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FOLIA OECOLOGICA – vol. 50, no. 1 (2023), doi: 10.2478/foecol-2023-0002
The role of soil and plant cover as drivers of soil macrofauna
of the Dnipro River oodplain ecosystems
1Oles Gonchar Dnipro National University, Gagarin av., 72, Dnipro, 49000, Ukraine
2
Ukraine
Abstract
2023. The role of soil and plant cover as drivers of
Folia Oecologica, 50 (1): 16–43.
Floodplain ecosystems are hotspots of biological diversity and perform important ecosystem functions
-
-
pal components of variation in soil physical properties and phytoindication assessments of environmental
Keywords
biodiversity, landscape management, nature conservation, phytoindication, temporal dynamics, zoolog-
ical diagnostics
Olga Kunakh1, Yulia Zhukova1, Volodymyr Yakovenko1, Olexander Zhukov2*
Introduction
Floodplain ecosystems play an important role in the func-
tioning of landscapes (, 2003). Ecosystems located
in river valleys perform important ecological functions.
-
species of plants and animals ( et al., 2005). A
-
tion, and animal communities is essential to understand-
-
ly related to the surrounding geospatial variations and
17
the nature of the rivers ( et al., 2021).
The high level of vertical and horizontal heterogeneity of
alluvial sediments, sedimentation regime, age of forma-
et al.,
2017). The physical properties of alluvial soils are subject
to considerable spatial variability. Geostatistical analysis
-
tation of the spatial variability of alluvial soils is an im-
portant prerequisite for the application of precision agri-
et al., 2005). The
the duration of the most important soil-forming processes
(et al., 2019; and , 1982).
-
ductive in nature ( and , 2007). In the steppe
et al., 2020), but they
provide a place of concentration of regional biodiversity
and diversity of soil cover ( et al., 2020). The
et al., 2018). These systems
et
al., 2020; et al., 2021), salt ( et al., 2015;
et al., 2020) and air regimes ( et
al., 2016) and the impact of erosion processes of varying
intensity ( et al., 2009;
-
and
2009). The anthropogenic impact on any component of
-
plain ecosystems (
ecosystems are sensitive to changes and processes that are
in other parts of the landscape and in a broad sense are a
et al., 2010).
-
action of soil, vegetation and soil biota ( et al.,
-
naturally or as a result of catastrophic events (et
possible based on an understanding of the interactions
soil cover ( et al., 2016; et al., 2022). The
important role in procedures of development of optimum
-
ecosystem ( et al., 2015; et
al., 2022).
Floodplain ecosystems have an important eco-
nomic value (
-
nities for human economic activity are severely limited.
Such locations become natural reserves of biological diversi-
ty (
-
plemented and biodiversity must be incorporated into man-
et al.,
2016). The application of these results to landscape man-
macrofauna. Typically, the soil macrofauna is not a popular
target for biodiversity conservation. When selecting species
for protection, an emotional component plays an important
Ukraine from the soil macrofauna only one species of ear th-
, 2009), one species of Chilo-
poda (, 2009) there are no species of spiders,
-
(, 2010; et al., 2020). The maintenance of a high
level of abundance and species diversity of soil animal com-
them ( and , 2021; et al., 2022).
-
ical properties and provides the formation of soil structure,
( et al., 2016; , 2000; and ,
1999). The number of species, population abundance and
biomass of soil macrofauna are declining in a range of eco-
systems from nature reserves to managed agro-ecosystems.
The main reason for this decline is the reduction of avail-
able organic matter and essential elements in the soil of the
agroecosystem (and , 1997).
soil macrofauna community diversity and the factors that
-
standing the patterns of functioning and sustainability of
, 2007; et al., 2010).
The diversity of animal communities depends on the his-
tory ( and , 2009; et al., 1996),
et al., 2015),
and factors of a neutral nature ( and , 2007;
et al., 2009; et al., 2021). The environ-
features of soil properties and soil regimes ( et al.,
2021; et al., 2019). The variability of soil chemical
properties determines habitat properties and the availability
of nutrients to animals and chemical elements that form the
protective structures of pedobionts ( et al., 2015;-
et al., 2022; et al., 2018; et al., 2019). An
-
cal properties of the soil (
ability and energy requirements for animals to move through
the soil. The physical properties of the soil also regulate the
aeration regime (
( et al., 2021), and the salinity regime ( et al., 2019;
( et al., 2019; et al., 2019).
The role of history in the formation of biodi-
versity patterns of soil macrofauna communities is ver y
18
-
ral patterns have a hierarchical multiscale structure
( et al., 2022; et al., 2021;
-
time is superimposed on the lack of series of data on the
-
the controlling factor. This approach proceeds from the
the community may have formed in a temporal pattern
that corresponds to the rhythm of the controlling factor.
If the temporal patterns of the factors do not correspond
(, 1996).
