(PDF) Старі кладовища як рефугіум степової флори на Півдні України

Moysiyenko І.І., Skobel N.O., Sudnik-Wójcikowska B., Dembicz I., Zachwatowicz M., Zakharova M.Ya

194

Теоретичні та прикладні питання

Old cemeteries as refuge of the steppe flora in Southern

Ukraine 

IVAN IVANOVYCH MOYSIYENKO

NADIIA OLEHIVNA SKOBEL

BARBARA SUDNIK-WÓJCIKOWSKA

IWONA DEMBICZ

MARIA ZACHWATOWICZ

MARYNA YAROSLAVIVNA ZAKHAROVA

VIKTORIA MYKOLAIVNA DZERKAL

MOYSIYENKO І.І., SKOBEL N.O., SUDNIK-WÓJCIKOWSKA B., DEMBICZ I.,

ZACHWATOWICZ M., ZAKHAROVA M.YA., DZERKAL V.M. (2021). Old cemeteries as

refuge of the steppe flora in Southern Ukraine. Chornomors’k. bot. z., 17 (3): 194–217.

doi: 10.32999/ksu1990-553X/2021-17-3-1

Changes in natural landscapes and economic activities lead to the loss of a large proportion

of the steppes. Recent studies (2008–2017) have demonstrated the high importance of old

cemeteries, that contribute to the preservation of steppe red-listed steppe species. The area

of the old cemeteries of the Lower Dnieper varies from 0.43 ha to 6.79 ha. The list of

vascular plants from 10 cemeteries of the Lower Dnieper region includes 388 species

belonging to 226 genera, 62 families, 3 classes and 2 divisions. The total number of species

within cemetery varied from 104 to 217 (on average 153 species). In old cemeteries of the

studied region, rare steppe species have survived, although steppe vegetation is preserved

where there are fragments of virgin and unploughed steppes around the cemeteries, near old

villages or cities. 22 protected vascular plant species were found in the studied old

cemeteries (5.65 % of the total species pool). Five of these are included in the Red Data

Book of Ukraine: Astragalus henningii, Stipa capillata, Stipa lessingiana, Stipa ucrainica,

Tulipa biebersteiniana. 17 species of vascular plants are included in the Red List of the

Kherson region: Amygdalus nana, Bellevalia sarmatica, Centaurea trichocephalla,

Convallaria majalis, Dianthus andrzejowskianus, Elytrigia pseudocaesia,

Ephedra distachya, Fraxinus excelsior, Iris halophila, Limonium platyphyllum, Linaria

macroura, Muscari neglectum, Peucedanum ruthenicum, Prangos odontalgica, Quercus

robur, Veronica capsellicarpa, Vinca herbacea.

Key words: Lower Dnieper, steppe, cultural heritage sites, vascular plants, flora,

sozophytes, in situ

МОЙСІЄНКО І.І., СКОБЕЛЬ Н.О., СУДНІК-ВОЙЦИКОВСЬКА Б., ДЕМБІЧ І., ЗАХВАТОВИЧ М.,

ЗАХАРОВА М.Я., ДЗЕРКАЛЬ В.М. (2021). Старі кладовища як рефугіум степової

флори на Півдні України. Чорноморськ. бот. ж., 17 (3): 194–217.

doi: 10.32999/ksu1990-553X/2021-17-3-1

© Moysiyenko І.І.1, Skobel N.O.1, Sudnik-Wójcikowska B.2, Dembicz I.2, Zachwatowicz M.3, Zakharova M.Ya.1,

Dzerkal V.M.1,4

1Kherson State University, 27 Universytetska Str., Kherson, 73000, Ukraine

2Department of Ecology and Environmental Conservation, Faculty of Biology, University of Warsaw, ul. Żwirki i

Wigury 101, 02-089 Warsaw, Poland

3Faculty of Geography and Regional Studies Chair of Physical Geography, Department of Geoecology, University of

Warsaw, ul. Krakowskie Przedmieście 30, 00-927 Warsaw, Poland

4National Natural Park “Nyzhnodniprovskyi” 136-a Universytetska Str., Kherson, 73000, Ukraine

e-mail: ivan.moysiyenko@gmail.com

Submitted 12 June 2021 Recommended by D. Dubyna Published 30 December 2021

Чорноморський ботанічний журнал – том 15, № 2 (2019)

Чорноморський ботанічний журнал – том 17, № 3 (2021)

Old cemeteries as refuge of the steppe flora in Southern Ukraine

195

Зміни природних ландшафтів та господарська діяльність призводить до втрати

великої частки степів. Недавні дослідження (протягом 2008–2017) продемонстрували

високе значення існування старих цвинтарів, які сприяють збереженню степових

созофітів. Площа старих цвинтарів Нижнього Дніпра коливається від 0.43 га до 6,79

га. У список флори судинних рослин входять 388 видів з 10 цвинтарів. Загальне

багатство видів варіювало від 104 до 217 (у середньому по видів 154 види). Флора

старих цвинтарів включає до 226 родів, 62 родини, 3 класи та 2 відділи. Зазвичай на

цвинтарях Нижнього Дніпра зустрічаються рідкісні види, які протягом тривалого

часу зберігаються на старих цвинтарях та сприяють збереженню виду in situ разом зі

степовим покривом, проте рослинний покрив зберігається не на всіх цвинтарях, а

лише на тих, які були закладені на цілинній та нерозораній ділянці степу, поблизу

старих сел чи міст. На старих цвинтарях знайдено 22 видів судинних рослин 5,65 %,

які підлягають охороні з них 5 видів рослин включені до Червоної книги України:

Astragalus henningii, Stipa capillata, Stipa lessingiana, Stipa ucrainica,

Tulipa biebersteiniana, та 17 видів судинних рослин, які включено до Червоного

Списку Херсонської області: Amygdalus nana, Bellevalia sarmatica, Centaurea

trichocephalla, Convallaria majalis, Dianthus andrzejowskianus, Elytrigia pseudocaesia,

Ephedra distachya, Fraxinus excelsior, Iris halophila, Limonium platyphyllum, Linaria

macroura, Muscari neglectum, Peucedanum ruthenicum, Prangos odontalgica, Quercus

robur, Veronica capsellicarpa, Vinca herbacea.

Ключові слова: Нижнє Дніпро, степ, об’єкти культурної спадщини,судинні рослини,

флора, созофіти, in situ

МОЙСИЕНКО И.И., СКОБЕЛЬ Н.О., СУДНИК-ВОЙЦИКОВСЬКАЯ Б., ДЕМБИЧ И.,

ЗАХВАТОВИЧ М., ЗАХАРОВА М.Я., ДЗЕРКАЛЬ В.Н. (2021). Старые кладбища как

рефугиум степной флоры Юга Украины. Черноморск. бот. ж., 17 (3): 194–217.

doi: 10.32999/ksu1990-553X/2021-17-3-1

Изменения ландшафтов и хозяйственная деятельность приводит к потере большой

части степей. Недавние исследования (в течение 2008–2017) продемонстрировали

высокое значение существования старых кладбищ, которые способствуют

сохранению степных созофитив. Площадь старых кладбищ Нижнего Днепра

колеблется от 0.43 га до 6,79 га. В список флоры сосудистых растений входят 388

видов из 10 кладбищ. Общее богатство видов варьировало от 104 до 217 (в среднем

по 154 вида). Флора старых кладбищ включает 226 родов, 62 семейства, 3 класса и 2

отдела. Обычно на кладбищах Нижнего Днепра встречаются редкие виды, которые в

течение длительного времени сохраняются на старых кладбищах и способствуют

сохранению вида in situ вместе со степным покровом, однако растительный покров

хранится не на всех кладбищах, а только на тех, которые были заложены в целинных

и не распаханных участках степи, вблизи старых сел или городов, На старых

кладбищах найдено 22 вида сосудистых растений 5,65 %, Из которых 5 видов

растений включены в Красную книгу Украины: Astragalus henningii, Stipa capillata,

Stipa lessingiana, Stipa ucrainica, Tulipa biebersteiniana, и 17 видов сосудистых

растений, включенных в Красную книгу Херсонской области: Amygdalus nana,

Bellevalia sarmatica, Centaurea trichocephalla, Convallaria majalis, Dianthus

andrzejowskianus, Elytrigia pseudocaesia, Ephedra distachya, Fraxinus excelsior, Iris

halophila, Limonium platyphyllum, Linaria macroura, Muscari neglectum, Peucedanum

ruthenicum, Prangos odontalgica, Quercus robur, Veronica capsellicarpa, Vinca herbacea.

