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Citation: Ruchin, A.B.; Egorov, L.V.;
Khapugin, A.A. Vertical Distribution
of Beetles (Coleoptera) in Pine
Forests in Central European Russia.
Diversity 2022,14, 622. https://
doi.org/10.3390/d14080622
Academic Editor: Luc Legal
Received: 14 July 2022
Accepted: 2 August 2022
Published: 3 August 2022
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diversity
Article
Vertical Distribution of Beetles (Coleoptera) in Pine Forests in
Central European Russia
Alexander B. Ruchin 1, * , Leonid V. Egorov 1,2 and Anatoliy A. Khapugin 1,3
1Joint Directorate of the Mordovia State Nature Reserve and National Park «Smolny», Saransk 430005,
Republic of Mordovia, Russia; platyscelis@mail.ru (L.V.E.); hapugin88@yandex.ru (A.A.K.)
2Prisursky State Nature Reserve, Cheboksary 428034, Chuvash Republic, Russia
3Institute of Environmental and Agricultural Biology (X-BIO), Tyumen State University,
Tyumen 625003, Russia
*Correspondence: ruchin.alexander@gmail.com; Tel.: +7-83445-296-35
Abstract:
Research on the Arthropoda distribution in temperate forest ecosystems has shown that
communities are distributed vertically in unequal numbers. However, this issue still has research gaps
in relation to the vertical stratification of Coleoptera in pine forests of the temperate zone. This study
was conducted in the Republic of Mordovia situated in central part of European Russia. We used
fermental traps to collect Coleoptera (the bait is fermenting beer with sugar and honey). The sampling
was conducted from May to September 2021 in five sites of pine forests. One hundred and twenty-five
species from 36 families were identified. The highest species richness was found in Nitidulidae and
Cerambycidae (19 species each), Elateridae (13), Curculionidae (7) and Scarabaeidae (6). The highest
number of species (84 species) was obtained at the height of 1.5 m, while the smallest species richness
(44 species) was found at the height of 12.0 m. At the height of 12 m, we also registered the minimum
number of specimens. Twenty-four species from 11 families were common to all the heights studied.
Cryptarcha strigata and Glischrochilus grandis preferred heights of 3 and 1.5 m. Cryptarcha undata and
Protaetia marmorata were more common at a height of 7 m with a frequency of 61.4–79.6% and 68.2–79.6%,
respectively. The absolute number of saproxyl and anthophilic beetle species was higher in the ground
layer and at the undergrowth level. These studies expand our understanding of the vertical distribution
of beetles in pine forests of the temperate zone of European Russia.
Keywords:
Insecta; species diversity; beer trap; saproxylic beetles; protected area; Mordovia State
Nature Reserve
1. Introduction
Among temperate ecosystems, pine forests have an outstanding richness of plant and
animal communities and biodiversity [
1
–
4
]. In the territory of Europe, coniferous forests
dominated by Pinus sylvestris L. are widely spread due to their high environmental and
economic importance (e.g., [
5
,
6
]). Although pine forests are generally characterized by
a lower biodiversity level in comparison with deciduous forests [
7
], P. sylvestris forms
habitats suitable for a large number of various organisms (e.g., [8,9]).
Forest systems are characterized by a high and spatially structured diversity of habi-
tats [
10
–
17
]. In recent years, the distribution of insects has been actively studied along ver-
tical gradients in forests of various zones, from tropical forests to temperate
forests [18–23]
.
In floodplain forests in Germany, alpha and beta diversity of both rare and common beetle
species in undergrowth and canopy differed significantly. It was shown that the species
richness is higher in the undergrowth layer than in the forest canopy for common species,
but it is lower for rare beetle species [
24
]. In a Sulawesi rainforest, 56% more common
species out of 701 Coleoptera showed no preference for layers. However, there were species
expressing obvious preferences for a certain height, 39% out of them were specific to the
forest canopy and only 5% for the ground layer [
25
]. At the height of 10.5 m, the abundance,
Diversity 2022,14, 622. https://doi.org/10.3390/d14080622 https://www.mdpi.com/journal/diversity
Diversity 2022,14, 622 2 of 21
species richness and biomass of Cetoniinae were higher in trap set. In this case, traps
should be located at least 4.5 m above the ground [
26
]. The average number of Dicerca
divaricata (Buprestidae) specimens in traps with attractants was statistically significantly
lower in the undergrowth layer than in any other positions of traps [
27
]. Collections of
Monochamus galloprovincialis (Cerambycidae) individuals in traps located in tree crowns
were larger than in traps situated at low heights [28].
Traps placed in the canopy of deciduous forest at the height of 13 m caught signifi-
cantly more specimens of Agrilus planipennis (Buprestidae) than those placed at the ground
level [
29
]. The Ips duplicatus (Curculionidae) abundance in pheromone traps was higher at
the height of 1.5–2.0 m than in traps at the ground level or at the height of 3.5–4.0 m [
30
].
Using bait traps located at various heights, Miller et al. [
31
] found certain preferences
for many Coleoptera species. Among them, Cnestus mutilatus (Curculionidae), Neoclytus
scutellaris, and Monochamus titillator (Cerambycidae) confined to heights of 18–23 m, while
Xylosandrus crassiusculus,Dendroctonus terebrans (Curculionidae) and Neoclytus acuminatus
(Cerambycidae) were more frequent at the height of 0.3–0.5 m. In forest systems of the East-
ern Canada, the Cerambycidae diversity was higher in the canopy than in the undergrowth
layer, whereas their abundance was higher in the undergrowth layer compared with the
canopy layer [
32
]. In the understory layer of beech–fir and oak forests of France, there
was a higher abundance and richness of Coleoptera species [
33
]. In temperate deciduous
forests in Japan, the abundance and composition of flying Coleoptera varied depending on
layers and seasons: Elateridae were most common in the lower layer, while Attelabidae
and Cantharidae were most common in the upper layer of the forest [34].
In this paper, we used beer-baited traps, placed at various heights in the tree canopy,
to study the occurrence frequency, abundance and diversity of Coleoptera species along
the vertical gradient in pine forests in the Central European Russia.
2. Materials and Methods
2.1. Research Area
The study was conducted in the Republic of Mordovia (Temnikov district, Mordovia
State Nature Reserve: 54.42–54.56
◦
N, 43.04–43.36
◦
E) (Figure 1). The studied protected area
covers 321.62 km
2
. According to the natural zoning, the study area is located on the border
of the zone of coniferous–deciduous forests and the forest-steppe. In the Mordovia State
Nature Reserve, forests cover 89.3% of the entire protected area. This forest type represents
the largest refugium for threatened insects in the region. Thus, pine forests provide a high
diversity of habitats [35,36].
Diversity 2022, 14, 622 3 of 22
Figure 1. Study sites in the Mordovia State Nature Reserve, European Russia.
The study was conducted in the green-moss pine forest. Pinus sylvestris L. forms the
canopy layer. The understory layer is quite thin, being represented by Betula pendula
Roth, and singular trees of Picea abies (L.) H.Karst. and Populus tremula L. In the under-
growth layer, the following shrubs are represented: Sorbus aucuparia L., Rubus idaeus L.
and Frangula alnus Mill. with a total percent cover of ca. 35%. The ground layer vegeta-
tion is dominated by Convallaria majalis L. (ca. 45%) with participation of Calamagrostis
epigejos (L.) Roth (ca. 5–7%). Other vascular plants of the ground layer are Dryopteris car-
thusiana (Vill.) H.P.Fuchs, Fragaria vesca L., Melampyrum pratense L., Vaccinium myrtillus L.,
Anthoxanthum odoratum L., Galium mollugo L., Agrostis gigantea Roth, Lysimachia europaea
(L.) U.Manns & Anderb., Luzula pilosa (L.) Willd. and Calamagrostis arundinacea (L.) Roth.
2.2. Sampling Procedures
Coleoptera were collected from May to September 2021, a period when activity of
insects is the highest. In each fragment of a pine forest, we collected samples at four
heights. In this case, traps were located at a distance of no more than 15 m from each
other. Between each fragment of a forest, where four traps were located at different
heights, there was a certain distance indicated below. This distance between sample sta-
tions was an attempt to ensure an independence between samples. In total, there were
five such forest fragments of the pine forest. Each fragment of a forest was a separate
habitat. Each part of the forest was located no less than 0.8 km from each other and by
representing an independent forest site. This distance is needed to obtain a representative
sampling size, since it was previously demonstrated that the distance between traps should
be at least 0.3 km [37].
In each sampling site, a set of four traps was installed on various heights (1.5, 3, 7
and 12 m above the ground) on branches of trees. We selected these heights to facilitate
Figure 1. Study sites in the Mordovia State Nature Reserve, European Russia.
Diversity 2022,14, 622 3 of 21
The study was conducted in the green-moss pine forest. Pinus sylvestris L. forms the
canopy layer. The understory layer is quite thin, being represented by Betula pendula Roth,
and singular trees of Picea abies (L.) H.Karst. and Populus tremula L. In the undergrowth
layer, the following shrubs are represented: Sorbus aucuparia L., Rubus idaeus L. and Frangula
alnus Mill. with a total percent cover of ca. 35%. The ground layer vegetation is dominated
by Convallaria majalis L. (ca. 45%) with participation of Calamagrostis epigejos (L.) Roth
(ca. 5–7%). Other vascular plants of the ground layer are Dryopteris carthusiana (Vill.)
H.P.Fuchs, Fragaria vesca L., Melampyrum pratense L., Vaccinium myrtillus L., Anthoxanthum
odoratum L., Galium mollugo L., Agrostis gigantea Roth, Lysimachia europaea (L.) U.Manns &
Anderb., Luzula pilosa (L.) Willd. and Calamagrostis arundinacea (L.) Roth.
2.2. Sampling Procedures
Coleoptera were collected from May to September 2021, a period when activity of
insects is the highest. In each fragment of a pine forest, we collected samples at four heights.
In this case, traps were located at a distance of no more than 15 m from each other. Between
each fragment of a forest, where four traps were located at different heights, there was a
certain distance indicated below. This distance between sample stations was an attempt to
ensure an independence between samples. In total, there were five such forest fragments of
the pine forest. Each fragment of a forest was a separate habitat. Each part of the forest was
located no less than 0.8 km from each other and by representing an independent forest site.
