the structure of herbaceous vegetation in our study plots, which may be useful for interpreting potential difference between spider assemblages, we measured vegetation cover in three groups according to the height of vegetation. Fifteen measurements of the vegetation cover were performed on each study plot from April to September, distinguishing them into three height categories: low (≤10 cm), medium (10–30 cm), and high (>30 cm) vegetation. Three measurement sites were randomly selected in the study plot each time the measurements were conducted. To facilitate the measurements, we used a wooden frame of 1 m2 divided into 16 smaller quadrats. In each quadrat, a vegetation cover index was estimated for each vegetation class based on a numerical scale: 0 for 0%, 1 for 1%–25%, 2 for 26%–50%, 3 for 51%–75%, and 4 for 76%–100%. Next, the mean value was calculated for the entire area of the frame and measurements from three selected sites were averaged.
Spiders were sampled within the study plots (defined as rectangles of about 20 × 40 m) in 1998 and 1999. From every surveyed plots, 19 samples were obtained: 8 in 1998 (1 in May, 2 in June, 2 in August, 2 in September, 1 in October) and 11 in 1999 (1 in June, 3 in July, 2 in August, 2 in September, 2 in October, 1 in November). The different number of samples collected in each month was due to weather conditions, as sampling was conducted exclusively on rain-free days to obtain representative and comparable material. Spiders were collected using a sweep net with a diameter of 35 cm. One sample consisted of four series, with 25 sweep net beats on herbaceous vegetation in each series (100 beats in total). Spider samples obtained during fieldwork were preserved in 75% ethanol and later identified in the laboratory to the species level. However, due to the immature stage of many individuals, identification was limited to genus or family for many specimens. The collected material was deposited at the Institute of Biological Sciences, University of Siedlce, Poland.
2.3. Statistical Analyses
The degree of herbaceous vegetation cover was compared in each height class separately, using Kruskal–Wallis tests. Next, Dunn’s post hoc tests were used to find differences between individual study plots. To assess sampling sufficiency, we calculated five species richness estimators (Chao1, Chao2, Jackknife1, Jackknife2, and Michaelis–Menten) using 100 randomizations [33]. Based on the obtained species richness estimators, we calculated sampling completeness, expressed as the proportion of observed species richness relative to the estimated total. Both adults and juveniles were included in this analysis to ensure the reliability of the species richness estimates [34]. G-tests were used to compare the proportion of families in plant-dwelling spider assemblages between primeval and managed stands for each type of forest. To compare the managed and primeval stands in each type of forest with respect to total species richness (i.e., the number of species found during the entire study period), rarefaction curves for the observed species richness were generated, accompanied by confidence intervals set at the 95% level, which were based on a bootstrap procedure with 100 replications. When the calculated confidence intervals do not overlap, it means that the species richness of the compared forests are statistically significantly different [35,36]. Generalized linear models (GLMs) were applied to evaluate the associations of the number of spider specimens and species recorded in each sample with forest type and forest management. In the model where the response variable was the number of spiders, the Gaussian error distribution and the log-link function were applied. In the model where the response variable was the number of species, the Gaussian error distribution and the identity link function were applied. Species richness in a given sample was determined based on all collected spider individuals irrespective of their age (adult or juvenile). The type of forest (ash–alder riparian forest, alder carr, oak–lime–hornbeam stand) and forest management (managed stands vs. primeval stands) were included as fixed categorical explanatory variables. All models included interactions between the aforementioned variables to detect possible differences between managed and unmanaged stands of the same forest type. Paired contrasts were performed to assess differences between the levels of explanatory variables, which showed significant associations with response variables. G-tests were performed using formulas prepared in Microsoft Excel, whereas Kruskal–Wallis tests and GLM analyses were performed in IBM SPSS Statistics 21.0 for Windows. Richness estimators and rarefaction curves were calculated in EstimateS version 9.1.0 [37].
3. Results
Vegetation cover varied among the study plots within all vegetation height categories analyzed: low layer (Kruskal–Wallis test, H5,90 = 27.17, p < 0.001), medium layer (Kruskal–Wallis test, H5,90 = 44.78, p < 0.001) and high layer (H5,90 = 46.10, p < 0.001). In general, the most complex herbaceous vegetation was found on both plots in the ash–alder riparian forest, while the least complex vegetation grew on plots located in the oak–lime–hornbeam forest. However, managed and primeval stands in particular forest types did not differ in vegetation cover across all analyzed height classes, except for low vegetation in the oak–lime–hornbeam stand (Figure 2).
A total of 7193 spiders were collected throughout the study period. The majority of the collected specimens were juvenile spiders (4534, 63%), especially in the managed ash–alder riparian forest and both oak–lime–hornbeam stands, where their abundance was three or more times higher than that of adult individuals (Table 2).
