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The European Zoological Journal
ISSN: (Print) (Online) Journal homepage: www.tandfonline.com/journals/tizo21
The influence of humidity gradient in a forest on
ground spider assemblages: a preliminary study
E. Ludwiczak, E. Topa, M. Nietupski, A. Kosewska & R. Rozwałka
To cite this article: E. Ludwiczak, E. Topa, M. Nietupski, A. Kosewska & R. Rozwałka (2024) The
influence of humidity gradient in a forest on ground spider assemblages: a preliminary study,
The European Zoological Journal, 91:2, 1104-1119, DOI: 10.1080/24750263.2024.2403537
To link to this article: https://doi.org/10.1080/24750263.2024.2403537
© 2024 The Author(s). Published by Informa
UK Limited, trading as Taylor & Francis
Group.
Published online: 07 Oct 2024.
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The inuence of humidity gradient in a forest on ground spider
assemblages: a preliminary study
E. LUDWICZAK
1
, E. TOPA
1
, M. NIETUPSKI
1
*, A. KOSEWSKA
1
, & R. ROZWAŁKA
2
1
Department of Entomology, Phytopathology and Molecular Diagnostics, University of Warmia and Mazury, Olsztyn,
Poland, and
2
Department of Zoology and Animal Ecology, University of Life Sciences, Lublin, Poland
(Received 22 January 2024; accepted 1 September 2024)
Abstract
A forest is a dynamic formation developed through evolutionary transformations that have progressed over a long time.
Bioindication can help us understand changes occurring in a forest. Spiders play a signicant role in biomonitoring. The
presence of stenotopic spiders is affected by several factors, of which water availability is vital. According to the assumptions
underlying water retention programs, water accumulated in retention reservoirs is expected to stimulate higher biodiversity of
habitats. An analysis of the effect of a moisture gradient on the appearance of epigeic spiders in the Dąbrówka Forest
Subdistrict has demonstrated the highest species diversity in the transect located closest to retention reservoirs (93 species).
The species diversity of epigeic spiders decreased with the growing distance from these water reservoirs (transect B: 84 species;
transect C: 70). Inverse relationships were determined with regard to the numbers of captured spiders (transect A: 927
specimens; B: 1134; C:1288). The spider species that dominated quantitatively was the forest species Pardosa lugubris, caught
most abundantly in the area assigned to transect C (76.28%). A higher humidity gradient of a habitat (transects A and B)
favors the occurrence of characteristic species with specic habitat and moisture requirements, which are ecologically valuable.
Keywords: Afforestation, Araneae, bioindicators, managed forests, species diversity,, water availability
Introduction
Forest biocenosis results from interactions between
humans and the natural environment that have evolved
over millennia (Grantham et al. 2020; Muigg & Tegel
2021). In the past, Poland’s territory was mainly cov-
ered by forests. As a result of historical, socio-economic
processes primarily aimed at achieving economic goals,
our country’s forests were transformed, and their total
area was reduced (Łoniewicz 1990; State Forests 2018;
National Forest Holding State Forests 2020). The main
cause of deforestation was the development of agricul-
ture. Initially, agricultural use of the land resulted in
only minor changes to the landscape (Tryjanowski
et al. 2011; Munteanu & Cocean 2020; Muigg &
Tegel 2021). It was not until the so-called agricultural
revolution (16th to 19th centuries) that permanent,
often irreversible changes occurred, including the long-
term deforestation of large areas (Tryjanowski et al.
2011). As a result, Poland’s forest cover, about 40% of
the country’s total area in the 17th century, decreased to
20% in 1945 (State Forests 2018).
In the face of a serious ecological threat, the concept of
environmental protection was developed, recognizing
the need to preserve forest ecosystems (Góral &
Rembisz 2017; Tahat et al. 2020; Krawczyk et al. 2021;
Mueller et al. 2021). Under suitable conditions, defor-
estation is, to some extent, a reversible process (Chazdon
et al. 2020). Therefore, the “National Programme for
Increasing the Forest Area of the Country” was launched
to increase the forest area in Poland. The main goal was
to improve the forest area to 30% by 2020 and 33% by
2050 (Ministry of Environmental Protection 1995). The
main course of action, other than planting stands of trees,
is to ensure optimal conditions for the growth and devel-
opment of forested areas.
*Correspondence: M. Nietupski, Department of Entomology, Phytopathology and Molecular Diagnostics, University of Warmia and Mazury in Olsztyn,
Prawocheńskiego 17, Olsztyn 10-719, Poland. Email: mariusz.nietupski@uwm.edu.pl
The European Zoological Journal, 2024, 1104–1119
https://doi.org/10.1080/24750263.2024.2403537
© 2024 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which
permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. The terms on which this article has been
published allow the posting of the Accepted Manuscript in a repository by the author(s) or with their consent.
Water is considered one of the most critical habi-
tat-forming factors, which directly shapes the stabi-
lity of forest ecosystems (Pierzgalski 2012;
Grajewski et al. 2013). However, this is not a one-
sided impact but a series of mutual interactions.
Water models forest ecosystems, while forests pos-
sess a high natural ability to retain water and are an
integral part of the global water cycle (Pierzgalski
2008; Liberacki et al. 2015; Food and Agriculture
Organization 2021). Forest ecosystems character-
ized by higher water retention capacity are taken
advantage of to counteract water shortages
(Pierzgalski 2012). Poland belongs to these
European countries that are the poorest in water
resources, necessitating measures to improve their
water retention capacity (Thier 2020).
For this reason, a national project called
“Increasing the country’s water retention capacity
and preventing oods and droughts in forest ecosys-
tems in lowland regions”. One of the main goals of
this project is to prevent water shortages
(Mioduszewski & Pierzgalski 2009; Kobus 2013).
The measures undertaken in the framework of this
project, known collectively as small water retention,
include a series of technical and non-technical solu-
tions. The underlying assumption is to improve the
water balance while protecting the natural value of
habitats (Zabrocka-Kostrubiec 2013).
Ongoing changes on former agricultural land that
has been reforested are evaluated based on the
response of living organisms (Košulič et al. 2021).
Spiders are used as model organisms in ecological
diversity research; therefore, arachnofauna plays
a signicant role in biomonitoring. Species-specic
traits of analyzed arthropods are increasingly scruti-
nized to gain better insight into how ecosystems
function. These organisms allow researchers to
determine the direction of ongoing changes in
terms of both biogeography and evolution (Barbaro
et al. 2005; Purchard et al. 2013; Barsoum et al.
2014; Benhadi-Marín et al. 2020; Vymazalová et al.
2021; Polchaninova & Marushchak 2023).