This distance is needed to obtain a representative sampling size, since it was previously
demonstrated that the distance between traps should be at least 0.3 km [37].
In each sampling site, a set of four traps was installed on various heights (1.5, 3, 7 and
12 m above the ground) on branches of trees. We selected these heights to facilitate the
manual installation of traps without the need to climb trees or to use special techniques
or equipment to install traps. To avoid a possible edge effect, we installed traps inside
the forest areas. The total sampling was equal to 176 trap exposures, where there were
9 repetitions at each height (while there were eight repetitions in one plot).
All collections were conducted by traps made by authors. Each trap consisted of a 5 L
plastic container with an opening on one side at a distance of 10 cm from the bottom [
37
].
For bait, we used the beer mixed with honey and sugar. The collected insects in plastic bags
containing 70% alcohol were delivered from forest sites to the laboratory. Then, the collected
samples were sorted and stored in alcohol. In total, 15,117 specimens were studied.
2.3. Identification and Taxonomic Status of Samples
Coleoptera families were classified according to Bouchard and co-authors [38], with ad-
ditions [
39
]. We took into account changes from the Catalog of Palaearctic
Coleoptera [40–46]
,
publications of Robertson and co-authors [
47
] for Cucujoidea, and Alonso-Zarazaga and
co-authors [
48
] for Curculionoidea. The nomenclature of beetles was standardized according
to the publications cited above, with addition of the Catalog of Palaearctic Coleoptera [
49
,
50
].
Years of description for some species are used according to Bousquet [51].
2.4. Data Analysis
We analyzed the number of trapped beetles per exposure time. Exposure time repre-
sented the period (counted in days) between installing a trap a taking the samples to be
analyzed; we collected a total of 44 exposure times for each height value. The occurrence
frequency is considered the ratio of the number of samples in which a taxon is present to
the total number of samples (counted in %). Saproxylic species were identified based on
approaches adopted in some publications [52–55]. We classified anthophilic beetles based
on our personal long-term observations.
The Jaccard index was used to assess the similarity of species composition between
applied heights. Those insects that could not be identified to the species levels were
discarded from the analyses. The diversity index of Shannon and the dominance index of
Simpson were calculated based on the collected data [56,57].
Diversity 2022,14, 622 4 of 21
All statistical analyses were performed using PAST 4.07 [
58
]. The ordination was
made using PCA (principal component analysis). Using this technique, we defined major
gradients in the spatial arrangement of the studied species selected for analysis. We used
the species abundance as response variables in PCA. For interpretation of the ordination
axes, groups of the used height were plotted onto the PCA ordination diagram as supple-
mentary environmental data. In the analysis, we included species represented by at least
100 specimens
collected over the sampling period. The coefficient of determination (R
2
, or
R-squared) was used according to Moore et al. [59].
3. Results
As a result of the conducted research, 125 beetle species from 36 families (Appendix A)
were identified. Some specimens from the families of Staphylinidae, Nitidulidae, Throsci-
dae, Scraptiidae, Laemophloeidae and Scirtidae were not identified to the species level.
The highest species richness was recorded in Nitidulidae and Cerambycidae (19 species
each), Elateridae (13 species), Curculionidae (7 species) and Scarabaeidae (6 species)
(Figures 2and 3)
. In total, these five families, along with the family Staphylinidae, in-
clude 95.8% of all identified specimens.
Diversity 2022, 14, 622 5 of 22
Figure 2. The proportion of species in five Coleoptera families collected using beer traps at various
heights (see Appendix A).
Figure 3. The proportion of specimens in five Coleoptera families collected using beer traps at
various heights.
Figure 2.
The proportion of species in five Coleoptera families collected using beer traps at various
heights (see Appendix A).
Cryptarcha strigata (Fabricius, 1787) (5809 specimens in total), Glischrochilus grandis
(Tournier, 1872) (3028 specimens), Protaetia marmorata (Fabricus, 1792) (765 specimens),
Rhagium mordax (De Geer, 1775) (615 specimens) and Glischrochilus hortensis (Geoffroy, 1785)
(519 specimens) had the highest number of specimens in traps.
Twenty-four species from 11 families were common to all studied heights (Gnathoncus
buyssoni Auzat, 1917; Quedius dilatatus (Fabricius, 1787); Cetonia aurata (Linnaeus, 1758);
Gnorimus variabilis (Linnaeus, 1758); P. marmorata;Dalopius marginatus (Linnaeus, 1758);
Attagenus schaefferi (Herbst, 1792); Ctesias serra (Fabricius, 1792); Trogoderma glabrum (Herbst,
1783); Thanasimus femoralis (Zetterstedt, 1828); C. strigata;Cryptarcha undata (G.-A. Olivier,
1790); G. grandis;G. hortensis;Glischrochilus quadripunctatus (Linnaeus, 1758); Soronia grisea
(Linnaeus, 1758); Halyzia sedecimguttata (Linnaeus, 1758); Litargus connexus (Geoffroy, 1785);
Leptura quadrifasciata Linnaeus, 1758; Leptura thoracica Creutzer, 1799; Lepturalia nigripes
Diversity 2022,14, 622 5 of 21
(De Geer, 1775); Pachyta quadrimaculata (Linnaeus, 1758); and Rh. mordax;Anisandrus dispar
(Fabricius, 1792)).
Diversity 2022, 14, 622 5 of 22
Figure 2. The proportion of species in five Coleoptera families collected using beer traps at various
heights (see Appendix A).
Figure 3. The proportion of specimens in five Coleoptera families collected using beer traps at
various heights.
Figure 3.
The proportion of specimens in five Coleoptera families collected using beer traps at
various heights.
The highest species richness (84 species) was found at the height of 1.5 m, and the
smallest number of species (44) was registered at the height of 12 m (Table 1). According
to the average number of specimens, the highest number of species was found at heights
of 1.5 m and 3 m (in average, 120 specimens). The minimum number of specimens was
recorded at the height of 12 m. The proportion of saproxylic beetles was lower at the height
of 1.5 m, while at other heights it is higher (Table 1). However, the absolute number of
saproxylic species was higher at the heights of 1.5 m and 3 m than at other heights. The
proportion of anthophilic species was also lower at the lower height. At the height of
12 m, the number of anthophilic species was the largest (Table 1). The absolute number of
saproxylic species at heights of 1.5 m and 3 m exceeded the number of saproxylic species at
other heights.
The Shannon and Simpson indices show the following results. At the height of 1.5 m,
the Shannon index reached its maximal value, while the Simpson index showed its minimal
value. Conversely, the minimal values of the Shannon index and maximal values of the
Simpson index were found at the height of 12 m. The intermediate values between these
values were obtained for other heights (Appendix A).
The values of the calculated Jaccard similarity index demonstrated certain differences
among the studied heights (Figure 4). The highest difference values were obtained between
heights of 1.5 m and 12 m. At the same time, differences were minimal between the heights
of 3 m and 7 m.
Diversity 2022,14, 622 6 of 21
Table 1.
The main parameters of Coleoptera individuals collected using beer traps at various heights.
Parameters Height (m)
1.5 3 7 12
Total of families 28 24 17 18
Total of individuals 5281 5273 3335 1228
Mean number of individuals per trap 120 120 76 28
Number of species (excluding unidentified insects) 84 69 59 44
Number of saproxylic species 56 52 48 33
Proportion (%) of saproxylic species 66.7 75.4 81.4 75.0
Number of anthophilic species 34 34 26 24
Proportion (%) of anthophilic species 40.5 49.3 44.1 54.5
Shannon index 2.24 1.91 1.85 1.81
Simpson index 0.20 0.26 0.30 0.32
Diversity 2022, 14, 622 7 of 22
Figure 4. The similarity of beetle species composition between four studied heights based on the
Jaccard index. We applied Ward’s method and Euclidean distance (cophenetic correlation coeffi-
cient = 0.771).
During the experiment, it was shown that only 11 Coleoptera species had a total
number of more than 100 specimens (P. marmorata, C. serra, C. strigata, C. undata, G. gran-
dis, G. hortensis, G. quadripunctatus, S. grisea, L. thoracica, Rh. mordax and A. dispar). We
calculated the preferred heights for them.
The distant position of C. strigata and G. grandis along Principal Component 1 (PC1)
in Figure 5 reflects their highest relative abundance in the studied pine forest. These
species indeed had 3.5–92.5-fold and 1.6–47.8-fold, respectively, higher abundance at all
heights in comparison with other species. According to the number of trapped speci-
mens, C. strigata seems to prefer the 3 m height, followed by 1.5 m, while its abundance
declined with an increase in the height of the trap location. The same tendency was found
for another of the most-abundant species, G. grandis, which was characterized by higher
abundance values at the heights of 3 m and 1.5 m; the abundance of this species was
much lower at 7 m, falling to an average value of 1.32 specimens per trap at 12 m (Figure
5).
Figure 4.
The similarity of beetle species composition between four studied heights based on
the Jaccard index. We applied Ward’s method and Euclidean distance (cophenetic correlation
coefficient = 0.771).
During the experiment, it was shown that only 11 Coleoptera species had a total
number of more than 100 specimens (P. marmorata,C. serra,C. strigata,C. undata,G. grandis,
G. hortensis,G. quadripunctatus,S. grisea,L. thoracica,Rh. mordax and A. dispar). We calculated
the preferred heights for them.
The distant position of C. strigata and G. grandis along Principal Component 1 (PC1) in
Figure 5reflects their highest relative abundance in the studied pine forest. These species
indeed had 3.5–92.5-fold and 1.6–47.8-fold, respectively, higher abundance at all heights in
comparison with other species. According to the number of trapped specimens, C. strigata
seems to prefer the 3 m height, followed by 1.5 m, while its abundance declined with an
increase in the height of the trap location. The same tendency was found for another of the
most-abundant species, G. grandis, which was characterized by higher abundance values at
Diversity 2022,14, 622 7 of 21
the heights of 3 m and 1.5 m; the abundance of this species was much lower at 7 m, falling
to an average value of 1.32 specimens per trap at 12 m (Figure 5).
