225
Abstract
Dry grassland ecosystems are highly fragmented in Ukraine and increasingly 
threatened from intensification of farming practices. Evaluation of the habitat 
conservation importance based on multitaxon studies allows for optimizing selec-
tion of priority sites. Botanical research of xerothermic habitats in the Kryvorizkyi 
Iron Ore Basin has been conducted since 2012, while spiders were investigated for 
the first time in 2017. In the four study sites, we recorded 265 vascular plant spe-
cies (23 under protection) and 95 spider species (19 are rare). Both vegetation and 
spider communities of the site Zelena Gully are well preserved and rich in rare 
and threatened species (18 plant and 11 spider species), which confirms its value 
as a priority site of the Emerald Network. Chervona Gully and the Slate Rocks 
should be included in the network. Frequent fires and intensive grazing in Khrys-
toforova Gully affect the spider diversity, decreasing the number of specialist and 
rare species, while early flowering plants benefit from this disturbance. Neverthe-
less, its vegetation is poorer than in Zelena and Chervona gullies. Inclusion in the 
Emerald Network is critically important to the investigated sites, since expansion 
of mining activities is constantly threatening natural habitats in industrial regions.
Iz vleček
Suha travišča so močno fragmentiran ekosistem v Ukrajini, ki jih močno 
ogroža intenzifikacija kmetijstva. Vrednotenje naravovarstvenega pomena na 
osnovi raziskav večih taksonov omogoča optimalno izbiro prioritetnih območij. 
Botanične raziskavekserotermnih habitatov na območju nahajališča železove rude 
Kryvorizkyi, potekajo od leta 2012, pajke pa smo prvič preučili leta 2017. Na 
štirih raziskovanih območjih smo zabeležili 265 rastlinskih vrst (23 zavarovanih) 
in 95 vrst pajkov. Vegetacija in združbe pajkov so v jarku Zelena dobro ohranjene 
s številnimi redkimi in ogroženimi vrstami (18 rastlin in 11 pajkov), kar potrjuje 
uvrstitev kot prioritetnega območja v omrežju Emerald, v katerega bi morali 
vključiti tudi jarek Chervona in skrilasto skalovje. Pogosti požari in intenzivna 
paša v jarku Khrystoforova vplivajo na pestrost pajkov ter zmanjšanje števila 
specialistov in redkih vrst, po drugi strani pa ta motnja ustreza zgodaj cvetočim 
rastlinam . Klub temu je vegetacija vrstno siromašnejša kot v jarkih Zelena in 
Chervona. Vključitev v omrežje Emerald je kritično pomembna za preučevana 
območja, saj širitev rudarskih aktivnosti predstavlja stalno grožnjo naravnim 
habitatom v industrijskih regijah.
Key words: conservation areas, 
protected plant species, spider fauna, 
steppe habitats, Ukraine.
Ključne besede: naravovarstvena 
območja, zavarovane rastlinske vrste, 
pajki, stepski habitati, Ukrajina.
Received: 31. 03. 2020
Revision received: 16. 12. 2020
Accepted: 28. 12. 2020
Co-ordinating Editor:  
Stephen Venn
1 V. N. Karazin Kharkiv National University, 4, Maidan Svobody, Kharkiv 61022, Ukraine. E-mail: n.polchaninova@karazin.ua, 
2 Kryvyi Rih Botanical Garden of the National Academy of Sciences of Ukraine, 50, Marshaka Str., Kryvyi Rih 50089, Ukraine. E-mail: kras.kbs.17@gmail.com
3 Kryvyi Rih State Pedagogical University, 54 Gagarin Av, Kryvyi Rih, 50086, Ukraine. E-mail: lisogor1981@gmail.com
*
 
Corresponding author
Nina Polchaninova1
,
*

, Olga Krasova2

, Liudmyla Lysohor3

 &  
Tatiana Atemasova1

DOI: 10.2478/hacq-2021-0003 20/1 • 2021, 225–242
Assessment of the conservation value 
of dry grassland habitats in the Inhulets 
River basin (Central Ukraine) based on 
vegetation and spider research

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Assessment of the conservation value of dry grassland habitats in the Inhulets 
River basin (Central Ukraine) based on vegetation and spider research
Introduction
The problem of biodiversity loss has become global and it 
continues to escalate under increasing pressure of indus-
trial society. Given high fragmentation of natural ecosys-
tems in the steppe zone of Ukraine, virtually every dry 
grassland area preserved in the agro-industrial landscape 
is of great conservation importance. These islands of na-
tive steppe biota are the refuges for many species, which 
can colonize abandoned and arable lands (Öberg et al. 
2008, Knappová et al. 2012), and thus promote biodi-
versity and ecosystem sustainability (Tamme et al. 2010, 
Poggio et al. 2010, Harlio et al. 2019).
A current mechanism of effective phyto- and habitat di-
versity conservation in Europe is the inclusion of natural 
vegetation refugia into Natura 2000, a network of nature 
protection areas in the territory of the EU. The Emerald 
Network serves the same purpose in non-EU countries. 
The theoretical basis for these networks is a habitat con-
cept formulated at the Berne Convention on Conservation 
of European Wildlife and Natural Habitats (Convention 
1979). In Ukraine, the Emerald Network has been devel-
oping since 2009; however, the dry grassland habitats were 
included in it only in 2016 (see Kuzemko et al. 2018).
It is advisable to assess the conservation value of a given 
area using multitaxon studies as they enable the devel-
opment of adequate measures for preserving biodiversity 
(Birkhofer et al. 2015, Harlio et al. 2019). Therefore, 
joint vegetation and invertebrate researches are valuable 
(Braschler & Baur 2016, Zao et al. 2018, Outhwaite et 
al. 2020), since the taxa responses to the same impact are 
far from being uniform (Niemelä & Baur 1998, Lafage et 
al. 2015, Polchaninova et al. 2019).
Spiders are known as indicators of environmental 
changes (Rodriguez-Artigas et al. 2016, Zografou et al. 
2017), habitat structure (Buchholz 2010), and manage-
ment (Batáry et al. 2012, Polchaninova et al. 2019) of 
dry grassland ecosystems. It is reasonable to use spiders in 
nature conservation studies, since they are numerous, di-
verse in ecological preferences, and inhabit all vegetation 
layers in various biotops (Perner & Malt 2003, Cardoso 
et al. 2004, Szmatona-T úri et al. 2019).
Criteria for the assessment of spider species vulnerability 
in Ukraine have not developed, and the order Araneae is 
absent from the National Red List. Some spiders are pro-
tected at the regional level, thus, six species are listed in the 
Red Data Book of Kharkiv Region, and 12 species in the 
Red Book of Donetsk Region (Tokarsky 2013, Zalevskiy 
& Bronskov 2017). Moreover, 80 species from the Ukrai-
nian Carpathians are included in the Carpathian Red List 
of Forest Habitats and Species (Gajdoš et al. 2014). Dry 
grassland spiders are studied relatively well in Left-Bank 
Ukraine (a region stretching east from the left bank of the 
Dnieper River to the state border). The faunistic and eco-
logical data were summarized in the Catalogue of spiders 
of Left-Bank Ukraine and its Addendum (Polchaninova & 
Prokopenko 2013, 2017), where “steppe” as a habitat was 
recorded for 370 species. Contrastingly, arachnological re-
search on the right-bank steppes of Ukraine is at the initial 
stage. The investigations were conducted only in some lo-
calities in Odeska (Deli, 2014), Mykolaivska (Polchaninova 
et al. 2017) and Khersonska (Prokopenko & Zhukov 2018, 
Iosypchuk 2019) oblasts. Prior to our studies, the inverte-
brate research in the Inhulets basin concerned only ground-
dwelling insects (Smetana &Smetana 2005). Preliminary 
data on spider species richness and xerothermic habitats of 
the area in question are given in Krasova et al. 2020. 
The aim of the present paper is (1) to compile a list of 
dry grassland habitats of the study area in accordance with 
the National Habitat Catalogue of Ukraine (Kuzemko et 
al. 2018); (2) to reveal rare species of the dry grassland 
flora and spider fauna; (3) to analyze specificity of spider 
assemblages in the study habitats; (4) to assess conserva-
tion value of the study sites based on both IPA criteria 
and diversity of spider communities.
Material and methods
Study area and sampled sites
The Ingulets River, the largest right tributary of the Dnieper 
lower reaches, has a length of 549 km. It crosses the Dnie-
per Upland and the Black Sea Lowland in the sub-meridian 
direction. Four smaller rivers, the Saksahan, the Zelena, the 
Zhovta and the Bokova, as well as a number of large gullies 
with permanent streams, flow into the Inhulets in its mid-
dle reaches. The natural hydrographic network has been 
partly destroyed during the ironstone mine workings.
In terms of geo-botanical regions, the area belongs 
mainly to the Buh-Dnieper (Kryvyi Rih) district of forb-
fescue-feather grass steppes, bairak forests (natural forests 
in the gullies in steppe and forest-steppe zones), and the 
vegetation of granite outcrops within the Black Sea–Azov 
steppe subprovince of the Pontic Steppe province of the 
Steppe subregion of Palearctic. The southern part of the 
study area is located in the same subprovince, in the Buh–
Inhul district of the fescue-feather grass steppes, steppe-
like meadows and the vegetation of limestone outcrops 
(Didukh & Shelyag-Sosonko 2003). Steppe habitats of 
today in the Inhulets basin have been preserved exclu-
sively as sloping biotopes in the gully and river systems. 
The research was conducted in four localities in the vi-
cinity of the town of Kryvyi Rih, Dnipropetrovska Oblast 
(Figure 1, 2):

