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315
ACTA ZOOLOGICA BULGARICA
Terrestrial Ecology and Behaviour
Research Article
*Corresponding author: shalva.barjadze@iliauni.edu.ge
Acta Zool. Bulg., 75 (3), September 2023: 315-330
Published online 28 August 2023
https://www.acta-zoologica-bulgarica.eu/2023/002693
Diversity and Ecology of the Soil- and Litter-dwelling
Invertebrates and the Plant Associations in the Habitats
of the Saskhori Limestone Quarry and Adjacent Areas,
Eastern Georgia, Caucasus
Zezva Asanidze1, Tea Arabuli2, Eter Maghradze2, Lado Shavadze2, Mariam Gogshelidze2,
Naia Modebadze2, Eleonora Kiria2, Nana Barnaveli3 & Shalva Barjadze2*
1Institute of Ecology, Ilia State University, Giorgi Tsereteli 1, 0162, Tbilisi, Georgia
2Institute of Zoology, Ilia State University, Giorgi Tsereteli 3, 0162, Tbilisi, Georgia
3Department of Biology, Ivane Javakishvili Tbilisi State University, Universiteti 13, 0186, Tbilisi, Georgia
Abstract: This case study is aimed to investigate the invertebrate fauna and distribution features of the faunal and
-
cent territories (Eastern Georgia, Caucasus). Totally, 122 species of invertebrates and 131 species of plants
were registered. The isopod species Chaetophiloscia hastata
Lucoppia burrowsi Mi-
crozetorchestes emeryi and Nothrus parvus
-
rated the diversity of the degraded landscapes (1) using combined data of the vegetation and invertebrate
diversity and (2) using only the diversity data of invertebrates. The latter analyses, along with the other
Key words: Semiarid landscapes; species richness; indicator species; vegetation; South Caucasus
Introduction
The Caucasus ecoregion is one of 36 worldwide wild-
species and ecosystem biodiversity, high endemism
and high risk of biodiversity loss ( et al. 2000,
-
gia has devised a National Biodiversity Strategy and
Action Plan of Development (National Biodiversity
Strategy and Action Plan – Georgia, NBSAP, 2014–
2020) that establishes high priorities for the develop-
ment of inventories of animal species and resources.
-
316
Asanidze Z., Arabuli T., Maghradze E., Shavadze L., Gogshelidze M., Modebadze N., Kiria E., Barnaveli N. & Barjadze S.
-
(
dwelling invertebrates such as springtails (Collem-
bola), arachnids (Arachnida: Araneae, Pseudoscor-
Isopoda) and beetles (Insecta: Coleoptera) have not
-
ries. The abovementioned invertebrates are widely
distributed and have an important role in the func-
tioning of ecosystems. They participate in the pro-
cesses of soil formation and most of them represent
bioindicators of heavy metals pollution in the soil
( & 2004,
& 2010).
The goal of this project was to conduct taxo-
nomic, faunistic and ecological investigations of
soil- and litter-dwelling invertebrates and their as-
sociations with vegetation in seven selected sites of
Materials and Methods
We placed sampling sites in various habitat types
represented on the territory of the Saskhori lime-
maximum representativeness of the sample for the
study area. The total number of sampling sites was
and structurally arranged for continuous sample col-
lection during the active phenological period of the
samples were collected in each selected site (sites
1-7). Isolation of invertebrates from the samples
was made using Berlese and Winkler’s extractors.
The isolated invertebrates were preserved in 70 %
-
malin were used for sampling invertebrates. Inver-
tebrates’ mounting and pinning process and species
-
evant methodologies for each animal group.
Studied invertebrate groups are mites, spiders,
harvestmen, pseudoscorpions, isopods, millipedes,
centipedes, collembolans and beetles.
Sampling sites were visited thirteen times. We
-
tiation of the invertebrate diversity between the de-
graded and natural habitats in the study territory.
Vegetation diversity was sampled in each sam-
pling site, in plots with a size of 5 × 5 m, in ac-
cordance with the sample size corresponding to the
level of plant diversity in the semiarid shrublands
in the Caucasus Ecoregion ( et al.
-
and July, when the species richness reaches its maxi-
mum in semiarid habitats. We sampled plant species
richness and abundance of each species of vascular
plant in the vegetation survey plots using the meth-
data on the plant species abundance were converted
into percentage data following the methodology ex-
plained by & (2013).
Flora of Geor-
gia ( &
modern plant taxonomic data were obtained from
recent literature ( et al. 2005, -
et al. 2018) and Web sources (
22006-2021).
-
&
European Nature Information System (EUNIS)
( et al. 2004).
Habitat and environmental characteristics of the
localities of the placement of the sample collection
-
khori and its adjacent areas are provided in Table 1.
In statistical analysis, we estimated the Shan-
non Diversity Index and Evenness of the vegetation
diversity and diversity of the studied invertebrate
groups for each sampling site. The estimated data
were averaged to the unit of the sampling site in or-
der to simplify the introduction of the results of the
was avoided in the recent step of the data analysis.
low within-group variability of the variables caused
by the close allocation of the sampling plots on the
site. We consider this approach as being correct due
Sørensen–Dice index (
used to compare the results of our study with the
diversity data obtained from the literature sources
concerning the diversity of the plant and inverte-
brate communities distributed on the territories of
index was converted into a percentage of the simi-
larity by multiplying its numeric values by 100.