Diversity 2022, 14, 622 8 of 22
Figure 5. The diagram of the principal component analysis ordination of the selected beetle species
in the Republic of Mordovia (European Russia) based on the number of specimens collected at
various heights of trap positions (1.5 m, 3 m, 7 m, 12 m). Designations: P.mar.—Protaetia marmorata;
C.ser.—Ctesias serra; C.str.—Cryptarcha strigata; C.und.—Cryptarcha undata; G.gra.—Glischrochilus
grandis; G.hort.—Glischrochilus hortensis; G.quad.—Glischrochilus quadripunctatus; S.gris.—Soronia
grisea; L.thor.—Leptura thoracica; Rh.mor.—Rhagium mordax; A.dis.—Anisandrus dispar. Groups of
heights of trap positions were plotted onto the ordination diagram as supplementary environ-
mental variables.
4. Discussion
This paper provides an idea of clusters of beetles captured using beer traps installed
at various heights in pine forests of the temperate zone of European Russia. Previously, it
was widely believed that temperate forests are less stratified than tropical forests because
of the smaller structural differences and seasonal changes in climatic conditions that force
many species to migrate between layers [60]. However, subsequent studies, as well as our
research, have shown that beetle communities in the canopy differ from communities in
the undergrowth. Longhorn and scarab beetles were studied by Touroult and Dalens [61]
using bait traps placed at three various heights: 3–5 m (undergrowth), 10–15 m and 25–30
m (under canopy). It was found that, for longhorn beetles, the abundance is higher in the
undergrowth, while the richness is much higher in the canopy. For scarab beetles,
abundance and richness are the same in all three layers. In temperate deciduous forests in
Japan, beetle abundance was higher in the canopy relative to the understory [62]. In On-
tario (Canada), a higher abundance of Coleoptera was found in the undergrowth of co-
niferous forest [32].
In the European part of Russia, the species richness of Coleoptera falling into traps is
significant. The number of species from this order actively flying to beer bait is extremely
high and usually amounts to more than 90% of the total of Coleoptera specimens [63]. In
these studies, we obtained similar results. As in other studies [37,63], this bait attracts
many Coleoptera species. However, there are species that can accidentally be caught by
the trap, which was demonstrated previously [63]. However, the number of such “acci-
dental” species is small and usually does not exceed 10%.
Our data showed that the abundance and species richness of Coleoptera in pine
forests are higher if the trap is set at heights of 1.5 m and 3 m. At the same time, the
Shannon index was higher, and the Simpson index was lower at the ground layer than in
Figure 5.
The diagram of the principal component analysis ordination of the selected beetle species in the
Republic of Mordovia (European Russia) based on the number of specimens collected at various heights
of trap positions (1.5 m, 3 m, 7 m, 12 m). Designations: P.mar.—Protaetia marmorata; C.ser.—Ctesias
serra; C.str.—Cryptarcha strigata; C.und.—Cryptarcha undata; G.gra.—Glischrochilus grandis; G.hort.—
Glischrochilus hortensis; G.quad.—Glischrochilus quadripunctatus; S.gris.—Soronia grisea; L.thor.—Leptura
thoracica; Rh.mor.—Rhagium mordax; A.dis.—Anisandrus dispar. Groups of heights of trap positions were
plotted onto the ordination diagram as supplementary environmental variables.
4. Discussion
This paper provides an idea of clusters of beetles captured using beer traps installed
at various heights in pine forests of the temperate zone of European Russia. Previously,
it was widely believed that temperate forests are less stratified than tropical forests be-
cause of the smaller structural differences and seasonal changes in climatic conditions that
force many species to migrate between layers [
60
]. However, subsequent studies, as well
as our research, have shown that beetle communities in the canopy differ from commu-
nities in the undergrowth. Longhorn and scarab beetles were studied by Touroult and
Dalens [
61
] using bait traps placed at three various heights: 3–5 m (undergrowth),
10–15 m
and
25–30 m
(under canopy). It was found that, for longhorn beetles, the abundance is
higher in the undergrowth, while the richness is much higher in the canopy. For scarab
beetles, abundance and richness are the same in all three layers. In temperate deciduous
forests in Japan, beetle abundance was higher in the canopy relative to the understory [
62
].
In Ontario (Canada), a higher abundance of Coleoptera was found in the undergrowth of
coniferous forest [32].
In the European part of Russia, the species richness of Coleoptera falling into traps is
significant. The number of species from this order actively flying to beer bait is extremely
high and usually amounts to more than 90% of the total of Coleoptera specimens [
63
].
In these studies, we obtained similar results. As in other studies [
37
,
63
], this bait attracts
many Coleoptera species. However, there are species that can accidentally be caught by the
trap, which was demonstrated previously [
63
]. However, the number of such “accidental”
species is small and usually does not exceed 10%.
Our data showed that the abundance and species richness of Coleoptera in pine forests
are higher if the trap is set at heights of 1.5 m and 3 m. At the same time, the Shannon
Diversity 2022,14, 622 8 of 21
index was higher, and the Simpson index was lower at the ground layer than in tree crowns
(at heights of 7 m and 12 m). The species richness at the height of 1.5 m was considerable.
Similar results were obtained in the studies of Ulyshen and Hanula [
64
]. Therefore, in the
crowns of trees of temperate pine forests, there is little species richness and the dominance
of one or more species.
The found high richness is also consistent with the life habits of some Coleoptera
families, for example, with their nutrition and reproduction, as well as with the interactions
in which they participate, such as predation and competition [
60
]. There are some families
and species that constantly fall into traps. For example, the families Elateridae, Ceram-
bycidae, Staphylinidae and Curculionidae, by number, prevailed at the height of 1.5 and
3 m, i.e., closer to the soil and at the level of the undergrowth. In other studies of vertical
distribution, they were also recorded in the same layer of tropical forest [
65
] and temperate
deciduous forests [
64
,
66
]. It is likely that for some families this is due to the fact that the
humidity in the undergrowth may be higher than in the canopy. For example, Staphylinidae
are predominantly found in habitats with higher humidity; in addition, the understory also
provides this family with resources for nutrition [
67
,
68
]. The species richness of Elateridae
was almost the same at heights of 1.5, 3 and 7 m. These species are mainly saproxylic and
often prefer a grassy layer and the level of undergrowth in forests [
53
,
69
]. Some species of
Scolytinae (Curculionidae) were associated with heights of 7 m to 21 m, while other species
are associated with the height of 1.2 m [70].
Representatives of such families as Chrysomelidae, Anthribidae, Oedemeridae, Mordelli-
dae, Melandryidae, and Endomychidae were found only in the undergrowth. The species
of these families are mainly phytophages (including anthophiles) and mycetophages [
71
–
73
].
Thus, herbaceous plants and woody fungi growing in the pine forest provide food resources
for these species. For families such as Histeridae and Silphidae, the undergrowth also rep-
resents more favorable living conditions. Representatives of these families are necrophages,
coprophages and sometimes nidicoles [73,74].
On the other hand, the species of the Cleridae family were distributed evenly over
the entire vertical gradient. They are associated with woody plants and are predators of
various species of xylophagous insects (Bostrichidae, Ptinidae, Scolitinae, Buprestidae and
Cerambycidae); therefore, they act as important regulators of their populations [
75
,
76
]. The
number and species richness of Dermestidae were almost the same at heights of 1.5, 3 and
7 m, while in the crowns of trees, these indicators decreased. The larvae of these beetles are
necrophages, and imagoes are often found on flowering plants [73,77].
During the experiment, 11 species were represented by more than 100 specimens
(P. marmorata,C. serra,C. strigata,C. undata,G. grandis,G. hortensis,G. quadripunctatus,
S. grisea,L. thoracica,Rh. mordax and A. dispar). The average occurrence frequency of
these species ranged from 11 to 100%. These species have a wide range, and they are
distributed in the most diverse types of forests, forest parks and squares [
78
–
82
]. For
C. strigata, it is shown that the abundance of the species is maximal at heights of 1.5 and
3 m with the greatest occurrence frequency at the same heights. Imagoes of this species
live near the flowing fermenting juice of Q. robur, where its larvae develop, and are rarely
caught on the leaking juice of P. tremula. It is in such habitats that these species thrive the
most [
83
]. G. grandis is common on the decaying tree sap of birches and oaks, where the
larva develops. Imago was also found on polypores and rotting berries [
84
,
85
]. Previously,
a high abundance of this species was recorded in pine forests with a well-developed
undergrowth of deciduous trees [
83
]. G. hortensis prefers lower layers of the forest (1.5 m).
P. marmorata was clearly confined to the highest layer of the forest.
Rh. mordax was captured in the highest number at the height of 1.5 m. Rh. mordax
larvae develop under the bark of dead coniferous and deciduous trees, and imagoes also
prefer small heights for habitat in the forest [
66
,
86
]. Scolytinae beetles prefer to inhabit
lower parts of the canopy layer, so most of them are trapped at the height of 2 m and
lower [
87
,
88
]. In our studies, among the Scolytinae, A. dispar had the largest number in
traps; its individuals clearly preferred the level of undergrowth, including in deciduous
Diversity 2022,14, 622 9 of 21
forests [
66
]. The saproxylic species L. thoracica did not have a clear preference in the pine
forest. However, it was previously noted that mass collections of this species occur in
places with a predominance in the birch stand and in the crowns of trees of various forest
types [
89
,
90
]. Larvae of this species are polyphages of deciduous trees and live in dead,
rotten wood of thick trunks (e.g., Populus,Betula,Tilia,Salix and Fagus) [91,92].
In our study, the relative number of saproxylic beetle species was lower at the height of
1.5 m, whereas at other heights it increased. In temperate forests, saproxylic beetles include
30% of all Coleoptera taxa [
93
]. Moreover, the biodiversity of such species is higher in forests
with an increase in dead wood, stumps, dead trees and coarse wood debris [
53
,
55
,
94
,
95
].
