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\begin{document}

\title{Life traits of ground beetle assemblages of spruce forests in
north-eastern Poland}



\author[1]{Mariusz Nietupski}%
\author[1]{Agnieszka Kosewska}%
\author[1]{Emilia Ludwiczak}%
\affil[1]{University of Warmia and Mazury in Olsztyn}%


\vspace{-1em}



  
  \date{May 22, 2024}


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\selectlanguage{english}
\begin{abstract}
Assemblages of epigeic ground beetles living in spruce forests in three
age ranges: 20-30 years (A); 40-50 years (B); 70-80 years (C) were
submitted for research. The investigations were carried out in spruce
forests in north-eastern Poland. In each age forest category, 4 plots
with 5 Barber traps were set up. Ground beetle assemblages were compared
in terms of their abundance, species richness, and the Shannon H' Index
value. Quantitative ecological description of carabids captured in the
analysed age-classes of spruce forests was performed, and the values of
the MIB (mean individual biomass) were calculated. To determine the
dependencies between the values of the MIB and abundance of Carabidae
trophic groups, the Spearman rank correlation coefficient was
calculated. The assemblages of ground beetles living in the spruce
forests in north-eastern Poland were characterised by quite large
species richness. Significant differences were determined in the number
of species richness between the Carabidae assemblages inhabiting the
analysed age ranges of spruce forests. The oldest spruce stands had a
smaller number of species and specimens of ground beetles as well as the
highest MIB values in comparison with the younger spruce forests (30 to
50 years old). It was found that high MIB values were positively
correlated with the presence of large ground beetle species with higher
moisture requirements. Lower values of the MIB index were due to the
presence of smaller, macropterous species, with the spring type of
breeding and associated with open areas.%
\end{abstract}%



\sloppy


\textbf{Introduction}

In the late 18\textsuperscript{th} century, forests in Poland covered
around 40\% of the country's area, and declined to 20.8\% in 1945, soon
after World War Two (Zaj\selectlanguage{polish}ą\selectlanguage{english}czkowski \emph{et al.} 2016). The undesirable
environmental changes caused by this process, as well as the great
demand for timber were the reasons why the forest cover in Poland was
enlarged to 27.0\% between 1945 and 1970. This increase was achieved
mainly by growing monoculture forest stands of Scots pine (\emph{Pinus
sylvestris} L.) and Norway spruce (\emph{Picea abies} (L.) H. Karst)
(Krawczyk 2014). The latter species in its natural range is one of the
main forest-forming species. It grows in the mountains of central and
southern Europe and also covers extensive areas in the north and east of
this continent. Nowadays, spruce in Poland occurs in two ranges:
north-eastern (on lowlands) and south-western (in the mountains), and
makes up around 6\% of all forests. The dying of spruce stands observed
in recent years may be caused by several factors. The following are
mentioned: decreasing soil moisture content, changes in soil chemistry
connected with acidification, and shortage of nutrients, e.g. magnesium
{[}Zwoli\selectlanguage{polish}ń\selectlanguage{english}ski 2003; Ma\selectlanguage{polish}ł\selectlanguage{english}ek 2014{]}. Another reason why spruce forests are
dying is the way they are grown, as a monoculture, where this species
forms dense single-storey stands; spruce a foreign species in Poland.
Monoculture spruce forests are a specific habitat, with poor litter, not
very rich understorey and groundcover, and extensive shading of the soil
surface {[}Holeksa and Szwagrzyk 2004{]}.

Descriptions of the state and condition of a variety of habitats often
include bioindicator methods. One of the groups of organisms used for
bioindication is the Carabidae family. Carabidae make very good
bioindicators, and are often used to evaluate changes occurring in
agricultural, woodland, and urban habitats {[}Niemel\selectlanguage{ngerman}ä et al. 1993, 2007;
Pizzolotto et al. 2018{]}. They are also one of the simple indicators
employed in descriptions of the development of succession in forests
{[}Szyszko 1983; 1991; Butterfield 1997; Sk\selectlanguage{polish}ł\selectlanguage{english}odowski 2009{]}. Assemblages
of ground beetles dwelling in pine forests are quite well recognised
{[}Schwerk and Szyszko 2007; Sk\selectlanguage{polish}ł\selectlanguage{english}odowski et al. 2018{]}. Numerous studies
provide the characteristics of ground beetle assemblages in pine forests
of different ages, growing on different types of soil and using
different forest management practices or having the understorey composed
of different deciduous species {[}Szyszko 1990; Jukes et al. 2001; Finch
2005; Taboada et al. 2008; Barsoum et al. 2014; Lange et al. 2014;
Kosewska et al. 2019{]}. Having explored the complex relationships
between ground beetle assemblages and the pine forests they inhabit, it
became possible to develop a synthetic indicator that describes these
dependences {[}Szyszko 1990{]}. The Mean Individual Biomass (MIB) of
carabid fauna has been proposed as an indicator of the stage of
succession in forests and is now employed in research and forest
management {[}Szyszko et al. 2000; Schwerk and Szyszko 2007; Instrukcja
ochrony lasu (Forest Protection Legislation) 2012; Sk\selectlanguage{polish}ł\selectlanguage{english}odowski et al.
2018{]}. Regarding spruce forests growing in North America, the
Fennoscandian Peninsula, and most of Europe, the research results
concerning this group of beetles have been well documented {[}Reeves et
al. 1983; Abildsnes and Tommer\selectlanguage{ngerman}äs 2000; Elek and Magura 2001; Jukes et
al. 2001; Koivula and Niemelä 2002; Lange et al. 2014{]}. In contrast,
we lack such reports on ground beetles in Polish spruce forests. Should
the species and quantitative composition of Carabidae assemblages in
spruce forests be known, such information might be useful for the
determination of the advancement of forest succession, and the
evaluation of the condition of a given forest habitat, especially in
view of the progressing death of spruce stands and risk of increase
presence of pests.