We used the method of the principal compo-
nent analysis (PCA) for the ordination of the veg-
etation and invertebrate diversity measured in the
sampling plots where PCs with the highest loads of
Diversity and Ecology of the Soil- and Litter-dwelling Invertebrates and the Plant Associations...
317
Table 1. Environmental characteristics of the localities of the placement of the sample collection sites on the territory
Site
ID Latitude Longitude
Geo-
graphic
position
Habitat Landscape feature
1 41.84584 44.51863 N
drought-resistant shrubbery (similar to ‘Pseudo-
-
Shrubland in the adjacent area to
2 41.84526 44.5173 N Extremely degraded habitat poor in vegetation Landscape with a bare surface of
3 41.84536 44.51661 E
Xero-thermophilous oak forest (similar to the
Quercus, Carpinus, Fraxi-
nus, Acer, Tilia, Ulmus, and related woodland
(habitat code: G 1. A)
Landscape with a degraded forest
in the adjacent area to the lime-
4 41.84706 44.51566 N
drought-resistant shrubbery (similar to ‘Pseudo-
-
Shrubland in the adjacent area to
5 41.84321 SE
land with unmixed crops grown by low-intensi-
ty agricultural methods” recognized by EUNIS
A landscape dominated by crop-
land – Almond garden
6 41.84655 44.50885 W
-
-
ciduous drought-resistant shrubbery (similar to
Degraded forest in small and dry
ravine adjacent to the limestone
7 S
-
riochloeto-Stipeto-Artemisieto steppes (similar
(Artemisia sp.) steppes recognized by EUNIS
The landscape of the abandoned
vegetation dominated by ruderal
and invasive plants
Fig. 1.
adjacent areas.
318
Asanidze Z., Arabuli T., Maghradze E., Shavadze L., Gogshelidze M., Modebadze N., Kiria E., Barnaveli N. & Barjadze S.
the analysed variables were used as the ordination
factors. PCA was done using abundance data of veg-
etation and invertebrate diversity.
We included united incidence data of plant and
invertebrate species in the cluster analysis and con-
ducted it using the algorithm of Unweighted Pair
and Jaccard similarity index as a method of dis-
tance measurement between the cluster groups. In
the graph visualizing the results of this analysis, we
also provided the average similarity of the diversity
of a particular surveyed site to the beta diversity or
task, we averaged the Jaccard similarity index of in-
dividual sites showing their similarity to the other
six sites and multiplied it by a hundred to convert it
into a percentage value of the similarity.
The means of the variables used in the cluster
analysis were compared using a two-tailed t-test be-
tween the cluster groups separated in the PCA. We
also employed an analysis of the similarity percent-
-
age percent contribution of separate plant and inver-
tebrate species in the formation of the communities
the extraction of limestone.
Indicator Species Analysis (ISA) was used for
the clearer attribution of the plant and invertebrate
species to the clusters of the natural and degraded
landscapes of the study area. In ISA, species indica-
-
cance of correlations of the species were calculated
using a randomization test with a bootstrap setting
-
er than 0.6 % were removed from the results of the
analysis to ensure the reliability of the results. The
analysis followed the methodology of &
&
Statistical analysis and illustration of the re-
sults were conducted using the software PAST v.
4.40 ( et al. 2001); SPSS v. 21 (
2013, & 2016).
Results
Taxonomic diversity
-
jacent territories, we registered 122 species of inver-
tebrates, of which 17 taxa of spiders; 8 centipedes,
millipedes and collembolans, 26 beetles, 4 isopods
and pseudoscorpions, 2 harvestmen and 45 oribatid
mites. Totally, 131 species of plants were recorded
in the studied territories, of which 18 species of
woody plants, 15 species of grasses, 2 sedges, 8 leg-
umes and 88 herbs (Appendix, Table A1).
The isopod Chaetophiloscia hastata Verho-
et al. 2023). Besides, three
oribatid mite species – Lucoppia burrowsi
Microzetorchestes emeryi and
Nothrus parvus
number of invertebrate species (52 species) was
found at sites Nos. 1 and 7, followed by 48, 46 and 45
species in sites Nos. 6, 4 and 3, respectively. In addi-
tion, 40 and 28 species were registered at sites Nos.
5 and 2, respectively (see Appendix, Table A2). Ex-
pectedly, invertebrate species richness was higher on
the landscapes including local natural habitats than in
-
tebrate species, i.e. Armadillidium vulgare (Latreille,
1804) and Trachelipus sp. (Isopoda), Phalangium
punctipes
Aleurodamaeus setosus (Berlese, 1883) and Schelori-
bates laevigatus-
mopolite species, were found in all sampling sites.
