Ecological Questions 30 (2019) 1: 9–19 http://dx.doi.org/10.12775/EQ.2019.001

Occurrence of cruciata in dry grassland (Festuco-Brometea LQ.RáDF]\FH 6WU]\ĪRZVNLH)RRWKLOOV

Tomasz Wójcik1*, Krystyna Towpasz2

1University of Rzeszow, Department of Agrobiology and Environmental Protection, =HOZHURZLF]D6W5]HV]yZ3RODQG 2-DJLHOORQLDQ8QLYHUVLW\'HSDUWPHQWRI3ODQW(FRORJ\.RSHUQLND.UDNyZ3RODQG *e-mail: [email protected] Received: 11 October 2018 / Accepted: 5 February 2019

Abstract. The study presents the results of phytosociological studies of a community with Gentiana cruciata. The investigations ZHUHFRQGXFWHGLQ.RáDF]\FH:LVáRND5LYHUYDOOH\ 6WU]\ĪRZVNLH)RRWKLOOV LQDQDUHDRIP2. The study consisted in taking SK\WRVRFLRORJLFDOUHOHYpVLHUHOHYpVZHUHWDNHQLQDQGUHOHYpVLQZLWKWKH%UDXQ%ODQTXHWPHWKRG7KHVSHFLHV population size was estimated at 2712 generative shoots. Gentiana cruciata grew in a community from the class Festuco-Brometea characterised by high floristic diversity and high species richness (H´ = 3.24). The entire community comprised 91 species (82 species RIYDVFXODUSODQWVDQGVSHFLHVRIPRVVHV ZKHUHDVVSHFLHV RQDYHUDJH ZHUHUHFRUGHGLQWKHUHOHYpV7KHPRVWQXPHURXV represented the class Festuco-Brometea (27 species) and Molinio-Arrhenatheretea (22 species). The highest abundance and constancy were exhibited by Centaurea scabiosa, Thymus pulegioides, Euphorbia cyparissias, Carex caryophyllea, and Gentiana cruciata from the class Festuco-Brometea as well as Origanum vulgare and Galium verum from the class Trifolio-Geranietea. The high natural value of the locality is indicated by the presence of protected species (Gentiana cruciata, Gentianella ciliata, Orobanche kochii) and rare species in the Carpathians (Thlaspi perfoliatum, Salvia verticillata, Carlina vulgaris, Brachypodium pinnatum, Viola hirta, Potentilla pusilla). After 30 years, the dry grassland with Gentiana cruciata is well preserved although it has been abandoned for 18 years. The appearance of shrubs (Prunus spinosa, Cornus sanguinea) indicates progressive secondary succession, which will lead in time to the development scrubs from the class Rhamno-Prunetea. Therefore, preservation of the area requires active protec- tion measures.

Keywords: threatened species, protected plants, phytosociology, habitat preferences, Western Carpathians.

1. Introduction arrangement. The tetramerous sessile flowers form a ca- pitulum at the stem apex, and 1-3 flowers grow in the axils Gentiana cruciata L. from the family is of the top leaves. The 2-3 cm long dirty-blue corolla has a perennial plant reaching a height of 10-50(60) cm. It usu- D WXEXODUFDPSDQXODWH VKDSH 3LĊNRĞ0LUNRZD 0LUHN ally has an unbranched erect stem with dense foliage and 2006).The plant flowers between July and August. The internodes that are substantially shorter than the leaves. flower is strongly protandrous. Gentiana cruciata is self- The 10 cm long and 2 cm wide stem leaves are elliptic to compatible but produces very few seeds in the absence lanceolate, fused into a sheath at the base, with decussate of pollination by bumblebees, hoverflies, and 3HWDQLGRX HW DO  .R]XKDURYD HW DO   7KH fruit is a 2 cm long wingless capsule. The seeds are light Ύഩcorresponding author. (on average 0.134 mg) and reach a size of 0.67 ± 0.12 10 Tomasz Wójcik, Krystyna Towpasz

 PP .OH\HUHWDO$OoLWHSHHWDO  the association Adonio-Brachypodietum pinnati (Matusz- 7KHIUXLWFRQWDLQVRQDYHUDJHVHHGV .pU\HWDO  kiewicz, 2001). Gentiana cruciata has a Eurasian range covering south- The aim of the investigations was to assess the floristic ern and central , Asia Minor, the , Turke- and phytocoenotic diversity of a community with Genti- VWDQ DQG ZHVWHUQ 6LEHULD +XOWpQ  )ULHV  =DMąF ana cruciata and to determine the conservation status and =DMąF ,Q3RODQGLWJURZVLQWKHPRXQWDLQVDQG transformation trends in this phytocoenosis. Additionally, highlands in the southern part of the country. Scattered lo- the population size was estimated based on the number of calities of this species have been reported from the lower generative shoots. 2GHUDQG9LVWXODDUHD0D]XU\/DNH'LVWULFW3RPHUDQLD DQG:LHONRSROVNDUHJLRQV =DMąF =DMąF  The species is a hemicryptophyte with high light and 2. Study area thermal requirements (Zarzycki et al., 2002). Most fre- quently, it grows on dry grasslands, dry meadows, pas- 7KHLQYHVWLJDWLRQVZHUHFRQGXFWHGLQ.RáDF]\FH:LVáRND tures, sun-exposed slopes, and roadsides. It prefers calci- 5LYHU YDOOH\ )LJ   7KH DUHD LV SDUW RI 6WU]\ĪRZVNLH um carbonate-containing clay or loess soils with neutral Foothills, which belongs to the Outer Western Carpathians or alkaline pH (pH 6.6-8.4). The species can be found on .RQGUDFNL 7KHDUHDRI6WU]\ĪRZVNLH)RRWKLOOVLV rendzinas and carbonate pararendzinas and less frequent- located on flysch formations, which are characterised by O\ RQ EURZQ VRLOV DQG FKHUQR]HPV %DEF]\ĔVND6HQGHN low content of calcium carbonate. This is reflected in the $QGU]HMF]XN  3LĊNRĞ0LUNRZD  0LUHN  local vegetation cover exhibiting a negligible proportion of 7UąEDHWDO Gentiana cruciata is a species charac- GU\KDELWDWFRPPXQLWLHV,QWKLVUHVSHFWWKH:LVáRND5LY- teristic for the class Festuco-Brometea and, regionally, for HUYDOOH\LVDQRXWVWDQGLQJDUHDZKHUHWKH.URVQROD\HUV

Figure 1. Localities of Gentiana cruciataLQWKH6WU]\ĪRZVNLH)RRWKLOOV Occurrence of Gentiana cruciata in dry grassland (Festuco-Brometea) in Kołaczyce (Strzyżowskie Foothills) 11

contain inerlayers of laminated white limestone referred IURPWKHPHDQQXPEHURIVSHFLHVSHUUHOHYpDQGWKHWRWDO WRDV-DVáRVKDOHV7KH[HURSKLORXVDQGFDOFLSKLORXVYHJ- number of species, whereas the total diversity was assessed etation developed on the shales is characterised by insular based on the Shannon-Wiener index (Pielou, 1974). Addi- occurrence in this part of the Carpathians (Towpasz, 1990; tionally, the size of the examined population was estimated :yMFLN 2FK\UD 7KHVWXG\ZDVFRQGXFWHGLQDQ based on the number of generative shoots. area located approximately 1 km north of the town centre. The observations were carried out on a plateau and slopes with varied exposure at an altitude of ca. 300 m a.s.l. The 4. Results dry grassland with an area of ca. 2000 m2 was located near various types of plant communities characterised by mosa- The Gentiana cruciata ORFDOLW\ LQ .RáDF]\FH KDV EHHQ ic-like spatial distribution. There were thermophilic scrubs known for 150 years (Towpasz, 1987). In 1982, it has from the class Rhamno-Prunetea, Larix decidua and Pinus been verified again by Towpasz (1987), who estimated the sylvestris plantings, balks with shrub vegetation, fresh and population size at several tens of specimens. This well- dry meadows from the association Arrhenatheretum ela- preserved locality has survived. 30 years ago Gentiana tioris, pastures from the association Lolio-Cynosuretum, cruciata grew in the association Arrhenatheretum elatio- wastelands, and arable fields. ris brizetosum mediae, in which the species characteristic RIIUHVKPHDGRZVSUHGRPLQDWHG UHOHYpVDQG7DEOH  The dry grassland with Gentiana cruciata developed on 3. Methods the plateau and the gently inclined (1-15°) slopes with var- ied (S, SW, N, NW) but mainly southern exposure. The The studies were conducted with the Braun-Blanquet community occupied an area of approximately 2000 m2 PHWKRG   7ZR SK\WRVRFLRORJLFDO UHOHYpV ZLWK DQ and exhibited distinct stratification. The highest layer was area of 50 m2DQGUHOHYpVZLWKDQDUHDRIP2 were composed of shrubs with a cover up to 25%. The herba- taken in 1982 and 2013, respectively. The collected mate- ceous plant cover was in the range of 95-100%. The lowest rial was compiled in a phytosociological table (Table 1) by layer comprised mosses, which accounted for 5-50% of calculation of the constancy and cover coefficient. The af- the cover. The community was characterised by high spe- filiation of the species to syntaxonomic units was adopted cies richness and great floristic diversity, as indicated by from Matuszkiewicz (2001). The nomenclature of vascular the high Shannon-Wiener diversity index H´ = 3.24. The plants and mosses followed those proposed by Mirek et al. SK\WRVRFLRORJLFDOUHOHYpVGHPRQVWUDWHGIURPWRVSH- (2002) and Ochyra et al. (2003), respectively. Protected cies (on average 45). In the entire community, there were species were enlisted based on the Regulation of the Min- 91 taxa, including 82 species and 9 moss LVWHURIWKH(QYLURQPHQW 5R]SRU]ąG]HQLH 7KHVSH- species (Table 1). cies richness of the analysed community was calculated

[12] Cover coeffcient Cover

275 47,7 61,4 54,5 90,9 34,1 47,7 1175

131,8 352,3 786,4 506,8 102,3 1140,9 1093,2 2454,5 Constancy 9 9 9 9 9 9 9 9 9 9 9 9 9 9 ,9 ,9 + + 25 2.3 1.1 100 100 ++++++ ++ . . r+ 50 10 10 10 10 5 3.2 3.2 2.2 1.2 1.2 1.2 3.4 +.2 1.2 +.2 +.2 1.2 307 305 303 303 299 300 SS-NWNW-S-SS 47 43 46 52 56 53 42 47 45 48 + 1.2 1.2 1.2 2.2 1.2 1.2+++++++++++ 1.2 1.2 1.2 1.2 + 1.2 1.2 2.3 1.3 2.3 2.3++ . . +++++++ 2.3 2.3 2.3 +.2 --2-105-21-15-1515 LQ.RáDF]\FH 15.06.2013 16.06.2013 + 1.1 + + 1.2 + 1.2 1.2 1.2 2.2 1.2 1.2++++++++++++++++ 2.2 + + 1.2 1.2++ . . +++1.21.2++++1.2++ + + + 1.2 3.2 3.2 3.2 3.2 2.2 3.2 2.2 2.2 2.2 2.2 2.2 2.2 2.2 2.2 3.2 2.2 Festuco-Brometea 55215----15-5251531020105

from the class 1982

8.07.

