85
Syntaxonomy of chalk outcrop vegetation 
of the order Thymo cretacei-Hyssopetalia 
cretacei
Abstract
The order Thymo cretacei-Hyssopetalia cretacei Didukh 1989 combines chalk 
outcrop plant communities of the southwestern Central Russian Upland. Its 
specificity can be attributed to a rather peculiar and heterogeneous flora with a 
large number of endemic species. The question about its origin has caused a lively 
discussion, which has been going on since the late nineteenth century. Since 1989 
works on the classification of these communities have frequently been carried 
out, but until today no unanimous decision could be reached. The purpose of 
our research was to conduct a critical analysis of the syntaxonomical structure 
of Cretaceous outcrop vegetation and to show its difference from the steppe 
vegetation of the class Festuco-Brometea Br.-Bl. et Tx. ex Soó 1947. The territory 
of our research covers the southwestern foothills of the Central Russian Upland 
and the Donetsk Range, located only within the steppe zone and characterised by 
Cenomanian chalk outcrops. In total 354 relevés were used for the analysis. The 
modified TWINSPAN classification was used for the analysis. Our research has 
shown that the order Thymo cretacei-Hyssopetalia cretacei includes twelve associa-
tions belonging to three alliances: Artemisio hololeucae-Hyssopion cretacei Romash-
chenko et al. 1996, Euphorbio cretophilae-Thymion cretacei Didukh 1989 and 
Centaureo carbonatae-Koelerion talievii Romashchenko et al. 1996.
Izvleček
V red Thymo cretacei-Hyssopetalia cretacei Didukh 1989 združujemo rastlinske 
združbe, ki jih najdemo na krednih izdankih v jugozahodnem Srednjeruskem 
višavju. Njegova specifičnost je posledica posebne in heterogene flore z velikim 
številom endemitov. Njen izvor je povzročil živahno diskusijo, ki traja že od konca 
19. stoletja. Številne raziskave o klasifikaciji teh združb so izvajali od leta 1989, 
vendar do danes ni prišlo do soglasnega pojasnila. Namen naše raziskave je naredi-
ti kritično sintaksonomsko analizo vegetacije krednih izdankov in prikazati razlike 
od stepske vegetacije razreda Festuco-Brometea Br.-Bl. et Tx. ex Soó 1947. Obmo-
čje raziskav obsega jugozahodna vznožja Srednjeruskega višavja in gorske verige 
Doneck, ki se nahaja znotraj stepske cone na cenomanijskih krednih izdankih. 
V analizi smo uporabili 354 vegetacijskih popisov in modificirano TWINSPAN 
klasifikacijsko metodo. Pokazali smo, da v red Thymo cretacei-Hyssopetalia cretacei 
uvrščamo 12 asociacij, ki pripadajo trem zvezam: Artemisio hololeucae-Hyssopion 
cretacei Romashchenko et al. 1996, Euphorbio cretophilae-Thymion cretacei Didukh 
1989 in Centaureo carbonatae-Koelerion talievii Romashchenko et al. 1996.
Keywords: Central Russian Upland, 
Hyssopus flora, lowered alpine plants 
classification. 
Ključne besede: Srednjerusko 
višavje, flora rodu Hyssopus, 
klasifikacija nizkih alpinskih rastlin.
Received: 14. 3. 2017
Revision received: 14. 7. 2017
Accepted: 10. 11. 2017
Co-ordinating Editor:  
Orsolya Valkó
1 M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, 2 Tereschenkivska St., 01601 Kyiv, Ukraine. E-mail: olgachusova@gmail.com
2 Karachay-Circassian State University, 29 Lenina St., 369200 Karachaevsk, Russia. E-mail: ondemina@yandex.ru
* Corresponding author
Yakiv Didukh1, Olga Chusova1,* & Olga Demina2
DOI: 10.1515/hacq-2017-001317/1 • 2018, 85–109
Y. Didukh, O. Chusova & O. Demina
Syntaxonomy of chalk outcrop vegetation of the order  
Thymo cretacei-Hyssopetalia cretacei
86
17/1 • 2018, 85–109
Introduction
The specificity of chalk outcrop flora and vegetation has 
been noted by botanists since the late nineteenth centu-
ry and caused a heated debate about its origins (Tsinger 
1885; Litvinov 1891, 1902, Taliev 1897, 1904, 1905, 
Pachos’kyy 1910, Kozo-Polyansky 1910, 1925, 1931, 
Szafer 1928, Kleopov 1930, Lavrenko 1932, Grosset 
1935, Hryn’ 1938). While Taliev (1905) regarded creta-
ceous flora as young and associated with human activi-
ties, most scientists accept the theory according to which 
its main core is relict and had formed long before the ice 
age. Because of the large number of endemic species con-
fined to Cretaceous outcrops, this flora was called “Hys-
sopus flora” (after its typical element Hyssopus cretaceus 
Dubj) (Kozo-Polyansky 1931). Also, Kozo-Polyansky 
showed that the flora of Cretaceous outcrops is hetero-
geneous and includes another component besides “Hys-
sopus flora”, which he called “lowered alpine plants” (the 
unit of periglacial Pleistocene steppe). Typical represent-
atives of this complex are Carex pediformis C. A. Mey., 
Carex humilis Leyss., Clausia aprica (Stephan) Trotsky, 
Alyssum lenense Adams and others. The core of this flora 
is formed by specific xerophytic endemic surface plants 
and squat plants confined to dry chalky substrates. 
They are characterised by grey stellate and weblike pu-
bescence or waxy coating (Hyssopus cretaceus, Linaria 
cretacea Spreng., Scrophularia cretacea Spreng., Silene 
cretacea Spreng), which shows the processes of specia-
tion in such conditions. Typical components of the flora 
are mountainous Mediterranean species, whose nearest 
occurence is in the Crimean Mountains (Euphorbia pe-
trophila C. A. Mey., Asperula tephrocarpa (incl. A. creta-
cea, A. supina), Silene supina M. Bieb., Polygala mediter-
ranea, Hedysarum ucrainicum (incl. H. tauricum)), and 
species of broader South European distribution (Heli-
anthemum canum s.l. (L.) Hornem., Jurinea stoechadifo-
lia (M. Bieb.) DC., Pimpinella tragium ssp. titanophylla 
(Woronow) Tutin, Linum czernjaevii Klok., Taraxacum 
serotinum (Waldst. & Kit.) Fisch., Scutellaria supina L.). 
On the other hand, there are Pontic-Siberian steppe 
species (Artemisia salsoloides Willd., Psephellus marschal-
lianus (Spreng.) K. Koch, Gypsophila altissima (incl. 
G. oligosperma), Cephalaria uralensis (Murray) Roem. & 
Schult., Astragalus albicaulis DC., Clausia aprica (Ste-
phan) Korn.-Trotzky, Hedysarum grandiflorum Pall., 
Alyssum lenense Adams, Androsace villosa (incl. A. kozo-
poljanskyi), Polygala sibirica L.) and even Central Asian 
species (Atrophaxis frutescens (L.) K. Koch, Artemisia 
salsoloides, Krascheninnikovia ceratoides (L.) Gueldenst.). 
Thus, the floristic composition of Cretaceous outcrops 
is rather peculiar and heterogeneous, and it vastly dif-
fers from zonal steppe vegetation. As for taxonomy, it is 
characterised by the presence of endemic races, which 
sometimes get species status (Dobrochayeva et al.1987, 
Klokov & Dobrochayeva 1974, Cvelev 1996).
The vegetation classification of the dominant basis 
is reflected in the works of Semenova-Tian-Shanskaya 
(1954), Golitsyn et al. (1963), Golitsyn (1965), Hryn’ 
(1973), Kondratyuk et al. (1988), Shelyag-Sosonko et al. 
(1991), Tkachenko et al. (1998) and Vakarenko & Duby-
na (2006). It was found that the main dominant species 
of these groups are Hyssopus cretaceous, Thymus calcareus, 
Asperula tephrocarpa, Scrophularia cretacea, Pimpinella 
tragium ssp. titanophylla, Artemisia hololeuca, A. sal-
soloides, Plantago maritima ssp. ciliata Printz, Jurinea 
stoechadifolia, Krascheninnikovia ceratoides, Silene creta-
cea and Festuca cretacea T. I. Popov & Proskor., which 
form together a variety of combinations. Cretaceous 
outcrop vegetation is quite different from zonal steppe 
vegetation (Didukh 1996). Based on the specific condi-
tions of substrate formation and apparent characteristics, 
H. Vysots’kyy (1915), B. Kozo-Polyanskyi (1931) and 
E. Lavrenko (1961) named it ”thymyannyky” (after the 
genus name Thymus). Ya. Didukh (1981) considered it an 
analogue of Mediterranean tomillares. 
The first floristic classification of Cretaceous vegeta-
tion was obtained by Ya. Didukh (1989). It was attrib-
uted to the order Thymo cretacei-Hyssopetalia cretacei 
Didukh 1989 and considered as part of the class Festu-
co-Brometea. Later, in 1996 and in virtue of additional 
studies, the chalky outcrop communities were assigned 
to the new class Helianthemo-Thymetea Romashchenko 
et al. 1996, in which there are three alliances: Artemi-
sio hololeucae-Hyssopion cretacei, Euphorbio cretophilae-
Thymion cretacei and Centaureo carbonatae-Koelerion 
talievii. After that studies have been conducted both in 
Ukraine (Didukh & Korotchenko 1996) and in Rus-
sia (Averinova 2005, 2011, 2012, 2014, Demina, 2011, 
2012, 2014, 2016; Poluyanov 2009, 2012, Sereda 2003, 
2008, 2009) that made some alterations to this clas-
sification. In the later construction EuroVegChecklist 
(Mucina et al. 2016), the order Hyssopetalia cretacei was 
included in the class Festuco-Brometea. Purpose of our 
research was to conduct a critical analysis of the syntaxo-
nomical structure of Cretaceous outcrop vegetation and 
to show its difference from the steppe vegetation of the 
class Festuco-Brometea.
Our key task was to critically analyse the classification 
of syntaxa described in previous years and to add novel 
information based on new data representing the whole 
research area within Ukraine and the Russian Federa-
tion.
Y. Didukh, O. Chusova & O. Demina
Syntaxonomy of chalk outcrop vegetation of the order  
Thymo cretacei-Hyssopetalia cretacei
87
17/1 • 2018, 85–109
Description of the research 
territory
The Central Russian Upland extends from Northwest to 
Southeast from the right bank of the Oka River to the 
Donetsk Range. In the West it is adjacent to the Dnieper 
Lowland and in the East to the Oka-Don Lowland; in the 
North its frontier follows the watershed of the rivers Desna, 
Oka and Don, and in the South it follows the watershed of 
the rivers Dnieper, Siversky Donets and Don (Karandeeva 
1957). The foothills of the Central Russian Upland are 
the plane inclined to the South and Southwest with eleva-
tions of about 190–200 m. It is deeply dissected by draws 
and ravines. In the South-East it borders with the Donetsk 
Range and the Siversky Donets valley. The Donetsk Range 
can be considered as foothills of the Central Russian Up-
land. It is located in the Donetsk region and the South of 
the Lugansk region, which belong to Ukraine and to the 
Rostov region of the Russian Federation. In the Donetsk 
Range, cretaceous outcrops appear only in the valleys of 
Siversky Donets and some of its feeders.
The south-western foothills of the Central Russian Up-
land are located within the forest-steppe and steppe zones 
and cover the territory of the Kharkiv, Donetsk and Lu-
hansk regions of Ukraine and Kursk and the Voronezh, 
Belgorod and Rostov regions of the Russian Federation 
(Baranov 1969). The research territory, 50° 52' 21" N, 
38° 4' 26" E to 49° 30' 44" N, 36° 41' 48" E and 49° 30' 
53" N, 43° 26' 04" E to 47° 27' 31" N, 40° 27' 30" E, 
covers the southwestern foothills of the Central Russian 
Upland and the Donetsk Range. It comprises only the 
steppe zone and is characterised by Cenomanian chalk 
outcrops. To the North, Cretaceous outcrops occur with-
in the forest-steppe zone, featuring hyper-carbonatophiles 
(Thymus calcareus Klokov & Des.-Shost., Inula ensifolia 
L., Polygala sibirica, Bupleurum falcatum L.), but in spe-
cies composition and structure, these communities are 
very different from Hyssopetalia cretacei (Golitsyin 1936, 
Honcharenko 2000).
The recent terrain of the Central Russian Upland is a 
result of the Neogene-Quaternary neotectonic surface 
emergence on the height of 200 meters, which was ac-
companied by tectonic and erosion-accumulative defor-
mation. Tectonic deformations are the result of structural 
heterogeneity of submerging basal complex, the presence 
of deep lateral and transverse crust fractures and inherited 
geological structure of the platform’s sedimentary sheath.
River valleys of the Siversky Donets and its feeders 
(Krasna, Aidar, Oskol) are wide, with steep right banks, 
which can be up to 100 m high. The right banks are dis-
sected by a system of ravines and gullies. The ravines are 
fairly long for the most part and sometimes have the 
shape of a “circus”. These formations indicate intense ero-
sion processes in the past and present. The left banks are 
gently sloping, with well-developed upland terraces.
The research area is under conditions of sub-conti-
nental (subarid) climate, characterised by large tempera-
ture amplitudes between winter and summer periods. 
The average annual temperature is 7–8 °C, the average 
January temperature ranges from −4 to −6 °C, the aver-
age July temperature from +21 to +24 °C. Annual pre-
cipitation is in the range of 450–500 mm in most parts 
of the research territory, ranging up to 550 mm in the 
northwestern part in the Russian Federation (Figure 1a). 
The central part of the reseach territory is characterised 
by considerable continentality and longer droughts in 
the summer period (Figure 1b). The most continental 
and most arid territory is the south-eastern part of the 
Donetsk Range (Figure 1c) with fairly irregular precipi-
tation of 400–420 mm per year. The largest is the spring 
and early summer rainfall, followed by a sharp decline in 
the second half of summer and a subsequent rise in win-
ter (Hrytsenko 2004, Weather in Ukraine, 2014).
Figure 1: Walter climographs based on climatic data from Hrytsenko (2004) and Weather in Ukraine (2014) for the northern (a), central (b) and 
southern (c) part of the research territory.
Slika 1: Walterjevi klimadiagrami na osnovi podatkov iz Hrytsenko (2004) in Weather in Ukraine (2014) za severni (a), srednji (b) in južni (c) del 
raziskovanega območja.
















           
 


  


  











           









           
  

	



  

	
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Y. Didukh, O. Chusova & O. Demina
Syntaxonomy of chalk outcrop vegetation of the order  
Thymo cretacei-Hyssopetalia cretacei
88
17/1 • 2018, 85–109
As the Walter climograph shows, such distribution 
determines a rather long (4–5 months) dry season, and 
its effect is further enhanced on the dry southern chalk 
slopes. This is one of the factors that determine the speci-
ficity of flora and vegetation. Under such conditions, 
black soils are formed in upland areas and rendzinas (sod-
calcareous soils) and regosols (deluvial-carbonate soils) on 
the slopes with the chalk outcrops.