The morphological feat ures of soils are very
run into hundreds and thousands of years ( et
al., 2014). The physical properties of soils may be close-
ly related to morphological features and have the same
temporal rhythm of variability (, 2011). Or they may
change very rapidly and be characterized by rhythmic
et al., 2015; et al., 2019; et al., 2018). The
rhythm of ecological processes, commensurate in time
to the phytoindicator assessments of environmental fac-
tors (
on the soil macrofauna community can model the role of
ecological drivers in the hierarchy of temporal patterns.
role of soil morphological and physical properties and
phytoindication assessments of environmental factors as
drivers of biological diversity of soil macrofauna of pro-
Materials and methods
Study sites
-
1990 ( et al., 2022). The area of the reserve is
changes in the relief on the territory of the reserve oc-
19
plant in the 1960s and the construction of Dniprodzer-
51.4 m above the sea level. Thus, after the construction of
the cascade of Dnipro reservoirs, the total rise in the level
-
-
also decreased.
Sampling design
(Fig. 1). The numbering of sampling plots is given in ac-
et al. (2019). The sam-
pling plots each consisted of 7 transects. Each transect
-
in such samples visible to the naked eye (macroscopic or-
ganisms) ( and , 2002; et al., 2003;
, 2016). Geobionts (large soil invertebrates that
permanently inhabit the soil) and geophiles (organisms
that live in the soil only for particular phases of their
li ve s) ( , 1992;
assessed. Samples consisted of single blocks of soil, 25 ×
the soil surface prior to taking the soil samples. The litter
stored in 4% formaldehyde ( et al., 2004).
its largest side along the largest local elevation gradient
The habitat type according to EUNIS (European Nature
Information System): G1.223 Southeast European Fraxi-
nus – Quercus – Alnus forests-
resource): Fluvic Calcic Mollic Gleysol (Loamic, Humic).
The habitat type according to EUNIS: G1.1112 Eastern
(Loamic, Protocalcic, Humic, Nechic).
(48°30’06’’N, 34°47’18’’E). The habitat type according
to EUNIS: G1.225 Sarmatic riverine [Quercus] forests.
to EUNIS: G1.225 Sarmatic riverine [Quercus] forests.
-
20
EUNIS: G1.225 Sarmatic riverine [Quercus] forests. The
-
Mollic Gleysol (Loamic, Humic).
Measurement of environmental indicators
-
-
average error of the measurement results of the device
a cross-sectional size of 2 cm2. Within each sampling
single repetition.
An HI 76305 sensor (Hanna Instruments,
-
total electrical conductivity of the soil, i.e. the combined
-
surement results of the device are presented in units of
soil salt concentration, i.e., g l–1. The comparison of HI
estimate the unit conversion factor as 1 dS m–1 = 155 mg
l–1 (and , 2002).
the method of dr y sieving according to Savinov (,
-
described by (2005) in his mono-
The phytoindication assessment of environ-
-
a size of 3 by 3 meters. The projective cover of plants
100 % (, 1973). Edaphic and climatic factors
can be assessed using phytoindication scales (,
2011). The edaphic phytoindication scales include the
total salt regime (Sl), the carbonate content in the soil
(Ca) and nitrogen content in the soil (Nt). The climat-
ic scales include the parameters of the thermal climate
(thermal regime, Tm), humidity (Om), crioclimate (Cr)
(Lc) is an indicator of the microclimate. The phytoindica-
by the ideal indicator method (, 2017). Phytoindi-
information about the projective cover of the herbaceous
layer.
Statistical methods
evaluated using multiple correspondence analysis. The
component analysis. The mentioned statistical analyses
Statistics. Data Analysis
Software System, 2014). The evaluation of alpha, beta,
and gamma diversity of soil macrofauna communities
entropart (
and , 2015) for a language and environment for
statistical computing R (, 2020). A canoni-
macrofauna community using the package ade4 (
and, 2007). The partitioning of the community
package vegan ( et al., 2018). Indicator value
the help of the package indicspecies (, 2013).
Results
Soil morphology
The studies of morphological properties of soils in the
-
-
ols (Table 1). By granulometric composition, the soils
-
-
in a layer 0–10 cm deep from the mineral soil surface.
The calcic horizon is characterized by the presence of
secondar y calcium carbonate detected by the hydrochlo-
ric acid test. The Protosalic properties are related to soil
solution carbonates precipitated in the soil. Diagnosis
of Protocalcic proper ties is based on their persistence
and a fairly marked amount in the soil. Nechic denotes
-
Thaptohumic denote the presence of buried horizons of
appropriate color.
The multiple correspondence analysis indi-
cated the relationship of soil properties in the studied
soil types (Fig. 3). The soils marginal in their prop-
21
Loamic Arenic Humic Ochric Protocalcic Nechic Thaptoochric Thaptohumic
16 16 Fluvic Calcic
Mollic Gleysol
Gleyic
Fluvisol
Gleyic
Fluvisol
Gleyic
Fluvisol
27
Gleyic
Fluvisol
Gleyic
Fluvisol
29
30 Fluvic Mollic
Gleysol
25, 26, 27, 29; soil types – Fluvisol and Gleysol, Humus layer/Granulometric composition – Loamic/Humic, Arenic/Ochric;
Mineralogy – Protocalcic and Other.