The study reported in this paper aimed to determine the quantitative and
species composition of epigeic assemblages of Carabidae dwelling in
spruce forests in north-eastern Poland. The hypothesis was that these
assemblages differ from one another depending on the age of the forest,
type of woodland habitat, and its humidity. It was assumed that older
forests would have:

- a greater share of large forest zoophagous species accompanied by a
decreasing percentage of open-space hemizoophagous species, - a higher
value of the MIB, - diminishing ground beetles species richness.

\textbf{Materials and methods}

\emph{Study area}

The research covered the area of 4 Forest Subdistricts in north-eastern
Poland: subdistrict Sosnowo (Forest District Skrwilno) in 2016 (plots:
A1, A2 -- area: 1.0 ha); subdistrict Zielony Dwór (Forest District
Gi\selectlanguage{polish}ż\selectlanguage{english}ycko) in 2015 (plots: B1, B2 -- area: 1.39 ha; A3, A4 -- area: 1.83
ha); subdistrict Maruny (Forest District Wipsowo) in 2018 (plots: B3,
B4: -- area: 3.00 ha) and subdistrict Przejmy (Forest District
Przasnysz) in 2015 (plots: C1, C2, C3, C4 -- area: 5.4 ha) (Figure 1).
Areas with a dominant share of spruce (70 -- 100 \%), representing 3 age
categories, were selected:

\begin{itemize}
\tightlist
\item
  A -- 20 - 30 years old (plots: A1, A2, A3, A4),
\item
  B -- 40 - 50 years old (plots: B1, B2, B3, B4),
\item
  C -- \emph{ca} 80 years old (plots: C1, C2, C3, C4).
\end{itemize}

More information about the research sites is given in table 1. In each
age category, 4 plots treated as replications were established. In each
research plot, 5 Barber traps were set up, which were plastic containers
measuring 9 cm in diameter and 13 cm in height. The traps were filled
with a a preservative liquid and submerged into the soil to be level
with the ground surface (Kotze et al. 2011). Traps in each plot were
spaced along a transect, at a distance of 10 m from each another. The
transects were placed in the central part of the spruce stands; the
distances between the transects and the arrangement of traps are shown
in Figure 1.

Twenty traps were set up in each age variant of spruce forest, that is,
60 traps in the entire experiment (3 variants x 4 replications x 5
traps). A trap served as an elementary sampling unit. Traps were placed
in the ground in spring (May), emptied every two weeks while
replenishing the preservative liquid, and the species of captured
beetles were identified (H\selectlanguage{polish}ů\selectlanguage{english}rka 1996). The traps were removed in autumn
(October), and the duration of their exposure (days) for each of the
research areas was as follows: Przejmy (135), Zielony Dw\selectlanguage{ngerman}ór (160),
Sosnowo (153) and Maruny (141). Moran's I index was calculated (SAM
v.4.0; Rangel et al. 2010) to measure spatial autocorrelation of carabid
abundance in the sampling plots of the studied areas. The distribution
of the abundance of carabid beetles along the spatial arrangement of
sampling plots was not autocorrelated (Moran's I = 0.264, p = 0.117).

\emph{Life traits of carabids}

Assemblages of ground beetles in the analysed age categories of spruce
forests were compared, taking into account the number of beetles,
species richness, and values of the Shannon H' species diversity index.
The species-identified beetles were classified in terms of body size and
food preferences, breeding strategy, preferred habitat, habitat moisture
content, and ability to fly (Meijer 1974, Lindroth 1985, 1986, H\selectlanguage{polish}ů\selectlanguage{english}rka
1996, Aleksandrowicz 2004) (Table 2). Regarding the ecological traits,
the captured Carabidae representatives were divided into large zoophages
(Lz) -- predatory species with a body length of more than 15 mm, small
zoophages (Sz) -- also predators but with a body length no more than 15
mm, and hemizoophages (Hz) -- ingesting mixed food. Concerning the
breeding strategy, the ground beetles were classified as spring (Sb) and
autumnal (Ab) types of breeding. Depending on habitat preferences, the
following groups of ground beetles were distinguished: forest (F), open
area (Oa), eurytopic (Eu), and wetland (Wet) species. According to the
moisture preferences, the distinguished groups of ground beetles were:
hygrophilous (H), mesophilous (M), and xerophilous (Xe). The ability to
fly was also considered, giving rise to the following division: able to
fly, with developed wings, i.e. macropterous species (Ma), species with
reduced wings, not flying, i.e. brachypterous (B), and dipterous (D),
i.e. those which can be found to present either of the forms of wing
development.

Analysis of ground beetle assemblages in terms of their ecological
characteristics can be useful in assessing the advancement of the forest
succession process. Assemblages of carabid beetles associated with early
phases of the forest succession progress are distinguished by a
relatively large share of small species with large dispersion power,
often connected with open space areas. In advanced stages of succession,
large, forest-type species with lesser dispersion power begin to appear
(Magura et al. 2002; Niemel\selectlanguage{ngerman}ä et al. 2007).