Ключевые слова: Нижний Днепр, степь, объекты культурного наследия, сосудистые

растения, флора, созофиты, in situ

In recent centuries, anthropogenic activity has led to significant losses of natural

habitats in the world [LÖKI et al., 2019; VICKERY et al., 2009]. Especially significant changes

occurred in steppe zone in the south of Ukraine, where the area of steppe vegetation decreased

forty-fold (steppe in the XXth centuries covered circa 40 % of the total territory of the country,

while today steppe remnants survived only on 1 % of this territory) [BURKOVSKYI et al.,

2013]. Recent studies demonstrated the high biodiversity of cultural heritage sites of

anthropogenic origin such [LÖKI et al., 2015, 2019a, b] as burial mounds or kurgans [DEÁK et

Moysiyenko І.І., Skobel N.O., Sudnik-Wójcikowska B., Dembicz I., Zachwatowicz M., Zakharova M.Ya

196

al., 2016, 2018, 2020, 2021; DEMBICZ et al., 2020; SUDNIK-WÓJCIKOWSKA, MOYSIYENKO,

2006; SUDNIK-WÓJCIKOWSKA et al., 2011; VALKÓ et al., 2018], sacred groves and forests

[BHAGWAT, RUTTE, 2006; BRANDT et al., 2013], old settlements [CELKA, 2011; DAYNEKO,

2019; MOYSIYENKO et al., 2015, 2018, 2019, 2020; MOYSIYENKO, DAYNEKO, 2019].

Another cultural heritage site that plays a significant role in biodiversity conservation

are the old cemeteries [BARRETT, BARRETT, 2001]. Nonetheless, sacred hills, caves and

islands and water bodies connected with religious beliefs like sacred rivers, lagoons and

springs are also common on several continents [VERSCHUUREN et al., 2010; AYSEL et al.,

2009]. Information on the role of cemeteries in biodiversity conservation is summarized in

Löki's article, which lists the groups of biodiversity and the most relevant publications

connected to them [LÖKI et al., 2015, 2019a, b]: «Cemeteries are important centers for the

preservation of lichens [HAWKSWORTH, MCMANUS, 1989], mosses [FUDALI, 2001], rare

mushrooms [BROWN et al., 2006], rare plants [HOLDEN, MCDONALD-MADDEN, 2017;

MOLNÁR et al, 2017; SIGIEL-DOPIERALA, JAGODZINSKI, 2011], trees that are much older than

the trees in the surrounding areas [GAO et al., 2013; PRESTON, 1972], invertebrates [ÖRSTAN,

2004; ÖRSTAN, KÖSEMEN, 2009; TAN, 2012; TAN et al., 2013], birds [ČANÁDY, 2017; KOCIAN

et al., 2003; LUNIAK, 1981; LUSSENHOP, 1977; PEARSON, 1915; VALLEJO et al., 2009;

VILLASEСOR, ESCOBAR, 2019] and bats [TREWHELLA et al., 2005]».

Numerous countries have recognized the natural and cultural value of cemeteries.

Significant differences in the role of cemeteries can vary depending on the location and land

use, biogeographical features and cultural traditions of the country [LÖKI et al., 2015, 2019a,

b]. World practice shows that cemeteries cover large areas [BHAGWAT, 2009; LÖKI et al.,

2015, 2019a, b] and usually do not have active anthropogenic activity and are key elements

for the preservation of natural habitats vegetation [BARRETT, BARRETT, 2001], therefore, the

flora of cemeteries is the subject of research in many countries around the world. Most case

studies (except reviews) are concentrated in Asia or Europe [LÖKI et al., 2015, 2019a, b].

Some botanical studies of cemeteries were conducted in Albania, Australia, Bangladesh,

Czech Republic, Germany, Great Britain, Greece, India, Israel, Japan Latvia, Morocco, New

Zealand, Pakistan, Romania, Russia, Slovakia, Slovenia, Tunisia, Turkey, USA [LÖKI et al.,

2015, 2019a, b].

In Europe, the flora of cemeteries is the best studied in Poland. Polish researchers most

often study old cemeteries in forests or areas of anthropogenic environments [GALERA et al.,

1993; LISOWSKA et al., 1994, NOWIŃSKA et al., 2010, 2019, 2020; SUDNIK-WÓJCIKOWSKA,

GALERA, 2005, TANAŚ, 2008; TRZASKOWSKA et al., 2013].

Studies of the flora of cemeteries in Ukraine are not very common. There are a few

special publications devoted to the spontaneous flora of cemeteries located in cities: Odessa

[GERASYMIUK, 2014; VASYLIEVA-NEMERTSALOVA 1996], Kropyvnytskyi [ARKUSHYNA, 2003

a, b, 2007] and Kherson [MOYSIYENKO, 1997]. The decorative flora of Kyiv cemeteries

[KUSHNYR, 2006, 2004; SUKHANOVA, 2010] and distribution of adventitious tree and shrub

plants from cemeteries in Donetsk [EREMENKO, 2013] was also studied. Cemeteries are an

integral part of cities, thus should be taken into account in studies focusing on urban flora.

Their flora is usually included in general publications of city floristic studies [BESARABCHUK,

VOLHIN, 2017; BURDA, HUMECH, 1988; HAMULIA, ZVIAHYNTSEVA, 2010; HUBAR, 2006;

MALTSEVA, 2019; MELNYK, 2001; MOYSIYENKO, 1997, 1999; VASYLIEVA-NEMERTSALOVA,

1996; ZAVIALOVA, 2010].

Rural cemeteries are generally unexplored. The role of cemeteries in the preservation

of steppe diversity has also not been studied in Ukraine.

In altered landscapes, historic burial sites, such as cemeteries, have the potential to

conserve biodiversity [LÖKI et al., 2019a, b]. Thus, in the steppe zone of Eurasia and in the

contact zones of the forest-steppe zone, mounds, cemeteries and settlements are often the last

safe havens of steppe vegetation in Eastern Europe [CREMENE et al., 2005; MOYSIYENKO et

Old cemeteries as refuge of the steppe flora in Southern Ukraine

197

al., 2014; MOYSIYENKO, SUDNIK-WÓJCIKOWSKA, 2006, 2009; SUDNIK-WÓJCIKOWSKA,

MOYSIYENKO, 2006, 2010, 2011]. Steppe vegetation in old cemeteries is a remnant of virgin

steppe landscapes, as most cemeteries were founded in 17–18th centuries, when the expansion

of natural habitats was continuous [MOYSIYENKO et al., 2017].

The purpose of our study was: 1) to establish the level of floristic richness of ancient

cemeteries on the example of 10 cemeteries of the Lower Dnieper; 2) to find out the

peculiarities of the structure of the flora of ancient cemeteries; 3) show the role of ancient

cemeteries in the preservation of steppe flora.

Study area

The characteristic features of the natural conditions of the Lower Dnieper region are

determined by its geographical location within the true steppe zone of the Eastern European

plain. The climate of the Lower Dnieper as a steppe zone is continental and is characterized

by a temperate-continental climate with mild snowless winters and hot dry summers. The

main features of this climate are formed under the influence of general and local climate-

forming factors. The surface of the territory is almost flat. There are no rivers. The average

duration of the frost-free period is 170–180 days. The total annual precipitation is below 350

mm; total summer rainfall is greater than total winter rainfall; maximum rainfall occurs in

June and July. Mean July temperature is +230C, mean temperature for January – not greater

than –30C; extreme temperatures: +390C in summer, –310C in winter The region is

characterized by low rainfall (350–420 mm per year) and intense solar radiation [MARYNYCH,

SHYSHCHENKO, 2005; MOYSIYENKO et al., 2006; MOYSIYENKO, DAYNEKO, 2019].