22
-
pling sites 25 and 27. The soils of sampling sites 16 and
-
maining soils had the property Loamic. The diagram
-
Soil properties
-
-
scribed 79.9% of the variation in traits. The principal
-
ed a unidirectional change in the soil penetration resis-
-
-
46.3% 14.9% 11.3% 7.4%
Soil penetration resistance (on depth, cm)
0–5 0.54 0.48 –0.31 –0.54
5–10 0.53 0.43 –0.41 –0.49
10–15 0.60 0.34 –0.51 –0.22
15–20 0.56 – –0.66 –
20–25 0.67 – –0.58 0.30
25–30 0.71 – –0.55 0.27
30–35 0.80 – –0.45 0.20
35–40 0.85 – –0.35 –
40–45 0.88 –0.11 –0.26 –
45–50 0.93 –0.11 –0.10 –
50–55 0.94 –0.10 – –
55–60 0.93 –0.15 – –
60–65 0.92 –0.17 0.22 –0.10
65–70 0.90 –0.17 0.28 –0.12
70–75 0.89 –0.17 0.30 –0.11
75–80 0.88 –0.19 0.33 –0.12
80–85 0.87 –0.17 0.37 –0.15
85–90 0.86 –0.18 0.39 –0.16
90–95 0.85 –0.17 0.39 –0.15
95–100 0.85 –0.17 0.39 –0.13
Other soil properties
Electrical conductivity, dSm/m – 0.45 0.39 –
Litter depth (cm) –0.32 – – –
Soil bulk density (g cm–3) –0.27 –0.29 0.14 –0.56
Aggregate fraction (mm)
7–10 – 0.90 0.19 –
5–7 – 0.74 0.18 0.23
3–5 0.66 – 0.13 0.42
2–3 0.70 –0.37 0.15 0.44
1–2 –0.16 –0.84 –0.28 –
0.5–1 –0.55 –0.66 –0.25 –
0.25–0.5 –0.58 –0.53 –0.28 –0.38
<0.25 –0.41 – – –0.63
23
-
cated an increase in the proportion of aggregate fractions
of 2–5 mm and a decrease in the proportion of aggre-
gate fractions of <0.25–2 mm. The principal component
2 described 14.9% of the trait variability. The positive
scores of this principal component indicated an increase
in the soil penetration resistance at 0–15 cm depth and
at 40 cm depth and deeper. The principal component 2
moisture, and soil bulk density. The positive PC2 scores
indicated an increase in the proportion of aggregate frac-
tions larger than 5 mm and a decrease in the proportion of
aggregate fractions of 0.25–3 mm. The principal compo-
nent 3 described 11.3% of the trait variation and indicated
the opposite dynamics of the soil penetration resistance
variability at 0–50 cm depth (the negative principal com-
ponent scores) and 60 cm and deeper (the positive prin-
sensitive to the variability in electrical conductivity, soil
moisture, and soil bulk density. The positive PC3 scores
indicated an increase in the proportion of aggregate frac-
tions larger than 2 mm and a decrease in the proportion of
-
pal component 4 described 7.4% of the trait variation and
indicated the opposite dynamics of the soil penetration
resistance variability at depths of 0–15 cm and 60–100
cm (the negative principal component scores) and 20–35
cm (the positive principal component scores). The prin-
moisture and soil bulk density. The positive PC4 scores
indicated an increase in the proportion of aggregate frac-
tions of size 2–7 mm and a decrease in the proportion of
aggregate fractions of size <0.25–0.5 mm.
The vegetation cover and phytoindication of ecological
regimes
-
of information about species composition of communities
-
-
favorable for acarbonatophiles, and the nitrogen content
for sub-heliophytes (plants of light forests and shrubber-
the shade). The climatic scales assessed conditions for the
scales that are sensitive to soil properties.
Soil macrofauna
The abundance of the soil macrofauna community ranged
from 135.9 ± 21.1 to 332.0 ± 34.9 ind. m–2
-
-
inosa trapezoides (Duges, 1828) and Aporrectodea rosea
-
-
Alpha diversity of the soil macrofauna commu-
in 95% of cases (Fig. 4). Gamma community diversity
ranged from 8.1 to 8.9.
Soil macrofauna community ordination
-
cies. According to the results of the preliminary detrend-
most adequate alternative as an ordination procedure. The
-
29.6% of the community variation (F = 11.2, p < 0.001).
-
nity variation (F = 11.3, p-
1.9% of the community variation (F = 5.3, p < 0.001).
The soil physical properties described 23.7% of commu-
nity variation (F = 41.8, p-
-
9.4% of community variation (F = 17.4, p < 0.001). The
-
F = 5.9, p < 0.001),
F = 1.8,
24
Ecological factor* 16 25 26 27 29
Hd 8.80 ± 0.06 7.29 ± 0.08 8.89 ± 0.06 8.50 ± 0.06 9.12 ± 0.08
fH 4.42 ± 0.04 4.11 ± 0.06 4.67 ± 0.06 4.71 ± 0.05 4.76 ± 0.06
Sl 6.13 ± 0.05 6.35 ± 0.04 6.36 ± 0.06 5.96 ± 0.05 6.55 ± 0.04
Ca 8.15 ± 0.06 8.28 ± 0.04 8.42 ± 0.06 7.86 ± 0.05 8.04 ± 0.04
Nt 8.38 ± 0.04 7.73 ± 0.04 8.65 ± 0.05 8.39 ± 0.05 8.37 ± 0.04
Ae 5.80 ± 0.02 5.90 ± 0.01 5.98 ± 0.02 6.05 ± 0.02 5.85 ± 0.02
Tm 9.76 ± 0.04 9.10 ± 0.06 9.66 ± 0.06 9.93 ± 0.04 9.80 ± 0.04
Om 12.21 ± 0.03 12.34 ± 0.03 12.15 ± 0.04 11.64 ± 0.03 12.01 ± 0.03
Cr 8.61 ± 0.06 8.58 ± 0.05 8.45 ± 0.07 8.65 ± 0.07 8.94 ± 0.06
Lc 7.51 ± 0.03 8.05 ± 0.03 7.48 ± 0.02 7.64 ± 0.03 7.49 ± 0.03
Table 3. Phytoindicator assessment of ecological factors for individual sampling sites: 16, 25, 26, 27, 29 (mean ± standard error)
Cr, cryoregime (average temperature of the coldest month); Lc, light regime.