The arachnofauna living in drained afforested
areas has not been thoroughly investigated yet;
hence, the present study has been launched in the
Dąbrówka Forest Subdistrict to ll in this gap in our
knowledge. The objective was to determine the cur-
rent status of assemblages of epigeic spiders on for-
mer farmland turned into woodland, submitted to
a small water retention program. The following
research hypotheses were made:
- the humidity gradient signicantly affects the
quantitative and qualitative composition of the
arachnofauna;
- an increase in habitat humidity leads to the appear-
ance of rare species with greater water demand.
Materials and methods
Study area
The study on the epigeic arachnofauna was con-
ducted in the northeastern part of Poland, in the
Dąbrówka Forest Subdistrict (UTM DE 66). The
research area, which is the property of the State
Treasury, is managed by a state administrative
unit, namely the State Forests (SF) National State
Holding (the Polish acronym: PGL LP).
The State Forests coordinated a nationwide project
to improve moisture conditions: “Increasing retention
possibilities and counteracting oods and droughts in
forest ecosystems in lowland areas”. As part of these
activities, the research area was covered by a land
improvement program entitled “Small retention pro-
gram for the Warmian-Masurian Voivodeship for
2006–2015” (Board of the Warmian-Masurian
Voivodeship 2007; Mioduszewski & Pierzgalski
2009). The design process (including planning drai-
nage systems and obtaining the necessary formal
permits) began in 2010, and the technical implemen-
tation of the task took place in 2011. The main task of
the drainage works was the construction of retention
reservoirs aimed at collecting water in periods of
excess water and the possibility of using it in periods
of drought (Olsztyn Forest District 2014).
Our experiment on epigeic spiders included three
study sites (I, II, III), situated on two sides of articial
water reservoirs created under the project mentioned
above (reservoir “a” and “b”), each having a dual,
irregular bowl and covering an area of 0.07 ha
(Forest Data Bank 2023). The study sites featured
similar habitat conditions. The whole observation
area was located on turfed and afforested former farm-
land. According to the Polish State Forests classica-
tion of habitats, study sites I and II were located on the
border of swamp mixed broadleaf forest (subdivision
07-05-1-01-40-k-00) and fresh mixed coniferous for-
est (subsection 07-05-1-01-40-d-00) (Forest Data
Bank 2023). The site I was located 30 meters from
reservoir “a” and 58 meters from reservoir’ b’. Site II
was 20 meters away from site I, 25 m from water
retention reservoir “a, 93 m from reservoir” b’, and
128 m from a sandy road. In line with the classica-
tion of forest habitats according to moisture condi-
tions, the mixed forest stand (4.10 ha) growing on
peat soils and transitional bogs was classied as
swampy and very wet (Forest Data Bank 2023).
This area was turfed and afforested; the undergrowth
was dominated by White willow (Salix alba L.), while
Humidity gradient and forest ground spiders 1105
the dominant species among the trees were Silver
birch (Betula pendula Roth) (22 years old) and Aspen
poplar (Populus tremula L.) (32 years old). The other
forest site, representing fresh mixed coniferous forest,
covered 18.28 ha, grew on typical rusty soil, and was
characterized by fresh moisture. The following four-
teen-year-old tree species represented the forest
canopy layer with its intermediate crown cover and
density: the dominant Scots pine (Pinus sylvestris L.),
Silver birch (Betula pendula Roth), and European
larch (Larix decidua Mill.). A group of admixture
species consisted of oak (Quercus L.), Black alder
(Alnus glutinosa (L.) Gaertn.), Small-leaved lime
(Tilia cordata Mill.), Field elm (Ulmus minor Mill.),
and Norway spruce (Picea abies (L.) H. Karst) (Forest
Data Bank 2023).
The third research site (III) was situated on the
border between the swamp mixed broadleaf forest,
which belonged to the same subdivision as in the
case of sites I and II, and the fresh mixed coniferous
forest (2.93 ha), located in another subdivision,
namely 07-05-1-01-40-a. This site was east of both
retention reservoirs (“a” and “b”). Site III was
located 100 m from Site I, 75 m from Site II, 47 m
from the sandy road, 26 m from retention reservoir
“a”, and 35 m from reservoir “b” (Forest Data Bank
2023). The broadleaf forest tree stand with
intermediate compactness and density, growing on
proper rusty soils, was composed of the dominant
Scots pine (Pinus sylvestris L.). Other species added
to the forest composition were Norway spruce (Picea
abies L.), oak (Quercus L.), Silver birch (Betula pen-
dula Roth), European larch (Larix decidua Mill.),
Black alder (Alnus glutinosa Gaertn.), Sycamore
(Acer pseudoplatanus L.) and Small-leaved lime
(Tilia cordata Mill) (Forest Data Bank 2023).
Data collection
The study was conducted from 27 April to
26 October (181 days) in 2012. It included contin-
uous insect captures. Three transects (A, B, and C)
were set in each research site (I, II, III). Five soil
traps were set up at 10-m intervals along each trans-
ect (3 sites × 3 transects × 5 traps). In total, obser-
vations were based on 45 traps (Figure 1).
Different moisture levels characterized the trans-
ects in all sites (I, II, and III). Transect A, the
wettest, was located in the mixed swamp forest.
Transect B, lying 20 m away from transect A, was
situated in an ecotone. Another 20 m away, in the
mixed broadleaf forest, there was the least wet trans-
ect, denoted as transect C (Figure 1) (Forest Data
Bank 2023).
study site
transects
water retention reservoirs
a, b
I, II, III
20 m
10 m
Soil trap arrangement diagram.
Olsztyn Forest District
soil traps
A, B, C
Legend
Poland
ABC
Figure 1. Approximate localization of the study sites (I, II, III), water retention reservoirs (a, b), and the transects set up in the research
(A–C). Source: Olsztyn Forest District.
1106 E. Ludwiczak et al.
A replication consisted of one trap from every
capture in a given research site. This study on epi-
geic arachnofauna involved using modied traps
composed of plastic containers with a capacity of
500 ml and a height of 12 cm dug into the soil.
The containers were lled up to 1/3 capacity with
ethylene glycol, buried at a level with the soil sur-
face, and emptied every 14 days.
For the soil moisture measurements in 2012, we
used an electronic soil hydrometer type LB-797
tted with two needle electrodes. This device
allowed us to take accurate measurements, which
we carried out four times (June, July, August, and
September). Over an area of 0.5 m
2
around each
trap, we took ve soil moisture measurements and
used them to calculate the average moisture content.
The highest average moisture content was deter-
mined in transect A (37%), lower in B (30%), and
the weakest in C (25%).
