Image - The Nikita Botanical Garden near Yalta in the Crimea.

Flora [Флора]. The vegetation of Ukraine evolved through long geologic epochs and through many developmental phases before it attained its present form. In the Paleocene, and particularly in the Eocene, Ukraine had a tropical and subtropical flora. Palms (Sabal ucrainica, Nipa burtini sive ucrainica, and other species) cinnamon trees (Cinnamonum ucrainicum), figs, laurels, eucalyptus, yacca trees (Podocarpus), banksias, sequoias, and other trees grew in Ukraine. In the mid-Oligocene Mediterranean plants began to spread gradually to Ukraine, including oleander, pomegranate, beech, maple, and poplar. In the Miocene the vegetation generally assumed a temperate broad-leaved or broad-leaved-coniferous character, with a preponderance of deciduous forms such as beech, oak, and walnut. Alongside these species grew the tulip tree, swamp cypress, sequoia, and pine. The laurel continued to flourish in the Crimea. Gradually the flora changed to temperate warm-climate vegetation. In the Pliocene most of Ukraine was covered with forest vegetation. Among the evergreens were species of pine, with an admixture of hemlock, spruce, fir, swamp cypress, and others; and among deciduous trees were birch, oak, hornbeam, maple, chestnut, walnut, and magnolia. The southern region was covered with steppe grasses and pigweed. At the end of the Pliocene the forest vegetation became impoverished and approached the vegetation of the early Anthropogenic period (although a small number of swamp cypress did survive).

As the climate grew colder in the Pleistocene, pine-birch forests spread through northern Ukraine. The impoverished deciduous forests retreated into areas that were suitable to their development and free from glaciers (the right bank of the Donets River, the Dnister Lowland, the southern slopes of the Crimea, and Caucasia). During the warmer interglacial periods the broad-leaved species spread out to form extensive deciduous forests in the forest-steppe belt. In certain areas swamps and meadows were inhabited by tundra species—the midget birch, midget willow, and dryas. In the Holocene there was a migration of forest vegetation. Pine and pine-birch forests predominated in the early Holocene. In the middle Holocene broad-leaved species from central and southern Europe—linden, elm, hazelnut, oak—began to spread into Ukraine and by the late Holocene began to force out the hornbeam and beech. The steppe vegetation in Southern Ukraine was enriched with Xerophyte migrants from the east, from the Aral-Caspian floral center, Caucasia, and the Balkans. With the development of a favorable climate and the leaching away of salts from the loess subsoil, the deciduous forests advanced into the humid steppe of Ukraine. They attained their maximum area about 5,000 years ago. Forests were always more widespread in Right-Bank Ukraine, particularly in regions of dissected relief, which hindered the development of agricultural plants. In Left-Bank Ukraine forests spread mainly along the banks of the Sula River, the Psol River, the Vorskla River, and the Donets River and in the higher regions of the Donets Ridge.

Human agricultural activity greatly altered the original vegetation of Ukraine. Almost all the steppe is under cultivation and devoted to agricultural species. Large areas of the forest belt have also been converted to agricultural use. Intense long-term logging in the Carpathian Mountains has diminished the protective influence of the forests, as frequent floods and increasing erosion have shown. Between 1814 and 1914 the forest area in Ukraine diminished by 30.5 percent. Only a few ancient tracts remain untouched by humans, and for their preservation they have been placed under state protection.

The present flora of Ukraine encompasses quite a large number of relicts from the Tertiary period and the Ice Age. It includes also migratory species from the north (coniferous forests, swamp vegetation), the west (deciduous trees), the south, the Crimea, and Caucasia (some deciduous trees and steppe plants), and the southwest and southeast (steppe vegetation, especially the plants of the saline soils). There are also a number of endemic forms in Ukraine, particularly on sands and rocks. The boundaries of the habitats of many plants and trees run through Ukraine: the northeastern boundary of the beech (Fagus silvatica) and silver fir (Abies alba); the eastern boundary of the sycamore maple (Acer pseudoplatanus), field maple (Acer campestre), linden (Tilia platyphyllos), and ash (Fraxinus excelsior); the southern boundary of the Scots pine (Pinus sylvestris); and the western boundary of the steppe cherry (Cerasus fruticosa).