Materials and Methods
For the analysis we used 354 relevés made by Ya. Didukh 
and O. Chusova in the Kharkiv, Donetsk and Lugansk 
region of Ukraine and by O. Demina in the Rostov re-
gion of the Russian Federation from 1987 to 2015. In 
addition, we used 51 relevés from the Kharkiv region 
kindly provided by O. Bezrodnova as well as 178 relevés 
published by K. Romashchenko et al. (1996), A. Poluy-
anov (2009, 2012), E. Averinova (2014) and M. Sereda 
(2008, 2009). To estimate the situation of Cretaceous 
outcrop vegetation in relation to other types, we used 
about 300 relevés of xerophytic herbaceous vegetation 
from the same territory. The plot sizes for relevés were 
from 25 to 100 m2. Unpublished releves were compiled 
in a TURBOVEG (Hennekens & Schaminee 2001) da-
tabase. Unpublished relevés sampled by Chusova (112 
relevés) were added to the GIVD World Warehouse in 
the Ukrainian Grassland Database (EU-UA-001). Un-
published releves sampled by Didukh (242 relevés) were 
added to the Vegetation of Lugansk region, Ukraine 
(EU-UA-008).
Floristic data were processed by using JUICE (Tichý 
2002). Clusterisation and syntaxa extraction were per-
formed using a modified TWINSPAN algorithm (mini-
mum number of groups – 2, pseudospecies cut level 
– 0 5 25; Simpson coefficient) (Roleček et al. 2009) im-
plemented in the software package JUICE (Tichý 2002). 
We also used the method of cluster analysis included in 
the STATISTICA 6.0 software package.
The diagnostic species of clusters were determined by 
the phi coefficient of fidelity (Chytrý et al. 2002). Us-
ing Fisher’s exact test at α = 0.05, species with a value of 
phi ≥ 0.3 were considered diagnostic, species with a value 
of phi ≥ 0.5 highly diagnostic. For dominant species, we 
adopted a projective coverage threshold of > 25% in the 
herbage and a 10% threshold of frequency.
We identified selected syntaxa on the basis of a critical 
analysis of publications (Didukh 1989, Romashchenko 
et al. 1996, Didukh & Korotchenko 1998, Sereda 2008, 
2009, Poluyanov 2009, 2012, Demina 2011, Averinova 
2014, Mucina et al. 2016). For the JUICE analysis, we 
have used the associations’ typical relevés to identify them.
Nomenclature of syntaxa follows the International 
Code of Phytosociological Nomenclature (Weber et al. 
2000); nomenclature of taxa is in accordance with the 
Euro+Med Plant base service (2006) or, in case of taxa 
not listed there, Flora Europaea (Tutin 1968, 1972, 1976, 
1980, 1993). We did use the name Hyssopus cretaceous 
Dubj. instead of Hyssopus officinalis L., though. Since the 
species of chalk outcrops are rather specific and narrowly 
local, many of them are endemic. However, in the Flora 
Europaea, these species are only listed as synonyms. Usu-
ally we have reduced them to synonyms that are accepted 
for Europe, but as for Hyssopus cretaceus, the situation is 
different. Within the research territory there are two sub-
species of Hyssopus officinalis, which both form associa-
tions of petrophytic type, but are quite different in terms 
of morphology and ecology. Hyssopus cretaceus is reduced 
in Flora Europaea and Euro+Med Plant to the synonym 
Hyssopus officinalis ssp. montanus (Jord. & Fourr.) Briq., 
which occurs only on the chalk substrate in communities 
of the order Thymo cretacei-Hyssopetalia cretacei. Another 
subspecies is sometimes found on sandy soils (Demina 
2012). Since Hyssopus officinalis ssp. montanus plays a very 
important role in the communities we are considering 
and in order to separate it from subspecies with a differ-
ent ecology, we decided to use the synonym here.
In the species composition analysis, it was taken into 
account that in our study region many species typical for 
Ukraine and southern regions are replaced by species that 
are similar in terms of environmental characteristics (e.g. 
Euphorbia cretophila Klokov and E. petrophila, Jurinea 
brachycephala Klokov and J. stoechadifolia, Genista tan-
aitica P. A. Smirn. and G. tinctoria L.).
Results and Discussion
Most communities of Cretaceous outcrops are character-
ised by small species variety and high diversity of coe-
noses. The presence of a large number of local endem-
ics among the dominant species is also characteristic for 
them. Some of the dominant species have diagnostic 
value at the level of associations, but the most typical spe-
cies of these groups (Thymus calcareus, Pimpinella tragium 
(Woronow) Tutin, Artemisia hololeuca, Hyssopus cretaceus) 
are characterised by a fairly wide range. These features are 
important when selecting syntaxa.
Different authors in various regions described four 
alliances (Didukh 1989, Romashchenko et al. 1996, 
Averinova 2014) and 21 associations (Didukh 1989, Ro-
mashchenko et al. 1996, Didukh & Korotchenko 1997, 
Sereda 2009, Poluyanov 2009, 2012, Demina 2011, 
2012, Averinova 2014). We conducted a critical analysis 
whereby it was established that the class Helianthemo-
Y. Didukh, O. Chusova & O. Demina
Syntaxonomy of chalk outcrop vegetation of the order  
Thymo cretacei-Hyssopetalia cretacei
89
17/1 • 2018, 85–109
tous steppes (“lowered alpine plants”) and falls into the 
two alliances Centaureo carbonatae-Koelerion talievii and 
Euphorbio cretophilae-Thymion cretacei. Cluster “B” cor-
responds to the alliance Artemisio hololeucae-Hyssopion 
cretacei and covers typical tomillares.
Consecutive number 1 2 3 4 5 6 7 8 9 10 11 12
15 12 20 107 29 19 24 14 15 26 15 46
Species richness
D. s. for ass. Artemisio nutantis-Plantaginetum salsae 
Plantago maritima 100 . 5 13 . . 4 7 . . 4 .
Artemisia nutans 80 8 . 3 . . 4 7 . . . .
Bassia prostrata 40 . 5 . . . . . . . 4 .
Convolvulus lineatus 53 . . 2 7 . 17 . . . 13 .
D. s. for ass. Erysimo cretacei-Festucetum cretacei 
Erysimum ucrainicum . 58 5 . . . . . . . . .
Melilotus officinalis . 75 . 10 . . 8 . . 4 . .
Festuca cretacea 20 100 40 8 . . . . . 4 . .
D. s. for ass. Lepidio meyeri-Scrophularietum cretacei 
Lepidium meyeri . . 45 . . . . . . . . .
Hedysarum cretaceum 13 . 69 . . . 8 . . . 8 .
Galium octonarium 7 33 85 3 3 . 33 21 . . 29 .
Artemisia absinthium . . 30 . . . 4 . . . . .
Poa compressa 13 42 30 10 10 . 17 . 13 20 4 20
Thesium arvense 7 33 30 11 28 . 17 21 40 . 25 .
D. s. for ass. Artemisio hololeucae-Polygaletum cretaceae 
Artemisia hololeuca 73 . 10 64 3 . 29 21 20 16 . .
D. s. for ass. Euphorbio cretophilae-Jurinetum brachycephalae 
Pinus sylvestris . . . . 7 . . . . . . .
Helianthemum canum . . . 2 59 . . 29 . 28 13 .
D. s. for ass. Genisto scythicae-Artemisietum salsoloides 
Genista scythica . . . . . 95 . . . . 4 .
Rhaponticoides taliewii . . . . . 53 . . . . . .
Leontodon biscutellifolius . . . . 14 53 . . . 4 . .
Linum tenuifolium 20 . 5 6 7 79 25 . . 4 8 .
Odontarrhena tortuosa . . . 3 28 84 25 14 . 12 17 26
Linaria vulgaris . . 5 1 . 21 . . . . . .
Consecutive number 1 2 3 4 5 6 7 8 9 10 11 12
15 12 20 107 29 19 24 14 15 26 15 46
Diplotaxis cretacea . . 5 4 . 47 . . . 12 . .
Polygala comosa . . . . . 21 . . . 4 . .
Linum hirsutum . . . 9 7 84 46 43 27 24 46 2
D. s. for ass. Hedysaro grandiflori-Centauretum sumensis 
Allium inaequale . . . . 10 . . 64 . 40 4 4
Hedysarum grandiflorum . . . 7 31 42 13 100 . 4 4 .
Thymelaea passerina . . . . . . . 43 . . . .
Asperula cynanchica . . . 3 3 53 4 93 . 4 4 48
Adonis vernalis . . . . . . . 50 . 8 8 13
Seseli annuum . . . . . . . 21 . . . .
Pimpinella saxifraga . . . . . . . 21 . . . .
Phlomis herba-venti . . . . . . . 21 . . . .
Salvia verticillata . . . 7 10 . 4 71 27 12 4 26
Aster amellus . . . . . . 4 36 . . 13 .
Asparagus officinalis . . . . 3 . . 21 . . 4 .
Caragana frutex . . . . 7 . 8 43 . 16 8 .
Centaurea orientalis . . . 2 7 . 29 43 . . 17 .
Echinops ritro 13 . 25 7 10 . 25 50 . 8 8 15
D. s. for ass. Androsacio koso-poljanskii-Caricetum humilis 
Androsace villosa subsp. 
koso-poljanskii 
. . . 6 . . . . 100 12 8 57
D. s. for ass. Bupleuro falcatae-Stipetum capillatae 
Bupleurum falcatum 13 . 10 23 45 . 21 86 20 76 42 59
Anisantha tectorum . . . . 3 . . . . 12 . .
Salvia nemorosa . . . . . . . . . 8 . .
Scabiosa ochroleuca 7 . . 11 17 . 8 43 40 84 21 48
D. s. for ass. Gypsophilo oligospermae-Campanuletum sibiricae 
Carex pediformis . . . . 3 . 4 . . . 29 .
Koeleria pyramidata . 8 20 3 24 . 4 . . 16 71 .
Table 1: Synoptic table of chalk outcrop vegetation communities with percentage frequencies.
Tabela 1: Sinoptična tabela rastlinskih združb krednih izdankov s frekvencami v odstotkih.
Figure 2: Tree diagram of chalk outcrop vegetation communities’ 
distribution (syntaxa numbering shown in Table 1). A – lowered 
alpine plants (Centaureo carbonatae-Koelerion talievii and Euphor-
bio cretophilae-Thymion cretacei); B –typical tomillares (Artemisio 
hololeucae-Hyssopion cretacei).
Slika 2: Dendrogram vegetacije krednih izdankov (oštevilčenje 
sintaksonov kot v Tabeli 1). A – nizke alpinske rastline (Centaureo 
carbonatae-Koelerion talievii in Euphorbio cretophilae-Thymion cretacei); 
B – tipični tomiljar (Artemisio hololeucae-Hyssopion cretacei).
Tymetea is represented by 14 associations within the terri-
tory of the south-western foothills of the Central Russian 
Upland. A tree diagram was built based on the assessment 
of species similarity. There are two clusters in this diagram 
(Figure 2). Cluster “A” covers communities of petrophy-




















 
           
Y. Didukh, O. Chusova & O. Demina
Syntaxonomy of chalk outcrop vegetation of the order  
Thymo cretacei-Hyssopetalia cretacei
90
17/1 • 2018, 85–109
Consecutive number 1 2 3 4 5 6 7 8 9 10 11 12
15 12 20 107 29 19 24 14 15 26 15 46
Nonea pulla . . . . . . 4 14 7 . 33 .
Bromopsis riparia . . . 6 3 11 17 57 33 12 75 11
Salvia nutans . . . 1 38 26 25 50 27 52 83 30
Galatella villosa . . . . . . . . . . 38 .
Veronica spicata . . . 3 3 . 21 14 . 8 29 .
Inula aspera . . 5 . . . 8 . 7 . 21 .
D. s. for ass. Carici humilis-Thymetum calcarei 
Astragalus austriacus 7 . . . . . . 43 33 4 29 41
Thalictrum minus . . . . 24 42 29 43 . 36 67 17
Oxytropis pilosa . . . 2 . . 4 . 7 . 17 7
Jurinea arachnoidea 7 . . 4 3 . 42 29 20 4 46 15
Viola rupestris . . . . . . . . . 4 . 35
Helianthemum nummu-
larium 
. . . . . . . 36 . 4 . 59
Asperula tinctoria . . . 2 . . . . . . 4 43
Anthericum ramosum . . . . 21 . . 21 7 8 . 54
Linum perenne . . . . . . . 14 . . . 26
Echium vulgare . . . . . . 4 . . 4 . 28
Vincetoxicum hirundinaria . . . . 3 . . . 7 36 8 50
Pilosella officinarum . . . . . . . . 7 . 4 26
Euphrasia pectinata . . . 5 . . 4 . 7 20 4 39
D. s. for alliance Artemisio hololeucae-Hyssopion cretacei
Scrophularia cretacea 33 58 65 45 21 11 17 7 13 16 . .
Hyssopus cretaceus 80 75 90 95 7 . 4 7 27 . . 2
Matthiola fragrans 100 . 25 50 10 . 17 . 20 . . 4
Linaria cretacea 20 . 10 11 14 . . . . . . .
D. s. for alliance Euphorbio cretophilae-Thymion cretacei
Artemisia salsoloides 20 . 85 36 34 100 79 7 . 8 17 .
Jurinea stoechadifolia . . . 1 86 58 . . . 12 . .
Euphorbia petrophila . . 30 5 93 100 8 . . 8 . .
Brassica elongata 13 . . 18 52 26 33 21 . 24 25 .
D. s. for alliance Centaureo carbonatae-Koelerion talievii
Stipa capillata 7 . 5 7 3 . 25 79 27 72 46 70
Campanula sibirica . . 5 7 59 42 42 71 20 60 88 61
Polygala sibirica . . . 9 3 . 17 36 47 24 58 61
Carex humilis . . . 4 14 . 17 79 27 60 67 87
Stipa pennata . . . 5 3 . 29 86 . 12 25 78
Onosma simplicissima 7 . . 23 59 53 58 86 27 64 46 67
Koeleria talievii 7 . . 9 14 . 29 50 13 52 13 61
Psephellus marschallianus . . 10 3 41 5 50 57 7 28 42 50
Teucrium polium 20 . . 36 66 100 96 100 47 76 96 .
D. s. for order Thymo cretacei-Hyssopetalia cretacei
Pimpinella tragium 73 92 90 91 83 95 83 57 93 56 50 37
Gypsophila oligosperma 60 50 70 62 76 89 83 93 67 84 79 85
Cephalaria uralensis 27 8 85 74 45 95 79 57 7 8 25 .
