Geographical distribution , habitats and modern state of Carlina cirsioides ( Asteraceae ) populations

The study of the current state and habitats of rare and threatened species of plants is a relevant task of ecological research. The paper presents the results of the study of systematic position, habitats and modern state of populations of a rare species included in the Red DataBook of Ukraine Carlina cirsioides Klokov (=C. acaulis L.). Carlina cirsioides is well-known to European phytosozologists as an endemic and relic species of the Flora of Ukraine. At the same time, the species independence of this taxon is not recognized by the authors of monographs on the Carlina genus and by the authors of “Flora Europaea”. Comparative morphological, chorological and ecological-coenotic analyses do not give sufficient reasons to consider C. cirsioides described by Klokov as a separate species, endemic and relic species of the Flora of Ukraine. According to the morphological traits, the specimens from the lowland part of Ukraine belong to the C. acaulis caulescens subspecies, which is distributed mostly in the lowlaand regions of Europe. Ecological-coenotic conditions of habitats of C. acaulis in the plains of Ukraine are different from those in the mountain regions and are close to the plains habitats of this species in Central Europe. Steppe communities of class FestucoBrometea, of which C. acaulis is component in the plains part of Ukraine, are close to xerothermic herbaceous communities of Central Europe; forest communities of Erico-Pinetea with C. cirsioides in Ukrainian Polissia and the North-East Poland are very сlose by floristic composition. The removal of separate species status of the plains populations of C. acaulis near the eastern border of the range does not at all downplay its sozological significance. Taking into account the low number of C. acaulis in the Volhynian-Podolian Upland and in the Polesian Lowland, all localities of this species in the lowland part of the range in Ukraine must be taken under protection in situ.


Introduction
Development of the scientific basis of the protection of floristic diversity is a relevant task of ecological surveys at the current stage. Special attention should be drawn to rare and threatened species of plants of the natural flonas. Providing protection to them is possible based on integrated in detail studies of geographic distribution, habitats and modern state of populations. In the conditions of intense anthropogenic pressure on the ecosystems, an urgent task, according to the Global Strategy of Plant Conservation (Heywood, 2009;Diekson, 2013), is monitoring the state of populations of endemic, relic and rare species, the ranges of which are represented by low number of small populations (Falk & Holsinger, 1991;Brigham & Schwartz, 2003;Berg et al., 2014;Corlett, 2016;Volis, 2016). Special attention in phytosozological aspect should be paid to the studies of spatial and age structures of populations of rare and extinct species of natural floras (Bowman et al., 2010;Quinones-Perez et al., 2014;Corlett, 2016;Volis, 2016).
An effective method of protection of floristic diversity is conservation of rare exceptional species within genera complexes. The genus Carlina L. (Asteraceae) is rich in rare species, including 28 species distributed in the Holarctic kingdom (Europe, North Africa, Siberia) from the Azores and Canary Islands to North-West China (Meusel & Kästner, 1994;Wahrmund et al., 2010). The flora of Europe includes 13 species of the Carlina genus (Webb, 1976), and the flora of Ukraine contains 5 species: C. acaulis L., C. biebersteinii Bernh. ex Hornem., C. vulgaris L., C. onopordifolia Besser ex Szafer, Kulcz. & Pawl. (=C. acanthifolia All.), C. cirsioides Klokov (Klokov, 1962). The two latter species are rare, in-cluded in the Red Data Book of Ukraine (Didukh et al., 2009). Furthermore, C. onopordifolia is included in the IUCN Red List (Melnyk, 2011), and C. cirsioides is assigned to the European Red List of Globally Threatened Animals and Plants (European …, 1991). Despite its high phytosozological status, Carlina cirsioides is not sufficiently studied in the systematic, chorological and ecological-coenotic aspects.
Furthermore, the systematic position of C. cirsioides is a subject of discussions. Klokov (1954), who described this species, identified it to endemics and relics of the Flora of Ukraine. Because this species is included to the European Red List (European …, 1991), it is one of the most wellknown endemic of Ukrainian flora in the European Scientific Community. At the same time, neither the authors of Flora Europaea (Webb, 1976) nor the authors of the monograph on the Carlina genus (Meusel & Kästner, 1994) have even mentioned this species in their data. To solve this contradiction, a comparative study of morphology, chorology, habitats of C. cirsioides together with the closely related species C. acaulis was needed.