1982 15.06. . + ++++ +++++ . . ++++ . + ++ . . ++++1.1++++. . . +. .++++.+ 1 2 345678910111213141516171819202122 1045 2151052-25 112- 5 4010101050153010305 5 105 + + 2.2 ++ + + +1.2+1.21.21.2++++++++ + + ++++ + 1.2+ + . +++++++++++++++++++ 1.2 + 2.2 1.2 3.2 2.2 3.2 1.2 50 50 25 25 25 2595 25 95 25 100 25 100 25 95 25 100 100 25 100 25 95 25 98 25 95 25 100 98 25 100 25 100 25 95 25 95 25 100 100 100 1.1 1.11.2 +++1.2+++++ 3.2 + 3.21.1+ 3.2 2.3 2.2 2.3 2.31.1 2.3 2.3 1.1 +.3 + 1.3 1.1 1.3 1.3 + 1.3 + 2.3 2.3 + 2.2 . .+++++.+..+r+ + 1.3 2.3 1.1 2.3 +1.1+ 1.3 2.3 + 1.3 1.1 1.1 1.1 1.2 + + 1.1 1.1 + + +.2 1.1 2.2 1.2 1.2 + 300 300 300 301 302 301 300 299 301 302 300 299 302 300 300 306 d . . 2.3 +.2 1.3 +.2 +.3 +.2 1.3 1.3 2.3 Gentiana cruciata d . . +.2 1.3 3.3 1.2 2.2 1.3 3.4 1.2 2.3 1.2 2.3 +.2 +.2 3.4 Festuco-Brometea 1XPEHURIUHOHYp month, year) Date (Day, $UHDRIUHOHYp>P@ $OWLWXGH>PDVO@ Exposure,QFOLQDWLRQ >ƒ@ &RYHURIVKUXEOD\HUE>@ &RYHURI+HUEOD\HUF>@ &RYHURIPRVVOD\HUG>@ Number of species SWChCl. SW scabiosa SCentaurea Thymus pulegioides S 29Euphorbia cyparissias S 45 caryophyllea Carex 38 -Gentiana cruciata 37 N 43Melampyrum arvense N 46Plantago media - 47Ranunculus polyanthemos 44 -Campanula glomerata 43 SPolygala comosa 41 48Abietinella abietina - 47 Homalothecium lutescens Sanguisorba minor Allium oleraceum Carlina vulgaris Hypericum perforatum Table 1. Community with Table

[13]

coeffcient Cover Cover

47,7 36,4 13,6 31,8 13,6 29,5 13,6 88,6 27,3

888,6 179,5 484,1 384,1 161,4 131,8 1222,7 1602,3 1047,7 Constancy 9 9 9 9 9 9 9 9 9 9 9 ,9 ,9 . III 22,7 . I 27,3 . + + III 20,4 +++ ....II . 3.2 2.2 2.3 3.2 1.2 1.2 1.2 + + +.....II + . + . . . + III 25 ...... +II +.++++.+ 2.2 2.2 1.2 2.2 1.2 2.2 2.2 2.2 . + 1.2 + + 1.2+ . 1.1 . +++++++ 1.2 + . 2.2 ...r.rr rrrrrrrr+. ++++r . . ++ .+.+++.++.+++ .+...... ++.II 2.2 2.2 2.2 2.2 1.2 1.2 2.1 1.2 1.1 1.1 2.2 2.1 2.2 15.06.2013 16.06.2013 +++++ . +++++++++ . . . + . ++++ . . . +++ .III ..+++..r...... II11,4

++++++1.11.1++++++++++ + . + . +++++ . .

1982

8.07.

1982 15.06. . .+ . . ++++ . + . . .+.1.2+..+. ...r.. . 1.1 2.2 2.2. + 1.2 . . .. +++2.21.21.2+++++1.2++ 2.2 . +++ . 2.2 . . +++++ . 2.2.+.. 3.2 3.2...... 2.2. +...+..+++.. 3.2 1.2 1.2 . 1.1+1.21.21.21.22.22.22.21.21.2+++++ .. . + . + + 2.2 + + + 1.2 1.2 + 1.2 + 1.2 1.2 + 1.2 1.2 1.2 + 2.2 2.2 + 2.2 3.3 3.2 1 2 345678910111213141516171819202122 + ...... + . .+...r... + 1.1 1.1 + + . .+. ++. . + . . .+.+. . + + + . ++++++++++++++++++ 1.1 1.1 2.2 + 1.1 1.1 1.2 2.2 1.2 3.32.22.12.22.21.22.21.21.2++1.1++++1.2+++++ 1.1.1.21.22.21.21.2+++++ 1.1 1.1 + + + + + d . . ..+.2.....1.2.+.2...... Trifolio-Geranietea Molinio- 1XPEHURIUHOHYp month, year) Date (Day, Potentilla pusilla Gentianella ciliata Ononis arvensis kochii Orobanche Thlaspi perfoliatum Rhodobryum ontariense ChCl. Origanum vulgare Galium verum Agrimonia eupatoria Medicago falcata Fragaria viridis medium Trifolium varia Coronilla hirta Viola Silene vulgaris Clinopodium vulgare ChCl. Arrhenatheretea Briza media elatius Arrhenatherum Lotus corniculatus Knautia arvensis cracca Vicia Festuca rubra Achillea millefolium

[14]

coeffcient Cover Cover

225

43,2 68,2 43,2 18,2 40,9 711,4 252,3 131,8 122,7 Constancy I 9,1 9 9 9 9 9 9 III 27,3 ,9 ,9 ,9 + + II 18,2 1.1 + .... ++++ .....I6,8 +.+.+ +....II ...... I9,1 ...+...I6,8 .. +++++ +.3+.3.1.3+.3...... + 1.2...... I6,8 + + . + . . II 36,4 ++++++++++ . . 1.1 ...... +.I6,8 + 1.1 1.1 + . + 1.1 . III 186,4 15.06.2013 16.06.2013 .+...... +++....II11,4 ...+...... I6,8 ++++++++++++ . ++ + . . . + + 1.2 + 2.2 + + 3.2 2.2 2.2 + ...... +...... I2,3

+.2 +.2 +.2 +.2 +.2 1.2 +.2 +.2 2.2 1.2 +.2 +.2 1.2 1.2 +.2 +.2 1.2 +.2 +.2

1982

8.07.

1982 15.06. . . +.3 +.3 1.3 +.3 +.3 . 1.3 1.3 1.3 +.3 . ..+++.+ .. 2.2 + .++++.+..+...... ++...... +...... + . . . . . +.2 . .+.+.. . .+ ++...... +...... + . . . .+. .+. . . +....+ ...... ++.. . + +1.3++++++++++++++ 1 2 345678910111213141516171819202122 + + ++ . ++++ . ++ . +++++++++ + + ++++++++ . ++++++ + . . . .+.+. .++.+..+ ++.+.....+... 1.1++1.21.21.2+....+ d d . . +.2 b . . ..++.... b . . + . + + + 2.2 . . b . .+.1.11.1++....1.1.1.11.1++1.12.1 b ++...... +++.+ Rhamno-Prunetea 1XPEHURIUHOHYp month, year) Date (Day, jacea Centaurea Linum catharticum Pimpinella saxifraga Leucanthemum vulgare Betonica officinalis Daucus carota Leontodon hispidus tomentosa Carex pratensis Tragopogon Galium mollugo Dactylis glomerata Plantago lanceolata pratense Trifolium ChCl. Prunus spinosa Prunus spinosa Cornus sanguinea Cornus sanguinea Rosa canina Rosa canina Crataegus ×macrocarpa Others species Plagiomnium cuspidatum pulchellum Eurhynchiastrum Cuscuta epithymum [15]

– coeffcient Cover Cover 6,8 22,3 15,9 31,8 20,4 22,7

Hieracium Cichorium Constancy Crepis biennis Crepis Other species Other 1(+, 2,3), 17(+, 2,3), 2(+, 2,3), sp. 1(+, 2,3); . +.2 . . I 6,8 Rubus + + 2.2 1.2 III 140,9 +.++II11,4 Solidago gigantea Carex transsilvanica Carex Ranunculus acris ...... II b 16(+, 2,3), 16(+, 2,3), 2(+, 2,3), 6(+, 4,5), 15(+, ..++.....II ++.++.+.+III Viburnum opulus Viburnum Festuca pratensis Hieracium umbellatum Senecio jacobaea 2(+, 2,3), b 13(+, 2,3), b 18(+, 2,3), Ulmus minor Poa pratensis . . +.2 . 1.2 . +.2 1.2 +.2 ...... +++..I9,1 +.2 +.2 +.2 +.2 III 129,5 +++++ . . . ++ . . . +III – 15.06.2013 16.06.2013 Frangula alnus 21(+.2, 4,5), 22(+.2, ....++...... 1.2+.+++ 8(+, 2,3), b 21(+, 4,5), 22(+, Molinio-Arrhenatheretea . +.2 . +.2 +.2 +.2. . . . ++++++ . . . + . . +.2.....+.2+.2+.2...... I . . +.2 +.2 +.2 . +.2 . . III ChCl. Brachypodium pinnatum

– 1982 8.07.

Rhamnus catharticus 1982 6(+.2, 2,3); –

Chaerophyllum aromaticum Chaerophyllum

2(+, 2,3). 15.06. . . +.2 ...... +...... ++.....I6,8 1 2 345678910111213141516171819202122 Festuco-Brometea d... d . . . . 1.2 1.2 . 1.2 d . . ..+.2...... +.2.. Salvia verticillata b 17(1,1, 22,7), Euphorbia esula Rhamno-Prunetea b...... + d ChCl. 7(+, 2,3), 2(+, 2,3), oiaovrara. . ..+++...... virgaurea 1XPEHURIUHOHYp month, year) Date (Day, Thuidium delicatulum Campyliadelphus chrysophyllum Solidago Medicago lupulinaEuphorbia platyphyllosMelilotus officinalis + .Pyrus pyraster + .Brachytecium albicans . . . Picris hieracioides . 1.2 ...... ae otn ...... ++.... montana Calamagrostis epigejos . +Carex . . . Plagiomnium affine 2(+, 2,3); intybus Fraxinus excelsior bauhinii Sporadic species: ChCl. 16 Tomasz Wójcik, Krystyna Towpasz