The floodplain of the Lower Dnieper is densely cut by numerous straits and branches.

Islands and lakes of various sizes and configurations are common. In the coastal strip of the

Lower Dnieper terrace-delta plain, coastal land, flooded by sea waters, spit and islands,

estuaries and lagoon lakes are situated. There are more than 140 lakes in the Lower Dnieper,

some of which are called estuaries: White Lake, Bezmen, Deaf Estuary, Pigeon Estuary, Mud,

Zburyiv Estuary, Kardashian Estuary, Burkut, Kokhansk, Oleshkiv Lakes and others.

[MARYNYCH, SHYSHCHENKO, 2005; MOYSIYENKO, DAYNEKO, 2019].

Regarding the geobotanical zoning, the Lower Dnipro region is located in three

districts of the Black Sea and Azov steppe sub-province of the Pontic steppe province of the

Steppe zone: Bug-Ingul district of grasses, grass meadows and vegetation of limestone

outcrops, Lower Dnipro district of sand steppes, sands and reed beds, Dnipro-Azov district of

grasses, wormwood-grass steppes and depression (pid – in Ukrainian) meadows

[GEOBOTANICAL ZONING …, 1977; MOYSIYENKO, DAYNEKO, 2019].

Floristic research was carried out in 10 old cemeteries (Fig. 1, Fig. 2). According to

the administrative and territorial division, the examined cemeteries are located in the

Beryslav, Henichesk, Skadovsk and Kherson districts (former Belozerka, Beryslav,

Velykooleksandrivka, Hola Prystan, Nyzhni Sirogozy districts) of Kherson region.

The area of the cemeteries varies from 0.43 ha to 6.79 ha (Table 1).

Material and methods

The study of the flora of 10 old cemeteries was conducted during 2008–2017 using

route-field method and literature data [SCHMIDT, 1980; SCHMIDT, 1984; TOLMACHEV, 1974;

SHELYAG-SOSONKO, DIDUKH, 1975].

We determined the date of the establishment of each cemetery indirectly - on the basis

of the date of foundation of the nearby villages. We searched for the date of establishing the

village in the literature [ISTORIIA…, 1972] and cartographic materials [MAP OF THE CRIMEA…,

1855; MAPS OF SCHUBERT, 1965; MAP OF THE KHERSON DISTRICT…, 1910].

Moysiyenko І.І., Skobel N.O., Sudnik-Wójcikowska B., Dembicz I., Zachwatowicz M., Zakharova M.Ya

198

Fig. 1. Map of the location of old cemeteries (OC). Explanation: OC1 – Dolmativka, OC2 – Ekonomiia

Ivanivka, 3 – Kurgan near khutir Balakshova, 4 – Nyzhni Torhai, 5 – Poniativka, 6 – Posad-Pokrovske, 7

– Stanislav, 8 – Tiahynka, 9 – Tokarivka, 10 – Tryfonivka.

After the research, the cemetery in Nyzhni Torhai was enlarged to 2.1638 ha (cadastral

number 6523883000: 02: 001: 0122), therefore we present floristic data related to the

examined area of 1.1 ha.

The study of each site was conducted at least 3 times during the growing season: spring,

summer and autumn. The data were compiled in Table A (Appendix 1) which contained the

following additional information about each taxon: its occurrence and abundance by

[MOYSIYENKO et al., 2006] (estimated according to a 3-point scale: 1 – sporadic, 2 –

infrequent, 3 – common) functional group (hs – habitat specialist, g – generalist), species life

form, species life span, status in the historical-geographical classification [KORNAŚ, 1981], the

number of old cemeteries the species occurs, and origin in the case of alien species. Alien

species were identified on the based on the publication of [PROTOPOPOVA, 1991]. Floristic

analyses was conducted in which species richness.

The following scale was used classification by [MOYSIYENKO et al., 2006] to assess the

frequency category of the species: I – rare (occurring in 1–2 cemeteries), II – relatively rare

(3–4), III – not rare (5–6), IV – relatively frequent (7–8), V – common (9–10).

To check what is the level of synanthropization of the flora of old settlements, we used

geographical-historical classification by [KORNAŚ, 1981] includes groups of adventive species

isolated by the time of drift (archaeophytes and kenophytes), degree of naturalization

(epecophytes, ephemerophytes, etc.), and groups of aboriginal species isolated on the basis of

resistance to anthropogenic impact (evapophytes, hemiapophytes, etc.), and assigned all

species to the following groups: indigenious species (non-synanthropic species, hemi-

apophytes, eu-apophytes, ekiophytes) and aliens or anthropophytes (archaeophytes, kenophytes

and ergasiophygophytes).

Old cemeteries as refuge of the steppe flora in Southern Ukraine

199

Fig. 2. The general view of some old cemeteries of the Lower Dnipro region: A – Abondanded cemeteries

(Ekonomiia Ivanivka); B – Invasion of Syringa vulgaris (Tokarivka); C – Used and abandended part of

cemeteries (Stanislav), D – Kurgan on cementeries (Tryfonivka), E – Cemetery among field (Kurgan

Khutir Balakshova), F – Cemeteries among village (Stanislav) (A-D by Ivan Moysiyenko; E, F – satellite

image of the system Google Earth).

To assess plant adaptation to certain ecological conditions we used classification of life

forms which includes certain groups as: hemicryptophytes, therophytes, geophytes,

phanerophytes, chamaephytes, hemicryptophytes-chamaephytes [RAUNKIAER, 1934]. Names of

plant species are given in Latin according to [MOSYAKIN, FEDORONCHUK, 1999], except for

species that are absent in the flora of Ukraine (the authors are listed according to the sources

in which they are mentioned). We have used Google Earth Pro [GOOGLE EARTH PRO, 2021]

and QGIS 3.16 Hannover [QGIS 3.16 HANNOVER 2021] to prepare mapping and spatial

analysis of the Lower Dniprо region.

Moysiyenko І.І., Skobel N.O., Sudnik-Wójcikowska B., Dembicz I., Zachwatowicz M., Zakharova M.Ya

200

Table 1

General information about old cemeteries

№

Name of the

cemetery

Location in the Kherson region

Year of

establishme

nt

Area, ha

Coordinates

1

Dolmativka

Kherson Region, Skadovsk District

(ex Velykooleskandrivka),

v. Dolmativka

1850-1855

3,17

46.220247

32.468254

2

Ekonomiia

Ivanivka

Kherson Region, Beryslav District

(ex Velykooleskandrivka)

1855-1865

0,43

47.097128

33.273068

3

Kurgan

near khutir

Balakshova

Kherson Region, Beryslav District

(ex Velykooleskandrivka)

1855-1865

0,7

47.098750

33.253558

4

Nyzhni Torhai

Kherson Region, Henichesk District

(ex Nyzhnsirohozy), v. Nyzhni Torhai

cadastral number 6523883000: 02:

001: 0122

1840

1,1

(2.1638)

46.789683

34.256028

5

Poniativka

Kherson Region, Kherson District, (ex

Bilozerka), v. Poniativka

1780

1,1

46.743071

32.903220

6

Posad-

Pokrovske

Kherson Region, Kherson District, (ex

Bilozerka), v. Posad-Pokrovske

1789

3,6

46.807617

32.271652

7

Stanislav

Kherson Region, Kherson District (ex

Bilozerka), v. Stanislav

1697

6,79

46.572933

32.150254

8

Tiahynka

Kherson Region, Beryslavsky District,

v. Tiahynka

1778

5,86

46.780492

33.062809

9

Tokarivka

Kherson Region, Kherson District (ex

Bilozerka), v. Tokarivka

1780

2,81

46.754950

32.974147

10

Tryfonivka

Kherson Region, Beryslav District (ex

Velykooleskandrivka),

v. Tryfonivka

1863

3,2

47.257042

33.524622

Results

1. Biodiversity of old cemeteries

The list of vascular flora includes 388 species of spontaneously growing plants

found in the 10 examined cemeteries. Total species richness ranged from 104 to 216 species

(average 153 species per 1 cemetery) (Fig. 3). The species belong to 226 genera, 62 families, 3

classes and 2 divisions.