Among the over 16,000 species of plants that are now found in Ukraine there are over 4,400 species of higher plants, 800 species of bryophytes, 1,000 species of lichen, close to 7,000 species of fungi and molds, over 3,000 species of continental algae, and 600 species of marine algae. Many foreign plants (various decorative plants and useful cultivated grasses) have also been introduced.

The close relation between vegetation and climate, relief, humidity, and soil is apparent in the organization of plant life in Ukraine into wide meridional belts stretching from the southwest to the northeast. Moving from north to south the belts are forest, forest-steppe, steppe, and Mediterranean. In the mountain regions of the Carpathian Mountains, Crimean Mountains, and Caucasia the vegetation is distributed in altitudinal belts.

The forest belt. The forest belt extends through northern and western Ukraine. Its southern border with the forest-steppe runs along the line Lviv-Kremenets-Zhytomyr-Kyiv-Nizhyn-Hlukhiv. This line also separates the podzolic soils of Polisia and the clayey chornozems of the forest-steppe. Within this belt lies a large island of forest-steppe on the Volhynia-Kholm Upland, circumscribed roughly by the line Kholm-Lutsk-Rivne-Mezhyrich-Kryvyn-Ostroh-Stoianiv-Belz. The forest belt can be subdivided into the western part, or the belt of Central European deciduous forests, and the northern belt of mixed forests in Polisia.

Central European deciduous forests cover the western part of Podilia, Roztochia, the Sian Lowland, Subcarpathia, and the Tysa Lowland. Here the forests contain a greater variety of trees than anywhere else in Ukraine. The beech is most characteristic of Western Ukraine, where the eastern boundary of its habitat is located. The beech grows in large numbers in Transcarpathia and the Roztochia region. The silver fir (Abies alba) is found in this belt also. It is widespread in the Carpathian Mountains and some areas of Subcarpathia and Roztochia (usually on the northern slopes) and rarer in the lowlands. The oak (Quercus sessiliflora, Q. pedunculata), linden (Tilia cordata), elm (Ulmus campestris), birch (Betula verrucosa), sycamore maple (Acer pseudoplatanus), Norway maple (A. platanoides), pine (Pinus silvestris), and spruce (Picea excelsa) are common here. The larch (Larix decidua) and English yew (Taxus baccata) (found in the Kniazh-Dvir reserve near Kolomyia) are rarer.

The forests of Polisia belong to the belt of Eastern European mixed forests. In the past Polisia formed a continuous forest-swamp landscape, but the wanton destruction of the forests diminished their area (eg, in 1861–1914 the area that forests occupied in Volhynia gubernia shrank from 41.9 percent to 25.4 percent). Today forests cover about 30 percent of Polisia. The distribution of vegetation depends on the type of soil and the relief. Many northern species are found here, particularly on the border with Belarus. Certain steppe forms have also found their way here (feather grass, adonis). Certain varieties of glacial flora (Lapland willow [Salix lapponum], etc) can also be found. The natural vegetation grows in forests, meadows, and swamps. The main forest trees are the pine (57.4 percent of the forest area), oak (21 percent), birch (10 percent), black alder (6 percent), European aspen (2 percent), and hornbeam (2 percent). Pine forests (bory) are widespread. They grow in weakly and moderately podzolized soils overlying deep sands. In places birches appear among the pines, and there is no undergrowth. These forests are indifferent to relief, humidity, and soil. Large tracts are covered with oak-pine forests (subory), which grow in sandy, slightly podzolized soils with a rich cover of bracken fern (Pteridium aquilinum). Depending on the depth of undersoil water, these forests are characterized as dry, fresh, moist, or wet. In the Zhytomyr area the yellow rhododendron is part of the undergrowth. Hornbeam-oak-pine woods (suhrudky) flourish on podzolized sandy soils underlain by clay. The herbaceous covering is diverse and contains bracken, blackberry (Rubus saxatilis), huckleberry, and so on. In eastern Polisia linden-oak-pine woods prevail. In the northern part of the Chernihiv region spruce forests abound, while numerous birch groves grow in the southern part. Oak-hornbeam woods (hrudy) are characteristic of podzolized clayey soils and have an abundant undergrowth. Oak-maple-linden woods are found in Left Bank Ukraine. Alder groves appear on peaty bog-gley soils, sometimes with an admixture of birch. Ash-alder and oak-hornbeam-alder woods are rarer. Bory and subory turn into swampy forests where the undersoil water rises close to the surface. Pine with an admixture of pilose birch is dominant in the less swampy forests, while a thick covering of sedges, reed grass, and cotton grass is found in the more swampy forests. Silver grass associations, sometimes richly colored by thyme, are dominant on poorly fixed sands. Areas that are somewhat lower are completely overgrown with heather.