Ecological traits
The results of the PCA showed that the abundance
of each individual invertebrate group had the great-
appeared to be an important factor for the ordination
as most of the surveyed sites were occupied by a
abundance of the invertebrate groups and the spe-
cies richness of the herbaceous plants had the high-
est eigenvalues of loading on PC1 and PC2 (Table
diversity of woody plants, grasses, sedges and herbs
from the vegetation components as well as the di-
versity of invertebrate species represented by the
Chordeumatida had the discriminative impact on the
PC, with eigenvalues > 20, explained 82.3 % of the
variation of the diversity of the vegetation and hab-
PCA separated vegetation and invertebrates’
diversity of the degraded habitats (Site 2 – limestone
-
(sites 1, 3, 4 and 6). Cluster A comprised closely al-
located sites that were surveyed on the natural land-
scapes of the study area, and Cluster B united sites
Diversity and Ecology of the Soil- and Litter-dwelling Invertebrates and the Plant Associations...
Table 2.
values of loadings are shown in bold font in the table
Variables PC 1 PC 2 PC 3 PC 4 PC 5 t-test
Woody plants 0.503 0.001 0.015 -0.005 0.003**
Grasses & Sedges -0.003 -0.684 -0.024 0.010 0.017 0.012*
Legumes 0.000 0.043 -0.013 -0.005 0.183
Herbs -0.042 -0.572 0.051 0.333 0.021*
Vegetation cover in the plot (%) 0.832 -0.037 0.127 -0.370 0.002**
Coleoptera 0.002 0.025 -0.488 0.003 -0.018 0.143
Isopoda 0.908 -0.027 -0.087 -0.018 -0.121 0.001**
Sarcoptiformes -0.045 0.400 0.036 0.082
Araneae -0.056 0.972 0.112 -0.152 -0.073 0.001**
Polydesmida 0.000 -0.060 -0.001 0.006 0.070 0.122
Julida -0.020 -0.075 0.514 -0.002 -0.120 0.038*
-0.568 0.062 -0.020 -0.011 0.017 0.034*
Pseudoscorpiones -0.006 0.781 0.102 0.112 0.431 0.013*
Entomobryomorpha 0.002 -0.018 0.104 -0.036 -0.158 0.332
Geophilomorpha 0.612 -0.042 0.018 -0.086 0.025 0.146
Chordeumatida -0.021 -0.521 0.016 -0.388 0.003 0.018*
0.132 -0.063 0.002 0.321 -0.018 0.233
* Variability of the variable between A and B cluster
** Variability of the variable between A and B cluster
Fig. 2.
320
Asanidze Z., Arabuli T., Maghradze E., Shavadze L., Gogshelidze M., Modebadze N., Kiria E., Barnaveli N. & Barjadze S.
surveyed on the degraded landscapes. Sites included
in Cluster B were more scattered than in Cluster A,
which indicates the existence of a weak relationship
of the diversity between the sites of the degraded
vegetation cover (P = 0.032) and species richness of
the vegetation groups (woody plant species, grasses,
or no variability of the abundance and incidence of
The results of the cluster analysis based on the
of the invertebrates and vegetation diversity (Den-
drogram A) and particularly just of invertebrate di-
versity (Dendrogram B), also separated the diversity
of the sites of the degraded landscapes from those
linked communities of the shrublands closer to each
other (Sites 1 and 4) and slightly separated them
from the communities of the oak-hornbeam (site 3)
and woodland of the dry ravine (site 6) and united it
in the group together. The results of the analysis re-
lated closer the plant and invertebrate communities
to those of the almond garden (site 5) rather than the
by the limestone mining (site 2); however, all these
communities clustered in one group. The results of
of a closer relation to the invertebrate diversity of
the woodlands (sites 3 and 6) and a lower level of
similarity of the invertebrates of shrublands (sites 1
and 4). It also showed the higher similarity of the in-
vertebrate diversity between the sites of the almond
Comparing the diversity of vegetation be-
tween the sites of the natural (PCA cluster A) and
degraded (PCA cluster B) landscapes, the Jaccard
value 0.548), showing rather low similarity of the
vegetation between the two clusters. In the case of
0.623) similarity between the clusters.
particular plant and invertebrate species had sig-
in communities in both natural and degraded land-
scapes. However, the top 20 species of plants and
invertebrates (Table 3) ranked the most common
-
tebrates in these landscapes. There were no promi-
nent plant species with stronger contribution in the
formation of the vegetation of degraded and natural
-
ences in the mean values of the species contribution
species (Table 3) had higher importance in the for-
mation of the vegetation community of natural land-
scapes; the last four species were most common in
the degraded landscapes.
There was a similar tendency in the results of
the invertebrate communities of the studied land-
scape types. The contribution values of the inver-
Fig. 3. Dendrograms of the relationship between the diversities of the surveyed sites. A. Dendrogram based on com-
bined data of the invertebrate and vegetation diversity. B. Dendrogram based only on the invertebrate diversities.
Diversity and Ecology of the Soil- and Litter-dwelling Invertebrates and the Plant Associations...
321
each other, which indicated that there were no prom-
inent dominant species forming the invertebrate
communities in the natural or degraded landscapes
The results of ISA associated 11 plant species
and 12 invertebrate species to the diversity of the
of invertebrates to the degraded landscapes (Table
4). The attributed species were the most representa-
tive of the linked ecotopes based on their ecology.
Georgia, occurring in dryland habitats having frag-
mented distribution patterns and occurring in the
eastern part of the country.