The assessment of meteorological conditions was
based on measurements and observations carried
out in the Meteorological Station in Olsztyn, located
at 133 m above sea level (Central Statistical Ofce
2013). The average annual temperature in year
2012 was + 7.5°C (January: 0.5°C, February:
2.8°C, March: 2.8°C, April: 7.8°C, May: 12.8°C,
June: 16.9°C, July: 18.2°C, August: 17.3°C,
September: 11.9°C, October: 8.8°C, November:
4°C, December 0.1ºC). Amounts of precipitations
also varied in the course of these 12 months,
increasing in the summer (January: 66 mm,
February: 37 mm, March: 34 mm, April: 73 mm,
May: 57 mm, June: 96 mm, July: 105 mm, August:
44 mm, September: 42 mm, October: 72 mm,
November: 51 mm, December: 23 mm) (Central
Statistical Ofce 2013).
Data analysis
The observations of Araneae were supported by the
quantitative method, including the analysis of the
number of specimens and species. The Shannon–
Wiener index (H’) analyzed the biological diversity
of captured bioindicators, while the Pielou index (J)
was used to determine the evenness of distribution
of the analyzed species. The ecological analysis was
performed considering the selected preferences:
forest (F), meadow (Md), hygrophilous (H), ther-
mophiles (T), and mesophiles (Me) (Hänggi et al.
1995; Rozwałka & Stachowicz 2021). The struc-
ture of dominance of the captured spiders was pre-
sented according to Górny & Grüm (1981). The
factorial analysis of the similarities of assemblages
of spiders captured in the researched area was
made using a nonmetric multidimensional scaling
(NMDS). The statistical signicance of the results
was veried by analyzing similarities (ANOSIM)
(Kenkel & Orloci 1986; Clarke 1993). The general-
ized linear model (GLM) was employed to investi-
gate the signicance of differences between the
assemblages of Araneae, which allowed us to deter-
mine the p-value. The tests of the importance of
effects in the model were carried out according to
the Wald statistics. In addition, an RDA analysis
was performed to evaluate the structure of assem-
blages of spiders (Ter Braak & Smilauer 1998).
Additionally, an analysis of indicator species
(IndVal) was performed to specify species charac-
teristics for individual transects (A, B, C) with the
“labdsv” package using R software (Dufrêne &
Legendre 1997; Roberts 1997). The statistical test
for this analysis was performed for each species
independently. Species indicator values were
expressed as percentages (from 0 to 100%). The
p-value determining statistical signicance was
determined based on permutations of the species’
position in the group. The statistical signicance of
indicator values was estimated using the Monte
Carlo permutations test. All statistical calculations
and their visual interpretation were conducted
using the following software programs: Statistica
13.3, Canoco 4.51, and PAST 2.01.
Results
During the observations, 3,349 spiders representing
125 species were caught (Table I). A total of 927
specimens were captured in three study sites (I, II,
II) along transect A. Along transect B, representing
the ecotone transect, 1,134 specimens of these
arthropods, classied to 84 species, were captured.
Most spiders were caught in the broadleaf forest (C)
(1,288), but the species richness there was the low-
est (70 species). The species accumulation curves
for each transect, shown in Figure 2, conrmed
adequate sampling effort. Differences in the number
of caught spider species and individuals between the
transects were signicant (Table II). The lowest
average number of caught Araneae individuals per
trap was found in transect A (6.44 specimens). For
comparison, this value was 7.88 in transect B and
8.94 (the highest) in transect C (Figure 3(a)). The
species richness of spiders displayed a contrary pat-
tern. The highest value (on average, 3.1 species
captured into one trap) was in transects A and
B. Signicantly fewer species were captured into
the traps along transect C (2.1 species on average)
(Figure 3(b)).
The calculated value of the Shannon - Wiener
index, helpful in assessing a given habitat’s
Humidity gradient and forest ground spiders 1107
Table I. Species composition, abundance, and ecological characteristics of ground-dwelling spiders occurring in the studied transects (A,
B, C). Ecological description: forest (F), meadow (Md), hygrophilous (H), thermophiles (T), mesophiles (Me).
Ecological Studied transects (A, B, C) in three study sites (I, II, III)
Family Species Description Abbreviation A B C
I II III I II III I II III
Liocranidae Agroeca brunnea F/Me Agr_brun 3 3 0 1 3 0 7 4 9
Liocranidae Agroeca proxima Md/T Agr_prox 4 0 1 1 3 1 0 0 1
Linyphiidae Agyneta afnis Md/Me Agy_aff 1 0 1 2 3 0 1 0 0
Linyphiidae Agyneta ramosa F/H Agy_ram 1 0 1 0 0 0 0 0 2
Linyphiidae Agyneta rurestris Md/Me Agy_rur 0 0 1 0 0 0 0 0 0
Linyphiidae Agyneta saxatilis Md/Me Agy_sax 0 1 0 1 1 0 0 3 0