\emph{Data analysis}

The mean individual biomass (MIB) of Carabidae was calculated using the
formula (Szyszko et al. 2000) which describes dependencies between the
body length and biomass of beetles. Evaluation of the distribution of
the abundance of a catch rate species and specimens of Carabidae as well
as the abundance of the distinguished trophic groups in the analysed
variants was supported by the Shapiro-Wilk W test. Data characterised by
unimodal distribution were assessed by applying the GLM (Generalised
Linear Model), which included Poisson's data distribution. Groups of
means not statistically different were assigned the same lettered index:
a, b (Bonferroni test). Data having normal distribution were examined by
ANOVA test. Spearman's correlation rank coefficient
(\emph{r\textsubscript{s}} ) was derived to determine the correlation
between values of the MIB index and the abundance of specific trophic
groups of Carabidae.

To visualize the differences of ground beetle assemblages living in the
analysed age variants of spruce forests, non-metric multidimensional
scaling (NMDS) was performed, using the Bray-Curtis similarity index.
This method enabled us to evaluate the ground beetle assemblages based
on the similarity between individual beetles and by analysing data
concerning the number of captured species and specimens. Samples with a
high degree of similarity are plotted close to one another in the NMDS
diagram. An assessment of the significance of differences between the
analysed assemblages using the NMDS method was performed using the
ANOSIM non-parametric statistical test (Clarke 1993). The ordination
methods were used to assess relationships between the presence of
Carabidae species and their specific ecological groups versus the
distinguished environmental variables (forest age, canopy, humidity,
fertility, MIB, and the number of treatments, such as stand cleaning and
thinning) (ter Braak and Smilauer 1998). The RDA method (Redundancy
Analysis) was used because the distribution of the analysed data was
linear (SD = 2.09). This type of analysis arranges samples in order
along a gradient represented by axes of the ordination diagram, in our
case based on data concerning the species composition of ground beetle
assemblages. The Jackknife 2 estimator was used for abundance data
(using EstimateS v. 9.1.0 statistical software) and the species
accumulation curves were calculated to assess the adequacy of the
sampling efficiency (Zahl 1977; Colwell 2005).

All statistical calculations and their graphical presentation were
supported by the following programmes: Statistica 13.3 (Shapiro-Wilk W
test, GLM, Spearman's correlation rank coefficient), Canoco 4.51 (RDA),
and Past 2.01 (NMDS and ANOSIM).

\textbf{Results}

During the study, 3,105 specimens belonging to 44 carabid species were
captured (Table 2). Species accumulation curves for individual transects
in the 3 years of the study confirmed that the sampling effort was
adequate (Figure 2). The dominant species in the analysed forest stands
was \emph{Carabus hortensis} , which composed 34.72 \% of the total
species captured. Other numerous species were: \emph{Pterostichus
niger}(14.98 \%), \emph{Pterostichus oblongopunctatus} (11.56
\%),\emph{Pterostichus melanarius} (9.15 \%), and \emph{Calathus
micropterus} (7.50 \%). The remaining species of ground beetles did not
exceed a 5 \% share in the examined assemblages. Most species (40) were
caught in the youngest spruce forest classified as A (up to 30 years
old), while in the 50-year-old spruce forest (B) there were 28 species,
and the 80-year-old spruce stands (C) were home to the smallest number
of ground beetle species, namely 22. The GLM analysis confirmed
significant differences in the average number of ground beetle species
caught between the three age categories of spruce stands.
The\emph{post-hoc} Bonferroni test arranged these average values into
three uniform groups (Figure 3a). The smallest average catch rate of
carabids was found in 80-year-old spruce stands (3.9 individuals on
average). Significantly more individuals were captured in younger spruce
forests (A, B), where their number was on a comparable level (Table 2,
Figure 3b). The highest MIB value was found in the assemblages
inhabiting the oldest stands (C) Figure 3c, Table 2).

In the three age categories of spruce stands, the dominant share of
ground beetles was composed of forest species, and the highest average
number of individuals representing this class of ground beetles was
noted in the youngest spruce forest (A) - 49.25 individuals on average
(Table 3). An increase in the age of the forest did not generate a
higher mean abundance of forest species of ground beetles (B - 39.85 and
C - 33.85). The second most numerous group consisted of eurytopic
species, most abundant in the 50-year-old forest (14.90) and 80-year-old
forest (7.10). The least numerous group of eurytopic species was
observed in the youngest forest (A). Open-area and wetland beetles
composed the least numerous groups, presenting a similar distribution in
all three age categories of forests. Our analysis of ground beetle
moisture preferences revealed that mesophilous beetles were the dominant
group, and they appeared in the highest number in the oldest stands (C).
The youngest spruce stands (A) comprised the highest percentage of
individuals classified as xerophilous ones as well as the ones
associated with higher moisture content in the soil. Concerning the
trophic preferences, the examined age categories of spruce stands were
populated by the prevalent share of predatory ground beetles. Large
zoophagous beetles were most frequently encountered in the 50-year-old
spruce forest (44.00). In the other two age categories (youngest and
oldest), the number of large zoophages was lower and similar in both
cases. The dominant type of breeding among the ground beetles was the
autumnal strategy, with the most numerous representatives found in the
50-year-old forest (B). As for the spring type of breeding, such species
were observed to gradually increase in number as the age of the examined
forest was lower. The most brachypterous individuals (36.70) appeared in
the 30-year-old spruce forest. The smallest number of beetles with no
ability to fly was found in the 50-year-old spruce forest (B). In this
forest, the dominant beetles were winged ones (macropterous and
dipterous) (Table 3).