The vast majority of species belong to the Magnoliophyta division (99.75 %).

Division Pinophyta (0.25 %) is represented by one family – Ephedraceae and one species –

Ephedra distachya. The following families were represented by the greatest number of taxa

(Fig. 4): Asteraceae (76), Poaceae (32), Fabaceae (27), Brassicaceae (24), Lamiaceae (20),

Caryophyllaceae (16), Rosaceae (16), Scrophulariaceae (15), Chenopodiaceae (15) and

Boraginaceae (11).

Genera leading by the number of species were: Veronica (9), Artemisia (8),

Astragalus (7), Euphorbia (7), Allium (6), Chenopodium (6), Galium (6), Centaurea (5),

Limonium (5), Medicago (5).

About half of the flora of cemeteries – 179 species (46 %) are plants that occur

sporadically (in 1-2 cemeteries – I frequency class). Most of them are planted on graves or

appear there temporarily (Fig. 5).

Old cemeteries as refuge of the steppe flora in Southern Ukraine

201

Fig. 3. The General number of species and the number of native and alien species in the flora of each of 10

cemeteries (the number of species is indicated at the top of the bar). Explanation: 1 – Dolmativka, 2 –

Ekonomiia Ivanivka, 3 – Kurgan near khutir Balakshova, 4 – Nyzhni Torhai, 5 – Poniativka, 6 – Posad-

Pokrovske, 7 – Stanislav, 8 – Tiahynka, 9 – Tokarivka, 10 – Tryfonivka.

Fig. 4. The most numerous families in the total flora of the 10 cemeteries studied.

The group of common species recorded on 9–10 of old cemeteries (frequency class V)

are 49 species, or 12.6 % of flora: Anisantha tectorum (10), Agropyron pectinatum (9),

Amaranthus retroflexus (9), Arenaria uralensis (10), Artemisia austriaca (10), Bromus

squarrosus (10), Buglossoides arvensis (10), Capsella bursa-pastoris (10), Chenopodium

album (10), Chondrilla juncea (9), Consolida paniculata (10), Convolvulus arvensis (10),

Coronilla varia (10), Descurainia sophia (9), Elytrigia repens (10), Falcaria vulgaris (10),

Festuca valesiaca (10), Galium aparine (10), Hemerocallis fulva (9), Holosteum umbellatum

Moysiyenko І.І., Skobel N.O., Sudnik-Wójcikowska B., Dembicz I., Zachwatowicz M., Zakharova M.Ya

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(10), Koeleria cristata (9), Lactuca serriola (10), Lamium amplexicaule (10), Medicago

falcata (10), Onopordum acanthium (9), Poa angustifolia (10), Poa bulbosa (10), Polygonum

aviculare (10), Potentilla argentea (10), Pterotheca sancta (10), Ranunculus oxyspermus (9),

Rosa canina (9), Salvia nemorosa (10), Senecio vernalis (10), Seseli tortuosum (10), Setaria

viridis (9), Sisymbrium loeselii (9), Stipa capillata (9), Syringa vulgaris (10), Taraxacum

erythrospermum (10), Tragopogon major (10), Trifolium arvense (10), Valerianella carinata

(10), Verbascum phoeniceum (10), Veronica arvensis (10), Veronica triphyllos (10), Veronica

verna (9), Vicia villosa (10), Viola kitaibeliana (10).

In the next 4 groups, there is a linear increase in the number of species with decreasing

frequency of occurrence. Thus, the most numerous class in terms of the frequency class I,

includes half of the species of flora of old cemeteries.

Fig. 5. Subdivision of the total flora of old cemeteries by frequency class (the total number of species in

each category is indicated at the top of the bar). Frequency classes: I – rare (1–2 cemeteries), II –

relatively rare (3–4 cemeteries), III – not rare (5–6 cemeteries), IV – relatively frequent (7–8 cemeteries),

V – common (9–10 cemeteries).

2. Spectrum of life forms

The spectrum of life forms of ancient cemeteries corresponds basically to that of the

flora of the Pontic grass steppe zone [SUDNIK-WÓJCIKOWSKA, MOYSIYENKO, 2006].

The dominant group of species in old cemeteries are therophytes (38.8 %) (Fig. 6).

The most frequent species therophytes in old cemeteries are: Anisantha tectorum, Arenaria

uralensis, Bromus squarrosus, Buglossoides arvensis, Capsella bursa-pastoris, Chenopodium

album, Consolida paniculata, Galium aparine, Holosteum umbellatum, Lactuca serriola,

Lamium amplexicaule, Polygonum aviculare, Pterotheca sancta, Senecio vernalis, Trifolium

arvense, Valerianella carinata, Veronica arvensis, Veronica triphyllos, Vicia villosa, Viola

kitaibeliana.

The second group by number of species are hemicryptophytes (33.4 %). Significant

representation of hemicryptophytes indicates the preservation of the flora of old cemeteries, as

they are the dominant group in natural steppe groups. The most frequent hemicryptophytes

species in old cemeteries were: Coronilla varia, Falcaria vulgaris, Festuca valesiaca,

Medicago falcata, Poa bulbosa, Potentilla argentea, Salvia nemorosa, Seseli tortuosum,

Taraxacum erythrospermum, Tragopogon major, Verbascum phoeniceum.

Geophytes and geophytes-hemicryptophytes occupy 3 positions in the life form

spectrum and include 56 species or 14.4 %. The most common species of this group are:

Ranunculus oxyspermus, Convolvulus arvensis, Elytrigia repens and Poa angustifolia.

Old cemeteries as refuge of the steppe flora in Southern Ukraine

203

Phanerophytes accounted for 9.3 % of the flora of investigated cemeteries. The most

common species of phanerophytes (nanophanerophytes) were shrubs: Rosa canina and

Syringa vulgaris. Phanerophytes are not typical for the steppe zone. Their relatively high

representation in cemeteries is due to the fact that they are often grown and run wild here.

The smallest number of species in the flora of old cemeteries were represented by

chamephytes (including hemicryptophytes-chamephytes), of which there were 16 species or

4.1 %. Among them, only Artemisia austriaca belongs to the V frequency class.

Fig. 6. Spectrum of life forms in the flora of old cemeteries.

3. Spectrum of species groups in the historical-geographical classification of plants

Most species of flora of old cemeteries were natives (254 species – 65 % of the flora).

As it is shown by Fig. 8, more than half of the native species are non-synanthropic plants –

135 species, or 35 % of the total number of species that are the plants not entering the habitats

altered by man, i.e. non-synanthropic, represented on the old cemeteries mostly by steppe

plants (Fig. 7). Non-synanthropes steppe species occurring with the highest frequency are:

Koeleria cristata, Stipa capillata, Festuca valesiaca. Non-synanthropic species, that are

“cemetery specific” is Sedum acre.

Depending on the level of transformation of the habitats into which they enter,

apophytes can be subdivided into two groups: eu-apophytes and hemi-apophytes. The most

frequent apophytes are as follows: Chenopodium album, Consolida paniculata, Convolvulus

arvensis, Elytrigia repens, Galium aparine, Polygonum aviculare, Senecio vernalis,

Tragopogon major, Trifolium arvense.

Significant number of local plants indicates a high level of preservation of vegetation

in old cemeteries. The contribution of native species in old cemeteries varied from 55.5 %

(Posad-Pokrovske) to 76.9 % (Kurgan near khutir Balakshova), depending on the area of the

cemetery that is subjected to intensive treatment.