Table 3.
invertebrates having the highest contribution to the formation of the vegetation community and invertebrate diversity
of the natural and degraded landscapes
Taxon Average
dissimilarity
Contribution
%
Cumulative
%
Mean – Natural
landscapes
Mean – Degraded
landscapes
Part A. Plant species
Stipa arabica 2.20 2.20 1.00 0.00
Brachypodium distachyon 2.20 4.40 1.00 0.00
Artemisia caucasica 1.62 1.80 6.20 0.75 0.00
Carex praecox 1.58 1.75 0.75 0.00
Astragalus hirtulus 1.46 1.62 0.75 0.00
Eremopyrum orientale 1.46 1.62 0.75 0.50
Lonicera caprifolium 1.46 1.62 12.81 0.75 0.00
Artemisia lercheana 1.46 1.62 14.42 0.75 0.00
Gentianella acuta 1.46 1.62 15.85 0.75 0.00
Daucus carota 1.43 17.28 0.75 0.00
Falcaria vulgaris 1.43 22.86 0.75 0.00
Aethusa cynapium 1.22 1.35 24.21 0.50 0.00
Setaria verticillata 1.22 1.35 25.56 0.50 0.00
Onobrychis miniata 1.22 1.35 0.50 0.00
Bupleurum exaltatum 1.27 1.41 20.10 0.00 1.00
Bunias orientalis 1.27 1.41 21.50 0.00 1.00
Centaurea solstitialis 1.27 1.41 0.00 1.00
Galium humifusum 1.22 1.35 28.21 0.00 1.00
Erysimum repandum 1.16 0.50 1.00
Part B. Habitat-specic invertebrates
Phalangium punctipes 1.16 0.00 1.00
Anthelephila pedestris 1.16 4.17 1.00 0.00
Armadilidium vulgare 1.62 0.25 1.00
Lithobius cronebergii 1.62 7.41 0.75 0.00
Tectocepheus velatus 0.81 1.46 8.87 0.75 0.00
Haplozetes tenuifusus 1.43 10.30 0.00 0.67
Crypticus quisquilius 1.43 11.73 0.00 0.67
Carabus adamsi 1.43 13.16 0.00 0.67
Trox sp. 1.43 0.00 0.67
Calathus melanocephalus 0.78 1.41 16.00 0.00 0.67
Brachinus crepitans 0.78 1.41 17.40 0.00 0.67
Pseudotorynorrhina sp. 0.74 1.34 18.74 0.00 0.67
Minunthozetes pseudofusiger 0.74 1.34 20.08 0.00 0.67
Lucoppia burrowsi 0.74 1.34 21.42 0.00 0.67
Zetorchestes micronychus 0.70 1.26 22.67 0.75 0.33
Liacarus brevilamellatus 1.25 0.25 0.67
Scytodes thoracica 1.24 25.17 0.75 0.33
Opilio sp. 1.24 26.40 0.75 0.33
322
Asanidze Z., Arabuli T., Maghradze E., Shavadze L., Gogshelidze M., Modebadze N., Kiria E., Barnaveli N. & Barjadze S.
Table 4.
Habitat-specic invertebrates Plant species
Taxa
Natural landscapes Degraded
Taxa
Natural landscapes Degraded
Indicator
values (%) P ≤ Indicator
values (%) P ≤ Indicator
values (%) P ≤ Indicator
values (%) P ≤
Anthelephila pedestris 0.88 0.001 - - Stipa capillata 0.001 - -
Lithobius erythrocephalus 0.76 0.001 - - Brachypodium dis-
tachyon 0.001 - -
Phalangium punctipes 0.62 0.050 0.85 0.001 Artemisia caucasica 0.86 0.001 - -
Armadilidium vulgare 0.87 0.001 0.71 0.01 Carex praecox 0.84 0.01 - -
Crypticus quisquilius - - 0.83 0.001 Eremopyrum orientale 0.72 0.03 0.60 0.05
Neobisium sp. 2 - - 0.82 0.001 Astragalus hirtulus 0.70 0.03 - -
Hister quadrimaculatus - - 0.8 0.001 Onobrychis angustifolia 0.66 0.03 - -
Trachelipus sp. - - 0.78 0.001 Daucus carota 0.64 0.05 - -
Lithobius ferganensis 0.001 - - Jasminum fruticans 0.60 0.05 - -
Carabus cf. maurus - - 0.76 0.01 Paliurus spina-Christi 0.60 0.05 - -
Chthonius sp. - - 0.68 0.01 Bupleurum exaltatum - - 0.88 0.001
Tegenaria sp. 0.73 0.001 0.63 0.03 Bunias orientalis - - 0.87 0.001
Agriotes sp. 0.72 0.001 0.63 0.03 Centaurea solstitialis - - 0.75 0.001
Opilio sp. 0.68 0.030 0.62 0.05 Carduus hamulosus - - 0.74 0.01
Dysdera sp. 0.65 0.050 0.6 0.05 Erysimum repandum - - 0.7 0.03
Geophilus sp. - - 0.6 0.05 Festuca varia 0.68 0.03 0.68 0.03
Gnaphosa sp. - - 0.6 0.05 Psephellus amblyolepis - - 0.64 0.05
Harpalus rupes 0.60 0.050 - - Scutellaria orientalis - - 0.63 0.05
Brachinus crepitans 0.60 0.050 - - Galium humifusum - - 0.63 0.05
Trochosa ruricola 0.60 0.050 - - Agropyron pectinatum - - 0.61 0.05
- - - - - Campanula hohenackeri - - 0.61 0.05
Diversity and Ecology of the Soil- and Litter-dwelling Invertebrates and the Plant Associations...