A total of 4303 specimens were determined to the species level, representing 103 species (Table 2). The highest species richness was found in the primeval alder carr (54 species), while the lowest was in the primeval oak–lime–hornbeam forest (30 species). Most species richness estimators indicated higher species richness than observed, especially in the managed oak–lime–hornbeam forest, where the sampling completeness calculated based on all estimators, except Michaelis–Menten, was the lowest of all study plots (Table 3).
Rarefaction curves showed that the total species richness remained similar across managed and unmanaged stands in all three analyzed forest types, but in the case of the ash–alder forest, the difference was close to significance (Figure 3).
The most abundant species in the managed oak–lime–hornbeam stand was Enoplognatha ovata (Clerck, 1757), while Pachygnatha listeri (Sundevall, 1830) was the most abundant species in the primeval oak–lime–hornbeam stand. The dominant species in the ash–alder riparian forest were Dolomedes fimbriatus (Clerck, 1757) for the plot located in the managed stand and Bathyphantes nigrinus (Westring, 1851) for the plot in the primeval stand. Pachygnatha clercki (Sundevall, 1830) was the species with the highest abundance in both alder carr plots (Table 2). The collected spiders belonged to 17 families. The most abundantly represented families were Linyphiidae for plots located in both oak–lime–hornbeam and primeval ash–alder riparian forests, and Tetragnathidae for plots located in both alder carr and managed ash–alder riparian forests (Table 2). The proportion of spider individuals from different families collected throughout the study period differed between the managed and primeval stands with regard to the ash–alder riparian forest (G-test = 49.45, p < 0.001, df = 14), while no such difference was found for the oak–lime-hornbeam forest (G-test = 20.45, p = 0.059, df = 12) and alder carr (G-test = 3.26, p = 0.993, df = 12).
GLM analysis showed that the number of spider individuals collected in particulars samples was associated with forest management; however, significant interaction between this variable and forest type was also found (Table 4).
Although more spider individuals were sampled in unmanaged stands compared to managed stands, considering specific forest types, a significant difference was found only in the alder carr (Figure 4). The species richness (assessed in individual samples) was significantly associated with the forest type; however, the interaction between forest management and forest type was also significant (Table 4). In general, the lowest number of species was revealed in samples from the oak–lime–hornbeam forest, while the highest number was found in alder carr. In the case of the oak–lime–hornbeam forest, significantly more species per sample were found in the managed stand compared to the primeval stand, while the opposite result was determined for the alder carr. No difference was found in the ash–alder riparian forest (Figure 5).
4. Discussion
Our hypothesis that spider species richness and abundance would be higher in primeval forests compared to managed forests was not unequivocally confirmed. Higher abundance of spiders and a higher number of species (both values recorded per sample) in primeval stands than in managed stands were found only in the alder carr, while higher species diversity in primeval stands was found only in the case of the oak–lime–hornbeam forest. On the other hand, no differences were found in total species richness between commercial and primeval stands in all forest types. The results of the study suggest that factors other than the location of the study plots (i.e., in primary or managed forest) influenced plant-dwelling spider assemblages.
Although no forestry operations were carried out on the study plots at the time of material collection, as a result of those previously carried out, plots in the managed forests were characterized by lower stand age (old trees were cut earlier) and less dead wood (which was removed) compared to plots in the primeval forest. The differences were particularly evident for the ash–alder forest, in which the managed stand was characterized by a low degree of canopy cover as a result of logging carried out years ago. This may explain the fact that while the two ash–alder riparian plots were similar in terms of the number of individuals captured and species identified, they differed in terms of the proportion of individuals belonging to specific families. Linyphiidae dominated the plot located in the BPN due to the high abundance of Bathypanthes nigrinus, Gongylidium rufipes (Linnaeus, 1758), and Helophora insignis (Blackwall, 1841), while Tetragnathidae dominated in the managed stand. In addition, Pisauridae, almost absent in the former plot, were abundant in the latter (mainly Dolomedes fimbriatus). In general, the typical forest species that dominated the plot in the national park were replaced by species showing preferences for open wetlands, as the canopy cover in the managed ash–alder riparian forest was only 40%. Our findings corroborate the results obtained by Oxbrough et al. [23], who showed that open space in a forest favors the occurrence of spider species normally absent there and may be positively linked to higher abundance and species richness. The effect of canopy cover was also demonstrated by Košulič et al. [24], who revealed that the highest species richness was associated with a partially open canopy, while rare and threatened species were more abundant in areas with a more open canopy, and the number of rare and threatened species was higher in areas with more open canopy. The above studies focused on epigeic spiders, but based on our research, it is clear that the impact of canopy cover (closely related to forest management) can also apply to plant-dwelling spider assemblages.