The objective of our investigation was determination of the systematic position, patterns of distribution, habitats and modern state of populations of C. cirsioides based on comparative-morphological, geographic and ecological-coenotic study.

Materials and methods
The study was based on the materials of field surveys performed in 2010-2020. Population of C. cirsioides was surveyed in the plains of Ukraine in Rivne, Zhytomyr, Lviv, Ivano-Frankivsk, Ternopil and Khmel-nytskyj regions. We studied herbarium samples of C. cirsioides kept in the herbariums of the M. G. Kholodny Institute of Botany of the National Academy of Sciences of Ukraine (KW), M. M. Gryshko National Botanical Garden (KWHA), Taras Shevchenko National University of Kyiv (KWU), Ivan Franko National University of Lviv (LW), National Museum of Natural History of the National Academy of Sciences of Ukraine (NAS) in Lviv (LWS), Institute of Ecology of Carpathians of the NAS of Ukraine (LWKS), and Rivne Regional Museum of Local Lore. Schematic range maps for the species were developed using the point method of cartographic image. For determination of ontogenetic spectrums of populations of C. cirsioides, we used diagnostic traits of the stages of the ontogenesis of the species described by Zelenchuk (1987). The account of density of the populations was made on 1 m 2 wide transects, within which 1х1 m 2 experimental plots were arranged.
Habitats and modern state of the populations. We provide phytocoenotic descriptions and modern stste of populations of C. cirsioides in the locations we surveyed.
Volhynian Polissia Habitat of C. cirsioides was found in 39th quarter of the Liubomyrsk Forestry near the Railway Station Liubomyrsk, which is allocated with forest cultures of Quercus robur L. The trees are around 50 years old, the average trunk diameter measuring 25 cm, and the average tree height equaling 18 m. The tree layer includes single trees of Carpinus betulus L., Populus tremula L., Pyrus communis L. The herb layer is thinned-out, and its projective cover equaled 50%. It was formed by Anthericum ramosum L., Betonica officinalis L., Campanula persicifolia L., Convallaria majalis L., Fragaria vesca L., Melampyrum nemorosum L., Maianthemum bifolium (L.) F. W. Schmidt, Digitalis grandiflora Mill. There we found only two single specimens of C. сirsioides.
Holohirka Tract is located north of Stinka village of Zolochiv district of Lviv region. The area of the site accounts for 150 m 2 . The soils are sodpodzolized. The vegetation is represented by formation of Elytrigieta intermedia -association between Elytrigia intermedia + Inula ensifolia.
Chortova Mountain is a Botanical Monument of National Significance, it is located between Pukiv and Babyntsi villages of Rohatyn district of Ivano-Frakivsk regiont. The area of the protected territory is 13 ha. The mountain is a butte with steep (20-30º) slopes, elevated 100-150 m above the surrounding surface. The soils are podzolized chornozems. The meadow-steppe vegetation is represented by formations of Brachypodieta pinnati and Cariceta humilis, which are represented by associations Brachypodium pinnatum + Carex humilis, Carex humilis + Anthericum ramosum. The projected area of the grass stand is 100%, that one of C. cirsioides measures from 15%. The herbaceous cover is also composed of Achillea millefolium L., Aconitum moldavicum Hacq., Adonis vernalis L., Anemone sylvestris L., Aster amellus L., Betonica officinalis L., Bromopsis inermis (Leuss.) Holub, Centaurea sanguineum L., Dactylis glomerata L., Elytrigia intermedia (Host) Nevski, Euphorbia cyparissias L., Kasova Mountain (Fig. 2) is located west of Bovshiv village of Halych district of Ivano-Frankivsk region. It is a steppe reserve within the Halych National Nature Park. The area of the reserve is 160 ha. The mountain stretches 4 km along the left slope of the Hnyla Lypa River and the Burshtyn Water Reserve. The mountain is 340 m above sea level and 100 m above the valley of the Hnyla Lypa River.