Thymus calcareus 60 . 5 87 97 100 92 86 93 68 33 96
Asperula tephrocarpa 73 . 5 90 93 5 88 14 60 52 50 2
Reseda lutea 7 33 5 28 52 74 58 71 13 20 58 9
Silene supina 33 . . 50 21 5 33 21 40 52 33 4
Linum pallasianum 27 . . 56 62 5 63 79 60 52 54 41
Polygala cretacea 20 33 . 36 14 . 63 57 13 36 25 26
Astragalus albicaulis 40 8 55 21 7 . 42 36 13 40 21 54
Genista tinctoria 27 . 5 23 7 . 21 36 . . 38 7
Consecutive number 1 2 3 4 5 6 7 8 9 10 11 12
15 12 20 107 29 19 24 14 15 26 15 46
Taraxacum serotinum 20 17 5 6 . . 13 29 . 4 8 .
Silene cretacea 20 . . 5 3 . . 14 . . . .
Orthantha lutea 33 25 . 20 21 . . 21 . 40 4 .
Hedysarum ucrainicum 7 . . 7 . . . . . . . .
Scutellaria supina . . . 2 3 21 8 . . . 4 .
Clausia aprica . . . . . . . . . . . 7
Syrenia cana . . . 2 . . . . . . . .
Linaria genistifolia . . . 2 3 . . . . 4 4 .
Schivereckia podolica . . . . 3 . . . . . . .
 D. s. for class Festuco-Brometea 
Festuca valesiaca . . 5 7 3 . 25 43 20 32 75 76
Euphorbia seguieriana 53 8 10 35 14 5 79 79 67 64 75 80
Galium ruthenicum . . . 2 7 . 29 . . 8 21 7
Hieracium virosum . . 5 1 3 . 13 . . 8 38 13
Euphorbia nicaeensis 13 8 10 4 7 . 21 . 13 16 46 .
Stachys recta . . . 3 3 . 29 36 7 32 54 22
Medicago falcata . 17 . 10 . . 21 43 40 16 29 35
Agropyron cristatum 13 . 5 1 14 . 25 . . 8 . .
Securigera varia . 25 . 6 7 . 29 50 33 12 42 13
Erysimum diffusum 7 . 5 7 10 21 38 43 13 36 17 35
Stipa pulcherrima . . . . . . 38 . 7 4 21 2
Festuca stricta . . . 9 . 5 21 . . 4 13 .
Elytrigia stipifolia 7 . . 3 . . . . 7 . . .
Stipa lessingiana . . . 8 3 21 38 14 . 20 13 .
Ajuga chamaepitys . . . . . . 8 . 7 . 8 13
Filipendula vulgaris . . . . . . 4 . . . 17 .
Elytrigia intermedia . . . . . . 17 . 7 4 13 22
Hieracium umbellatum . . . . 3 . . . . 12 4 13
Viola ambigua . . . . 10 . 4 . . 16 17 9
Pilosella echioides . . . . . . . . 7 4 4 11
Astragalus onobrychis 7 . . 3 . . 4 29 13 . 17 4
Potentilla humifusa . . . . . . 4 . 7 4 25 28
Rhaponticoides ruthenica . . . . . . 8 . . . 8 13
Onobrychis arenaria . . . 1 . . 4 29 7 4 21 17
Plantago media . . . . . . 4 . . . 17 9
Plantago lanceolata . . . 2 3 . . . . 4 8 .
Achillea setacea . . . . . . . . . 8 17 9
Eryngium campestre . . . . 3 . 13 . 7 4 . .
Cota tinctoria . 8 5 1 3 11 8 . 7 . 4 .
Achillea millefolium . . . . . . 13 . 7 4 4 .
Astragalus subuliformis 7 . . 8 . . 13 . . 12 4 .
Potentilla incana . . . . . . 4 . 7 . 13 11
Potentilla recta . . . . 14 . . . 7 8 8 .
Linum austriacum . . . 2 . . 25 . . . 17 .
Centaurea stoebe . . . 1 . . . . . 4 . 4
Cleistogenes serotina . . . . 3 . 4 . . 12 17 .
Hypericum elegans . . . . 7 . 8 21 . 4 17 .
Potentilla patula . . . . . . 4 . . 16 17 .
Stipa zalesskii . . . . . . . . 13 4 4 .
Other species
Helichrysum arenarium 7 . 60 11 41 . 46 . . 44 50 .
Ephedra distachya . . 5 . 3 . 4 . . . 4 .
Cytisus ruthenicus . . . 2 10 . 17 . . 12 21 7
Y. Didukh, O. Chusova & O. Demina
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Consecutive number 1 2 3 4 5 6 7 8 9 10 11 12
15 12 20 107 29 19 24 14 15 26 15 46
Melampyrum arvense . . . 8 3 . 13 29 . 4 8 7
Centaurea diffusa . . . . 3 . 4 21 . . 13 .
Atraphaxis frutescens . . . 5 . . . . . . . .
Elytrigia repens 7 . . . . . . . . 4 13 .
Verbascum lychnitis . . . 2 . . . . . 8 4 9
Artemisia campestris . . . . . . . . . . 8 9
Seseli libanotis . . 10 2 . . . . . 4 . .
Melica transsilvanica . . 20 . . . 4 . . . 4 .
Poa angustifolia . . 20 1 . . 4 . . 16 13 .
Krascheninnikovia cera-
toides 
. . 20 . . . . . . . . .
Hypericum perforatum . . . . . . . . . 20 13 .
Cichorium intybus . 17 . . . . . . 7 4 4 .
Lotus corniculatus . . . . . . . . . 4 . .
Gypsophila fastigiata . . . 4 . . . . 20 8 . .
Marrubium peregrinum . . . . . . 4 . . 12 13 .
Poa bulbosa . . 5 1 . . . . . 20 4 4
Odontites vulgaris . . . 4 . . 13 . . . . 13
Cotinus coggygria . . . . 3 . . . . 4 . .
Calamagrostis epigeios . . . . . . 8 . . 12 8 .
Gypsophila paniculata . . . 2 . . 4 . . . 4 .
Artemisia austriaca . . 15 1 . . . . . 12 8 .
Associations: 1. Artemisio nutantis-Plantaginetum salsae; 2. Erysimo 
cretacei-Festucetum cretacei; 3. Lepidiomeyeri-Scrophularietumcre-
tacei; 4. Artemisio hololeucae-Polygaletum cretaceae; 5. Euphorbio 
cretophilae-Jurinetum brachycephalae; 6. Genisto scythicae-Artemi-
sietum salsoloides; 7. Pimpinello titanophillae-Artemisietum salsoloi-
des; 8. Hedysaro grandiflori-Centauretum carbonatae; 9. Androsacio 
koso-poljanskii-Caricetum humilis; 10. Bupleuro falcatae-Stipetum 
capillatae; 11. Gypsophilo oligospermae-Campanuletum sibiricae; 
12. Carici humilis-Thymetum calcarei
Classification scheme of chalk outcrop vegetation 
 communities.
Ord. Thymo cretacei-Hyssopetalia cretacei 
All. Artemisio hololeucae-Hyssopion cretacei 
1 Ass. Artemisio nutantis-Plantaginetum salsae Didukh 
1989
2 Ass. Erysimo cretacei-Festucetum cretacei Demina 
2011
3 Ass. Lepidio meyeri-Scrophularietum cretacei Demi-
na 2012 (syn. Hedysaro cretacei-Melicetum transsil-
vanicae Demina 2011)
4 Ass. Artemisio hololeucae-Polygaletum cretaceae Di-
dukh 1989 (syn. Thymo cretacei-Hyssopetum cretacei 
Didukh 1989, Onosmo tanaiticae-Androsacietum ko-
zo-poljanskii Didukh et al. 1996, Matthiolo fragran-
si-Atraphaxietum frutescens Demina 2011, Polygalo 
sibiricae-Hyssopetum cretacei Poluyanov 2012)
All. Euphorbio cretophilae-Thymion cretacei Didukh 1989
5 Ass. Euphorbio cretophilae-Jurinetum brachycepha-
lae Didukh 1989 (syn. Jurineo brachycephalae-
Helianthemetum cretophilae Romashchenko et al. 
1996; Jurineo brachycephalae-Koelerietum talievii 
Romashchenko et al. 1996, Scrophulario cretacei-
Helianthemetum cretacei Didukh et al. 1996)
6 Ass. Genisto scythicae-Artemisietum salsoloides Sere-
da 2009
All. Centaureo carbonatae-Koelerion talievii (syn. Carici 
humilis-Thymion calcarei Averinova 2014)
7 Ass. Pimpinello titanophillae-Artemisietum salsoloi-
des Didukh 1989
8 Ass. Hedysaro grandiflori-Psephelletum carbonatae 
Averinova 2014 corr. Didukh et al. nom. corr. hoc 
loco (original name: Hedysaro grandiflori-Centaure-
tum sumensis Averinova 2014: 37–47)
9 Ass. Androsacio koso-poljanskii-Caricetum humilis 
Korotchenko & Didukh 1998
10 Ass. Bupleuro falcatae-Stipetum capillatae
11 Ass. Gypsophilo oligospermae-Campanuletum sibiri-
cae Romashchenko et al. 1996 (syn. Scrophulario 
cretacei-Helianthemetum cretacei Romashchenko et 
al. 1996)
12 Ass. Carici humilis-Thymetum calcarei Poluyanov 
2009
The alliance Artemisio hololeucae-Hyssopion cretacei.
Typical tomillares of this alliance are represented by 
communities dominated be chamaephyte subshrubs 
(Thymus calcareus, Hyssopus cretaceus, Pimpinella tragium 
and Artemisia hololeuca). The communities occupy steep 
(15–45°) slopes up to 30–50 m, with mostly southern 
and sometimes eastern exposure. Substrate is a loose, 
moving chalk. They are confined to the right river banks 
and are absent in the upland watershed areas. Charac-
teristic features are the processes of denudation and su-
perficial erosion resulting in feeble soil formation or its 
absence. The plant cover of these communities is quite 
patchy (30%). Diagnostic species of these communities 
are, besides the dominant, Asperula tephrocarpa Popov & 
Chrshan., Linum pallasianum Schult, Matthiola fragrans 
and  Scrophularia cretacea.
1. Artemisio nutantis-Plantaginetum salsae 
Diagnostic species: Plantago maritima Printz, Artemisia 
nutans Willd., Bassia prostrata (L.) Beck, Convolvulus 
lineatus L.
Dominant species: Plantago maritima, Artemisia nutans, 
rarely Artemisia hololeuca, Hyssopus cretaceus.
Among the described associations, this one occupies 
the most saline substrates. Communities are encoun-
Y. Didukh, O. Chusova & O. Demina
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tered at the foot of the chalk slopes and at their bottom 
(10–20°) on deluvial, sufficiently coherent sediments, 
characterised by carbonate-sulfate soil salinisation. Indi-
cators of the salinity are Artemisia nutans, Plantago ma-
ritima, Convolvulus lineatus, rarer Bassia prostrata. These 
are also diagnostic species of this association, which is 
prevalent in the eastern part of the Luhansk region, on 
the banks of Siversky Donets’ left feeders (Aydar, Derkul, 
Kamyshna).
2. Erysimo cretacei-Festucetum cretacei 
Diagnostic species: Erysimum ucrainicum Desf., Festuca 
cretacea T.I. Popov & Proskor.
Dominant species: Festuca cretacea, Hyssopus cretaceus, 
Pimpinella tragium.
The community is abundant in the Rostov region. The 
association is characterised by a limited species composi-
tion. Apart from the dominant species mentioned above, 
obligate erodiophiles (species associated with soil ero-
sion) with powerful caudex are present (Scrophularia cre-
tacea, Gypsophila oligosperma Krasnova) as well as facul-
tative species with a wide range (Melilotus officinalis (L.) 
Lam., Poa compressa L., Reseda lutea L., Securigera varia 
(L.) Lassen) (Demina 2012).
3. Lepidio meyeri-Scrophularietum cretacei 
Diagnostic species: Hedysarum cretaceum DC., Lepidium 
meyerii Walp., Hedysarum cretaceum, Artemisia salso-
loides.
Dominant species: Artemisia salsoloides, Hedysarum creta-
ceum, rarely Hyssopus cretaceus, Gypsophila oligosperma, 
Thymus calcareus, Scrophularia cretacea.
These communities are present on chalk outcrops on 
the right bank of the Middle Don valley. The association 
includes two subassociations. Lepidio meyeri-Scrophula-
rietum cretacei typicum is formed at the bottom of the 
slopes on dense delluvial with signs of carbonate-sulfate 
salinity. Presence of Lepidium meyerii and sometimes 
Krascheninnikovia ceratoides is characteristic. Only four 
species (Hyssopus cretaceus, Artemisia salsoloides, Lepidi-
um meyeri, Pimpinella tragium) have significant roles in 
coenosis formation (Demina 2012, 2016).
The subassociation L. m.-S. c. hedysaretosum cretacei 
is restricted to the upper part of slopes with northern 
aspect. It is formed on chalk washed-off regosols. These 
are the most “humid” communities, which are confined 
to northern slopes. Erodiophiles like Melica transsilvani-
ca, Poa compressa and Thesium arvense have a significant 
share, whereas typical xerophytes are absent. Hedysarum 
cretaceum and Artemisia salsoloides are most frquent in 
these communities (Demina 2012, 2016). 
4. Artemisio hololeucae-Polygaletum cretaceae 
Diagnostic species: Artemisia hololeuca, Hyssopus cretaceus, 
Matthiola fragrans.
Dominant species: Artemisia hololeuca, Artemisia salsoloides, 
Festuca cretacea, Pimpinella tragium, Thymus calcareus.
These are the most common communities of the alli-
ance. They are confined to Cretaceous eroded steep slopes 
(15–45°) on the right banks of the Siversky Donets basin 
with different aspects. These communities are sometimes 
covered by lime rock of the Upper Eocene in the valleys 
of the Nagolnaya and Kamyshnaya rivers (Demina 2012). 
The communities of the association have a low plant cov-
er (50%) and a diverse species composition with the same 
dominant species as mentioned before. For this reason we 
have selected five associations with rather poor diagnostic 
blocks. We consider them variants or facies with specific 
patterns of distribution, primarily with respect to expo-
sure and steepness of slopes and to structure, mobility and 
density of the substrate. For example, Hyssopus cretaceus 
and Pimpinella tragium are dominant mostly on moving 
loose substrate on the northern slopes, while the southern 
slopes with compressed and strong substrate are covered 
by Artemisia spp. Under such conditions Thymus calca-
reus also dominates. It is a characteristic species of the as-
sociations of the next alliance. An ongoing involvement 
of Asperula tephrocarpa, Linum pallasianum and Polygala 
cretacea Kotov is also characteristic.
The alliance Euphorbio cretophilae-Thymion cretacei in-
cludes communities formed on looser substrate with evi-
dent organic content both on slight (20%) and on steeper 
slopes. The common name “jurineiniky” used for the 
communities of this alliance indicates the occurrence of 
Jurinea spp. (J. stoechadifolia s.l., J. arachnoidea Bunge), 
which are diagnostic species of this association. Euphor-
bia petrophila, Helianthemum canum (L.) Hornem. and 
Thymus calcareus subshrubs also contribute to the com-
munities of this alliance. Artemisia salsoloides is domi-
nant on the steeper rocky slopes. With approximately 
40–60%, the plant cover of these communities is higher 
than in the previous alliance.