However, in the case of plant-dwelling spiders, herbaceous vegetation can play a key role in shaping their assemblages. It is primarily the vegetation that determines the heterogeneity of a habitat. Many studies showed a positive relationship between habitat complexity and species diversity or species richness [38]. A more complex habitat provides more diverse niches, which means that more species can occur in a given habitat [31]. In addition, vegetation provides shelter, sufficient humidity, abundant prey, and structures on which spiders can build their webs [39,40,41,42,43,44]. In general, vegetation height, vegetation cover or volume, and plant diversity are positively correlated with spider abundance, species diversity, and the number of spider species [31,39,43,45]. In our study, primeval and managed plots in a given forest type did not differ in terms of vegetation complexity, expressed as the degree of cover by vegetation in different height classes, except for low vegetation in the oak–lime–hornbeam forest. This may be the reason, among other things, for the lack of major differences between primary and managed stands, clearly indicating that either of them substantially differs in terms of spider abundance and species richness. On the other hand, significant differences in the degree of herbaceous vegetation cover were found between different forest types. It seems likely that the smallest number of species found in the oak–lime–hornbeam stand may result, among other things, from much less complex vegetation in the two plots located in this forest type compared to the others.
Finding the highest species richness in alder carr shows that it may play an important role as a biodiversity hotspot. This may be because it is primarily a very diverse habitat in terms of structure and microclimatic conditions (e.g., moisture), which guarantees a diverse spectrum of habitat niches [20]. On the other hand, the two plots located in the alder carr differed significantly in the number of individuals collected. It seems that the higher number of spiders collected in the primary alder carr is due to the fact that the trees growing here are older and larger than in the managed alder carr. This makes hummocks formed by their roots larger, which provides more space and shelter for spiders.
An analogous study from the Białowieża Forest, but concerning spiders inhabiting tree branches, showed no differences between plots located in primeval and managed forests in the total number of species found [16]. The above study also showed no differences in the number of individuals collected. However, in the case of the number of species per sample, a higher number in primeval stands was observed [16]. It is also worth mentioning the results of the study conducted, among others, on the same plots, showing that managed forests reduce the occurrence of rare and threatened species [15].
5. Conclusions
Our research did not show a clear negative impact of forest management on plant-dwelling spider assemblages, and the results varied depending on the forest type. On the one hand, a higher abundance of spider individuals and a higher number of species (per sample) were found in the primeval alder carr compared to the managed alder carr. On the other hand, the managed oak–lime–hornbeam forest was characterized by a higher number of species (per sample) compared to the primeval oak–lime–hornbeam stand. In contrast, the impact of forest management was most evident in the case of the ash–alder riparian forest, where the managed stand was characterized by low canopy cover as a result of logging carried out years ago, which is likely to have resulted in differences in family composition.
Author Contributions
Conceptualization, M.S. and T.S.; methodology, M.S. and T.S.; formal analysis, T.S.; investigation, M.S. and T.S.; writing—original draft preparation, T.S.; writing—review and editing, M.S., T.S. and B.P.; visualization, B.P.; project administration, M.S.; funding acquisition, M.S. All authors have read and agreed to the published version of the manuscript.
Funding
This research was funded by University of Siedlce.
Data Availability Statement
The raw data supporting the conclusions of this article will be made available by the authors on request.
Acknowledgments
We thank the authorities of the Białowieża National Park for their help during our research. The authors have reviewed and edited the output and take full responsibility for the content of this publication.
Conflicts of Interest
The authors declare no conflicts of interest. The funder had no role in the design of the study; in the collection, analyses, interpretation of data; in the writing of the manuscript; and in the decision to publish the results.
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Figure 1.
Spatial distribution of the study plots (black dots) in managed forests and the Białowieża National Park. Abbreviations: M OLH—managed oak–lime–hornbeam forest, P OLH—primeval oak–lime–hornbeam forest, M RF—managed ash–alder riparian forest, P RF—primeval ash–alder riparian forest, M AC—managed alder carr, P AC—primeval alder carr.
Figure 1.
Spatial distribution of the study plots (black dots) in managed forests and the Białowieża National Park. Abbreviations: M OLH—managed oak–lime–hornbeam forest, P OLH—primeval oak–lime–hornbeam forest, M RF—managed ash–alder riparian forest, P RF—primeval ash–alder riparian forest, M AC—managed alder carr, P AC—primeval alder carr.
Figure 2.
Cover of (A) low vegetation (≤10 cm), (B) medium vegetation (10–30 cm), and (C) high vegetation (>30 cm) on study plots (median with 25–75% quartile ranges). Different letters indicate significant differences between plots. Abbreviations: P—primeval stand, M—managed stand, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr.
Figure 2.
Cover of (A) low vegetation (≤10 cm), (B) medium vegetation (10–30 cm), and (C) high vegetation (>30 cm) on study plots (median with 25–75% quartile ranges). Different letters indicate significant differences between plots. Abbreviations: P—primeval stand, M—managed stand, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr.
Figure 3.
Individual-based rarefaction curves (solid) with 95% confidence limits (dashed curves) comparing species richness in managed (black) and primeval (red) stands in: (A) lime–oak–hornbeam forest, (B) ash–alder riparian forest, and (C) alder carr.