The population of C. cirsioides on the Kasova Mountain is large, comprising over 967 specimens, which occupy the area of about 1.5 ha in the central part of the mountain.
The Hora Hareva Tract of 9.6 ha area is located near Korostovychi village of Halych disrict of Ivano-Frankivsk region. Near the foothill, the herbaceous layer is represented by the formation of Brachypodieta pinnati with association between Brachypodium pinnatum + Festuca valesiaca. The projected area of herbaceous layer is 90%, and that of C. cirsioides < 1%. The dominants are Brachypodium pinnatum (L.) P. Beauv (50%) and Festuca valesiaca Gaudin (30%). The vegetation also includes Adonis vernalis L., Anthericum ramosum L., Asperula cynanchica L., Briza media L., Centaurea scabiosa L., Pulsatilla grandis Wender, Salvia verticillata L., Scabiosa ochroleuca L., Teucrium chamaedrys L. Because forbs, particularly grasses, have significantly overgrown the slopes of the mountain, rare species, particularly C. cirsioides, may soon vanish. The population of C. cirsioides in the Kamin Tract is small, with around 138 indiviuals counted. The plants grow singly and in small groups of 5-12 individuals/m 2 .
The Adamove Zaimysko Tract is located in the lands of Radoshiv village of the Council of Iziaslav district of Khemelnytska region, near Plishchyn village of Shepetivsky district of Khmelnytsk region. The area accounts for 7.5 ha. The soils are sod-podzolized, loamy-sandy.
The growing location is confined to the ecotone between oak-hornbeam forest and meadows in the valley of the ravine. At the edge of the forest, Betula verrucosa Ehrh grows, young undergrowth of which is massively invading the neighbouring meadows. There are separate trees of Quercus robur L., Cerasus avium (L.) Moench., Salix caprea L., and dense thickets of Crataegus monogyna Jacq., Prunus spinosa L., Rosa canina L., Rubus caesius L. and Chamaecytisus ruthenicus D.

Discussion
Carlina cirsioides Klok. was described by Klokov (1954) using a herbarium sample collected by А. H. Rаkochy in 1901 from the outskirts of Bilychi village near Kyiv (the sample is stored in the Komarov Botanical Institute of the Russian Academy of Sciences in Saint Petersburg).
According to the first description by Klokov (1954) and his description in "The Flora of USSR" (Klokov, 1962), C. cirsioides is monocarpic, biennial plant, endemic and relic plant of the Ukrainian flora. Its root system is represented by a strong central root which goes 30-40 cm deep. The stems are singular, around 5-55 cm (up to 1 m) long, usually welldeveloped, rarely shortened, erect, regular, spiny-wooly, thin felt-like until bloom, dirty-purple under the indumenta, with one large petiole.The leaves are arranged into basal rosette and are located along the stem, lightgreen, hard, petiolate, with brownish dirty purple, the petiole which is only slightly enlarged at the base and three times shorter than the lamina, with no spines on the margins, elongated-elliptic to elliptic on the outline, of around 11-45 cm long and 7-15 cm wide, notably woolly on both sides during bloom, quite densely woolly to sparsely woolly at a young age; there are 4-10 pairs of lateral segments ot leaf lobes, about 10-68 mm long and 6-50 mm wide, pinnate and partly separated, with elongatedlancete spiky lobes, serrated (together with the apical part) on the margins.