Linyphiidae Allomengea scopigera Md/H All_sco 0 0 2 0 0 0 0 0 0
Linyphiidae Allomengea vidua Md/H All_vid 3 11 0 0 2 1 0 0 0
Lycosidae Alopecosa cuneata Md/Me Alo_cune 12 14 6 47 22 7 5 1 1
Lycosidae Alopecosa pulverulenta Md/Me Alo_pulv 14 32 6 42 32 3 0 0 0
Linyphiidae Anguliphantes angulipalpis F/Me Ang_angu 0 0 0 0 0 0 0 1 1
Hahniidae Antistea elegans Md/H Ant_eleg 3 0 1 1 1 1 0 0 0
Linyphiidae Araeoncus humilis Md/Me Ara_hum 0 0 0 0 0 0 0 0 1
Araneidae Araneus quadratus Md/Me Ara_quad 0 0 0 0 0 1 0 0 0
Linyphiidae Bathyphantes
approximatus
Md/H Bat_appro 1 0 0 0 0 0 0 0 0
Linyphiidae Bathyphantes gracilis Md/Me Bat_grac 3 3 1 0 3 5 0 0 0
Linyphiidae Bathyphantes nigrinus F/H Bat_nigri 0 1 0 0 1 0 0 1 0
Linyphiidae Bathyphantes parvulus Md/Me Bat_parv 6 8 1 4 8 2 3 3 5
Linyphiidae Centromerita bicolor Md/Me Cen_bic 1 0 2 8 1 1 1 0 1
Linyphiidae Centromerus sylvaticus F/Me Cen_sylva 15 10 27 18 11 10 13 3 3
Linyphiidae Ceratinella brevipes F/H Cer_brev 1 1 0 0 1 0 0 0 0
Linyphiidae Ceratinella brevis F/Me Cer_revis 0 1 0 0 0 1 0 0 4
Araneidae Cercidia prominens Md/Me Cer_promi 0 1 0 0 0 0 0 0 0
Clubionidae Clubiona reclusa Md/Me Clu_reclu 1 1 0 0 0 0 0 0 0
Clubionidae Clubiona subtilis F/Me Clu_sub 1 0 0 0 0 0 0 0 0
Clubionidae Clubiona terrestris F/Me Clu_terr 0 0 0 0 0 0 0 0 2
Linyphiidae Dicymbium nigrum F/Me Dic_nigr 0 1 2 3 3 2 1 0 1
Linyphiidae Dicymbium brevisetosum Md/H Dic_nigr_brev 0 1 1 1 0 1 0 1 0
Linyphiidae Dicymbium tibiale F/Me Dic_tib 1 1 2 0 0 0 1 0 0
Linyphiidae Diplocephalus latifrons F/Me Dip_lati 0 0 0 0 0 0 0 1 0
Linyphiidae Diplocephalus picinus F/Me Dip_pici 0 1 2 0 0 0 2 1 7
Linyphiidae Diplostyla concolor F/Me Dip_conc 1 0 1 0 0 0 1 0 1
Gnaphosidae Drassyllus lutetianus F/H Dra_lute 1 0 0 0 1 0 0 0 0
Gnaphosidae Drassyllus praecus Md/T Dra_prae 0 2 0 2 1 0 0 0 0
Gnaphosidae Drassyllus pusillus Md/Me Dra_pusi 3 2 3 1 5 2 2 0 3
Linyphiidae Erigone atra Md/Me Eri_atra 0 0 2 2 5 4 0 0 3
Linyphiidae Erigone dentipalpis Md/Me Eri_dent 0 0 6 8 1 6 0 0 4
Linyphiidae Erigonella hiemalis F/Me Eri_hiem 0 0 1 0 3 2 2 1 4
Linyphiidae Erigonella ignobilis Md/H Eri_igno 1 0 0 0 1 0 0 0 0
Mimetidae Ero furcata F/Me Ero_furc 0 1 0 0 0 0 0 1 0
Salticidae Euophrys frontalis Md/Me Euo_front 2 0 1 0 1 0 2 0 2
Salticidae Evarcha arcuata Md/Me Eva_arcu 0 0 1 1 0 0 0 0 0
Linyphiidae Floronia bucculenata F/Me Flo_bucc 1 0 0 0 0 0 0 0 1
Linyphiidae Gonatium rubens F/Me Gon_rub 7 1 6 0 2 0 0 0 1
Linyphiidae Gongylidiellum murcidum Md/H Gon_murci 0 0 0 0 0 1 1 0 0
Hahniidae Hahnia nava Md/T Hah_nava 1 0 0 1 1 0 0 0 0
Gnaphosidae Haplodrassus signifer Md/Me Hap_signi 0 0 0 2 0 0 0 0 0
Gnaphosidae Haplodrassus silvestris F/Me Hap_silv 0 0 0 0 0 0 4 0 0
Gnaphosidae Haplodrassus soerenseni F/Me Hap_soer 0 0 0 0 0 0 1 0 0
Salticidae Heliophanus avipes Md/Me Hel_avi 0 0 0 1 0 0 0 0 0
Linyphiidae Hylyphantes graminicola Md/H Hyl_grami 0 0 0 0 0 0 1 0 0
Araneidae Hypsosinga sanguinea Md/Me Hyp_sang 2 1 0 2 0 0 0 0 0
Linyphiidae Kaestneria pullata Md/H Kae_pull 1 3 0 0 0 0 0 0 0
Linyphiidae Linyphia triangularis F/Me Lin_trian 1 0 0 0 0 1 0 0 1
Linyphiidae Mansuphantes mansuetus F/Me Man_mans 0 0 0 0 0 0 0 1 0
(Continued )
1108 E. Ludwiczak et al.
Table I. (Continued).
Ecological Studied transects (A, B, C) in three study sites (I, II, III)
Family Species Description Abbreviation A B C
I II III I II III I II III
Tetragnathidae Metellina mengei F/Me Met_meng 0 1 0 0 0 0 0 0 0
Gnaphosidae Micaria pulicaria Md/Me Mic_pulic 4 4 0 4 6 0 1 0 0
Linyphiidae Micrargus apertus F/Me Mic_aper 1 1 0 0 0 0 0 0 0
Linyphiidae Micrargus herbigradus F/Me Mic_herb 0 1 0 0 0 0 0 0 1
Linyphiidae Microlinyphia pusilla Md/Me Mic_pusi 0 0 0 0 0 1 0 0 0
Linyphiidae Microneta viaria F/Me Mic_viar 1 2 2 0 0 0 2 6 17
Salticidae Neon reticulatus F/Me Neo_reti 0 0 0 0 0 0 0 0 1
Linyphiidae Neriene clathrata F/Me Ner_clath 1 0 0 0 0 0 2 0 2
Linyphiidae Neriene montana F/Me Ner_mont 0 0 0 0 0 0 0 0 3
Linyphiidae Neriene peltata F/Me Ner_pelt 0 0 0 0 0 0 0 0 1
Linyphiidae Obscuriphantes obscurus F/Me Obs_obsc 0 0 0 0 0 0 0 2 0
Linyphiidae Oedothorax apicatus Md/Me Oed_apic 0 0 1 0 0 0 0 0 0
Linyphiidae Oedothorax retusus Md/H Oed_retu 1 0 4 0 0 1 1 0 1
Thomisidae Ozyptila atomaria F/Me Ozy_atom 0 0 0 0 1 0 0 0 0
Thomisidae Ozyptila praticola F/Me Ozy_prati 1 0 0 0 0 0 0 0 0
Thomisidae Ozyptila trux Md/H Ozy_trux 5 6 1 2 3 4 0 0 0
Tetragnathidae Pachygnatha clercki Md/H Pach_cle 3 2 2 0 0 1 0 0 1
Tetragnathidae Pachygnatha degeeri Md/T Pach_deg 3 0 2 28 30 21 0 0 2
Tetragnathidae Pachygnatha listeri F/Me Pach_list 0 2 0 1 1 0 6 0 5
Linyphiidae Palliduphantes pallidus F/Me Pall_alut 1 1 0 0 1 0 0 0 1
Lycosidae Pardosa amentata Md/H Par_amen 3 1 25 2 2 0 1 0 0
Lycosidae Pardosa lugubris F/Me Par_lugu 63 20 34 97 38 44 505 151 296
Lycosidae Pardosa paludicola Md/H Par_palu 4 6 0 1 2 1 0 1 0
Lycosidae Pardosa palustris Md/Me Par_palus 3 1 0 5 2 0 0 0 0