5. Euphorbio cretophilae-Jurinetum brachycephalae
Diagnostic species: Euphorbia petrophila, Helianthemum 
canum, Jurinea stoechadifolia.
Dominant species: Helianthemum canum, Jurinea stoecha-
difolia, Thymus calcareus.
This association is common in the South of the research 
territory, on the right banks of the Siversky Donets River 
and its left tributaries. The communities occur on steep 
and moderately steep slopes (up to 15°) in areas adjacent 
to upland and hilltops. Its communities are characterised 
by the dominance of the orthotropic chasmophyte Ju-
Y. Didukh, O. Chusova & O. Demina
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rinea stoechadifolia, among which the plagiotropic Heli-
anthemum canum and Thymus calcareus occur. The plant 
cover can reach 70%.
6. Genisto scythicae-Artemisietum salsoloides 
Diagnostic species: Odontarrhena tortuosa (Waldst. & Kit. 
ex Willd.) C. A. Mey., Artemisia salsoloides, Rhaponti-
coides taliewii, Diplotaxis cretacea Kotov, Euphorbia 
petrophila, Genista scythica Pacz., Jurinea stoechadifolia, 
Leontodon biscutellifolius DC., Linaria vulgaris Mill., 
Linum hirsutum, Linum tenuifolium L., Reseda lutea.
Dominant species: Thymus calcareus, Genista scythica, Ju-
rinea stoechadifolia, Artemisia salsoloides.
Communities are gappy (plant cover 40–60%) and 
confined to the middle and upper parts of river valleys 
slopes (with 30–45° steepness), typically with southern, 
southeastern or southwestern exposure.
It is a quite rare association with specific diagnostic spe-
cies. Outcrops along the Tuzlov River in the Rostov region 
are the only location of petrophytic communities with 
occurrences of Genista scythica in Russia (Sereda 2008). 
In Ukraine Genista scythica grows only on the Donetsk 
Ridge and does not occur within the research territory.
The communities of the alliance Centaureo carbonatae-
Koelerion talievii are confined to places with chalk depos-
its and outcrops. Unlike in case of the alliance mentioned 
above, carbonaceous rocks are immobile and very dense, 
but the soil is not formed (regosols) or undifferentiated 
(rendzinas). The plant cover is much denser, ranging from 
30 to 70%. Dominant species are Thymus calcareus, Carex 
humilis, C. pediformis, Onosma simplicissima, Androsace 
villosa (Ovcz.) Fed. and Hedysarum grandiflorum. Typi-
cal for cretaceous communities are the following species: 
Linum pallasianum L., L. hirsutum, Teucrium polium L., 
Brassica elongata Ehrh., Thymus calcareus, Onosma simpli-
cissima, Silene supina, Psephellus marschallianus (Spreng.) 
K. Koch. Steppe species (Carex humilis, Stipa capillata 
L.) are also part of the floristic composition (Didukh & 
Chusova 2014). However, there are no such typical car-
bonatophiles as Hyssopus cretaceus, Scrophularia cretacea 
and Matthiola fragrans, and occurrence of Artemisia holo-
leuca and A. salsoloides is limited. 
This peculiarity indicates that it may be included in the 
order Stipo pulcherrimae-Festucetalia pallentis, but to prove 
this argument, it is necessary to analyse larger samples 
from other regions. Given that similar groups dominated 
by Jurinea stoechadifolia, Pimpinella tragium ssp. titano-
phylla and Thymus dimorphus Klokov & Des.-Shost. are 
distributed more widely, particularly on limestone in the 
steppe zone of Ukraine and Moldova, the final solution 
for this issue would be possible through broader analysis. 
As for now, the presence and dominance of hyper-car-
bonatophiles and carbonatophiles (Helianthemum canum, 
Thymus calcareus, Brassica elongata, Euphorbia petrophila, 
Koeleria talievii, Hedysarum grandiflorum, Asperula tephro-
carpa) and the minor role of grass and typical steppe spe-
cies of the order Festucetalia valesiacae (Medicago falcata 
L., Agrimonia eupatoria L., Veronica incana L., Ranunculus 
polyanthemos L., Achillea millefolium L., Potentilla argentea 
L., Falcaria vulgaris Bernh., Plantago media L., Elytrigia 
intermedia (Host) Nevski, Carex praecox Schreb., Tragop-
ogon dubius (Jacq.) Vollm.) suggest to subsume these com-
munities in the order Thymo cretacei-Hyssopetalia cretacei.
An important feature of this alliance is the quite strong 
floristic variability of its communities. They are based on 
the diagnostic species of the alliance, but in some associa-
tions, the diagnostic species are not clearly marked. That 
is why it is quite difficult to distinguish such communities 
solely by the floristic composition.
7. Pimpinello titanophillae-Artemisietum salsoloides
Diagnostic species: Artemisia salsoloides, Jurinea arachnoi-
dea, Linum austriacum L., Polygala cretacea.
Dominant species: Artemisia salsoloides, Hedysarum creta-
ceum, Onosma simplicissima L., Thymus calcareus.
The communities are visually differentiated from a 
background of gray aspect due to the dominance of tall 
(60 cm), dense dark green stands of A. salsoloides. They are 
sporadically abundant in the middle of gentle (20°) slopes 
along the banks of left tributaries (Krasna, Aidar, Derkul, 
Kamyshna) of the Siversky Donets.
8. Hedysaro grandiflori-Psephelletum carbonatae
Diagnostic species: Adonis vernalis L., Allium inaequale 
Janka, Asperula cynanchica L., Carex humilis, Hedysa-
rum grandiflorum, Phlomis herba-venti (Willd.) Maire 
ex De Filipps, Pimpinella saxifraga L., Salvia verticillata 
L., Stipa capillata, Stipa pennata L., Thymelea passerina 
(L.) Coss. & Germ.
Dominant species: Carex humilis, Festuca valesiaca Gaud-
in, Gypsophila oligosperma, Thymus calcareus.
As Averinova (2014) notes, communities are distrib-
uted on chalk outcrops of relatively gentle (2–10°) slopes 
of ravines and river valleys with mainly southern expo-
sure. The substrate is a chalky fine earth with broken rock. 
The association may be diagnosed by the presence of a 
number of species. Its communities are described from 
the Belgorod region (Russian Federation).
The choice of Psephellus sumensis (Kalen.) Greuter. as 
diagnostic species is false because the race distinguished 
by M. Klokov is psamophytic and bound to boreal sand 
and pine forest communities. It is replaced by Psephellus 
carbonatus (Klokov) Greuteron on calcareous soil. Both 
species are included in the complex P. marschallianus s.l. 
Y. Didukh, O. Chusova & O. Demina
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9. Androsacio koso-poljanskii-Caricetum humilis
Diagnostic species: Androsace villosa.
Dominant species: Androsace villosa, Artemisia hololeuca, 
Carex humilis, Thymus calcareus.
The association is confined to rather steep (30–50°) 
slopes with different exposures. Its communities occur on 
a feebly (up to several centimetres) developed rendzina 
high in carbonate. They are characterised by a high spe-
cies richness and the presence of species with well-devel-
oped sod (Carex humilis, Festuca valesiaca).
10. Bupleuro falcatae-Stipetum capillatae
Diagnostic species: Carex humilis, Koeleria talievii Lavren-
ko, Poa bulbosa L., Vincetoxicum hirundinaria Medik.
Dominant species: Carex humilis, Stipa capillata, Thymus 
calcareus.
Communities occur at the bottom of gentle slopes and 
in depressions of slopes with moderate soil moisture. The 
association is contiguous to communities of the order 
Festucetalia valesiacae. Its communities are diagnosed by 
the dominance of Stipa capillata with species of the or-
der Thymo cretacei-Hyssopetalia cretacei being present. The 
plant cover is more than 60%.
11. Gypsophilo oligospermae-Campanuletum sibiricae
Diagnostic species: Bromopsis riparia (Rehmann) Holub, 
Campanula sibirica L., Carex pediformis, Galatella vil-
losa (L.) Rchb. f., Hieracium virosum Pall., Inula aspera 
Poir., Koeleria pyramidata (Lam.) P. Beauv., Nonea 
pulla DC., Salvia nutans L., Veronica incana, Veronica 
spicata L.
Dominant species: Astragalus albicaulis DC., Bromopsis 
riparia, Carex humilis, Psephellus marschallianus, Gala-
tella villosa, Festuca valesiaca, Gypsophila oligosperma, 
Helianthemum canum, Onosma simplicissima, Salvia 
nutans, Stipa capillata, Stipa pennata, Thymus calcareus.
The communities develop on slightly eroded chalk 
slopes and hilltops of the chalk mountains, on steppi-
fied sites with Cretaceous basis. The association is quite 
common and widespread. In general the plant cover may 
reach 80% due to the presence of Carex humilis and rep-
resentatives of the Poaceae family. The well-developed 
vegetation cover prevents surface erosion. The communi-
ties are observed to have a high species diversity. Stipa 
spp. or other prevalent grasses act as dominant species. 
However, due to the general xerophytism and signifi-
cant contribution of carbonatophiles, these communities 
belong to the order Thymo cretacei-Hyssopetalia cretacei. 
The association most commonly occupies areas tran-
sitional to the steppe vegetation adjacent to upland or 
steep (15–30°) southern slopes. In such places the plant 
cover contributes to soil accumulation. This association 
also includes communities with dominance of Carex 
pediformis, which is an arctic-boreal species. It is on the 
southern edge of its area here and is found only in a few 
exclaves. The coenotic range of the communities is quite 
narrow (Golitsyin 1958, Didukh & Chusova 2014).
12. Carici humilis-Thymetum calcarei 
Diagnostic species: Anthericum ramosum L., Bupleurum 
falcatum, Carex humilis, Euphrasia pectinata Ten., He-
lianthemum nummularium (L.) Mill., Pilosella officina-
rum Vaill., Vincetoxicum hirundinaria, Viola rupestris 
F.W. Schmidt
Dominant species: Androsace villosa, Thymus calcareus.
This association combines communities that are typical 
for the Upper Oskol Region on slightly sodded chalky 
slopes with dominance of calciphilous shrubs and sub-
shrubs. Its communities often occupy the most elevated 
relief elements, which are well warmed up, i.e. the upper 
and middle parts of steep (up to 40°) slopes with exposure 
to the South. They are less common on northern slopes 
and gentle hilltops (Poluyanov 2012).
To estimate separated syntaxa in the general classifica-
tion system of xerophilous grass-undershrub vegetation, 
we have processed the entire array of relevés  and built a 
tree diagram on the basis of species composition similar-
ity. As a result, there are six allocated clusters (Figure 3), 
two of which (1 and 2) correspond to alliances of the or-
der Thymo cretacei-Hyssopetalia cretacei. Relevés of steppe 
vegetation (order Festucetalia valesiacae) matched the 
clusters 3–5 and relevés of sandy steppes (class Koelerio-
Figure 3: Distribution of Cretaceous outcrop vegetation in relation to 
the steppe vegetation. A – Ord. Festucetalia valesiacae; B – Cl. Koelerio-
Corynephoretea (all. Festucion vaginatae); C – Ord. Thymo cretacei-
Hyssopetalia cretacei.
Slika 3: Razporeditev vegetacije krednih izdankov glede na step-
sko vegetacijo. A – red Festucetalia valesiacae; B – razred Koelerio-
Corynephoretea (all. Festucion vaginatae); C – red Thymo cretacei-Hysso-
petalia cretacei.


















 
     

Y. Didukh, O. Chusova & O. Demina
Syntaxonomy of chalk outcrop vegetation of the order  
Thymo cretacei-Hyssopetalia cretacei
95
17/1 • 2018, 85–109
Corynephoretea) cluster 6. Thus, this analysis shows that 
the communities of the order Thymo cretacei-Hyssopetalia 
cretacei differ from the Festucetalia valesiacae communi-
ties as a Koelerio-Corynephoretea in their floristic com-
position. All these communities have in common the 
predominance of xerophilous species; however, grasses 
(Festuca, Stipa) dominate in Festucetalia valesiacae and 
Koelerio-Corynephoretea communities, whereas chas-
mophilous plants are dominant in the order Thymo creta-
cei-Hyssopetalia cretacei.
The position of Cretaceous outcrop vegetation com-
munities within the modern system of syntaxonomical 
classification is quite debatable and depends on the in-
terpretation of the size of different syntaxa. Consider-
ing Festuco-Brometea in a fairly wide range, it should be 
noted that calciphyte communities are common in the 
steppe zone and have been in direct contact with zonal 
vegetation so that some species of the class Festuco-Bro-
metea are present in those coenoses. But in general, these 
events are quite random with low class constancy (I–II). 
Inclusion of the order Thymo cretacei-Hyssopetalia creta-
cei into the class Festuco-Brometea can be considered as 
substitution of Stipo pulcherrimae-Festucetalia pallentis 
Pop 1968 in the East. However, the difference in ecologi-
cal conditions between them is that the first is common 
on dense carbonates (gypsum, dolomite, limestone) and 
the latter – on idiomorphic soils with different densities 
– often on loose and mobile regosols. In this respect, the 
Thymo cretacei-Hyssopetalia cretacei communities may be 
regarded as similar to Koelerio-Corynephoretea Klika in 
Klika et Novák 1941, and Stipo pulcherrimae-Festuceta-
lia pallentis as an analogue of Sedo-Scleranthetea Br.-Bl 
1955. They are similar in structure and growing con-
ditions to the Mediterranean communities of the class 
Drypidetea spinosae Quezel 1964, where Mucina et al. 
(2016) included Crimean-Caucasian syntaxa of Onosmo 
polyphyllae-Ptilostemetalia Korzhenevsky 1990. But the 
location and status of this order has not been fully es-
tablished yet. All of the above shows that a number of 
syntaxonomical issues are still unresolved and require 
broader discussion.
Acknowledgement
The autors are very grateful to Dr. Orsolya Valkó for ed-
iting and handling the manuscript, to Dr. Aiko Huckauf 
and Andriy Mosyakin for English proofreading and lin-
guistic edition. We also thank two anonymous reviewers 
for the valuable comments and recommendations.
Conclusions
We provided a comprehensive syntaxonomic survey of 
Cretaceous outcrop vegetation from south-western foot-
hills of the Central Russian Upland. Based on relevés 
from Ukraine and the Russian Federation, we could con-
clude that all communities belong to the order Thymo 
cretacei-Hyssopetalia cretacei and are clearly separated from 
other steppe communities. The order Thymo cretacei-Hys-
sopetalia cretacei includes twelve associations belonging to 
the three alliances Artemisio hololeucae-Hyssopion cretacei, 
Euphorbio cretophilae-Thymion cretacei and Centaureo car-
bonatae-Koelerion talievii.