Fig. 4. Alpha, beta, and gamma diversity of soil macrofauna communities.
25
Species CCA1,
Radj
2=10.2% CCA2,
Radj
2 = 7.1% CCA3, Radj
2 = 6.7% CCA4, Radj
2 = 2.1%
F = 73.2, p < 0.001 F = 50.8, p < 0.001 F = 47.8, p < 0.001 F = 15.4, p < 0.001
Agriotes lineatus 0.06 –1.04 0.79 –0.04
Agrotis segetum –0.77 0.12 0.08 –0.09
Agrypnus murinus 0.76 0.55 0.02 –0.55
Amara familiaris 0.49 –1.24 0.72 –0.08
Amara similata 0.44 1.25 0.85 0.14
Ampedus balteatus –0.31 –0.77 –2.67 0.27
Amphimallon solstitiale 0.36 1.18 0.55 –0.12
Aporrectodea rosea –1.23 0.28 0.13 0.18
Aporrectodea trapezoides 0.47 0.41 0.12 0.02
Asilidae sp.1 –0.81 –0.64 –0.21 –0.43
Athous haemorrhoidalis 0.39 –0.24 –0.20 –0.08
Cardiophorus rupes 0.36 –1.01 0.28 –0.09
Cepaea vindobonensis 0.32 –1.36 0.75 –0.14
Chrysolina fastuosa 0.23 –0.68 0.48 0.66
Cochlicopa lubrica –0.11 –0.03 –0.04 0.78
Dendrobaena octaedra 0.60 0.64 0.07 –0.53
Dendroxena quadrimaculata 0.80 1.06 –0.04 –0.97
Enchytraeus sp. 1 –1.42 0.27 –0.25 –0.22
Forcula auricularia 0.40 –0.95 0.65 –0.05
Geophilus proximus 0.22 0.28 0.33 0.57
Isomira murina 0.21 –0.80 –0.98 0.03
Lithobius aeruginosus 0.27 0.61 –0.31 –0.67
Lithobius curtipes 0.07 0.56 0.14 –0.54
Lumbricidae sp. 0.44 –1.18 0.65 –0.28
Megaphyllum rossicum –0.62 –0.56 0.52 –0.27
Megaphyllum sjaelandicum –1.02 –0.03 0.33 0.06
Melolontha melolontha 0.42 –0.23 0.05 0.16
Octodrilus transpadanus –0.22 –0.01 –1.65 –0.15
Othius angustus 1.10 0.95 0.00 –0.77
Otiorhynchus ligustici 0.13 –0.81 0.40 –0.19
Pachymerium ferrugineum –0.21 –0.94 0.51 –0.27
Pardosa lugubris –1.42 0.14 0.06 –0.55
Platydracus fulvipes –0.21 –0.94 0.62 –0.11
Polydesmus inconstans –1.33 0.31 0.35 0.30
Polyphylla fullo 0.48 –0.63 0.30 –0.24
Prosternon tessellatum 0.18 0.09 0.42 1.16
Pterostichus ovoideus 0.67 1.06 –0.16 –0.93
Rhagio scolopaceus –1.19 –0.65 0.46 –0.45
Rhipidia uniseriata –0.30 –0.30 –2.46 0.08
Serica brunnea 0.07 –0.80 –1.23 0.08
Tabanus bromius –0.10 –0.03 –0.87 –0.29
Thereva nobilitata –0.97 –0.35 –0.32 0.00
Tipula lunata 0.38 0.41 –0.57 0.51
Trachelipus rathkii 0.38 0.04 0.22 –0.04
Xerolycosa miniata 0.43 –0.78 0.50 0.37
26
p
morphological and physical property variability.
-
nities of sampling site (sampling plot) 26 from all others
-
of sampling site 26. High aeration, high carbonate and
these species preferred the living conditions observed
in sampling site 26. Such species include Pardosa lugu-
bris, Polydesmus inconstans, Enchytraeus sp., Rhagio
scolopaceus (larvae), Aporrectodea rosea, Megaphyllum
sjaelandicum and Thereva nobilitata (la r vae).