To determine the correlations between MIB values in the analysed age
ranges of spruce forests and the abundance of the identified ecological
groups of ground beetles, values of Spearman's rank correlation
coefficient (\emph{r\textsubscript{s}} ) were calculated. Higher MIB
values were significantly positively correlated with the ascending age
of spruce forests (\emph{r\textsubscript{s}} =0.26), and with the
occurrence of large zoophagous beetles (\emph{r\textsubscript{s}} =0.34)
and the group of ground beetles having a high demand for moisture, i.e.
hygrophilous (\emph{r\textsubscript{s}} =0.38) and wetland species
(\emph{r\textsubscript{s}} =0.29) (Table 3). Low values of the MIB index
were caused by the presence of numerous small zoophagous beetles
(\emph{r\textsubscript{s}} =-0.87). A significant correlation, in this
case, was also found with the numerous occurrence of open area
(\emph{r\textsubscript{s}} =-0.61), macropterous
(\emph{r\textsubscript{s}} =-0.58) and xerophilous
(\emph{r\textsubscript{s}} =-0.58) beetles, and spring breeders
(\emph{r\textsubscript{s}} =-0.55). Higher values of the Shannon H'
index were also correlated with a decrease in the MIB index values
(\emph{r\textsubscript{s}} =-0.80) (Table 3).

The ANOSIM analysis indicated significant differences (R=0.31,
p\textless{}0.01) between the carabid assemblages inhabiting the three
age ranges of spruce stands; this significant differences are shown on
the NMDS diagram (Figure 4). More information about the ground beetle
assemblages is shown on the RDA diagram (Figure 5). The RDA analysis
presents the distribution of ground beetle species and indicates their
ecological groups along environmental gradients, represented by two main
canonical axes, describing 67.8 \% of the variance. The redundancy
analysis RDA showed that the B objects (40-50 years old), situated in
two different forest habitats (also different in terms of humidity) are
grouped in one part of the ordination diagram. Similar regularity is
observed for most objects located in stand A (20-30 years old), which
are grouped in the upper, left-hand quarter of the diagram and correlate
with the 2nd ordination axis, in addition to which they are also
localised in different types of forest habitat. Our analysis of this
part of the diagram also indicates that the high shading of soil
(canopy) was correlated with the presence of open area species (Oa),
small zoophagous species (Sz) with the spring type of breeding (Sb), and
xerophilous (Xe) species.

Further analysis of the RDA diagram shows that the first ordination axis
correlated with the presence of large zoophagous species (Lz), which is
also linked to high values of the MIB. This axis is also strongly
correlated with habitat humidity and to the presence of hygrophilous
species (H) and species associated with wet areas (Wet),
e.g.\emph{Carabus granulatus} , \emph{Oxypselaphus obscurus} . The
fertility of the habitat was not correlated with the first two
ordination axes describing most of the ground beetle assemblages.

\textbf{Discussion}

The species richness obtained in this study on ground beetles in spruce
forests was relatively high, although it needs to be added that this is
the total of ground beetles caught in 4 localities situated in 2
macroregions of Poland (the Masurian Lakeland, and the
Che\selectlanguage{polish}ł\selectlanguage{english}mi\selectlanguage{polish}ń\selectlanguage{english}sko-Dobrzy\selectlanguage{polish}ń\selectlanguage{english}skie lake District) (Kondracki 2011). Considering the
above, the number of species caught was similar to the reported number
of species in Finland (Koivula and Niemel\selectlanguage{ngerman}ä 2002, Koivula et al. 2019),
Canada (Niemelä et al. 1993), Slovakia (Macko 2016), and Germany (Lange
et al. 2014). There were differences in the average number of ground
beetle species in the examined spruce forests, with a tendency towards
smaller species richness in older stands (Figure 3a). The same tendency
is seen in the number of caught beetles.

The NMDS analysis showed significant differences between ground beetle
assemblages from the three age classes of spruce stands (Figure 4).
Qualitative and quantitative changes in the assemblages of ground
beetles identified in the older stands (B and C) were mainly due to a
more numerous presence of large forest (\emph{C. hortensis} , \emph{C.
violaceus} ) and eurytopic species (\emph{P. melanarius} , \emph{C.
nemoralis} ). The frequent occurrence of \emph{C. hortensis} is not an
unusual finding. This species, in Scandinavia, is characteristic of
deciduous and mixed forests, and in central Europe it appears in large
numbers in coniferous forests (Lindroth 1985). At the same time,\emph{C.
hortensis} is considered to be typical of mature tree stands (Niemelä et
al. 2007). The more numerous appearance of \emph{C. violaceus} in spruce
stands over 80 years old is worth noting in the context of the observed
decline in its presence in Polish forests. An interesting finding is the
frequent presence of \emph{P. melanarius} and\emph{C. nemoralis} in the
spruce forests covered by our study. These are eurytopic species, which
also appear in forests. However, they belong to the group of deciduous
forest specialists, characteristic of native forests (Magura et al.
2000). \emph{P. melanarius} is not a species that lives in abundance in
spruce forests (Lynikiene 2006, Schwerk and Szyszko 2007), but it is
more abundant in younger forests (Tarwacki 2012).

In the youngest age range (up to 30 years), the dominant ground beetles
were typical forest zoophagous species with poor dispersal power
(brachypterous): \emph{C. micropterus} and \emph{C. arvensis} . \emph{P.
oblongopunctatus} also appeared in large numbers (Table 2). The two
former species were described by Magura et al. (2000), in a study on
ground beetle assemblages in managed and unmanaged spruce plantations,
as species associated with the presence of deciduous trees. These
authors also suggested that tree thinning in managed spruce plantations
induces the growth of a leafy understory, which leads to the
accumulation of leafy litter. In such habitats, the frequent presence
of\emph{P. oblongopunctatus} has been noted. The spruce forests which
were in the youngest age class in our research were also distinguished
by the presence of Carabidae species classified as an open area species
with the spring type of breeding, xerophilous, small zoophagous, and
hemizoophagous. These groups are shown in the ordination diagram (RDA)
and are associated with the objects in this age class of spruce forests
(A).

