PHYTOGEOGRAPHY


In S.W. Asia phytogeographical regions are so striking that an attempt has been made to assign as many species as possible to phytogeographical elements (cf. p. 14). As previously stated, Turkey is the meeting ground of three phytogeographical regions: Euro-Siberian, Mediterranean and Irano-Turanian. In the field these can usually be readily recognised by their different vegetational aspects, reflecting differences in climate; but they are more fundamentally, if less obviously, based on floristic differences, including endemism. Accounts of the concepts and methods used in the recognition and delimitation of phytogeographical regions in Europe and the Near East are given by Braun-Blanquet, Eig, and Zohary. [*Cf. M. Zohary, 1962, 1963, for discussion and further references. (See p. 26.)]

These concepts, with some modification, have been adopted by the editor. It should be made clear, however, that areas have been referred to particular phytogeographical divisions on the basis of their present-day vegetation and floristic composition, not on an inferred 'historical' climax no longer present in the area. The editor has not committed himself to mapping the boundaries of the regions in Turkey. Our knowledge of the vegetation is still very incomplete, and, as floristic analysis is at the root of phytogeographical division, we shall obviously be in a better position to map phytogeographical areas when the Flora is completed than we are now. Nevertheless, although there is considerable difference of opinion about the limits of the regions, and the treatment of enclaves and transitional areas, the editor has little doubt about the reality of the three phytogeographical regions and their nuclei. Their approximate boundaries will be outlined in the following pages with reference to Map 2, and a short account given of their vegetation and floristic characteristics.

The nomenclature of some of the species mentioned in this account may need some alteration when their genera come to be revised for the Flora. It should be pointed out that the spot maps referred to below show the general distribution of a species but convey little (if any) idea of the relative abundance of the species in different areas. There is a tendency for common plants to be less collected than rare ones, so that only field observations can provide information on abundance or dominance; it would be helpful if collectors would include such information on their herbarium labels. When a plant is referred to as characteristic of a particular region in Turkey, it should not be assumed that it is confined to it.

The account of the phytogeographical areas is largely based on the field observations of the editor, supplemented by distributional information gleaned from Volume 1, and by information given in the works cited on pp. 25-26.


Euro-Siberian Region

This territory extends along most of North Anatolia (approximating to the area so-named on Map 2) and in a narrower strip along the Black Sea coast of Turkey -in-Europe (Istranca Daglari to the Belgrad forest near the Bosporus). The relatively humid climate puts little strain on the plant and is reflected in the predominantly mesophytic vegetation, deciduous forest being the usual climax in the middle and lower regions; phanerophytes and hemicryptophytes are abundant in the region. There is, however, an interrupted chain of Mediterranean enclaves (mainly consisting of sclerophyll scrub) that extends along much of the [16] Black Sea coast; such Mediterranean enclaves are poorly developed in Lazistan (cf. p. 20).

In Anatolia the whole of the Euro-Siberian belt is probably best referred to the Euxine province, an area which covers much of Georgia and the Caucasus, the Istranca mountains of European Turkey and south-east Bulgaria, and, in a modified form, probably the mountainous part of the Crimea. Excluding the Crimea, the province falls into two distinct parts: an eastern part comprising the Caucasus, Colchis (Western Georgia), and Lazistan; and a western, generally drier, part consisting of Paphlagonia, Bithynia, and the Istranca range.

In Turkey-in-Europe one can safely say that the Istranca area belongs to the Euxine province. Destruction of the vegetation and lack of information make it very difficult to decide to which phytogeographical areas most of the rest of Turkish Thrace should be referred. Many of the typical Euxine species are apparently absent from the wooded area that covers the hills that stretch from Tekir Daglari towards the Ergene river. It seems quite possible that this area belongs to the Balkan province of the Euro-Siberian region, and is floristically connected with Greek Thrace. The plains of the central area are largely cultivated, but have a steppe-like aspect. They are certainly not Irano-Turanian, and may represent an extension of the Sarmatian province of the Euro-Siberian region, which is well developed in the Danube valley. European Turkey needs attention by the collector, taxonomist, and ecologist, in order that its relationships can be firmly established - and urgently, before its native vegetation is further destroyed.