The class Festuco-Brometea was represented by 5. Discussion and conclusion VSHFLHVZLWKKDYLQJFRQVWDQF\GHJUHH9*UHDWLP- portance in the community structure and high abundance were achieved by Centaurea scabiosa, Thymus pule- Dry grasslands, whose occurrence is determined by the gioides, Euphorbia cyparissias, Carex caryophyllea, Gen- presence of calcium carbonate in the substrate, are ex- tiana cruciata, and Melampyrum arvense, which were the tremely rare communities in the Polish Carpathians. To co-dominant species. From the class Trifolio-Geranietea, date, they have been described from e.g. the Pieniny there were 10 characteristic species, with Origanum vu- 0RXQWDLQV .DĨPLHUF]DNRZD   %XNRZVNLH )RRW- gare and Galium verum reaching high coverage. The class KLOOV 7UąEDHWDO DQG3U]HP\VNLH)RRWKLOOV 6]F- Molinio-Arrhenathereta was represented by 24 species. ]HEOHZVND -DQHFNL.XFKDU]\N.XFKDU]\N Eleven of these species were characterised by the highest 6]DU\7UąEDHWDO 7KHDUHDVRI.DUSDFNLH constancy degree, but a majority had low abundance and Foothills is characterised by calcium carbonate-poor flysch grew in small clusters. The exceptions were Briza media, formations, hence the absence of well-developed dry grass- Arrhenatherum elatius, Lotus corniculatus, Festuca rubra, lands Festuco-Brometea in this region. Investigations car- and Betonica officinalisZKLFKLQVRPHUHOHYpVFRQVWLWXWHG ried out in recent years have demonstrated the presence of an important complement and achieved significant cover- small grassland stands on slopes and plateaus of river val- age in the phytocoenosis. The class Rhamno-Prunetea was leys developed on inerlayers of laminated white limestone represented by 8 species, but a majority of these species FDOOHG -DVáR VKDOHV ;HURSKLORXV YHJHWDWLRQ VWDQGV KDYH occurred sporadically, with the exception of Prunus spi- EHHQUHSRUWHGIURP6WU]\ĪRZVNLH)RRWKLOOVLH:LHORSROND nosa and Cornus sanguinea. There were also 22 species 5LYHUYDOOH\ :yMFLN 3LąWHN DQG:LVáRND5LYHU with undetermined syntaxonomic affiliation. YDOOH\ :yMFLN 2FK\UD:yMFLN DQGIURP The community with Gentiana cruciata exhibited high '\QRZVNLH)RRWKLOOV =LDMD :yMFLN:yMFLN =L- species diversity. There were no expansive or clearly domi- aja, 2015). They are difficult to classify into lower syn- nant species. Noteworthy, there was a rich moss layer com- taxonomic units due to the large proportion of xerothermic posed of 9 moss species, i.e. Rhodobryum ontariense, Abi- species from the class Festuco-Brometea, fringe species etinella abietina, Homalothecium lutescens, Plagiomnium from the class Trifolio-Geranietea, and meadow species cuspidatum, Eurhynchiastrum pulchellum, Thuidium deli- from the class Molinio-Arrhenatheretea. This indicates an catulum, Campyliadelphus chrysophyllum, Brachytecium intermediate nature of these communities. albicans, and Plagiomnium affine. The high natural value 7KH SK\WRFRHQRVLV REVHUYHG LQ .RáDF]\FH KDV EHHQ of the locality was emphasised by the presence of protected described as a community from the class Festuco-Bro- species (Gentiana cruciata, Gentianella ciliata, Orobanche metea. However, it should be emphasized that 30 years kochii) and rare species in the Carpathians (Thlaspi perfo- ago it was similar to the association Arrhenatheretum liatum, Salvia verticillata, Carlina vulgaris, Brachypodium elatioris brizetosum mediae. Its xerothermic character pinnatum, Viola hirta, Potentilla pusilla). Noteworthy was is highlighted by the well-developed moss layer and the also the occurrence of a rare moss species Rhodobryum on- relatively small proportion of clumping grasses. The ad- tariense, which has been reported from only several locali- jacent grassland stands with the rare moss species Rho- WLHVLQ3RODQG :yMFLN 2FK\UD(OOLVHWDO  dobryum ontariense :yMFLN  2FK\UD  (OOLV HW The pH measurements from 1982 showed a value of al., 2016), which comprised abundant Gentiana cruciata 7.5, which was also confirmed in 2013. In the first study SODQWVZHUHFODVVLILHGVLPLODUO\%HVLGHV.RáDF]\FHGen- period, regular grazing was carried out in the locality with tiana cruciataLVSUHVHQWLQWZRORFDOLWLHVLQ6WU]\ĪRZVNLH Gentiana cruciata. In 2000, the land use was discontinued Foothills (Fig. 1). In Nawsie, it grows on a slope descend- (information provided by the owner of the land), which ing to the Wielopolka River valley in the association Arh- resulted in a progressive process of succession. It is evi- henatheretum elatioris, which is characterised by a high denced by the presence of scrub species from the class drying degree and a large proportion of thermophilic spe- Rhamno-Prunetea, in particular Prunus spinosa and Cor- FLHV :yMFLN 3LąWHN ,QWXUQLQ%XNRZDWKHDQD- nus sanguinea. The Gentiana cruciata population is highly O\VHGVSHFLHVRFFXUVRQVWHHSVORSHVLQWKH:LVáRND5LY- abundant and well preserved – it counts 2712 generative er valley in a community with Brachypodium pinnatum shoots. However, its further survival will require active from the class Festuco-Brometea :yMFLN   ,Q WKH protection measures (mowing, grazing) and introduction neighbouring mesoregion, i.e. Dynowskie Foothills, Gen- of monitoring. tiana cruciata was noted sporadically in a fringe commu- nity from the class Trifolio-Geranietea =LDMD :yMFLN 2014). In Bukowskie Foothills, it was present in commu- nity stands with Bromus erectus and in a community with Brachypodium pinnatum 7UąEDHWDO 7KHVSHFLHV Occurrence of Gentiana cruciata in dry grassland (Festuco-Brometea) in Kołaczyce (Strzyżowskie Foothills) 17

is more frequent in Przemyskie Foothills, where it was tum and in a community with Bromus erectus. The former reported from a community with Brachypodium pinna- community comprised 197 species in total and there were tum 6]F]HEOHZVND  -DQHFNL  .XFKDU]\N   VSHFLHV RQ DYHUDJH   LQ UHOHYpV 7KHVH YDOXHV 7UąEDHWDO DVVRFLDWLRQArrhenatheretum elatioris in the latter community were 103 and 25-43 (on average brizetosum mediae %DUDEDV].UDVQ\ DQGDVVRFLD- 33), respectively. Plant communities comprising the ana- tion Arrhenatheretum elatioris centauretosum scabiosae lysed species are characterised by a high Shannon-Wiener .XFKDU]\N 6]DU\ %HVLGHVWKH)RRWKLOOVGen- GLYHUVLW\LQGH[DVFRQILUPHGE\WKHUHVXOWIRU.RáDF]\FH tiana cruciata has been reported mainly from the Pien- (H´ = 3.24). Similar values of this index were record- iny Mountains, where it grows in associations Origano- ed in the community with Brachypodium pinnatum Brachypodietum .DĨPLHUF]DNRZD DQGAnthyllidi- (H´ = 3.22) and those noted in the community with Bromus Trifolietum montani .DĨPLHUF]DNRZDHWDO 7KH erectusZHUHVOLJKWO\ORZHU +   7UąEDHWDO  species grows in similar habitats on the other side of the Investigations conducted by many authors indicate dif- Carpathians. In , it is present in associations Sca- ferent sizes of Gentiana cruciata populations. In Switzer- bioso ochroleucae-Brachypodietum pinnati, Polygalo ma- land and , they range from 1 to 337 individuals, but joris-Brachypodietum pinnati, and Carici albae-Brometum most populations are small and do not exceed 12 individu- monocladi from the class Festuco-Brometea (Hegedüšová DOV .pU\HWDO ,Q3RPRU]H=DFKRGQLHWKHSRSXOD-  âNRGRYi   ,Q WXUQ LQ WKH &]HFK 5HSXEOLF LW LV tion size ranges from 30 to 111 individuals (Piotrowska, a diagnostic species for the association Anthoxantho odo- 2010), whereas the abundance of the species some locali- rati-Agrostietum tenuis from the alliance Cynosurion cris- WLHVLQ:\Ī\QDĝOąVND8SODQGLVHVWLPDWHGDWVHYHUDOWKRX- tati (Chytrý, 2010). VDQGJHQHUDWLYHVKRRWV %DEF]\ĔVND6HQGHN $QGU]HMF- Besides the Carpathian localities, phytocoenoses with zuk, 1997). With its size of 2712 generative shoots, the the presence of Gentiana cruciata have been reported from ORFDOLW\LQ.RáDF]\FHUHSUHVHQWVDODUJHSRSXODWLRQLQFRP- WKHXSODQGSDUWRIWKHFRXQWU\,Q:\Ī\QDĝOąVND8SODQG SDULVRQWRWKHVWXG\UHSRUWHGE\.pU\HWDO   the species grows in floristically rich dry grasslands from After 30 years of research, the dry grassland with Gen- the association Cirsio-Brachypodion %DEF]\ĔVND6HQGHN tiana cruciata is well preserved; however, it has been aban- $QGU]HMF]XN FRPPXQLW\ZLWKBromus erectus, doned for 18 years. The progressive secondary succession community Centaurea scabiosa-Agrimonia eupatoria, and is evidenced by the appearance of shrubs (Prunus spino- association Adonio-Brachypodietum pinnati arrhenathere- sa, Cornus sanguinea) accounting for 25% of coverage tosum %DEF]\ĔVND6HQGHN 7KHFRPPXQLW\Cen- LQVRPHUHOHYpV7KLVZLOOOHDGLQWLPHWRWKHGHYHORSPHQW taurea scabiosa-Agrimonia eupatoria is most similar to the scrubs from the class Rhamno-Prunetea, as indicated by SK\WRFRHQRVLV SUHVHQW LQ .RáDF]\FH LQ WHUPV RI WKH IOR- RWKHUDXWKRUV $ODUGHWDO%DUDEDV].UDVQ\ ULVWLFFRPSRVLWLRQ,Q:\Ī\QD.UDNRZVNR&]ĊVWRFKRZVND ']ZRQNR /RVWHU.RPSDáD%ąED %ąED  Upland, Gentiana cruciata was found in dry grasslands Intensive growth of shrubs is observed in close proximity from the class Festuco-Brometea %DEF]\ĔVND6HQGHNHW to the analysed grassland. An additional threat is the affor- al., 2014). Localities from lowland regions have been re- estation of grass areas, e.g. in a surrounding area, where ported as well. In Pomorze Zachodnie, Gentiana cruciata Pinus sylvestris and Larix decidua had been planted on the was reported from the association Adonio-Brachypodietum dry grassland. Further preservation of this valuable and pinnati and from degraded grassland stands associated rare phytocoenosis in the Carpathians requires active pro- with meadows dominated by dry Arrhenatherum elatius tection measures consisting in grazing or extensive mow- and Calamagrostis epigejos (Piotrowska, 2010). In turn, LQJDVZHOODVUHPRYDORIWUHHVDQGVKUXEV %ąED  LQ3RMH]LHU]H&KHáPLĔVNLHWKHVSHFLHVZDVIRXQGLQ GU\ grasslands from the class Festuco-Brometea, association Arrhenatheretum elatioris, and initial community from the Acknowledgments classes Robinietea and Artemisietea vulgaris .DPLĔVNLHW al., 2018). The authors are grateful to prof. dr hab. Ryszard Ochyra Gentiana cruciata usually grows in floristically rich IRULGHQWLILFDWLRQRIPRVVHVSURIGUKDE(OĪELHWD.XWDIRU communities. In the analysed locality, there were in total identification of Viola hirta, and dr hab. Renata Piwowarc- VSHFLHVDQGWKHSK\WRVRFLRORJLFDOUHOHYpVVKRZHG zyk for identification of Orobanche kochii. 56 species (on average 45). The results of this study are in good agreement with literature reports. 139 species were reported in the association Adonio-Brachypodietum pin- References nati DQG RQDYHUDJH ZHUHQRWHGLQUHOHYpV 3L- RWURZVND 6LPLODUUHVXOWVZHUHSUHVHQWHGE\7UąED $ODUG'&KDEUHULH2'XWRLW'5RFKH3 /DQJORLV( et al. (2012) in a community with Brachypodium pinna- 2005, Patterns of secondary succession in calcareous 18 Tomasz Wójcik, Krystyna Towpasz