Figure 3.
Individual-based rarefaction curves (solid) with 95% confidence limits (dashed curves) comparing species richness in managed (black) and primeval (red) stands in: (A) lime–oak–hornbeam forest, (B) ash–alder riparian forest, and (C) alder carr.
Figure 4.
The number of spider individuals (least squares means with 95% confidence limits) recorded in a single sample in analyzed forest plots. Asterisks show where primeval and managed plots differ significantly in given forest types. Abbreviations: AF—all forest types, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr.
Figure 4.
The number of spider individuals (least squares means with 95% confidence limits) recorded in a single sample in analyzed forest plots. Asterisks show where primeval and managed plots differ significantly in given forest types. Abbreviations: AF—all forest types, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr.
Figure 5.
The number of spider species (least squares means with 95% confidence limits) recorded in a single sample in analyzed forest plots. Asterisks show where primeval and managed plots differ significantly in given forest types. Abbreviations: AF—all forest types, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr.
Figure 5.
The number of spider species (least squares means with 95% confidence limits) recorded in a single sample in analyzed forest plots. Asterisks show where primeval and managed plots differ significantly in given forest types. Abbreviations: AF—all forest types, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr.
Table 1.
Characteristics of the study plots in the Białowieża Forest. Abbreviations: M—managed stand, P—primeval stand, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr.
Table 1.
Characteristics of the study plots in the Białowieża Forest. Abbreviations: M—managed stand, P—primeval stand, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr.
| Plot | Main Tree Species | Tree Stand Structure | Herbaceous Vegetation |
|---|---|---|---|
| M OLH | Quercus robur L., Populus tremula L., Carpinus betulus L. | Most trees were of similar age (around 70 years). Older trees were cut down and dead or dying trees were removed. Dense canopy cover (about 95%). | Anemone nemorosa L., Aegopodium podagraria L., Galeobdolon luteum (L.) L., Impatiens noli-tangere L. |
| P OLH | Carpinus betulus, Quercus robur, Picea abies (L.) H. Karst, Pinus silvestris L., Tilia cordata Mill. | Trees were diverse in size and age (up to 200 years). Considerable volume of dead wood. Canopy cover of about 70%. | Anemone nemorosa, Stellaria holostea L., Hepatica nobilis Mill, Oxalis acetosella L., Maianthemum bifolium (L.) F. W. Schmidt, Calamagrostis arundinacea (L.) Roth., Pteridium aquilinum (L.) Kuhn, Convallaria majalis L., Vaccinium myrtillus L., Trientalis europaea (L.) U. Manns & Anderb. |
| M RF | Fraxinus excelsior L., Alnus glutinosa (L.) Gaertn. | The oldest, thickest trees were cut down here, which caused a significant disturbance of the forest structure and thinning of the stand. Age of oldest trees were up to 50 years. Small amount of dead wood. Canopy cover of about 40%. | Rubus caesius L., Urtica dioica L., Aegopodium podagraria, Filipendula ulmaria (L.) Maxim., Lysimachia vulgaris L., L. nummularia L. |
| P RF | Alnus glutinosa, Fraxinus excelsior, Picea abies. | Trees were diverse in size and age (mainly 120 years, with the oldest trees up to 200 years old). Considerable volume of dead wood. Canopy cover of about 80%. | Ficaria verna huds., Impatiens noli-tangere, Mercurialis perennis L., Aegopodium podagraria, Lamium maculatum L., Urtica dioica. |
| M AC | Alnus glutinosa, Fraxinus excelsior, Picea abies. | After the oldest trees were cut down, the stand regenerated from regrowth. The hummock–hollow structure of the forest floor was preserved, although the hummocks were smaller than in the BNP. Trees at 70 years of age. Canopy cover of about 70%. | Filipendula ulmaria, Solanum dulcamara L., Carex elongate L., Galium palustre L., Iris pseudacorus L., Urtica dioica, Maianthemum bifolium, Oxalis acetosella. |
| P AC | Alnus glutinosa Fraxinus excelsior Picea abies | Hummock–hollow structure in the forest floor. The hummocks were large, which was related to the age and size of the root systems of the trees that formed them. Trees were diverse in size and age (averaging at 120 years, with the oldest trees up to 200 years old). Considerable volume of dead wood. Canopy cover of about 70%. | Carex elongata, Thelypteris palustris Schott, Solanum dulcamara, Lycopus europaeus L., Carex pseudocyperus L., Cicuta virosa L., Galium palustre, Iris pseudacorus, Scutellaria hastifolia L., Sium latifolium L., Peucedanum palustre (L.) Moench, Phalaris arundinacea L., Maianthemum bifolium, Oxalis acetosella. |
Table 2.