Most of the Euxine province below the tree line is covered with forest, or by scrub where the forest has been destroyed. In the lower parts the forest is mainly deciduous, often associated with evergreen shrubs, but in the higher parts conifers increase or even predominate. The following species may be considered characteristic for much of the Euxine province below the tree line. Trees and shrubs:

Abies nordmanniana (Map 5), Acer campestre, A. trautvetteri, Ainus glutinosa, Buxus sempervirens, Carpinus betulus, C. orientalis, Castanea sativa, Cornus australis, Corylus avellana, C. colurna, Crataegus microphylla, Fagus orientalis (frequently dominant). Daphne pontica, Fraxinus excelsior, Hedera colchica, Hypericum androsaemum, Laurocerasus officinalis, Mespilus germanica, Ostrya carpinifolia, Pinus nigra subsp. pallasiana (Map 8), P. sylvestris (Map 7), Quercus petraea- Q. dschorocensis group, Q. hartwissiana, Q. pedunculiflora, Rhododendronluteum, R. ponticum, Smilax excelsa, Sorbus torminalis, Staphylea pinnata, Tilia rubra. Herbs: Argyrolobium calycinum, Astrantia maxima, Calamintha grandiflora, Cardamine bulbifera, Circaea lutetiana, Epimedium pubigerum, Euphorbia amygdaloides, Galium odoratum, Helleborus orientalis (Map 18), Lathyrus aureus, Pyrola spp.. Ranunculus brutius, Salvia glutinosa, S. forskahlei, Trachystemon orientate, Valeriana alliariifolia.

In the eastern part (Lazistan) many species occur that do not extend further west. These include the following trees and shrubs: Acer cappadocicum, Alnus barbata, Betula medwedewii. Daphne glomerata, Diospyros lotus, Phillyrea decora, Picea excelsa (Map 6), Quercus pontica, Rhamnus imeritanus. Rhododendron caucasicum, R. smirnovii, R. ungernii, Sorbus subfusca. Herbs: Chamaesciadium acaule (monotypic), Draba hispida. Geranium psilostemon, Hypericum bupleuroides, Lathyrus roseus, Lilium ponticum, Lycopodium spp., Pachyphragma macrophyllum (monotypic), Papaver lateritium. Primula cortusifolia. The alpine flora [17] is closely connected with that of the Caucasus, and shows little floristic connection with the rest of Turkey.

The western part of the Euxine province (Paphlagonia and Bithynia), shows a progressive decrease in the Euxine species concentrated in Lazistan and West Caucasia, and at the same time an infiltration of species centred in the Balkans or Central Europe, such as Clematis viticella (Map 23), Lilium martagon, Tilia tomentosa and Quercus frainetto. The alpine flora shows many links with the Irano-Turanian region and even with the oreo-Mediterranean flora of Turkey.

It has been pointed out by Czeczott (1932) that several characteristic species of the Euxine province are absent from Paphlagonia (cf. Polystichum lonchitis, Map 4; Actaea spicata. Map 23). More collecting is needed to confirm this distributional gap.

Enclaves of Euxine vegetation occur further south - on Kaz Dagi (Mysia) and Murad Dag (Phrygia); some Euro-Siberian species reach the Anti-Taurus and many more the Amanus (for example, Cardamine lazica. Map 26); others penetrate as far south as the Lebanon.

Characteristic crops of the Euxine province are Zea mays, Corylus asellana and C. maxima. Camellia sinensis. Citrus fruits and Diospyros lotus are cultivated in Lazistan. In Lazistan the Euxine province is rather abruptly demarcated from the Irano-Turanian region to the south. Further west this is not the case. The intermediate belt, much of it covered by Pinus nigra subsp. pallasiana forest (Map 8) or deciduous oak scrub, is relatively wide in Bithynia and Paphlagonia, and is complicated by the intrusion of Mediterranean vegetation into Bithynia with elements penetrating still further east. Only field studies can elucidate the position here.

In North-West Anatolia, where Mediterranean and Euxine vegetation integrade, considerable areas are now dominated by Paliurus aculeatus.

The connections between the western part of the Euxine province and Europe have already been mentioned. The flora of Lazistan is intimately connected with that of adjacent western Georgia, including the western half of the main Caucasus range, particularly Abkhazya (all parts of the Euxine province). Endemism is high in this area as a whole, and many of the endemics hold isolated taxonomic positions. The flora of the montane coniferous forests and the alpine zone show particularly close links with the same zones in the Caucasus and even with the mountains of Central Europe (cf. Polygala alpestris. Map 30).

