Author: Mária Höhn
a . deals with the geographical distribution of plant species around the World and the principles that govern species’ occurrence in space.
„Where do species live?”
- origin and area of distribution:
Autochtonous species: A species is autochtonous (i.e., native) in the biogeographical region of the Carpathian basin if it has been naturally present starting with the last big historical climate- and vegetation changes in the Holocene, and its occurrence is demonstrably not the consequence of human activity.
Area of distribution: the geographical distribution range of a species. Each species can be unequivocally identified with its area.
b. analyses the species composition of a geographical region,
„Which species occur within a certain geographical region?”
- floristics → floristic units, floristic regions
The central object of study in Phytogeography is the species.
The species concept defined formerly in the Introduction may be extended within the framework of Phytogeography. The species is a group of individuals which share many similar external and internal features, originate from one common ancestor and can actually or potentially interbreed and produce fertile offsprings. Each species has a well delimited range of distribution, i.e., each species has its own area.
Further important terms used in Phytogeography are: geoelements, flora, floristic kingdoms (realms)
The geographical distribution range of a species, which has boundaries and can be mapped.
Taxon area : could mean the area of any higher taxonomical unit, thus families and genera may have their own area of distribution (e.g., the Betulaceae and Orchidaceae families or the Bambusa genus have well-defined geographical ranges).
- Major types of the areas of distribution:
1. continuous (connected)
e.g. Fraxinus excelsior (common ash) (Figure 1), Crataegus monogyna (common hawthorn)
Fig. 1. The continuous area of Fraxinus excelsior in Europe
2. discontinuous (not connected):
2a. the disjunct (scattered) areas contain small portions of the geographical range widely separated from the main area
e.c. Pinus sylvestris (Scots pine),
- 2b. the disperse area consists of small fragments more or less isolated from each other. Relict species may often show such distribution.
e.c. Pinus nigra (European black pine)
Fig. 2. The disperse area of Pinus nigra
The area of any species is strongly connected with the evolutionary processes that lead to (i.e., speciation and with the species’ natural history.
The process of speciation takes a long time (millions of years), and it is the consequence of several, usually nonlinear collective events.
The most important events that may launch speciation are:
Isolation can be: geographical, ecological (when individuals live in different environmental conditions) and sexual (reproductive).
These processes may result in the formation of a new, endemic species with limited geographic distribution.
e.g. Dianthus serotinus (Figure 3) and Seseli leucospermum are Pannonian endemic species with a small area, limited to the Carpathian basin.
Fig. 3. Dianthus serotinus
Other speciation events:
Stabilization makes it possible for some species to become widespread.
e.g. Eurasian species like Salix alba (white willow), or Cosmopolitan species like Phragmites australis (common reed).
7. shrinkage of the geographic range, followed by isolation: it is usually triggered by fundamental changes in macroclimate and/or paleogeographical conditions; species’ geographic range may shrink, fragment or shift, during which ecological conditions across the range may change.
→ This may give rise to relict species, only able to survive in small, restricted areas where environmental conditions typical to the former macroclimate can persist longer.
Relict species of the last Ice Age from the Carpathian basin are e.g Comarum palustre (Potentilla palustris), from the Carpathians e.g., Pinus cembra.
A species surviving in a small area for a long enough time will likely go through new speciation events. New traits will be acquired in addition to the existing ones (which can also disappear), giving rise to relict endemic species.
e.g. Sesleria sadleriana, Seseli leucospermum in the Carpathian basin
Fig. 4. Sesleria sadleriana on the dolomite hills of the Mt. Sas-hegy, Budapest (Photo: Antal Halász)
The formation of new species is a long and complex evolutionary process. There are several ways in which new species can evolve:
- Allopatric speciation: The main driving factor is fragmentation or split of a former distribution area followed by isolation of the group of individuals from the fragmented territories. Generally in consequence of a geographic barrier that arises, e.g. mountain building, the isolated groups of individuals will evolve under different selective pressure and new mutations will appear and later they might be selected.