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Assessment of the conservation value of dry grassland habitats in the Inhulets 
River basin (Central Ukraine) based on vegetation and spider research
Figure 1: Map of collecting localities. 1. Chervona Gully, 2. Slate 
Rocks, 3. Khrystoforova Gully, 4. Zelena Gully.
Slika 1: Karta preučevanih lokalitet. 1. jarek Chervona, 2. skrilaste 
skale, 3. jarek Khrystoforova, 4. jarek Zelena.
Figure 2: Map of the sample plots 
in the study sites. 1 – Chervona 
Gully: 1.1 – forb-fescue-feather 
grass steppe, 1.2 – steppe scrub, 
1.3 – meadow steppe; 2 – Slate 
Rocks, stony slope; 3 – Khrystoforo-
va Gully: 3.1 – forb-fescue-feather 
grass steppe, 3.2 – petrophytic steppe 
on granite bedrock, 3.3 – meadow 
steppe; 4 – Zelena Gully: 4.1 – forb-
fescue-feather grass steppe, 
4.2 – steppe scrub, 4.3 – meadow 
steppe, 4.4 – petrophytic steppe on 
limestone bedrock.
Slika 2: Karta vzorčnih ploskev na 
preučevanih mestih. 1 – jarek Cher-
vona: 1.1 – travnata stepa z zelišči, 
bilnicami in bodalicami, 1.2 – stepa 
z grmišči, 1.3 – ravniška stepa; 
2 – skrilaste skale, kamnito pobočje; 
3 – jarek Khrystoforova: 3.1 – trav-
nata stepa z zelišči, bilnicami in 
bodalicami, 3.2 – petrofitska stepa 
na granitni podlagi, 3.3 – travniška 
stepa; 4 – jarek Zelena: 4.1 – trav-
nata stepa z zelišči, bilnicami in 
bodalicami, 4.2 – stepa z grmišči, 
4.3 – travniška stepa, 4.4 – petrofit-
ska stepa na apneniški podlagi.

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Assessment of the conservation value of dry grassland habitats in the Inhulets 
River basin (Central Ukraine) based on vegetation and spider research
1. Kryvyi Rih T own, Chervona Gully, geographic coordi-
nates of the central part of the study site 48°06'15.5"N 
33°31'02.8"E (Figure 3). The middle part of the gully 
with a chain of small quartzite rocks has the status of 
a geological preserve of local importance; an area of 
28.3 ha has a status of a landscape preserve of national 
importance. The gully is located on the outskirts of 
the town and borders an iron ore quarry and its waste 
banks. A stream runs along the gully bottom, the 
alongside meadows are partly used as a cattle pasture. 
The gully is crossed by paths but in general the vegeta-
tion is not trampled.
2. Kryvyi Rih Town, Slate Rocks, 47°56'27.8"N 33°23' 
18.5"E (Figure 4). Geological monument of nature of 
local importance. It is a chain of amphibolite and slate 
rocks 20 m high and 250 m long, preserved in the urban 
ecosystem. The site is often visited by holidaymakers.
3. Kudasheve Village, Khrystoforova Gully, 47°57'46.5"N 
33°04'14.4"E( Figure 5). A shallow gully with gentle 
slopes and small patches of granite outcrops on the 
lower slopes. The gully is used as a pasture and often 
experiences chaotic burning. In the study year, nearly 
the entire gully was burnt in early spring.
4. Poltavka Village, Zelena Gully, 47°45'11.0"N 33°11' 
21.3"E (Figure 6). The gully stretches across the border 
of the forb-fescue-feather grass and fescue-feather grass 
steppe subzones. Carbonate substrata (limestone) surface 
on the lower parts of north-facing slopes and provide 
accumulation of thick diluvium along the bottom. The 
 investigated part of the gully is rarely visited by people.
Figure 3: Chervona Gully, May 2017.  
The gully is adjacent to the iron ore quarry 
(background). Photo: Nina Polchaninova.
Slika 3: Jarek Chervona, maj 2017. Jarek je v 
bližini izkopa železove rude (v ozadju).  
Foto: Nina Polchaninova.
Figure 4: Slate Rocks, May 2017.  
Photo: Nina Polchaninova.
Slika 4: Skrilaste skale, maj 2017.  
Foto: Nina Polchaninova.

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River basin (Central Ukraine) based on vegetation and spider research
Figure 5: Khrystoforova Gully, south-facing 
slope with granite outcrops. May 2017, a 
month after the fire. Photo: Nina Polchaninova.
Slika 5: Jarek Khrystoforova, južno pobočje z 
granitnimi izdanki. Maj 2017, mesec po požaru. 
Foto: Nina Polchaninova.
Figure 6: Zelena Gully, June 2017.  
Photo: Nina Polchaninova.
Slika 6: Jarek Zelena, junij 2017.  
Foto: Nina Polchaninova.
Vegetation sampling and habitat 
classification
In the course of the field studies in 2012–2017, a total 
of 122 geobotanical descriptions were made: Chervona 
Gully (n=34); Slate Rocks (n=27); Khrystoforova Gully 
(n=22); Zelena Gully (n=39). Sizes of sample plots de-
pended on the type of habitat and ranged from 1×1 m 
(rocky vegetation with mosses and lichens) to 5×5 m 
(grassy communities and steppe scrub) (Pott 2011). Clas-
sification of the study habitats was adopted from the 
“National Habitat Catalogue of Ukraine” (Kuzemko et 
al. 2018). The habitats were identified to the level of veg -
etation associations (Figure 2, Table 1).
Spider collection
Spiders were collected in May–June 2017 by standard 
collecting methods: sweep netting, pitfall trapping and by 
hand. To cover habitat diversity of each site, we arranged 
sample plots at the gully bottoms and on the slopes of 
various expositions (Figure 2). At each sample plot, eight 
traps (300 ml plastic caps) were set up in a line, at a 
distance 10 m from each other. A 4% formalin solution 
served as preservative. The traps were exposed for 33 days.
A total of 11 plots were sampled at four sites (Figure 2), 
their vegetation characteristics and syntaxonomy is given 
in Table 1: Chervona Gully: ChG1 (1.1) – forb-fescue-