323
Discussion
The ecological part of our study clearly demonstrat-
diversity of the natural and degraded landscapes of
test of both plant and invertebrate groups as vari-
vegetation and invertebrate diversity between natu-
ral and degraded landscapes. However, only some
groups of plants and invertebrate show a high im-
pact on the separation of the clusters of the natural
and degraded landscapes; this is an expected result
as these groups include species of diverse ecological
which are linked to either natural or degraded land-
-
-
ever, it also shows that the contribution values of the
species included in the ranked list (Table 3) are not
The results of the PCA also indicate the absence of
their communities. It can be explained by the sam-
data.
The results of ISA create a clearer picture of
invertebrate diversity in the natural and degraded
landscapes of the study area by focusing on the most
representative species of these landscapes (Table 4).
The plant species aggregated to the sites of land-
scapes degraded due to anthropogenic activity (such
as limestone mining and gardening) are Bupleurum
exaltatum, Bunias orientalis, Centaurea solstitialis,
Carduus hamulosus, Erysimum repandum, Festuca
varia, Psephellus amblyolepis, Scutellaria orienta-
lis, Galium humifusum, Agropyron pectinatum and
Campanula hohenackeri. The invertebrate species
characteristic for these degraded habitats are: Geo-
philus sp. (centipede), Chthonius sp., Neobisium sp.
2 (false scorpions), Dysdera sp., Gnaphosa sp. (spi-
ders), Opilio sp., Phalangium punctipes (harvest-
men), Armadilidium vulgare, Trachelipus sp. (iso-
pods), Carabus cf. maurus, Crypticus quisquilius
and Hister quadrimaculatus (beetles).
-
card’s index shows that the diversity of the plant
and invertebrate communities of the degraded land-
scapes (sites 2, 5 and 7) is higher, with values of the
sites of the study area) than compared to the natural
landscapes. It means that the sites of degraded land-
scapes are inhabited by trivial species and the sites
of the natural landscapes support higher proportion
result as the species with the strong physiological
cosmopolite species, which are well adapted to a
wide range of environmental conditions (
& 2006, et al. 2006, -
&
these species, especially plants, gain a high rank
which determines the reliability of the results of
-
are cosmopolite species occurring in a wide range of
habitat and landscape types; however, they are also
characteristic for the habitats of Georgia, as they are
associated with the semiarid vegetation communi-
ties having fragmented distribution and occurring
in the eastern part of the country. These species are
the assimilation of degraded habitats. Their discov-
ery is one of the main results of our study as these
species form the diversity of the primary succession
on the degraded landscape with limestone-rich sub-
the entire South Caucasus.
& (2014) describes only
the taxonomic diversity of the plant communities
in Georgia. This information allows us to estimate
the similarity of the diversity of the plant commu-
-
pany and their adjacent habitats in the eastern part
of Georgia. According to this source, the vegeta-
-
ries at Dedophlistskaro, which are far east locality
in the country, and Gardabani, which is located in
the southeast part of Georgia, is dominated by the
-
bery, with fragments of the xerothermophilous oak
forest. These are the most widespread vegetation
of the dry types of woodlands, which occur in the
& (2014) associate 126
plant species with the Dedophlistskaro and 168
324
Asanidze Z., Arabuli T., Maghradze E., Shavadze L., Gogshelidze M., Modebadze N., Kiria E., Barnaveli N. & Barjadze S.
species with the Gardabani dry type of woodlands,
from which 54 species (36.7 %) are shared between
the communities of these two sites. The similar
(36.7 %) with the communities of Dedophlistskaro
(out of 110 species registered in similar plant com-
munities during our study). The comparison also
shows that all plant species associated with the
results of ISA (Table 4) are also represented in the
-
listskaro and Gardabani. This result once again
supports out observations on the successfulness of
-
tion for the occupation of the degraded landscape
3) are widely distributed species and they should
-
distributed species are dominating in the degraded
landscapes due their easer adaptation to harsh envi-
ronmental conditions.
-
It is highly desirable to continue with a repetitive
study in the future years in order to observe the dy-
namics of changes in the vegetation and invertebrate
diversity.
In addition, it should be mentioned that orib-
atid mite communities on post-industrial sand and
manganese tailing sites, reclaimed areas, natural
meadows and forests in the Chiatura region were
-
et al. 2013). Based on Sorensen-Dice index,
the similarity between oribatid mite complexes of
mining area is 35.11 %. All shared oribatid species
are widely distributed and, in our opinion, the deg-
radation of the natural habitats might be the reason
for their high similarity.
Acknowledgments: The investigation was supported by the
project Investigation of soil biodiversity in Saskhori Quarry
with the participation of local school pupils funded by the
Quarry Life Award, Heidelberg Cement, Georgia. We are
-
-
respectively.
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Appendix
Table A1.