grasslands: can we distinguish the influence of former +HJHGãRYi. âNRGRYi, HGV 9HJHWiFLD6ORY- land uses from present vegetation data? Basic and Ap- HQVND5DVWOLQQpVSRORþHQVWYi6ORYHQVND7UDYLQQR plied Ecology 6: 161-173. E\OLQQiYHJHWiFLD>9HJHWDFLRQRI6ORYDNLD3ODQWFRP- $OoLWHSH((UNHQ6*OEDJ) g]]DPEDN0( PXQLWLHVRI6ORYDNLD*UDVVODQGYHJHWDWLRQ@6ORYDN Remarks using the SEM method species seeds of Gen- Akademy of Sciences, Bratislava. tiana occurring in . Planta Daninha 35: 1-9. +XOWpQ( )ULHV0$WODVRI1RUWK(XURSHDQYDV- %DEF]\ĔVND6HQGHN% $QGU]HMF]XN,Gentiana FXODUSODQWVQRUWKRIWKH7URSLFRI&DQFHU,,.RHOW] cruciata/LQWKHQHLJKERXUKRRGRI7DUQRZVNLH*yU\ 6FLHQWLILF%RRNV.|QLJVWHLQ Natura Silesiae Superioris 1: 33-42. .DPLĔVNL'3LHUQLN$ *UREHOVNL:0RUSKR- %DEF]\ĔVND6HQGHN%3UREOHP\ILWRJHRJUDILF]QH logical variability of Gentiana cruciata L. from differ- L V\QWDNVRQRPLF]QH NVHURWHUPyZ :\Ī\Q\ ĝOąVNLHM HQWKDELWDWV>LQ@WK&RQJUHVRIWKH(XURSHDQ9HJ- >3K\WRJHRJUDSKLF DQG V\QWD[RQRPLF SUREOHPV RI WKH HWDWLRQ 6XUYH\  0D\  :URFáDZ 3RODQG 6LOHVLDQ 8SODQG [HURWKHUPV@:\GDZQLFWZR 8QLZHU- 9HJHWDWLRQ VXUYH\  \HDUV DIWHU WKH SXEOLFDWLRQ RI V\WHWXĝOąVNLHJR.DWRZLFH Braun-Blanquet’s textbook – new challenges and con- %DEF]\ĔVND6HQGHN%%áRĔVND$ 6NRZURQHN, cepts. Book of Abstracts. Botanical Garden University New localities of Gentiana cruciata (Gentianaceae) RI:URFáDZ:URFáDZ LQWKH.UDNyZ&]ĊVWRFKRZD8SODQG)UDJPHQWD)OR- .DĨPLHUF]DNRZD 5  ;HURWKHUPLF JUDVVODQGV DQG ristica et Geobotanica Polonica 21(1): 67-76. shrubs of the Pieniny National Park. Studia Naturae %DUDEDV].UDVQ\ %  =UyĪQLFRZDQLH URĞOLQQRĞFL 49: 277-296. L VXNFHVMD ZWyUQD QD RGáRJDFK ZLHONRSRZLHU]FKQLR- .DĨPLHUF]DNRZD5=DU]\FNL-:UyEHO, 9RQþLQD G., Z\FK 3RJyU]D 3U]HP\VNLHJR >9HJHWDWLRQGLIIHUHQWLD- 2004, Meadows, pastures and wet habitat communi- tion and secondary succession on abandoned agricul- ties of the Pieniny National Park. Studia Naturae 49: WXUDOJUDQGDUHDVLQ3U]HP\ĞO)RRWKLOOV 6RXWK(DVWHUQ 195-251. 3RODQG @,QVW\WXW%RWDQLNLLP:6]DIHUD3ROVND$ND- .pU\00DWWKLHV' )LVFKHU07KHHIIHFWRI GHPLD1DXN.UDNyZ plant population size on the interactions between the %ąED:&KDQJHVLQWKHVWUXFWXUHDQGIORULVWLFFRP- rare plant Gentiana cruciata and its specialized herbi- position of the limestone grassland after cutting trees vore Maculinea rebeli. Journal of Ecology 89: 418-427. and shrubs and mowing. Acta Societatis Botanicorum .OH\HU0%HNNHU50.QHYHO,&%DNNHU-37KRPS- Poloniae 72(1): 61-69. VRQ.6RQQHQVFKHLQ03RVFKORG39DQ*URHQHQ- Braun-Blanquet J., 1964, Pflanzensoziologie. Grundzüge GDHO -0 .OLPHã / .OLPHãRYi - .ORW] 6 5XVFK GHU 9HJHWDWLRQVNXQGH :LHQ 6SULQJHU KWWSG[GRL G.M., Hermy M., Adriaens D., Boedeltje G., Bossuyt org/10.1007/978-3-7091-8110-2). %'DQQHPDQQ$(QGHOV3*|W]HQEHUJHU/+RGJ- &K\WUê 0 HG   9HJHWDFH ýHVNp UHSXEOLN\  VRQ-*-DFNHO$..KQ,.XQ]PDQQ'2]LQJD 7UDYLQQiDNHĜtþNRYiYHJHWDFH>9HJHWDWLRQRIWKH&]HFK :$ 5|PHUPDQQ & 6WDGOHU 0 6FKOHJHOPLOFK - 5HSXEOLF*UDVVODQGDQG+HDWKODQG9HJHWDWLRQ@$F- Steendam H.J., Tackenberg O., Wilmann B., Cornelis- ademia, Praha. VHQ-+&(ULNVVRQ2*DUQLHU( 3HFR% ']ZRQNR= /RVWHU6&KDQJHVLQVSHFLHVFRP- The LEDA Traitbase: a database of life-history traits position of an abandoned limestone grassland in rela- of Northwest European flora. Journal of Ecology 96: tion to climatic conditions. Polish Journal of Ecology 1266-1274. 59(4): 687-698. .RPSDáD%ąED$ %ąED:7KHVSRQWDQHRXVVXF- (OOLV /7 $VWKDQD $. 6ULYDVWDYD 3 2PDU , 5DZ- cession in a sand-pit – the role of life history traits and DW .. 6DKX9&DQR 0- &RVWD '3'LDV (0 species habitat preferences. Polish Journal of Ecology 'LDV'RV6DQWRV16LOYD-%)HGRVRY9(.R]KLQ 61(1): 13-22. M.N., Ignatova E.A., Germano S.R., Golovina E.O., .RQGUDFNL-*HRJUDILDUHJLRQDOQD3ROVNL>5HJLRQDO *UHPPHQ1-0,RQ56WHIDQXW69RQ.RQUDW0 JHRJUDSK\RI3RODQG@3:1:DUV]DZD -LPHQH]066XiUH]*0.LHEDFKHU7/HERXYLHU .R]XKDURYD (.$QFKHY 0(  3RSRY 37KH M., Long D.G., Maity D., Ochyra R., Parnikoza I., pollination ecology of Gentiana cruciata (Gentianace- 3OiãHN9)LDORYi/6NRXSi=3RSRQHVVL6$OHIIL ae) – specifics of a Bulgarian population in compari- 06DERYOMHYLü066DERYOMHYLü$'6DKD3$]L] sion to Dutch populations. Nordic Journal of Botany 01 6DZLFNL - 6XOHLPDQ 0 6XQ %< 9iĖD - 23(3): 365-372. :yMFLN7

ests”. Rocznik Przemyski, Nauki Przyrodnicze 45(5): 7RZSDV] .  *HRERWDQLFDO GHVFULSWLRQ RI WKH 65-79. 6WU]\ĪyZ )RRWKLOOV =HV]\W\ 1DXNRZH 8QLZHUV\WHWX .XFKDU]\N67KH[HURWKHUPLFJUDVVODQGZLWKWKH -DJLHOORĔVNLHJR5R]SUDZ\KDELOLWDF\MQH snowdrop windflower Anemone sylvestris L. on the 7UąED&:RODĔVNL3 2NOHMHZLF].&RPPXQL- 3U]HP\VNLH)RRWKLOOV 6(3RODQG &KURĔP\3U]\URGĊ ties with Brachypodium pinnatum and Bromus erectus 2MF]\VWą   in the Wiar and the San valleys. Annales Universitatis Matuszkiewicz W., 2001, Przewodnik do oznaczania zbi- 0DULDH&XULH6NáRGRZVND6HFWLR&   RURZLVNURĞOLQQ\FK3ROVNL>$JXLGHIRULGHQWLILFDWLRQRI :yMFLN72FFXUUHQFHRIGentiana cruciata (Genti- SODQWFRPPXQLWLHVLQ3RODQG@3:1:DUV]DZD anaceae) in a community with Brachypodium pinnatum 0LUHN=3LĊNRĞ0LUNRZD+=DMąF$ =DMąF0 (Festuco-Brometea LQ%XNRZD 3RJyU]H6WU]\ĪRZVNLH Flowering Plants and Pteridophytes of . A Check- foothills). Fragmenta Floristica et Geobotanica Polon- list. W. Szafer Institute of Botany, Polish Academy of ica 25(2): 205-215. 6FLHQFHV.UDNyZ :yMFLN 7  2FK\UD 5  7KH ILUVW ORFDOLW\ RI WKH 2FK\UD 5 ĩDUQRZLHF -  %HGQDUHN2FK\UD +  Ontario rose moss Rhodobryum ontariense in the Car- Census catalogue of Polish mosses. W. Szafer Institute SDWKLDQ)RRWKLOOV&KURĔP\3U]\URGĊ2MF]\VWą   RI%RWDQ\3ROLVK$FDGHP\RI6FLHQFHV.UDNyZ 130-140. 3HWDQLGRX 7 1LMV -&0  2RVWHUPHLMHU -*%  :yMFLN7 3LąWHN.1HZORFDOLW\RIGentiana Pollination ecology and constraints on seed set of the cruciata/LQWKH6WU]\ĪRZVNLH)RRWKLOOV :HVWHUQ&DU- rare perennial Gentiana cruciata L. in The . pathians). Steciana 19(2): 67-73. Acta Botanica Neerlandica 44(1): 55-74. :yMFLN7 =LDMD03ODQWFRPPXQLWLHVRI.DPL- 3LĊNRĞ0LUNRZD +  0LUHN =  )ORUD 3ROVNL HQLHFKLOOLQ'\QyZ)RRWKLOOV :HVWHUQ&DUSDWKLDQV  5RĞOLQ\FKURQLRQH>3ROLVKIORUD3URWHFWHGSDQWV@0XO- Parki Narodowe i Rezerwaty Przyrody 34(2): 57-74. tico Oficyna Wydawnicza, Warszawa. =DMąF$  =DMąF 0 HGV   'LVWULEXWLRQ DWODV RI Pielou E.C., 1974, Population and community ecology: vascular plants in Poland. Laboratory of Computer principles and methods. CRC Press, . Chorology, Institute of Botany, Jagiellonian Univer- Piotrowska J., 2010, Evaluation of preservation degree VLW\.UDNyZ within selected populations of Gentiana cruciata =DMąF 0  =DMąF $  7KH JHRJUDSKLFDO HOHPHQW in Western Pomerania. Folia Pomeranae Universitatis of native flora of Poland. Laboratory of Computer Technologiae Stetinensis 278(14): 57-74. Chorology, Institute of Botany, Jagiellonian Univer- 5R]SRU]ąG]HQLH0LQLVWUDĝURGRZLVND]GQLDSDĨG]LHUQLND VLW\.UDNyZ  Z VSUDZLH RFKURQ\ JDWXQNRZHM URĞOLQ ']8 =DU]\FNL.7U]FLĔVND7DFLN+5yĪDĔVNL:6]HOąJ= SR] >5HJXODWLRQRIWKH0LQLVWHURI(QYL- :RáHN- .RU]HQLDN8(FRORJLFDOLQGLFDWRU ronment of 9 October 2014 on the plant species protec- values of vascular plants of Poland. W. Szafer Institute WLRQ@:DUV]DZD RI%RWDQ\3ROLVK$FDGHP\RI6FLHQFHV.UDNyZ 6]F]HEOHZVND$ -DQHFNL-;HURWKHUPLFYHJHWD- =LDMD0 :yMFLN77KHUPRSKLOLFSODQWFRPPX- WLRQRIWKHKLOOVQHDUàXF]\FHDQG-DNVPDQLFHLQWKH QLWLHVLQ1DWXUDVLWH³àąNLQDG:RMNyZNą´3/+ YLFLQLW\RI3U]HP\ĞO :HVWHUQ2SROH 2FKURQD3U]\- 180051 – Podkarpacie Province. Annales Universitatis rody 56: 79-89. 0DULDH&XULH6NáRGRZVND6HFWLR&   7RZSDV].7KHYDVFXODUSODQWVRIWKH6WU]\ĪyZ)RRW- KLOOV=HV]\W\1DXNRZH8QLZHUV\WHWX-DJLHOORĔVNLHJR Prace Botaniczne 16: 1-160.