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River basin (Central Ukraine) based on vegetation and spider research
Table 1: Xerothermic habitats of the study sites. Numbers and letters in bold indicate habitat location on the map (Figure 2) and 
abbreviations in the text. For the explanations, see Material and Methods.
Tabela 1: Kserotermni habitati na preučevanih mestih. Številke in črke, podarjene krepko, predstavljajo lokacijo habitata na karti 
(Slika 2) in okrajšave v besedilu. Za razlago glej poglavje Material in metode.
Syntaxa after
Prodrome of the vegetation of 
Ukraine
Habitats after
National Habitat Catalogue of 
Ukraine
Habitats after
EUNIS classification
adapted for Ukraine
Festuco-Brometea Br.-Bl. et Tx. ex Soo 1947,
T anaceto achilleifolii-Stipetalia lessingianae Lysenko et Mucina 
in Mucina et al. 2016,
Poo bulbosae-Stipion graniticolae Vynokurov 2014,
Potentillo incanae-Seselietum pallasii Vynokurov 2014 
(3.2, Kh2)
Potentillo arenariae-Linion czerniaevii Krasova et Smetana 
1999,
Lino tenuifolii-Jurineetum brachycephalae Krasova et Smetana 
1999 (4.4, ZG4)
Festucetalia valesiacae Soó 1947,
Festucion valesiacae Klika 1931,
Thalictro mini-Spiraeetum hypericifoliae Vynokurov 2014  
(1.3, ChG3),
Elytrigio trichophorae-Poetum angustifoliae Osychniuk et al.  
ex Solomakha 1995 (3.3, KhG3; 4.3, ZG3)
T anaceto achilleifolii-Stipetalia lessingianae Lysenko et 
Mucinain Mucina et al. 2016,
Stipo lessingianae-Salvion nutantis Vynokurov 2014
Stipo lessingianae-Salvietum nutantis Vynokurov 2014  
(1.1, ChG1; 3.1, KhG1)
T anaceto millefolii-Galatellion villosae Vynokurov in 
Kolomiichuk et Vynokurov 2016
Ephedro distachyae-Stipetum capillatae Kolomiychuk et 
Vynokurov 2016 (4.1, ZG1)
T1. Dry grassy habitats
T1.2 Petrophytic steppes
T1.2.1 Petrophytic steppes on 
siliceous substrata
T1.2.2.‘ б’ Petrophytic steppes on 
carbonate substrata of the Black 
Sea region
T1.3 Meadow steppes
T1.3.2 Meadow steppes on 
chernozems
T1.4 T rue forb-fescue-feather 
grass and fescue-feather grass 
steppes
T1.4. a T rue forb- fescue-feather 
grass and fescue-feather grass 
steppes of the steppe zone 
E1. Dry grasslands
E1.11 Euro-Siberian rock 
debris swards
E1.2
(no equivalent)
E1.23 Meso-xerophile 
subcontinental
meadow-steppes
E1.2D Ponto-Sarmatic steppes
Rhamno-Prunetea Rivas Goday et Borja Carbonellex 
Tüxen1961,
Prunetalia spinosae Tüxen1952,
Prunion fruticosae Tx. 1952, 
Amygdalo nanae-Spiraeetum hypericifoliae Fitsailo 2008  
(1.2, ChG2; 4.2, ZG2)
‘ Ч’4. Deciduous shrubs
‘ Ч’4.2 Steppe scrub
F3.247 Ponto-sarmatic 
deciduous thickets
Sedo-Scleranthetea Braun-Blanquet 1955, Trifolio arvensis-
Festucetalia ovinae Moravec 1967, Sempervivo-Sedion 
(Braun-Blanquet 1955) Müller 1961
Sempervivo ruthenicae-Sedetum ruprechtii Didukh et Kontar 
1998 (2, SR)
K1. Siliceous rocks
K1.3 Siliceous rocks and screes of 
the Ukrainian crystalline shield
H3.19 Lowland northern- and 
middle-European siliceous cliffs
feather grass steppe on a south-facing slope, ChG2 (1.2) – 
steppe scrub on a north-facing slope, ChG3 (1.3) – mead-
ow steppe at the gully bottom. Slate Rocks: SR (2) – steep 
stony slope with sparse vegetation. Khrystoforova Gully: 
KhG1 (3.1) – forb-fescue-feather grass steppe on a north-
facing slope, KhG2 (3.2) – petrophyticsteppe on granite 
bedrock on the lower slope, Kh3 (3.3) – meadow steppe at 
the gully bottom. Zelena Gully: ZG1 (4.1) – forb-fescue-
feather grass steppe on a south-facing slope, ZG2 (4.2) 
– steppe scrub on the upper slope, ZG3 (4.3) – meadow 
steppe at the gully bottom, ZG4 (4.4) – petrophytic steppe 
on limestone bedrock on a north-facing slope. 
In addition, we set up four lines of traps in Chervona 
Gully from 9 to 29 September 2017. Since the study plot 

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River basin (Central Ukraine) based on vegetation and spider research
1.1 was burnt, we chose another plot with forb-fescue-
feather grass vegetation and added steppe plots on silicate 
bedrock and old mound. The steppe scrub was sampled at 
the same plot (ChG2).
Herb-dwelling spiders were collected on 4–6 May, 
8–10 June and 9 September by sweep netting. Spiders 
were swept near the trap lines and further along the slopes 
and/or gully bottoms. At least three samples of 50 sweeps 
were taken from each habitat.
Data analysis
We used criterion A (presence of threatened plant spe-
cies) to evaluate botanical importance of the study sites 
(Anderson 2002, Onyshchenko 2017). We also followed 
criterion C (the presence of habitats from the Revised 
Annex I from the Resolution 4 of the Bern Convention 
2019) to categorize the habitat conservation status and 
threat of its degradation under changing of grassland 
management, high recreation pressure, frequent burning 
and/or afforestation. We assessed floristic richness of the 
study habitats and compared them using Sørensen simi-
larity index (Magurran 1991).
We defined spider species rarity based on their general 
geographic distribution (WSC 2020), distribution in the 
East European Plain (Mikhailov 2013, Polchaninova & 
Prokopenko 2019; Nentwig et al. 2020), and habitat 
distribution within the steppe zone of Eastern Europe 
(Polchaninova & Prokopenko 2013, 2017, Ponomarev 
2017). The spider species list is arranged as a table in-
dicating localities and habitats; autumnal collection is 
compiled in a separate column (Appendix, Table 2). We 
mentioned several species as juveniles, if adults from a 
certain locality were not recorded. The spider taxonomy 
follows WSC (2020); adult individuals were identified us-
ing Nentwig et al. 2020.
Given species habitat preferences, seven ecological 
groups of spiders were distinguished: g – generalists,  
f/gr – inhabiting open forests, forest edges and mesic 
grasslands, f/dgr – same forest habitats and dry grass-
lands, gr – grasslands of various types, mgr –wet and/or 
mesic grasslands, dgr – dry grasslands, st – steppe spe-
cialists, and a group of unspecified species (un) with data 
deficiency.
Comparison of the spider faunas of investigated 
habitats was based on the total list of species collected 
in May–June in the four study localities. Sørensen index 
was taken as a similarity measure (Magurran 1991). For 
convenience, we pooled the habitats SR, KhG2 and ZG4 
into one group of “stony slopes”. Hand collection and 
netting were used for faunistic assessment only, since 
they give a great number of juvenile spiders that cannot 
be identified to species. The results of autumnal pitfall 
trapping were included in the general species list.
The alpha (Shannon index) and beta (Whittaker index) 
diversity of spider assemblages (i.e. the measures that take 
into account the number of individuals of each species in 
a sample) were estimated based on May–June pitfall trap 
catches at each study plot separately. Only adult spiders, 
in total 576 individuals, were identified and counted. De-
trended Correspondence Analysis (DCA) ordination of 
the study plots was also based on the species composition 
and activity density (ind./1 trap-day) of ground-dwelling 
spiders. Species recorded as singletons were excluded 
from the DCA procedure. All calculations were made in 
the program PAST, differences between the index values 
were estimated by means of permutation test. Statistical 
significance was set at p < 0.05. (Hammer et al. 2001).
Results
Vegetation and habitats
The vegetation communities described in our studies 
belong to eight associations, seven units, five orders and 
three classes (Dubyna et al. 2019). In terms of habitat 
diversity, they correspond to three higher taxa: Т – grassy 
habitats, ‘ Ч’ – shrubby habitats, and Ҡ – habitats of stony 
outcrops (Table 1).
Communities of the associations Stipo lessingianae-
Salvietum nutantis and Ephedro distachyae-Stipetum capil-
latae occupy vast areas on the gentle slopes of all gullies, 
while the meadow steppe association Elytrigio trichopho-
rae-Poetum angustifoliae was described only from the bot-
toms of Zelena and Khrystoforova gullies. An extrazonal 
meadow steppe association Thalictro mini-Spiraeetum hy-
pericifoliae is formed near the foothills with silicate out-
crops in mesic conditions. In our studies, this association 
was only found at the bottom of Chervona Gully. Steppe 
scrub (Amygdalo nanae-Spiraeetum hypericifoliae) is found 
as small patches in Chervona and Zelena gullies.
A total of 265 vascular plant species were recorded from 
11 habitats in the four study sites. The richest was the flo-
ra of Zelena Gully – 179 species: an association Amygdalo 
nanae-Spiraeetum hypericifoliae (habitat ZG2, Table 1) 
was represented by 107 species, Lino tenuifolii-Jurineetum 
brachycephalae (ZB4) by 99 species, Ephedro distachyae-
Stipetum capillatae (ZG1) by 91 species, and Elytrigio 
trichophorae-Poetum angustifoliae (ZG3) by 84 species. 
The floristic richness of Chervona Gully was slightly low-
er, 172 species, but there, only three habitats were investi-
gated: Stipo lessingianae-Salvietum nutantis (ChG1) – 116 
species, Amygdalo nanae-Spiraeetum hypericifoliae (ChG2) 
– 99 species, and Thalictro mini-Spiraeetum hypericifoliae 