The Euxine province (particularly the eastern part) more closely resembles the Hyrcanian province of North Iran and adjacent Talysh than any other part of the Euro-Siberian region. The two areas (separated by the lower Aras valley) are often treated as a single province (the Hyrcano-Euxine province), and share many leading species, such as Acer cappadocicum, Fagus orientalis, Laurocerasus officinalis and Petrocarya fraxinifolia. On the other hand the Hyrcanian forests contain several deciduous trees (for example, Parrotia persica, Ainus subcordata, Gleditschia caspica) absent from the Euxine forests, and lack some important evergreen genera (for example, Abies, Pinus, Picea, Rhododendron) that are characteristic of the Euxine. Each of these two areas contains numerous endemics.


Mediterranean Region

This very natural region is adopted in its narrow sense to cover all areas bordering [18] the Mediterranean with the exception of some arid parts of North Africa. It reaches its eastern limits in the Anti-Taurus, Amanus and Lebanon. In Turkey all the Mediterranean vegetation belongs to the East Mediterranean province, which is treated here as extending from the eastern half of Italy to the Lebanon. A large number of geophytes, therophytes and suffrutescent chamaephytes is characteristic of the region, although sclerophyll vegetation dominates the landscape.

In Turkey-in-Europe the Mediterranean territory is surprisingly small, being confined to the southern part, mainly the Gallipoli peninsula. In Anatolia it covers most of what we have delimited as West and South Anatolia (cf. Map 2) and extends as a series of enclaves along much of the Black Sea coast. In West Anatolia the transition between the Mediterranean, Euro-Siberian, and Irano- Turanian regions is gradual, but in South Anatolia (where only Mediterranean and Irano-Turanian regions are involved) relatively abrupt.

Although West and South Anatolia share many leading species, there are considerable floristic differences; these are most noticeable at higher altitudes and in regard to endemism, which is considerably higher in the Taurus than it is in the West. In South Anatolia, the flora of the Amanus differs markedly from that of the drier Cilician Taurus and Anti-Taurus.

Macchie, dominated by evergreen shrubs, covers much of the Mediterranean territory of Turkey below 1000 or 1200 m. On deeper soils, or where there has been less interference with the natural climax, forests prevail. The leading woody species include Arbutus andrachne, Calycotome villosa, Carpinus betulus, Celtis australis, Ceratonia siliqua, Cistus creticus (Map 28), C. salviifolius (Map 29), Cotinus coggyria. Daphne sericea. Erica verticillata, Fontanesia phillyreoides (South Anatolia), Juniperus oxycedrus (Map 14), Laurus nobilis, Myrtus communis, Olea europaea var. oleaster, Phillyrea media, Pistacia lentiscus, P. terebinthus (incl. P. palaestina), Pinus brutia (Map 9; often a dominant forest tree), P. pinea (Map 10), Rhamnus palaestinus, Quercus cocci f era (Q. calliprinos), Q. aegilops, Q. haas, Q. libani, Q. infectoria subsp. boissieri, Smilax aspera and Styrax officinalis. A remarkable forest of Liquidambar orientalis covers the flood plain near Koycegiz in Caria.

In many places the macchie has been degraded and replaced by phrygana.

Here Cistus creticus (Map 28), C. salviifolius (Map 29), Lavandula stoechas, Poterium spinosum and Thymbra spicata are often among the leading species. By the sides of streams or in dried-up river beds the following are characteristic: Ainus orientalis (South and South-West Anatolia), Nerium oleander, Platanus orientalis, Vitex agnus-castus, and the lianas Vitis sylvestris and Smilax excelsa. Above 1000 or 1200 m (corresponding to sporadic winter snowline) the Mediterranean region is largely dominated by conifers. Pinus nigra subsp. pallasiana (having its maximum extent in North-West Anatolia) is the most widespread species, but in the Taurus its place is often taken by Cedrus libani (Map 6) and Abies cilicica (Map 5). In Turkey, native Cupressus sempervirens (Map 11) and Juniperus drupacea (Map 12) are confined to the mountains of South Anatolia; Juniperus excelsa (Map 17) often forms the tree-line. In North-West Anatolia, Pinus nigra descends lower than it does in the Taurus, and is often accompanied by Cistus laurifolius (Map 28) which dominates considerable areas after the pine has been destroyed by fire. This Cistus (otherwise a West Mediterranean plant) is characteristic of a considerable area of North and West Anatolia in territory [19] that is apparently transitional between the Mediterranean and Euro-Siberian or Irano-Turanian regions.