Some native species are cultivated in cemeteries and sometimes they run wild from

cultivation (ekiophytes). The most common of them are: Convallaria majalis, Euphorbia

cyparissias, Ficaria calthifolia, Fraxinus excelsior, Ligustrum vulgare, Muscari neglectum,

Quercus robur, Sedum sexangulare, Sempervivum ruthenicum, Ulmus laevis, Ulmus minor.

Alien species occur in all old cemeteries (a total of 134 species, 35 % of flora), and

their share ranges from 23.1 % (Kurgan near khutir Balakshova) to 44.5 % (Posad-

Pokrovske). Significant number of species indicate the threat of possible spread of these alien

species in the future.

Moysiyenko І.І., Skobel N.O., Sudnik-Wójcikowska B., Dembicz I., Zachwatowicz M., Zakharova M.Ya

204

Among anthropophytes, archeophytes predominate over kenophytes. There are 48

species of kenophytes, or 12.6 % of the flora of old cemeteries. The most common of

kenophytes belonging to the V class of frequency are1 species: Amaranthus retroflexus. The

majority of kenophytes belong to the frequency class I (20 species) and occur rarely in

individual cemeteries.

There are 54 species of archeophytes (14 % of the total flora). The total number of

archaeophytes in cemeteries was greater than the sum of kenophytes. The most common

archeophytes belonging to the frequency class V, were 13 species: Anisantha tectorum,

Bromus squarrosus, Buglossoides arvensis, Capsella bursa-pastoris, Descurainia sophia,

Lactuca serriola, Lamium amplexicaule, Onopordum acanthium, Setaria viridis,

Sisymbrium loeselii, Veronica arvensis, Veronica triphyllos, Vicia villosa.

Among anthropophytes, there is a significant percentage of ergasiophygophytes, plants

that are grown on graves and appear temporarily in cemeteries. The most common "fugitives

from culture" are: Alcea rosea, Calendula officinalis, Centaurea dealbata, Gaillardia

pulchella, Hemerocallis fulva, Sedum reflexum, Syringa vulgaris,Iris × hybrida, Verbesina

encelioides.

Fig. 7. Historical-geographical classification of the total flora of old cemeteries.

Discussion

Despite the relatively small size of the old cemeteries, they are characterized by a

fairly high level of floristic richness of vascular plants. The flora of old cemeteries represent

7.6 % of the flora of Ukraine, which includes 5 100 species [MOSYAKIN, FEDORONCHUK,

1999] and 19.2 % of the flora of the Northern Black Sea coast, which includes 2 025 species

[MOYSIYENKO, 2013].

The size of cemeteries affects the quantitative characteristics of local flora (number of

species). Cemetery size, spatial isolation and current use can affect the floristicdiversity of

cemeteries. Оn the other hand, the qualitative features of flora probably depend more on other

factors (human activity and location) than on cemetery size. The location of a cemetery

affects the species specificity of the flora. This statement applies in particular to abandoned

cemeteries, which are characterized by lower species similarity and a higher proportion of

native plants in comparison to cemeteries that are still in use [NOWIŃSKA et al., 2020].

Steppe vegetation is not preserved in all cemeteries, but only in those that were laid on

the virgin and unploughed part of the steppe, near old villages or towns. If the cemetery is

Old cemeteries as refuge of the steppe flora in Southern Ukraine

205

already laid on the plowed area (newly created) or transferred to another area, it does not

proper for the existence of steppe plants” or something similar. A clear indicator of the

conservation value of old cemeteries is the share of sozophytes in the flora [MOYSIYENKO et

al., 2017]. In the old cemeteries in the Lower Dnieper region, there are rare steppe species that

have survived (in situ) for a long time. It is possible because the burial places in Ukraine are

held in esteem, and any maintenance activities activities are traditionally limited to the care of

graves. On the other hand, spontaneous vegetation in the vicinity of graves and wastelands is

not destroyed [MOYSIYENKO et al., 2017]. Currently used cemeteries are richer in species and

more diverse than abandoned ones, because human activities (burial and systematic economic

practices introduction of ornamental species) usually contribute to the short-term emergence

of random species [NOWIŃSKA et al., 2020]. The relatively high proportion of kenophytes is

related to their spontaneous spread in favorable (corresponding) habitats.

A number of unfavorable anthropogenic factors in old cemeteries lead to a decrease in

the share of biodiversity. Fires, grazing, littering, and cultural traditions of burial and care of

graves in cemeteries lead to the loss of steppe cover due to the appearance of deliberately

introduced ornamental plants and other adventitious species in cemeteries, which are usually

planted by locals around graves, however the presence of a tombstone or other tombstones

affects the escape of lichens and bryoflora [FUDALI, 2001]. The location of the cemetery near

the settlements affects the species specificity of the flora [NOWIŃSKA et al., 2020].

The presence of typical steppe species, such as Festuca valesiaca, Koeleria cristata,

Stipa capillata and a large proportion of natural non-synanthropic species, compared to

invasive ones, indicates a relatively good state of preservation of steppe vegetation in old

cemeteries in the natural state in situ.

In the natural steppe flora, therophytes are usually less numerous than the

hemicryptophytes. The high number of therophytes in the flora of the old cemeteries of the

Lower Dnieper region is due to less favorable conditions in the south of Ukraine (temperate-

continental climate) and anthropogenic influences. Also, the loss of steppe cover may be due

to the appearance in cemeteries of purposefully introduced ornamental plants, which are

usually planted by the local population, on graves, as well as the spontaneous spread of

anthropophytes. The contribution of native species in old cemeteries varied from 55.5 %

(Posad-Pokrovske) to 76.9 % (Kurgan near khutir Balakshova), which indicates that the flora

of cemeteries is largely transformed by man. These numbers are really high suggesting that

old cemeteries are very important habitats for steppe plants.

The specificity of the flora of cemeteries is a significant percentage of plants that

found themselves in this territory as a result of escaping from cultivation (ergasiophygophytes

and ekiophytes). This is due to the peculiarities of the care of cemeteries in the South of

Ukraine, which consists in the intensive planting of cemeteries with ornamental plants, which

are represented by both non-native and local plants. From the point of view of preservation of

a natural vegetation cover widespread cultivation of plants in cemeteries has two

consequences. Wild non-native plants have a negative effect by competing with local plants.

In particular, large areas of cemeteries are occupied by thickets Syringa vulgaris, Ailanthus

altissima, Lycium barbatum. Native woody plants (trees, such as Fraxinus excelsior, Quercus

robur, and shrubs, as Ligustrum vulgare), which are not characteristic of steppe vegetation,

but also are cultivated in cemeteries and escape from cultivation, may have a negative impact

on steppe vegetation on cemeteries and in vicinity.

Moysiyenko І.І., Skobel N.O., Sudnik-Wójcikowska B., Dembicz I., Zachwatowicz M., Zakharova M.Ya

206

Fig. 8. Rare species of some old cemeteries of the Lower Dnipro – A Stipa capillata; B – Stipa lessingiana;

C – Stipa ucrainica; D – Limonium platyphyllum; E – Muscari neglectum; F– Vinca herbacea; G – Ephedra

distachya; H – Dianthus andrzejowskianus (A, B, C included in the Red Data Book of Ukraine; D, E, F, G,

H – Included in the Red List of the Kherson region, all photos by Moysiyenko І.І.).

On the other hand, local plants spontaneously growing in old cemeteries are also

grown near burials (especially beautiful steppe plants, including Stipa capillata, Ficaria

calthifolia, Iris pumila, Ornithogalum kochii, Vinca herbacea, Viola odorata). Thanks to this,

they have a chance to spread and survive.

It is possible that some beautiful flowering local plants were not specially planted near

the burials, but appeared there spontaneously, and were not destroyed during the clearing of

the burials from wild plants. Such plants include Asparagus officinalis, Potentilla recta,

Salvia nemorosa. Aboriginal plants, which are not typical for steppe, but are often cultivated

Old cemeteries as refuge of the steppe flora in Southern Ukraine

207

on cemeteries. We have classified some of these species as ekiophytes, including such rare

plants as: Convallaria majalis, Fraxinus excelsior, Muscari neglectum, Quercus robur.