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River basin (Central Ukraine) based on vegetation and spider research
(ChG3) – 63 species. The flora of Khrystoforova Gully, 
comprising 119 recorded species, was the poorest. It num-
bered 84 species in Stipo lessingianae-Salvietum nutantis 
association (KhG1), 48 species in Elytrigio trichophorae-
Poetum angustifoliae (KhG3), and only 15 species in Po-
tentillo incanae-Seselietum pallasii (KhG2). The associa-
tion Sempervivo ruthenicae-Sedetum ruprechtii on the Slate 
Rocks (SR) comprised 50 recorded species.
Cluster analysis based on the sintaxon floristic compo-
sition showed the highest similarity of the associations 
Stipo lessingianae-Salvietum nutantis in Chervona and 
Khrystoforova gullies (Sørensen ind. 75%) (Figure 7, 
habitats ChG1, KhG2). They form one cluster with the 
associations of forb-fescue feather grass, petrophytic, 
meadow steppes and the steppe scrub of Zelena Gully. 
Other associations do not create a separate group but 
consequently join the first one. The most different was 
the association Potentillo incanae-Seselietum pallasii on the 
granite substrate in Khrystoforova Gully with the very 
low similarity level of 12%.
Spiders
A total of 95 spider species from 20 families were record-
ed from the dry grassland habitats of the study area (Ap-
pendix, Table 2). Of these, 55 species were found from 
the stony slopes, 58 species from the forb-fescue-feather 
grass steppe, 51 species from the steppe scrub, and 47 spe-
cies from the meadow steppes at the gully bottoms. Spi-
der species richness was highest in the forb-fescue-feather 
grass steppe and steppe scrub (30–34 species in each lo-
cality), as well as in the meadow steppe in Zelena Gully 
(31 species). The most variable was the number of species 
on the stony slopes (from 12 to 24, Appendix, Table 2). 
The spring-summer spider fauna of Zelena Gully was the 
most species-rich, while that of Khrystoforova Gully was 
the poorest (62 and 48 species, respectively). In Chervona 
Gully, 55 species were found in May–June and only an 
additional nine species in autumn.
The spider faunistic similarity was highest at the 
two plots of forb-fescue-feather grass steppe (ChG1/
ZG1, K
s 
= 0.60) and lowest at the plots of stony slopes 
(K
s  
= 0.27–0.33) (Figure 8). A comparison of the spider 
species composition of various habitats within one site 
showed highest K
s
 values in Zelena Gully (ZG1/ZG2 – 
0.56, ZG1/ZG4 – 0.56, and ZG2/ZG3 – 0.58). At other 
sites, it ranged from 0.27 (KhG2/KhG3) to 0.52 (ChG1/
ChG2), being lowest in Khrystoforova Gully (Figure 8).
The stony slopes were the richest in the number of rare 
spider species (12 out of 19) while the meadow steppes 
were the poorest (3 species). Seven rare species were 
found in the forb-fescue-feather grass steppes and four 
species in the steppe scrub. Zelena Gully hosted ten rare 
species, Chervona Gully eight species, and Khrystoforova 
Gally only three species. Species distribution by ecologi-
cal groups was as follows: 7 generalists, 19 forest-grassland 
species, 9 forest-dry grassland species, 9 grassland species, 
29 dry grassland species, 19 steppe specialists and 5 spe-
cies with unclear preferences. The number of steppe spe-
cialists decreased from 11 in Chervona and Zelena gullies 
to four in the Khrystoforova one (Appendix, Table 2).
The ground-dwelling spider assemblage of the Slate 
Rocks was the most diverse, showing significant differenc-
es with the five studied assemblages out of ten (Figure 9). 
Among the five, there were spiders of the two forb-fescue-
feather grass steppe habitats and the two meadow ones, as 
well as spiders of petrophytic steppe in the Khrystoforova 
Gully. The Shannon indices of other spider assemblages 
did not differ so pronouncedly, still they were higher at 
both plots of steppe scrub and in the petrophytic steppe 
in Zelena Gully.
Beta-diversity (Whittaker index) of the ground-dwell-
ing spiders of the study localities decreased as follows: 
Figure 7: Dendrogram of floristic similarity of the vegetation of the 
study habitats (Sørensen index). For abbreviations, see Table 1.
Slika 7: Dendrogram floristične podobnosti vegetacije preučevanih 
habitatov (Sørensenov indeks). Za okrajšave glej Tabela 1.
The investigated sites host 23 vascular plant species that 
have been added to the Red Lists of various ranks or cat-
egorized as being under threat (Appendix, T able 1). In the 
Red Data Book of Ukraine (Didukh 2009), nine plant 
species recorded from the study area were assigned to the 
category vulnerable, two species – nearly threaten nine 
species – least concern, and two species of data deficiency. 
Zelena Gully was the richest in the number of protected 
species (18), ten species were recorded from Chervona 
Gully, eight species from Khrystoforova Gully and only 
three species from the Slate Rocks. In terms of habitat 
preferences, the distribution of rare species was as fol-
lows: Т1.2.1 – 0, Т1.2.2.‘ б’ – 9, Т1.3 – 4, Т1.4.a – 15,  
‘ Ч’4.2 – 5, and Ҡ1.3 – 2 species (Appendix, Table1).

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Assessment of the conservation value of dry grassland habitats in the Inhulets 
River basin (Central Ukraine) based on vegetation and spider research
 Zelena Gully (1.327), Chervona Gully (1.002), and 
Khrystoforova Gully (0.846). In terms of habitat pref-
erences, spiders of meadow steppes showed low beta di-
versity (K
W 
= 0.588) while in the other habitats, it was 
equally high (stony slopes – 1.100, steppe scrub – 1.071, 
forb-fescue-feather grass steppes – 1.069).
DCA ordination of the study habitats based on species 
and individual abundances of the ground-dwelling 
spiders showed a grouping of the forb-fescue-feather 
grass and petrophytic steppes of Chervona and Zelena 
gullies, a middle position of the steppe scrub and meadow 
steppes of the same gullies, and location of the three 
Khrystoforova Gully plots on the highest extremities of 
Axis 1 (Figure 10). The Slate Rocks and the granites in 
Khrystoforova Gully differed along Axis 2, being at the 
maximal distance from the meadows of Chervona and 
Zelena gullies.
Figure 8: Similarity of the spider species composition of various habi-
tats (Sørensen index values: 0.27–0.33 – green line, 0.36–0.42 – blue 
line, 0.49–0.52 – black line, 0.56–0.60 – red line. For abbreviations, 
see T able 1. 
Slika 8: Podobnost vrstne sestave pajkov v različnih habitatih (vrednos-
ti Sørensenovega indeksa: 0,27–0,33 – zelena črta, 0,36–0,42 – modra 
črta, 0,49–0,52 – črna črta, 0,56–0,60 – rdeča črta. Za okrajšave glej 
Tabela 1.
Figure 9: Species diversity of the ground-dwelling spider assemblages 
of the study plots (Shannon index). Whiskers indicate min and max 
bootstrap values. For abbreviations, see Table 1.
Slika 9: Vrstna pestrost združb pajkov, ki prebivajo na tleh na 
preučevanih ploskvah (Shannonov indeks). Ročaji predstavljajo mini-
malne in maksimalne vrednosti, dobljenih z zankanjem. Za okrajšave 
glej T abelo 1.
Figure 10: Detrended Correspondence Analysis ordination of the 
study habitats based on the ground-dwelling spider species composi-
tion and individual numbers. For abbreviations, see Table 1.
Slika 10: DCA ordinacija preučevanih habitatov na osnovi vrstne 
sestave talnih pajkov in števila osebkov. Za okrajšave glej Tabelo 1.
Discussion
Xerothermic habitats of the middle reaches of the Inhu-
lets River basin belong to the three main types: dry grassy 
habitats, deciduous shrubs and siliceous rocks. All of 
them are under protection of the Resolution 4 of the Bern 
Convention (Revised Annex I 2019). The habitats Т1.2, 
Т1.3, and Т1.4 correspond to the category E1.2 Perennial 
calcareous grasslands and basic steppes, the habitats ‘ Ч’ to 
F3.247 Ponto-Sarmatic deciduous thickets, and the habi-
tats ‘K’ to H3.1 Acid siliceous inland cliffs. In the frame 
of the Habitats Directive (Council Directive 1992), the 
investigated meadow, petrophytic and true steppes belong 