The above ecological description of species composing assemblages of
ground beetles is typical of forests at an early stage of forest
succession (Buddle et al. 2006, Sk\selectlanguage{polish}ł\selectlanguage{english}odowski 2017, Kosewska et al. 2018).
It was only in the youngest spruce stands (besides the typical forest
species that appeared in all tree stands, i.e. \emph{Amara brunnea} )
that open area species of the genus \emph{Amara} were detected
(\emph{Amara bifrons} , \emph{Amara communis,} and A\emph{mara similata}
). These species are connected with open areas (meadows, agrocenoses,
etc.), and have uniquely strong dispersal power owing to their ability
to fly (Hurka 1996). A similar relationship was determined concerning
two species of the genus \emph{Harpalus} : \emph{Harpalus rufipes} and
\emph{Harpalus tardus} . This group of hemizoophagous species associated
with open areas is capable of inhabiting coniferous forests at the early
stages of succession, which may be explained by their ability to find
plant food in these habitats (Kosewska et al. 2019). The conditions
which are suitable for the development of these species disappear as a
forest grows older (disappearance of plant food). The number of
early-succession species decreases when the tree canopy closes, that is
when the forest is 20 to 30 years old (Niemel\selectlanguage{ngerman}ä et al. 2007).

One of the most popular indicators used to identify stages in the
succession of forests is the mean individual biomass (MIB) index
(Szyszko 1983, Szyszko et al. 2000, Cárdenas and Hidalgo 2007, Jelaska
et al. 2011). The results obtained in our study indicate a growth in the
MIB values determined for ground beetles in older spruce forests (Figure
3c). The calculated values of Spearman's rank correlation coefficient
(Table 3) indicate that high MIB values are related to the presence of
species classified as large zoophages, having high moisture requirements
(hygrophilous and wetland species). Low MIB values were significantly
correlated with the higher species richness (Shannon H') of Carabidae,
the frequent presence of small zoophagous, xerophilous species, and open
area species with high dispersal capability (macropterous). The presence
of species having specific ecological features in younger and older age
classes of spruce forests is closely similar to the situation described
by many other authors (Szyszko 1983, Magura et al. 2002, Niemelä et al.
2007, Sk\selectlanguage{polish}ł\selectlanguage{english}odowski 2009). A certain divergence consists in the decreasing
abundance within the species typical of forest habitats with the growing
age of the spruce forests, which we observed in this study. The most
probable reason was the frequent occurrence in the youngest forests
(class A) of \emph{C. micropterus} and \emph{C. arvensis} (Table 2).
This might be attributed to the fact that all the research sites are
habitats typically inhabited by the forest species of ground beetles,
and the more frequent presence of certain species in one of the age
classes of forests might be due to some specific habitat-related
characteristics (e.g. the type of microhabitats) (Pearce and Venier
2006).

\textbf{Conclusions}

Assemblages of ground beetles dwelling in spruce forests in the
north-eastern part of Poland are characterised by quite high species
richness. In all age classes of spruce forests, the most numerous
carabids were those classified as large, forest zoophages. However,
significant differences were determined between the assemblages of
ground beetles living in the three age classes of spruce forests. The
oldest forests (around 80 years old) were the sites where the smallest
number of carabid species and specimens were captured, but these were
species characteristic for forests at an advanced stage of succession,
which translated into the highest MIB values. The 30- to 50-year-old
spruce forests (objects A and B) were characterised by higher species
richness and a greater number of ground beetles, accompanied by lower
MIB values. The calculated values of Spearman's rank correlation
coefficient reveal that high MIB values are positively correlated with
the presence of large Carabidae species that require more moisture.
Lower MIB values are associated with the presence of smaller,
macropterous species with the spring strategy of breeding, associated
with open areas.

\textbf{Acknowledgements:} We would like to thank Magdalena Obidzi\selectlanguage{polish}ń\selectlanguage{english}ska,
Joanna Skurapowicz, Maciej G\selectlanguage{polish}ą\selectlanguage{english}ga\selectlanguage{polish}ł\selectlanguage{english}a, Jacek Maczyszyn for help with
material collection. We are grateful to Dr Robert Lee for improving the
English. The results presented in this paper were obtained as a part of
comprehensive study financed by the University of Warmia and Mazury in
Olsztyn, Faculty of Agriculture and Forestry, Departament of Entomology,
Phytopathology and Molecular Diagnostic.

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Table 1. Description and characteristics of spruce forests\selectlanguage{english}
\begin{longtable}[]{@{}llllllllll@{}}
\toprule
Age variant & District & Plot & Age {[}years{]} & Height above sea level
{[}m{]} & Soil & Type of forest & Share of spruce {[}\%{]} & Canopy
{[}\%{]} & No of treatments *\tabularnewline
\midrule
\endhead
A & Sosnowo & A1 & 28 & 135 & haplic brunic arenosol & mixed fresh
coniferous forest & 100 & 100 & 1\tabularnewline
& & A2 & 28 & 135 & haplic brunic arenosol & mixed fresh coniferous
forest & 100 & 100 & 1\tabularnewline
& Zielony Dw\selectlanguage{ngerman}ór & A3 & 28 & 151 & haplic luvisol & fresh forest & 80 &
100 & 1\tabularnewline
& & A4 & 28 & 151 & haplic luvisol & fresh forest & 80 & 100 &
1\tabularnewline
B & Zielony Dw\selectlanguage{ngerman}ór & B1 & 43 & 150 & haplic luvisol & fresh forest & 80 &
90 & 2\tabularnewline
& & B2 & 43 & 150 & haplic luvisol & fresh forest & 80 & 90 &
2\tabularnewline
& Maruny & B3 & 48 & 139 & albic brunic arenosol & mixed fresh forest &
90 & 90 & 2\tabularnewline
& & B4 & 48 & 139 & albic brunic arenosol & mixed fresh forest & 90 & 90
& 2\tabularnewline
C & Przejmy & C1 & 78 & 120 & mucky soils & moist forest & 70 & 80 &
7\tabularnewline
& & C2 & 78 & 120 & mucky soils & moist forest & 70 & 80 &
7\tabularnewline
& & C3 & 78 & 119 & mucky soils & moist forest & 70 & 80 &
7\tabularnewline
& & C4 & 78 & 119 & mucky soils & moist forest & 70 & 80 &
7\tabularnewline
\bottomrule
\end{longtable}