For example, Vodka and Cizek [
78
] found that the saproxylic species richness was higher in
the undergrowth than in the canopy at the edge of the forest, while the opposite situation
was observed in depths of the forest. Preisser et al. [
96
] revealed that most of the studied
families are more numerous in traps at the ground level. Previous studies [
97
] showed that
the number of saproxyl beetle species was higher in well-illuminated forests and correlated
positively with temperature variability. Therefore, exposure to the sun (high temperature)
promotes the saproxyl beetle diversity due to the higher microclimatic heterogeneity within
and between dead wood logs. Of considerable importance for saproxylic species in the
ground layer of the forest are the composition of tree species, dead wood, fallen trees and
remains of stumps. Under specific conditions, for example, the increased temperature on
the forest ground layer, this can lead to an increase in the insect species richness of this
forest layer [
98
–
101
]. This is confirmed in studies by Weiss et al. [
12
], who proved that in
temperate forests, an increase in the proportion of saprophages near the ground layer is
caused by the accumulation of dead organic matter on the forest floor.
Our research confirmed earlier studies in deciduous forests [
66
]. Differences in the
vertical distribution of saproxylic species may be due to differences in bait sensitivity,
behavior, dispersal ability, microclimate, ecological interactions or spatial heterogeneity
of the nutrition quality and quantity [
94
,
102
]. For example, the difference in the vertical
structure of the longhorn species community correlated with a high number of flowering
plants, a high leaf area index, as well as with a high level of light intensity and steep relief
in pine forests [
32
]. At the same time, a higher density of snags and the amount of felled
wood are often recorded in such areas, which are important variables for the development
of Cerambycidae larvae [
103
]. Since most adult Cerambycidae are good fliers and spend a
lot of time feeding on flowers away from sites, which they inhabit, it is possible that adults
were more active in searching for flowering plants (almost all Cerambycidae we found were
anthophiles). In our study, in absolute terms, the number of saproxylic species at heights of
1.5 and 3 m exceeded the analogous indicator at other heights. A good herbaceous cover of
perennial grasses and a shrubby tier are a good food base for anthophilic species.
Earlier, using the example of five different forest areas with a predominance of indi-
vidual species of trees, it was shown that the highest species richness is observed in pine
forests, and the lowest one in oak forests [
83
]. When comparing the results obtained with
other studies in deciduous forests [
66
], this pattern persists. As a result, 92 Coleoptera
species belonging to 26 families were identified [
66
]. One hundred twenty-five species
from 36 families were obtained in pine forests. It is possible that a more significant biodi-
versity of Coleoptera in pine forests is associated with the microclimate of temperate pine
forests, characteristics of undergrowth and grassy layer, humidity, sunlight penetration
and other conditions. For example, in the Mediterranean pine forests in summer, the light
transmission of tree crowns is expressed to a greater extent. In addition, the soil moisture
in the undergrowth was lower than outside the areas where shrubs grow. Despite the
soil moisture decrease, in pine forests and in treeless areas, shrubs serve as safe places for
plant development due to the buffering of the microclimate in the summer period [
104
].
In Western Hungary, the middle part and the shrub layer of the pine forest played a key role
in maintaining a specific microclimate in forests having a close canopy. Such conditions are
favorable for sensitive taxa of forest specialists (for example, some forest grasses, epiphytic
mosses and lichens) [
105
]. At the same time, deciduous forests with high crown closeness
Diversity 2022,14, 622 10 of 21
contribute to the reduction in undergrowth and poor herbaceous cover, which ultimately
reduces the biodiversity of Coleoptera [66].
5. Conclusions
The research of beetles at various heights in temperate pine forests of the European
Russia allowed us to find 125 species from 36 families. The highest species richness
was revealed for Nitidulidae and Cerambycidae (19 species each), Elateridae (13 species),
Curculionidae (7 species) and Scarabaeidae (6 species). Eleven species had the maximal
abundance of individuals in traps, while twenty-four beetle species were common in all
studied heights. According to the absolute and average number of specimens, the number
of families and species, superiority was observed at heights of 1.5 m and 3 m. The minimal
number of specimens and the smallest species richness of beetles were found at the height
of 12 m. The highest differences in the Jaccard similarity index were obtained between
minimal, 1.5 m, and maximal, 12 m, heights. The values of the Shannon index were higher
at the ground layer compared to the tree crown, while values of the Simpson index were
characterized by the opposite trend. C. strigata and G. grandis had clear height preferences
(3 and 1.5 m). The specimen number of these species decreased with increasing height
of the trap position. C. undata and P. marmorata were more common at a height of 7 m
with a frequency of 61.4–79.6% and 68.2–79.6%, respectively. The absolute number of
saproxylic and anthophilic species was higher at the ground and undergrowth levels of
the forest. This study showed that there are still little data on the vertical distribution of
arthropod species inhabiting various forests of the temperate zone. To better understand
such regularities, we propose using a set of various methods of data collection, expand the
taxonomic composition of the studied arthropod groups, and increase the sampling efforts,
including the number of studied sites per research.
Author Contributions:
Conceptualization, A.B.R. and L.V.E.; methodology, A.B.R. and A.K; software,
A.A.K.; validation, A.B.R. and L.V.E.; formal analysis, A.B.R. and L.V.E.; investigation, A.B.R.; resources,
A.B.R.; data curation, L.V.E.; writing—original draft preparation, A.B.R. and A.K; writing—review
and editing, L.V.E.; visualization, L.V.E.; supervision, A.B.R.; project administration, A.B.R.; funding
acquisition, A.B.R. All authors have read and agreed to the published version of the manuscript.
Funding: This research was funded by Russian Science Foundation, grant number 22-14-00026.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: Not applicable.
Conflicts of Interest: Not applicable.
Diversity 2022,14, 622 11 of 21
Appendix A
Table A1.
The mean number (M), total number of species (T) and the occurrence frequency of beetle specimens (O, %) collected using beer traps at various heights in
temperate forests, Republic of Mordovia, European Russia.
Taxon Name 1.5 m 3 m 7 m 12 m
T M O T M O T M O T M O
Carabidae
Calathus micropterus (Duftschmid, 1812) 1 0.02 2.27
Harpalus signaticornis (Duftschmid, 1812) 1 0.02 2.27
Histeridae
Gnathoncus buyssoni Auzat, 1917 22 0.50 36.36 16 0.36 29.55 16 0.36 18.18 8 0.18 11.36
Margarinotus striola (C.R. Sahlberg, 1819) 24 0.54 22.73 1 0.02 2.27
Platysoma deplanatum (Gyllenhal, 1808) 2 0.05 4.55 2 0.05 4.55
Platysoma elongatum (Thunberg, 1787) 17 0.39 27.27 1 0.02 2.27 1 0.02 2.27
Platysoma lineare Erichson, 1834 20 0.45 27.27 2 0.05 4.55
Silphidae
Necrodes littoralis (Linnaeus, 1758) 7 0.16 13.64 6 0.14 9.09 4 0.09 9.09
Nicrophorus vespilloides Herbst, 1783 3 0.07 2.27
Oiceoptoma thoracicum (Linnaeus, 1758) 12 0.27 18.18 1 0.02 2.27 1 0.02 2.27
Staphylinidae
Staphylinidae sp. 775 17.61 95.45 291 6.61 79.55 151 3.43 75.00 22 0.50 29.55
Euplectus sp. 1 0.02 2.27
Quedius dilatatus (Fabricius, 1787) 5 0.11 6.82 22 0.50 15.91 15 0.34 13.63 2 0.05 4.55
Scarabaeidae
Cetonia aurata (Linnaeus, 1758) 3 0.07 6.82 4 0.09 9.09 1 0.02 2.27 1 0.02 2.27
Gnorimus variabilis (Linnaeus, 1758) 2 0.05 2.27 3 0.07 4.55 1 0.02 2.27 1 0.02 2.27
Oxythyrea funesta (Poda von Neuhaus, 1761) 1 0.02 2.27
Protaetia fieberi (Kraatz, 1880) 4 0.09 9.09 4 0.09 9.09 6 0.14 9.09
Diversity 2022,14, 622 12 of 21
Table A1. Cont.
Taxon Name 1.5 m 3 m 7 m 12 m
T M O T M O T M O T M O
Protaetia marmorata (Fabricus, 1792) 100 2.27 68.18 219 4.98 68.18 307 6.98 79.55 139 3.16 72.73
Potosia cuprea volhyniensis (Gory & Percheron, 1833) 14 0.32 22.73 12 0.27 25.00 5 0.11 11.36
Scirtidae
Contacyphon sp. 1 0.02 2.27
Contacyphon padi (Linnaeus, 1758) 1 0.02 2.27
Contacyphon pubescens (Fabricius, 1792) 1 0.02 2.27
Throscidae
Trixagus sp. 1 0.02 2.27 1 0.02 2.27
Elateridae
Agrypnus murinus (Linnaeus, 1758) 2 0.05 4.55
Ampedus balteatus (Linnaeus, 1758) 5 0.11 9.09 1 0.02 2.27 1 0.02 2.27
Ampedus cinnabarinus (Eschscholtz, 1829) 1 0.02 2.27
Ampedus nigroflavus (Goeze, 1777) 2 0.05 4.55 2 0.05 4.55
Ampedus pomorum (Herbst, 1784) 2 0.05 4.55 3 0.07 6.82
Ampedus praeustus (Fabricius, 1792) 1 0.02 2.27
Athous subfuscus (O.F. Müller, 1764) 1 0.02 2.27 1 0.02 2.27
Dalopius marginatus (Linnaeus, 1758) 1 0.02 2.27 4 0.09 4.55 3 0.07 4.55 1 0.02 2.27
Denticollis borealis (Paykull, 1800) 1 0.02 2.27
Elater ferrugineus Linnaeus, 1758 1 0.02 2.27 1 0.02 2.27
Melanotus castanipes (Paykull, 1800) 3 0.07 6.82 1 0.02 2.27
Prosternon tesselatum (Linnaeus, 1758) 23 0.52 15.91 5 0.11 9.09 2 0.05 4.55
Selatosomus aeneus (Linnaeus, 1758) 1 0.02 2.27 1 0.02 2.27 1 0.02 2.27
Buprestidae
Dircaea quadriguttata (Paykull, 1798) 1 0.02 2.27
Diversity 2022,14, 622 13 of 21
Table A1. Cont.
