THE
CHANGE OF ZOOCOENOTIC CHARACTERISTICS OF GROUND-BEETLE COMMUNITIES UNDER THE
INFLUENCE OF GRAZING HORSE ON FLOODPLAIN MEADOWS IN BELARUS
Leonora KHOTKO
«Scientific-practical Center
of the National Academy of Sciences of Belarus for biological resources»,
Akademicheskaya str., 27, Minsk, 20072, Belarus, e-mail: khotko1@yandex.ru
Key
words: ground beetles, dominance, meadows, abundance, dominant species.
Abstract: The investigations on the change
of the degree of ground beetle environmental similarity, of the
degree of the relative habitat timing and species dominance were carried out in
the Minsk district in 2011. During the study period 5198 traps/days were
processed, 6024 specimens of ground-beetles were determined. In addition, it is established that during
the growing season the number of species in grazed plots compared with initial
ones did not change - 34-51 species. Only the composition of species changed. As a result of studying species
composition and zoocoenotic indicators of ground beetle
communities 83 species living on floodplain meadows were revealed. Of the
ecological groups selected on relation to humidity the greatest number of
species on floodplain meadows presented mesophiles, and after grazing I,
obviously, with rainfall gigrophiles were. On the basis of biotopic
distribution we took these species to 8 ecological groups. Field species were
characterized by the most number, sometimes their abundance reached 54 %. The
majority of ground-beetle species (82,3 %) bred in the spring and in the
beginning of summer. Species of multiseasonal phenological group weren't
numerous (5,9 % – 26,5 %), and summer and summer-and-autumn groups were random
(2,6 % - 11,8 %). It was explained by migration of “nearwater” mesophile
species, which made a considerable part in fauna, from a spring high water to
adjacent biotopes.
1. Introduction
Ground-beetles are an important component of soil
fauna and occupy one of the leading positions in all land biogeocenoses, both
on number of species, and by quantity of
individuals in populations. They are a quite large group of coleoptera in a systematic respect and one of the most important groups in practical terms. They
occur everywhere and have a high diversity of species (about 308 described
species in the fauna of Belarus). Ground-beetles are
sensitive enough to anthropogenic pressure, which makes them
rather expressive and convenient material for biocenotic research and selection
of biological indicators of ecosystem state
among them. The complexes of
ground-beetles on meadows of the forest zone are studied
well enough [1-3].
In Belarus studied the complexes of ground-beetles on
meadows [4-7]. There are stud farms for breeding horses in Belarus. It is necessary to graze horses for feeding on meadows, including floodplain ones. Studies on the influence of grazing horses on the
ground-beetle communities in Belarus were not previously
carried out.
The purpose of this work is to determine consequences of the influence of grazing horses on ground-beetle communities.
2. Materials
and methods of the research
The
objects of the research were ground-beetles, as the most numerous on species
and quantity group of Coleoptera, living on floodplain meadows. To avoid the
influence of the seasonal changes in composition and abundance on the results
of the research samples were selected every two weeks during the growing
season. Ground-beetles were collected by the methods adopted in zoological
studies – with the help of soil traps. The degree of environmental similarity
of grazed meadows by the ground-beetle complexes was calculated according to [8]:
√c(a+b)∙100Psc
2ab
,
where c is the
number of common species; a and b - the number of species in comparable
communities; Psc - the amount of the lesser of two indicators of each
species abundance.
The degree of relative biotopical attachment of the
species was determined by ratio of the difference between the percentage of
species in a given biotope and its percentage in all investigated habitats to the sum of these shares [9].
The degree of species dominance
was calculated according to [10].
During the study period 5198 traps/days
were processed, 6024 specimens of ground-beetles were determined. On the grazed
meadow we used the so-called gentle method of grazing horses. It lied in the
facts that in the spring, almost to the end of May horses were not put out on a
meadow. These data were like an original starting point of the ground-beetle community. Then horses were
grazing only after 22 days later, in the summer at the end of May – mid June
(grazing I), when the grass grew back up to 15 cm. The second grazing started
after 40 days, and the third one only after 55 days. In the autumn, starting
from the end of August, horses did not graze.