Lycosidae Pardosa prativaga Md/Me Par_prat 42 69 22 21 30 19 0 0 1
Lycosidae Pardosa pullata Md/Me Par_pull 41 43 4 81 132 35 0 0 1
Lycosidae Pardosa riparia Md/H Par_rip 0 1 0 0 2 0 0 0 0
Philodromidae Philodromus collinus F/Me Phi_coll 1 0 0 0 0 0 0 0 0
Phrurolithidae Phrurolithus festivus Md/Me Phr_fest 0 1 0 0 1 5 2 0 3
Lycosidae Pirata piraticus Md/H Pir_pira 0 2 0 0 1 0 0 0 0
Lycosidae Pirata piscatorius Md/H Pir_pisca 0 0 0 0 0 1 0 0 0
Lycosidae Piratula uliginosa Md/H Pir_ulig 0 0 0 0 1 0 0 0 0
Lycosidae Piratula hygrophila Md/H Pir_hygr 0 0 0 0 0 1 0 0 1
Linyphiidae Pocadicnemis juncea Md/Me Poc_jun 3 6 1 0 3 5 1 0 3
Linyphiidae Pocadicnemis pumila Md/Me Poc_pum 0 1 1 1 0 1 0 0 0
Theridiidae Robertus arundineti F/H Rob_arun 0 0 1 0 0 2 0 0 0
Theridiidae Robertus lividus Md/Me Rob_livi 3 1 1 0 1 2 0 0 0
Linyphiidae Silometopus reussi Md/H Sil_reu 0 0 0 1 0 0 0 0 0
Linyphiidae Stemonyphantes lineatus F/Me Ste_line 0 0 0 0 0 2 0 0 0
Linyphiidae Tallusia experta F/H Tal_expe 2 0 1 0 1 0 0 0 0
Linyphiidae Tenuiphantes avipes F/Me Tan_avi 0 0 0 0 0 0 0 0 1
Linyphiidae Tapinocyba insecta F/Me Tap_ins 1 4 1 0 1 0 4 1 2
Linyphiidae Tapinocyba pallens F/Me Tap_pall 0 0 1 0 0 0 1 0 0
Linyphiidae Tapinopa longidens F/Me Tap_longi 1 0 0 0 0 0 0 0 0
Linyphiidae Taranucnus setosus Md/H Tar_set 1 0 0 0 0 0 0 0 0
Linyphiidae Tenuiphantes mengei F/Me Ten_men 16 15 3 6 1 18 13 8 7
Linyphiidae Tenuiphantes tenebricola F/Me Ten_tene 0 0 0 0 0 0 0 2 1
Tetragnathidae Tetragnatha extensa Md/Me Tet_exte 0 0 0 1 0 0 0 0 0
Philodromidae Thanatus arenarius Md/T Tha_aren 0 0 0 0 1 0 0 0 0
Philodromidae Thanatus striatus Md/H Tha_stria 0 1 0 0 1 0 0 0 0
Linyphiidae Tiso vagans Md/Me Tis_vag 0 0 0 2 0 1 0 0 0
Lycosidae Trochosa ruricola Md/Me Tro_ruri 17 13 11 29 11 12 4 1 6
Lycosidae Trochosa spinipalpis Md/H Tro_spin 4 15 2 4 3 0 0 0 0
Lycosidae Trochosa terricola F/Me Tro_terri 19 14 7 15 18 10 19 12 21
Linyphiidae Walckenaeria acuminata F/Me Wal_acu 0 1 0 0 0 1 0 0 0
Linyphiidae Walckenaeria alticeps F/H Wal_alti 0 1 0 0 1 0 0 0 3
Linyphiidae Walckenaeria antica F/Me Wal_anti 1 1 0 0 0 2 1 3 0
(Continued )
Humidity gradient and forest ground spiders 1109
biodiversity, proved to be the highest in transect
A (H’ = 3.4) (Table I). It was lower (H’ = 2.99) in
the ecotone’s transect (B) and the weakest in
transect C (H’ = 1.49). Differences in these values
of H’ between the transects, calculated for each
trap as a replication, were statistically signicant
(Table II, Figure 3(c)). An identical relationship
was determined for the computed values of the
Pielou evenness index (J’), which expresses the
ratio of actual diversity to maximum diversity (A,
J’ = 0.75; B, J’ = 0.67; C, J’ = 0.35) (Table I, Table
II, Figure 3(c)).
The species that occurred most numerously was
Pardosa lugubris (Walck.), with 1,248 individuals
captured during the study (Table I). This species
appeared in the highest number in transect C (952
specimens), while fewer individuals were caught in
the mixed forest (transect A − 117 specimens) and
in the ecotone (transect B − 179 individuals).
The second most abundant species was Pardosa pull-
ata (Clerck) (337 indiv.), which was most numerous
in the ecotone (transect B − 248 indiv.). The third
most numerous species (204 indiv.) in the research
transects was Pardosa prativaga (L. Koch). Other
species which appeared in considerable numbers
were: Trochosa terricola Thorell (135 indiv.),
Alopecosa pulverulenta (Clerck) (129), Alopecosa
cuneata (Clerck) (115), Centromerus sylvaticus
(Blackwall) (110) and Trochosa ruricola (De Geer)
(104). P. lugubris is a species that dominated quan-
titatively in transect C, creating a group of super-
dominants that composed 73.91% of all this
assemblage (Table III). The most numerous species
in transect A was P. prativaga (14.35%) and
P. lugubris (12.62%), composed of the class of eudo-
minants. In the ecotone (B), a class of eudominants
also comprised P. lugubris (15.79%) and P. pullata
(21.88%). In transect C, no species could be classi-
ed as eudominant or dominant. In more moist
transects A and B, the dominant species group was
represented by 3 and 4 species, respectively.
Transect A was also characterized by an even dis-
tribution of species in the classes of subdominants
and recedents, as well as the most numerous species
group, which did not exceed 1% of the total number
of spiders (subrecedents). For transect B, a similar
distribution of species within dominance classes was
observed, whereas transect C showed clear imbal-
ances in the composition of dominance classes
(Table III).
The analyzed transects differed signicantly in the
species composition of the spider assemblages,
which was conrmed by the applied nonmetric mul-
tidimensional scaling (NMDS). The results were
Table I. (Continued).
Ecological Studied transects (A, B, C) in three study sites (I, II, III)