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Consecutive number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21
Total vegetation cover (%) 60 15 40 20 50 45 20 50 40 25 70 70 50 40 20 30 50 60 40 50 50
Slope (°) 10 5 5 25 30 45 25 5 15 25 30 30 20 45
Aspect (°) 5 90 180 135 180 135 90 180 180 45 90 90 180 90
Species richness 14 11 15 19 17 15 15 13 11 14 17 9 13 13 11 14 20 16 15 17 16
D. s. of ass. Artemisio nutantis-Plantaginetum salsae  
Plantago maritima 3 1 4 4 3 1 2 2 1 1 . . . . . . . . . . .
Artemisia nutans 4 2 3 2 1 1 1 1 1 1 . . . . . . . . . . .
Bassia prostrata 1 1 . . . . . . . . . . . . . . . . . . .
Convolvulus lineatus 1 1 1 1 . . . . . . . . . . . . . . . . .
D. s. of ass. Artemisio hololeucae-Polygaletum cretacei 
Artemisia hololeuca . . . 1 1 1 3 3 3 3 1 2 4 3 3 3 4 3 4 4 3
D. s. of alliance Artemisio hololeucae-Hyssopion cretacei 
Scrophularia cretacea . . . . . 1 1 . . 1 . . 1 2 2 2 . 1 . 1 .
Hyssopus cretaceus . . 1 . 3 3 2 2 3 2 1 3 2 2 2 3 2 1 1 3 1
Matthiola fragrans 1 1 1 1 1 1 1 1 1 1 1 . . 1 1 2 1 1 1 1 .
Linaria cretacea . . . 1 1 . 1 . . . . . . . . . . . 2 . .
D. s. of alliance Euphorbio cretophilae-Thymion cretacei
Brassica elongata . . . 1 . . . . . . . . . . . . . . . . 1
Artemisia salsoloides . . . 1 . . . . 2 . 4 2 . 2 . . . . . 1 .
D. s. of alliance Centaureo carbonatae-Koelerion talievii  
Polygala sibirica . . . . . . . . . . . . . 1 . . . . . . .
Carex humilis . . . . . . . . . . . . . . . 1 1 . . . .
Onosma simplicissima . . . . . . . . . . . . . . . 2 1 1 . 1 1
Koeleria talievii . . . 1 . . . . . . . . . . . . . . . . .
Teucrium polium . 1 . . . . . . 1 . 2 . . . 1 . . . 2 1 1
D. s. of order Thymo cretacei-Hyssopetalia cretacei 
Pimpinella tragium 1 1 . 1 1 3 . 1 1 1 1 5 2 5 2 1 1 1 2 . 1
Gypsophila oligosperma . 1 . . 1 1 1 . . 1 1 . . . . . . 1 2 1 1
Cephalaria uralensis . . . . . 1 . . . . 1 2 1 2 . 2 . 1 2 1 1
Thymus calcareus . . . 1 1 2 . . 1 1 4 1 3 3 2 2 1 3 2 1 4
Asperula tephrocarpa 1 . 1 1 1 2 1 . 1 1 1 2 2 2 . 1 2 1 1 1 1
Reseda lutea . . . . . . . . . . 1 . . . 1 . 1 . 1 1 .
Silene supina . . . . 1 1 . . . 1 2 1 . . . . 2 1 1 1 .
Linum pallasianum . . . 1 1 . . . . . 2 . 1 1 1 2 2 1 2 1 1
Polygala cretacea . . . . . . 1 1 . 1 1 . 1 2 1 1 1 1 . 1 .
Astragalus albicaulis 1 1 . . . 1 . . . . . . . . . . 1 . . 1 1
Genista tinctoria 1 1 . 1 . . 1 . . . . . 1 . . . 1 . . . 1
Taraxacum serotinum . . . 1 . . . . . . . . . . . . . . . . .
Silene cretacea . . . . . . 1 . . 1 . . . . . . . . . . .
Orthantha lutea . 1 . 1 . . . 1 . . . . . . . 1 . . . . .
Hedysarum ucrainicum . . . . . . . 1 . . . . . . . . . . 2 . .
D. s. of class Festuco-Brometea 
Euphorbia seguieriana 1 . 1 1 . 1 . 1 . . 1 . . . . . . . . . .
Euphorbia nicaeensis . . . . . 1 . . . . . . . . . . . . . . .
Medicago falcata . . . . . . . . . . . . . . . . 1 . . . .
Astragalus onobrychis . . 1 . . . . . . . . . . . . . . . . . .
Plantago lanceolata . . . . . . . . . . . . 1 . . . . . . . .
Appendix 1a: Characteristic table of syntaxa of the alliance Artemisio hololeucae-Hyssopion cretacei.
Priloga 1a: Tabela sintaksonov zveze Artemisio hololeucae-Hyssopion cretacei.
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Consecutive number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21
Another species    
Festuca cretacea . . . 1 . . 1 1 . . . . . . . . . 4 . 2 .
Hedysarum cretaceum . . . . 1 . . . . 2 . . . . . . . . . . .
Galium octonarium . . 1 . . . . . . . . . . . . . . . . . 1
Poa compressa . . 1 . . . 1 . . . . . . . . . . . . . .
Helianthemum canum . . . . . . . . . . . . . . . . . . . . 1
Linum tenuifolium 1 . . . 1 . . . 1 . . . . . . . . . . . .
Odontarrhena tortuosa . . . . . . . . . . . . 1 . . . . . . . .
Diplotaxis cretacea . . . . . . . . . . . . . . . . . . 1 . .
Hedysarum grandiflorum . . . . . . . . . . . . . . . . 3 . . . 1
Androsace villosa . . . . . . . . . . . . . . . . 2 . . . .
Bupleurum falcatum . . . . . . . 1 . . . . . . . . . . . . .
Scabiosa ochroleuca . . . . . . . . . . . . . . . . 1 . . . .
Koeleria pyramidata . . . . . . . . . . . . . . . . 1 . . . .
Astragalus austriacus . . 1 . . . . . . . . . . . . . . . . . .
Jurinea arachnoidea . . . . 1 . . . . . . . . . . . . . . . .
Helichrysum arenarium . . . 1 . . . . . . 1 . . . . . . 1 . . .
Agropyron cristatum 1 . . . 1 . . . . . . . . . . . . . . . .
Melampyrum arvense . . . . . . . . . . . . . . . . . 1 . . .
Elytrigia repens . . 1 . . . . . . . . . . . . . . . . . .
Cytisus ruthenicus . . . . . . . . . . . . . 1 . . . . . . .
Galium humifusum . . 1 . . . . . . . . . . . . . . . . . .
Echinops ritro . . 1 . 1 . . . . . . . . . . . . . . . .
Festuca stricta . . . . . . . . . . . 1 . . . . . . . . .
Erysimum diffusum 1 . . . . . . . . . 1 . . . . . . . . . .
Agrostis capillaris 1 . 3 . . . . . . . . . . . . . . . . . .
Melilotus albus . . . . . . . 1 . . . . . . . . . . . . .
Puccinellia distans . . . . . . 1 . . . . . . . . . . . . . .
Gypsophila fastigiata . . . . . . . . . . . . . . 1 1 1 . . . .
Gypsophila paniculata . . . . . . . . . . . . 1 . . . . . . . .
Localities: 01 – vil. Stril’tsivka, Milovs’kyy district; Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 02 – vil. Stril’tsivka, Milovs’kyy district; Lugansk reg.; Ukraine; 27.06.1987 (Ya. Didukh)
Localities: 03 – Bilovodsk, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 04 – right bank of Aidar River, vil. Rybiancevo, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 05 – vil. Kamyshivka, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 06 – Horodysche, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 07 – vil. Stril’tsivka, Milovs’kyy district;Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 08 – Serebryanka, Donetsk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 09 – Bilovodsk, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 10 – vil. Stril’tsivka, Milovs’kyy district;Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 11 – Derkul River, Markivka, Lugansk reg.; Ukraine; 3.07.1987 (Ya. Didukh)
Localities: 12 – vil. Razdollya, Lugansk reg.; Ukraine; 16.07.2009 (L. Borovyk)
Localities: 13 – Nova Duvanka, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 14 – vil. Zorykivka, Lugansk reg.; Ukraine; 24.06.2009 (L. Borovyk)
Localities: 15 – Dubovyy ravine, Milovs’kyy district; Lugansk reg.; Ukraine; 22.06.2008 (L. Borovyk)
Localities: 16 – vil. Stril’tsivka, Milovs’kyy district; Lugansk reg.; Ukraine; 23.08.2011 (L. Borovyk)
Localities: 17 – “Striltsivskyi steppe” (Luhansk Natural Reserve); Lugansk reg.; Ukraine; 6.07.2012 (L. Borovyk)
Localities: 18 – Derkul River, Markivka, Lugansk reg.; Ukraine; 3.07.1987 (Ya. Didukh)
Localities: 19 – Krasna River, Svatovo, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 20 – Bilovodsk, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 21 – Oskil River, Dvorichna, Kharkiv reg.; Ukraine; 06.1995 (Ya. Didukh)
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Appendix 1a (extension): Characteristic table of syntaxa of the alliance Artemisio hololeucae-Hyssopion cretacei.
Priloga 1a (nadaljevanje): Tabela sintaksonov zveze Artemisio hololeucae-Hyssopion cretacei.
Relevé number 1 2 3 4 5 6 7 8 9 10 11 21 12 13 22 14 23 15 18 19 24
Species richness 8 6 4 21 7 16 6 52 31 4 11 14 10 10 15 13 16 11 10 12 15
Slope (°) 50 70 70 7 5 3 3 5 15
Aspect (°) 45 45 225 250 204 280 289 253 174
Total vegetation cover (%) 30 30 5 30 25 30 15 65 55 25 15 70 12 10 70 10 45 15 25 30 60
D. s. of ass. Erysimo cretacei-Festucetum cretacei  
Erysimum ucranicum 1 1 1 1 1 1 . . . . . . . . . . . . . . .
Melilotus officinalis 1 1 . 1 . 1 1 1 1 . . . . . . . . . . . .
Festuca cretacea 1 2 1 2 1 2 1 4 4 1 1 . . 1 . 1 . . . 1 .
D. s. of ass. Lepidio meyeri-Scrophularietum cretacei 
Lepidium meyeri . . . . . . . . . . 2 . 2 2 . 2 . 2 1 1 .
Hedysarum cretaceum . . . . . . . . . . . 4 . . 4 . 4 . 2 3 4
Galium octonarium . . . 1 . . . 1 1 . 2 1 1 1 1 2 1 2 2 1 1
Artemisia absinthium . . . . . . . . . . . 1 . . 1 . 2 . . . 1
Poa compressa . . . . 1 . . 2 1 . . 1 1 . 1 . 1 . . . 1
Thesium arvense . . . . . 1 . . 1 . . 1 . . 1 . 1 . . 1 1
D. s. of alliance Artemisio hololeucae-Hyssopion cretacei
Scrophularia cretacea 2 2 2 . . 2 2 . . 2 . . 2 1 1 1 1 2 2 . .
Hyssopus officinalis 4 2 1 2 4 4 4 . . 4 2 1 3 2 1 2 1 2 3 2 1
Matthiola fragrans . . . . . . . . . . . . . 1 . 1 . 1 . . .
Linaria cretacea . . . . . . . . . . . . 1 . . . . . . . .
D. s. of alliance Euphorbio cretophilae-Thymion cretacei
Artemisia salsoloides . . . . . . . . . . 2 5 2 2 2 2 2 2 4 4 4
D. s. of alliance Centaureo carbonatae-Koelerion talievii
Stipa capillata . . . . . . . . . . . . . . . . . . . . .
Campanula sibirica . . . . . . . . . . . . . . . . . . . . 1
D. s. of order Thymo cretacei-Hyssopetalia cretacei
Pimpinella tragium 2 4 . 4 1 2 2 2 4 4 2 1 3 2 2 2 2 2 2 2 1
Gypsophila oligosperma 1 . . 2 . 1 . . 1 . 2 4 . . 2 . 4 . 1 2 2
Cephalaria uralensis . . . . . . . . . . 2 1 1 1 . 1 . 1 2 2 1
Reseda lutea . . . 1 . 1 . . 1 . . . . . . . . . . . .
Polygala cretacea . . . 1 . . . 2 1 . . . . . . . . . . . .
Astragalus albicaulis . . . . . . . . . . 1 . 1 1 . 1 . 1 . . .
Taraxacum serotinum . . . . 1 . . . . . . . . . . . . . . . .
Orthantha lutea . . . . . 1 1 . 1 . . . . . . . . . . . .
D. s. of class Festuco-Brometea 
Festuca valesiaca . . . . . . . . . . . . . . 1 . 1 . . . .
Hieracium virosum . . . . . . . . . . . 1 . . . . . . . . .
Medicago falcata . . . . . . . 1 1 . . . . . . . . . . . .
Achillea nobilis . . . . . . . . . . . . . . . . 1 . . . .
Another species    
Artemisia nutans 2 . . . . . . . . . . . . . . . . . . . .
Euphorbia petrophila . . . . . . . . . . . . . . . . . . 1 1 .
Koeleria pyramidata . . . . . . . . . . . . . . 1 . 1 . . . 1
Helichrysum arenarium . . . . . . . . . . 1 1 . . 1 1 1 1 . 1 1
Echinops ritro . . . . . . . . . . 1 . . . . 1 . 1 . . .
Melica transsilvanica . . . . . . . . . . . 1 . . 1 . 1 . . . 1
Poa angustifolia . . . . . . . . . . . 1 . . 1 . 1 . . . 1
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Relevé number 1 2 3 4 5 6 7 8 9 10 11 21 12 13 22 14 23 15 18 19 24
Securigera varia . . . 1 . . . 2 1 . . . . . . . . . . . .
Cichorium intybus . . . . . 1 . . 1 . . . . . . . . . . . .
Poa bulbosa . . . . . . . . . . . . . . . 1 . . . . .