- Peripatric speciation: A limited number of individuals situated at the edge of the species area of distribution, may often encounter extreme environmental conditions which favour the start of speciation processes. Individuals are selected against these conditions and are prevented from exchanging genes with the main population. As a result, new genetic, and morphological traits will start to fixate due to isolation from the rest of the individuals.
This kind of speciation is often the consequence of the so called founder effect. when a very small number of individuals will establish in a new territory, usually far from the main population and this cause a big loss in genetic variability.
- Parapatric speciation: The process is similar to the peripatric speciation, but there is no obvious geographic barrier to gene flow. Groups of individuals may occupy a new habitat (e.g. under increasing demographic pressure), and will more likely mate with their neighbors of the same habitat. As a result, the successful types and their offspring will be selected in the new habitat.
- Sympatric speciation: Is triggered when ecological or reproductive isolation occurs between two spatially close populations or individuals that share the same area of distribution (e.g. changes in flower size and/or morphology as a result of adaption to different pollinators). It is a rare event because frequent backcrossing swipes out the emergent differences.
Fig: 5. Forms of speciation
(Source: http://evolution.berkeley.edu/evosite/evo101/VC1aModesSpeciation.shtml)
Vicariance: Speciation events can produce vicariance. (vicarius <latin> means supply, surrogate, substitute).
Vicariant taxa are closely related (may be species or families), and exist in separate geographical areas or under different ecological conditions. They are thought to have originated from a common population that was divided by geographical or ecological barriers.
a. Geographical vicariants are closely related species found in similar habitats, but different geographical regions. E.g. the species of the Helleborus genus or of the Cedrus genus.
b. Ecological vicariants are closely related species that have different ecological requirements, but may share the same area of distribution.
The Rhododendron species of the European alpine region are ecological vicariants because Rh. hirsutum grows on calcareous substrate and Rh. ferrugineum prefers silicious substrate. The same is true for two Asplenium species: the wall-rue (A. ruta muraria) grows on limestone, whereas the northern spleenwort (A. septentrionale) inhabits volcanic basalt and andesite cliffs.
Fig. 6. Distribution map of the geographical vicariant Cedrus species
(Source: http://www.sciencedirect.com/science/article/pii/S037811270200539X )
Pseudovicariance :
- Species or genera that belong to distant taxonomical groups (i.e., distant relatives) substitute each other in similar habitats within different geographic regions
e.g. Euphorbiaceae (grow only in African deserts) – Cactaceae (grow only in American deserts)
Figure 7. Pseudovicariance Euphorbia and Opuntia (Pócs T.)
- groups of species that have similar geographic distribution (occur in the the same area)
Within the Carpathian basin, the most common geoelements are:
Eurasian elements (eua): species commonly found in the temperate zone of Europe and Asia, e.g. Betula pendula, Pinus sylvestris, Ranunculus acris, Salix alba; their proportion among native species of the Carpathian Basin is ~22,5%.
Figure 8: Distribution map of Betula pendula.
European elements (eur): species widespread in Europe (up to the Ural Mountains and includingAsia Minor and the Middle East); their proportion among native species of the Carpathian Basin is ~14,4%; e.g. Corylus avellana, Quercus robur, Picea abies;
Central European elements (ceu): e.g. Acer platanoides, Fagus sylvatica; several forest- and grassland species;
Figure 9: Distribution map of Fagus sylvatica.
The proportion of the European and Central European elements among native species of the Carpathian Basin is higher than 20%
Figure 10: Distribution map of Adonis vernalis.