Family Species Description Abbreviation A B C
I II III I II III I II III
Linyphiidae Walckenaeria atrotibialis F/Me Wal_atro 2 2 2 0 0 4 2 5 1
Linyphiidae Walckenaeria dysderoides F/Me Wal_dysd 0 1 0 0 0 0 0 0 0
Linyphiidae Walckenaeria nudipalpis F/H Wal_nudi 1 0 0 0 0 0 0 0 0
Linyphiidae Walckenaeria unicornis F/H Wal_unic 1 3 2 0 0 2 0 0 1
Lycosidae Xerolycosa miniata Md/T Xer_min 0 0 2 3 7 0 0 0 0
Thomisidae Xysticus audax F/T Xys_aud 0 0 0 1 0 0 0 0 0
Thomisidae Xysticus cristatus Md/Me Xys_crist 0 1 1 0 1 1 0 0 0
Thomisidae Xysticus kochi Md/Me Xys_koch 0 0 0 1 0 0 0 0 0
Thomisidae Xysticus ulmi Md/H Xys_ulmi 3 1 2 0 0 1 1 0 0
Gnaphosidae Zelotes electus Md/T Zel_elec 0 0 0 1 0 0 0 0 0
Gnaphosidae Zelotes latreillei Md/Me Zel_latr 1 6 5 4 1 3 4 0 0
Miturgidae Zora nemoralis Md/T Zor_nem 0 0 0 0 0 0 0 1 0
Miturgidae Zora spinimana Md/Me Zor_spin 4 1 5 0 0 1 0 5 9
Total Individuals 350 353 224 460 420 254 618 220 450
927 1134 1288
Total species 62 60 52 44 55 48 35 26 49
93 84 70
Shannon H’ Log Base 2.718 3,197 3,158 3,208 2,703 2,789 3,085 1,03 1,519 1,802
3,403 2,987 1,485
Pielou J’ 0,775 0,771 0,812 0,714 0,696 0,797 0,29 0,466 0,463
0,751 0,674 0,35
1110 E. Ludwiczak et al.
plotted in a diagram (Figure 4), illustrating the simi-
larities and differences in the species composition
and number of spiders in the analyzed transects.
The ANOSIM assays conrmed signicant differ-
ences between the analyzed assemblages of spiders (R:
0.96; p: 0.0001), which occupy distant positions from
each other on the ordination diagram (Figure 4).
The assemblages of spiders living along the ana-
lyzed transects were submitted to an ecological
analysis, distinguishing the species characteristic for
forests and meadows. Transect A, situated on peat-
bog soils and transient peatlands, was populated
mainly by meadow species (55.8%), although
a large percentage of forest species (44.2%) was
also observed (Figures 5(a), 5(b)). In the ecotone
zone (transect B), meadow species were dominant
(Md − 62.7%), while typical forest species were
fewer (37.3%) than in transect A, located further
away from the broadleaf forest. The third research
transect was dominated by forest spider species
(83.3%), and the meadow species settling in this
site composed 16.7% of the whole community.
Another ecological division is applied to the
described assemblages of spider species associated
with very wet areas (H) or areas well exposed to
sunlight (T). The third group in this division
included ubiquitous species highly tolerant to differ-
ent moisture levels in a habitat (Me). The represen-
tatives of this group were prevalent in the three
analyzed research areas, being the most numerous
in transect C (95.4%) (Figure 6(a)). Many smaller
species were caught in the ecotone (B − 81.9%) and
very wet transect (A– 74.0%). Thermophilic species
most numerously settled in the ecotone between the
broadleaf and mixed forest (B − 8.9%), while com-
posing 4.5% in the mixed forest and only 2.4% in
transect C (Figure 6(b)). The presence of hygrophi-
lous species was consistent with the distribution of
the moisture gradient. This group was most numer-
ous (20.6%) in transect A, which was classied as
swampy and very wet according to the adopted clas-
sication of moisture levels. It was followed by the
ecotone transect (B − 9.2%) and the broadleaf forest
(C − 2.2%) (Figure 6(c)).
More detailed relationships between the ana-
lyzed assemblages of spiders versus the distin-
guished environmental variables are shown in the
RDA diagram (Figure 7). This diagram displays
the differentiation of assemblages of these arthro-
pods within the analyzed habitats. The chart
Figure 2. Expected number of spider species caught in studied
transects (A, B, and C) using the Jackknife 2 estimator (±SD) of
species richness.
Table II. Results of the GLM test of signicance (Wald statistic)
of the tested transects (A, B, C) on the abundance and species
richness of spiders and abundance of its ecological groups.
df Wald Statistic p
Abundance 2 58.39 0.00
Richness 2 35.92 0.00
Shannon H’ 2 6.74 0.03
Forest (F) 2 696.79 0.00
Meadow (Md) 2 388.84 0.00
Hygrophilous (H) 2 97.06 0.00
Thermophiles (T) 2 82.70 0.00
Mesophiles (Me) 2 129.62 0.00
Humidity gradient and forest ground spiders 1111
shows the distribution of captured spiders and
their ecological groups versus the environmental
gradients represented by two principal canonical
axes, describing 76.0% of the variance. It also
considers the location of the analyzed transects
relative to the distinguished environmental vari-
able (moisture). The factor differentiating the
described assemblages of spiders is the moisture
gradient, which groups hygrophilous species of
transect A in the upper left-hand quarter of the
RDA diagram. The rst ordination axis, describ-
ing 74,1% of the variance, strongly correlates with
transect C, forest, and mesophilic species. The
transect located in the ecotone (B) occupies the
Figure 3. Mean values of abundance (3a), species richness (3b), and Shannon H’ (3c) of spider assemblages in the three transects (A, B,
C) (vertical lines denote SE).
Table III. The most abundant spider species were collected in three transects (A, B, C). The Górny and Grüm (1981) the scale of
dominance was used.
Dominance Class
Transect A Transect B Transect C
Superdominant species (>30%)
Pardosa lugubris (73.91%)
Eudominant species (10–30%)
Pardosa prativaga (14.35%) Pardosa pullata (21.88%)
Pardosa lugubris (12.62%) Pardosa lugubris (15.79%)
Pachygnatha degeeri (6.97%)
Alopecosa pulverulenta (6.79%)
Alopecosa cuneata (6.70%)
Pardosa prativaga (6.17%)
Dominant species (5–10%)
Pardosa pullata (9.49%)
Alopecosa pulverulenta (5.61%)
Centromerus sylvaticus (5.61%)
Subdominant species (2–5%)
Trochosa ruricola (4.42%) Trochosa ruricola (4.59%) Trochosa terricola (4.04%)
Trochosa terricola (4.31%) Trochosa terricola (3.79%) Tenuiphantes mengei (2.17%)
Tenuiphantes mengei (3.67%) Centromerus sylvaticus (3.44%)
Alopecosa cuneata (3.45%) Tenuiphantes mengei (2.21%)
Pardosa amentata (3.13%)
Trochosa spinipalpis (2.27%)
Recedent species (1–2%)
Bathyphantes parvulus (1.62%) Erigone dentipalpis (1.32%) Microneta viaria (1.94%)
Allomengea vidua (1.51%) Bathyphantes parvulus (1.24%) Agroeca brunnea (1.55%)
Gonatium rubens (1.51%) Centromerus sylvaticus (1.48%)
Ozyptila trux (1.29%) Zora spinimana (1.09%)
Zelotes latreillei (1.29%)
Pardosa paludicola (1.08%)
Pocadicnemis juncea (1.08%)
Zora spinimana (1.08%)
Subrecedent species (<1%)
The remaining 74 species (20.59%) The remaining 72 species (19.14%) The remaining 63 species (13.82%)
1112 E. Ludwiczak et al.
Figure 4. Diagram of nonmetric multidimensional scaling (NMDS) for spider assemblages about the studied transects (A, B, and C).