* Treatments: cleaning; thinning

Table 2. Abundance, ecological classification and species name
abbreviations used in the RDA analysis of ground beetles in age variants
of spruce forests\selectlanguage{english}
\begin{longtable}[]{@{}lllllllllllllll@{}}
\toprule
& Abbre-viations & Ecological characteristic & Individuals / Percentage
& Individuals / Percentage & Individuals / Percentage & Individuals /
Percentage & Individuals / Percentage & Individuals / Percentage &
Individuals / Percentage & Individuals / Percentage & Individuals /
Percentage & Individuals / Percentage & Individuals / Percentage &
Individuals / Percentage\tabularnewline
\midrule
\endhead
Species & & & A* & A* & A* & A* & B & B & B & B & C & C & C &
C\tabularnewline
& & & A1, A2 & A1, A2 & A3, A4 & A3, A4 & B1, B2 & B1, B2 & B3, B4 & B3,
B4 & C1, C2 & C1, C2 & C3, C4 & C3, C4\tabularnewline
& & & n & \% & n & \% & n & \% & n & \% & n & \% & n & \%\tabularnewline
\emph{Amara bifrons} (Gyllenhal,1810) & A\_bifr & Oa/Xe/Hz/Ab/Ma & 2 &
0.3 & & & & & & & & & &\tabularnewline
\emph{A. brunnea} (Gyllenhal, 1810) & A\_bru & F/M/Hz/Ab/Ma & 4 & 0.6 &
& & & & & & 4 & 0.8 & 2 & 0.5\tabularnewline
\emph{A. communis} (Panzer, 1797) & A\_com & Oa/M/Hz/Sb/Ma & 1 & 0.1 & &
& & & & & & & &\tabularnewline
\emph{A. similata} (Gyllenhal,1810) & A\_simi & Oa/M/Hz/Sb/Ma & 1 & 0.1
& & & & & & & & & &\tabularnewline
\emph{Badister lacertosus} Sturm, 1815 & Ba\_lace & F/M/Sz/Sb/Ma & & & &
& 2 & 0.4 & & & 1 & 0.2 & &\tabularnewline
\emph{Calathus ambiguus} (Paykull,1790) & Ca\_amb & Oa/Xe/Sz/Ab/Ma & 1 &
0.1 & & & & & & & & & &\tabularnewline
\emph{C. erratus} (Sahlberg, 1827) & Ca\_err & F/Xe/Sz/Ab/D & 2 & 0.3 &
& & & & & & & & &\tabularnewline
\emph{C. fuscipes} Goeze, 1777 & Ca\_fusc & Oa/M/Sz/Ab/B & 26 & 3.9 & &
& & & 3 & 0.5 & & & &\tabularnewline
\emph{C. melanocephalus} (Linnaeus, 1758) & Ca\_mela & Oa/M/Sz/Ab/B & 3
& 0.4 & & & & & & & 9 & 1.9 & 13 & 3.4\tabularnewline
\emph{C. micropterus} (Duftschmid, 1812) & Ca\_micr & F/M/Sz/Ab/B & 181
& 26.3 & & & 2 & 0.4 & 14 & 2.3 & 19 & 4.0 & 17 & 4.5\tabularnewline
\emph{Carabus arvensis} Herbst,1784 & C\_arv & F/Xe/Lz/Sb/B & 89 & 12.9
& & & & & & & 3 & 0.6 & &\tabularnewline
\emph{C. cancellatus} Illiger,1798 & C\_canc & Eu/M/Lz/Sb/B & 3 & 0.4 &
18 & 4.1 & 12 & 2.3 & & & 3 & 0.6 & 3 & 0.8\tabularnewline
\emph{C. convexus} Fabricius, 1775 & C\_conv & F/Xe/Lz/Sb/B & 12 & 1.8 &
& & & & & & & & &\tabularnewline
\emph{C. coriaceus} Linnaeus, 1758 & C\_coria & F/M/Lz/Ab/B & & & & & &
& 1 & 0.2 & & & &\tabularnewline
\emph{C. glabratus} Paykull, 1790 & C\_glab & F/M/Lz/Ab/B & 3 & 0.4 & &
& & & 1 & 0.2 & 4 & 0.8 & 6 & 1.6\tabularnewline
\emph{C. granulatus} Linnaeus, 1758 & C\_gran & Wet/H/Lz/Sb/B & 1 & 0.1
& 12 & 2.8 & 4 & 0.8 & 1 & 0.2 & & & 5 & 1.3\tabularnewline
\emph{C. hortensis} Linnaeus, 1758 & C\_hort & F/M/Lz/Ab/B & 118 & 17.2
& 215 & 49.0 & 236 & 46.0 & 108 & 18.0 & 229 & 48.0 & 172 &
45.0\tabularnewline
\emph{C. marginalis} Fabricius, 1794 & C\_marg & F/M/Lz/Sb/B & 2 & 0.3 &
& & & & 5 & 0.8 & & & &\tabularnewline
\emph{C. nemoralis} O.F.Muller, 1764 & C\_nemo & Eu/M/Lz/Sb/B & 1 & 0.1
& 19 & 4.4 & 35 & 6.8 & 23 & 3.8 & 28 & 5.9 & 19 & 5.0\tabularnewline