N Plant Group Species
Sites of the sample collection
(Site ID)
1234567
1 Woody plants Acer laetum - --
2 Woody plants Carpinus orientalis - - ---
3 Woody plants Cotinus coggygria - ---
4 Woody plants Cotoneaster integerrimus -----
5 Woody plants Crataegus orientalis - - ---
6 Woody plants Ephedra procera - - ---
7 Woody plants Jasminum fruticans - - ----
8 Woody plants Juniperus communis
(syn. J. oblonga) - - ---
Woody plants Ligustrum vulgare ------
10 Woody plants Lonicera caprifolium - ---
11 Woody plants Paliurus spina - Christi - - ---
12 Woody plants Prunus spinosa ----- -
13 Woody plants Prunus dulcis (syn. Amygdalus communis) /
cultivated - - ---
14 Woody plants Quercus petraea subsp. iberica ------
15 Woody plants Rhamnus pallasii ------
16 Woody plants Rosa cannina - - ----
17 Woody plants Rosa spinosissima ------
18 Woody plants Swida australis - - ----
Grasses Agropyron pectinatum - - -
20 Grasses Bothriochloa ischaemum ------
21 Grasses Brachipodium sylvaticum - - ----
326
Asanidze Z., Arabuli T., Maghradze E., Shavadze L., Gogshelidze M., Modebadze N., Kiria E., Barnaveli N. & Barjadze S.
22 Grasses Brachypodium distachyon ------
23 Grasses Bromus squarrosu ------
24 Grasses Calamagrostis glomerata (Syn C. georgica) --- -
25 Grasses Dactylis glomerata - - ---
26 Grasses Elytrigia repens - - ---
27 Grasses Eremopyrum orientale - ---
28 Grasses Festuca varia -- - - -
Grasses Lolium temulentum ------
30 Grasses Phleum pratense - - ----
31 Grasses Setaria verticillata ----
32 Grasses Stipa arabica - - ---
33 Grasses Stipa capillata ----- -
34 Sedges Carex capitata ------
35 Sedges Carex praecox ----- -
36 Legumes Astragalus cyri ------
37 Legumes Astragalus hirtulus - - - - -
38 Legumes Astragalus falcatus - - - - -
Legumes Lathyrus tuberosus -----
40 Legumes Melilotus ocinalis - - ----
41 Legumes Onobrychis miniata -----
42 Legumes Onobrychis angustifolia - - ---
43 Legumes Trifolium pratense - - ---
44 Herbs Adonis parviora ----- -
45 Herbs Aegopodium podagraria - - ----
46 Herbs Aethusa cynapium -----
47 Herbs Agrimonia eupatoria ----- -
48 Herbs Ajuga chia - - ----
Herbs Allium albidum ------
50 Herbs Allium atroviolaceum - - ----
51 Herbs Anacamptis pyramidalis ------
52 Herbs Anchusa ocinalis - - - -
53 Herbs Anthemis candidissima - - ----
54 Herbs Arctium lappa - - - - -
55 Herbs Artemisia caucasica ----- -
56 Herbs Artemisia lercheana - ---
57 Herbs Asparagus verticillatus ----- -
58 Herbs Atriplex patula ------
Herbs Barbarea vulgaris ------
60 Herbs Bunias orientalis - - - -
61 Herbs Bupleurum exaltatum - -
62 Herbs Campanula bononiensis ------
63 Herbs Campanula hohenackeri -- - - -
64 Herbs Carduus hamulosus - - - -
65 Herbs Carthamus lanatus - - -
66 Herbs Centaurea solstitialis - - ---
67 Herbs Cerinthe minor --- -
68 Herbs Cichorium intybus ------
Herbs Cirsium arvense -----
70 Herbs Clinopodium vulgare ----- -
71 Herbs Consolida orientalis - - ---
72 Herbs Convolvulus cantabrica --- - -
73 Herbs Crepis pannonica ------
74 Herbs Cynoglossum ocinale - - - -
75 Herbs Daucus carota - - -
76 Herbs Dianthus crinitus ----- -
Diversity and Ecology of the Soil- and Litter-dwelling Invertebrates and the Plant Associations...