In areas transitional between the Mediterranean and Irano-Turanian regions, Pinus nigra is particularly abundant, and encloses the plateau on its north, west, and south sides. Quercus infectoria subsp. boissieri is often a leading plant in the scrub belt. Other species characteristic of this intermediate territory are Pistacia atlantica and several woody Rosaceae (including Pyrus elaeagrifolia, Prunus microcarpa and Amygdalus orientalis). In the Anti-Taurus the Mediterranean and Irano-Turanian communities are inter-digitated rather than blended, apparently being dependent on the exposure of the slope.

Above the tree-line (at c. 1700 m in the Taurus) the mountains of the Mediterranean region are usually dominated by various cushion communities in which spiny species of Astragalus, Acantholimon, and Onobrychis cornuta often play a leading part and afford protection for weaker associates. Much of this flora is probably of Irano-Turanian origin.

The Mediterranean enclaves on the Black Sea coast (cf. p. 16) contain the following woody species: Arbutus andrachne, A. unedo, Cercis siliquastrum (Bithynia), Cistus creticus (Map 28), C. salviifolius (Map 29), Erica arborea, Jasminum fruticans, Myrtus communis, Laurus nobilis, Pinus brutia (Map 9), P. pinea (nr. Trabzon and Borcka), Pistacia terebinthus (incl. P. palaestina), Phillyrea media, Platanus orientalis (by streams), Quercus ilex, Spartium junceum and Vitex agnus-castus. The belt is a narrow one, and occurs intermittently from sea level to 200 or 300 m, particularly on thin soils and southern exposures. Some intermingling of Mediterranean and Euxine elements occurs. These coastal Mediterranean enclaves are lacking in many species that are characteristic of macchie or phrygana in West Anatolia, presumably due, at least in part, to colder winters. On the other hand. Erica arborea and Quercus ilex are commoner in the Mediterranean enclaves of the Black Sea coast than they are in West or South Anatolia; they are, in fact, more abundant in the West Mediterranean province than they are in the East, and extend into Atlantic Europe which the Black Sea coast climatically resembles.

Some Mediterranean elements occur in sheltered habitats on the south side of the North Anatolian range, and may even form local, impoverished enclaves.

Many of these elements are common in the north coast enclaves. There are, however, a few other species (for example, Cedrus libani, Fontanesia phillyreoides, Quercus coccifera) that occur elsewhere in Anatolia only in the South and West.

Crops grown in the Mediterranean region of Turkey include Olea europaea, Vitis vinifera (mainly in the West), Triticum spp., Citrus fruits (mainly south-west and south), Ficus carica. Legumes (Cicer, Lens, Medicago, Phaseolus, Pisum, Vicia, etc.), Gossypium in the West and on the Cilician plain, and Musa on the south coast.

Many authors have included the whole of North Anatolia in the Mediterranean region. To the editor this seems unjustified. The coastal Mediterranean vegetation is better treated as enclavic in the Euxine Province. It is atypical, impoverished, often invasive and almost lacking in endemic species, suggesting that it may be of relatively recent origin in this area; destruction of Euxine forest appears to have greatly facilitated its spread.

Some infiltration of Mediterranean elements extends to the lower parts of [20] South-East Anatolia, but this is not so pronounced as it is in Iraqi Kurdistan and in West Iran.

The Mediterranean flora of West Anatolia shows many affinities with that of the East Aegean islands and even (especially at higher altitudes) with the Greek mainland. Further exploration of the West Anatolian peninsulas will no doubt add more Aegean species to the Turkish list. South-West Anatolia (particularly the Marmaris peninsula and Lycia) show relationships with the southern islands of the Dodecanese (for example, Rodhos, Karpathos) and with Crete, besides, of course, being intimately connected with the main Taurus range to the east.

The flora of the Amanus is related on the one hand with the Cilician Taurus and Anti-Taurus, on the other with the coastal ranges of Latakia and Lebanon.