\emph{C. violaceus} Linnaeus, 1758 & C\_viol & F/M/Lz/Ab/B & 10 & 1.5 &
& & & & 11 & 1.8 & 11 & 2.3 & 15 & 3.9\tabularnewline
\emph{Clivina fossor} (Linnaeus,1758) & Cl\_fos & Oa/M/Sz/Sb/D & & & & &
1 & 0.2 & & & & & &\tabularnewline
\emph{Cychrus caraboides} (Linnaeus, 1758) & Cy\_cara & F/M/Lz/Ab/B & &
& 12 & 2.8 & 8 & 1.6 & & & 10 & 2.1 & 2 & 0.5\tabularnewline
\emph{Harpalus laevipes} Zetterstedt, 1828 & H\_leav & F/M/Hz/Sb/Ma & 1
& 0.1 & 5 & 1.1 & 3 & 0.6 & 5 & 0.8 & 10 & 2.1 & 8 & 2.1\tabularnewline
\emph{H. rufipes} (De Geer, 1774) & H\_rufipe & Oa/M/Hz/Ab/Ma & 4 & 0.6
& & & & & 3 & 0.5 & & & &\tabularnewline
\emph{H. tardus} (Panzer, 1797) & H\_tar & Oa/Xe/Hz/Sb/Ma & 3 & 0.4 & &
& & & 2 & 0.3 & & & &\tabularnewline
\emph{Leistus rufomarginatus}(Duftschmid, 1812) & L\_rufo & F/M/Sz/Ab/B
& 2 & 0.3 & & & & & & & 2 & 0.4 & &\tabularnewline
\emph{L. terminatus} (Hellwig, 1770) & L\_term & Wet/H/Sz/Ab/B & & & 1 &
0.2 & & & 1 & 0.2 & 1 & 0.2 & 1 & 0.3\tabularnewline
\emph{Limodromus assimilis} (Paykull, 1790) & Li\_ass & F/H/Sz/Sb/Ma & &
& 11 & 2.5 & 10 & 2.0 & & & 10 & 2.1 & 9 & 2.4\tabularnewline
\emph{Loricera pilicornis} (Fabricius,1775) & Lo\_pili & Wet/H/Sz/Sb/Ma
& Wet/H/Sz/Sb/Ma & & 1 & 0.2 & & & & & & & &\tabularnewline
\emph{Nebria brevicollis} (Fabricius, 1792) & N\_brev & Eu/M/Sz/Ab/Ma &
& & 1 & 0.2 & & & 55 & 9.0 & 8 & 1.7 & &\tabularnewline
\emph{Notiophilus biguttatus} (Fabricius, 1779) & No\_big & F/M/Sz/Sb/D
& 18 & 2.7 & & & 2 & 0.4 & 2 & 0.3 & 1 & 0.2 & 1 & 0.3\tabularnewline
\emph{N. palustris} (Duftschmid, 1812) & No\_pal & Oa/M/Sz/Sb/B & 1 &
0.1 & & & & & 8 & 1.3 & & & &\tabularnewline
\emph{Ophonus rufibarbis} (Fabricius,1792) & Op\_rufi & Eu/M/Sz/Ab/Ma &
& & 1 & 0.2 & & & & & & & &\tabularnewline
\emph{Oxypselaphus obscurus} (Herbst, 1784) & Ox\_ob & F/H/Sz/Sb/Ma & &
& 1 & 0.2 & & & 1 & 0.2 & & & &\tabularnewline
\emph{Patrobus atrorufus} (Strom,1768) & Pa\_atr & Wet/H/Sz/Ab/B & & & 4
& 0.9 & & & & & & & &\tabularnewline
\emph{Poecilus lepidus} (Leske,1785) & Po\_lep & Oa/Ks/Sz/Sb/D & 3 & 0.4
& & & & & & & & & &\tabularnewline
\emph{P. versicolor} (Sturm, 1824) & Po\_ver & Oa/M/Sz/Sb/Ma & 1 & 0.1 &
& & 1 & 0.2 & 2 & 0.3 & 3 & 0.6 & 6 & 1.6\tabularnewline
\emph{Pterostichus anthracinus} (Illiger,1798) & P\_anth & Wet/H/Sz/Sb/D
& 1 & 0.1 & & & 1 & 0.2 & & & & & &\tabularnewline
\emph{P. melanarius} (Illiger, 1798) & P\_mela & Eu/M/Lz/Ab/D & & & 30 &
6.9 & 55 & 11.0 & 118 & 19.0 & 41 & 8.6 & 40 & 11.0\tabularnewline
\emph{P. niger} (Schaller, 1783) & P\_nige & F/M/Lz/Ab/Ma & 86 & 12.5 &
66 & 15.0 & 87 & 17.0 & 175 & 29.0 & 22 & 4.6 & 29 & 7.6\tabularnewline
\emph{P. oblongopunctatus} (Fabricius, 1787) & P\_obl & F/M/Sz/Sb/Ma &
109 & 15.8 & 38 & 8.7 & 52 & 10.0 & 70 & 11.0 & 58 & 12.0 & 32 &
8.4\tabularnewline
\emph{P. strenuus} (Panzer, 1797) & P\_str & F/H/Sz/Sb/D & & & & & & & 2
& 0.3 & & & &\tabularnewline
\emph{P. vernalis} (Panzer,1796) & P\_ver & Oa/H/Sz/Sb/Ma & & & 1 & 0.2
& & & & & & & &\tabularnewline
\emph{Syntomus truncatellus} (Linnaeus,1761) & Sy\_tru & Oa/Xe/Sz/Sb/B &
1 & 0.1 & & & & & & & & & &\tabularnewline
Individuals - Total & & & 690 & 100 & 436 & 100 & 511 & 100 & 611 & 100
& 477 & 100 & 380 & 100\tabularnewline
\bottomrule
\end{longtable}