The prevailing forests are broken up by dry meadows, which have developed where the trees have been cut down. The vegetation in the meadows is diverse: mat-grass, sedges, tufted hair-grass. Flood meadows along the Polisian river valleys are rich in variety and yield large harvests of high-quality hay.

Bogs are most characteristic of Polisia. Most of them are eutrophic bogs located in low-lying areas and rich in mineral salts. They are covered with sedges and sedge-hypnum communities. Many bogs are mesotrophic (transitional) and are located on watersheds and sandy terraces. Oligotrophic bogs, which are elevated and poor in mineral salts, are common in western Polisia. Their vegetation is sparse. Low swamp pines grow in clusters.

Little Polisia is a unique part of Polisia. It is dissected by slow-flowing streams with swampy valleys and is dotted with bory and subory. The region is bounded by the Volhynia-Kholm Upland in the north, the Podolian Upland in the southeast, and Roztochia in the west. In the east it merges with Polisia through the Ostroh-Slavuta Lowland. Pine and deciduous-pine woods, without beech, are abundant in Little Polisia. The meadows are covered with sedges and various grasses. The swamps are often quite deep and rich in turf deposits.

The forest-steppe belt. The forest-steppe, which extends south of Polisia and east of the deciduous forests of the Central European variety in Western Ukraine (Soviet scholars subsume this territory under the forest-steppe), is the Middle Dnipro subprovince of the East European forest-steppe province. In the south the forest-steppe merges with the steppe belt. The boundary between them is indistinct (many forest islands located in its vicinity have been cut down) and is defined in various ways, usually as the line Chişinau-Kropyvnytskyi-Kremenchuk-Krasnohrad-Zmiiv-Valuiky. The soils of the forest-steppe are deep chornozems, parts of which have changed under the influence of forests to degraded chornozems and gray forest podzolized soils. Some forest-steppe islands are found in the forest belt (in the Volhynia-Kholm Upland) and in the steppe belt (in the Donets Ridge). A narrow forest-steppe belt appears in the mountain foothills and is connected with the altitudinal zonality of mountain vegetation.

In the forest-steppe the elevated, more-dissected right banks of the rivers, watersheds, hilly areas, ravines, and gullies are covered with forests. The oak is the predominant tree of the forest-steppe, covering almost half of its surface, but only a few large oak groves remain—the Chornyi Lis Forest in Kirovohrad oblast and several groves in Khmelnytskyi oblast. West of the Dnipro River hornbeam trees grow among the oaks. Ash, elm, linden, maple, and beech (the last only in Podilia) are also found in the forest-steppe. The underbrush consists mainly of hazel (Corylus avellana), small-leaved field maple (Acer campestre), and spindletree (Euonymus verrucosus). Under these are found certain ferns and many types of flowering herbs. There are also hornbeam woods with very limited vegetation. As one moves east through the forest-steppe, western species become rarer: the beech does not grow beyond western Podilia, and hornbeam is rare in the forest-steppe of the Left Bank, where oak-maple-linden woods are widespread. The river valleys contain mostly oak, ash, elm, black poplar, and willow. Pine woods, mixed woods, and hypnum-sedge and sedge-sphagnum swamps are common on the sandy river terraces, particularly on the left bank of the Dnipro River. In the ravines and gullies the so-called ravine (bairak) forests are found. They consist of oak and hornbeam, but also of ash, maple, and linden and sometimes birch and European aspen. The underbrush consists of wild guelder rose (snowball; Viburnum opulus), wayfaring tree (V. lantana), European dogwood (Cornus sanguinea), and hazel. The deep ravines of Podilia have a peculiar vegetation.