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River basin (Central Ukraine) based on vegetation and spider research
to the priority habitats 62C0* Ponto-Sarmatic steppes, 
the shrub steppes to 40A0* Subcontinental peri-Pannon-
ic scrub, and the siliceous outcrops to 8230 Siliceous rock 
with pioneer vegetation of the Sedo-Scleranthion or of the 
Sedo albi-Veronicion dillenii.
Comparison of floristic composition of the vegetation 
of the study habitats showed the highest similarity (75 
%) of the associations Stipo lessingianae-Salvietum nutan-
tis in Khrystoforova and Chervona gullies located at the 
north of the study area. In Zelena Gully, the vegetation of 
the same associations differed significantly due to the site 
specificity. In the case of steppe scrub with a dominance 
of Caragana frutex (L.) K. Koch (ass. Amygdalo nanae-
Spiraeetum hypericifoliae), the difference is determined by 
edaphic conditions: carbonate bedrocks in Zelena Gully 
and siliceous bedrocks in Chervona Gully. The associa-
tion Elytrigio trichophorae-Poetum angustifoliae, formed 
at the bottom of Zelena Gully, was rich in calcareous 
grassland species, while that of the Khrystoforova Gully 
included a number of ruderal species appeared due to hu-
man economic activity.
The extremely poor flora of Potentillo incanae-Sese-
lietum pallasii association (15 species) can be explained 
by the specificity of its habitat. This vegetation is formed 
on shallow eluvial sediments on the surface in the cracks 
of granite boulders. Compared with the vegetation of 
granite bedrocks, the communities of Slate Rocks are 
richer due to more favourable edaphic conditions (50 
species).
The spider fauna of the study sites is characteristic of 
xerothermic habitats. Dry grassland and steppe species 
comprise together 50% of the fauna; open forest–dry 
grassland inhabitants add a further 9%. The two stud-
ied gullies, Chervona and Zelena, are well preserved and 
have a high conservation value. The local communities 
can serve as a pattern of spider species composition and 
habitat preferences in the south of the forb-fescue-feather 
grass steppe subzone (Polchaninova 2012). Khrystoforova 
Gully is highly disturbed by continuous grazing and un-
controlled fires. The spider community responds to these 
impacts with a decrease in the total species richness and 
the number of rare species and steppe specialists. In this 
Figure 11: Gymnospermium odessanum and Allium lineare on the Slate Rocks. Photo: Olga Krasova.
Slika 11: Gymnospermium odessanum in Allium lineare na skrilastem skalovju. Foto: Olga Krasova.

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River basin (Central Ukraine) based on vegetation and spider research
gully, spider assemblages of the same habitat type differ 
from those in Zelena and Chervona gullies (Figure 8).
At the same time, frequent fires that destroy a thick lit-
ter layer seem to enable a high number and abundance in 
flowering early-spring plant species (Adonis wolgensis, A. 
vernalis, Bulbocodium versicolor). Previously we observed 
a similar pattern in Zelena Gully: a comparison of geo-
botanical descriptions taken before and after the fire con-
firmed an increase of the early-blossoming plant abun-
dance at the fire site. Khrystoforova Gully hosts eight 
protected plant species.
Spider research revealed 19 rare species (Appendix, 
Table 2). For Harpactea azovensis and Zelotes eugeni, this 
is the most northwestern known locality; these species’ 
ranges stretch from the Dnipropetrovska Oblast and 
Crimea of Ukraine to Rostov-on-Don Oblast of Russia 
(Kovblyk & Kastrygina 2015). For Dysdera hungarica, 
this is the northernmost locality on the East European 
Plain. Gnaphosa opaca has been recorded in Ukraine from 
the Dnipropetrovska and Mykolaivska (NP unpublished) 
oblasts only. Micaria bosmansi has not been found west of 
the Dnipropetrovska Oblast. Euryopis saukea is a North 
American invasive species, now patchily distributed in 
the Palearctic nemoral belt. The other 13 species in this 
list are regionally rare. Mainly they are common inhabit-
ants of the dry grasslands and/or open forests in Southern 
Ukraine and in the South East of European Russia and 
spread north via azonal habitats (Polchaninova & Proko-
penko 2013, Ponomarev 2017).
In the basin of the Inhulets middle reaches, the largest 
number of rare plant and spider species are associated 
with the forb-fescue-feather grass and petrophytic 
calcareous steppes. Zonal forb-fescue-feather grass 
steppes cover vast areas at all study sites and host 15 rare 
plant species and seven spider species. In the petrophytic 
steppes, which are patchy and smaller, the proportion 
is inverse, with more spider species (11) and less plant 
species (9).
Stony slopes of various bedrocks have long been 
known as hotspots of regionally rare and specialist 
species (Sobko 1972, Buchar & Růžička 2002) even if 
they are of anthropogenic origin and/or experience high 
recreation pressure (Řezáč et al. 2015). Calcareous and 
crystalline outcrops in the East European Plain create 
mountain-like conditions, which favor the formation of 
a biota markedly different from that of the surrounding 
zonal landscapes and contribute to the spread of southern 
species to the north (Kontar 2001, Polchaninova & 
Prokonenko 2013). In our study, the Slate Rocks and 
the limestone slopes in Zelena Gully hosted the highest 
proportion of rare spiders (17% and 29% respectively). 
Moreover, despite the small area and high recreation 
pressure, the Slate Rocks preserve populations of a 
plethora of protected plants. They are a refuge for a 
narrow-range endemic species of the crystalline outcrops 
of the south of the Dnieper Upland, Sedum borissovae 
Balk. (Sobko 1972), and the northernmost detached 
locality of Gymnospermium odessanum (Figure 11). The 
narrow broken strips of the forb-fescue-feather grass 
steppe on the rocks’ flattened upper part are a habitat of 
Tulipa hypanica and Stipa ucrainica. Among the species 
of the genus Stipa of the plain Ukraine, S. ucrainica is 
one of the most sensitive to anthropogenic impacts. In 
the area in hand, its assemblages were recorded in small 
fragments only. 
Interestingly, the spider species richness in various 
habitats followed floristic richness, being higher in the 
forb-fescue-feather grass steppe and steppe scrub and 
lower in the meadow steppe. Both were the poorest on 
the granite and slate outcrops, while the petrophytic 
steppe on limestone soil was rich in plant but poor in 
spider species.
The valley of the Inhulets middle reaches, along with 
the gullies flowing into it, was added to the “Shadow 
list” (NGO lobby for new conservation areas) and then 
to the list of the Emerald Network sites of Ukraine (No. 
UA0000310, Updated list 2019). Unfortunately, only 
Zelena Gully has got the “Emerald status”, since its flora 
has a high number of protected plant species (18). There 
are endemic species of calcareous steppes that determine 
uniqueness of this site. To our mind, steppe gullies and 
crystalline rocks of the Saksahan River basin, the Inhulets 
largest tributary, are also sites of conservation concern. 
Given coenotic and habitat diversity of the Slate Rocks 
and Chervona Gully, located in its valley, this area re-
quires protection in accordance with European standards. 
If it receives Emerald status, it may be excluded from the 
plans of mining expansion, which is currently the greatest 
threat to natural habitats in the study area. Moreover, this 
status would give legal grounds to control recreation and 
grazing intensity and recommend prescribed burning to 
help avoid uncontrolled fires.
Conclusion
Botanical and arachnological evaluations of priority areas 
complement each other and make the assessment more 
profound. Rich vegetation and spider communities with 
rare and specialist species in Zelena Gully confirm its 
high conservation status. The Slate Rocks and Chervona 
Gully are recommended to be included into the Emerald 
Network, since they host well-preserved natural habitats 
listed in the Resolution 4 of the Bern Convention as well 
as a number of protected and rare species.