3. Results and discussion
Since a fresh
grass contains a lot of potassium in the spring (under the snow), it cannot be
used for horses. So they were put out on a meadow just at the end of May. From
late April to late May the meadow served as a starting point for counting
ground-beetles. During this period 35 species of ground-beetles represented 18
genera on the floodplain meadow were found. Among genera Harpalus (6 species)
and Bembidion (5 species) were the most numerous on the number of species. In
addition, it is established that during the growing season the number of
species in grazed plots compared with initial ones did not change - 34-51 species. Only the
composition of species changed. So, at the beginning spring-and-summer rare
species were present. They disappeared during the subsequent period - Elaphrus cupreus, Broscus cephalotes, Bembidion
lampros, Amara spreta, Harpalus anxius, Ophonus nitidulus. All of them were found to be widely distributed
on the territory of Belarus, except Ophonus
nitidulus, which was found only in northern and central parts of Belarus.
Three species (Poecilus cupreus, Bembidion properans and Amara aenea) prevailed here (Table 1). Loricera pilicornis (3,5%) and Harpalus
rufipes (1,6%) were subdominants. The rest 30 species belonged to rare
ones. The causes of rarity and a local occurrence of certain species of
ground-beetles in the involved area have a different basis.
Table 1 The degree of
dominance of the dominant species on the meadows after grazing horses
|
Dominants |
Biotopes |
|||
|
initial area |
grazing I |
grazing II |
grazing III |
|
|
Poecilus cupreus (L.) |
79,0 |
49,0 |
10,7 |
40,6 |
|
Bembidion properans (Steph.) |
5,2 |
14,9 |
5,4 |
18,4 |
|
Amara aenea
(Deg.) |
4,9 |
12,0 |
- |
6,5 |
|
Loricera pilicornis (F.) |
- |
5,7 |
- |
- |
|
Harpalus rufipes (Deg.) |
- |
- |
- |
5,7 |
|
Calathus fuscipes (Gz.) |
- |
- |
- |
4,5 |
Some of them were located
on the boundary of its geographic range, while others were local, others lived
in specific habitats, locally occurring in the territory. Finally, a number of
species is poorly explored and data on their biotopical attachment and geographic range are
very fragmentary.
Seven groups, of which field (44,2 %) and forest (23,5 %)
species dominated, were selected on their biotopical attachment. It was
explained by the proximity of the fields with various cultures and the
forests (Table 2). On phenology of development
three groups were selected, among which
spring-and-summer species dominated, making 82,3 %. In relation to
soil humidity five groups were selected. Among them mesophiles (35,3 %) prevailed.
Table 2 The
abundance of species on biotopical attachment, phenology
of development and relation to soil humidity in biotopes
after grazing horses
|
Groups |
Biotopes |
|||
|
initial area |
grazing I |
grazing II |
grazing III |
|
|
Abundance of species on biotopical attachment (%) |
||||
|
Field |
44,2 |
37,9 |
54,0 |
45,5 |
|
Forest |
23,5 |
24,3 |
26,0 |
27,3 |
|
Meadow-and-field |
14,7 |
10,8 |
8,0 |
6,1 |
|
Meadow |
5,9 |
5,4 |
2,0 |
9,1 |
|
Meadow-and-forest |
5,9 |
5,4 |
4,0 |
3,0 |
|
Everybiont |
2,9 |
2,7 |
2,0 |
3,0 |
|
Swamp |
2,9 |
5,4 |
2,0 |
3,0 |
|
Meadow-and-swamp |
- |
8,1 |
2,0 |
3,0 |
|
Abundance of species on phenology of development (%) |
||||
|
Spring-and-summer |
82,3 |
76,3 |
58,8 |
61,7 |
|
Summer-and-autumn |
11,8 |
7,9 |
5,9 |
5,9 |
|
All-season |
5,9 |
13,2 |
23,5 |
26,5 |
|
Summer |
- |
2,6 |
11,8 |
5,9 |
|
Abundance of species on relation to soil humidity (%) |
||||
|
Mesophiles |
35,3 |
26,3 |
47,1 |
44,1 |
|
Mesogygrophiles |
26,5 |
23,7 |
17,6 |
14,7 |
|
Gygrophiles |
14,7 |
36,8 |
11,8 |
14,7 |
|
Xerophiles |
17,6 |
7,9 |
9,8 |
11,8 |
|
Mesoxerophiles |
5,9 |
5,3 |
13,7 |
14,7 |
After grazing I the quantity of ground-beetle species
did not change (38 species from 17 genera, among which Pterostichus (7 species) and Carabus (5
species) were the most numerous on the
number of species, i.e. there was a change of genera
by the number of species. Among them rare, spring-and-summer
species Pterostichus anthracinus, Pt. minor, Pt. diligens, Pt. aterrimus, Pt. vernalis, from Carabus - C. menetriesi and C. clathratus were found.