-
rofauna communities of sampling site 25 (positive scores)
and sampling site 29 (negative scores). The positive scores
-
moclimate values and soil nitrogen content. The negative
values and a higher level of soil solution salinity. Such
species as Pterostichus ovoideus, Dendroxena quadrimac-
ulata, Othius angustus, Dendrobaena octaedra, Lithobius
aeruginosus, and L. curtipes occurred more frequently
species such as Cardiophorus rupes (larvae), Amphimal-
lon solstitiale (larvae), Agriotes lineatus (larvae), Amara
similata (larvae), A. familiaris (lar vae) and Cepaea vindo-
bonensis.
and Gleysols. Fluvisols had the Arenic property more fre-
quently, and Gleysols had the Loamic property more fre-
quently. Fluvisols had a more alkaline soil solution reac-
tion, and Gleysols had a more acidic soil solution reaction.
The community of sampling site 25 can be dif-
3. The negative values of principal components 1 and 2,
-
-
ized by the presence of Protocalcic and Loamic properties.
Ampe-
dus balteatus (larvae), Rhipidia uniseriate (larvae), Serica
brunnea (larvae), Octodrilus transpadanus, Isomira muri-
na (larvae) and Tabanus bromius (l arva e).
-
fauna community of sampling site 27 from all others. This
salinity of soil solution and carbonate content in the soil.
Species such as Prosternon tessellatum (larvae), Cochlico-
pa lubrica, Geophilus proximus, Chrysolina fastuosa (lar-
vae), Tipula lunata (larvae), and Polydesmus inconstans
Indicator value analysis of soil macrofauna species
-
ical soil properties and soil macrofauna species (Table 5).
Only one species (Cochlicopa lubrica)
sampling site type. Species such as Agrypnus murinus
(larvae), Dendrobaena octaedra, Dendroxena quadrimac-
ulat a, Lithob ius aeru gin osu s, Othius angu stu s, an d Pteros-
tichus ovoideus
-
cies as Ampedus balteatus (larvae), Octodrilus transpada-
nus, Rhipidia uniseriate (larvae). The unique indicators of
Enchytraeus sp., Pardosa lugubris. No unique indicators
of this sampling site are also indicators of sampling sites
-
pling sites. The unique indicator species of sampling site 26
Agriotes lineatus (larvae), Amara familiaris (larva e),
Amara similata (larvae), Amphimallon solstitiale ( la r vae) ,
Cardiophorus rupes (larvae), Cepaea vindobonensis,
Forcula auricularia, Otiorhynchus ligustici (larvae) and
Polyphylla fullo (larva e).
-
fauna communities of sampling site 25 (positive scores) and
sampling site 29 (negative scores). The positive scores of the
-
mate values and soil nitrogen content. The negative scores of
higher level of soil solution salinity. Such species as Pteros-
tichus ovoideus, Dendroxena quadrimaculata, Othius an-
gustus, Dendrobaena octaedra, Lithobius aeruginosus, and
L. curtipes
Car-
diophorus rupes (larvae), Amphimallon solstitiale ( la r vae),
Agriotes lineatus (lar vae), Amara similata (larvae), A. famil-
iaris (larvae) and Cepaea vindobonensis.
and Gleysols. Fluvisols had the Arenic property more fre-
quently, and Gleysols had the Loamic property more fre-
quently. Fluvisols had a more alkaline soil solution reac-
tion, and Gleysols had a more acidic soil solution reaction.
The community of sampling site 25 can be dif-
3. The negative values of principal components 1 and 2,
-
-
ized by the presence of Protocalcic and Loamic properties.
Ampe-
dus balteatus (larvae), Rhipidia uniseriate (larvae), Serica
brunnea (larvae), Octodrilus transpadanus, Isomira muri-
na (larvae) and Tabanus bromius (l arva e).
-
fauna community of sampling site 27 from all others. This
salinity of soil solution and carbonate content in the soil.
Species such as Prosternon tessellatum (larvae), Cochlico-
pa lubrica, Geophilus proximus, Chrysolina fastuosa (lar-
vae), Tipula lunata (larvae), and Polydesmus inconstans
27
-
regime; Ca – carbonate content in soil; Nt – nitrogen content in soil; Ae – soil aeration; Tm – ther mal climate; Om – climate
-
28
Indicator value analysis of soil macrofauna species
categorical soil properties and soil macrofauna species
(Table 5). Only one species (Cochlicopa lubrica-
Agr ypnus
murinus (larvae), Dendrobaena octaedra, Dendroxena
quadrimaculata, Lithobius aeruginosus, Othius angus-
tus, and Pterostichus ovoideus
sampling site 16. The unique indicators of sampling site
Ampedus balteatus (larvae), Oc-
todrilus transpadanus, Rhipidia uniseriate (larvae). The
Asilidae sp. (larvae), Enchytraeus sp., Pardosa lugubris.
The indicator species of this sampling site are also indi-
cators of sampling sites 26 or 29, less frequently in com-
Agriotes lineatus (l arva e),
Amara familiaris (larvae), Amara similata (larvae), Amphi-
mallon solstitiale (larvae), Cardiophorus rupes ( la r vae) ,
Cepaea vindobonensis, Forcula auricularia, Otiorhyn-
chus ligustici (larvae) and Polyphylla fullo (l arva e).
the total number of species in the analysis). The Fluvisol
-
(48.9%) and the Loamic/Humic properties indicators
-
Discussion
-
soils. These are morphological characteristics of soils,
physical properties of soils, and phytoindication assess-
ments of environmental factors. Each of these groups of
chosen set of predictors of the soil macrofauna commu-
dynamics of ecological conditions. The intrinsic time of
variability of the physical properties of the soil has a dura-
tion of days, months, years. The intrinsic time of variabil-
ity of phytoindication assessments of environmental fac-
tors has duration of some years or decades. The intrinsic
time of variability of morphological properties of soil has
duration of some years,dozens of years, centuries.