Spiders collected on herbaceous vegetation in managed and primeval stands in the Białowieża Forest. Families and species are in alphabetical order. In the rows where the names of the families are shown, only their percentage contribution is given. In other cases, the number of adult/juvenile individuals is given. Abbreviations: M—managed stand, P—primeval stand, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr, ad./juv.—the number of adult/juvenile spider individuals, un.—individuals determined only to the family level.
Table 2.
Spiders collected on herbaceous vegetation in managed and primeval stands in the Białowieża Forest. Families and species are in alphabetical order. In the rows where the names of the families are shown, only their percentage contribution is given. In other cases, the number of adult/juvenile individuals is given. Abbreviations: M—managed stand, P—primeval stand, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr, ad./juv.—the number of adult/juvenile spider individuals, un.—individuals determined only to the family level.
| Family/Genus/Species | M OLH ad./juv. | P OLH ad./juv. | M RF ad./juv. | P RF ad./juv. | M AC ad./juv. | P AC ad./juv. |
|---|---|---|---|---|---|---|
| Family Agelenidae | 0.08% | |||||
| Coelotes atropos (Walckener, 1830) | 0/1 | |||||
| Family Anyphaenidae | 2.71% | 0.39% | 1.36% | 1.52% | 0.70% | 0.85% |
| Anyphaena accentuata (Walckener, 1802) | 1/31 | 0/4 | 0/6 | 8/6 | 1/15 | 5/15 |
| Family Araneidae | 7.04% | 3.33% | 3.23% | 1.06% | 3.51% | 1.40% |
| Agalenatea redii (Scopoli, 1763) | 0/1 | 0/2 | ||||
| Araneidae un. | 0/8 | 0/2 | 0/2 | |||
| Araneus alsine (Walckener, 1802) | 0/2 | |||||
| Araneus diadematus Clerck, 1757 | 0/5 | 0/3 | 0/1 | 1/2 | 0/1 | |
| Araneus marmoreus Clerck, 1757 | 0/1 | 3/2 | 2/0 | 1/0 | ||
| Araneus quadratus Clerck, 1757 | 2/3 | |||||
| Araneus sp. Clerck, 1757 | 0/4 | 0/2 | 0/1 | |||
| Araneus sturmi (Hahn, 1831) | 0/2 | 0/3 | 0/1 | |||
| Araniella cucurbitina (Clerck, 1757) | 1/0 | 1/0 | 3/0 | 1/0 | ||
| Araniella sp. Chamberlin & Ivie, 1942 | 0/13 | 0/1 | 0/3 | 0/11 | 0/4 | |
| Cyclosa conica (Pallas, 1772) | 2/48 | 2/24 | 2/4 | 2/8 | 1/5 | 3/10 |
| Hypsosinga pygmaea (Sundevall, 1831) | 0/2 | |||||
| Mangora acalypha (Walckener, 1802) | 0/2 | 2/8 | 1/0 | |||
| Nuctenea umbratical (Clerck, 1757) | 0/1 | |||||
| Family Clubionidae | 1.36% | 1.57% | 1.62% | 2.35% | 1.76% | 2.01% |
| Clubiona comta C. L. Koch, 1839 | 1/0 | |||||
| Clubiona lutescens Westring, 1851 | 1/0 | 1/0 | 11/1 | 1/0 | 3/0 | |
| Clubiona pallidula (Clerck, 1757) | 1/0 | |||||
| Clubiona sp. Latreille, 1804 | 0/14 | 0/15 | 0/18 | 0/18 | 0/14 | 1/29 |
| Clubiona stagnatilis Kulczyński, 1897 | 1/0 | |||||
| Family Dictynidae | 0.08% | 0.20% | 0.09% | |||
| Dictyna arundinacea (Linnaeus, 1758) | 1/0 | |||||
| Dictyna sp. Sundevall, 1833 | 0/1 | 0/1 | 0/1 | |||
| Family Dysderidae | 0.10% | |||||
| Harpactea sp. Bristowe, 1939 | 0/1 | |||||
| Family Linyphiidae | 44.53% | 43.25% | 18.20% | 57.42% | 30.33% | 32.76% |
| Agyneta affinis (Kulczyński, 1898) | 1/0 | |||||
| Agyneta rurestris (C. L. Koch, 1836) | 1/0 | 1/0 | ||||
| Agyneta sp. Blackwall, 1859 | 0/1 | |||||
| Bathyphantes approximatus (O. Pickard-Cambridge, 1871) | 6/0 | 3/0 | ||||
| Bathyphantes gracilis (Blackwall, 1841) | 1/0 | 3/0 | ||||