Taxon Name 1.5 m 3 m 7 m 12 m
T M O T M O T M O T M O
Cantharidae
Cantharis nigricans O.F. Müller, 1776 2 0.05 4.55 1 0.02 2.27
Cantharis pellucida Fabricius, 1792 1 0.02 2.27
Rhagonycha nigripes (W. Redtenbacher, 1842) 2 0.05 4.55
Dermestidae
Attagenus schaefferi (Herbst, 1792) 8 0.18 9.09 1 0.02 2.27 6 0.14 9.09 1 0.02 2.27
Ctesias serra (Fabricius, 1792) 24 0.54 15.91 73 1.66 22.73 81 1.84 20.45 37 0.84 22.73
Dermestes lardarius Linnaeus, 1758 1 0.02 2.27
Megatoma undata (Linnaeus, 1758) 4 0.09 9.09 3 0.07 6.82
Trogoderma glabrum (Herbst, 1783) 12 0.27 15.91 14 0.32 20.45 19 0.43 18.18 1 0.02 2.27
Cleridae
Thanasimus femoralis (Zetterstedt, 1828) 1 0.02 2.27 2 0.05 4.55 1 0.02 2.27 1 0.02 2.27
Thanasimus formicarius (Linnaeus, 1758) 4 0.09 9.09
Melyridae
Aplocnemus nigricornis (Fabricius, 1792) 2 0.05 4.55 1 0.02 2.27
Dasytes fusculus (Illiger, 1801) 1 0.02 2.27
Dasytes niger (Linnaeus, 1761) 1 0.02 2.27
Malachius bipustulatus (Linnaeus, 1758) 2 0.05
Erotylidae
Dacne bipustulata (Thunberg, 1781) 1 0.02 2.27
Triplax russica (Linnaeus, 1758) 1 0.02 2.27 1 0.02 2.27
Monotomidae
Rhizophagus bipustulatus (Fabricius, 1792) 1 0.02 2.27
Rhizophagus cribratus (Gyllenhal, 1827) 1 0.02 2.27
Rhizophagus fenestralis (Linnaeus, 1758) 8 0.18 9.09 9 0.20 13.63 6 0.14 4.55
Diversity 2022,14, 622 14 of 21
Table A1. Cont.
Taxon Name 1.5 m 3 m 7 m 12 m
T M O T M O T M O T M O
Nitidulidae
Carpophilus hemipterus (Linnaeus, 1758) 1 0.02 2.27 1 0.02 2.27
Carpophilus marginellus Motschulsky, 1858 1 0.02 2.27 1 0.02 2.27
Cryptarcha strigata (Fabricius, 1787) 1476 33.55 100 2074 47.14 97.72 1608 24.27 95.45 651 14.80 90.91
Cryptarcha undata (G.-A. Olivier, 1790) 75 1.70 61.36 162 3.68 77.27 123 2.80 79.55 63 1.43 68.18
Cychramus luteus (Fabricius, 1787) 22 0.50 18.18 9 0.20 11.36
Cychramus variegatus (Herbst, 1792) 2 0.05 4.55 1 0.02 2.27
Epuraea sp. 161 3.66 79.55 90 2.05 63.64 26 0.59 27.27 8 0.18 13.63
Epuraea guttata (G.-A. Olivier, 1811) 4 0.09 9.09 3 0.07 6.82 1 0.02 2.27
Glischrochilus grandis (Tournier, 1872) 1136 25.82 68.18 1345 30.57 54.55 489 11.11 43.18 58 1.32 38.64
Glischrochilus hortensis (Geoffroy, 1785) 277 6.30 56.82 173 3.93 56.82 62 1.41 27.27 7 0.16 13.63
Glischrochilus quadriguttatus (Fabricius, 1777) 48 1.09 6.82 3 0.07 6.82 1 0.02 2.27
Glischrochilus quadripunctatus (Linnaeus, 1758) 95 2.16 40.91 41 0.93 27.27 15 0.34 13.63 19 0.43 6.82
Glischrochilus quadrisignatus (Say, 1835) 2 0.05 4.55 2 0.05 4.55 1 0.02 2.27
Ipidia binotata Reitter, 1875 1 0.02 2.27 1 0.02 2.27
Nitidula bipunctata (Linnaeus, 1758) 2 0.05 2.27
Omosita depressa (Linnaeus, 1758) 3 0.07 6.82
Omosita discoidea (Fabricius, 1775) 2 0.05 4.55
Pityophagus ferrugineus (Linnaeus, 1761) 1 0.02 2.27
Soronia grisea (Linnaeus, 1758) 85 1.93 50.00 131 2.98 54.55 60 1.36 40.91 22 0.50 31.82
Soronia punctatissima (Illiger, 1794) 6 0.14 9.09 1 0.02 2.27
Silvanidae
Silvanus bidentatus (Fabricius, 1792) 2 0.05 4.55
Cucujidae
Pediacus depressus (Herbst, 1797) 31 0.70 27.27 9 0.20 20.45 1 0.02 2.27
Diversity 2022,14, 622 15 of 21
Table A1. Cont.
Taxon Name 1.5 m 3 m 7 m 12 m
T M O T M O T M O T M O
Laemophloeidae
Cryptolestes sp. 1 0.02 2.27
Endomychidae
Mycetina cruciata (Schaller, 1783) 1 0.02 2.27
Coccinellidae
Anatis ocellata (Linnaeus, 1758) 3 0.07 4.55 4 0.09 6.82
Calvia quatuordecimguttata (Linnaeus, 1758) 1 0.02 2.27 1 0.02 2.27 1 0.02 2.27
Halyzia sedecimguttata (Linnaeus, 1758) 1 0.02 2.27 7 0.16 15.91 5 0.11 11.36 1 0.02 2.27
Harmonia quadripunctata (Pontoppidan, 1763) 1 0.02 2.27
Mycetophagidae
Litargus connexus (Geoffroy, 1785) 5 0.11 9.09 6 0.14 11.36 1 0.02 2.27 1 0.02 2.27
Mycetophagus piceus (Fabricius, 1777) 1 0.02 2.27
Mycetophagus quadripustulatus (Linnaeus, 1761) 1 0.02 2.27
Ciidae
Orthocis alni (Gyllenhal, 1813) 1 0.02 2.27
Melandryidae
Dircaea quadriguttata (Paykull, 1798) 1 0.02 2.27
Orchesia fasciata (Illiger, 1798) 4 0.09 6.82
Mordellidae
Mordellistena humeralis (Linnaeus, 1758) 1 0.02 2.27
Tomoxia bucephala A. Costa, 1854 1 0.02 2.27
Tenebrionidae
Bolitophagus reticulatus (Linnaeus, 1767) 1 0.02 2.27
Pseudocistela ceramboides (Linnaeus, 1758) 1 0.02 2.27
Diversity 2022,14, 622 16 of 21
Table A1. Cont.
Taxon Name 1.5 m 3 m 7 m 12 m
T M O T M O T M O T M O
Oedemeridae
Chrysanthia geniculata W.L.E. Schmidt, 1846 1 0.02 2.27
Chrysanthia viridissima (Linnaeus, 1758) 5 0.11 6.82
Oedemera lurida (Marsham, 1802) 1 0.02 2.27
Pythidae
Pytho depressus (Linnaeus, 1767) 1 0.02 2.27
Pyrochroidae
Schizotus pectinicornis (Linnaeus, 1758) 1 0.02 2.27
Anthicidae
Notoxus monoceros (Linnaeus, 1761) 1 0.02 2.27
Scraptiidae
Anaspis sp. 3 0.07 6.82
Cerambycidae
Anastrangalia reyi (Heyden, 1889) 1 0.02 2.27
Aromia moschata (Linnaeus, 1758) 2 0.05 4.55 1 0.02 2.27 1 0.02 2.27
Cortodera femorata (Fabricius, 1787) 1 0.02 2.27
Dinoptera collaris (Linnaeus, 1758) 1 0.02 2.27 1 0.02 2.27
Leptura quadrifasciata Linnaeus, 1758 34 0.77 31.82 17 0.39 20.45 9 0.20 6.82 1 0.02 2.27
Leptura thoracica Creutzer, 1799 111 2.52 38.64 128 2.91 40.91 116 2.64 34.09 77 1.75 36.36
Lepturalia nigripes (De Geer, 1775) 5 0.11 9.09 2 0.05 4.55 2 0.05 4.55 1 0.02 2.27
Molorchus minor (Linnaeus, 1758) 1 0.02 2.27 1 0.02 2.27
Necydalis major Linnaeus, 1758 5 0.11 6.82 3 0.07 4.55 4 0.09 9.09
Obrium cantharinum (Linnaeus, 1767) 1 0.02 2.27
Pachyta quadrimaculata (Linnaeus, 1758) 4 0.09 9.09 8 0.18 13.63 4 0.09 6.82 1 0.02 2.27
Phymatodes testaceus (Linnaeus, 1758) 1 0.02 2.27
Diversity 2022,14, 622 17 of 21
Table A1. Cont.
Taxon Name 1.5 m 3 m 7 m 12 m
T M O T M O T M O T M O
Plagionotus detritus (Linnaeus, 1758) 1 0.02 2.27
Rhagium inquisitor (Linnaeus, 1758) 10 0.23 9.09 6 0.14 9.09 1 0.02 2.27
Rhagium mordax (De Geer, 1775) 389 8.84 52.27 169 3.84 40.91 46 1.05 34.09 11 0.25 15.91
Rutpela maculata (Poda von Neuhaus, 1761) 1 0.02 2.27
Stenocorus meridianus (Linnaeus, 1758) 1 0.02 2.27
Stenurella melanura (Linnaeus, 1758) 1 0.02 2.27
Stictoleptura maculicornis (De Geer, 1775) 1 0.02 2.27 1 0.02 2.27 1 0.02 2.27
Chrysomelidae
Cryptocephalus flavipes Fabricius, 1781 1 0.02 2.27
Phyllotreta atra (Fabricius, 1775) 1 0.02 2.27
Anthribidae
Gonotropis dorsalis (Gyllenhal, 1813) 1 0.02 2.27
Tropideres albirostris (Schaller, 1783) 2 0.05 4.55
Nemonychidae
Cimberis attelaboides (Fabricius, 1787) 1 0.02 2.27 2 0.05 4.55 6 0.14 11.36
Curculionidae
Anisandrus dispar (Fabricius, 1792) 149 3.39 20.45 152 3.45 25.00 104 2.36 15.91 58 0.32 11.36
Anthonomus phyllocola (Herbst, 1795) 1 0.02 2.27
Hylastes brunneus (Erichson, 1836) 1 0.02 2.27
Phyllobius argentatus (Linnaeus, 1758) 1 0.02 2.27
Polydrusus cervinus (Linnaeus, 1758) 1 0.02 2.27
Tomicus minor (Hartig, 1834) 1 0.02 2.27
Xyleborinus saxesenii (Ratzeburg, 1837) 1 0.02 2.27
Total of individuals 5281 5273 3335 1228
Diversity 2022,14, 622 18 of 21
References
1.