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Assessment of the conservation value of dry grassland habitats in the Inhulets 
River basin (Central Ukraine) based on vegetation and spider research
Acknowledgements
The authors are thankful to Dr. Stephen Venn as the 
 Coordinating Editor and the two anonymous Reviewers 
for the valuable comments on the initial draft. The lin-
guistic proofreading made by Dr. Stephen Venn is also 
deeply acknowledged.
Nina Polchaninova  https://orcid.org/0000-0003-4605-8788
Olga Krasova  https://orcid.org/0000-0003-3035-5614
Liudmyla Lysohor  https://orcid.org/0000-0002-1949-1394
Tatiana Atemasova  https://orcid.org/0000-0002-7527-5143
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River basin (Central Ukraine) based on vegetation and spider research
Species
A(i)
A(iii)
A(iv)
IUCN RL
RDBU
Sites and habitats
Adonis vernalis L. . . . . NT ChG (1.1–1.3); KhG (3.1); ZG (4.1–4.4)
Adonis wolgensis Stevenex DC. . . . . NT ChG (1.1–1.3); KhG (3.1);  ZG (4.1–4.3)
Allium lineare L. . . . . LC SR (K1.3)
Astragalus henningii (Steven) Klokov . . . . VU ZG (T1.4.a)
Astragalus dasyanthus Pall. . . . . DD ChG (1.1, 1.2); KhG (3.1)
Astragalus odessanus Besser . . . . VU ZG (T1.4.a)
Bulbocodium versicolor (Ker-Gawl.) Spreng. . . . . LC KhG (3.1); ZG (4.1)
Caragana scythica Pojark. . . + . VU ZG (4.1)
Cytisus graniticus Rehmann . + . . VU ZG (4.3)
Crocus reticulatus Steven ex Adams . . . . LC KhG (3.1); ZG (4.1)
Elymus stipifolius (T rautv.) Melderis + LC LC ZG (4.3)
Eremogone cephalotes (Bieb.) Fenzl . + . . VU ZG (4.1)
Genista scythica Pacz. . + . . LC ZG (4.3)
Gymnospermium odessanum (DC.) Takht. . . + . VU SR (K1.3)
Iris pontica Zapal. . . + . VU ZG (T1.4.a)
Ornithogalum boucheanum (Kunth) Asch. . . . . LC ChG (T1.3.2)
Pulsatilla pratensis (L.) Mill. . . . . LC ChG (1.3); KhG (3.1); ZG (4.3)
Stipa asperella Klokov et Ossycznjuk . . . . DD ZG (4.3)
Stipa capillata L. . . . . LC ChG (1.1, 1.2); KhG (3.1); ZG (4.1–4.3)
Stipa lessingiana T rin. etRupr. + . . LC LC ChG (1.1, 1.2); KhG (3.1); ZG (4.1–4.3)
Stipa pulcherrima K. Koch . . . . VU ChG (T1.4.a)
Stipa ucrainica P . Smirn. . . . . LC ChG (T1.4.a), SR (T1.4a); ZG (T1.4.a)
Tulipa hypanica Klokov et Zoz . + . . VU ChG (T1.4.a); SR (T1.4a); ZG (T1.2.2.‘ б’)
Total species 2 4 3 2 23
Appendix
Table 1: Rare vascular plant species at the study sites / habitats in the basin of the Inhulets middle reaches 
Tabela 1: Redke rastlinske vrste na preučevanih rastiščih/habitatih v srednjem teku reke Inhulets.
Abbreviations: A(i) – species with categories CR and VU in the IUCN database, version 2020-2; A(iii) – species with catego-
ries “endangered” and “vulnerable” in the Red Data Book of Ukraine (Didukh 2009) that are considered national endemics of 
Ukraine; A(iv) –  limited range species with categories “endangered” and “vulnerable” in the Red Data Book of Ukraine that are 
not national endemics; IUCN RL – species included in the IUCN Red List of threatened species (IUCN version 2020); RDBU 
– species listed in the Red Data Book of Ukraine. Categories: DD – data deficient, i.e. insufficient information for a proper assess-
ment of conservation status to be made; LC – least concern, unlikely to become extinct in the near future; NT ‒ near threatened, 
close to being at high risk of extinction in the near future; VU –vulnerable, likely to become endangered unless the circumstances 
that are threatening their survival and reproduction improve.
Sites and habitats. SR – Kryvyi Rih Town, Slate Rocks; ChG – Kryvyi Rih Town, Chervona Gully: ChG1 – forb-fescue-
feather grass steppe, ChG2 – steppe scrubs, ChG3 – meadow steppe, ChGa – autumnal collection in various habitats; 
KhG – Kudasheve Vil., Khrystoforova Gully:  KhG1 ‒ forb- fescue-feather grass steppe, KhG2 ‒ petrophytic steppe on granite 
bedrock, KhG3 ‒  meadow steppe; ZG – Poltavka Vil., Zelena Gully: ZG1 ‒ forb- fescue-feather grass steppe, ZG2 ‒ steppe 
scrubs, ZG3 ‒ meadow steppe, ZG4 ‒ petrophytic steppe on limestone bedrock. (Figures 1, 2). If a species was not found at the 
sample plots marked on the map, we indicated the type of habitat at the study site where it was recorded.

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Assessment of the conservation value of dry grassland habitats in the Inhulets 
River basin (Central Ukraine) based on vegetation and spider research
Families/species
R
HP
Localities/habitats
SR ChG1 ChG2 ChG3 ChGa KhG1 KhG2 KhG3 ZG1 ZG2 ZG3 ZG4
Fam. Agelenidae (2)
Eratigena agrestis (Walckenaer, 1802) f/dgr . . . . 9 . . . . . . .
T egenaria lapicidinarum Spassky, 1934 f/gr . . . . . . 1 . . . . .
Fam. Araneidae (9)
Agalenatea redii (Scopoli, 1763) gr 2 2 1 1 2 . . 5 2 . 1
Araneus quadratus Clerck, 1757 mgr . . 2 3 5 . . 5 . 3 .
Argiope bruennichi (Scopoli, 1772) gr . 1 1 3 8 . . 4 . 2 3 .
Cyclosa oculata (Walckenaer, 1802) f/dgr . 1 . . 3 . . 1 . . .
Gibbaranea bituberculata (Walckenaer, 1802) f/gr 1 . 6 1 . . . 2 3 2 . .
Larinioides cornutus (Clerck, 1757) mgr . . . . . . . 2 . . ..
L. suspicax   (O. Pickard-Cambridge, 1876) mgr . . . . . . . . . 2 2 .
Mangora acalypha (Walckenaer, 1802) g 3 1 6 8 . 2 1 6 5 4 9 .
Neoscona adianta (Walckenaer, 1802) dgr . 2 . 2 . 6 . . 3 1 . .
Fam. Cheiracanthiidae (1)
Cheiracanthium pennyi O. Pickard-Cambridge, 1873 dgr . 1 1 . . . . . . . . .
Fam. Dictynidae (3)
Brigittea latens (Fabricus, 1775) st . 1 . . . . . . 1 . . 2
Dictyna arundinacea (Linnaeus, 1758) f/gr . . 2 . . . . . . 1 .
Lathys stigmatisata (Menge, 1869)
r
f/dgr
1 . . . . . . . . . . .
Fam. Dysderidae (3)
Dysdera hungarica Kulczyński, 1897
r
un
. . . . . . . . . . . 1
Harpactea azovensis Charitonov, 1956*
r
f/dgr
4 . . . . 1 1 . . . . .
Harpactea rubicunda (C. L. Koch, 1838)* f/gr 1 . 1 . . . . . . . . .
Fam. Eresidae (1)
Eresus kollari Rossi, 1846 dgr . . 1 . 21 . . . . . . .
Fam. Gnaphosidae (19)
Berlandina cinerea (Menge, 1872)* dgr 6 1 1 . . . . . 7 2 1 3
Civizelotes caucasius (L. Koch, 1866)
lr
dgr
. 1 . . 1 . . . . . . .
Table 2: Spider species composition and individual numbers at the study sites / habitats in the basin of the Inhulets middle reaches. May‒June and September 2017
Tabela 2: Vrstna sestava in število osebkov pajkov. Maj-junij in september 2017.