Localities: 01 – right bank of the Don River, Rostov reg.; Russia; 07.2009 (M. Sereda)
Localities: 02 – right bank of the Don River, Rostov reg.; Russia; 07.2009 (M. Sereda)
Localities: 03 – right bank of the Don River, Rostov reg.; Russia; 07.2009 (M. Sereda)
Localities: 04 – Verkhnedonskoy district, 3 km west of the village Demidovskiy, Chayka, Rostov reg.; Russia; 27.06.2009  
(O. Demina)
Localities: 05 – Verkhnedonskoy district, vicinity of village Stogovskoy, Rostov reg.; Russia; 02.06.2008 (O. Demina)
Localities: 06 – Verkhnedonskoy district, vicinity of village Stogovskoy, Belogorskoe hole, Rostov reg.; Russia; 04.06.2009 
(O. Demina)
Localities: 07 – Verkhnedonskoy district, 1km north of the village Stogovskoy, Rostov reg.; Russia; 05.06.2009 (O. Demina)
Localities: 08 – Verkhnedonskoy district, 3 km west from of the Demidovskiy, Chayka, Rostov reg.; Russia; 27.06.2009 
(O. Demina)
Localities: 09 – Chertkovskiy district, 2 km north of the village Mankovo-Kalitvenskoe, Rostov reg.; Russia; 07.06.2009 
(O. Demina)
Localities: 10 – Verkhnedonskoy district, vicinity of village Stogovskoy, Belogorskoe hole, Rostov reg.; Russia; 04.06.2009 
(O. Demina)
Localities: 11 – Zatonsky, Rostov reg.; Russia; 25.08.2014 (O. Demina)
Localities: 21 – Sholokhovskiy district, vicinities of village Zatonskiy, Rostov reg.; Russia; 06.07.2009 (O. Demina)
Localities: 12 – Zatonsky, Rostov reg.; Russia; 25.08.2014 (O. Demina)
Localities: 13 – Zatonsky, Rostov reg.; Russia; 25.08.2014 (O. Demina)
Localities: 22 – Sholokhovskiy district, vicinities of village Zatonskiy, Rostov reg.; Russia; 06.07.2009 (O. Demina)
Localities: 14 – Zatonsky, Rostov reg.; Russia; 25.08.2014 (O. Demina)
Localities: 23 – Sholokhovskiy district, vicinities of village Zatonskiy, Rostov reg.; Russia; 06.07.2009 (O. Demina)
Localities: 15 – Zatonsky, Rostov reg.; Russia; 25.08.2014 (O. Demina)
Localities: 18 – Zatonsky, Rostov reg.; Russia; 25.08.2014 (O. Demina)
Localities: 19 – Zatonsky, Rostov reg.; Russia; 25.08.2014 (O. Demina)
Localities: 24 – Sholokhovskiy district, vicinities of village Zatonskiy, Rostov reg.; Russia; 06.07.2009 (O. Demina)
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Consecutive number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
Total vegetation cover (%) 20 70 80 60 70 60 20 60 31 20 50 60 60 50 50 40 60 60 50 50
Slope (°) 30 15 10 20 8 20 5 40 40 40 30 30 40 30 30 45 40 40
Aspect (°) 135 155 180 225 315 180 135 315 180 135 135 135 135 225 135 180 225 180
Species richness 16 24 19 12 18 15 19 20 29 19 19 18 19 17 19 15 19 19 20 19
D. s. of ass. Euphorbio cretophilae-Jurinetum brachycephalae  
Helianthemum canum 2 . 2 4 . 1 . 4 1 3 . . . . . . . . . .
D. s. of ass. Genisto scythicae-Artemisietum salsoloides 
Genista scythica . . . . . . . . . . 2 1 2 3 1 2 2 2 3 2
Rhaponticoides taliewii . . . . . . . . . . 1 . 1 . 1 1 . 1 1 1
Leontodon biscutellifolius . . . . . . . . . . . 1 . . 1 1 1 . 1 .
Linum tenuifolium . 1 . . . . . . . . . 1 1 . 1 1 1 1 1 1
Odontarrhena tortuosa . 2 . . . . . . . . 1 1 1 1 2 2 1 2 1 1
Linaria vulgaris . . . . . . . . . . . 1 . . 1 . . . 1 .
Diplotaxis cretacea . . . . . . . . . . . . 1 . 1 . 1 1 . 1
Polygala comosa . . . . . . . . . . . . 1 . . . 1 . . 1
Linum hirsutum . . . . . . . . . . 2 2 1 1 1 1 1 1 1 1
D. s. of alliance Artemisio hololeucae-Hyssopion cretacei
Scrophularia cretacea 1 . . . . . . . . 1 . . . . 1 . 1 . . .
Hyssopus cretaceus 1 . . . . . . . . 1 . . . . . . . . . .
Matthiola fragrans . 1 . . . . . . . . . . . . . . . . . .
Linaria cretacea . . 1 1 1 . . . . . . . . . . . . . . .
D. s. of alliance Euphorbio cretophilae-Thymion cretacei
Jurinea stoechadifolia 1 4 4 2 3 4 4 3 1 2 1 3 1 2 3 . 2 1 . 2
Euphorbia petrophila 1 2 1 1 1 1 1 2 1 2 2 2 1 2 2 1 2 2 2 1
Artemisia salsoloides 1 3 . . 3 1 . . . . 2 2 3 2 3 1 2 2 2 2
Brassica elongata 1 . . . 1 1 . 2 1 1 . . . . . . 1 . 1 .
D. s. of alliance Centaureo carbonatae-Koelerion talievii
Campanula sibirica . 1 . . 1 1 1 . 1 . . 1 . . . . 1 . . 1
Carex humilis . . 2 . . . . . 2 1 . . . . . . . . . .
Stipa pennata . . 1 . . . . . . . . . . . . . . . . .
Onosma simplicissima 2 . 2 1 . . . 1 1 1 2 . . . 1 . . 1 2 1
Koeleria talievii . . . . . . 1 . 2 . . . . . . . . . . .
Psephellus marschallianus . 1 . . . . 1 1 1 . . . . . . . . . . .
Teucrium polium . 1 2 . 1 1 1 . 1 1 1 2 2 1 1 1 1 1 1 1
D. s. of order Thymo cretacei-Hyssopetalia cretacei
Pimpinella tragium 1 1 . 1 1 3 1 1 1 1 3 1 2 2 2 2 3 2 1 2
Gypsophila oligosperma 1 1 3 1 1 1 2 . 1 2 1 1 1 1 1 1 1 1 1 1
Cephalaria uralensis . 1 1 1 . . . 1 . . 1 1 1 2 1 1 1 1 1 .
Thymus calcareus 4 4 2 3 5 . 1 2 4 4 1 2 2 2 2 3 3 3 2 2
Asperula tephrocarpa 1 1 1 3 2 1 1 1 1 1 . . . . . . . . . .
Reseda lutea . 1 1 . . 1 . . . . 2 1 1 1 1 . 1 1 2 1
Silene supina . 1 . . . . . 1 . . . . . . . . . . . .
Linum pallasianum 2 1 2 . . . . 1 . 2 . . . . . . . . . .
Orthantha lutea 1 . . . . . 1 . 1 1 . . . . . . . . . .
Scutellaria supina . . . . . . 1 . . . . . 1 . . . . . 1 .
Linaria genistifolia . . . . . . . . 1 . . . . . . . . . . .
Appendix 1b: Characteristic table of syntaxa of the alliance Euphorbio cretophilae-Thymion cretacei.
Priloga 1b: Tabela sintaksonov zveze Euphorbio cretophilae-Thymion cretacei.
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Localities: 01 – vil. Zakotnoe, Donetsk reg.; Ukraine; 08.1994 (K. Romashchenko)
Localities: 02 – vil. Nyzhnye, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 03 – “Kreidova flora”, Donetsk reg.; Ukraine; 08.2016 (O. Chusova)
Localities: 04 – “Kreidova flora”, Donetsk reg.; Ukraine; 08.2016 (O. Chusova)
Localities: 05 – “Kreidova flora”, Donetsk reg.; Ukraine; 08.2016 (O. Chusova)
Localities: 06 – vil. Kryva Luka, Donetsk reg.; Ukraine; 22.06.1987 (Ya. Didukh)
Localities: 07 – vil. Bogorodychnoe, Donetsk reg.; Ukraine; 08.1994 (K. Romashchenko)
Localities: 08 – vil. Kryva Luka, Donetsk reg.; Ukraine; 22.06.1987 (Ya. Didukh)
Localities: 09 – vil. Bogorodychnoe, Donetsk reg.; Ukraine; 08.1994 (K. Romashchenko)
Localities: 10 – “Kreidova flora”, Donetsk reg.; Ukraine; 28.08.1994 (K. Romashchenko)
Localities: 11 – right bank of Tuzlov River, vil. Lysogorka, Rostov reg.; Russia; 4.06.2002 (M. Sereda)
Localities: 12 – right bank of Tuzlov River, vil. Lysogorka, Rostov reg.; Russia; 4.06.2002 (M. Sereda)
Localities: 13 – right bank of Tuzlov River, vil. Lysogorka, Rostov reg.; Russia; 4.06.2002 (M. Sereda)
Localities: 14 – right bank of Tuzlov River, vil. Lysogorka, Rostov reg.; Russia; 4.06.2002 (M. Sereda)
Localities: 15 – right bank of Tuzlov River, vil. Lysogorka, Rostov reg.; Russia; 4.06.2002 (M. Sereda)
Localities: 16 – right bank of Tuzlov River, vil. Lysogorka, Rostov reg.; Russia; 4.06.2002 (M. Sereda)
Localities: 17 – right bank of Tuzlov River, vil. Lysogorka, Rostov reg.; Russia; 4.06.2002 (M. Sereda)
Localities: 18 – right bank of Tuzlov River, vil. Lysogorka, Rostov reg.; Russia; 4.06.2002 (M. Sereda)
Localities: 19 – right bank of Tuzlov River, vil. Lysogorka, Rostov reg.; Russia; 4.06.2002 (M. Sereda)
Localities: 20 – right bank of Tuzlov River, vil. Lysogorka, Rostov reg.; Russia; 4.06.2002 (M. Sereda)
Consecutive number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
D. s. of class Festuco-Brometea 
Euphorbia seguieriana . . . . . . 1 1 1 1 . . . . . . . . . .
Galium ruthenicum . . . . . . 1 . . . . . . . . . . . . .
Potentilla recta . . . . . . 1 . 1 . . . . . . . . . . .
Astragalus sulcatus . . . . . . . . 1 . . . . . . . . . . .
Adonis volgensis . . . . . . . . 1 . . . . . . . . . . .
Another species    
Polygala cretacea . . . . 1 . . . . . . . . . . . . . . .
Convolvulus lineatus . 1 . . . . . . . . . . . . . . . . . .
Thesium arvense . 1 1 . . . . . . . . . . . . . . . . .
Artemisia hololeuca . . . . . . . . . . . . . . . . . . . .
Allium inaequale . . . . . . 1 . 1 . . . . . . . . . . .
Hedysarum grandiflorum 1 1 1 . . . . . . . 1 1 . 1 1 . . 1 . 1
Asperula cynanchica . . . . . . . . . . 1 . . . . . . 1 1 .
Salvia verticillata . . . . . . . 1 . . . . . . . . . . . .
Caragana frutex . . . . . . . 1 . . . . . . . . . . . .
Centaurea orientalis . 1 . . . . . . . . . . . . . . . . . .
Bupleurum falcatum . . . 1 1 . 1 . 1 . . . . . . . . . . .
Anisantha tectorum . . . . . . . . 1 . . . . . . . . . . .
Scabiosa ochroleuca 1 . . . . . 1 . 1 1 . . . . . . . . . .
Koeleria pyramidata . . . . . 1 . 1 . . . . . . . . . . . .
Bromopsis riparia . . . . . . . 1 . . . . . . . 1 . . . .
Salvia nutans . . . . . . . 1 1 . . . . . . . 1 1 . .
Thalictrum minus . 1 . . 1 . . 1 . . . 1 1 1 . . . . . 1
Helichrysum arenarium . . 1 1 1 1 . . 1 . . . . . . . . . . .
Cytisus ruthenicus . . . . . . 1 . 1 . . . . . . . . . . .
Viola ambigua . . 1 . 1 . . . . . . . . . . . . . . .
Echinops ritro . . . . . . . 2 . . . . . . . . . . . .
Stipa lessingiana . . . . . . . . . . 1 . . 1 . . . . . .
Erysimum diffusum . . . . . . . . . . . . 1 1 . 1 . . . .
Ephedra distachya . 1 . . . . . . . . . . . . . . . . . .
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Consecutive number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30
Total vegetation cover (%) 70 70 60 90 60 65 65 70 50 70 40 60 50 65 65 45 50 60 80 40 15 40 40 60 10 10 60 80 40
Slope (º) 15 45 45 20 5 10 20 25 30 20 10 5 10 7 7 5 7 5 10 30 25 30 7 25 30 30 5 20
Aspect (°) 45 45 45 270 5 90 90 90 5 180 225 180 180 270 315 135 225 135 180 180 90 180 180 135 45 225 135
Species richness 28 25 23 22 22 23 27 20 31 19 26 46 31 32 37 40 52 41 74 48 15 11 20 12 18 22 11 21 23 15
D.s. ass. Pimpinello titanophillae-Artemisietum salsoloides 
Artemisia salsoloides 5 1 4 1 3 5 . . 3 4 . . . . 1 . . . . . . . . . . . . . . .
Polygala cretacea 1 1 1 . 1 . . . . . 1 1 . . 1 . 1 . . 1 1 . . . . . . 1 . .
Linum hirsutum 1 2 . . 2 . . 1 . . . . . . 1 1 1 . . . 1 . . 1 . . . . 1 .
D.s. ass. Hedysaro grandiflori-Psephelletum carbonatae
Allium inaequale . . . . . . . . . . . 1 1 2 . 1 2 1 1 2 . . . . . . . . . .
Hedysarum grandiflorum . . . 4 . . . 3 . . 2 3 1 1 3 1 1 1 3 1 . . . . . . . . . .
Thymelaea passerina . . . . . . . . . . 1 . 1 1 . . . . 1 . . . . . . . . . . .
Asperula cynanchica . . . . . . . . . . 1 1 1 1 . 1 1 1 1 1 . . . . . . . . . .
Adonis vernalis . . . . . . . . . . . 1 . 1 1 1 . . 3 2 . . . . . . . . . .
Seseli annuum . . . . . . . . . . . . . . . 1 . . 1 . . . . . . . . . . .
Pimpinella saxifraga . . . . . . . . . . . 1 . . 1 . . . 1 . . . . . . . . . . .
Phlomis herba-venti . . . . . . . . . . . 1 . . . . . . 1 1 . . . . . . . . . .
Salvia verticillata . . . . . . . . . . 1 . 1 1 . 1 1 . 1 1 . . 1 . . . . . . .
Aster amellus . . . . . . . . . . . . . . . 1 1 . . . . . . . . . . . . .
Asparagus officinalis . . . . . . . . . . . . . . . . 1 . 1 1 . . . . . . . . . .
Caragana frutex . . . . . . . . . 1 . 1 . . . . 1 1 . 2 . . . . . . . . . .
Centaurea orientalis . 1 1 1 . . . 1 . . . . . . 2 . 1 . 1 1 . . . . . . . . . .
D.s. ass. Androsacio koso-poljanskii-Caricetum humilis 
Androsace villosa . . . . . . . . . . . . . . . . . . . . 3 2 3 4 3 4 2 3 4 4
D. s. of alliance Artemisio hololeucae-Hyssopion cretacei
Artemisia hololeuca . . . . . . . . 1 . . 3 . . 3 . 1 . . . 3 . . . 4 . . 3 . .
Scrophularia cretacea . . 1 . . . . . . . . . . . 1 . . . . . 2 . . . . . . 2 . .
Hyssopus cretaceus . . . . . . . . . . . 3 . . . . . . . . . . . . . . 1 . . .