Ecological Questions 30 (2019) 1: 21–33 http://dx.doi.org/10.12775/EQ.2019.005

The vegetation of the cretaceous outcrops of Novhorod-Siverskyi Polesie loess “islands” () and the new locality of Gentiana cruciata L.

Oleksandr Lukash1,*, Iryna Miroshnyk2, Oleksandr Yakovenko1, Svitlana Strilets2

1Department of Ecology and Nature Conservation, 2Department of Languages and Methodology, T. Shevchenko National University “Chernihiv Collegium”, Hetman Polubotko Str., 53, 14013, Chernihiv, Ukraine, *e-mail: [email protected] Received: 21 December 2018 / Accepted: 22 February 2019

Abstract. The vegetation of the cretaceous outcrops of Novhorod-Siverskyi Polesie loess “islands” is represented by the grasslands communities of the Artemisietea vulgaris Lohmeyer et al. in Tx. ex von Rochow1951, Festuco-Brometea%U%OHW7[H[6Ry Trifolio-Geranietea sanguinei T. Müller 1962 classes and shrubby phytocoenoses of the Robinietea-XUNRH[+DGDþHW6RIURQ class. The structure and composition of the plant communities are influenced by the degree of anthropogenic influence (both in the past and present) on the ecosystems of cretaceous outcrops. The determining anthropogenic factors contributing to the formation of the ruderal communities were chalk mining and gardening. The influence of erosive processes is manifested in the spatial delimitation of plant communities of various syntaxonomic belongings. The anthropogenic successional communities: semiruderal grasslands and herblands of the immoral and subboreal zones of Europe (Convolvulo-Agropyretum repentis )HOI|OG\  Poo compressae- Tussilaginetum farfarae R. Tx. 1931) and scrub communities of temperate Europe, represented by the Elytrigio repentis-Robinietum Smetana 2002 phytocoenoses typical for Steppe zones, prevail. Semixerothermic communities were found on the steep slopes of the cretaceous outcrops (eastern and southwestern expositions) – early successional stages with a significant amount of the characteristic species of the Festuco-Brometea%U%OHW7[H[6RyFODVV2QWKHFUHWDFHRXVRXWFURSVRI1RYKRURG6LYHUVN\L3ROHVLHORHVV “islands” the communities of Trifolion medii T. Müller 1962 (meso-subxerophytic fringe phytocoenoses on nutrient-poor but base-rich soils at lower altitudes of temperate Western and Central Europe) is localized on the slopes of the eastern and southeastern parts of the expositions which do not undergo anthropogenic influence and are separated by erosion forms. Such conditions were favorable for preserving the Gentiana cruciata L. relict species in this locality. The population of this species was found in the area of 50 m2 in the Trifolio medii-Agrimonietum Th. Müller 1962 association community and represented by two compact groups of individuals (the area of 0.5 m2 each) and individual plants. In order to preserve the habitat of this rare species, it is worth creating here a reserve.

Keywords: Polesie, cretaceous outcrops, loess “islands”, plant communities, syntaxonomy, successional stages, rare plant species, anthropogenic impact.

1. Introduction and phytoindicative study of the plant communities of cretaceous outcrops of the Desna River right bank was Among other Polesie districts, Novhorod-Siverskyi Polesie FDUULHGRXWLQWKHRXWVNLUWVRIWKH.DPLQ3XVKNDUL5RKLYND is characterized by the considerable dismemberment and villages of the Novhorod-Siverskyi district, Chernihiv outcrop of indigenous sediments. This imposes an imprint UHJLRQ 6DYRQ  /\VHQNR   2XU UHVHDUFK FRYHUHG on the vegetation cover of the territory. The ecological the area located to the south of the earlier studies: in the 22 Oleksandr Lukash, Iryna Miroshnyk, Oleksandr Yakovenko, Svitlana Strilets

outskirts of the Putyvsk, Yukhnove and Horky villages. and its right bank tributaries, gullies, ravines cut the whole The goal of our study was to determine the composition thickness of anthropogenic and paleogene sediments and of plant communities of cretaceous outcrops of Novhorod- penetrate deeply into the thickness of cretaceous sediments. Siverskyi Polesie, to find out their syntaxonomic affiliation In the thickness of cretaceous sediments in the right bank and floristic features. The working hypotheses of the re- of the slopes of the Desna valley dark gray marls, lime- search: the cretaceous outcrops of Novhorod-Siverskyi stone glauconitic sands and white chalk are outcropped. Polesie are the potential places of formation of the xero- The surface of cretaceous sediments carries the marks of thermic and semixerothermic grass communities and the intense erosive dismemberment. The variation of absolute habitats of rare plant species, characterizing these commu- marks is from 125 to 165 m (Marynich, 1968). nities. One of such species is Gentiana cruciata L. – the In some parts paleogene sediments are partially blurred object of the population modelling the spatial interactions and anthropogenic sediments lie directly on chalky rocks. between plants and insects (Clarke et al., 1998). The lower layer of the anthropogenic strata is moraine. Moraine often lays the slopes of the right tributaries of the Desna River and its large gullies. The valleys of the right 2. Study area bank tributaries of the Desna River divide the explored area into several loess “islands”: Rohovskyi, Novhorod- The investigated area (Fig. 1A) covers the zone with the Siverskyi, Blystovitskyi, Ponornytskyi. The last ones are largest dismemberment and and outcrop of root sediments the continuation of the loess “islands” of Chernihiv Polesie, along the right bank of the meridial part of the Desna River which are characterized by a greater degree of synanthropy with the outcrops of indigenous sediments from the right of the vegetation cover (Lukash et al., 2018). bank of the lower part of the Sudost river to the turn of In the hypsometric plane, the territory is an elevated the Desna River to the southwest. The Desna River valley plain, absolute markings of which exceed 200 m. The ex-

Figure 1. Location of the cretaceous outcrops of Novhorod-Siverskyi Polesie loess “islands” (the square marked as “A”) Occurrence of Gentiana cruciata in dry grassland (Festuco-Brometea) in Kołaczyce (Strzyżowskie Foothills) 23

cess of the Desna is 100-105 m. The proximity of the deep The group of semiruderal alliances local base of erosion and the fact that the loess cover is Alliance:Convolvulo arvensis-Agropyrion repentis *|RUV easily eroded is the main cause of the exclusive develop- 1967 ment of the ravine and gully network, the density of which Association: Convolvulo-Agropyretum repentis )HOI|OG\ exceeds 1 km / km2. The meadow-steppe and synanthropic (1942) 1943 vegetation is well represented on the slopes of the ravines Association: Falcario vulgaris-Agropyretum repentis and gullies. 0OOHUHW*|UV Association: Poo compressae-Tussilaginetum farfarae R. Tx. 1931 3. Material and methods Class: Festuco-Brometea%U%OHW7[H[6Ry The group of orders of sub-xeric steppic grasslands The materials for the article were collected during the Order: Brachypodietalia pinnati.RUQHFN field research of the loess “islands” of Novhorod-Siverskyi Iinitial community: Chamaecytisus ruthenicus-Aster amel- Polesie during 2006-2018. The field study of the vegetation lus >Cirsio-Brachypodion pinnati+DGDþHW.OLNDLQ.OLND ZDV FDUULHG RXW E\ JHRERWDQLFDO PHWKRGV .RUFKDKLQ HW+DGDþ Molinio-Arrhenatheretea Tx. 1937+Tri- 2012). The vegetation descriptions were carried out during folio-Geranietea sanguinei70OOHU@ theoptimum of vegetation period in the areas of 30-50 m2. Iinitial community Elytrigia intermedia-Salvia pratensis The exposition and steepness of the slopes, the general >Cirsio-Brachypodion pinnati+DGDþHW.OLNDLQ.OLNDHW projective coverage of the vegetation community and the +DGDþ Molinio-Arrhenatheretea Tx. 1937 + Trifo- coverage of each species were noted Cover abundance lio-Geranietea sanguinei70OOHU@ scale is the following: + – up to 1%, 1 – 1-5%, 2 – 6-15, Initial community Origanum vulgare purum >Cirsio- 3 – 16-25%, 4 – 26-50%, 5>50%. 25 phytosociological Brachypodion pinnati+DGDþHW.OLNDLQ.OLNDHW+DGDþ UHOHYpVZHUHWDNHQ6\QWD[DZHUHLGHQWLILHGDFFRUGLQJWR 1944 + Molinio-Arrhenatheretea Tx. 1937 + Trifolio-Ge- Mucina et al. (2016), Matuszkiewicz (2001) (for natural ranietea sanguinei70OOHU@ vegetation), Solomakha et al. (1992) (for synanthropic veg- Class: Trifolio-Geranietea sanguinei T. Müller 1962 etation), Brzeg (2005) (for Trifolio-Geranietea sanguinei Order: Origanetalia T. Müller 1962 communities). Syntaxa names are ordered according to Alliance: Trifolion medii T. Müller 1962 Mucina et al. (2016). The successional stages of vegeta- Association: Trifolio medii-Agrimonietum Th. Müller 1962 tion are named by the dominant species. Class: Robinietea-XUNRH[+DGDþHW6RIURQ Order: Chelidonio-Robinietalia+DGDþHW6RIURQ Alliance: Balloto nigrae-Robinion pseudoacaciae +DGDþ 4. Results and discussion et Sofron 1980 Association: Elytrigio repentis-Robinietum Smetana 2002. A generalized scheme of the vegetation of the cretaceous 5HOHYpV 7DEOH EHORQJWRWKHConvolvulo ar- outcrops of Novhorod-Siverskyi Polesie loess “islands” is vensis-Agropyrion repentis association from the Agropyre- the following: talia intermedio-repentis order of the Artemisietea vulgaris Class: Artemisietea vulgaris Lohmeyer et al. in Tx. ex von class. The cenoses of the Convolvulo-Agropyretum repentis Rochow1951 association occupy the largest areas on the cretaceous sedi- Order: Agropyretalia intermedio-repentis70OOHUHW*|UV ments (Fig. 2B).