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River basin (Central Ukraine) based on vegetation and spider research
Families/species
R
HP
Localities/habitats
SR ChG1 ChG2 ChG3 ChGa KhG1 KhG2 KhG3 ZG1 ZG2 ZG3 ZG4
Drassodes lapidosus (Walckenaer, 1802)*
lr
f/dgr
. . . . . . . . . 1 . 1
Drassyllus praeficus (L. Koch, 1866)* f/dgr 1 1 1 4 . 5 1 . . 1 1 1
D. pusillus (C. L. Koch, 1833)* gr . . . . . . . . . . 2 .
D. vinealis (Kulczyn'ski, 1897) st . . . . . . . . 1 . . .
Gnaphosa lucifuga (Walckenaer, 1802) st . . . . . . 1 . . . . .
G. opaca Herman, 1897*  r st 5 1 . . . . . . . 3 1
G. taurica Thorell, 1875* lr st 2 2 . . . . . . . ..
Haplodrassus bohemicus Miller & Buchar, 1977* f/gr 4 . . 1 . 22 1 14 1 6 11 3
H. dalmatensis (L. Koch, 1866)* lr st 1 1 . . . . . . 1 . . .
H. kulczynskii Lohmander, 1942* dgr 1 1 . . . . . . . 1 . .
H. minor (O. Pickard-Cambridge, 1879)*
r
st
. . . . . . . . . . 3 .
H. signifer( C. L. Koch, 1839)* f/gr . . . . . . . . 1 . . 1
Micaria bosmansi Kovblyuk & Nadolny, 2008*
r
un
. . . 2 . . . . . . . .
Nomisia aussereri (L. Koch, 1872):
lr
st
. . . . 1 . . . . . . .
Z. electus (C. L. Koch, 1839)* f/gr 1 . 2 . . 3 1 . . 4 3 .
Z. eugenei Kovblyuk 2009* r un . . . . . 1 1 . . . . 1
Z. longipes (L. Koch, 1866) dgr . . . . 11 . . . . . . .
Z. segrex (Simon, 1878):* r st 1 . . . . . . . . . . 1
Fam. Linyphiidae (4)
Agyneta rurestris (C. L. Koch, 1836); g . . . . . 1 . . . . . .
Ipa terrenus (L. Koch, 1879) r st . . . . . . . . 1 . . .
Linyphia tenuipalpis Simon, 1884 f/gr . . . . . 7 . . . . . .
Stemonyphantes lineatus (Linnaeus, 1758)* dgr . 1 . . . . . . . 2 . .
Fam. Liocranidae (2)
Agroeca cuprea Menge, 1873* dgr . . . . 4 . . . . 1 7 .
Agroeca lusatica (L. Koch,1875)* f/gr . . . . . 1 . 1 . . . .
Fam. Lycosidae (11)
Alopecosa cuneata (Clerck,1757)* gr . . 2 1 . . . . . . . .
A. cursor (Hahn, 1831)* st 11 17 7 . . 4 . . 18 15 12 31
A. farinosa (Herman, 1879)* gr . . . . . 5 . 3 . . 1 .
A. kovblyuki Nadolny & Ponomarev, 2012* dgr . . . . 2 1 3 2 . . . .
A. pulverulenta (Clerck, 1757)* f/gr 2 10 1 . . . 6 . 8 32 .
A. schmidti (Hahn, 1835) dgr . . . . . . . . . 1 . .
A. solitaria (Herman, 1879) dgr 1 . . 7 . . . . . . .
Geolycosa vultuosa (C. L. Koch, 1838)
r
un
. . . . 1 . . . . . . .
Pardosa agrestis (Westring, 1861) gr . . . . . . . 1 . . . .
Trochosa robusta (Simon, 1876)* f/gr 1 2 . . 1 5 . . 3 .
Xerolycosa miniata (C.L. Koch, 1834)* dgr 1 . . 2 . 16 . 13 . 1 . .
Fam. Miturgidae (1)
Zora pardalis* dgr . . . 1 1 . . 1 . . 2 .
Fam. Oxyoidae (2)
Oxyopes heterophthalmus (Latreille, 1804) dgr . 2 . . . . . . 5 1 . 2
O. lineatus Latreille, 1806 f/gr . 2 . . 4 . . . 6 . . 2
Fam. Philodromidae (4)
Philodromus cespitum (Walckenaer, 1802) g . . 2 . . . . . . . . .
Rhyzodromus histrio (Latreille, 1819) . .. . . . 1 . . . . . 1 .
Thanatus arenarius L. Koch, 1872* dgr . 8 4 13 . 9 1 4 13 17 4 4
Tibellus oblongus (Walckenaer, 1802) f/gr . . . 1 . . . 1 1 . . .
Fam. Pisauridae (1)
Pisaura novicia (L. Koch, 1878) f/gr . . . . . . . . 1 . 1 .
Fam. Salticidae (11)
Aelurillus v-insignitus (Clerck, 1757)* dgr 4 . . . . 1 . . . 1 . .
Asianellus festivus (C. L. Koch, 1834)* dgr . 1 . . . 1 . . 3 . 1 .
Euophrys frontalis (Walckenaer, 1802)* dgr . . . . . . . . . 1 1 .
Evarcha arcuata (Clerck, 1757) f/gr . . 1 1 . . . . . . . .
Heliophanus cupreus (Walckenaer, 1802) g . . 1 1 . . . . 1 1 1 .
H. flavipes (Hahn, 1832) gr 1 2 1 . . . . 1 4 3 . .
H. lineiventris Simon, 1868 lr st . . . . . . . . . . . 1
Pellenes nigrociliatus (Simon 1875) r st . . 1 . . . . . . . . 1
Philaeus chrysops (Poda, 1761) st 1 2 . 8 . . . . . . .
Sibianor aurocinctus (Ohlert, 1865) dgr . . . . . 1 . . . . . .
T alavera aequipes O. Pickard-Cambridge, 1871) st 1 . . . . . . . . . . .
Fam. T etragnathidae (1)
Pachygnatha degeeri Sundevall, 1830 f/gr . . . . . . . . . . 1 .
Fam. Theridiidae (6)
Crustulina guttata (Wider, 1834) g . . 1 . . . . . . . . .
Enoplognatha thoracica (Hahn, 1833) dgr . . 1 . . . . . . . . .