Continental elements (con): species widespread in the continental steppe and wooded steppe interior regions of Eurasia; their proportion among native species of the Carpathian Basin is ~11% e.g.. Carex humilis, Prunus tenella, Adonis vernalis, Acer tataricum
Fig. 11. A: Prunus tenella, B: Acer tataricum
Pontic-Pannonian elements (pop): common species of the Southern Russian (Ukrainian) lowland (north of the Black Sea) and the Pannonian Plain
e.g. Linum flavum, Crambe tataria
Pontic –Mediterranean elements (pom): species widespread in the Black Sea region and the Crimean Peninsula towards the Caucasus
e.g. Cotinus coggygria, Prunus mahaleb
Fig. 12. Cotinus coggygria (A), Prunus mahaleb (B)
Turanian elements (tur): semi-desert species of the Caspian Sea and Aral Lake regions, mainly in alkali flat inlands
e.g. Lepidium crassifolium, Kochia prostrata, Eurotia ceratiodes
Submediterranean elements (sme): distributed in the deciduous forest zone of the Iberian Peninsula, Southern France, Southern Alps, the Apennine and the Northern part of the Balkan Peninsula, excluding the real evergreen Mediterranean zone; their proportion among inland species is ~18%
e.g. Castanea sativa, Cornus mas, Quercus pubescens
Fig. 13. A: Downy oak (Quercus pubescens) with a Scarce swallowtail butterfly ( Iphiclides podalirius) ; B: Downy oak habit
Central-European Mediterranean elements (cem): similar to the former, but also distributed in the Southern region of Central Europe
e.g. Fumana procumbens, Colutea arborescens
Atlantic Mediterranean elements (atm): species found in the moderate climate stretching between the Mediterranean Sea and Western Europe
e.g. Tamus communis, Ruscus aculeatus, Hedera helix, Ilex aquifolium
Fig. 14. Tamus communis (A), Ilex aquifolium (B)
Balkanian elements (bal): characteristic species of the Balkan Peninsula. Difference exist between Western Balkanian (or Illyr) and Eastern Balkanian (or Moezian) species; their proportion among native species of the Carpathian Basin is more than 4%
e.g. Tilia argentea, Ranunculus illyricus, Syringa vulgaris
Balkan- Caucasian element (BaC): characteristic species of Southeastern Europe, from the Balkan peninsula to the Caucasus
e.g. Carpinus orientalis,
Fig. 15. Carpinus orientalis – supposed to be a relict species of warmer period
Boreal (northern) elements (bor): characteristic species of the coniferous forests and swamps of the northern taiga region; most of these species are relicts of the last Ice Age in our country.
e.g. Comarum palustre (Potentilla palustris), Vaccinium oxycoccus,
Fig. 16. Vaccinium oxycoccus
Alpine elements (alp): plants of the Alps, and the Carpathians, and of the northern taiga forests and tundra e.g. Alnus viridis, Arabis alpina.
Circumpolar elements (cir): species which are widespread in the whole Northern temperate region around the Globe (am-bor+eua); their proportion among the Carpathian Basin flora is ~ 6, 8%
e.g. Rubus idaeus, Juniperus communis, Vaccinium myrtillus, Viburnum opulus
Fig. 17. Vaccinium myrtillus
Endemic elements (end):
a. Pannonian endemic species: occur exclusively in Hungary and the border-lands (i.e., in the Pannonian region); their proportion in the Hungarian flora is ~2%
e.g. Thlaspi jankae, Dianthus serotinus, Centaurea sadleriana, Linum dolomiticum
Fig. 18. Linum dolomiticum (Photo credit: László Udvardy)
b. Carpathian elements (crp): endemic plants of the Carpathian Mountains
e.g. Dentaria glandulosa, Crocus heuffelianus
Fig. 19. Crocus heuffelianus
Cosmopolitan elements (cos): species widespread on all continents. These are e.g., certain ancient ferns (Pteridium aquilinum, Equisetum arvense), aquatic plants inhabiting freshwaters around the world (Lemna minor, Phragmites australis) and species that spread due to human activity (e.g.Galinsoga parviflora). Their proportion among the native species of the Carpathian basin is around 6,5%.