17 species, which did not occur on an initial area, appeared.
Of these 9 species (Blethisa multipunctata,
Pterostichus anthracinus, Pt. minor,
Pt. diligens, Pt. aterrimus, Agonum duftschmidi, A. viduum, Amara eurynota,
Badister peltatus) were not found
more on meadows. They were rare or locally common species.
Four species dominated, i.e. conditions improved for species prevailing on the number.
This is Poecilus cupreus, Bembidion properans, Amara aenea and Loricera
pilicornis. By reducing the
number of the first dominant the dominance
degree of the other three dominants
increased. Loricera
pilicornis (a subdominant on an initial area) went into a
dominant after grazing I.
Poecilus versicolor
(2,5 %), Harpalus rufipes (2,2), Pterostichus nigrita (1,9) and Carabus
menetriesi (1,8 %) were subdominants. The
other 30 species were rare.
8 groups were selected on biotopical attachment, of which field and forest species
also dominated. Small
amounts of meadow-and-swamp species (8,1 %), which were not on an initial area, appeared.
Four groups were selected on phenology
of development, among which spring-and-summer species dominated making
76,3 %. Summer species, which
were not in the spring, appeared in connection with the onset of summer. 5 groups
were selected on relation to soil humidity as well, of which
mesophiles (35,3 %) dominated. It was
21 common species between an initial area and grazing I. The degree of environmental
similarity among them was quite high
and amounted to 62,4 %.
After grazing II in the summer (from June till the end of August) the number of species (51) and genera (21) also did
not change. The composition of genera
including a large number of species changed. They were Amara (10 species), Harpalus (8) and
Carabus (5 species). The rest genera contained from 1
to 4 species. Within the genus Amara rare (80 %), spring-and-summer (70 %), field (70 %) species (A. famelica, A. familiaris, A. tibialis; of the genus Harpalus
- H. signaticornis and H. xanthopus winkleri) were
found.
22 new species appeared: there were no species on an
initial area
after grazing I. They belonged mainly to forest species (41 %) preferring mesophilic
conditions, extended in all seasons of year (since spring till autumn) or only
in the spring and in the summer. Thus two circumstances could play a role.
First, feeding horses favorably affected some field, rare species of
ground-beetles (Calathus fuscipes, C.
ambiguus, melanocephalus, C. halensis, Agonum dolens, Amara plebeja, A.
famelica, A. familiaris, ovata, A. similata, A. tibialis, A. bifrons, A.
apricaria, Bradycellus caucasicus, Harpalus tardus, Microlestes minutulus),
for which better microclimatic conditions were created. Secondly, though forest
species (Notiophilus palustris, Carabus hortensis, Epaphius secalis,
Pterostichus niger, Pt. melanarius, Calathus melanocephalus, Amara ovata, A.
similata, Harpalus smaragdinus) fly badly enough, they use flight for
settlement of young beetles, which emerged from a pupa and lost ability to
flight in the process of maturing. This leads to a well-known proverb, which
may be paraphrased and applied to ground-beetles: the born to run can't fly. Of
these 22 species A. tibialis and H. smaragdinus didn't appear more on
meadows. The first species preferred fields and consequently was very rare on
meadows, and the second one more often occurred in the young light forests.