Brygadyrenko, V.V. Effect of canopy density on litter invertebrate community structure in pine forests. Ekológia
2016
,35, 90–102.
[CrossRef]
2.
Connor, S.E.; Araújo, J.; Boski, T.; Gomes, A.; Gomes, S.D.; Leira, M.; Freitas, M.C.; Andrade, C.; Morales-Molino, C.;
Franco-Múgica, F.; et al. Drought, fire and grazing precursors to large-scale pine forest decline. Divers. Distrib.
2021
,27,
1138–1151. [CrossRef]
3.
Popkova, T.V.; Zryanin, V.A.; Ruchin, A.B. The ant fauna (Hymenoptera: Formicidae) of the Mordovia State Nature Reserve,
Russia. Nat. Conserv. Res. 2021,6, 45–57. [CrossRef]
4.
Ivanova, N.V.; Shashkov, M.P.; Shanin, V.N. Study of pine forest stand structure in the Priosko-Terrasny State Nature Biosphere
Reserve (Russia) based on aerial photography by quad-rocopter. Nat. Conserv. Res. 2021,6, 1–14. [CrossRef]
5. Ganopoulos, I.; Aravanopoulos, F.; Madesis, P.; Pasentsis, K.; Bosmali, I.; Ouzounis, C.; Tsaftaris, A. Taxonomic identification of
Mediterranean pines and their hybrids based on the high resolution melting (HRM) and trnL approaches: From cytoplasmic
inheritance to timber tracing. PLoS ONE 2013,8, e60945. [CrossRef] [PubMed]
6.
Houston Durrant, T.; de Rigo, D.; Caudullo, G. Pinus sylvestrisin Europe: Distribution, habitat, usage and threats. In European
Atlas of Forest Tree Species; San-Miguel-Ayanz, J., de Rigo, D., Caudullo, G., Houston Durrant, T., Mauri, A., Eds.; Publications
Office of the European Union: Luxembourg, 2016; pp. 132–133.
7.
Horák, J.; Materna, J.; Halda, J.P.; Mladenovi´c, S.; Bogusch, P.; Pech, P. Biodiversity in remnants of naturalmountain forests under
conservation-oriented management. Sci. Rep. 2019,9, 89. [CrossRef] [PubMed]
8.
Lust, N.; Muys, B.; Nachtergale, L. Increase of biodiversity in homogeneous Scots pine stands by anecologically diversified
management. Biodivers. Conserv. 1998,7, 249–260. [CrossRef]
9.
Urbanavichus, G.P.; Urbanavichene, I.N.; Vondrák, J.; Ismailov, A.B. Epiphytic lichen biota of Prielbrusie National Park (Northern
Caucasus, Russia). Nat. Conserv. Res. 2021,6, 77–94. [CrossRef]
10.
Filimonova, L.V. Vegetation dynamics in the Kostomuksha State Nature Reserve (Russia) and surroundings against changes in
the natural environment during the Holocene. Nat. Conserv. Res. 2021,6(Suppl. S1), 98–115. [CrossRef]
11.
Ba¸skent, E.Z. Assessment and valuation of key ecosystem services provided by two forest ecosystems in Turkey. J. Environ.
Manag. 2021,285, 112135. [CrossRef]
12.
Weiss, M.; Didham, R.K.; Procházka, J.; Schlaghamerský, J.; Basset, Y.; Odegaard, F.; Tichechkin, A.; Schmidl, J.; Floren, A.;
Curletti, G.; et al. Saproxylic beetles in tropical and temperate forests—A standardized comparison of vertical stratification
patterns. For. Ecol. Manag. 2019,444, 50–58. [CrossRef]
13.
Teshome, M.; Asfaw, Z.; Mohammed, M. Pattern of functional diversity along the elevation gradient in the dry evergreen
Afromontane forest of Hararghe Highland, Southeast Ethiopia. Biosyst. Divers. 2020,28, 257–264. [CrossRef]
14.
Sabattini, J.A.; Sabattini, R.A.; Cian, J.C.; Sabattini, I.A. Carbon stock in subtropical native forests in a South American Protected
Area. Nat. Conserv. Res. 2021,6, 66–79. [CrossRef]
15.
Ivanov, D.G.; Kotlov, I.P.; Minayeva, T.Y.; Kurbatova, J.A. Estimation of carbon dioxide fluxes on a ridge-hollow bog complex
using a high resolution orthophotoplan. Nat. Conserv. Res. 2021,6, 16–28. [CrossRef]
16.
Kim, J. Subdivision design and landscape structure: Case study of The Woodlands, Texas, US. Urban For. Urban Green.
2019
,38,
232–241. [CrossRef]
17.
Kharitonova, A.O.; Kharitonova, T.I. The effect of landscape pattern on the 2010 wildfire spread in the Mordovia State Nature
Reserve, Russia. Nat. Conserv. Res. 2021,6, 29–41. [CrossRef]
18.
Kirstová, M.; Pyszko, P.; Šipoš, J.; Drozd, P.; Koˇcárek, P. Vertical distribution of earwigs (Dermaptera: Forficulidae) in a temperate
lowland forest, based on sampling with a mobile aerial lift platform. Entomol. Sci. 2017,20, 57–64. [CrossRef]
19.
Giovanni, F.; Mei, M.; Cerretti, P. Vertical stratification of selected Hymenoptera in a remnant forest of the Po Plain (Italy,
Lombardy) (Hymenoptera: Ampulicidae, Crabronidae, Sphecidae). Fragm. Entomol. 2017,49, 71–77. [CrossRef]
20.
Dvoˇrák, L.; Dvoˇráková, K.; Obo ˇna, J.; Ruchin, A.B. Selected Diptera families caught with beer traps in the Republic of Mordovia
(Russia). Nat. Conserv. Res. 2020,5, 65–77. [CrossRef]
21.
Ruchin, A.B. Seasonal dynamics and spatial distribution of lepidopterans in selected locations in Mordovia, Russia. Biodiversitas
2021,22, 2569–2575. [CrossRef]
22.
Urban-Mead, K.R.; Muñiz, P.; Gillung, J.; Espinoza, A.; Fordyce, R.; van Dyke, M.; McArt, S.H.; Danforth, B.N. Bees in the trees:
Diverse spring fauna in temperate forest edge canopies. For. Ecol. Manag. 2021,482, 118903. [CrossRef]
23.
Makarkin, V.N.; Ruchin, A.B. Materials on the Neuroptera and Raphidioptera fauna in Mordovia and adjacent regions of
European Russia. Proc. Mordovia State Nat. Reserve 2020,24, 161–181.
24.
Haack, N.; Borges, P.A.V.; Grimm-Seyfarth, A.; Schlegel, M.; Wirth, C.; Bernhard, D.; Brunk, I.; Henle, K.; Pereira, H.M. Response
of common and rare beetle species to tree species and vertical stratification in a floodplain forest. Insects
2022
,13, 161. [CrossRef]
25.
Davis, A.J.; Sutton, S.L.; Brendell, M.J.D. Vertical distribution of beetles in a tropical rainforest in Sulawesi: The role of the canopy
in contributing to biodiversity. Sepilok Bull. 2011,13, 59–83.
26.
Puker, A.; Ferreira, K.R.; Correa, C.M.A. Sampling flower chafer beetles (coleoptera: Cetoniidae) in the Amazon rainforest: The
role of bait types and trap installation heights get access arrow. Environ. Entomol. 2020,49, 1096–1104. [CrossRef]
Diversity 2022,14, 622 19 of 21
27.
Sweeney, J.; Hughes, C.; Webster, V.; Kostanowicz, C.; Webster, R.; Mayo, P.; Allison, J.D. Impact of horizontal edge–interior and
vertical canopy–understory gradients on the abundance and diversity of bark and woodboring beetles in survey traps. Insects
2020,11, 573. [CrossRef]
28. Foit, J.; ˇ
Cermák, V.; Gaar, V.; Hradil, K.; Nový, V.; Rolincová, P. New insights into the life history of Monochamus galloprovincialis
can enhance surveillance strategies for the pinewood nematode. J. Pest Sci. 2019,92, 1203–1215. [CrossRef]
29.
Francese, J.A.; Oliver, J.B.; Fraser, I.; Lance, D.R.; Youssef, N.; Sawyer, A.J.; Mastro, V.C. Influence of trap placement and design on
capture of the emerald ash borer (Coleoptera: Buprestidae). J. Econ. Entomol. 2008,101, 1831–1837. [CrossRef]
30.
Chen, G.; Zhang, Q.H.; Wang, Y.; Liu, G.T.; Zhou, X.; Niu, J.; Schlyter, F. Catching Ips duplicatus (Sahlberg) (Coleoptera:
Scolytidae) with pheromone-baited traps: Optimal trap type, colour, height and distance to infestation. Pest Manag. Sci.
2010
,66,
213–219. [CrossRef]
31.
Miller, D.R.; Crowe, C.M.; Sweeney, J.D. Trap height affects catches of bark and woodboring beetles (Coleoptera: Curculionidae,
Cerambycidae) in baited multiple-funnel traps in Southeastern United States. J. Econ. Entomol. 2020,113, 273–280. [CrossRef]
32.