The soil types and soil morphological properties
can also be indicated by soil macrofauna species. The ba-
so their role as indicators is of particular interest. Our re-
A. rosea, and the indicators of Gleysols are
A. trapezoides and the epigean
D. octaedra
ecologically diverse. Species that have a similar ecologi-
cal optimum are combined into ecological groups (-
-
ferent ecological niches, so the ecological niche is charac-
terized not only by an optimum, but also by an ecological
best indicators. Fluvisol indicators include both epigeic
species (Agrotis segetum (larvae), Pardosa lugubris,
Polydesmus inconstans, Rhipidia uniseriate (larvae),
Thereva nobilitata (larvae), Megaphyllum sjaelandicum)
and endogeic species (Ampedus balteatus (larvae), Pros-
ternon tessellatum (larvae), Rhagio scolopaceus (larvae),
Serica brunnea (larvae), Enchytraeus sp.). The same is
species (Amara familiaris, Dendroxena quadrimaculata,
L. aeruginosus, Lithobius curtipes, Othius angustus (lar-
vae), Pterostichus ovoideus, Tipula lunata (larvae), Tra -
chelipus rathkii) and endogeic species (larval stages of
Agrypnus murinus, Athous haemorrhoidalis, Melolontha
melolontha, Polyphylla fullo).
The ecological groups of soil animal species
consequence of the activity of elementary soil processes
species that are indicators of the granulometric composi-
-
the ground beetle Amara familiaris prefers sandy soils
(, 1977). The beetle Amara similata prefers damp
, 2014).
This species is part of the group that indicates pioneer
sandbars ( et al., 2005). A. lineatus
(-
ic matter ( et al., 2018). This species prefers
et al., 2012). In our
density than loam soils. The larvae of Amphimallon sol-
stitiale 1952).
Also inhabitants of sandy soils are the larvae of Cardio-
phorus rupes (, 1978). The mollusk C. vindobon-
ensis
( et al., 2004), but in river valleys it is usually
-
posits ( , 2009).
The larvae of Melolontha melolontha prefer
loamy soils (Octodri-
lus transpadanus
of soil galleries ( and , 2010; and
29
Species Sampling site Soil Granulometry Mineralogy
16 25 26 27 29 Fluvisol Gleysol Arenic Loamic Non Protoc. Protocalcic
Agriotes lineatus 0 0 0 0 1 – – 1 0 1 0
Agrotis segetum 1 0 1 1 0 1 0 1 0 1 0
Agrypnus murinus 1 0 0 0 0 0 1 – – – –
Amara familiaris 0 0 0 0 1 0 1 1 0 1 0
Amara similata 0 0 0 0 1 – – 1 0 1 0
Ampedus balteatus 0 1 0 0 0 1 0 – – 0 1
Amphimallon solstitiale 0 0 0 0 1 – – 1 0 1 0
Aporrectodea rosea 0 0 1 1 0 1 0 – – – –
Aporrectodea trapezoides 1 0 0 1 1 0 1 – – – –
Asilidae sp. 0 0 1 0 0 – – 1 0 – –
Athous haemorrhoidalis 1 1 0 0 1 0 1 0 1 – –
Cardiophorus rupes 0 0 0 0 1 – – 1 0 1 0
Cepaea vindobonensis 0 0 0 0 1 – – 1 0 1 0
Chrysolina fastuosa 0 0 0 1 1 – – – – – –
Cochlicopa lubrica – – – – – – – – – – –
Dendrobaena octaedra 1 0 0 0 0 0 1 0 1 1 0
Dendroxena quadrimaculata 1 0 0 0 0 0 1 0 1 1 0
Enchytraeus sp. 0 0 1 0 0 1 0 1 0 – –
Forcula auricularia 0 0 0 0 1 – – 1 0 1 0
Geophilus proximus 1 0 0 1 1 – – – – 1 0
Isomira murina 0 1 0 0 1 – – – – 0 1
Lithobius aeruginosus 1 0 0 0 0 0 1 0 1 1 0
Lithobius curtipes 1 0 1 0 0 0 1 – – 1 0
Lumbricidae sp. 0 0 0 0 1 0 1 1 0 1 0
Megaphyllum rossicum 0 0 1 0 1 – – 1 0 1 0
Megaphyllum sjaelandicum 0 0 1 1 0 1 0 1 0 1 0
Melolontha melolontha 1 0 0 1 1 0 1 0 1 1 0
Octodrilus transpadanus 0 1 0 0 0 – – 0 1 0 1
Othius angustus 1 0 0 0 0 0 1 0 1 1 0
Otiorhynchus ligustici 0 0 0 0 1 – – 1 0 1 0
Pachymerium ferrugineum 0 0 1 0 1 – – 1 0 1 0
Pardosa lugubris 0 0 1 0 0 1 0 1 0 1 0
Platydracus fulvipes 0 0 1 0 1 – – 1 0 1 0
Polydesmus inconstans 0 0 1 1 0 1 0 1 0 – –
Polyphylla fullo 0 0 0 0 1 0 1 1 0 1 0
Prosternon tessellatum 0 0 0 1 1 1 0 – – – –
Pterostichus ovoideus 1 0 0 0 0 0 1 0 1 1 0
Rhagio scolopaceus 0 0 1 0 1 1 0 1 0 1 0
Rhipidia uniseriata 0 1 0 0 0 1 0 0 1 0 1
Serica brunnea 0 1 0 0 1 1 0 – – 0 1
Tabanus bromius 1 1 1 0 1 – – – – 0 1
Thereva nobilitata 0 1 1 0 0 1 0 1 0 – –
Tipula lunata 1 1 0 1 0 0 1 0 1 0 1
Trachelipus rathkii 1 0 0 1 1 0 1 – – 1 0
Xerolycosa miniata 0 0 0 1 1 – – – – 1 0
Protoc., Protocalcic.