The capitulum is about 4-5 cm long (with simple inner leaves of the involucre) and about 9-13 cm in diameter with outward bent horizontal leaves, the capitulum disk (without leaves) is 4.0-7.5 cm in diameter; the upper leaves which support the capitulum partly resemble the basal and stem ones, up to 20 cm long and 9 cm wide, but mostly smaller, with broad petioles and feathery-ramified spines below the middle, transitional in shape and sizes to leaves of the involucre of the outer row; the latter are sessile, with ramified spines on the sides, slightly enlarged or narrowed on the apices, spine-sharpened, in general elongated on the outline, 14-50 mm long, 7-15 mm wide, mostly dark brown; their spines are divided into 2-8 parts; the middle leaves are ovoid or narrow lancet, 11-25 mm long and 2.5-5.0 mm wide (at the base), thinly sharpened, brownish on the outer side, notably woolly, the broader and shorter of these leaves above the middle have notable dense spines on the sides; the inner leaves are linear or narrow linear, 33-37 mm long and 2-5 mm wide; slightly broadened on the sides above the middle, glossy and yellowish in this region; brownish-dark purple below the middle on the outer side, almost the same on the inner side as above, sharpened at the apices; the coronas are 11-13 mm long, with with 1.2-1.7 mm long yellowish teeth. The achenes are oblong, 3.8-6.0 mm long, 1.5-2.0 mm wide, dark grey, densely hairy, with a circle of up to 2.5 mm long hairs at the apex, the papus is 14-17 mm long and its hairs are stuck together in bundles of 5-7.  Klokov (1954) summarizes the analysis of morphological traits of C. cirsioides as a new species by noting that "our species is notably different from Carlina acaulis L., except for distinct developed stems, other shape of leaves and more distinguished indumenta on them, and also much finer capitula".
Are there enough such small traits to identify a new species? -Obviously not. Therefore, it is not by chance that this taxon has not been identified as a separate in the monographs on Carlina (Meusel & Kästner, 1994). While not recognizing the species C. cirsioides, they identified specimens with distinct stems to subspecies C. acaulis ssp. сaulescens (Lam.) Schriber et Martens. Unlike the stemless or short-stemmed subspecies C. acaulis acaulis (Fig. 3), C. acaulis caulescens (Fig. 4.) is a plant composed of well developed flower-bearing shoots of 20-40 (up to 90) cm height, with no leaves or with pre-stem leaves. Unlike C. acaulis acaulis, distributed in the mountains of Europe (Pyrenees, Apennine Mountains, Balkans, Alps, the Carpathias), C. acaulis caulescens grows not only in mountains, but also in the lowlands of Europe. Therefore, in the Hercynian massifs of Europe, this subspecies is distributed in the Bourgogne Upland, the Rhenish Massif, Thuringian Forest (Meusel & Kästner, 1994).
The author of description of Carlina species in "Flora Europaea" (Webb, 1976) also distinguishes two subspecies of C. acaulis. C. acaulis caulescens in sense of the monographs on the genus Carlina corresponds to subspecies C. acaulis simplex (Waldst. et Kit) Nyman. with ramified or regular stems. He did not recognize the species independence of Сarlina cirsioides. Hegi (1987) distinguishes two subspecies of C. acaulis, and within subspecies C. acaulis acaulis, he distinguishes form var. caulescens Lam., which is distinct by developed stems of up to 30 cm long and the rosette of leaves located around the inflorescence. Another subspecies of this species, Carlina acaulis aygreta Waldst. et Kit (=C. acaulis simplex (Waldst. et Kit), has developed stems and no rosette around the inflorescence.
In the flora of Romania (Nyarady, 1964), three forms of C. acaulis are described: typical var. typical Beek -stemless or short-stemmed plant with a rosette of leaves; var. alpine Beek. l.c. (var. caulifera Bauneg.)with developed 10-40 (50) cm stems and rosette of leaves around the capitulum; var. aqgregata W. et K. -with developed stems and singular leaves on them. In Poland, C. acaulis is common in mountain regions (Carpathians, Sudetes), and also in Małopolska Upland. In other lowland territories (Wielkopolska, Kujawy, Masurian Lake district, Pomerania) it occurs rarely. At the same time the populations are represented mostly by subspecies C. acaulis caulescens (Wasilowska & Wolkowski, 2009).
The review we provided indicates that C. acaulis is a complex syngameon composed of mostly highland stemless or short-stemmed subspecies C. acaulis acaulis and mostly lowland subspecies C. acaulis caulescens. Morphological traits of the species C. cirsioides described by Klokov completely coincide with the traits of the latter subspecies. The analysis of herbarium samples of C. cirsioides kept in the Herbarium of the M. G. Kholodny Institute of Botany of the National Academy of Sciences of Ukraine showed that there are plants both having welldeveloped stems with rosettes of leaves around the inflorescences or without them, and stemless plants.