Vance, C.C.; Kirby, K.R.; Malcolm, J.R.; Smith, S.M. Community composition of longhorned beetles (Coleoptera: Cerambycidae) in
the canopy and understorey of sugar maple and white pine stands in south-central Ontario. Environ. Entomol.
2003
,32, 1066–1074.
[CrossRef]
33.
Bouget, C.; Brin, A.; Brustel, H. Exploring the “last biotic frontier”: Are temperate forest canopies special for saproxylic beetles?
For. Ecol. Manag. 2011,261, 211–220. [CrossRef]
34.
Leksono, A.S.; Takada, K.; Koji, S.; Nakagoshi, N.; Anggraeni, T.; Nakamura, K. Vertical and seasonal distribution of flying beetles
in a suburban temperate deciduous forest collected by water pan trap. Insect Sci. 2005,12, 199–206. [CrossRef]
35.
Ruchin, A.B.; Khapugin, A.A. Red Data Book Invertebrates in a Protected Area of European Russia. Acta Zool. Acad. Sci. Hung.
2019,65, 349–370. [CrossRef]
36.
Ruchin, A.B.; Makarkin, N.V. Neuroptera and Raphidioptera in the Mordovia State Nature Reserve. Nat. Conserv. Res.
2017
,2,
38–46. [CrossRef]
37.
Ruchin, A.B.; Egorov, L.V.; Khapugin, A.A.; Vikhrev, N.E.; Esin, M.N. The use of simple crown traps for the insects collection. Nat.
Conserv. Res. 2020,5, 87–108. [CrossRef]
38.
Bouchard, P.; Bousquet, Y.; Davies, A.E.; Alonso-Zarazaga, M.A.; Lawrence, J.F.; Lyal, C.H.C.; Newton, A.F.; Ried, C.A.M.; Schmitt,
M.; ´
Slipi´nski, S.A.; et al. Family-group names in Coleoptera (Insecta). ZooKeys 2011,88, 1–972. [CrossRef]
39.
Bouchard, P.; Bousquet, Y. Additions and corrections to “Family-group names in Coleoptera (Insecta)”. ZooKeys
2020
,922, 65–139.
[CrossRef]
40.
Löbl, I.; Smetana, A. (Eds.) Catalogue of Palaearctic Coleoptera. In Curculionoidea I; Apollo Books: Stenstrup, Denmark, 2011;
Volume 7, p. 373.
41.
Löbl, I.; Smetana, A. (Eds.) Catalogue of Palaearctic Coleoptera. In Curculionoidea II; Apollo Books: Stenstrup, Denmark, 2013;
Volume 8, p. 707.
42.
Löbl, I.; Löbl, D. (Eds.) Catalogue of Palaearctic Coleoptera. Revised and Updated Version. Hydrophiloidea–Staphylinoidea; Brill: Leiden,
The Netherlands; Boston, MA, USA, 2015; Volume 2/1, p. 1702.
43.
Löbl, I.; Löbl, D. (Eds.) Catalogue of Palaearctic Coleoptera. Revised and Updated Version. Scarabaeoidea–Scirtoidea–Dascilloidea–
Buprestoidea–Byrrhoidea; Brill: Leiden, The Netherlands; Boston, MA, USA, 2016; Volume 3, p. 983.
44.
Löbl, I.; Löbl, D. (Eds.) Catalogue of Palaearctic Coleoptera. Revised and Updated Version. Archostemata–Adephaga–Myxophaga; Brill:
Leiden, The Netherlands; Boston, MA, USA, 2017; Volume 1, p. 1443.
45.
Iwan, D.; Löbl, I. (Eds.) Catalogue of Palaearctic Coleoptera. Revised and Updated Second Edition. Tenebrionoidea; Brill: Leiden,
The Netherlands; Boston, MA, USA, 2020; Volume 5, p. 945.
46.
Danilevsky, M. (Ed.) Catalogue of Palaearctic Coleoptera. Updated and Revised Second Edition. Chrysomeloidea I (Vesperidae, Disteniidae,
Cerambycidae); Brill: Leiden, The Netherlands; Boston, MA, USA, 2020; Volume 6/1, p. 712.
47.
Robertson, J.; ´
Slipi´nski, A.; Moulton, M.; Shockley, F.W.; Giorgi, A.; Lord, N.P.; McKenna, D.D.; Tomaszewska, W.; Forrester, J.;
Miller, K.B.; et al. Phylogeny and classification of Cucujoidea and the recognition of a new superfamily Coccinelloidea (Coleoptera:
Cucujiformia). Syst. Entomol. 2015,40, 745–778. [CrossRef]
48.
Alonso-Zarazaga, M.A.; Barrios, H.; Borovec, R.; Bouchard, P.; Caldara, R.; Colonnelli, E.; Gültekin, L.; Hlaváˇc, P.; Korotyaev, B.;
Lyal, C.H.C.; et al. Cooperative Catalogue of Palaearctic Coleoptera Curculionoidea. Monogr. Electrón. SEA 2017,8, 1–729.
49.
Löbl, I.; Smetana, A. (Eds.) Catalogue of Palaearctic Coleoptera. Elateroidea–Derodontoidea–Bos-trichoidea–Lymexyloidea–Cleroidea–
Cucujoidea; Apollo Books: Stenstrup, Denmark, 2007; Volume 4, p. 935.
50.
Löbl, I.; Smetana, A. (Eds.) Catalogue of Palaearctic Coleoptera. Chrysomeloidae; Apollo Books: Stenstrup, Denmark, 2010; Volume 6,
p. 924.
51.
Bousquet, Y. Litteratura Coleopterologica (1758–1900): A guide to selected books related to the taxonomy of Coleoptera with
publication dates and notes. ZooKeys 2016,583, 1–776. [CrossRef]
52.
Speight, M.C.D. Saproxylic invertebrates and their conservation. In Nature and Environment Series 42; Council of Europe:
Strasbourg, France, 1989; pp. 1–79.
53.
Lachat, T.; Wermelinger, B.; Gossner, M.M.; Bussler, H.; Isacsson, G.; Müller, J. Saproxylic beetles as indicator species for
dead-wood amount and temperature in European beech forests. Ecol. Indic. 2012,23, 323–331. [CrossRef]
Diversity 2022,14, 622 20 of 21
54.
Carpaneto, G.M.; Baviera, C.; Biscaccianti, A.B.; Brandmayr, P.; Mazzei, A.; Mason, F.; Battistoni, A.; Teofili, C.; Rondinini, C.;
Fattorini, S.; et al. A red list of Italian saproxylic beetles: Taxonomic overview, ecological features and conservation issues
(Coleoptera). Fragm. Entomol. 2015,47, 53–126. [CrossRef]
55.
Papis, M.; Mokrzycki, T. Saproxylic beetles (Coleoptera) of the strictly protected area Bukowa Góra in the Roztocza´nski National
Park. For. Res. Pap. 2015,76, 229–239. [CrossRef]
56. Shannon, C.E. A mathematical theory of communication. Bell Syst. Tech. J. 1948,27, 379–423, 623–659. [CrossRef]
57. Magurran, A.E. Ecological Diversity and Its Measurement; Chapman & Hall: London, UK, 1996; p. 179.
58.
Hammer, Ø.; Harper, D.A.T.; Ryan, P.D. PAST: Paleontological statistics soft-ware package for education and data analysis.
Palaeontol. Electron. 2001,4, 9.
59.
Moore, D.S.; Notz, W.I.; Flinger, M.A. The Basic Practice of Statistics, 6th ed.; W. H. Freeman and Company: New York, NY, USA,
2013; p. 745.
60.
Ulyshen, M.D. Arthropod vertical stratification in temperate deciduous forests: Implications for conservation-oriented manage-
ment. For. Ecol. Manag. 2011,261, 1479–1489. [CrossRef]
61.
Touroult, J.; Dalens, P.H. Beetles vertical stratification in French Guiana’ forests: Study using aerial fruit traps. In Coléoptères de
Guyane; ACOREP-France: Paris, France, 2012; pp. 16–24.
62.
Hirao, T.; Murakami, M.; Kashizaki, A. Importance of the understory stratum to entomofaunal diversity in a temperate deciduous
forest. Ecol. Res. 2009,24, 263–272. [CrossRef]
63.
Ruchin, A.B.; Egorov, L.V.; Khapugin, A.A. Usage of fermental traps for studying the species diversity of Coleoptera. Insects
2021
,
12, 407. [CrossRef]
64.
Ulyshen, M.D.; Hanula, J.L. A comparison of the beetle (Coleoptera) fauna captured at two heights above the ground in a North
American temperate deciduous forest. Am. Midl. Nat. 2007,158, 260–278. [CrossRef]
65.
Kato, M.; Inque, T.; Hamid, A.A.; Nagamitsu, T.; Merdek, M.B.; Nona, A.R.; Itino, T.; Yamane, S.; Yumoto, T. Seasonality and
vertical structure of light-attracted insect communities in a dipterocarp forest in Sarawak. Res. Popul. Ecol.
1995
,37, 59–79.
[CrossRef]
66.
Ruchin, A.B.; Egorov, L.V. Vertical stratification of beetles in deciduous forest communities in the Centre of European Russia.
Diversity 2021,13, 508. [CrossRef]
67.
Lange, M.; Türke, M.; Pašali´c, E.; Boch, S.; Hessenmöller, D.; Müller, J.; Prati, D.; Socher, S.A.; Fischer, M.; Weisser, W.W.; et al.
Effects of forest management on ground-dwelling beetles (Coleoptera; Carabidae,Staphylinidae) in Central Europe are mainly
mediated by changes in forest structure. For. Ecol. Manag. 2014,329, 166–176. [CrossRef]
68.
Porhajašová, J.I.; Babošová, M.; Noskoviˇc, J.; Ondrišík, P. Long-term developments and biodiversity in carabid and staphylinid
(Coleoptera: Carabidae and Staphylinidae) fauna during the application of organic fertilizers under agroecosystem conditions. Pol.
J. Environ. Stud. 2018,27, 2229–2235. [CrossRef]
69.