p < 0.05)
30
anecic species (, 1977; et al., 1976).
stream dynamics, resulting in erosion and sedimentation
processes. Sandy soils usually have a thin humus horizon,
-
zon ( et al., 2007; et al., 2011).
The list of species that are indicators of sandy soils
carbonate horizon. This is consistent, since the carbon-
number of the indicator species of carbonate horizon is very
Ampedus balteatus
or forest litter (, 1978), therefore, the indicator role of
the carbonate horizon may be as a consequence of the pecu-
cover. These larvae prefer sandy soils ( and -
-
systems. Our results indicate that this species is an indicator
Octodrilus transpadanus is also an in-
dicator of Protocalcic horizon. Increased calcium content in
the soil is a factor in soil structure ( and
, 2007). In turn, soil structure is necessary to main-
et al., 2017),
and calcium is necessary for the normal digestive process of
et al., 2015; et al., 2003).
the duration of the process of their genesis is measured in
centuries ( et al., 2014; et al., 2020). The
every spring ( 1950). Due to construction of a
-
gime of the river has changed dramatically ( et
-
-
et al., 2009). The de-
supply of organic matter to the surface horizons and grad-
ually leads to soil depletion ( et al., 2016).
-
-
the result of the interaction of the main soil-forming fac-
tors: relief, soil-forming material, climate, communities of
living organisms and time (c, 1883; ,
2012; , 1941; et al., 2006).
The physical properties of soil depend on the fac-
tors of soil formation (-
variability of soil physical properties is represented by tem-
content in alluvial soils at the ecosystem scale is deter-
by other soil factors such as soil str ucture, microrelief,
rock content ( et al., 2016).
Plant communities are subject to successional
dynamics. The duration of successional changes of plant
community is centuries (, 1936; , 1974).
-
-
et al., 2011; et al., 2002). Nevertheless, the
nature of interrelation of plants and ecological processes
(et al., 2021; et al., 2019). Ob-
viously, such an approach is sensitive to the ecological
-
namics of the plant community. The species composition
of plant phytocenoses and changes in the quantitative
studying and monitoring environmental changes. Simi-
methods of estimating nitrogen content in soil. For oth-
er ecological properties, direct and phytoindication esti-
researchers to prefer the direct measurements of environ-
mental properties ( et al., 2021). Obviously,
-
ent aspects of the dynamics of environmental properties.
at the moment of measuring the property. The phytoin-
dication assessment characterizes the regime of environ-
dynamics of the plant community.
About 1/3 of the variability of the soil macrofauna
-
ly random, due to the action of unmeasured factors in the
study, the action of measured factors at the ecosystem level,
or be the result of causes of a neutral nature. Among the
sets of measured environmental factors considered, the soil
physical properties are the most important driver of changes
in the soil macrofauna community. The soil morphological
some proportion of the community variation, but the larg-
31
physical properties. The result obtained can be interpreted
-
the dimensionality of the feature space and solve the prob-
the intra-ecosystem variability of soil physical properties is
inter-ecosystem level. Thus, it can be assumed that some
-
el. Moran’s Eigenvector Maps and related methods for the
spatial multiscale analysis of ecological data can be used
factors (et al., 2008; et al., 2006).
physical properties of the soil and environmental factors
-
soil macrofauna community is the aeration regime and the
presence of calcium in the soil. A soil str ucture, represent-
ed by aggregates and intra-aggregate and inter-aggregate
pore space (, 2017; and ,
2021), is considered the main indicator of soil physical con-
-
tions (, 2004; et al., 1994; , 2009;
et al., 2022). In turn, the soil aggregate structure
is able to provide an optimal respiratory regime for soil bi-
ota (
soil penetration resistance characterize the availability of
soil space for plant roots and soil animals to penetrate. For
many plants, a soil penetration resistance value of 3 MPa
is the limiting value that roots can overcome. If the value
are 0.13 MPa and 0.195 MPa, respectively ( and ,
2018). The construction of soil galleries by animals is a very
energy-consuming process. The opposite feature should
also be noted: the constructed galleries should have some
-
-
-
by both density and soil penetration resistance. Therefore,
these indicators are information-valuable predictors of soil
macrofauna community structure.