| Bathyphantes nigrinus (Westring, 1851) | 39/0 | 8/0 | 12/0 | 239/0 | 45/0 | 103/0 |
| Bathyphantes parvulus (Westring, 1851) | 2/0 | |||||
| Bathyphantes sp. Menge, 1866 | 0/8 | 0/15 | 0/5 | 0/31 | 0/9 | |
| Centromerus sylvaticus (Blackwall, 1841) | 1/0 | |||||
| Ceratinella brevis (Wider, 1834) | 1/0 | |||||
| Diplocephalus picinus (Blackwall, 1841) | 1/0 | |||||
| Dismodicus bifrons (Blackwall, 1841) | 2/0 | 5/0 | ||||
| Drapetisca socialis (Sundevall, 1833) | 1/0 | 1/1 | ||||
| Entelecara congenera (O. Pickard-Cambridge, 1879) | 1/0 | |||||
| Entelecara media Kulczyński, 1887 | 1/0 | |||||
| Erigone atra Blackwall, 1833 | 2/0 | 2/0 | ||||
| Erigone dentipalpis (Wider, 1834) | 1/0 | |||||
| Floronia bucculenta (Clerck, 1757) | 1/0 | 4/0 | 2/1 | 10/0 | ||
| Gonatium rubellum (Blackwall, 1841) | 6/2 | 9/1 | ||||
| Gongylidiellum murcidium Simon, 1884 | 1/0 | 1/0 | ||||
| Gongylidium rufipes (Linnaeus, 1758) | 1/0 | 18/67 | 25/112 | 9/10 | 32/23 | |
| Helophora insignis (Blackwall, 1841) | 52/53 | 0/3 | 1/0 | 65/58 | 9/0 | 40/8 |
| Hypomma bituberculatum (Wider, 1834) | 3/0 | |||||
| Kaestneria dorsalis (Wider, 1834) | 1/2 | |||||
| Kaestneria pullata (O. Pickard-Cambridge, 1863) | 1/0 | |||||
| Linyphia hortensis Sundevall, 1830 | 1/0 | 3/0 | 9/1 | 4/1 | 4/0 | 3/0 |
| Linyphia sp. Latreille, 1804 | 0/26 | 0/61 | ||||
| Linyphia triangularis (Clerck, 1757) | 38/0 | 29/1 | 35/5 | 16/21 | 22/2 | 79/9 |
| Linyphiidae un. | 0/142 | 0/194 | 0/4 | 0/6 | ||
| Maso sundevalli (Westring, 1851) | 6/0 | |||||
| Microlinyphia pusilla (Sundevall, 1830) | 1/13 | 23/18 | 3/0 | 14/1 | ||
| Neriene clathrata (Sundevall, 1830) | 1/0 | 1/0 | 1/2 | 1/0 | 1/0 | |
| Neriene emphana (Walckenaer, 1841) | 1/0 | 0/1 | 3/0 | 1/0 | ||
| Neriene montana (Clerck, 1757) | 2/0 | 5/0 | 15/0 | 28/0 | 78/0 | |
| Neriene peltata (Wider, 1834) | 13/1 | 2/3 | 5/47 | 6/4 | 14/42 | |
| Neriene radiata (Walckenaer, 1841) | 1/2 | 1/0 | ||||
| Neriene sp. Blackwall, 1833 | 0/25 | 0/59 | 0/9 | 0/21 | 0/9 | 0/9 |
| Obscuriphantes obscurus (Blackwall, 1841) | 3/0 | |||||
| Oedothorax apicatus (Blackwall, 1850) | 1/0 | |||||
| Oedothorax gibbosus (Kulczyński, 1882) | 4/0 | |||||
| Oedothorax retusus (Westring, 1851) | 1/0 | 1/0 | ||||
| Oryphantes angulatus (O. Pickard-Cambridge, 1881) | 1/0 | |||||
| Pityohyphantes phrygianus (C. L. Koch, 1836) | 1/2 | |||||
| Porrhomma pygmaeum (Blackwall, 1834) | 1/0 | 2/0 | 63/0 | 16/5 | ||
| Savignia frontata Blackwall, 1833 | 1/0 | 1/0 | ||||
| Tenuiphantes alacris (Blackwall, 1853) | 1/0 | 15/0 | ||||
| Tenuiphantes cristatus (Menge, 1866) | 2/0 | 3/0 | 1/0 | 2/0 | 1/0 | |
| Tenuiphantes mengei (Kulczyński, 1887) | 1/0 | |||||
| Tenuiphantes sp. Saaristo & Tanasevitch 1996 | 0/8 | 0/14 | ||||
| Tenuiphantes tenebricola (Wider, 1834) | 3/0 | 1/0 | ||||
| Thyreostenius parasiticus (Westring, 1851) | 1/0 | |||||
| Trematocephalus cristatus (Wider, 1834) | 3/117 | 0/9 | 0/11 | 0/29 | 13/0 | 2/3 |
| Family Lycosidae | 4.08% | 0.12% | ||||
| Pardosa amentata (Clerck, 1757) | 1/45 | |||||
| Pardosa prativaga (L. Koch, 1870) | 1/1 | |||||
| Piratula hygrophila (Thorell, 1872) | 2/0 | |||||
| Family Mimetidae | 0.08% | 0.49% | 0.08% | |||
| Ero furcata (Villers, 1789) | 1/0 | 4/0 | 1/0 | |||
| Ero sp. (C. L. Koch, 1836) | 0/1 | |||||
| Family Philodromidae | 1.02% | 9.30% | 0.17% | 0.15% | 0.59% | 0.30% |