Some rare plants, grown in cemeteries do not show a tendency to go wild and are

found only in culture. Among them Betula borysthenica Klokov, Paeonia tenuifolia L., which

are included in the Red Data Book of Ukraine [RED DATA BOOK, 2009] and Anemonoides

sylvestris L., Stachys germanica L. – included in the Red List of Kherson region

[CHERVONYI …, 2013]. Plants that last only in the place of cultivation are not included in the

list of species.

However, 65 % of all species found in old cemeteries are native plants. More than half

of this group, 35 % of the total number of species are non-synanthropic plants, which are

represented in old cemeteries, mainly steppe plants. The good preservation of natural flora is

also indicated by the presence of protected plants in old cemeteries. In total, we found 22

species (5.65 %) of vascular plants that are subject to protection in 10 cemeteries (Fig.8).

Among them are 5 species of plants are included in the Red Data Book of Ukraine [RED

DATA BOOK, 2009]: Astragalus henningii, Stipa capillata, S. lessingiana, S. ucrainica, Tulipa

biebersteiniana and 17 species of vascular plants are included in the Red List of the Kherson

region [CHERVONYI …, 2013]: Amygdalus nana, Bellevalia sarmatica, Centaurea

trichocephala, Convallaria majalis, Dianthus andrzejowskianus, Elytrigia pseudocaesia,

Ephedra distachya, Fraxinus excelsior, Iris halophila, Limonium platyphyllum, Linaria

macroura, Muscari neglectum, Peucedanum ruthenicum, Prangos odontalgica, Quercus

robur, Veronica capsellicarpa, Vinca herbacea.

The preservation of the natural vegetation cover in cemeteries is supported by the

sacred status of cemeteries that are places where economic activities are not allowed. This is

evidenced by the attitude to the steppe areas in cemeteries, which were formed in places with

destroyed tombstones. In the middle of the 20th century, for atheistic reasons, cemeteries

were destroyed to destroy old crosses. Such places exist today in the form of plain meadows

(this is where the most typical steppe vegetation is presented). Repeated burials were not

performed.

The results obtained emphasize the floristic specificity and value of old cemeteries.

They could play an important role in steppe phytodiversity conservation, and give perspective

for the future steppe restoration actions.

Conclusions

The flora of old cemeteries in the south of Ukraine shows several specificity, its

manifestations are:

a) The share of native flora – approx. 2/3 of the flora, is similar to the typical flora of

cities, but among native plants, typical apophytes constitute only 20 %, and species of natural

and semi-natural habitats definitely dominate.

b) The dominant share of therophytes, both native and alien, in the flora of old

cemeteries indicates instability of habitats.

c) The share of species with woody shoots – phanerophytes is small, about 9.2 %, of

which native trees and shrubs account for only 1/3. This reflects the location in the forestless

zone (steppes).

d) Typical for cemeteries is the relatively large share of species that go wild (over

11%), both native and foreign. As might be expected, they are mostly ornamental plants. The

cemetery may prove to be a "relay station" for some prairie species with sumptuous flowers.

The cultivation of native (equally beautiful) steppe species should be promoted.

e) Old Ukrainian cemeteries, which survived the communist times and are now

extensively used, should be subject to special protection as monuments and because they

become an enclave for steppe species returning here. Among them are species that are legally

protected and listed in the Red Books.

Moysiyenko І.І., Skobel N.O., Sudnik-Wójcikowska B., Dembicz I., Zachwatowicz M., Zakharova M.Ya

208

Acknowledgments

This research is supported by the Swedish Science Council (Vetenskapsrådet) project N 2012-06112

and by National Research Foundation of Ukraine «Grass habitats of Ukraine of European importance: current

status, the extent of losses and the conservation strategy in the context of global climate change and

anthropogenic transformation of the environment» (registration № 0120U104763).

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212

Apendix 1.

Cheklist of the flora of old cemeteries

№

Name of species

Cemetery Dolmativka

Cemetery Ekonomiia Ivanivka

Cemetery near khutir

Balakshova

Cemetery Nyzhni Torhai

Cemetery Poniativka

Cemetery Posad- Pokrovske

Cemetery Stanislav

Cemetery Tiahynka

Cemetery Tokarivka

Cemetery Tryfonivka

Species occurrence

Functional group

Status in the Ukrainian flora

Life form

Life span

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

1

Achillea leptophylla M.Bieb.

1

2

HS

Ns

h

P

2

Achillea micranthoides Klokov

2

1

HS

Ns

h

P

3

Achillea nobilis L.

1

2

4

HS

Ns

h

P

4

Achillea pannonica Scheele

1

2

1

7

G

Ns

h

P

5

Achillea setacea Waldst. & Kit.

1

2

3

5

G

Ha

hg

P

6

Aegilops cylindrica Host

2

1

5

G

Ha

t

A

7

Aesculus hippocastanum L.

1

G

Eg

mf

P

8

Agropyron pectinatum (M.Bieb.) P.Beauv.

2

3

1

3

1

2

9

HS

Ha

h

P

9

Ailanthus altissima (Mill.) Swingle

1

2

1

4

G

Ke

mf

P

10

Ajuga chia Schreb.

1

G

Ha

hg

A/B

11

Alcea rosea L.

1

6

G

Eg

h

B/P

12

Allium cepa L.

1

G

Eg

g

P

13

Allium guttatum Steven

2

1

2

4

HS

Ns

g

P

14

Allium inaequale Janka

1

HS

Ns

g

P

15

Allium paczoskianum Tuzs.

1

2

1

5

HS

Ns

g

P

16

Allium paniculatum L.

2

1

2

HS

Ns

g

P

17

Allium rotundum L.

2

1

2

HS

Ns

g

P

18

Alopecurus pratensis L.

1

G

Ns

g

P

19

Alyssum desertorum Stapf.

3

1

2

1

2

7

G

Ha

t

A

20

Alyssum hirsutum M.Bieb.

1

3

G

Ha

t

A

21

Amaranthus albus L.

1

2

1

7

G

Ke

t

A

22

Amaranthus blitoides S.Watson

1

4

G

Ke

t

A

23

Amaranthus retroflexus L.

1

2

1

9

G

Ke

t

A

24

Ambrosia artemisiifolia L.

1

7

G

Ke

t

A

25

Amygdalus nana L.

1

HS

Ns

nf

P

26

Androsace elongata L.

1

2

3

HS

Ns

t

A

27

Anethum graveolens L.

1

G

Eg

t

A

28

Anisantha sterilis (L.) Nevski

1

3

G

Ar

t

A

29

Anisantha tectorum (L.) Nevski

2

3

1

10

G

Ar

t

A

30

Anthemis ruthenica M.Bieb.

3

2

1

2

1

6

G

Ap

t

A

31

Anthriscus cerefolium (L.) Hoffm.

2

1

6

G

Ap

t

A

32

Arabidopsis thaliana (L.) Heynh.

1

2

G

Ke

t

A

33

Arctium lappa L.

1

G

Ap

h

B

34

Arctium tomentosum Mill.

1

G

Ap

h

B

35

Arenaria uralensis Pall. ex Spreng.

3

1

3

2

3

2

1

3

10

G

Ha

t

A

36

Armeniaca vulgaris Lam.

1

5

G

Ke

mf

P

37

Armoracia rusticana P.Gaertn., B.Mey. & Scherb.

1

G

Ke

g

P

38

Artemisia absinthium L.

2

1

2

1

2

6

G

Ar

c

P

39

Artemisia annua L.

1

G

Ke

t

A

40

Artemisia austriaca Jacq.

2

3

2

3

2

3

10

G

Ha

c

P

41

Artemisia santonica L.

2

1

G

Ns

c

P

42

Artemisia vulgaris L.

1

G

Ap

hc

P

43

Artemisia lerchiana Weber

1

3

HS

Ha

h

P

44

Artemisia pontica L.

2

3

2

G

Ns

h

P

45

Asparagus officinalis L.