Adventive elements (adv): Alien (i.e., non-native) species introduced to Europe after the last big climatic changes (800 B.C.), which are able to spread spontaneously in natural ecosystems. These species were introduced intentionally or accidentally due to human activity.
Most of the adventive species are weeds. Their proportion among species of the Carpathian basin exceeds 12%, but this number increases continuously as a result of increasing human impact and of the climate change.
Types of adventives elements:
a. Archaeophytes: Ancient weeds introduced to the Carpathian Basin along with early agricultural crops. Their introduction precedes the European discovery of America (the XVIth century).
e.g. ancient weeds of grain crops, such as Papaver rhoeas, Centaurea cyanus, Agrostemma githago
b. Neophytes: Species introduced after the European discovery of America, starting with the XVIth century
North American species such as Robinia pseudoacacia, Acer negundo, Amorpha fruticosa, Solidago canadensis were mostly brought in intentionally by humans, while others such as Ambrosia artemisiifolia and Amaranthus retroflexus were introduced accidentally.
Of the Neophytes introduced to Europe from Eastern Asia, the most common is the Heaven tree (Ailanthus altissima). Heracleum mantegazzianum and H. sosnowski originate from the Caucasus.
Fig. 20. The Giant hogweed (Heracleum sosnowski) poses high risk to humans, causing photosensitivity and severe sunburns following contact with skin.
- A comprehensive list of all plant species that occur within a region;
A region’s flora can be characterized according to the:
- number of species
- proportion of different taxonomical groups
- distribution or proportion of different geoelements
- number of native species or proportion of non-native elements
Based on these features, the flora of two regions can be more or less similar, and can also be very different from other region’s flora. The floristic map of the World was drawn based on these existing similarities and/ or differences, which outlined well-defined floristic units of various sizes.
The floristic units of the worlds’ flora, in decreasing order of geographic scale, are:
- floristic kingdom: continent-sized
- floristic region
- floristic province
- floristic district (and floristic districts can be further divided in even smaller units)
We will examine later on some examples.
The present-day floristic map of the World is the result of a long-term evolutionary history of biota during geological times.
The earliest concept that provided support for present-day distribution of biota was Wegener’s theory (1912) on continental drift. Building upon it, the theory of global plate tectonics formulated between 1967-1970 explains better the historical continental rearrangements. The continental shields floating on the surface of the liquid magma of the Earth is composed of large tectonic plates. These plates are being moved by convectional streams produced inside the magma. When the tectonic plates move, they either slide apart or slide over each other,e.g. the floor of the Atlantic Ocean enlarges because the American shields diverge from the Eurasian and the African shields; the Pacific plate is being crushed under the Eurasian plate. Such events are accompanied by violent earthquakes and volcanic activities,as we all know for example in the Japanese archipelago or the Philippines.
Fig. 21: A: The Earth’s continental shields and B: cross section of the Earth http://www.geography.learnontheinternet.co.uk/topics/structureofearth.htm
Around 230 million years ago, there was one old supercontinent on Earth called Pangaea, which was surrounded by the ancient Thethys Ocean. Following the break-up of Pangaea, continents gradually diverged from each other, which forced their fauna and flora to evolve independently. The more ancient the split between two continents, the more different their fauna and flora is. E.g., Australia and Antarctica separated first from Pangaea (about 195 million years ago). Antarctica migrated towards the poles and life on the continent went extinct, while the flora and fauna of Australia became isolated from the rest of the Earth’ biota. As a result, a large number of taxonomic groups are unique to Australia and do not occur elsewhere, or are only represented by distant relatives on other continents, such as the Eucalyptus species, some Protea species and the marsupial mammals.
In contrast, members of the Rose family as well as placental mammals completely lack from Australia. These groups developed after the Australian and Antarctic continents separated. A similar split occurred between South America and the African continent, which also have very different biota. The last unbroken part of Pangaea constitute today the continents of the Northern hemisphere. The Eurasian and North American continents were connected even during the last Ice Age, several tens of thousands years ago, when ice sheet covered the Bering Strait.