The folders under the name "Carlina acaulis" in the Herbarium of the M. G. Kholodny Institute of Botany of the National Academy of Sciences of Ukraine, along with stemless and short-stemmed plants also contain plants with well-developed stems which morphologically do not differ from the samples stored in the folders titled "C. cirsioides". The difference between them is in geographical distribution. The first were collected in the Carpathians, whereas the latter come from the plains part of Ukraine. Therefore, a logical question is why highland plants are identified to C. acaulis, while the lowland plants to C. cirsioides. It would be worth rejecting the species name C. cirsioides and use the priority name C. acaulis for plants from the entire territory of Ukraine. By the way, all the samples of the plants initially collected in the lowlands, starting from the collections of Montrezor (1898), were identified as C. acaulis caulescens, as documented on the labels. They were transferred to C. cirsioides after revision by Klokov (1954), as indicated in his notа critica. Because the plants from the plains territory of Ukraine occur only under the name C. cirsioides in all the editions of the Red Book of Ukraine (Didukh, 2009) andthe International Red Lists (European..., 1991), it is acceptable to use this name as C. acaulis synonym. Klokov (1954;1962) describing the new species C. cirsioides, identifies it to biennial monocarpic plants. Fol-lowing Klokov, this species was related to monocarpic by Shishkin & Bobrov (1962), Belousova et al. (1979) and other authors. Based on the observations on the plant in the conditions of nature and cultivation, Zelenchuk (1987) came to a conclusion that C. cirsioides is perennial polycarpic plant. Our multi-years monitoring of growth and development of this species in the natural and cultivation conditions confirms this point of view.
The data provided above do not allow the population of the lowland part of Ukraine to be identified as a separate species C. cirsioides. According to all morphological traits it is C. acaulis. Therefore, the statement of Klokov (1962) that C. cirsioides is a separate species, endemic and relic plant of the flora of Ukraine is erroneous. Neither has the relic nature of it been confirmed by phytogeographical data. There are no reliccharacteristic types of disjunct distribution in the range of this "species". The Ukrainian lowland populations adjoin directly to the Carpathian part of the range of C. acaulis and plains populations of this species in Poland. In general, C. acaulis is a Central European species, the range of which mostly includes mountain regions of Europe (Pyrenees, Alps, Apennine Mountains, Jura Mountains, Balcans, Carpathias and the neighbouring plains of the Hercynian massifs of Europe (Bourgogne Upland, the Rhenish Massif, Thuringian Forest), Poland (Pomerania, Masurian Lake district, Wielkopolska, Kujawy, Małopolska Upland) and Ukraine (Vohlynian-Podolian Upland and the Polesian Lowland).
In Ukraine C. acaulis is chiefly distributed in the Carpathians (Fig. 5), and plains (Fig. 1), this species grows near the north-east border of its range and is quite rare. Its range border runs along the edge of the Polesian Lowland near Derazhne villige of Kostopil district of Rivne region, Zhytomyr, Kyiv and Brovary, from where it makes a turn toward the south, passing near Berdychiv, Vinnytsia, Khmelnytsky, and Pidvolochysk.

Fig. 5. Geographical distribution of Carlina acaulis acaulis in Ukrainian Carpathians
According to the data of Klokov (1962), the new species of the Carlina genus which he described, grows in thinned-out forests, dry meadows and steppe slopes. Our detailed analysis of it habitat revealed that in the Podoilina Upland it is confined mostly to meadow steppes on the slopes of the buttes of south, west and east exposition, where it grows in calciumrich soils (rendzinas).
The species is a component of formations Therefore, in the plains of Ukraine, C. acaulis is mostly a component of steppe communities of Festuco-Brometea class, and much rarer the forest communities of Erico-Pinetea class and meadow-forest ecotones.