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River basin (Central Ukraine) based on vegetation and spider research
Families/species
R
HP
Localities/habitats
SR ChG1 ChG2 ChG3 ChGa KhG1 KhG2 KhG3 ZG1 ZG2 ZG3 ZG4
Drassodes lapidosus (Walckenaer, 1802)*
lr
f/dgr
. . . . . . . . . 1 . 1
Drassyllus praeficus (L. Koch, 1866)* f/dgr 1 1 1 4 . 5 1 . . 1 1 1
D. pusillus (C. L. Koch, 1833)* gr . . . . . . . . . . 2 .
D. vinealis (Kulczyn'ski, 1897) st . . . . . . . . 1 . . .
Gnaphosa lucifuga (Walckenaer, 1802) st . . . . . . 1 . . . . .
G. opaca Herman, 1897*  r st 5 1 . . . . . . . 3 1
G. taurica Thorell, 1875* lr st 2 2 . . . . . . . ..
Haplodrassus bohemicus Miller & Buchar, 1977* f/gr 4 . . 1 . 22 1 14 1 6 11 3
H. dalmatensis (L. Koch, 1866)* lr st 1 1 . . . . . . 1 . . .
H. kulczynskii Lohmander, 1942* dgr 1 1 . . . . . . . 1 . .
H. minor (O. Pickard-Cambridge, 1879)*
r
st
. . . . . . . . . . 3 .
H. signifer( C. L. Koch, 1839)* f/gr . . . . . . . . 1 . . 1
Micaria bosmansi Kovblyuk & Nadolny, 2008*
r
un
. . . 2 . . . . . . . .
Nomisia aussereri (L. Koch, 1872):
lr
st
. . . . 1 . . . . . . .
Z. electus (C. L. Koch, 1839)* f/gr 1 . 2 . . 3 1 . . 4 3 .
Z. eugenei Kovblyuk 2009* r un . . . . . 1 1 . . . . 1
Z. longipes (L. Koch, 1866) dgr . . . . 11 . . . . . . .
Z. segrex (Simon, 1878):* r st 1 . . . . . . . . . . 1
Fam. Linyphiidae (4)
Agyneta rurestris (C. L. Koch, 1836); g . . . . . 1 . . . . . .
Ipa terrenus (L. Koch, 1879) r st . . . . . . . . 1 . . .
Linyphia tenuipalpis Simon, 1884 f/gr . . . . . 7 . . . . . .
Stemonyphantes lineatus (Linnaeus, 1758)* dgr . 1 . . . . . . . 2 . .
Fam. Liocranidae (2)
Agroeca cuprea Menge, 1873* dgr . . . . 4 . . . . 1 7 .
Agroeca lusatica (L. Koch,1875)* f/gr . . . . . 1 . 1 . . . .
Fam. Lycosidae (11)
Alopecosa cuneata (Clerck,1757)* gr . . 2 1 . . . . . . . .
A. cursor (Hahn, 1831)* st 11 17 7 . . 4 . . 18 15 12 31
A. farinosa (Herman, 1879)* gr . . . . . 5 . 3 . . 1 .
A. kovblyuki Nadolny & Ponomarev, 2012* dgr . . . . 2 1 3 2 . . . .
A. pulverulenta (Clerck, 1757)* f/gr 2 10 1 . . . 6 . 8 32 .
A. schmidti (Hahn, 1835) dgr . . . . . . . . . 1 . .
A. solitaria (Herman, 1879) dgr 1 . . 7 . . . . . . .
Geolycosa vultuosa (C. L. Koch, 1838)
r
un
. . . . 1 . . . . . . .
Pardosa agrestis (Westring, 1861) gr . . . . . . . 1 . . . .
Trochosa robusta (Simon, 1876)* f/gr 1 2 . . 1 5 . . 3 .
Xerolycosa miniata (C.L. Koch, 1834)* dgr 1 . . 2 . 16 . 13 . 1 . .
Fam. Miturgidae (1)
Zora pardalis* dgr . . . 1 1 . . 1 . . 2 .
Fam. Oxyoidae (2)
Oxyopes heterophthalmus (Latreille, 1804) dgr . 2 . . . . . . 5 1 . 2
O. lineatus Latreille, 1806 f/gr . 2 . . 4 . . . 6 . . 2
Fam. Philodromidae (4)
Philodromus cespitum (Walckenaer, 1802) g . . 2 . . . . . . . . .
Rhyzodromus histrio (Latreille, 1819) . .. . . . 1 . . . . . 1 .
Thanatus arenarius L. Koch, 1872* dgr . 8 4 13 . 9 1 4 13 17 4 4
Tibellus oblongus (Walckenaer, 1802) f/gr . . . 1 . . . 1 1 . . .
Fam. Pisauridae (1)
Pisaura novicia (L. Koch, 1878) f/gr . . . . . . . . 1 . 1 .
Fam. Salticidae (11)
Aelurillus v-insignitus (Clerck, 1757)* dgr 4 . . . . 1 . . . 1 . .
Asianellus festivus (C. L. Koch, 1834)* dgr . 1 . . . 1 . . 3 . 1 .
Euophrys frontalis (Walckenaer, 1802)* dgr . . . . . . . . . 1 1 .
Evarcha arcuata (Clerck, 1757) f/gr . . 1 1 . . . . . . . .
Heliophanus cupreus (Walckenaer, 1802) g . . 1 1 . . . . 1 1 1 .
H. flavipes (Hahn, 1832) gr 1 2 1 . . . . 1 4 3 . .
H. lineiventris Simon, 1868 lr st . . . . . . . . . . . 1
Pellenes nigrociliatus (Simon 1875) r st . . 1 . . . . . . . . 1
Philaeus chrysops (Poda, 1761) st 1 2 . 8 . . . . . . .
Sibianor aurocinctus (Ohlert, 1865) dgr . . . . . 1 . . . . . .
T alavera aequipes O. Pickard-Cambridge, 1871) st 1 . . . . . . . . . . .
Fam. T etragnathidae (1)
Pachygnatha degeeri Sundevall, 1830 f/gr . . . . . . . . . . 1 .
Fam. Theridiidae (6)
Crustulina guttata (Wider, 1834) g . . 1 . . . . . . . . .
Enoplognatha thoracica (Hahn, 1833) dgr . . 1 . . . . . . . . .

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River basin (Central Ukraine) based on vegetation and spider research
Families/species
R
HP
Localities/habitats
SR ChG1 ChG2 ChG3 ChGa KhG1 KhG2 KhG3 ZG1 ZG2 ZG3 ZG4
Euryopis saukea Levi, 1951 r f/dgr . . . . . 1 . . . . . .
Phylloneta impressa (L. Koch, 1881) dgr 1 1 . 1 . . . . 1 1 . .
Robertus arundineti (O. Pickard-Cambridge, 1871) gr . . . . . . . . . 1 . .
Simitidion simile (C. L. Koch, 1836) dgr . . 3 1 . . . . 2 4 . .
Fam. Thomisidae (10)
Heriaeus oblongus Simon, 1918 st . . . . . 1 . . 1 . . 1
Misumena vatia (Clerck, 1757) f/gr . . 1 1 . . . . 2 1 2 .
Ozyptila scabricula (Westring, 1851)* dgr . . . 1 . . . . . 3 1 .
Runcinia grammica (C. L. Koch, 1837) st . . . 1 1 . . . 5 3 1 2
Spiracme striatipes (L. Koch, 1870): dgr . . . . 2 . . . . . . .
Thomisus onustus Walckenaer, 1805 dgr 1 5 1 . . 2 1 . 4 3 . 6
Xysticus acerbus Thorell, 1872* dgr 1 6 4 . 2 . . 2 1 2 1
X. cristatus (Clerck, 1757)* f/gr . . . . . 1 . . . 2 1 .
X. kochi Thorell, 1872* f/dgr 8 4 2 7 . 2 . 3 4 3 2 3
Xysticus marmoratus Thorell, 1875 st . 1 . . 6 . 1 . . . . .
Fam. Titanoecidae (1)
Titanoeca veteranica Herman, 1879* lr st 6 3 1 . . . . . . . . .
Fam. Uloboridae (1)
Uloborus walckenaerius Latreille, 1806 dgr . 2 . . . . . . 4 . . 3
Fam. Zodariidae (1)
Zodarion thoni Nosek, 1905 f/dgr . . . . . . . . . . . 1
Total species 24 30 32 25 19 25 12 18 30 34 31 24
Abbreviations: R ‒ rarity: r – rare, lr ‒ locally rare, wide-spread species are not mentioned; HP – habitat preference (after Polchaninova & Prokopenko 2013; Ponomarev 2017 
and personal data): g – generalist, f/gr – open forests and grasslands, f/dgr – open forests and dry grasslands, gr –grasslands of various types, mgr – mesic and/or wet grasslands, 
dgr ‒ dry grasslands, st – steppe specialist, un – unspecified. * - species used in the DCA procedure. For the site/habitat code, see Table 1.