The largest floristic units have witnessed an evolutionary history of 100 000 years, when the natural history of the continents took shape. The largest floristic units are therefore the continental-scale floristic kingdoms.
Fig. 22. Floristic map of the Earth with the floristic realms
1. Holarctis: comprises the whole Northern Hemisphere of the Earth, with the Arctic, Boreal and Temperate regions (the extratropical regions), including the Mediterranean strip of North Africa.
Characteristic taxa: e.g. Pinaceae, Betulaceae, Fagaceae, Saxifragaceae, Caryophyllaceae, Ranunculaceae
2. Paleotropis: Africa South of Sahara, Madagascar, South-East Asia: India, South of the Jangtze river in China, Taiwan and the Indonesian islands.
Characteristic taxa: e.g. Orchidaceae, Araliaceae, Moraceae, Coffea, Oryza, Musa, Citrus
3. Neotropis: South and Central America, up to Florida
Characteristic taxa: e.g. Bromeliaceae, Rubiaceae (Hevea), some species of Orchidaceae (Vanilla), Solanaceae (Capsicum, Lycopersicon, Solanum tuberosum), Agave, Cactaceae, Phaseolus
4. Capensis: South Africa, the Cape Colony
Characteristic taxa: Proteaceae, Freesia, Acacia, Gerbera, Clivia, Pelargonium
5. Australis: the Mediterranean regions of Australia, the desert and Tasmania
Characteristic taxa: Eucalyptus, Acacia, Proteaceae (Banksia), Xanthorrhoeaceae
6. Antarctis: the Southern part of Chile, Patagonia, Fireland and South of Cook Strait in New Zealand
Characteristic taxa: Fitzroya, Nothofagus, Podocarpus, Agathis
The origin of economically important plants from different floristic kingdoms of the Earth has been described by many scientists. The hotspots of these plants correspond to the hotspots of natural flora diversity.
The Russian botanist and geneticist Nikolai Vavilov (1951) has defined eight gene centres, i.e., centers of diversity and origin of domesticated plants. Later, Zsukovszkij and the Hungarian Rajmund Rapaics delimited further three gene centers: the European-Siberian, the Northern-American and the Australian centers.
Fig. 23. Vavilov’s gene centers according (see also the pdf. file annexed)
The European flora belongs to the Holarctic floristic kingdom and includes several floristic regions.
Fig. 24. Map of the main floristic regions of Europe
The present-day flora of the Carpathian Basin developed during the last ten thousands of years, when the last Ice Age (the Pleistocene) came to end and the Holocene warming began. The history of biota can be disentangled with the help of palynology (the analysis of pollen composition), and using geological and paleobotanical evidences. Bálint Zólyomi, Magda Járainé Komlódi, Pal Sümegi have, among others, brought substantial contributions to the Hungarian palynological science.
10. 9. 1. The post-glacial vegetation history of the Carpathian Basin
The postglacial vegetation history of the Carpathian Basin is the story of the last 13.000-10.000 years, driven by the warming climate following the end of the ice age. This type of long-term, non-repeatable vegetation history is termed secular succession.
The vegetation of the Carpathian basin at the end of the Pleistocene and beginning of the Holocene was extremely varied. The Transdanubian and Middle Mountains were covered by taiga forests, and the lowland (Mezőföld, Hortobágy and the region between the Duna and Tisza rivers) was dominated by herbaceous cold steppe vegetation.
During the Holocene warming, 5 more important climatic cycles followed each other:
1. Preboreal age: Pine-birch age11000-9000 B.C. (equivalent to the ancient Stone Age)
- characterized by a mosaic of cold taiga with pine and birch forests and cold hardy steppe-like vegetation in some parts of the Great Plain (Hortobágy, Mezőföld, Duna-Tisza köze region)
2. Boreal age: Hazel age 9000-7500 B.C. (equivalent to the Mesolithic Age)
- the warming started but the climate remained cold and mostly continental. Hazel scrublands and broadleaf forests covered most of Hungary, the oak began migrating in.