The steppe in the forest-steppe belt is now almost entirely under cultivation. At one time it was a grassy, colored, broad-leaved steppe. Its well-watered soil supports the following meadow-steppe vegetation: low sedge (Carex humilis), sheep fescue grass (Festuca sulcata), narrow-leaved bent grass (Agrostis tenuifolia), feather grasses (Stipa capillata and Stipa joannis stenophylla), representatives of the colored northern herbs such as yellow bedstraw (Galium verum), meadow sage (Salvia pratensis), pedicularis (Pedicularis comosa), and steppe varieties of clover. The thickets consist of blackthorn (Prunus spinosa), steppe cherry (Erasus fruticosa), wild rose (Rosa spp.), and Ruthenian broom (Cytisus ruthenicus). In the southern part cut-leaved meadow sweet (Spiraea crenifolia) occurs as well. The left-bank terraces of the Dnipro provide a habitat for holophytes (salt plants). Low swamps are frequent in the floodplains along the left-bank tributaries of the Dnipro River.

In the forest-steppe of the Volhynia-Kholm Upland the forests are concentrated mainly along the southern and northern edges. The belt in the middle of the upland is almost forestless. Pine-oak and hornbeam groves are prevalent. Sometimes oak-pine subory occur, frequently containing hornbeam. The meadows along river plains have survived.

The forest-steppe of the Tysa Lowland in Transcarpathia consists of oak forests, flood meadows, and steppe plant varieties, which migrated there from the Danube River valley. Natural vegetation in the form of small oak and hornbeam groves has been preserved in this cultivated lowland only here and there on steep slopes.

The Donets Ridge, which by its nature belongs to the forest-steppe, is covered with various fescue-grass and feather-grass steppe vegetation and an admixture of ravine and watered-valley forests of oak, linden, and ash. Hornbeam and a number of grassy plants typical of deciduous forests grow in isolated clusters here.

The forest-steppe in the Crimea extends to the north and east of the Crimean Mountains. It consists of alternating strips of steppe and forest. The forests contain various oaks—pubescent, rock, and common (Quercus pubescens, Q. petraea, Q. robur)—and small-leaved field maple (Acer campestre), pear, and other species. The region abounds in thickets (European dogwood, hawthorn, blackthorn, buckthorn).

In the Subcaucasian forest-steppe, meadow-steppe fields are common along the line Kuban RiverKrasnodar–mouth of the Laba–Piatigorsk–Groznyi. They are dotted by islands of woods containing oak (Quercus pedunculata, Q. sessiliflora), hornbeam, ash, elm (Ulmus campestris), maple, pear, apple, and other species. The undergrowth consists of hazel, hawthorn, yellow rhododendron, and so on. The forests of the Stavropol Upland also belong to the forest-steppe. Of the feather grasses one finds Stipa joannis and Stipa pulcherima. On the periphery oak-ash forests with some hornbeam trees appear. The birch tree is rare.

The steppe belt. This belt extends southward from the forest-steppe belt to the foothills of the Crimean Mountains and the Caucasus. It is part of the Black SeaSea of Azov subprovince of the Pontic-Pannonian Steppe province. The steppe belt can be subdivided into several smaller belts according to climatic and soil conditions.