327
77 Herbs Dictamnus caucasicus ------
78 Herbs Echinops ruthenicus - - --
Herbs Echium russicum ----- -
80 Herbs Eryngium caeruleum ------
81 Herbs Erysimum repandum - - ----
82 Herbs Erysimum hieracifolium -- - -
83 Herbs Euphorbia seguieriana - - -- -
84 Herbs Euphrasia pectinata - - -- -
85 Herbs Falcaria vulgaris - - --
86 Herbs Galatella villosa ----- -
87 Herbs Galinsoga parviora -----
88 Herbs Galium album ------
Herbs Galium humifusum - - ----
Herbs Gentiana macrophylla ----- -
Herbs Gentianella acuta - - ---
Herbs Glaucium corniculatum -----
Herbs Globularia trichosantha - - ---
Herbs Gypsophila stevenii - - --
Herbs Helichrysum plinthocalyx ------
Herbs Hypericum perforatum ---- -
Herbs Inula aspera - - ----
Herbs Isatis costata ------
Herbs Lactuca serriola - - ---
100 Herbs Lapsana intermedia ------
101 Herbs Linum austriacum ----- -
102 Herbs Lycopsis arvensis ----- -
103 Herbs Mentha arvensis ------
104 Herbs Melampyrum arvense ----- -
105 Herbs Origanum vulgare - - ---
106 Herbs Ornithogalum pyrenaicum ------
107 Herbs Papaver arenarium ------
108 Herbs Petrorhagia prolifera ---- - -
Herbs Polygonum aviculare ----- -
110 Herbs Prunella vulgaris - - ---
111 Herbs Psephellus amblyolepis - - - -
112 Herbs Psephellus carthalinicus ----- -
113 Herbs Ranunculus repens ------
114 Herbs Rhinanthus minor ------
115 Herbs Rubia tinctorum - - - - -
116 Herbs Salvia viridis - - ---
117 Herbs Sanguisorba ocinalis ------
118 Herbs Scorzonera idae - - - - -
Herbs Scutellaria orientalis - ----
120 Herbs Senecio vernalis ----- -
121 Herbs Silybum marianum ----- -
122 Herbs Sonchus arvensis - --
123 Herbs Teucrium nuchense - - ---
124 Herbs Teucrium polium - - ---
125 Herbs Traagopogon serotinus ----- -
126 Herbs Veronica ocinalis ----- -
127 Herbs Viola rupestris ----- -
128 Herbs Xanthium spinosum ------
Herbs Xeranthemum annuum - - ----
130 Herbs Ziziphora acinos - - - - -
131 Herbs Urtica dioica - - - - -
328
Asanidze Z., Arabuli T., Maghradze E., Shavadze L., Gogshelidze M., Modebadze N., Kiria E., Barnaveli N. & Barjadze S.
Table A2.
N Family Species Sites of the sample collection
(Site ID)
Coleoptera 1 2 3 4 5 6 7
1 Anthicidae Anthelephila pedestris -----
2 Carabidae Brachinus crepitans (L., 1758) - - - - -
3 Carabidae Badister sodalis (Duftschmid, 1812) - - - - - -
4 Carabidae Calathus melanocephalus (L., 1758) - - - - -
5 Carabidae Carabus adamsi ---- -
6 Carabidae Carabus cf. maurus (Adams, 1817) - - - -
7 Carabidae Harpalus rupes (DeGeer, 1774) - - - - - -
8 Carabidae Olisthopus sp. -----
Chrysomelidae Podagrica sp. ------
10 Coccinellidae Coccinella septempunctata (L., 1758) - - - - - -
11 Coccinellidae Coccinella novemnotata ------
12 Curculionidae Anthonomus sp. - - ----
13 Curculionidae Otiorhynchus sp. - - - - -
14 Curculionidae Polydrusus sp. - - - ---
15 Elateridae Agriotes sp. - - - -
16 Histeridae Hister quadrimaculatus (L., 1758) ---
17 Forcula sp. - - - - - -
18 Lucanidae Dorcus parallelipipedus (L., 1758) ------
Scarabaeidae Amphimallon solstitiale (L., 1758) - - - - - -
20 Scarabaeidae Oryctes sp. ------
21 Scarabaeidae protaetia sp. - - - - -
22 Silphidae Silpha cf. obscura L., 1758 - - - - - -
23 Tenebrionidae Dendarus sp. ----- -
24 Tenebrionidae Crypticus quisquilius (L., 1760) - - - - - -
25 Tenebrionidae Eleodes sp. ------
26 Trogidae Trox sp. - - - - -
Geophilomorpha 1 2 3 4 5 6 7
27 Dignathodontidae Henia cf. brevis ------
28 Geophilidae Clinopodes caucasicus - - ----
Geophilidae Geophilus sp. ----
Lithobiomorpha 1 2 3 4 5 6 7
30 Lithobiidae Lithobius sp. ------
31 Lithobiidae Lithobius aeruginosus L. Koch, 1862 - - - - -
32 Lithobiidae Lithobius erythrocephalus C.L. Koch, 1847 - - - - - -
33 Lithobiidae Lithobius ferganensis - - - -
Scutigeromorpha 1234567
34 Scutigeridae Scutigera coleoptrata L., 1758 - -----
Entomobryomorpha 1234567
35 Orchesella cf. cincta (L., 1758) -
36 Entomobryidae Lepidocyrtus lignorum - ---
37 Entomobryidae Entomobrya cf. quinquelineatus ------
38 Entomobryidae Seira domestica (Nicolet, 1842) - - - - - -
Tomoceridae Tomocerus vulgaris (Tullberg, 1871) - - ----
40 Isotomidae Isotomurus sp. --- -
Diversity and Ecology of the Soil- and Litter-dwelling Invertebrates and the Plant Associations...