- xerothermic (adapted to dry and hot climate) continental species immigrated to the Great Plain from the Sarmatian region
e.g. Salvia nutans, Adonis volgensis, Crambe tataria
Fig. 25: Salvia nutans (A), Crambe tataria (B)
3. Atlantic age: age of mixed oak forests 7500-5000 B.C. (equivalent to the Neolithic Age)
- characterized by a warmer, more moderate climate; closed broadleaf forests developed;
- oak forests were dominant and gallery forests followed the larger rivers
- humans started exerting a strong impact on the natural environment through deforestation and agricultural activities.
4. Subboreal age: Beech I. age 5000-2500 B.C. (equivalent to the Bronze Age, and the early Iron Age)
- the climate turned cold again, the beech forests retracted to low elevations, reaching the Great Plain
5. Subatlantic age: Beech II. age 2500 B.C. –present (between the Iron Age and present)
- the climate became more continental and the amount of the yearly rainfall decreased
- the beech forests disappearedcontracted from the lowlands and hillsides towards higher elevations in the middle mountains.
In this period the formation of the present vegetation belts of East-Central Europe began:
1. forest- steppe (lowland)
2. broadleaf (deciduous) forests (Middle Mts. and the Transdanubian region)
10. 9. 2. Floristic subdivisions of the Hungarian flora
Fig. 26. The floristic map of Hungary
The floristic positioning of the Carpathian Basin,:
— Holarctis — Northern Temperate Floristic Kingdom
— Europa Centralis — Central European Floristic region
A; Pannonicum — the Pannonian Floristic Province
This floristic province together with its floristic regions covers almost the entire region of Hungary.
I. The Matricum floristic district — the Northern Middle Mountains: From Sátor Mountains (Zemplén Mountains) to Gödöllő Hills and Nagymaros-Visegrád Mountains.
Small floristical units:
- Tokajense — the southern part of Sátor Mountains (Zemplén Mountains), the Tokaj Mountains-Hegyalja foothills
- Tornense — the Torna Karst : From Aggtelek Mountains to Boldva valley, Gömör Hills and Cserehát at Abaúj.
- Borsodense — the Bükk Mountain at Borsod, between Sajó and Tarna valley, and Szarvaskõ.
- Agriense — the andesite mountains of Mátra, the sandstone mountains of Medves and the basalt mountains of Karancs, stretching to the valley of Zagyva.
- Neogradense — the hillside of Cserhát, Gödöllő Hills and the andesite Börzsöny Mountain, except its Southern part.
- Visegradense — the andesite mountains of Nagymaros, Visegrád and Szentendre and the limestone-dolomite-sandstone mountain of Naszály which shows some likeness to the Pilisense floristic unit.
II. Bakonyicum floristic district — the Transdanubian Middle Mountains: from Naszály to Keszthely Mountains
Small floristical units:
1. Pilisense — The limestone and dolomite Pilis, Gerecse and Buda Mountains, and the sandstone Hárshegy Mountain.
2. Vesprimense — The Vértes and Velence Mountains and Bakony Mountains towards Sümeg including isolated mountains of Vas county (e.g. Ság) and the hills of Pannonhalma.
3. Balatonicum — The Balaton Uplands from the isolated mountains of Fejér county to Keszthely Mountains and the isolated mountains of Tapolca Basin and Somló.
III. Praenoricum floristic district — Western Transdanubia (the the Alpine foothills) from Sopron and Lajta Mountains to Göcsej.
Small floristical units:
1. Laitaicum— Lajta Mountains , the hills of Fertő region in on Hungarian side, near Sopron.
2. Castriferreicum — The Alpine foothills from Sopron to Őrség, Vas ridge and Kemenes hills.
3. Petovicum —Southern Styria, stretching to Nortwestern Zala (Göcsej: from Lenti to Zalalövõ).
IV. Eupannonicum floristic district — the Little and Great Hungarian Plains
Small floristical units:
1. Arrabonicum — the Little Hungarian Plain : the Hanság, Szigetköz, Fertő-region, Sokoróalja and the sand land of Győr–Dereg.