Figure 5. Percentages of spider habitat types (forest 5a, meadow 5b) in the analyzed transects (A, B, C) (vertical lines indicate SE).
Figure 6. Percentages of spider habitat types (mesophiles 6a, thermophiles 6b, and hygrophilous 6c) in the analyzed transects (A, B, C)
(vertical lines indicate SE).
Humidity gradient and forest ground spiders 1113
left-hand lower quarter of the ordination diagram
and correlates with meadow (Me) and thermophi-
lics (T) species.
An indicator species analysis (IndVal) was used to
identify signicant (p < 0.05) spider species for
studying transects representing different habitats.
The IndVal method distinguished seven species
characteristics for transect A (Pardosa prativaga,
Gonatium rubens, Allomengea vidua, Trochosa spini-
palpis, Pardosa amentata, Pachygnatha clercki,
Kaestneria pullata), 6 for transect B (Pardosa pullata,
Pachygnatha degeeri, Trochosa ruricola, Alopecosa pul-
verulenta, Alopecosa cuneata, Erigone atra) and 6 for
transect C (Pardosa lugubris, Agroeca brunnea,
Microneta viaria, Pachygnatha listeri, Diplocephalus
picinus, Haplodrassus silvestris) (Table IV).
Discussion
Water is the fundamental factor in maintaining the
ecological balance in nature (Pierzgalski 2008;
European Environment Agency 2015; Gavrilescu
2021). The underlying assumption of water retention
Figure 7. Diagram of the redundancy analysis (RDA) presenting the correlations between the spider’s ecological groups, studied transects,
and the selected habitat factors (moisture). All abbreviations are as given in the methodology section and Table I.
Table IV. Values of the IndVal indicator of the spider species
were signicantly characteristic (p < 0.05) for the study of
transects.
IndVal pFrequency
Transect A
Pardosa prativaga 17.66 0.001 70
Gonatium rubens 5.72 0.003 13
Allomengea vidua 5.15 0.007 11
Trochosa spinipalpis 4.17 0.016 12
Pardosa amentata 4.15 0.047 11
Pachygnatha clercki 3.78 0.019 9
Kaestneria pullata 2.78 0.035 4
Transect B
Pardosa pullata 21.46 0.001 67
Pachygnatha degeeri 12.76 0.001 27
Trochosa ruricola 10.07 0.014 56
Alopecosa pulverulenta 7.88 0.012 33
Alopecosa cuneata 5.97 0.027 26
Erigone atra 4.30 0.011 12
Transect C
Pardosa lugubris 29.14 0.001 135
Agroeca brunnea 6.48 0.008 23
Microneta viaria 6.37 0.006 16
Pachygnatha listeri 4.58 0.016 13
Diplocephalus picinus 4.27 0.012 11
Haplodrassus silvestris 2.78 0.036 4
1114 E. Ludwiczak et al.
programs is to create water reservoirs, which – by
increasing the heterogeneity of landscapes – can stimu-
late higher biodiversity of ora and fauna in an ecosys-
tem (Mioduszewski & Pierzgalski 2009; Mioduszewski
et al. 2015; Futter 2019) and can make ecosystems
more resilient to shocks (Bellone et al. 2017). The
inuence of water on species diversity has been the
subject of numerous studies. Observations on arthro-
pods have shown that building new water reservoirs in
forest ecosystems can inuence the number of species
living in a given habitat and their number, particularly
the share of rare species with more signicant moisture
requirements, often precious (Thiele 1977;
Jędryczkowski & Kupryjanowicz 2005; Nietupski
et al. 2006; Ludwiczak et al. 2020, 2022).
During this study, conducted in the Dąbrówka
Forest Subdistrict, 3,349 specimens of Araneae were
caught. They were assigned to 125 species (Table I),
corresponding to 14.74% of the spider species in
Poland (https://www.araneae.nmbe.ch/ 2023).
Spiders are a species-rich group of animals, of which
nearly half are species typical of forest areas (Oxbrough
& Ziesche 2013). Forest species were also found to
dominate in the studied forest area where a small
water retention program was conducted; they com-
posed 83.3% in transect C, 37.3% in transect B, and
42.2% in transect A (Table I).
The importance of transformations of habitats to the
presence of these arthropods has also been studied
extensively (Ziesche 2008; Hsieh & Linsenmair 2011;
Oxbrough & Ziesche 2013; Košulič et al. 2016; Šipoš
et al. 2017; Bishop et al. 2018; Gallé et al. 2021). The
main objective of the research conducted in the
Olsztyn Forest District has been to determine the effect
of the distance to articially created water reservoirs on
the biodiversity of captured spiders. In their observa-
tions, (Czyżyk & Porter 2018) demonstrated that the
highest species diversity of Carabidae appeared in
a zone nearest to water reservoirs (at a 5-m distance).
Likewise, a study concerning ground beetles con-
ducted in the Dąbrówka Forest Subdistrict proved
the highest species diversity in a research area situated
in the nearest proximity to the water reservoirs and the
ecotone (Ludwiczak et al. 2020, 2022). The qualita-
tive analysis of the captured Araneae showed that the
species diversity decreased as the distance to the water
ponds increased. Most species (93) were caught in
transect A (Table I, Figure 3(b)). A short distance
from forest water reservoirs directly inuences the
greater diversity of ora and fauna found in the area
(Czyżyk & Porter 2018). These results are consistent
with the report delivered by Jędryczkowski and
Kupryjanowicz (2005), who demonstrated that the
species diversity of Carabidae increased directly and
proportionally to the increase in the moisture content
of the habitat. Valuable fauna species were also
detected during studies conducted in the same area
(Staręga et al. 2002). For example, species classied
as endangered ones (Drassyllus praecus (L. Koch),
Micrargus apertus (O. Pickard-Cambridge), Thanatus
arenarius L. Koch, Thanatus striatus CL Koch) were
caught near the water retention reservoirs (transects
A and B) (Table I). M. apertus deserves attention as
this is a new site where it appeared, previously not
indicated in the territory of the Warmińsko-
Mazurskie Province (Wiśniewski et al. 2018).
Moreover, this species was previously characterized
as associated with various types of forests and shrubs.
In our research, it was caught in a meadow adjacent to
the created retention reservoirs. However, the rare and
thermophilic D. praecus, found mainly in dry areas
(Rozwałka & Stachowicz 2021), in our research
occurred in transects with higher water resources
(transects A and B). In transect C, with the least
water resources, D. praecus did not occur (Table I).
This may indicate potential positive consequences of
introducing retention programs on ecological diver-
sity, which we have also shown in previous studies
(Wiśniewski et al. 2018; Ludwiczak et al. 2020,
2022). The occurrence of stenotopic Araneae is
affected by various factors (Purchard et al. 2013).