Numerous Euro-Siberian elements in the Amanus demonstrate its connections with the Euxine province. The links between the South Anatolian flora and Cyprus are much less pronounced than might be expected from the proximity of the two areas. There are close floristic connections between the Mediterranean and Irano-Turanian regions.


Irano-Turanian Region

This region is by far the largest of the three in Turkey, and, apart from a few enclaves, is confined to Central and East Anatolia (cf. Erysimum crassipes. Map 27). Although so large and rich in herbaceous and sufiruticose species, it is far less well understood than the Mediterranean and Euro-Siberian regions, largely due to the difficulties of identification in several genera which play an important part in its vegetation. These include Acantholimon, Artemisia, Astragalus, Bromus, Rosaceae (Pomoideae, Prunoideae), Quercus and Stipa. Until these genera are revised and the distribution of their species becomes better known, it is difficult to name even the dominants in some widespread communities.

In the broad sense the Irano-Turanian region extends from Inner Anatolia southwards to Palestine and eastwards to Mongolia. According to the division of Zohary (1963), Inner Anatolia belongs to the Irano-Turanian subregion; this area includes much of the Syrian Desert, North Iraq (Kurdistan), most of Iran (except for the Caspian strip and southernmost area), most of W. Pakistan, most of Afghanistan, part of the Aralo-Caspian deserts (the Turanian province), and the western Tien Shan. The subregion also includes the Hauts Plateaux of North Africa (Mauritanian province), which lie between the Irano-Turanian and Saharo-Arabian regions.

Zohary refers Inner Anatolia to two separate provinces of the Irano-Turanian subregion: the Mesopotamian province, which includes Turkish Mesopotamia and much of the Syrian Desert; and the Irano-Anatolian province, which covers the rest of Inner Anatolia and the Irano-Turanian parts of Iran and Afghanistan.

This seems to the editor to be too heterogeneous an area to be treated as a single province, but we must defer any decision on the subdivision of this very interesting region until more information is available. It should be pointed out, however, that Anatolia is on the periphery of the Irano-Turanian region, and that in this area it meets and intergrades with the two adjacent regions. Many of the genera so characteristic of the region further east (Acantholimon, Acanthophyllum, Calligonum, Cousinia, Ferula, Eremostachys, etc.) have relatively few species in Anatolia; on the other hand, there are other groups which have their main centres of development in the Irano-Turanian part of Anatolia (for example, Achillea[21] Santolinoideae, Aethionema, Alyssum Sect. Gamosepalum, Isatis Sect. Isatis, the pinnate-leaved group in Salvia Sect. Eusphace). As already mentioned (p. 21), the floristic links between this region and the Mediterranean region are very close in Anatolia.

Except at high altitudes, the Irano-Turanian region in Turkey divides into two major vegetational areas: (1) a wide peripheral area of deciduous scrub and even park-like forest, which, according to Louis (1939) and Walter (1956b), represents an originally forested area; (2) inner areas of treeless, 'true' steppe, of which the largest is the Central Anatolian steppe, centred on Tuz Gold but extending, with some interruptions, as far as Eskisehir, Karaman, Nigde, Sivas, and Cankiri; other areas of treeless steppe occupy much of Mesopotamia and several parts of East Anatolia, notably round Malatya, from Erzincan to Erzurum, from Gümüsane to Bayburt, and from Lake Van to the Aras valley. 'Mountain steppe' occurs in the highlands of East Anatolia.

In the treeless steppes of Central Anatolia, Artemisia fragrans is often a leading species, replaced in some areas by the closely allied A. spicata. Characteristic species are Achillea santolina, Euphorbia tinctoria, Globularia orientalis, Isatis glauca (sensu lato), Linum hirsutum subsp. anatolicum, Moltkia aurea, Noaea mucronata, Peganum harmala, Phlomis armeniaca, Poa bulbosa, Teucrium orientate, and several species of Stipa and spiny Astragalus. Other species are endemic to the area (for example, Consolida stenocarpa (Map 20) and Delphinium venulosum (Map 19)).

Round Tuz Gölü the halophytic vegetation includes many Chenopodiaceae of Iranian or Trans-Caspian affinities, and several distinctive endemics in genera which are not usually associated with halophytic habitats.