2

1

7

HS

Ns

g

P

46

Asparagus verticillatus L.

1

G

Ns

g

P

47

Asperugo procumbens L.

1

3

1

3

G

Ap

t

A

48

Asperula cynanchica L.

1

HS

Ns

h

P

49

Astragalus cicer L.

1

HS

Ns

h

P

50

Astragalus henningii (Steven) Klokov

2

1

HS

Ns

h

P

51

Astragalus onobrychis L.

3

1

2

HS

Ns

h

P

52

Astragalus pallescens M.Bieb.

1

HS

Ns

hc

P

53

Astragalus ucrainicus Popov & Klokov

1

2

HS

Ns

h

P

54

Astragalus varius S.G.Gmel.

1

HS

Ns

hc

P

55

Astragalus corniculatus M. Bieb.

1

HS

Ns

h

P

56

Asyneuma canescens (Waldst. & Kit.) Griseb. &

Schenk

1

HS

Ns

h

P

57

Atriplex patula L.

1

G

Ap

t

A

58

Atriplex oblongifolia Waldst. & Kit.

1

3

1

5

G

Ha

t

A

59

Atriplex prostrata Boucher ex DC.

1

G

Ha

t

A

60

Atriplex sagittata Borkh.

2

1

G

Ar

t

A

61

Atriplex tatarica L.

1

2

1

2

7

G

Ke

t

A

62

Ballota nigra L.

1

2

6

G

Ar

hc

P

63

Bassia sedoides Asch.

2

1

5

G

Ha

t

A

64

Bellevalia sarmatica (Pall. ex Georgi) Woronow

1

HS

Ns

g

P

65

Berberis vulgaris L.

1

G

Ns

nf

P

Old cemeteries as refuge of the steppe flora in Southern Ukraine

213

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

66

Berteroa incana (L.) DC.

3

1

2

G

Ap

th

A/B

67

Borago officinalis L.

1

G

Eg

h

A

68

Bromopsis inermis (Leyss.) Holub

2

3

2

3

7

G

Ha

hg

P

69

Bromus hordeaceus L.

2

1

G

Ap

th

A/B

70

Bromus squarrosus L.

2

3

2

3

2

1

10

G

Ar

t

A

71

Bromus japonicus Thunb.

2

1

G

Ha

t

A

72

Buglossoides arvensis (L.) I.M.Johnst.

3

1

2

1

2

3

2

3

2

10

G

Ar

t

P

73

Calamagrostis epigeios (L.) Roth

1

G

Ap

h

P

74

Calendula officinalis L.

1

2

1

7

G

Eg

t

A

75

Camelina microcarpa Andrz. ex DC.

2

1

6

G

Ar

th

A/B

76

Campsis radicans (L.) Seem.

1

2

G

Eg

nf

P

77

Cannabis sativa L.

1

G

Ke

t

A

78

Capsella bursa-pastoris (L.) Medik.

3

1

3

2

3

2

10

G

Ar

th

A/B

79

Cardaria draba (L.) Desv.

3

2

3

2

3

2

1

8

G

Ke

hg

P

80

Carduus nutans L.

2

1

G

Ar

th

A/B

81

Carduus uncinatus M.Bieb.

1

2

1

2

1

8

HS

Ns

th

A/B

82

Carduus acanthoides L.

1

2

G

Ar

th

A/B

83

Carex melanostachya M.Bieb. ex Willd.

3

1

2

4

HS

Ns

g

P

84

Carex praecox Schreb.

3

2

7

G

Ha

hg

P

85

Carex stenophylla Wahlenb.

2

1

3

2

1

3

8

HS

Ns

hg

P

86

Carex supina Willd. ex Wahlenb.

2

1

HS

Ns

hg

P

87

Centaurea adpressa Ledeb.

1

HS

Ns

h

P

88

Centaurea dealbata Willd.

1

6

G

Eg

h

P

89

Centaurea trichocephalla M.Bieb. ex Willd.

2

1

HS

Ns

h

P

90

Centaurea diffusa Lam.

1

3

1

2

1

3

8

G

Ke

th

A/B

91

Centaurea salonitana Vis.

1

2

HS

Ha

h

P

92

Cerastium tomentosum L.

1

G

Eg

c

P

93

Cerastium ucrainicum (Kleopw) Klokov

3

1

3

G

Ns

t

A

94

Cerasus vulgaris Mill.

2

1

2

3

2

1

7

G

Eg

mf

P

95

Chenopodium album L.

2

3

2

3

2

10

G

Ap

t

A

96

Chenopodium glaucum L.

1

G

Ha

t

P

97

Chenopodium opulifolium Schrad. ex W.D.J.Koch

& Ziz

1

2

G

Ar

t

A

98

Chenopodium preissmannii Murr

2

1

2

G

Ke

t

A

99

Chenopodium striatiforme Murr

1

6

G

Ke

t

A

100

Chenopodium strictum Roth

1

G

Ke

t

A

95

Chenopodium glaucum L.

1

G

Ha

t

P

96

Chenopodium strictum Roth

1

G

Ke

t

A

97

Chenopodium × preissmannii Murr

2

1

2

G

Ke

t

A

98

Chenopodium album L.

2

3

2

3

2

10

G

Ap

t

A

97

Chenopodium × preissmannii Murr

2

1

2

G

Ke

t

A

98

Chenopodium album L.

2

3

2

3

2

10

G

Ap

t

A

99

Chenopodium opulifolium Schrad. ex W.D.J.Koch

& Ziz

1

2

G

Ar

t

A

100

Chenopodium striatiforme J.Murr

1

6

G

Ke

t

A

101

Chondrilla juncea L.

2

1

2

1

2

1

3

9

G

Ha

h

P

102

Chondrilla latifolia M.Bieb.

1

5

G

Ha

h

P

103

Chorispora tenella (Pall.) DC.

1

2

1

4

G

Ke

t

A

104

Cichorium intybus L.

2

1

2

6

G

Ar

h

P

105

Cirsium vulgare (Savi) Ten.

1

2

G

Ap

th

A/B

106

Cirsium setosum (Willd.) Besser ex M.Bieb.

2

1

2

5

G

Ap

th

A/B

107

Cleistogenes bulgarica (Bornm.) Keng

2

1

2

HS

Ns

h

P

108

Conium maculatum L.

2

G

Ar

h

B/P

109

Consolida ajacis (L.) Schur

1

2

G

Ke

t

A

110

Consolida orientalis (J.Gay) Schrödinger

1

G

Ke

t

A

111

Consolida paniculata (Host) Schur

1

2

1

2

1

2

1

10

G

Ap

t

A

112

Convallaria majalis L.

1

4

G

Ek

g

P

113

Convolvulus arvensis L.

3

1

2

3

10

G

Ap

hg

P

114

Convolvulus lineatus L.

3

2

HS

Ns

h

P

115

Conyza canadensis (L.) Cronquist

2

1

2

1

2

8

G

Ke

th

A/B

116

Coronilla varia L.

1

2

3

1

2

10

G

Ha

h

P

117

Cosmos bipinnatus Cav.

1

2

G

Eg

t

A

118

Crataegus monogyna Jacq.

1

2

G

Ns

nf

P

119

Crepis ramosissima d'Urv.

2

1

2

1

5

G

Ha

t

A

120

Crepis rhoeadifolia M.Bieb.

1

2

1

5

G

Ap

t

A

121

Cuscuta approximata Bab.

1

3

HS

Ns

t

A

122

Cuscuta campestris Yunck.

1

4

G

Ke

t

A

123

Cynodon dactylon (L.) Pers.

1

2

3

G

Ap

hg

P

124

Cynoglossum officinale L.

1

3

G

Ar

h

P

125

Descurainia sophia (L.) Webb ex Prantl

2

1

2

3

2

1

9

G

Ar

t

A/B

126

Dianthus andrzejowskianus (Zapal.) Kulcz.

1

HS

Ns

h

P

127

Dianthus carbonatus Klokov

1

4

HS

Ns

h

P

128

Echium vulgare L.