Košulič et al. showed that differences in greater open-
ness of tree canopy, ensuring better insolation, lead to
a more intricate nature of the habitat and thus affect
epigeic organisms (Košulič et al. 2016). The inuence
of an increase in the shade and moisture levels on
spiders has also been scrutinized by Entling et al.
(Entling et al. 2007), (Ziesche & Roth 2013), and
Cernecká et al. (2019). The analyzed arachnids’ spe-
cies decreased in transect B (84 species) (Table I,
Figure 3(b)). Ecotone areas, lying on the border
between two habitats, are an excellent place for the
foraging and reproduction by insects (Szyszko 2002),
which is conrmed by the study conducted in the
Dąbrówka Forest Subdistrict in 2012 (Table I,
Figure 3). High species diversity of Araneae in eco-
tones has also been proven by Košulič et al. (2016),
Oleszczuk (Oleszczuk 2017), and Stańska et al.
(2018). An inverse relationship, compared to the spe-
cies diversity, was noted when analyzing the abun-
dance of Araneae (transect A: 927 indiv.; transect B:
1134 indiv.; transect C: 1288 indiv.) (Table I,
Figure 3(a)). The observed quantitative and qualita-
tive differences in spider assemblages among the ana-
lyzed transects were signicant (Table II, Figure 4),
also conrmed by the NMDS analysis (Figure 4).
Among the captured arachnofauna, the most
numerous were cosmopolitan species of the genus
Pardosa. The most numerous species in this group
were P. lugubris (1248 indiv.), P. pullata (338 indiv.)
Humidity gradient and forest ground spiders 1115
and P. prativaga (204 indiv.). The comparison of
Araneae dominance along the transects differed in
the distance to the water reservoirs, showing con-
siderable disproportions. The abundance of the for-
est species P. lugubris, dominant in the spider
assemblages, increased directly and proportionally
to the distance to the water reservoirs (transect A:
117 indiv., transect B: 179 indiv., transect C: 952
indiv.). The dominance of the forest species in
transect A may suggest its migratory abilities, for
example, in search of food. In transect C, this spe-
cies achieved the rank of a superdominant
(73.91%). As no eudominants or dominants were
determined in this site, the dominance structure in
the area covered with the fresh mixed broadleaf
forest was sharper. P. lugubris was also dominant in
naturally renewed Scots pine stands (Topa et al.
2021). The dominance of a single species with
high abundance can implicate certain disorders in
a habitat. Disproportions among the shares of the
analyzed species in each community intensied at
a further distance of the analyzed transects from the
water retention reservoirs, which may be associated
with a change in habitat conditions, particularly its
moisture level (Table III). Transect A, nearest the
water reservoirs, was characterized by the species’
richest group of recedents (8 species) and subrece-
dents (74 species). A high number of species with
a relatively small abundance does not undermine
their role in the analyzed biocenosis. On the con-
trary, the number of residents and precedents is
typical of relatively stable ecosystems characterized
by an adequately developed internal structure
(Trojan 1998).
The IndVal analysis specied which spider species
are characteristic of particular habitats (transect A, B,
C) (Table IV). According to Staręga et al. (2002),
P. prativaga is a species typical of open areas (includ-
ing meadows, fallow lands, and dry peat bogs). Our
research conrmed this species is strongly associated
with the most open transect A (Table IV). In the
remaining transects, which were characterized by
more excellent shade, this species was not character-
istic, and only one individual of this species was
caught in the forest area (Tables I, IV). Particularly
noteworthy in transect A are the species characteristic
of this area – A. vidua and K. pullata, which are very
rare, according to Rozwałka and Stachowicz (2021)
(Table IV). This may conrm the benecial effect of
retaining water in forests on shaping biodiversity,
especially naturally valuable species. Our observa-
tions showed ubiquitous species dominated in trans-
ect B, and P. pullata was a species characteristic of the
ecotone zone (transect B). Rozwałka and Stachowicz
(2021) characterize this species as frequently encoun-
tered and heliophilous. Its preferences were con-
rmed in transect C shaded by the tree canopy
(only one individual) (Tables I, IV). The most char-
acteristic species for transect C was P. lugubris,
a common species. However, due to problems with
its diagnosis, Rozwałka and Stachowicz (2021) draw
attention to the need to verify its occurrence cor-
rectly. The species associated mainly with dry ground
(e.g., forests, thickets, orchards, and mid-eld trees)
and also occurred in its typical location during the
research in the Dąbrówka Forest District (the least
humid transect C). However, this species was also
recorded in moderately moist areas (Rozwałka &
Stachowicz 2021). In our observations, it occurs in
large numbers in transects B and A. This may indi-
cate that the mosaic of habitats created after the
introduction of retention reservoirs is benecial for
this species. This knowledge may be helpful when
implementing new forest management practices to
ensure these habitats’ stability, durability, and
diversity.
Summary
Heterogeneity in microhabitat structure is believed
to be the driving force behind animal diversity.
Nature’s heterogeneity, measured at many unique
scales, inuences the structure of Araneae commu-
nities. In turn, the sensitivity of spiders to habitat
conditions facilitates the interpretation of the com-
plex ecology of habitats covered by the small water
retention program. The species diversity of epigeic
spiders is mainly dependent on soil moisture.
Increasing the distance from forest water reservoirs
reduces the species diversity of caught arachnofauna
and affects their numbers. Our research shows that
the created retention reservoirs are a refuge for rare
and previously undetected species with higher
humidity preferences, at risk of extinction
(D. praecus, M. apertus, T. arenarius, T. striatus).
Restoring the natural retention properties of ecosys-
tems through, among others, creating retention
reservoirs can increase the diversity of environmen-
tal fauna. However, only based on further observa-
tions can it be assessed whether these changes are
short-term and reversible or not.
Funding
The results presented in this paper were obtained as
a part of a comprehensive study nanced by the
University of Warmia and Mazury in Olsztyn,
1116 E. Ludwiczak et al.
Faculty of Agriculture and Forestry, Department of
Entomology Phytopathology and Molecular
Diagnostics, no [30.610.010-110].
Disclosure statement
No potential conict of interest was reported by the
author(s).
Authors’ contributions
Investigation – E.L., E.T., M.N., A.K., conceptuali-
zation – E.T., M.N., A.K., data curation – E.L., E.T.,
M.N., A.K., methodology – E.L., E.T., M.N., A.K.,
analysis – E.L., M.N., A.K., R.R., writing−original
design – E.L., M.N., A.K., visualization – E.L., M.
N., A.K., writing−review and editing – E.L., M.N.,
A.K.
ORCID
E. Ludwiczak http://orcid.org/0000-0002-6307-1522
E. Topa http://orcid.org/0000-0002-4691-2051
M. Nietupski http://orcid.org/0000-0001-6509-4579
A. Kosewska http://orcid.org/0000-0003-2711-4457
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