1969 [24]

Figure 2. Mapping of the vegetation of the cretaceous outcrops of Novhorod-Siverskyi Polesie loess “islands”. Syntaxon (1-8): 1 – Falcario vulgaris – Agropyretum repentis, 2 – Convolvulo arvensis – Agropyretum repentis, 3 – Poo compressae-Tussilaginetum farfarae, 4 – Chamaecytisus ruthenicus – Aster amellus, 5 – Elytri- gia intermedia–Salvia pratensis, 6 – Origanum vulgare purum, 7 – Trifolio medii-Agrimonietum, 8 – Elytrigio repentis-Robinietum; 9 – the habitat of Gentiana cruciata L. [25] .... SE SE S NE W ȿ ...... +.. +++++ ...... 21121422++. . . 1 . ++++ . . . . + . . . . + . 2+1 ...... +...... 32...... +...... ++...... 555...... +1+...... ++1...... +11 .+++..34+++.+++++++...... +535434354. .+. . .+. . .3343 00000000000000105000201040354550 41+...++...... +...... 0000000000000050000000000 ++...+..+.+.+...... ++. +.+.++..+. . .+++. +3...... +...... +. ++132.+1342+++...... 40 45 40 40 3635 42 20 48 45 2 40 1 35 2 40 30 30 10 25 30 30 30 5 36 10 42 0 36 0 25 50 0 42 0 50 35 40 30 45 40 35 35 40 10 5 20 15 10 5 90 95 95 90 95 100 50 70 65 85 70 80 75 85 45 50 25 25 20 90 90 25 20 35 20 @ 0 @ 2 Relevé 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 *Elytrigia intermedia Campanula bononiensis Ch., *D. All. Cirsio- Ch., *D. Brachypodion pinnati Aster amellus Iris aphilla Poa angustifolia Cerastium arvense &K '$VV3RRFRPSUHVVDH farfarae –Tussilaginetum farfara Tussilago stolonifera *Agrostis *Poa compressa *Ranunculus repens *Bunias orientalis Ch. Ass. Convolvulo- repentis Agropyretum Convolvulus arvensis Ch. All. Convolvulo arvensis- Ch. Agropyrion repentis Agropyretalia Ch. O. intermedio-repentis, Artemisietea vulgaris Ch. Cl. inermis Bromopsis Elytrigia repens $UHD>P Syntaxon 1 2 3 4 5 6 7 8 Calamagrostis epigeios Calamagrostis 6KUXEOD\HUFRYHU>@ Ass. Falcario Ch., *D. vulgaris-Agropyretum repentis Falcaria vulgaris +HUEOD\HUFRYHU>@ 0RVVHVFRYHU>@ Exposition of the slope N N SE SE NE E W S W S NE S SE E E SW SW E SW 6ORSHVWHHSQHVV> Table 1. Floristic structure of plant communities the cretaceous outcrops Novhorod-Siverskyi Polesie loess “islands” Table [26] ...... +.+..... +.+. . +.+...... +++++...... ++...+++.+++.+...+...... +++..+...... +..+...... +...... 1++...... ++...... +.....+.+++...... + ...... + . . . . +++++++ ...... ++.....+...... +...... +...++ ...... +++++...... ++...... ++++.+.+...... +...... +...+...... ++.++...... +...... ++..+...... ++++...... +++++.+...... + .+.+.....+....+..++...... +...... +...+.+...+++...+...... ++.+...... +...... +.....+.+++...... +...... ++...... ++....+....+.++...... +1+...... +.+.+++..+...... 1+.. .+.+.++.+. .+...... +.....+...+..++ . .+...... +....+.+...+.... +.+...... +...++.+++..... +.+++...++.....+++...... +.+....+...... +++++...... subsp Relevé 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 Trifolium prantense Trifolium Tragopogon orientalis Tragopogon Linum flavum Thalictrum simplex Ch. O. Brachypodietalia pinnati Anthemis tinctoria Carex praecox Carex subtinctoria Rumex acetosa Campanula glomerata Asparagus officinalis Campanula sibirica Echium russicum Carex humilis Carex Salvia verticillata Scabiosa ochroleuca Brachypodium pinnatum Thymus tschernjajevii rupestris Viola Ch. Cl. Festuco-Brometea Anthyllis macrocephala Thymus marschallianus Euphorbia cyparissias Filipendula vulgaris Gentiana cruciata Plantago media Poa bulbosa Stachys recta Stachys recta Veronica spicata Veronica Ch. Cl. Molinio- Arrhenatheretea elatius Arrhenatherum Campanula patula Phleum pratense Plantago lanceolata Ranunculus acris Leucanthemum vulgare Leucanthemum vulgare [27] ...... ++++ 1.... 1 .+.++...... +1...... 1..++...... ++...... ++...... + . . . . . +++++++ ..+++...+ .....++..+1...... +++.....+.... .+...... ++. . .+++1+32++.+ .+. . . . .+.+. .+1..2....+..++....++...... ++...... ++. ...++...... +..+...+...... ++...++...++..+..+...... +. .+.++.. .+.+.++.++..+...... +..+++...... +..+...... +....+++. ...+++...... +....++...... +...... +.1+.... .+.+. . .+.+ . . .+. .+.+1 .+...+.....+...... +.+ ...... +..+.. .+...... +.....++.+++.... .+...... +.+...++...... +. . .+. . .+++.+. . . .+++++. 2 . 3+4 . . + . . . ++++ . . . . ++ +.3+++.++...... +..... +.+22..+..+...+++..31.... +++++ . + . +++ +.1....+...... +++2211.... +.+...... +. .+++. .+.++ +.... (b) Relevé 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 Clinopodium vulgare Clinopodium vulgare *Centaurea jacea *Centaurea *Frangula alnus Ch., *D. Ass. Trifolio medii- Trifolio Ass. Ch., *D. Agrimonietum Agrimonia eupatoria Fragaria viridis Galium verum Securigera varia *Daucus carota *Daucus carota *Medicago falcata *Festuca pratensis Trifolium medium Trifolium Galeopsis bifida Origanum vulgare Origanum vulgare *Pimpinella saxifraga *Solidago virgaurea Ch., *D. All. Trifolion medii Trifolion All. Ch., *D. *Achillea submillefolium *Campanula rotundifolia Carex hirta Carex Verbascum lychnitis Verbascum Dactylis glomerata Galium mollugo Knautia arvensis D. Ass. Elytrigio repentis- Robinietum Artemisia vulgaris *Lathyrus pratensis Vicia sepium Vicia Vicia cracca Vicia Veronica chamaedrys Veronica Ch., *D. O. Origanetalia Ch., *D. Cl. Trifolio-Geranietea sanguinei Astragalus cicer [28] ...... 3334 ++++ ++++ .+.+. .+...... +...+...... +..+.1++1 .+...... +...... +...... ++++ ...... +++ ..+...... +...... ++.. ...+..+...... +..1 ...... 1+11 ..+..+.+++...... ++.+ .+...... +...+..+..+1+...... ++.5...... 23+..32...... +.+....+...... ++++ .+.+1...... +...... +121 ...4+...... +...... +.... .+...... ++ ...... 1+ .....+...... ++...... +...... ++.+ ...... +++ ...... +.++ ...... +1 ...... 11.....1+.+ ...... +.++ +.112+.+ + +. . . . .++.+.+.+.+.. . +.+. .+.+.+. .++. . .++. . ++2.....+..+..+..++...... +...... ++....+....+..... ++ . 1++ . . .. + .+.+...... ++...... + . . ++++ . . . + . . ɫ (b) (b) (c) (b) (b) (b) (c) Relevé 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 Caragana arborescens Caragana arborescens Centaurea diffusa Centaurea Ballota nigra Anthriscus sylvestris Artemisia absinthium Dianthus fischeri Acer tataricum D. O. Chelidonio-Robinietalia D. Cl. Robinietea Acer negundo Acer negundo Cichorium intybus Taraxacum officinale Taraxacum Centaurea phrygia Centaurea Chamaecytisus ruthenicus Berteroa incana Berteroa Robinia pseudoacacia Robinia pseudoacacia Artemisia scoparia Lactuca serriola Artemisia marschalliana D. All. Balloto nigrae- D. Robinion pseudoacaciae Chenopodium album Equisetum pratense Eryngium planum Echium vulgare Echium vulgare Chelidonium majus Fallopia dumetorum Geum urbanum Geranium robertianum Lapsana communis Sisymbrium loeselii Swida sanquinea species Other Agrimonia pilosa Allium paczoskianum Humulus lupulus [29] ...... Chamaecytisus ruthenicus– iverskyi district, between the 4- , 7, 25 – 16.06.2018 Elytrigio repentis-Robinietum 14 – the Chernihiv region, Novhorod-Siverskyi ...... , 8 – compressae-Tussilaginetum farfarae compressae-Tussilaginetum , 3 – Poo ....+1.+...... +.+. . Trifolio medii-Agrimonietum Trifolio 7 – ...... 1...... +.+.+. .++.+..1. Origanum vulgare purum, Origanum vulgare Convolvulo arvensis-Agropyretum repentis Convolvulo arvensis-Agropyretum , 6 – , 2 – .+.++.....+..+.+.+..+...... +..+...++.....+..+...... +...+..+.+...... ++...... +...... +++..+.+...... +...... +.+..+...... +. . . .+.+.+ ..+...... +....+...... +....+...... +...... +...... +.+....+....+...... +1. . . .+.++.+.+1.+. .+...... +...+....+1...... +.+...... +.1+...++....+.....+.... .+.++...+.+.++.+...... 1.+....++.....+.2...+.... +.+1++.+.....++.+..1+.... +..++...++....+...... +.2+4..2...... +.+...... +.+.....+.....+..+...... +.+++..+...... +...... 1-11, 15-19, 22-25 – the Chernihiv region, Novhorod-Siverskyi district, Putyvsk village; 12 Novhorod-S 1-11, 1-11, 18-23 – O. Lukash, 12, 16, 17, 24 – O. Yakovenko, O. Lukash, 13-15, 25 – I. Miroshnyk, S. Strilets. Yakovenko, 18-23 – O. Lukash, 12, 16, 17, 24 1-11, (b) Elytrigia intermedia–Salvia pratensis 5- – Falcario vulgaris-Agropyretum repentis – Falcario vulgaris-Agropyretum sp. 1 Relevé 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 – dominant species of the initial community. 1-9 – 16.08.2006, 10-17.09.2012, 11-24.07.2016, 12, 16, 17-25.07.2017, 13-15 – 16.06.2018, 22, 23 24.07.2016, 24 25.07.201 1-9 – 16.08.2006, 10-17.09.2012, 11-24.07.2016, Trifolium campestre campestre Trifolium Salvia pratensis Rubus caesius Sedum ruprechtii Senecio jacobaea Thalictrum aquilegifolium Pyrus communis Potentilla impolita Psammophiliella muralis Euphrasia stricta Melilotus officinalis Nonea rossica Odontites vulgaris Phlomis tuberosa Polytrichum Euphorbia stricta Pastinaca sylvestris Leontodon hispidus Libanotis intermedia Melilotus albus Helichrysum arenarium Helichrysum arenarium Lavatera thuringiaca Hieracium umbellatum Hypericum perforatum Note: Syntaxon: Locality of relevés: and Putyvsk villages; 13 – the Chernihiv region, Novhorod-Siverskyi district, between Horky Horbove Yukhnove district, Putyvsk village; 20-21 – the Chernihiv region, Novhorod-Siverskyi between Horky and Horbove villages. Date: Authors of relevés: Aster amellus, 30 Oleksandr Lukash, Iryna Miroshnyk, Oleksandr Yakovenko, Svitlana Strilets