* A: 20-30 years; B: 40-50 years; C: 70 - 80 years

Hz- hemizoophages, Sz- small zoophages, Lz- large zoophages, Ab- autumn
breeders, Sb- spring breeders, F- forest, Oa- open area, Eu- eurytopic,
Wet - wetland, M- mesophilous, Xe- xerophilous, H- hygrophilous, Ma-
macropteric, B- brachipteric, D- dipteric

Table 3. Results of statistical tests (GLM, ANOVA) and values of
Spearman's rank correlation coefficient for species richness, abundance,
diversity and MIB values of Carabidae ecological groups within age
classes of spruce forests\selectlanguage{english}
\begin{longtable}[]{@{}llllllll@{}}
\toprule
Specifications & Means for the analysed age & Means for the analysed age
& Means for the analysed age & Wald's statistic & ANOVA \emph{F} value &
\emph{p} & \emph{r}\textsubscript{s} Spearman p \textless{}
0.05\tabularnewline
\midrule
\endhead
& classes of spruce forests & classes of spruce forests & classes of
spruce forests & & & &\tabularnewline
& A & B & C & & & &\tabularnewline
Individuals & 5.12 & 5.10 & 3.90 & 45.65 & - & 0.00 & n. s.
*\tabularnewline
Species & 2.78 & 2.43 & 1.50 & 84.75 & - & 0.00 & n. s.\tabularnewline
Shannon H' & 0.78 & 0.75 & 0.71 & 0.07 & - & 0.96 & -0.80\tabularnewline
MIB (mg) & 273.82 & 286.67 & 341.84 & 173.1 & - & 0.00 &
-\tabularnewline
* Eu & 3.65 & 14.90 & 7.10 & 138.18 & - & 0.00 & n. s.\tabularnewline
F & 49.25 & 39.85 & 33.85 & 58.29 & - & 0.00 & n. s.\tabularnewline
Oa & 2.40 & 1.00 & 1.55 & 11.51 & - & 0.00 & -0.61\tabularnewline
Wet & 1.00 & 0.35 & 0.35 & 9.08 & - & 0.01 & 0.29\tabularnewline
H & 1.65 & 1.00 & 1.30 & 3.17 & - & 0.21 & 0.38\tabularnewline
Xe & 5.65 & 0.10 & 0.15 & 69.01 & - & 0.00 & -0.58\tabularnewline
M & 49.00 & 55.00 & 41.40 & - & 3.34 & 0.04 & n. s.\tabularnewline
Hz & 1.05 & 0.65 & 1.20 & 3.26 & - & 0.20 & n. s.\tabularnewline
Lz & 34.85 & 44.00 & 32.10 & 41.70 & - & 0.00 & 0.34\tabularnewline
Sz & 20.40 & 11.45 & 9.55 & 93.60 & - & 0.00 & -0.87\tabularnewline
Ab & 38.60 & 43.90 & 32.85 & 31.61 & - & 0.00 & n.s.\tabularnewline
Sb & 17.70 & 12.20 & 10.00 & 46.33 & - & 0.00 & -0.55\tabularnewline
B & 36.70 & 23.65 & 28.60 & 57.85 & - & 0.00 & n.s.\tabularnewline
D & 2.70 & 9.05 & 4.15 & 76.28 & - & 0.00 & n.s.\tabularnewline
Ma & 16.90 & 23.40 & 10.10 & 100.78 & - & 0.00 & -0.58\tabularnewline
Age of forest & \textless{} 80 & \textless{} 50 & \textless{} 30 & - & -
& - & 0.26\tabularnewline
(years) & & & & & & &\tabularnewline
\bottomrule
\end{longtable}

* n. s. non sagnificant; ** - explanations of the abbreviations is given
in Table 2\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

Figure 1. Location of the study area, with distribution of sampling
sites and transect of traps\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}

Figure 2. Expected number of carabid species caught in spruce forests
using the Jackknife estimator (\selectlanguage{ngerman}±SD) of species richness\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}

Figure 3. Mean values of species richness (1a), abundance (1b) and MIB
(1c) of ground beetles in the three age classes of spruce forests (*
Vertical lines denote SE; a, b\ldots{} - means indicated by the same
letter do not differ according to the Bonferroni test)\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image4/image4}

\end{center}
\end{figure}

Figure 4. Diagram of non-metric multidimensional scaling (NMDS) for
carabid assemblages in relation to age classes of spruce forests (A:
20-30 years; B: 40-50 years; C: 70-80 years)\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image5/image5}

\end{center}
\end{figure}

Figure 5. Diagram of redundance analysis (RDA) illustrating dependences
between captured species of Carabidae, their ecological groups, and
environmental variables (MIB, humidity, no of treatments, fertility,
canopy and forest age) (Abbreviations are given in Table 2)

\selectlanguage{english}
\FloatBarrier
\end{document}