The composition of the steppe of East Anatolia is even less known than that of the Centre. Artemisia fragrans is still a leading species, although other species occur, such as A. orientalis and A. araratica; the genus flowers in late summer and needs extensive collecting. In the mountains, Bromus tomentellus is widespread; spiny cushion plants increase with altitude, mainly Astragalus subgen. Tragacantha, Acantholimon spp. and Onobrychis cornuta. In Kurdistan the mountain steppe is often dominated by giant Umbelliferae (Ferula, Prangos, etc.). As elsewhere in Inner Anatolia, many of the commonest perennials in the steppe are unpalatable to grazing animals.

On the Armenian Highlands (North Armenia) a different type of steppe occurs in which Stipa plays an important role and spiny cushion plants are poorly represented. Although predominantly Irano-Turanian in its present-day floristic composition, this area (which extends from Erzurum to Lake Sevan in Soviet Armenia) is believed by Walter (1956a) to have supported Euro-Siberian vegetation in historical times; even today, forests of Pinus sylvestris (Map 7) occur in favourable habitats (for example, near Sarikamis), and Euxine species of Delphinium and Aconitum grow near streams.

The lower, milder 'outer' plateau of Mesopotamia may be the only part of Turkey where Artemisia herba-alba grows - a dominant species in the Syrian Desert and one which extends westwards throughout the Mauretanian province.

Some species characteristic of Mesopotamia include: Consolida tomentosa subsp. oligantha (Map 22), Eryngium noeanum, Hypericum laeve, Papaver stylatum (Map 24), Phlomis kurdica and Salvia spinosa. Most of the plains are now extensively cultivated, so that it is difficult to get an idea of their natural vegetation. [22] We have already referred to the broad girdle of Pinus nigra subsp. pallasiana forest which borders Central Anatolia in the north, west and south. Where this forest meets the oak scrub, which is the most abundant type of vegetation on the periphery of the Central Anatolian steppes, it is usually associated with a largely Irano-Turanian ground flora. This Irano-Turanian scrub is best developed in the north and west. The leading woody species are often Quercus pubescens (especially in the north and west), Q. infectoria sensu lato (mainly in the west and south) and Q. cerris. Frequent associates are Juniperus oxycedrus (Map 14), J. excelsa (Map 17) or foetidissima (Map 16), Pistacia atlantica (north and west), Berberis crataegina, and various woody Rosaceae (Pyrus elaeagrifolia, P. amygdaliformis, Prunus microcarpa, Amygdalus orientalis, Crataegus orientalis, etc.).

Most of East Anatolia that is not covered by steppe or mountain cushion communities is occupied by scrub or even park-like forest often dominated by juniper and deciduous oaks. Leading species include Juniperus excelsa, Quercus infectoria (mostly subsp. boissieri), Q. libani, Q. longipes (concentrated in the south-east), Q. persica subsp. brantii (southerly in its distribution) and Q. pubescens; Acer cinerascens, Pistacia khinjuk, Rhamnus kurdicus and Sorbus persica grow in Kurdistan. This woody vegetation has suffered much destruction by man.

As in much of the Irano-Turanian region, the arboreal vegetation in South and East Anatolia has both an upper and a lower (apparently natural) altitudinal limit, the lower presumably determined by aridity, the upper by exposure.

The following crops are characteristic of the Irano-Turanian region in Anatolia: Triticum and Hordeum spp., Papaver somniferum, Prunus armeniaca, Pyrus communis, Vitis vinifera (Central Anatolia), Citrullus and Cucumis spp. Enclaves of Irano-Turanian steppe vegetation (probably secondary) occur in the Mediterranean territory of West Anatolia, especially near Burdur, Denizli and in many parts of the vilayets of Kutahya and Usak. The intergradation between the Irano-Turanian vegetation and that of the Euro-Siberian and Mediterranean regions has already been mentioned under those sections.

In general, the Irano-Turanian flora in Turkey, with the exception of Mesopotamia, is probably most closely related to that of Transcaucasia, North-West and West Iran, and North Iraq. Many of the endemics are related to other Anatolian species, some of which are Irano-Turanian, others Mediterranean.

The relationship between the Irano-Turanian and Mediterranean flora is particularly close in the 'contact zone' of South and West Anatolia, and on the south- western slopes of the mountains which extend from Turkish Kurdistan south- eastwards to Siraz. This strip (which must be included in the Irano-Turanian region) is roughly demarcated by such endemic 'Kurdo-Luric' species as Nepeta macrosiphon and Phlomis bruguieri. Within this area the Mediterranean element seems to be more strongly represented in North Iraq and West Iran than it is in South-East Anatolia, but the vilayets of Siirt and Hakkari require more exploration before this can be verified. The flora of Turkish Mesopotamia is much closer to that of the northern part of the Syrian Desert.