Matthiola fragrans . 1 . . . . . . . . . . . . . . . . . . . 2 1 . . . . . . .
D. s. of alliance Euphorbio cretophilae-Thymion cretacei
Brassica elongata . . . . . . . . 1 . 1 . . 1 . . . . . . . . . . . . . . . .
D. s. of alliance Centaureo carbonatae-Koelerion talievii
Stipa capillata 1 . . . . . . 1 . . 1 1 1 1 1 . 1 1 3 2 . . . . . 2 . . 1 1
Campanula sibirica . 1 . . 1 1 1 1 1 . 1 . 1 1 . 1 1 1 1 1 1 . . . . 1 . 1 . .
Polygala sibirica . 1 . . . . . . 1 . 1 . . . 1 . . . . . . . 2 . 1 1 . 1 . .
Carex humilis . . . . . . . . 1 . . 1 1 5 4 4 4 3 3 3 . . . . . 4 . 3 4 1
Stipa pennata . . . . . . . . . . 2 1 1 1 1 1 . 1 . 3 . . . . . . . . . .
Onosma simplicissima 1 1 . . 3 . . 1 1 . 2 1 . 1 3 3 . 1 3 2 . . . . . . . 2 1 .
Koeleria talievii . . . . . . . . . . 1 1 . 1 . . 1 1 1 . . . . . . . 1 . . 1
Psephellus marschallianus 1 . . . 2 . 1 . . . . . 1 . 1 1 1 1 1 1 . . 1 . . . . . . .
Teucrium polium 1 1 2 1 3 1 1 1 1 1 1 1 1 1 3 1 1 2 1 1 . 1 1 . 1 1 . 1 1 .
D. s. of order Thymo cretacei-Hyssopetalia cretacei
Pimpinella tragium . 3 1 . . 1 4 1 1 1 1 1 1 1 . . 1 . . . 2 1 1 1 2 1 2 2 . 1
Gypsophila oligosperma 2 3 1 1 1 1 . . 1 . 1 . 1 1 1 1 1 4 1 1 . . 1 1 1 1 1 . 1 .
Cephalaria uralensis 1 . 1 . 1 . . . 1 1 2 . 2 1 . . 1 1 1 . . 1 . . . . . . . .
Thymus calcareus 1 4 1 1 . 3 3 3 3 . 3 1 4 1 3 2 1 1 3 . 3 3 4 4 1 1 2 3 . 1
Asperula tephrocarpa 1 1 1 . 2 1 2 3 1 1 . 1 . . . . 1 . . . 2 . 2 1 . 1 1 . . .
Reseda lutea . 1 . . . . . 1 . . 1 1 1 . 3 . 1 1 1 . . 1 . 1 . . . . . .
Appendix 1c: Characteristic table of syntaxa of the alliance Centaureo carbonatae-Koelerion talievii.
Priloga 1c: Tabela sintaksonov zveze Centaureo carbonatae-Koelerion talievii.
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Consecutive number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30
Silene supina 1 . 1 . . . 1 . . . 1 1 . . . . . . . . . . 1 . 1 1 . . 1 .
Linum pallasianum . . . . 2 . . 1 1 . 1 2 . . 2 1 1 1 1 1 2 2 1 1 . 1 . 2 . .
Astragalus albicaulis 2 1 2 . 1 . . 1 1 . 1 1 . . 3 . . . 3 . 1 . . . . . . . . .
Genista tinctoria 1 2 1 . . . . . 1 . . 1 . . . . 1 1 1 1 . . . . . . . . . .
Taraxacum serotinum . . . . . . . . . . . . . 1 . . . . 1 . . . . . . . . . . .
Hedysarum cretaceum                . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Helianthemum canum . . . . . . . . . . . . 1 1 . 1 . . 1 . . . . . . . . . . .
Linum tenuifolium 1 . 1 . . 1 1 . 1 1 . . . . . . . . . . . . . . . . . . . .
Odontarrhena tortuosa . . . . 2 1 . . . . . 1 . . 1 . . . . . . . . . . . . . . .
Bupleurum falcatum . 1 . . 2 1 . . 1 . 1 . 1 2 1 1 1 . 1 1 . . . . 1 . . 1 . .
Jurinea arachnoidea 1 1 . 1 . . 1 . . 1 1 . . . . 1 . . . . . . 1 . . . . . . .
D. s. of class Festuco-Brometea
Festuca valesiaca 3 . . 5 2 1 . . 1 3 . . . . 4 1 1 1 1 . . . . 1 . 2 . . . 1
Euphorbia seguieriana 1 1 1 1 3 2 1 1 1 1 1 . 1 1 . 1 1 1 1 1 . . . 1 1 1 1 1 1 1
Euphorbia nicaeensis . . . . . 1 . . 1 . . . . . . . . . . . . 1 . . . . . . . .
Stachys recta . 1 1 . . . . . . . . . . 1 . . 1 1 1 1 . . . . 1 . . . . .
Medicago falcata 1 . . . 2 . 1 . . . . . . . . 1 1 1 1 1 . . 1 . 1 1 1 . 2 .
Astragalus onobrychis . . . . . . 1 . . . . 1 . . . . . . 1 1 . . 1 . . . . . . .
Onobrychis arenaria . . . . . . . . . . . . . . . 1 . . 1 . . . . . . . . . 1 .
Scabiosa ochroleuca . . . . . . . . . . . . 1 1 . 1 . 1 1 . 2 . . . . . . 1 . 1
Bromopsis riparia . . . . . . 1 1 . . 1 . 1 1 . . 1 1 . . . . . 1 . 1 1 . . .
Salvia nutans . . . . 2 2 1 . . 3 . 1 . . 1 . 1 1 1 2 . . . . . . . 1 1 1
Veronica spicata . . . . . . . . 1 1 . . . . . . . 1 . 1 . . . . . . . . . .
Astragalus austriacus . . . . . . . . . . . . . 1 . . 1 1 1 1 . . 1 . 1 1 . . 1 .
Thalictrum minus 1 . . 1 1 . . 1 1 . . 1 1 2 . . . 1 1 1 . . . . . . . . . .
Another species
Helichrysum arenarium 1 1 1 1 . 1 . . 1 . . . . . . . . . . . . . . . . . . . . .
Agropyron cristatum 1 . 1 3 . . 1 . . 1 . . . . . . . . . . . . . . . . . . . .
Galium octonarium 1 2 2 3 . 1 . . 1 2 . . . . . 1 1 . . 1 . . . . . . . . . .
Thesium arvense 1 1 . . . . 1 . 1 . . . . . . . . . 1 . . . 1 1 1 1 . . 1 .
Melampyrum arvense . . . 1 3 . . . 1 . . . . . . 1 1 . 1 . . . . . . . . . . .
Elytrigia intermedia 1 3 4 . . . . . . . . . . . . . . . . . . . . . . 2 . . . .
Echinops ritro . . . . . 1 . . . . . . . . . 1 . . 1 1 . . . . . . . . . .
Stipa lessingiana . . . . . . . . . 3 . 1 . . 1 . . . . . . . . . . . . . . .
Stipa pulcherrima . . . 2 . 1 1 . . 2 . . . . . . . . . . . . . . . . . . 4 .
Securigera varia . . 1 1 . . . . . . . . . . 1 . 1 1 1 1 . . 2 . 1 1 . . . .
Erysimum diffusum 1 . . 1 1 . 1 . . 1 1 . . 1 . 1 1 . . 1 . . . . . . 1 . . 1
Gypsophila fastigiata . . . . . . . . . . . . . . . . . . . . 2 1 . . . . . 1 . .
Helianthemum nummu-
larium . . . . . . . . . . . . . . . 1 . . 1 1 . . . . . . . . . .
Anthericum ramosum . . . . . . . . . . . . . . . 1 . . 1 2 . . . . . . . . 1 .
Stipa zalesskii . . . . . . . . . . . . . . . . . . . . 2 . . . . . . 2 . .
Localities: 11 – vil. Baranikivka, Lugansk reg.; Ukraine; 07.1987 (Ya. Didukh)
Localities: 12 – vil. Baranikivka, Lugansk reg.; Ukraine; 07.1987 (Ya. Didukh)
Localities: 13 – vil. Baranikivka, Lugansk reg.; Ukraine; 07.1987 (Ya. Didukh)
Localities: 14 – 5 km South of the Bilovodsk, Lugansk reg.; Ukraine; 07.1987 (Ya. Didukh)
Localities: 15 – vil. Nyzhnya Duvanka, Lugansk reg.; Ukraine; 13.06.2013 (O. Chusova)
Localities: 16 – right bank of Derkul River, Lugansk reg.; Ukraine; 07.1987 (Ya. Didukh)
Localities: 17 – right bank of Kamyshna River, Lugansk reg.; Ukraine; 07.1987 (Ya. Didukh)
Localities: 18 – right bank of Kobylka River, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 19 – vil. Stril’tsivka, Milovs’kyy district; Lugansk reg.; Ukraine; 07.1987 (Ya. Didukh)
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Localities: 10 – 5 km South of the Bilovodsk, Lugansk reg.; Ukraine; 07.1987 (Ya. Didukh)
Localities: 11 – Oskol River, Belogorie Reserve, Belgorod reg.; Russia; 4.08.2012 (E. Averinova)
Localities: 12 – border between Rossoshinsky and Kantemirovsky districts, Voronezh reg.; Russia; 1.08.1983 (A. Grigorevskaya*)
Localities: 13 – vil Makeshkino, Belgorod reg.; Russia; 1.08.2012 (E. Averinova)
Localities: 14 – vil Makeshkino, Belgorod reg.; Russia; 1.08.2012 (E. Averinova)
Localities: 15 – 0.5 km north-west of the Veidelevka, Belgorod reg.; Russia; 24.08.2012 (E. Averinova)
Localities: 16 – Oskol River, Belogorie Reserve, Belgorod reg.; Russia; 3.08.2012 (E. Averinova)
Localities: 17 – 1 km south-east of the Roven’ki Belgorod reg.; Russia; 6.08.2011 (E. Averinova)
Localities: 18 – 1 km west of the Berezhnyi Farm, Belgorod reg.; Russia; 9.08.2011 (E. Averinova)
Localities: 19 – Oskol River, Belogorie Reserve, Belgorod reg.; Russia; 25.08. 1983 (A. Grigorevskaya*)
Localities: 20 – Oskol River, Belogorie Reserve, Belgorod reg.; Russia; 4.08.2012 (E. Averinova)
Localities: 21 – vil. Kamyanka, Kharkiv reg.; Ukraine; 27.05. 2012 (O. Bezrodnova)
Localities: 22 – vil. Dvorichna, Kharkiv reg.; Ukraine; 26.05. 2012 (O. Bezrodnova)
Localities: 23 – Rogan’, Kharkiv reg.; Ukraine; 14.06.1996 (Ya. Didukh)
Localities: 24 – Rogan’, Kharkiv reg.; Ukraine; 14.06.1996 (Ya. Didukh)
Localities: 25 – vil. Yaichne, Kharkiv reg.; Ukraine; 06.1996 (Ya. Didukh)
Localities: 26 – south of the Rogan’, Kharkiv reg.; Ukraine; 05.1995 (Ya. Didukh)
Localities: 27 – vil. Staromelovoe, Kursk reg.; Russia; 2.08. 2005 (A. Poluyanov)
Localities: 28 – vil. Kamyanka, Kharkiv reg.; Ukraine; 27.05. 2012 (O. Bezrodnova)
Localities: 29 – Pechenigy, Kharkiv reg.; Ukraine; 15.06.1996 (Ya. Didukh)
Localities: 30 – vil. Kamyanka, Kharkiv reg.; Ukraine; 21.08. 2012 (O. Bezrodnova)
* – relevés were taken from Averinova (2014).
Appendix 1c (extension): Characteristic table of syntaxa of the alliance Centaureo carbonatae-Koelerion talievii.
Priloga 1c (nadaljevanje): Tabela sintaksonov zveze Centaureo carbonatae-Koelerion talievii.
Consecutive number 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
Total vegetation cover (%) 70 70 50 45 30 75 30 30 50 60 40 50 45 70 70 85 65 60 80 60 50 65 40 70 50 20 75 50 40 60
Slope (°) 10 15 40 10 5 15 10 15 40 10 10 25 7 22 10 15 20 22 10 20 15 15 20
Aspect (°) 70 45 180 90 5 90 90 5 5 270 5 180 45 45 135 225 45 180 180 180 180 180 180
Species richness 17 25 31 31 25 24 18 14 26 21 30 22 26 25 36 33 27 27 40 25 24 31 30 28 26 27 29 23 27 29
D. s. of ass. Bupleuro falcatae-Stipetum capillatae 
Bupleurum falcatum 1 1 1 1 1 1 1 1 1 . . 1 1 . . . . 1 . . 1 . 1 1 1 1 1 1 1 1
Anisantha tectorum . . 1 1 . . 1 . . . . . . . . . . . . . . . . . . . . . . .
Scabiosa ochroleuca 1 1 1 1 1 . 1 1 1 3 . . . . . . . . . 2 . . 1 1 1 1 1 1 1 .
D. s. of ass. Gypsophilo oligospermae-Campanuletum sibiricae 
Carex pediformis . . . . . . . . . . 3 3 3 2 . . . . . . . . . . . . . . . .
Koeleria pyramidata . . . . . . . . . 1 2 1 2 2 1 1 . . . 3 . . . . . . . . . .
Nonea pulla . . . . . . . . . . . . 1 1 . . . 1 . 2 . . . . . . . . . .
Bromopsis riparia . . . . . . . . . 2 2 . 3 4 1 2 . 1 1 . 1 . . 2 . . 1 . . .
Salvia nutans . . 1 1 1 1 1 2 1 . 1 . 3 2 1 2 1 1 3 3 . 1 . . 1 . 1 . . 1
Galatella villosa . . . . . . . . . . 4 . 1 1 . 4 . . 1 2 . . . . . . . . . .
Linum hirsutum 1 1 . 1 . . . . . . 1 2 2 2 1 . 2 . . . . . . . . . . . . .
D. s. of ass. Carici humilis-Thymetum calcarei 
Androsace villosa . . . 1 . . . . . . . . . . . . 1 3 . . 4 1 3 4 . . 3 . . .
Astragalus austriacus . . 1 . . . . . . . . . . . . . 2 1 . . 1 1 . . . . . . . 1
Thalictrum minus . 1 . . 1 1 . 1 1 . . . 1 1 1 . 2 1 1 . . 1 . . . 1 1 . . .
Oxytropis pilosa . . . . . . . . . . . . . . . 1 . . . 1 . . 1 . . . . . . 1
Jurinea arachnoidea . . . . . . . . . . 2 . . . 1 1 2 . 1 . . . . . . 1 . . . .
Viola rupestris . . . . . . . . . . . . . . . . . . . . 1 . 1 . . . . 1 1 1
Helianthemum nummularium . . . . . . . . 1 . . . . . . . . . . . 1 1 1 . 1 1 1 . 1 .