| Philodromus collinus (C. L. Koch, 1835) | 3/0 | 1/0 | ||||
| Philodromus dispar Walckenaer, 1826 | 1/0 | 1/0 | ||||
| Philodromus emarginatus (Schrank, 1803) | 1/0 | |||||
| Philodromus praedatus (O. Pickard-Cambridge, 1871) | 1/0 | |||||
| Philodromus sp. Walckenaer, 1826 | 0/9 | 0/91 | 0/2 | 0/1 | 0/4 | 0/4 |
| Tibellus oblongus (Walckenaer, 1802) | 1/0 | 1/0 | ||||
| Family Pisauridae | 0.76% | 12.59% | 0.30% | 2.58% | 0.85% | |
| Dolomedes fimbriatus (Clerck, 1757) | 1/2 | 7/137 | 0/4 | 3/19 | 3/10 | |
| Pisaura mirabilis (Clercka, 1757) | 0/6 | 0/4 | 1/0 | |||
| Family Salticidae | 0.17% | 0.23% | ||||
| Pseudicius encarpatus (Walckenaer, 1802) | 1/0 | 1/0 | ||||
| Salticidae un. | 0/1 | |||||
| Salticus cingulatus (Panzer, 1797) | 1/0 | |||||
| Family Segestriidae | 0.09% | 0.06% | ||||
| Segestria senoculata (Linnaeus, 1758) | 0/1 | 0/1 | ||||
| Family Tetragnathidae | 22.22% | 33.46% | 51.28% | 31.97% | 57.14% | 58.22% |
| Metellina mengei (Blackwall, 1869) | 8/0 | 34/0 | 5/1 | 12/0 | 0/21 | 12/34 |
| Metellina merianae (Scopoli, 1763) | 1/0 | |||||
| Metellina segmentata (Clerck, 1757) | 9/0 | 1/0 | 17/0 | 10/0 | 19/15 | 21/9 |
| Metellina sp. Chamberlin & Ivie, 1941 | 0/68 | 0/213 | 0/24 | 0/66 | 0/5 | 0/7 |
| Pachygnatha clercki Sundevall, 1823 | 39/20 | 67/13 | 135/4 | |||
| Pachygnatha degeeri Sundevall, 1830 | 2/0 | |||||
| Pachygnatha listeri Sundevall, 1830 | 62/5 | 71/0 | 21/13 | 129/32 | 18/27 | 66/26 |
| Pachygnatha sp. Sundevall, 1823 | 0/16 | 0/3 | ||||
| Tetragnatha dearmata Thorell, 1873 | 1/0 | 5/0 | 21/0 | 31/0 | ||
| Tetragnatha extensa (Linnaeus, 1758) | 1/0 | 0/0 | ||||
| Tetragnatha montana Simon, 1874 | 1/0 | 26/0 | 19/0 | 51/0 | 52/0 | |
| Tetragnatha pinicola L. Koch, 1870 | 1/- | |||||
| Tetragnatha sp. Latreille, 1804 | 0/93 | 0/19 | 0/436 | 0/148 | 0/231 | 9/547 |
| Family Theridiidae | 14.50% | 5.48% | 4.08% | 3.33% | 1.64% | 2.25% |
| Enoplognatha ovata (Clerck, 1757) | 49/100 | 13/34 | 28/12 | 18/21 | 8/4 | 23/0 |
| Episinus angulatus (Blackwall, 1836) | 0/7 | 1/1 | 0/1 | |||
| Neottiura bimaculata (Linnaeus, 1767) | 4/0 | 0/1 | ||||
| Paidiscura pallens (Blackwall, 1834) | 1/0 | 1/0 | 2/0 | |||
| Parasteatoda lunata (Clerck, 1757) | 1/0 | |||||
| Platnickina tincta (Walckenaer, 1802) | 0/1 | 1/0 | ||||
| Robertus neglectus (O. Pickard-Cambridge, 1871) | 13/0 | |||||
| Rougathodes instabilis (O. Pickard-Cambridge, 1871) | 1/0 | |||||
| Theridion sp. Walckenaer, 1805 | 0/6 | 0/2 | 0/3 | 0/9 | ||
| Theridion varians Hahn, 1833 | 2/0 | 1/0 | 1/0 | |||
| Family Theridiosomatidae | 0.30% | 0.12% | 0.49% | |||
| Theridiosoma gemmosum (L. Koch, 1877) | 1/3 | 1/0 | 8/0 | |||
| Family Thomisidae | 5.60% | 2.45% | 3.06% | 1.52% | 1.41% | 0.67% |
| Diaea dorsata (Fabricius, 1777) | 0/50 | 0/10 | 0/3 | 1/11 | 1/7 | 1/7 |
| Heriaeus graminicola (Doleschall, 1852) | 1/1 | |||||
| Misumena vatia (Clerck, 1757) | 0/1 | 0/18 | ||||
| Ozyptila sp. Simon, 1864 | 0/9 | 0/3 | ||||
| Psammitis sabulosa (Hahn, 1832) | 1/0 | |||||
| Xysticus cristatus (Clerck, 1757) | 2/0 | 1/0 | ||||
| Xysticus sp. C. L. Koch, 1835 | 0/6 | 0/12 | 0/13 | 0/5 | 0/2 | 0/2 |
| Xysticus ulmi (Hahn, 1831) | 1/0 | 1/0 | ||||
| Total no. of individuals | 301/878 | 217/805 | 259/917 | 637/683 | 429/425 | 816/826 |
| Total no. of species | 40 | 30 | 50 | 43 | 46 | 54 |
| No. of common species | 19 | 24 | 38 | |||
Table 3.