1

3

G

Ar

h

B

129

Elaeagnus angustifolia L.

1

3

G

Ke

mf

P

130

Elisanthe viscosa (L.) Rupr.

1

G

Ns

h

P

131

Elytrigia intermedia (Host) Nevski

2

1

G

Ha

hg

P

132

Elytrigia pseudocaesia (Pacz.) Prokud.

2

1

G

Ns

g

P

133

Elytrigia repens (L.) Nevski

3

1

3

10

G

Ap

hg

P

134

Ephedra distachya L.

1

4

HS

Ns

nf

P

135

Eragrostis minor Host

2

1

2

5

G

Ke

t

A

136

Erodium neilreichii Janka

1

G

Ns

th

A/B

Moysiyenko І.І., Skobel N.O., Sudnik-Wójcikowska B., Dembicz I., Zachwatowicz M., Zakharova M.Ya

214

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

137

Erodium ciconium (L.) L'Hér.

1

G

Ha

t

A

138

Erodium cicutarium (L.) L'Hér.

3

1

2

3

1

2

8

G

Ap

th

A/B

139

Erophila verna (L.) DC.

1

2

1

6

G

Ha

th

A/B

140

Erucastrum armoracioides (Czern. ex Turcz.)

Cruchet

2

1

HS

Ha

th

A/B

141

Eryngium campestre L.

2

3

2

7

G

Ha

th

A/B

142

Erysimum diffusum Ehrh.

1

2

G

Ha

th

A/B

143

Erysimum repandum L.

1

G

Ar

t

A

144

Euphorbia cyparissias L.

1

G

Ha

h

P

145

Euphorbia glareosa Pall. ex M.Bieb.

1

HS

Ns

h

P

146

Euphorbia marginata Pursh

1

2

G

Eg

t

A

147

Euphorbia virgata Waldst. & Kit.

2

1

2

8

G

Ha

h

P

148

Euphorbia agraria M.Bieb.

3

1

2

7

G

Ha

h

P

149

Euphorbia leptocaula Boiss.

2

3

1

3

HS

Ns

h

P

150

Euphorbia seguieriana Neck.

3

2

6

HS

Ns

h

P

151

Falcaria vulgaris Bernh.

2

3

2

3

1

2

10

G

Ha

h

B/P

152

Festuca valesiaca Schleich. ex Gaudin

2

1

3

2

3

2

3

10

HS

Ns

h

P

153

Ficaria calthifolia Rchb.

1

4

G

Ek

g

P

154

Filago arvensis L.

1

G

Ha

th

A/B

155

Fraxinus excelsior L.

1

2

G

Ek

mf

P

156

Fraxinus pennsylvanica Marshall

1

4

G

Ke

mf

P

157

Fumaria schleicheri Soy.-Will.

1

2

G

Ar

t

A

158

Fumaria vaillantii Loisel.

1

G

Ar

t

A

159

Gagea bulbifera (Pall.) Salisb.

1

HS

Ns

g

P

160

Gagea pusilla (F.W.Schmid) Sweet

1

2

1

2

4

HS

Ns

g

P

161

Gaillardia pulchella Foug.

2

1

2

1

8

G

Eg

h

A/P

162

Galatella biflora (L.) Nees

3

2

1

3

HS

Ns

h

P

163

Galatella villosa (L.) Rchb.f.

2

1

2

HS

Ns

h

P

164

Galium aparine L.

2

3

1

3

2

3

2

10

G

Ap

t

A

165

Galium humifusum M.Bieb.

2

1

2

6

G

Ha

h

P

166

Galium ruthenicum Willd.

2

3

2

3

HS

Ns

h

P

167

Galium spurium L.

1

2

4

G

Ar

t

A

168

Galium verum L.

3

1

3

G

Ha

h

P

169

Galium volhynicum Pobed.

2

1

2

HS

Ns

h

P

170

Geranium pusillum L.

2

1

2

3

2

6

G

Ar

th

A/B

171

Glaucium corniculatum (L.) Curtis

1

G

Ap

t

A/B

172

Gleditsia triacanthos L.

1

4

G

Ke

mf

P

173

Goniolimon besserianum (Schult. ex Rchb.) Kusn.

1

2

HS

Ns

h

P

174

Grindelia squarrosa (Pursh) Dunal

1

2

G

Ke

h

P

175

Gypsophila paniculata L.

2

1

HS

Ns

h

P

176

Helianthus tuberosus L.

1

G

Ke

g

P

177

Helianthus x laetiflorus Pers.

1

G

Ke

g

P

178

Helianthus annuus L.

1

2

G

Eg

t

A

179

Helichrysum arenarium (L.) Moench

1

2

HS

Ns

h

P

180

Heliotropium europaeum L.

1

2

5

G

Ap

t

A

181

Hemerocallis fulva (L.) L.

1

2

1

2

1

2

1

9

G

Eg

g

P

182

Herniaria besseri Fisch. ex Hornem.

1

2

1

6

HS

Ns

c

A/B/P

183

Holosteum umbellatum L.

3

2

3

2

3

2

10

G

Ha

t

A

184

Hordeum murinum L.

2

1

2

5

G

Ar

t

A

185

Hyoscyamus niger L.

1

2

G

Ar

th

A/B

186

Hypericum elegans Stephan ex Willd.

2

1

2

1

5

HS

Ns

h

P

187

Inula britannica L.

1

2

3

G

Ha

hg

P

188

Inula germanica L.

2

1

HS

Ns

h

P

189

Inula oculus-christi L.

1

HS

Ns

h

P

190

Iris germanica Mix

2

1

7

G

Eg

g

P

191

Iris halophila Pall.

2

1

HS

Ns

g

P

192

Iris pumila L.

3

1

2

1

3

2

7

HS

Ns

g

P

193

Iva xanthiifolia Nutt.

1

2

1

3

G

Ke

t

A

194

Jakobaea borysthenica (DC.) B. Nord. & Greuter.

1

HS

Ns

h

P

195

Jacobaea vulgaris Gaertn.

1

3

G

Ha

h

P

196

Juglans regia L.

1

2

4

G

Ke

mf

P

197

Jurinea multiflora (L.) B.Fedtsch.

2

1

2

HS

Ns

h

P

198

Kochia prostrata (L.) chrad.

1

2

1

3

2

6

HS

Ns

c

P

199

Kochia scoparia (L.) Schrad.

2

1

2

G

Ke

t

A

200

Koeleria cristata (L.) Pers.

2

1

3

2

3

2

9

HS

Ns

h

P

201

Kohlrauschia prolifera (L.) Kunth

1

HS

Ns

th

A

202

Lactuca saligna L.

1

G

Ha

th

A/B

203

Lactuca serriola L.

1

2

3

2

3

10

G

Ar

th

A/B

204

Lactuca tatarica (L.) C.A.Mey.

1

2

1

4

G

Ap

hg

P

205

Lamium amplexicaule L.

3

2

3

2

3

2

10

G

Ar

t

A

206

Lamium purpureum L.

1

G

Ar

th

A/B

207

Lappula patula (Lehm.) Asch. ex Gürke

1

G

Ke

th

A/B

208

Lappula squarrosa (Retz.) Dumort.

1

2

G

Ar

th

A/B

209

Lathyrus tuberosus L.

2

3

G

Ar

hg

P

210

Lavatera thuringiaca L.

1

G

Ns

h

P

211

Leonurus glaucescens Bunge

1

3

G

Ha

h

P

212

Lepidium perfoliatum L.

1

2

1

7

G

Ar

t

A/B

213

Lepidium ruderale L.

1

2

G

Ar

th

A/B

214

Leymus ramosus (Trin.) Tzvelev

3

1

G

Ha

g

P

215

Ligustrum vulgare L.

1

G

Ek

nf

P

216

Limonium hypanicum Klokov

2

1

2

HS

Ns

h

P

217

Limonium platyphyllum Lincz.

1

2

HS

Ns

Старі кладовища як рефугіум степової флори на Півдні України

Abstract and Figures