Asperula tinctoria . . . . . . . . . . . . . . . . . . . . . . 1 . . . 1 1 . 1
Anthericum ramosum . . . . . . . . . . . . . . . . . . . . . 3 1 . 1 2 1 . 3 .
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Consecutive number 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
Linum perenne . . . . . . . . . . . . . . . . . . . . . . . . 1 . . 1 . .
Vincetoxicum hirundinaria . . . . . . . . . . . . . . . . . 1 . . . 1 . . 1 1 . 1 1 1
Euphrasia pectinata . 1 1 . 1 . . . . . . . . . . . . . . . 1 . . 1 . . 1 . 1 1
D. s. of alliance Artemisio hololeucae-Hyssopion cretacei
Artemisia hololeuca . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . .
D. s. of alliance Euphorbio cretophilae-Thymion cretacei
Brassica elongata . 1 . 1 . 1 . . . 1 2 2 . 1 . . . . . . . . . . . . . . . .
Jurinea stoechadifolia . . . . . 2 1 . 1 . . . . . . . . . . . . . . . . . . . . .
Euphorbia petrophila . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . .
D. s. of allliance Centaureo carbonatae-Koelerion talievii
Stipa capillata . 1 1 1 . . 4 4 4 1 2 . 2 5 4 . 1 . . 3 . . 1 . 1 1 1 1 1 3
Campanula sibirica 1 1 1 1 1 1 . . . 2 1 1 2 2 1 2 . 1 1 3 1 . 1 1 1 1 . 1 1 .
Polygala sibirica . . . 1 1 . . . . . 1 . 1 . 1 1 1 1 . 1 . . 1 1 1 1 . . 1 .
Carex humilis . . 1 1 4 2 1 . . 4 . 1 . . 5 2 4 4 . 4 2 2 3 2 3 3 2 3 3 3
Stipa pennata . . . . . . . . . . . . . . . 3 . . . . 1 3 1 1 . . . . . 3
Onosma simplicissima 2 . 1 2 1 . . 1 1 . . 2 . . 1 2 2 1 1 . . 3 2 . 3 3 1 2 2 .
Koeleria talievii . 1 1 1 1 1 . . 1 . . . . . . . . . . . 1 . . 1 . . . 1 . 1
Psephellus marschallianus . . . . 1 2 1 . 1 . . . . . 1 . 1 1 1 . 1 1 1 . . 1 1 . 1 1
Teucrium polium 1 . 1 1 . 2 1 1 1 2 2 2 2 1 1 1 1 1 1 . . . . . . . . . . .
D. s. of order Thymo cretacei-Hyssopetalia cretacei
Pimpinella tragium 1 1 1 1 . 1 . . . . . 2 . . 1 1 . 1 1 . 1 . 1 1 . 1 1 . . .
Gypsophila oligosperma 1 2 1 1 1 . . . . 2 . 1 . 1 . . . . . 1 1 1 1 1 1 1 1 1 1 1
Cephalaria uralensis . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . .
Thymus calcareus 4 4 3 4 2 1 . . . 2 . 4 . . 1 . 1 1 . . 1 1 2 2 3 1 4 3 1 1
Asperula tephrocarpa 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 . . . . . . . . . . .
Reseda lutea 1 1 1 1 . . . . . 2 2 3 1 1 . 1 . . 1 2 . . . . . . . . . .
Silene supina . . 1 . . . . . . . 2 1 1 1 . . . . . 1 . . . . . . . . . .
Linum pallasianum 1 1 2 2 1 . . . . 2 2 1 3 1 1 2 . 1 1 . . . 1 . . . 1 . . 1
Polygala cretacea 1 1 . . . . . . . . . . . . 1 . 2 . . . . 1 . . . 1 . . . .
Astragalus albicaulis 1 1 1 1 . . . . . . . . . . . . . . . . . 1 . . 1 2 . . 1 2
Genista tinctoria . . . . . . . . . . 2 . 1 . . 3 . . . 3 . . . . . . . . . .
Orthantha lutea 1 1 1 1 1 1 1 . 1 . . . . . . . . . . . . . . . . . . . . .
Helianthemum canum . 1 1 1 . . . . . . . 4 . . . . . . . . . . . . . . . . . .
Odontarrhena tortuosa . . . . 1 . . . . . . 3 . . 1 . . . . . . 1 . . . . . . . 1
Diplotaxis cretacea . . . . . 1 1 1 . . . . . . . . . . . . . . . . . . . . . .
Artemisia salsoloides . . . . . . . . . . . . . . 2 . . . . . . . . . . . . . . .
Plantago maritima . . . . . . . . . . . . . . . 2 . . . . . . . . . . . . . .
D. s. of class Festuco-Brometea
Festuca valesiaca . 1 2 . . 1 . . 1 . 4 1 4 3 1 2 3 1 1 3 . 2 1 1 . . 2 1 1 1
Euphorbia seguieriana . 1 . 1 1 1 . . 2 3 . 2 1 2 1 1 2 1 1 2 . 1 1 1 1 1 1 1 1 1
Hieracium virosum . . . . . . . . . . 1 . . . . 1 . . . 2 . . . . . . . . 1 .
Euphorbia nicaeensis . . . 1 . . . . . . 3 . . . 1 2 . . . . . . . . . . . . . .
Stachys recta . . 1 1 1 . . . . . 2 . 2 2 . . 2 . 1 . . . . . 1 . . . . .
Medicago falcata . . 1 . . . . . . . 1 . . . . . 2 . 2 . 1 . 1 . . . . 1 1 .
Potentilla humifusa . . . . . . . . . . . . 1 1 . . 1 1 . . . 1 1 1 . 1 . . . .
Rhaponticoides ruthenica . . . . . . . . . . . . . . . 2 1 . . . . . . . . 1 . . 1 .
Onobrychis arenaria . 2 . . . . . . . . 1 . . . . 1 . . 1 . 1 . . 1 . . . 1 . .
Plantago media . . . . . . . . . . 3 . . . . . . . . . . . . 1 . . . . . .
Achillea setacea . . 1 . . . . . . . 1 . . . . 1 . . . . . . . . . . . . . .
Potentilla incana . . . . . . . . . . . . . . 1 1 . . . . . . . . . . . 2 . 2
Potentilla recta . . . . . 1 . . 1 . . . . . . 1 . . 1 . . . . . . . . . . .
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Consecutive number 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
Iris pumila . . . . . . . . . . . . 1 . . . . . 1 . . . . . . . . . . .
Linum austriacum . . . . . . . . . . . . . . . 1 1 . 1 . . . . . . . . . . .
Adonis volgensis . . 1 . . . . . . . . . . . 1 . . . . . . . . . . . . . . .
Hypericum elegans . . . . . 1 . . . . . . 1 . 1 . . . . . . . . . . . . . . .
Veronica incana . . . . . . . . . . 2 . . 1 . 2 . . . 1 . . . . . . . . . .
Klasea radiata . . . . . . . . . . . . . . . 1 . . . 1 . . . . . . . . . .
Another species  
Allium inaequale . . . 1 1 1 1 1 1 2 . . . . . . . . . . . . . . . . . 1 . .
Hedysarum grandiflorum . . . . . . . . . 2 . . . . . . . 1 . . . . . . . . . . . .
Asperula cynanchica . . . . . . . . . . . . . . . . . . . . 1 1 . . . 1 . . 1 .
Adonis vernalis . . . . . . . . . . . . . . . . 1 1 . . . 1 . . 1 . . . . .
Salvia verticillata 1 . . . . . . . . . . . . . . . . . . . . . 1 . 1 . . . . .
Inula aspera . . . . . . . . . . . . . . . 2 . 1 . . . . . . . . . . . .
Caragana frutex . . . . . 1 . . 1 . . . . . . . . . . . . . . . . . . . . .
Centaurea orientalis . . . . . . . . . . . . . . . . . . 3 . . . . . . . . . . .
Pilosella officinarum . . . . . . . . . . . . . . . . . . . . 1 . 1 . 1 . . . . 1
Bassia prostrata . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . .
Festuca cretacea . . . . 2 . . . . . . . . . . . . . . . . . . . . . . . . .
Hedysarum cretaceum . . . . . . . . . . . . . . . 2 1 . . . . . . . . . . . . .
Galium octonarium . . . . . . . . . . . . . . 1 . . 1 1 . . . . . . . . . . .
Poa compressa . . 1 1 . . . . 1 . . . . . 1 . . . . . . . . 1 . . 1 . . .
Thesium arvense . . . . . . . . . . . 1 . . 1 . 1 1 1 . . . . . . . . . . .
Helichrysum arenarium . . . 1 1 1 1 . 1 3 2 . 1 . 1 1 . . . 1 . . . . . . . . . .
Agropyron cristatum . . . . . 1 . 2 . . . . . . . . . . . . . . . . . . . . . .
Melampyrum arvense . . . . . . . . . . . . . . . 1 . . . . . . . . . 1 . . 1 .
Cytisus ruthenicus . . . . . . . . . . . . . . 1 . . . . . . 1 . . . . . . . .
Centaurea diffusa . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . .
Viola ambigua . . . . . . . . . 2 . . . . . 1 2 1 . . . . . . 1 . . . . .
Hypericum perforatum . . . . . . 1 . 1 . 1 . . . . . . . . . . . . . . . . . . .
Rhamnus cathartica . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . 1 .
Polygonatum odoratum . . . . . . . . . . . . . . . . . . . . . 1 . . . 1 . . . .
Hieracium umbellatum . . 1 . 1 . . . . . . . . . . . . 1 . . 1 . . 1 . . . . . .
Echinops ritro . . . . . . . . . . . . . . . 2 1 . . . . . . . 1 . . . . .
Stipa pulcherrima . . . . . . . . . . . . . . 1 . . 2 5 . . . . . . . . . . .
Festuca stricta . . . . . . . . . . . . . . . 2 1 . 2 . . . . . . . . . . .
Astragalus subuliformis . . . . . 1 1 1 . . . . . . . . . . . . . . . . . . . . . .
Securigera varia . . 1 . . . . . . . 2 . . 1 . . . . 1 2 . . 1 . . . . . . .
Erysimum diffusum . 1 1 1 . 1 . . . . . . . . . . . . 1 . . 1 . . . 1 . 1 . 1
Jacobaea vulgaris . . . . . 1 1 . . . . . . . . 1 1 . . . . . . . . . . . . .
Silene chlorantha . . . . . 1 1 . . . . . . . . . . . . . . . . . . . . . . .
Gypsophila fastigiata . . . . . . . 1 . . . . . . . . . 1 . . . . . . . . . . . .
Potentilla patula . . . . . . . . 2 2 . . . . . . . . . . . . . . . . . . . .
Poa bulbosa . . . . . 1 . 1 . . . . . . . . . . 1 . . . . . . . . . . 1
Localities: 21 – right bank of Oskol River, vil. Znamenka, Kharkiv reg.; Ukraine; 21.08.1994 (K. Romashchenko)
Localities: 22 – right bank of Oskol River, vil. Znamenka, Kharkiv reg.; Ukraine; 21.08.1994 (K. Romashchenko)
Localities: 23 – right bank of Oskol River, vil. Znamenka, Kharkiv reg.; Ukraine; 9.08.1994 (K. Romashchenko)
Localities: 24 – right bank of Oskol River, vil. Topolya, Kharkiv reg.; Ukraine; 08.1994 (K. Romashchenko)
Localities: 25 – right bank of Oskol River, vil. Nyzhnya Melnitsa, Kharkiv reg.; Ukraine; 08.1994 (K. Romashchenko)
Localities: 26 – vil. Lavrentievka, Donetsk reg.; Ukraine; 1.08.1994 (K. Romashchenko)
Localities: 27 – vil. Kryva Luka, Donetsk reg.; Ukraine; 26.08.1994 (K. Romashchenko)
Localities: 28 – vil. Kryva Luka, Donetsk reg.; Ukraine; 26.08.1994 (K. Romashchenko)
Y. Didukh, O. Chusova & O. Demina
Syntaxonomy of chalk outcrop vegetation of the order  
Thymo cretacei-Hyssopetalia cretacei
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17/1 • 2018, 85–109
Localities: 29 – vil. Kryva Luka, Donetsk reg.; Ukraine; 26.08.1994 (K. Romashchenko)
Localities: 30 – vil. Preobrazhenne, Lugansk reg.; Ukraine; 11.07.2013 (O. Chusova)
Localities: 31 – vil. Preobrazhenne, Lugansk reg.; Ukraine; 14.06.2013 (O. Chusova)
Localities: 32 – vil. Fomovka, Lugansk reg.; Ukraine; 14.06.2013 (Ya. Didukh)
Localities: 33 – vil. Fomovka, Lugansk reg.; Ukraine; 14.06.2013 (Ya. Didukh)
Localities: 34 – vil. Fomovka, Lugansk reg.; Ukraine; 14.06.2013 (Ya. Didukh)
Localities: 35 – 5 km South of the Bilovodsk, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 36 – vil. Preobrazhenne, Lugansk reg.; Ukraine; 14.06.2013 (O. Chusova)
Localities: 37 – right bank of Volchya River, vil. Ohrymivka, Kharkiv reg.; Ukraine; 06.1995 (Ya. Didukh)
Localities: 38 – right bank of Oskol River, vil. Dvorichna, Kharkiv reg.; Ukraine; 06.1995 (Ya. Didukh)
Localities: 39 – vil. Kalynivka, Lugansk reg.; Ukraine; 06.1987 (Ya. Didukh)
Localities: 40 – vil. Preobrazhenne, Lugansk reg.; Ukraine; 14.06.2013 (O. Chusova)
Localities: 41 – 1 km South of vil. Aleksandrovka, Kursk reg.; Russia; 4.09. 2005 (A. Poluyanov)
Localities: 42 – right bank of Oskol River, 1 km North of vil. Kun’e, Kursk reg.; Russia; 11.05. 2005 (A. Poluyanov)
Localities: 43 – 6 km South of vil. Novomelovoe, Kursk reg.; Russia; 1.08. 2005 (A. Poluyanov)
Localities: 44 – 1 km South of vil. Aleksandrovka, Kursk reg.; Russia; 4.09. 2005 (A. Poluyanov)
Localities: 45 – 0.5 km East of vil. Midlle Apochka, Kursk reg.; Russia; 4.09.2005 (A. Poluyanov)
Localities: 46 – 2 km North of vil. Midlle Apochka, Kursk reg.; Russia; 5.09.2005 (A. Poluyanov)
Localities: 47 – 6 km South of vil. Novomelovoe, Kursk reg.; Russia; 1.08.2005 (A. Poluyanov)
Localities: 48 – 1 km North of vil. Nizhnedorozhnoe, Kursk reg.; Russia; 4.08.2005 (A. Poluyanov)
Localities: 49 – 2 km North of vil. Midlle Apochka, Kursk reg.; Russia; 5.09.2005 (A. Poluyanov)
Localities: 50 – left bank of Bystrik River, 1 km East of vil. Beketovo, Kursk reg.; Russia; 27.05.2006 (A. Poluyanov)