Mountain habitats of C. acaulis significantly differ from the plains ones. In the Carpathians, C. acaulis is a component of mountain meadows, confined to warm slopes covered with dense peat formations of matgrass Nardus stricta, wood small-reed Calamagrostis epigejos and bonfire grasses, reaching the altitude of 2,000 m (Malynovskyi, 1980). Simular habitats are in Alps, where it grows in subalpine meadow with domination of Bromus erectus Huds, Nardus stricta L., Festuca valesiaca Gaudin (Hegi, 1987).
In Sudets, C. acaulis grows in mountain pastures and is a component of of the plant community Molinio-Arrhenathereta class (Nadolna, 2013).
In the lowland regions of the Central Europe, the habitats of C. acaulis are very different from the mountain ones. There, the species is confined mostly to xerophilous herbaceous communities on calcium-rich soils. Such communities are reported for the Vosges, Schwarzwald, Bavarian Forest, Swabian Alb, Swabian-Bavarian Plateaau (Meusel & Kastner, 1994).
Comparison of ecological-coenotic conditions of habitats of C. acaulis revealed significant differences between plains and mountain ecotopes. In the Carpathians and Alps, this species is a component of mountain meadows, while in the plains regions of Europe it is confined mostly to meadow steppes in the east of the range and xerothermal herbaceous communities in the western part. These communities s are floristically close and confined to calcium-rich soils. Forest communities with C. acaulis in the Polesian Lowland are close to the adjacent ecotopes of the North-East Poland. Thus, ecological-coenotic habitat conditions of the plains part of Ukraine are not specific, and rather are typical for all the plains part of the range of C. acaulis.
In general, the data on morphology, chorology and phytocoenology do not allow us to consider C. cirsioides described by Klokov a separate species, endemic and relic plant of the Flora of Ukraine. Plains localities of this "species" in Ukraine may conditionally be considered geographic population of C. acaulis. Identification of this population to C. acaulis does not in any way reduce its phytosozoological value at all, the reason being is that xerothermal meadow communities involving C. acaulis are rare in plains throughout Europe and this species has been included in the lists of species which need protection in France and Poland (Boron et al., 2019;Strzemski et al., 2019).
Analysis of the abundance of the population of C. acaulis in the plain part of its range in Ukraine showed that near the eastern border of the range this species is extremely rare. A count of the number of specimens in populations of C. acaulis in the Podolian Upland (Table) revealed that they comprise 3 to 2,000 individuals. The population was the lowest on the Makitra Mountain (Lviv region) -only three specimens. In five of 14 surveyed populations, there are several dozens of specimens -25 to 90, or several hundreds of specimens -135 to 630 specimens (Table). And only in two local populations -on the Chortova Mountain in Ivano-Frankivsk region and the Zarvanytsia Mountain in Ternopil region, did the population accouns for around 1,000 specimens in each (Table). Therefore, the abundance of the populations is extremely low. Average density of populations equaled 3-5 specimens/m 2 , maximal -20 specimens/m 2 .
The data we obtained correlate with the materials of accounts of populations of this species conducted in the Burshtyn Opillia (Ivano-Frankivsk region) in 2009-2011 by Dmytrash, Shumska (2011). The number of specimens from nine populations they surveyed was 25 to 5,000 specimens. Outside the Podolian Upland, on the border of the Polesian Lowland, only singular individuals of C. acaulis were found.
Carlina cirsioides (=C. acaulis) within Vohlynia-Podillia was noted by high parameters of seed productivity. According to the data of Zelenchuk (1987), the actual seed productivity of this species accounted for 270 seeds per individual, the seed yield from area of 25 m 2 was 8,800 seeds, and the seed germination rate equaled 70-80%.
In the conditions of good seeding of the habitats, dense herbaceous cover with dominance of grasses is an obstruction to seeds, preventing them from entering the mineral part of soil. Therefore, in germinationfavourable conditions, only a small portion of seeds is introduced to the soil. The parameter of dying of seddlings is fairly high, related to deficiency of moisture in the soil during summer. A total of 42-50% seddlings die (Zelenchuk & Zelenchuk, 1987).