The results indicate that moisture and aeration of
and the higher the soil moisture, the less air available for
breathing conditions for soil animals. Also importantly, the
moisture regime marks the most impor tant gradient of soil
important gradient (CCA2) marks the trophicity factor. The
CCA3 is also noted to be marked by a phytoindicative in-
ideas of O. L. Belgard (, 1971, 1950) on the typol-
intensity are considered as the main structuring gradients.
Our results indicate that these structuring factors also act at
the level of soil macrofauna.
Conclusion
-
-
The indicators of soil types and morphological properties
A. rosea is indicator of
A. trapezoides and
the epigean D. octaedra are indicators of Gleaysols. Mor-
phological characteristics of soils, physical properties of
soils, and phytoindication estimates of environmental fac-
physical properties of soils are the most important driver of
variation in the soil macrofauna community. The morpho-
logical features of soils and phytoindication scales are also
-
ical properties.
Acknowledgements
-
-
-
ers for helping us to improve earlier versions of this paper.
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Received July 8, 2022
Accepted October 29, 2022
Properties 16 25 26 27 29
Soil penetration resistance, MPa (on depth, cm)
0–5 0.76 ± 0.02 0.65 ± 0.01 0.97 ± 0.03 1.83 ± 0.01 1.29 ± 0.02
5–10 0.81 ± 0.02 0.87 ± 0.01 1.17 ± 0.03 2.10 ± 0.05 1.50 ± 0.04
10–15 0.81 ± 0.02 0.95 ± 0.01 1.54 ± 0.05 1.96 ± 0.05 1.46 ± 0.03
15–20 0.86 ± 0.02 1.37 ± 0.02 2.04 ± 0.06 1.86 ± 0.04 1.45 ± 0.02
20–25 0.81 ± 0.03 1.25 ± 0.02 2.54 ± 0.06 1.97 ± 0.04 1.51 ± 0.03
25–30 0.79 ± 0.02 1.40 ± 0.02 3.03 ± 0.07 2.29 ± 0.05 1.85 ± 0.04
30–35 0.75 ± 0.02 1.52 ± 0.03 3.47 ± 0.10 2.55 ± 0.07 2.44 ± 0.05
35–40 0.81 ± 0.02 1.48 ± 0.03 3.74 ± 0.12 2.96 ± 0.09 3.07 ± 0.08
40– 45 1.04 ± 0.03 1.72 ± 0.04 4.33 ± 0.13 3.34 ± 0.10 3.83 ± 0.10
45–50 1.32 ± 0.03 1.78 ± 0.04 4.84 ± 0.13 3.77 ± 0.08 5.02 ± 0.11
50–55 1.79 ± 0.05 1.83 ± 0.03 5.36 ± 0.14 4.17 ± 0.07 6.18 ± 0.13
55–60 2.21 ± 0.05 2.04 ± 0.03 5.91 ± 0.14 4.30 ± 0.07 7.36 ± 0.13
60– 65 2.52 ± 0.05 2.29 ± 0.05 6.24 ± 0.16 4.38 ± 0.07 8.10 ± 0.11
65–70 2.86 ± 0.08 2.46 ± 0.05 6.56 ± 0.14 4.61 ± 0.05 8.70 ± 0.10
70–75 3.03 ± 0.09 2.59 ± 0.04 6.73 ± 0.16 4.76 ± 0.04 9.05 ± 0.09
75–80 3.18 ± 0.07 2.91 ± 0.05 6.71 ± 0.16 4.81 ± 0.04 9.40 ± 0.06
80–85 3.44 ± 0.09 2.88 ± 0.06 6.54 ± 0.16 4.89 ± 0.03 9.64 ± 0.03
85–90 3.50 ± 0.09 2.95 ± 0.07 6.59 ± 0.16 4.93 ± 0.02 9.88 ± 0.02
90–95 3.58 ± 0.08 3.01 ± 0.06 6.53 ± 0.18 4.95 ± 0.02 9.94 ± 0.01
95–100 3.69 ± 0.09 3.08 ± 0.05 6.70 ± 0.19 4.96 ± 0.01 9.96 ± 0.01
Other soil properties
Electrical conductivity (dSm m–2) 0.54 ± 0.06 0.14 ± 0.01 0.13 ± 0.01 0.25 ± 0.02 0.22 ± 0.01
Litter depth (cm) 2.40 ± 0.07 3.14 ± 0.05 2.04 ± 0.14 2.10 ± 0.03 2.43 ± 0.04
Bulk density (g cm–3) 1.07 ± 0.01 1.13 ± 0.01 0.91 ± 0.01 1.02 ± 0.01 1.12 ± 0.00
Aggregate fraction, mm (in %)
7–10 10.39 ± 0.20 4.10 ± 0.14 7.94 ± 0.20 10.75 ± 0.21 7.77 ± 0.11
5–7 11.93 ± 0.22 8.77 ± 0.18 10.34 ± 0.21 12.08 ± 0.17 9.97 ± 0.08
3–5 18.12 ± 0.26 14.17 ± 0.27 23.60 ± 0.40 19.19 ± 0.28 21.60 ± 0.18
2–3 14.36 ± 0.21 15.85 ± 0.20 28.04 ± 0.48 14.61 ± 0.23 23.47 ± 0.26
1–2 8.40 ± 0.23 19.53 ± 0.31 13.55 ± 0.49 8.44 ± 0.22 11.94 ± 0.17