The northern part of the steppe is a more humid fescue-feather-grass colored steppe or grassy-meadow steppe. It is characterized by an abundance of thick, rough grasses, such as fescue (Festuca sulcata), koeleria (Koeleria gracilis), feather grasses (Stipa lessingiana, S. capillata, and on slopes S. stenophylla), and dicotyledons, such as the fernleaf peony (Paeonia tenuifolia), pheasant's eye (Adonis volgensis), and knapweed (Centaurea trinervia). There are many broad-leaved grasses with creeping rhizomes: bromegrass (Bromus erectus riparius), bluegrass (Poa pratensis), couch grass (Agropyron glaucum). Various tumbleweeds—sea cabbage (Crambe tataria), sea pink (Statice latifolia), Cossack tumbleweed (Phlomis pungens)—flourish in the steppe. Mosses such as Tortula ruralis grow in the gaps of the sod cover. Steppe copses contain blackthorn (Prunus steposa), caragana (Caragana frutex), broom (Cytisus), and almond (Amygdalus nana). Oak groves are found on the slopes of ravines. The original vegetation of the steppe is preserved on the following reserves: the Ukrainian Steppe Nature Reserve (including Khomutivskyi Steppe and Kamiani Mohyly Nature Reserve), and Striltsivskyi Steppe.

Farther south, down to the Black Sea and the Sea of Azov, lies the dry grassy steppe or the narrow-leaved, fescue-feather-grass steppe. It grows on southern chornozems and dark chestnut soils. Among the grasses, fescue and feather grasses (Stipa capillata, S. lessingiana, and S. ucrainica) predominate. Grass varieties are few and consist of xeric species: pheasant's-eye, ferula (Ferula caspica, F. orientalis), Limonium sareptanum, and others.

Many ephemeral annuals and perennials flower in the spring: jagged chickweed (Holosteum umbellatum), whitlow grass (Erophila verna), medwort alisson (Alyssum desertorum), androsace (Androsace elongata), forget-me-not (Myosotis micrantha), and tulip (Tulipa schrenkii, T. biebersteiniana). The original vegetation of the steppe is protected on the Askaniia-Nova Biosphere Reserve.

Meadow vegetation occurs in the river floodplains, and includes: meadow foxtail (Alopecurus pratensis), quack grass (Agropyron repens), epigean reedgrass (Calamagrostis epigeios), and sedge (Carex schreberi), and such dicotyledons as the buttercup (Ranunculus acer), bird's-foot trefoil (Lotus corniculatus), and clover (Trifolium). The humid, low-lying meadows produce various sedges and grasses (Beckmania eruciformis, Poa palustris). Small woods of oak, elm, black poplar, and alder grow in the valleys of the larger rivers. Swamp vegetation can also be found in the river floodplains and on the shores of limans and lakes, which are periodically flooded; such vegetation includes the reed (Phragmites communis), broad-leaved cattail (Typha latifolia), the great bullrush (Scirpus lacustris), and sweet flag (Acorus calamus). Floodplains cover large areas along the great rivers of Ukraine. The best-known among them is the Velykyi Luh of the Dnipro River with its ravine woods, terrace groves, reeds, and cattails. It once played a significant role in Cossack history, but is now submerged under the Kakhovka Reservoir.

Depressions known as pody, which are common on the watersheds between the Boh River and the Dnipro River and between the Dnipro River and the Molochna River, contain a meadow and meadow-halophytic vegetation. The sandy steppe from Nova Kakhovka to the Black Sea supports typical psammophile (sand-loving) vegetation (Festuca beckeri, etc).

In the south, in the Syvash Lake region and along the coast of the Black Sea, there is a strip of salinized chestnut soils, which supports a grassy sagebrush steppe. This is a transitional form between steppe and desert vegetation. Here we find fescue grasses, sometimes feather grass Stipa capillata, and an abundance of Crimean and Austrian wormwood (Artemisia taurica, A. austriaca). On the seacoast halophilic plants grow in solonets soils. Jointed glasswort (Salicornia herbacea), sea-blite (Suaeda), and sea pink (Statice suffruticosa) are dominant. To the south of the steppe the southern chornozem plains of the Crimea were once covered with fescue–feather-grass steppes. The ordinary chornozems farther south supported steppes of various grasses, fescues, and feather grasses. Now these lands are cultivated, but some remnants of the steppe continue to flourish on the slopes of ravines. The Tarkhankut Peninsula and the Kerch Peninsula with their rocky and gravelly soils are covered with a shrub steppe consisting of thyme, germander (Teucrium), Crimean skullcap (Scutellaria taurica), cliff lucerne (Medicago rupestris), various feather grasses, fescue grasses, and others.