Poduromorpha 1 2 3 4 5 6 7
41 Neanuridae Anurida sp. ------
Symphypleona 1 2 3 4 5 6 7
42 Sminthuridae Sminthurus viridis (L, 1758) - - - -
43 Phalangiidae Opilio sp. - - -
44 Phalangiidae Phalangium punctipes
Isopoda
45 Armadillidiidae Armadillidium vulgare (Latreille, 1804)
46 Trachelipodidae Trachelipus sp.
47 Philosciidae Chaetophiloscia hastata - - ---
48 Ligiidae Ligidium sp. - - ---
Chordeumatida 1 2 3 4 5 6 7
Anthroleucosoma-
tidae
Metamastigophorophyllon cf. torsivum
- ----
50 Anthroleucosoma-
tidae
Pseudoagellophorella eskovi
-----
Polydesmida 1 234567
51 Paradoxosomatidae Strongylosoma sp. - - - - - -
Julida 1 2 34567
52 Julidae Omobrachyiulus caucasicus (Karsch, 1881) - - - - - -
53 Julidae Byzantorhopalum rossicum - - ----
54 Julidae Cylindroiulus sp. - - - - - -
55 Julidae Leptoiulus tanymorphus - - --
56 Julidae Syrioiulus adsharicus ---- -
Pseudoscorpiones 1 2 3 4567
57 Neobisiidae Neobisium sp. 1 -
58 Neobisiidae Neobisium sp. 2 - - --
Atemnidae Atemnus politus (Simon, 1878) ------
60 Chthoniidae Chthonius sp. - - --
Araneae 1 2 3 4 5 6 7
61 Agelenidae Tegenaria sp. - - -
62 Araneidae Araneus sp. - - - - - -
63 Clubionidae Clubiona sp. - -
64 Dysderidae Dysdera cf. spasskyi - --
65 Dysderidae Dysdera sp.
66 Gnaphosidae Gnaphosa sp. - -
67 Gnaphosidae Nomisia exornata - - - -
68 Lycosidae Hogna radiata (Latreille, 1817) - - - ---
Lycosidae Trochosa ruricola (De Geer, 1778) - - - -
70 Lycosidae Trochosa sp. ----
71 Lycosidae Thanatus sp. - - ---
72 Salticidae Euophrys frontalis (Walckenaer, 1802) ------
73 Scytodidae Scytodes thoracica (Latreille, 1802) - --
74 Sparassidae Micrommata virescens (Clerck, 1757) - - - - - -
75 Theridiidae Steatoda nobilis (Thorell, 1875) - - - - - -
76 Thomisidae Ozyptila sp. -
77 Thomisidae Xysticus kochi Thorell, 1872 - -
Oribatida 1 2 3 4 5 6 7
78 Nothridae Nothrus parvus ------
330
Asanidze Z., Arabuli T., Maghradze E., Shavadze L., Gogshelidze M., Modebadze N., Kiria E., Barnaveli N. & Barjadze S.
Crotoniidae Camisia horrida (Hermann, 1804) - - - - -
80 Crotoniidae Camisia lapponica -----
81 Euphthiracaridae Acrotritia ardua (Koch, 1841) - - - - - -
82 Phthiracaridae Phthiracarus lentulus (Koch, 1841) ------
83 Phthiracaridae Steganacarus carinatus (Koch, 1841) -
84 Ramusella clavipectinata -----
85 Oppiella (Rhinoppia) hygrophila -----
86 Hermanniellidae Hermanniella punctulata - - - -
87 Liacaridae Dorycranosus splendens -- -
88 Liacaridae Dorycranosus ovatus - - ----
Liacaridae Liacarus brevilamellatus --- -
Liacaridae Liacarus coracinus (Koch, 1841) - - ----
Gymnodamaeidae Arthrodamaeus femoratus (C.L.Koch, 1840) - - -
Aleurodamaeidae Aleurodamaeus setosus (Berlese, 1883)
Damaeidae Metabelba monilipeda ----- -
Damaeidae Metabelba agelliseta ------
Haplozetidae Haplozetes tenuifusus ---- -
Scutoverticidae Scutovertex minutus (Koch, 1835) - - ----
Plateremaeidae Lopheremaeus mirabilis ------
Amerobelbidae Amerobelba decedens ----- -
Neoliodidae Liodes theleproctus (Hermann, 1804) - - - -
100 Neoliodidae Poroliodes farinosus ------
101 Ceratozetidae Trichoribates naltschicki - - -
102 Ceratoppiidae Ceratoppia bipilis (Hermann, 1804) -
103 Zetorchestidae Microzetorchestes emeryi -----
104 Zetorchestidae Zetorchestes micronychus (Berlese, 1883) - - -
105 Xenillidae Xenillus tegeocranus (Hermann, 1804) -
106 Oribatella berlesei ------
107 Gustaviidae Gustavia microcephala (Nicolet, 1855) ---
108 Scheloribatidae Scheloribates laevigatus (Koch, 1835)
Phenopelopidae Eupelops acromios (Hermann, 1804) - ---
110 Phenopelopidae Eupelops torulosus - - ---
111 Phenopelopidae Eupelops occultus (Koch, 1835) - -----
112 Phenopelopidae Peloptulus phaenotus (Koch, 1841) -----
113 Lucoppia burrowsi -----
114 Oribatula tibialis (Nicolet, 1855) -
115 Oribatula (Zygoribatula) cognata --
116 Zygoribatula exilis (Nicolet, 1855) -----
117 Punctoribatidae Punctoribates punctum -
118 Punctoribatidae Minunthozetes pseudofusiger -----
Carabodidae Austrocarabodes foliaceisetus georgiensis -
vanidze &Weigmann, 2007 -----
120 Tectocepheidae Tectocepheus velatus - ---
121 Galumnidae Pergalumna nervosa ------
122 Galumnidae Galumna alata (Hermann, 1804) - -