After grazing II the quantity of dominants decreased to 2 - Poecilus cupreus (10,7 %), Bembidion
properans (5,4 %). Hence, conditions for dominants became worse. Loricera pilicornis was a subdominant
(1,5 %). The other 48 species were rare.
8 groups, of which field (54 %) and forest (26 %) species
also dominated, were selected on biotopical
attachment. Meadow-and-swamp species, which were not on an initial
area, appeared. 4 groups, among which spring-and-summer species again
dominated, were selected on phenology of development, making 58,8 %. With
approach of autumn more multiseasonal (23,5 %) species appeared, the quantity
of summer-and-autumn species became much less (5,9 %). 5 groups, of which mesophiles also dominated, were selected on relation to soil humidity. More mesoxerophile species appeared in connection
with the reduction of rainfall. There were 21 general species between grazing I
and grazing II. The degree of environmental similarity among them was high
enough making 62,7 %.
After grazing III in the end of August the quantity of
species (34) and genera (19) did not change in comparison with ground-beetles
after grazing II. The structure of genera containing a considerable quantity of
species changed a little. They were Amara (8 species), Harpalus and Calathus
(on 3 species). The other genera contained 1-2 species.
10 new
species, which weren't on an initial area, appeared after grazing I and grazing
II. It was mainly rare species, a half of which was made by field species. The
third part of species belonged to multiseasonal ones, and the rest – to
spring-and-summer species, late in the development.
After grazing III the quantity of dominants increased to
5 and almost reached an initial level
after grazing I, excluding Loricera
pilicornis (4,0 %), which became a subdominant. Two species – H. rufipes (5,7 %) and C. fuscipes (4,5 %) – were added to dominants. The quantity of subdominants increased
to 4. Hence conditions for ground-beetles improved after grazing III. The other
25 species were rare.
8 groups, of which field (45,5 %) and forest (27,3 %)
species dominated, were selected on biotopical attachment. It
was explained by the proximity of the fields with various cultures and the
forests (see Table 2). 4 groups, among which spring-and-summer species
dominated, were selected on phenology of development, making 61,7 %. 5 groups,
from which mesophiles dominated (44,1 %), were selected on relation to soil
humidity. There were 23 general species between grazing II and grazing III. The
degree of environmental similarity among them was low and amounted 37,1 %.
Horses did not graze any more as in the late autumn level of nitrates increased
in the grass due to an insufficient insolation and rainy weather.
If to spend a comparison of ground-beetle communities
after three grazing one could conclude that the sparing mode of grazing horses
didn't lead ground- beetle communities to a large reorganization in their
structure. The quantity of species remained unchanged throughout all growing
season. The structure of species was not changing long time, remained high
enough and only by the autumn made the third part of initial forms, due to the
fact that many species already finished the development by the autumn. The
species of field, forest and everybiont groups were steady enough to grazing
horses; their abundance remained at the same level throughout the whole season.
Meadow-and-field, meadow-and-forest, meadow, swamp and meadow-and-swamp groups,
which number was insignificant, were the most sensitive to grazing. Some groups
(meadow and forest) increased their number by the autumn, and others such as
meadow-and-field, meadow-and-forest and meadow-and-swamp decreased.
4. Conclusions
Thus, as a result of
studying zoocoenotic indicators of ground-
beetle communities 83 species living on floodplain meadows
were revealed. Of the ecological groups selected on relation to humidity the
greatest number of species on floodplain meadows presented mesophiles, and
after grazing I, obviously, with rainfall gigrophiles were. On the basis of
biotopic distribution we took these species to 8 ecological groups. Field
species were characterized by the most number, sometimes their abundance
reached 54 %. The majority of ground-beetle species (82,3 %) bred in the spring
and in the beginning of summer. Species of multiseasonal phenological group
weren't numerous (5,9 % – 26,5 %), and summer and summer-and-autumn groups were
random (2,6 % - 11,8 %). It was
explained by migration of “nearwater” mesophile species, which made a
considerable part in fauna, from a spring high water to adjacent biotopes.
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