Ruchin, A.B.; Egorov, L.V.; Semishin, G.B. Fauna of click beetles (Coleoptera: Elateridae) in the interfluve of Rivers Moksha and
Sura, Republic of Mordovia, Russia. Biodiversitas 2018,19, 1352–1365. [CrossRef]
70.
Procházka, J.; Cizek, L.; Schlaghamerský, J. Vertical stratification of scolytine beetles in temperate forests. Insect Conserv. Divers.
2018,11, 534–544. [CrossRef]
71.
Nikitsky, N.B.; Legalov, A.A. The ant-like stone beetles (Scydmaenidae) and the fungus weevils (Anthribidae) of the Moscow
region. Euroasian Entomol. J. 2016,15, 219–227.
72.
Zemoglyadchuk, A.V.; Ruchin, A.B.; Egorov, L.V. An annotated checklist of the tumbling flower beetles (Coleoptera, Mordellidae)
of the Republic of Mordovia, with a short review of the family in European Russia. Entomol. Rev.
2020
,100, 771–787. [CrossRef]
73.
Ruchin, A.B.; Egorov, L.V.; Polumordvinov, O.A. Coleoptera of the Penza region, Russia based on fermental crown trap.
Biodiversitas 2021,22, 1946–1960. [CrossRef]
74.
Bajerlein, D.; Matuszewski, S.; Konwersk, S. Insect succession on carrion: Seasonality, habitat preference and residency of histerid
beetles (Coleoptera: Histeridae) visiting pig carrion exposed in various Forests (Western Poland). Pol. J. Ecol. 2011,59, 787–797.
75.
Markovi´c, ˇ
C.; Stojanovi´c, A. Fauna of phloemo-xylophagous insects, their parasitoids and predators on Ulmus minor in Serbia.
Biologia 2012,67, 584–589. [CrossRef]
76.
Krivets, S.A.; Kerchev, I.A. Insects inhabiting the galleries of the four-eyed fir bark beetle Polygraphus proximus Blandf. (Coleoptera,
Curculionidae: Scolytinae) in Siberia. Entomol. Rev. 2016,96, 545–558. [CrossRef]
77.
Kadej, M. Larva and pupa of Ctesias (s. str.) serra (Fabricius, 1792) with remarks on biology and economic importance, and larval
comparison of co-occurring genera (Coleoptera, Dermestidae). ZooKeys 2018,758, 115–135. [CrossRef]
78.
Vodka, Š.; Cizek, L. The effects of edge-interior and understorey-canopy gradients on the distribution of saproxylic beetles in a
temperate lowland forest. For. Ecol. Manag. 2013,304, 33–41. [CrossRef]
79.
Ruchin, A.B.; Egorov, L.V. Fauna of longicorn beetles (Coleoptera: Cerambycidae) of Mordovia. Russ. Entomol. J.
2018
,27, 161–177.
[CrossRef]
80.
Egorov, L.V.; Ruchin, A.B.; Semenov, V.B.; Semionenkov, O.I.; Semishin, G.B. Checklist of the Coleoptera of Mordovia State Nature
Reserve, Russia. ZooKeys 2020,962, 13–122. [CrossRef]
81.
Egorov, L.V.; Ruchin, A.B.; Semishin, G.B. Some data on the Coleoptera fauna of the Mordovia State Nature Reserve. Report 10.
Proc. Mordovia State Nat. Reserve 2021,26, 96–128.
82.
Ruzzier, E.; Galli, A.; Bani, L. Monitoring exotic beetles with inexpensive attractants: A case study. Insects
2021
,12, 462. [CrossRef]
Diversity 2022,14, 622 21 of 21
83.
Ruchin, A.B.; Egorov, L.V.; Khapugin, A.A. Seasonal activity of Coleoptera attracted by fermental crown traps in forest ecosystems
of Central Russia. Ecol. Quest. 2021,32, 37–53. [CrossRef]
84.
Alekseev, V.I.; Nikitsky, N.B. Rare and new for the fauna of the Baltic States beetles (Coleoptera) from the Kaliningrad Region.
Acta Zool. Litu. 2008,18, 254–259. [CrossRef]
85.
Laso´n, A.; Holly, M. Glischrochilus grandis Tournier, 1872—New species of beetle for the Polish fauna and new data on the
occurrence of genus Glischrochilus Reitter, 1873 (Coleoptera: Nitidulidae: Cryptarchinae). Acta Entomol. Sil. 2015,23, 1–4.
86.
Gorshkova, V.P.; Volodchenko, A.N. The specific assemblage structure of longhorn beetles (Coleoptera, Cerambycidae) in
floodplain forests of the western part of Saratov oblast. Biol. Bull. 2016,43, 1416–1421. [CrossRef]
87.
Sheehan, T.N.; Ulyshen, M.D.; Horn, S.; Hoebeke, E.R. Vertical and horizontal distribution of bark and woodboring beetles by
feeding guild: Is there an optimal trap location for detection? J. Pest Sci. 2019,92, 327–341. [CrossRef]
88.
Holuša, J.; Fiala, T.; Foit, J. Ambrosia beetles prefer closed canopies: A case study in oak forests in Central Europe. Forests
2021
,
12, 1223. [CrossRef]
89.
Danilevsky, M.L.; Ruchin, A.B.; Egorov, L.V. Mass collection of two rare longicorn-species (Coleoptera, Cerambycidae) in Central
Russia. Humanit. Space 2019,8, 1179–1183.
90.
Ruchin, A.B.; Egorov, L.V.; MacGowan, I.; Makarkin, V.N.; Antropov, A.V.; Gornostaev, N.G.; Khapugin, A.A.; Dvoˇrák, L.; Esin,
M.N. Post-fire insect fauna explored by crown fermental traps in forests of the European Russia. Sci. Rep.
2021
,11, 21334.
[CrossRef]
91.
Sama, G. Atlas of the Cerambycidae of Europe and the Mediterranean Area. Northern, Western, Central and Eastern Europe British Isles and
Continental Europe from France (Excl. Corsica) to Scandinavia and Urals. Kabourek, Zlín; Coronet Books Incorporated: London, UK,
2002; Volume 1, p. 173.
92.
Karpi´nski, L.; Szczepa ´nski, W.T.; Boldgiv, B.; Walczak, M. New data on the longhorn bee-tles of Mongolia with particular
emphasis on the genus Eodorcadion Breuning, 1947 (Coleoptera, Cerambycidae). ZooKeys 2018,739, 107–150. [CrossRef]
93.
Haeler, E.; Bergamini, A.; Blaser, S.; Ginzler, C.; Hindenlang, K.; Keller, C.; Kiebacher, T.; Kormann, U.G.; Scheidegger, C.; Schmidt,
R.; et al. Saproxylic species are linked to the amount and isolation of dead wood across spatial scales in a beech forest. Landsc.
Ecol. 2021,36, 89–104. [CrossRef]
94.
Ekström, A.L.; Bergmark, P.; Hekkala, A.M. Can multifunctional forest landscapes sustain a high diversity of saproxylic beetles?
For. Ecol. Manag. 2021,490, 119107. [CrossRef]
95.
Parisi, F.; Innangi, M.; Tognetti, R.; Lombardi, F.; Chirici, G.; Marchetti, M. Forest stand structure and coarse woody debris
determine the biodiversity of beetle communities in Mediterranean mountain beech forests. Glob. Ecol. Conserv.
2021
,28, e01637.
[CrossRef]
96.
Preisser, E.; Smith, D.C.; Lowman, M.D. Canopy and ground level insect distribution in a temperate forest. Selbyana
1998
,19,
141–146.
97.
Lettenmaier, L.; Seibold, S.; Bässler, C.; Brandl, R.; Gruppe, A.; Müller, J.; Hagge, J. Beetle diversity is higher in sunny forests due
to higher microclimatic heterogeneity in deadwood. Oecologia 2022,198, 825–834. [CrossRef]
98.
Pakeman, R.J.; Stockan, J.A. Drivers of carabid functional diversity: Abiotic environment, plant functional traits, or plant
functional diversity? Ecology 2014,95, 1213–1224. [CrossRef]
99.
Penone, C.; Allan, E.; Soliveres, S.; Felipe-Lucia, M.R.; Gossner, M.M.; Seibold, S.; Simons, N.K.; Schall, P.; Van Der Plas, F.;
Manning, P.; et al. Specialisation and diversity of multiple trophic groups are promoted by different forest features. Ecol. Lett.
2019,22, 170–180. [CrossRef]
100.
Schneider, A.; Blick, T.; Köhler, F.; Pauls, S.U.; Römbke, J.; Zub, P.; Dorow, W.H.O. Animal diversity in beech forests—An analysis
of 30 years of intense faunistic research in Hessian strict forest reserves. For. Ecol. Manag. 2021,499, 119564. [CrossRef]
101.
Tinya, F.; Kovács, B.; Bidló, A.; Dima, B.; Király, I.; Kutszegi, G.; Lakatos, F.; Mag, Z.; Márialigeti, S.; Nascimbene, J.; et al.
Environmental drivers of forest biodiversity in temperate mixed forests—A multi-taxon approach. Sci. Total Environ.
2021
,
795, 148720. [CrossRef]
102.
Maguire, D.Y.; Robert, K.; Brochu, K.; Larrivée, M.; Buddle, C.M.; Wheeler, T.A. Vertical stratification of beetles (Coleoptera) and
flies (Diptera) in temperate forest canopies. Environ. Entomol. 2014,43, 9–17. [CrossRef]
103.
Gosling, D.C.L.; Gosling, N.M. An annotated list of the Cerambycidae of Michigan (Coleoptera) Part II, the subfamilies Lepturinae
and Lamiinae. Great Lakes Entomol. 1977,10, 1–37.
104.
Prévosto, B.; Helluy, M.; Gavinet, J.; Fernandez, C.; Balandier, P. Microclimate in Mediterranean pine forests: What is the influence
of the shrub layer? Agric. For. Meteorol. 2020,282–283, 107856. [CrossRef]
105.
Kovács, B.; Tinya, F.; Ódor, P. Stand structural drivers of microclimate in mature temperate mixed forests. Agric. For. Meteorol.
2017,234–235, 11–21. [CrossRef]