The Convolvulo-Agropyretum repentis phytocoenoses vulgaris Moench, Plantago media L., Poa bulbosa L., Poa were formed in the areas covered with a 0.5-1.0-meter compressa L., Stachys recta L., Veronica spicata L. in par- layer of loess sediments. These areas were used by the ticular. There are the characteristic species of the Brachy- locals 20-25 years ago as kitchen gardens, and now they podietalia pinnati order and the Cirsio-Brachypodion pin- are experienced such types of anthropogenic influence as natiDOOLDQFHLQDOOWKHILYHUHOHYpV+RZHYHUWKHVWUXFWXUH cattle grazing and recreational load. The communities are and (or) composition of the described communities does formed by such species as Elytrigia repens (L.) Nevski not allow to clearly refer them to one or another associa- and Convolvulus arvensis L., sometimes Calamagrostis tion of the alliance mentioned above. Note that there is epigeios (L.) Roth. co-predominates, Cerastium arvense a number of species according to which the Molinio-Ar- L., Elytrigia intermedia, Poa angustifolia L., P. compres- rhenatheretea and Trifolio-Geranietea sanguinei classes sa L. Bromopsis inermis (Leyss.) Holub are singly found. DUHGLDJQRVHGLQUHOHYpV3UREDEO\WKHSK\WRFRHQR- The species of the Molinio-Arrhenatheretea and Festuco- ses described by us are the communities at early stages of -Brometea classes take part in this association communities successions with domination of Chamaecytisus ruthenicus formation. )LVFK([:Rá .OiVNRYiDQGAster amellus/ UHOHYpV Consequently, the Convolvulo-Agropyretum repentis 15-16), Elytrigia intermedia (Host) Nevski and Salvia ruderal communities described by us, which are characte- pratensis/ UHOHYp DVZHOODVOriganum vulgare L. rized by a high participation of biennial or perennial plants, UHOHYpV $SURMHFWLYHFRYHUDJHRIWKHGRPLQDQWVLV are mostly a succession stage, which is replaced by the 15-25% with a total projective cover of 20-50%. As part community of Sisymbrion officinalis (Class Papaveretea of these communities, there is a number of rare for Polesie rhoeadis). species that are situated in this region on the northern bor- The communities, belonging to the Falcario vulgaris- der of distribution. For example: Aster amellus L., Carex -Agropyretum repentis association, occur on the northern praecox Schreb, Echium russicum J.F. Gmel, Iris aphilla slopes near the Putyvsk village (Fig. 2B). These phytoco- L. and Linum flavum L., Salvia verticillata L., Phlomis tu- enoses occupy small “islets” between erosive dams. They berosa L. and others. Thus, the recorded xerothermophilic are diagnosed by the dominant species of Falcaria vulgaris communities have an environmental significance and are Bernh. and Bunias orientalis L. the objects of monitoring researches. The foot of cretaceous outcrops is occupied by the ;HURWKHUPLF DQG VHPL[HURWKHUPLF VWHSSH FRPPXQL- phytocoenoses belonging to the Poo compressae-Tussila- ties in Polesie are rare. For the Western Polesie within ginetumDVVRFLDWLRQ )LJ$%7DEOHUHOHYpV  WKHERUGHUVRI%HODUXV3RODQGDQG8NUDLQH )LMDáNRZVNL As a rule they were formed in the places of industrial and et al., 2002), as well as the “Prybuzhskoe Polesie” Bio- spontaneous mining of chalk. It should be noted that in- sphere Reserve (Demyanchik, 2006), the presence of xe- dustrial mining of chalk was carried out near the Putyvsk rothermic grass communities of the Festuco-Brometea village in the 70’s-90’s years of the twentieth century. class on the cretaceous sediments is indicated. Such com- Unauthorized local chalk mining is continued to the pre- munities are not mentioned for the Polesie National Park sent time. Tussilago farfara L. (the characteristic species %DU\ODHWDOĝZLĊV DQG3ROHVLH1DWXUDO5H- of the association) dominates in all of the described are- VHUYH 9RURE\RYHWDO :LWKLQ8NUDLQLDQ 6RXWK- as. These communities are also differentiated by Agrostis ern) Polesie the Festuco-Brometea steppe communities stolonifera L., Poa compressa L. and Ranunculus repens are known in Zhytomyr Polesie near the rivers, where L. Depending on the time of phytocoenoses formation the crystalline sediments are lying off (Onishchenko, 2006). areas of the Poo compressae-Tussilaginetum association But they have not been studied in detail. The communities vary by the number of species. In the areas, where indus- of the Festuco-Brometea class in the Briansk region (Rus- WULDOPLQLQJRIFKDONVWRSSHG\HDUVDJR 7DEOHUHOHYp sia) within the boundaries of the loess plateau landscapes 14), 34 species were recorded, among which rhizome pe- in the western spurs of the Middle Russian Highlands, rennials predominate, as well as tree and shrub species (on where they are on the northern border of their habitat (Bu- the level of the grassy tier). lokhov, 2001, 2009) have been investigated to the great- In the absence of anthropogenic pressure, grass com- est extent. Within Eastern Polesie these are the closest munities with traits of steppe phytocoenoses were formed xerothermic phytocoenoses in the cretaceous sediments to on the steep slopes of the cretaceous outcrops of the eas- the communities described by us. On the territory of the WHUQDQGVRXWKZHVWH[SRVLWLRQV )LJ%7DEOHUHOHYpV Briansk region, in the composition of the Festuco-Brom- 15-19). This is evidenced by the presence of the characte- etea class one Festucetalia valesiacae 6Ry  RUGHU ristic species of the Festuco-Brometea class in the descri- Cirsio-Brachypodion pinnati +DGDþHW.OLNDLQ.OLNDHW bed areas, Anthyllis macrocephala Wender, Brachypodium +DGDþDOOLDQFHZLWKRQHPoo compressae-Onobry- pinnatum (Huds.) P.Beauv., Campanula glomerata L., Ca- chidoetum arenariae Bulokhov 1990 association (Bulok- rex humilis Leyss., Euphorbia cyparissias L., Filipendula hov, 2001) was established. The diagnostic species of this Occurrence of Gentiana cruciata in dry grassland (Festuco-Brometea) in Kołaczyce (Strzyżowskie Foothills) 31

association are Onobrychis arenaria .LW '&DQGPoa We believe that the Elytrigio repentis-Robinietum phy- compressa/7KHFRPPXQLWLHVRIWKLVDVVRFLDWLRQɫDQEH tocoenoses are the last stage of the overgrowth of the cre- found in small sections on the steep eroded slopes of river taceous outcrops slopes in the succession series: ruderal valleys and gullies on the complex of ravine-gully gray phytocenoses of nutrient-demanding short-lived winter forest soils spread by chalk. The comparison of phytocoe- annual grasses on sandy anthropogenic soils (Sisymbrion notic data makes it possible to note that the communities officinalis Tx. et al. ex von Rochow 1951) Æ semiru- described by A. Bulokhov (2001, 2009) differ from the deral grasslands and herblands (Convolvulo-Agropyretum xerothermic phytocoenoses on the cretaceous sediments repentis)HOI|OG\   Æ ruderal shrub communi- of Novhorod-Siverskyi Polesie with a greater representa- ties Elytrigio repentis-Robinietum Smetana 2002). tion of steppe species. In chalk outcrops in the community of the Trifolio me- ,QUHOHYpVDQG 7DEOH WDNHGRXWDWWKHFKDON dii-Agrimonietum association in the area of 50 m2 (Fig. 2A; outcrops near the Horky village, the characteristic species 7DEOHUHOHYp WKHGentiana cruciata population was (Astragalus cicer L., Clinopodium vulgare L., Securigera determined. The middle density of the population was 0.34 varia (L.) Lassen, Origanum vulgare, Verbascum lychnitis individuals/m2. It is represented by two compact groups L.) and diagnostic species (Frangula alnus Mill., Medicago of 3 and 4 generative individuals, the area of 0.5 m2 each, falcata L., Pimpinella saxifraga L.., Solidago virgaurea L.) and 10 juvenile individual plants. The plants of other on- of the Trifolio-Geranietea class and the Origanetalia order togenetic states have not been found. Thus, the population were identified. The described community is referred to the of G. cruciata in a new locality can be characterized as Trifolion medii alliance due to the presence of a number incompletely limbed with a left-sided spectrum: juvenile of diagnostic species, among which are the species of plants predominate. In Fig. 3 the spatial structure of the G. the specified alliance: Agrimonia eupatoria L., Galium cruciata population is represented. mollugo L., Trifolium medium L., Vicia sepium L., and G. cruciata – a European-Southwest Asian forest- the typical species of the Molinio-Arrhenatheretea class steppe relict species, included in the Red Books of the (Achillea submillefolium .ORNRYHW.U\W]NDCampanula Republic of (Skuratovich, 2015) and the Briansk rotundifolia L., Dactylis glomerata L., Knautia arvensis region (Evstigneev, 2004). This species is very rare for the (L.) Coult., Lathyrus pratensis L., Veronica chamaedrys Eastern Polesie. The nearest to the identified place is the L., Vicia cracca L. et al.). Its belonging to the group of the “Markovsk mountains” (the Briansk region) – the richest neutrophilic associations shows the presence of Geranium in the Eastern Polesie center of the calcephalous flora. For sylvaticum, Medicago falcata L., Securigera varia. A. today, our discovery of G. cruciata is the first and only Brzeg (2005) points out the characteristic (Agrimonia eu- one for Novhorod-Siverskyi Polesie. It should be noted, patoria, Trifolium medium L.) and differential (Centaurea that G. cruciata is a diagnostic species of the Festuco- jacea L., Daucus carota L., Festuca pratensis+XGV Po- Brometea%U%OHW7[H[6RyFODVV+RZHYHUWKH tentilla reptans L.) species for the Trifolio medii-Agrimo- conditions of the determined location (open slope of the nietum association. All these species, except the last one, eastern exposition, close occurrence of carbonate rocks) were recorded in the phytocoenoses described by us. That are typical for the location of this species. For compari- LVZK\ZHUHIHUUHGWKHVHFRPPXQLWLHVIURPUHOHYpVDQG son, in Western Europe (in particular, in Poland), the lo- 21 to the specified association. cations of G. cruciata were found in xerothermic grass- In the outskirts of the Putyvsk village on the slopes of land on the southern and south-western slopes of the river the cretaceous outcrops scrub communities of temperate valleys in the communities that are characterized by high Europe, represented by non-typical for Polesie synanthrop- proportions of species of the Festuco-Brometea, Molinio- ic Elytrigio repentis-Robinietum phytocenoses, which are Arrhenatheretea, Trifolio-Geranietea sanguinei and Rham- characteristic of the Steppe zone of Ukraine, were formed no-Prunetea FODVVHV :yMFLN  3LąWHN  :yMFLN )LJ%7DEOHUHOHYpV 7KHVHFRPPXQLWLHVDUH 2018). In Western Pomerania the population of this spe- formed by Robinia pseudoacacia L. (3-4 m high) with an cies was found in Adonido-Brachypodietum pinnati com- admixture of Acer negundo L. The reason for referring the munities for which G. cruciata is a characteristic species identified communities to the corresponding association is (Piotrowska, 2010). It is worth noting that the populations the presence of the diagnostic species (Chenopodium al- of G. cruciata in western localities, in comparison with bum L., Lactuca serriola L., Taraxacum officinale Wigg. the populations in Novhorod-Siverskyi Polesie, are larger Aggr.) of the Balloto nigrae-Robinion pseudoacaciae in size, more numerous and denser. For example, in Br- +DGDþHW6RIURQDVVRFLDWLRQDQGWKHGLDJQRVWLFVSH- wice population 183 individuals of G. cruciata were found cies (Elytrigia repens (L.) Nevski with a 20-40% projective in the area 1200 m2 (Piotrowska, 2010). 1107 individuals covering, as well as Artemisia vulgaris L., Carex hirta L., of G. cruciataZHUHIRXQGLQ8QLVáDZORFDOLW\WKHKLJKHVW Galeopsis bifida Boenn. Humulus lupulus L.) of the Ely- frequency and density was 0.339 individuals/m2 .UDVLFND trigio repentis-Robinietum association. .RUF]\ĔVNDHWDO 32 Oleksandr Lukash, Iryna Miroshnyk, Oleksandr Yakovenko, Svitlana Strilets