Mediterranean vegetation. A narrow zone of Mediterranean vegetation extends along the Crimean southern shore and the Caucasian coast of the Black Sea. The typical forests in this zone consist of pubescent oak, tall juniper, Crimean pine (Pinus pallasiana), and Stankevich pine (P. Stankevichi). The undergrowth consists of arbutus (Arbutus andrachne), Ruscus ponticus, and other species. Today the Mediterranean vegetation is different from what it was: wild plants have been displaced by vineyards, orchards, and decorative gardens.

Mountain vegetation. The Carpathian Mountains are a subprovince of the Central European deciduous province. They contain a large number of West European varieties (mainly beech, Carpathian hornbeam, common and rock oak, linden, and maple), a number of endemic species, and some representatives of north Balkan, Mediterranean, and Euro-Siberian taiga and alpine vegetation. The diverse climatic zones in the mountains have reproduced an altitudinal distribution of plants in several belts. The foothills and the lower slopes up to 500–600 m above sea level constitute the mixed-forest belt consisting primarily of oak (Quercus robur, Q. sessiliflora), as well as pine, fir, hornbeam, beech, sharp-leaved maple, and large-leaved linden. At higher elevations the proportion of beech and coniferous species increases. At 600–1,200 m lies the belt of beech and mixed beech-fir and spruce forests. In Subcarpathia this forest belt faces north and contains mostly coniferous trees, but in Transcarpathia the belt is occupied by large beech forests. Spruce (Picea excelsa) forests are dominant only at higher elevations. The upper limit of the forests reaches 1,500–1,600 m. Above this line lie the subalpine and alpine belts, with thick scrub of dwarf pine (Pinus mughus), Siberian juniper, mountain alder (Alnus viridis), and Carpathian rhododendron. Extensive mountain meadows (polonyny) and clearings at this elevation are used for pastures in summer. The alpine belt (above 1,800–1,850 m) contains arnica (Arnica moutana), gentians, various orchids, etc; the ground is covered with white mosses and matted lichens at higher elevations.

The Crimean Mountains, which are a subprovince of the Euxine province of the Mediterranean forest region, are remarkable for their developed forests and rich vegetation. In the Tertiary period they belonged to the east Mediterranean landmass, on which the ancient Mediterranean vegetation established itself. Northern varieties are fewer here. The mountains are covered with forests of oak, beech, hornbeam, and other trees. Near the upper boundary of the beech forests there are small tracts of pine. The mountains are topped with pastures, called yaily, with short, thick grasses.

The western and southern foothills of the Caucasus Mountains are covered with thick, Mediterranean evergreen vegetation. Above the juniper and juniper-oak belt (with some Caucasian beech [Fagus orientalis]) the relatively low coastal mountain range is forestless and covered with mountain-steppe or mountain-meadow vegetation. Above this vegetation belt is a belt of fescues and feather grasses. Strong winds are responsible for drastically lowering the boundary of forest vegetation. Farther south, at elevations up to 2,000 m, there are deciduous forests of oak, beech, hornbeam, maple, and other trees and mixed forests containing the Caucasian fir (Abies nordmanniana) and spruce (Picea orientalis). Above 2,000 m lies a subalpine belt, with high mountain meadows, and an alpine belt. Beyond 3,000 m the bare cliffs are covered with permanent snow.

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Hryhorii Makhiv, Edvard Zharsky

[This article originally appeared in the Encyclopedia of Ukraine, vol. 1 (1984).]

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