Moderate cattle grazing (animals break up the turf with their hooves) would obviously have helped the seeds to enter the mineral part of soil, where conditions for growth and development of sprouts are optimum. Thus, grazing cattle would provide the conditions of intensive replenishment of populations with new generations of individuals.
Significant damage to populations of C. acaulis near the Eastern border of the range is caused by preparation of medicinal raw materials from its herbs. This species is used in both official and folk medicine, especially in Germany and Poland (Link et al., 2014;Strzemski et al., 2019). In general, populations of C. acaulis near the eastern border of its range in Volhynia-Podillia are mainly homeostatic (Table), indicating that coenotypical conditions of meadow steppes which cover the slopes of the buttes meet the ecological needs of C. acaulis. Beyond the borders of the existing localities, there is not a sufficient amount of ecotopes favourable for the development of invasive populations, though the seeds there are easily distributed to large distances. In Vohlyni-Podilla, there was found one invasive population on the Makitra Mountain (Lviv region), consisting of only three specimens. As mentioned above, in the Polesian Lowland, only single individuals of this species occur.
Populations of C. cirsioides (=C. acaulis) have sufficient protection in the plains of Ukraine. They are protected in the Halytsky National Nature Park (Ivano-Frankivsk region), in Medobory Nature Reserve (Ternopil region), Botanical monuments of the National Importance: Lysa Hora Mountain and Sypuha Mountain (Lviv region), Chortova and Kasova Mountains (Ivano-Frankivsk region), Botanical Reserve of Local Significance Mohyla Mountain and the Kurylyha Mountain (Ternopil region), in the Adamove Zaimysko Tract (Khemelnytsky region).
Following our scientific recommendations, a botanical reserve Kurylyha Mountain was created in Ternopil region. Taking into account the high phytosozological value of C. acaulis near the eastern border of the range, in the future all populations of this species in Volhynia-Podillia and Polissia should be taken under protection.

Conclusions
Carlina cirsioides Klokov (Asteraceae), a well-known endemic plant of the Flora of Ukraine, is actually not an endemic of our Flora. Comparative-morphological and geographical analyses revealed that the populations of this "species" are identified to Carlina acaulis L. and have no significant differences from the populations in the plains regions of Central Europe which belong to subspecies C. acaulis сaulescens. Diagnostic traits according to which C. cirsioides was described are so insufficient that they provide no reasons for distinguishing it as a species different from C. acaulis.
Identification of populations from Volhynia-Podillia and Polissia to relic plants of the Flora of Ukraine is erroneous because they are not separate and directly adjoin to the Carpathian part of the range of C. acaulis and its populations in Poland.
Ecological-coenotic conditions of habitats of C. acaulis in the plains of Ukraine are different from the Carpathian ecotopes and close to plains regions of Central Europe. Meadow steppe formations of Brachypodieta pinnati, Bromopsideta inermis, Cariceta humilis, Festuceta valesiacae, including C. acaulis, in Vohlynia-Podillia, according to ecologicalcoenotic conditions and floristic composition are close to xerotherm herbaceous communities of Central Europe, and open pine and mixed forests are habitats of C. acaulis in Ukraine and North-East Poland.
Analysis of age structure of populations of C. acaulis in the plains of Ukraine revealed that most of them are homeostatic with right-sided spectra of ontogenetic states. A minimum was seen in juvenile and immature specimens, which is related to obstructions for seeds to enter the mineral soil due to the high level of development peat and significant rate of death of sprouts under the dense herbaceous layer. The share of invasive populations is small due to absence of favourable ecological-coenotic conditions outside the existing localities of C. acaulis.
Despite the removal of separate species status of the population in the plains of Ukraine due to the correction of their identification from endemic species C. cirsioides to the broadly-distributed European species C. acaulis, their phytosozoological importance remained unchanged.
C. acaulis is rare in the Ukrainian plains because of its narrow ecologic-coenotic amplitude and negative anthropogenic impact on habitats and populations (forestation of slopes, collection of plants as medicinal raw material and bouquets). To protect the gene fund of C. acaulis, all populations of C. acaulis near the eastern border of the range of the species should be taken under protection.