Figure 3. Spatial structure of the G. cruciata population in the Novhorod-Siverskyi locality. Denotations: j – juvenile individuals, g – generative individuals

The G. cruciata locality in Novgorod-Siverskyi Polesie the cretaceous outcrops slopes during the succession from loess “islands” is not under conservation. Therefore, in or- the ruderal vegetation of nutrient-demanding short-lived der to preserve the habitat of this rare species, it is worth winter annual grasses on sandy anthropogenic soils to creating here a reserve. Robinia groves with weedy understorey on loamy dry soils. The initial semixerothermic communities (Chamae- cytisus ruthenicus-Aster amellus, Elytrigia intermedia- 5. Conclusions Salvia pratensis, Origanum vulgare purum) are close to the phytocoenoses of the Festuco-Brometea class by the The structure and composition of the vegetation commu- species composition. Natural meso-subxerophytic fringe nities of the cretaceous outcrops of Novhorod-Siverskyi vegetation on nutrient-poor but base-rich soils (Trifoli- Polesie loess “islands” is influenced by the degree of an- on medii T. Müller 1962) are represented fragmentarily. thropogenic influence (both in the past and present) on They do not suffer from the anthropogenic pressure and the ecosystems. The determining anthropogenic factors are separated by erosive forms. Such conditions were contributing to the formation of the ruderal communities favorable for preserving the relict Gentiana cruciata L. were chalk mining and gardening. The influence of erosive species in this locality. The population of this species processes is manifested in the spatial delimitation of plant was found in the area of 50 m2 in the community of the communities of various syntaxonomic belongings. Trifolio medii-Agrimonietum Th. Müller 1962 association In the vegetation cover of the cretaceous outcrops of and represented by two compact groups of individuals (the Novhorod-Siverskyi Polesie loess “islands” semiruderal area of 0.5 m2 each) and individual plants. In order to pre- grasslands and herblands phytocoenoses of the nemoral serve the habitat of this rare species, it is worth creating and subboreal zones of Europe belonging to the Convolvulo here a reserve. arvensis-Agropyrion UHSHQWLV *|RUV  DVVRFLDWLRQ predominate. The Elytrigio repentis-Robinietum Smetana 2002 phytocenoses is the last stage of the overgrowth of Occurrence of Gentiana cruciata in dry grassland (Festuco-Brometea) in Kołaczyce (Strzyżowskie Foothills) 33

References *XHUUD$6(UPDNRY19DODFKRYLþ06FKDPLQüH- H.J., Lysenko T., Didukh Y.P., Pignatti S., Rodwell J.S., Capelo J., Weber H.E., Solomeshch A., Dimopoulos P., %DU\áD5 8UEDQ'(NRV\VWHP\áąNRZH>0HDGRZ $JXLDU&+HQQHNHQV60 7LFKê/9HJHWDWLRQ HFRV\VWHPV@>LQ@65DGZDQD HG 3ROHVNL3DUN1DU- of Europe: hierarchical floristic classification system of RGRZ\ >3ROHVLH 1DWLRQDO 3DUN@ :\GDZQLFWZR 025- vascular plant, bryophyte, lichen, and algal communi- POL, Lublin: 201-215. WLHV$SSOLHG9HJHWDWLRQ6FLHQFH 6  KWWSV %U]HJ$=HVSRá\NVHURWHUPRILOQ\FK]LRáRURĞOLRNUD- onlinelibrary.wiley.com/doi/epdf/10.1111/avsc.12257). jkowych z klasy Trifolio-Geranietea sanguinei Th. Mül- Marynich A.M. (ed.), 1968, Physical and geographical zon- ler Z3ROVFH>;HURWKHUPRSKLORXVIRUEIULQJHVDQG LQJ RI WKH 8NUDLQLDQ 665 3XEOLVKLQJ +RXVH RI .LHY forest edge communities of the class Trifolio-Gerani- 8QLYHUVLW\.LHY etea sanguinei Th. Müller  LQ 3RODQG@ %RJXFNL 2QLVFKHQNR9$)ORULVWLFFODVVLILFDWLRQRIWKH8NUDL- :\GDZQLFWZR1DXNRZH3R]QDĔ QLDQ3ROHVLHYHJHWDWLRQ>LQ@7/$QGULHQNR HG 3K\- Bulokhov A.D., 2001, Grass vegetation of the South-West- todiversity of the Ukrainian Polesie and its conservation. ern Non-Black Earth Region of Russia. Publishing 3K\WRVRFLRFHQWUH.\LY House of Bryansk State University, Bryansk. Piotrowska J., 2010, Ocena stanu zachowania wybranych Bulokhov A.D., 2009, Typology of meadows of the Bryansk SRSXODFML JRU\F]NL NU]\ĞRZHM Gentiana cruciata) na region. Publishing House of Bryansk State University, 3RPRU]X =DFKRGQLP >(YDOXDWLRQ RI SUHVHUYDWLRQ GH- Bryansk. gree within selected populations of Gentiana cruciata Clarke R.T., Thomas J.A., Elmes G.W., Wardlaw J.C., Mun- LQ:HVWHUQ3RPHUDQLD@)ROLD3RPHUDQDH8QLYHUVLWDWLV JXLUD0/ +RFKEHUJ0(3RSXODWLRQPRGHO- Technologiae Stetinensis, Agric., Aliment., Pisc., Zoo- ling of the spatial interactions between Maculinea rebeli tech. 278(14): 57-74. their initial foodplant Gentiana cruciata and Myrmica ants 6DYRQ$3  /\VHQNR +0  (FRORJLFDO DQG SK\- within a site. Journal of Insect Conservation 2(1): 29-37. toindicative characteristic of plant communities of cre- 'HP\DQFKLN97³3ULEX]KVNR\H3ROHVLH´%LRVSKHUH taceous outcrops of Novhorod-Siverskyi Polesie. Eco- Reserve, I.I. Lishtvan (ed.). PH Academy, Brest. logical state of the Chernihiv region and ways of its Evstigneev O.I., 2004, Gentiana cruciata/>LQ@LQ@ ,0 )LMDáNRZVNL ' $QGULHQNR 7 2Q\V]HQNR : ĝZLĊV ) .DFKDQRYVN\0(1LNLIRURY9,3DUIHQRY HGV 7KH  8UEDQ '  (NRV\VWHP\ OąGRZH ± OHĞQH NVH- Red Book of the Republic of Belarus. Plants. Petrus rotermiczne i synantropijne Polesia (Flora i fitocenozy) Brovka Belorussian Encyclopedia, Minsk: 184-185. >7HUUHVWULDO DQG IRUHVW [HURWKHUPLF DQG V\QDQWKURSLF 6RORPDNKD9$.RVW\ORY29 6KHOLDK6RVRQNR6\QDQWKURSLF Zasoby populacji Gentiana cruciata/QDSU]\NáDG]LH HFRV\VWHPV@>LQ@65DGZDQD HG 3ROHVNL3DUN1DU- PXUDZ\ NVHURWHUPLF]QHM Z 8QLVáDZLX >5HVRXUFHV RI RGRZ\ >3ROHVLH 1DWLRQDO 3DUN@ :\GDZQLFWZR 025- Gentiana cruciata L. population of kerothermic grass- POL, Lublin: 216-230. ODQGVLQ8QLVáDZ@(NRORJLDL7HFKQLND   9RURE\RY2FFXUUHQFHRIGentiana cruciata (Genti- ]ELRURZLVNURĞOLQQ\FK3ROVNL>*XLGHIRUGHWHUPLQDWLRQ anaceae) in a community with Brachypo-dium pinnatum RI3ROLVKSODQWFRPPXQLWLHV@:\GDZQLFWZR1DXNRZH (Festuco-Brometea LQ%XNRZD 3RJyU]H6WU]\ĪRZVNLH PWN, Warszawa. IRRWKLOOV @)UDJPHQWD)ORULVWLFDHW*HRERWDQLFD3RORQL- 0XFLQD / %XOWPDQQ + 'LHU‰HQ . 7KHXULOODW -3 ca 25(2): 205-215. 5DXV7ýDUQL$âXPEHURYi.:LOOQHU:'HQJOHU :yMFLN7 3LąWHN.1HZORFDOLW\RIGentiana - *DUFtD 5* &K\WUê 0 +iMHN 0 'L 3LHWUR 5 cruciata/LQWKH6WU]\ĪRZVNLH)RRWKLOOV :HVWHUQ&DU- ,DNXVKHQNR'3DOODV-'DQLɺOV)-$%HUJPHLHU( pathians). Steciana 19(2): 67-73.