2. Colocense — the loess cultural landscape of Mezõföld, the Danube valley (the plain of Solt) and the Turjánregion. The sand-dunes of Németkér and Vajta belong to the Praematricum.
3. Praematricum — the Danube–Tisza interfluve : inland sand-dunes mainly with calcareous, rarely acidic soils; alkali lakes and swamps with high salt content in the Great Plain; the sandy region of Tiszazug and Cserkeszőlő.
4. Crisicum — the Transtisza region (Körös-region): the interfluves of Sajó–Zagyva, the Jászság and Heves-Borsod ridges, Hortobágy, Nagykunság, Körös-region, Sárrét, the Tisza and Maros valleys, the loess ridge of Békés and Csanád and the Northern part of Bánság.
5. Nyírségense — The sandy area of Nyírség, ± acidic or slightly calcareous soils, birch swamps.
6. Samicum — the Northern Great Plain (Bodrogköz, Rétköz, Tiszahát, the Szatmár Plain and Érmellék).
7. Titelicum — the Southern Great Plain : the Dráva plain, the island of Mohács and beyond the country borders Bácska, Slavonia and the plains of Southern Bánság.
V. Praeillyricum floristic district — Southern Transdanubia
Small floristical units:
1. Saladiense — the Zala hills from the Eastern-Southeastern Zala to Csurgó, Balaton, Keszthely Mountains, Bakony foothills, the Little Plain and the Tapolca Basin.
2. Somogyicum — the sandy area of Inner Somogy with Nagyberek at Balaton, the Marcal ridge and Zselic. The last one is a transition zone to the Sopianicum floristic unit
3. Kaposense — Outer Somogy , loesscultural landscape, with characteristics of the Great Plain.
4. Sopianicum — The Mecsek with its isolated mountains and the TolnaRidge.
Three other floristic provinces bordering Hungary cross the country borders.
B. Illyricum — Western Balkan floristic province
I. Croaticum — Dinaric Alps floristic district
II. Slavonicum — the low elevation mountains between Drava and Sava rivers; in Hungary, they the hills along the Drava valley from Csurgó and Zákány to Őrtilos, the Villányi Mountains and Szársomlyó.
C. Alpicum — Eastern Alps floristic province
I. Noricum floristic district
1. Ceticum — the Sopron and Kőszeg Mountains,Vashegy at Felsőcsatár.
2. Stiriacum — the Hungarian side of the Vend region, Southwest to Szentgotthárd.
D. Carpaticum occidentale — Western Carpathian floristic province
I. Praecarpaticum — Northern Carpathian floristic district
1. Cassovicum — the Kassa region; in Hungary Telkibánya and the Füzér region in Northern Sátor (Zemplén) Mountains.
- Kárpáti Z., Terpó A. 1971: Alkalmazott növényföldrajz. Mezőgazdasági kiadó. Budapest.
- Simon T. (szerk.) 2001: Növényföldrajz, Társulástan és Ökológia. Tankönyvkiadó, Budapest.
- Turcsányi G. (szerk.) 1998: Mezőgazdasági növénytan. Mezőgazdasági Szaktudás Kiadó, Budapest. 2. kiadás
- Járainé Komlódi M. (főszerk.) 2007: Pannon enciklopédia. Urbis Könyvkiadó. Budapest.
- Jakab G 2011: Negyedidőszaki makrobotanika. Geolitera. Szeged.
Az "Angol és magyar nyelvű, digitális tananyagok fejlesztése a BCE kertészettudományi kar kertészmérnök és multiple degree hallgatói számára" pályázat a TÁMOP-4.1.2.A/1-11/1-2011-0028 pályázati projektek támogatásával készült.