Species richness estimates and sampling completeness (expressed as percentage in brackets) for plant -dwelling spider assemblages in managed and primeval stands in the Białowieża Forest. Abbreviations: M—managed stand, P—primeval stand, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr, SD—standard deviation.
Table 3.
Species richness estimates and sampling completeness (expressed as percentage in brackets) for plant -dwelling spider assemblages in managed and primeval stands in the Białowieża Forest. Abbreviations: M—managed stand, P—primeval stand, OLH—oak–lime–hornbeam forest, RF—ash–alder riparian forest, AC—alder carr, SD—standard deviation.
| M OLH | P OLH | M RF | P RF | M AC | P AC | |
|---|---|---|---|---|---|---|
| Observed species richness | 40 | 30 | 50 | 43 | 46 | 54 |
| Estimates | ||||||
| Chao1 ± SD | 113 ± 54 (35%) | 76 ± 30 (39%) | 61 ± 8 (82%) | 64 ± 16 (67%) | 91 ± 33 (51%) | 90 ± 24 (60%) |
| Chao2 ± SD | 155 ± 95 (26%) | 54 ± 19 (56%) | 62 ± 8 (81%) | 67 ± 17 (64%) | 65 ± 11 (71%) | 87 ± 20 (62%) |
| Jackknife1 ± SD | 61 ± 6 (66%) | 42 ± 3 (71%) | 65 ± 5 (77%) | 58 ± 4 (74%) | 64 ± 5 (72%) | 75 ± 5 (72%) |
| Jackknife2 | 79 (51%) | 51 (59%) | 71 (70%) | 68 (63%) | 73 (63%) | 87 (62%) |
| Michaelis–Menten | 46 (87%) | 39 (77%) | 61 (82%) | 49 (88%) | 56 (82%) | 61 (89%) |
Table 4.
Results of generalized linear models assessing the effect of forest type and forest management on spider abundance and species richness.
Table 4.
Results of generalized linear models assessing the effect of forest type and forest management on spider abundance and species richness.
| Effect | Wald χ2 | df | p |
|---|---|---|---|
| Spider abundance | |||
| Intercept | 6711.09 | 1 | <0.001 |
| Forest type | 1.18 | 2 | 0.572 |
| Managed/Primeval | 4.30 | 1 | 0.038 |
| Forest type x Managed/Primeval | 10.02 | 2 | 0.007 |
| Number of species | |||
| Intercept | 1380.93 | 1 | <0.001 |
| Forest type | 57.27 | 2 | <0.001 |
| Managed/Primeval | 0.001 | 1 | 0.973 |
| Forest type x Managed/Primeval | 14.71 | 2 | 0.001 |
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Stańska, M.; Stański, T.; Patoleta, B. Plant-Dwelling Spider Assemblages in Managed and Protected Primeval Deciduous Stands of Białowieża Forest, Poland. Forests 2025, 16, 1093. https://doi.org/10.3390/f16071093
AMA Style
Stańska M, Stański T, Patoleta B. Plant-Dwelling Spider Assemblages in Managed and Protected Primeval Deciduous Stands of Białowieża Forest, Poland. Forests. 2025; 16(7):1093. https://doi.org/10.3390/f16071093
Chicago/Turabian StyleStańska, Marzena, Tomasz Stański, and Barbara Patoleta. 2025. "Plant-Dwelling Spider Assemblages in Managed and Protected Primeval Deciduous Stands of Białowieża Forest, Poland" Forests 16, no. 7: 1093. https://doi.org/10.3390/f16071093
APA StyleStańska, M., Stański, T., & Patoleta, B. (2025). Plant-Dwelling Spider Assemblages in Managed and Protected Primeval Deciduous Stands of Białowieża Forest, Poland. Forests, 16(7), 1093. https://doi.org/10.3390/f16071093
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