The steppe of the Armenian Highlands has already been referred to. This area, despite its overall Irano-Turanian character and connections with the Euxine flora, shows some interesting links with the European steppes (or forest-steppes) of the Sarmatian (including Pontic) province, which, on floristic grounds, is best assigned to the Euro-Siberian region. These Sarmatian elements include Salvia [23] nemorosa. Adonis wolgensis and Echium rubrum. Other Sarmatian elements are scattered in other parts of the Irano-Turanian region of Turkey, but seem to have passed almost unrecognised.


Phytogeographical elements

An element, as it is used in this Flora, is a taxon confined or centred in one phytogeographical region (or province). A species whose maximum abundance is clearly in one region is treated as an element of that region. Such an element may occur locally in communities of another region, or in association with other elements of its own region, forming enclaves isolated from the main area. In the Euxine province, for instance, if we accept a broad interpretation of it as approximating to North Anatolia on Map 2, there is a narrow, interrupted strip of Mediterranean enclaves along the Black Sea coast. In this area, a plant growing at Sinop will probably be a Mediterranean element, even though it is growing within the general area of the Euxine province of the Euro-Siberian region.

Similarly, isolated Euxine elements occur in the Amanus, which is part of the Mediterranean region.

Acute problems are posed by high mountain plants, particularly in the Taurus; here an Irano-Turanian flora appears to have penetrated from the interior and mingled with an alpine flora that may well be of 'oreo-Mediterranean' origin.

Phytosociological studies may be able to settle this issue; the editor has usually refrained from treating such plants as elements of a particular region, even though the area in which they grow (at least in the Taurus and West Anatolia) is included in the Mediterranean region.

Numerous species are not cited as regional elements for other reasons: because they are widely distributed in two or more regions; because they are apparently limited to transitional areas; and because information is inadequate. Endemics are treated as particular elements on the basis of the vegetation in which they grow, not on considerations concerning their probable evolutionary history.

The concept of phytogeographical elements used in the Flora is a broad one, but that is consistent with our uneven state of knowledge. A more detailed analysis would not be possible without much additional information. This is a first and no doubt faltering step to set the Turkish flora in the broader framework of phytogeographical relationships in Europe and South-West Asia.


Endemism

A survey of the 902 species accepted in Volume 1 shows that about 20 per cent of the species are endemic to Turkey. When subspecies or varieties are considered, the percentage rises to about 24. About a third of these endemics are at present known from only one locality - generally in the mountains.

Endemics are scattered throughout the country, but are almost absent from Turkey-in-Europe. The largest number of endemics occurs in the Irano-Turanian region (in which the most widespread species are concentrated) and the Mediterranean region. Many genera well-developed in Anatolia (for example, Alyssum, Asyneuma, Alkanna, Rhamnus, Salvia, Sideritis, Verbascum) contain closely related endemics that 'replace' one another in different areas. The floristic links between the Mediterranean and Irano-Turanian regions are much closer than between the flora of either of these regions and the Euro-Siberian region. Many genera are well represented in both and have endemics in each region. A rather large [24] number of endemics, however, is apparently confined to areas where adjacent regions meet or intergrade - a feature which deserves further study.

The endemics show definite areas of concentration throughout the country, predominating in the mountainous parts of South and East Anatolia. The highest concentration is in the Cilician Taurus, followed by the Lycian Taurus and the mountains round Erzincan; Anatolian endemics are numerous, however, throughout the 'Anatolian Diagonal' (cf. Papaver triniifolium. Map 24); Kurdistan (Delphinium carduchorum (Map 19)), the Kastamonu-Cankiri area, and the Amanus. It should be borne in mind that this distributional pattern must be considered with two factors in mind: uneven collecting, and the fact that eastern-most Anatolia is a frontier zone which does not coincide with phytogeographical boundaries. There is no doubt, for instance, that endemism in Lazistan would be much higher if the area under review included the adjacent mountains of Georgia.

It should not be assumed that the pattern of endemism briefly described above will apply to the whole flora.


General Literature